ANIMALS

FARM

OF
NUTRITION

THE

"The

THE

MACMILLAN

NEW

YORK

COMPANY

BOSTON

DALLAS

CHICAGO

"

ATLANTA
"

SAN

MACMILLAN

FRANCISCO

"

LONDON

"

Limited

CO.,

BOMBAY

CALCUTTA

"
"

MELBOURNE

THE

MACMILLAN

CO.

TORONTO

OF

CANADA,

Ltd.

NUTRITION

THE

OF

FARM

ANIMALS

BY

HENRY

PRENTISS

DIRECTOR

OF

THE

PENNSYLVANIA

ARMSBY,

INSTITUTE

NUTRITION,

COLLEGE;

UNITED

EXPERT

FOREIGN

ACADEMY

OF
OF

THE

NUTRITION

MEMBER,
TURE
AGRICUL-

SWEDEN

COMPANY

MACMILLAN
1917
All

IN

DEPARTMENT

STATES

AGRICULTURE;
ROYAL

ANIMAL

OF

STATE

Ph.D.,

rights

reserved

OF
ANIMAL
OF

LL.D.
THE

Copyright,

By

Set

up

COMPANY.

MACMILLAN

THE

and

1917,

Published

electrotyped.

JUN

21

S.

Cushing
Norwood,

Co.

1917.

1917

Nortooofi
J.

June,

$regs
Berwick

"

"

Mass.,

U.S.A.

,0
"GI.A470024

Smith

Co.

PREFACE

The

animals

which

in

manner

presented

is

impart practical

of

of

principles
and

chief

and

of

But
of this

to

while

means

of

and

chemical

because

shall

which

the

Accordingly,

the

with

natural

the

and
seeks

with
of

be

which

points,

fullness

aim

expressed

in

almost

science

in

the
half

the

the

aspects

significance
convinced

that

training

greater
laid

be

the

upon

practice
and

importance

real

somewhat

underlie

collegiate

as

pline
disci-

well

as

deal

primarily

farm

animals,

to

animal

of
to

firm

dogmatic

mere

trie*

the

to

the

husbandry,
practice.

It

and,

statements,
of

citation

literature

even

evidence

experimental
of

the

limitations

opportunities

for

further

fundamental

the

discuss

is

feeding
persuasion

century

United

of

specific details

present

stock

of

of

indicatesomething

and

is

to

nutrition

field

complete

to

successful

based

the

attempts

broader

attempting

Its
upon

as

knowledge

present

results

business

intrinsic

made

the

governing

may

sufficient

liminary
pre-

course,

the

wisely

may

their

volume

present

far

important

study

student.

only secondarily

not

with

the

is

as

information

an

which

laws

to

the

so

regarded

believes

He

be

with

although
upon

thus

the

avoid

to

is nevertheless

receive

from

to

be

cording
ac-

purpose

stock, the

minimize

writer

of

may

laws

distinguished

as

live

to

to

desirable.

contribute

of

information

of

farm

differ

prime

agricultural colleges

our

account

on

subject

the

the

physiological

both

feeding,

related

sometimes

they

than

the

of

of

naturally

familiarity

disposed

is

nutrition

If

likely

nature

as

the

acquisition of practical skill.

of

procedure

emphasis

feeding

upon

subject,

students

the

different

the

of the

aspect

for

that

no

view.

is

laid

the

subject, and
by

will

the

experience, particularly

of the

student

the

of

be

will

stress

of

nutrition

partake

to

subject
in

end

skill in

to

the

the

to

ultimate

the

to

the

ago

consciously

of

by

the
the

States, Professor

truth
father
Samuel

vestigati
in-

principles
or

sciously
uncon-

so

pithily

of

tural
agriculWilliam

Vi

PREFACE

Johnson,

that,

advanced

and

master,

the

knows,

the

he

more

guide

to

training

of
a

Bulletin

of

part

Agriculture

and

Agriculture";

in

indebted
named
The

Carnegie

The

F.

"

A.

Institution

Davis

Company,

43,
The

John

44,

and

W.

B.

Company,

inary
prelim-

elementary

an

net

similar

values

for

of

the

sion
permisstuffs

compilations
He

Feeding."
for

in

use

feeding

extensive

and

of

of American

Morrison

and

XVIII,

Department

Cyclopedia

their

Feeds

Chapter

the

for
"

energy

upon
"

in

Secretary

of

use

"

College,
191

of

Figure

Washington,

is

the

wise
likecuts

Figure
2,

3,

24.

17.

6, 7, 8, 16, 19,

Figures

15,

4,

29,

20,

and

22.

31,

32,

^3"

45.
Saunders

Wiley

May,

Figures

Company,

Macmillan

The

State

than

Ginn

37,

of

degree

States

following publishers

the

to

is the

rather

Honorable

United

Henry

the

of

edition

fifteenth

the

The

to

Bailey's

Appendix

the

in

contained

certain

Company

from

of

tables

the

base

to

the

Messrs.

to

student

reproduce,

to

of

459

of material

XV

Chapter

and

physics.

Macmillan

the

theory

true

reader, including

obligations

No.

to

and

permission

for

Agriculture,

"A

the

assumes

part

is under

author

The

economically

more

for

of the

chemistry

of

knowledge

the

on

he

more

practice."

and

farmer

the

for

directly

The

is the

sively,
deeply, comprehen-

that

intended

is

book

short, the

In

successful

more

farmer

farming.

the

and

work,"

the

more

think,

can

he

can

The

do.

the

equal,

which

his

be

must

can

he

clearly

of

theory

successful

advantageously
surest

the

complete

more

and

qualifications being

"Other

7.

Sons,

Pa.,

Company,

Figures

Inc., Figures

18

and

1,5,
40.

and

14.

34,

CONTENTS
PAGE

vii

Introduction

PART

MATERIALS

THE

NUTRITION

OF

CHAPTER

The

Components

Plants

of

Animals

i.

Dry

matter;

"

2.

The

carbohydrates

"

3.

Fats

"

4.

The

"

5.

The

"

6.

Sundry

proteins

3
7

bodies

related

ash

organic matter;

"

and

and

The

Lipoids

24

non-proteins

36
.

ingredients

39

CHAPTER

The

Composition

"
"
"

"

1.

2.

Animal

3.
4.

The

Animals

tissues

and

and

organs

of

composition

of feeding stuffs

the

and

as

66

OF

NUTRITION

III

Resorption

1.

organs

2.

The

chemistry

of

"

77

digestion
of

77

digestion
.

"3. Resorption
4.

61
.

II

PROCESSES

The

The

whole

"

42

45

animal

composition

CHAPTER

"

Stuffs

42

THE

Digestion

Feeding

of

cell

The

The

of

II

PART

"

16

The

determination

feces

of

89
105

digestibility
vii

in

CONTEXTS

Vlll

CHAPTER

IV
PAGE

Circulation,

Respiration,

123

Circulation

"
" 2. Respiration
" 3. Excretion
1.

Excretion

and

123
132

139

CHAPTER

Metabolism

"
"

1.
2.

144

General

Enzyms

agents

as

144

in metabolism
.

of the

carbohydrates

152

The

metabolism

of the

160

The

metabolism

of the

simpleproteins
nucleoproteins

6. The

metabolism

of the fats

"
" 4.
" 5.
"
" 8.

Functions

of ash

ingredients

182

192

"

1.

General

"

2.

Methods

"

3. The

"

4.

The

"

5.

of
Significance

conception
of

balance
balance

192

investigation
of matter

202

216
.

results

241

FEED

III

REQUIREMENTS

CHAPTER

proteinkatabolism

"2.

The

energy

"3-

Conditions

1.

VII

Katabolism

The

"

194

of energy

THE

Fasting

VI

Nutrition

of

PART

The

178

of the nutrients

CHAPTER

Balance

168
171

7. Metabolism

The

148

metabolism

3. The

"

conception

katabolism

249

in
in

fasting
fasting

the fasting
katabolism
affecting

251

256

258

ix

CONTENTS

CHAPTER

VIII
PAGE

Maintenance

"

The

"

Energy

1.

"

2.

Maintenance

"

3. Factors

"

4. The

energy

Requirements

.267

for maintenance

values

Net

requirements

271

animals

of farm

the maintenance
affecting

relation of the

requirement
requirement to
.

maintenance

.280
.

temperature

308

CHAPTER

1.

" 2.
" 3.

IX

(Continued) The Requirements

proteinrequirements for maintenance
The ash requirementsfor maintenance
Accessorysubstances

Maintenance

"

"

1.

2.

Fattening

"
"

Matter

of

The

CHAPTER
The

Mature

of

.313
.332

348

Animals

350

Composition of the increase in fattening

Feed requirementsfor fattening

.350
359

XI

Growth

371

"

1.

General

"

2.

The

utilization of feed in

The

feed

" 3.

of

nature

growth

371

growth

requirementsfor

381

growth

CHAPTER

396

XII

Production

424

"

1.

"

2.

factor in meat

"

3. Feeding for meat

production

"

4. Influence

Nature
The

of meat

animal

as

production

424

production

428
444

of external conditions

453

CHAPTER
Milk

313

CHAPTER

Meat

304

external

XIII

Production

459

"

1.

The

physiologyof

"

2.

The

animal

milk

production
production
milk production
" 3. The influence of environment
on
" 4. The utilization of feed in milk production
"5. Feeding.for milk production
as

factor in milk

459
470

478
488
500

CONTENTS

CHAPTER

XIV
PAGE

Production

Work

531

1.

The

physiologyof work

"

2.

The

of the
efficiency

"

3. Feed

"

production

body

as

THE

FEED

CHAPTER

"

1.

Roughages,

"

2.

Roots,
The

"

1.

"

2.

" 3.

tubers

and

Values

578
579

Feeding

of

1.

" 2.
" 3.

XVI

Stuffs

591

Direct

comparisonsof feedingstuffs
Relative values based on compositionand digestibility
Conditions affecting
digestibility

Values

General

The

Feeding

of

values

Stuffs

computation of

Production

values

net

as

of

energy

"

values

regards protein

Rations

Feeding standards
Feed requirements
Method
of computation

597
601

630
630

regardsenergy

as

591

XVII

considerations

Production

Computation

"

572

fruits

CHAPTER
The

XV

concentrates

Production

" 1.
" 2.
" 3.
" 4.

SUPPLY

571

CHAPTER
The

IV

fodders

coarse

or

CHAPTER
Relative

560

Stuffs

Feeding

" 3.

544

requirementsfor work

PART

The

531

motor

Net

energy

values

634

....

667

....

678

XVIII

689
689
691
697

CONTENTS

XI

APPENDIX
PAGE

Estimated

Protein

Table

Energy

and

Maintenance

I.

per

Table

day

and

Animals

cattle

and

711

horses,

head

and

711
of

swine,

and

sheep

per

head

711

for

Requirements

III.

of

requirements

day

Farm

of

requirements

Maintenance

II.

Table

Requirements

fattening

all

growth

in

species

"

maintenance

with

considerable

no

addition

to

"

the

requirement

712
....

Table

IV.

Table
Table

Requirements

for

growth

V.

Requirements

for

milk

VI.

Requirements

for

work

with

no

considerable

tening
fat-

712

production

7i4
....

Average

Dry
Values

Matter,
of

Digestible

Table

VII.

Values

Table

VIII.

Values

Table

IX.

Values

Table

X.

Mineral

Protein

Stuffs

Feeding

production

the

by

7i4

Energy

Net

and

horse

per

ioo

Pounds

7i4

ioo

pounds

for

ruminants

7i5

per

ioo

pounds

for

the

per

ioo

pounds

for

swine

per

horse

721
722
....

of

elements

pounds

of

dry

stuffs

feeding

per

"

ioo

substance

723

REFERENCES

The
the

full-face
numbered

numbers

paragraphs

in
and

in

parenthesis
not

to

pages.

the

body

of

the

text

refer

to

ILLUSTRATIONS

FIG.

PAGE

i.

Different

2.

One

end

3.

Part

of

4.

Fat

5.

Scheme

of

types
of

cells

muscle

composing

the

body

43

fiber

50

fiber
.

cells

5i

in muscles

52

......

of

fat cell

Successive

6-8.
9.

muscle

Sheep's

58
in the

stages

formation

of

tissue

adipose

stomach

59

'

80

10.

Stomach

and

11.

Stomach

of

12.

Intestines

13.

Ccecum

of

14.

Section

of villi

duodenum

of

horse

81

....

hog

of

81

.......

cattle

84

horse
.

15.

Steer

16.

Blood

17.

Diagram

18.

Scheme

19.

Relation

in

85

105

digestion

stall

113

corpuscles
of

124

heart

mammalian

125

....

of

circulation

of

blood

127

....

of

cells

to

blood

20.

Main

21.

Alveoli

of

22.

Section

of two

23.

Diagrammatic

24.

Scheme

25.

Original Regnault-Reiset

29.

The

30.

Horse

31.

Lavoisier's

32.

Section

^^.

The

34.

Dulong's

35.

The

36.

Rubner's

37.

The

38.

Rate

131

of

133

scheme

133

of

metabolism

182

closed-circuit

Regnault-Reiset

Pettenkofer

130

alveoli

26.
27.

lymphatics

lung

of

28.

and

trunks

lymphatic

Scheme

vessels

respiration apparatus

apparatus

Pettenkofer

210

used

as

by

Zuntz

211

respiration apparatus

respiration apparatus,

Mockern

209

apparatus

213

explanatory

sketch

213

respiration apparatus
for

equipped
ice

experiments

214

with

Zuntz

apparatus

215

calorimeter

222

.....

of bomb

calorimeter

224

....

Zuntz

tread

power

calorimeter

water

respiration

237

calorimeter

at

The

Pennsylvania

State

calorimeter

marbling
of

226

dynamometer

gain

of

protein

238
239

356

meat
......

of

College

per

1000

pounds
xiii

live

weight

379

ILLUSTRATIONS

XIV

39."Rate
40.

of

gain

of

energy

Lobule

of

milk

gland

Alveoli

of

milk

gland

per

1000

43.

44.

Structure

of

milk

398

gland
463

Partial

section

of

wheat

Partial

section

of

oat

Partial

section

of

maize

grain

582

grain
.

45.

weight

463

42.

live

462

41.

pounds

584

kernel

588

INTRODUCTION

and

indirectly

materials

the

essential

an

of

of
smaller

portion

stored

in

up

in

legumes,

value

animal

in

into

forms

human

plays

both

in

increasing

an

for

which

conversion

the

into

human

With

the

of

growth

important
supply through

the

performed

one

with

hand,
laws

as

part

in

power

of

the

place

the

also

in

is

the

utilization

maximum

governing

to

yet

as

products

largely
the

greater

or

of

inedible

its aspects.

farm

and

of

an

as

of

conserving
soil

efficiency.

XV

to

tory
fac-

agency

body.

function

nutrition

animal

destined
all

the

work
In

seem

of

the

by

replace

in

the

industry.
to

degree

no

conversion

motor,

suggestion

no

and
of

utilized

motors

animal

knowledge

the

contain

of

inedible

possible

by-products

of

of
a

the

be

ample,
ex-

grasses

non-agricultural population
this

man's

function

production

of the

that

intimate

that,

is

are

for

they

agriculture and

may

as

for

the

the

which

agriculture,are

can

inanimate

food, there

take

can

role

only

products contribute

and

fibers

textiles, while

our

which

these

energy

substitution

however,

Vegetable

extent.

fibers

important

an

of

farm

ucts
prod-

wheat,

as

agriculture is

animal

that

clothing

of

respects,

less

and

that

proteins

essential

possible

as

matter

of

The
of

of

means

fact

such

energy

systems

system

It is true

supply

our

But

whose

all

economical

inedible

the

nutrition, while

man.

proportion

body.

animal

fill

permanent

large

as

to

as

the

directly available

the

human

for

food

the

crops,

of

in

important

so

is

crops

for

of

or

society is to
directly,because
by

regarding
well-recognized

directly

in

products

energy

both

in

true

two-thirds

unavailable

be

man

consists

distinctivelyfood

than

more

solar

the

farmer's

the

Even

direct

of

farmer

function

of

animal

this

is

It

nutrition

into

vegetable

his

function

that

farm.

the

the

concern

the

Particularly is

animals.

may

for
of

part

conversion

use.

nutrition

indirectly because

"

furnish

of

of

problems

The

This

shall

requires,
of

the

animals,

food

the

products

possible
farm

it is increasingl

on

be
the

mental
fundaso

that

xvi

INTRODUCTION

transformation

the

and,

the other

on

effected with the least

hand, the

so
ability

greatest economic

the

secure

be

may

possiblewaste,
laws

apply these

to

return, since

it must

as

to

be

never

in agricultureis not
forgotten that the criterion of success
a
maximum
production but a maximum
profit. It is with the
former
portion of this complex problem that the present work

attempts
Without
and

primarilyto deal.
enteringinto the controversy
be

guiding principle,
may
including the
maintain

entire

of the feed

materials

crude

nutrition

mechanist, the

the

of

regarded
complex

activities of the

the

vitalist

animal, whatever

the

its

physico-chemical
process,

as

of

reactions

by

which

into substances

converted

are

between

cells

the
suited

capable of being
In other
built up into livingstructures.
words, the study of
nutrition is a study of the chemistryand physics of the changes
through which the crude products of the soil yieldanimal tissues
hand
and
the one
secretions on
or
excretory products on the
to

the

body

or

other.
dealt with the food as a supply of
investigators
and
matter, dividingit into inorganicand organic constituents
the latter between
the nitrogenous and
distinguishing
among
In other words, they studied
the
non-nitrogenoussubstances.
as
problems of biological
problems of nutrition substantially
fundamental
far to
went
investigations
chemistry. Rubner's
shift the emphasis to the physicalside of the problem. It has
tially
to be clearly
come
recognized that the animal body is essenearlier

The

of

the

more

or

for the conversion

of energy
a mechanism
while
into motion
of its feed
chemical
energy
energy
material
of energy-containing
less incidentally
a
reserve

transformer

stored

may

be

this

capacityof

secretions

up

"

which

the

can

animal

part of the

be

utilized for human

body

matter

to

and

store

up

energy

in itself
of the

It is

food.
or

feed

in its
it

sumes
con-

a
as
significance
gives the animal its economic
of the food supply. Its value in this respect depends
proportionof its feed which it is able thus to set

which
conserver

upon
aside

the
"

i.e. upon

the

balance

between

the

income

and

outgo

this point of view that

and it is from
of energy
undertakes
to present the nutrition of farm
the present volume
falls into
this standpoint,the subject naturally
animals.
From
of matter

four

and

principaldivisions.

"

PART

THE

MATERIALS

OF

NUTRITION

NUTRITION

FARM

OF
ANIMALS

CHAPTER

COMPONENTS

THE

"
1.

Dry

animal
The

matter.

two

at

weight.
2.

well

incidental

or

for

known

needed
as

plant

or

food

animal
with

solutes

Finally,

its

Water

study

is

and
are

an

action

is
In

It

its

it is

more

regarded
animals.

or

the

The

and

of

in

than

of

pressures

physiologicalprocesses.
of

living

supplied

to

live

not

the

The

stock.

because
water

be

to

matter.

chiefly with

important.
3

the

carrier

osmotic

the

chemistry

more

is

ism
organ-

tissues

and

solvent

transformations,

economically

too

reflection

part of the

supporting

as

factor

essential

an

as

dissociating electrolytes appears

to

lose

to

ceases

little

very

nutrition, therefore, deals

and

material

the

function, lending elasticity

acts

important
in

the

matter.

living beings is

to

essential

as

usually abundantly

of animal

ordinarily

while

products

waste

related

supply
fundamentally any

matter,

dry

or

matter.

be

plants

supply

mechanical

strength.

intimately

of

water

ingredient.
a

dry
to

means

it

plant

consisting solely of

as

the

course,
no

water

until

regarded

component

the

it has

materials

of the

very

that

other

combined

and

water

water

require mention,

show

any

by

adequate

an

to

to

is, of
is

Water

"

is

weight

residue

the

Water.

necessity

in

loss

The

accidental

consisting of

boiling point of

the

while

water,

the

composing

ordinarily separated by maintaining

are

above

or

Ash

Matter;

material
as

ANIMALS

AND

Organic

The

"

regarded

be

may

PLANTS

OF

Matter;

Dry

i.

but

the

dry

this
because

is

Organic

3.

matter.

ANIMALS

FARM

OF

NUTRITION

By the action of

"

dry matter of plantsor animals

one
being converted into
portions,
the

into

two

perature,
high tem-

be

may

separated

the gaseous

state,

solid form.

in the

Following
to
older nomenclature, it is customary
distinguishthese
gredient
ash," ininorganic,"or
organic and
portionsas
the other

while
the

behind

remains

terms, however,

The
since

supposed
Organic matter

the

during oxidation

be volatilized

be said
may
of the organism,

commonly used
carbon compounds
and

this definition is inexact

even

once

is

term

broadlyequivalentto
but

may

the

in which

sense

misnomers,

extent

some

sharp distinction exists as was

organic and inorganiccompounds.

between

be

to

are

such

no

in the

"

"

"

"

two

to

at

oxygen

or

the

may

element

same

in the ash

appear

accordingto circumstances.
For example, the element
sulphur is an essential ingredient
in air part of the
these are
of the proteins. When
burned
sulphur escapes in the gaseous form, but a part also combines
in the ash as sulphates.
with any bases present and appears
Even
the element carbon, distinctive of so-called organicmatter,
may
appear in part in the ash of the plantor animal in the form of
when

carbonates
radicles.
be

an

These

the bases of the ash

examples serve

integral
part
and

matter

to

of the molecules

yet appear

in

are

that

show

after incineration

element

an

make

which

of the acid

excess

in the

up

the

ash.

may

organic
it

Thus

recentlybeen shown that the phosphorus of wheat bran

and
other
feeding stuffs is present chieflyin the form of a
complex carbon compound, yet when these materials are burned
in the ash in the form of phosphates.
the phosphorus appears
of the elements
is usuallyregarded as consisting
Organic matter
phorus
carbon, hydrogen, oxygen, nitrogen and sulphur,phosbut doubtless other
being sometimes added to the list,

has

elements

like

the structure
4.

potassium, sodium, chlorin,etc., also

of the

Subdivision

of

"

"

organic

organic

organiccompounds found
is very
necessary
only the

great.

For

the

into

enter

molecules.

matter.

"

The

in the animal

present

purpose,

number

body

or

ual
of individin the

however,

plant

it is

not

separatelyeach individual substance but

into which
of the important groups
general properties
to

consider

they may be classified.

The organicconstituents

of the

body

may

be subdivided

into

COMPONENTS

THE

PLANTS

OF

ANIMALS

AND

Under the former

non-nitrogenousand nitrogenoussubstances.
the fats,the organicacids and
included the carbohydrates,
are
The
clude
various other minor
nitrogenous substances ingroups.
stances
the proteinsand a varietyof simplernitrogenous sub-

sometimes

classed

togetheras

non-proteins.In

the

the

following sections these various groups will be considered as

for an
far as is requisite
intelligent
study of their behavior in
that the reader has already
the animal body, it being assumed

knowledge of
physical.
some

Mineral

5.

found

their

matter,

in the ash

and

both
generalproperties,

ash. "To

or

what

extent

reckoned

commonly

as

chemical and

the elements

the mineral

ments,
ele-

namely, potassium,sodium, calcium, magnesium, iron,

phosphorus,sulphur,chlorin,silicon,
etc., are actuallypresent
in the livingplant or animal as electrolytes
and to what extent

of complex organicmolecules, it is at present impossible

ingredients

as

term
at

with

definiteness.

ordinary usage the

ash is equivalent
to the residue remaining after incineration
low a temperature as possible,
usuallynot exceeding a
to

as

state

any

In

dull red heat.

The

proportionof

ash in

ordinary feedingstuffs varies

siderably
con-

accordingto the kind of plant,the portionof the plant

used (seeds,stems, leaves, roots, etc.),the maturity of the
plantand various other conditions. Wolff givesthe followingas
generalaverages for the proportionof ash in the dry matter :
"

Cereal crops

Leguminous
Oil plants
.

The

crops
.

varies
proportion

In the

About

in the straw

most

animal, the presence of ash is most

two-thirds of the dry

fat) consists of ash.

Ash

matter

is

by

no

and

least in the

evident in the bones.

of the clean bone

means

grain.

absent

(freefrom

from

the soft

gredient
body, however, of which it forms an essential inThe
proportionvaries in different organs, but as
the body, inclusive of the skeleton,
rough general average

tissues of the

contains

about

3.5

per

cent

of ash in

the

fresh

substance,

NUTRITION

about

equivalentto

FARM

OF

of the

cent

7.1 per

ANIMALS

dry

matter.

The

portion
pro-

is greater in the young

than in the
dry
and greater in the lean than in the fat condition.
animal
lows:
more
important elements found in the ash are as folof ash

mature

The

to

matter

"

metal

is

indispensableto plant growth and is

in the active,growing
found in all parts of the plant,but especially
such
parts. In the animal body it is found abundantly in the tissues,
the muscles, glands,nerves, etc., while the fluids (blood,plasma,
as
small amounts
of it.
lymph, etc.)contain relatively
Unlike potassium,sodium is not indispensable
Sodium.
to plant
growth, although it apparentlyis useful to the plantunder some
ditions.
conand
in the stems
leaves of plants,
It is found especially
although not so abundantly as potassium. Seeds contain but little
in the fluids,
of it. In the animal body it is especiallyabundant
littlepotassium.
which, as justnoted, contain relatively
for the growth
Calcium.
Like potassium, calcium is necessary
in the leaves and stems
of plants. It is found
of plants
especially
in the seeds.
It appears
and to a much
less extent
to be equally
in
and
and
is
essential to the animal
found
all parts
organs of the body.
Its most
of the skeleton,
the
striking
use, however, is in the formation
mineral portion of which (81) consists chieflyof calcium phosphate
and carbonate.
Both these compounds being scarcelyat all soluble
in water, they are well adapted to form the framework
of the body.
In the skeletons of the higher animals calcium phosphate is the chief
Potassium.

This

"

"

"

mineral

ingredient,while

which
Crustacea, the shell,

in

the

lower

animals

like shellfish and

correspondsto the bones of domestic

mals,
ani-

contains

chieflycalcium carbonate.
Magnesium.
Magnesium is also one of the elements essential
for plantgrowth. It is found throughout the plantin smaller amounts
than calcium,but is more
abundant
than the latter in the seeds and
In the animal
to aid in seed formation.
seems
body, magnesium
in
much
smaller
but
amounts.
usuallyaccompanies calcium,
Iron.
A small amount
of iron is requiredby the higher plants
for the formation of the green coloringmatter
(chlorophyl)by means
of which
mal,
they assimilate the carbon dioxid of the air. In the aniiron in small quantity is necessary
for the formation
of the red
(haemoglobin)of the blood which is the agent for
coloringmatter
conveying the oxygen of the air to the tissues. While, therefore,but
of iron is requiredby either plants or animals,1
a very small amount
"

"

it is nevertheless essential
1

It is estimated

to

that the blood

the most

fundamental

of

man

an

adult

processes

contains about 3 grams

of life.
of iron.

by chlorophyl-b
earingplants,and the great
vegetabletissue is composed either of carbohydratesor
dioxid

of carbon
of

mass

of their
The

nearly related derivatives.

carbohydrateshave

common

more

Starch is familiar

time.

tubers,and cellulose
forms, in the woody fiber

presence

in
of

since almost

known

has been

long

plants. Of the sugars, cane

times,while the
prehistoric

other sugars in
The
familiar fact.

for

mealy portionof grainsand

and linen and, in impure
cotton

of this and

honey, etc.,is

been known

in the

to us

in certain

sugar

ANIMALS

FARM

OF

NUTRITION

plant juices,in
more

common

fruits,

sweet

sugars

were

quiteearlyin the historyof chemistry.

The
7. Classification.
carbohydrates contain
hydrogen
in exactlythe proportions
to form water, and their
and oxygen
this fact, although compounds exist
is derived
from
name
and
of oxygen
of hydrogen to one
atoms
which contain two
yet are not carbohydrates,such, for example, as acetic acid,
the simple
C2H4O2. The simplestof the carbohydratesare
By
exactlydesignatedas the monosaccharids.
sugars, more
of water, the monosaccharids
with elimination
polymerization,
classified
yieldmore
complex carbohydrateswhich are conveniently
and polysaccharids.
as
di-,tri-,
identified

separatedand

"

Monosaccharids,or simplesugars
8.

Composition.

the

by

general formula

formula
nine

known

are

carbon

and

atoms

monosaccharids

The

"

Cn H2" On.
whose

which,

may

Substances

molecules
from

represented
having this general
be

from

contain

chemical

one

to

point of view,

dehyde,
carbohydrates. The simplestof these is formalto be the first step in
CH20, which is believed by many
the synthesis
of carbohydratesby the green plant. Only the
the
as
C6 and C5 compounds, however, known
respectively
of importance in their
hexose and pentose carbohydrates,
are
be called

may

relations to nutrition.
9.
are

an

Hexoses.

"

The

most

important hexose

monosaccharids

dextrose,levulose,galactoseand mannose.
Dextrose,^-glucose,or grape sugar, is generallyregarded as
aldose

of the hexatomic
Sorbite
Dextrose

alcohol sorbite.

CH2OH:

CH2OH-

(CH

"

(CH

OH)4- CH2OH
"

OH)4-

CHO

It

and

levulose

cane

and

Galactose

in small

also

of

isomers

this

with
in

amounts

compound

known.

are

of

isomers

are

mannose

juicesof plants along

the

Sixteen

of which

possible,twelve

are

ANIMALS

AND

is found

and

sugar,

mammals.

of

blood

the

universallyin

almost

occurs

PLANTS

OF

COMPONENTS

THE

dextrose, occurring in

di- or polysaccharids.
as
only in combination
Levulose, or fruit sugar, is a ketose of sorbite, having the
formula
CH2OH(CH
OH)3-CO-CH2OH,
eight isomers
nature

"

possible.
being theoretically
in
honey.
plant juicesand
and

readily diffusible
found

those

in

nature

have
are

polarizedlight. Thus
right-handed rotation
They reduce an alkaline
of

fermented
and

in

less sweet

or

dextrose

water
taste.

in

and
All

opticallyactive, rotating the plane

dextrose, as its name
implies,has a

and

levulose

left-handed

salts,especiallyof

of metallic

solution

rotation.

qualitativetest for
of quantitativedetermination.
a means
as
They are
by yeast, yieldingas the chief products ethyl alcohol

and

them

more

with

all soluble

are

this fact is utilized both

and

copper,

mixed

occurs

monosaccharids

hexose

The

It

as

dioxid.

carbon

ing
simple sugars, correspondbut
the hexoses
to
C5H10O5.
having the general formula
aldoses.
Like the hexoses, they
Those
are
occurring in nature
reduce
metallic
able
oxids, but unlike them
they are not fermentby yeast.
10.

Pentoses.

"

The

pentoses

are

By the hydrolysis of gum-arabic or cherry gum,

there is produced dextro-rotatory arabinose
(/-arabinose). LevoThe
(d-arabinose)has been prepared artificially.
rotatory arabinose
form
racemic
inactive or
(i-arabinose)has been found in human
Arabinose.

urine

in small

"

amounts.

there is produced a
By the hydrolysis of wood
Xylose.
gum
known
The
as
levo-rotatoryform
/-xylose.
dextro-rotatory pentose
of the same
(J-xylose)is obtained in the hydrolysis of certain
sugar
nucleo-proteins,the pentose group
seeming to be a constituent of
of those compounds.
the molecule
"

Rhamnose

is

derivative

of the

pentose

replaced by
hydrogen has been
widely in the vegetable kingdom.

of

sugars

methyl.

It

in which
occurs

an

atom

somewhat

NUTRITION

IO

OF

FARM

ANIMALS

Glucosids
monosaccharids

only occur in the free state but

with a great varietyof substances in the
also in combination
so-called glucosids.The glucosidsreadilyundergo hydrolytic
either by the
cleavageinto their two (or more) constituents,
For example, the
action of chemical reagents or of enzyms.
amygdalin of the bitter almond yieldstwo molecules of dextrose,
of hydrocyanic acid,and cerebron,
of benzaldehyd and one
one
of the brain,splits
a constituent
up into cerebronic acid,sphinor less familiar glucosids
gosinand galactose.Among other more
be mentioned
salicin,
saponin,phloridzinand digitalin.
may
The

11.

not

Disaccharids
12.
as

The

hexose

group.

The

"

polymers or anhydridsof
of two

union

the

to

by

hexose

the

another

From

glucosidsof
Sucrose.

juicesof
the

sap

number

By

of

Sucrose,or

"

cane

familiar

more

the

reverse

C12H22O11 +

by

H20.

writers

some

has

utilized

are

commercial

as

it is present in

amounts

heat, aided by

be

may

dilute acid

with

brought about,

large

alkali,or by

or

molecule

and

levulose.

rotates

while,owing
greater than

to

the

is formed

to

sucrose

molecule

C6H12O6 -f-C6H12O6

the rotatory power

dextrose, the mixture

in the

of

yield one

plane of polarizedlight to

the fact that

that of

molecule

one

of water

one

C12H22O11 + H2O

which

sented
repre-

of certain enzyms,
notably the invertase of yeast,
of the
of the general reaction
for the formation

each of dextrose

Sucrose

be

may

plants.

disaccharids

combining

long
be-

probably been longest

carbohydrates. It is found in the
sorghum, in the sugar beet and in
sugar,

smaller

In

the action of

action

formation

equationCeHi206 + CeHi206
point of view they are termed

the sugar cane

and
of the maple, all of which

the

two

their

one

present known

at

of sugar.

sources

and

of

the elimination

only disaccharids

group

may

the monosaccharids.

of the

known

The

be

regarded
monosaccharids, formed by the

of the latter with

molecules

of water.

molecule

the

disaccharids

of

the

of levulose

equal parts

foregoing reaction

right,

rotates

is

of the
to

the

THE

left.
has

COMPONENTS

On

account

been

called

extended

to

OF

of this

PLANTS

inversion and

the

milk

Lactose, or

"

cane

of this term

use

sugar

has

been

of

monosaccharids,
than

It is less soluble

taste, having

in

this

ent
ingredi-

it may
be broken
of water, into two molecules

case

and

sucrose

characteristic

sucrose,

molecule

grittyfeel

is

Like

one

di-

monosaccharids.

sugar,

of mammals.

the addition

of

II

designatein general the hydrolyticcleavageof

of the milk
up, with

ANIMALS

fact,this breaking up of

saccharids into their constituent

Lactose.

AND

dextrose

and

therefore less

in the mouth.

It is

galactose.

sweet

the

to

found

not

in

plants.
Maltose.

the

By

"

action

duringthe germinationof
of

animals, a disaccharid

of certain

seeds

and

known

as

ferments

maltose

tract
digestive
is produced. It is

abundantly in malt, whence

hydrolyzedyieldstwo molecules

when

13.
soluble

General

properties.
"

in water

The

starch

also in the

therefore present
sugar

upon

its

This

name.

of dextrose.

disaccharids

are

crystalline,

and

active.
Sucrose does not reduce
optically
alkaline copper
but lactose and maltose
do.
The
an
solution,
disaccharids are not fermentable.
Any cases in which they are
action
apparently fermented are found to be precededby some
inverts

which

breaks

or

up

the

disaccharids

into

their

stituent
con-

monosaccharids.

Trisaccharids
14.

the

By

elimination
the

as

molecules

of C6Hi206

molecules

of

with
be

One

is present in the sugar beet,the cotton

in wheat.
Upon hydrolysisit yieldsone

ramnose,

barleyand
each

of three

of water, there may

Ci8H320i6, called a trisaccharid.

two

compound

known
in

of

union

dextrose,levulose and

the

formed

such,
seed,
cule
mole-

galactose.

Polysaccharids
Chemical

sacchari
like the dipolysaccharids,
are
anhydrids,but are formed by the combination of
molecules of the monosaccharids
and have a correspondmany
ingly
high molecular weight. The generalformula of the hexose
the value of n doubtless varying
polysaccharidsis (C6H10O5)n,
15.

structure.

"

The

NUTRITION

12

through

wide

FARM

OF

but

range,

the

ANIMALS

dividua
weights of the indetermined.
finally
stances
usuallyamorphous sub-

molecular

have not been

polysaccharids
The
tasteless and
are
polysaccharids
less
are
more
or
which, with the exceptionof cellulose,
active but in generalare
soluble in water.
They are optically
not diffusible through membranes.
They are hydrolyzedeasily,
especially
by the action of heat and acids and by enzyms, yielding
monosaccharids.
ultimately
In addition to their common
they are designatedby
names,
of which
derived from the monosaccharids
out
terms
they are
built up.
Thus
starch,which is an anhydrid of dextrose and
is a dextran.
yieldsonly this sugar upon hydrolysis,
Similarly,
there are
arabans, xylans,etc.,
levulans,galactans,
mannans,
yieldingthe correspondingsugars when hydrolyzed. In the
to
it is customary
the
same
distinguishbetween
manner,

hexosans, derived

anhydridsof
16.

The

those which

the

from

hexoses, and

the

pentosans, the

pentoses.

hexosans.
are

the

most

"

This

of
group
in the
abundant

carbohydratesincludes
vegetablekingdom and

of the greatest significance

of nutriment
as
sources
animals,viz.,starch,the dextrins and gums, and
its various

derivatives.

It will be

important hexosans

more

convenient

to

in the order

somewhat

for

man

and

cellulose and

consider
of their

the
sistance
re-

to solvents.

17.

Cellulose.

Cellulose

"

constitutes

the

basis of the cell

plantsand is also found in certain lower animals (tunicates). Clean cotton consists of nearly pure cellulose,each
fiber being a singlecell from which
the contents
(protoplasm)
of paper
have nearlydisappeared. Linen and the best qualities
other examples of nearly pure
cellulose. A
are
crystalline
walls of

form

has also been

Cellulose
to

described.

is insoluble in

reagents in

water

and

comparativelyresistant

general. Plants, however, contain

enzyms

which are able to bringit into solution in the processes

(cytases)
of plantgrowth, and apparentlythese enzyms
play some
part
dissolved
It
also
in its digestion
animals.
attacked
and
is
by
solves
by some
speciesof bacteria. Concentrated
sulphuricacid disit,and the solution,on dilution and boiling,
undergoes
dextrose.
Cellulose is therefore a dextran.
hydrolysis,
yielding
Its molecular
weight is unknown.

COMPONENTS

THE

in

cellulose

instead

in the

further

fact that

of

in the

yieldingonly dextrose,as
produces a variety of both hexose
former
mannose
includinggalactose,
dextrose,and
and

While

plant,the

the
analysis,

(109) and
19.

whether
true

cellulose

in the

hydrolysis,
of

case

to

xylose. The

"

found

are

loses
hemicellu-

of

both in the

extent

as

feedingstuffs

"

crude fiber

"

(110).
of

advancing maturity they become

thickened,not only by the formation

of hemicelluloses

is

of

plant, the cell walls consist

In

"

union

the framework

method

"

hexosans

chemical

or

greater or less

extract
nitrogen-free

cellulose,

the
sugars,
levulose,as well as

constitutes

serve

the young
cellulose.
With

nearlypure

and

the

pentose

and

In the conventional

hemicelluloses

Lignin.

and

in mixture

hemicelluloses

material.

reserve

13

regarded,therefore,as containingboth

pentosans, but

uncertain.
the

the latter arabinose

be

must

ANIMALS

These

"

dilute acids and

with

AND

polysaccharidsdiffer from true

being hydrolyzed by comparatively short boiling

Hemicelluloses.

18.

PLANTS

OF

of additional cellulose and

"

but

of numerous
by the deposition
incrusting
substances,"the most
important group of which has received
the collective name
of lignin. These substances contain a considerably
of
carbon than cellulose (54 to 60
higher percentage
be separatedfrom the latter by oxidizing
per cent) and may
agents. The substances of the ligningroup contain methoxyl
(" O CH3) and ethoxyl (" O C2H5) groups in considerable
are
amount, and by some
regardedas substituted celluloses.
so-called "crude
The
fiber.
20. Crude
fiber"
(109) of
plantscontains most of the cellulose and ligninof the cell walls
"

"

"

and

in addition

of carbon
to be

third group
the cutin group1
whose percentage
is stillhigher(60-75Per cent). Cutin appears
"

"

indigestible.
Starch

is

of the most

and

important
of the vegetablecarbohydrates. In the growth of plants,
starch is formed in the green leaves by the aid of light,
and is
In the mature
the first visible product of assimilation.
plant,
it is stored up in largequantities
in the seed or in the tuber to
supply the needs of the new
grains,
plant. Hence the common
well
as
potatoes, are rich in
corn, wheat, oats, barley,etc., as
21.

Starch.

starch and

legumes

"

form

contain
1

one

commercial

sources

it in less amounts

Compare Konig:

Landw.

Vers.

common

of it.
but

The

still

seeds of

most

abundantly.

Stat.,65 (1907), 55.

In

NUTRITION

14

OF

ANIMALS

FARM

It is not

the oil seeds it is replacedby oil.

found

in the animal

body.

plantsin the form of microscopicgranules,

form for each species,
which have a peculiar
so that we
speak
may
These grainsconsist of
of the starches rather than of starch.
of a varietyof cellulose inclosing
a
surroundingenvelope consisting
Starch

and

soluble substance

more

treated with much

When

lose.

in

occurs

bursts

the

envelope and

substances

or

hot

water

known

as

granuthe starch grainswells

the enclosed

granulose dissolves,

less hydration.
or
probably after undergoingmore
be hydrolyzedreadilyby dilute acids or alkalies
Starch may
or
by heat. The final product of its hydrolysisis dextrose,
in an
which
glucose or
impure form constitutes commercial
As alreadynoted,
Starch is therefore a dextran.
starch sugar.
certain enzyms,
notably those formed in germinating seeds
and others secreted in the digestive
tract of animals, act upon
Starch is also
starch readilywith the productionof maltose.
acted upon
speciesof bacteria with the formation of
by some
and in some
cases
lactic,butyricand other acids,methan
drogen
hy22.

Galactans.

"

Galactans

plants,other feedingstuffs
from
23.

in leguminou
particularly
being comparativelyfree
more

occur

them.
Inulin.

The

"

roots

of the

artichoke,dahlia,dandelion,

chicoryand other compositecontain instead of starch a quite

similar carbohydrate,
inulin,which on hydrolysis
yieldslevulose
instead of

it is a levulan.
dextrose,i.e.,

hydrolysisof starch a series of

intermediate
ill-defined,
compounds is produced, collectively
called dextrins.
Commercial
dextrin is made
by heatingmoist
starch to about 2350 Fahrenheit.
It is likewise produced in the
cooking of starchymaterials,the brown crust of bread, for
example,consisting
largelyof dextrin. Various dextrins have
been
separated and described, but it seems
questionable
whether
the investigators
have worked
with definite chemical
tively
For the present, it seems
individuals.
wiser to speak collecof the dextrins as intermediate
productsbetween starch
and the simplerdi- and mono-saccharids.
a

24.

The

25.

Glycogen.

less

dextrins.

degreein

"

"

In

In

the

the liver and

other parts of the

muscles

body, there

of

animals, and

is found

to

in rather

NUTRITION

"

Fats

3.

OF

Related

and

ANIMALS

FARM

Bodies

Lipoids

the

"

the rather vague term

fat-like substances,there are included,besides
Classification.

28.
or

of chemical

largenumber

"

Under

"

individuals

of varied

lipoids,"
fats,a
complex
(with the

true

and

Chemically, these substances

characterized by containing
are
exceptionof the cholesterins)
the higherones
radicles of the so-called fatty acids,principally
of the series. Physically,the lipoidshave been defined,principally
from the standpointof the physiological
chemist, as
substances which are soluble in organic solvents,such as ether,
alcohol,chloroform or benzol. This latter definition,
however,
such
be
includes substances,
the cholesterins,
which would
as
excluded
by the chemical definition justgiven. For the present
molecular

structure.

the

be conveniently
principal
lipoids
grouped under
may
five heads : (1)fats,(2) waxes,
(4)phosphatids
(3)cholesterins,
or phospholipins,
(5) cerebrosids or galactolipins.
purpose,

The
29.

Occurrence.

animals

contain

It is

"

seldom
rise

may

the

as

below
falling
high as fortyin

material

reserve

familiar fact that

inconsiderable

not

Fats

the very
of the body and

of
adiposetissue (94) consisting

called

more

or

less filledwith

tissue

are

found

fat.
in all

Larger

or

of

amount

six in the very

the bodies

fat,the

lean animal

centage
per-

while it

fat animal.

The

is contained

in what

cells of connective
smaller

of

fat is
is

tissue
of adipose

amounts

in
parts of the body but especially

the subcutaneous

the tissues surroundingthe intestines,

tissues,
in fat animals, in the muscles.
and, particularly

plants,fats

In

all parts of the

usuallyless abundant.
stored
plant but are especially
are

where

They
up

occur

in the

in

seeds,

material

which
is metabolized
as
reserve
they serve
seeds, like those of cotton, flax
during germination. Some
and rape, contain fat so abundantly that they are commercial
of oil. In the plant,'
the fat is not depositedin special
sources
tissues but is usually distributed through the protoplasm of
the

cell.

various
free

Both

animal

and

simple fats,often

fattyacids.

vegetable

fats

containingalso

are

small

mixtures

of

amounts

of

COMPONENTS

THE

Molecular

30.

structure.

AND

PLANTS

OF

The

"

ANIMALS

17

fats

simple neutral

are

tri-

that is, they are

esters
of the triatomic alcohol
glycerids,
of
fatty acids, the hydrogen atoms
glycerolwith monobasic
the three hydroxyIs being replacedby the acid radicles. Their
the
generalformula is as follows,Ri, R2 and R3 representing

which
radicles,

acid

OH
CH2
fat CH2
ORi

Glycerol
Neutral

fattyacids

The

unsaturated.

CnH2n02
lower

the two

acetic acids.
each

carbon

saturated

and

OH

OR2

to

"

CH2
CH2

"

"

have

acids of

of which

being united

same

OH

OR3

into the saturated

the

mula
general forthe aliphatic
series,
formic

as

of these acids

the

and

the

familiar

are

generalformula

The

"

fattyacids

the normal

are

members

atom

CH

"

be divided

may

The

CH

"

be the

not

may

or

may

is

adjacentones

and

as

follows,

by

single

bond.

CH3-(CH2)n

-COOH

saturated acids contained in the animal fats

principal
and palmiticacid,C16H32O2. Besides
stearic acid,C18H36O2,

The
are

two

also found in small amounts.

these two, however, others are
In butter fat,especially,
several of the lower acids of the series
ones
present, the principal

are

beingbutyric,C4H8O2, caproic,

C8Hi602, capric,C10H20O2,lauric,C12H24O2
C6Hi202, caprylic,
and

myristic,C14H28O2.
found also higheracids of

In

the

the

same

body

fats there

have

been

arachnic
series,
particularly

acid,C20H40O2.
The
in

unsaturated

containingtwo

instead
than

of

one

in

or

and

the saturated

abundant

fattyacids
more

differ from

carbon

acids.

animal

CH3-(CH2)7-CH

united

Of the unsaturated

fats is oleic
=

acids

by two bonds
containingless hydrogen

atoms

consequentlyin

the saturated

acids,the

most

acid, having the formula

CH-(CH2)7-COOH

eruic acid of rape oil also belongsto this series,

and the
of linseed oil and other dryingoils belongs
linoleicacid,C18H32O2,
The

to

related

series of unsaturated

CnH2n_402 with

double

unions

of the

of carbon

general formula

atoms.

noteworthyfact that nearlyall the fattyacids occurring

animal body contain an even
number
of carbon atoms,

It is a
in the

two

acids

reactions.

Chemical

31.

the chemical

Of

"

ANIMALS

FARM

OF

NUTRITION

reactions

of the

is that known
importance physiologically
as
more
or
hydrolysis. It consists of
strictly
as
saponification,
a cleavage and
hydrationof the molecule, yieldingglyceroland
of this reaction
familiar instance
is in
fatty acids. The most
For example, if tri-stearin is acted
the process of soap making.

fats,the

of

one

most

is

final result

by potassium hydrate the

by the followingequation:
upon

represented

as

"

+ (KOH)3
C3H5(C18H3502)3
Potassium

Tristearin

In
soap,

(KC18H3502)3+ C3H803

hydrate

tristearate

Potassium

Glycerol

reaction,the alkali salt of the fattyacid, that is,a

is obtained.
siderably
By the action of water at temperatures conresult is reached
ioo" C, essentially
the same
above

this

instead

except that the free acid is obtained

of the

salt.

The

of acids,
decompositionmay also be effected by means
which probably act as catalyzers.
is the hydrolysis
of fat by
Of most importancephysiologically
Such enzyms
of enzyms.
are
means
produced by certain plants
and are also found in various digestive
juices,notably in the
same

of the

secretion

of

general name
to

appears
of low
may

be

the

lipases.The

reversible

molecular

effect

These

pancreas.

enzyms

hydrolysisof

reaction,at
In

weight.
of

have

other

fats

least with

words,

the

glyceroland the
an
equilibriumat

certain

the

by enzyms
glycerids

the

same

enzym

the combination

cleavage
glycerid
the
reaction in either
fattyacid,
or

received

case

of

reaching

stage.

general propertiesare
Their specific
to all the fats.
common
gravityis in all cases
less than one, so that they float on water.
They have a fatty
feel and leave a permanent
grease spot on paper or fabric. They
is soluble to a not
almost insoluble in water, althoughwater
are
in fats. They are readilysoluble in ether,
inconsiderable extent
of them
in petroleum ether,
benzol, carbon disulphidand most
but only sparinglyin alcohol.
The
melting point of the fatty acids increases with the
cult
molecular
weight. The exact melting point of a fat is diffifor the three common
to determine, but
glyceridsand
be stated approximately as
the correspondingacids it may
32.

Physical properties.

follows

"

"

Certain

COMPONENTS

THE

OF

PLANTS

Melting

AND

ANIMALS

Points

Oleic acid

-40 to -50 C.
140 C.

Palmitin

630 to 650 C.

Olein
.

Palmitic

fats
The
the

62. 6" C.

acid

Stearin

71.60 C.

Stearic acid

71.50C.

distinction is

19

commonly

made

between

fats and

oils,the

ordinary temperatures and the oils liquid.

the proportionin which
difference depends largelyupon
various simplefats are
taining
present. Olein and other fats conunsaturated
acids are
usuallyliquidat room
ature
temperbeing solid

and

at

their presence

increases the softness of the fat.

The

fattyacids of highermolecular weight are volatile only

at comparativelyhigh temperatures and
at reduced
pressure.
Those of lower molecular
ter
weight,notablythose contained in butof steam
and their
fat,can be readilydistilled in a current
butter fat from other animal
to distinguish
proportionserves
fats.

importantphysicalproperty of the fats,which, however,

by no means
peculiarto them, is that of forming what is

An
is

known

as

an

emulsion.

Fat

is said

to

be

emulsified

when, in

the

liquidstate, it is distributed in minute dropletsor globules

other liquid
throughout some
; for example, if fat be violently
shaken
is

not

and

with

water

permanent,

risingto

certain

other

an

emulsion

is formed.

Such

however, the fat droplets

very

the surface.
substances

prevent this

The

an

soon

of small

presence
dissolved in the water,

emulsion

coalescing
amounts

of

however, will

separationand give rise to a permanent emulsion.

The most
substance producing this effect is soap.
common
Certain gums
and also proteinslikewise serve
to retain fat
in the emulsified state.
The most
familiar example of such an
emulsion
is milk, the fat being held in suspensionin this case
by the action of the proteinsof the milk. This effect of various
substances in retaining
fat in the emulsified form depends upon
their effect upon
the surface tension of the contact
tween
layerbefat and water, but a full discussion of this point would
be out of placein this connection.
As has already been stated,the reserve
33. Native
fats.
fats of the animal body are triglycerids,
oleic
of stearic,
chiefly
"

NUTRITION

20

OF

FARM

ANIMALS

and

of esters of lauric,
palmiticacids,althoughsmall quantities
myristicand arachnic acids and frequentlyfree fattyacids are
also found, as well as minute
of esters
amounts
of the higher
Since stearin and palmitin are
alcohols,coloringmatter, etc.
solid at ordinarytemperatures, while olein is liquid,the consistency
of a fat depends largelyupon
the proportion of olein
which it contains and varies not only between
different species
of animals but often in different parts of the body of the same
animal.

The

fats of cold-blooded

than those of warm-blooded

lower

at

animals

animals

and

contain

olein

more

therefore remain

liquid

temperatures.

The

vegetablefats contain a greater varietyof fatty acids

than the animal
fats,notably unsaturated acids like linoleic
and eruic,as well as oxy-acidsand esters of the higheralcohols
(waxes),while the so-called crude fat,or ether extract (108)of
vegetablematerials contains a great varietyof ether-soluble
of
substances,includingwaxes, resins,chlorophyl,etc., some
which are but remotely related to the true fats.
34. Elementary composition.
The
three principaltriglycin formula
erids,stearin,palmitin and olein,while differing
and molecular weight,differ but little in their elementarycomposition,
the following
table shows :
as
"

"

Table

i.

"

Composition

of

Triglycerids

Triolein

Carbon

77-31

Hydrogen
Oxygen

10.85

11.84

Total

100.00

Naturally,therefore,the composition
animal

fats varies but

different
Schulze
gave

speciesof
and

either
little,
animals.

Reinecke1

upon
the followingresults.
1

Landw.

The
the

of the

ordinarymixed

in different individuals
classic

in

investigationsof

compositionof

Vers. Stat.,9 (1867),97-

or

animal

fats

Table

Benedict

and

2.

to

be

Composition

"

fat

Table

average

PLANTS

Osterberg1 obtained

of human

The

OF

COMPONENTS

THE

3.

carbon

76.5 per

Animal

of

the

ANIMALS

21

Fats

followingfor

the

position
com-

commonly

sidered
con-

"

Composition

"

AND

content

of

Human

of animal

Fat

fat is

cent.

Waxes
35.

In

popular

usage,

upon

their obvious

well-known

feel

based

the distinction between

fats and

waxes

is
the

substances having
physicalproperties,
called
oils
fats
or
being
according to their
greasy
are
consistencyat ordinary temperatures while the waxes
solid,can
be kneaded
and lack largelyor wholly the greasy feel.
1

Amer.

Jour. Physiol.,4 (1901),69.

NUTRITION

2 2

OF

FARM

ANIMALS

defined as fattyacid esters of alcohols other

are
Chemically,waxes
while the fats have already been defined as the fatty
than glycerol,
mon
esters of glycerol. This distinction is far from
according with comsubstances
Under
it many
as
popularly known
waxes
usage.
in
for
and
beeswax.
wax
as
are
f
ats,
example,
Japan
technically
part
materials ordinarily
On the other hand, numerous
regarded as oils or
familiar bodies
One of the most
be designatedas waxes.
fats must
of this class is spermaceti,
commonly regarded as a fat,which consists
chieflyof the palmiticester of cetylalcohol,CH3(CH2)i4CH2OH, and
would also be regarded as a
oil,which contains no glycerids,
sperm
wool
cluding
infat is chemicallya mixture of waxes,
Similarly
liquidwax.
of cholesterin and

the stearic esters

isocholesterin.

Beeswax

containingthe palmiticester of myricyl

The
secretion
of the anal glands of
alcohol,CH3(CH2)28CH2OH.
certain birds contains esters of octodeckylalcohol,C18H37OH.
is likewise in part

true

wax,

Cholesterins
36.

residue
of this group are found in the nonsaponifiable
fats.
In the animal organism they are
found widely

Substances

of various
distributed

through

normal

the

constituents

tissues in small
of

in wool

protoplasm.

amounts

As

and

are

ently
appar-

just noted, they

are

fats in combination

with stearic acid.

especiallyabundant
in
also
They are
widely distributed
plants. Their exact constitution
is stillunknown, but they contain a singlealcohol hydroxyl and apparently
Their
the
is
to
formula
belong
C27H44OH,
terpene group.
From
chemical
the
latter.
the
or
C27H460H, more
point
probably
of view, they are entirelyunrelated to the other groups
classified as
but biologically
their functions appear
to be closelyrelated
lipoids,
to those

of the other

ether-soluble

cell constituents.

Phosphatidsor Phospholipins
Quite closelyrelated to the fats are the
substances
known
which are
as
lecithins,
sometimes, although
called
Like
the fats, the lecithins
inexactly,
phosphorized fats.
37.

Lecithins.

"

of

glycerol.They differ from the fats in that

only two of the hydroxylsof the glycerolare replacedby fatty
acid radicles,the third being replaced by phosphoric acid
which is also in combination
with the nitrogenousbase cholin,
are

esters

derivative

addition

to

nitrogen.

of

glycol. The lecithins,therefore, contain, in

carbon, hydrogen and oxygen, both phosphorus and

NUTRITION

24

"
40.

Importance.
of the

matter

"

4.

animal

The

ANIMALS

Proteins

far the

By

body,
the

belonging to
these

FARM

OF

largershare

aside

organic

from

well-defined

stances
fat, consists of subof the proteins,
group

the results of

compounds, according to

of the

analysesrecorded

subsequentpages (98),making up from 17.5 to 21 per cent

These
of the fat-free body.
substances
characteristic of
are
of plants. Biologithe animal body, as the carbohydratesare
cally,
both
they are of prime importance to
plantsand animals,
since they form
the basis of the cytoplasm and nucleus
of
every livingcell.
on

41.

Nomenclature.
has

molecule
and

even

Accordingly,the
substances

"

until

The

has been

been

for

but

of

structure

quite recentlybeen

yet has
basis

chemical

very

almost

the

tein
pro-

known
entirelyun-

partiallyunraveled.

scientific classification of these

As

of

therefore,
necessity,
the nomenclature
hitherto followed has been based chiefly
their physicalproperties,
their solubilities
on
more
particularly
and coagulationtemperatures.
Naturally,such a classification
has been far from satisfactory
and this has been the more
true
of the difficulty
of accuratelyseparating the different
account
on
or
proteinseither by precipitation
crystallization.
there has existed a great and confusingdiversity
Accordingly,
in the nomenclature
of the proteins,
and uniformityis stillfar
from having been reached.
desirable
For the present, it seems
to

follow

lacking.

the classification and

matter

nomenclature

which

has

been

adopted provisionally
Society1
by the American
Physiological
This
and
the American
Society of BiologicalChemists.2
nomenclature
rejectsentirelythe term proteidas ambiguous
of the wide diversity
in its use, and employs protein
account
on
a
as
general term to signifythe group of substances which,
accordingto the nomenclature
adopted by the Association of
American
AgriculturalCollegesand Experiment Stations in
1898,3was called proteids. In other words, proteinunder the
new
plan excludes altogetherthe non-protein nitrogenous
substances
1
2
3

of

plants and animals.

Physiol. Soc, Amer. Jour. Physiol.,21 (1908),xxvii.

Soc. Biol. Chemists, 1, 142.
Proceedings, Amer.
U. S. Dept. Agr., Office of Expt. Stas.,Bui. 65, pp. 11 7-123.

Proceedings,Amer.

The

OF

COMPONENTS

THE

this

proteinsin

2.

3.

defined

Simple proteins are

acids

their

or

to

some

"

molecule

other

upon

contain

which

yielding only
gated
hydrolysis. Conjucule
the
protein mole-

those

as

derivatives

those

proteins are
united

into

25

Simple proteins
Conjugated proteins
Derived proteins

1.

amino

subdivided

are

sense

ANIMALS

AND

PLANTS

or

molecules

otherwise

than

proteinsare the productsof the hydrolytic

of
and include a wide range
cleavage of the proteinmolecule
altered proteinto the peptids.
substances,from slightly
42.
Physical properties. In the dry state, the proteinsare
tinted substances.
in generalwhite or slightly
They are usually
of them
have also been obtained in
amorphous, but a number
in nature.
found crystallized
form and some
the crystalline
are
soluble in water, others only in salt
of the proteinsare
Some
solutions or in acids or alkalies. They are insoluble in- most
other ordinarysolvents.
The proteinsbelong to the class of colloids,
i.e.,
they do not
and are claimed to have no osmotic
diffuse through membranes
free from electrolytes.Colloids in generalexist
pressure when
known
a
in two forms, a liquidform, technically
a sol,and
as
solid form called a gel,the difference being well illustrated by
colloid is distributed
the familiar substance
a
gelatin. When
through water so as to form an apparent solution the latter is
to be regarded
known
the proteins
are
as
a hydrosol. Whether
as

salt.

Derived

"

as

soluble in water,

or

whether

realitya suspension,has
shown, however, that

been

their apparent solution

It has
discussed.
much

these solutions

are

conductors

of

is in

been

electricity

they are true solutions. It may

be said, however, that no
sharp boundary exists between a
of
but that an indefinite number
true solution and a suspension
of convenience,
intermediate
stages is possible.As a matter
however, we may speak of solutions of the proteins.
from their solutions
be precipitated
Different proteinsmay
acids,alkalies and metallic
by various reagents, particularly
has been largelyused
salts. Ammonium
sulphate,especially,
of
for the purpose
of separatingdifferent proteinsby means
fractional precipitation.

and

it has been

concluded

that

26

NUTRITION

OF

FARM

ANIMALS

43.

Coagulation.

An

"

is that of

coagulation.

egg albumin

be heated

of

important property
instance,if

For

550 C.

solution

the

proteins
ordinary-

of

the albumin

beginsto separate
in an insoluble form and at about 6o" C. the precipitation
is
complete. This change differs from the change in the case of
gelatinsolutions from liquidto solid in being irreversible,
i.e.,
be changed back to the soluble form.
coagulated proteincannot
It should be noted that this change is entirely
distinct from the
of proteinsby means
of ammonium
precipitation
sulphatefor
of the change is unknown, but it
example. The exact nature
would

All forms

of

the chemical
obtained

be in part chemical

to

seem

standingpass
"

proteinappear

sense

from

to

of the word.

solutions
more

form.

to

or

The

be

subjectto coagulationin
the precipitated
proteins

Thus

at first in the colloidal form

are

less

in character.

rapidlyinto
is true

same

but

on

"

the

of the

natured
decoagulatedor
solid proteinslike

fibrin,etc. The coagulatedproteinsare insoluble in water

and salt solutions,
but may
be dissolved in acids or alkalies.
The
44.

Composition.

simpleproteins

The

pounds
simpleproteinsdiffer from the comin the previous sections in containing,
considered
in
addition to carbon, hydrogen and oxygen,
the elements
gen
nitroand sulphur. Notwithstandingthe considerable variation
in the properties
of the different simpleproteins
and the notable

differences which

"

have

been

shown

to

exist in their chemical

structure, their

elementary composition differs but little.

Cohnheim
quotes the followingfiguresfrom Michel for the
compositionof serum
albumin, which is in many
respects a
animal
typical
protein.
1

Carbon

53-o8

Hydrogen
Nitrogen
Sulphur
Oxygen

7.10
iS-93
1.90
21.99
100.00

The
as

variations

stated

by

in the
*

Cohnheim

Chemie

The

der

elements
percentages of the principal

and

by

Plimmer

Eiweisskorper,2d Ed.,

Chemical

Constitution

of the

are

"

p. 151.

Proteins,Part I, p.

2.

COMPONENTS

THE

AND

PLANTS

OF

ANIMALS

27

Plimmer

5i-55%

Carbon

Nitrogen
Hydrogen

15-17%
7%

Sulphur

0.4-2.5%

0.4

rule,the vegetableproteinscontain
of nitrogen than do the animal
proteins.
As

45.

the

proteinsis

very

large,but

latter

given concordant
weight of 16,666, from
the

pressure.2

For

weight of

the

result

results

which

the

for edestin

to

like those

mum
indicatinga minihas been computed

of determinations

5000

and

8000

figure10,166

is

These

14,500.

has

within

marked

minimum

been

figuresare

comparatively few

justmade

of its osmotic

For

albumin,
ever,
of value, how-

of the

years,
the limits of our

lar
molecu-

obtained.

reported,for

as
chiefly
showing the complex nature

Up

the

of this result has

globinof haemoglobin,a

between

albumin,

of

haemoglobin (a conjugated protein)by

of

reportedas

of the

Determinations

made.

of

weights are

determinations
satisfactory

C758Hi203Ni96FeS2. Confirmation

the formula

5378, and

very
been

structure

their molecular

have

molecular

serum

no

have

structure

methods

been

yet

as

"

and

molecular

The

proteins.

complex

very

magnitude

molecular
two

of the

Structure

higherpercentage

egg

cule.
proteinmole-

generalstatements
knowledge of the

proteins. The masterly researches of

Emil
Fischer, however, and especiallyhis creation of new
experimentalmethods, have resulted in a very great advance
in knowledge,and to-day,thanks to his labors and those of a
in applying and improving his
of investigators
large number
methods, we possess a fairlydefinite generalconceptionof the
of
of the protein molecule.
As in the investigation
structure
been
lines of attack have
chemical compounds in general,two
the
followed, viz., a study of the products resulting from
to synthesizethe
and attempts
splitting
up of the molecule
compositionand
compound from simplersubstances of known
chemical

nature

of the

structure.
1

Four

to

five per

cent

in keratins.

Zentbl. Physiol.,
21, 73".

28

NUTRITION

Hydrolysis of proteins.

46.

"

when
undergo hydrolysis

by various

or

FARM

OF

enzyms

ANIMALS

The

acted upon
such as the

simple proteinsreadily

by strong acids or alkalies,

pepsin of the gastricjuice,

pancreaticjuice,etc. These various agents

effect a succession of cleavages and hydrationsresultingin a
creasing
series of products of decreasingmolecular
complexity and inmodified proteins
solubility,
rangingfrom very slightly
through the so-called proteoses and peptones to still simpler
trypsinof

the

the

substances.
the

When

hydrolysis,
acid hydrolysis,
of the simpleproteinsis pushed as
especially
far as possible,
there result a number
of comparativelysimple
for all
substances which are qualitatively
the same
crystalline
proteinswith a few exceptions,
although the proportionsof the
from
various products obtained
different proteinsvary
terially.
maIt is believed,therefore,
that the proteinmolecule
is
built up of these final products of hydrolysis,
the so-called
buildingstones."
These primary cleavageproducts of the simpleproteinsare
Cleavage products of proteins.

47.

"

"

all

amino

glycinor
:

acids.

of the first of them

One

be isolated

was

mula
acid, representedby the followingfor-

aminoacetic

"

CH3

CH2

"

NH2

COOH

COOH

Acetic acid

The

to

Glycin

other

cleavageproductsof the simple proteinsmay be

regarded as derived from glycinby the replacement of one
atom
of hydrogen in the CH2 group
by various atomic groupings.
In all of them the NH2
position
occupiesthe same
group
in the molecule
COOH
in glycin,the
relative to the group
as
so-called a position. The
atomic grouping

I
CH

"

CO

is therefore

common

general chemical
which

they

are

NH2
OH

to all of these bodies

behavior
derived.

as

well

as

and

determines

that of the

their

proteinsfrom

COMPONENTS

THE

The

into

amino

two

acids derived

from

classes ; the monamino

NH2

containing one
NH2

two

group,
To these

groups.

the
has

proteins may be divided

acids,of which glycin is typical,
and the diamino acids,containing
there

are

be added

to

certain

acid.
Glycin,C2H5NO2, or aminoacetic
COOH
CH2
(NH2)
Alanin, C3H7NO2, or a-aminopropionicacid.
COOH
CH3
CH(NH2)
"

"

"

3.

29

Monoaminomonocarboxylic acids

"

2.

ANIMALS

gives the followinglist of

amino
acids which have been identified with certaintyamong
cleavage products of the proteins. The presence of others
been claimed by several investigators.
A.

1.

AND

the

Plimmer

heterocycliccompounds.
the

PLANTS

OF

Valin, C5HnN02,

a-aminoisovalerianic

or

acid.

CH3\
CH

CH(NH2)

"

COOH

"

CH3/
4.

Leucin, C6Hi3N02

a-aminoisocaproicacid.

or

CH3\
CH

CH2

"

CH(NH2)

"

COOH

"

CH3/
5.

Isoleucin,C6Hi3N02,

or

acid.
a-amino-|3-methyl-/3-ethyi-propionic

CH3v
CH

CH(NH2)

"

COOH

"

C2H5/
acid.
Phenylalanin,C9HnN02, or jS-phenyl-a-aminopropionic
COOH
CH2
C6H5
CH(NH2)
or
/3-parahydroxyphenyl-a-aminopropionic
7. Tyrosin, C9HnN03,
acid.
COOH
HO
C6H4
CH2
CH(NH2)
acid.
8. Serin, C3H7NO3, or /3-hydroxy-a:-aminopropionic
COOH
CH(NH2)
CH2(OH)
dicysteine,or di- (/3-thio-a-aminopro9. Cystin, C6Hi2N204S2, or
HOOC
S
S
COOH
CH2
CH2
pionicacid)
CH(NH2)
CH(NH2)
6.

"

"

"

'

"

"

"

"

"

"

"

"

11.

"

"

"

"

Monoaminodicarboxylic acids

B.
10.

"

acid.
Asparticacid,C4H7N04, or a aminosuccinic
HOOC
COOH
CH2
CH(NH2)
Glutamic
acid,C5H9NOi, or a-aminoglutaricacid.
HOOC
COOH
CH2
CH2
CH(NH2)
1

The

Chemical

"

"

"

"

Constitution

"

"

"

of the Proteins, Part

I, 2d Ed.,

1912.

NUTRITION

30

OF

Arginin,C6Hi4N402,
HN

"-amino-y-guanidinvalerianic

or

Lysin, CoHi4N202
H2N

"

"

or

CH2

a,

CH2

CH2

"

"

CH2

CH(NH2)

"

COOH

"

e-diaminocaproic acid.

CH2

"

CH2

"

CH2

"

CH(NH2)

"

COOH

"

Heterocyclic
compounds

D.

Histidin,C6H9N302,

14.

acid.

C/NH2
NH

13.

ANIMALS

Diaminomonocarboxylic acids

C.
12.

FARM

or

acid.
/3-imidazol-a-aminopropionic
CH

^
N

NH

CH(NH2)
-pyrrolidin
carboxylicacid

CH

Prolin,C5H9N02,

15.

or

CH2

CH2

"

"

COOH.

"

CH2

"

CH2

CH

"

COOH

/
NH

16.

Oxyprolin,or oxypyrrolidinecarboxylicacid.

17.

QH9N03
acid.
Tryptophan, CnHi2N202, or /3-indol-a-aminopropionic
C

CH2

"

CH(NH2)

"

"

COOH

/x
CH

C6H4

NH
48.

have

Synthesis of proteins.
shown

of the

of

one

that the amino

acids which

Fischer

"

and
the

result from

others

cleavage

simpleproteins may combine with each other, the NH2

of the other with the
unitingwith the COOH
group

elimination
of amino
exact

Peptids.

"

of

one

tnolecule

water.

As

as

many

18 molecules

in this way,
although the
less
of the resultingcompounds is still more
or

acids have

structure

of

been

combined

uncertain.
The

compounds of the
have received
artificially

amino
the

di-,tri-,
etc., being used
acid molecules
entering into

acids which

generalname
to

the

indicate

prepared
peptids,the prefixes

have
of

been

the number

compound.

The

of amino
term

poly-

Table

The

4.

Products

Cleavage

"

results shown

ANIMALS

FARM

OF

NUTRITION

32

in the

of

Proteins

foregoingtable

typical. In

are

proteins,certain amino acids have not been found at

For example, no glycinhas been found among
the products of
of gliadin,
the hydrolysis
casein and no lysin
or
zein,albumin
those of gliadin
or zein.
Furthermore, the proportionof
among
all.

few

the various

cleavage products
of the most

one
proteins,

acid which
of wheat

to

abundant

more

51.

Classification.
to

The

ranges

littleover

enter

sign +

determined.

determined

into

quite variable

in egg albumin, and is notably

in vegetable than in animal
proteins.
9 per

cent

the present purpose,

it

seems

full description
of the various

that the substance was

signifies
blank
simply indicates that

but does not

in the different

strikinginstances being that of glutamic

from nearly44 per cent in the gliadin

For

"

is

show

that it

was

not

present but
the

was

ingredient in

present.

fluous
super-

simplepro-

quantitatively
not
question was

not

THE

teins.

The

COMPONENTS

principal
groups

designatedas

"

into which

are

Albumins

ANIMALS

they are

33

subdivided

are

simple proteins soluble

coagulableby heat.
they include the albumins

serum.

AND

These

"

PLANTS

and

water

follows:

Albumins.

a.

OF

have

also been

Besides

the

of blood

found

in pure
familiar egg albumin,
and

serum

in small

of milk
in

amounts

great varietyof

pea,
b.

seeds,includingthose of wheat, rye, barley,

vetch, soybean and cowpea.
Globulins.
The globulinsare
simple proteins insoluble
"

in pure water
but soluble in neutral solutions of salts of
bases
with strong acids. Globulins
found in the
are

and

the blood

and

serum

in the muscles

and

other

strong

lymph

organs,

but

they appear to be especiallycharacteristic of the vegetable

in a largenumber
kingdom, occurringin considerable amounts
of seeds.
Osborne1
gives a list of 15 globulinsoccurringin 24
different speciesof seeds and enumerates
additional species
12
which
contain
have
globulinsto which no distinctive names
yet been given.
Glutelins.
These are defined as simpleproteinsinsoluble
c.
in all neutral solvents but readilysoluble in very dilute acids
and alkalies. The only well-defined members
of this group
at
the gluteninof wheat and the oryzenin of
are
present known
to believe that similar prorice,although there seems
reason
teins
"

exist in the seeds of other cereals.

d.

Prolamins,or alcohol-soluble proteins. The typicalmember

of this group
is the gliadinof wheat
and
the name
has
been applied by some
authors to the entire group, but the
term
prolamins,proposed by Osborne, seems
preferable. The
in
prolaminsare soluble
relatively
strong alcohol (70-80percent)
"

but insoluble in water, absolute alcohol and other neutral solvents.

They are characteristic of the seeds of the cereals,
the

principalprolamins being the gliadinof wheat and

hordein of barley,the zein of maize and the bynin of
e.

Albuminoids.

This

"

name,

formerlyused

to

rye, the
malt.

able
consider-

with proteins,is now

practicallysynonymous
applied to two groups of nitrogenous substances which have
been otherwise designatedas the collagens,
and
or
gelatinoids,
the keratins.
Albuminoids
defined as simpleproteinswhich
are
extent

possess

as

the
essentially
1

The

same

chemical

structure

Vegetable Proteins,p. 78.

as

the

other

34

NUTRITION

proteinsbut

are

characterized
form

They

neutral solvents.

appendages and

its

the

in all
by great insolubility
principal
organicconstituents

definition does

is,however,

artificialderivative

the

not

also been

constitutes the

which
provide for gelatin,
of collagen. Besides gelatin

organicbasis

of

and
cartilage
the ligaments;

of

epidermal tissues

such

the keratins

of the

chondrin,

are

bone; elastin,the characteristic component

and

ing
cover-

called sclero-

of this group

important members

more

which
collagen,

or

and of their external

have

hence

proteins. This
an

ANIMALS

of animals

of the skeletal structures

and

FARM

OF

hair,wool,

as

feathers,horns, hoofs,etc.
The
52.

Nucleoproteins.

conjugatedproteins
"

In

of classification here

the scheme

"

These
defined as follows :
are
(41),the nucleoproteins
proteinsare especiallycharacteristic of the nucleus of the
cules
vegetableand animal cell (74). They consist of proteinmoleof the compounds known
united with one
as
or
more
ing
These are
nucleic acids.
complex organiccompounds containalso a xanthin group."
a phosphoricacid radicle and
The simpleproteinsof the nucleoproteins
apparentlymay be
of the
and belong to various groups
of quite diverse nature
specialinterest of the nucleoproteins
simple proteins. The
attaches to the nucleic acids enteringinto their composition.
These
acids.
53. Nucleic
compounds contain in addition
the element
to carbon, hydrogen, nitrogen and
phorus.
phosoxygen
Their constitution has not yet been fullyworked
out,
but their cleavage yieldsfour classes of products,viz.,
followed

"

i.
2.

3.
4.

According to
the

the recent

acid

nucleic

Xanthin, or purin,bases
Pyrimidin bases
A pentose carbohydrate
Phosphoric acid

molecule

of
investigations
may

be

regarded

combinations
nucleosids,or glucosid-like
with
nucleosid

of

purin or pyrimidinbase. By
with phosphoric acid there is
a

Finally,the

most

common

nucleic

Levene

acids

and

others,

built up

as

pentose

from

drate
carbohy-

the union

of such

formed

nucleotid.

are

tetranucleotids.

which

OF

COMPONENTS

THE

always

seem

to contain

PLANTS

both

AND

ANIMALS

purin and

35

pyrimidinnucleo-

sids.
54.

Glycoproteins.
of the

The

"

"

glycoproteinsare defined

proteinmolecule

with

substance

other

containing a carbohydrate group

The
principalcompounds of this

than

are

group

as

pounds
Com-

substances

or

nucleic

the mucins

and

acid.
the

mucoids."
55.

Phosphoproteins.

"

These

are

the

defined

compounds of
yet undefined,phosphorusas

protein molecule with some, as

containing substance other than a nucleic acid or lecithin.
The
of the most
casein,or rather caseinogen,of milk is one
familiar and important of this group.
56.
The
haemoglobins are compounds of
Haemoglobins.
similar substance,
the protein molecule with haematin or some
"

and

constitute the red

57.

with

of the

"

protein molecule

lecithins.
The

58.

of the blood.

coloringmatter
Lecithoproteins.
Compounds
derived

proteins

protein derivatives.

Primary
formed

through

"

Derivatives

hydrolyticchanges which

of

involve

protein apparently
only slight

alterations of the molecule.

Proteans.

Insoluble

"

products which

apparently result

from

the

incipientaction of water, very dilute acids or enzyms.

Metaproteins. Products of the further action of acids and alkalies
whereby the molecule is so far altered as to form products soluble in
"

very
group

weak

acids

and

alkalies but

will thus include the familiar

insoluble
"acid

in neutral

proteins"and

fluids.

This

"alkali proteins,"

the salts of

proteinswith acids.
Coagulatedproteins. Insoluble products which result from (1)the
action of heat on
their solutions,
the
or (2) the action of alcohols on
protein.
59. Secondary protein derivatives.
of the further
Products
hydrolyticcleavageof the proteinmolecule.
Soluble
in water, uncoagulated by heat, and
Proteoses.
cipitated
prezinc
or
by saturating their solutions with ammonium
sulphate.
Peptones. Soluble in water, uncoagulated by heat but not precipitated
by saturatingtheir solutions with ammonium
sulphate.
of two
Peptids. Definitelycharacterized combinations
or
more
amino
the
acids,
carboxyl group of one being united with the amino
of
the
other
with the elimination of a molecule of water
group
(48)
not

"

"

"

"

"

36

NUTRITION

"
60.

and

Occurrence.

"

contain

animals

considerable

Non-proteins

5. The

addition

In
a

to

the

proteins,both

great varietyand
of

amounts

ANIMALS

FARM

OF

plants
relatively

sometimes

nitrogenous compounds of the

the

most

of such

pecially
compounds, esfrom an early day, it was
in feedingstuffs,
known
was
relatively
cant
insignifipresent were
long assumed that the amounts
in regarding all
involved
and that no material error
was
the nitrogenof a feedingstuff as existingin the form of protein.
Accordingly,the total nitrogenmultipliedby the conventional
crude protein was
taken as
factor 6.25 and designatedas
of the material.
The
representingthe true protein content

While

diverse nature.

occurrence

"

researches of

Scheibler,E. Schulze and

however, showed
was
were

that

found
very

one-third

"

that

this

was

Kellner

far from

nitrogenous substances

widely distributed

that

even

of the

seventies,

being

the

other

than

sometimes

as

case.

It

protein
much

as

nitrogen of feeding
These
results
stuffs existed in these non-proteincompounds.
and it
have been fullyconfirmed
by subsequentinvestigations
between
these
to distinguish
has therefore become
necessary
and the true proteins.
substances
these nitrogGeneral
61. Definition.
properties. While
enous
varied
compounds other than protein are of the most
less
nature, they all differ from the proteinsin having a much
Complex molecular structure.
Many are comparativelysimple,
of them
substances,most
crystalline
readilysoluble in water
and diffusible,
and they appear
inferior in nutritive
distinctly
value to the proteins. It is a matter
of practical
convenience,
have
them
to
which
to distinguish
a group
name
therefore,
by
and for this purpose
the term non-proteins
has been proposed.
It is,of course, a contraction for non-proteinnitrogenoussubstances
and means
simply substances which contain nitrogen
but are not proteins. It therefore includes a great varietyof
for
a
cover
compounds and may be considered as in a sense
The
our
more
ignorance of their exact nature.
important
of
non-proteinsare:
groups
or

one-half

and

in the

total

"

"

The

nitrogenousmuscle extractives
The nitrogenouslipoids

THE

COMPONENTS

The

OF

PLANTS

AND

ANIMALS

37

nitrogenousglucosids
and organicbases

Alkaloids
Amino

acids and

Nitrates and

62.

The

muscle

muscle
bases

extractives

includes

and

more

important nitrogenous

creatin,creatinin

are

and

the

purin

"

number

of

contained

noted

(37-39), the lipoid

compounds, classed as phosphatids
As

lipoids.

the lecithins.

are

The

"

which
contain
cerebrosids,
with fatty acid radicles.

this group

salts

hypoxanthin.

Nitrogenous

group

ammonium

extractives.

xanthinand

63.

amids

nitrogenous group

The

The

familiar members

most

actual

either in the animal

in combination
of

stances
of these sub-

amounts

plant are small and

important fraction of

or

their

nitrogen does not constitute any

the total nitrogen of the body or of the feed.
64.

Alkaloids

and

organic bases.

Alkaloids

"

are

tively
compara-

in

agricultural
plants,the seeds of the lupineforming
the principalexception. The
the other
organic bases, on
hand, appear to be somewhat
widely distributed. In addition
"
hexon bases
to the so-called
derived
arginin,lysinand histidin,
from the proteins and nucleo-proteins,
the bases cholin,
and stachydrinhave been found in a variety
betain,trigonellin
of plants.
rare

"

65.

The

Nitrogenous glucosids.
"

characteristic

of

substances

the

of this group
They contain

vegetable kingdom.
a
nitrogenouscompounds coupled with simple sugars.
The nitrogenousglucosidsdo not appear
to be especially
dant
abunin the ordinary feeding stuffs of domestic
animals
and
where they do occur
rather by their specific
are
distinguished
physiologicaleffects than by their nutritive value in the ordinary
are

varietyof

sense.

which

have

66.

Amino

the most

Schulze1

E.

been

found

acids

abundant

and

forms

mentions

in various
amids.

"

bodies

seven

of this class

plants.
These

substances

of

are

by

far

non-proteinin vegetablematerials.
The first one
in 1805, in asparto be discovered was
asparagin,
agus
this
and
substance has since been found in a large
shoots,
number
of plants or parts of plants. Glutamin, a second
in plants.
amid, is also of frequentoccurrence
1

Jour. Landw.,

52

(1904),305.

38

NUTRITION

OF

FARM

ANIMALS

the amids of aspartic

and
Asparagin and glutamin are respectively
which
constituents of the proteinmolecule.
are
glutamicacids,both of
COOH

COOH

COOH

COOH

CH2

CH2

CH2

CH2

CH2

CH2

CH

"

CH

NH2

"

NH2

COOH

CO

Aspartic acid

CH

NH2

"

"

CH

NH2

COOH

less

or

thus

NH2

Asparagin

Glutamic

It has

"

used

commonly

that the term

about

come

as

"

NH2

Glutamin

amids

has been

for the
generaldesignation

nitrogenoussubstances
however, is unfortunate.
class of chemical

CO
acid

in

found
The

substances

feedingstuffs.

word

amid

denotes

more

tein
non-pro-

The

usage,

distinct

of which

only asparaginand glutamin

be
in
to
common
especially
plants,while the
appear
latter contain a varietyof nitrogenoussubstances
which
are
not amids at all. The general
term
proposedabove,
non-protein
therefore,seems
preferable.
In addition to asparaginand glutamin there have been found
of the cleavageproductsof the
in feedingstuffs a largenumber
ten amino
proteins. E. Schulze **" enumerates
acids,viz.,valin,
leucin,isoleucin,phenylalanin,tyrosin,prolin,tryptophan,
besides the purin bases xanthin,hyarginin,
lysinand histidin,

poxanthin,adenin and guanin, as

and carnin,as having been isolated
and

Hart

Bentley

of the water-soluble
as

amounted

amino
to

Occurrence.
relation
be in

to

from

acids

found

well
from

guanidin,allantoin
various vegetablematerials.

as

that from

50

cent

to

20

per cent.

evidentlystand in a close
to the proteinmetabolism
of the plant. They appear
of protein
part intermediate productsin the synthesis
"

These

nitrates and

substances

ammonium

in part to be formed in
for their translocation and

salts and

cleavageof proteinsnecessary
resynthesis
duringthe processes of growth. They

are

especially

2
Ztschr. Physiol.Chem., 45 (1905),38.
(1904),305.
4
Jour. Biol. Chem., 22 (1915), 477Physiol.Chem., 47 (1906),507.

Jour. f. Landw.,

3Ztschr.

70 per

isted
nitrogenof a varietyof feedingstuffs exwhile the amid nitrogenproper
or
peptids,

only 10

the

to

52

not
great variety of ingredients

Similarly,plantscontain
strictlybelonging to any

of

aggregate, these substances

do not

food

ANIMALS

FARM

OF

NUTRITION

40

the

four

main

often add

feeding stuffs,but, on
cases
considerablymodify

they may in some

of
the activity
or

the

the various processes of nutrition

these
results of feeding. Until recently

the actual

have

feeding stuffs

of

received

other

their

the

the potential

greatlyto

of

value

In

groups.

hand,

palatability
and

affect

so

gredien
secondaryincomparativelylittle

attention.

Organic acids.

69.

Aside

"

the small

from

of free

amounts

fattyacids occurringin most native fats,both animal and vegetable

found
(29,33),the acids of this series are seldom or never
In those feeding stuffs which
in native feedingstuffs.
have
undergone bacterial fermentation,however, notablyin the case
less acetic and butyric acids occur, but the
of silage,
more
or
is lactic acid,
principalacid product of such fermentations
C3H603. The same
acids, along with formic and propionic
of ethyl aldehyde,likewise result
amounts
acids and minute
of the carbohydratesof the
from the bacterial fermentation
and thus constitute a not unfeed in the paunch of ruminants
important
portion of the non-nitrogenousmaterial resorbed
from the feed (128-132). The
principalorganic acids found
in native feedingstuffs are
malic, tartaric,citric and oxalic,
usuallyas the potassium,sodium or calcium salts.
70.

of

Ethereal

complex

oils in their

in

oils.

molecular

The

so-called ethereal oils are

of

resembling the true

tilled
which can
readilybe disexamplesare the so-called

Familiar

steam.

peppermint,lemon, anise,and
they have any direct nutritive

to the flavor and

add

odor

of

the

stimulate

to

the

hay, for

ethereal

oils. To

and

71.

the

sense

food
of

cases

The

the

of

lieved
be-

agreeable

vorable
part its fa-

in

erties
prop-

ethereal

oils

in

and

general.

"

What

is called

feedingstuff depends largelyupon

smell of a great varietyof substances
or

are

resembling in
class

same

they

whose deleterious
garlic,
dairyproductsis so well known.

Flavoring substances

flavor of
on

the flavor of

some

doubtless

substances

belong the oils of mustard, onion

effect upon

in

known

but

themselves

digestiveprocesses.

example,
dietetic effect,
is due to

good

value

of feeds and

aroma

It is not

the like.

oils of

that

substances

nature, somewhat

physicalpropertiesbut

current

"

the

the action
either

con-

tained

foods,

bitter
of

or

whose

and

properties
The

known.

the

from

aside

gums,

is

palatability
direct

any

little

feed,

these

upon

nutritive

is
cated,
indi-

factor

will

etc.,

ingredients,

accessory

effect

etc.,

already

as

of

nothing

or

important

an

cooking.

resins,

waxes,

of

case

variety

great

of

the

during

effects

palatability

dependent

that

fact

contain

41

in

especially

physiological

and

flavor

usually

are

while

substances,

astringent

ANIMALS

developed

or

feeds

stock

oils,

AND

or,

added

artificially

ethereal

Besides

PLANTS

originally

material

the

in

human

OF

COMPONENTS

THE

in
in

appear

trition
nu-

later

discussions.
Vitamins

72.

been

yet

devoted
rather

their

ill-defined

and

are

properties
in

and

their

growth

few

past
of

group

however,

considered
maintenance

the

vitamins

chemical

Much

substances.

attention

has

"

during

investigators
substances,

Growth

by

food
others

known
and

by
may

relations

(438,

to

years

498,

important

an

constituents

effects

therefore
to

738).

by

substances.

growth
their

called

but

the

more

some

These

rather

be

as

than

by

ately
appropri-

requirements

for

CHAPTER

THE

OF

COMPOSITION

ANIMALS

"
Definition.

73.

of all life.

unit

exist.

can

relation

molecule
in its

seen

in which
As

united
to

cell

form

to

as

that

of the

cells

the

such

74.

cell

Structure

body,

of

distributed

of form

in

the

Fig.

cells.

of

from
cell
a

the

cells

of labor

and

the
The

extreme.

likened
of the

single citizen

forms

of animal

i.

The

"

remainder

of

part
the

nucleus.

All forms

entire active
of

dissolublyassociated

and

consists

cell

typical

is the
often

of

separate

It

should

is not

be

The

cell.

more

cell

the

was

as

well

but

as

animal,

through

by Huxley

understood, however,

chemical

not

protoplasm is often
cell,i.e.,to cytoplasm plus

manifested

called

or

name

part of the

and

the
The

nucleus.

somewhat

to

serves

life,vegetable

with

which

protoplasm,

of life.

the

of

that

to

distinct
membrane,
surroundings. Sometimes
a
wall, is developed, especiallyin plants, although this is

necessary

or

are

an

been

are

being

its

applied to

of

singlecell.

numbered

diverse

or

than

of cells

cells

reach

the

complex

different

the

has

same

vital functions

functions

few

to

number

among

which
cytoplasm, within
peripheral portion of the cytoplasm is

compact

does

the

proteins for example.

organisms (protozoa)

organization

individual

living matter

performed by

are

biological

somewhat

physiological division

an

represented

are

of life

nation, in which

or

one

in unicellular

of

STUFFS

the

as

which

atom

higher organisms

highly specialized.

are

of

differentiation

organization of
state

the

individual, the various

while

corresponding

in

plant that

scale

the

In

defined

or

less extent

or

by myriads,

the

FEEDING

simplest
regarded as bearing

functions

one

groups.

be
form

simplest form

in

greater

such

all the

ascend

we

cell may

be

might

OF

Cell

It is the

the animal

to

organic
It is

It

AND

The

i.

The

"

II

biological

the

are

the activities

physical
the

that

word

It is

term.

42

in-

basis
toplasm
pro-

struc-

COMPOSITION

ture

OF

ANIMALS

substance.

than

rather

to

common

kinds

are

all

OF

AND

FEEDING

STUFFS

Moreover, there is

cells,but

as

many

not

43

one

toplasm,
pro-

protoplasmsas

there

of cells.

the
sharp differentiation between
those
of the cytoplasm. The
functions of the nucleus and
concerned in cell reproduction,
to be especially
nucleus appears
is

There

Fig.

1.

or

less

Different types of cell composing the body.

in Health and Disease.)

"

the formation
nucleus

more

of

of

new

continued

an

of

at

least

beginning with a division of the

and being followed by a division

minimum

function
the

cytoplasm, on

of it is essential

amount

existence of the nucleus.

the labors of the

of the

nutrition of the cell,and

to

the
the

the present purpose,

further discussion of those finer
For

it is unnecessary
to attempt a
details of the structure
of the cell which

by

Horse

cell

existingc"U
main
of its cytoplasm. The
the other hand, seems
to be
presence

(Hadley, The

have

been

and physiologist.
histologist

worked

out

NUTRITION

44

75.

FARM

OF

Composition of protoplasm.

ANIMALS

The

"

chemical

constitution

of

protoplasm is unknown,
partlybecause it is undoubtedly
living
because of its instability
and the impossibil
very complex but chiefly
of isolating
it without at the same
time destroying
its life. Moreover, it doubtless varies materiallyin cells of
different types. The
with each
proteins,perhaps combined
other
into
giant molecules," undoubtedly constitute the
basis and predominatingingredient
of protoplasm,but certain
and cholesterins),
ash ingredients
(lecithins
lipoids
(electrolytes),
and perhaps glycogen and other carbohydrates,in addition,
"

of course, to water, appear

In
to be also essential constituents.
the cytoplasm,the simpleproteins(41) seem
to predominate,

while the nucleus

is

characterized by the presence

of
especially
the nucleoproteins
(52).
As already indicated,the protoplasm
cell wall.
76. The
often developsa cell wall.
So far as
the speciesof
concerns
feed for farm
be said
as
animals, it may
plants which serve
that a vegetable cell is always surrounded
by a cell wall the
basic ingredient
of which
is the carbohydratecellulose,
stance
suba
found in the bodies of the higher animals.
not
In the young
and growing parts of plants,
the cell wall is thin
and consists substantially
of cellulose only. In certain parts
of plants,such as the cotyledonsand endosperms of seeds or
"

the tissues of succulent

tubers,the

and

roots

cell wall

remains

in mature
tissue. In other parts of
comparativelythin even
the plant,on the contrary, it becomes
thickened by
very much
of additional cellulose and especially
the deposition
of substances

other
to

be

than

cellulose.

These

other

essentially
carbohydratesor

general kinds.

The

which

readilyattacked

pure

of
The

are

more

cellulose and

reserve

second

material

substances, which
their derivatives,
are

first of these

of two

(18),

is the hemicelluloses

by hydrolyzingagents than
constitute to a large extent
a deposit

which
and

appear

include

both

and

hexosans

pentosans.

ligninand
belonging
cutin groups
(19,20),which serve to impartstrengthand rigidity
less impermeabilityto the cell wall.
or
along with more
They
in older plants as comabundant
pared
are, therefore,particularly
with

consists of substances

younger

support the plant,such

thickened

and

ones

cell wall is

as
seen

in those

the stem.
in wood.

the

to

organs

The
A

which

extreme

few

serve

form

of the

to

of the

numerous

AND

vegetable cells

of

forms

ANIMALS

OF

COMPOSITION

in

illustrated

are

STUFFS

FEEDING

OF

45

Figs. 43-45

of

Chapter XV.
Cell enclosures.

77.

cytoplasm and

of the
a

variety of

entered

has

which

the cell and

products of

of waste

cells have

many

may
is on

consist of food
its way

protoplasm, or,
activityon their
of

the power

to

in cells

on

material

porated
being incor-

the other

way

hand,

creted
being ex-

to

surroundingmedium.

the cell into the

from

observed

are

of

cell

essential constituents

the

designated as subsidiaryingredients

These

into the molecules

to

there

nucleus

substances

other

cell enclosures.

or

addition

In

"

Moreover,

storingup surplusfood,especially

material.
Such material
non-nitrogenoussubstances,as reserve
is not usuallyregarded as constitutinga part of the protoplasm
but as being simply included in it mechanically.
in the animal is fat,which
cell enclosure
The
most
common
is contained in large quantityin certain connective tissue cells
and
the

constitutes

in

also store

the

reserve

fuel material

of the animal
much

carbohydrates (glycogen) being

amount.
While
some
important groups

storage

limited

the

up

of

large amounts

of fat in their
materials

predominatingreserve

of

body,
more

plants

seeds,nevertheless

vegetablekingdom
carbohydratesof the

in the

carbohydrates,including the reserve

Starch is found in
cell wall and, as a cell enclosure,starch.
in seeds and in
abundant
all parts of plants,but is especially
stance
of this subthe starchyroots and tubers,where large amounts
Illustrations of plant cells containing
stored up.
are
starch are afforded by Figs. 43-45 of Chapter XV.
are

Both
and

because

the

animal

of the

chemical

composition of

the

cell wall

enclosures,carbohydrates are quantitatively

the predominating ingredientsof most
plants,while
of the cell

nature

cells and

"

tissues

2.

are

Animal

proteidor fattyin
chiefly
Tissues

and

character.

Organs

only do the cells of higheranimals

show
extreme
differentiation of form and function, but cells
having the same
generalnature and office are associated together
called tissues,
such as nerve
to form what
tissue,muscular
are
tissue,connective tissue and the like,each serving its own
These
specificpurpose.
tissues,again, are grouped together
78.

Classification.

"

Not

46

NUTRITION

form

to

such

organs,

liver and

the

as

AMIMALS

the

muscles, heart, lungs,stomach,

plex
performingits specialpart in the com-

as

like,each

interplayof

FARM

OF

for the lifeof the

activities necessary

organism

whole.

Since this is not

treatise

to
anatomy, it is unnecessary
consider in detail all the diverse types of tissue or all the various
a

on

body. It is desirable,however, that the

notion of the chemical
student of nutrition should acquiresome
make-up of the various parts of the body. For this purpose it
the followingscheme, based chiefly
will be convenient
to use
of tissues,
the functions performed by the different groups
on
the distinction between
tissues
which
extent
ignoresto some
and organs and which does not pretend to be an exact or exhaustive
making

organs

the

up

classification.
lage,
supportingtissues,
bone, tendon,cartiincluding
ligament,elastic tissue,etc.
tissues of motion, including the muscular
Second : The
First

The

tissues and
Third
and

the
The

and
respiration
:

The
The

Fifth:

tissue,those

the

or

system.

nervous

alimentation,includingthe tissues
in digestion,resorption,circulation,

tissues of
concerned

organs

Fourth

tissues

nerve

excretion.

epidermaltissues.
reserve

tissues, including,besides

tissues in which

glycogen is

more

or

adipose

dantly
less abun-

stored.
The
79.

animals

Intercellular

supportingtissues

substance.

certain tissues show

"

an

the bodies

In

of the

development

enormous

higher
of the

substance, so that the cells,instead of

in a mass
of noncloselyadjoiningeach other, are imbedded
cellular material which may vary greatlyin consistency.Sometimes

so-called intercellular

this intercellular substance

it usuallycontains

greater

or

is

entirelyhomogeneous

less number

of fibers imbedded

but
in

of
By virtue of the specialproperties
the intercellular substance, tissues of this sort perform primarily
mechanical
functions,maintainingthe form of the body
while the cells
or
servingto connect and support other tissues,

homogeneous

themselves

serve

basis.

to produce and
principally

substance.

The

organicbasis

maintain

the intercellular

of the latter is the

group

48

NUTRITION

OF

FARM

the total ash of the entire animal

of the

carcass

of the

animal
81.

Bone

83.01 per

in the bones.

of the ash

cent

gen
Of the total nitro-

eight of these animals 18.04 Per cent

bones.
Correspondingdata for the entire

in the

recorded.

not

are

and

of

carcasses

contained

was

contained

was

ANIMALS

ash.

But

"

is quitevariable,that of

while

the

compositionof bone itself

the bone ash is notablyconstant
in
even
predominant ingredientis tri-calcic

species. The
phosphate but it contains,also,calcium carbonate as well as
phosphates and carbonates of magnesium and other bases.
The average
compositiongiven by Zalesky' is as follows :
different

"

Table

5.

Composition

"

of

Bone

Ash

of

Species

Different

Turtle

%
Calcium

phosphate
phosphate
.

85.98
1.36

Magnesium
Calcium

combined
.

dioxid

5-27

analysesby

Gabriel2

yielded the

following

"

Table

6.32

....

detailed

More

results

C02

CI, Fl
Carbon

with

Neumeister

2Ztschr.

6.

"

; Lehrbuch

Physiol.Chem.,

Composition

der
18

of

Bone

Ash

PhysiologischenChemie, 1897,
(1894),257.

p.

456.

ANIMALS

OF

COMPOSITION

OF

AND

FEEDING

STUFFS

49

of

magnesium, sodium, potassium, carbon

of bone
to be as essential ingredients
dioxid and chlorin appear
ash as its calcium or phosphorus.
The

small

82.

Cartilage, ligament, tendon, elastic

of

the
in

down
end

amounts

tissue.

of the
cartilaginousground work
into bone.
the embryo is converted
of bones

surfaces

at

skeleton

all

laid

as

the
particular,
which in
cartilage,
adjoining bones,
In

of

jointconsist

Not

"

connecting link between

as, for example, the cartilageconnecting the ribs with
At
breast bone, thus allowing a limited degree of motion.
jointsproper, the bones are held in place,and the direction
extent
of their motions
limited,by the ligaments, while
other

which

muscles

the

forms

cases

body

attached

are

impart motion

to

serve

the bones

to

the

to

by

various

the
the
and
the

parts of

of tendons.

means

the intercellular substance of the

cases
many
these fibers
contains
fibers of elastin. When

In

supporting tissue
abundant

are

the

ligaments and tendons of the

A strikinginstance is
joints,which are almost inextensible.
afforded by the elastic band
(Ligamentum nucha) which runs
along the back of the neck of quadrupeds and supports the
Another
weight of the head.
example is furnished by the layer
tissue is elastic in contrast

of elastic tissue

gives them
blood

pumped by
"

The

in the walls

contained

certain

the

to

degree of resilience

of the

arteries which

to the pressure

of the

the heart.

"

sue,
organic portion of all these forms of supporting tisof
like the organicportion of the bones, consists essentially
different proteinsbelonging to the group
of albuminoids.
Connective

83.

tissue.

This

"

is

name

sometimes

applied

of

supporting tissue,since they also

to connect
the various organs of the body. In a more
serve
ordinary and limited sense, however, it is used to designate a
form of supporting tissue of which the most
familiar example
is the tissue lyingbetween
the skin and the underlyingmuscles,
to all the

or

various

lean meat,

forms

and

servingto

careful examination

shows

tissue is continuous

with

between

the

them

that
other
and

singlemuscles
and

connect

them.

of connective

tissue extends

into muscular

bundles

or

connect

them

together. A

this subcutaneous

similar
serves

Not

at

only

same

so,

but

time

extends
to

limit
de-

this sheath

itself,
dividing it
these againinto secondary

into the muscle

fasciculi and

connective

tissue which
the

more

NUTRITION

5"
The

fasciculi.

tissue of the interior of the

connective

unites at the ends and

is continuous

of which

means

form

the muscles

muscle

of connective

tendons, by

attached

are

the

to

(Fig.2).
of connective

similar sheath

with

already mentioned, viz., the

tissue

bones

ANIMALS

FARM

OF

of* the

the internal organs

them, forming

tissue surrounds
and

body

framework

extends

which

active tissues of these organs

as

into

supports the

well

as

the blood

lymphatics,etc.,so that it may be

said in a broad general way
that the body of a
higheranimal consists of a varietyof active tissues
and organs contained in and supportedby connective
tissue and the other forms of supportingtissue
alreadydescribed.
Like other forms of supportingtissue,
the connective
tissue consists of cells which have produced a
of intercellular substance,
relatively
largeamount
vessels,nerves,

If
Fig.
One

2.

of

end

fiber.

muscle

and
(Hough
Sedgwick,
The

Human

Mechanism.)

which

in connective
it is

however,
of fat

tissue consists

composed

of

also store

may

collagen. Cells
within

up

ically,
Chem-

fibers.

of connective

themselves

tissue,

largeamounts

(94).
Tissues

84.

chieflyof

The

animal

muscles.

"

Both

of motion
the

external

of

movements

an

and

those of the internal organs are effected by means

of the muscles, and the muscular
tissue is preeminently the
tissue of motion.
of the entire

nearly all

Moreover, the muscles

of the

mass

body

of

the

proteincontained in
The compositionof muscle
carcass.
is of special
interest.
85.

Structure

muscular

of muscles.

"

The

make

lean

the

up
animal

edible

and muscular

largepart

and

furnish

portion of

the

therefore,
tissue,

smallest anatomical

element

of

tissue is the

singlemuscle fiber. This is a highlyspecialized

and greatlyelongated,thread-like cell one
and a half inches
to one
It is enlong and having a diameter of from .0004 to .004 inch.
closed
in

nuclei.
"

are

very

thin

transparent

membrane

and

contains

many

hundreds or even
thousands
largenumber of these fibers
bound
togetherto form a fasciculus,the fibers running lengthwise
each
and overlapping
other,being generallyshorter than the
A

"

surrounded

by

its

bundles, each with

again being united to
also to

serves

in

connective

is

of

the lean meat

86.

various

lipoidsand

the

in greater

less

or

such

But

protoplasm.

tive
ac-

ing
considerof

the

limitation of the

entire

is

muscular

fibers,and
with

that

Since,however, the
muscular

tissue,
nerves,
less of the

Fig.

the nutrition

muscle

system,

structures, and

rather than

little suited

we

not
are

as

merely for

productionof

the

"

"

meat

muscle-fibers.

contains

variety of tissues
sense

narrower

"

lymph vessels,etc. (85)

connective
with

more

or

fluid

contents

of the

latter

composition is likelyto be variable.

especiallyof fat animals, contains
of fat

(Hough

concerned, therefore,with the

of the lean meat
whole
i.e.,

tissue in the

blood and

of

Part

"

fiber.

all

with

of the ultimate
lean

3.

Human'Mechanism

build

requiredto

composition of the muscles

aside from

rational from

In

animal, material

all

in

amounts

present purpose.

its accessory
the muscle

to

the

electrolytes

standpoint,is

anatomical

amount

the

extractives

tissue,however

muscular

the

to

accessory,

protein, meat

the

up

the

elements

structural

of water,
found

up

may

it may
probably be
tially
general wray that it consists essen-

as

said in

are

producing motion,

other

muscle

of the

moreover,

and

If

"

limited

be

fibers which

muscular

the

muscles.

of

tissue

muscular

agents in

to

lymphatics

and

material.

Composition

ultimate

an

connective

tuting
consti-

far from

animal, are

an

homogeneous

term

bundles

of fat.

largeror smaller accumulations

Evidently, then, the muscle as a whole,
the collective muscles
more
even
making

and

The

muscles.

blood

contain

term

tissue,these

vessels,nerves
abundantly supplied,and,

carry

51

united

the individual
the

STUFFS

previousparagraph (83),
into largerfasciculi,

tissue and

envelope of

form

muscle

the

which

with

FEEDING

OF

stated

connective

or

tissue

AND

as
fasciculi,

These

fasciculus.
are

ANIMALS

OF

COMPOSITION

even

"

it is evident

that its

Moreover, the lean meat,

a

considerable

after all the fat tissue which

and

variable

it is practicable

ever,
This fat, howseparate mechanically has been removed.
forms no part of the muscle proper but is simply a deposit

NUTRITION

52
of

individual

the

between
size from

the

largermasses
the scalpel.It

with

lean

between
of

content

fat,and

which

be

may

is necessary
in the commercial

meat

bundles

fibers and

muscular

of

masses

muscle

the

between

(94) developed

tissue

in minute

It is contained

material.

reserve

ANIMALS

FARM

OF

adipose
even

or

in

only
differing
trimmed

off

or

moved
re-

fore,
theredistinguish,

to

sense, with

its varying

in the stricter scientific sense,

lean meat

i.e.,the fat-free

The

muscle.

of the latter may

position
com-

be

tained
ascer-

by actually removing
ordinary trimmed

either

the fat from

the

and
of a solvent
by means
veniently,
conanalyzing the residue or, more
by analyzing the fresh
ically,
and removing the fat arithmetmeat
position
i.e.,by calculatingthe commeat

"

of the fat-free muscle.

Composition of fat-free muscle.

87.
The

composition of

lean meat

of butchers'

"

the

cuts

fat-free
has been

determined

Fig.

4.

Fat

"

cells in muscle.

(Bailey'sCyclopedia
Agriculture.)
meat

of the

Table

7.

"

of

entire

Average

1
2

can
Ameri-

by Henneberg, Kern and

Wattenberg1 for two old sheep and
six younger
ones
ranging from 6J to
termined
28 months
old, and Jordan2 has dethe compositionof the lean

carcasses

Composition

of four

of

steers.

Fat-free

Lean

Jour. Landw., 26 (1878), 549; 28 (il 5i),289.

Maine
Expt. Sta. Rpt. 1895, H, 36.

Meat

of

Sheep

8.

Table

ANIMALS

OF

COMPOSITION

of

Fat-free

No.

Water

53
Steers

of

Old
Average
All
of

No.

Meat

Mos.

32

No.

STUFFS

Lean

Old

Mos.

22

No.

77.61

76.60

78.01

77.18

77-35

21.37

22.30

20.94

21.77

21.60

(by

Total

nitrogenous matter
difference)

Ash

OF

Composition

Average

"

FEEDING

AND

1.05

1. 10

1.02

...

1.

1.05

100.00

100.00

100.00

05

100.00

figuresof the foregoingtables indicate but very slight

of the
differences in the composition of the fat-free lean meat
content
different animals, aside from
a
slightlygreater water
is 95 per cent total
in that of the sheep. An approximateaverage
stance.
and 5 per cent ash in the dry, fat-free subnitrogenousmatter
The

"n the

of

course

investigations
upon

merous
nutrition,nu-

human

analyses have been made of the various commercial cuts

in general confirm
the foregoingfigures and
which
of meat
the
show
relativelysmall differences in this respect between
different cuts.
88.

Elementary

composition

of

fat-free

meat.

"

The

lowing
fol-

and Langbein, and Aranalysesby Rubner, Stohmann

gutinsky show the ultimate composition of ash-free muscular

prolonged extraction

tissue after
Table

9.

Composition

"

Fat-

of

Carbon

Rubner

and

and

Argutinsky

Kohler

Langbein

....

muscular
The

ether

Hydro-

gen

Nitro-

52.02

7-30

52-33

7-30

Ash-free

gen

"

Muscular

Oxygen

16.30
16.36
16.15

53-40

Stohmann

hens.

with

24.32

Tissue

Heat
of
Combustion
Gram.
per
Cals.

5.6561
5.6409

24.22

investigatedthe elementary composition of the

tissue of cattle, sheep, swine, horses, rabbits and
material was
prepared with much care, the fat being
has

Ztschr. Physiol.Chem.,

31

(1901),479.

NUTRITION

54
removed

fully as

as

The

ether.

OF

FARM

ANIMALS

possibleby prolonged

residual fat which

extraction

be removed

cannot

with

in this way

determined

and a
by Dornmeyer's digestionmethod
in the analytical
results.
correction made
The

was

are

Table

his averages

io.

"

for the fat- and

Composition

of

Fat-

and

ash-free substance

Ash-free

Lean

responding
cor-

lowing
fol:

"

Meat

Heat
Combus-

of

tion
per

Gram.
Cals.

Cattle

Sheep

5.6776
5-6387

Swine

5-6758

Horse

5-5990

Rabbit

5.6166
5-6173

Hen

All
were

the

samples
found, except

which

an

accounts

tested

were

for

glycogen,but only

traces

horseflesh,for the two samples of

of 3.65 per cent was
obtained,a result which

average

in

for the low

the

figurefor nitrogen.

The

tissues

of alimentation

grouped the
and tissues directlyconcerned
supplying food to
organs
the organism,with its distribution through the body, and with
cludes
of waste
the removal
productsof cell activity. That is,it inthe organs
of digestion,
circulation,
tion
respiraresorption,
and excretion,
which constitute what are ordinarily
spoken
of the
animals.
So far as most
of as the entrails of slaughtered
familiar internal organs
concerned
of the animal
are
they may
sues
be considered as made
of the classes of tisup to a largeextent
In addition, however, the internal
already considered.
distinct type of tissue,
include a somewhat
viz.,glandular
organs
gestive
tissue,which plays an especially
important part in the difor
processes, while it is also of the highestsignificance
89.

other

Definition.

"

Under

bodily functions.

this

heading

may
with

be

56

NUTRITION

Table

12.

Percentage

"

between

OF

ANIMALS

FARM

Distribution
Carcass

Ash,

of
and

Protein

and

Fat

Offal

Protein

Ash

(NX

6.25)

Fat

calf

Fat

In

carcass

In offal
.

Half-fatox
In

carcass

In offal

Fat

ox

In

carcass

In offal

lamb

Fat
In

carcass

In offal

Store

sheep

In

carcass

In offal

Half-fatold sheep
In

carcass

In offal
.

sheep

Fat
In

carcass

In offal
.

Extra-fatsheep
In

carcass

In offal

Store pig
In

carcass

In offal

Fat

pig

In

carcass

In offal

Mean
In

of all
carcass

In offal
100.

100.0

1 00.0

ANIMALS

OF

COMPOSITION

OF

AND

FEEDING

STUFFS

57

that the offal contained

It thus appears
relativelymore
ash
than
less
From
and
fat
the
and water
and
carcass.
the

weightsof

and

carcass

respectively,
offal,
may

on

the

the

ash

and

average
23

per

39
cent

per

from

12,

of the

cent

of the

which

it

protein,28 per

the

two

appears

of

cent

fat of the entire animal

was

in the offal.

contained

Epidermal
91.

computed

between
ingredients

the percentage distribution of the

in Table
with the results shown
that

be

protein
the

Functions.

The

"

tissues

epidermis,or

outer

layer of the skin,

the
layers of cells of which those nearer
alive and capable of multiplicationwhile towards
true skin are
to flattened,
surface they are
the outer
gradually transformed
as
a
protectivelayer and gradually
horny scales which serve
slough off. Both the epidermisand the protectivecovering of
animals, hair, wool, feathers,etc., as well as the hoofs
and horns, correspondingto the nails in man, are modified forms
of epidermal tissue,their characteristic
ingredientsbeing the
class of albuminoids
designatedas keratins (51 e).
consists of

numerous

"

"

phur
Except for their high and variable sultion
content, the keratins differ little in elementary composibut they are much
resistant
from the simple proteins,
more
chemical reagents, being,for example, insoluble in alkalies in

92.

to

the

Composition.

cold

and

unattacked

of

by

either

pepsin or

outer

followingtable shows
the more
important epidermal tissues
Table

Epidermis of

13.

man

Hair
.

Nails
Horn

of

Hoof

of horse

Pure

dry
dry

Pure

....

cow

wool
.

wool
.

trypsin.

These

covering of the animal.

the elementary compositionof some

well for the

propertiesfit them
The

"

"

Composition

of

"

Epidermal

Tissues

58

OF

NUTRITION

The
Food

93.

storage.

foregoingparagraphsmay
the working machinery

tissues

reserve

classes

The

"

ANIMALS

FARM

be said in

of tissue considered
a

generalway

to

in the

constitute

of the

body. They are composed of

activities
the organism through specific
cells which either serve
sort or anof their protoplasm,as by producing motion of one
other,
other prodor
ucts,
transmittingstimuli or secreting
enzyms
of the extraordinarydevelopment
which, by means
or
of their intercellular

substance,support and protect the various

body as a whole.
cells have
the
(77),many
previouslystated, however
of storingup surplusfood in the form of cell enclosures,
power
especiallyas fat or glycogen, which apparently constitute
serve
no
simply repart of the protoplasm itself but which are
of the

organs
As

This

material.

is

less

or

more

certain tissues show

they

this property to
properly be spoken of as

may

of

true

all

cells,but

marked

degree so that
preeminentlythe reserve
a

tissues.
tissue.

Adipose

94.

form

of

of

material

reserve

in the form

Fat

cytoplasmof

adipose tissue,in

tissue is

reserve

which

in the animal

constitutes

most

portant
im-

which

the

the great

body.

dropletsmay

of minute

and

familiar

most

consists of fat and

stored material
store

The

"

be

depositedin

the

than minute
all body cellsbut the presence of more
in normal cells of muscles,
amounts

glands,etc.,is unusual,
.g ^^ icularly
in certain cells

nerves,

"".-.Nucleu
ro

op

asm.

-^

of the connective

"

Fat

drop.

'

Cell-membrane.

of
,-rFlG'
" ~~.pckeme

tissue that

the

of visible fat
largeaccumulations
tne
m
body take place. At the
these cells present no special
outset

fat cell,

characters,but in a well-nourished
animal
globules of fat begin to
of Histology.)
in them, the cells enaccumulate
large,
and finally
the globulesof fat coalesce into larger ones
is reduced
to
the cell substance
a
mere
envelope, cytoplasm
(Bohm, Davidorf, Huber, Text

and

being pushed to
the cell occupiedby

nucleus

volume
thus

Book

of

loaded

with fat constitute

one

fat.

side and
Masses

almost

the

of connective

adipose tissue.

whole
tissue

COMPOSITION

The

increase of

place in
an

OF

two

ways

in

animal
animal

in

condition
as

1 :

OF

FEEDING

STUFFS

59

cells
in

fx

ameter
di-

emaciated

an

/x in

60

an

ordinary thrifty
to

in

ft

much

as

fat

very

The

corresponding
volumes, therefore,

relative
are

20

and

200

animal.

fat

about

to

AND

adiposetissue,accordingto Waters and Bell,1takes

of new
cells and second by
: first by the formation
size of existing
cells as the storage of fat proceeds.

increase in the

observed
They
ranging from

ANIMALS

27

There

: 1000.

regionsin

two

are

in which
fat tends
particular
to
accumulate, viz.,in the

subcutaneous
tissue and
tissue

Fig.

connective

in the

connective

ternal
surrounding the inthat
organs, especially

of the

and

mesentery

tum,
omen-

although all the

forms

of

looser

connective

tissue,
including,as already noted,
the connective
tissue lying
between

and

within

muscles, may

serve

the

for

the

storage of fat.

Fig.

of

95.

Composition
What
adipose tissue.

is

"

here

called

adipose tissue

is

commonly spoken of as
fat,but it is evident that
only a portion of it is fat
in the strict sense,

the

mainder
re-

nective
consistingof conmade
tissue,
up of
albuminoids, or collagens,
their actogether with
companying

Fig.
Figs.

water.

this

It is

nitrogenous material
1

Proceedings,Soc.

6-8.

"

Successive

of adipose tissue.
The

Prom.

Human

8
tion
stages in the forma-

(Hough

Mechanism.)

Agri. Science, 1909,

pp.

20-24.

and

Sedgwick,

6o

NUTRITION

which

forms
lard

making
It
vary
cells

"

the

OF

"

cracklings

FARM

when

ANIMALS

the fat is melted

out

as

in

tallow.

or

composition of adipose tissue must

accordingto the extent to which the depositionof fat in the
has been
carried.
When
the cells are
enlarged and well
is evident

filled with

that

the

in the fattened

fat,as

animal, the percentage

of fat

protein,water and ash correspondingly

there has been little depositionof fat,or when
When
fat
low.
by starvation,the fat
previouslypresent has been withdrawn
will be low and
the percentage .of protein,water
and
content
ash will be high. The following
figuresreportedby Beythien for
the extremes
of composition of the adipose tissue of commercial
to give a general idea of the comserve
position
beef,pork and mutton
of such depositsin well-fed animals.
Table

that

high and

will be

14.

Range

"

of

of

Composition

of

Adipose

Tissue

of

Commercial

Meat

Maximum

Water

Fat

Nitrogenous

matter

Ash

As

illustration of the

an

adipose tissue in different regions of the

figuresfound by Henneberg, Kern
average
the composition of the fat tissues of five

Table

15.

"

Composition

of

Adipose

composition of
body the following
and Wattenberg for

in the

variations

Tissue

lambs

of

SubcutaFat

neous
%

Water
....

Fat

Fat-free dry matter

may

Different

Kidney
Fat

84.49

4-36
93-89

4-5i

i-75

11.00

100.00

100.00

be

sented.
pre-

Regions

Intestinal
Fat

5.82
92.15
2.03
100.00

figuresreported by
for the composition of the kidney fat of a steer
other

the

At

had

received

and

was

in

the

condition

reduced

very

for about

ration

submaintenance

stand

extreme

Protein

storage of

limited

material

reserve

cent

4-59 Per

cent

cent

the

to

of

in the form

occur

may

in
carbohydrateglycogen,especially

the

which

tions
large accumulamore
contains,a much

addition

sometimes

body

the

of fat which

In

"

bridge
Trow-

eleven months

81.42 per
9.60 per

Fat

Glycogen storage.

"

Water

96.

STUFFS

FEEDING

OF

AND

ANIMALS

OF

COMPOSITION

in the

and

the muscles

liver.
that the liver of the average
man
may
and
the
muscles
of
glycogen
approximately150 grams
estimates

Neumeister
store

and

up

total of about

weight of

the liver of

of

the total amount

be

1200

the liver and

glycogen in

of

body.

pound steer at 16 pounds

pounds, and assuming a content

800

at

making a
Estimating the

amount,

same

for the entire

grams

300

of the muscles
cent

approximately the

tissues

other

one

glycogen

of 0,4 per cent

contained in the

and
of

10

in the

body

that
per

cles,
mus-

would

would
but naturallythis amount
grams,
different times accordingto the conditions of

approximately2200
greatly at

vary

exercise.

feedingand
"

3.

97.

Composition

The

of

Composition of entire body.

of individual
the animal

body

Animal

the

of

of many

Whole

of the great number

already discovered in
methods
quantitative

and of the lack of accurate

for the determination

In view

"

compounds

chemical

as

them,

any

complete and

tailed
de-

compositionof the body as a whole is

termine
be done is to dethat can
manifestly impossible. The most
of compounds
the proportionsof the principal
groups
of the

estimate

enumerated
been

made

matter,

as

in

Chapter

Several

different times.

at

well

as

the fat

while sometimes
1

I.

Proc.

Amer.

content

In

investigationshave
and dry
of them
water

such
all

of the latter,have

determinations

Soc. Animal

Nutrition,

of the total
1910,

p. 13.

been

termined
de-

nitrogen

62

or

NUTRITION

of the ash
a

the gross

Table

or

of both

OF

have

FARM

been

ANIMALS

added.

generalidea may be reached

compositionof the body.

16.

"

Composition

of

Entire

From

of what

Bodies

of

such

might

Animals

gations
investibe called

"

Weight

By difference.
By difference in soft parts.

Includes feathers.

By difference in bones.

Empty

64

OF

NUTRITION

FARM

ANIMALS

foregoingfiguresthat the most abundant

singleconstituent,although one which is subject to marked
80 in
variations,is water, its percentage ranging from over
pigs to 37 in a very fat sheep. Only in this latter
very young
from

It appears

case

and

the percentage of water

the greatest variations are
Relatively,
others

two

that of fat.
as

would

be

Table

18.

the

does

expected,since

"

Composition

of

fat is the

Fat-free

in the

material

reserve

Body

fall below

"

Empty

fat,

of the

Weight

body and
instance

be stored up

may

48 per cent,

or,

on

in

FEEDING

65

STUFFS

reaching in
largequantities,

the other

hand,

be almost

may

one

ing
lack-

fasted animal.

of fat-free

Composition

OF

AND

fed or
insufficiently

in the
98.

ANIMALS

OF

COMPOSITION

body.

Since the

"

adipose tissue

a storage of reserve
body represents substantially
material (93,94) temporarily set aside from
the physiological
activities of the organism, a better idea of the composition
of the working machinery of the body is obtained by computing

of the animal

its

compositionfat-free
this is

When
free

body

as

in Table

18.

compositionof

that the

done, it appears

less variable than that of the

is much

being due to variations

depends chieflyupon the

the

in turn

precedingtable

these few

shows.

So

far

as

of the

age

whole,

content,

animal, as

be concluded

can

however, the fat-free bodies

cases,

as

in the water

the chief difference

which

body

the fat-

of mature

from

cattle

less water
than
to contain three to four per cent
appear
those of mature
sheep or swine. In the case of geese, the percentage
of water
of the relatively
is probably low on account

would

small
99.

the

in the feathers.

amount

Composition of fat- and ash-free

viz.,in
foregoinginvestigations,

hlet's and

three of

Chaniewski's,the

dry

matter.

Lawes

In

"

some

of

Gilbert's,Soxmined
detertotal nitrogenwas
and

proteinhas been calculated by multiplyingby

the factor 6.25. These
experimentspermit a computation of
the percentage of nitrogen contained in the fat-free dry matter
which
in the other experiments has been
regarded as protein.
and

the

For

example, in the

are

as

follows

of Lawes

case

and Gilbert's fat calf the

figures

"

Per cent

Total

dry

matter

34.9

Ash

3.9

..........

Fat
Fat-

Total

2.537

-1-

15.7

1^3
and

ash-free

dry

matter

19-2

15.7

nitrogen

16.16%

2.537

nitrogen in

ash-

and

fat-free

dry

matter.

The
in

which

Table

computation for all

publisheddata permit it

results of such

19.

the

of the
are

experiments

contained

in

66

NUTRITION

Table

19.

OF

Nitrogen

"

Fat-

in

ANIMALS

FARM

Ash-free

and

Dry

Matter

Nitrogen
Fat- and
Ash-free
Dry
Matter

in

Half-fat

and

Gilbert

16.30

ox

Fat

ox

16.01

Fat

lamb

16.05

Store

Lawes

16.16

calf

Fat

Half-fat
Fat

16.08

sheep

16.14

sheep

16.04

sheep

Extra-fat

16.15

sheep

Store pig
Fat

16.15
16.05

pig

16. II

Average

Soxhlet

Swine

13.08

Swine

I3-03

Swine

12-44

Average

12.85

Geese

16.06

Geese

16.07

Geese

16.05

Chaniewski

16.06

Average

With

exceptionof Soxhlet's experiments,the percentage

nitrogenapproximates closelyto that of animal proteins.

of

the

If account
used

in

be taken
these

of the fact that the ether-extraction

does
investigations

fat from dried animal

the
has

not

completely

remove

method
the

that
appears justified
fat contained in the animal body
conclusion

the
tissue,

organicmatter other than

the compositionof protein.
substantially
"

100.

body,
in the

4.

Groups
the

vast

plant,as

The
of

Composition

ingredients.
"

number
well

of

as

for the determination

of

As

in the

of many

case

of the animal

compounds found
of accurate
methods
quantitative
of them, renders it practically

single chemical

the lack

Stuffs

Feeding

ANIMALS

OF

COMPOSITION

to be content
necessary
plant substances into

AND

in most

OF

FEEDING

with

67

STUFFS

separationof the
few major groups
gredient
a
or
sub-groups of inAs
ordinarilycarried out, feeding stuffs analysis
of these categories,
viz.,water, ash, protein,
recognizesseven
tract.
non-protein,ether extract, crude fiber and nitrogen-freeex101.

Water.

"

The

amount

cases

of

water

in

feeding stuff

is

weight which the substance

above
the boilingpoint of water.
at a temperature
There
is also a possibility,
however, of a loss of other volatile
besides water, while,on the other hand, some
substances
matter
and thus increase in weight,especially
tend to absorb oxygen
dried in air at a high temperature.
The
mination
deterexact
when
of water
and dry matter, therefore,
is by no means
an
easy problem, but the results obtained by the ordinarymethods
for almost
all purposes
related to stock
exact
are
sufficiently
feeding. Commonly, the residue is weighed and regarded as
of water
being obtained by difference.
dry matter, the amount
Ash.
In the ordinary feeding stuffs analysis,ash is
102.
equivalentto the residue left after complete incineration of the
substance in air or oxygen,
the process being carried out at as
low a temperature as practicablein order to avoid volatilization
commonly
undergoes

inferred

from

the loss of

"

of part of the alkalies present.

ments
distinguishbetween those elewhich
which
and those
were
originally
present as electrolytes
in
combination
has
were
organic
alreadybeen pointed out (5),as has
also the fact that certain elements, notably sulphur and phosphorus,
of
are
only partiallyrecovered in the ash by the ordinary method
preparation. As the study of the functions of the ash ingredients
progresses,
it may
the
be anticipatedthat we
shall come
to determine
involved
in the way
several elements
most
appropriateto each rather
than simply to determine
the ash as a whole.
That

this method

103.

Nitrogenous

for the

fails entirelyto

constituents.

"

As

yet

no

methods

exist

quantitativeseparationof the nitrogenous constituents

from the other ingredientsof plants. While much
labor has
been expended upon a study of the individual proteins
of a comparatively
few vegetable materials, and while in some
instances
it is possibleto state with approximate accuracy
the amounts
of the several proteins present, nevertheless the only available
methods
for the determination
of the nitrogenous compounds

68

of

NUTRITION

in

feeding stuffs

determination

OF

FARM

ANIMALS

indirect

general are

based

ones

upon

of their characteristic element

nitrogen.
the method
of feedingstuffs anal104. Crude
ysis
protein.
of Henneberg and
inherited from the early investigations
of total
Stohmann, the proteinis estimated from the amount
nitrogenupon two assumptions: first,that all the proteins
In

"

contain

16 per

cent

of

nitrogenand, second, that

all the nitrogen

On the basis
feedingstuffs exists in the proteinform.
of these assumptions,the proteinis,of course, equal to total
nitrogenmultipliedby 6.25. The proteinas thus determined
is designatedas crude proteinto indicate the approximate nature
of

of the determination.

by Scheibler,E. Schulze, Kellner

Subsequent investigations
in many
and others have
shown
the presence
feeding stuffs
of relatively
of non-proteinnitrogenouscompounds,
large amounts
the
at least between
so that it is desirable to distinguish
nitrogenpresent as true proteinand that present in the simpler
the
general term
compounds grouped under
non-protein
all
stuffs
and
of
for
scientific
(60-67)
analyses feeding
purposes
should at least make
this distinction. Logically,
too, the term
crude protein should be dropped altogether,but when, as in
this is impracticable,
should
the case
of the older analyses,
care
be taken to retain the adjective,
protein
reservingthe term
for use in the sense
given it in the next paragraph.
As a means
of effecting
105. True protein.
an
approximate
of
the
from
true
the
other
nitrogenous
separation
protein
compounds present in plants,advantage is taken of the fact
,

"

"

"

of the latter class of substances

that

most

An

aqueous

soluble in water.

feeding stuff,therefore,contains
by far the largershare of its non-protein. Such an extract,
however, contains also any water-soluble
proteins existing
in the substance.
in part by coagulation
These are removed
by
the
the
addition
in
heating,i.e.,
by boiling solution,and
part by
of some
reagent with which they form insoluble compounds.
Various

of

are

extract

substances

have

present officialmethod

and

but
the
for this purpose
tigations,
of analysis,
Stutzer's invesbased upon

been

used

hydrate as
copper
feedingstuff is boiled with
uses

the

the soluble

matter

residue is

precipitant.In practice,
added
water, the precipitant
the

filtered off.

The

nitrogen of the

regardedas being proteinnitrogenand

soluble
in-

from

ANIMALS

OF

COMPOSITION

AND

OF

FEEDING

by 6.25 (or some

by multiplication
of proteinis calculated.
amount
It is obvious
a

of

that this method

conventional

employed in a
probably includes

method

somewhat

is

69

agreed factor)the

other

it

STUFFS

stantiall
determiningprotein is suband that the adjectivetrue

Pickwickian

The

sense.

result

proteinsof the feed but may also

include other insoluble nitrogenous compounds.
106.
The non-proteinin feedingstuffs analysis
Non-protein.
in
includes all the nitrogenous compounds
which
remain
all of the

"

solution when

the material

is treated

for the determination

of

in the

manner

scribed
just de-

protein. The

nitrogen may
in the solution but ordinarilyit is obtained
be determined
by
subtractingthe proteinnitrogen from the total nitrogen. The
conventional
difference,
multipliedby some
factor,equals the
non-protein. Obviously, the non-protein is a heterogeneous
mixture, varying as between different feeding stuffs and even
in the

same

feedingstuff

grown

or

harvested

under

different

conditions.

Evidently the accuracy with which

in a feedingstuff are determined
the proteinand the non-protein
with which
the protein
depends not only upon the accuracy
and non-proteinnitrogen can
be separated and determined
but
also on the correctness
of the factors used for converting nitrogen
into proteinor non-protein respectively.
For proteinthe usual factor has been 6.25 as alreadystated,
based upon the assumption of 16 per cent of nitrogenin average
protein. As was stated in Chapter I (44),however, different
proteinsvary in their nitrogen content, and in particularthe
vegetableproteinsrun higher in nitrogen than the animal proteins,
of
which
is,
equivalentto a smaller conversion
course,
107.

factor.

Nitrogen

But

factors.

while it is

"

easilyshown

incorrect in many
cases, it is not so
There
in the
is a rather wide range
individual vegetable proteins,while

that the present factor is

easy to find a substitute.

of the
nitrogen content
most
tain
feeding stuffs contwo
or
more
proteinsin unknown
proportions. Moreover,
the proteins of the majority of feeding stuffs,
especiallyof the

roughages,have
x

not

yet been

separatedand

studied.

suggested the use of the factor 5.7 for the

majorityof cereal grains and leguminous seeds,5.5 for the oil
Ritthausen

has

1Landw.

Vers.

Stat.,47 (1896), 391.

NUTRITION

70
seeds

and

for

lupines,and

rape
various classes of human

proposed the
from

for

6.0

soybean,white bean, and

For

FARM

OF

and

ANIMALS

barley,maize, buckwheat,
other brassicas.

foods,Atwater and Bryant l have

followingfactors for the computation of protein

protein nitrogen:

"

Animal

foods

Wheat,

rye, barley and their manufactured

and
rice, and
oats, buckwheat

6.25

products

Maize,
products
Dried

seeds of

5.70

their manufactured
6.00

legumes

6.25
5.65
5.80

Vegetables
Fruits

feedingstuffs

For

there

be

to

seems

whose

proteinshave not yet

for changing from
reason

no

been

studied,

the

present

usage.

With

the

more
non-proteinsthe case is even
perplexingin
view of the greater varietyof substances
included under
this
of their nitrogen content.
and
the wide
term
The
range
writer
has used tentatively
4.7, the factor for asparagin(66),
of the most
one
widely distributed substances of this class,but
the choice of this factor is substantially
arbitrary.
'

108.

Crude

fat.

Ether

extract.

the fat content

of

by means
possibleof the
been

used

of feeds

solvent

some

other

which

such

as

methods

based

are

ingredients.A

for this purpose,

The

"

for
its

upon

dissolves

as

terminin
de-

tion
extrac-

little

as

varietyof solvents has

carbon
disulphid,carbon

most
tetrachlorid,
petroleum ether and the like,but the one
"
commonly employed is ethylether,or the so-called sulphuric
"

ether

commonly

All the various

used

more

obtained

an

solvents

besides neutral
less of the

as

other

used, however, remove

stances
sub-

fats and

or
fattyacids,includingmore
tract
complex lipoids.In particularthe ether exfrom coarse
fodders contains a varietyof waxes,
tively
well as the chlorophylof the leaves,and a rela-

resins,
etc.,as
small proportion of
to

anaesthetic.

designatethe

extracted

since ether is the reagent

1

true

fats.

It

is

material

used, as
ordinarily

customary,
"

as
"

crude
ether

Storrs (Conn.) Agr. Expt. Sta.,Rpt., 12, 79.

fat

fore,
there"

or,
extract."

FARM

OF

NUTRITION

72

ANIMALS

consists to a considerable
extract
nitrogen-free
of carbohydratesof one
sort or another.
extent
Indeed, it has
been designatedby the latter name,
but the use of
sometimes
is misleadingand undesirable.
the word in this sense
includes not only hexose but also
extract
The nitrogen-free
these latter substances being,therefore,
pentose carbohydrates,
divided between
by the ordinarymethod of feeding stuffs analysis,
of
fiber and
Some
extract.
the crude
nitrogen-free
be determined
these various carbohydratescan
separatelywith
tunately
a reasonable
degree of accuracy, while others,includingunforless approxior
starch,can be determined
only more
mately.
is far from consisting
That the nitrogen-free
extract
by
exclusivelyof carbohydrates has been strikinglyshown
but

as

Stone.1

rule the

He

determined

the

of the

content

various

classes of

carbohydratesin samples of wheat and maize as accuratelyas

in both cases
and found that the sum
was
considerably
possible
determined
of nitrogen-free
extract
less than the amount
as
Much
by the conventional method.
greater differences in this
respect have been shown to exist in roughages.
The compositionand characteristic
111.
of feeding stuffs.
Classes
of the principal
classes of feedingstuffs are considered
cussion
in Chapter XV, but it seems
desirable to anticipate
that disof indicating
the three major classes
here to the extent
into which
the feeding stuffs are
commonly divided. This
classification is based
primarily on botanical characteristics
with which, however, are
associated correspondingdifferences
in chemical composition.
Concentrates or concentrated feeds. As
the* name
implies,
these are
in a
feeding stuffs which contain much nutriment
small bulk.
They include primarilythe grainsand other seeds
wide
rived
of technical by-products dea
and, secondarily,
range
from them
well as certain by-productsof animal origin.
as
low content
Chemically,they are characterized by their relatively
of crude fiber,
in certa'in byzero
ranging from practically
produc
to perhaps 10 or
12
siderable
per cent in grainshaving a conproportionof hulls, like oats or buckwheat, and in
certain by-products.
Coarse fodders or roughage. Botanically,
these consist of
the vegetativeorgans of the plant,i. e., substantially
of stalks
"

"

"

Jour. Amer.

Chem.

Soc,

19

(1897),183.

OF

COMPOSITION

and

leaves.

They

either

As

larger

Roots

and

the

tubers.

These

roughages.
that

of

fiber

and

soluble.

Their
the

dry

in

concentrates

They

large

proportion

might

difficultly

this

matter,

be

the

containing

briefly

crude

other

characterized

limits.
feeds

fresh

hand,

relatively

ingredients

fiber,

and

ingredients.

contain
the

age
foracterized
char-

are

bulky

soluble

respect

of

wide

quite

stuffs

on

of

they

are

73

forms

of

within

they

in

resembling

water,

other

percentage

feeding

"

of

and

concentrates

of

STUFFS

Chemically,

varies

proportion

FEEDING

straw

high

naturally
with

OF

dried.

or

relatively

however,

contain

hay,

ensiled

their

compared

AND

include

fresh,

by
which,

ANIMALS

large
or

resembles
little

which
as

centage
per-

ensiled

are

dilute

crude

easily
centrates.
con-

PART

THE

PROCESSES

II

OF

NUTRITION

CHAPTER

AND

DIGESTION

Part

first

The

112.

those

differences

body

which

familiar

before

modification
need

be

not

they
of

which

products
that

profound

into

meat

of

of

digestion,

113.

the

in

ameba

comes

in

forms

its

particle,forming

cavity becomes
the

entrance

the

evolution

matter,
of which

thus

that

canal

Organs
The

surface

outer

cavity

residues

permanent

feed

is the

giving
of the

and

one,

the

provision

of

as

the

the

the

the

is

organs

same

higher animals

is

of
a

in its

seen

ameba.

When

the

reverse

the

cess.
pro-

temporary

serving

opening
exit

sion
depres-

digestive

temporary

The

around

scale, this

separate

form

77

of waste.

typical

the

tion
diges-

finally closes

in

is the

appendages.

rejected by

exit

grain

particle of feed,

serves
are

one

and

hay

digestion

like

which

which

the

convinces

process

its

of

with

with

Digestion

of

process

contact

animals

constituting

with

in

conclusion.

animals, special tissues

together

slightly higher
a

of

"

this

lowest

animals

animals

In

in

organism.

this

convert

to

unicellular

Undigested

organ.

work,

considerable

them

of

stances
sub-

example,

animal

reach

to

out

the

for

the

way,

ingredients

many

given farm

first step

The

plan.

form

in

necessary

alimentary

i.

General

simplest

this

the

"

used

of

animal

general

undergo

must

feeds

all but

In

for
or

are

The

milk.

apart

set

are

changes

feed.

the

plain that,

manufacture

they

or

it is

be

the

in

contains

can

the

and

are,

chemist, however,

simple comparison

which

in

particulars

of herbivora

cellulose

and

considered

some

former

latter, and

starch

like

One

The

one.

the

in

found

not

nourish

to

show

to

feed

the

facts

The

"

incidentally

serves

every

RESORPTION

nutrition.

between

it

to

in

step

served

have

III

next

for

for

step
the

in

waste

digestive apparatus,

development,

consisting

78
of

OF

NUTRITION

ANIMALS

FARM

cavityor cavities communicating with the external world

by two openings,one for the receptionof feed and the other
of waste.
for the rejection
In domestic
animals, the digestive
is largeand of very complex structure, but in all cases
tract
it
is built upon
the generalplan justoutlined.
Always, from the
ameba
the inner surface of the digestive
cavityis
up to man,
morphologically
simply a continuation of the external surface
of the body, turned in as one
quently,
might a glove finger. Consethe material contained
in the digestive
cavity,strictly
is
stilloutside
the body.1
speaking,
essential part of the digestive
as
an
Finally,
apparatus, there
must
be such organs as the cilia,
tentacles,
proboscis,
lips,
etc.,
is
and
which
feed
introduced
into
the
grasped
by
digestive
cavity,
and likewise means
by which it may be mechanicallyground
of digestion,
to fit it for the process
as, for example, the teeth
of mammals, the billsand gizzards
of birds,etc.
a

the present purpose,

it is unnecessary
into any
to enter
of the anatomy of the digestive
elaborate consideration
organs,
since we
concerned
chemical
with
the
rather
than
are
chiefly
For

the

physicalprocesses

confined

to

of domestic

folds and
114.

and
digestion,
of
general description

this section

be

the

diverticula.

Digestive fluids and

enzyms.

"

In

the

ever
ameba, what-

changes are effected in the substances which it

feed are
accomplishedby the cells of the introverted
or
by their secretions. As the digestiveapparatus
complicated,however,

more

may

digestiveorgans
In these animals, the digestive
apparatus
tube
various
as
a
briefly
having
enlargements,

very
animals.

be described

may

of

place and certain groups

which act
digestive
juices

takes

surface
becomes

division of cellular labor

of cells

as

takes

produce the
In the higheranimals,
upon the feed.
these cells become
the numerous
secretingglands which are
essential part of the organs
of digestion.The
an
principal
active agents in digestionare certain enzyms
secreted by these
in the higher
glands, the more
important digestiveenzyms
animals being:
The
i.
amylases,ptyalin(in the saliva)and amylopsin (in
the pancreatic
juice),
actingupon starch.
are

set

apart

to

"

see

For
R.

more

Meade

complete
Smith, The

discussion

of the

Physiology of

of the digestive apparatus

Animals, pp. 203-226.

development

the Domestic

8o

NUTRITION

alimentarycanal.

OF

The

FARM

horse

ANIMALS

and

hog, as well as carnivorous

animals
like the dog and cat, have a singlestomach
cavity,
while ruminants, such as cattle,
sheep and goats, have a soin the farm
called compound stomach
consisting,
animals, of
four divisions,called respectively
the rumen,
or
paunch, the

Fig.

9.

Sheep's stomach.

"

(Hagemann,

Anatomie

und

Physiologieder

Haus-Saugetiere.)
I, Rumen.

Reticulum.

2,

reticulum,the
In

true

5, Duodenum.

manifolds, and

the

6, Esophagus.

abomasum,

first three divisions of the ruminant

or

esophagus in which the

fermentation
preliminaryto

compound

the

even

while

they

so-called

have

stomachs.

distinct parts may

portion,the fundus

having

domestic

fourth division is a stomach

fowls

the base of the neck

esophagus at
hog,

In

sense.

In

but
the

similar dilatation

constitutes

singlestomachs
a
singlecavity,are

case

of

the

horse

be

of the true

stomach.

In

of

the crop.
of the horse
in
three

distinguished,
viz.,the left
regionand the pyloricregion,the

the functions

stomach

of the

undergoes a softeningand
while only the
gastric
digestion,

Moreover,
and

or

4, Abomasum.

regarded as dilatations

in the strict
the

omasum,

realitythe

to be

feed

Omasum.

stomach.

true

are

3,

the

or

two
case

reality

quite
cardiac

latter
of the

DIGESTION

Fig.

io.

Stomach

"

and

AND

duodenum

8l

RESORPTION

of horse.

(Hagemann, Anatomie

und

Physiologie der Haus-Saugetiere.)

C, Cardiac portion. M, Fundus.

Sch.,Esophagus.

hog the

cardiac

of the

organ

F, Pyloric region. D, Duodenum.

portionis comparativelysmall and the remainder

is to be regarded as constituting
the stomach

proper.
117.

feeds

Rumination.
may

directly
although
sum,
of

matter

and

liquid

(Hagemann,

Ana-

water

abomaas

fact, they

reach

to

seem

ruminant,

quite

pass

the

to

the

In

"

all four

divisions of the stomach.

The

bulky feeds,

more

however, fail

through
canal

but

This

is

pass

esophageal
the

enter

and

rumen

case

the

to

reticulum.

especiallythe

because

nant
rumi-

the

masticates

its feed

FlG-

imperfectlyat the
time of eating. In the
reticulum and especially

".

"

Stomach

of hog.

very

in

the

capacious
for

some

rumen,

time

in

1-4,

Fundus.

2, Cardiac

portion. 5, Pylorus.

8, Duodenum.

the
contact

partiallymasticated
with

the

saliva

feed
and

mains
re-

such

82

NUTRITION

portionof
softened

and

large

so

material

the drink

that

and

The

liquidor
directlyto the

FARM

ANIMALS

reaches this stomach

prepared for further

it always contains
a
the

when

feed

new

and

action.

is

thoroughly

The

considerable

swallowed

is

is

rumen

of

amount
more

or

less

alreadyin the rumen

by the peristaltic
of the latter,thus tending to prolong its stay.
finelycohiminuted
portionsprobably pass on

completelymixed
action

as

OF

with

omasum,

that

manifolds, and

or

abomasum,

the

but

undergoes the process of rumination.

After the animal
has completed feeding,and if it is left in
quiet,small portionsof the feed are raised again to the mouth
from
the rumen
and reticulum by contraction of these organs,
aided by the action of the abdominal
muscles and of the diaphragm,
second
and
swallowed
time.
a
thoroughly chewed
This
time they pass
considerable
to
a
extent, though not
the esophageal canal and enter
the third stomach,
entirely,
the bulk

the

of the feed

manifolds, and

from

this pass

into

the

fourth

or

true

stomach.
The

ruminants

are

animals

speed and cunning to

it is an advantage to them

on

amounts

of feed and

from

escape

then

in the

wild

state

their enemies.

to

be able to

to

retire to

Hence

hastilylarge

consume

some

depend

safe concealment

digestion.Rumination
also enables these animals
to utilize more
thoroughly coarse
menting
rough fodders,the long stay in the paunch softeningand ferthe material and helpingespecially
solve
to destroyor disthe carbohydratesof the cell walls and thus render the
cell contents
fluids.
accessible to the digestive
118. The
membrane
The mucous
gastricjuice.
liningthe
true stomach
contains numerous
simpleglands(tubularglands)
in appearance
in different portionsof the stomach.
differing
Those of the fundus regioncontain two
kinds of secreting
cells,
The
chief
and
cells.
commonly designatedas
parietal
cells similar to those
chief
glandsof the pyloricend contain
of the fundus
occasional
glands, but only an
parietal
cell. The
their
cells
essential
secrete
as
parietal
product
chief
cells produce the enzym
hydrochloricacid. The
of pepsin called pepsinogen.
pepsin, or rather a precursor
The mixed
secretion of these different glands constitutes the
gastricjuice,which is a thin, clear acid liquid having a
to

remasticate

which

and

prepare

it for

"

"

"

"

"

"

"

"

"

"

"

gravityof
specific
about

of

of

hydrochloricacid

and

feed

the

to

of

acid

proteins,especiallyproteoses and
into polypeptids.1The
extent
chloric
hydro-

some

also

gastricjuice has

the

likewise

and

action

and

1.01

derived

into

peptones, and

of
containinga maximum
action of the pepsin
The combined
the gastricjuice splitsthe proteins

1.005 to
of solids.

2.5 per cent

83

RESORPTION

AND

DIGESTION

dissolve

to

serves

septic
anti-

important

an

more

or

less of

the ash of the feed.

digestiveaction on proteins,the gastric

which
brings about the coagulation
juice contains an enzym
the rennet
of the caseinogen of milk
ferment, or chymosin.
investigators,
chymosin is identical with
According to some
to be in favor of its
pepsin, but the weight of opinion seems
In

addition

its

to

"

independentexistence.
119.

The

of

passage

feed

the

from

stomach.

The

"

lower

muscle
is closed by a sphincter
end of the stomach
posterior
called the pylorus,which prevents the ingestedfeed from passing
or

being

or

forced

in the

When
between

of

course

the acid

by

of

contractions

the

gastricdigestion,however,

reaction

of the stomach

the

stomach.

the difference

and

contents

normally prevailsin

which

the

kaline
the al-

duodenum

level,the pylorus relaxes and allows the

certain

it

into

reaction

reaches

alimentary canal,the duodenum,

division of the

into the next

of the stomach

contraction

to

press

istaltic
per-

portionof its acid

partialneutralization of
the pylorusto close
the duodenal
which results causes
contents
again until the alkaline reaction is restored,when the cyclemay
be repeated.
into

contents

The

the

mechanism

Cannon

in

animals

is also

proof

of this is

The

duodenum.

of

this process

carnivora,but

it may

true
substantially

be

has

presumed

of

especiallystudied

been

that what

is true

by

of these

herbivora,although experimental

lacking.

protein and carbohydratesundergo considerable

digestion in the stomach, it is evident that one
important
the stomach
function which
performs is that of a receptacle
which prevents too rapid passage
of the feed into the duodenum
and at the same
time initiates chemical
changes and prepares
While

both

By prolonged peptic digestion in

it is not

believed that

this

occurs

vitro amino

in natural

acids may

digestion.

also be

produced

but

84

NUTRITION

FARM

OF

ANIMALS

vigorousaction of the intestinal enzyms.

Moreover, the settingfree of cell contents
by the fermentation
of the cell walls of vegetablefeeds,as well as the liberation of
the fat of animal feeds by the solution of the proteinof the
accessible to the
adipose tissue,render these materials more
action of the digestive
juices.
On leavingthe stomach
120. The small intestine.
through
which may
the pylorus,
the feed enters the small intestine,
briefly
the feed for the

more

"

Fig.

12.

be described

"

Intestines of cattle.

Die Rindviehzucht.)
(Leisering,

Its average
tube.
long,comparativelynarrow
lengthis,accordingto Colin,about nine times that of the body
in the horse,sixteen times in the ox and sheep and eleven times
in the hog. It is suspended in the abdominal
flection
cavityby a reof the peritoneum called the mesentery, and as shown
in Fig.12 is much
into
It is commonly subdivided
convoluted.
duodenum, jejunum and ileum.
121.

the

The

as

coecum.

tract
digestive

ccecum,

which

"

pass,

From

the small intestine the contents

of

ileo-ccecal valve, into

the

through the

is a diverticulum

of the

canal,the point
digestive

DIGESTION

from

of entrance

AND

85

RESORPTION

the small intestine and

that of exit into the

togetherand in the upper part of the ccecum.

Anatomically, it might almost be called a second stomach.
Its functions,however, resemble
those of the first stomach
of
colon

being near

ruminants

and

of

those

not

the

true

stomach, the feed

stagnating,so
in
speak,
and

ccecum

to

the

going
under-

extensive fermentation

and

putrefaction.The
size of the ccecum,
in a general way,
varies

inverselyas
of

that

the

ach.
stom-

Thus
horse

in the

it

is

very

large,
having about
16 per

of the

cent

total
the

capacity of
canal.
digestive

In

the

other

hand, it has

only about
cent

the

on

ox,

per

in

the

than

2.5

and

sheep less
per cent

of the total

capacityand
hog about

in the
5.5 per

cent.

122.

The

intestine.

large
The

"

Fig.

alimentarycanal
continued

from

ccecum

the

as

13.

"

is

Coecum
comparee

of horse.

(Colin,Physiologie

des Animaux.)

the

which, as its name

largeintestine,

implies,is generally

than
of greater diameter
the small intestine but
shorter.
It is subdivided
into the colon and the rectum
serves

rather

as

resorbent than

as

digestive
organ.

The

also
and
colon

86

NUTRITION

ANIMALS

FARM

OF

enormously developedin the horse,having about 45 per cent

of the total capacityof the digestive
tract, and serves, like the
the digestion
of the less soluble portions
to continue
ccecum,
is

of the feed.

stomach, the glands which

the gastric
secrete
juiceare located in the mucous
liningof the
of the intestines,
In the case
the glands which supply
organ.
like the salivary
the various digestive
glandsof the mouth,
juices,
123.

The

lie in part
of the most

pancreas.

"

the

In

entirelyoutside

the

alimentarycanal

proper.
This is

important of these is the pancreas.

gland located near the stomach, liver and duodenum,

openinginto

and

by

exit with

common

large

its duct

that of the

(cattle,
swine).
like that of the salivary
secretory action of the pancreas,
gastricglands,is intermittent,the gland being active only

bile duct
The

the latter either

One

(horse,sheep),or somewhat

down

lower

feed is present in the duodenum.

The pancreaticjuiceis a clear, viscid

when

reaction due

to

its content

liquid,having

of sodium

salts.

an

kaline
al-

It contains

and
neighborhoodof eightto ten per cent of solid matter
has a specific
gravityof approximately1.030. It differs from
other digestive
juicesin containinga relatively
largeamount
of all the other digestive
the
of protein. As in the case
fluids,
of which the pancreatic
essential active ingredients
are
enzyms,
viz.,a protease, trypsin,
acting
juicecontains three in particular,
the proteins,
an
amylase,amylopsin,actingupon starch,
upon
of
and a lipase,
steapsin,acting upon fats. Small amounts
In the juice
chymosin and of a lactase have also been found.
if not the other
secreted by the pancreas, the trypsinat least,
as
exists in the form of a pro-enzym,
which
trypsinogen,
enzyms,
is converted
into trypsin (" activated ") after the secretion
in the

reaches
124.

located

the duodenum.
The

liver.

"

This, the largestgland in the body, is

immediately below the diaphragm and lies chieflyon

small in ruminants
the rightside of the body. It is relatively
and largein the hog.
The liver has other important functions in nutrition,
will
as
in Chapter V, but as related to digestion
the
it secretes
appear
bile. This fluid,produced by the hepatic cells,passes from
which
unite to form
small ducts,
them into the bile capillaries,
and constitutingthe bile duct.
In
the latter finally
coalescing

88

NUTRITION

FARM

OF

ANIMALS

neutralization

namely by
prevented from

and

the pepprecipitation,
sin
is
the
of
the
digesting
pancreatic
enzyms
juiceand bile,an action which would otherwise take place,since
these enzyms
to be substantially
protein in their nature.1
appear
two

ways,

the

In

small

intestine,the neutralized

are

subjected to

the

bile and

through

by

the

movements

of

the

bring

of the

movements
peristaltic

also

serve

to

action

pancreatic juice,
the intestinal juice,while they are
moved
along
successive divisions of the small and largeintestines

of the

means

and

the combined

of the stomach

contents

them

mix

to

the

intimate

into

contents

latter.

of
with

contact

These

the

intestines

the

intestinal

walls.
.

The

fats of the

feed, under

the action

of the

steapsinof the
pancreaticjuice, undergo a cleavage into glycerol and fatty
acids and this change is considerablyaccelerated
by the bile,
also aids in emulsifyingthe fats and so exposing vastly
which
surface to the action
of the enzyms.
The
more
fatty acids
thus

free unite

set

to

greater

the

pancreatic juiceand

and

free

fattyacids

carbonate.

The

pointed out

in

are

or

less extent

with

the alkali of

soaps, while both soaps

soluble in bile in the presence
of sodium

bile,forming

presence

Chapter

of soaps in solution also aids,as was

emulsion
I, in producing a permanent

of the fats.

Starch, if any escapes digestionin the stomach, is acted upon

in the same
manner
by the pancreatic amylopsin substantially
as
more
by the ptyalin of the saliva but much
energetically,
yieldingmaltose, while both maltose and any other disaccharid
present in the feed

are

acted

upon

by

the invertases

juice,yieldingmonosaccharids.
proteins which escape digestionin

Any
likewise

the

proteoses

and

peptones

the

of the intestina

stomach, and

resultingfrom

gestion
peptic di-

more
hydrolyzed by trypsin and erepsin much
than by pepsin and yield not only proteoses and
energetically
series of progressivelysimpler polypeptones, but a whole
peptids and finallyare largelyor wholly split up into their
are

constituent
1

The

mixed

in the

amino

acids.

foregoing statements
in the

laboratory.

intestine is somewhat

The

describe
actual

what

takes

importance

in doubt.

the materials
are
place when
precipitationof the pepsin

of the

"
127.

Chemistry

The

2.

Digestion

chemical

89

RESORPTION

AND

DIGESTION

Digestion

of

process.

The

foregoing paragraphs
facts
ing
regardgeneral

"

chieflywith those more

for
the organs of digestionwhich are necessary
and
of their functions and only incidentally
dealt

have

with
to

the

digestionis

in outline

of feed

conversion

first step in the

in the
body substances,and specifically
of vegetableinto animal
the conversion

into

animals

standing
under-

time to revert
It is now
processes involved.
at the beginning of the chapter,namely,
made

the chemical

the statement

that

an

stances
sub-

of farm

case

substances.

this
and from
These, however, are chemical transformations
of the digestive
point of view a knowledge of the structure
chieflyas an aid to the understanding
apparatus is of significance
In taking up this aspect of the subject,it
of these processes.
the three chief groups

to consider

will be convenient

of nutrients

separately.
of carbohydrates
digestion

The

By far the largerproportionof the carbohydratescontained

in the feed of farm

animals

consists of

especially
polysaccharids,

starch,cellulose and the various pentosans and hexo-pentosans.

and lactose,
The disaccharids,
sucrose
probably stand
especially
of
in importance, while comparatively small amounts
next
monosaccharids

Cellulose.

128.

assumed

of this

incorrectness

cellulose of

The

"

Haubner
indigestible.

be

to

consumed.

are

feeding stuffs
was

assumption and

to

was

long

the first to show

the

that

the

prove

even

pulp and sawdust, as well

quent
that of ordinary feeds,was
as
digestedby cattle. The subse(158, 707)
investigationsof Henneberg and Stohmann
showed
that the crude fiber of feeding stuffs was
digestedto a
cellulose of such

substances

considerable

by cattle
proved this

extent

have

as

paper

and
to

sheep,and
be true

not

later

only of ruminants

varying degree of other animals, both

omnivora, includingdomestic fowls. Even man
but

to

digestingthe

tenderer

forms

of

cellulose

periments
digestionex-

to

herbivora
is
a

and

capable of

considerable

extent.

None
have

of the

digestive
enzyms

been shown

to

have

any

of the

higheranimals,however,

action upon

cellulose and

the small

OF

NUTRITION

90

ANIMALS

FARM

found in some
(cytases)
cellulose-dissolving
enzyms
for its solution,so
feeds appear
quite inadequate to account
vestigat
of its digestion
that the manner
was
long a puzzle. The inthe digestive
in
of Wildt l in 1874 upon
process
sheep,however, showed, as Zuntz 2 subsequentlypointed out,
amounts

the solution of cellulose

that
the
the

of

in
chiefly

occurs

the feed

alimentarycanal where
paunch of the ruminant

stagnates,
"

in the

and

those

ccecum

portionsof
that
and

is,in
colon.

alreadyadvanced that the

due to a process of
solution of cellulose in the digestive
tract was
fermentation,and this hypothesisalso served to explainthe
of methane
tract.
and hydrogen in the digestive
Tappresence
have
been the first to show
to
perimentally
expeiner,3however, seems
of cellulose in the digestive
that the disappearance
organisms
tract is effected by a fermentation
brought about by the microinhabitingthe alimentarycanal.
Tappeiner'sconclusions have been fullyconfirmed by more
recent
notably those of Markoff 4 in Zuntz's
investigations,
that the consumption of
while Kellner 5 has shown
laboratory,
increase in
crude fiber (strawpulp) by cattle causes
a marked
In the light of these
the amount
of methane
eliminated.
be regardedas established that the disappearance
results it may
of cellulose during its passage
through the alimentary canal of
in the sense
of a simple hydroherbivora is not due to a digestion
but to a destructive
like that of starch or protein,
lyticcleavage,
The products of this fermentation
fermentation.
are large
tities
quanof hydroand small amounts
of carbon dioxid and methane
gen,
which are excreted,and various organicacids of the aliphatic
fact tended

This

series which

to

confirm

combine

fluids.
digestive

The

with

the view

of the saliva

the alkalies

salts thus formed

the sole contribution which

are

or

other

resorbed

tute
and consti-

to the

nutrition of

cellulose makes

to be acetic and
acids formed appear
body. The principal
butyric,although others are also present. In ruminants, the
is the capaciousfirst stomach,
chief seat of this fermentation
while in the horse, with his relatively
small,simple stomach,
and
it takes placeprincipally
ccecum
or wholly in the enormous

the

colon.
1

Jour. Landw.,

(1874),1.
(1884),52.

Biol.,20
Biochem.
Ztschr.,34 (1911), 211 ;
Landw.
Vers. Stat.,53 (1900), 193,

3Ztschr.
4

22

57

Landw.

(1913), 1.

300.

Jahrb.,8 (1879),101.

DIGESTION

AND

The

RESORPTION

91

widely distributed
in the vegetable kingdom and appear
to be contained
chiefly
or
wholly in the cell walls of plants,probably in combination
with
hexosans.
If the ordinary
to
a
greater or less extent
methods
of feeding stuffs analysisare followed,both the crude
contain them
fiber and nitrogen-free
extract
(109,110).
the first to show
that they were
Stone,1 who was
digestible,
129.

Pentosans.

found

"

pentosans

are

of about 60 for the pentosans

percentage digestibility
in the ordinary feed of the rabbit.
Later,2 in conjunction with
a

Jones,
from

he

48

showed

to

that

per cent
in mixed
rations

while

90

results in

different

samples of roughages
of the pentosans were
digestedby sheep,
the corresponding figureswere
from 46

Weiske3

cent.

to 71 per

in

14

about

the

time obtained

same

similar

with

sheep and rabbits. The digestibility

of pentosans has been fullyconfirmed by later experiments.
while pentosans are
But
at least disappear in
or
digestible,
the digestivetract, the manner
of their digestionis not certainly
known.
the
have been
Up to
present time no enzyms
discovered
either in the digestiveorgans
or
elsewhere,which
have been proved to be capableof hydrolyzingthem.
On the
other hand, however, the pentosans are
attacked
by bacteria
much
like other
carbohydrates and yield similar products,
experiments

the
especially
to

are

acids of the

considerable

series.
aliphatic

extent

subject to

That

the pentosans
the methane
tion
fermenta-

in the

clear from Kellner's investigations

digestivetract seems
gested
pulp (128) in which over one- third of the diupon straw
consisted of pentosans, so that it is difficult
organicmatter
resist the conclusion that these,as well as the cellulose,
went
underof similar
fermentation.
Moreover, in a large number
,

to

experiments,the
general

to

way

methane
be

fermentation

has

the

proportionalto

been

total

found

in

digestiblecrude

fiber and

nitrogen-freeextract, including the pentosans. Of

these results do not preclude the possibility
of a hydrolys
of the pentosans in the digestivetract, converting them

course

into pentose sugars,

but

such

place.

the

process

yet there is

Amer.

3Ztschr.

no

direct evidence

If it does not, then

digestionof the pentosans

those from
1

takes

as

the

products of

substantiallythe

are

that

same

as

cellulose.

Chem.

Jour., 14 (1892), 9.
Physiol.Chem., 20 (1895),489.

Agricultural Science, 7 (1893), 6.

NUTRITION

92
130.

Hemicelluloses.

pentosans

the cell wall

of

whole.

No

hold

animal

enzyms

galactans,levulans,etc., or

ANIMALS

is

true

also for the

so-called

the

"

What

"

to

appears

FARM

OF

are

which

of the
specifically
reserve
carbohydrates

(18)

hemicelluloses
known

which

break

up

as

"

hydrolyze the

their

union, if it

exists,with the pentosans, but nevertheless these substances

tract of herbivora.
disappear in part in the digestive
Pending

knowledge on this point, the assumption seems

that they as well as the pentosans undergo bacterial
warranted
the same
fermentation
and yieldsubstantially
products.
exact

more

131.

Starch.

The

"

first agent

to

act

starch

upon

is the

with the other

ptyalin of the saliva (115). As is the case
but its effects and the
been isolated,
ptyalinhas never
enzyms,
conditions
studied,
governing its action have been extensively
in part owing to the ease
be procured.
with which saliva can
The most
important of these conditions are that ptyalinacts
in the neighborhood of 400 C, that is,at about
most
efficiently
blood
trace

temperature, in
of free acid

or

neutral

solution,while

alkali inhibits its action.

To

more

than

acids

or

kalies
al-

even
though they show an acid
proteins,
alkaline reaction to ordinaryindicators,
or
ptyalinis much less
sensitive and it is also less sensitive to organicthan to inorganic
acids.
In brief,the action of ptyalinis inhibited by a very low
concentration
of either hydrogen or hydroxyl ions.
The action of ptyalinon starch consists of a succession of
in the formation of the various
cleavagesand hydrationsresulting
With
cooked
dextrins (24)and finallyof sugar.
starch,the
action is fairly
slowly,
rapid; upon raw starch ptyalinacts more
the rate varying somewhat
with the kind of starch and being
apparentlydetermined by the degreeof resistance of the cellulose
gous
envelopeof the starch grains. Chemically,the action is analoto that of acids,but is less vigorousand is not carried so far.
The
action of acids yieldsdextrose
a final product; that of
as
ptyalinis usuallystated to stop with the productionof maltose.1
The
is necessarily
action of ptyalinin the mouth
very brief.
In the stomach
the feed comes
with the gastric
into contact
juice containingfree hydrochloricacid. At first,this acid
with the proteinscontained
in the feed, but as soon
combines

combined

with

Carlson and Luckhart

(Amer. Jour. Physiol.,23 (1908-9),
149) state that both
ptyalin and amylopsin produce dextrose from starch.
1

DIGESTION

RESORPTION

AND

93

of free acid

accumulates, or to speak more

ceeds
exactly,as soon as the concentration of the hydrogen ions exsmall limit,the action of the ferment is not only
a certain
inhibited,but the ptyalinis digestedby the pepsin.
This, however, is far from happening immediately upon the
The secretion of the gastric
entry of the feed into the stomach.
Moreover, the contents
juicerequiresa certain length of time.
of the stomach
semi-solid rather than liquidand while the
are
than

more

as

trace

of the stomach

contractions

muscular

extent, this effect is less than

some

of

sections
been

animals

given show

which

to

the feed

mix

stomach

the

have

feeds
be

to

to

Frozen

is often assumed.

variously colored
of

the contents

to

serve

distinctly

ingestionof feed. Furthermore,

the gastric juice is secreted
only in the pyloric portion of
is required, therefore,for the
the stomach
(116). Time
and
acid to penetrate and
mass
quently
conseacidify the whole
stratified some

the

after the

time

of

action

the

continue

ptyalin may

for

siderable
con-

period.
Extensive
investigations,especiallyby Ellenberger and
Hofmeister, upon gastricdigestionin the horse and hog have
demonstrated

that in these animals

the stomach

of the

the starch

upon

action of the saliva in

the

feed

plays

an

important

part in digestion. In the horse (116),the left end of the stomach

is

dilation of the

reallya

esophagus.

the entire surface of the stomach

the

In

is lined with

hog, while nearly

mucous

membrane,

the stomach
peptic glands. When
it is evident that the action of the
is filledwith feed,therefore,
hydrochloric acid will begin along the walls of the fundus of

the left-hand end contains

the stomach

no

tents.
only graduallyspread to the rest of the conIt is true that lactic fermentation
usuallysets in during
this period,aiding to acidifythe stomach
contents
but, as
already stated,ptyalin is less sensitive to organic than to
inorganicacids. It has been found that the solution of starch

may

and

continue

five hours

to

both

are

even

feed remains

in the
more

in contact

time, the
alkaline by
slightly
animals

greater

(115). It

horse

may

less
and

favorable

extent

as

much

as

four

or

ditions
ruminants, the consalivaryaction,since the

hog.
for

for
In

with the saliva in the

contents

the

or

of this stomach

rumen

for

siderable
con-

being maintained

of saliva secreted by these

largeamount
be assumed, therefore,in spite of the

OF

NUTRITION

94

fact that the saliva of ruminants

a

of
digestion

considerable

ANIMALS

FARM

that
but littleptyalin,

contains

is effected.

starch

duodenum, any starch not digested in the stomach,

well as any
as
dextrins,etc., produced by the action of the
ptyalin,are subjectedto the action of the amylopsin of the
is
if not identical with ptyalin,
pancreaticjuice. This enzym,
In

the

As
to act* more
energetically.
very similar to it but appears
of ptyalin,
in the case
the final product of its action is maltose.1
the digestion
from
resulting
of starch is more
convenientlyconsidered along with that of
other disaccharids in a succeedingparagraph.
The
of
starch.
132. Fermentation
organisms producing
in the digestive
believed
tract
fermentation
the methane
were
to act
and not
by Tappeiner to attack cellulose specifically
other carbohydrates. As regards ruminants, however,
upon

further fate of the maltose

The

"

this has

shown

been

to be

an

G. Kiihn's

In

error.

extensive

respiration
experimentswith cattle upon the formation of fats
added
of starch were
from carbohydrates,
considerable amounts
to basal rations.
cretion
Invariablythis resulted in an increased exof methane.

the other

hand,

Moreover, there
or

more

no

was

in the

less decrease

fiber

digested,
showing

that the additional

had

its

starch

in

source

the

increase,but

This

consumed.

of crude

amount

methane

on

must

have

conclusion

is

by the fact that the total.excretion of methane was

of the digestedcrude
quite closelyproportionalto the sum
four parts of
fiber and nitrogen-free
extract.
On the average
confirmed

methane

produced for each

were

parts of starch

hundred

one

3
have
fully
subsequent investigations
these results,
confirmed
although givinga lower average, viz.,
hundred
parts of digestible
3.07 parts of methane
per one
that
shown
have
starch.
Moreover, Kellner's investigations

digested. Kellner's

the

methane

fermentation

confined

is not

to

cellulose and

that,as alreadyindicated,the complex of compounds

grouped under the head of nitrogen-freeextract, includingthe
His experiments also show
sugars, is subjectto this process.
that the proportionof methane
variable,
produced is somewhat
depending upon conditions not yet fullyinvestigated.
starch but

As
1

alreadystated (128),the

See footnote

3Landw.

Vers.

on

p. 92.

chief

Kellner;

Stat.,53 (1900),423.

seat

Landw.

of fermentation
Vers. Stat.,44

in the

(1894),257.

96

NUTRITION

pancreaticjuice. The two

explainedin Chapter I, the
of

the formation

small

fats contain

native

presence

fat.

hand

go

of free

emulsion.

amounts

in the natural

dissolved

ANIMALS

processes

permanent

FARM

OF

in hand.

fattyacids

As

favors

As

there

noted, most

of such

acids

which

Furthermore, there

exist
to

seems

be

cleavageof fat into fattyacids and

good evidence that some
gestion
glyceroltakes place in the stomach of carnivora, while the diof proteinin the stomach
helpsto liberate any enclosed
fat.

fats

acid

the

When

pancreaticjuice,the molecules
and in this way
the
saponified
emulsion.
which
tend

fat
splits

with

the

alkaline

of the free acid in solution

of fat is broken

mass

up

into

are
an

steapsinof the pancreaticjuice,

glyceroland fattyacids,would obviously

into

to aid in the

contact

of the

action

The

in

come

emulsification,
while,on

the other

hand,

the

of surface exposedby the

the amount
latter,
by vastlyincreasing
fats,tends to aid the action of the enzyms.

The

of fat is accomplished
saponification
essentially
by the lipasesteapsinof the pancreaticjuice.
is facilitated by the previous
As justnoted, the saponification
while the presence of the bile is also an important
emulsification,
135.

Saponification.
"

factor.

that the presence

of bile is necessary
to
steapsin,while it has also been shown that the

It is claimed

activate

the

cleavageof the fats is much

the effect beingascribed to
presence
power

accelerated

the presence of bile,

the lecithins which it contains.
The

by

of bile also assists in the process

of dissolvinglarge quantitiesof

their calcium
bile aids in

and

some

of

digestionby
fatty acids and

its
of

It appears
also that the
magnesium soaps.
of the fat,but justhow
in the resorption
way

clear.

is not
Fats

do

tract

not

and

have

be fermented

seem

to

when

administered

been

extent

in the

cattle in the form

tive
diges-

of emulsions

had

produce no effect upon the excretion

When
stances
ingiven in substance, they have in some
the effect of diminishingthe excretion of that gas.

The

digestion
of the proteinsand non-proteins

of methane.

found

to

to any

to

digestionthe proteinsare first

subjectedin the stomach to the action of the pepsin and drochl
hyacid of the gastric
juice.
136.

Peptic digestion.
"

In

DIGESTION

AND

RESORPTION

97

productsof peptic digestionare

longing
usually substances beof derived proteins (58, 59). The
to the group
first
product or products are substances called metaproteins,or,
according to the older terminology,syntonin or acid proteins.
By stillfurther action there is formed a succession of proteoses,
and from these,by subsequent cleavage,peptones.
edly
Undoubtfrom
the products resulting
peptic digestion contain a
The

of chemical

individuals but for the present purpose

it is sufficient to say that the action of pepsin and hydrochloric
acid gives rise to the formation of a series of progressively

largenumber

soluble and more

diffusible substances.
more
simpler,
the action extends in the main only as
natural digestion,
the productionof peptones, although polypeptidsseem
as
be

also formed

to

some

extent.

Amino

acids

are

not

In

far
to

found

products of natural peptic digestion,

althoughthey
be produced by the long continued
action of pepsin-hydrochloric
may
acid in artificialdigestion.
The conjugated proteinsare split
into their two constituents
and the proteinportion is then acted upon like other proteins.
The gastricjuice has no action upon
the nucleic acids of the
among

the

nucleoproteins.
In

the

duodenum, the proteins

and the products of their peptic digestion
are
subjected to the
action of the trypsinof the pancreaticjuice. This is produced
137.

Tryptic digestion.
"

in the pancreas

in the form

of

called
pro-fermentor zymogen,
juicein the duodenum
pancreatic
a

presence of
the glandsof the latter to the

trypsinogen. The
stimulates

testinal
production of the incontain a substance,

has found to
juicewhich Pawlow
which activates the trypsinogen,
or
enterokinase,
in
trypsin,

The
studied
pancreas

action
in
or

unknown

some

of

converts

it into

manner.

like
trypsin,

that of

laboratory experiments
with its secretion

as

pepsin,has

either with

obtained

from

been

largely

extracts

of the

fistulae.

sin,
Tryp-

in a neutral or alkaline solution,

acts upon
especially
teins
proin
the
same
manner
as
a
pepsin,
causing
substantially
hydrolyticcleavageand producing at first proteoses and peptones.
It acts much
than
more
pepsin,however,
energetically
and carries the cleavagemuch
further.
The action of pepsin
substantially
stops with the production of peptones. Trypsin,
the other hand, produces a relatively
of the
on
largeamount
H

98

NUTRITION

simple amino

acids out

up.

leaves

most

considerable

but which

found
them

the

Even

on

FARM

OF

ANIMALS

proteinmolecule is built
prolonged action of trypsin,however,
the

of which

residue in which

acids

free amino

no

hydrolysiswith strong

mineral

acids

are

yields

in abundance.

by trypsinin the
much
more
energetically.
of a proteolyticenzym
in
The
138. Erepsin.
presence
cells of the small intestine
the intestinal juiceand in the epithelial
does not act on
has alreadybeen noted (135) This enzym
unaltered proteins,with the exceptionof casein,but it hyyieldingcrystalline
drolyzesproteoses and peptones energetically,
cleavage products. It is of specialinterest to note that,
according to Cohnheim,1 erepsinis capable of effectingthe
tacked
cleavage of that part of the proteinmolecule which is not atby pepsin and trypsin and that in artificial digestion
into comparatively
experiments almost complete conversion
short
productsmay be obtained in a relatively
simplecrystalline
to
Conjugated proteinsseem
as
manner
same
by pepsinbut

be acted

upon

"

time.
139.

Extent

of

protein cleavage in

natural

digestion.
"

that the
by Ktihne and Chittenden
other products
action of trypsinupon
proteinsyieldedamong
substances as leucin and tyrosin,
such simplecrystalline
paratively
comlittle importance was
attached to the fact from the
as
interesting
physiological
standpoint. While the fact was
it
throwing lightupon the chemical structure of the proteins,
the soluble proteoses and
believed that in natural digestion
was
from
the digestive
tract by
promptly removed
peptones were
and that at most
but a small proportionof the feed
resorption
profound cleavage. This belief was
protein underwent
any
the stronger because it was
believed that only proteinsor their
altered derivatives,
the proteoses and peptones, could
slightly
of the organism for proteins. Whatever
supply the demands
into crystalline
broken down
protein was
products was looked
wasted.
With
the progress of investigation,
however,
as
upon
of hyit has become
clear that the processes
increasingly
important
drolyticcleavagego further and play a much more
part than was
formerlysupposed. While it is admitted that
soluble proteins,may be resorbed,the weight
peptones, or even
When

it

was

first shown

Ztschr. Physiol.Chem.

49

(1906),64;

51

(1907),415.

RESORPTION

AND

DIGESTION

99

fact,proteinsare largelyresorbed in the form of comparativelysimple cleavage products ;

not
necessarilyin every case as simple amino acids but at least
in the form of comparativelysimple peptids.
tion
The nucleic acids derived from the peptic or tryptic digesof

opinionis that, as

of

matter

testinal
nucleoproteinsare splitby the nucleases of the inthese
and
again by the
juice into mononucleotids
secretion into nucleosids (53). No
nucleotidases of the same

of the

attacking the latter class of compounds are

digestiveenzyms
extent
by intestinal bacteria
known, but they are splitto some
Furthermore, it has
into pentoses and purinor pyrimidinbases.
found

been

that extracts

of the intestinal

membrane

mucous

cells)possess the power of bringing about the same

(epithelial
testina
of the incleavages which are accomplishedby the enzyms
juice,and in addition are able to splitthe resulting
It appears,
into pentose and base.
then, that
final digestiveproducts of the nucleic acids are, as in the

nucleosids

the
case

phoric
relativelysimple substances,viz.,phossimpleproteins,
acid,pentoses, and purin and pyrimidinbases.
Attention has alreadybeen
of proteins.
140.
Putrefaction
called,in connection with the digestionof the carbohydrates,
The
the bacterial flora of the digestivetract.
to
drates,
carbohywas
as
shown, are acted upon chieflyby the organisms
rivative
Proteins and their defermentation.
producing the methane
the other hand, have been shown
by Kellner to
on
contribute practically
nothing to this fermentation in the case
of cattle. They are, however, especially
subject to the action
action of such
of the organisms producing putrefaction.The
acid
organismsis preventedin the stomach by the hydrochloric
of the gastric
more
juice. In the small intestine they become
the feed reaches the lower portion,while
as
active,especially
testine
lessens again as the lower portionof the largeintheir activity
of water
is reached,owing to the progressive
resorption

of the

"

from
the

The

the intestinal contents.

putrefactionare

derived
The

from

the

aromatic

and

ammonia

characteristic
certain

products of

compounds
proteins(47).

aromatic

heterocyclic
components of the
productsof putrefaction(indolsand

phenols)

found in part in the feces but are

large part resorbed.
are
They cannot, however, be utilized by the organism but, on the
in

contrary, are

poisonous and

are

therefore combined

with

other

which

substances
unite with
are

OF

NUTRITION

IOO

render

sulphatesto

them

the urine furnishes

innocuous.

The

In

particular,
they
which
ether-sulphates
of these

amount

convenient

ANIMALS

the so-called

form

in the urine.

excreted

FARM

substances

index to the extent

in

of intestinal

putrefaction.
141.

non-proteins.

The

"

As

106), the non-proteinsconstitute

(61,
ordinarilydetermined
of nitrogenoussubstances
a
group

of which
soluble in water, many
closelyrelated to the final products of

are

identical with

or

digestionof

the

the

proteins.Accordingly,they have generallybeen

be ready for resorption
without further action by
juicesand therefore to be entirely
digestible.
It has been

is by

assumed
the

shown, however, that,in ruminants

there.
In

no

the
least,
of waterresorption

so

This has several consequences.

the first place,it appears
that

readilyattacked

are

more

are

the true

proteinsof

therefore,tends
In the second
matter

means

to

stimulates

and

utilized

the feed.

The

the

of these
a

supply

abundant

these soluble

by

the

presence

protect the proteinsfrom

place,an

digestive

at

simpleas the mere

In the capacious first stomach
soluble substances.
animals, the non-proteinsplay an important role as
of nitrogenousfood for the organisms which are
so
matter

to

compounds
organisms than
of non-proteins,

bacterial decomposition.

supplyof soluble nitrogenous

and activityof the organisms
multiplication
abundant

of
extensive fermentation
bringsabout a more
the carbohydratesof the feed, as is evidenced
by an increase
hydrates
in the methane
given off and in the proportionof the carboapparentlydigested.
well made
to be fairly
out that the nitrogen
Third, it seems
which
these organisms assimilate is utilized to build up their
protoplasmand thus,by a sort of symbiosis,becomes a source
of proteinto their host.
It has been claimed that this bacterial
but the evidence on
which
this claim
proteinis indigestible,
and there seems
is based is capableof a different interpretation
for believing
be acted on in the
that it may
to be good reason
like other prostomach
and intestines by the digestive
teins
enzyms
of
of proteinto the body. Some
and serve
as
a
source
the evidence in favor of this view is presentedin a subsequent
discussion of the nutritive value of the non-proteins
(786-789).
and

so

AND

DIGESTION

The

RESORPTION

digestionof

IOI

ash

whose
action has been considered
digestiveenzyms
in the foregoing pages
bring about extensive chemical
of
changes in the organic nutrients of feeding stuffs by means
into the nutritive processes
which they are prepared to enter
in the tissues. At the same
time, the so-called " inorganic
gestion
ingredientsof feed are also prepared for resorption,but the di-

various

The

"

of these
than

that of the

142.

Sulphur

substances

its union

from

less

extensivelystudied

organic nutrients.
and

regards the sulphur of

not
that this element is separated
appear
carbon, nitrogen and hydrogen in the processes
phosphorus.

it does
proteins,

the

been

has

with

As

"

tained
proteindigestion. The sulphur of the proteins is confar
in the amino-acid
as
known, is
cystin,which, so
resorbed without further change. As regardsthe phosphorus of
the nucleo-proteins,
opinions stilldiffer as to whether it is split
of digestionor resorbed,still
off as phosphoric acid in the course
in
To
what
extent
organic combination, as a nucleosid.
other ash ingredients
taken up, like sulphurand phosphorus,
are
in organiccombination, it is difficult to say, but that such resorption

of

"

"

takes
143.

placeis

Electrolytes.
"

feeds which

are

present

to

As
as

be

regardedas probable.
those

regards

ash

ingredients of
be assumed

it may
electrolytes,

that

they are brought into solution to a greater or less extent by the

hydrochloricacid of the gastricjuice,but how much reprecipiin the more
tation may
occur
intestine it is difficult to say.
of the ash ingredientsof

related
the

problems of

considered

up

the

to

ash

The

whole

that it

metabolism

can

be

more

profitably

in that connection.

Solution

of

largerpart

well) are

of the

subject of the digestion

timately
feedingstuffs,however, is so inthe question of the paths of excretion and to

Summary
144.

less alkaline contents

or

of changesin digestion

nutrients.

"

The

substances

of the feed of domestic

animals

which

(and of

make
man

of them,
Some
comparatively insoluble in water.
such as cellulose and the fats,may
be regarded as practically
entirelyinsoluble. Others, like starch and the proteins,are

as

OF

NUTRITION

102

FARM

ANIMALS

small amounts
of soluble proteins
While
sparinglyso.
of soluble carbohydratesoccur,
and somewhat
largerquantities
One
play but a subordinate role in nutrition.
they ordinarily
obvious result of the chemical
changes brought about by the
of the digestivetract is to
and organized ferments
enzyms
very

these

convert

insoluble

substances

into

soluble

ones.

Thus

yields sugar, cellulose the organic acids, fats form

acids. It was
and proteinyieldspeptones and amino
soaps
natural, therefore,that digestionshould be looked upon as
of solution and
compared to the preparationof
a
process
of various
in a pharmaceutical
extracts
laboratoryby means
starch

solvents.

digestive
juicesis important,since
the animal, like the plant,absorbs its real food substances
of the
The
solution.
in aqueous
mere
dissolving
substantially
of the feeds,however, is far from being the only or
ingredients
the chief function of the digestive
juices,as is clearly
even
indicated,for example,by the existence of a coagulatingenzym
the soluble casein,or the preslike chymosin,which precipitates
ence
which attack substances already
of the various invertases,
The

solvent

action of the

soluble.
145.

Colloids

converted

into

crystalloids. The
"

principal

called colloids.
belong to the class of substances
Gelatin is a typicalcolloid as are, indeed, all the proteinsand
abundant
the more
carbohydrates,while the sugars, organic
acids,etc.,are classed as crystalloids.
tween
As related to digestion,
the most
important distinction beis the difference in the osmotic
colloids and crystalloids
of which
of their solutions by virtue
crystalloids
pressures
This diffusidiffuse readilythrough permeable membranes.
of the digested
bility
plays an importantpart in the resorption
in the
material into the blood and lymph current, as will appear
the only factor concerned.
next
section,although it is by no means
nutrients

review

of the chemical

changes which

tion
placein digesmolecular simplification.
cleavage by which

take

they are all in the direction of

They are substantially
processes of
Such
smaller ones.
largemolecules are splitinto two or more
a
change,however, is in the direction from the colloid to the
final products of digestionare
The
condition.
crystalloid
shows

that

OF

NUTRITION

104

bearing the

same

different individuals
animal

The
and

but

name

of the

FARM

ANIMALS

derived
same

from

different

species,the

being toxic to another.

both
body proteins,then, are specific

speciesor
proteinsof one

often

and

structure

differ in both

respects from

as

to

composition

those of the feed.

nourish the organism they must

that the latter may
be
converted
into the specific
plished
proteinsof the body. This is accomIn order

through their cleavagein the digestivetract into their

"
which
the body may
constituent
then
building stones
reassemble
to form
proteinsconstructed accordingto its own
only so, but the proteinsof different
specific
pattern. Not
tissues or even
cells must
be regarded as specific.The body
proteinsare built not after a singlepattern but after numerous
It is only by a very
if not
ones.
extensive,even
complete
of the feed proteinsthat
(139),breaking down of the structure
for the body to build up out of the fragments
it becomes
possible
Its protein molethe various proteinswhich
it requires.
cules
have a different architecture
from
those of the plant."
This fact throws
an
interesting
lightupon the coagulationof
the soluble casein of milk in the stomach.
Although present
in soluble form, it is not
a
body proteinand its coagulation
to retain it in the digestive
tract and give the proteolytic
serves
an
opportunity to break it up 'into its constituent
enzyms
"

"

amino

acids.

What

is

of the proteinsis likewise true of

true
strikingly
other nutrients.
The
not
digestivecleavages serve
merely,
render
and
them
soluble
to
diffusible
or
perhaps not chiefly,
but to reduce the molecular complexes to forms which the body
cells can
assimilate.
The
carbohydrates,e.g., are converted
into monosaccharids, even
the somewhat
largermolecules of
the disaccharids appearing to be too largeor to have an
suitable
unfor the body cells to use.
molecular
In
structure
generalthe complex compounds of the feed are splitup by the
of the digestive
fluids into their constituent
atomic
enzyms
which supply the material out
groupingsor
buildingstones
of which
the body by selection and rearrangement builds up
the proteins,carbohydratesand fats peculiarto itself,
and the
so

"

value

of

feed

"

of the
the nature
and amounts
upon
it yieldsin digestion
rather than upon
which it contains.

depends
cleavageproducts which
the

substances
specific

DIGESTION

"
148.

Resorption

3.

Definition.

AND

As

"

was

RESORPTION

"

stated

The
at

I05

Feces
the

beginning of this
in the alimentary

chapter (113),the digested feed contained

canal is really
outside the body, justas in the case
of the ameba.
In order to enter
the body, the digested material must
pass
through or be taken up by the cells surrounding the digestive
cavity. The process by which the products of the digestionof
the feed are transferred from the digestive
ing
organs to the circulatmedia
(blood and lymph) of the body is called resorption.
149.
The
membrane
Epithelium. Villi.
inner,or mucous,
of the digestive tract
bears on
its surface a layer of epi"

Epithelium
of villus.
"

Central
vessel

chyleof villus.

Artery of
villus.

Chyle-vessel.

Fig.

14.

"

Section of villi. (Bohm,

thelial cells,
more
is

or

closelyunderlaid
lymph vessels. It

Davidorf, Huber, Text

Book

of

Histology.)

less resemblingthose
with
is these

agents in resorption.

network

liningthe mouth, which

and
of blood
capillaries

cells which
epithelial

are

the active

106

OF

NUTRITION

ANIMALS

FARM

higheranimals the extent of resorbingsurface is greatly

of the interior surface of the
increased by certain projections
In the

small intestine known

the villi.

as

Those

shaped protuberancesof the inner

They are covered,like all parts of the
cells just described,which
epithelial

round

are

surface

of

intestinal

the

intestine.

surface,with the

underlaid

are

club-

or

by

cate
deli-

minute
membrane, beneath which are found numerous
a
cular
muslayer of smooth (involuntary)
capillaryblood-vessels,
fibers and
villus ends

vessel called

considerable

shows

In

nerves.

of each

center

section
longitudinal
and

takes place in
resorption
of water

amount

the

lacteal,
belongingto the lymphatic

absent in the stomach

villiare

Although some

of

network

Figure 14

system.
The

and

in the

the

more

or

of three

largeintestine.

stomach, and while

tation
less of the fermen-

products are resorbed in the largeintestine,the

intestine is the special
resorptive
organ.
150.

Mechanism

are
digestion

substances

of

Since

resorption.
"

apparentlydirected

into soluble and

toward

diffusible

villi.

small

the

of
processes
the conversion of feed

forms, it was

quitenatural

In
resorptionshould be regardedas an osmotic process.
cells and other tissues between
this conceptionof it,the epithelial
the cavityof the digestive
organs and the blood and lymph
vessels constituted a membrane
through which osmosis took
the contents
were
place. On the one side of this membrane
of the digestive
tion,
tract, containing the soluble products of digeswhile on the other side were
the blood and lymph, containing
of these products. Under
these conditions,
little or none
osmosis was
assumed
nutrients
to set in and transfer the digested
and lymph.
to the blood
Undoubtedly osmosis plays an important part in resorption,
but its effects are profoundlymodified
by the propertiesof the
resorbingcells of the intestinal epitheliumin ways which as yet
but very partially
are
understood, and resorptioncan
by no
be explainedby a simple analogy with the parchment
means
dialyzingtube of the laboratory.

that

Differences

in the

intercellular substance

cells and of the

permeabilityof the epithelial
for the various

dissolved

substances

in the

digestivetract doubtless play their part in bringing about the phenomena

of selective resorption,
of the
while variations in the affinity
cell colloids for

water

may

be

assumed

to

influence

the

resorp-

tion of that substance

however,
chemical

for

which

well

as

as

of salts.

Notable

of the intestinal
towards

among
and

the present, it is necessary

that resorptionis a function

For

There

are

other

07

facts,

to

epitheliumin

the blood

offer any
physicopresent
these is the predominant permeability

it is difficult at

explanation.

intestinal lumen

RESORPTION

AND

DIGESTION

to

direction,
viz.,from
lymph vessels.
one

with

be content

the

the

ment
state-

of the

livingepithelial
cells,
explains nothing but
although such a statement, of course,
that it is impossible
at present to form an adequate
simply means
of the process.
conception of the intimate mechanism
cretion.
seResorption might be characterized brieflyas a reverse
In secretion the active cells of a gland take materials
from
the blood
into the specific
them
or
lymph, transform
and then eject the latter
substances characteristic of the cells,
into the duct of the gland. The
cells of the digestive
epithelial
tract, on the other hand, take up digested materials from the
of the alimentary canal, modify them
less
contents
more
or
chemically and transmit the products to the blood or lymph.
151.

Paths

of

Most

resorption.
"

of the resorbed

substances

cells to the blood in the capillaries

epithelial
and thus finally
through the portalvein (182) to the liver.
hydrates,
This is true of the cleavage products of the proteins,of carboFats, on the other
organic acids, salts and water.
hand, enter the circulation chieflyor wholly through the lymph
in the form of minute
dropletswhich are secreted by the epitheliu
into the lacteals of the villi,
whence
they pass through
the lymphatics to the thoracic duct (186).
seem

to

152.

pass

from

Chemical

the

changes

in

resorption.
"

It

is

somewhat

of
products of digestion,
especially
the proteinsand fats,undergo rather extensive chemical
changes
cells
the
of
in the epithelial
during
resorption.This
process
in Chapter V but may
question is considered more
particularly
be briefly
referred to here for the sake of completeness.
Proteins.
In
digestion the proteins yield comparatively
simple cleavage products. It has been maintained, especially
and his school, that these cleavage products
by Abderhalden
cells into serum
are
resynthesizedin the epithelial
albumin,
which is regarded as the common
of all the body proteins.
source
This view has been based chieflyon the failure to detect amino
acids or other protein cleavage products in the blood coming

generallybelieved

"

that

the

108

NUTRITION

FARM

OF

ANIMALS

during the heightof proteinresorption.

Folin and Denis and also Van
Slyke and Meyer have recently
delicate tests show
demonstrated, however, that sufficiently
of such products in the portalblood in amounts
the presence
as
large as could be expectedin view of the gradualnature of
and of the largevolume
of blood passand resorption
digestion
ing
The
prevailingopinion
through the intestinal capillaries.
to be at present that the digestive
seems
products of the proteins
littlemodification
before
enteringinto
undergo relatively

from

the intestines

even

the circulation.

has been the subject

resorption
have
maintained
of heated
physiologists
controversy. Some
of emulsified
that it is chiefly
a physical
process ; that globules
cells and excreted
feed fat are taken up bodilyby the epithelial
again by them into the lacteals. This view is based largely
which
of apobservations
show the presence
on
microscopical
parently
of
in
unaltered
fat globules
the intestinal emulsion
cells and in the lymph of the
the protoplasmof the epithelial
of fat. Other no less eminent
lacteals after the ingestion
ologists,
physihave
held
that
fats
resorbed
not
as
are
however,
stoutly
unaltered but only after cleavageinto glycerol
and fattyacids
which are soluble in bile (135),and that the fat
(ortheir salts),
cells are
the product of a
globulesobserved in the epithelial
resynthesis.At present the weight of scientific opinion is
stronglyin favor of this latter view. It is perhaps true that
unaltered fat dropletsmay
be taken up by the epithelial
cells
Fats.

but

"

that

The

mechanism

any

considerable

of fat

amount

of fat is resorbed

in this

fashion is to say the least very questionable.

At any rate, the digestedfat reaches the lacteals almost
in the form
fat the lacteals
found

to

are

contain

of

fat,so

that after

filled with

milky
relatively
largeamounts
a

meal
fluid

tirely
en-

containingmuch
and the lymph is

of neutral

fats.

It

is clear,
then,that the resorbed soaps and fattyacids are speedily
synthesizedto fat again. This syntheticpower is stillfurther
and

demonstrated
strikingly
by the fact that free fattyacids
are
readilydigested but are transmitted to the lacteals in the
form
of the corresponding
neutral fats,having evidentlybeen
combined
in the process
of resorption,
with glycerol
although
the source
from which
the body derives its glycerol
is stilluncertain.
Evidently,then, from this point of view, nothing

DIGESTION

in the way

stands

AND

RESORPTION

09

of the

suppositionthat the digestedfats are

completely splitup into glyceroland fatty acids in the process
of digestion
and synthesizedagain in the epithelial
cells,
although,
of
the
other
it
does
that
not
such
is the
on
hand,
course,
prove
case.

The

153.

forward

feces.

the contents

As

"

through the

small

of the

tract
digestive

move

and

large intestines they become

and more
material
more
impoverished in digestible
progressively
and also,in the lower portionof the largeintestine,
are
deprived

of part of their water, so that there accumulates

in the rectum
less solid residue which is voided at intervals as the
a more
or
feces.
feces

The

(198)and
feces

are

whose

feces

example, feces
feed residues

of

be

to

Even

The

"

obvious

produced

fact that the

in the

carnivora,
wholly digestible,
meat
diet,for

almost

On

man.

either absent

are

an

also in

continue

only.

remainder

excretory product.

an

it is evident

feces,while

excretory product

an

feed consists of substances

normal

amounts

as

excretory product is most

an

but

both

regarded as

feed residue.

The

154.

be

to

are

pure
in which

undigested

entirelyor present in

minimal

fastinganimal continues to produce

separatedfrom the
empty loop of the intestine,
the digestivetract, soon
fillsup with fecal-like
a

material.
The

excretory ingredientsof the feces include

unresorbed

digestivejuices and their decompositionproducts, intestinal

cells and cell fragments,leucocytes
worn-out
epithelial
mucus,
and excretions of the intestinal mucosa.
Especiallynotable
the latter

among

the

salts of calcium

are

phosphates of

include

155.

inconsiderable

not

The

feces

as

diet of man,
and to
feed of herbivorous
amounts
one

reason

in the feces.

extract," and

which
escape

are

other

The

feces also

organisms.
intestinal micro-

ordinary mixed
degree the ordinary

The

"

marked

more

relativelyconsiderable

either

or
indigestible
digestionand therefore

insoluble

ash

like

which

ingredients,
may

be

for

reappear

lignin,cutin,the
non-fatty ingredients of the

these,some,

Among
the

residue.

animals, contains

another

chlorophyl and

magnesium.

proportion of

feed

much

of materials
or

and

calcium

and of iron and in herbivora

waxes,
"

ether

regarded

NUTRITION

HO

OF

FARM

ANIMALS

Of more
importance, however, are
wholly indigestible.
such carbohydratesas cellulose and the various hemicelluloses,
the levulans, galactans,mannans,
pentosans, etc., which
be
be
said to
practicallyonly partiallydigestible.
may
By this is not meant, of course, that one molecule of cellulose,
less digestible
per se than another, but only that
e.g., is any
of fact
part of the cellulose of ordinaryfeeds does as a matter
digestion,largelybecause the length of time during
escape
which it is exposed to the action of the organisms which attack
it is insufficient to allow of its complete solution.
The feces of
as

herbivorous

animals, therefore,contain

amounts

of these

verse
di-

carbohydratesvarying with the character of the feed and

of the fermentation
the activity
tract.
processes in the digestive
in the roughages,
Since these compounds are especially
abundant
the feces from these feeds are bulky and especially
rich in undigested
"

crude fiber."

Other

of vegetablefeeding stuffs,
ingredients,
particularly
of any lack of the apaccount
not on
partially
propriat
escape digestion
but
because
are
digestiveenzyms
they
mechanically
from
the
latter.
action
of
the
If
protected
granules of starch,
contained
within a cell which has not been ruptured
e.g., are
during the mastication of the feed,the cell wall tends to protect
them from the action of the digestive
and they may escape
juices,
to
digestionalthough per se entirelydigestible.The extent
which such a nutrient will actuallybe digested,
therefore,will
depend to a considerable degree upon whether the cellulose of
the cell wall is attacked and destroyedby the organisms of the
is true
of starch in this respect is
alimentary canal. What
or
obviouslytrue of all cell enclosures,and explainswhy more
less matter
be rejected
in the feces.
intrinsically
digestible
may
For

like reason,

seeds which

escape

mastication

are

but

perfectl
im-

insoluble
digested,being protectedby the relatively
seed coats.
cellulose itself may
be so impregnated
Similarly,
with ligninand cutin substances that the
crude fiber
may
be attacked only with difficulty
not
at all by the methane
or
"

"

fermentation.
the possibility
of a misFinally,there is to be considered
and resorption.In heavy rations,
proportionbetween digestion
substances
which
are
especially,
actually digestedmay
haps
perthe
insufficient
contact
with
resorptionthrough
escape

NUTRITION

112

OF

FARM

ANIMALS

subsequentparagraph (160)showed that out of

each ioo
of proteinin the clover hay eaten
by the animal
grams
The digestiwere
apparentlydigestedand resorbed.
bility
53 grams
of the proteinin this case, therefore,
is said to be 53 per
in this sense
is obviously a conception
cent.
Digestibility
distinct from that of rapidityor ease
of digestion.A
entirely
effects and
feed may
have no
injuriousnor disagreeable
yet
have a low percentage digestibilitystraw, for example.
may
alone
determine
Neither
does the percentage digestibility
the effect produced by a feed.
Two
feeds may
gestibl
be equallydiin

described

"

and

yet

one

may

be

more

valuable

than

the other because

its

be used to better advantage by

can
digestedmatter
the indigestible
the body. Nevertheless, it is clear that
portionof the feed can make no- contribution to the nutrition
of the body. The first step, therefore,
althoughby no means
the last one, in comparing the values of different feeds or rations
is to determine
as
accuratelyas possiblewhat proportionof
each ingredientis capable of digestion.In other words, we
shall seek to add to the qualitative
knowledge of the processes
of digestion
and resorptionoutlined in the precedingsections
a quantitative
knowledge of the extent of digestion.
158.

Method

of

digestion experiments.

"

The

percentage

of feedingstuffs can, as a rule,be determined

only
digestibility
called digestionexSuch trials are
periments,
by trial with an animal.
and

they are made

digestibility.
The method
is substantially
that originatedby Henneberg
in their earlyinvestigations
and Stohmann
(707). Since it is
the substances
obviouslyimpossibleto collect and measure
digestedand resorbed from the feed,it is necessary to have
to an
indirect method, viz.,
what portions
to determine
recourse
of the feed are
not
digestedand compute by difference the
amounts
digested. As already stated (155),the undigested
portionsof the feed are excreted in the feces. A digestion
experimentreallyconsists in determiningas exactlyas may be
how much
of the feed is thus rejected,
any portionsof it which
disappearduringits passage through the alimentary canal being
regardedas digested.
brief outline of the way in which
will aid in understandingjustwhat is meant
by

If the
matter

feces consisted

would

be very

only of undigested feed residues,the

simple,but they also

include

greater or

RESORPTION

AND

DIGESTION

113

proportionof excretory products (154). In the case

herbivora, the proportionof the latter is relativelysmall

of

less

and

digestionexperiment as ordinarilyconducted is neglected,

it being assumed, in other words, that the feces are
equivalent
method
is
The
same
to undigestedfeed residues.
pursued in
digestion experimentswith swine, although in these animals
the proportionof excretory products in the feces is larger. The
thus introduced into digestionexperiments is not negligible,
error
as
regardscertain ingredients.It will be convenient,
especially
of digestionexperiments
however, to take up first the methods
and
and to consider later the nature
as
ordinarilyconducted
the presence of the
introduced by neglecting
of the errors
extent
in the

excretory

products.

Time

159.

that

Fig.

15.

"

Steer

required
a

for

digestion experiments.

digestionexperiment

in digestion stall.

periodin which the feed to

daily as in
weighed amounts
purpose

of

preceded by

(Bailey'sCyclopedia of

nary

for the

be

removing

be

is fed
investigated
the

is

sential
es-

prelimi-

Agriculture.)
in the

same

experiment. This is
digestivetract residues

the actual

from

American

It

"

NUTRITION

114
of

previousfeeds

OF

FARM

ANIMALS

also of

uniform a rate of
as
establishing
excretion of feces as practicable.In the case of ruminants, such
a preliminary
periodshould extend over one or two weeks, while
less. In the succeeding
with swine it may
be made
somewhat
digestionexperiment proper, the same
feeding is continued
and the feces are collected quantitatively
for a number
of days
(sevento ten or more),in order to eliminate the error due to the
of the excretion from
the
irregularity
day to day. From
weightsof feeds and feces and their compositionas determined
the digestibility
is computed as illustrated in the
by analysis,
followingparagraphs.
and

160.

fed 3.7
A steer was
Example of a digestion experiment.
kilograms of clover hay per day for three weeks.
During the last
ten days of this time, the average
weight of the dailyfeces was 5.662
kilograms. Samples of each were
analyzed and found to contain
the followingpercentages of dry matter.
"

Clover

hay

84.97 Per
22.36 per

Feces
The

cent
cent

contained 3.144 kilograms of

kilograms of hay, therefore,
while
the
dry
5.662 kilograms of feces excreted contained
The difference,1.877 kilograms,
only 1.267 kilogramsof dry matter.
which did not appear in the feces,is regarded as having been digested
This amount
is 59.7 per cent of the 3.144 kilograms
by the steer.
of the dry
eaten ; we
say, then, that the percentage digestibility
of this hay was
called its
matter
is sometimes
59.7 and this number
digestioncoefficient
In precisely
gredien
the same
of each inthe percentage digestibility
way
be computed from the results of analyses of the hay
may
of
the
and
feces,which in this case gave the followingresults :
3.7

matter

''

' '

"

Feces

Hay

Water
Ash

Protein

Non-protein
Crude

fiber

Nitrogen-freeextract
Ether

extract

All the nitrogen of

assumption which,
does

not

as

the feces is here

will

affect the method

appear

of

assumed

to exist in the

later,is far from

computation.

being

true

of protein,an
(166), but which

form

DIGESTION

These
excreted

AND

RESORPTION

115

figures,together with the weights of hay

per day, yieldthe followingresults :

eaten

and

feces

"

Table

20.

"

Results

of

161.

Digestion

Experiment

The
method
Digestibilityof concentrates.
just outlined
for determining the digestibility
of a roughage or of a total ration
is in conceptionvery simple. The determination
of the digestibility
is somewhat
of concentrates
more
by herbivora
complicated, since
be made
the sole feed of these animals.
they cannot
They must,
of a roughage whose
amount
therefore,be fed along with a known
animal
is likewise determined
in a preceddigestibility
by the same
ing
the digestibility
of the total ration
or
followingperiod. From
and the known
of the roughage that of the concentrate
digestibility
is obtained
of
second
calculation
a
by means
by difference.
used
the
in
the
steer
same
Thus,
experiment of the preceding paragraph
received per day in a subsequent period the same
amount,
3.7
of
clover
and
in
addition
of
meal.
maize
kilograms,
hay
4 kilograms
The average
excretion
of
feces
this
mixed ration was
on
daily
8.715
kilograms. An analysisof the clover hay used in this periodshowed
but slightvariations from
that of the preceding period. The
position
com"

of

the maize

Water
Ash

Protein
....

Non-protein
Crude

fiber

Nitrogen-freeextract
Ether

extract
.

meal

and

of the feces

was

"

Il6

NUTRITION

OF

ANIMALS

FARM

in the total ration,computed

contained
digestiblematter
in the first part
shown
as
exactlyas in the previousexample, was
of the
that the digestibility
of Table 21.
If,now, it be assumed
unaltered by the addition of the maize meal, it is
clover hay was
matter
tein,
(propossibleto compute how much of each kind of digestible
in
the total ration was
crude fiber,nitrogen-free
extract, etc.)
have come
the
derived from the hay ;
remainder, therefore,must
from the maize meal and by comparison with the total amounts
is computed.
present in the latter the percentage digestibility
of the digestibility
of a conthat the determination
It is evident
centrate

The

than that of

is less accurate

in this way

feed which

can

be

of the roughage
givenby itself. The assumption that the digestibility
over,
Morecorrect.
is not changed is unproved and probably not strictly
and
in
likewise
all
the
this
source
errors
arisingfrom
any errors
assignedto
weighing and analysisare, by the method of calculation,
The

the concentrate.
thus

writer has

introduced

may

be

shown
very

that the range of uncertainty

It will evidently be
wide.

to roughage is least and

greatest when the proportionof concentrate
which are present
of the concentrate
will affect most
those ingredients

in the smallest
In

extreme

proportion,such

cases,

absurd

or
negativedigestibility

162.

Laboratory

as

crude fiber and

results

are

sometimes

often ether extract.

obtained, such

digestibility
greater than
of

determination

100

per

as

cent.

digestibility. Actual
"

digestionexperimentsupon animals accordingto the method

requirespecialfacilities
justoutlined,while simplein principle,
It would
and a considerable expenditureof time.
obviously
be very desirable to possess methods
by which the action of the
fluids of the body could be imitated in the laboratory
digestive
and
the digestibility
of feeds thus determined
in a simpler
and more
manner.
expeditious
Numerous
to solve this problem,
attempts have been made
but as yet a satisfactory
has been
worked
method
out
only
for protein,while attempts to devise similar methods
for the
non-nitrogenousingredientsof feeding stuffs have not yet
successful. The
method
for protein is based
proven
upon
m
ade
and
Stockhardt
suggestions
long ago by
by Hofmeister,
but was
first put into practical
form by Stutzer.2
It consists
in treatingthe feed with a solution of pepsin and hydrochloric
acid under specified
conditions and determiningthe undissolved
this and the
nitrogenin the residue. The difference between
1

Amer.

Jour. Sci.,29 (1885), 355.

Jour. Landw.,

28

(1880), 195 and 435.

Table

21.

"

Computation

Digestibility

the

of

117'

RESORPTION

AND

DIGESTION

of

tO

CO

c-i d
W

"

vO
CO

tO

i"-

vO

r~~

OO

co

vO
t-

to

on
o

co

ON

CO

rj-

on

00

vO

t-~

"

g0
Tf

O^

CO
CO

rf

to

vO

00

ON

co

On

to

co

ON

to

"*

"M

to

l"-

On

"*

co

CO
CO

oWni

On

to

co

t"-

00

to

lO

tJ-

CN

to

to

ON

CO

""*"

vd

COO

co

vq

ON

"N

""*" Ov

o
-1-

vO

a
H

lO

..g

to

to

to

Ov

co

s^".
"S

T3
"u

"^
"

co

to
"u

60

bO
.;h

60

co

.NO

+-"

u
60
-t-"

__

"u

-0

el

cu

Concentrate

Il8

NUTRITION

FARM

OF

ANIMALS

total

of
nitrogenof the feed represents, of course, the amount
fore,
nitrogenousmatter which has been dissolved and which, thereis regardedas digestible.
G. Kiihn 3 and others between
Comparisons by Kellner,1Pfeiffer,2
the natural and artificialdigestionof protein have shown
that the
of the presence
former method
in the
gives lower results on account
feces of nitrogenousexcretory products (154, 158),but that when a
for the latter in the

correction is made
page

(166) the results

of

the

indicated

manner

two

methods

In other words, the method

agreement.
shows with a good degree of accuracy
the true

on

show

quent
subse-

tial
substan-

of artificialdigestion
as

compared with the

(163,167)of the protein.

apparent digestibility
Influence

163.

of

excretory products

on

apparent

bility.
digesti-

conducted
digestion
experimentas ordinarily
sults
ignores the presence in the feces of excretory products,the reobtained by its use will necessarily
stances
be too low, since subreckoned
which really
are
are
as
undigestedingredients
such.
affected by this
not
Obviously, the ingredientsmost
Since the

"

will be

error

those

hand, are contained in

proportionand which, on the other

which,

the feed in the smallest

hand,

are

the

on

one

abundant

relativelymost

the

among

excretory

the ordinary
ingredientsare,
scheme
of feedingstuffs analysisis followed,ash, ether
As
and
extract
regards the crude
nitrogenous substances.
is absent, since obviously
fiber,on the other hand, this error
crude fiber is included among
the excretory products,and it
no
the same
seems
probable that substantially
thing is true of
the nitrogen-free
extract.

products in the

164.

feces.

when

These

Digestibility of

ash

ingredients.
"

Certain

ash

gredien
in-

phorus,
iron, calcium, magnesium and phosparticularly
are
largelyor wholly excreted from the body in the
feces (199). Furthermore,the resorption
of the ash ingredients
of the digestive
juicesmay not be complete and these residues
may

be

added

to

the ash

content

of the feces.

The

ordinary

digestionexperiment,therefore,affords little information as to

the ash ingredients
of the feed are actually
the extent
to which
1

Centbl. Agr. Chem., 9 (1880),763.

Jour. Landw., 33 (1885),149; 34 (1886),425.
3Landw.
Vers. Stat.,44 (1894), 188.

OF

NUTRITION

120

pepsin-insoluble
nitrogenof

FARM

ANIMALS

feeds

the

quantitatively
in the feces,where it may
be regardedas representing
indigestible
feed protein,
while the pepsin-soluble
nitrogenof the feces
is contained in the excretory products,
part of which are protein
(mucus, epithelium,etc.)and part non-protein(residuesof
digestivefluids,etc.). An
approximate correction for the
also be comof nitrogenousexcretory products may
amount
puted
by the use of Pfeiffer's factor of 0.4 gram nitrogenper
100
digesteddry matter.
grams
167.

Apparent

appears

digestibility. When

the

"

results of the

dinary
or-

digestionexperiment are corrected,in the manner

just
outlined,for the nitrogenousexcretory products in the feces
we
get an approximation to the true percentage digestibility
of the protein,
while, as regardsthe carbohydrates,the error,
at least for herbivora.
as has been shown, is probablynot
serious,
There
is another
of looking at the matter, however.
way
The intestinal products found in the feces are, in effect,
part
of the cost of digestingthe feed.
wear
They represent the
and tear
of the digestive
The
tween
difference,
then, beorgans.
feed and feces will show the net gain to the animal from
the digestion
of the feed,that is,it will show how much
more
proteins,
carbohydrates,
etc., the body has at its disposalthan
it would have had if the feed had not been given. From
this
as
point of view, we may speak of the digestibility
ordinarily
determined
the apparent digestibility,
and regard it as a
as
of the matter
at least)
measure
(approximately
gained by the
body from the consumption of the feed. For many
purposes,
the apparent digestibility
paring
therefore,
givesa better basis for comthe values of feedingstuffs than does the real digestibility.
l
from this point of view that Atwater
It was
proposed the use
of the term
the equivalentof what is here called
as
availability
apparent digestibility.
"

"

"

168.

Composition

of

digested

crude

fiber.

"

The

crude

fiber

(109)consists of the cellulose of the plant togetherwith

of pentosans and of lignin
and other incrusting
varyingamounts
substances,the ratio of which to the cellulose increases with
the maturityof the plant. Cellulose itselfseems
to be attacked
and

dissolved

rumen

and

with

comparative ease

the ccecum,
1

Rpt. Conn.

and

the

same

by
is

organisms of
probably true of

the

(Storrs)Expt. Sta., 1897,

p.

156.

the
the

pentosans, but ligninappears

and

RESORPTION

AND

DIGESTION

to

1 21

be much

less

readilydigested

incrustingmaterials not at all. As a

tion
computation based on the elementarycomposi-

of the other

some

a
consequence,
of the crude

of the feces

respectively
shows
digestible

fiber of the feed and

is
the percentage of the former which
the digestedportionto have approximatelythe ultimate sition
compoof cellulose.
and heat of combustion

and

on

This

by

the

be

cannot

equivalentto saying that the digested

only of cellulose. The variations between
experiments show

in individual

results

and

means

no

fiber consists

crude
the

is

and

case

doubtless

the crude

of
ingredients

other

it is nevertheless

extent, but

fiber

of the
which

it

to

some

of combustion
of the

energy

will
as
supply for the bodily activities,

appear

measure

clearlylater.

more

169.
a

attacked

is the heat

chief constituent

can

are

that the cellulose is the

evident

digested. Neither
digestedportionin any sense

less of the pentosans

or

more

this

clearly that

Composition
for crude

fiber but

extract.

"

somewhat

shown

involving certain assumptions, it has been

has
extract
digestedportionof the nitrogen-free
approximately the composition and heat of combustion

that the

starch
does

or

cellulose.

this fact

serve

less than

Even
to

fix with

any

By

ployed
principlewith that emmore
complicatedin its

identical in

calculation

difference

details and

digested nitrogen-free

of

in the

case

of crude

also
of

fiber

definiteness the nutritive

the

nitrogenstances
of feeding stuffs includes a great variety of subfree extract
of which, like starch,are
digestedin the
(110),some
of the word while many
others,like the hemisense
narrower
value

of the

digestedportion.

know

We

that

digested.
The data as to the compositionof the digestedportionindicate,
but on account
it is true, that it consists chieflyof carbohydrates,
of the small range of ultimate compositionshown by these
afforded of the specific
substances no indications are
drates
carbohycelluloses,
pentosans, etc., are

170.

present.
Digestible carbohydrates.

crude fiber and

C6Hi0O5, it
of

the

ultimate

an

values

fermented

has

"

rather

Since

both

than

the

have
extract
digestednitrogen-free
compositioncorrespondingto the

become

feedingstuffs

formula

estimating the nutritive

add togetherthe digestible
portions

customary
to

digested
mately
approxi-

in

ANIMALS

FARM

OF

NUTRITION

122

"

the
as
designate the sum
digestible
the
early expericarbohydrates." The practice dates from
ments
in
but
the
of HennebergandStohmann,
lightof our present
knowledge has little justification.
in composition
In the first place, as just stated, the agreement
The
essential point,however,
is but approximate and variable.
is that a digestionexperiment can
show
simply that a certain
of a certain ultimate
of material
amount
composition has failed
in the feces of the animal, and
to reappear
by itself affords no
the changes which
it has undergone nor
information
to
as
as
of the products actually resorbed.
As a matter
to the nature
of fact, a large share
of the
digested portion of these two
especiallyin the case of ruminants, has been fermented
groups,
rather than digested. A considerable
proportion of it has been
of these

two

and

groups

to

"

excreted

only
the

in

the

171.

term

crude

of the

fiber have

heat

of

The

products

been

than
presence

of

feces

that
in

the

of

average

per

the

former

Ether
Ether

and

those

few

determinations

of

extract

of ether

the
on

the

reported by

are

soluble

of such

excretory

comparison

approximations.
have
a higher heat

to

heats

were

extract

Digested ether
Landw.

to

ether

The

as

for pure

that

extract

feces

of

of combustion

of the
account
on
hay fed, doubtless
of the indigestiblewaxes,
etc., while
of the digested portion was
combustion

The

of five trials

the accuracy

found

the

gram.

but

regarded

was

computed heat of
distinctlylower than
Cals.

methane

similar

extract

digested

in

be

must

the

9.5

the

(165) interferes with

of the

and

determinations

No

"

ether

made,

presence

its results

extract

extract.

digested

combustion

Kellner.1

and

ether

Digested

composition

dioxid

carbon

as

Such
being
organic acids has been resorbed.
nomer.
digestiblecarbohydrates is a palpable mis-

of

residue

case,

form

gaseous

"

of

fats, which

of combustion

average
per

gram

"

hay

9.194

....

9.824 Cals.

of feces
extract
.

Vers.

Cals.

....

Stat., 47

8.322 Cals.

(1896), 301.

about
on

the

CHAPTER

CIRCULATION,

RESPIRATION

"
172.

Distribution

which

the

of the

organism

of

that

must

not

the

take

only

distributed

receive

the

in

substances
is the

accomplished

by

by Harvey
173.

The

serves

blood.

it

blood,

into

resorbed

feed

them

the

to

products
be

to

or

The

of their

of

clear

processes

the

which

The

the

they

chief

of

must

cells may

vehicle

resorbed

circulation

of

processes

myriad

the

but

being

of

nutrients

distribution
the

is

covered
blood, dis-

and

be

used

carrier

cells but

and

oxygen

fluid

highly complex
only the

not

the tissues

of the

transmitting
away

the

of the

body

carrying

in other

such

parts

very
proper

reader
X

thickish, somewhat

dark

to

Under

red.

5), and

07

the

the

slightly salt

It is somewhat

it is

suspension
The

corpuscles.

latter

comprehension
is

attempted

is referred
and

XI, and

to

of

of the
the

here.

Hough

for further

For
and
details

123

outlines

phenomena
a

of

The

consist

of

are

functions

as

and

elementary

seems

of the
sion,
discus-

Mechanism,

Human

larger treatises

two

the white

metabolism

complete

more

Sedgwick's
to the

of these

per

number

vast

red

general consideration

21

to

seen
a

and

heavier

about

contains

microscope
in

viscid
taste

corpuscles,or erythrocytes,and
leucocytes.

as

the

peculiar odor,

045-1.

bodies,

of nutrition

Chapters IX,

is

fluid,the plasma, holding

corpuscles, or

for

1.

solids.

kinds, known

Only

of its

In

they

purpose

chapter but

requires.

animals

bright

(sp. gr.

small, solid

to

higher

varying from

essential

(113).

the

by

the

necessary

faint but

of total

body

each

familiar

activity to

of the

water

cent

their

nutrition

excreted.

fluid,having
than

the

proper

indirectly, and

purposes,

ingredients

blood

color

This

equally

for

1621.

"

variety of

body
serve

the

that

of

in

shall

which

means

the

preceding

transferred, directly or

are

of

the

digestive changes by

prepared

are

into

up

through it, so

this distribution

feed

formed

taken

EXCRETION1

The

"

outside

place

resorption described
be

nutrients.

products
be

AND

Circulation

i.

ingredients of the

order

IV

on

physiology.

NUTRITION

124

174.

blood

Red

of the

numerous

but thicker
of

To

In

at

to

by

are

Their

5 J millions

number

more

like

have

diameter

millimeter

coin

is enormous,

cubic

per

far the

round

they are

man

in the center, and

edges than

0.0060-0.0085 millimeter.

estimated

These

"

kinds.

ANIMALS

FARM

corpuscles.

two

the

at

OF

being

of blood.

opacityof the blood are due.

of animal are peculiar
of each species
to it,both
corpuscles
to shape and size,but
as
the color and

them
The

their generalcharacteristics
the

are

of most

in all.

same

Those

animalsaresmaller

than those of

corpuscleis

Each

man.

cell,having

but

nucleus

containing
its characteristic ingredient
the conjugatedprotein
which
haemoglobinto

no
as

the red color of the blood

is due.

Haemoglobin is a
substance
and
crystalline
it has

Reichert

by
Fig.
Above

16.

Blood

"

corpuscles.

haemoglobinof

shown
nine red corpuscles,highly magnified;
below, less highly magnified, the appearance
of the blood soon
after being drawn.
(Hough and
Sedgwick, The Human
Mechanism.)

White

blood

colorless,nucleated
blood but which, by
to

through

pass

lymph

the

corpuscles.
"

which

cells

of connective

spaces

of

walls

each

cies
spe-

confined

not

are

the

as

able

are

the

and

vessels

so-called

the

to

movements,
blood

the

tissue

the

form and
crystalline
properties.
white
The
corpusclesare

of ameboid

means

that

has its specific

of animal

are

175.

recentlybeen shown

"

ing
wander-

cells."

They have important functions, especiallyin

protectingthe body from disease,but need not be further
considered
176.

here.

Blood

platelets.

the blood
in

diameter

"

contains
from

In

addition
minute

more

the white

corpusclesand
the coagulationof the blood.

are

the two
nucleated

kinds

are

much

thought to

more

of

puscles,
cor-

cells,ranging

millimeter,called

0.0003-0.0005

or
platelets,
thrombocytes. They

than

to

blood

abundant

be concerned

in

AND

RESPIRATION

CIRCULATION,

EXCRETION

125

sides
complex fluid contains,beof water, a great variety of substances,
about
90 per cent
the most
prominent of which are the proteins,of which two
serum
globulinsand
are
recognized,viz.,two or more
groups
the so-called serum
albumin, which is probably not a single
also
Plasma
individual.
contains
chemical
approximately
of dextrose,from 0.1 to as much
1.0
as
0.1-0.15
per
per cent
form
in
soluble
of fat,usually
cent
(243),a great variety
some
in part waste
of so-called extractives which
are
products of cell
action,and about 1 per cent of mineral ingredients.
177.

Blood

178.

Coagulation.

plasma.

"

This

"

When

very

blood

is drawn

from

the

body

it

The
usually coagulatesor clots within a few minutes.
lating
coagusubstance
and its coagulation
is a globulincalled fibrinogen
is an enzymatic reaction
brought about
bin,
by a ferment, throm-

believed
from

derived
blood
a

very

be

to

the

platelets by
complicated
The

process.

ulated
coag-

protein

stitutes
con-

the so-called
blood

fibrin,which
self
entangles within itthe corpuscles,
iar
producing the familblood clot.
the clot is very
the dry blood
amounts

bulky
fibrin

only 0.2-

to
cent

per

0.3

While

of the

weight of the blood.

179.

The

Tljeblood
to

the
a

body
most

organ,

Figure

heart."

Fig.

"

of mammalian

Diagram

heart.

uted
is distrib-

all parts of
of
by means

auricle.
ventricle,
b, Right ventricle,
c. Left
a, Left
auricle.
arteries.
Pulmonary
/, Aorta.
d, Right
gg,
veins.
(Smith, Physiology of the Domestic
op.. Pulmonary

Animals.)

interesting
the heart, which
17

17.

shows

is

in

effect

diagrammaticallythe

living force

structure

of the

pump.
mam-

126

NUTRITION

ANIMALS

the
substantially

is

heart, which

malian

FARM

OF

all farm

in

-same

animals.
It is divided

half,and

by
of

each

into
imperviouspartition
these is subdivided
by a cross
an

each

chambers, communicating with

two

the

dividingwall.

are

known

The

left

rightand

the

as

and

upper

other

smaller

rightand left
into
partition
by a valve in

of these

auricles,and

the

divisions
lower

and

largeras the rightand left ventricles. Into these cavities of the

whose mouths
closed
heart open several largeblood vessels,
are
valves

with

auricles and

out

the heart

arteries and
and

of

blood

The

"

is

tissue and

in
originate

and
ventricle,

from

the heart subdivide

the

strong, elastic

the force with

heart.

Their

which

the

walls consist

a middle
tissue,

layer

teries
layerof epithelium. The ar(h,Fig.18),which receives the blood
as
they extend farther and farther

and

minute

with

called

are

throw

off branches

of which

called

are

to

the various

arterioles,
finally

the

capillaries.
are
Capillaries. The capillaries

181.

blood

more

body

inner

the aorta

the left

ending in

the

by

an

from

organs,

withstand

to

them

into

tubes

as

layer of elastic and connective

of muscular

into the

the blood

conduct

of the

organs

described

be

may

pumped

outer

an

only flow

can

vessels which

the various

to

contractile walls,

blood

the blood

of the ventricles.

Arteries.

180.

from

arranged that

so

"

vessels

form

the

walls

are

which

penetrate all the

connectinglink
thin and

exceedinglyminute
tissues of the body and

the arteries and

between

and through them

delicate,

Their

veins.

the nutritive matters

of the blood pass out into the tissues while the waste
ucts
prodof cell activity
In
the tissues into the blood.
pass from
of the posterior
Fig. 1 8, n represents the capillaries
part of the
t those of the kidneys,
body, o those of the stomach and intestines,

p those of the liver,and

the

body.

The

of the anterior part of

capillaries
graduallyunite again into larger

vessels,the veins,which
and lungs.

those

the blood

convey

back

to

the heart
g

182.

Veins.

to the arteries

"

veins

but with

of the blood
of the

The

on

are

weaker

them

cross

and

non-elastic

being slight,owing

between
capillaries

fact that their total

tubular vessels somewhat

them

section

and

similar

walls,the
to

the

tion
interposi-

the arteries and

is greater than

sure
pres-

to

the

that of the

128

OF

NUTRITION

ANIMALS

FARM

pulmonary artery, c, which divides into two

of the rightand left lungs,
leading to the capillaries

ventricle into the

branches
The

d, d.

is providedwith

The
to

by

to this blood

entrance
a

which

valve

vessel,like

prevents the

that

of the

others,

of the blood.

return

returns
blood, after passingthrough the lung capillaries,
the left auricle,
/, through the pulmonary veins,represented

sends
contracting,

auricle then

The

e.

the

blood

into

the^

powerfullyand
ventricle,
g, which, in its turn, contracts
expelsthe blood into one -largevessel,the aorta, h. The aorta,
after leavingthe heart,divides into two branches, i and j,
soon

left

forming the arteries which carry

repeatedlysubdivide,
it returns
whence
the blood to the arterioles and capillaries,
again through the veins to the rightside of the heart.
The passage of the blood from the left side of the heart through
and back to the right side is called the
the body capillaries
systemic circulation ; that from the right side
greater or
the pulmonary circulation
of the heart through the lung capillaries,
and these

The

of the

appearance

the

in

blood

and

veins

arteries is

dark, cherry-red
color,but after it has passed through the lungs and is sent out
the heart

by

former

the

In

different.
strikingly

the arteries it has

to

the

is called venous,

it has

veins

brightscarlet

arterial blood.
latter,

The

color.
An

exception

rule, that the arteries carry brightred blood and

the veins dark, is found in the pulmonary circulation,
where,
this

to

leadingfrom the heart to the lungs carry

venous
blood, and those leadingfrom the lungs to the heart,
is adhered
arterial. Nevertheless, the general nomenclature
of course,

to, and

the vessels

the former
conduct

to

speak of

a
reality

the

constitutes

hose

The

pumped

it is not

it.

uncommon

blood, or of the blood stream, suggesting

brook or river,the circulation is not in

to

into

hose

the force pump

powerful

is driven
were

While

toward

teries
Ar-

the flow of the

drives the blood

water

"

flow of this sort but resembles

water

hose.

of circulation.

analogy

an

the latter veins.

from the heart, veins

the blood

Mechanics

184.

called arteries and

are

into the arteries

left open

the water

contraction

muscular

into the hose

The
heart
force pump.
the arteries correspondto the

by

and

by

of

rather the movement

by
the

would

of

successive
pump.

issue in

the

ventricle

impulses,as
If the
a

end

the

of the

series of spurts

EXCRETION

AND

RESPIRATION

CIRCULATION,

the strokes of the pump.

than the hose
of smaller diameter

correspondingto
nozzle

is converted

outflow

into

steady

129

By the addition

this intermittent
resistance of

The

stream.

of

of the water
gives rise to a pressure
the passage
stretches the walls of the hose, and their elastic force

the nozzle
which

to

between

the flow

maintains

Substantiallythe

same
are

the arteries terminate

may

resistance

caused

channels

tends

arteries

which,

to

the immediate

to

be

compared
and

which

by

these

minute

pressure

in the

steady
In other words,
through the capillaries.
of the blood through the
of the motion

cause

weakened

The

the nozzle of the hose.

produces a

in the

like the pressure

of
is the elasticity
capillaries
become

to

in
capillaries

the flow of the blood

hold it back

of blood

movement

the

elastic while

walls of the arteries

The

the strokes of the pump.

conditions exist in the body.

and

hose, causes

If the

arterial walls.

the

lose their tone

or

become

hardened

latter
as

in

the driving force is lessened and the

old age (arteriosclerosis),
blood to the
return
circulation slows down, since the veins can
by the
heart only as fast as it is forced through the capillaries

therefore,
pressure in the arteries,
tem.
important indicator of the activityof the circulatorysysThe
veins serve
system, the
substantiallyas a return

arterial pressure.
is

an

blood

The

being pushed along them by the residual pressure from the

by the expansion of the
capillaries,
perhaps aided somewhat
flow.
auricle of the heart, while valves prevent any backward
As compared with the arterial pressure, therefore,the blood

blood

in the veins is low.

pressure
185.

The

but

lymph.
are

"

imbedded

more

spaces

lymph

which

or

between

(83) leaving spaces

These

body

The

gether
closelypacked tosue
tisless looselyin connective
them
spaces).
(intercellular

cells are

not

colorless transparent fluid called the

the cells
in which
is the real nutritive medium
contain

branes
of osmosis through their outer memit,by means
and perhaps in other ways, the cells derive the substances
requiredfor their vital activities and into it they discharge

live.

From

the

products of their action.

lymph in its turn stands in relation to the blood, from
While
it is separatedby the thin walls of the capillaries.

waste

The
which

the minute
to

capillaries
penetrate all

all parts of the

body,

it should

the tissues and

be understood

convey

blood

that the cir-

NUTRITION

13"

is

culatoryapparatus
delicate walls

OF

of the

FARM

closed

ANIMALS

Even

the

very

continuous

and

the blood

system.

are
capillaries

with

into direct contact

thin

the

livingcells,except,
of course, those liningthe blood
vessels. The accompanying
lations
rediagram (Fig.19) illustrates schematicallythe anatomical
of the cells,intercellular spaces, capillaries
and lymphatics
minute
A representing
viding
subdia
artery, or arteriole,
reunited
into capillaries
which
the small
to form
are
walls the nutritive substances
vein V.
Through the capillary
contained in the blood pass, partlyby osmosis
and partlyby
does

not

come

Fig. 19.

"

Relation

of cells to blood

Sedgwick, The

into the lymph to

filtration,
products of cell action pass
blood
186.

and

are

carried

Lymphatics.
another

unite like the

set

"

of

lymphatics. (Hough and

Mechanism.)

vessels and

Human

maintain
in the

its

stock,while the

oppositedirection

waste

into the

off.
In

the

minute

there origintercellular spaces

inates
vessels,the lymphatics,which

to form
capillaries
largerones (L in Fig. 19) and
finallyform two main lymphatic trunks, the thoracic duct
and the small lymphatictrunk,which empty into the great veins
the heart. The lacteals of the intestinal villi,
near
throughwhich
the fats are chieflyresorbed,belong to the lymphaticsystem.

RESPIRATION

CIRCULATION,

In the

lymph

lymphaticsthere

from

is

AND

continuous

the tissues towards

the main

EXCRETION

slow

131

of the

movement

trunks,the lymphatics,

like the

veins,being provided at intervals with valves preventing

flow.
This lymph
a backward
flow
is sustained
in part by a
slightlygreater pressure in the
lymphatic spaces but largelyby
of breaththe rhythmic motions
ing,
and is aided
by muscular
activity.Thus, in addition to the
exchange of substances between
the lymph and the blood through
the walls of the capillaries,
there
is

of

general movement
the
lymph itself over
a

the cells which

the

tends

the

surface

facilitate

to

exchangesbetween

of

it and

the

protoplasm.
187.
"

The

of
activity

tissues varies

the various

different times.

at

muscle, for example,

at

tion.
circula-

of

Adjustment

and

rest

is

sometimes

actively contracting.
a

greater

or

times
some-

less

quently,
Conse-

supply

of food material and of oxygen

will
suffice accordingto circumstances,

supply needs to be
regulatedaccordingly.
This
regulation is effected in
two
First,
substantially
ways.
when
the cells of any particular
tissue increase their activitythey
and
consume
more
give
oxygen
Fig. 20.
off more
waste
products than be- (in white).
fore,tending to produce a relative The Human
and

the blood

"

1^11

rr

of
deficiency

the

of the other

bring about
of
by means

an
a

one

and

in the

increase
nerve

an

lymphatic
and

trunks

Sedgwick,

Mechanism.)

cess
ex-

lymph
in

Main

(Hough

and

blood.

the heart

stimulus, so

action

that

These

conditions

(194),probably

the amount

of blood

OF

NUTRITION

132

passing through the heart

supply of it reaches the
be a partialshunting of
of the body to another as
for a largeramount.
This
of the middle

or

muscular

allow

ANIMALS

is increased
active
the

is

and

tissues.

blood

abundant

more

Second, there

supply

of organs

set

one

from
or

of the
When

nerves.

may

region

one

another

accomplishedthrough the

coat

the so-called vaso-motor

is called for in the

FARM

calls

agency

controlled by
arterioles,
a

largersupplyof

blood

muscles, for example, these fibers relax and

the arterioles to

offered

the blood

to

force blood

into the

for this there is

organs,

upon

enlarge,thus reducing the resistance

flow and allowingthe arterial pressure
to
more
capillaries
rapidly. To compensate

contraction

of the arterioles of the internal

of the abdominal
minishe
especially
resultingin a diorgans,
blood supply. The effect of the performance of work
discussed in Chapter XVI
(721),is possibly
digestion,

connected

with

this effect upon

the blood flow.
hearty meal the arterioles of the

On

the other

hand, after a
digestivetract
blood vessels tend to contract
and
relax,while the superficial
the blood supply is partially
diverted from the surface tissues
This power
of the body to regulate
to the internal organs.
the supply of blood to different regionsis of special
importance,
will appear
later (321),
in connection with the regulation
of
as
the body temperature.
"
188.

The

described
from
and

are

"

By

means

precedingsection

the feed and

lacteals

Respiration

supply.

oxygen

in the

2.

taken

distributed

the nutritive

by

the

the

processes
materials derived

the intestinal

various

189.

blood
blood

capillaries

tissues and

supply of feed materials to

cells,
however, is a supply of oxygen, and this another
those of respiration,
to
are
engaged in furnishing
for transmission
to
through another set of capillaries

Equally necessary

with

to

up

of

the

cells.

living

set of

gans,
or-

the blood
the

cells.

from the air to the

transfer of oxygen
is effected in the lungs,which, with the heart and large
The

lungs.

"

The

ity
vessels,fillthe cavityof the thorax,or chest. This cavis enclosed on
the sides by the ribs and their connections,
forming the chest walls,and is separatedfrom the abdominal
cavity,
containingthe digestive
organs, by a strong, arched,mus-

RESPIRATION

CIRCULATION,

AND

EXCRETION

133

cular

toward
the
convex
partition,
chest, the diaphragm. The air
the lungs through the
enters
trachea, or windpipe, from the
mouth
and nostrils. The trachea,
after reaching the chest,divides
into two
branches, or bronchi,
one
leading to the right and one
to the left lung. Each
bronchus
subdivides
repeatedly into a
of
multitude
fine
tubes, the

smallest of which

called bronchioles

are

(little
bronchi), each of
finallyends in an alveolus,

which

the inner surface of which

Fig.

is much

21.

ckens, Form

increased

the

of

form

the
In

by being arranged

Fig.

pits

represents

21,

air cells.

scale

Fig.

Figure

section of

cross

22.

or

"

air

Section of two

22

two

in

"

Alveoli of lung.
und

wirthschaftlichen

Leben

der

(WilLand-

Hausthiere.)

cells.

bronchiolus,aa
shows

two

alveoli and

diagrammaticallyon

alveoli.

alveoli.

(Hough

Mechanism.)

and

Sedgwick, The

Human

bb

large

NUTRITION

134

walls of the trachea

The

rings which

and

bronchi

from
and

surrounded

are

consist of

cavities of the

lungs are
tissue

this connective

pulmonary artery and

which
of capillaries
alveoli in

direct

covers

diaphragm, the
with a liquid.
190.

of

Mechanics

themselves

out

of the chest
and

found

that the minute

their walls elastic.

larger branches
pulmonary vein, connected
by

are

air

the

of
a

In

the
work
net-

are

breathing.

In

"

breathing,the

lungs

of
essentially
passiverole,the movement
of them
being effected by changes in the capacity
about
the
of
the
phragm
diamovements
brought
by

play

air into and

extensible and

so

spread out over the inside of the

Each
with their liningmembrane.
contact
in a double-walled
sack, the pleura,one wall
the other the chest walls and
the lungs and
the two
narrow
being filled
cavity between

is enclosed

of which

cartilaginous

collapsing.The alveoli and

bound
togetherby connective

consistinglargelyof elastic fibers

tissue

lung

FARM

prevent them

bronchioles

ANIMALS

OF

an

ribs.

Since the

tion
toward
the chest its contracdiaphragm is convex
tends to pull the apex of the dome
toward
the abdomen,
of the chest cavityand by pressure
thus increasing
the volume
walls.
When
the digestive
on
organs distendingthe abdominal
the diaphragm relaxes again the volume
of the chest is reduced
and the abdominal
walls return
to their former
position. This
type of breathingis what is called abdominal
breathing.
The ribs pass obliquelyaround
the chest from
the spine to
of
the breast bone
the intercostal
(sternum). By means

muscles, located

between

turning on their attachments

of
increasingthe diameter
and

from

chest

front

cavity.
breathing.
The

to

This

back

and

type of

the

them,
to

the

the
chest

ribs

can

spine and
both

from

be

elevated,

sternum,

thus

side

side

to

increasingthe capacityof the

breathingis called costal,or rib,
so

types of breathing are

By
ordinarilycombined.
their jointaction the size of the closed pleuralcavity containing
forces
the lungs is increased and the atmospheric pressure
air into the extensible
alveoli of the lungs,so that the
more
latter expand along with the chest cavity,the whole constituting
the act of inspiration,
the diaphragm
or
breathing in. When
and the intercostal muscles relax,the elasticity
of the
two

136

NUTRITION

converted

FARM

OF

ANIMALS

wholly into oxyhemoglobin, the amount

of the oxygen
taken up ranging from eight to twelve volume
The color of haemoglobinis a dark red or purplish,
per cent.
while that of oxyhemoglobin is brightscarlet. To this difference
partly or

color is due

of

venous

and

the

marked

difference in appearance

tween
be-

arterial blood.
The

is very
respiration
of breathing
commonly applied to the mechanical
processes
just described or to the exchange of gases in the lungs. In
which
realityall these are preliminaryto the real respiration,
takes place in the tissues. The
vital processes
in the body
cells consist,broadly speaking,as will appear
in detail in the
next
chapter,of a series of oxidations. The requisite
oxygen
is necessarily
drawn from the lymph in which
the cellsexist (185),
while
the carbon dioxid produced by oxidation is discharged
into it. The
lymph, therefore,tends continuallyto become
192.

of

Respiration

richer in carbon

justdescribed

tissues.

dioxid and
the blood

"

term

in oxygen.

poorer

takes up

In

the

manner

in the

lungs and acts

carrier through the body. Through the capillary
blood
as
a
vessels of the body generally,
therefore,there are continually
passing red blood corpusclescharged with looselycombined
the other side of the capillary
wall is a fluid
oxygen, while on
is relatively
(thelymph) in which the partial
pressure of oxygen
low.
of oxygen
and haemoglobin
Accordingly,the combination
is dissociated
into the
the

lymph

time the

same

in the
from

as

to

greater

less extent

requiredto supply the

excess

of carbon

opposite direction into

the

or

oxygen

neighborhood of

and

passes

needs of the cells. At

dioxid in the

the blood

oxygen

and

lymph

is thus

passes

removed

the cell.1 It is this continual

sumption
con-

and
of oxygen
elimination of carbon dioxid by the
cells which
while the
constitutes the real act of respiration,

of
lungs and the elaborate mechanism
for
breathingand of the blood corpusclesare simply means
providingoxygen to the cells and taking away carbon dioxid.
That the movements
of breathingare not an essential part of
is strikingly
shown
respiration
by the fact that it is perfectly
possible
by suitable devices to maintain oxygenationof the blood

complex

In

these

exchanges, as

large part, its effects

cells.

of the

structure

are

no

in other
doubt

similar
modified

ones,

by

while
the

diffusion

doubtless

plays a
living

specialpropertiesof the

of

of oxygen

foregoingthat the amount

the

consumed

of this element

relativelyinactive

they

lymph

but

little. As

take

the

apparent

When

the

by

up

the

upon

cells.

body

amount

they

are

little oxygen
correspondingly

up

ered
in the latter is low-

the tension of oxygen

and

the

by

ever.
what-

It is

taken

in the first instance

lungs depends

137

movements
respiratory

"

blood in the

from

of any

EXCRETION

Respiration regulated by cell activity.

193.

from

in the absence

animal

an

AND

RESPIRATION

CIRCULATION,

is less dissociation

there

consequence

of

oxyhemoglobin in the blood and the corpuscles tend to

less oxygen
fore
and thereto the lungs stillcarryingmore
return
or
less. On the other hand, as
capableof taking up relatively
the
active they consume
the tissues become
more
more
oxygen,
extensivelydissociated
oxyhemoglobin in the corpusclesis more
back to the lungs relatively
and the corpusclestend to come
exhausted of oxygen
and ready to take up the maximum
amount.
Any considerable degree of tissue activity,however, calls for
a more
rapid supply of oxygen than can be providedfor in this
stimulus
and this need is met
to the heart, causing
by a nerve
way
it to beat faster and more
powerfully,thus increasingthe
the

and

arterial pressure

therefore

the

of blood

amount

passing

given time. In these two ways

in the lungs is very accurately
of oxygen
absorbed
the amount
adjusted to the needs of the organism. It is impossibleto
stimulate
the body oxidations by a free supply of air as, for
example, by deep and rapid breathing,as one might blow up a
fire with a bellows,or to get more
intense combustion
placing
by reair with pure oxygen.
In the body such additional air
reaches
the fire. Each
never
or
corpuscleis a receptacle
oxygen
and if
of oxygen
which can
carry only a definite amount
it comes
back to the source
stillpartlyfilledit takes up so much
the less on its next
trip,or if it travels slowly it is less efficient
through

if it

than

tissues
the

the

can

194.
tration

no

be affected

handle.

more

stream

just used

of

from

the

which

rhythm

is true, of course,

carries

at

So, too, the

the

by increasingthe ventilation of
is by
of water
delivered by a pump

the amount

of the

of
respiration

frequently.The

more

Regulation

stream

returns

lungs than

the volume

the

in
capillaries

least
amount

as

much

the

of

water

is taken.

breathing.

only

on

The

condition

the pump
available in the

water

of oxygen

the

"

as

illusr

that
can

lungs

138
must
a

least

at

equal

with

true

the

that the

varying

rate

the tissues.

It is

of ventilation of the

lungs

activityof the

but is
activities,
which, as everyone

of all these
work

ANIMALS

requiredby

the amount

familiar observation

varies

FA*RM

OF

NUTRITION

body

This

cells.

familiar in the

most

case

is
of

knows, promptly increases

the rate and depth of breathing,
that severe
such as
so
exercise,
sources
rerapidrunning,for example,bringsinto play all the reserve
muscular

the

of

breathing mechanism.

already stated
(190),the muscles which are used in breathingare ordinarily
controlled from the so-called
and it is
center
respiratory
that
the regulationis effected.
through this center
If, for
example, an animal be suppliedwith air largelydiluted with
indifferent gas, such as nitrogenor hydrogen, the partial
some
As

"

of the oxygen

pressure

in the alveoli is

of the blood
Such
and

"

is

so

reduced

moglobin
that the hae-

with oxygen.

saturated
only partially

stimulates
of oxygen
center
the respiratory
deficiency
crease
active breathingand a correspondinginproduces more
in the rapidity
with which the air in the alveoli (residual
a

air)is

renewed.

Under

ordinary conditions,however, the stimulus to the

in the blood but an
center
is not a lack of oxygen
respiratory
of carbon
As
dioxid.
has already been implied,the
excess
not
lungs serve
only for the absorptionof oxygen but for the
of the
elimination
carbon
dioxid produced by respiration,
which
of the lymph to the blood and thence to
passes by way
the air in the alveoli
of the
tends
Even

to

tissues

increase

the

of the

by

which

lungs. Any
carbon

more

dioxid

of this substance

content

in the

increase

tivity
ac-

produced

is

in the

blood.

slightincrease,however, promptly stimulates the

and so causes
of the muscles
center
respiratory
greater activity
in deeperand to some
extent more
concerned,resulting
especially
the ventilation of the lungs
rapid breathing. By this means
is augmented and so provisionis made
of a
for the removal
a

very

greater amount
It is

of carbon

plain,however,
alone

dioxid.

that

gases between

simpleincrease

in the

lung

tilation
ven-

is not

sufficient,
except in a limited degree,to carry
carbon dioxid from
the tissues.
Along with the

more
away
increased ventilation there

of the blood

current

the

which

lungsand

must

be

an

rapidity

by which the transfer of

lymph takes place. Accordingly,

is the medium
the

increase in the

RESPIRATION

CIRCULATION,
find that

we

substantiallythe

EXCRETION

139

stimuli which

same

breathingalso stimulate

active

AND

the heart

cause

action and

more

vice

Lack

of oxygen
of carbon dioxid are
the two
excess
or
factors in regulatingthe breathingrhythm but by no

the

only

They

ones.

the

however,

are,

of

ones

versa.

cipal
prinmeans

tance
impor-

most

in the present connection.

Gaseous

195.

the

exchange

exchange
of gases

the

through

between

skin.

the air and

addition

In

"

the

blood

to

which

in the

lungs,a similar process takes place,though to a

smaller extent, through the skin.
The true skin,underlying
the cuticle or scarf-skin,
is penetrated by capillaryblood
the blood
vessels,and in its passage through these capillaries
carbon
dioxid and takes up some
fusion
gives off some
by difoxygen
through the skin. The amounts
given off and taken
small compared with the corresponding amounts
in the
up are
and must
be taken into
lungs,but still are not inconsiderable,
goes
much

on

in accurate

account

experimentalwork.
" 3. Excretion

196.

Excretory products.

activities of the
which

must

with

products and

formed.

For

oxidations

from

the

with

already implied,
the formation

to

the cells and

dischargedfrom

will be concerned
these

cells lead

body

be removed

ultimately be

As

"

the

of

nature

some

the

The
more

of the steps

the present, it suffices

to

say

of

by

vital

products

of which

some

body.

the

must

chapter
important of
next

which

that

they are

the

gradual

of

non-nitrogenousmaterial taking place in the cells

to the production of carbon
dioxid and
give rise substantially
water, while the proteinsand related substances yieldin addition
certain comparativelysimple nitrogenous substances of which
the most
is urea.
abundant
In addition to these substances,
more

or

197.

less of the mineral

Excretion

of carbon

section,the carbon

dioxid

also
ingredients
dioxid.

"

As

produced by

pass into the excreta.

stated in the previous
the

tissue

respiration

by way of the lymph into the blood and is excreted

through the lungsand to a minor degreethrough the skin. In

passes

the blood

the carbon

dioxid is carried

by both the corpuscles

and the plasma, but chiefly(two-thirdsor more) by the latter,
in combination
with proteinsand haemoglobins,but especially

OF

NUTRITION

140

the

with

alkalies.

in the

As

ANIMALS

of oxygen,

case

the amount

in the blood

dioxid contained

carbon

FARM

depends
surroundingmedium.

upon

the

of

partial

of this gas in the

Since the tension
of the carbon dioxid in the alveolar air is less than that in

pressure

the blood

alveolar

of the

the
capillaries,

the former..

carbon

If the air

dioxid

passes

stationarythe
continue
until an
reached.
would
equilibriumwas
process
Since the air is being continuallyrenewed
by breathing,the
tension of carbon dioxid in it is kept permanently lower than
from

the latter

to

that in the blood

and

dioxid from

carbon

there

were

is,therefore,a continual

the blood

to

the alveolar

of

passage

air.

of this

anism
tendency to equilibriumthat the mechcreased
Infor the regulationof breathing is set in motion.
tissue respiration
carbon dioxid into the
more
discharges
a
more
blood, where its tension increases. This causes
rapid
It is

by

means

diffusion of the gas into the alveolar air and

carbon dioxid tension also,so that with an

lung ventilation

carbon

would

dioxid

level

be raised.

Even

to

raise its

unchanged

of both

rate

of

the alveolar

slightrise in
dioxid tension in the blood, however, as already
the carbon
and stimulates
center
stated,acts promptly upon the respiratory
in an
increased lung
the muscles of breathing,resulting
ventilation and consequently a more
rapid excretion. At the
time the rapidityof circulation is increased and in these
same

air and

the blood

the

tends

very

the level of carbon dioxid tension in the blood and

ways
in the alveolar air is maintained
On the other
very constant.
two

hand, if the lung ventilation

may

falls below

be

so

the normal

increased,as
artificially

by

of oxygen,
the carbon dioxid
use
by
facilitated that the amount
in the blood

artificial respiration
or
excretion

be

the

and

the movements

of

breathingmay

be

temporarilysuspended (apncea).
198.

Excretion

of

nitrogenous products.

other

"

The

urea

and

nitrogenousproducts of cell action, like the non-nitrogenous

products,pass ultimatelyinto the blood. In its course
through the body the blood passes through a capillary
system
in two bean-shaped organs, the kidneys,indicated by / in Fig.
1 8, situated in the abdominal
cavityon either side of the spine
the loins. In these organs
the urine is being continually
near
secreted,passing thence through the ureters into the bladder
from

whence

it is voided

at

intervals.

RESPIRATION

CIRCULATION,

The

chief stimulus

is the water

EXCRETION

141

the secretion of water

to

by the kidneys
blood, the kidneys acting as regulators

of the

content

AND

of this

important factor and eliminating

more
or less water
the blood contains a largeror smaller percentage of it.
as
As regards the excretion of dissolved matter, very interesting
relations exist. With one
important exception (hippuricacid)
the kidneys do not manufacture
the excretory products. Their
the composition of the blood
essential function is to maintain
For each substance
constant.
capableof being excreted at all
in the urine

there

the blood above

the urine.

exists

which

it begins to pass
the normal
excretory

For

foreignsubstances,this
minute

very

certain minimum

minimum

the

blood.

per

cent, for sodium

dextrose

For

products,as

substances
limit is

the

chlorid

o-6

in

through the kidneysinto

approaches

of these

amounts

concentration

that

zero,

be

can

retained

So

etc.

for

as

is,only

approximately
cent,

per

well

in

0-2-0-3

long

as

the percentage of one

of these substances in the blood does not
exceed its own
of it is excreted through the
particularlimit,none

kidneys. On

the

an

of the

kidneys
on

hand,

slight rise above

excretion of the substance

causes

valve

other

has

tank.

has its

been

likened

It should

to

concerned.
the

be added

minimum,. independent to

own

This

working of

that each
a

this limit

an

function
overflow

substance
particular
largedegree of

all the others.

The
not

so

functions

of the

simpleas

those of

the concentration
urine
work

of

than

in the blood.

In

is not

and

the other

can

be detected
addition

pressure

and

inconsiderable.

nitrogenouswaste

substances

other

substances
by transferring

this work

In

overflow

an

of the excreted

higher osmotic

in this respect are

valve for the reason
that

kidneys,however,

words,

from

is greater in the
the kidney does its

fluid of lower

to

fluid

the

expenditure of energy in
This is notably true of urea
products,of which only traces

in the blood.

nitrogenous substances excreted in the

urine there are
present in the feces,as already noted (154),
excretory products which represent a certain fraction of the
and
of urea
organic body waste.
Finally, small amounts
other nitrogenous substances
excreted
in the perspiration.
are
199.
of
ash
Excretion
Being non-volatile
ingredients.
the ash ingredientsare
excreted
chieflythrough the feces or
to

the

"

NUTRITION

142
urine

the intestines

accordingas
of

path

excretion,although they

also in the
The

ANIMALS

kidneys form

or

contained

are

the normal

to

small

tent
ex-

perspiration.

intestines

mineral

FARM

OF

usual

the

are

of excretion

path

substances,notably iron,calcium

magnesium. To these must be added

bivora phosphoric acid, which, under

in

and
the

for certain

to

extent

some

of the

case

her-

ordinary conditions,is
excreted in the feces. The urine of herbivora,especially
when
in more
or
they consume
roughage freely,
generalterms when
the basic predominate over
the acid ingredients
of the ash, is
of phosphoricacid.
alkaline and contains but minute
amounts
On the other hand, during fastingor upon
ration having an
a
acid

ash,the urine

have

may

acid urine of carnivora

acid.

The

urine

sulphur,chlorin
200.

Excretion

omnivora,

or

normal

is the

and

acid reaction and

an

contain

may

vehicle

then, like the

for the

phosphoric
excretion

of

the alkalies.

of water.

The

"

motions

of air in and

of

out

the

of removing from the body more

or
lungs are the means
less incidentally
of water
by simple evaporalarge amounts
tion.
in
The presence
the
miliar
of water
air
is a faexpired
vapor
cold surface or in
on
a
fact,shown by its condensation
of its sweat-glands,
cold air. The skin likewise acts, by means
as

channel

for the removal

being
form

of the

"

of water

from

the system, considerable

continuallyevaporatingfrom the skin in the

stances
perspiration."Under certain circumis so rapid as to give rise to the
the excretion of water
formation of visible drops (sweating).
The
than
ox

in

amounts

insensible

of water

sometimes

are

in these two

excreted

realized.

For

ordinary temperature and

through the lungs and skin
twenty-fourhours, the amount
at

of

upon

and

water

in the

case

larger

are

example, a thousand pound

crete
on
lightfeed may easilyexeightto ten pounds of water
depending to a considerable

the temperature and amount

surroundingair. The feces also contain
extent

ways

of herbivorous

of the

movement

of

large percentage

animals

thus

the amount

eliminated

is very considerable.
is excreted in the
water
Finally,

for the

urine,servingas

which
nitrogenousproductsof cell activity

through this
in

channel.

part upon

The

amount

the amount

of

water

consumed,

thus

are

solvent
removed

excreted

in part upon

pends
dethe

CHAPTER

METABOLISM

"
201.

Assimilation

defined

(73)
of

the

the

such

Every
from

these

to

which

cell

direct

of

are

the

blood

consisting

feed

and

is
the

for
from

202.

Definition

that

the

removal

of

the

are

so

the

blood

of

and

is the

simple

supplied

cesses
protained
con-

series

"

of

It

molecular
blood

the

of

is clear

extensive

culation
cir-

cell oxygen

digestive tract

cell

them

puscles
cor-

of

mechanism

from

remove

the

to

tissue and

each

products

144

by

mals
ani-

material

resorptive

material

seat

the

materials

solution

lung capillaries. The

metabolism.

higher
In

lymph.

into

of

qualitativelyuniform
is

which

the

nutritive

that

products

waste

in

the

transmit

oxygen

of excretion

body

the

comparatively

waste

life of

organs

the

that

current

the

of

and

water

replace

to

fulfilled

are

nutritive

and

away

organs

seldom

continued

without

continually distributingto

carrying

plasm
proto-

mation
transfor-

is essential

dioxid,

the

heterogeneous

substances

lungs

their

they

appear,

gradually brought

lymph

through

various

the

(146, 147), while

structure

from

are

cleavages,
of

the

of oxygen

carbon

to

necessary

of

digestion, the

of molecular

of

and

rived
de-

energy,

simple compounds.

circulation

in the

and

will

as

conditions

Both

the

solutes

presence

oxidized

and, second,

by

multifarious

of

constituents

and

first,a supply of material

its activities.

living protoplasm

the

of

seat

been

oxidations, nevertheless, they yield products

ultimately

consumed

to

of

The

while,

and

things, then,

already

It is the

expenditure

an

down

forms.

comparatively

Two

cell has

life.

is the

of cell enclosures

other

are

other

cells which

breaking

changes

primarily

The

"

of

activity requires

or

into

excretion.

organism.

the

itself

Conception

biologicalunit

body

of

activities

and

the

as

General

i.

action
from

to

the

from

chemical

the

the

body.
going
foretrans-

METABOLISM

formations.

hand, molecules of resorbed digestion

products are being assimilated by the body cells and built up
of their protoplasm, while, on
the other
into the structure
hand, molecules of protoplasm or of cell enclosures are being
and oxidized,yieldingfinally
broken down
the relatively
simple
one

products.

excretory
The

On

the

HS

is

metabolism

term

of the chemical

commonly

changes

used

which

the

to

designatethe

constituents

of

tality
to-

the

feed

of their conversion
into
undergo in the course
the corresponding excretory products. Similarly,one
may
restricted
of
in
the
of
metabolism
sense
a
more
speak
single
ingredientsof the feed, as of the proteins,carbohydrates or
the chemical
fats,protein metabolism, for example, signifying
changes undergone by the digestionproducts of the proteins
of the feed during their assimilation and subsequent transformation
into excretory products. The
adjective metabolic is

resorbed

also used
203.

describe these chemical

to

Anabolism

and

katabolism.

changes.
The

metabolism,
as
just defined,includes processes of two distinct kinds, viz.,
molecules of sugars, organic acids,amino
those by which
acids,
into
built
in
the
more
complex compounds
body
etc., are
up
and those by which these complex compounds are broken down
again into simpler substances and finallyinto the excretory
"

term

products.
The buildingup metabolism
has received the name
anabolism,
while the breakingdown
oxidative phase is called katabolism.
or
Any change in the direction of greater molecular complexity
is spoken of as an anabolic change, while one
in the direction of
greater molecular simplicityis a katabolic change.
It must

always

be

inferred

from

precedes katabolism.

constituents
On

not

by

any

what

Neither

has been
is the

of
process
instances are known

means

the contrary, many

at various
stages by anabolic

said

that

anabolism

of cell
breaking
uninterrupted katabolism.
in which
it is interrupted
down

While
changes of one sort or another.
the general direction of the change is towards
there
simplification,
eddies in the current.
are
Moreover, it is by no means
probable that
all the resorbed
substances
are
actuallybuilt up into protoplasm
before being katabolized.
It is true that,to the best of our
edge,
knowlthe metabolic
take place within
the cells but it approcesses
pears
that
the
of material
unlikely
relatively
somelarge amounts

146

times katabolized

firstbecome

must

ANIMALS

FARM

OF

NUTRITION

integralparts

of the

protoplasm.

words, it is probable that the cells have the power to kataba

olize substances present within them but not structurally
part of
In other

them.
The foregoing
conception
Synthetic processes in the body.
of metabolism
impliesthat the body has power to carry out extensive
that the course
chemical syntheses,contrary to the idea stillcurrent
the buildingup of
of chemical change in the organic world is toward
complex compounds in the plant and their breaking down to simpler
in the animal.
Synthethicchemical changes were
long regarded
ones
in the animal
while
the
reactions
as
peculiarto the vegetablekingdom,
supposed to be exclusivelyanalytic. The first synbody were
thetic
of
the formation
action to be recognized in the animal was
hippuricacid from benzoic acid,discovered by Keller and Wohler in
attention.
attracted
wide
More
recent
logical
physio1824, and which
isolated
have shown that this is by no means
an
investigations
executed in the animal
case, but that synthesesin great variety are
animal
and vegetable
body. No such sharp distinction between
laws
formerly supposed. The fundamental
organisms exists as was
for both and both execute
the same
of metabolism
are
syntheticas
It is only the specialsyntheticactivity
well as analyticprocesses.
of the chlorophylin green
mental
plants which tends to obscure this fundalikeness. The conception,then, that the digestivecleavages
supply to the body cells comparatively simple "building stones"
which are synthesized to produce the complex ingredientsof cells
and tissues is quite in harmony with our
generalknowledge of the
204.

nature

"

of metabolism.

205.

Metabolism
as

whole

Oxygen
blood

and

reacts

productsof
are

and

and

oxidative
may

"

be characterized

is introduced
with

analytic.

into

the feed

or

the

Metabolism

garded
re-

chemicallyas an
system through

tissue materials

or

with

dation.
oxithe
the

their

breakingdown, and the final excretory products

either completely oxidized substances,like carbon
dioxid
approaching this condition, like
water, or substances

urea, etc.
From
a

different pointof view, metabolism

as a whole
slightly
be characterized as an analyticas opposed to a synthetic
may
The
of
the formation
generaltendency is toward
process.
of dextrose or
For example, the molecule
simplermolecules.
levulose

contains

24

atoms

and

fats, respectively,
155,
molecules

of carbon

dioxid and

those of the three

167
water

and

173

most

atoms,

from
resulting

mon
com-

while

their

me

the
tab-

METABOLISM

147

olism contain

each.
Even
but 3 atoms
of protein which are resorbed from
the

exceptions,much

few

result from

which

cleavageproducts

digestivetract

complex than

more

the

final

with

are,

products

their metabolism.

Metabolism

206.

the

gradual

process.

While

"

metabolism

justbeen characterized as an oxidative process, and is often

looselyspoken of as a burning of the feed or tissue ingredients,
it is in fact radically
different from what is commonly
stood
underThe building up and breaking down
of
by these terms.
is a gradual,
materials in metabolism
a step by step,process.
i.e.,
has

Metabolism

is the

of the chemical

through which
the life of the cells is manifested.
These
reactions,however,
and even
in the
differ from tissue to tissue and from cell to cell,
time
cell from
to time, and
the resultingproducts are
same
sum

numerous
correspondingly

supplied

the cells

to

metabolism

excreted

as

varied.

and
the

by

reactions

blood

from

the

Between

and

the final
there

body

the nutrients

products of
innumerable

are

intermediate

products,comparatively few of which, in all probability,

have been recognized. We know
the firstand last terms

of the series and

thus

able

are

it were, the algebraic

of the singlefactors making up

to

of the

changes,but
the so-called intermediarymetabolism
in the changes,we
tissues concerned
know
that they are numerous.
we
sum

Furthermore, while metabolism

highly oxidized products,it

direct union

of oxygen
step processes of which
of partial oxidations.

with

as

measure,

as

well

of the

as

though
largelyignorant, al-

are

results in the

does

not

feed

it is made

specific

consist

formation

of

primarilyin

the

materials,i.e.,the step by
up

do

not

consist of

series

The

primary processes of metabolism

of the nature
of cleavagesand hydrationsand it is only
are
the comparativelysimple molecules resulting
from these which
unite directlywith oxygen.
of
Correspondingly, the extent
metabolism
of functional activity
is determined
by the amount
of the various
in

cells and

not,

fire,
by the supply of
notion

increased

voluntaryincrease

in the rate

material

except

so

far

exertion.

to be oxidized
as

increased

in the

case

(193)

oxygen

an

more

that

as

The

somewhat

proportionof
and

of direct oxidation

in the air or

oxygen

depth of breathingmay

in the

body is without

breathing involves

mon
coma

cause

foundation,

increased

cular
mus-

148

NUTRITION

207.

OF

ANIMALS

FARM

As

implied at the opening

of this chapter,the vital activities of the body are essentially
transformations
of energy.
The
livingbody is continually
doing work upon its surroundingsand continually
loosingheat
and the mainto them and the energy for the productionof work
tenance
of the body temperature is derived,as alreadystated,
from

Purpose

the transformation

the substances
the

metabolism.

of

broken

of the chemical

This

process.

will be

occasion

but
detail,
conceptionthat the final end

matter

is the

least its

at

in

being indeed
is familiarly,
if not
that

statement

the

consider

aim

and

of metabolism

is to supply

that the demand

factor
controlling

transformations
are

to

for the vital activities and

energy

contained

this aspect of the

it is important at the outset
to grasp the

in

later

fact
the

altogetheraccurately,expressed in
feed is the fuel of the body.
There

energy

down, this transformation

of the whole

essence

"

in all its processes.

which, if not synonymous

of energy

for

ergy
en-

It is these

with

life,

objectivemanifestation.

while it is essential to hold fast to this broad

But

ception
generalconof metabolism, it is also importantto understand
clearly
that the processes by which this end is reached are exceedingly
cussion
complex. A volume would be requiredfor any adequate disof existing
even
knowledge regardingthe details of the

metabolic

processes.

Such

discussion

the scope
in this chapteris

lies outside

All that is attempted

of the present work.
to outline the metabolism
of the principal
groups

stances
of feed sub-

subsequent consideration of
their values as sources
and energy
of matter
to the body, to
indicate the functions which
they perform in the buildingup
and maintenance
ities.
of the organism and the support of its activand,

"

or

as

2.

preliminaryto

Enzyms

of

views

seems

208.

metabolism

are

been

seen

to

in

that

in

Metabolism

play so largea part, even if a more

the current
conceptionsof the processes
of
the
brief
outline
a
prevailing

called for.

Extracellular

tract

Agents

as

Enzym action has come

less hypothetical
one,

those which

enzyms.
are

most

"

of the digestive
enzyms
familiar in physiology. As has

(114),the digestionof

The

all three of

the chief classes

METABOLISM

149

of feed

ingredientsis brought about

agency

and

effected

is often

Thus

maltose

while

in successive

different enzyms
the saliva converts

by

ptyalin of

the

stages.

their

largelyor wholly by
starch

into

of the latter into dextrose

the further conversion

Quite similar are

by the maltase of the intestine.
the successive actions of pepsin,trypsinand erepsinon the proteins.

is effected

In all these cases,

well

as

in the

even

germinating seeds,the

of the diastase of

produce them

the cells which

from

as

familiar

more

act

enzyms

and

at

case

tance
dis-

have, therefore,

called extracellular enzyms.

been

Intracellular

209.

obvious
acted

enzyms.

"

the fact that

From

the

most

of enzym
action were
those in which the ferment
to
distance from the cells producingit,enzyms
came

cases

at

belonged in a different
that of living cells. A sharp distinction was
category from
drawn
between
unorganized substances,acting substantially
chemical reagents,and organisms producing chemical
changes
as
whose

substances

be

regardedas

by

virtue of their life. The

afforded

action of the yeast plantupon sugar

of this distinction.
shown
It was

typicalexample

action

which
was
(invertase)
capableof
independentlyof the action of the yeast cell,,

that yeast secreted

an

enzym

invertingsucrose
while, on the other hand, the alcoholic
saccharids

held

was

be

to

of

fermentation

of the

vital function

mono-

livingyeast

cells.

Buchner, however,
could

there
which

be

enzym.

monosaccharids
zymase.

1897, showed
from

extracted

yeast

that
a

by suitable

means

(zymase)

substance

simple sugars exactly like yeast in the

of any livingorganism whatever; i.e.,
it acted as an
It became
evident,then, that the yeast cell ferments

fermented

absence

in

The

the

only

it is alive but

because

not

because

essential difference between

it contains

the yeast fermentatio

diastase or by

that, for example, produced by

the invertase of yeast is that the enzym
normally acts within
the cell which produces it. Later it was
shown
that what is
and

true

of the

caused

yeast fermentation

by the lactic

acid

which

can

is true

bacillus.

It, too, is due

to

an

cellular
intra-

separated from the cell and act

gard
independently. Investigatorsare inclined,therefore,to reof
of which,
all fermentation
the work
as
some
enzyms,
like the digestive
excreted by the cells and may
are
enzyms,
enzym

be

also of the fermentation

NUTRITION

150
act

at

while

OF

considerable

others

ANIMALS

FARM

distance

from

their

point

their effect within

normally produce

of

origin,

the secreting

cell.
210.

Intracellular

in the

enzyms

body.

"

Still more

recently

in all parts of the animal

the presence
of intracellular enzyms
body has been recognized. It has been shown that a very

considerable
in the

body

varietyof reactions which

may

action of extracts

also be

brought about

of various tissues and

are

known

outside the

under

organs

take

place
body by the

to

conditions

sequently
apparentlyexcludingthe action of any livingorganisms. Conthey have been ascribed to the action of enzyms
originally
present in the cells,and the reactions in the body
have
been
The
regarded as due to these same
enzyms.
idea of intracellular enzyms
has thus been extended to account
for the metabolic
activities of the organism,and this explanation
has been very generally
According
acceptedby physiologists.
metabolic
to this view, the body cells bring about
changes
in
substantially
the lactic acid
enzyms

This

which

phase of
unanimity

the

same

way

as

viz.,by
bacillus,
act

the

the

upon

subject

cells of yeast or of
the formation
of appropriate
do

the

substances
is

to

be

metabolized.

one
comparatively new
been
and
has by no means
to individual cases
as
reached,but of the value of the generalconceptionas a working
hypothesisthere can be littlequestion.
The word explanationis used above, of course, in a limited
It is not known
how the cell producesenzyms,
with
sense.
nor
acts.
degree of certaintyhow an enzym
Nevertheless,
any
if confirmed, is a real explanationas far as it
this hypothesis,
gether
goes, in that it enables related phenomena to be grouped tofrom a broader standpoint,
will be apparent from the
as
followingparagraphs.

211.

Enzym

reactions

reversible.

said to be reversible when

accordingto

the conditions.

it may

"

progress

chemical

reaction

is

in either direction

gen
example,if a mixture of hydroand iodin in molecular
proportionsbe heated to 448"C.
hydrogen iodid is produced. If, however, hydrogen iodid be
heated to the same
temperature it yieldshydrogen and iodin.
The reaction between
these two elements,then, is represented
by the equation
H2 + I2 ^
(HI)2
For

FARM

OF

NUTRITION

152

Reversibilityof metabolic

212.

ANIMALS

reactions.

It would

"

appear,

which are believed

of the intracellular enzyms
be
to play such an
important part in metabolism
may
and that the metabolic processes
syntheticas well as analytic,

then, that the action

be conceived

may

The
of

as

accelerated

now

state

of

idea

complex

and

that

retarded

now

each

of reversible chemical

changes,is an
and
simplicity,

of

one

the

attractive
there

another

or

and

comparative

that it contains

little doubt

be

to

seems

in its domain

substance

its breadth

in

one

zyms.1
by appropriateen-

body thus exists in a

accordingas
equilibrium,

cell of

chemical
constantly shifting

the concentration

tions,
reac-

will prove an important aid to research.

yet, however, it is to be regarded as a probablehypothesis
of truth and

elements
As

rather than

"

3.

as

The

fullyestablished
Metabolism
The

213.

Glycogenic

of

hexose

function

fact.

the

Carbohydrates

carbohydrates
of

liver.

the

The

"

monosac-

dextrose)produced in the digestive

(principally
cleavage
of the carbohydratesare resorbed chiefly
or whollyby the blood
unite into the
of the intestines. These
capillaries
capillaries
portal vein leading to the liver,where it subdivides into a
capillarysystem in which the blood is brought into intimate
charids

it passes
the cells of that organ and from whence
by way of the hepaticvein into the posteriorvena
cava, thus
enteringthe generalcirculation (182).
contact

with

proportionof dextrose found in the blood of the general

circulation is remarkably constant, and
if any
considerable
neys
be introduced
it is promptly excreted through the kidexcess
(198). On the other hand, the supply of carbohydrates
be more
less intermittent
from the digestivetract may
or
or
for
that there is evidentlyneed
some
so
tory
regulafluctuating,
in
mechanism
of sugar by excretion
to prevent a waste
in two
the urine.
This regulationis effected chiefly
localities,
The

viz.,in the
1

that
For

That
enzym

muscles

syntheses can
reactions

and

be eSected
in

general are

by the

into dextrose.

on
a

function

of enzyms

agency

reversible

example, the ferment maltase, acting

but isomaltose, and it is claimed
that

isomaltose

The

in the liver.

is

seems

of the liver
established,but

questioned by good

dextrose, is stated
different enzym

is

to

authorities.

produce

requiredto

not

tose
mal-

reconvert

METABOLISM

the
in this respect was
considered
appropriately

earliest

be discovered

to

and

may

53
be

first.

being freely resorbed from the digestive

in the liver,
tract, it undergoes dehydration and polymerization
yielding the polysaccharidglycogen (25),which is stored up
trose
in the liver cells. If,on the other hand, the resorptionof dexthe supply
from the intestines is insufficient to maintain
in the blood, glycogen previously formed
undergo the
may
rise to a proreverse
duction
process of hydration and cleavage,giving
This regulatoryactivity,discovered
by
of dextrose.
in 1853, by which carbohydrates are held back
Claude Bernard
of the body, is called the
released according to the demands
or
is

dextrose

When

glycogenicfunction

214.

be likened

has other

to

storage reservoir by which

is controlled.
of

Mechanism

this function

drates
later,as respects the digestedcarbohy-

aspects, as will appear

the liver may
the flow of a stream

While

of the liver.

regulation.
"

It is of interest

to

note

illustrates two of
carbohydrate metabolism
the general conceptions formulated
on
precedingpages.
First,the formation of glycogen is a syntheticreaction. The
porarily
comparatively simple molecules of dextrose are built up teminto the more
complex molecules of the polysaccharid.
In other words, almost the first step in carbohydratemetabolism
is an anabolic change (203).
cogen
Second, the process of the formation and destruction of glyaction
reis susceptibleof explanationas a reversible enzym
that the conversion of glycogen into
(212). It is known
be exwhich may
tracted
dextrose is effected by an enzym
or
enzyms
be similar
must
from the liver and which, it would seem,
of the latter,
The action of one
to those of the digestivetract.
maltase, however, is claimed to be reversible (211),and one is
cogen
naturallytempted to infer that the synthesisof the liver glyage,
which bringsabout its cleavis effected by the same
enzym
although experimentalproofthat such is the case is lacking.
According to this hypothesis,the changes taking place in the
be representedby the equation
liver would
that

this

phase

of

n(C6Hi206)^_
An

excess

of dextrose

C6nHi0nO5n + n(H20)

in the

blood

would

displacingthe point of equilibriumin

have

the

effect of

the direction of the for-

NUTRITION

154
mation

of

OF

ANIMALS

FARM

of dextrose would have the

deficiency
contrary effect. If it be supposed further that the glycogen
formed
combines
with the protoplasm of the liver
as
soon
as
cells,
forming compounds which withdraw a considerable portion
after the analof it from the sphereof action of the enzym,
ogy
of the precipitation
of an insoluble compound, we
have a
if chiefly
scheme
of the chemical
even
plausible,
hypothetical,
mechanism
of the process.
Whether
not it adequatelyrepor
resents

glycogen,while

the actual
of the
be

may
215.

facts,it
manner

appliedto

Muscle

in which
metabolic

glycogen.

the liver has been

the

the liver excluded

from

at

may

"

While

least
the

serve

as

concrete

conceptionof

enzym

lustra
iltion
ac-

processes.

the

glycogenicfunction of
extensive
investigation,

subjectof very
of glycogenis by no means
the presence
confined to this organ.
constituent of animal
to be a normal
Indeed, glycogen seems
in practiprotoplasm. It is found in greater or less amounts
cally
all tissues,
abundant
where
being particularly
rapid cell
is taking place,as in embryonic tissues or in
multiplication
It is estimated
that in an animal in
rapidlygrowing tumors.
condition roughly one-half of the glycogen of the body
normal
is contained in the liver. Of the other half by far the larger
proportionis found in the muscles (96).
The glycogen of the muscles
(and other organs)is not simply
glycogenwhich has been formed in the liver and transportedto
the muscles, but is produced independentlyfrom the dextrose
of the blood,apparently in much
in the
the same
manner
as
liver. That this is true is shown
by the fact that glycogen is
still formed
in the
muscles
when, by surgicalinterference
the blood is prevented from passingthrough the
(Eck fistula),
liver. In fact,the formation
of glycogen in the muscles, etc.,
to be the primary process, while the liver serves
rather
appears
be eliminated
without
as
a
secondary reservoir which may
With
the generalcarbohydratemetabolism.
seriously
affecting
from

the

the dextrose
circulation,

resorbed

the

animal

into glycogenand the

tract is stillconverted
digestive
is stillable to digestconsiderable quantities
of carbohydrates

without
Even
sugar

the appearance

in the normal

is not

the conversion

of sugar

in the urine.

plus
animal,however, the power to disposeof surIf largequantities
unlimited.
of sugar are consumed,
into glycogen,togetherwith the normal katabolism,

METABOLISM

excretion
an
pace with the resorptionand there occurs
"
the so-called
in the urine
alimentaryglycosuria." The

keep

not

may

of sugar

"

which

of sugar

amount

without

be resorbed

can

glycosuria, i.e.,the limit of tolerance

kind of sugar, being highestwith dextrose

carbohydrates of

the

has been
sugar

in

Carbohydrates formed

216.

seen,

supply
from

varies

"

the

body.

and

muscles

feed, the

their relation

In

"

liver act,

as

sort

the

in the

the

(220)

feed, however, by
these

from

body

other materials.

on
exclusively

meat

or

fat,shows

normal

bohydrates
car-

of metabolism
tured
manufac-

carnivorous

the

The

body.

the

from

substances

in

results

means

no

course
appear to be essential to the normal
if they are absent from the feed,they are

and

with

of storage reservoir or regulatorof the

of carbohyThe
total withdrawal
to the blood.
drates
as

disappearance of

e.g., fed

the

producing alimentary

for sugar

"

to

55

animal,

percentage

blood, while its liver and muscles contain a

normal amount
of glycogen. It is true that in such an
ment
experithe
contained
in
meat
small quantitiesof glycogen are
is entirelyinsignificant
as
pared
comconsumed, but their amount
of dextrose

in its

with

the

of
quantities

dextrose

which

there is

to

reason

produced and katabolized in the organism. This

dextrose must
obviously have its origineither in the proteinsor
believe

are

the fats.

be

thus

utilized will be

the metabolism
217.

of the two

Which

is the

source

considered

of

fat.

The

"

both

whether

can

with

in connection

later

(235,253).

of those substances

Formation

or

of

transformations

mutual

of the
glycogen tend to keep the dextrose content
blood approximately constant, while holding a supply of readily
available carbohydrate material at hand to meet
promptly any
be
sudden
demand.
of carbohydrates which can
The amount
it is
disposed of in this way is,however, limited. For man
sugar

and

estimated

at

(96).
excess

about

300

and

grams

for cattle at about

It is evident,then, that if the feed contains

of

carbohydratesover

the needs

capacity to store them up as glycogen will

A surplus of carbohydratesover
the amount
of in this way
of fat,which

is
may

in the cells of connective

in those

appliedby

stored up

be
tissue

immediatelybeneath

the

through

soon

of the

grams
kilo-

2
a

body

manent
per-

the

be exhausted.

which

organism

can

to

be disposed
the

duction
pro-

in very large amounts

the body, but especially

the skin and

about

the abdominal

156

NUTRITION

ANIMALS

the adipose tissue (94).

constituting
of non-nitrogenous
material
a
reserve

organs,

later if need

mobilized
Of

FARM

OF

well

fats,as

of the

as

tissue

which

stitutes
con-

be

may

arises.

chemistry of

the

This

the

of

conversion
or

organ

organs

carbohydratesinto

where

it is

effected,

knowledge is still meager, but the fact of such a change

is undisputed and
it is perhaps the most
notable
example
of a synthetic and
anabolic
in the animal
body.
process
The
the quantitative
evidence for this fact and
physiological
relations of the process
be taken up more
conveniently
may
later (249).
218. Katabolism
of carbohydrates.
The physiological
nifican
sigour

"

of the dextrose
muscles
not

and

liver appears
in the lightof a more

in the

body, but
Of

rather

as

of the blood

and

the

clearlywhen

most

glycogen of the

regarded,

they are

or
temporary storage of matter
carriers of energy for the physiological

less

these

which

the most
processes,
vastly predominatesover all others,is the

performanceof

by

what

processes.

the

muscles, external

work

is in the main

and

internal,but

obvious

one,

is true

also of the subordinate

true

of

work
cular
mus-

forms

of

glandularand cellular activity.The former, therefore,

may
be taken as typical.
In the performance of muscular
work, as will appear later,
there is a rapid katabolism
of non-nitrogenousmaterial and
of carbohydrates,
it would appear, in the form
especially
largely,
of dextrose.
The
resultingimpoverishment of the blood in
dextrose causes
of stored up glycogeninto dextrose
a conversion
to supply the lack.
If the view of the formation
of glycogen
which regardsit as a reversible reaction may
be accepted,we
equilibriumbetween the dextrose
may
say that the chemical
and the glycogen is disturbed by the removal
of the former
is continued,the
As long as the work
during muscular work.
of glycogeninto dextrose also continues,
process of conversion
and by prolongedwork it is possible
to reduce the glycogencontent
of

animal

an

It should

to

very

low

limit.

be

clearlyunderstood that the foregoingis only a

highlyschematic view of the chemistryof muscular contraction
of the carbohydrates. Some further
as related to the katabolism
consideration is given in Chapter XIV
plicated
(630) to the very comchemical

mechanism

of the process.

METABOLISM

219.

Intermediary
of

katabolism

katabolism.

57

Regarding the intermediary

"

the

is known.
carbohydrates, not very much
It appears
probable,however, that dextrose undergoes preliminary
of
with
the
formation
cleavage
glyceric aldehyde,
pyruvic aldehyde (methyl glyoxal)and either lactic or pyruvic
acid,which is then further oxidized to acetic acid,carbon dioxid
and
water.
facts,including especiallythose derived
Many
from a study of the fermentation
of sugar, seem
to point to the
tributed
Lactic acid is also widely dispossibilityof such reactions.
in the body, although its presence
is also susceptible
of explanation as arisingin the katabolism
of protein (233),
it has been shown
that lactic acid may
and, moreover,
give rise
to
body. Accordingly,
glycogen or dextrose in the animal
transformations
of glycogen and
these changes, like the mutual
be conceived
of as constituting
series of rea
versible
dextrose,may
reactions.
(Dextrose)

^ CH2OH(CHOH)4CHO

Glycogen

(Glycericaldehyde)

^t 2CH2OH

CH2OH(CHOH)4CHO

CHOH

"

"

CHO

(Pyruvic aldehyde)

CH2OH

CHOH

"

CHO-H20

"

CH3

"

CO

"

CHO

(Lactic acid)

CH3

CO

"

CHO

"

H20

CH3CHOH

COOH

"

or

(Pyruvic acid)

CH

"

The

CO

"

CHO

conversion

"

of dextrose

CH3CO

"

COOH

into lactic acid

is

nearly iso-

the
process, the resultinglactic acid containing almost
the dextrose.
of chemical
amount
as
If, then,
energy

thermic
same

these

cleavagesoccur in
obviously preparatory
portion of the
principal

the katabolism
to

energy

The

The

the
man,

the

actual

and

of the
The

amounts

oxidation

which

in

the

pentosecarbohydrates
treated

constitute the chief

carnivora

feed of

carbohydratesthey are

is liberated.

foregoingparagraphs have

hexoses,which

of

so

far

as

of

carbohydrate supply of

the latter

herbivora,however,

of various

of the metabolism

hydrates.
carbo-

consume

contains

which
pentose carbohydrates

erable
also considare

in

part

158

OF

NUTRITION

FARM

ANIMALS

the feed duringits passage

or at least disappearfrom
digestible,
through the alimentarycanal.
In general,
it may
be stated that the
220. Pentose
sugars.
arabinose and xylose),whether
pentose sugars (in particular
administered
by the stomach or injectedinto the blood, are at
oxidized in the body. The pentoses differ from
least partially
in the fact that the limit of tolerance in the
the hexoses chiefly
of hexose carbohyamounts
blood (215) is lower.
Excessive
drates
"

cause

excretion

an

of

sugar

in the urine.

effect is produced by the pentoses, but much

are
relatively,
requiredto bringit about.

The

smaller

have found
Most, although not all,investigators

same

quantities,
increase

an

the ingestion
the liver consequent upon
of
pentoses, but in every case it has been the ordinaryC6 glycogen,
in the

glycogen of

that
indicating
221.

the effect is an

Pentosans.

pentose sugars

or

"

indirect

one.

The

their

investigationsupon
derivatives justreferred

the

soluble

to have

shown

that

they are to a greater or less extent assimilable. The pentose

in
the
feed
of
exist
to a
herbivora,however,
carbohydrates
They are chiefly
very limited extent, if at all,in this form.
polysaccharids,
being either pure pentosans or combinations
In discussing
of pentosans and hexosans.
the nutritive value
of these pentosans, it seems
assumed
that
to have been frequently
they are converted into pentoses duringdigestion.As a matter
of

fact,however, there is no

direct evidence

while Kellner's results

that such is the case,

to believe that they

(129)afford reason
are
largelyfermented
along with cellulose,yielding,besides
products,chieflyorganic acids. If this is the case,
gaseous
farm animals do not acquirefrom
their feed any considerable
of pentoses and conclusions drawn
from experiments
amounts
with the pentose sugars regardingthe nutritive value of these
substances
are
inapplicableto ordinary stock feeds. Their
true value in the latter would
be simply that of the productsof
their fermentation.
The
222.

Formation

in

organicacids

digestion.
"

As

was

shown

in

Chapter

(128-130,132),a considerable proportionof both the hexose

and pentose carbohydratescontained
in the feed of herbivora
undergoes fermentation in the digestivetract, giving rise,in
IV

160

NUTRITION

FARM

OF

ANIMALS

productsof proteinkatabolism which will be considered in the

following section, contains also non-nitrogenousmaterials,
presumably arisingfrom the incompletekatabolism of ingredients
In the urine of

of the feed.

and

man

of the carnivora

these

or
chiefly
wholly such as might
of proteins(phenolsand other
be derived from the katabolism
and
their amount
is comparatively
compounds of the aromatic series),
In the urine of herbivora,particularly
of
small.
erable,
considruminants, however, their quantity is relatively
very
and it seems
impossibleto regard any largeportionof
them as products of proteinkatabolism.
225. Origin.
Apparently these non-nitrogenousorganic
in some
substances originate
from the roughages. Their
way
sists
proportionin the urine is relatively
largewhen the ration conof roughage,and the addition of such feeding
exclusively

substances
non-nitrogenous

are

"

stuffs to

basal ration

causes

marked

increase in their amount,

vestiga
have been inconcentrates
as

the other

hand, such
do not produce this effect to any very considerable
fixed
extent.
to bear no
seems
Furthermore, their amount
relation to the proteinof the feed.
of the
When
the amount
latter ingredient
is small, the total organicmatter
of the urine
has in some
exceeded the digested
cases
proteinof the feed,thus
demonstratingthat a portion at least of the non-nitrogenous
As
have had some
other source.
urinary constituents must
of
the proportionof proteinin the feed increases,the amount
nitrogenousproductsin the urine likewise increases,while that
of the non-nitrogenous
constant,
products appears to be more
The
that the ratio of urinarynitrogento carbon increases.
so
most
to be that the
plausibleexplanationof these facts seems
of the non-nitrogesubstances in questionare derived from some
nous
ingredientsof the roughages,but from what ones, or
what is the nature
of the products,we are stillignorant.
while, on

"

4.

The

Metabolism

of

the

Simple

Proteins

Anabolism
226.

Synthesis

of

proteins from

digestive products.

"

The

paratively
simple proteins are resorbed (139, 152) in the form of comsimple cleavageproducts; largelyas amino acids
but in part perhaps as more
less complex polypeptids. Out
or

161

METABOLISM

body builds
specific
proteinswhich are peculiarto
of these substances

the

chemical

up the great varietyof

itself and which
differ in

proteinsof the feed,

those of the vegetable kingdom
(147). This
of building animal
proteinsfrom the fragments of
process
conspicuous example at once of
vegetableproteinsis the most
of the animal
the syntheticpowers
organism and of the object
and
properties
from
especially

of the
227.

from

structure

the

cleavage.
digestive
of

Seat

As

protein synthesis.
"

regardsthe place where

synthesisof proteinsoccurs, opinions are divided. Until

most
experimentershave not been able to detect the
recently,
products of digestive
cleavagewith certaintyin the blood,either
in the general circulation or in the portalvein,and the current
view has been, therefore,that of Abderhalden, viz.,that the
this

"

"

buildingstones

the

of

proteinsare

cells of the intestine and

epithelial
in particularserum
albumin
"

that

to

serve

nourishment

as

Various

of greater or
with the aid of more

Denis1

and Van
that

less

in amounts

amino

acids

that

to

may

seem

pass

be found

methods

be doubled

have

to

through

shown
the

tissues

that

was

likewise
acids in

and that the increase affects the blood

of amino

amounts

blood of

of the
has

acids from

The evidence
livinganimals.
in favor of the
seems
therefore,
decisively
into which

and

beyond
resorbing

and

all
practically

circulatorysystem and not that

only, while Abel,3 by a diffusion method,
considerable

body.

Folin

(152). The latter experimenters have

after meat
feedingthe proportionof amino

the blood may

of the entire

secure

of the

in the blood

for

account

the blood

to

on

reportedthe presence in
of non-proteinnitrogen

chemical

Slykeand Meyer2

sufficient

administered

amounts

refined

question
epithelium unchanged and

shown

proteintissues

investigators,
however, have

the blood
and

the

to

passed

are

"

synthesizedin the
the resulting
proteins

portalvein

been

able

ing
the circulat-

the present
view that the

at

to

time,
ments
frag-

the

proteinmolecule is splitduring digestion

and serve
pass without material change into the blood current
from which the proteins,
both of the blood
as
a common
source
and the various tissues,
built up and that every livingcell,
are
each
1
3

in its

own

measure,

has

this anabolic power.

Jour. Biol. Chem., 11 (1912), 87.

Jour. Pharmacol, and Expt'l Therap.,
M

Jour. Biol. Chem., 12 (1912),399.

(1914),
275.

62

NUTRITION

OF

FARM

ANIMALS

Katabolism
228.

Nitrogenous end

products.

"

The

total katabolism

of

proteinsresults in the elimination of all their nitrogen

throughthe kidneys in the form of the various relatively
simple
the
found
in
urine.
Of
the
nitrogenous
products
crystalline
is the most
and
of
the
man
carnivora,urea
excretory products
prominent,while others,such as uric acid,creatin,creatinin,
ammonia, etc., are of subordinate importance quantitatively.
and
Traces
of hippuricacid are also found in the urine of man
in that
carnivora,while it is present in relatively
largeamounts
of herbivora
along with considerable quantitiesof ammonia
and apparentlybut little urea.
of the urine of mammals
other
The nitrogenousingredients
from the
than those justmentioned
either derived chiefly
are
whose
will be considered
metabolism
later,
nucleoproteins,
to call for no
are
as
or
special
present in such small amounts
from
consideration
the present very
generalpoint of view.
it should be noted for completenessthat a small amount
Finally,
and
of nitrogenousproducts is eliminated in the perspiration
also that from one
point of view the incompletelykatabolized
nitrogenousexcretory products of the feces (154)may also be
regardedas productsof proteinkatabolism.
the

dicarbamid,CO (NH2)2, is the chief nitrogenousproduct

of the simple proteinsin carnivora and omnivora.
of the katabolism
In human
urine from 8o to 90 per cent of the nitrogenis ordinarily
be considerably
present in this form, although the proportion may
diminished
under specialconditions,
notably on a low proteindiet.
Urea, however, is not simply splitoff as such from the proteinsas
Urea,

or

earlier schematic

some

statements

have

sometimes

been

taken

to

is ammonium
antecedent of urea
carbonate,
imply. The immediate
which undergoes a dehydration in the liver or elsewhere, while there
is brought to the
is evidence in favor of the view that the ammonia
liver in the form of ammonium
lactate. At any rate it is an accepted
of the nitrogen of the simpleproteinspasses
fact that most, if not all,
That
to excretion as urea.1
through the ammonia
stage on its way
from ammonia
the formation of urea
function of
is not exclusively
a
the liver is shown by the fact that it stillcontinues
when
this organ
is excluded from the circulation by means
of an Eck fistula.
1

which

It has been, shown

splitsoff the

that the liver,

kidneys and other organs contain an enzym
from
guanidin group
arginin(47) producing urea and ornithin.

163

METABOLISM

Hippuric
urine

acid

in small

of mammals

but

is

amounts

normal

constituent

of

the

is

in that of herbivora.
especiallyabundant
is the result of a synthesis(204). When
Its formation
benzoic acid
other
or
compounds
containing the benzoyl radicle are introduced
into the circulation they are
paired with glycin,one of the cleavage
the
of
in
products
proteins, the kidneys and excreted
as
hippuric
is
acid, which, chemically, benzoyl-glycin,or benzamidoacetic
acid,
COOH.
The
normal
of
small quan(C6H5 CO)NHCH2
presence
tities
of hippuricacid in the urine arises from the fact that the putrefaction
of the proteinsin the intestines yieldscompounds containing
the benzoyl radicle which
resorbed
and
combine
with glycin to
are
form hippuricacid.
But a small proportionof the hippuric acid produced
however.
of
Most
it
by herbivora can be thus accounted
for;
its originto the roughages consumed
to owe
by these animals,
appears
especiallythose derived from plants of the gramineae,while, on the
other hand, concentrates
do not seem
its amount.
to increase
parentl
Apits formation
bears some
relation to some
of the ingredients
of the cell walls,but to what
in particularis not clear.
ones
"

"

229.

The

non-nitrogenous

nitrogenous products
of the

oxidation

residue.

eliminated

protein molecule

of considerable

amounts

of carbon

in

In

"

the

gives

addition

urine,

rise to

dioxid

and

the

the
water,

to

the

complete
production
which

are

excreted

channels as those derived from the

through the same
of carbohydrates or fats. To put the matter
katabolism
in the
while
the urinary products account
for all the
reverse
way,
nitrogen of the protein, they contain but a relativelysmall
This is clearlyshown
part of its carbon, hydrogen and oxygen.
of these elements
in 100
by comparing the average amounts
in the urea
parts of protein wTith the quantities contained
corresponding to the total nitrogen of the protein. Disregarding
the sulphur of the proteins,the results of such a computation
are

as

follows

"

Proteins

Carbon
.

Hydrogen
Oxygen

Nitrogen

Urea

Residue

53-o

6.86

46.16

7.0

2.29

4.71

24.0

9.14

14.86

26.0

16.00

100.

34-29

65-7I

64

FARM

OF

NUTRITION

ANIMALS

siderably
abstractingthe elements of urea, there remains conhalf the hydrogen and oxygen
of the proteinand
over
A substantially
the largerpart of its carbon.
similar result is
of the other nitrogenous metabolic
reached in case
products.
off of these products from the proteinsleaves a
The splitting

After

non-nitrogenous residue.
Two

generalstages
be distinguished.
in the katabolism
of the proteinsmay
The
firstis a hydrolysis
by which the proteinsare splitup into their
230.

stages of protein katabolism.

Two

constituent
the amino

amino

acids.

second

The

acids in which

the

"

is

nitrogenof

deaminization

these acids is

of

splitoff

ammonia.

as

231.

of the

Protein

The

hydrolysis.
"

body proteinsis

first stage in the katabolism

less
or
hydrolyticcleavage, more
similar to that effected in digestion
and like the latter brought
about by enzyms,
contained in the body
which in this case
are
cells
the intracellular enzyms
(209).
The truth of this view is attested by the facts that the presence
a

"

of proteases in almost
body has been demonstrated

they effect a rapidsolution

all of the tissues and

and

that under

of the

organs

proper

conditions

proteins the so-called

confirmation
is afforded by the known
autolysis. Further
facts regardingthe transformation
of one
proteininto another
in the body, while finally
the productionin the organism of
of the cleavageproducts of the proteins,
some
presumably as
shown.
productsof katabolism, may be indirectly
232.

Is

katabolism

of the tissue

protein hydrolysis

"

reversible

process

"

If the

of

age
body proteinsis initiated by an enzymatic cleavin the body cells,
the reverse
this is precisely
of the synthetic
is
action by which
believed
it
that body proteinsare
built up out of the products of digestive
cleavage(226),and
the

question at

reversible enzym

once

arises whether

Protein

the

be

have

reaction,analogous to

suggestedas occurringin the case

the generalnature
of which may

It

we

freelyadmitted

that

to

do here with
which

has

been

carbohydrates(214),
representedby the formula

of the
be

Amino

acids

of
proof of the reversibility
action of proteolytic
is as yet lacking,
such phenomena
enzyms
the formation
of the plasteins
discovered by Okunew
as
must

that

165

METABOLISM

and

by the action of pepallegedformation of paraneuclein

sin
Robertson
due
to
as
by
being
apparently
tion
adsorpreported
ties
phenomena.1 On the other hand, however, many authorithe

teristic
as
a
regard reversibility
general characof enzym
action and mere
negative evidence cannot, of
disprovethis belief. At any rate the conception of

course,
a

inclined

are

to

reversible reaction between

acids of the blood

the amino

and

lymph and the proteins of the cells affords a comparatively

simple and unforced explanation of the facts outlined in the
fluence
foregoingparagraphs, as well as of others relatingto the inof the supply of feed protein on metabolism
which will
it may
be observed
be considered later (402). In particular,
all the amino
acids
that, accordingto this view, by no means
resorbed. into

the

blood

would

stream

undergo synthesisto

acid supply were

that, especiallyif the amino
a large part of them
liberal,
might pass directlyto the second
stage of proteinkatabolism,viz.,deaminization.
Finally,since the proportionsof the singleamino acids supplied
from the digestive
must
tract
conceive,not of
vary, one
a
singlereaction between protein and amino acid,but, speaking

proteinsbut

broadly,of
there

amino

are

233.

as

the amino
related

seems

reactions

as

acids concerned.

Deaminization.

katabolism

independent reversible

many

"

second

The

to be the

generalstage

off of the NH2

splitting

of

protein

group

from

or
acids,the productsbeing the corresponding
closely

non-nitrogenousorganic acids

the

on

hand

one

and

also,it would appear, is a case

of enzym
action,althoughthe discoveryof deaminizingenzyms
and its biological
importance
in various tissues is comparatively recent
is stillto some
extent
speculative.
the
of the amino
The ammonia
deaminization
resultingfrom
ammonia

the other.

on

acids is believed
which

it is

resorbed

excess

In this way

the

of the immediate

of urea,
not

into

exclusively

any amino acids

of the body cells

nitrogenof
demands

cretory
promptly converted into exdisposedof, while the larger part of

protein building material

their

antecedent

rapidlyconverted, chieflyalthough
(228).

for

be the immediate

to

in the liver
in

This

products and so
carbon and hydrogen
1
2

Compare

Rohonyi
Compare Bayliss;

Biochem.

The

Nature

is

remains

in

series of substances

Ztschr., 53 (1913), 179.

Action
of Enzym
(1908),Chapter V.

66

FARM

OF

NUTRITION

ANIMALS

gether
less close relation to the fatty acids and toor
bearinga more
of the
non-nitrogenousresidue
constitutingthe
proteins(229).
It is important to note that these non-nitrogenous
products
of the original
contain the largershare of the chemical
energy
gestive
the ammonia
carryingoff but little and both the diproteins,
cleavageand the deaminization being nearly isothermic
of proteins,
therefore,
The cleavage and deaminization
processes.
"

"

necessarilyinvolve

do not

destruction

of their nutritive

of nitrogen in the
given amount
the total destruction
urine is not to be regarded as indicating
of protein. It has ceased to exist as
amount
of a corresponding
residue is made
but its non-nitrogenous
protein,
up of substances
which are closelyrelated chemicallyto both the carbohydrates
be katabolized to supply
and fats,and which, like these,may
and

value

excretion

the

of

energy.
234.

body

reversible.

Deaminization
appears

to be

whether

in this

an

reversible
So

far

in the

evidence

is any

that the reaction

is

one.

direct

as

such

concerned, no
andEmbden

Since deaminization

quire
enzymatic reaction,it is natural to incase, as in the other enzymatic reactions

already considered,there
a

"

and

are
experimentswith deaminizingenzyms
evidence has been produced, but Knoop

demonstrated

have

Schmitz2

mental
fact of funda-

acids may
in metabolism, viz.,that amino
significance
monium
be formed in the body from ketonic or hydroxy acids and amture
manufacsalts. In other words, the animal body can
of the proteins,
at least of the
some
buildingstones
and from the latter presumably the proteinsthemselves
(226),
salts of the correspondingketonic or
of the ammonium
out
of this comparatively
hydroxy acids. The full significance
recent
discoveryis not yet fullyapparent. The questionof the
"

"

considered

salts will be

utilization of ammonium

later.

In

important fact is that these results indicate

series of reactions,
that the reaction,
or
by which deaminization
that the whole process of protein
takes place is reversible,
so
be represented
metabolism
schematicallyas follows :
may
this connection

the

"

_.._...

Proteins
1

Ztschr.

[ Organic acids

j
_"
acids
\
Zt.
^
"

^ Ammo

Physiol.Chem.,

67

(1910),489.

{ Ammonia

Biochem.

Ztschr.,29 (1910), 423.

68

OF

metabolism

may

carbohydrate
thus

FARM

NUTRITION

ANIMALS

schematicallyexpressed

be

"

Glycogen$

Dextrose

"^\
Lower

Proteins

Amino

acids

"$"

fattyacids

CO2 and H20

-"

"

I NH3
Formation

236.

of fat from

""

proteins.
"

Urea

Since the

non-nitrogenous
consist largely

productsof proteinkatabolism appear to

of comparativelysimplesubstances closelyrelated to the lower
least of
and since some
members
of the fatty acid series,
at
be synthesized
in all probability
to
these may
carbohydrates,
while the latter can
undoubtedlygive rise to fats,it is natural
conclude
that the non-nitrogenous products of protein
to
katabolism
of fat,either by direct synserve
as
a
source
thesis
may
of the simplerfatty acid chains or possiblyby way
of
the carbohydrates. The
conclusion
is one
which
has been
in its favor
of the earlier evidence
hotly debated and much
be inconclusive.
has been shown
The
dence
to
experimentalevibe

may
a

discussion

of the

present, it may

proteinseems
the

of animal

sources

suffice

to

say

that

fat

fat thus

formed

in connection

that

under

with

(247-249). For

the formation

probable,but
altogether
of

amount

conveniently considered

more

on

normal

the

of fat from

the other hand

conditions

is

usuallyunimportant.
5. The

Metabolism

metabolism

of the

"
The

of

the

Nucleoproteins

with the excepconjugatedproteins,

tion
terest.
of the nucleoproteins,
offers few features of specialinIn general
it may be said that they are split
up into their
constituents during digestionand that the cleavageproducts
metabolic
the same
sumed
undergo substantially
changes as if conof
In the case
form.
by the animal in the uncombined
of the nucleic
the nucleoproteins,
however, the metabolism
consideration.
acid portion of the molecule
calls for more
specific

Anabolism
237.

Fate

extensive

of

digestiveproducts.

"

The

nucleic acids

enzymatic cleavagesin digestion(139),the

undergo
products

169

METABOLISM

phosphoricacid,
being essentially
By analogy with
pentoses, and purin and pyrimidin bases.
the simple proteins,
one
might expect, therefore,to find that
the circulation

passinginto

fragments

these

acid molecule

nucleic

of the

rebuilt into

are

of which they constitute such

in the body cells,
nucleoproteins
ingredient(75).
an
indispensable
of such a synthesis,
The occurrence
however, has been seriously
gestion
questioned. One argument against it is the fact that the insults
of nucleoproteins,
more
or
purin bases, respecifically
in a prompt excretion in the urine of end productsof their
katabolism
which, although it has not been proved to be quantitative,
excreted on
a
is certainly
large,while the amounts
It has been
purin free diet are small and notably uniform.
purins,i.e.,
exogenous
argued, therefore,that the so-called
"

"

the nucleic acid constituents

Preciselythe

acids in the cells.

the

serving

without

excreted

and

katabolized

from

derived

to

simply

feed,are

nucleic

rebuild

argument might be made,

same

their

of the simpleproteinsfrom
however, against the synthesis

cleavageproducts,since in this
prompt

causes

of the end
238.

differs from

rather than

of

feed

increase in the

It is true

"

of the

that

in the

that

simple proteinsin
from

its

cretion
ex-

proteins
of nucleo-

that the formation

of the molecule

supply

the

fragments l

the stricter sense, while it has been

build up nucleic acids out
body can

synthesisin
that

demonstrated

an

almost

reconstruction

also

quantitativeincrease
(402).
product, urea

and

Autogenesis.

latter is

case

the

supply containing

purins,pyrimidins nor

neither

striking instances of this is

in the development of the embryo of birds and insects.

pentoses.
seen

The

eggs

One

contain

of

the

none
practically

yet the bodies

amounts

most

of the

of nucleic acid.

of the substances

young

animals

tioned,
justmen-

contain

is the
Equally significant

case

normal
of the

sucklingmammal,which receives in the milk a food very poor

in purins,pyrimidinsand pentoses, yet which maintains a rapid
with its accompanying formation
growth and cell multiplication
and Mendel's extensive
of nucleoproteins.So, too, in Osborne
2
normal
the nutritive values of the proteins,
investigations
upon
1

The

of the formation
possibility

of proteinsfrom
of nutrition.

under ordinary conditions

Carnegie Institution of Washington,

Publication

ammonia
No.

nificanc
(234) is of littlesig-

156, p. 85.

OF

NUTRITION

170

ANIMALS

FARM

growth of rats through two generationswas secured on purinfeed.

Another
fact pointingin the same
and pyrimidin-free
to requirea supply
direction is that the body does not appear
but
build up its
can
of phosphorus in organic combination
organiccompounds from phosphates(258).
capacity of

the

absence

entire

may

the

when
in

recombined

be

cleavage products.

view

In

"

of

produce nucleoproteinsin the

it
constituent
buildingstones
may
the latter are suppliedin the feed they

body

of their

supposed that

be

from

Regeneration

239.

to

"

the

"

cells somewhat

as

the

are

amino

acids of the
No
seem,

simpleproteins?
be given to
can
positiveanswer

question. It would
the autogenesisof the

this

however, that the first steps in

of pentoses and
of the
be the formation
must
nucleoproteins
purin and pyrimidin bases, i.e.,of preciselythose substances
the digestive
which result from
cleavages. Even though it be
assumed
that,in the former case, they are produced within the
cells where
they are further synthesizedto nucleic acid, it is
substances
not
brought to the
altogetherclear why the same
should not be available for the same
cell by the blood current
at least,it seems
perfectlypossibleto
Provisionally,
purpose.
contained
buildingstones
regard the entire stock of these
derived from the feed or produced by the
in the body, whether
body cells,as potentiallyavailable for the regenerationof
From
this point of view, the increased excretion
nucleic acids.
be conresults from
their ingestionwould
sidered
of purins which
"

as

the blood
which
acids

and

follows

consequence

"

of their increased

concentration

in

analogous to the increased excretion of urea

the ingestionof simple proteinsor of amino

as

(402).
Katabolism

240.

Cleavages.

"

The

katabolism

of

the

nucleic

of the

acids

simple proteins.
of the latter,
As in the case
the firstgeneralstage of the
consists of a series of enzymatic cleavages. These
process
cleavagesare quite analogous to those of the simple proteins
and yieldas final products the comparativelysimple building
of the nucleic acids.
Since it is to be supposed that
stones
the autogenesis(238) of these compounds is via these same
bears

close

generalresemblance

to

that

"

"

METABOLISM

"

"

buildingstones
of

case

it would

that

appear

simple proteins,a complex

the

have

we

of

here,as

reversible

71

in the
enzym

reactions.

Phosphoric acid
Nucleic

acid

Pentose
Purin

Deaminization.

241.

the nucleic

off from

phosphoric acid

is,of

acids

of this substance

stock

The

"

in the

added

course,

The

body.

which

is

split
the general

to

pentose may

be presumed

possiblybuilt up into a hexose.

The
bases, on the other hand, like the amino acids derived
from the proteins,undergo, as the second general stage of their
and oxidation.
The
katabolism, an enzymatic deaminization
to

NH2

be

are

groups

or

splitoff

ammonia

as

and

converted

into urea,

ring formations are largelyunbroken, the principal

and
products of purin katabolism being uric acid in man

while
end

katabolized

pyrimidin bases

or

the

allantoin

other

in most

pyrimidin bases

mammals.

little is known.

242.

along with

Synthesis

acid is the

of

the end
uric

The

and

complete, however, purin

the urine

Of

the katabolism
deaminization

pyrimidin

bases

of the
is

never

appearing in

products of katabolism.

acid.

birds

In

"

principalnitrogenous constituent

and

uric
reptiles,

of the semi-solid

portion of its nitrogen can have

existed as preformed purins in the feed,it is evident that these
animals
must
synthesize uric acid. This synthesisappears
the antecedents probablybeing lactic
to take placein the liver,
urine.

acid and

Since

no

considerable

urea.

"

6.

Metabolism

The

of

the

Fats

Anabolism

considering the resorption

shown
of the fats (152) it was
that,while the products of
their digestion are
glyceroland fatty acids (or their salts),
after resorptiononly neutral fats have been recognized in the
epithelialcells and in the lymph of the intestinal lacteals.
The cleavage of the fats in digestionis reversed in the epithelial
cells. It seems
altogetherplausibleto ascribe this resynthesis
since the action
to the action of an
intracellular lipase,
the more
243.

Resynthesis

of

feed

fat.

"

In

of

lipasehas

been

to the blood

of

and

sometimes

processes

the material

case

(211).

of the process

of the metabolic

one

cases

of
is

transmitted

of fats.
substantially
contained in the lymph in the

are

in this state

from

pass

vein.

subclavian

the

some

consists

current

digestedfats

The

as

In either

of indifference.

matter

be reversible in

to

be classed

resorptionor

form

shown

ANIMALS

be regarded as part
resynthesis

this

Whether

FARM

OF

NUTRITION

172

The

the thoracic duct into the blood

blood

much

containingas

as

itself,
however, although
of fat, does not
per cent

be well established

to

sort

and

244.

from

may

Storage

that the fat of the blood

is soluble

which

of combination
which

fats,and

the fat

globulesenteringit
of the
from the thoracic duct do not long persist. The nature
change is stilluncertain ; by some, it has been regarded as a
cleavageinto fattyacids and glyceroland by others as a union
the nature
of the change it seems
.with proteins. But whatever
normally carry

emulsified

emulsified

in

be called for convenience

of

fat.

"

liberal

and

water
"

exists in

some

diffusible

soluble fat."

supply of

fat to the blood

the

digestivetract may give rise to a storage of reserve

fat in the adiposetissues (94)of the body. It is to be presumed
that this deposition
of reserve
fat is substantially
a reversal of
tion
the process, whatever
it is,by which it was
brought into soluin the blood, the
soluble fat
of the latter passinginto
the cells and being there reconverted
into the emulsified form
and so givingrise to the globulescharacteristic of fat cells.
245. Formation
The fats depositedin the
of cell lipoids.
terial,
maas
adipose tissues,
alreadyimplied,are a store of reserve
laid aside temporarilyfrom the body metabolism
when
the feed supply is more
needs.
than adequate for immediate
The
various
more
complex lipoids(37-39, 75), however
lecithins and
other
(cholesterins,
phosphatids, cerebrosids,
of protoplasm and to
to be essential ingredients
etc.),
appear
functions in the cell. All these substances
perform specific
have
their basis fatty acid molecules
as
coupled with other
and it is a reasonable assumption that the former are
groups
derived from
the
soluble fat
of the blood and synthesized
in the cells into the specific
lipoidsas required.
"

"

"

"

246.
serve

Manufacture
as

source

dependent upon

"

of

of

the

fats.

"

But

while

the

feed

fats

may

body fats,the organism is by no means

former for its supply of these substances,

METABOLISM

but

the

to

facture
already been indicated (217,236), manufats from other ingredientsof its feed.
view, first propounded by Liebig in 1843, was
contrary

has

as

may,

This

73

led

and
opinionthen prevailing

to

livelycontroversy

lief.
beresolved in favor of the newer
which, however, was definitely
mals,
aniin case of herbivorous
Indeed, the feed fats,especially
of body fat,
are
usuallyof subordinate importance as sources
in the body.
a large share of the latter being produced de now

explainsin part the generaluniformityof composition

steer
of the body fat of each species. The
produces beef
identical rations
fat on substantially
fat and the sheep mutton
largelybecause the fat depositedin the body is derived only
of it beingproduced by the
in small part from the feed fat,most
metabolic activities of the body cells. The seat of this
specific
synthetic production of fat,however, as well as the manner
stillunknown.
in which it is depositedin the reserve
are
tissues,
This fact

The
247.

of animal fat

sources

evidence.

Experimental

The

"

of animal

sources

fat

feed fat
already indicated. Aside from whatever
the body can
be stored up in the adipose tissues,
produce
may
the carbohydratesof the feed (217) and in all probfat from
ability
from
the non-nitrogenous residue of the proteins(236).
In view of the historic interest attachingto the long controversy
this question,however, as well as of its intrinsic importance,
over

have

been

appropriate.
experimentalevidence seems
of body fat was
never
That the feed fat is a source
seriously
of Liebig's
contention that
the correctness
questioned. When
outline of the

an

the animal
it

body
was

fat had

also manufacture

can

assumed

that

the

source

been

strated,
demon-

of this new-formed

carbohydratesof the feed and this

nology,
for years the accepted view.
Following Liebig's termiwas
the proteins
materials,"
were
designatedas the
plastic
and fats were
while the carbohydrates
servingto build up tissue,
of heat and of fat.
materials,"serving as sources
respiratory

fat

was

to

in the

be found

"

"

248.

Fat

from

facts
the animal

protein.
"

pointing

body, but
1

to

Several

the formation

Carl Voit

Ztschr.

was

earlier

served
investigatorsob-

proteinin
addistinctly

of fat from

the first to

Biol.,5 (1869),79-169.

OF

NUTRITION

174

FARM

ANIMALS

the belief that

proteinconstitutes an important source

of animal fat,this conclusion being based largely
the famous
on
in
respiration
experimentsof Pettenkofer and Voit at Munich
freed from visible fat as carefully
which a dog was
fed lean meat
as
practicableapproach to a
possible(thisbeing the nearest
protein diet) and the balance of nitrogen and carbon
pure
in many
The results showed
cases
(287,292) determined.
a
of
carbon
animal
retention
by the
greater than correspondedto
the quantity of proteingained,and this difference was
preted,
interdescribed
in Chapter VI
according to the methods
(293) as showing a productionof fat.
Pettenkofer
and Voit's experiments were
long accepted as
1
conclusive until Pfliigersubjectedthem
cism,
critito destructive
of material errors
in the estimates
showing the possibility
vocate

of the carbon

It

of both
be

scarcelyneed

fat is not

formed

feed and

visible

excreta.

said that this result does

not

prove

that

from

protein,but simply that Pettenkofer and

Voit's experiments fail to demonstrate
it. Of later experiments
on

the

of

subject,a

small

number

seem

to

show

clearlythe

mation
for-

of fat from

after every
protein,even
allowance
has been made
for the objections
raised by Pfliiger
in his criticisms of the experiments. A number
of negative
results have, it is true, also been reported,
but naturallynegative
results are of much
less value than positive
ones.
Moreover, the indirect evidence in favor of the possibility
of the formation
of fat from
sive.
concluproteinseems
practically
As alreadystated (235),it has been established beyond
reasonable
doubt
that carbohydratesmay
be produced from
sarily
proteinin the body. If this is true, however, it almost necesinvolves the possibility
of the formation of fats from protein,
since carbohydratesare undoubtedly a source
of fat.
249.

Fat
went

formation

from

amount

carbohydrates.

further

than

to

Pettenkofer

"

and

ever,
Voit,2how-

the
demonstrate, as they believed,

of fat from

protein. Their experimentsincluded a

in which
number
ration
of
added
to
a
carbohydrateswere
protein (lean meat). Assuming with Henneberg 3 that ioo
of protein might yield51.4 grams
of fat,they computed
grams
that all the fat produced by the animal in these experiments
1

Arch.

51 (1892),229.
Physiol.(Pfliiger),

2Ztschr. Biol.,9 (1873),435.

Landw.

Vers. Stat.,10

(1868),455.

176

OF

NUTRITION

FARM

ANIMALS

the case
it came
to be more
as
particularly
clearlyrecognizedthat Henneberg's estimate of a production
in all probof proteinwas
of fat from 100 grams
of 51.4 grams
ability
after it was
shown
that what
too high,and especially
had been regardedas digestedproteinin many
of these experiments
N
X
consisted
in
of
much
6.25)
(i.e.,
digestible
part
of
simpler nitrogenous compounds. The
ready formation
fat by the hog rendered this animal
suitable subjectfor
a very
this
on
experiment,and the great majority of investigations
animal
have
supported the view that fat is produced from
but similar results upon other species
have not
carbohydrates,
been lacking,
while respirationexperiments upon
swine, geese,
t
he
and
extensive
especially
dogs,
investigations
by G. Kiihn l
cattle have completed the demonstration.2
upon
In the lightof all these results,
of fat from
the formation
admitted,
carbohydratesin the animal body is now
universally
from
its
while
production
proteinis still questionedby a few
and in any case
is of little economic
that we
so
significance,
have come
back by a curious reversal of views almost to Liebig'sclassification of the nutrients into plasticand respiratory.
This conclusion appliesspecifically
hexose carbohydrates,
to the pure
In many
starch.
of the experiments
particularly
cited,however, the non-nitrogenousmaterial digestedby the
the feed.

animal

This

was

consisted

to

of uncertain
crude

fiber

present any
it may

and

suffice

as

chemical

nature

included

nitrogen-freeextract.
to

say

here

that both of

of those

extent

in

the

Postponing

discussion of the nutritive value

show
particular
serve

inconsiderable

not

stances
subterms

for

the

of these groups,

3
in
investigations
the pentosans, may
them, including

that

Kellner's

of fat.

sources

Katabolism
250.

Body fat

aside from

reserve.

"

The

its mechanical

of

Kellner

Compare

Landw.

Landw.

Vers.

the author's
Vers.

Stat. ; 44

in the

body.

observation

the

shows

Nutrition,

great

In the lack of

that this

(1894),257.

Principles of Animal

Stat.;51 (1900).

sues,
adiposetis-

stored fat of the

functions,constitutes

material
energy-yielding
adequate feed supply,common

reserve
an

pp.

165-184.

METABOLISM

is drawn

reserve

as

77

for the support of the internal activities

of energy for the performanceof
source

upon

body and

of the

external work.
of

Mobilization

251.

fat.

reserve

In order

"

that the stored

of the body it
be used for the general metabolism
fat may
first be transferred from the adipose tissue cells to the
must

it is needed.

localitieswhere

by its reconversion into

"

Presumably this is accomplished

soluble fat

the walls of the cells into the

"

and

its passage

blood, that is,by

through

reversal of

laid down.
Since the transfer of
the process by which it was
fat through the epithelial
cells in resorptionis effected by a

hydrolyticcleavage (152),one is tempted to,imagine a similar

reversible enzymatic process in this case.
Direct evidence of
this is lacking,
but apparentlysuch a cleavagetakes place somewhere
at an
sulting
earlystage in the katabolism of the fats,the reof dextrose.
From
glycerol
perhaps servingas a source
that point on the katabolism is that of the fattyacids.
252.

at the

Oxidation

p carbon

atom.

"

The

oxidation of the

unsaturated, to carbon dioxid

and water, like the other katabolic processes alreadyconsidered,
The researches of Knoop, Embden,
is a step by step process.
1 have
rendered it highlyprobable,if not cerDakin and others
tain,
of the normal satuat least in the case
that the oxidation,
rated
at the second
(i.e.,
acids,begins at the /3 carbon atom

fattyacids,either

saturated

carbon

the COOH

from

atom

off of two

carbon

atoms

and

dioxid, water

at

or

group)and
a

time.

The

results in the

products

fatty acid containing two

splitting

are

carbon

less carbon

and with which the same

one
original
process
of erosion is repeated.
in
If it be true that the fattyacids thus undergo katabolism
the body by stages of two carbon atoms
each, and particularly
be built up
if it may
be regarded as probablethat they may
from
again in a similar manner
simpleratomic chains, there
is afforded a plausible
explanationof the rather strikingfact
that nearlyall of these compounds found in the animal body
than

atoms

contain
This
lower

an

the

even

scheme

number
does

acids of the

I7-47N

atoms.

providefor the oxidation of the three

acetic and formic, and in
series,
propionic,
not

Dakin, Oxidations

Compare,

of carbon

and

Reductions

in the Animal

Body,

191 2, pp.

178

NUTRITION

fact,while these acids

body,
253.

the chemical

Formation

FARM

OF

known

are

mechanism

of

ANIMALS

to

be

freelyoxidized

of the process

in the

is little understood.

In

discussing
the probability
of the formation
of carbohydratesfrom proteins
it
often
that
their
out
be
(235), was pointed
originmight
It was
ascribed to either proteinsor fats or both.
there shown
that in many
the probabilities
tion
cases
stronglyfavored a formafrom proteins. In other instances,however, the probabilities
that
fats giverise to carbohydrates.
seem
equallystrong
In particular,
experiments upon phloridzindiabetes of the
than could be
dog have shown the productionof more
sugar
formed
from the quantity of protein katabolized
during the
same
time, while the stock of glycogen in the animals experimented
and muscular
had been so exhausted by fasting
work
on
that it seems
wise
scarcelypossibleto interpretthe results otherthan as showing the formation of sugar from fat. It should
be added, however, that it has been seriously
questionedwhether
the conditions of the experiments were
controlled
sufficiently
carbohydrates

from

fats.

"

'

to

the conclusions

warrant

The
have
also
from

low values for the

respiratory
quotient(296)which
been reported in some
for hibernating
animals have
cases
been interpreted
of carbohydrates
as indicating
a production
very

fat.
be

In
an

of fat into sugar, there must

viously
obevolution
absorptionof oxygen with no corresponding

the conversion

of carbon
the

drawn.

dioxid,the tendency of which

respiratory
quotient. The

oxidation

of

fat

low
as
figures
fastinganimal
demonstrate

In
is 0.7.
0.3 have been

as

increased.

be

to

lower

value of the latter for the direct

hibernating animals, however,

while the weight of the
reported,

While

the formation

would

facts,of course, do not

from fat, they are
quite

these

of sugar

and
to indicate a
seem
interpretation
ing
The more
recent
experimentson hibernatstorage of oxygen.
animals,however, have failed to give such low quotientsas
obtained by earlier observers.
were

compatible with

"
254.
are

7. Metabolism

Certain

found

that

chemical

wholly or

of

elements

Ash

Ingredients

of the

body

in part in their ash when

and

of the feed

these materials

METABOLISM

79

though,
spoken of as ash ingredients,alas
already pointed out (3,5), this does not necessarily
"
inorganic
imply that they existed in the originalmaterial in
and

burned

are

therefore

are

"

elements

of these

Most

combination.

are

as

essential

to

the

gen,
carbon, nitroorganiccompounds,"
hydrogen and oxygen of the so-called
although unfortunately the laws regulating their metabolism
ments
less extensively studied.
these elehave been
much
Among
sulphur and phosphorus are of specialimportance in
vital processes

the

as

abundant

more

elements
"

this connection.

Sulphur
contain
small
feeding stuffs may
of sulphur in the form of sulphates,by far the greater
amounts
part of this element in the feed of animals exists in organic
Such, for instance,are the allylsulphid(CsHs^S,
compounds.
of the genus allium,and
contained in garlicand other members
the allyl
sulphocyanat,C3H5
CNS, found in mustard and other
of the cruciferse.
riers
Ordinarily,however, the chief cargenera
in feeding stuffs and
of organic sulphur, both
animals,
255.

Sources.

While

"

"

the

are

contain

which
proteins,

acid

di-amino

the element

in the form

of the

cystin (47).

question whether the animal body

from
build up its sulphur compounds
can
inorganic sulphur
been investigated.
to have
does not appear
256.

Katabolism.

The

katabolism

"

of

The

the

cystin component

of

proteinspresumably

that of the other

as
general course
the proteins by hyamino
acids, i.e.,it is split off from
of the
One
drolyticcleavage and subsequently deaminized.
of cystin appears
to be
taurin,
products of the katabolism
CH2
CH2
NH2
SO3H, contained in the taurocholic acid of the
the latter comthe extent, therefore,to which
bile. To
pound
in the lower intestine,
it carries small
escapes resorption
of sulphur into the feces. Both
amounts
cystin and taurin,
however, are readily oxidized in the body, the largerpart of
their sulphur taking ultimatelythe form of sulphuricacid and
being excreted in the urine. The sulphuric acid of the urine
follows

"

from
and

the

same

"

"

exists in

the

combination

in

part with

aromatic

radicles derived

putrefactionof the proteins in the lower

in part

with

bases.

In

human

urine

about

intestine

one-fifth

of

l8o

NUTRITION

"

the

FARM

sulphur exists in a
neutral sulphur,the

total

known

OF

as

ANIMALS

less

completely oxidized form

and originof which
is
nature

obscure.

Phosphorus
The

phosphorussupply of

the

body
in the four forms indicated in Chapter
is received substantially
I (5),viz.,as phosphates,as
phosphatids,as phospho- and
and as phytin. Of these the various
nucleo-proteins
organic
forms usuallypredominate.
It appears
probable,however, that all these various forms of
phosphorus are resorbed into the blood stream in the form of
phosphoric acid. Of the phosphates ingestedas such this is
for believing
There seems
that the
certainlytrue.
good reason
phosphoric acid radicle contained in the nucleic acid of the
nucleoproteinsis quite completely splitoff by the digestive
and reaches the blood as phosphoric acid (139),
and
enzyms
the same
thing is presumably true of the phosphoproteins.
The phosphatidsare probably acted on by the lipasesof the
acid resulting
digestive
tract,but whether the glycerophosphoric
from their cleavageis further split
The ready
up is unknown.
cleavageof phytin in seeds would suggest that probably its
phosphorus also is resorbed as phosphoricacid.
258. Anabolism
The animal body contains
and katabolism.
a largestore of phosphorus in the
inorganic form, especially
257.

Forms

ingested.

"

"

"

"

"

in the skeleton.
the resorbed
The

body

For

the maintenance

phosphoricacid
also

"

or

increase

of this store

is

naturallyavailable.
pounds,
contains,however, organicphosphorus com-

which, although less

in

amount

than

the

inorganic,

of the

lief
for the vital functions. The behighestsignificance
that the phosphorus supply of the body is resorbed chiefly
in the form
of phosphoric acid necessarily
implies,therefore,
that the organism is able to utilize inorganic
phosphorusfor the
synthesisof nucleic acids,phosphoproteins,
phosphatids,etc.,
are

and

the

belief
power

experimental

(497).
of

the

In

evidence

this respect,as

organism

appears

strongly in favor of this

in many
others,the synthetic
to be greater than was
long
is

supposed.
Little is known
in the

regardingthe

katabolism

of the

by the phosphorus
phosphatids,
nucleoproteins,

course

followed

181

METABOLISM

Ultimately it takes the form of phosphoricacid and is

mediate
interexcreted in the feces or urine (199),but whether
any
compounds are formed is not known.
etc.

Other elements

important
than the two justconsidered,littleis known
regardingtheir
in the ordinary sense, i.e.,
of the chemical changes
katabolism
exist in
which
they may
they undergo in the body. That
feedingstuffs in organicas well as in inorganicforms is probable.
in the animal body is
That they enter into organiccombination
likewise

to

be assumed

iodin of the

and

of
to

serves

the

only

haemoglobin
and

sodium

Both

"

animals

few

and

the

potassium
in

considerable

extent

as

inconsiderable

not

consume

and

condiment

is often

is less than

this

probablethat

salt,althoughit appears

common

actuallynecessary

amount

in

ordinary foods and feeding stuffs and

farm

and

man

amounts

of

iron

less

no

positivelyknown

potassium.

in the

contained

addition

are
ingredients

thyroidglands.

Sodium

259.

is

but

of the

like that

instances

are

so-called ash

the other

While

that

the
Bab-

supposed.

without access
for over
a year
cock, e.g., was able to keep cows
to salt,
except that contained in their feed,without any obvious
illconsequences.

potassium and

Both

with

sodium,

well

as

of

the latter in the form

as

salt,are

the

chlorin

excreted

bined
com-

in the

urine.
Calcium

260.

magnesium.

"

maintenance

in the soft tissues

in

in

elements,calcium

These

especiallyimportant

particular,are

growth and

and

their relations

to

the

but they are not lacking

skeleton,
they perform important functions.

of the

also,where

As noted
little is known.
intermediarymetabolism
in Chapter IV (199),the normal
path of excretion of calcium,
testine,
and
of magnesium, is through the lower into some
extent
is
of these elements
that the apparent digestibility
so

Of

no

their

measure

body

of the

amount

actuallyresorbed and utilized

in the

processes.

long controversy has been carried on

be resorbed and
the question whether
inorganiciron may
261.

whether

Iron.

it can

"

be utilized for the

synthesisof

the

over

if

so

haemoglobins

l82

NUTRITION

OF

and

for other purposes

in the

ANIMALS

FARM

Both

body.

however,
questions,

is
regarded as settled in the affirmative. The matter
of importance in its relations to the use of iron in medicine but
in stock feeding.
is of no special
significance
The
excretion of iron takes place almost
exclusively
through
may

be

this fact led to

the intestines and

inorganiciron

262.

General

metabolism

Functions

8.

scheme

found

Fig.

in each

23.

"

of

the

of metabolism.

of the several

paragraphs,it was

that

be resorbed.

cannot

"

the earlier conclusion

classes

Nutrients
"

In

of nutrients

that the main

Diagrammatic scheme

consideringthe
in the foregoing

features of the process

of metabolism.

might be conceived of in accordance with the ideas

of a complex of reversible
suggested in " 2 (210-212)as consisting
reactions accelerated or retarded by intracellular enzyms.
By combining the equationsused to represent those reactions,it
lowing
possibleto take a further step and formulate the folappears
which
scheme for the total metabolism
highlygeneralized
may

case

serve

to

show

the

interrelations between

the

metabolism

84

FARM

OF

NUTRITION

ANIMALS

the radiant energy of the sun

of their various constituents,
to be

plantstransform
energy

into the chemical

yieldedup by

organism through the processes

conceptionhas become a familiar one

of metabolism.

latter to the animal

This

emphasis has
Second,

laid upon

been

the

and

much

it in recent

the feed suppliesthe

years.
materials
specific

requiredfor

of the
buildingand maintaining all the complex structures
it is the source
i.e.,
functioning,
body and for their harmonious
of structural and repairmaterial.
which make
fats and mineral ingredients
That the proteins,
of the body (99,280)
up by far the largerpart of the dry matter
stration,
derived ultimatelyfrom the feed needs no specialdemonare
substances present in the
but the importanceof many
For
tends to be overlooked.
amounts
body in only minute
example,the enzyms of the body, both extra- and intra-cellular,
form no considerable portionof its mass,
yet they are essential
to

its vital activities.

So, too,

the

various

hormones

and

cretions
se-

glands,while ignoredin the broad scheme

essential to the vital processes.
are
justpresented,
feed must
supply material for the production of

of the ductless
of metabolism

Clearly,the
these and

other similar substances.

material will
energy-yielding
substances
suffice to support lifein the absence of those specific
in order that the machinery of conversion
which are necessary
of coal under the
shall operate properly,much
amount
as
no
In

other

words,

boiler will enable

no

electric

an

of

amount

plant to furnish

normal

amount

of the generator is defective.

example,if tryptophan is necessary for the formation of

of

if the insulation

current

internal

essential

secretion,a

diet

lacking that

For
some

substance,

might furnish,would fail to support the

organismpermanently unless the body can manufacture
phan
tryptohowever

much

from

energy

it

other substances.

The animal
is a very importantone.
qualification
in its feed
the presence
is very far from being dependent upon
of all the varied chemical compounds requiredfor its operation.
is the case.
As has appeared in preIndeed, quite the reverse
vious
t
he
actual
resorbed
substances
are
sections,
comparatively
The

latter

simple and uniform and

a great varietyof chemical
What

is necessary

upon

them

the animal

body

executes

and synthetic.
changes,both analytic

is that the resorbed

feed shall include

sub-

185

METABOLISM

which

the

body

can

manufacture

the

compounds

it requires.
of

Functions

264.

the

once

of which

out

stances

the

familiar

most

The

proteins furnish at
strikingexample of this

proteins.
"

and

the most

of the feed.

dual function
Since the

proteinsmay be katabolized in the body with the

of products (carbon dioxid,water, urea,
etc.) containing

formation

either
that found

available

no

energy

but

or

fraction of

small

in the

originalproteins,it is clear that the latter

carriers of energy.
In fact,it has been shown
as
to be
serve
animal in normal
possibleto maintain a carnivorous
activity
for an indefinite time on a diet containing
nothing
substantially
but

proteinas

But

of energy.

source

also

proteins serve

water, the

buildingmaterial. Aside from

working machinery of the body is composed largelyof
as

at least of the specialsubstances

proteins,while very many
are
already mentioned
nitrogenousand probably derived from
the proteins. These
protein tissues and other substances must
be built up in the growing animal and maintained
in the mature
only proteinsor their cleavage products
one, and for this purpose
a
nd
be
in the feed is indistheir presence
pensable.
can
utilized,

point which sometimes

portionof protein may not
A

but

as

also

causes

only

perplexityis
serve

as

structural

for the vital processes,

yield energy

that the

so

same

material

that in estimating

the energy suppliedby a feedingstuff that of its protein

is included.
well as that of its other ingredients
The difficulty
when

disappears,however,
portion of proteindoes
time.

If

it is remembered

perform both

not

that

any

these functions

given
at

the

of proteinin the feed of a mature

animal
gram
is used for structural purposes
it practically
takes the place of
of tissue protein,while the latter is katabolized
an
equal amount
same

and

the
yields substantially

have

been

available

been katabolized
has been
some

from

instead.

may

as

its store

of energy,
the katabolic process but at

itself be

protein and katabolized

serve

of energy
would
as
of feed protein had that

amount

the gram
The latter,
with

temporarily set aside from

later time

amount

same

The
of energy.
illustration.
an

in

replacedby another gram of feed

its turn, liberating
the corresponding

repairingof
The

old wood

wooden
taken

out

building may
to

make

way

86

NUTRITION

well as
as
material,
immediately required, may

for

FARM

OF

new

ANIMALS

wood over
that
surplusof new
used
fuel for
as
indifferently

any
be

of the young
animal, in which
plex
comproteinis permanentlyset aside for growth,is a triflemore

warming the building.The

the
substantially

but

Functions

265.

of fats.

"

case

considerations

same

In

the

case

is the

hold

good.

of the fats the energyFats

and obvious one.

predominant
form
of fuel,containingmuch
concentrated
more
are
a
energy
unit
of
other
nutrients.
than
the
They supply much
any
per
well adapted for the
in a small bulk and are, therefore,
energy
storage of reserve
energy in the body.
The fats and closelyrelated bodies (the lipoids),
however,
also important and
are
apparently essential constituents of
important
protoplasm (75). The
lipoids,therefore, have
structural functions and an
adequate supply of them in the
interest
body is indispensable.From this point of view, some
of investigators
attaches to the results obtained by a number
that a certain minimum
who
claim to have shown
supply of
for growing animals.
especially
lipoidsin the feed is essential,
The evidence,however, is negativeevidence,i. e., experimental
diet.
In view
animals failed to grow normally on a lipoid-free
of the positive
results obtained by Osborne and Mendel,1 as well
of the fact that both the simplefats and the phosphatids,
at
as
least,can be synthesized
freelyin the organism,and takinginto
consideration
of the body in
the extensive syntheticpower
general,it is difficult to believe that the presence of lipoidsin
and more
have
recent
the feed is indispensable,
investigations
of the observed facts (498) On
afforded a different explanation
bearing function

the other

also that the fats appear,

the productionof milk fat.

growth
favor
266.
more

While
the

that the lecithins stimulate

hand, it has been shown

and

Functions

of

other

hand,

be

consumed

certain

limits,to

The

carbohydrateseven
carriers of energy.
serve
as
chiefly
per unit than fats,they can, on
in largerquantities
and they

carbohydrates.

than the fats

distinctly
containingless energy

within

"

of the energy
in the diet of
the presence of carbohydrates

practically
supply the greater part
man

and of farm

(dextrose)in the blood

be

on
chiefly

While

animals.

of

account
1

this appears
to
lymph is essential,
their ready availability
as fuel material.

and

Jour. Biol. Chem.,

12

(1912),81.

METABOLISM

The

carbohydratesseem,

in relation
to

fastingor

to

however,

the katabolism

have

to

of fats.

function
specific

the

When

87

is

body

pelled
com-

draw

its energy
supply chieflyfrom the fats,as in
in diabetes (inwhich the power
of katabolizing
carbohydrates
is

lost),or

diet,the katabolism

when

of the fats fails

of

amounts

katabolic

carbohydratesare
be

to

beta-oxybutyricacid

product

aceton

from

complete
well

as

excreted

are

absent

and

as

the
siderable
con-

normal
the ab-

unoxidized

in the

urine.
267.

Non-nitrogenous
that both

appears

the fats
in

specialpurposes

serve

nutrients

that their chief function

ordinary rations
functions

they

as

are

carriers

in the

is to

of

general.

While

"

it thus

and carbohydrates may

(or lipoids)
the body, it is,nevertheless,clear
Their

supply energy.

abundant

so

in

energy

that
any

amounts

in

compared with their

which
specificpurposes
as

amply provided for. As related to

of farm
in particular,it is of special
the nutrition
animals
interest to note
that not only the fats and carbohydrates digested
such but also the products of the bacterial fermentation
as
serve

body

of the insoluble

are

carbohydratesare

available

as

sources

of

energy.
268.

Functions

of ash

ingredients.
"

While

the

nous
non-nitroge-

chieflyas carriers of energy

organicnutrients serve
only in a minor degree to provide the compounds necessary

and
for

the

performance of specificbodily functions,the so-called ash

in this respect. They
ingredientsrepresent the other extreme
available energy into the organism but,
introduce practically
no
the other hand, they are not only essential structural comon
ponents
of the body tissues but likewise supply and maintain
certain

conditions

indispensableto

the

performance

of

the

bodilyfunctions.
The

structural

importance of the ash ingredientsis most

of the skeleton,which, in the higher animanifest in the case
mals,
of calcium
and
contains relatively
large amounts
phoric
phosand
acid and
small
quantitiesof magnesium, sodium
carbonic acid (81) which
chanical
meimpart to it certain necessary
qualitiesof strength and rigidity. The necessityfor
in that of
a
supply of these substances in the feed, especially
growing animals, is too obvious to require discussion. The
ash ingredients,
however, have other equallyimportant functions

88

NUTRITION

providingthe

in

OF

conditions for the chemical

necessary

activities of the various

Osmotic

269.

pressure.

membranes.
in the

lymph,

order

In

protoplasm of

various

tissues

lymph which constitutes their

(185)and from which they are
of semipermeable
partake of the nature
maintain

to

normal

tions
condi-

the cells the osmotic

therefore that of the blood

and

cal
physi-

the

cell walls which

separatedby

cells of the

The

"

nutritive environment

immediate

and

tissues.

from

their nourishment

draw

ANIMALS

FARM

pressure of the
which it is derived,

from

maintained

be

The
osmotic
approximately constant.
of the blood is stated to be approximately about
8
pressure
contained in
atmospheres, due largelyto the ash ingredients
solution.
With
an
tration
adequate supply in the feed the concenof mineral matter
in the blood is regulatedchiefly
by
of
the excretory activityof the kidneys. Thus, in the case
sodium
chlorid,for example, it is estimated that the blood of
contains approximately30 grams
of this substance,
an
man
average
of which
hardly half a gram is excreted daily when
must

is consumed.

none

is

excess

promptly

four hours.

What

other diffusible
270.

in the

Ionic

If,however, salt is added

excreted
is

true

concentration.

course

of salt in this

of
ingredients
"

diet,the

of the next

twenty-

respect is

also of

true

the blood.
The

variqus salts

are

contained

body largelyin
however,

In such soludilute aqueous

solution.
tions,
it is believed that salts are largelydissociated

ions, a dilute solution of

into their constituent

for

in the

the

to

example,containingin

addition

salt,

common

unchanged NaCl
of calcium sulphatethe ions Ca and
the ions Na and CI, one
SO4, etc. Acids are similarlydissociated,
yieldinghydrogen
alkalies
ions
ions
+ SO4), while
(H2SO4 ^H
yield OH
of these ions have been shown
to
Some
(KOH ^ K + OH)
effects on certain cellular activities.
For example,
have specific
bility
a frogmuscle
kept in 0.7 per cent NaCl solution retains its irritafor one
two
or
days. In a solution of a non-electrolyte,
to

some

like sugar,

muscle

asparagin,etc.,having the

soon

solution

loses its

same

but
irritability,

other

it

apparent that

hand,

Na

the

effect is due

ions alone

cause

to

of other
of other
the

pressure,

be added

if NaCl

regains it. Since a number

while chlorids
effect,
produce the same
it is

osmotic

Na

long continued

to

sodium
metals
ions.

the
the
salts

do not,
On the

rhythmic con-

189

METABOLISM

muscles,which, however, is suspended by the presence

ions
like
of
Ca
certain
solution
in the
or
(not all)dyad
Mg.
actions
of
ions
other examples of such antagonistic
Numerous
known, such as those observed by Loeb, for example, in
are
be said that cell
In generalit may
the development of the egg.
suitable
other things upon
a
activities are
dependent among
of various
elements in their surroundings,
ionic concentration
fact that the so-called physiand it is a strikingand interesting
ological
tionally
be kept funcsalt solutions in which livingorgans may
active for a longer or shorter time contain the various
traction of

approximately the

salts in

same

proportionsas

found

are

in

water.

sea

example of the influence of ionic concentration is

of the digestive enzyms.
afforded in the case
Ptyalin, for
of hydrogen ions.
example, is sensitive to a very slightexcess
of
active in the presence
Pepsin, on the other hand, is most
hydrogen ions, while trypsinacts best in the presence of an
Another

"

foregoingtopic and constitutingindeed

of

is that of the maintenance

fluid

in

is neutral
of H

excess

no

Closely connected

neutrality.

of

Maintenance

271.

the

ions.

of OH

excess

nor

the

chemical

of OH

ions,an

equivalentto acidityand
It has

been

shown

neutralityin

that

an

specialcase of it,
the body fluids. A
a

it

when

sense

the blood

serum,

contains

being
alkalinity.

of the former

excess

of the latter to

excess

with

as

of
representative

body fluids,is very nearly neutral,its content of H and

ions being approximately 0.4 X io-7 and
OH
7.2 X io-7,
equivalentto about 0.000012
i.e.,it has an alkalinity
gram
the

NaOH
The

per

liter.1

body

katabolism

is

continuallyproducing acids,especially

carbonic,phosphoric and sulphuricacids (256,259),which

These
acids are in
tend to increase the acidityof the blood.
produced in the katabolism of
part neutralized by the ammonia
of
protein (233),but it has been shown by the investigations
L. J. Henderson
that the salts of the blood serum,
especially
the sodium
phosphates and bicarbonates,play an important
part in maintainingits neutrality. They are present in such
1

Blood

is

commonly

said to be alkaline because

litmus.

ordinary indicators,such as
of the true alkalinityor
measure

Such

acidity.

it gives

an

alkaline reaction to

reaction, however, gives

no

definite

NUTRITION

190

OF

ANIMALS

FARM

proportionsthat their solution possesses nearly the maximum

while they also,
capacity for the preservationof neutrality,
of elimination of
the phosphates,serve
a means
as
particularly
of acid through the kidneys in the form of the acid
an
excess
phosphatesof the urine.
The three generalfunctions
272.
Other
functions
of ash.
exhaust the listof offices performed
justenumerated by no means
gredien
by the ash ingredients.Iron, for example, is an essential inof the red blood
of haemoglobin,the coloringmatter
is distributed
which is the vehicle by which
corpuscles,
oxygen
throughout the body (191). Although contained in the body
in relatively
minute
of
this element is,therefore,
one
amounts,
prime necessity.Iodin appears to be an essential ingredient
of the thyroidglands,and although we
are
ignorant of its
functions it is known
that the absence of these glands,or
exact
their failure to function, gives rise to serious disturbances
cate
indito
seem
(goitre,myxcedema). Recent
investigations
and
that manganese
boron, and perhaps other elements
heretofore
not
have
regarded as essential,may
important
in
functions
as
catalysts plants and perhaps, therefore,in
animals
also,although this is at present a conjecture. It is
likewise possiblethat other elements present in small amounts
later be shown
functions.
to have physiological
may
tends to make
Its very familiarity
273. Functions
of water.
"

"

us

overlook

the

strikingnature

it is impossiblein the absence

of the fact that

of

If

water.

life as

we

know

protoplasm may

be

regarded as a collodial solution,one may almost say that life

is possible
only in aqueous solutions.
Some
for this are fairly
obvious.
The phenomena of
reasons
and
osmotic
ionization,for example, whose
tance
imporpressure
has just been indicated,are
solution phenomena.
substantially
It is possible
fundamental
also that there are more
for this striking
fact.
reasons
Certainlythe largershare of our
present chemical knowledge relates to the chemistry of either
solutions or gases, two
states
resemblingeach other
aqueous
in many
most

respects and

readily,if indeed

Moreover, it has been

which

water

dependent

upon

is not

in which

it

chemical
takes

ever

shown

that

commonly

the presence

action

place in

seems

to

occur

the solid state.

reactions,at least,in
regarded as concerned, are
some

"of minute

amounts

of this sub-

CHAPTER

THE

BALANCE

"
The

274.

power

which

may
c
lose
fairly

what
a

Conception

prime

motor.

"

The

livinganimal

prime motor
; that is,it generates
for its own
operationand is able to produce a surplus
be applied to do external
In particular,
work.
a
analogy may be drawn between the animal body and

known

are

is known

what

constitutes

as

NUTRITION

OF

General

i.

animal

VI

as

internal combustion

as

In such motors,

motors.

alcohol,etc.)is burned in
(gas,gasoline,
engineitselfand its available chemical energy

fuel

the

in part into
similar manner

by

the

and

motion
the

processes

work

noted

cylinderof

is transformed
In

somewhat

cells of the

body

circulation

are

the oxygen
introduced
through the
for the various activities of the organism.

with

lungs,and yieldenergy
be

part into heat.

compounds suppliedto the

of digestion,
resorptionand

katabolized,combine
It should

in

the

that these activities include

ternal
merely exdone by the animal but likewise a varietyof internal
that of circulation,
as
digestion,
respiration,
resorption,

work, such
secretion,etc.

In

always in operation,even

other
when

not

words, the animal

performing no
When

in

machine

is

external work.

chanical
operation,a me(a gasolineengine,for example) consumes
prime motor
two
things. First,the material of which the working parts are
that ultimatelyrepairs
so
composed is graduallyworn
away
in amount
and second, fuel is consumed
are
ing
dependnecessary,
the work done.
the same
Substantially
thing is true
upon
of the animal body.
be regarded as
The
working machinery of the body may
of water, ash and
chinery,
composed essentially
protein. This malike that of the engine,is continually
wearingout ; that
katabolized and
is being continually
is,the proteinin particular

275.

Expenditure by the body.

IQ2

"

THE

BALANCE

OF

NUTRITION

193

ties
products of its oxidation excreted. In addition,the activiof the body, like those of the engine,require a supply of
fuel material
containing available chemical energy equivalent
For this purpose
the body utilizes
to be done.
to the work

the

in the first instance

and

tissues.

of

the

As

body

"

the

substances

contained

in its

Chapter V, all the organicingredients

undergo
protein, fat and carbohydrates

shown

in

"

and
katabolism, giving rise to carbon dioxid,water
simple nitrogenous products,accompanied by
into other forms.
energy
storehouse
of chemical energy as

of their chemical

words, the body

cells

own

is

tively
comparaformation
trans-

In

other

well

as

is contained

of the

body
particularlyin its fat,and to a minor degree in its glycogen,
while the body protein,although it likewise yieldsenergy when
enous
katabolized,especiallythrough the oxidation of its non-nitrogresidue (229),usually plays a small part quantitatively.
The
The fat of the body constitutes its great reserve
of energy.
be compared, for the
material in the body may
store of reserve
sake of illustration,
to the gasolinein the tank of an
automobile,
with the difference,
or
however, that the body derives more
mechanism.

This

stored-upenergy

of the material

less energy
from the combustion
the engineitself.
276.

The

feed.

Neither

"

long depend entirelyupon

disaster.

Sooner

or

the automobile
its

later it must

own

stock
obtain

nor

(protein)of

the animal

of material

can

without

side.
suppliesfrom the outare
obviously of two

suppliesrequiredin both cases

classes,
corresponding to the two classes of materials consumed
be brieflydesignated
in the operation of the machine, and may
as
repairmaterial and fuel.
In the automobile, parts of the machinery, the tires,
etc., as
of the same
be replacedby new
kind,
out must
ones
they wear
The

ing
gasolinetank must be filledat intervals and the workof the animal
The case
be suitablylubricated.
parts must
be suppliedin
is preciselysimilar. In the first place,it must
its feed with materials from which, by the processes of digestion

while the

atomic groupings
it can
the particular
secure
resorption,
etc.)which will exactly
(amino acids,peptids,ash ingredients,

and

protoplasm and replacethose eliminated by the vital

its
also derive from
activities. In the second place it must
feed molecules which it may, accordingto circumstances,break

fitinto its

NUTRITION

194

at
(katabolize)

down

as

up
to

the

carry

such

analogy
the

build

and

sets

In

do

We

with

broad
not

generalway, of

need

material.

course,

do work

to

Similarly,we

of

maintenance

is needed

than

namely,
various

the

in
mere

the

as

functions

2.

body

is evident

exhibits

two

prime
nutrition,tendingto
tained
body gains,is mainas

these

between

this fact is perfectly

obvious.
that

us

the horse

or

cow

to

the fact that those

go on in a state of so-called rest

their support, so that the mere

calls for

an

expenditureof feed.
nutrition is something

study of
general knowledge of
or

It

"

the balance

scientific

feedingstuffs

"

of

osmotic

yieldmilk unless supplied

good the resultingloss of body

of the

quantitative measure

the nutritive

278.

animal

an

of

familiar with

are

operationsof the body which

for
likewise requirematerial

more

its feed

ingredients

its actions

the

teach
or

make

to

in

Whether

to
physiologist

the

those

falls away
depends upon
of activities.

sufficient feed

What

from

several ash

that

concerned

it.

increase

long continue

cannot

the

expenditure.

foregoingconsiderations

up
or

two

and

income

tendingto destroyit,and

motor,

obtain

must

Finally,

working conditions of
and the like,somewhat

necessary

those
activities,

of

sets

of

Balance

the

from

step further,it

store

or

be lubricated.

enginemust
277.

of their energy
(fat,glycogen)for future use.

ionic concentration

pressure,

ANIMALS

for the sake

proportionsof

and

amounts

will maintain

as

once

of energy

reserve

FARM

OF

facts ;
which
the

these familiar
extent

to

their

of the

Methods

contribute
singleingredients
body under varyingconditions.

Investigation

of

Investigation of details

to

of

metabolism.

attackingthe problem just stated

"

One

method

by investigatingthe
details of the metabolic
In the study of metabolism
processes.
(includingthe chemical changes in digestionand resorption)
of the feed
to follow the various
the attempt is made
ingredients
through the body and to trace in detail how, where and to what
extent
or
they contribute to the maintenance
growth of tissue
or
supply energy for the use of the organism. Such studies are
of fundamental
importance. They reveal to us how the animal
mechanism

operates.

When

carried

is

to

their ultimate

con-

THE

NUTRITION

OF

BALANCE

95

complete knowledge of
the chemical ingredientsfound in feeding stuffs,they will make
of nutrition as a physicoaccount
it possible
to give an exhaustive
zation
to say that the realiIt is hardly necessary
chemical process.
when

elusion,and

accompanied by

of this ideal lies in the distant

279.

effect.

nutritive

Total

future.
students

Meanwhile,

"

of

stock

different aspect
plex
of the subject,
viz.,in the aggregate effect of the varied and com-

feeding are

interested

metabolic
the stock of

any

due

in

processes

somewhat

energy

in the

increasing

body.

Is the

given regimen maintaining itself and making

is the animal
doing work or yieldingmilk or

growth, or
other productsat the expense
a questionof balance.
to

reducing,maintainingor

and of chemical

matter

body under

primarily in

of its

own

This

tissues?
of the

Is the income

dently
is evi-

body equal

its outgo?
280.

The

schematic

dependent upon

body.

balance

"

between

The

idea

of the

constructive

and

organism

as

destructive

of the conception
by means
specific
of the schematic body,which
regards the body of the animal,
of ash, protein and
aside from water, as consistingessentially
for
of glycogen so small that it may
fat,togetherwith an amount

activities may

be made

more

be neglected.
purposes
many
for this conceptionis found
The justification

in the data

tained
con-

Chapter II," 3, regardingthe composition of the animal

there
It will be recalled that in the investigations
as
a whole.
determined
directly,the
recorded the water, ash and fat were
of these and the total weight of the
the sum
difference between
of fat- and ash-free dry
animal, of course, showing the amount
the total nitrogen contained
in which
In those cases
matter.
also determined, it appeared (99) that, with
in the body was
one
exception,the percentage of nitrogenin this fat- and ash-free
closelyapproximated that in the animal proteins.
dry matter
of glycogen and other substances
In other words, the amount
is so small as to be
included in the fat- and ash-free dry matter
in

be considered
to consist essentially
negligibleand the latter may
of protein.
this point of view, it is evident that the effect of any
From
feedingstuff or ration in causing a gain or preventing a loss of
tive
ash, proteinand fat (and glycogen)shows its aggregate nutribe
of the body may
effect. Or, since the organic matter

196

NUTRITION

looked

OF

ANIMALS

in the lightof stored energy, a still simpler expression

upon
of the nutritive effect may
be obtained by determining

the effect of the feed upon

energy in the body.1

Experiments directed
of

FARM

and

the store

to

of

proteinand

the determination

of chemical

of the

gain or

loss

of energy

by the body have been of two general

kinds, viz.,comparative slaughtertests and what are called
balance experiments. Both have played an important role in
the study of nutrition.
matter

Live

281.

weight

as

measure

of nutritive

effect.

"

the

At

questionarises whether the simple

of weighing an experimentalanimal is not
and obvious method
the aggregate effect of a ration,
sufficient to determine
without
for any elaborate experimentaldevices.
the necessity
The answer
the object
to this questiondepends largely
upon
undertaken
of the experiment. If it be one
to answer
a
mercial
comthe increase in live weightduringa considerable
question,
period,when determined with the necessary precautions,
may
results
of
be entirely
the
obtained.
If,
adequate as a measure
for example, the question under investigation
is the relative
of two methods
of fattening
cattle,the gainsmade by a
profits
considerable number
of animals, togetherwith the judgment of
the market
regardingthe qualityof the finished animals, will
is to be preferred.The
determine
which method
substantially
would not only be
of more
elaborate experimentalmethods
use
needless refinement
but might actuallyinterfere with the
a
settlement
of the economic
So, too, in the
questioninvolved.
stock or in milk production,the general
handling of young
and condition of the animals,togetherwith the gain
appearance
in live weight or the yieldof milk, furnishes a sufficiently
curate
acindication of the practical
results obtained, provided a
sufficient number
of individuals be employed.
is to study some
If,however,the purpose of the investigation
of nutrition,
principles
questionrelatingto the fundamental
outset, however, the

very

To

make

to show

maintained

the

demonstration

that
and

the stock

that

absolutelycomplete, of course,
different kind of proteinin

of each

all the energy

containing material derived

it would
the
from

body
the

be

sary
neces-

had
feed

been
was

to the organism. Usually, however,

actuallycapable of yielding up its energy
its stock of
that if the body maintains
be assumed
on
a mixed
diet,it may
especially
maintained, while no considerable storage
protein,each particularkind is practically
of unavailable energy
in the body has been recognized.

BALANCE

THE

OF

NUTRITION

197

such, for example, as the relative values of the carbohydrates

and fats,the changes of live weight are of littlevalue as indicators.
For this there are two principalreasons.
in live weight.
In the first place the
282.
Fluctuations
from day to day,
live weight of an animal fluctuates considerably
taken under what seem
when
to be identical conditions,
even
of variations in the weight of the contents
chieflyon account
of the digestivetract.
This is true of all animals,but especially
of herbivora on account
of their comparativelybulky feed,and
"

reaches

For

the

in ruminants.

extreme

example,

which

had

been

receivingdaily for two

of 6.35 Kgs. of timothy hay and which
a
was
under
uniform
conditions as possiblewas
as
kept
weighed daily 24
hours after watering. On February 19 he weighed 419.0 Kgs. and
March
the same
6 practically
on
mediate
418.6 Kgs. The interamount,
follows:
weights,however, were
as
months

uniform

steer

ration

"

February
February
February
February
February
February
February
February
February
February

Kgs.
431.6 Kgs.
Kgs.
431.0
440.6 Kgs.
Kgs.
431.2
444.8 Kgs.
427.6 Kgs.

19

419.0

20
21
22

23
24

25
26

March

Kgs.
437.8 Kgs.
436.0 Kgs.
437.2 Kgs.

March

443.0

March

March

March

March

It is evident

427.9

27
28

that conclusions

Kgs.
428.4 Kgs.
433.4 Kgs.
436.8 Kgs.
418.6 Kgs.
based

comparison of single
weighings would have been entirelyuntrustworthy. Thus
a
parison
comof the live weight of February
6 would
19 with that of March
have led to the conclusion that the animal
tained.
mainwas
being practically
on
If,however, the initial weight had chanced
to be taken
shown
6 would have
February 20, a comparison with that of March
the
a loss of 13 Kgs., while
other hand, a comparison of February
on
with
March
would
shown
have
Even
aver19
a gain of 17.8 Kgs.
5
upon

198
aging two

or

it reduces the
of the average

ANIMALS

FARM

OF

NUTRITION

dailyweighings,as is often done, while

eliminate it. For example, a comparison

three successive
error

of

does

not

February

with

19-21

if each

that of March
be

3-5

shows

a
day later,viz.,
gain of 8.7 Kgs., while
average
March
the
and
shows
a loss of 4.8
4-6,
comparison
February 20-22
of singleweighings averaged, the
Kgs. By increasingthe number
nated,
uncertaintymay, of course, be further reduced but not entirelyelimieven
varying materially,as is illustrated
ten-day averages
by the followingfigures.

February
February
February
February
A

24-March

inclusive,
435.3 Kgs.
25-March 6, inclusive,432.7 Kgs.
26-March
432.6 Kgs.
7, inclusive,
27-March 8, inclusive,
Kgs.
434.2

similar reduction

number

of animals

of the

combined

the fluctuations
each

taken

5,

error

into

in the

be obtained

may
a

group

which

singleanimals

by the

is treated

tending to

of

use
as

an

dividual
in-

balance

other.

These

gain in

fluctuations

are

such

to

as

preclude the

use

of the

live

of the nutritive effect in exact

weight as a measure
scientific investigations,
while it is evident that they must
also
of commercial
be considered in the planning and interpreting
experiments,as well as in judgingthe effects of rations in practice.
Such
considerable
a
experiments should extend over

length of time and include

while the

considerable

of

animals,

comparisonsare based should be the

of as many
singleweighingsas possible.
average
In the second place,even
283. Composition of increase.
it possibleto ascertain
the gain or loss in weight by the
were
of the digestive
body tissues proper, exclusive of the contents
the empty weight, the compositionof the material
tract, i.e.,
increase of a kilogram in
An
gained would stillbe unknown.
tissue weight,for example,might consist chiefly
of adiposetissue
containing10 or 12 per cent of water (95),or it might be largely
muscular
tissue with 75 or 80 per cent of water
(87). Moreover,
aside from
the difference in water
content, the dry matter
of adipose tissue carries more
than that of
chemical
energy
muscular
tissue,so that a gain of a kilogram in the former
would
be equivalentto the storage of seven
case
or
eight times
much
the
in the latter. Finally,
as
as
a knowledge of
energy
weightson

which

number

"

NUTRITION

200

care

OF

ANIMALS

FARM

skill exercised in the choice of the check

and

animal.

The

sumed
as-

compositionof the two animals cannot in the nature

In a short
of thingsbe proved and is very unlikelyto be absolute.
tively
experiment,therefore,the error thus possiblyintroduced may be relalarge. Its importance diminishes the greater the increase
the longer the period covered
the originalweight, i.e.,
made
over
by the experiment. Furthermore, an experiment by this method
of additional check
be divided into periods only by the use
can
animals, involving additional assumptions as to identityof composition
these subdivisions,
for the reason
at different times, while even
be fairlylong. Finally,the method
ous,
is laborijust stated, must
The different parts of the
especiallywith the larger animals.
be separated,the weight of each part accuratelydetermined,
must
carcass
allowance
for
mechanical
losses
and
due
avoiding
making
be
A correct
sample of each part must
evaporation of water.
taken promptly and at once
treated as to preclude any changes
so
of a
previous to its analysis. The task of analyzing the carcass
the degree of
that of a steer, with
hog or sheep, and still more
is not one
to be undertaken
requiredin a scientific investigation
accuracy
lightly.
identityof

285.
test

balance

The

attempts

contained

to

in

experiment,on
and

experiment.

determine

the

body

the

outgo and

at

the
two

"

The

weightsof

comparativeslaughter
the several

different times.

ingredients

The

balance

come
contrary, consists of a comparison of inthe original
does not attempt to determine

body. If I know that I have a balance of $50 in

bank at the beginningof the month
and $150 at the end, it is
clear that I have gained $100 in the meantime.
This is the
of the comparativeslaughter
On the other hand,
test.
principle
if I know
that my depositsduring the month
were
$500 and my
drafts $400, I am
equallysure that I have gained $100, even
if I do not know
whether my balance at the beginningwas
$50
of the balance experiment. If,
or
$500. This is the principle
for example,a steer digests
of proteinout of his daily
750 grams
ration and if the amount
of nitrogenousproducts excreted in
of protein,
it
that he has katabolized 500 grams
24 hours shows
is evident that his original
stock of protein,
whatever
its amount
have been,has been increased by 250 grams.
sons
By comparimay
based on the same
plicated
comgeneral principle,
although more
of the body's
decrease
to details,the increase
as
or
stock of fat,glycogen,ash and water
of chemical energy may
or
stock in the

THE

be determined.

described

are

The

BALANCE

OF

methods
specific

in the two

NUTRITION

201

used

for such

comparisons

followingsections.

great advantage of the balance

experiment is the comparatively

it requires. A periodsufficiently
mination
long for the deterof the digestibility
of a ration (159) is in general sufficient
also for a balance
while
for
the
d
tion
eterminaexperiment,
requisite
of the respiratory
the
of
heat
products or
produced twenty-four
hours suffice,
and even
this short period may
to forty-eight
be divided
into a number
of subperiodsof a few hours each.
For this reason,
and
the animal
is not injured in the process,
also because
repeated
be made
the same
on
experiments may
subject,so that the effect of
various rations or conditions may
be compared on the same
individual,
of comparative slaughtertests necessarily
while the method
involves
between
different animals.
two
comparisons
On the other hand, the complete balance experiment requireselaborate
and
while
expensive apparatus,
opinions as to the relative
of labor involved in the two
amount
classes of experiments would
perhaps depend largelyupon the previous experience of the experimenter.
the
balance
shows
the
amounts
Furthermore,
experiment
only
of the constituent
etc.
fat,
gained or lost. It
protein,
groups
affords no opportunity to subdivide these and determine
the fate of
singlechemical compounds nor does it give any clue to the particular
regionof the body where the gains have been deposited.
short

time

which

"

286.

The
"

used

balance
as

of nutrition.

the titleof this

the balance between

body

"

income

and

"

The

phrase

chapter refers in
outgo of

matter

"

balance

of

generalway

and

energy

determined

trition
nu-

to

in the

of the balance experiment.

by the methods
of course,
the comparative slaughtertest, if comLogically,
bined
with determinations
of the feed consumed, may
also be.
regarded as a balance experiment. In it the income of the body
and the resulting
gain are determined, leavingthe outgo to be
inferred,while in a balance experiment in the technical sense,
the income
and outgo are
determined
and the gain is inferred.
Nevertheless,the latter type of experiment has played so large
of nutrition,
a part in the study of the balance
both for physiological
and for agricultural
that
clear
a
conception of
purposes,
its methods
and postulatesis essential for a comprehension of
of the results to be considered
in subsequent chapters.
many
The
be conveniently considered
under
the two
subject may

heads

as

of the balance -of matter

and

the balance

of energy.

OF

NUTRITION

202

" 3.

Balance

The

The

gain

or

ANIMALS

FARM

of

loss

Matter

of protein

protein which fails to

be stored up in the body is not excreted as proteinbut in the
The gain or
form of the various products of its katabolism.
be determined
loss of protein,therefore,cannot
by a direct
comparison of its income and outgo because there is no outgo
of proteinas such.
Since,however, the proteinof the schematic
body (280)is equivalentto total nitrogenousmatter, the gain or
be inferred from
that of its characteristic
loss of proteinmay
parison
element, nitrogen,and this is readilyascertained by a comof
of the total nitrogen
the feed with the total nitrogen
of the excreta, i.e.,
by a determination of the nitrogenbalance..
In Chapter V (228) it
288. Free
nitrogen not excreted.
stated that all the nitrogenof the protein katabolized is
was
and other organiccompounds which are exfound in the urea
creted
in the urine.
Obviously this is a point of fundamental
importance. If nitrogen leaves the body only as combined
nitrogenin the urine and in the feed residues and nitrogenous
in the feces,it is a comparatively
excretory products found
to compare
the income
and outgo.
If,however,
simplematter
287.

nitrogen balance.

The

"

Feed

"

the

metabolic

processes

or

the

fermentations

of

the

feed

in

then the nitrogen

tract yieldalso gaseous
nitrogen,
digestive
of the respiratory
products must also be determined, a task of
small difficulty.
no
The questionof the excretion of gaseous
nitrogenhas been
of investigation
and controversy.
the subjectof a vast amount
have been
two
Substantially
generalmethods of experimentation
and outgo of combined
followed,viz.,a comparison of the income
nitrogen and direct investigationof the respiratory
products,and the results of both have been in substantial
The
force of the great number
of experiaccord.
cumulative
ments
the

in

outgo

which

of combined

substantial

nitrogen

equality

between

been

observed

has

income
under

and
tions
condi-

of any considerable gainor

precludedthe possibility
with the fact that the very careful
loss of body protein,together
of recent
and accurate
investigations
years upon the respiratory
to
excretion of free nitrogenhave givennegativeresults,
amount

which

demonstration

that

nitrogen

Determination

289.

NUTRITION

leaves

203

the

body

of

balance.

nitrogen

There

"

balance

nitrogen,a determination
of the
requiressimply a determination

element

contained

excretion

only

in

the

in the visible excreta.

form

combined

OF

BALANCE

THE

of gaseous

in the feed and

is

this end

in the

being no

of the
amounts

visible excreta.

attained
alreadypartially

in

nitrogen
of this
dently
Evi-

periment
digestionexin addition
to provide
(158). It is only necessary
collection and analysisof the urine and, in
for the quantitative
experiments, of the perspirationand of the epidermal
very accurate
excreta, in order to obtain data for a comparison of the
the same
income
and outgo of nitrogen,and
precautions as to
length of period,uniformity of feeding,etc.,which are necessary
in a digestionexperiment, suffice also to render the results of a
balance experiment representative.
a

The digestion
nitrogen balance experiment.
clover hay used as an example in Chapter III (160)
also to illustrate the nature
of a nitrogen balance
serve
ment.
experimay
contained
In that experiment the hay consumed
daily
3.144
of
and
the
feces
while
the
1.267 Kgs.,
dry matter
daily
Kgs.
average
of
the
urine
for
dailyweight
Analysis showed
5.449 Kgs.
9 days was
the followingpercentages of nitrogen:
290.

of

Example
experiment with

"

"

In
In

dry
dry

matter

of

matter

of feces

%
2.240%
1.074%

hay

2.271

In fresh urine

found
to contain
(hair,dandruff,etc.)were
balance
of
The
daily nitrogen
1.87 grams
nitrogen per day.
be
accordingly computed as follows,showing a loss from the body
may
which, of course, must be placed in the income column to complete the

The

brushings of

the steer

balance.
Table

22.

Example

"

of

Nitrogen

Balance

Otttgo
Grms.

Nitrogen
Nitrogen
Nitrogen
Nitrogen
Nitrogen

in

hay

in feces
in urine
in

brushings
lost from body
.

28.40

58.50
1.87

88.77

NUTRITION

204

291.

of

Computation

FARM

OF

protein.
"

ANIMALS

The

matic
conceptionof the scheexperimentsare based
the animal
as
consisting

body (280) upon which balance

of
regardsthe total nitrogenousmatter
of protein.All the vast number
of other substances
substantially
containingthis element which have been identified as constituents
of the body are
in amount
as
insignificant
compared with the
of proteinwhich
it contains.
Accordingly,a gain
great mass
loss of nitrogenis interpreted
loss of
or
as
a gain or
signifying
of the latter may
be computed from
proteinand the amount
the former justas the proteinof a feedingstuff is computed from
its proteinnitrogen,
it beingonly necessary
to fix upon
a suitable
i. e., to know
factor or factors,
the average percentage of nitrogen
in body protein.
the results of

From

analyses of entire bodies of animals cited in

Chapter II, the average
nitrogencontent of the fat- and ash-free dry
matter
was
computed (99) to be :
"

In Lawes

and

Gilbert's

experiments
experiments

Chaniewski's

In

It is

probable that

still contained
extraction

some

in both

16.11%
16.06 %

the

supposedly fat-free matter

that
fat, it having been subsequently shown

with ether does not

cases

remove

the last traces

of it from

mal
ani-

tissues.
varianalyses (88)of the fat- and ash-free lean meat of ous
after correction for the glycogencontent
of the horse flesh,
species,

Kohler's

show

an

of Lawes

of 16.64 per cent.

Since the material
average nitrogen content
and Gilbert's and of Chaniewski's
cluded
experiments doubtless in-

residual fat and

non-nitrogenoussubstance,and
body is contained in the
it appears
muscular tissues,
to regard Kohler's
justifiable
figuresas
with
substantial
the
representing
position
elementary comaverage
accuracy
of body protein as a whole,especially
since they are the results
of direct analysiswhile the others are derived from slaughterexperiments
some

since the

largershare of the

in which

the limits of

other

proteinof the

error

are

somewhat

wide.

that average body

Assuming, on the basis of Kohler's results,
proteincontains 16.64 Per cent of nitrogen,the corresponding
proteinfactor is 6.0,and the gain or loss of nitrogenobserved in
balance experiment multiplied
a nitrogen
by this factor givesthe
gain or loss of protein. This is,of course, an approximation,
since proteinis not the only nitrogenoussubstance contained

in

the

since

and

body

OF

BALANCE

THE

NUTRITION

all the

not

205

animal

proteins contain

exactly 16.67 Per cent of nitrogen,but the

in most
far as it relates
cases
so
insignificant
On

this

basis,the

daily17.37

6.0

the results of
can

be

income

between

the balance

an

and

grams
experiment in

The

gain

foregoingexample was losing

body protein. Evidently
which
a gain of nitrogenoccurs
of

same

way.

offat and

loss

or

question of

outgo.

the
precisely

in

computed

the

to

is

in the

steer

104.22

involved

error

glycogen

By a method
quite similar in
to that just described for protein,it is possibleto compute
principle
bined
approximatelythe gain or loss of body fat from the comincome
and outgo of nitrogenand carbon, while if the
292.

The

balance

of

balance.

carbon

"

of oxygen
also be determined
can
made
and
exact
considerablymore

hydrogen and

computation

be

may

the
may

include

glycogen also. The experimental methods, however,

much
elaborate than those required for a
are
more
necessarily
simpledetermination of the nitrogenbalance,since it is evident
that, in addition

excreta, it is necessary
in

contained

the

determine

to

included,the hydrogen

the

of

of the

hydrogen

feed,the

and

is
it

outset

given in

and

is to be

oxygen

and

the

also be ascertained.

employed for these

succeeding paragraph (297),but at the

desirable to confine attention

seems

dioxid

excreted

water

of oxygen
taken up from the air must
outline of the experimental methods

purposes

of this element

amount

amount

An

of the visible

feed and

excreta, viz., carbon

gaseous

if the balance

methane, while

of the

the carbon

to

to

the

principles

involved.
293.

Computation

of

gain

or

loss of fat.

-*-

According to

the

body (280) on which the whole

of the balance experiment is based,substantially
scheme
all the
in the two forms of
carbon of the body is regarded as existing
protein and fat. Evidentlyif a comparison of the income and
outgo of carbon shows a gain of that element it can, according
to the fundamental
assumption, have been only in one or the
ever,
other or both of these two forms.
The nitrogenbalance,howconception

shows
of

of

the

the

schematic

amount

protein is known.

of

protein gained
If the carbon

and

the

of the

carbon

tent
con-

protein gained

206

NUTRITION

subtracted

be

have

can

steer

on

is

as

Table

gain

of

complete

follows

23.

the

of

statement

Carbon

and

respirationexperiment
and
carbon
nitrogen

Balances

of

400

16619
4357

timothy hay

grms.

linseed

grms.

feces

grms.

urine

37 grms.

56.4

21.9

Grms.

4730

grms.

142

grms.

methan

Outgo
Grms.

33-5

1428.7

32.4

124.2

8.0

i-3

dioxid
.

by body

Income
Grms.

2831.7
172.6

brushings
carbon

Gain

meal

Steer

Carbon

Outgo

Income
Grms.

grms.

sponding
corre-

Nitrogen

6988

the

readilycomputed.
In

"

remainder

"

Nitrogen

"

of fat and
be

can

balance.

carbon

carbon, the

of

in the form

of this substance

Example

balances

total

gained only

been
amount

294.

the

from

ANIMALS

FARM

OF

1290.2
106.6

11.

....

78.3

46.6

78.3

3004.3

The

3004.3

nitrogenbalance shows that the animal gained 11.1 X 6.0

of protein. According to Kohler's
66.6 grams
results (88), the
of
cattle
contains
of
quently,
Consecarbon.
protein
average
52.54 per cent
the proteingained in this experiment contained
66.6 X .5254
of carbon.
The
total gain of carbon, however, as
35.0 grams
shown by the carbon balance, was
than this,viz.,
more
46.6 grams, and
we
accordinglyhave the following:
=

"

Total

gain of carbon
in proteingained
Carbon

35-Q grams

Carbon

The

46.6 grams

gained as fat

elementary composition of

animal

fat

1.6 grams

shown

in

Chapter
A
(34) to be very uniform, averaging 76.5 per cent of carbon.
gain of 0.765 gram of carbon in the form of fat,therefore,is equivalent
of
of
to a gain of one
of
fat
carbon
to
or
a
one
gain
gram
gram
of fat, and accordinglythe gain of 11.6 grams
of carbon
1. 3 1 grams
in the form
of fat shows
a
gain by the animal of 11.6 -"" 0.765, or
the same
of
of fat. Substantially
method
11. 6 X
1. 31
15.2 grams
I

was

208

NUTRITION

0.7 literof carbon

OF

FARM

ANIMALS

dioxid.

it may
be computed that
Similarly,
if proteinof average
compositionbe oxidized to urea, carbon
dioxid and water, the respiratory
quotient will be approximately
0.8, although in realitythe quotientfor protein varies
accordingto the nature of the nitrogenousproductsformed and
of carbon
from
the amount
oxidation to
thereby withdrawn
dioxid.
carbon
Ordinarily,however, the proportion of the
exchange of the body due to the katabolism of protein
gaseous
is comparativelysmall,so that if,for example, the respiratory
quotientcloselyapproaches 1.0, it is clear that the katabolism
be chiefly
that of carbohydrates,
must
while if,on
the other
hand, its value approaches 0.7, it is equallyevident that the
katabolism
be chiefly
that of fat. Values for the respiramust
tory
quotientintermediate between these extremes
imply that
the katabolism
is in part that of fats (orproteins)
and in part
that of carbohydrates.
The respiratory
affords no
information
quotient of course
mination
regardingthe balance between income and outgo but its deterof the
gives valuable information as to the nature
material which is being katabolized in the body, particularly
in short periods.
297.

The

respiration apparatus.

"

determination

of the

exchange of an animal, such as is necessary in order to

formulate
of
the complete balance of matter, requiresthe use
form of special
some
as a respiration
apparatus.
apparatus known
In its simplestand earliest form
the respiration
apparatus
of known
consisted of a closed chamber
capacity,such as was
used by Crawford, Mayow,
Black, Priestly,Lavoisier and
others in their earlyexperiments. The animal was
placed in
the hermetically
sealed apparatus and the changes in the composition
of the enclosed air which
were
brought about by its
determined.
were
respiration
Evidently,however, the method,
while charmingly simple,is open
to objections.The
oxygen
of the air is graduallyconsumed, while the carbon
dioxid and
if the experiaccumulate.
Even
other productsof respiration
ment

gaseous

be broken
is made
while
Two

no

under
very
obvious

off before

fatal results

to

the animal

varying and increasinglyabnormal

long trials are possible.
methods

suggest themselves; either

of
to

ensue,

it

conditions,

at
once
avoiding this difficulty
absorb the products of respira-

BALANCE

THE

NUTRITION

OF

209

replacethe oxygen consumed or to conduct a current

of air through the apparatus. Correspondingly,
two
different
spectively
reapparatus have been evolved, known
types of respiration
tion and

or

from

the

closed

the

as

of the

names

circuit and

who
investigators

practicable
appliances,as

into

Pettenkofer apparatus.
into

of

animal

an

Each

intended

those

open

the

and
Regnault-Reiset

of these two

to

types may

be

the
divided
sub-

determine

those which

and

circuit apparatus,
first developedthem

the total gaseous exchange

take account
only of the

pulmonary exchange.
In the closed
298. The
Regnault-Reisetapparatus.
cuit,
cirtakes place in
or
Regnault-Reiset
apparatus, respiration
.

"

RESPIRATION

CHAMBER

0 used

M,0

",

N
0

deficient

^produced

introduced

CO0

ab"orbed
/Na OH

deficient
"

24

"

HjO

ROTARY

absorbed bu.

014
\

\Ca[OHl,)

"1

Fig

BLOWER

h.scu

respiration
(Atwater and Benedict,
apparatus.
Institution
Publication
No. 42.)
of
Washington,
Carnegie

Scheme

confined

of closed circuit

of any
tween
exchange beair,the possibility
it and the outside atmosphere being carefullyguarded
(a blower, for instance)
against. By suitable mechanical means
the confined air is kept in circulation over
suitable absorbents
which
dioxid given off, while
and carbon
take up the water
a

volume

of

consumed
the oxygen
is
of the compressedgas.
is shown

plus any

in

replacedfrom a gasometer
The
generalscheme for

Fig. 24.

The

increase

increase in the amount

in

or

such

weight

of carbon

cylinder
an

paratus
ap-

sorbents
of the ab-

dioxid and

NUTRITION

2IO

water

minus

any

methane
and

air at

the

contained

or

may
close

in the

the amounts

the amount

of fresh oxygen
contained in the air

of the oxygen
the quantityof this element

increase

of the apparatus shows

Any

apparatus shows

produced,while

of these substances

ANIMALS

FARM

in the air of the

contained

admitted

OF

hydrogen

excreted

accumulates

absorbed.

in

the

be determined

paratus
apof
contained
the
analysis

of the

The

by an
experiment.

amount

apparatus should,of course, remain

if the apparatus is

working properly.

of

nitrogen

unchanged

THE

BALANCE

OF

NUTRITION

211

"

:3
J3

3
G
CuO
o

P4

fa

FARM

OF

NUTRITION

212

ANIMALS

If the entire

respiratory
exchange is to be determined, the
representedin the
subjectis placed in the respirationchamber
gation,
diagram. If only the pulmonary exchange is under investichamber
is replacedby a mask
piece
mouththe respiration
or
or
even
by a suitable cannula inserted in the trachea.
Regnault-Reisetapparatus 1 is shown in
investigators
subsequently devised a largerone
Fig. 25. The same
of experimentsupon
in which they made
animals of various
a number
speciesincludingsheep, calves,swine and fowls. In theory this is
the most
nical
techperfectform of respirationapparatus, but numerous
The

of the

originalform

difficultiesarise in its
but Atwater
size suitable for

and

Benedict

which

man

was

later forms

Various

use.
2

have

the first to construct

were

capable of

been
one

vised
deof

high degree of accuracy.

respirationapparatus of

has constructed
a
Quite recently Zuntz
domestic
this type for experiments on
animals, a section of which
in Fig. 26, while for the determination
of the pulmonary
is shown
4
has devised a so-called "Universal"
exchange,Benedict
respiration

apparatus.

299.
open

The

Pettenkofer

the

In

"

Pettenkofer, or

circuit,respirationapparatus, the subjectbreathes

continuous

measured

of water, carbon
after passingthe
much

apparatus.

current

dioxid and

of

atmosphericair

is determined

methane

of
animal, the difference,

of each gas the

for small animals

subjecthas

whose

added.

the entire amount

in

content

before and

course, showing how

In an apparatus suitable

of carbon

dioxid

and

in the

mined,
be deterincoming and outgoingair current
may
but in the largerforms it is necessary
the air
to measure
and make
current
small samples,so that
analysesupon relatively
the analytical
terminat
errors
are
multiplied
by a largefactor,while a dewater

of the

oxygen

balance

has

not

yet been found

apparatus is shown
as

practicable.The generalscheme of such an

in the diagram,Fig.27.
As in the case
of the Regnault-Reiset
be replacedby a mask,
apparatus, the respirationchamber
may
of the pulmonary
mouthpiece or cannula for the investigation
exchange.
1

Ann.

Carnegie Institution

Landw.

4Deut.

de Chem.

et de

Physique,3"me Series,26,
of Washington,

Publication

Jahrb.,44 (1913),776.
Arch. Klin. Med., 107 (1912), 156.

299.

No.

42

(1905).

BALANCE

THE

The

first

form
practicable
1

by Pettenkofer
shown

for

Fig. 28.

in

OF

of open

NUTRITION

circuit apparatus

"

CHAMBER
9

"

COz

27.

Henneberg
G.

Kiihn

in

use

and

and

ABSORBED

of Pettenkofer

respirationapparatus.

readilybuilt of any desired size led to its

animals, notably by
agricultural
investigations
upon
Stohmann
Stohmann
at
at Gottingen,
Leipzig and

the fact that it could

extensive

METER
ME1

J^OSCO^

CH+
OXIDJZED

Scheme

"

"

J"

NzO"5COz
ABSORBED

METEP.

PRODUCED

METER.

Fig.

devised

was

is
Its generalappearance
experiments on man.
The
comparative simplicityof its operationand

RESPIRATION
"

213

be

later Kellner

Fig.

28.

"

at

Pettenkofer

Explanatory sketch.
(Atwater, U.
Stations,Bulletin No. 21.)
1

Ann.

Mockern.

Chem.

S.

respirationapparatus.

Department

of

Agriculture, Office of Experiment

Pharm., Suppl. Bd. II, p.

1.

NUTRITION

214

OF

FARM

ANIMALS

The

principleof the Pettenkofer apparatus has also been very

extensivelyused for the investigationof the pulmonary exchange,
the development of
to whom
especially
by Zuntz and his associates,
this form is largelydue.
Figure 30 shows a horse equipped with a
tracheal cannula for experiments with this type of apparatus.
ing
Owthe fact that the excretory
of air,as is the
times their volume
to

used, the results

Fig.

are

of oxygen

the amount

given

much

consumed

when

case

sharper

and

as

not

are

gases

as

many

respirationchamber

it is

well

diluted with

possibleto
that

is

determine

of carbon

dioxid

off.

29.

300.

"

The

can
respiration
(Bailey'sCyclopedia of Ameriapparatus.
Agriculture.)

Mockern

Investigation of pulmonary
a

determination

either with

of the

exchange.

gaseous

"

For

exchange

the

in

many

the

poses
pur-

lungs,

Regnault-Reisetor the Pettenkofer type of

of the total exchange
to determinations
apparatus, is preferable
in a respiration
chamber.
The
former method
is especially
adapted for short experiments. By its use, it is possibleto
of the metabolism, the
trace
sharplychanges in the amount
of
quotient, etc., produced by the administration
respiratory
feed substances,drugs, etc., by experimentallesions,and especially
would
often be
by work,
changes whose amounts
for
relatively
very small as compared with the total excretion
in the respiration
chamber
and which,
24 hours as measured
if they did not escape detection altogether,
could not
therefore,
logically.
chronobe as accurately determined
either quantitatively
or
to continue
On the other hand, it is impracticable
its use through longperiods, a day, e.g.,
and since it takes
"

"

"

2l6

of the excretion

account

no

OF

NUTRITION

of

the total balance

ANIMALS

through the

skin and

the

alimentary
pute
possibleto comcarbon, hydrogen and oxygen by its
that

methods

indirect

canal, it is only by

FARM

it is

use.

301.

of ash

and

of water

Balance

apparatus of either type

of water

the

by

vapor

The

ingredients.
"

to

serves

determine

well

subjectas

piration
res-

the

cretion
ex-

that of carbon

as

other gases and thus, in connection

analyses of the feed and visible excreta,

with

dioxid and

the

sary
neces-

establish the

to

less difficulty
more
or
hydrogen. Unfortunately,
is experiencedin determining accurately the hydrogen balance
of water
in the
to condensation
owing in part to the liability
of organic
apparatus and in part to the fact that the amount
of the animal
hydrogen actuallyenteringinto the metabolism
and
of water
consumed
is small as compared with the amounts
simply evaporatedagain.
of course, with the possibleexception
The
ash ingredients,
of sulphur,
in the visible
all leave the animal
of minute
amounts

gain or

loss of

of these

the balance

and

excreta

determined

the

accordingto

same

elements

therefore

may

as
principles

the balance

be

of

nitrogen.

" 4.
302.

animal

of

nutrition

Energy

of

includes

energy.

the

Since

also,correspondingto the dual function of

but

matter

"

of energy
(207),the
with the income
and

is

the

(263),with

study

Balance

transformer
a
essentially
nutrition is not only concerned

body

outgo of
feed

of

Balance

balance

The

loss of energy
by the body. The
is a method
of investigating
of energy

gain

of the balance

or

important problems of nutrition which has been

and
which
has proved
especiallydeveloped in recent
years
its consideration,
Before
fruitful of results.
entering upon
of the

some

however,

brief review

related
as
energetics
superfluous.

to

of

some

of the

elementary concepts of

physiological
processes

may

not

prove

Elementaryprinciples
303.
used

Energy.

without

"

any

Up

to

this

point the

precisedefinition.

In

word
a

energy

has

been

study of
specific

the

NUTRITION

OF

BALANCE

THE

217

important to have as definite

It is
by the term.
a
conception as possibleof what is meant
not
altogethereasy to give a simple generaldefinition of energy,
be
that given by Noyes * may
for the present purpose
but
"
erties
adopted,viz., That which gives rise to changes in the propof energy,

balance

and

of bodies

to

the

be

to

power

however,

the present purpose,

For

it is

however,

readilyapprehended from

more

such

produce

changes."

the conception of energy

illustrations than

may

from

nitions.
defi-

conveniently approached from the side

A moving body is capableof producing certain
of mechanics.
The
effects by virtue of its motion.
fallingweight of a pile
sistance
against the redriver,for example, forces the pile downward
time produces heat at
of the ground and at the same
the point of impact. The
projectilefired from a sixteen inch
the cohesive
ship overcomes
gun strikingthe side of the armored
force of the armor
plate and deforms or penetrates it,while
The blows of
of heat.
the blow also gives rise to an evolution
raise the iron
the blacksmith, if rapid and heavy enough, may
his anvil to a red heat.
Accordingly,it is said that a moving
on
The

be

subject may

body possesses

energy

called kinetic energy,

in the form

or

energy

of motion.

body suspended above the earth

of contact
ground, and at the moment
If

of kinetic

certain amount

something which

its fall from

other

was

the
which
energy,
virtue of its position,
may
of

form

with

which
not

the earth possesses

was

energy

generatedduring
This

of motion.

body possessedprevious to
be called

gravitation
energy.
ing
relation is illustrated by a swinging pendulum. Dursame
which it posthe downward
swing, the gravitation
sessed
energy
when at its highestpoint is converted into kinetic energy,

its fall by
The

energy

is set free it falls to the

while

when

expendingwork, which is stored

be liberated again as kinetic energy

of

conscious
energy,

which it possesses is reconverted

it rises the kinetic energy
lifta weightwe are
When
into gravitation
we
energy.

to

up

as

when

gravitation
the weight

falls.
304.

motion
accurate

Forms

of

the rate of
general,whenever
familiar if less
body is increased (or,to use a more
of energy.

"

whenever
expression,
1

General

In

motion

is

produced)it

Principlesof Physical Science, 1902.

is

to

be

2l8

NUTRITION

in
as
inferred,

the

OF

of the

case

FARM

ANIMALS

falling
body

or

the

pendulum, that

the kinetic energy produced has been derived from some

other
In the examples thus far given this other form
form of energy.
of energy

however, this is

cylinderof
the motion
the

the

not

steam

of the

case.

The

familiar

expanding

instances,
in the

steam

of its heat to produce

engineparts with some
piston. The electric current in the wire sets

of the motor

armature

In many

gravitation
energy.

was

in revolution.

The

gasolinein the cylinderof an engine produces motion

and chemical
engine as well as heat. Heat, electricity
all be sources
of kinetic energy
may
of heat energy, electrical energy and

The

manifestations

but

its forms

Of

"

Magnetic

energy

7.

Chemical

energy

3.

Cohesion

8.

Heat

4.

Volume

9.

Radiant

5.

Electrical energy

energy
energy

most

in the animal
305.

energy,

chemical

energy

energy

and

in

of energy.

speakingof

various forms
may

in turn

of energy
be converted

is converted

heat

energy

of

ergy
en-

body.

Transformations

assumed

energy

importance in consideringthe balance

"

As

is illustrated

examples given in the previousparagraphs,and

Heat

is inferred.

energy

2.

those of

are

chemical

6.

these, kinetic

action

therefore the existence

Kinetic energy
Gravitation energy

i.

of the

of energy are of the most

varied character
be convenientlygrouped under the following

may

generalheads

and

of

combustion

changes in the
capableof mutual

energy
are

into motion

into heat when

as

animal

by

has

the

been

body, the

transformations.

in the heat

engine. Motion
a moving body is retarded
with another body. When

by friction or stopped by contact

its chemical energy is converted into
gasolineis burned freely,
heat, but when it is exploded in the cylinderof an engine it
be stored in the
This motion in turn may
yieldsalso motion.
form of gravitationenergy
in a lifted weight,or as cohesion
in a coiled spring,or it may
trical
of elecbe made
a
source
energy
which in its turn givesrise to the radiant energy
energy
of lightin the filament of a lamp.
In brief,
all the physical
phenomena of the universe of which
take cognizancecan
of changes of
we
be described in terms
can

either

energy

to

as

OF

BALANCE

THE

form

NUTRITION

and
intensity,

or

to identifythe concepts, of
physicists

that the former

maintain

Without

the latter.

it will be convenient
familiar

can

be

follow

for

this fact has led

and

matter

into

this debated

present purpose

our

and energy
regarding matter
entities.
although indissolublyconnected
The

306.

of

course

conservation

of energy.

"

energy is converted into heat energy

of heat obtained
is-always the same

employed
heat

be

in

it is found
no

effectingthe conversion.

question,
the

more

distinct

two

unit

of

of kinetic

that the

amount

what

the process
Similarly,if a unit of

matter

into kinetic energy

the amount
of the latter
is always the same
is always equal to
and moreover

the

quantity of kinetic
heat is produced.

What

When

as

to

converted

obtained

of

some

energy and
fullyinterpretedin terms

entering here
to

219

energy

which

disappearswhen

unit

one

is true

of heat energy and kinetic energy in this respect

has been shown to be true of all the forms of energy.
Not only
are
they convertible into each other but there is no loss or gain
of energy

in the conversion.

When

quantity of energy of one

form disappearsan equivalentquantity simultaneously
appears
in some
somewhere
This great generaliother form or forms.
zation,
perhaps the most
important in the history of physical
of energy,
the law of the conservation
as
science,is known
or
first
law of energetics. It was
the
first clearlyand distinctly
formulated
by Mayer in 1842 and since that time has been
verified by a great number
of the most
exact
experiments and
forms the basis of modern
conceptions of physicalprocesses.
In substance, it asserts
that the total energy
of the universe
far as man
knows
it is a constant
as
tinual
quantity, subjectto con-

changes of
That

the law

form

but neither

of the conservation

created

nor

of energy

destroyed.
applies to the
was
exceedingly
experimentally

taking placein the body of the animal

probable, a priori,and has been demonstrated
by the researches of Rubner upon dogs, of Laulanie on various
animals, of Atwater, Benedict, Lusk and their associates upon
and
of Armsby
men
and
cattle.1
Fries upon
The
tance
imporof this fact in relation to the study of energy
changes in
processes

the
1

body

is obvious.

Compare the writer's Principlesof Animal

Expt. Sta., Bui. 126.

Nutrition,pp.

263-268 and

Penna.

OF

NUTRITION

2 20

Heat

307.

unique.

energy

FARM

ANIMALS

In

"

respect heat

one

Other

energy

forms

of energy
in
are
into heat but there is

occupies peculiarposition.
generalreadilyand completelytransformed
known
method
no
by which heat can be completelytransformed
a

into other

kinetic energy.
Whatever
portion of
the heat is thus transformed
obeys the law of the conservation

of energy
308.

forms, such

but
Units

as

part of it always remains

of

energy.

Quantities of

"

by convertingthem into the same

with some
quantityof the same
as

Since

of heat.1

in the form

form

form

measured
are
energy
and
comparing them

of energy

sumed
asarbitrarily

unit.

of
quantities

of mass, space and

the fundamental
as

kinetic energy
time, a unit based
unit

of

can
on

energy.

be

expressedin

terms

these concepts is taken

so-called
The
C. G. S.

unit is the erg. An erg is a quantity

(centimeter-gram-second)
of energy equal to twice the kinetic energy
possessedby a mass
o
f
of one gram moving with a velocity one centimeter
per second.
Since this is a very small quantity,a unit called the joule,equal
measurements
to ten million ergs, is often employed in practical
of energy, that is,i joule io7 ergs.
For purposes
where a still
largerunit is desired the kilo-joule
equal to one thousand joules
is also employed.
In practice,
however, heat is the form of energy which generally
lends itself most
determination
and,
readily to exact
since other forms of energy are easilyconverted into heat, units
The
of heat are extensively
employed in the study of energy.
=

is the calorie,
which
unit for this purpose
quantityof heat requiredto raise the temperature of one
of water
one
degree centigrade.2

most

common

The

foregoingis known
largerquantitiesof

Where

as

the

heat

small,or

are

to

gram

calorie
the

be measured

is the
gram

(cal.).
large,

This is,of course,

one
aspect of the second law of energetics. Theoretically,
perfectheat engine with a lower temperature limit of absolute zero would convert
heat completely into kinetic energy.
Since, however, we can neither obtain the
of absolute
construct
zero
nor
a
temperature
perfectheat engine, this theoretical
less remotely approached in
limit
which
is
be
more
or
conception
simply a
may
but
attained.
never
practice
2 Since
tion
the specific
heat of water
varies at different temperatures, an exact defini1

of the calorie must

specify the temperature at which

differs in this respect but the preferable unit is the mean
hundredth
water

of the amount

from o" to ioo" C.

of heat

required to

Practice

it is measured.

calorie,which

raise the temperature

of

is
one

one

gram

one-

of

THE

BALANCE

NUTRITION

OF

equal to
kilogramcalorie (Cal.),

221

thousand

small

calories,
is employed, while for still largerquantitiesthe Therm, equal
In the following
be used.
thousand
to one
largecalories,
may
the large,or kilogram,calorie,
calorie signifies
the term
pages
or

one

expresslystated.
are
gravitation
energy

unless the contrary is

units

Certain

of

frequentlyused,
important ones
being the

the

especiallyin

also

more
mechanics,
and
the foot pound.
The
meter, the kilogram meter
gram
is the energy
meter
required to raise a weight of one gram
gram
gram
verticallythrough one meter in oppositionto gravity,the kilogram
kilometer
is the energy required to raise a weight of one
quired
through one meter, and the foot pound is the energy reto raise a weight of one
pound through one foot. Since
the earth's
the force of gravity varies at different points on
surface these units as thus defined are not invariable.
Taking the
force of gravity at sea level,
however, as equal to 980.5
average

dynes,the

these various

relations between
OF

EQUIVALENCE

UNITS

OF

units

are

as

follows

ENERGY

Kilogram
Calories

meter

gram

1
1

kilogram meter
foot pound

calorie

Calorie

309.

io-6

0.002344

0.000324

O.OOI

Measurement

measured

are

0.2344

of

in
chiefly

the temperature

of

heat

two

some

energy.

"

Quantities of heat

viz.,by their effects in raising

substance or in changing its state of
ways,

for measuring quantitiesof heat

aggregation. Instruments
called calorimeters,
heat measurers.
are
i.e.,
In the first method, as alreadyimpliedin the definition of the
calorie (308),water
is ordinarilyused as the calorimetric substance.1
For example, if the quantity of heat to be measured

be transferred

can

the

kilogram of water, and if

dent
is thereby raised 20 C, it is evitemperature of the water
that the quantityof heat imparted to it is two large caloOther

the most

without

substances than water

convenient.

may,

loss to

of course,

be employed, but water

is

usually

NUTRITION

222

ries.

calorimeter

Such

calorimeter.

varietyof forms
two

essential
or

OF

FARM

constructed

after this

calorimeters

according

requirements are

radiation

have

the

to

ANIMALS

been

that any

calorimeter

Lavoisier's ice calorimeter.

is one

used
extensively
simple form of
of heat

water

great

in view.
of heat

escape

The

by

duction
con-

preventableor measurable
be accurately
determined.
is caused to expend itself in

changing the physicalstate of some

in meltingice or in evaporatingsome

"

in

shall be either

that the temperature increase

In the second method
the heat

31.

devised

special
purpose

and

Fig.

is
principle

for certain

volatile

ice calorimeter

used

by

ice

of Physiology.)

and has been

of calorimeter

classes of work.

example,

liquid. The

Book

(Schaefer,Text

of the oldest forms

as, for

substance

Figure 31

shows

The

Lavoisier.

source

is

placed in the central vessel and imparts its heat to

the surroundingice,while the access
heat is
of any extraneous

preventedby

the outside

In Lavoisier's calorimeter

by collectingand
accurate

ice

method

jacketof

ice.

the amount

of ice melted

weighing the resultingwater,

of measurement

undergoes when

converted

is based
into water.

upon

but

was
a

determined
much

the contraction

more

which

NUTRITION

224

ANIMALS

FARM

OF

l..,.".

i:":i,i.n:,

""

"
,.

f'iG. 32.

"

Section

of bomb

calorimeter.

;"i
"'";

(Atwater, U.

S. Department

Office of Experiment Stations,Bulletin No.

21.)

of Agriculture,

THE

is transformed

OF

BALANCE

into heat

by its

NUTRITION

almost

225

instantaneous

The
in oxygen.
to this question is found
answer
called the law of initialand final states.

This

law

solelyupon
at all upon

is

is that in any

of
independentsystem the amount
during a change in the system depends

transformed

energy

burning

in what

the initial and

the

rapidityof

final states

the

of the

system and

not

the kind
upon
number
of the intermediate stages through which it passes.
or
Although this law is true in the generalform here stated,it was

originally
propounded
with

start

starch and

quantitiesof carbon
energy

converted

whether
oxygen

of

related

as

oxygen

whether

to

and

dioxid and

into heat

end

reactions.

with

water, the

the

through

energy

it be

subjectedto

intermediate

necessary

between

obtain

the

we

of chemical

amount

or

slow oxidation in the tissues

to

and

stages like maltose,

dextrose,lactic acid,etc.,etc., as

simply

If

corresponding

in the soil ; whether carbon

dioxid
the immediate
products of the action or whether

passes

to

chemical

plantburied

are

is

nor

other forms is the same, no matter

be burned
almost instantaneously
in pure

the starch
or

transformation

determine

in the

the

body

water

the starch

glycogen,

of the animal.

It

difference in chemical

the system in its initial and in its final state

of energy
transformed
amount
during the

change.
313.

method
animal

Measurement
of

measuring the
motion

as

into

of kinetic

energy

energy

energy.

"

liberated

The

by

most

common

machine

is its conversion,actuallyor

or

an

virtually,

gravitationenergy, which is measured

by the units given
of energy
on
a previous page
(308). In case of small amounts
a
weight may be actuallylifted,the product of weight into
distance givingthe number
of gram
centimeters or foot pounds
of energy
expended. In other cases, the subject may
pull
againsta resistance produced,for example,by the friction of a
form
of spring
brake, the traction being measured
by some
balance.
In this case
of fact,
the kinetic energy is,as a matter
converted
into heat, but the tractive pull multipliedby the
distance gives the equivalentnumber
In
of gravitationunits.
stillanother form the subjectvirtually
liftshis own
weightby
the
inclined
of
tread
the
a
climbing
body weight
plane
power,
multiplied
by the distance multipliedby the sine of the angle
of ascent equaling
the units of gravitation
energy to be measured.
Q

226

NUTRITION

Figure 33 shows a form

work experimentsupon
Another

method

convertingit

into

for

OF

FARM

ANIMALS

of this apparatus
horses.

measuring

electrical energy

used

kinetic

by

for

Zuntz

consists in

energy

by causing the subjectto

work

of

against the resistance

field.

magnetic

The

of current

amount

thus erated
genin
measured

be

can

electrical

units, or, as has

been done by Atwater, Benedict
and others,the electrical
"

be converted into
energy may
in calories.
heat and measured

The

body'sincome
Gross

"

314.

can

of energy.

energy

Only chemical

be

utilized.
in

stated

energy

As

"

was

the

introductory
this chapter,
paragraphs
the animal body resembles an

of

internal combustion

motor

in

mechanism

for

the

being

conversion

of

of certain

energy

chemical

the

compounds

T3

in

contained

the

kinetic energy.
the balance

feed

into

In considering

come
in-

between

outgo of energy,

and

is essential

to

it

recognize a

pointof resemblance,

further

viz.,that neither the animal

nor

the

other

than

There

is no

animal
way

motor

chemical

energy.

evidence

that the

body

can

of the

any

of energy,
or

utilize

can

such

use

other
as

solar

in any
forms

tricity
heat,elecradiation

reach it from

which

engine can

energy

Gross

energy.

NUTRITION

environment, any
of

is not

body

can

than the
of

or

gasoline

electric current.

an

but the

derive its

only source

supply.

of energy

income

The

"

227

more

fallingwater
merely a source

the animal

which

315.

its

the energy

use

Chemical
from

OF

BALANCE

THE

be

may

tained,
ascer-

tained
contherefore,by determining the chemical
energy
in the various
compounds present in the feed in the
manner
already indicated (309),viz.,by convertingit into heat
of a suitable
of the latter by means
and measuring the amount
of chemical energy is
calorimeter.
In other words, the income
In order to
of the feed.
measured
by the heat of combustion
of energy
avoid the implicationthat this is the total amount
the
to use
with the feed (311),it will be convenient
associated

term

the feed is

heat when

as

Since the chemical

that this is
chemical

simply
of

energy

It is

the

does

anthracite

coal, with
would

gram,

gas, with

the

upon
utilize.

Heats

and

not

the heat

of

of combustion

combustion.

it is the

produced by

7.9 Cals.

some

feeding stuffs,while hydrogen

most

of

more

34 Cals. per gram

than

"

The

feed value

Obviously,the

depends not only upon its content

proportion of the latter which
of

that

to the animal.

stillhigherin the list.

stand

but

heat

outrank

pressed
usuallyex-

that the gross energy of

its nutritive value.
Otherwise,

measure
a

into

point out

to

heat of combustion

substance

316.

not

use

fested
mani-

understood,however,
clearly

feeding stuffs and

is of

is

its amount

of convenience

matter

scarcelynecessary

feed

would

which

their combustion

of

of

energy

of energy

amount

completely oxidized.
a feeding stuff is converted

of measurement,
for purposes
in heat units. It should be

heat

per

the

equivalentto

as

gross energy

heats

of gross energy

the

body

of combustion

can

of

determined.
been
organic substances have
a
Atwater 1 in 1895 published a compilation of results upon
of compounds of importance in nutrition,Fries 2
largenumber
and
Benedict
extensive list,and
has prepared a rather more

great

Osborne

variety

have

of

the heats of combustion

determined

of nineteen

vegetableproteins.
S. Dept. Agr., Office Expt. Stas.,Bull.
S. Dept. Agr., Bur. Anim.
Indus., Bui.

U.

U.

Jour. Biol. Chem.,

(1907),

119.

21

94

(1895).
(1907).

228

OF

NUTRITION

The

followingtabulation

FARM

ANIMALS

give a generalidea of
the gross energy
of some
of the more
important substances
concerned
in nutrition.
It should
be specially
noted that the
figuresgiven are in most instances simplyapproximate averages.
Table

Animal

protein
Vegetable protein

24.

"

may

serve

to

Gross

Approximate

Energy

....

Carbohydrates

....

Sucrose
Animal

fats

Butter

fat

Vegetable fats

....

Ether

extracts

of seeds

Ether

extracts

of

roughages

followingexamples taken from the work of Kellner and of

to give a generalidea of the gross
Armsby and Fries will serve
of common
that the
feedingstuffs. It will be observed
energy
is relatively
instances.
small in most
range of variation
The

Table

Roughage
Timothy

25.

hay
hay
Mixed
hay
Alfalfa hay
Meadow
hay
Red

clover

Corn

Oat

stover
straw

Wheat

straw

Straw

pulp

"

Gross

Energy

of

Feeding

Stuffs

THE

BALANCE

OF

NUTRITION

229

Concentrates

meal

Corn

Hominy
Wheat

....

chop

bran
.

Grain

mixture

No.

ia

Grain

mixture

No.

2b

molasses

Beet

Starch

oil

Peanut

Wheat

gluten

a. Wheat
bran, 14.28 per
meal, 42.86 per cent.
b. Corn
meal, 60 per cent
10

per

cent;

meal, 42.86

corn

crushed

oats, 30

per

cent

old process

cent;

per

old process

linseed

linseed meal,

cent.

Some

data

also available

are

chieflyfrom

derived

Table

26.

Gross

"

energy

gross

of the

feeding stuffs. The following averages

Kellner's
investigations.

of

digestednutrients

regardingthe

Energy

of

are

Nutrients

Digested

Per
100
Pounds
Therms.

Protein

Crude

gluten

of wheat

Protein
"

assumed

average

193.O
of

extract
Nitrogen-free

of starch

Peanut
Total

extract

of

hay

192.O

189.8

hay

377-5

....

oil

400.I

organicmatter

In the
used

258.6

fiber

Nitrogen-freeextract
Ether

271.O

are,

of

roughage

202.9

computation of energy balance, the factors commonly

and for body
for body protein 5.7 Cals. per gram

fat 9.5 Cals. per gram,

although Kohler's average
former is slightly
lower,viz.,5.628 Cals. (88).
The
Chemical

energy

for either of two

outgo of chemical

suppliedin
reasons:

the

the

energy

feed

first,the

for

may

escape

substances

unused

carrying it

NUTRITION

230

may

be

FARM

OF

fail to be

incorporatedinto
incompletelykatabolized.
The

317.

feces.

Since

"

ANIMALS

the

body, or second,they may

greater

of most
feedingstuffs
organicmatter
by farm animals and so does
(148),a considerable amount
proper
in the

escapes

feces,while

(154) contain

carry

conversion
of farm

animals

fails of

suitable

sorption
digestionand reinto the body
enter

not

of unused

feed

energy

excretory products which

which has failed of
energy
The

body.

constitutes

very

chemical

in
precautions,

be

they
plete
com-

of the feces

energy

considerable

Its amount
total outgo of energy.
can
of feeding stuffs by burning a
the case

with

proportion of the

the

chemical

in the

less

or

item

in their

determined

as

in

sample, after drying

calorimeter and measuring the

heat evolved.
Combustible

318.

gases.

"

The

combustible

gases
also carry off

by fermentation in the digestivetract

of unused
chemical
large amounts
energy,
being preciselyanalogous to that in
way
of the feces

matter

These

except that it escapes

well

in

the

produced
relatively

loss in

this

the

undigested
invisible products.

separatedfrom the other gaseous

of making a direct determination
for the purpose
excreta
of their
The
of carbon
in
and hydrogen excreted
amounts
energy.
them, however, can be determined with the aid of the respiration
ane
apparatus and on the well-founded assumption that only methand hydrogen are produced the amount
of each excreted
gases

cannot

be

may

be

gases

of
being known, the amount
off can
be readilycomputed.

carry
319.

calculated.

Products

of

of

The

heats

of combustion
chemical

incomplete katabolism.

of both

energy

which

The

heat of

"

these

they
bustion
com-

of heat
substance,as alreadydefined,is the amount
evolved when
it is completelyoxidized,that is,in the case
of
substances
it is
ordinarilyoccurringin feeding stuffs,when
burned
to C02, H20, N2, and
in the
S03. If the katabolism
a

body stops short

energy

of these end

transformed

is

products,the quantityof chemical

clearlyless than the gross energy of the

substance
the

by an amount
equal to the
incompletelyoxidized products.

The

proteins of the feed constitute the

of this sort.

and

heat

part of its

All

the

of combustion

most

nitrogenof the
carbon, hydrogen and oxygen,

of

stance
important indigestedprotein
are

excreted

in

OF

NUTRITION

232

FARM

ANIMALS

and

measured
as
by their heats of combustion.
If,
gaseous,
be subtracted
from
then, the gross energy of the total excreta
the gross energy
of the feed, the remainder
shows
how
much
of the chemical

of the feed can

be metabolized, that is,
energy
into other forms in the organism. To this difference,

converted

metabolizable

the term

Metabolizable
energy
in the

has

energy

energy
of the feed minus

be

been

applied.
brieflydefined

may
the gross energy

of the feed and

total ration

follows

as

were

Energy offeed
6988 grms.
400

and

excreta

the
:

metabolizable

illustrate the
content

energy

of

energy

the

"

timothy hay

Cals.

27,727

linseed meal

grms.

gross
Thus

of the excreta.

experiment cited previously (294) to

of determiningthe balance of matter, the

method

the

as

181

Cals.

29,538 Cals.

Energy of excreta
16,619 grms.
4357
142

grms.

feces

*4j243

Cals.

urine

12

Cals.

methane

grms.

10

1896 Cals.

....

Total
Metabolizable

It should
assertion

12,189 Cals.

energy

be observed

whatever

Cals.

17,349

as

to

that

the

foregoingdefinition
into which

the forms

makes

no

the metabolizable

degree to which the

of
has been of service to the organism. Some
transformation
be retained in the body in a gain
the energy, for example, may
of fat or protein,as in the illustration just given,i.e.,
it may
be temporarily set aside as a reserve
to be used later,but it is
and
still capable of transformation
therefore
into other forms
has been

energy

constitutes

transformed

nor

as

to

the

On the other
energy.
tion
substance
capable of oxida-

part of the metabolizable

hand, the feed might contain

some

in the

was

value
body but of no physiological
simply burned to get rid of it. The heat

might
would

be

entirely useless

to

be part of the metabolizable

the

to

thus

animal, yet
energy

it and

generated

this

of the feed.

liberated as heat in the

energy
fermentation
constitutes
part of the metabolizable
althoughit does not enter into the tissue metabolism.
the
similarly,

which

energy
what
Some-

methane
energy,

Metab-

BALANCE

THE

OF

NUTRITION

233

olizable energy means

simply energy capable of transformation
It is the maximum
in the body.
quantity which the feed can
contribute
does

the

to

changes

energy

the method

from

apparent

the

nutritive

necessarilymeasure

not

in

organism. That
is indeed

value

it

ciently
suffi-

for its determination.

used

example already given shows, this does not requireany

of the gain or loss by the animal, but, like a
measurement
itself simply with the feed and
digestionexperiment, concerns
the

As

the excreta.
323.

for

Synonyms

metabolizable

energy.

Two

"

other

frequently employed with substantiallythe same

metabolizable
availas
significance
energy, viz.,fuel value and able
terms

are

energy.

Fuel

value.

of the feed is evidently

energy
into heat in the body. Since a

The

"

metabolizable

capable of conversion
all of it is actually thus
considerable portion and sometimes
is usuallyexpressedas a matter
converted,and since its amount
of convenience in heat units,the term fuel value (or physiological
heat value) has come
into use as synonymous
with metabolizable
energy.

The

advantage of brevity,but has also certain

disadvantages. In conjunction with the unit of measurement
employed, it has a tendency to suggest that the purpose of
the feed is to supply heat energy
and
that it is of value
in proportion as it can
do this,which
is far from
being the
Moreover, there appears

case.

due

has the

term

the fact that the

to

in relation

"

The

its value

the

The

less

it

can

is used
"

fuel value

of

furnish when

energy,
as

different

sense

of

boiler

feedingstuff,on

oxidized

confusion

coal,for example,
it liberates when
burned,

fuel under

as

of

in

the gross energy

of its metabolizable

sense

which

if used

danger of

some

of heat which

amount

fuel value

be

term

same

to
therefore,
corresponds,

to
i.e.,
plant.

or

fuels.

the total

means

and

to

to

or

feedingstuff,
in a heating

the other

hand,

is the amount

it is in the

in

of heat

more
body,i.e.,

incompletely.

Available

energy.

"

much

more

available
employment of the term
German
Nutzwert"
Physiologischer
"

metabolizable

This

unfortunate
energy,
in the

usage

is the

equivalent to the
here assigned
sense

dates back
to Rubner's
usage
of the replacement values of nutrients in 1882investigations
to

energy.

NUTRITION

234

OF

FARM

ANIMALS

In the
1885 and to his isodynamic values based upon them.
lightof the knowledge available at that time, this use of the
but as will appear
later (369),
it has
term
was
perhaps justified,
since been
feed is

part of the metabolizable

that

shown

virtuallyavailable

of the
energy
alone,while only

for heat

production
the remainder
be used for general body purposes.
If the
can
of the term
is to be continued,therefore,
available energy
use
it becomes
ability,
to
distinguish two
degrees of availnecessary
using,for instance,the term gross available energy as
and net available energy
equivalent to metabolizable
energy
able
which is availto signifythat part of the metabolizable
energy
for other

than heat production.

purposes
the writer's judgment, simplicityand clearness

In

will be

of tion
concept
he
discontinuing
altogether use of the

promoted by

available

and
metabolizable
employing the term
energy
perhaps fuel value provided the latter is understood

term

energy, or
with
the proper

to designate that portion of

restrictions,

of the feed
gross energy
the animal
organism.
for

Factors

324.

the

have

metabolizable
with

computed
Table

27.

capableof
energy.

of

diet

the

the

by

of

man

in

and

Rubner,

"

factors

use

of
be

may

degreeof accuracy.1

Metabolizable

for

transformation

proposed

energy

considerable

Factors

"

is

metabolizable

subsequentlyAtwater,
which

which

the

Energy

of

Human

Atwater

Rubner
Per

gram

Food

digested

nutrients

Per

total
gram
nutrients

Cals.

Protein

4.0

Carbohydrates

4.0

Fats

8.9

The

use

of these

results for carnivora.

to

the

factors

same

They

nutrients
digestible

have

yieldsapproximatelycorrect
been appliedalso
sometimes

of the feed of herbivora

but

without

sufficient warrant.
1

Compare

the writer's

Principlesof

Animal

Nutrition,

pp.

272-281.

THE

The
325.

of

kinetic

and

combustion

energy.

after these losses have

been met,

his

the whole

although on

at

each

described

in

in

work

and

drawing a load,
simply lifting

or

successive

step, a portion,

into kinetic energy.

The
in accordance
be measured

may

maining
re-

small percentage, of his total

relatively
is
i.e.,
energy is expended in moving objects,

of chemical

converted

energy

in part into mechanical

body

own

The

to

the metabolizable
i.e.,

performs work, whether

rider,operating a tread power

carrying a
the weight of

losses due

of the fuel.

animal

an

income

tible
feces,urine,combusoff chiefly unused
carry

comparable with

are

be converted
energy, may
in part into heat.

from the body

the illustration of the internal

to

recur

235

The

"

leakage or incompletecombustion

When

heat

excreta

they

motor,

NUTRITION

and

cutaneous

energy.1 To

chemical

OF

outgo of work

Outgo
gases

BALANCE

kinetic energy
thus produced
with the generalmethods

previous paragraph (313),usuallyby conversion

into gravitationenergy and measurement
in gravitationunits,
the gram meter, kilogram meter
foot pound.
i.e.,
or
326.

of

Outgo

heat.

experience teaches
of

and

men

the

total

that
be

outgo of heat

represents

of chemical

income

which

continuallytakingplace from

of animals

very

the bodies

considerable
It has

energy.

common

been

share of

computed

if the heat

produced by the average healthy man

from escapingfrom the body it would in

prevented

day

raise it to

of

month

is

The

"

pasteurizing
temperature,

the

at

approximatelyto
327.

Animal

the heat

that of

meltingcast
The

"

single
course

be raised

iron.

great variety of animal

have

been devised for the purpose

of measuring
of
production
livinganimals have been of three

general types, which

calorimeters

be

may

latent heat calorimeters

Water

rate, the temperature would

same

calorimeters.

calorimeters which

while in the

could

are

and

designatedas
emission

those

water

calorimeters,

calorimeters.

in which

the heat

is

imparted

known

quantity of water, the rise of temperature of which

is measured, i.e.,
of
they employ the first of the two methods
to

The

heat

which

in the next

they also

paragraph.

carry

off is included

in the total outgo

of heat

sidered
con-

236

NUTRITION

measuring heat
be

may
to

subdivided

heat

into

those

which

it is taken

of the

physicalstate
employed an

which

the

in

and

it to

calorimetric

ice calorimeter

effect

in his

has

been

but

calorimeters

at

be

heat

to

in

the

walls

not

to be

in

walls

remains

keepingthe
when

same

in the

first case,

as

is

is the

same

producing heat

unit of time

be

conditions

that the
so
resistance,
in

If, now,

external

radiation

animal

it is introduced

at

the

of the

them

with

between

for

heat

of

source

the

creasing
in-

the

of

temperature

second

heat,

first

one,

the
to

To
at

same

The

apparatus.

remove

water

vapor

as

the so-called

"

rate

same

as

the

that

that

and

was

rate

the

of
the

electric

the animal

produced by

measured.
indirectly

prevent

the

and

first case,

of heat

All

animal

must

calorimeters

used

necessarilybe provided

loss of heat

temperature

in the air current,

as

that

at

which

it

ventilatingair current, however,

from

of this water, of course,

of heat

the

in the

amount

is thus

ventilation.

tends

from

it is concluded

"

leaves

closed receptacle

established

the same,

respirationcalorimeter.
experiments of any length

with

of heat

source

temperature

substituted

as

The
for

measure

if it appears
that,
equilibriumis reached,the temperature of the walls is

an

heat

the

As

will be radiated

example,

sense

temperature, it will tend

constant

container.

constant.

for

animal,

lations
re-

type of

strict

directlyto

serve

rapidityuntil a balance is
and
heat
production and

radiation

the

of the

the

the rate of heat procompare

duction
but they may
be used indirectly
eter
principleof the emission calorim-

kept at

room

rises,however, heat

walls

an

Lavoisier

experimentswith

to

be illustrated as follows :
If a known
may
for example)be placedin
(an electric resistance,
located

the

to

quantities. The

measure

Thus

of

in

change

upon
production. This

said not

do

all,i.e.,
they

quantitiesof heat but only

by different sources,
to

method

little used.

calorimeters may

Emission

called flow

experiments

and heat
respiration
calorimeter,however, is not well suited
and

is imparted

of water.

substance.

between

animals

heat

those

current

eters
calorim-

of the second

use

viz.,causing

measurement,

Water

by

up

calorimeters make

heat

(309).

of water

stationarymass

ANIMALS

FARM

previouslydescribed

in
calorimeters,
Latent

OF

latent

the chamber

absorbs

heat of

and

the evaporation

correspondingamount
evaporation of water.
"

BALANCE

THE

OF

Either, therefore,evaporation
air in the chamber

the
more

course

also

include

combustible

34.

Dulong's

"

The

may
mass

thus

water,

introducing

carbon

animal

dioxid

of

perhaps

(Schaefer, Text

becomes

then

and

the

the

Book

and

Physiology).

combination

calorimeter

and

con-

oxygen

of

the

of

hence

piration
res-

has been

respirationcalorimeter.

of animal
to

serve

of water

various

calorimeter

apparatus

apparatus

source

excreted, and

water

apparatus
called

of

determinations

gases

The

sumed.

with

saturated

prevented by keeping

of water
conditions,or the amount
be determined.
must
ventilatingair current
is followed,it is a relatively
simple matter

in the

If the latter

Fig.

be

must

237

less abnormal

or

carried away
to

NUTRITION

used

by Dulong

heat, the

construction

illustrate the form

is used.

This

modifications,
notably
1

Smithsonian

2N.

Y. Med.

in 1822

in his

of which

of calorimeter

type

of
investigation
is shown

in which

of calorimeter

in the

United

Contributions
to Knowledge,
Jour., 49 (1889), 342.

has

States
No.

23

in
a

Fig. 34,

stationary

been

used

by Wood,1
(1

the

Ott

in
2

238

NUTRITION

OF

FARM

ANIMALS

ANIMALS

FARM

OF

NUTRITION

240

pensation
interestingform of emission calorimeter is the so-called comcalorimeter,in which the heat produced by the subjectis
balanced
against that produced, for example, by burning pure hydrogen

An

by

or

electrical resistance in

an

Calorimeters

have

of this type

been

preciselysimilar chamber.
described by Haldane, ' Bohr
a

recently by Tangl.3

and

328.
in the

Summary.

followingtabular

income

and

outgo

contained

energy

329.
upon

The

"

steer

the

energy

computed

also
may
balance.
In

work,

so

Table

28.

as

energy

for which

the metabolizable

energy
external

an

this
no

well

the

as

balance.

particularform

The

"

same

of

experiment

balance
and
nitrogen and carbon
(290,294, 322) have already been

to

serve

that

be summarized
may
showing the several items of

statement

of energy
in each.

of

Example

foregoing facts

illustrate the determination

experiment

the

energy

had

Energy

Balance

to

animal
be

of the

performed no

measured

in that

form.

1
2

"

Daily

Jour. Physiol. (London), 16 (1894),123.

Skand.
Arch. Physiol., 14 (1903),398.

of

Biochem.

Steer

Ztschr., 53 (1913),21.

BALANCE

THE

NUTRITION

OF

241

According to the conception of the schematic body (280),

of 608 Cals. was
that energy
to the extent
these figuresshow
stored up in the body as the chemical energy of either protein,
no
fat or glycogen. Assuming that there was
change in the
glycogen content of the animal, the nitrogenand carbon balance
showed
a
computed storage of 66.6 grams of protein and 15.2
of protein is
chemical energy
of fat (294). The average
grams
The
and that of fat 9.5 Cals. per gram.
5.7 Cals. per gram
of energy stored up in the fat and proteingainedby the
amounts
therefore be computed as follows :
steer can
In

protein 5.7 Cals.

In fat

66.6

9.5 Cals. X

15.2

Total
Found

from

balance

energy

found

by

that in this
appears
a direct determination

computed
within

It

carbon

524

Cals.

608

Cals.

experiment

the

gain of

energy

of energy
of the total amount
per cent
of the nitrogen
that determinations
is evident
0.3

balance

the other may

heat productionof

on

from

Cals.

balance and that

of the energy
carbon
of nitrogen and
agreed

balance

the

84 Cals.,or

involved.
and

from

144

84 Cals.

Difference

It thus

380 Cals.

the

on
serve
an

determinations

as

of the energy balance

check, and also that the

and

hand

one

mutual

be

computed quiteaccurately
of the nitrogen and carbon balances (indirect
animal

may

calorimetry.)

" 5.

Significance

of

Results

nent
played a very promiand agricultural
role in both physiological
investigation.
Having considered in the foregoingpages the generalmethods
of the balance
cance
experiment, a brief consideration of the signifiof the results obtained by their use as compared with those
Studies of the balance

reached
330.

of nutrition have

by other methods
Comparison

with

seems

called for.

metabolism

investigations.
"

The

used as an
with a steer
experiments like the one
illustration in previous paragraphs show, within the limits of
experimental error, the loss or the storage of chemical energy
results

of

NUTRITION

242

from the
resulting
in what
lost

energy

or

is
therefore,

OF

of

use

FARM

certain feed

contained.

gainedwas

or

ration and

mately
approxi-

fat,glycogen)the
(protein,

of material

kind

ANIMALS

adapted to determine

The

balance

experiment,

the total nutritive effect of

if the

given substance,while

comparativeslaughtertest be
regardedas a form of balance experiment(284)the particular
mined.
organs or tissues in which gain or loss took placecan be detera

balance

The

into the
however,affords no insight
experiment,
which
the
observed
tritive
nuby
is broughtabout.
For example,balance experiments

details of the chemical mechanism

result

have

fat and

that starch may

serve
the amount
quantitatively

demonstrated

have

shown

as

source

of

of fat formed

givenweight of starch. As appliedto known chemical

definite and of the
compounds,such a result as this is perfectly
information as to the
no
highestvalue,but it givesabsolutely

from

intermediate steps of fat

formation,either in the

processes

of

or metabolism.
resorption
digestion,

On the other

of the intermediary
olism,
metabhand,investigations
main

like those whose

have

Chapter V,
They have demonstrated
various
not

anabolisms

attemptedto

results have

been
necessarily
and

deal

to a

of the

some

katabolisms

been

outlined

in

largeextent qualitative.
steps through which the

occur,

but

as

rule have

with quantitative
directly
questions.1

Naturallythe foregoing
comparisonis neither comprehensivenor
exclusive. It aims simply to pointout a broad generaldistinction
which in reality
shade
between two types of nutrition investigation
into each other.

Balance

experimentshave

sometimes

been

characterized,

"
certain half contemptuous implication,
as
with the body." The characterization is a good

with

is unwarranted.
implication
of the balance
accurate

finished

of the
conduct

It is perfectly
true,as

experimentpointout, that,for

record of the income

productswould
operationsof
of such

an

of

raw

bookkeeping
but

one

critics

some

example,the most

materials

outgo of

and

of itself givea very incompletenotion

great factoryand that the successful

enterprise
requiresas

intimate

of the more
of some
For a summary
important of these methods
Dakin, Oxidations and Reductions in the Animal Body, Chapter III.
1

the

knowlcompare

THE

edge
and

BALANCE

NUTRITION

OF

243

possible of the functions of each separate machine

of the changes undergone by the materials submitted
to
as

its action.

even
fairlybe presumed, however, that these critics,
of
such a
fullest knowledge of the technical details
it as
conduct
business
to
a
factory,would hardly undertake
of the stock purchased and
enterprisewithout keeping account
the output realized,
i.e.,
exactly the sort of bookkeeping which
the balance experiment attempts for the animal
body. The
truth is that both types of investigation
are
equally necessary
of the other.
The balance
and each aids in the interpretation
experiment has been especiallyprominent in the past, while at
vestiga
to inpresent attention is being directed to a greater extent
of the intermediary metabolism, but neither can

It may
with the

The

331.
"

"

the other

to

say

balance

I have

no

experiment

alreadyindicated,the

As

need

of thee."
in

methods

agriculturalinvestigations.
of the balance

experiment

in

agriculturalinvestigations.
quite largelyapplied
have been made, in the majority of cases,
Such investigations
with single chemical
not
compounds, but with feeding stuffs
in such an
rations as a whole, and the effect observed
periment
exor
is obviously a summation
of the effects of all the
The
fore,
ingredientscontained in the feed consumed.
result,therethe total nutritive
while entirelyadequate to determine
material experimented with is less capable
effect of the particular
obtained
of generalizationthan
with a single chemical
one
compound like starch or fat,and from this point of view may
be regarded as being in a sense
even
empirical. A comprehensive
plies,
knowledge of the nutritive value of a feeding stuff imof the kinds and amounts
of chemical
a determination
first,
compounds contained in it and, second, a determination of the
exact
physiologicalfunctions of each. Obviously, however,
such a complete determination
of the nutritive value of any
of feeding stuffs is a work requiringa vast
considerable number
for
therefore,
expenditureof time and labor. One justification,
the
ance
of determiningsummarily by a balmethod
short-cut
experiment the effect of a feeding stuff or ration is that it

have

been

"

"

appears

data

possibleto

which

secure

in this way

be put to
feedingstuffs and rations.
can

within

practicaluse
Moreover,

in the
it is to

reasonable

time

comparison of
be anticipated

OF

NUTRITION

244

the conclusions

that
drawn

from

the

even

The

of

value

is far

feeding,however,
of

determination

although

the

from

logical
physioand

firmed
con-

experiment.

being

nutritive

of

stock

to

the

to

values

summary

feeding stuffs,

in that field.

III, the

in Part

relation

limited

service

important

material

checked

experiment in

total

rendered

it has

will become

As

balance

the

be

finallyto

of the balance

the methods

of any
chemical
and

elaborate

most

will need
investigations

by

ANIMALS

the nutritive value

to

as

FARM

feed

requirements
for animals for various purposes,
well as the generalphysioas
logical
of maintenance, growth,
laws governing the processes
milk production,the performance of work, etc., can
fattening,
be successfully
the aid of balance
studied only with
ments,
experiapparent

experiments are of
general scientific value independent of the particularfeeding
stuff used.
A strikingillustration of the importance of such
tained
is afforded
farm animals
on
investigations
by the results oband

the

results

and

Zuntz

by

energy
(365-370). The

of the feed

these animals
and

man

or

of the

marked

have
investigations
yielded important
physiology.
332.

should

Comparison

with

be

that

observed

based
do
energy
so-called

not

on

regards the
work

of

the

methods

determination

lation
assimibetween

to

make
"

digestion

subjects and

thus

contributions

to

the
parative
com-

Finally, it
feeding experiments

of exact

of the

garding
re-

character

served

have
"

practical experiments.

differ in their ultimate

"

differences

as

lattei

others

digestionand

of the so-called
the

and

Kellner

in the

carnivora

inconspicuous in

were

such

digestiveprocesses

factors

certain

prominent
which

and

in

associates,
by

expenditureof

the

of the feed

his

obtained

balance

"

and

of matter
from

logicalbasis

those

of

"

ure
practical experiments. In both cases, the measof the nutritive value of a feeding stuff,of the influence of
of the animal
as
a food
changed conditions,or of the efficiency

producer,is

the

in the

accuracy
determined.
The

as

measure

effect upon
the animal.
and
degree of detail with
reasons

of nutritive

(281-283),while

the

by changes in

weight of

effect is

the live

weight

than

more

the

considered

50

to measure

animal

lies

that

already been

effect have

that the attempt

the

difference

which

inadequacy of

experience of

demonstrated
sufficiently

effects

for the

The

or

by

has
years
nutritive
the gross

BALANCE

THE

yielded

product
each

other

or

Only
or

upon

the

fails
which
the

to

practice

laborious
of
those

of

give

permit

refinements
reveal

to

OF

stock

results
of

the

methods

NUTRITION

which

245

are

formulation
of

the

of

investigation

fundamental

laws

depends.

upon

with

general

balance

physiological

feeding

consistent

ciples.
prin-

experiment
can

which

be
the

relied
cessful
suc-

VII

CHAPTER

Significance.

333.

of feed

and

for

is liberated

energy

(201, 207)

animal

in

study

the

of

demands

fasting

opportunity

an

vital

digestion and

of

The

protoplasm.

pairment
im-

or

to
complicated
un-

processes

resorption

or

by

requirements of growth, fattening or reproduction.

qualitativeand quantitative knowledge of the expenditure

the
A

of energy

and

matter

important

an

of the

Fat.

is

fact

It

"

is

is

the

familiar

of

equally
active

and

familiar.

fail

lives

from

in which

the

as

well
nature

exact

material.

The

fasting

to

feed

nents
compo-

for the

or

perform

analysis of the
by the
of the

this

fed
insufficiently

mal
ani-

outgo

These

fully

been

firmed
con-

of well-fed

of balance
from

Obviously,

of fat.

reserve

carcasses

results

fore
be-

emaciation

have

observation

as

of

converse

extreme

its

body

the

that

their functions.

upon

is the

formation

and

The

stage of

largely

common

by comparative
animals

fuel

reach

may

tissues

of fasted

fat

that

conception

reserve

fasting animal

conclusions

of

supply

principal
yield energy

disposing of surplus feed,

fat, and

loses

the

All

"

katabolized

be

may

of

store

obviously

fasting organism.

method

body's
fat

katabolized.

body

of the

support

ascertaining the

is

purposes.

Substances

334.

fasting animal, then,

the

by

towards

step

for various

necessary

the

the

fundamental

the

by the functions

of

which

of material

lack

by

of rest, therefore, affords

state

of

integrity

the

water

by

processes

the

at

physiologicalactivities of the body

for a considerable
period, although, of

ultimately halted

are

of

metabolic

time

If sufficient

itself.

body

for

absence

the

continue

may

they

course,

of the

in the

that

supported

be

can

supplied, those

be

oxygen

fact

familiar

animal

materials

of the

expense

It is

"

life of the

the

KATABOLISM

FASTING

THE

the

and

experiments

body

been

has

determined.

Carbohydrates.
or

less

"

In

non-nitrogenous

addition
matter

to

fat

in the
249

the

body

form

of

contains

glycogen

more

in

the

ANIMALS

FARM

OF

NUTRITION

25"

this
days of fasting,
is indicated by
as
store of carbohydratesis also drawn
upon,
the fact that the respiratoryquotient tends to approach unity,
becomes
of glycogen katabolized
while later the amount
very
the

During

muscles.

liver and

first few

small.
Protein.

Balance

"

that

conclusion

the

chieflyof
is also

fat

togetherwith

continual

experiments, however, while confirming

the loss of tissue in fastingusuallyconsists
that there

show
carbohydrates,
of body proteinand

some

katabolism

sponding
corre-

the
urinary nitrogen. While
energy
expended by the fasting animal is derived chieflyfrom the
of non-nitrogenous material, the functional
breaking down
of a
the katabolism
activities of the body necessarilyinvolve
of

excretion

minimum

certain

of

amount

protein.

Finally,in addition to those groups of substances

eral
whose
katabolism
yieldsenergy to the body, the so-called minelements,or ash, of the body take part in the processes of
and
katabolism
are
continuouslyexcreted in the urine of the
Ash.

"

fastinganimal.
The foregoingfacts

well illustrated

are

Table

The

29.

results of a
average
fasted for from
two
to

No.

Omitting

one

"

follows

Average

of Inanition

Influence

Publication

as

were

77

number
seven

Katabolism

of

Fasting

in which

small

of
secutive
con-

Carnegie Institution

Men

of

(1907),pp. 456-464.
case

tigations
inves-

"

Metabolism

on

Benedict's

The

inanition.
upon
experiments in which men

days

by

gain of glycogen

was

observed.

Washington,

"

of the

functions

structural

Fasting

in

normally small.

katabolism

Protein

335.

251

Katabolism

Protein

The

i.

KATABOLISM

FASTING

THE

In

"

view

is of

proteins(264), it

of the
portance
im-

some

ment
inquirewhat proportion of the total energy requireis suppliedby these substances.
This aspect of the subject has been considered especially
by
to

E.

has

Voit,1 who

compiled and

the results of

discussed

siderable
con-

of
experiments upon fasting. While some
based on estimates,they are
his computations are
sufficiently
features of the fasting
the main
to outline definitely
accurate
katabolism.
They show that in what may be spoken of as the
normal
fastinganimal, in which the influence of the previous
feeding has disappearedand in which, on the other hand, the
has not been exhausted, the proteinkatabolism
fat reserve
plies
suptotal
of
the
small
transformed,
a rather
proportion
energy
in
the
the percentage with dogs, e.g., ranging
majorityof cases
of

number

between

and

10

17.

It is not
variable.
true,
Fasting protein katabolism
been looselystated on the basis of
however, as has sometimes
336.

"

C. Voit's

experiments (338), that

becomes

fastinganimal

tends

its amount

to

of

of

diminish

an

Benedict's

in

the total

urinarynitrogen upon

was

Table

30.

one

adequate

short time.
of

amount

of

On

body fat,

the progress

of

"

"

Katabolism

Protein

the

fasting. For
fastingexperiments (Table 29),
the several days of the experiment

with

example,

protein katabolism

within

constant

contrary, in the presence

the

Ztschr. Biol.,41

of

Fasting

(1901),167.

Man

"

Benedict

NUTRITION

252

Influence

337.

likewise showed
olism in

of

FARM

OF

body

ANIMALS

compilation (335)
protein to total katabindividuals,
considerablyas between
fat.

"

E.

Voit's

that the ratio of

clearly

fastingmay

vary
the relative amount

of fat contained
in the body.
depending
available as fuel,the amount
of
So long as body fat is readily
small,but if the animal is
proteinkatabolized remains relatively
of fat becomes
much
deficient in fat,or if its content
originally
reduced
more
proteinis katabolized to make up
during fasting,
on

for the

deficiency.

body is less than that of protein,

is relatively
while in fastingits exhaustion
more
rapid. There comes
when
the supply of non-nitrogenous material to the
a time, therefore,
this happens, the protein katabolism
tissues begins to flag. When
fuel material
beginsto increase ; that is,when the supply of reserve
Usually,the

store

of fat in the

low, the organism begins to

runs

use

more

of the

protein of

its tissues

that this occurs

of energy, and Voit 1 has shown
whenever
the ratio of fat to protein remaining in the body fallsbelow a certain

as

source

well nourished,this rise in the

originally
protein katabolism occurs
only shortlybefore death, from which it
mals
has received the name
premortal rise. In the case of very fat anithe
this point may
other hand, in a
be reached, while, on
never
increase steadilyfrom the
lean animal the proteinkatabolism
may
beginning of the fasting. The following three experiments
very
fat dog and a lean rabbit,cited by
a fat guinea pig,a medium
upon
Voit from Rubner's experiments,serve
to illustrate these three types
of fastingkatabolism.

limit.

Table

If the

31.

"

animal

Fasting

was

Protein

Katabolism

of

Fat,

Animals

Ztschr.

Biol.,41 (iqoi), 502.

Medium

and

Thin

338.

Influence

KATABOLISM

FASTING

THE

253

previous protein feeding.

of

experimentsof Carl Voit *

in
the proteinkatabolism

"

The

classic

fastingdogs have shown

the early days of fastingmay

upon

that

vary
consumed.
of proteinpreviously

widely according to the amount

the fasting follows a high protein ration, the protein
When
be relatively
the first day of fastingmay
katabolism
large,
on
which
is
falls to a comparativelylow level
but it soon
mately
approxithe

whatever

same

well illustrated

by the

animal, which have been

experiments
Table

32.

"

Protein

initial ration.

the

following results,all
fully confirmed by

Katabolism

of

Fasting

behavior

This

the

upon

"

same

sequent
sub-

numerous

Dog

is

Voit

Furthermore, the high proteinkatabolism which is observed

during the first two or three days of fastingafter high protein
feeding is accompanied by a relativelysmaller katabolism
of fat. Thus, in the first of the foregoingexperiments respiration
made
the second, fifth and eighth days with
trials were
on
the followingresults :
"

Ztschr.

Biol.,2 (1866), 307.

Computed

from

Voit's figuresfor

urea.

Table

33.

Total

"

FARM

OF

NUTRITION

254

ANIMALS

Katabolism

Fasting

oe

Dog

Protein
Urinary
Nitrogen

Fat

Grams

Second

day
Fifth day
Eighth day

Katabolism
Per
Total
Cent
of
Katabolism

Katabo-

lized

in

Grams

1.6

86

26.2

5-7

103

12.7

4-7

99

11.

of what
takes place in
Obviously, we have here the reverse
of fat for
the later days of fasting,viz.,a gradual substitution
the

proteinas

readilyavailable supply of

is reduced.

Doubtless

be

marked

stillmore
katabolism
339.

was

the

effect

equivalentto

Physiological

would

the first day of

on

minimum

the latter in the

28.1
of

have

been

when
fasting,
of

grams

body

found

to

the protein

nitrogen.
The

protein.
"

facts

corded
re-

in the

previousparagraphs render it evident that the

lowest
level of protein katabolism
is not
attained
necessarily
during complete fasting. Although the proteinkatabolism of
reaches a comparativelylow level which
animal
a fasting
soon
be greatly
changes but slowly,nevertheless its amount
may
of protein previously
affected,on the one hand by the amount
consumed, and on the other hand by the stock of non-nitrogenous
material
(fat and glycogen) contained in the body. While
to
normally some
10
17 per
in complete fasting
is derived

rise to twice

may

this amount

cent

from
on

of the

energy

metabolized

protein (335),the proportion

day followingheavy protein

feeding,or to almost 100 per cent in case of an animal whose

stock of body fat is exhausted.
it is evident that
In such cases
part of the protein is katabolized
ing
simply for the sake of supplysince the smaller

energy,

be called
are

at

latter
normal
The
amount

energy,

normal

amounts

katabolized

in what

of the
condition
average
all the vital
least sufficient to maintain
a

proceeding

or

for

considerable

time

in

may
animal
fasting

functions,the

substantially

manner.

protein katabolism
being so dependent on the
of
non-nitrogenous material available as a source
question naturallyarises whether by supplyingan

level of
of
the

256
be

NUTRITION

the

synthesizedby

FARM

OF

ANIMALS

fore,
very well be, therein the
protein katabolism

might

It

organism.
unavoidable

that the minimum

of

for certain
nitrogenousfeed is due to such a demand
acids or other groupings,and only in part or not at all
amino
of cell proteinsas a condition
of
to a necessary
breaking down
protoplasmic activity.Moreover, it is quite conceivable that
absence

both

of these views

be

may

true

; that

part of the

minimum

destruction
of cell
protein katabolism
represents a necessary
protoplasm in the performance of its functions,while the other
down
for the sake of securing
part represents protein broken
certain constituents
for specific
purposes.
There
will be occasion
these possibilities
further
to consider
in discussing
the proteinrequirement for maintenance
(398).

" 2.
341.
feed

Internal

and

variety
the

Of

Energy

work.

doing
former,

342.

the

is still

(632),while
various glandsare
existence

in the term

Measure

of

fastinganimal,
in the

mechanical

the

all the

in

are

undergo
may
s
ince
it
does
work
no
ultimately,

furnishes
or

of what
343.

recorded
in

measure

is often

Relative
in

"

and
fasting,

form

of internal

of heat.

The

determination

in

of the

expended

constancy

of energy

regardingthe
the way

nature

in which

dicated
in-

chemical

surroundings of

fastinganimal
energy
called the basal

work

In

energy
transformations,but

A
a

"

of
performed by means
the fats,carbohydrates

the

upon

work.

internal

forms

numerous

the

These

work.

previous paragraph

animal, it assumes
heat produced by

excretory

performance is essential
animal, may be conveniently

various

utilized

maintenance

due

derived from the katabolism

of
energy
and
in the tissues.
proteins contained
thus

of

work

muscular

secretory and

expended

energy

chemical.

and

the

great

typicalof the latter.

of the

internal

receivingno

carrying on

most

whose
bodily activities,

summarized

animal

an

prominent is the
respiration,together with

the continued

to

work

Fasting

in

of

body

tonus

activities of the
various

The

"

both
activities,

the

and

muscular

Katabolism

external

no

of internal

circulation
of

The

state

the

of the

of rest, therefore,

internal

in

work,

metabolism.
katabolism.

"

of the material

The

results

katabolized

fat,carbohydrates and protein

mutually replaceeach

may
one

other is

the

or

KATABOLISM

FASTING

THE

other

fuel material

as

available,render

most

according as

it evident

of the

in the katabolism

controllingfactor

257

that the

fastingbody is the

for energy for the performanceof the internal work

must
hardly have failed to suggest that this demand

demand
can

constant
relatively

in the

such is in fact the

That
number

individual under

same

has been

maintained

to

day

for the
more

seems

or

from

derived

The

constant.

requiresapproximatelythe

like conditions.

demonstrated

experiments. While not

the fastingkatabolism, expressedin terms
value in proportipnas
to approach a uniform
are

be

by a large
mathematically invariable,

case

of

conditions

and

same

quantityof

tends

of energy,

experimental
fasting organism
the

from

energy

day

of its necessary
internal work, but
less indifferent as
this energy
is
to whether

performance

the katabolism

of

fats,carbohydratesor proteins.

example, in Voit's experiment cited in the previoussection to

illustrate the interrelations of proteinand fat katabolism (Table 33),
the computed energy of the proteinand fat katabolized on each of
which
it
in the following table,from
the three days was
shown
as
when
that the total energy katabolism, especially
computed
appears
the
on
approximatelythe same
per kilogram of live weight, was
different days.
For

Table

34.

"

Energy

Katabolism

Live
Weight

Second

day
Fifth day
Eighth day

The

katabolism
likewise

344.

Fasting

Energy

Energy

from

from

Protein

Fat

proteinkatabolism

of Benedict's

Dog

Total
Energy

Cals.

Cals.

Cals.

32.87
31.67

289.3

816.9

1106.2

142.2

978.5

1120.7

33-66
35-38

3Q-54

117.

942.4

1059.6

34-70

have

been

fastingmen

results

fasting
as

gards
re-

(337)and

(334).

Energy expenditure in fasting a measure

In the fasting animal
in
requirement.
s

Cals.

of the

alreadyconsidered

investigations
upon

"

Total
Energy
Kg.
per
Live
Weight

Kgs.

thing is true of Rubner's determinations

of a rabbit,a dog and a guinea pig,whose

same

the

of

of
a

tenance
main-

state

of

258

NUTRITION

complete rest

and

activities

evidentlyreduced

with

are

animal

an

of life.

continuance

the

is

external

moderate

at

performed at

ANIMALS

FARM

OF

the

temperature, the vital

to

the

minimum

Since

the

internal
of the

expense

compatible
work

of such

chemical

energy-

tissues,the body's stock of energy is being

up
equivalent to the internal
constantly depleted by an amount
be made
done and this loss of energy
work
must
good from the
in its

stored

is to be maintained.

feed if the animal

of the

total katabolism

The

constant
relatively

fastinganimal, as expressed in

its heat

of the amount
of energy
production, is therefore the measure
vital activities of
expended in carrying on the fundamental
of the minimum
the body, and consequently
quantitywhich must

suppliedin

be

" 3.

Affecting

Conditions
Size

345.

and

matter

animal.

of

ration.

maintenance

produce

That

"

heat

more

Fasting

the

Katabolism

largeanimals
than

smaller

katabolize

ones,

and

more

therefore

periment
maintenance, needs no specialproof. Exshows, however, that the difference is not proportional
to size or
weight, but that small animals have a more
than
intense katabolism
large ones, its amount
being approximately
the
of
to
proportional
body surface,which,
course, is
relatively
greater in the smaller animal.
feed for

requiremore

The

relation

body surface
(citedby Rubner)

by Bergmann
2

Rubner

but

of the

to

seems

to

have

made

question,determining

the

been first suggested

to have
appears
in 1852 and later by Miintz * in 1878,
the first,
quantitativeinvestigation

fastingkatabolism

of six

dogs whose

kilograms.
the fasting
While not mathematically constant, the ratio between
and surface showed
katabolism
a close approximation to uniformity,
number
of investigat
and the same
fact has been verified by a considerable
cepted.
exceptions, and is now
generallyacalthough with some
Moreover, it has been shown 3 to be approximately true not
only of animals of the same
speciesbut of animals ranging in size

weights ranged

from

to

man

346.

is

Ann.

3E.

3 to

domestic

Computation

familiar

from

24

fowls
of

and

including also cold blooded

katabolism

fact that the

surfaces

Inst. Agron., Ill, p. 59.

Voit; Ztschr. Biol.,41 (1901),113.

per

unit

of

of solids of the
2

Ztschr.

animals.

surface.
same

"

It

shape,

Biol.,19 (1883),535.

of those which
i.e.,

material

the

that

error

it
slightly,

varies but

be

may

of animals

surfaces

body

tional
propor-

Since

their volumes.

of

powers

specificgravity of animals
without

259

similar figures,
are
geometrically

are

two-thirds

the

to

KATABOLISM

FASTING

THE

the
said

of the

shape are proportionalto the two-thirds powers of their

mula,
weights. This relation may be expressedby the followingforproposed by Meeh,1 in which W equals the weight in
same

which

The

of

value

the

factor

shape.

same

kWl

horse

the

for

constant

Moulton

Trowbridge,

is 9.02.

Hecker

of the

for all animals

is constant

k is

centimeters, and

in square

surface

the

grams,

as

reportedby

Haigh

and

termined
de-

have

cattle of
for 35 Hereford-Shorthorn
of fatness,
various ages from birth up and in various conditions
using the empty weight as a basis. Dividing the animals into
the

they found

groups

Table

Animals

animals

two

Values

"

of

"

Cattle

Beef

for

'
.

9.92

condition
old

18 months

years

values

followingaverage

in medium

animals

Fat
Fat

35.

thin animals

and

Young

of

value

the

old

or

or

9.41

less

8.57

7.65

more

investigatorsthe empty weight of cattle

constitutes the followingpercentages of their live weight :
According

to

these

"

Table

36.

the

aid

Live

92~94

Weight
per cent

91

89-90
87-89

foregoing factors

of the

cattle,and

reasonable

be

accuracy.

perhaps of
computed
It is

other

types,

unit

per

total katabolism

the

of

as

determined

body surface

Ztschr.

live

Biol.,15 (1879), 425.

by
with

apparent that comparisons based

weight instead of the empty weight would

cattle.
for thin and
medium
substantiallyaccurate
the

similar data exist for other

1

of

cattle

experiment may

be

Percentage

cattle

Thin

upon

as

cattle

Fat

Medium

of beef

Weight

cattle

Show

With

Empty

"

speciesof
2

farm

Mo.

also
No

animals.

Expt. Sta., Research

Bui.

18.

260

NUTRITION

The

be the

variation

shown

has

for k

cattle is somewhat

more

the two-thirds

to

five-ninths
values

347.

gives the

of

Computation
desirable

is often

weights
be made

the

been

and

the katabolism

surface

determinations

of

none

of

the

of different
to

some

rather

the

than

body

with

of

dently
evi-

body

surface

of

exception of
farm animals, so that it is

on

the

the

katabolism

of the

of

comparison
type, however,

species and

same

results

It

"

comparisons should

in many
to express
cases
impracticable
latter per unit of surface.
For purposes

individuals

weight.

of animals

experimental

weight. Such
basis of body

cattle

corresponding

standard

to

made, howTever,and

of beef

the

13.40.

katabolism

compute

actual

and

the horse

to

on

Few

have

animals

closest agreement,

compare

standard

convenient

weight.

to

or

the

by making

for fat cattle the

weight,while

11.86
being respectively

of k

eliminated

basis of

of the empty

power

power

if this be

computation, an average value of

very closelyapproximating those actually
finds that the body surface of thin and medium
than
closelyproportional to the five-eighths

likewise

He

observed.

the groups of cattle

of
proportion fat to active tissue,

in the
that

weight the
gives results

fat-free empty
10.34

ANIMALS

of the difference between

principalcause

to
appears
and Moulton

FARM

OF

between
least

at

approximate results may be secured on the assumption that

the animals
to be compared are
geometricallysimilar,so that
their bpdy surfaces are
substantiallyproportionalto the twothirds powers

of their

1283 pounds was found

It
(374) of 8671 Cals.
such

result to

other

to

between

the surfaces
and

1000*:

1283%

would

therefore

this way

have

is often

weight

of

pounds,
katabolism
in proportion
1000

For

weights.

example, a steer weighing

computed fastingkatabolism
of convenience

matter

pounds.

1000

thingsbeingequal,would
to

of the two

its smaller

animals

the

fastingkatabolism

be

8671

it is easy

to

individuals

Cals.

the

compute

without

the

be

steer

have

surface.

would

to

pute
com-

ing
weighsmaller

ratio

The

approximately

of the smaller

(f!nrf)!7345
=

relative katabolism

animal

Cals.

In

ferent
of dif-

necessity of expressingit

per

unit of surface.

Of course,
as
particular,

not

of the

same

such

comparison is only an approximation. In

has just been shown
(346),different animals are
shape. The young animal differs in conformaa

Jour. Biol. Chem.,

24

(1916),299.

FASTING

THE

tion from

the older

beef

and

steer

similar.

the fat from

e.g.,

are

and

thin

and

one

the

being geometrically

of the

species,types

be of much

the

far from

determinations

in different

would
of

dairy cow,

Additional

weight

to

and

one

261

KATABOLISM

relation

ages

of

surface

of domestic

interest but in their absence

mals
ani-

the method

to give
probably be assumed
and is certainlymore
accurate
a
than a simple computation in proportion to weight.
Muscular
348.
activity. As was implied in the introductory
in greater detail
section of Chapter VI (274),and as will appear
in Chapter XIV, muscular
work is done at the expense of energy
derived from the katabolism
of body substance, and
other
no
The
singlefactor so largelyinfluences the total katabolism.

comparison just outlined

may
fair approximationto the truth

"

represents the demands

minimum

which
fastingkatabolism
life processes
is
indispensable

the

complete muscular
when
animal, even
more

or

rest.
at

rest

of

only in a state of
It is rarelythe case, however, that an
in the ordinary sense, does not
tain
main-

less muscular

exhibited

tension

or

execute

more

or

less

tions
mo-

of various

when
parts of the body, all of which, even
penditure
apparently slight,involve in the aggregate considerable exof energy.
Zuntz

and

Hagemann,1
in which

horse

a
upon
flies in the

in the

chamber

for

example, report

the uneasiness
of the

metabolism.

caused

experiment
respiration
of a few
the
by
presence
a

caused

apparatus
2

found

an

increase

the

of

10

excretion

per
of

hourly
Johansson
when
man
fasting
simply lying in bed (awake)
by
to be 24.94 grams
as
compared with 20.72 grams when all the muscles
relaxed as possible. Benedict
and Carpenter 3 have
as perfectly
were
of men
during sleepwith that of the same
compared the metabolism
subjectslyingquietlyin bed immediately after waking. In the three
which
cases
they regard as strictly
comparable the increase in the
heat production during the waking period ranged from
5.8 to 15.2
Benedict
and Talbot,4in experiments
per cent, averaging11. 4 per cent.
muscular
f
ound
that
noticeable
even
infants,
activity
scarcely
upon
marked
effect
the
dioxid
carbon
on
produced a most
excretion,and
Benedict
and Pratt 5 have noted
similar results with dogs.
cent

dioxid

carbon

1
2
3
4

Landw.

Jahrb., 23 (1894),161.
Skand.
Arch. Physiol.,8 (1898), 85.
Carnegie Institution of Washington, Publication
Amer.
Jour. Diseases of Children, 4 (1912), 129.
Jour. Biol. Chem., 15 (1913), 1.

No.

126

(1910),p.

241.

262

OF

NUTRITION

ANIMALS

FARM

have
such
movements
a
comparatively insignificant
strikingeffect upon the total katabolism, it is evident that the
be an
of muscular
amount
activitymust
important factor in
determining the relative energy requirements of two animals
katabolism
in a state of absolute
even
though their minimum
be identical.
In experiments of any
considerable
rest may
duration
normal
or
on
animals, it is impossible to avoid more
in this incidental muscular
less expenditureof energy
work,

Since

periodsof
349.
is

the
to make
difficulty
experimentcomparable in this respect.

it is often

while

an

and

Standing
the

Considerable

lying.
"

the

requiredduring

position of

of

matter

hours

waking

members

different

to

of

muscular

maintain

the

different

exertion

the

relative

This

body.

is

pecially
es-

or
an
standing. It has been shown that a man
carbon
when
dioxid
animal
notably more
standing excretes
than when
restingor lyingdown and produces correspondingly

true

observed

of

Differences

heat.

more

of

in

and

man

much

as

of 30 to 40

as

25

per

have

cent

been

It
per cent, in cattle.
lies down
for twelve hours

or

more

evident,then, that if one animal

and another
for only eight hours during the twenty-four, the
former
will,other thingsbeing equal, requireless feed energy
is

for actual
350.

than

maintenance

External

the latter.

temperature.

generalclass known as warm

whose
bodies during health

"

Farm

animals

belong to

that

blooded,or homoiothermic, animals,

maintain

nearly constant

ture
tempera-

surroundings. The
heat
is being continuallygeneratedby the
animal
so-called
katabolism
going on in the body, while on the other hand the
animal
is continuallyimpartingheat to its surroundings in four
principal
tion
ways : viz.,by conduction, by radiation,by evaporawhich

"

"

of water, and
Since
the

that of their usual

is higherthan

as

the animal
maintenance

the existence

of

some

the sensible

heat

of the excreta.

ally,
producing and losingheat continuof a constant
body temperature implies
of which
regulativemechanism
by means

is both

the

production and emission of heat may be adjustedto each

other.
This adjustment is effected in generalin two ways which
be called,
respectively,
physicaland chemical regulation.
may
351. Physical regulation of body
Changes
temperature.
in the temperature of its surroundings,in the relative humidity
of the air,etc.,tend to produce the same
effect upon
the animal
"

264

NUTRITION

of the

OF

FARM

ANIMALS

be effected
can
entirelyclosed,nothing more
toward
in this manner
maintaining its temperature, and if the
weather
continues
be
to grow
must
colder,the fire in the room
increased.
Similarly,if the external cooling effect upon the
animal
becomes
the limits of physical adjustment,
to exceed
so great as
fuel material
is katabolized,that is,more
heat
more
of maintaining the body temperature
is produced. This method
is commonly called chemical
regulation.
are

room

Mechanism

353.

of

chemical

The

regulation.
"

chemical

is probably effected largelythrough muscular

action,
regulation
or
by an increase in the muscular
by visible motion
tonus,
increased
involves
heat production. This
either of which
an
has been clearlyshown
and probablyapplies
to be true of man
also

other

to

Above

animals.

be

the
in

slightincrease
risingtemperature, probably due
to

appears

in the various
354.

Critical temperature.

Above

of

temperature

for

50 C.
3o"-35",and

; for the

about

and

of Von

man

for

the

der Heide

varied

production with

additional

energy

pended
ex-

physicalregulation.

or

temperature

begins to

be

at

which

the

supplemented

called the critical ture.1

temperathe radiatingcapacity of the

the

regulationis

productionis

there

been

this temperature
is varied to meet

body surface
it,this method
the heat

to

heat

the

The

"

physicalregulationgives way
by the chemical regulationhas

the

to

of

processes

critical temperature,

varying conditions ; below

largelyexhausted and therefore

to

the

meet

need.

The

critical

wearing ordinaryclothingappears to be
dog it is about 200 C, for the guinea pig
hog, accordingto the results of Tangl 2
and Klein,3about 2o"-23"C.

tate
Any conditions tendingto facilithe escape of heat from the body will obviously act like a fall of
Wind, for example, by removing the layerof partially
temperature.
warmed
air next
heat more
to remove
to the skin, tends
rapidlyfrom
the body, so that the cold is felt more
severelyon a windy day, while,
is modified
the other hand, the effect of a high temperature
on
by
wind.
A high percentage humidity of the air on a warm
day hinders
355.

Other

the removal
than

that
2

The

term

of heat

dry

heat.

refers,of

of the animal
Biochem.

thermal

conditions.

"

by evaporation, so that a
Cold moist air,on
the
course,

to

the

moist
other

heat

is

more
ing
tryfacilitates
hand,

temperature of the surroundings and

itself.

Ztschr., 44 (1912), 252.

Ibid.,55 (1913),195.

not

to

FASTING

THE

of heat

the escape

clothing,hair

of the
a

dry

The

cold.

direct

thermal

that

cold is

damp

of the

rays

sun

more

impart

may

than

severe
a

considerable

peratures
temmoderating
those
of
hand, increasing
and, on
high temperatures.
of
should
critical
be strictly
one
a
then,
speak
accurate,

body,

the

To

body by increasingthe conducting power

fur,so

or

to the

of heat

amount

the

from

265

KATABOLISM

thus

the effects of low

other

of the

environment

animal

than

rather

simply

of

critical

temperature.
Influence

356.

is apparent from
the
katabolism of the fastinganimal

katabolism.

on

It

"

foregoingfacts that the energy

is affected by the external temperature and other thermal conditions
than *has been frequently
to a considerablyless extent
that every fall in external
true
imagined. It is by no means
temperature results in
for the sake of heat
would

increased

an

for
production,

; that

should

is,we

fact that

external

the katabolism

have

life without

the

the

case

verse
con-

the

productionof

heat

it either remains

unchanged
determined
by

is not

temperature
would

was

be reduced

katabolism,which

in terms.

at the critical temperature and

minimum

extent

the

that

is a contradiction
The

were

animal

true, viz.,that every rise in the external

decrease in the katabolism,
cause
a corresponding

finally,when
equivalentto that of the body,
to zero

if this

in the

arlso be

temperature would
so

katabolism

or

animal

that above

increases

the needs

an

reaches
that

shows
slightly

of the

point

that its

organism for

heat

such, since these diminish as the temperature rises. As a

of fact,the productionof heat in the body is not the
matter
but merely an
incident of it. Heat
is
purpose of katabolism
as

the form

which

the chemical

energy

of the katabolized

material

takes after it has served its purposes in the vital processes, and
the nearly constant
heat production above
the critical temperature
is

simply

requiredfor
constant

and

temperature.
of.

due

to

the fact that

the internal work

cannot

Heat

of the

be reduced
is

simply by raisingthe external

excretum
to be gotten rid
an
essentially

Incidentally,in warm-blooded

maintain

the

body temperature

of the vital

quantityof energy
body is approximately
the

animals, it
necessary

serves

also

for the normal

to

formance
per-

functions,but above the critical temperature

there is a surplus over
the amount
required for this
which is disposedof by the processes of physicalregupurpose

266

described.

already

lation

sinks

therefore,

energy

requirement.

body

within

At

low

it

as

actual

is

of

the

animal

body

At

the

the

heat

stimulate

very
overcome

to

low

high

rises.

perature
tem-

processes
below

only

this

the

influence

The

of

regulation

"

foregoing

is

paragraph

temperatures

possible

heat.

and
which

which

case

of

about

the

as

fall.

to

An

the

sity
inten-

cold-blooded

fall

further

heat

in

production,

cold.

from

reverse

the

heat

reduces

diminishes

process
are

however,

therefore
is

begins

bringing

rise,

regulation

produce

to

the

in

regulation

slight

very

and

does

perishes

physical

unable

however,

further

temperatures

circle"

it

the

chemical

of

temperature

sinks,

speedily

is

body

as

again

of

possibilities

temperature,

production

katabolism

"vicious

the

exactly

possibilities

by

and

temperature

animal

the

body

which

the

the

animal

temperature

external

katabolic

produced,

the

the

and

katabolism

very

When

rise

of

temperature

that

the

temperature.

in

that

so

abstracted

the

of

temperatures

lowering

that

the

limits.

exhausted,

be

may

when

external

described

certain

very

the

is

ANIMALS

only

point

extremes

temperature

only

fast

more

of

Effects

is

heat

does

point,

357.

this

and

FARM

It

below

stimulated

are

so

OF

NUTRITION

the

speedily

has

leads

production,
that

to

the

body

shown

been

of

place.

the

exhausted,

heat

converse

take

may

to

giving

occurring
animal

at

being

CHAPTER

MAINTENANCE

materials
him

by

But

for human

the

REQUIREMENTS

"

Feed

is

they

may

yield products

food

and

clothingor

performance

much

ENERGY

of maintenance.

that

either

animals

THE

"

Definition

358.

VIII

of mechanical

that

supplied to farm
useful

they

to

may

man

as
serve

work.

first be

supplied with enough

the shafting,
to keep in motion
belting and machinery
power
in generalbefore any product can
be turned out, so the animal
be provided with sufficient feed to maintain
mechanism
must
essential to life before any continued
the processes
production
is called
is possible. The
amount
required for this purpose
ration
of the particular animal.
the maintenance
It is the
quantity necessary
simply to support the animal when doing
work and yieldingno material product. A balance experino
ment
with

as

factory must

animal

ration
receivingpreciselya maintenance
reveal an
would
exact
and outgo of
equalitybetween income
ash, nitrogen,carbon, hydrogen and energy, showing that the
body was neither gaining nor losingprotein,fat,carbohydrates
an

mineral

elements.

this

From

of

view, maintenance
of labile equilibrium between
as
a state
might be characterized
of metabolism
the anabolic and katabolic processes
(203).

nor

The

word

example, by

signifythe

means

total

horse in order

calf in order to make

the idea

is sometimes

maintenance
to

sense

normal

to

minimum

total feed of the horse

used
of

amount

perform his

growth.

that, in its technical

the

point

sense,

requiredsimply

other
popularlyin anfeed required,for
dailywork or by a

It is important to grasp

the
to

maintenance
maintain

ration

life. The

would, from this point of view, be

tenance
regarded as consistingof two portions; one of them the mainration,which if fed by itself would just support the
horse at rest or the calf without
growth, and the other the
is
work
of which
productive portion of the ration,by means
or

calf

267

268

NUTRITION

done

growth

or

the

running,
to

made.

To

while

the

the finished

out

Significance of

might
to

first

at

seem

the

the illustration of the factory,

to

recur

additional

ANIMALS

FARM

ration

maintenance

turn

359.

It

OF

keeps
feed

the

machinery

empty

furnishes

the

sary
neces-

power

product.
maintenance

thought
of

determination

that
the

ration
not

much

in

practice.
"

importance

maintenance

ration.

taches
at-

The

return
kept on such a ration yieldsno direct economic
and hence
simple maintenance
feedingis to be avoided, so far
while if it appears
desirable to practiceit the observatio
as
possible,
of the skilled stockman, especially
if supplemented
by occasional weighings,will usuallysuffice to determine whether
the end is being attained.
not
or
Nevertheless,the subjecthas
both for practiceand for science.
much
significance
fraction of the feed actually consumed
A very considerable
the average
on
by farm animals
probably fully one-half

animal

"

"

requiredsimply for
feed bill is expended

is

for
its

maintenance.

But

if half of the farmer's

for

maintenance, it is clearlyimportant
how
him
to know
something of the laws of maintenance,
different animals, how
requirements vary as between
they
"

affected

are

how

different

etc.,
"

as

by

the

conditions

under

which

animals

are

kept,

feeding stuffs

well

as

to

in value for maintenance,

compare
understand
the principlesgoverning the

productionof meat, milk,

or

work

from

the

other

half of his

feed.
360.

Bearing

on

interpretation of

feeding

experiments.

"

point of view of the experimenter a knowledge of

the maintenance
requirement is likewise of great importance.
rational study of the laws of nutrition,it is plainly
In any
inadmissible
to attempt
to establish
general principlesby a
viz.,production,
comparison of the feed with one of its effects,
From

the

while

ignoring entirely its other effect,viz., maintenance.

ing
misleadFailure to appreciate this fact is responsiblefor many
deductions
from feedingexperiments in the past.
the results of such experiIt has been quite usual to compare
ments
the
feed
consumed
ratio
of
to
product
by computing
yielded i.e.,either the feed consumed
per pound of gain
made
the gain produced per pound of feed consumed.
Such
or
a
comparison, however, may give an entirelydistorted idea of
the real teachings of an
experiment. Suppose, for example,
"

MAINTENANCE
that in

REQUIREMENTS

ENERGY

-THE

of
fatteningexperiment the quantities

rations consumed

and

gainsmade

the

were

First

Daily

Ration

21.0

i.olb.

gain

:-

Second

18.0 lb.

Feedeaten

different

two

follows

as

269

Ration

lb.

1.5 lb-

to proCompared in the way justindicated,the feed required

duce
and
one
pound of gainwas 18 pounds
14 poundsrespectively

the second ration appears to have been superior to the first

bv about 29 per cent.
If,however, it was found that in each
requiredsimply to maintain
case
12
pounds of the feed were

or

animal, a

the

comparisonis obtained,viz.

different

very

Second

Fikst
Ration

Ration
Lb.

Lb.

Feedeaten

Expended in maintenance
"

Surplusleft for production

Daily gain

pound

have

been

the

second
to

the
was

the fact that

to

of

more

has
in

to
principle
applies

recentlybeen shown in a
experiments upon dairy

for

Mo.

of
and

the side

obtained,not with

all

very

any

more

than
efficiently

Clearly,a

maintenance.

advantage :
be added

that

feeding as
productive

strikingmanner

cows.

the

that the second

of

cases

on

show

by Eckles
knowledge of

logical
interpretation
-

""j;""
tr"
to
Corresponding

the dual function of the feed

bodily activities and

growth,maintenance

to

not

ration is essentialto any

results.
of experimental

Requirements

on

different animals

the maintenance

361.

6.0

two

kind of

same

6.0

been

with

but

i'"

have

the economic
simply demonstrates
largerconsumption of feed. It scarcelyneed

the

I2
"

is thus shown

but
the first,
a

I20

higher nutritive value but

if the
it was
eaten.
Similarly,

does
feed,the experiment
assimilatedhis feed
animal digested
or

same

2I-0

any

supposedto

different rations

pound

economic advantage

due

not

be

rations per

two

The

same.

oregoingresults
two

of gain

real value of the

simply

18.0

"""""""

Surplusfeed per
The

(263) as a source of energy for he

for the
substances necessary
specific
repairof the tissues, the maintenance

Exp. Sta., Research

Bui. No.

2.

requirements of

the animal

rations

purpose

view:

for that

and

the values

be

may

ANIMALS

of

considered

feedingstuffs

from

and

points of

two

"

inquirehow much energy is

quiescentanimal and what amounts
rations can
supply in forms available

First,we
the
feeds and

may

Second,
of each
due

FARM

OF

NUTRITION

270

ask what
may
be suppliedin the feed

we

continual

the

which

to be considered

need

to

of

katabolism

rather
the proteins,
or
particularly
and
them, and the ash ingredients
mines

port
supvarious

for this purpose.

specificmaterials and how much

must

to

to

necessary
of it the

make

the

good

losses

substances.

body

acids

the amino

perhaps the

composing

so-called

in this respect, the

vita-

body

other

of

is

It

parently
ap-

manufacturing
possessinglargepowers
from
those
suppliedin ordinaryfeeding stuffs.
ingredients
these two
It will be convenient
to consider
generalclasses of
maintenance
requirements in the order named, the present
chapter dealingwith the energy requirements.
362.

Mutual

replacement

cussion$in Chapter V,

of

organic

of the functions

nutrients.

of the

sary
neces-

The

"

dis-

principalgroups

that, besides certain

(262-267)showed
organic nutrients
values
of particularchemical
as
sources
compounds,
specific
they all serve as carriers of chemical energy for the needs of the
organism. It would be anticipated,therefore,that the various
digestiblenutrients might mutually replaceeach other or the
of the body, and numerous
experimentshave shown
ingredients
of

that such
Fats

is indeed

fed

the loss of

to

the

case.

diminish

previouslyfastinganimal
a

similar

result.

As

has

suspend

be substituted

body fat,while carbohydrates may

the feed fat with

or

for

shown

alreadybeen

(337),body protein may replacebody fat in the katabolism of

mal
the fasting
animal, while when proteinis given to such an anithe non-nitrogenousportion of the molecule
as
a
serves
for body
of energy to the organismand can
be substituted
source
fat.

On

the

other

hand,

an

excess

of feed

the

protein above

by
requirementcan be replacedby fats or particularly
and likewise by organicacids.
carbohydrates,

minimum

the
In
of

brief,the

animal

organism manifests
of the
as
regards the nature
flexibility

remarkable

material

degree

which

the small
utilize for its energy
Aside
from
metabolism.
of protein required,
activities of the
the metabolic

it

can

imum
min-

body

OF

NUTRITION

272
olizable

of the
energy
determined
in the manner

were

follows

as

Table

37.

and

hay

ANIMALS

the

illustrated

Net

of

Hay

Period

Difference per lb. dry

LIZABLE

Eaten

Energy

Pro-

Gain
duced
Energy

of Period
of the

this amount

animal

9812

6.17

5768

8064

-2296

4.04

3776

1748

2028

433

502

2028

hay, therefore,computed
in

dog

exactlyas

that

to

energy

The
in

of the

energy
steer.

Only

268

hay

502

could

Cals.

were

no

be

means

energy

from

the

effect of the

supposed

2028

added

-s- 4.04

hay

case

of

502
contained
=

Cals. per pound of

all of the metabolizable

utilized
used

the insufficient

net

the

of 935

the amount

Clearly,therefore,
by

matter.

Cals.

is,they contributed

the

per
metabolizable

to

loss of

preceding paragraph, was

the
But
pound of dry matter.

Cals.

by the
place

the

Cals. ;
its maintenance.

by

towards

9544

diminished

III

of

Cals.

added
hay (water-free),

of

Hay

of

matter

pounds

4.04

ration

dry

Heat

935

The

Timothy

of

Cals.

hay

the

Value

Metabo-

10.21

Difference

body

in

of

Pounds

production per day,

Chapter VI (322,329),

Energy

ter
Mat-

Dry

Period

heat

"

Determination

"

FARM

for

for

maintenance

this purpose,

in

from
the katabolism
of
previously derived
the fat and protein of the body, while the remaining 433 Cals.
in the body, but resulted simply in
indeed, metabolized
were,
The
increasingthe heat production by this amount.2
tion
proporof the metabolizable
which
available
of
this
was
hay
energy
for maintenance, then, was
The
going
fore502 -r 935 =53-7%result is typicalof a large number
have
of others which
reached
in experiments on
various
been
species of animals.
1

Since

of

energy

of course
fed, the gains were
negative,
chemical
lost from the body.
was
i.e.,
energy
2 Since
and
income
gains of energy
are
computed from the difference between
of the table necessarily agree with those of the
outgo, the figuresof the last column
facts.
two
preceding ones.
They simply present a different aspect of the same

submaintenance

rations

were

MAINTENANCE

Even
ora,

ENERGY

THE

"

REQUIREMENTS

273

of pure, or nearly pure, nutrients fed to carnivof

values are
less than their content
their maintenance
in the

case

metabolizable

energy.

Feed

365.

heat

increases

consumption

From

production.
"

experiment just cited

furnishes a good illustration of the important fact that the
consumption of feed tends to increase the heat production of
sier.
the body. This is an observation as old as the time of Lavoithe oxygen
That investigatorobserved
consumption of
to increase materially
(about 37 per cent) after a meal,
a man
vestigat
inof subsequent experiments by numerous
and a multitude
and on various speciesof animals have
firmed
fullyconincreased metabolism
these earlier results,
so that the fact of an
consequent on the ingestionof feed is fully established.
of Zuntz and his associates l
It is especially
to the investigations
of this fact and the recognitionof its
that the demonstration
in relation to the nutritive values of feeding stuffs
significance
a

different point
slightly

of

view,

the

is due.
of
example, Zuntz and Hageman,2 on the average of a number
experiments in which the respiratoryexchange of a horse shortly
hours
and some
before feedingin the morning, shortlyafter feeding,
of the Zuntz
form of the Pettenkofer
determined
later was
by means
gram
the followingresults,
computed per kiloapparatus (299),obtained
live weight per minute.
For

Table

38.

Heat

"

Production

by

Horse

Computed
Heat
Pro-

Oxygen

duction

Consumed

Cubic
Centimeters

Gram-calories

Fasting

3-339

16.929

after feeding
23 minutes
3^ hours after feeding

3.648

18.510
18.787

3-704

The

effect has been

same

by Armsby
directlyby
1

means

Fries

in which

of the

the heat

with

cattle in

production

calorimeter.
respiration

the writer's Principles

of Animal
Nutrition,
Jahrb., 27 (1898), Erganzbd. Ill, 282.
Jour. Agr. Research, 3 (1915), 435.

Compare

2Landw.
3

and

invariablyobserved

pp.

was

Thus

ments
experimined
deterin

377_385-

an

to

the

in different

steer

same

Table

39.

periodsthe results
Production

Heat

"

of alfalfa

different amounts

three

experiment in which

ANIMALS

FARM

OF

NUTRITION

274

by

as

were

Dry
of

hay

fed

were

follows

"

Steer

Heat

Matter
Feed
per
Day

duced
Proper

Day

Cals.

Period

11272

10388

Period 3
Period 5

7754

respirationexperiments on fattening cattle have shown

feed is added
effect is produced when
to a basal ration.
that the same
added
wheat
to a lightfattening ration 1 the
when
Thus
gluten was
the balance of matter
heat production as calculated (329) from
was
Kellner's

as

follows

Table

40.

Heat

"

Production

by

Ox

an

Dry
of

Heat

Matter
Feed
per
Day

duced
Proper

Day

Cals.

Period

1, Basal

Period 4, Same

I934I

24007

gluten

taught,in
"isodynamic replacement," that
For

of

ration

it was

years

many

the nutrients
with man,
of metabolizable
energy.
as

well

(367e),have
horses and

as

were

Rubner's
recent

of

true

that if the experiment be made

animal

there is in each

the metabolizable
Thus
was

the average
fed gave the
1

Landw.

Die

case

an

in
own

ones

is true

that what

cattle is also

with

of value

still more

shown

accordance

with

Rubner's

carnivora, and

theory
presumably

proportion to their content

later investigations,2
ever,
howby

of the

nearlypure

Lusk

and

his

feedingstuffs
nutrients

associates

consumed

fed to

by

dogs, viz.,

the critical temperature for the

increase in the heat production,so that
above

is

available for maintenance.

only partially
of two
of Rubner's
experimentsin which lean meat
followingresults as compared with the fastingstate
energy

Vers.

Gesetze

Stat.,53 (1900),130-131.

des Energieverbrauchs

bei der Ernahrung,

1902.

MAINTENANCE

"

Table

41.

"

THE

ENERGY

Heat

Production

REQUIREMENTS

by

275

Dog

Daily
Heat
Production
gram
Kiloper
Live
Weight

Cals.

Fasting

5I-50

Fed

70.55

efficient in stimulatingthe heat productionbut

especially
effect although to a much
fats and carbohydratesproduce the same
less degree.

Proteins

are

The

366.

specificdynamic action.

"

The

effect of the various

nutrients,notably of protein,in raisingthe heat production of

the animal above the fastinglevel,as in the experiments just
their specific
cited,has been called by Rubner
dynamic action.
1
divides
Kellner
has proposed a different terminology. He
of the feed into thermic and dynamic
the metabolizable
energy
Thermic
equivalent to Rubner's
specific
energy,
energy.
dynamic action, signifiesthat portion of the metabolizable
is of value
which
to the organism only as
a
source
energy
of heat.
Dynamic energy, equivalent to net energy as defined
in a subsequent paragraph (370),on
the other hand, is that

portion of the metabolizable energy which

the performance of the vital functions.
of increased

Causes

367.

of feed

sets

manifested

not
a.

Mechanical

heat

can

production.

"

be

The

utilized for

tion
consump-

operation (orincreases)a varietyof activities

by the fastingorganism.
in

Work.

"

not

inconsiderable

amount

of

activityis expended by farm animals and especially

by the herbivora in the prehension and mastication of their
in moving it through the alimentary canal.
feed and
Since
work
muscular
involves an expenditure of energy, all of which,
in the case
of internal work, finally
takes the form of heat (342),
the mechanical
of digestionis a considerable
factor in
work
of farm animals, although Armsby
increasingthe metabolism
and Fries2 have presentedreasons
for believingthat peristalsis
muscular

1
2

Ernahrung landw. Nutzt., 6th Ed.,

Jour. Agr. Research, 3 (1915), 479.

p. 105.

276

OF

NUTRITION

in cattle does

not

contribute

FARM

ANIMALS

very largelyto the increased

the consumption of feed.

heat

production consequent on
b. Glandular
required
activity. The increased metabolism
and
for
fluids
the
excretion of
for the secretion of the digestive
"

metabolic

productsis also to be
productionconsequent

reckoned

among

the

of

causes

the

ingestionof feed.
The
extensive
Fermentations.
fermentations,especially
c.
the methane
fermentation,occurring in the digestivetract of
evolution of
herbivora
(128-130, 132) result in a considerable
determinations
of its amount
heat.
No
entirelysatisfactory
*
but Von
der Heide, Klein and Zuntz
have yet been reported,
Markoff's
mentation
ferfrom
experiments that the methane
compute
of 4.374 Cals. of
in cattle gives rise to the evolution
heat per cubic centimeter of methane, equivalentto 6.07 Cals.
the heat

on

"

per gram.
d. Intermediary metabolism.

chemical

The

"

changes which

undergo during digestion and resorption and

in the intermediary metabolism
(compare Chapters
especially
invoked
heat
been
to explain the increased
III and V) have
productionconsequent on the consumption of feed,particularly
nutrients

the

protein,but

of

of these reactions
Direct

e.

Lusk

and

apparently without
seeming

stimulus

to

be

Recent

"

the

upon
action, togetherwith earlier

have

gone
Lusk, the

far towards

most

substantiallyisothermic.

to metabolism.

his associates

sufficient warrant,

clearingup

by
investigations
the
cause
specificdynamic
experiments by Gigon,3

the

of

subject. According

to

carbohydratesand fats is to be explained

done by C. Voit in 1881, viz.,as the direct
was
as
substantially
effect of a greater supply of non-nitrogenous material
to the
of mass
action.
The
a
case
as
products of
cells,
i.e.,
virtually
the hydroxy
proteinkatabolism, on the contrary, particularly
the deaminization
of the amino
from
and keto-acids
resulting
action of

(233), act
nitrogenous matter

acids

as

direct

stimuli

to

the

katabolism

of

non-

body cells.
mechanical
That
these actions play their part, along with
heat
work
and fermentations,in bringing about the increased
1

Landw.

in the

Jahrb.,44 (1913).
795-

13 (1912), 27, 155,

185; 20 (1915), 555Jour. Biol. Chem., 12 (1012), 349;
Arch. Inter. Medicine, 12 (1913), 485.
Proc. Internat. Cong. Hygiene, 1913.
Jour.
Amer.
Med.
Asso., 63 (1914),824.
3 Skand.
Arch. Physiol, 21 (1909), 35* ; Arch. Physiol.(Pfliiger),
140 (191 1),548.
2

MAINTENANCE

ENERGY

THE

"

REQUIREMENTS

277

consumption of feed by herbivora

be doubted.
Besides
cannot
proteins,carbohydrates and fats,
however, the feed of herbivora contains a great varietyof other
substances and the results upon steers obtained
by Armsby and

productionresultingfrom

Fries

stimuli

as
acting specifically

muscular

the minor

to

these

that among

indicate

to

seem

the

be

may

pounds
com-

the cell metabolism

to

or

animal.

of the

movements

there

Among the
notably true of

feeding stuffs examined, this appeared to be

alfalfa hay and maize meal.
Work
of digestion."
The
368.
expenditure of energy
by the body which results from the ingestionof feed has been
somewhat
loosely,and perhaps not altogether fortunately,
be obwork
of digestion." While
there may
jections
designated as
and while it must
be interpretedtoo
not
to the term
lective
colit may
nevertheless serve
useful purpose
as
a
a
literally,
expressionfor the energy cost to the organism of all the
various processes involved in the digestionand assimilation of
is equal, of course,
the feed.
Its total amount
to the extra
heat produced above
that generated by the fasting animal.
Rubner's
specific
dynamic action,or Kellner's thermal energy,
is equivalentto the
work of digestion in this broad meaning.
The considerations presentedin the previous paragraphs
of the factors of the
work
of digestion
to indicate some
serve
"

"

"

"

"

"

and

render

it evident

mechanical
one,

part.

In

sense.

while with

that

similar difference is

subordinate

play a

369.

no

this

carnivora

it

are

all work

means

factor

is

in the

important
apparently plays a small

shown
strikingly

digestivefermentations,which
but

by

herbivora

and

man

it is

"

in the

extensive

very

an

case

of the

in ruminants

role in other animals.

Significance of

of

expenditure

energy

in

feed

sumption.
con-

Whatever

the part played by various factors in

the increase of metabolism
due to feed ingestion,the existence
"

of that increase and

is

fullyestablished

the consequent
fact which has

the value of the feed

Recurring
with
has

an

to

once

internal
obtain

it is
level,

more

as

to

source

supply

evident
1

of energy.

the

combustion

its

an

heat

production
important bearing upon

augmented

of

comparison of the animal body

motor
(274),if a gasolineengine
fuel by hoisting it from
lower'
a

that the energy

spent in this way

Jour. Agr. Research,

(1915), 479.

diminishes

278
to

just that

can

deliver

other

in

same

delivered

forms

the

of

if the

as

at

FARM

ANIMALS

quantity of

the

extent

virtuallythe
as

OF

NUTRITION

which

energy

work,

that

so

the

of

content

energy

diminished

cylinderwere

the

effect

the

by

engine
is

gasoline
the

same

amount.

In
"

similar
precisely

"

of

work

digestion

of the

general purposes

like the chemical

energy,

part

way, the energy

and eliminated
as

at least is

be eaten

be hoisted.

The

bolizable energy
of energy
which

Net

which

like that with

(364),the
of
towards
The

the

the

or

its

cited

The

use.

was

net

By

waste

of

amount

net

external work

or

the

of balance

illustration in

called the net energy

result

value

pound

the meta-

experiments
a previousparagraph
the heat production

upon
of energy
which
it contributes
the body may
be determined.

the net

502 Cals. per

be

the

to

feedingstuff

energy

from
the

obtain

to

means

an

amount

latter result has been

it shows

to

the

be made

must

performanceof

as

maintenance

because

by

effect of

animal

an

said

the

serve

performance of the
to its maintenance)
body (i.e.,

the

"

used

steer

not

engine must
tutes
doing, however, consti-

so

contribute

can

values.

energy

be

for
gasoline

spent in

internal work
of
necessary
the
to such processes
or
as
fat.
or
storage of meat
370.

the

of the feed in order

it

does

the so-called

of the feces,for example, since

energy
for necessary
The feed must
purposes.

energy
deduction

a
virtually

heat

It cannot

body.

expended
assimilated,
justas

and

expended in

of

of

dry

be

regardsenergy obtained
the hay in the illustration
as

energy might
minus
the work
of
Net

matter.

as metabolizable
defined,therefore,
digestion,the latter term, of course,

value of the substance

energy

in the
being understood
as
general sense
equivalent to the
already indicated
very
additional
heat production caused
by the consumption of
the feed.
Stated

in

different way, the net

slightly
is the energy remainingafter the

feedingstuff

in the
energy
the processes

have been
deductions
One

various

excreta

incident

deducted

to

value of a
energy
losses of chemical

the

also the energy

consumption of

expended in

and

the

material

of these
its gross energy.
The amount
different feeding stuffs.
naturallyvaries as between

containingmuch

from

digestiblematter, readilymasticated

exertinglittle stimulatingeffect

on

the

metabolic

and

processes

280

"

2.

of

maintenance

preservationof

animal

an

the

in the

contained

pages, however,
the maintenance

of

store

body,

ANIMALS

and

of

much

live

weight

in the

strict
and

only a

ration

the

recorded

ration is derived

Animals

maintenance.

of

from

"

the
signifies

sense

potentialenergy

which

effects this is

in
appear
information

will

As

Farm

of

matter

ration.

maintenance

really a

and

maintenance

True

FARM

Requirements

Maintenance

The

372.

The

OF

NUTRITION

subsequent
regarding
in which

experiments

its effect in
sufficiencyof the ration was
In experiments on
maintaining the live weight of the animal.
considerable period of
and
animals
mature
a
extending over
is involved, especially
time, it is unlikelythat any gross error

the criterion of the

of

nitrogen balance show

the
on
adequate. In short periods,

if determinations
be

supply to

the

the
other

animals,the live

in experiments on young
especially
is a notoriously untrustworthy guide.
but in young
animals
for this are familiar,
and

factor

tendency

to

into

enters

of young
animals.
increase of tissue is
over

is

growth

consideration.

Waters
marked

so

the demand

for

rations

15

immature

cattle

just sufficient

to

impulse to
apparentlytake precedence

maintenance,
weight and continue
time

on

characteristics
this

that

it may

important
known, the

very

is well

marked

shown

that

maintain
its
even
may
of skeleton for a considerable
Some

has

and
to

that

an

increase

maintain

their

live weight.

animal
in size
ration.

sub-maintenance

fed for considerable

were

hand,
weight

general reasons

another
As

of the most

one

The

protein

periods on
Under

these

height, in depth of chest

dent
time, however, there was" an evioff in the amount
of fat tissue,both as judged by the eye
falling
and as shown
and by the chemical
by the appearance
compositionof
the carcass.
in the size
a reduction
studies,too, showed
Histological
fat
of the fat cells and analysesof the adipose tissue showed
a lower
and higher water
and protein content
than in check animals.
What
occurred
of
fat
to supply energy,
was
evidentlya consumption
body
while at the same
time
an
approximately equal weight of protein
low energy
of the relatively
tissue was
produced, which, on account
value of protein and of the relatively
of water
panying
accomlargeamount
than did
smaller quantity of energy
it,representeda much

conditions,the animals continued

and length of head.
At the same

the fat tissue which

1

Soc. Prom.

to

disappeared.

Agr. Sci.,Proc.

grow

In

in

other

29th Annual

words

Meeting

the

rations

(1908), p.

71.

were

MAINTENANCE

THE

"

ENERGY

281

REQUIREMENTS

rations.
It
reallybut only apparently maintenance
hardly correct to say that in these experiments growth

is

not

the

at

of the

of the

expense

would

case

fat of the

be

that

the

tissues.
increase

A
of

more

perhaps

was

tained
main-

exact

ment
state-

protein tissue and

the loss of fat.

Presumably this effect would

in more
marked
mature
animals,in which the true maintenance
would
doubtless approach each
the live weight maintenance
over
closelywhen measured
long periods.
masked

water

Methods

373.

The

"

of

determining the maintenance

obvious

most

for

method

less

be

and

other

requirement.

determining the

maintenance

of trial. It consists of
requirement of an animal is the method
of feed until constancy
of live weight is
varying the amount
tween
attained or until a balance experiment shows
equilibrium beThis method,
and outgo of matter
and energy.
income
considerable
if extended
over
a
length of time, is particularly
of the live weight maintenance.
adapted to the determination
When
tested by the more
refined method
of the balance
periment,
exdent
however, such a ration will only rarely and by accibe found

to

be

exactlya

there will be revealed

for which

correction

more

be

must

maintenance

true

less

or

ration.

ally
Usu-

gain or loss by the body

applied.
comparison, like that used

in

previous paragraph (364),to illustrate the determination

net energy
values,between the effects of two different amounts

of

A second

method

consists of

of the
Such
the

same

comparison

net

value

energy

determine
in the

feedingstuff

case

Table

the energy
cited were

42.

"

not

ration upon
the balance
of
only affords the means

of the feed

consumed

but

requirement of the animal.

as

follows

Determination

of

Maintenance

Pounds

Period

Difference per lb. of dry matter

of

hay

to

serves

The

results

Metabolizable

Energy

Cals.

Heat

Gain

of

Pro-

duced Energy
Cals.

Cals.

9544

9812

-268

6.17

5768

8064

-2296

4.04

3776

1748

2028

10.21

Difference

also

Requirement

Dry

computing

"

Matter
Hay
of
Eaten

Period

of energy.

or

935

433

502

282

NUTRITION

OF

FARM

ANIMALS

of the

hay
The

permitteda

502 Cals.
the animal
loss from

loss to

would

Each

pound

from

the

of

dry matter

body by

zero

lb. and

0.53
would
have
=

an

exact

been

10.21

for maintenance

0.53

fashion

Cals.

ration

reduce

To

of 268

this
-r

502

regards energy

as

pounds

the metabolizable

lb. still

10.21

addition

10.74

of

of

energy

the

hay.
required

was

268

9544
374.

of 268

maintenance

the loss of energy

ration

obviouslyrequirethe

preciselysimilar

In

decreased

of

Computation

Cal.

-f-g-Jf10042
=

fasting katabolism.

the

Another

"

since it
comparison,however, is of greater significance,
affords results of more
to bring out
generalvalue and also serves
the net energy
values of feeding
clearlythe relations between
and the maintenance
the fastingkatabolism
requirement.
stuffs,
In the foregoingexperimentseach pound of hay withdrawn
of

method

from

ration

the

caused

heat

the

production to

decrease

by

withdrawn
from Period 3
If,then, all the hay were
and the animal reduced to the fastingstate,the heat production,
in other words the fastingkatabolism,would be
or
433

Cals.

8064
The

(433 X 6.17)

5392 Cals.

also be computed from

the losses of
result may
of each pound
The
withdrawal
suffered by the animal.
same

energy
of hay

increased

this

loss

by

502

Cals.

The

withdrawal

increase
6.17 pounds of Period 3, therefore,would
the loss by 502 X 6.17
3096 Cals., making a total loss
In other
of 5392
Cals.,equal to the fasting katabolism.
words, by such a comparison as the foregoingit is possibleto
it is scarcely
the fastingkatabolism,which
determine
indirectly
to determine
directly.
practicable
ing
in Chapter VII
It was
shown
(344),however, that the fastof the maintenance
katabolism
is the measure
requirement.
of all the

To
to

supply in

losses in the

equal

additional
which

of energy,
of metabolizable
amount

amount

an
(i.e.,
fastingkatabolism,

amount

to

offset

consumption of the
work of digestion.

the

the
i.e.,

an

excreta

the

to

his feed

be necessary
after deducting the

of this illustration it would

the steer

maintain

the

5372

Cals.,plus a

additional

feed would

heat

energy),
sufficient

production
inevitablyoccasion,

MAINTENANCE

of

the work

the

between

difference

the

But

digestionis

the

net

283

REQUIREMENTS

ENERGY

THE

"

metabolizable

energy

energy

and

(370). Consequently
tenance
saying that the main-

is equivalent to
foregoingstatement
requirement of the steer was
5392 Cals. of net energy.
Each
pound of this particularhay had a net energy value of
there would be
the animal, therefore,
To maintain
502 Cals.
required5392 -5- 502
10.74 lb.,as previouslycomputed.

the

of

Manner

375.

stating

requirement.

maintenance

the

"

requirementof an animal, such as

Evidently the maintenance
be stated in a variety
the steer of the foregoing illustration,
may
of digestibl
in terms
of weight of feed, of amounts
of ways
of net energy.
or
nutrients,of metabolizable
energy
concerned, it
So far as the results of a singleexperiment are
is adopted,
of statement
little difference which
makes
manner
of expressingthe same
since they are all simply different ways
it is desired to make
facts. When
generalstatements, however,
there are very manifest advantages in statingthe maintenance
"

ration
It

of net energy.

in terms

in

shown

was

Chapter VII

(343) that the fastingkatabo-

under uniform
constant
might be regarded as practically
conditions.
requirement for
Consequently the net
energy
is equally constant, and in the foregoingexample
maintenance
of 5392 Cals. would
value
energy
any ration having a net
lism

been

have

since

But
as
can

well

as

ration.

maintenance
the

the

net

values

energy

proportionof

of different

their metabolizable

be utilized for maintenance, may vary

range, the weight of feed or the amount

energy

which

will suffice

the kind of material fed.

to

paragraphs (380, 381) that

about

6.0 Therms

results of

Armsby

of net
and

which

energy

through

able
consider-

of metabolizable

will vary with

example, it is shown in subsequent

maintain

For

feedingstuffs,

energy

an

animal

thousand-pound
for maintenance.

Fries' determinations

of

net

requires

steer

From

the
values

energy

in timothy
(760),it is easy to compute that to supply this amount
hay with a net energy value of 48.63 Therms
per 100
would
pounds of dry matter
require 6.0 -f- 0.4863
12.34
pounds of dry matter, but that if mixed hay with a net energy
of
value of 43.37 Therms
pounds were used,the amount
per 1000
would
be 6.0 -f- 0.4337
13.83 pounds.
dry matter
necessary
The quantitiesof metabolizable energy contained in these main=

284

OF

NUTRITION

rations would

tenance

followingstatement,
mixed

two
Table

43.

likewise be
in which

rations

Examples

"

FARM

of

used

ANIMALS

is shown
as
different,
included

are

the

by

for further illustration

experimenters.

same

Maintenance

in the

Rations

for

iooo-Pound

Steer

Net
Energy

Material

Therms

Timothy hay
Mixed
hay

6.0

and

meal

Corn

Mixed

6.0

mixed

hay, 2 : 1
hay, 2 :

alfalfa

grain and

6.0
6.0

requirement in terms of net energy

By statingthe maintenance
is obtained
for an animal, or a single
for
a singlevalue
average
is a general expression of its mainclass of animals, which
tenance
a
of the parrequirement irrespective(substantially)
ticular
feed or feeds which
be
used
while
to
a
satisfyit,
may
statement

in

feed must

also

of

terms

of weight of
or
energy
particularkind of feed to which it

metabolizable

specify the

applies. The greater convenience

the computation of actual rations
which

the net

be estimated
of

of

the

former

is evident.

To

method
the

extent

values of feedingstuffs are known

energy
it is possible
endless
to make
up an almost

combinations

which

will furnish

the amount

followed

far

all be

will

maintenance

for

or

to
can

variety
rations,i.e.,

requiredby the animal.

energy
of expression
In the following
will be
paragraphs, this method
so

few

have

Modified

of the

practicable,
although unfortunatelycomparatively

determinations

maintenance
376.

as

requirementsfor
energy
reportedexcept in the case of cattle.

of the

yet been

conception of

energy

requirement.

state

work,

study
which

fastinganimal makes it evident

requirement outlined in
energy
in its application
to
modification

conceptionof the
Chapter VII requiressome
the actual feedingof animals.
As was
there shown
(344),the
in

"

of the

the

animal

net

as
conditions,especially
regardsmuscular

influence the katabolism

that

of net

of absolute

heat

muscular

productionof the fasting

be regardedas
rest may

MAINTENANCE

"

THE

ENERGY

285

REQUIREMENTS

for its internal

measuring the quantity of energy indispensable
of fact,however, absolute muscular
As a matter
work.
rest
for any
considerable length of time, at
be maintained
cannot
least during the waking hours, even
by voluntaryeffort. The
horse or ox when
at rest in the ordinary sense, i.e.,
when
doing
external work, is still expending a not inconsiderable amount
no
in muscular
activities of various sorts, some
of energy
indicated in "3 of the same
were
chapter (348). In

of which

particular,

(349) that standingas compared with lyingcauses

increase in the heat production, especially
in
a
very marked
the case
of cattle.
When, therefore,the heat production of
such an animal in the fastingstate is taken as a measure
of the
requiredfor its maintenance, it does not represent a
energy
of absolute rest but simply with one
of relativelyless
state
in the
activity. The
requirement for maintenance
energy
economic
includes not
sense
only the absolute minimum
quired
refor the internal work
but also the amount
expended in
stated

it was

various

of incidental muscular

forms

but
physiologically

unnecessary

Moreover,

less variable

individual

between

as

at

and

to

unavoidable

are

in

are

and

sense

practically.

of this incidental work

different individuals

is

more

in the

or
same

tenance
times, the energy requirement for mainfixed,constant
quantity whose exact value can

determined, but
is

which

different

is not
be

the amount

since

work

show

variable

the range

to determine

one.

of variation

general average

The

purpose

which

value

may

of

investigation
be expected

for the conditions

of

ordinarypractice.
The
377.

Net

carnivora

maintenance

energy

requirementof

requirement.

swine, having

"

With

swine

animals

such

as

man,

comparatively simple digestive

and
apparatus
feed, the
consuming relativelyconcentrated
fastingenergy expenditure can be determined without special
tively
difficulty
by deprivingthe restinganimal of feed during a relashort period and measuring the katabolism
with the
aid of a respiration apparatus
The
total
calorimeter.
or
of heat
amount
either directly or
produced, determined
furnishes the measure
by calculation,
of the energy expenditure
and

or

therefore of the

net

energy

requirement

for maintenance.

286

NUTRITION

Such

experimentmust,

an

of course,

ANIMALS

be made

(354) for

the critical temperature

above

FARM

OF

at

temperature

the

wise
animal, since otherthe heat produced would
be greater than that corresponding
internal work
to the necessary
by the additional amount
the body temperature.
to maintain
necessary
determinations
of the fastingkatabolism
Numerous
of man
of
the
smaller
such
and
the dog, cat, rabbit,guinea
as
animals,
pig,etc.,are on record,but the only experiments of this sort
farm animals
those of Meissl,Strohmer
and Lorenz l
are
upon
and of Tangl 2 upon
swine.
Meissl's determinations
200 C, a ternmade
at about
were
is well above
perature which, accordingto Tangl's later results,
the critical temperature for mature
swine.
In Tangl's experiments

the animals
no

paper

statements

Excluding
below

the

live

Table

44.

critical
in

pounds
weight,were
too

per

of

made

as

"

lying; in

temperature,

follows

Meissl's

this

on

proportion

Net

time

point.
Tangl's experiments which

are

those

of the

most

spent

to

the

were

ently
appar-

results,computed

the two-thirds

of the

power

"

Energy

Maintenance

for

of

Swine

Energy

Net
per

100

Live
per

MeissVs

Swine

H1

Swine

H2

Pounds
Weight
Day

Therms

experiments.

1.283
1.244

Average

I.266

TangVs experiments.
Two

mature

animals

at

i6"-i7"C

at

220 C

I.220

I.224

growing animals

Two
at

200 C

at

230 C

I.307
1.226

at26"C

I.270

Average

Ztschr.

1.249

Biol.,22 (1886), 63.

Biochem.

Ztschr., 44 (1912), 252.

288

NUTRITION

Table

45.

OF

ANIMALS

Maintenance

Daily

"

FARM

Metabolizable

energy

100

per

Rations

of

lb. live

weight
2.558 Therms
0.897 Therm

Maximum
Minimum

Average of all
Average omitting lowest
Average omittinglowest

the

with

Comparison

379.

and

highest
two
highest
.

and

net

energy.

Therms

1.5

Therms

10

Therms

1.474

the average

On

"

1-534

of all

corded
respirationexperiments on fattening swine which are rein Chapter XVII
olizable
(761), 78.14 per cent of the metabbe computed to have
been
lized
utisupplied may
energy
for maintenance
be assumed
to
plus gain. If this may

represent approximately the

energy
rations

available

amounts

of net

for

46.

Daily

"

Computed

net

"

Maintenance
energy

Rations

differ
on

1.998 Therms
of all

for

requirement of net energy as thus computed

indicated
periments
by the exgreatlyfrom the amount

fastinganimals

(377),but

the

380.

Net

maintenance

energy

requirementof

requirement.

In

"

the

of

katabolism

range

in

manner
striking

cattle
of

case

the
directly

to determine
it is hardlypracticable

by measuring the

enormous

in

singleexperimentsshows
further investigation.

The

Therms

i-x99

the results of the

the need

Therm

0.701

Average

not

Swine

of

lb. live weight

100

per

Minimum

averarge

metabolizable

the

Maximum

The

the

foregoingmaintenance
pounds live weight, the following

100

energy

of

percentage

maintenance,

contained, per

Table

does

Swine

the

ruminants,

net

energy

quirement
re-

fastinganimal.

Prolonged fasting would be required to free the voluminous

and
complicated digestiveorgans of these animals from feed
residues,ff this could be accomplishedat all,and it would be
that point was
when
difficult to determine
reached, while it is

questionablewhether
regarded

as

normal.

the results

on

such

an

animal

could

be

MAINTENANCE

Table

47.

Net

"

"

Energy

ENERGY

THE

Requirement

Corrected

to

12

hours

for

Maintenance

standing

Year

1902

1903
1904

1905
1905

1906
1906
1907
1907

1908

1909

1911

1912

Average of all
Average,omitting alfalfa meal
Average of expts. with roughage
.......

only x
Average

of

expts. with

mixed

rations
.

Omitting experiment on
U

289

REQUIREMENTS

alfalfa meal.

of

Cattle

NUTRITION

290

FARM

OF

ANIMALS

of such
animal
an
fastingkatabolism
be computed in the manner
already described
comparison of two periods on different amounts
The

however,
a
(374) from

may,

of the

same

for maintenance.
ration,both being less than that necessary
nine
Twenty-three experiments of this sort, on
of which
tive
different steers, only one
was
fat,in which the relametabolism
of the animals
when
standing and when lying
puted
was
determined, have been made by Armsby and Fries.1 Comof
lb.,in proportionto the two- thirds power
per iooo
the live weight (347) and corrected 2 to a uniform
period of 12
hours standing out of the 24, the net energy requirements were

feed, or

shown

as

plan

in Table

have

other

No

47.

this
precisely

experimentson

yet been

reported.
if a few seemingly extreme
Even
results,like those of 1902
and
The
wide
a
191 2 be excluded, the figures show
range.
show
rations
of roughage and
trials with mixed
concentrates
the whole
somewhat
on
higher results than those with roughage
only,but the experiments are hardly numerous
enough to
show
whether
this difference is significant.
381.

Net

in

energy

of earlier

number

of net

amounts

experiments

based

have

by
4

small

The

in which

by

corrected

approximate
of

means

for upon

metabolizable

Eight

Buls.
2

3
4

74,

of them
ioi,

In the

for maintenance

number

of

Of

those

important are

respirationexperiments
the

less accuracy.

or

interest.

and

energy

have

been

the
on

by

G.

rations

were

the animals

fed;
were

respirationapparatus

required for

S.

rent,
long cur-

investigations,
Kiihn and by

productive rations,and

U.

which

the later

basis of results obtained

reported.

on

subsequent ones,3

maintenance

Pettenkofer

the

rations

maintenance

gains or losses of proteinand fat by

determined
and

historic

far the most

in which

Henneberg and Stohmann,

considerable

chieflyan

now

Kellner

as

more

considerable

"

record

on

in actual

feeding standard

Wolff's

well

as

also

are

contained
energy
be computed with

of cattle may
The earlyexperiments of
was

rations.

maintenance

maintenance

Dept.

Agr.,

Bur.

128.

described
in Jour. Agr. Research, 3 (1915), 454.
manner
Compare Penna. Expt. Sta., Bui. 42 (1898),pp. 8-21.
Landw.
Vers. Sta., 53 (1900), pp. 6-16,
Reported by Kellner:

in

other

in this way
was

Anim.

puted.
com-

Indus,,

MAINTENANCE

In

addition
the live

upon

ENERGY

REQUIREMENTS

291

these

respirationexperiments, investigations
of cattle made
weight maintenance
by the writer,1
to

by Haecker,2and

by

THE

"

by

Evvard

have

been

discussed

elsewhere

the writer.

Table

48.

"

Net

Energy

in

Daily

Maintenance

the purpose
of computing
values of these rations it seems
For

that the
1

Penna.

Thesis

same

the

Rations

approximate

permissibleto

of

Cattle

net

energy

assume

proportion of their metabolizable

2 Minn.
Expt. Sta., Bui. 42 (1898).
Expt. Sta., Bui.
degree of M. S., University of Missouri, 1909.
4U. S. Dept. Agr., Bur. Anim.
Indus., Bui. 143, 44-46.
5
Omitting the experiment on alfalfa meal.
6
Giving each experiment equal weight.

for

sionally
provienergy
79.

NUTRITION

292

available

was

for maintenance

of alfalfa

ANIMALS

in the

as

rations

mixed

of the

case

of 55

percentage

hays

sive
exclu-

Fries,1viz.,52.8 per

and

investigatedby Armsby

the

For

cent.

FARM

OF

has

been

sumed.
as-

experiments the net energy was computed

nations
investigator. Eckles 2 has also reported five determiof the live weight maintenance
of dry cows
in which
In Evvard's

by

that

the

net

values

energy
estimated
from

the

of

medium

Kellner's

and

although if
be

the

It

in the

that

6000

medium

with

the

(375) and

of

is indeed

The
382.

metabolizable

kinds

of
the

is

is

rations

The
of the

Cals.,

viz.,6 181

state

61 13 Cals.

but

the data

feeds

are

too

trustworthy average.

maintenance
must

the actual
maintenance
energy
used
has

by
The

evidentlydistinctly

ration

of

mature

the

supply,on

age,
aver-

pounds live

this average
of feedingstuffs

weight
ration,as

contained

in

already been

well

the

as

it,will

vary

pointed out

obvious.
sufficiently

maintenance

requirementofsheep

in maintenance
energy
ration of sheep are
regardingthe maintenance

those

two

Cals.

5934

to

Metabolizable

than

Fries'

results of the

net

That

constitute

quantitiesof

energy
than
the

and

Armsby

per thousand
energy
considerable
variations
from

cases.

to

cattle

condition

Cals. of

weight,although with
in individual

is reduced

unfattened

statement

then,
to

net

apparently exceptionalresult obtained

one

the

permit

48,

ment
require-

energy

net

that

required

in

animals,

greater than

about

found

different

omitted, the

appears,
cattle in thin

were

Table

less
distinctly

individual

average
requirement of fat

to

net

the

of

maintenance

few

is

requirement
16

the

in the maintenance
energy
experiments is somewhat
greater,

weight

Evvard

rations

mean

estimated

average

consumed

comparison.
thin animals, the estimated

maintenance

experiments. The
experimenters, on
live

for

maintenance

average

of

determinations

being included
For

rations

computed averages.3
in
computations are shown

the

Fries'

and

Armsby

mixed

the

the writer's

results of

The

of

for cattle and

no

rations.
much

experiments have

Jour. Agr. Research, 3 (1915),484-485.

Mo.
Bui. 7, p. 120.
Expt. Sta., Research
3U. S. Dept. Agr., Farmers'
Bui. 346, p.
2

15.

"

Data

dant
less abunbeen

re-

MAINTENANCE

THE

"

ENERGY

REQUIREMENTS

293

ported in which the net energy requiredfor maintenance, i.e.,

the fastingkatabolism, has been determined.
by
sheep have been made
Respiration experiments upon
l
in 1867-1868 on
two
animals,by
Henneberg and Stohmann
2
in 1872 upon
two
animals,
Henneberg, Fleisher and Miiller
4
3
in 1899 on one
animal, and by Kellner
by Hagemann
upon
ance
balthe
these
of
the
With
animal.
were
third,
one
exception
experiments with a Pettenkofer apparatus and included
far as reported. The
of energy
direct determinations
so
no
metabolism
third investigationcomprised a
digestion and
creta
experiment in which the energy of the feed and the visible exdetermined
was
directly and also 42 determinations
with the Zuntz
of the pulmonary respiration
type of apparatus.
(299).

experimentsthere are
foregoingrespiration
in which
the
number
of digestion experiments by Wolff
a
live weight of the animals
was
approximatelymaintained,5and
In addition

Henry

to

the

reports a series of experiments by Carlyle and

Klein-

rations proin which various mixed

breeding ewes
duced
head
in
animals
of
0.16
an
dailygain
pound per
average
in
weight.
averaging 145 pounds
of the rations
In these experiments the metabolizable
energy
the digestibleorganic
be computed approximately from
may
described in Chapter XVII
in the manner
(774). In
matter
the respiration
experimentsa correction for the gain or loss by
in the case
of Kellner' s experiments
be made
the animal may
as
cattle (381),while an approximate correction for the gain in
on
also be made.
the live weight experimentsmay
The
results,computed per 100 lb. live weight, are shown in
heinz

with

Table

49.

Net

383.

stated, no
for

direct

of

unsatisfactorydata
metabolizable

energy

are

sheep

available

for

of maintenance

are

on

energy
record and

rations.

ready
al-

As

"

net

computing

Beitrage, etc., pp. 68-286.

(1873-74), H, 145.
Jahresber. Agr. Chem., 16-17
3 Arch.
(Anat. u.) Physiol. ; 1899, Suppl., p. 138.
4
Die Ernahrung. landw. Nutzt., 6th Ed., p. 422.
5
Compare U. S. Dept. of Agr., Bur. Anim. Indus., Bui.

sheep.

of the

determinations

the maintenance

of

rations

in maintenance

energy

quired
re-

only

it from

the

the

fore-

For

Neue

Feeds

and

Feeding, 10th Ed.,

p.

482.

143

(1912), pp.

49~5i-

however,
going experiments,
of cattle,that
in the case
as

ANIMALS

FARM

OF

NUTRITION

294

it may
about

be

permissibleto assume,
52.8 per cent of the metab-

of roughage and
55 per
energy
available for maintenance.
was

olizable
rations

cent

of that

The

results

of mixed
of

putation
com-

of the
this basis are contained in the last column
upon
following table. They possess a certain degree of interest,
although obviously they are of uncertain value.
Table

49.

"

Energy

Rations

Maintenance

Daily

in

Per

Sheep

of

Lb.

100

Live

Metabolizable

Weight

Net

Energy

Energy

Respirationexperiments
Ffenneberg and Stohmann
Henneberg, Fleischer and

Therms

Therms
.

1-475

Muller

"779

I.420

.781

Kellner

I. no

.611

Hagemann

1.282

.705

1.322

.719

1.634

.863

1.72s

.863

1.513

.832

1.624

.853

1.368

.791

Average
Live weight experiments

Wolff, 1871, 6 experiments

Wolff, 1892-1893, 8 experiments
Carlyle and Klienheinz
.

Average
Average

384.

cattle.

of all

Comparison

to compare

of

sheep with cattle.

the maintenance

Since

"

requirement

It is of
of

some

sheep

with

interest
that

of

sheep is a much smaller animal than the steer

it naturallyrequiresrelatively
feed in proportionto its
more
weight (345), as the foregoingfiguresshow to be the case.
As compared with a steer weighing 1000
ing
pounds, ten sheep weigheach
for
would
100
pounds
require
maintenance, according
the
to
foregoingestimates,about 30 per cent more
energy.
If,however, the comparison be made in proportion to the twoof the live weight, i.e.,
in proporthirds power
substantially
tion
to the body surface (347),a very different result is reached,
the maintenance
ration of the sheep as thus computed amounting
to

the

little more

than

60

per

cent

of that of cattle.

296

NUTRITION

rations is

the various

on

FARM

OF

computed

ANIMALS

the results of five previous

from

by combining them in various

consumed.
ratios according to the proportionsof oats, hay and straw
have
been
heat
the
this is subtracted
From
produced in
computed to
of mastication),
the digestionof the feed (not includingthe work
due
to internal
the remainder
showing, of course, the katabolism
the net energy requirement.1
work, i.e.,
balance

similar rations

experimentson

eight experiments compared the

total estimated
heat production per day, the computed energy
ference
expenditurecaused by the consumption of the feed,and by difthe energy
the net
state,i.e.,
expenditurein the fasting
Table

requirementfor

energy
Table

51.

In

the

Energy

Net

"

while

of 3.97 Therms

weight.

Zuntz

represents
1

For

of Animal

maintenance.

Requirement

more

in

for

standard

Maintenance

ration

of

Horse

of 6

kgs. of oats,
computed
average

(or seven) of hay, the

periods was
4.33
per day in three winter
it
reaches
the
minimum
period
singlesummer

six

fastingkatabolism
Therms,

the

experiments with

of straw, and

one

for

shows

51

per
and

head, or 4.08 Therms

Hagemann consider

approximatelythe
complete

account

Nutrition, pp. 386-387.

of

minimum
the

method,

pounds live
per 1000
that the latter amount
requirement for

compare

the writer's

the inPrinciples

MAINTENANCE

ENERGY

THE

"

REQUIREMENTS

297

regard the higherfiguresobtained in the winter

experiments a3 indicatinga stimulation of the heat production
by the low temperature to which the animal was exposed ; i.e.,
made below the critical
they consider that the experiments were
temperature. The notably higher results obtained with lighter
rations they ascribe to a similar cause, viz.,that the heat arising
from the work of digestion,
sary
togetherwith that due to the necesand

ternal work

(fastingkatabolism),was

work

internal

body temperature.
be confessed that,in view
It must
of the horse as compared
low figurefor the fastingkatabolism
the fact should

the results of

of

of total

applicableto

that

ration

may

various
the

to

seems

on

are

record

requiredfor

"

in which
the maintenance

mined
sheep may be deterby varying the quantity
and

income

outgo

stancy
con-

or

is

exert

greater stimulus upon

case

with

considerable

excess

consumed

be

the

by

of

income
appear

and

an

in

show

may

that

maintenance

expended

Chapter VII

outgo

to

actual
and

horse

his muscular

phlegmaticruminants,

more

over

activities noted

loss,i.e.,
may

nor

rations.

is not fully
attained,but this method
active ment,
the horse.
Owing to his more
tempera-

minor

balance

it with

compare

determined.

has been

weight

activitythan is the
so

experiments

of live

feed

to

was

maintenance

good degree of accuracy

given until equalitybetween

of feed

short

rations of cattle and

maintenance

with

from

unusually
nearly the physiological
tenance,
required for mainenergy

matter
digestible

of the horse
The

animal

interest

in

energy

number

considerable
amount

more

24-hour experiments.

Metabolizable

386.

perament
active,temwith cattle this relatively
is rather surprising,
and

the

net

be of much

it would

and

of

minimum

the economic

of the

that it is derived

be overlooked

not

periods in which it is probable that

quiet. It perhaps represents more

the

to

the

maintain

than

insufficient

show

in

the

(348), while
neither gain

the ration

is

tenance
main-

ration.

of avoiding this
method
One
Wolffs determinations.
and determining the true maintenance
ration is that
difficulty
*
work
employed by Wolff in his extensive investigations
upon
productionby the horse (670,779). In these experimentsthe
a.

"

Compare the writer's Principlesof

Animal

Nutrition, pp. 53i_535-

298

NUTRITION

horse

performed

adjusted

OF

measured

in different
the

feed

when

the

live

case

for

unchanged
urinary nitrogen did
of the

performed
heavier

feed

on

one,

consumed

basal
the

This

of the

show
done

animal

of

work

within

so

as

to

stantially
sub-

when

the

consequence
comparing the work

that which

the

remained

period and

(637). By

with

was

considered

was

increase

an

} which

nearly as possiblein

as

considerable

established

was

be

to

weight

ration
ratio

of work

consumed.

not

work

additional

ANIMALS

amount

periodsas

equilibriumwith
be the

FARM

could

done

to

be done

with

the additional

the limits of

error

of the

method, this being the prime objectof the experiments. This

to coma
pute
being determined, however, it was
simple matter
the

amount

while
ration

of feed

correspondingto

the difference between

evidentlywas

the

the maintenance

the

total work

latter and

ration.

From

the

performed,
total

the total

tenance,
(inclusiveof crude fiber)requiredfor mainthus computed by Wolff, the equivalentamounts
of
as
metabolizable
also be
required for maintenance
energy
may
computed approximatelyby the use of Zuntz and Hagemann's
factor of 3.96 Cals. per gram
(776).
In Wolff's earlier experiments and
in those
later ones
in
which
approximately equal proportions of hay and grain were
ration was
found
be approxito
consumed, the maintenance
mately
total nutrients
4200
grams
per 500 kgs. live weight,
In those later experiments (including
equivalentto 16.63 Therms.
the results of similar investigations
as
computed
reby Grandeau
by Wolff) in which a largerproportion of grainwas
fed, the total nutrients required for maintenance
ranged from
3600 to 3800 grams, equivalent to from 14.26 to 15.05 Therms.
In other words, the amount
of metabolizable
required
energy
for maintenance
varied with the proportion of roughage present,
would
be anticipatedfrom
the results with cattle reas
corded
on
previous pages.
b. Zuntz
and
From
a respiration
periment
exHagemann's results.
at the GottingenExperiment Station,
and HageZuntz
nutrients
digestible

"

mann
1

compute
Wolff's

the

metabolizable

energy

of

the

maintenance

made
with a sweep-power
also as a
experiments were
arranged to serve
The actual measurements
of the work performed, except in the later
dynamometer.
experiments, proved to be too low, but Wolff believes them to be relatively correct,
that the ratio between
the work
measured
and the additional feed required to
so
as
the basis of computation.
still serve
produce it may
as

MAINTENANCE

of

ration

by subtracting from

horse

the

REQUIREMENTS

ENERGY

THE

"

carbohydrateequivalentof
.gainedby the animal, disregardingthe
nutrients

the

effect of the feed.

total

digested
the proteinand fat
possiblestimulating

this way, they find for the maintenance

nutrients per head, equivatotal digested
lent

In

ration 2955.4

the

299

grams
Therms
Therms
to 11.70
or 1 2.1
pounds live weight,
per 1000
This difference is ascribed
result notablv lower than Wolff's.

and

Zuntz

by

Hagemann

rations, the work

Wolff's

the

to

digestionof

of

fiber in

as
ingredient

this

timated
es-

nearly accounting for the differ-

(777)very

by them

of crude

largercontent

ence.
*

1878-1879 attempted to
ration of the horse by startingwith
determine the maintenance
insufficientration and gradually
increasingit until an equiliban
rium
feed and live weight was
between
secured, seeking in this
feed (392).
to eliminate the stimulatingeffect of excess
manner
Muntz's

c.

experiments."Miintz

made

trials were

The

their work
found

He

which

that

may

ration

equal to A

estimated

be

able energy per 1000

pounds live
sufficient for maintenance.

in

addition

Wolff's

weight was

to

results,fed five cab horses

ration

and LeClerc,2

in connection

experimentsmentioned

the

than

more
slightly

Grandeau

"

and

ration

of metabohz-

Therms

12.1

and LeClerc's results.

Grandeau

d.

work

of the

contain

to

pany,
Com-

previousexperiments.

from

being known

ration

of the Paris Omnibus

the horses

on

in

with

kgs. of hay

of 8

during a total of 14 periodsof a month each (one to five periods

of the
for each animal) during each of which the digestibility
ration

was

results per
Total

determined.

On

day and head

were

nutrients (fat X 2.4)

digestible

Equivalent metabolizable energy

at

"

the
periods,

of all the

the average
follows
as

"

2783-7 grams

"

"

3.96 Cals.

per

gram

11.03

Daily gain in weight

Average live weight

393 -6

foregoing ration, which

The
than

"-*9

maintenance

metabolizable
1

Annales

L'alimentation

per

1000

de l'Institut National

du Cheval

de

kg.
kgs.

evidently somewhat

ration, is equivalentto

energy

was

Therms

pounds
Agronomique,

live
Tome

Trait, 1883, III.

13.

more

Therms

weight. This
3,

1876-1879.

of
is

3""

NUTRITION

materiallyless
with hay and

than

and

Zuntz
The

OF

obtained

was

about

the

same

followingsummary

by

as

regardingthe

metaboliz-

the

shows

maintenance

of variation

range

the

experiments
that found
by him and by
containingmuch grain.

of the data

required for

energy
considerable

ANIMALS

in Wolff's earlier

for rations

Hagemann

able

FARM

by

which

varying proportions of

horse

is

only partiallyexplicable
tained
grain and roughage con-

in the rations.
Table

52.

Maintenance

"

Rations

of

Horse

the

Experimenter

Wolff
Wolff

and

Grandeau

and

LeClerc

and Hagemann

Zuntz

Muntz
Grandeau

and

LeClerc
.

Wolff

387.

Metabolizable

The

"

net

requiredfor

energy

equals of course
Hagemann
compute,

in the

be

thousand

there
the

Therms

4.1

the

per

temperature

net

maintenance, as

other

with

This Zuntz

already described

manner

live

pounds

those

of their

lower

was

requirement.

energy

fastingkatabolism.

pointed out, however,

external

with

compared

or

weight.

twelve
Therms
2.5 of

the
l
on

and

(385) to
As

was

experiments in which

the amount

of feed less

also notably the case

higher results. The latter was
gave
earlier experiments in which
fed.
still lighter
rations were
On

mals,
ani-

in

of the eight most

satisfactoryexperiments out of
average
Horse II the total katabolism
1 1.027
per day and head was

upon

hay.

ration

consistingof

Computed

in

the

3.5

same

Kgs.
manner

and
of oats, 0.5 of straw
in Table
as
51, the

digestionof this ration is equal to

of 7244 Cals.,equivalent to
a
This
of surface.
centimeter
is a higher
Cals.,per square
140.3
in Table
figure than any of those contained
51, although the total
katabolism
not
was
notably different.
in
expenditure of energy
which
leaves
3782 Cals.,

Landw.

the

remainder

Jahrb.,18,

27, Ergzbd. Ill, 356-357.

MAINTENANCE

ENERGY

THE

"

REQUIREMENTS

301

the amount
of
conclude, therefore,that when
is small the lack is comheat liberated by the digestivework
pensated
of body tissue.
for by an increased katabolism
Their
final result is that their animal
requiredper head at least 11.00
his body temperature.
In other
of heat to maintain
Therms
which
of metabolizable
words, this is the minimum
energy
be contained
in a maintenance
must
ration,since if less be
although the ration supply the requisiteamount
present, even
of net energy,
body tissue would still be katabolized for the
authors

The

production of the necessary heat. Computed per thousand

tenance
pounds live weight, Zuntz and Hagemann's estimated mainrequirementis :
"

Net

energy

Additional

for internal

work

requiredfor

heat

Total

metabolizable

In

computing

horse

energy

ration

7.8 Therms

production
required
for the

11.9

of the

maintenance

actual

Therms

rest, it is necessary,

at

only

not

energy

the other

hand

computing work

but

no

such

need

allowance

rations.

production.

heat

ordinarilybe made

horse when

The

olizable
sufficient metab-

it contains

also whether

support the necessary

to

of heat

excess

to consider
according to these figures,
fastit suppliesnet energy equal to the ing

whether

katabolism

an

4.1 Therms

(compare Chapter XIV),

at

in

is producing

work

and

On

ing
during the work-

expenditure of feed energy for the sake of heat

productionwould be called for, while any ordinary working
ration would
tabolizable
probably contain a considerable surplus of mehours

no

hours

388.

of

maintenance

during the

demand

of rest.

The

the

the

over

energy

Net

maintenance

requirementof fowls

requirements.

energy

"

Gerhartz

has

measured

of a number
requirement of fowls by means
of respiration
experiments with the Regnault-Reisettype
has also comHe
puted
two
fastinghens.
apparatus (298) upon
the fastingkatabolism
of respiration
from a number
periments
exnet

energy

in which

the animals

the total metabolism

that
1

Landw.

were

computed

fed
to

by subtracting from
have

Jahrb.,46 (1914),797.

been

due

to

the

NUTRITION

3"2

of the

OF

feed

FARM

ANIMALS

i.e.,
by substantiallythe same
eral
genmethods
used by Zuntz
and Hagemann
for the horse (385).
of body
His results,computed per thousand
square centimeters
surface and also per 5 pounds live weight in proportion to the
follows :
as
" power of the latter,were
consumption

"

"

Table

53.

Net

"

Energy

Maintenance

for

Hens

of

Per
s
Pounds
Live
Weight

In

fastingexperiments
Minimum
when
not
laying
Average when not laying
Average when laying
.

from

Computed
Minimum

Average when
when

It would

57-oi

experimentswith

when

Average

Cals.

laying
not laying
laying

76.75
91-45

feed

not

from

55-13

66.58
93-91

the

tenance
mainfigures that the average
requirement of a 5-pound hen may be estimated at approximate
while
in
of
minimum
muscular
tivity
acperiods
72 Cals.,
fall as low as 56 Cals.
it may
The
much
higher figure
in the periods when
the hen
was
(93 Cals.) obtained
laying
does

represent maintenance

not

energy

appear

expended

in the formation

of the egg.
of the hen per unit

animals, the katabolism

when
computed per unit of
from
that of other species.
389.

hartz

Metabolizable
also

determined

the live
amounts
cm.

body

simply,but

weight

surface

in

energy

the

of metabolizable

also the

with

all small

is

high,but

As

weight

it does

differ

not

maintenance

rations.

and

energy

to

have

Rest

period
Molting period
f

Average

"

Ger-

tain
requiredto mainthe corresponding
computes
been

per

surface.

Brooding period

greatly

of feed

amount

of his fowls

includes

102

Cals.

107

Cals.

'"

Cals

Cals!

95 Cals.

iooo

sq.

OF

NUTRITION

3"4

used

indiscriminatelyor

Affecting

Factors

ANIMALS

computations of

for exact

cases.

Maintenance

the

which

conditions

Certain

basis

ill individual

the effects of feeds

" 3.

as

FARM

affect

the

Requirement

expenditure of

energy

of net
fastinganimal, and therefore the amount
energy
been
discussed
in Chapter
requiredfor maintenance, have already
be more
VII (345-357) while a few others may
conveniently
considered at this point.
the

391.

Temperament.

expending

The

"

in

may
very
his
maintenance
for

by the quieterand more

question that those

as

of different animals

vaguely

"

to

of muscular

amounts

in

Thus

"

individuality are

Armsby

between
which
due

There
the

to

maintenance

ascribed

are

be little

can

quirements
re-

somewhat

largeextent

to

ing
vary-

activity.
*

Fries' determinations

of the maintenance

(380)the two animals designated as A

and a scrub, the
were
a pure-bredbeef animal
respectively
less dairyblood and being of a decidedly
or
having more
difference
than
the animal
The
A.
nervous
disposition

requirement of
and

and

tinually
con-

ments
moveunnecessary
the amount
of energy
compared with that required

phlegmaticanimal.
differences

is

of

variety
materiallyincrease

energy

which

needed

restless animal

nervous,

latter
more

cattle

in the

requirements of
comparison, may

the

two

be

reasonably ascribed

animals,as

shown

by
to

the following

this difference

in temperament.

Table

55.

"

Net

Energy

Requirement

for

Maintenance

Scrub
Steer
B

Year

Therms

1905

"

"

1906

6.052
6.305

6.067

1907

Average

6. 141

U.

S. Dept. Agr., Bur.

Anim.

Indus., Bui. 128 (1911),

p. 53.

MAINTENANCE

Like

the

ENERGY

THE

"

conditions

external

temperament,

affect the

REQUIREMENTS

any
of muscular

305

tending to

activitywill also tend to affect

the maintenance
requirement. The steer confined in a stall,
take less muscular
for example, may
exercise,and therefore
requireless energy for maintenance, than one simply confined
animal
to a pen
or
comfortably bedded and
open yard. The
will
of his time lying down
to
spend much
thereby induced
than
consume
a smaller
proportionof his feed for maintenance
conditions.
one
kept under less comfortable
Any sort of
is likelyto be paid for by increased muscular
excitement
tivity
acincreased
and correspondingly
consumption of feed for
degree

maintenance.
392.

The

plane of nutrition.

It is somewhat

"

lieved
generallybe-

of feed necessary for maintenance

varies
of nutrition on which
the animal is kept. By

that the amount

with

the

plane

this is meant

that

some

time will

than

similar

an

animal

which

has

has been

sparsely fed and is in

less reduced
condition.
more
a
or
Thus, Waters 1 writes :
"
Apparently the animal
organism when
kept for a long
tenance
periodof time on a low nutritive plane,as in the case of mainbasis than when
economical
animals, gets on a more
fed. For example, if we
reduce the feed of an
more
liberally
animal that has been previouslyliberally
nourished to a point
where

for

month

or

equilibrium will
animal

may

later

increase

which

highly fed for

of feed for maintenance

requirea largeramount'
animal

been

there is

more

be
in

loss in

small

established

weight,an
subsequentlythe

and

weight, the quantity and

quality of

the feed

Thus
ration that was
ficient
insufa
remaining the same.
to sustain live weight at first may
be capable later of
maintaining the animal at a stationarybody weight,and still
later of causing an increase in weight. Digestion experiments

with
the

number

of animals

indicate that

part of this is due

complete digestionof the feed by the animal on a

nutritive plane,but so far as the experimentshave thus far
more

there

in the

does

degree to

not

which

seem

to

been

have

the feed has been

to

low
gressed
pro-

sufficient increase

digestedto

account

for all the increased

in the ration noted." 2

efficiency
sults
Comparativelylittle experimental confirmation of these rehas as yet been published,although respirationexperi-

Proc. Soc. Prom.

X

Agr. Sci.,1908, p. 96.

Compare Chapter XVI, "

(722).

306

NUTRITION

ments

dogs by

on

hauser2

OF

Kleinert

FARM

and

ANIMALS

by

Schlossmann

Mursch-

and

direction.
point in the same
made
Observations
by Zuntz and Hagemann 3 on the horse
In a series of respiration
suggestivein this connection.
appear
the common
observation that
experiments they have confirmed
lates
a
surplus of feed above the maintenance
requirement stimuthe muscular
activityand restlessness of this animal, so
that

ration

may
animal

the

give rise
A

to

seem

similar

to

no

be

more
considerably

than

sufficient to maintain

when

standing quietly in the stall and yet

increase in weight under
ordinary conditions.

effect of
stimulating

the feed upon

the minor
of cattle,
expeciallywhile standing,seems

movements

muscular
to

dicated
be in-

by the experiments of Armsby and Fries (367 e). It

tenance
possiblethat part, at least,of the diminution of the maindue
have
been
requirementobserved by Waters
may

seems

voluntaryrestriction of motion on the part of the animals

the low nutritive plane.
on
In attempting to determine
experimentallythe minimum
maintenance
of
requirementit is evidentlythe safer method
tenance
with
the horse, to approach the mainprocedure,especially
point by graduallyincreasing
a sub-maintenance
ration,
to

as

in Miintz's
and

Armsby
reduction
393.

of

the

experiments on
Fries

on

cattle

horse

(374)rather

(386 c) and
than

by

the

those

of

gradual

supermaintenance ration.
That

fat animals

have

relatively
greater
maintenance
than
thin
be
to
ones
seems
fairly
requirement
well established for cattle by the experiments of Kellner
and
of Evvard, the results of which are recorded
in Table
48 (381).
One
obvious reason
why the maintenance
requirement per
head
animal
in

Fattening.

"

should

be greater for a fattened animal

than for the same
before fattening
effort expended
is the greater muscular

standing,due

and

to

the greater

weight to
experiments upon the

be

supported.

Zuntz

horse carryingweight
Hagemann, in
its back, found that this increase was
on
proportionalto the
of weight added
amount
(665). If this be true generally,then
that portion of the metabolism
due to standing will increase
Fries'
and
In Armsby
more
rapidly than the body surface.
experimentson unfatted cattle,however, the energy expendi1

Ztschr.

3Landw.

s Biochem.
Biol.,61 (1913),346.
Jahrb. 27 (1898),Ergzbd. Ill, 211, 236.

Ztschr., 53 (1013),265.

MAINTENANCE

ENERGY

THE

"

REQUIREMENTS

307

to only about
standing 12 hours amounted
15 per
The
increase in the maintenance
of the total daily metabolism.
cent
requirement per unit of surface which is indicated by
Kellner's results is considerablygreater than would be computed

due

ture

to

the

this basis and

on

becoming greater

difference
394.

Age.

is

naturallysmaller
the

of

account

on

the animals

head

per

fatter.

grew

of

requirement

maintenance

The

"

as

fat animals, the

of Evvard's

is true

same

that of

than

an

there

Whether

difference in size.

mal
aniyoung
older animal

is any

ment
requirement, that is,in the requirecomputed to uniform weight or surface,is not altogether
farm
animals
results on
being on record.
clear,few specific
results on
Evvard's
higher than
yearlings(381) are somewhat
the

difference in

with

obtained

been

have

which

of those

most

relative

tle,
cat-

mature

figuresdo not relate to the same

individuals at different ages.
Armsby and Fries x in a series of
animals
two
respirationcalorimeter experiments upon the same
although,of

in three

these

with their full-blood

years found
in the maintenance
requirementas a

successive
decrease

marked

course,

steer

yearling

three-yearold, when corrected to a uniform number of

hours standing and computed in proportionto the two-thirds
and

as

power

hand,

of

the

scrub

steer,

on

the

other

requirementwas

of the maintenance

distinct decrease

no

the

With

weight.

observed.
extensive data

Somewhat
of

man

at different ages.

are

on

regardingthe metabolism
these by Tigerstedt2 seems

record
of

summary

clearlythat the metabolism

per unit of surface diminishes,
although,not very rapidly,from youth to maturity. In view of the
these results are
slow growth of man,
comparable to such
relatively
as
might be obtained during the first six to twelve months of the life
factory
animals and for these ages there are no satisof ordinary domestic
determinations
of the maintenance
requirement.

to

show

If it be
animal

true

that

the maintenance

relatively
greater than

is

requirement of

that of

an

young

this may
to the
extent

older one,

fairlybe presumed to be due to a considerable

animals,
activityusually exhibited by young
greater muscular
which, as already pointed out, notably increases the body
katabolism.
S. Dept. Agr., Bur.

U.

Nagel's Handbuch

Anim.

Indus., Bui. 128.

der Physiologie des Menschen,

I, 469.

308
"

NUTRITION

The

4.

Relation

of

one

other

factors

to warrant

which

is

(350-357) it

shown

relations

quirement,
re-

involved

seem

critical temperature.

the

In

"

the fasting
katabupon
for the fasting
animal
there

that

thermal
(or more
strictly,
critical temperature," at which

"

the

to

the unavoidable

justbalances

that

vironment),
en-

the

the necessary
fastingkatabolism
loss by radiation,
conduction and

the

body temperature is just maintained.

has a surplus of
temperature, the fastinganimal
it gets rid of by means
of the physicalregulation.

evaporation,so
this

heat which
the

Below

is but

temperature

production incidental

Above

exposed

of temperature

was

called
heat

is

affect its maintenance

complicated

influence

certain external

animal

an

may

consumption lowers

the
discussing
olism

which

separate discussion.

Feed

395.

Requirement

Temperature

to

the somewhat

ANIMALS

Maintenance

the

the temperature

among

FARM

External

to

While

OF

critical temperature,

the other

on

hand,

its katabolism

is increased
of the

the
i.e.,
has

As

energy
been

results in

the
increasing
is

animal

an

in

body

in the

beyond that necessary for the internal work

order to supply the necessary
of heat;
amount
expenditure is augmented.
shown
(365),however, the consumption of feed

the necessary

addition

productionof

it has
fed,therefore,

fastingstate, the

to

heat

heat

two

of heat

sources

from
resulting

internal work

of the

obvious
of its

that

the

maintain

to

critical

greater the
which

the animal

the result of feed

as

the

its

of feed

external

critical point,so

"

is

lowered.

consumed

temperature

that

animals

can

of

definite amount

which, togetherwith

the

dent
inci-

work

of

being
consumption,it is
more

heat

greater coolingeffect
katabolize

body
In other words,
Furthermore, the
is the

the lower
fall without

receivingheavy

feedingcan withstand more

maintenance.
Conversely, for any
there will be

"

so-called

beingcompelledto
body temperature.

temperature

amount

withstand

can

surroundingswithout

substance
"

body

as
first,

body, and second, in

this,the heat generatedby the

these
digestion." Under
conditions,with
the

When

the katabolism

to

produced in

animal.

an

cold than

point to

reaching the
rations

those

on

in productive

simple

particulartemperature
of any given feed the consumption
katabolism
requiredfor the internal

MAINTENANCE

THE

"

work, will give rise

just sufficient

to

the

balance

to

ENERGY

REQUIREMENTS

production of

the

unavoidable

309
of heat

amount

an

loss of heat

the

to

surroundings.
influence

This

the

of the

quantity of feed

following tabulation

different

temperatures

of

Rubner's

and

is well

results

illustrated

by

dog

at

upon

different

consuming

of

amounts

meat.

Table

56.

Influence

"

External

of

Temperature

Heat

on

Production

The

of feed

required to justoffset the coolingeffect

of
of a low temperature
might be called the critical amount
It will obviously be less the greater
feed for that temperature.

proportionof

the
as

amount

its metabolizable

energy

which

is

dissipated

heat.
For

example,

feeds necessary

in
for

of different
discussingthe relative amounts
maintenance
(375) it was stated that either
hay or 9.07 lb. of mixed grain and alfalfa

13.83 lb. of mixed

hay would yieldapproximately6.0
would

therefore

pound

steer.

The

in these two

gain

nor

are

be

be

of

net

energy,

for

ration

alfalfa

and

viz. :
different,

rations,the animal

finallyconverted

1000-

"

Therms

10.69 Therms

hay

all the

and

contained

energy

12.01

maintenance

lose energy

feed would

of metabolizable

amounts

hay
grain and

lb. mixed

Since both

maintenance

rations,however, would

13.83 lb. mixed

9.07

constitute

Therms

metabolizable
into

heat

would

neither

energy

of the

in both

cases.

The

animal
his

the exclusive

on

Therms

disposal1.32
could

his

Net

energy

critical temperature
and

than

heat

the other

and

temperature without

lower

have

at

ingly
accord-

drawing on

for fuel.

body

396.

hay ration,therefore,would
more

withstand

ANIMALS

FARM

OF

NUTRITION

310

below

critical

which

correspondsto

temperature.
the

consumed, in accordance with

the previous paragraphs, only part
of feed

kind

out

in

able

energy

to

serves

maintain

animal.

the

Down

"

particularamount
the facts
of

the

brought

metaboliz-

remainder

The

"

the

to

is

"

work
of digestion and converted
virtuallyexpended in the
comes
into heat, and this heat, since not needed
by the animal, beand is gotten rid of.
excretum
ternal
an
If,however, the ex-

temperature falls below

the critical

point the

is dif-.

case

ingestionof feed is justas useful

other
heat from
for keeping the body warm.
source
as
any
these conditions,therefore,
all the metabolizable
Under
energy
be of use.
of the feed may
Part of it (the net energy)is used
directlyfor supporting the necessary internal work of the body,
tissue
while the remainder
prevents the necessityof katabolizing
for the sake of heat production and is therefore
indirectly
from the consumption
In other words, the heat resulting
of use.

ferent.

Heat

resultingfrom

of feed

may

be

have

be

to

external

the

substituted

obtained

the

digestivework

the

metabolizable

the

by

falls

temperature

for

heat

that all the heat

required for

this purpose,
of the ration is of use

is

energy
to prevent loss of energy
all of it appears
to be net
energy.

in
are

kept

6.0 Therms

support

from

at

of net

the

energy

internal work

falls to

10.69 Therms

for the

the critical temperature

ration

and

is necessary

tration
illus-

ings
surround-

supplieswill

body, and the heat thus produced

while the
the body temperature,

which

of the

be

used

of the

maintain

internal work

an

as

the

point at

to

therefore

fastinganimal, the
to

will be expended in
energy
that the external temperature
now,

fasting katabolism
this
instead of 6.0 Therms, i.e.,at which
a

directly
in-

or
directly

which

just sufficient to maintain

of metabolizable
remaining 4.69 Therms
superfluous heat production. Suppose,

necessary

body

the

When

obviously all

of mixed

will be

heat

the

otherwise

produced by

grain and alfalfa hay used

the previousparagraph be fed to a steer whose

if the ration

Thus,

of tissue.

katabolism
low

so

would

which

the

would
amount

be
of

The
body temperature.
body stillyields6.0 Therms, as before,
the

normal

NUTRITION

312

To

the

put

when

that
critical

external

in

summer

be

may

insufficient

available

less
for

for

swine

the

(377)

be

is

less

when

fed

than

that

therefore,

of

reported
fattening

wide

for

it contains

to

the

meet

mand
de-

the

temperature

experiments
the
the

of
animals

supplying
as

U.

source

S.

Dept.

much

are

net

digestion
so

that,

energy

"

63

"

is

gains

of the

very

due

feed

for
at

the

sufficient

somewhat

No

exact

of

of heat

ently,
Appar-

rather
a

and

for maintenance

low,

relatively
of feeding,

therefore
is also

of heat.

Agr., Bur.

Anim.

Indus.,

Bui.

108

of

sake

olism,
katab-

ordinary conditions
surplus

out
with-

in the

is

source

be

may

the

reported.

ruminants

that

ration

maintenance.
been

made.

were

F., the

ferences
dif-

to

shown

increase

no

by

energy

be

have

comparisons

have

under

producing

part

maintenance

below

of

tenance
main-

the

on

may

Fries

of

showed

also

tures,
tempera-

confirmed

experiments

and

of

temperature

heat,

(378)

likely

actual

exposure

katabolism

series

is

of metabolizable

the

temperatures

of

work

amount

these

(560 F.)

critical
"

single

lower

at

of

the

increased

animals,

low

be

to

seems

of swine

an

rations

on

the

neighborhood

of
A

increase

amount

below

these

to

possible that

which

ture
tempera-

(68"-73"F.),

high

Exposure
belief

seems

at

critical

concentrates,

on

to

by Armsby

cattle

the

digestion by

influence

in the

considerably

lower

ample

It also

indication

even

this

the

upon

in

very

ration

the

for maintenance

it fails

wholly

expected

swine, and

on

production.

be

temperature

heat

large

ration

because

not

that

in

or

maintenance

the

temperatures

while

because

energy

observed

Experiments

any

winter,

sider
con-

comparatively high,

is sufficient

of ruminants.

swine.1

in the

reduced

is

comparatively

chiefly

results

variations

required

at

of

may

of

ration
the

maintenance

horse

showed

likewise

expenditure

especially
to

Hagemann

heat.

Tangl's experiments
while

in

but

energy

the

which

ration

and

Zuntz

ordinary

for

temperature

that, for example,

ANIMALS

way,

the

receiving

so

any

another

in

matter

FARM

OF

(1908),

pp.

84-86.

CHAPTER

IX

{Continued)

MAINTENANCE

"

THE

OF

REQUIREMENTS

MATTER

As

pointed out

was

in the introduction

the

in

presence

previous Chapter

requirement is a twofold

(361),the maintenance
for the

to the

feed

of

one,

adequate amounts
an
adequate supply

of

calling
certain

of energy.
specificsubstances as well as for
in some
The former phase of maintenance
of its broader aspects
stances
forms the subject of the present Chapter. These
specificsubof this discussion as
be grouped for the purpose
may
proteins or their constituents,ash ingredientsand accessory
constituents.

"

The

i.

Nature

398.

of

Requirements

protein

"

Maintenance
As

was

shown

in

VII

of

protein as the necessary

and
tear
(Rubner's " wear
that

molecule
the

requirement.

for

(340),the loss of protein which the fastingbody

be rebe interpreted
in two
garded
First,it may
may
ways.
of a certain amount
to the complete breakingdown
as due

Chapter
suffers

Protein

certain

may

body, so

protein may

atomic

of cell activity
accompaniment
quota). Second, it may be supposed
in
the
contained
protein
groupings
"

functioningof
indispensablefor the normal
that, if they are not contained in the feed, body
be katabolized
for the sake of obtainingthem.
be

either case, it is clear that what

the feed must
supply in
order to maintain
the body in nitrogen equilibrium is not,
In

strictly
speaking,protein as such, but materials whose digestive
and proportions of the constituent
cleavagewill yield certain amounts
the requirements
amino
acids.
On the first hypothesis,
stantially
subfor the different
be determined
would
buildingstones
in the body tissues
by the quantitiesof each existing
"

"

313

OF

NUTRITION

314

ANIMALS

FARM

According to the second hypothesis, it might

contained
be presumed that only certain of the atomic
groups
in protein,
need to be supplied.
such as tryptophan, e.g., would
not
Moreover, it appears
unlikelythat both hypotheses may
both as a whole
be true and that body proteinis katabolized
of obtaining certain amino
acids.
If
and at times as a means
all the
of the
such is the case, substantially
buildingstones
be manufactured
in the body,
proteins,so far as they cannot
of certain
be supplied in the feed, but relativelymore
must
particularones
might be requiredthan would be indicated by
the make-up of the body proteins. Finally,it seems
to be fairly
katabolized.

"

established

well

manufactured

that

of the

some

the

in

it becomes

even

clear

more

called,is reallyan

amino

399.

required for

so

Amino

acids

knowledge

of the

still meager,

but

and

amino

acid

amino

acids

be

can

certainlytrue of
arginin. If such be the
that the protein requirement,
requirement.

body. This
perhaps of prolin and

glycin and
case,

least

at

"

is almost

maintenance.

actual

Our

"

is
requirement for maintenance
shown
that a supply of tryptophan

acid

it has been

of

of

for the maintenance

tyrosin is necessary
while lysinis dispensable.
nitrogenequilibrium,

probably

Willcock

and

Hopkins, *

for

example,
maize, which contains neither tryptophan
supporting neither growth nor maintenance

that

found

mice,

zein

of

capable of

lysin,was

nor

in

the

while

tion
the addi-

of

tryptophan diminished
although it did not altogether stop
the loss of nitrogen from
the body.
similar
Henriques 2 obtained
It is to the
although less strikingresults in experiments with rats.
work

of Osborne
of

showed

the

better

results
a

similar results

mice.

was

the most

tryptophan
as

the

while

and

secured.

Furthermore,

conclusive demonstration

maintenance.

They

capable
inprotein was
the addition
of tryptophan
tenance
in two
cases
complete main4
has reported
Miss Wheeler

sole

with

for

Osborne

source

and

of

Mendel

have

shown

the deficiencies

the ration

to

zein

owe

we

obtained

were

long time
on

that

necessity of

maintaining rats,

for

that

Mendel

conclusivelythat
of

much

and

of zein may
be compensated for by the addition
of other proteins containing the lacking amino
acids.

Jour. Physiol. (London), 35 (1906),88.

2Ztschr.

Physiol. Chem., 60 (1909), 108.

Carnegie Institution of Washington, Publication
Chem., 13 (1912), 233; 17 (1914), 3254
Jour. Exp. Zoology, 15 (1913), 209.
3

No.

156 (191 1); Jour. Biol.

MAINTENANCE

"

REQUIREMENTS

OF

MATTER

315

the other

hand, they have also shown that lysinis not essential

animals
proteinmaintenance, they having been able to maintain

On

to

for

which
long periodson rations containingas their sole proteingliadin,
contains no lysinbut does contain tryptophan.

What

has been

shown

regarding the necessityof tryptophan

of lysin for maintenance
and the dispensability
doubtless
may
for other protein constituents,so that ultimately
to be true
prove
it may
be possibleto estimate the relative maintenance
values of proteins on
the basis of their chemical constitution.
At present, however, this is far from
The
being the case.
of the proteins,particularlythose of the
constitution of many
roughages,is known
inadequatelyor not at all,while the specific
amino
acid requirements for maintenance
have stillto be worked
out
and may
conceivably vary as between different species.
400.

Relative

values

of

facts

regarding the variations

proteins which are recorded

proteins for

maintenance.

in the constitution

"

The

of the different

Chapter I (50) render it evident

that these substances may
be of quite unequal value as sources
of amino acids to the organism. Thus, accordingto the data
there tabulated,gliadinand zein would be about
three or four
times as valuable
acid
of the amino
as
legumin as a source
prolin,while on the other hand legumin would be 2J times as
cereal
valuable as wheat
of lysin. The
glutenin as a source
proteins,especiallythose of wheat, are notably rich in glutamic
acid and therefore relatively
in other constituents.
If,
poorer
is in realityan
then, the protein requirement for maintenance
amino
acid requirement it would
that these various proseem
teins
be of unequal value for that purpose.
must
As
regards singleproteins,the experimentalevidence just
and
cited strongly supports this presumption,while Osborne
Mendel

have

likewise shown

in

the existence of distinct differences

lactalbumin, casein edestin,gliadinand milk

It must
of rats.
not be forgotten,
proteinsfor the maintenance
and domestic
animals ordinarilyconsume
however, that both man
be presumed that
of proteins,so that it may
a mixture
deficiencies or excesses
of particular" building stones
pensate
comin the values

of

"

the

for each other to.a greater or less extent.

On the whole
statement
seems
justifiedthat while distinct differences
1

Jour. Biol. Chem.,

22

(1915),241.

316

have

for maintenance

value

in many
to
cases
known
differences

be

ANITvL\LS

different

proteinsfrom

mixed

between

FARM

OF

NUTRITION

been

less than

shown

in their chemical

so

greatly to
to be

to

desired,the only

follow

established

in the belief that

but
so

great

to

as

such

anticipatedfrom

the

further

As

the

matter

investigations

for the present

open
and deal with the tein
custom
prothe consciousness
as
a whole, with
course

feeding stuffs and rations

that it is of unequal nutritive

of

regards their
exist,they appear
as

constitution.

necessity,then, pending
practical

seems

to

be

might

of

be

sources

value

differences

in different
in all

not

are

materials

probability

seriouslyinvalidate the generalusefulness of the

results.

proteinkatabolism

Influenceoffeed supply on
401.

minimum

The

(339)

minimum

fastinganimal

below

cannot

feed

of

which
be

The

protein.
"

physiological

protein katabolism
evidentlyconstitutes

of

the

reduced

limit to the necessary

supply of feed
if any, above
be
this minimum
must

protein,but what
suppliedin order

the

lower

surplus,
to

secure

is stillan unsettled question. That

protein maintenance
the amount
of feed protein necessary
for maintenance
atively
is relsmall has been fullydemonstrated.
It appears
to be
well established
diet containing an
also that on
abundance
a
of non-nitrogenousnutrients,especially
of carbohydrates, a
supply of protein materially less than the proteinkatabolism
of the
the needs
duringcomplete fastingis sufficient to meet
little or no greater
organism, while it is possiblethat an amount
actual

than

that

consumed

katabolized

when

abundance

of

will suffice.

Fats

to be distinctly
less efficient than
appear
at the minimum.
keeping the protein katabolism

is the

case

advanced
to
amount

of

carbohydratesis

fullymade out, although Landergren l has

of carbohydratesis essential
explanationthat a minimum

has
the

carbohydrates in
Preciselywhy this

not

been

the chemical

is not

and that when

a sufficient
processes of metabolism
supplied in the feed,protein is katabolized for the sake

producing carbohydrates,

diet nitrogen katabolism

1

with

the

result

that

is increased.

Jahresber. Tier. Chem.,

32

(1903), 685.

on

low

protein

MAINTENANCE

OF

REQUIREMENTS

"

MATTER

317

Chapter VII (335-338),however, make

be affected by the
it evident that the protein katabolism
may
of both proteinand non-nitrogenousmaterial
amount
available
in the body.
For a correct
periments
interpretationof the results of exthe maintenance
fore,
requirement of protein,thereupon
of the feed supply upon
the
a knowledge of the influence
proteinkatabolism is essential.
While a relatively
402.
small
Surplus protein katabolized.
in the presence
of an
protein is sufficient,
quantity of digestible
abundant
the body in
supply of fuel material, to maintain
nitrogen equilibrium,an increase of the feed protein above
this minimum
does not result in any large or long-continued
creases
gain of protein tissue by the mature
animal, but simply inthe protein katabolism, as
is shown
by the prompt
of a correspondingamount
of nitrogen in the urine.
appearance
facts recorded

The

in

"

Table

The

57.

fact

Protein

"

demonstrated

was

in collaboration
tenkofer

the

at first with

more

Sheep

of

than

Bischoff

in

animals

numerous

Gesetze

Published

der

but

of

man

and

on
investigations

Day

per

Head

and

by C.
ago
later alone and with

fiftyyears

and

experiments on carnivorous
have
subsequent investigations

only of these
Of

Katabolism

that

of herbivorous
herbivora

Pet-

almost

merable
innu-

it is true

not

animals, and
shown

Voit

animals

in which

the

as

well.
nitro-

i860.
Ernahrung des Fleischfressers,
der Chemie
und Pharmacie
and the Zeitschrift
chieflyin the Annalen
See also Voit, "Physiologie des Stoffwechsels," in Hermann's
Handfur Biologic
buch der Physiologie.

318
gen

NUTRITION

excretion

OF

has been

sheep were
To
barleymeal.

hay
periodsvarying amounts
glutin (of lupins)or of
from

digested

the

latter increased

of

of

flesh
with

protein cannot
obvious
sufficiently
the weakling cannot
feeding him upon a
size

by

of the

very

mature

animal

basal

an

ample.1
ex-

ration

of

protein in the form of conmeal.

A comparison of the nitrogen
the urinary nitrogen shows
that the
with
substantially
parallel

converted

abundant

as

in the intermediate

That

"

the

mere

from

meat

added

large storingup of
daily experience. The

cause

be

were

protein limited.

of

serve

57 may
and 7

nearly pure

diminished

Utilization

403.

this ration

ration

and

ANIMALS

determined, Table
fed in periods

Two

and

FARM

those

into

the former.

tion
consumpit is indeed
muscles

of

of the athlete

by

increased
in
diet,nor the small man
protein supply. The proteintissues

have

reached

their natural

limit

of size

and

consequently the capacity of the body to store up protein

is limited.
Beyond the minimum
requiredto make good the
katabolism
in the cells,protein can
be utilized in
necessary
such an animal
fore
only to a small extent as protein,and it is thererapidlykatabolized,its nitrogen appearing in the urine as
and
other familiar
end products. Nor
is the situation
urea
essentiallydifferent in the growing or the milk-producing
animal.

these

While

animals

able

are

to

utilize

erable
consid-

of feed

protein,yet the limit to this utilization

is set by the normal
of growth of the protein tissues or
rate
the capacity of the mammary
the
glands to manufacture
casein and
other
Any surplus of proproteinsof the milk.
tein
amounts

what

over

be

can

used

preciselyas is a surplus over

animal.
unproductivemature
Chapter XIII, " 4.)
404.

Protein
of

as

of

source

protein,however,

for

this

purpose
small
very

the

is katabolized

of the

demand

(Compare Chapter XI, "

energy.

is not

to

"

be

This

increased

regarded as

and

olism
katab-

the

total

surplus of
acids
protein, the amino
digestion are in
(233), their nitrogen being excreted
large part deaminized
chieflyas urea, while a non-nitrogenous residue is left which
contains the largerportion of the chemical
energy of the protein

loss of

so

which

it represents and

much

feed

Henneberg

material.

the presence
resultingfrom its

and

In

is in condition
Pfeiffer;Jour. Landw.,

to

38

be

of

oxidized

(1890), 215.

as

fuel

OF

NUTRITION

320

FARM

ANIMALS

protein supplied in the feed,while

for each level of protein stock a certain supply in the feed is
that is, the protein requirement for maintenance
necessary
With
carnivora
varies.
a
on
largelyprotein diet the adjustment
the
take
of the body to
to
protein supply seems
place
of herbivora,however, the adrather promptly. In the case
justment
to be more
gradual, possibly owing to the
appears
relativelylargesupply of non-nitrogenous ingredientsin their
feed,and apparently some
gain of protein may continue for a
considerable
time, although when expressedas a percentage of
either the total feed protein or of the body protein the gain is
small.
relatively
according to the

of

amount

"

407.

The

Effect

prime

of

deficiency of

demand

of the

of its vital
its own
upon
the laws of

non-nitrogenous

organism
functions,and

is for energy
if necessary

tissues for this purpose.

be
can
protein metabolism

into consideration

the energy

No

nutrients.

"

for

the formance
perit will draw

conception of
without
taking

clear

reached

relations.

alreadybeen shown, the proteinsor at least the cleavage

products of their digestionreadilyundergo a process of deaminization by which their nitrogenis split
off and excreted,leaving
As has

of
is available
a
source
as
non-nitrogenous residue which
however, the proportion of energy derived
Ordinarily,
energy.
from
the katabolism
of protein is relativelysmall, the nonsource.
nitrogenous nutrients constitutingits principal
But if,with an
of protein in the feed just sufficient
amount
the non-nitrogenousnutrients,
to sustain nitrogen equilibrium,
that
especiallythe carbohydrates, be so reduced in amount

the

total

supply is insufficient for maintenance,

energy
but
to make
only is body fat drawn
upon
up the deficit,
also increases,so
that a supply of
protein katabolism
nutrient
and

The

which

loss of

previously adequate
body protein occurs.

effect is

nutrients

was

naturallymost

marked

when

became

not

the
this

insufficient

the non-nitrogenous

altogether. For example, Voit and Korkunoff found that when

a dog was
supply of carbohydrates,
given an abundant
protein equivalentto about 4.5 grams of nitrogen was sufficient
him in nitrogen equilibrium. But when a similar amount
to maintain
are

withdrawn

^tschr.

Biol.,32 (1895),67.

MAINTENANCE

given without non-nitrogenous

protein was
and about
entirelyinsufficient for this purpose
was
required to attain protein maintenance, as
:

Table

three

it

times

the

32

nutrients

of

shows

MATTER

OF

REQUIREMENTS

"

proved

as

much

followingtable

"

58.

"

Effect

Supply

Protein

of

on

Katabolism

Protein

of

Dog

Feed

Nothing
Extracted

(grams)

meat

100

140

165

185
200

230

360

......

410

360

The

results furnish
between

also

proteinkatabolism

proteinsupply and

demonstrated

than

more

of BischorT and

Voit

of the

strikingillustration
30

years

earlier

by

which

classic

the

rendering it evident

(402),while

relations
interesting
had

been

experiments
quantity

that the

proteinrequiredto produce nitrogen equilibriumwhen fed alone

demand
of the body.
the minimum
very far from representing

of

What

is

so

nutrients

also in less degree in

holds

good
relative deficiency.If a portion
are

of

in the total absence

true
strikingly

withdrawn

from

of

mixed

is

enous
non-nitrog-

case

of their

trients
non-nitrogenous nuolism
ration,the protein katab-

the

usuallyincreases.
408.
on

the

Effect

of

surplus

non-nitrogenous

converse

diminish
deficient

of the statement

of

the katabolism
in energy,

of the

nutrients.

be

contrary, non-nitrogenous nutrients

ration,they tend to
As regards rations
the

of

this

If,

"

added

to

protein.

is,of

course,

precedingparagraph

only
that

and

katabolism,
rations

the

of

addition
a

ration

experiments

cattle

on

which

rations

Table

ration

59.

"

in

with

Effect

of

to

Starch

the

proteintends

level.

The

of Kellner's

starch

cause

effect

respiration

added

was

basal

to

fattening.

some

the

urinary nitrogen upon

augmented

on

of

lower

those

the

the

upon

that

found

the minimum

sufficient to

compares

that

been

in the

even

of carbohydrates,
particular,

which

themselves

were

following table

basal

than

proteinkatabolism
for example,by
illustrated,

is well

The

or, in

protein

submaintenance

or

it has

the

reduce

to

surplus of fat
containingmore
a

increase the

to

equivalent. But

are

supermaintenance rations

of

case

statements

two

tends

maintenance

regards

as

ANIMALS

FARM

materials

of these

the withdrawal

to

OF

NUTRITION

322

Protein

ration

"

Cattle

Katabolism

of

Daily

Nitrogen

Urinary

OxD
OxF
OxG
OxH
Ox

It has

only by

likewise
the

been

shown

fats and

the

this effect is

soluble

produced

not

hexose

carbohydrates,
but likewise by the pentoses and,
as
in the case
of herbivorous
animals,by those ill-known ingredients
of feedingstuffs,
especiallyof the crude fiber and the nitrogenfree extract, which
disappear in the passage of the feed through
the alimentary canal
which
and
are
commonly spoken of as
also the organic acids,
being digested. This statement
covers
whether
resultingfrom the fermentation of the carbohydrates
such

or

starch

contained
409.

should

true

and

the sugars,

in the feed.

Protein

be

by

that

katabolism

depends

clearlyunderstood

surplus of fat

or

that

even

chiefly
in

on

the

carbohydrates the dependence

1

Landw.

Vers.

Stat.,53 (1900).

supply.

"

It

of a
presence
of the protein

MAINTENANCE

katabolism
the

most

323

protein supply still holds true.

liberal supply of non-nitrogenous nutrients
the

upon

and
splitting-off

prevent the

proteinwhich
reduces

MATTER

OF

REQUIREMENTS

"

below

it somewhat

cannot

of the

nitrogenof surplus
previousparagraphs,but simply

excretion

illustrated in

was

Even

the level which

it would

otherwise

that extent, it helps to bring about, and probably

tioned
prolong somewhat, the temporary storage of protein mena previous page
on
(406)and thus to bring the animal

reach.
to

To

a higher plane of
upon
the
It is clear from

is

be

to

protein nutrition.
that

foregoingstatements
of between

conceived

tinction
sharp dis-

no

and
insufficiency

an

of non-nitrogenousnutrients,but
sufficiency

rather

tendency
diminish the protein katabolism,a
on
in
less marked
or
accordingto their abundance
tendency more
that no
It is not
to be understood
the ration.
nitrogenous
for
fuel
is
katabolized
material
as
long as sufficient
purposes
non-nitrogenousnutrients are present to supply the demands
a

the part of the latter to

for energy,

prevent

can

that

nor

the

Protein
410.

Minimum

use

by

the

body.

requirementsoffarm animals

and

optimum

protein requirements
it is

of

katabolism

possibleconstructive

largestquantitiesof the latter

of its
protein suppliedin excess

the

even

of

protein.
"

of the different

between
important to distinguish

In

the
considering

speciesof farm animals,

On
two
points of view.

be sought to determine
the least amount
hand, it may
of feed protein upon
which the protein tissues of the animal can
be to
be maintained.
On the other hand, the endeavor
may
of protein to supply
the most
formulate
advantageous amount
the

one

when

an

amount

may

minimum.

actuallyto be maintained for a time and this

very possiblybe greater than the physiological
first point of view, however, is plainly the
The

fundamental

one

determined
be
a

is

animal

and

should

the lower

first receive consideration.

limit of

ing
Hav-

protein supply,it will then

the advantages,if any, of

possibleto consider intelligently

surplus.
In considering
the

determination
individual

or

results of

of the minimum

species,it

experiments directed toward the

of feed proteinrequired by any

is essential to

bear

in mind

the facts

NUTRITION

324

katabolism

which

The

411.

of

influence

the

regarding

OF

have

plane

FARM

the

feed

just been

of

protein

ANIMALS

supply

upon

the

has

been

protein

considered.

nutrition.

"

It

shown

previous paragraphs that

itself more
or less promptly to

the

in its feed

by

the latter to be the least amount

necessary

for the

protein katabolism
adjusts
the supply in the feed.
A surplus
the minimum
above
requirement, while causing a small storage
in raisingthe plane of proteinnutrition
of protein,results chiefly
until income
and
and
so
increasingthe katabolism
outgo of
into equilibrium. The
mere
fact, therefore,
nitrogen come
is in equilibriumwith a certain supply of protein
that an animal
in

livingupon
412.

also been

supply

shown

depends
the

body,

proves
maintenance

but

that

not

of

non-nitrogenous

the

of
sufficiency

only

upon
within
also,

non-nitrogenous nutrients
abundant
an
supply of the
the fastingkatabolism, or
sufficient minimum

to

be

of it is diverted

more

for

former

amount

an

perhaps

reach
use

"

It

has

supplied with

is reduced

required to

nutrients.

of progiven amount
tein
the plane of protein nutrition
of
certain limits,upon
the amount
of

for maintenance.

nitrogenousmaterials
seems

of the

animal, since it may be

high plane of proteinnutrition.
unnecessarily

an

The

means

no

as

of

the
of

protein. With
proteinequalto

less,appears

even

to

be

supply of nonlargersupply of feed protein

As

the

and
more
equilibrium because
fuel,so that in the total absence
of protein must
a
large excess

non-nitrogenous nutrients
be fed before equilibrium between
and outgo of nitrogen
income
is reached.
In interpretingexperiments or formulating a
maintenance
ration,therefore,it is not sufficient to consider
of protein,but account
also be taken of
must
simply the amount
the supply of non-nitrogenous materials,and
only when the
of the ration is ample for maintenance
content
net energy
can
the protein
that a loss of body protein shows
it be concluded

supply to
413.
of

Value

of

non-protein.

"

The

crude

proteinof

the feed

protein but a great

venience
variety of other nitrogenous substances, grouped for conthe designation non-protein. In considering
under
the results of experiments upon
the protein requirements of
whether
it is necessary
these animals,therefore,
to determine
the

farm

be insufficient.

true

animals

includes

proteinshould

be

not

only

true

the basis of

comparison or

whether

MAINTENANCE

the

REQUIREMENTS

"

non-proteinhas

tissues of the

body.

It appears

to have

value

some

OF

for

MATTER

325

maintaining the protein

been demonstrated

by recent experimental
results,
tory
especiallyby
Kellner,Morgen, and the Laborafor AgriculturalResearch
in Copenhagen, that the nonprotein
of ordinary feedingstuffs is available for the maintenance
of ruminants, probably indirectlythrough a
conversion
to
of
protein by means
micro-organismsin the digestive tract
have
(141). On the other hand, investigations
thus far failed
to demonstrate
that non-protein has any
material
value for
other species or for production purposes
(786-789). In the
computation of rations for productive feeding, therefore,it
those

appears

the

desirable

for

digestibletrue

implies,however,

of

the present

to

consider

protein,ignoring
that

the

results

of

non-protein. This
the
experiments upon

proteinrequirement shall be expressed in

This

will have

ordinarilyonly

the

the

same

manner.

effects :

First,it will make the protein requirement

is.
really
Suppose, for example, that
appear
series of trials in which
the
ratio of digestiblenon-protein to
a
digestible
proteinis 1 : 10 shows that nitrogenequilibriumis reached
with a ration
supplying 500 grams
protein.
nonprotein and
50 grams
Regarding the true proteinonly,the maintenance
ment
requireis 500 grams,
while includingthe non-protein it is 550
grams.
In the second
will be largelycompenplace,however, this error
sated
for when
the actual computation of rations is also based on
the
in the case
true protein. Thus
ration
justsupposed,if a maintenance
be computed from
feed or mixture
in which
the ratio of nonprotein
any
is
the
in
the
to protein
as
same
experiments from which the
maintenance
it
is
obvious
final
was
that the same
deduced,viz.,1 : 10,
result will be reached
whether
the maintenance
be
sidered
conrequirement
of true proteinor 550 grams
to be 500 grams
of crude protein.
Only when the proportionof non-proteinto true proteinvaries widely
from that existing
in the rations used in determiningthe protein requirement
will any significant
arise in computing rations.
error
two

smaller

than

it

In the results considered

on

succeedingpages,

both

the crude

protein and true protein of the rations are stated when

are
given in the reports of the experiments.
414.

Computation

to unit

weight.

"

It

was

shown

in

these

ter
Chap-

is
(345) that the energy requirement for maintenance
substantially
proportional to the body surface of the animal.
VII

326

comparisons of

similar

No

does

certain
of the

the

protein requirement appear

protein
Since, however, the minimum

made.

been

have

ANIMALS

FARM

OF

NUTRITION

not

represent

quirement
re-

for

but
for
energy
normal
functioning

demand

to

required for the

specificsubstances
plausibleto suppose that
body, it seems

will

its amount

of active tissue than upon the body

upon the mass
be the case, the protein requirement may,
If such
surface.
be computed in
for practical
with sufficient accuracy
purposes,

depend rather

proportionto
the

the

live

weight

and

that

course

cattle.

For

is followed

in

succeedingparagraphs.
Protein

415.

requirement

it is impracticable
to ascertain

of
the

"

obvious

reasons

of ruminants

fasting katabolism

estimating their maintenance

requirement
but by a comparison of the recorded
as
ments
experiregardsprotein,
small amounts
of feed
in which the nitrogenbalance upon
it is possible
to fix approximately the limit
has been determined
of an abundant
in the presence
of proteinsupply below which, even
a loss of body protein
nutrients,
supplyof non-nitrogenous
as

basis

for

occurs.

tenance
the energy
requirement for maininvestigations
upon
in Chapter VIII (381)only those of Kellner
summarized
and
the live weight experiments of the writer,together with
the early results of Henneberg and
Stohmann, afford data
protein requirement. While protein
regarding the minimum
maintenance
was
probably secured in the remaining instances
that a surplus of protein
there is no sufficient evidence
to show
not being consumed
was
(402,411). In addition to the foregoing,
the investigations
by the Laboratory for AgriculturalResearch
in Copenhagen 1 upon
the proteinrequirements for milk production
data
the
tenance
mainafford
also
to
as
(586)
approximate
the
in experiments upon
requirement,and Fingerling,2
esting
protein requirements of growing calves (471),obtained intertenance.
indications regarding the quantity requiredfor main-

Of the

The
upon

lowest

which

recorded

amounts

per

nitrogenequilibriumwas

1000

pounds

reached

were

live
0.21

weight
pound

fra den Kgl. Veterinaer

of Landbohojskoles Laboratorium
Denmark-Beretning
Translated
Forsog. 6ode, 1906, and 63de, 1907, Kobenhavn.
de
Rationelle
du Betail.
Compte Rendu
by Mallevre, Societe de l'Alimentation
et i2eme
1 1 erne
Congres.
2 Landw.
Vers. Stat.,76 (1911), 1.
1

for landokonomiske

328

NUTRITION

OF

ANIMALS

FARM

growth of wool is considerablygreater than

after all,
that of cattle for the growth of hair,the absolute difference,
does not add very greatlyto the total maintenance
requirement.
of sheep call
standards
the
maintenance
for
The
current
feeding

quirement of sheep

for

1.

lb. of

0-1.6

for the

digestiblecrude

protein

per

pounds live
Stohmann's
early

1000

weight,apparentlyupon the basis of Henneberg and

protein or 1.041b. true
experiments (382) in which
1.3 lb. of crude
proteinproduced but a slightgain of body protein in addition to the
be littledoubt, however, that HenneThere
berg
can
growth of the wool.
and Stohmann's
sheep received a surplus of protein above the
actual maintenance

requirement.

digestionand metabolism
experimentsby Schulze
and Marcker,1 decidedlysmaller amounts
of protein proved sufficient
of 6 experiments in
to maintain
nitrogen equilibrium, the average
which
observed
loss of body proteinwas
no
being 0.653 lb. digestible
crude proteinper 1000
is
It
live
weight.
evident, then,
pounds
much
be reduced
that the protein supply of sheep can
below
the
fed in Henneberg and Stohmann's
ing
amount
experiments without leadto a loss of body protein.
In

series of

20

data regarding the minimum

satisfactory
ment
requirein
of sheep are afforded
by Katayama's2 investigations,
of nearly pure protein (" aleuronat
amounts
")
increasing

The

most

which

added

protein,consistingof
The proteinin every case
hay, oat straw, starch and cane sugar.
of starch,so that
substituted for a corresponding amount
was
the total energy of the ration remained
substantially
unchanged.
On the average
of two animals, 0.41 lb. digestible
true
protein
the
sufficient to maintain
pounds live weight was
per 1000
nitrogen balance.
Since, however, the growth of wool must
have gone on, with a corresponding storage of nitrogen,there
were

to

been

have

must

tissues of the
If to
per
that

an

ration

very

in

poor

equivalentloss

of

proteinby

the

active

body.

the minimum

lb.
of 0.41 pound there be added
0.14
live weight for the growth of wool, it appears
of
protein requirement for the maintenance

pounds

1000

the minimum

sheep

mature

to

the

basal

body

note

is in the
that

tissues

is

neighborhood

the

actual

of

0.55

maintenance

apparently quite

as

low

lb.

It

esting
is inter-

requirement
as
relatively

cattle.
1

Wolff

Landw.

; Die

Ernahrung

Vers.

der landwirtschaftlichen

Stat.,69 (1908), 321.

Nutztiere,

p. 300,

for
for

MAINTENANCE

Protein

417.

of the

an

the

OF

REQUIREMENTS

requirement

fastingkatabolism

(377)gave
weight for
with

"

of

swine.

of swine

MATTER

The

"

recorded

329

determinations
in

Chapter VIII
of 0.48 lb. per
thousand
pounds live
average
of swine, although
fastingprotein katabolism

considerable

range

in the individual

results.

McCollum

olism
figuresfor the protein katabof
of swine receivingno protein but fed liberal amounts
of twelve
experiments being 0.26 lb. per
starch,the mean
1000
pounds live weight with a range of 0.14 lb. -0.33 lb.
these are sufficient
as
Whether, however, such small amounts
above
them
is
for actual maintenance, or if not, what
excess
has not been certainlydetermined.
necessary,
has

reportedconsiderablylower

experiments of Von d. Heide and Klein (378),in one of

ration was
fed to three young
which
an
approximate maintenance
The
material gain of proteinby the animals.
a
swine, there was
of nitrogen
amounts
actuallykatabolized,however, as shown by the amount
excreted
in the urine,were
follows for the three animalsas
together:
In

the

"

ration of swine
Dietrich,2 in his experiments upon maintenance
found
that
(378),
proteinper thousand
0.70 to 0.84 lb. of digestible
live
pounds
weight sufficed to produce nitrogenequilibriumin two
periodsfollowingan eight-dayfastingperiod,but that about the same
amounts
(0.80 to 0.90)previousto the fastingperiodwere
insufficient,
and
while in two trials in which
lb.
.06
sumed
con1
were
respectively
0.94
reached.
was
proteinmaintenance

418.

by

Protein

Grandeau

requirement of the horse.

and

LeClerc
1
2

described

in

Expt. Sta.,Research Bui. 21.

Ills.Expt. Sta., Bui. 163 (1913).
Wis.

In

the

experiments
Chapter VIII (386 d),
"

OF

NUTRITION

33"

of the

nitrogen balance

the

FARM

horses

ANIMALS

determined

was

during six

of
periods. The following table shows the amounts
proteinand of non-protein nitrogen digested in each period,
the small losses in epithelial
tissue
the urinary nitrogen,and
hoofs,hair,etc.):
(epidermis,
of

the

"

6i.

Table

Balance

Nitrogen

"

of

Horses

Omitting the results upon horse No. 3 in March, when the

exceptionallylow, the other five periods
protein was
digestible
show
the

an

average
crude
average

,1b.per
true

1000

daily gain of nitrogen of 1.33 grams,

or
protein digested was
235 grams,

pounds

live

weight,equivalentto

about

while

0.59
0.50 lb.

protein.

419.

The

optimum of protein.
"

The

data of the

foregoing
paragraphs

strikinguniformity in the minimum

animals
speciesof domestic
proteinrequirement of the principal
with perhaps the exception of the hog when
mature, 0.4 to 0.6
tain
lb. per 1000
pounds live weight apparently sufficingto mainnitrogenequilibriumunder favorable conditions.
be clearlyunderstood, however, that this figure
It should
limit.
It purless accuratelydetermined
or
ports
represents a more
seem

to

be
animal

be

to

the

reduced

indicate

below

amount

eventual

without

body, however,

proteinsupply above

the

which

the

proteinsupply
protein starvation.

adjust itself to
minimum, using some

may

not
can-

The

of
range
of it to increase
wide

MAINTENANCE

the stock of

proteinin

fuel material.

An

OF

REQUIREMENTS

"

the

body

and

MATTER

the
katabolizing

331
remainder

increase of the

protein supply above the

after a
minimum
tenance
relativelyshort time, the maincauses,
of the body protein at a higher level (411). The practical
in
actual
maintenance
is
far less as to the least
question
of protein which
be used than as to the most
amount
may
advantageous level of protein nutrition;that is, as to the
optimum of protein.
in connection
This question has been warmly debated
with
human
nutrition,having been brought to the fore especially
of Chittenden
and his associates.1
by the investigations
as

On

the whole

it cannot

be

said

that

considerable

surplus

the minimum
protein over
requirement for maintenance
that is,the maintenance
of protein nutrition on a high plane
has been proved to be of any
material advantage in the

of

"

"

maintenance

either

of

men

covering several

or

of

domestic

animals

during

low
periods
a
proteindiet through years or generationswould show a different
result is at present largelya matter
of speculation. It is to be
remarked, however, that the particularpoint under discussion
is the protein requirement of the mature
organism. That a
of protein in the diet of a growing animal
have
deficiency
may
disastrous results is clear.
If,however, the habitual food supply
of a race
of men
of animals is low in protein,the
or
a
group
with the mature, and
are
likelyto share this deficiency
young
it seems
not
impossible that this is an important factor in the
of certain races
of men
allegedphysicalinferiority
livingon a
low protein diet. This consideration warns
to exercise care
us
in this
420.

months.

Whether

continued

of the breedingherd.
respect in the management
In the
Digestibilityof low protein rations.
"

maintenance
of the

feedingof
ration

farm

must

animals, the
also

be

matter

considered.

of the
It

has

actual

ibility
digestbeen

of protein in the
(723, 724) that a relative deficiency
ration tends to depress the apparent digestibility
of both the
protein and non-nitrogenousnutrients,especiallyin the case of
ruminants.
A maintenance
ration for these animals
containing
the minimum
of
amount
protein together with the quantity
of non-nitrogenous nutrients required to maintain
the energy
supply,would have a nutritive ratio (709)computed in the ordishown

Economy
Physiological

in Nutrition.

NUTRITION

332

FARM

OF

ANIMALS

of approximately i': 12.

On such
ration
a
way,
be some
loss of digestibility
and
would, in all probability,

there

nary

an

crease
in-

proteinby 50 per cent might perhaps effect a gain in

would
than offset the increased
which
more
digestibility
cost,
if any.
Indeed, unless feeding stuffs especiallypoor in protein
often be difficult,
it desirable,to
even
were
are
used, it may
of a maintenance
reduce
the protein content
ration to the low
level of absolute necessity.
of its

" 2.
421.

The

Ash

Ash

Requirements

Maintenance

for

ingredients indispensable.

the so-called mineral

That

"

supply of
of protein

ash

ingredients,as well as
and of energy yieldingmaterials,is necessary for the growth and
maintenance
of animals
has been fullyrecognized since the
time of Liebig, and
was
strikinglydemonstrated
by the well*
and
of Lunin,2 which
known
showed
experiments of Forster
that animals
suppliedonly with ash-free feed perished even
than

sooner

deprivedof

when

of the

Some

or

for these

reasons

of the functions
where

it

for both

all feed.
facts

of the nutrients

that, besides

was

shown

the

skeleton

and

the

indicated

were

in

their

soft

structural

tissues,the

in the body fluids is essential to

ingredients
of the vital processes.
Aside
and regulation
uses

of

singleelements, such

of the

and

pressure

of the various

the

422.

Ash
and

appear

at

than
ingredients

example,
content

causes

of the

relativelypoor
1

of feed

content

Ztschr.

an

as

large.
"

mixed

particularlythe

what
ash

ions,and,
neutrality.

preservationof

from

body
of

the

relative

Most

in ash

Biol.,9 (1873), 297.

specific

rations

contain

rations
much
*

of the

latter,

tain
animals, conof
largeramounts

made

production,for
the ash
upon
up of materials
than

larger amounts

Ztschr.

tration
concen-

of farm

sightto be much
body requires. Milk
even

tissues

however,
feedingstuffs,

exceptionallylarge drain

body, yet

the

fluids and

case
specific

first
the

of ash
presence
the maintenance

iron,fluorin,iodin,etc., three
there
ingredientsas a whole were
in the

osmotic

(268-272),
importance

as

general functions of the ash

mentioned, viz.,the maintenance
normal

Chapter

in the discussion

Physiol. Chem.,

are

(1881),31.

MAINTENANCE

in the milk

found

of
Kellner

for

62.

cow

"

Zuntz

MATTER

333

parison
gives the followingcomration recommended
by

ingredientsin a
producing 22 pounds of milk dailywith

of the milk

ash content

Table

produced.

ash

the

OF

REQUIREMENTS

"

Comparison

yield:
of

Ash

the

"

Content

of

Ration

and

of

Milk

Comparisons like the foregoinghave tended to confirm the

somewhat
spects
prevalent idea that rations adequate in other reof ash ingredients.
be assumed
to contain a sufficiency
may
but
This is doubtless true of animals livingin a state of nature
it is a questionableassumption under the artificialconditions
farm animals are subjected,as when
to which
receiving
many
of technical
of some
or
excess
an
singlegrain like Indian corn
when

by-products,or
423.

Ash

stimulated

to

ingredients digestible.
"

tion.
high degree of produc-

It is true

that of this relatively

in ordinary rations,
a very
largesupply of mineral matter
of certain elements, is found
considerable fraction,especially
in the feces,and this fact has led to their being regarded as
stated in Chapters III and
relativelyindigestible.As was
arises
IV (164, 199),however, this apparentlylow digestibility

from

the fact that

path

of excretion

the intestinal tract

for certain

constitutes

the normal

elements,notably, in the

case

of

has
Forster
herbivora, for calcium and phosphorus. Thus
shown that in the dog the calcium of the feed is largelyresorbed
1

Jahrb. Deut.

Landw.

Gesell.,1912,

570.

NUTRITION

334

in the
while

OF

FARM

part of the intestine

upper
more

where

less of it is excreted

or

ANIMALS

the contents

again in

acid,

are

the lower

intestine.

it is

While

of the
impossible,therefore,to determine
by means
much
of such ingredients
ordinary digestion experiment how
have
and
excreted
actually been resorbed
again and what
proportionhas escaped digestion,it appears safe to conclude
that at least

in the

resorbed

upper
rations as

of certain
the

considerable

Contrast

There

is

The

nutrients.

has been

and

that

dissolved and

the

insufficiency

is
ingredients

not

due

to

latter.

between

obvious

an

of them

digestivetract
regards mineral

of the
indigestibility

424.

share

organic

and

inorganic nutrients.

distinction between

former

be

may

said

"

organic and
to

inorganic
destroyed in the

be

of dextrose or of
performance of their functions. A molecule
for example, can
stearin,
yieldenergy to the body only by being
splitup and oxidized step by step to carbon dioxid and water.
The
is similar with protein so far as it is used for fuel purcase
poses
and even
its specific
functions seem
the cleavage
to involve
and oxidation
of its molecules.
With the electrolytes
contained
in the body the case
is different. A
of disodium
molecule
phosphate,for example (or its ions),is not destroyedby the
performance of its functions in maintainingneutralityno matter
how

long

chlorid
blood

it

that

serves

contributes

serum

as

it escapes
arise.

its quota

long

from

as

purpose

it remains

as

the

body

Losses
425.

Causes

of mature

will
which

the

be

various

course,

"

the

osmotic

dissolved

will the

of the
pressure
in that fluid. Only

need

for

fresh

supply

of ash

are

organs.

In

the demand

for

eliminated

is determined

in

the

from

body

growing animals there is in

ash ingredients
for structural

both for the buildingup of the skeleton

purposes,
in this case
degree of the soft tissues,but even

requirement

of sodium

molecule

So

elements

through the excretory

addition,of

the

far,therefore,as the maintenance

is concerned, the magnitude of the ash requirement
the
d
etermined
rate
at
substantially
by

of loss.

animals

to

and

largedegree by

and
the

the

to

total

rate

of

less

ash
cretion.
ex-

336

NUTRITION

of the

cells and

OF

FARM

ANIMALS

the

important part played by the ash

ingredientsin maintaining this neutrality.
Acidosis.
A varietyof chemical
normal
a.
processes, both
and
in the body which
tend
its
to disturb
occur
pathological,
neutralityeither by the addition of acid or the giving off of
alkali so as
known
to produce the condition
as
acidosis,by

body

to

"

which
A

is meant
well-known

diabetes.

relative

example

The

of

perverted

katabolism

certain

of

metabolism

by

extent

an

insoluble calcium

salts of the

ingredientsin

of ammonia

means

of acid

sources

accumulation

example

is the form

fatty acids

the feces.

are

be consumed
may
in fermented
materials

neutralized

juice,which

by

in the

milk

as

is

tralized
neu-

from

the

results
finally
of infantile acidosis

intestines and

It has been

in

in the

results in the formation

suggested

so

that

exceptionallyrich

exist in the normal

First, acids

products or

results

derived

gradual

failure of young
animals
to thrive on
in
be due
cause.
part to the same
may

Several

is that observed

of this disease

of fat in the food

excess

radicles.

oxybutyric acid (266),which

Another

coma.

in which

basic

of

proteinbut whose

in the diabetic

basic

over

pathologicalacidosis

production of large amounts

to

of acid

excess

of
moves
re-

the

in fat

organism.

such, either in natural

like silage. These
acids

the alkalies of the saliva

or

of the

pancreatic

thus

from
the body
temporarily withdrawn
fluids.
After resorption,however, the resultingalkali salts of
the more
acids are
common
readilyoxidized,yielding carbon
dioxid
and
and
water
restoring to the body fluids the bases
of some
Small
amounts
previously withdrawn.
acids,such as
tartaric and malic,however, tend to escape oxidization
and to be
excreted
in the urine,carrying a corresponding amount
of base
with them.
Oxalic acid and its salts are oxidized with difficulty
and
tend to impoverish the body in calcium
by the formation
of the

are

insoluble

calcium

injuriousto
while in

mature

fermentation

oxalate.

horses

ruminants

and
it

This

swine

seems

acid

and
to be

to

is liable to be
young

pecially
es-

ruminants,

largelydestroyed by

in the first stomach.

a
are
Second, the fermentations in the paunch of ruminants
of large amounts
of organic acids which, like those consource
tained
in the feed,may
withdrawal
from the
cause
a temporary
body fluids of alkali which is later restored when the salts are

katabolized.

MAINTENANCE

REQUIREMENTS

"

Third, the considerable

by

herbivora

makes

very

of

amount

OF

MATTER

337

hippuricacid produced

considerable

draft upon the organism

cited by
experiments by Diakow

Thus, in four
of
equivalent to from J to | of the total excess
Zuntz, it was
bases over
inorganicacids in the urine.
of the proteins,nucleo-proteins
Fourth, in the katabolism
other compounds
and
containing sulphur and phosphorus,
and phosphoric
these elements are largelyoxidized to sulphuric
acids. The sulphur of one
position
pound of protein having the comof serum
albumin, for example, if fullyoxidized,would
ounce
centrated
yield the equivalent of nearly one
by weight of consulphuric acid. High protein rations, therefore,
for bases.

tend to

bringabout

b. Neutralization
is

loss of bases from

of acids.

"

In

body.

all these various

ways

there

neutralityof the body

fluids and towards the establishment
of an acidosis,
to prevent
which
the acids must
be neutralized.
The
significanceof
this was
first shown
ferred
by the experiments of Lunin already reshowed
that the life of animals fed on
to (421), which
ash-free feed could be considerably
prolonged by the addition
of sodium
to neutralize the acids produced in the
carbonate
body. Normally, this neutralization is accomplished in two
general ways.
of acid may
be combined
with the ammonia
First,an excess
a

which

constant

is

tendency

produced

to

from

disturb

the

the

the amino

acids in the katabolism

of

into urea
in the
protein (233) and is subsequentlyconverted
liver. A
be diverted
part of this ammonia, however, may
from this course
and utilized to neutralize acids,the resulting
ammonium
salts being excreted in the urine in place of a corresponding
amount

of

the lower

in
putrefactions

The

urea.

ammonia

arisingfrom

the

the same
(140) may serve
A small quantity of ammonium
salts,
arisingfrom the
purpose.
neutralization of the acids produced especiallyin the protein
katabolism, is normally found in the urine,while the feeding
of inorganicacids or their injectioninto the blood stream, or a
acidosis,
pathological
greatlyincrease their amount.
]
may

On

the basis of

that the
to

intestines

earlyexperiments upon rabbits it has been taught

of ammonia
is peculiar
abilityto neutralize acids by means

carnivora

and

omnivora

and

is present to

very

limited extent

in

338
herbivora.
metabolic

It

seems

is

species. It should

is often

on

consequentlyrelativelyless ammonia

that

case

phase

the

shown

various

that the

available than

be

may

in

animals.

of carnivorous

Another

difference in the

olism
proteinmetablow plane,and
relatively

remembered, however,
animals

distinction between

fundamental

be

of herbivorous

ANIMALS

later investigationshave

should exist,and

such

no

FARM

prioriunlikelythat such

processes

there

that

the

OF

NUTRITION

of the

which

matter,

has

tion,
received littleconsidera-

long-continuedpresence of ammonium
of
have
salts in the body may
an
injuriouseffect. The possibility
injury through acid rations in this way could hardly be determined
of experiments,covering,if possible,
the whole life
except by means
cycleof the animal.
that
is the possibility

Second, an
with

the

has

blood

and
that

shown

disposedof by

present in the body.

bases

firstinstance,those
of the

be

of acid may

excess

fixed

the

contained
other

these

salts

These

Henderson,

phosphates

already noted,

as

present in the blood

are

in the

are,

and

in the carbonates

fluids.

combination

in

serum

of acids may
be
proportionsthat relativelylargeamounts
the reaction
disposedof in this way without materiallyaltering
such

of the blood.
c.

Excretion

in the
of

an

urine

body

of acids.

does

possess
shows
of Henderson
or

less of the
it

as

of base.

considerable
that

the

It is

of acids

The

skeleton

of bases

in the

of bases
as

familiar

produced

the excretion
fact that

kidneys

are

able

the bases
urine

and

to

separate

of the

the

work

degree of acidity. The

phosphoricacid from
acid phosphates in the

amount
428.

neutralization

involve
not, however,necessarily

equivalentamount
may

The

"

more

creting
blood, ex-

retaininga

responding
cor-

in the blood.
reserve

of ash

ingredients.
"

The

The larger
body fluids,
however, is limited.
part of the ash of the body is contained in the skeleton,which
constitutes
of basic phosphates and
a
large reserve
relatively
be drawn
carbonates which may
to supplement the supply
upon
in the blood.
This fact has an important bearing on the question
likewise
of the necessary ash supply in the feed,while it must
in experimental work.
be taken into account
Long-continued
maintenance
abnormal
feeds or under
conditions favoring
on
the
acid production in the body may
result in extractingfrom
to
of mineral
matter
even
body comparatively large amounts
store

MAINTENANCE

the

OF

REQUIREMENTS

"

MATTER

339

extent, apparently, of
while

the other

on

enable

may

losses,if not too extensive,to be made good.

is of specialimportance is that these fluctuations

such

point which

The

conditions,
bringingabout pathological
hand, normal feedingand conditions

of the ash content

of the skeleton affect the ash

as

whole.

composition of the bone ash as

II (81)remains practically
when
constant
even
given in
the skeleton has been greatly impoverished in total ash.
In
particularthis has been shown to be true not only of the calcium
and
gredient
phosphoric acid of the bones but also of the minor insuch as carbonic acid,magnesium and even
sodium.
A draft upon
the skeleton for sodium, for example, could be
of total bone ash
met
only by the mobilization of an amount
sult
containingthe requisitequantity of sodium, and this would reIt

was

found

by
Chapter

that the

Aron

throwing into the circulation relatively

largeamounts
of calcium
and phosphoricacid for which there may
be no requirement,
thus raisingthe percentage of these ingredients
above the normal limit and leadingto their excretion.
in

Maintenance
429.

Relation

indicate

that

chieflyon
for
in

by

one
excess

the

of the

the
reason

of

the

ash

the

"

The

of the various

amounts
or

balance

foregoingparagraphs clearly
requirements for maintenance
depend

feed.

to

of ash

another,

body's

excretory organs,

thrown

are

and

needs
and

ash

ingredientswhich,

into
are

the

circulation

therefore

furthermore, that

removed

the

nature

feed

the relative proportions

consumed, particularly
of its ash elements,is an important factor in determiningthese

losses.
430.

Deficiencies

in

ash

ingredients.
"

Some

feedingstuffs

especiallydeficient
in particularelements.
and familiar example
The most
striking
of this is maize.
According to Henry and Morrison * average
maize contains about 1.8 per cent of total ash,while its lime content
is only 0.02
per cent and that of soda only 0.04 per cent.
Some
dients.
by-productfeeds are similarly
poor in particularingreObviouslysuch feeds are not by themselves well adapted
contain

little total
relatively

Feeds

ash

and

are

and Feeding, 15th Ed., p. 672.

FARM

OF

NUTRITION

34-0

ANIMALS

growing or milkinganimals,in which a storage

the simple maintenance
of
For
product occurs.
for

which

however,

the small

whether

body. If
containing no
the

amount

the rate

maintenance
whether

far

so

whatever
as

how

largelyon
Acid

431.

and

is that which

of that
It is

"

is

being lost from

zero, a feedingstuff
to be adequate for

is concerned.
to be

In

general,
containing

regardedas

body

for maintenance

pends
de-

functions

termine
de-

which

element.

usually considered

that the

of feed in the respect justmentioned

of neutralityin the
to the maintenance

form

it bears

Feeding stuffs or rations containingin assimilable

much
sulphuror phosphorus,for example, tend to cause
phuric
of sulproductionin the body of correspondingamounts
and
phosphoric acids which must be neutralized. On
other hand, feedingstuffs containinglargeproportions
of

body

the

ash.

to

seem

it affects those

basic

imals,
an-

important relation

most

the

lime

ash element

some

of excretion

the rate

would

ration is

or

of

insufficient amount

an

which

that substance

feedingstuff

at

this loss could be reduced

lime

mature

the question
topicunder discussion,
of lime present, for example, is adequate

is the

depends upon

of ash in the

fluids.

the bases

tend

to have

the

oppositeeffect.

The

relation of acid

an
has,therefore,
importantbearingupon the
of a feedingstuff for ash maintenance.1
suitability
Feeding stuffs differ widely in this respect. In generalit

to basic

elements

be

may

said

phosphorus
amounts

and

low

that

the

concentrates

and

sulphur, little calcium

of potassium and
sodium, while
those of better quality,
rich in
are
in sulphur and
phosphorus. A

these differences

as

relativelymuch
and
only moderate
the roughages, especially

contain

calcium
definite

related to the maintenance

of

and

alkalies
of

measure

neutralityin

by converting the percentages of the

several ash ingredientsinto chemical equivalents.
Alfalfa hay, for example,accordingto Henry and Morrison,2
the

body

contains

in

the
1

is obtained

in

one

kilogram the

first column

of

amounts

the

in such

comparisons

as

well

as

shown
ingredients

followingstatement.

Evidently the sulphur and phosphorus present

included

of ash

the elements

in organic combination

present in the form

Dividing
must

In the older ash analyses the sulphuric acid represents only that part
sulphur remaining after the material has been ashed, which, as is now
known,
a

small
2

part of the

Feeds

and

total sulphur.
Feeding, 15th Ed., p. 672.

be

lytes.
of electroof the
is but

MAINTENANCE

OF

REQUIREMENTS

"

MATTER

341

equivalent weight gives the gram

in the last two
columns, showing that a
equivalentsshown
kilogram of this feed contains 1.300 gram equivalentsof excess
of each

the amount

by

its

base.

The

results of
Forbes

by

sort

equivalentsof

considerable

number

contained

in Table

are

bases

and

of
X

computationsof this
of the Appendix, the

meters
being expressed in cubic centisolution.
of normal
The table shows clearlythat some
like the hays, for example, contain a considerable
feedingstuffs,
of basic

excess

which

is

considerable
432.

laid

on

while
ingredients,

elements.

forming

The

specialcase

ratio of

others have

excess

an

phosphorus

to

of the ratio of acids

to

of acid-

calcium, too,
bases, shows

variations.

Significanceof acidityin
this distinction

Phosphoric acid is regarded

replacedby basic elements.
2

acids

Ohio Expt. Sta., Bui. 255.

between
as

ash.

"

Much

stress

feeding stuffs

neutralized when

two

with

of its hydrogen

had

been

acid
atoms

or

are

OF

NUTRITION

342

of domestic

that

all

regardsmaintenance.
extent
providedwith a
through its abilityto
and through the power

fact

point is

neutralize

and

that

important

an

basic

over

one

elements

hardlywarranted, especially
that the body is to a certain

of defense

means

against excessive acids

them

of ammonia
by means
kidneys to separate acids and
of acid-formingelements
an
excess

of the
that

show

is sufficient to

bases

seems

The

as

nutrition

of acid

excess

be avoided

in the diet should

human

the

Doubtless

animals.

assumption

the

but

of both

in discussions

alkaline ash

ANIMALS

FARM

necessarilyinjurious. It is only when

is so largeas to exceed the capacityof these regulative
the excess
and
when
it therefore
begins to draw on the
arrangements
it causes
the producfixed bases of .the body, or possiblywhen
tion
of
that
it
becomes
of
of largequantities ammonia,
a source
danger.
in the feed

of

ratio

Alkali

433.

is not

ash.

to sodium

the ratio of
have
It

potassium
important bearing on

an

was

sorbed

there

stated

into

the

blood

is

through the kidneys, it

sodium,

so

that

ration

effects if continued
in the

sodium

carry

in the

of

in other
well

rickets

to

sodium.

of normal

with
the

the

the stock

of

it has

been

be

that fodders

that

resultingin the disease known

(Rachitis)usuallyshow a misproportion of potassium
x
cites the following
Zuntz
comparisons of the ash
Along
hay with that of hays causing the disease.
of the bones

greater ratio of

somewhat

injurioushays

alkali

on

indirect

impoverished
phosphoricacid,as

would

found

as

less

fact

malnutrition

cause

serious

draft

and

In

or

tend
may
loss of sodium

have

skeleton,so that the bones

especiallycalcium
ingredients,

sodium.

as

Such

cause

of the

ash

more

draft,as already said (428),

of the total
corresponding amount

Such

skeleton.

it

in the former

might

to

with

along

latter.

appear,

long enough

the solution

involves

rich
relatively

impoverish the body

from
the body, it would

previousparagraphs,
stuff
a feeding
may
the body.
ash from

in

surplusof potassium salts repromptly disposed of by excretion

may

to

in

the losses of

while

that

indicated

As

"

ratio, as

show

appears

Jahrb. Deut.

calcium,

striking difference in the

very
from
the following table to which
cow's

milk

have

"

to

for
correspondingfigures

comparison :

phosphoricacid

Landw.

Gesell.,1912,

p. 577.

been

added

for

NUTRITION

344

FARM

OF

ANIMALS

to say, since

to determine

how

of any

in the

consists of

there is at present no way

found
ash ingredient
particular
much

how

and

undigested material

is to be

feces

much

regardedas

an

cretory
ex-

product. This being the case, a table like that of the

Appendix can give only a general and approximate idea of the
of its basic and acid
total quantityof ash or of the balance
of the alkalies in the materials actuallyresorbed
elements
or
enteringinto the metabolism.
Influenceof supply on excretion.

and

paragraphs (429-433) of the

ingredientsactually resorbed
potassium

and

to losses of calcium
to

considerable

the

the ash

phosphoricacid,thus possiblyaffecting

extent

Adding

excreta.

proportionsof

and

nature

their excretion.
For
ample,
exupon
and this in turn
to a loss of sodium

lead

may

is necessary
discussed in previous

the influence

consideration

into

take

to

Moreover, it

"

to

ratio of acid

the

facts that

this the

basic

to

more

or

elements

in

less of the

produced in the body may be neutralized by ammonia

instead of by fixed bases (427 b),and that the kidneyshave the
in some
speciesat least,to separate acids from bases
power,
of acid phosphates,it
to excretion
(427 c),leadingespecially
fall considerably
short
is evident that the data of the table may
tenance
of representingthe actual value of the feed as regardsmainacids

of the ash
435.

The

that
and

balance.

balance.

ash

the value

of conclusions

must
feedingstuffs concerned
as
particularly
regards farm
or

less

from

the

more

or

be

animals.

probable deductions

previousparagraphsregardingthe
and

need

effect of

herbivora

to
a

be

checked

feeding
can

be

stuff

by
or

with

render

composition
less

conclusions

the

of ash in the

ash

are

outlined

experiments. The
on

of the

problematical,

the facts

functions

it evident

of income

balance

Such

from

direct

ration

determined

the

to

as

drawn

outgo of ash elements

more

considerations

These

"

body
actual

maintenance

certaintyonly by

in

means

of

of

and outgo of all the ash elements,

comparisons of the income
contained
in feed,
of the amounts
i.e. by determinations
j
urine
feces and
(metabolism experiments)or by comparative
parative
(comanalyses of carefullyselected test and control animals
slaughtertests).

direct

Data
some

of this sort
tentative

for

conclusions

mature

may

animals
be

drawn

are

from

very

scanty

but

experiments by

MAINTENANCE

Diakow

Cochrane,2each

and

the

345

Diakow's

singlesteer.

periments
ex-

periods on mixed rations containing

hay. In Cochrane's experiments alfalfa hay constituted
and submainsole feed, supermaintenance, maintenance
in

contained
Under

those

urine

showed

which

instance

every

an

in

rather constant

of

getting rid
in

retained

was

of

excess

the

considerable

total ration

of bases.

excess

only the feces but likewise the

bases over
acids,i.e.,the animal

conditions,not

engaged

Accordingly,the

being consumed.

rations

tenance

was

on

MATTER

four

include

much

OF

REQUIREMENTS

"

bases.

excess

body

composition even

The

residue

net

consisted of basic material

in the

case

of Diakow's

of

nearly

animal.

mature

clusion
then,these experimentsconfirm the conthat with rations containing a large proportion of
of
to fear losses either specifically
roughage,there is no reason
fixed bases or in generalof total ash.
Such would almost always
be the case
with the ordinary maintenance
rations of cattle,
sheep and horses. Swine, on the other hand, if maintained
entirelyon grain, might very well receive rations not well

So far

balanced

as

which
that
if it

they

go,

to indicate
Chapter XI (492-496)seem
for maintenance
the ordinarygrain ration,especially
even
consists largelyof maize, should have its ash composition
are

discussed

of in the

stillto be

body without
investigated.In

bases excreted under

experiments, it
excess

of acid

the stock

acid ash

an

the extent

herbivora,and

on

in

effect of

The

corrected.

care

observations

experiments and

regards ash, and

as

to

which

drawing

would

elements

seem

likelythat

even

and
a

ash, has

of

largeamounts

surplus

Cochrane's
considerable

might be neutralized without

drawing

body.

experiments, the minimum

lb. calcium

of

its reserves

on

be taken

aciditycan

the conditions of Diakow's

of fixed bases in the

In Diakow's

such

view of the

rations of

in the mixed

quantitiesof 0.115

and 0.045 lb. phosphorusin the feed per

live weight sufficed to support not inconsiderable

pounds
gains by the
1000

of 0.147 lb. calcium

experiments,a minimum
pounds also resulted in a gain, while 0.039 lb.
per 1000
In Henneberg's
phosphoruswas justsufficientfor maintenance.
3
of cattle,however, disthe maintenance
investigations
upon

body.

Landw.

In Cochrane's

2 Penna.
Inst, of An. Nutr., unpublished results.
Jahrb.,44 (1913),833.
1
Beitrage,etc., Heft, 1 (i860),p. 113.

346

NUTRITION

OF

FARM

ANIMALS

smaller amounts, viz.,

lb.
tinctly
0.090 lb. of calcium and 0.021
x
of phosphorusprovedadequatefor maintenance. Weiske
found
that a mature
of ash ingredients
sheep gainedsmall amounts
of
meadow
on
a ration
hay containing,
pounds live
per 1000
lb.
calcium
and
lbweight,0.179
phosphorus.
0.045
of ash
tenance,
As
deficiencies.
436. Correction
regards mainit seems
clear that the ash requirement
is a qualitative
rather than a quantitative
that
the
it
is
proportions
one; i.e.,
of ash ingredients
far more
than the total amounts
that are
important. If,then,there is reason to fear that the ash supply
in the ration is inadequatefor maintenance,any measures
taken to remedy this must be directed chiefly
toward the correction
of the misproportion
between different ingredients
and
to an increase of their total quantity.
only secondarily
of effecting
One method
such a correction is by the direct
of
mineral
In attempting to correct deficiencies
addition
matter.
in this way, however,the simple
addition of more
ash
"

the ration may

not be
also to take into account
the nature

material

to

effective. It is necessary
of the defects to be made

good. Maize, for example,has alreadybeen instanced (430)

stuff peculiarly
low in ash, the exclusive use of
as
a feeding
lead to a loss of
which, even for maintenance,might readily
ash from the body. Maize is especially
deficient in calcium and
its exclusive

use

would

be liable to

cause

loss of this element.

supply additional calcium,however,by the

addition of such materials as calcium phosphate or sulphate
would not help the situation materially
because the ash would
stillremain acid and thus capableof causinga loss of fixed
bases irrespective
of the additional amount
of calcium present.
The

On

to

attempt

the other hand

by
only

the

the addition of calcium

of

use

chalk
precipitated

suppliesadditional
which

leads to

demonstrated

in the

calcium

numerous

or

wood

bonate,
car-

ashes,not

the acid

dition
con-

element,

has been

well

as

on
growing swine
experiments

referred to in

Chapter XI (496). The correction of the ash

hays causingmalnutrition of the bones,like
instanced by Zuntz
quirements.
(433),presents quite different reThe very injurious
tained
Brandenburg hay, e.g., conthe following
:
percentages of ash ingredients

compositionof
those

of

remedies

but

loss of that

in the form

"

Landw.

Jahrb.,9 (1880),290.

MAINTENANCE

Table

64.

"

OF

REQUIREMENTS

"

Ash

Hay

of

Causing

MATTER

Rickets

Gram
Per

347

Equivalents

Cent
Acid

CaO

Base

MgO

0.693
0.308

K20

1.762

0-3737

Na20

0.082

O.0264

SO3

(Estimated

from

protein)

0.2470

O.1526

2.650

0.0668

P205

0.380

0-0535

CI

0.722

0.2037
0.7997

0.3240

Such
and

would

hay

is

deficient
relatively

presumably

be

improved by

phosphate, while the addition

since
probably be unnecessary,
basic

acid

over

ingredients. In

in calcium

and

in calcium

of

and

phosphorus

the addition

calcium

carbonate

the

hay contains an
addition,however,

phosphorus,it

shows

of calcium

would

excess

to

of

ficiency
its de-

misproportion
of potassium to sodium, which, as already explained,would
tend to increase the excretion of calcium
phosphate unless
sodium salts,particularly
the chlorid,
added.
were
In

the mixed

rations

herbivora,however, direct addition

is seldom likelyto be necessary
of mineral matter
unless roughage
of abnormal
is
the
quality
employed. Usually
surplusof
bases in forage crops will more
than balance
the surplus of
acid elements in the concentrates
salt
used, while the common
will more
than balance any
given as a condiment
ordinarily
of potassium in the rations.
Unusual
probable excess
rations,
such perhaps as very heavy grain rations,those containing an
unusual
proportionof protein, or those made up of unusual
feeds may
form an exception to this general rule and require
In the case
of swine, such a balancing
specialconsideration.
of one
feedingstuff against another as regardsash ingredients
is less practicable,and the securingof a proper balance needs
of the
attention.
determinations
more
Until, however, more
actual ash balance
of different species on
different classes of
rations

are

made

of

(435) it

is

hardly possibleto

state

with

any

348

NUTRITION

ANIMALS

FARM

OF

definiteness what

required in
wide

proportionsof the different ash elements are

maintenance
rations and
the whole
subject offers

field for

investigation.
" 3.

Leaving

Accessory

of account

out

of

the fact that the

and
energy
matter, the proteinsand
source

Substances

proteinsare a minor
consideringonly the requirements for
ash

of the feed

elements

substantiallyfor

required

are

make

losses of the

to
maintenance, i.e.,
good
of the body, especiallyif the ash content
structural
elements
of the body fluids be included
under
this designation. Recent
however, have revealed the presence in the feed
investigations,

of minute

of substances

amounts

different relation

nutrition

to

Vitamins.

substances

the

diet of so-called

seed coats
the

known

nutritional

have

been

that many

sense

on

rice.

by

the excessive

which

was

It has

been
use

the
Substantially

bear

to

may

as

beri

beri.

It has

the

these

been

rice and

shown

"

disease

same

experiments on these
imperfectknowledge of the subjectis

by

the
of

been

use

of

rough

shown
a

animals

on

rice until

that

rational

rice

or

that

disease,resultingfrom a preponderancein
"
rice from
which
the
polished rice,i.e.,
It is a tropicaldisease onlyin
removed.
inhabitants of the tropicssubsist largely
shown
that it can
be produced in Europe
of this grain.

it is to

It has

sory
acces-

of the

cause

(polyneuritis)
may
in
animals, especially fowls,by an exclusive diet

in

quite

called for

be

first called to

through investigationsinto

tropicaldisease
this is

Attention

"

and

appear

substances.

convenience,accessory
437.

which

in

that

shown

of
most

polished
of

our

due.

beri beri may

and especially
dietary,
by

of other

have

man

animals

be induced

be

prevented

the substitution

ments
grains for polished rice. Experithat a subject fed on
polished

the

point of death may be restored to normal

condition in a short time by the administration
of small amounts
of an aqueous
of rice bran, the improvement being so
extract
The
rapid as to appear almost miraculous.
generallyaccepted
of a waterexplanationis that the bran contains a small amount
soluble

nearly at

substance
of the

or

substances

body,

the

lack

tioning
funcfor the normal
necessary
of which
in polishedrice gives

MAINTENANCE

rise

disease.

the

to

notably yeast,

themselves

has

has

who

Funk,1

as

well

to

the

the

the

as

notably by

character

have

to

certain

and

Hart

fats, such

growth.

That

vitamins

of

this

Funk

maintenance

and

of

is

thought

and

scurvy

pellagra,

likewise

are

absence

and

of

Mendel.

permanently
from

the

secured

supporting

fact

the

from

that

norma]

from

rations

on

liver

otherwise

ration

It

with

cod

yolks,

egg

(or substances) differs

apparent

what
some-

associated

are

from

of

be

due

regarding growth,

there

fat, the fat

substance

but

investigation,
It

by Osborne

that

apparently

may

as

observed

incapable

seems

isolated

vitamins.

been

butter

it

line

effects.

Very interesting facts of

"

whose

renders

adequate

in these

have

oil, etc., substances

been

yet

such

demonstrated
as

curative

same

this

substances,

of vitamins.

name

McCollum,

been

in

349

other

poisoning of swine,

substances.

similar

the

not

diseases,

cotton-seed

Growth

seems

general

of diets deficient

use

438.

have

MATTER

of

extracts

prominent

nutritional

other

that

been

them

given

Aqueous

OF

capable of producing

are

substances

The

REQUIREMENTS

"

which

it is absent.
The

interesting results

very

Steenbock

and

balanced
the

to

the

play

when

in

maintenance

discussion
deferred

directed
of

to

their

until

that
Ergeb.

Wis.

McCollum,

rations

properly

made

from

up

these

their

Physiol.,

of farm

nutrition
two

nutritive

functions

Expt.

for

of live

subject

the

all of them

investigations upon
been

in

continued

reproduction, although
for the

have

fed

cows

ordinary criteria but

important part

an

rations

Most

with

by Hart,

of

specific differences
and

obtained

single plants (wheat, oats, maize) suggest that

similar
the
the vitamins
to
or
growth substances

substances

These

Humphrey

according

products

may

three

or

effect

as

seemed

fested
mani-

regards growth
to

accessory

be

quate
fairlyade-

in

to

substances, however,

growth

nutrition

may

is considered.
13

years

weight.

relation

(1913),

Sta., Research

animals.

125.

Bui., No.

17

(191 1).

and

further

therefore

be

CHAPTER

feed

in

of

excess

of

stored

are

and

in the

up

have

which

however,

well

as

of

can

studied

purpose

of the

of the

fattening

feed

the

440.

with

and

first

instance,

maintenance

the

from

Fattening,

part

of the

and

while

feed

the

general

The
nature

it makes

which

upon

for discussion

aspects

it is

requirements
animal.

mature

duction
pro-

in

the

Just

for

maintenance
from

lost

amounts

must

the

support

quantities of
depend, in the
the
body during

the

as

be

fattening

supplied

in

excess

will

process

of

depend

gain made.
obvious
first step in considering the feed
requirements of
fatteninganimal, therefore, is a study of the composition of

primarily
The

to

its

consider

"

quantities which

the

fasting, so

is derived

meat,

in the

its economic

required

upon

of

demands

requirements.

energy

since

maturity.

animals

the

importance,

essential

grades

duction
pro-

Such

animals.

supply

an

of such

production.

meat

Fattening

matter

and

process

which

katabolized

examples

full

is to

be

duction
pro-

formed
trans-

of work.

economic

advantage

supply, leaving

connection

forms

Chapter

present

be

may

of mature

meat

immature

on

better

to

of great

better

feed

it may

familiar

yet reached

the

maintenance,

performance

world's

the

consumes

flesh, fat, milk, etc.,

fattening

growth,

least

at

for

surplus

as

most

the

not

as

practiced largely
be

in the

animal

an

secreted, or

or

and

by
of

share

larger

animals

body

it is true, is not

fattening,
the

products

simplest

is afforded

The

results.

expended

of the

When

"

required simply

material

its energy

One

that
sort

some

into

surplus feed.

of

Disposal

439.

ANIMALS

MATURE

OF

FATTENING

THE

on

the

and

amount

the

of

composition

the increase.

"
441.

i.

Composition

Increase

chapters

have

chiefly
rendered

of

Increase

the

fat.

"

The

it evident
350

Fattening

in

discussions
that

the

chief

in

previous

function

of

weights
to

of the

were

approximately

compute

exclusive

offal parts

the

of the

of the contents

ANIMALS

FARM

OF

NUTRITION

352

recorded, so
weight of

with

the

it is

possible

fat-free

body,

and of the wool.

tract
digestive
periods
75 and 82 day fattening

comparisonsbased on
swine were
reportedby Soxhlet

Similar

that

in 1881

and

the

results

fatteningexperiments (16 to 37 days) on geese by B.

in 1884, the primary
Schulze 2 in 1882 and by Chaniewski3
of animal
fat.
objectof each case being a study of the sources
Friske4 in 1909 and Pfeiffer and Friske5 in 191 1, in a study of the
animals during a fatteningperiodof
gain of proteinby mature
of partialanalyses
about 100 days, likewise reporteda number
of mature
by Henneberg, Kern
sheep similar to those made
of short

and

Wattenberg.

Table

65.

Composition

"

Increase

of

Live

in

Composition
Average
Age
of
Animal

Weight

of

in

Fattening

Energy
Content

Increase

of

Increase,

Water

Ash

Protein

Fat

Calories
Lb.
per

Cattle
Lawes

Gilbert

and

4 years

24.64

1.47

7.69

66.20

3051

20.13

2-34

7-13

70.40

3218

25.80

6.64

67.56

3083

20.30

5-23
1.68

74-47

3344

91.87

4002

Sheep
Lawes

Gilbert

and

years

and

Kern
Henneberg,
Wattenberg

Fat

2 j

Very

fat

Last

stage of fattening

Friske
Pfeiffer and

Friske
.

years

2! years
2 I years

6-45

15-07

4 years

12.03

32 years

64.33

72.90

3531

7.11

28.56

1415

6.44

71.50

3247

24-45

1401

53-59

2485

58.39
61.46

2602

Swine
Lawes

and

Gilbert (Average)
......

Soxhlet

"

Swine

No.

Swine

No.

2
.

16^
i6|

mos.
mos.

22.00

0.06

58.96

3-i7

13.42

35-99

3.62

6.80

37.06

1. 21

3-34

24.15

i-37

3.02

Geese

Schulze

mos.

Chaniewski

Centbl. Agr. Chem., 10 (1881),674.

2Landw.

Jahrb.,11 (1882), 57.

6

Ztschr.

Landw.

Ibid., 74 (1911), 409.

2726

Biol.,20 (1884),179.
Vers. Stat.,71 (1909),441.

THE

The

they

FATTENING

results

OF

MATURE

ANIMALS

353

of these

comparative slaughter tests, so far as

composition of the increase,are summarized
65, which includes also the computed energy content

relate to the

in Table

of the increase.
443.

Respiration experiments.
animals

mature
tests
as

from

appears

the summary

far the most

Respirationexperiments on
the results of slaughter
proteinto fat in the increase,

fully confirmed

'regardsthe proportionof

as

By

have

"

of Table

66.

extensive

respiration
experimentsare those made
x
Station
Experiment
by G. Kiihn and by Kellner
on
mature
fatteningcattle. Of the 60 reported experiments in
which
there was
a
gain of both protein and fat,only 3 show less
than 70 per cent of fat in the total organic matter
gained and only
mainder
Rejecting these 6 and grouping the re3, a percentage above
95.
the
of
fat
according to
givesthe results shown
percentage
at

the Moeckern

in the first five lines of the table.

these

To

earlier experimentsby

Henneberg, Fleischer
and by Meissl
The
swine.
on
gains of ash
in these experiments.
determined

and

Table

66.

Proportions

"

Protein

of

and

Fat

and

added

are

in

Range
of
Percentage
Fat
in
of
Organic
Matter
Gained

Kellner

Experiments on

"

cattle

the results of

Miiller
of water

Fattening

sheep

on

not

were

Increase

Average
Composition
Organic
of
Matter
Gain
of
Total
Protein

Fat

Group

70-74.99

26.25

73-75

Group
Group

II

75-79-99

23-30

III

17.17

Group

IV

80-84.99
85-89.99

76.70
82.83
87-45

Group

90-94.99

Henneberg, Fleischer and

on

"

Lorenz

"

91.94

periments
Ex-

sheep
and

Meissl, Strohmer

Miiller

12-55
8.06

4.26

9r-74

90.25

ments
Experi-

swine

on

Animal

No.

9-75

Animal

No.

Animal

No.

10.67
16.39

89-33
83.61

Animal

No.

15.16

84.84

1
2

Landw.

Vers. Stat.,44 (1894), 370;

53 (1900), 1.
Jahresber, Agr. Chem., 16-17 (1876),II, 145.
2

Ztschr. Biol.,22 (1886), 63.

the

Both

slaughter tests
animal
is, as its
is

adipose

ANIMALS

and

respiration experiments
that

demonstrate

which

FARM

OF

NUTRITION

354

the

the

comparative

fatteningof

mature

implies,largely a production of fat,

and
internal
depositedchieflyin the sub-cutaneous
name

tissue and

to

limited

also in the

extent

muscles.

comparative slaughtertests show a large storage of

but the organic matter
water
chiefly
gained in every case was
the average
of all the foregoingexperiments by both
fat.
On
stored up in the
methods, the composition of the organicmatter
follows :
as
body of the fatteninganimal was
few

of the

"

Table

67.

Average

"

Composition

of

Organic

Gained

Matter

in

Fattening

Minimum

64-57%
4-26%

Fat
.

Protein

444.

however,
chieflyof
small

This

that while
fat there

actual

which

the

It is

of mineral

matter.

gain of proteinin
from

appears

metabolism

over

"

amounts

The

foregoing data also show,

the gain of dry matter
in fatteningconsists
is also a gain of more
less proteinand of
or
The

gain of protein.

The

Table

not

was

includes both

68, which

experiments,most
composition of the

cases

some

of which
increase

are

has

inconsiderable.

slaughtertests

identical with
been

those

and
from

calculated.

of these

experiments
considerable
the feed contained
a
surplus of protein
that necessary
for maintenance.
Such a surplusof protein,
probably safe

shown

to

that

assume

in most

(406),has a tendency to produce

limited storage of protein,which probably consists
a somewhat
in an increase of the contents
of the cells or of the protein
held in solution in the body fluids rather than in an increase of the,
structural elements
of the body.
The observed gain,therefore,
as

may
or

was

in

represent in part
in the soluble
small

Chapter

actual

an

is

horn, epidermis, etc.,

increase in the cell protoplasm

tively
body, while in addition,a relaaccounted
for by the growth of hoof,

protein of

amount

IX

of

the

the

cattle

and

swine.

Moreover,

FATTENING

THE

OF

ANIMALS

MATURE

355

accomplished largelyby an increase

of existingcells there appears
to be also an
in the fat-content
of cells in the adipose tissue,and the
increase in the number
be assumed
to require a supply of protein.
latter process may
The
adipose
protein contained in one pound of subcutaneous
be
tissue of average
equivalent to the
composition would
storage of about 0.045 lb. of protein. Obviously,however, the
count
growth of epidermal and adipose tissue can but partiallyacof these instances
for the observed gain of proteinin many
and apparentlya distinct increase of the nitrogenous tissue in
be admitted, averaging, in these experiments,
fattening must
about
pounds live weight or about
0.2
pound per day and 1000
5.5 per cent of the total increase in live weight.
while the

of fat is

laying on

Table

68.

Gain

"

of

Protein

Animals

Mature

by

AverCharacter
Experiment

age

of

Live
Weight

Cattle
Kiihn

and

Metabolism

Kellner

Sheep

and

Weiske

Pfeiffer and

and

Kern

and

Kgs.

Grams

667

82.0

Per
Live

1000

Wt.

O.123

Mul-

ler

Henneberg,
berg
Henneberg

Per
Head

of

Fleischer

Henneberg,

Daily
Gain
Protein

Metabolism

34-2

8.50

O.248

Metabolism

54-8

9.24

O.169

36.45

9.346

Watten-

Pfeiffer

Slaughter

48.3

Metabolism

43-5

Metabolism

Kalb

38.5

Friske

Metabolism

Slaughter

35-2

Pfeiffer and

Metabolism

36.7

Slaughter

36.7

Friske

35-2

Swine

Soxhlet

Slaughter

"7-5
70.0

Meissl

Metabolism

104.0

125.0
{ 140.0

Geese

Schulze

Slaughter
1

The

nitrogen of the wool

The

same

animals

were

is not
used

included

3-9

in the gain.

also in the slaughter tests.

356

NUTRITION

Influence

445.
meat.

ANIMALS

composition of the lean

the

fattening on

pose
fatteningconsists largelyin an increase of adiin the ordinary sense, it has an
important effect
the composition of lean meat
in the commercial
of the muscle
tissue proper
that
(fat-free
upon

tissue
upon
and

sense

FARM

While

"

both

of

OF

meat).
Percentage of fat.
is by no means
meat
fibers
only muscular
lean

What

"

Fig.

which

37.

The

"

themselves

Porterhouse
steak from
marbling of meat.
(Illinois
Experiment Station.)

degrees of magnitude
the

the

found

of intramuscular
in the so-called

cause

between

prime

the

steer.

tissue of all
bundles

muscle

and

tensive
(86). Fattening, especiallyinmarked

fat in the lean

"

of connective

masses

separate muscles

fattening,
may

Such

commonly spoken of as lean

fat,since the term includes not
with
the relatively
little fat

free from

they contain,but

between

is

meat,

increase
as

in the

is evident

to

storage
the

eye

marbling."

analyses of lean

meat

as

are

recorded

confirm

the

dence
evi-

of the eye in this respect. A summary

of the results on
this point has been
following
The
given by the writer * elsewhere.
from
that
example taken
illustrate the point in question.
1

U.

S.

Dept. Agr., Bur. Anim.

publication may

Indus., Bui. 108 (1908),p.

serve

33.

to

THE

Table

FATTENING

69.

Fat

"

OF

Fresh

in

MATURE

Lean

ANIMALS

Meat

Leyder

"

357

and

Pyro

Very
Fat
Cow

Neck

Leg

2.8

5-8

Flank

8.8

Tenderloin

12.9

Similarly,Braman
in lean

from

meat

found

medium

the
fat

following percentages

(common)

and

of fat

well-fattened

(prime)steer.
Table

70.

Fat

"

in

Fresh

Lean

Meat

Braman

"

Porterhouse

12.71
6.66

Round
.

in making such
practicaldifficulty
comparisons arises in the
of
the
fat sursample. Obviously, the subcutaneous
preparation
rounding
A

the

should

meat

of fat found

be

discarded

between

the

and

the

same

is true

of the

largemasses
just what part of
of a fat animal
the adipose tissue scattered through the meat
should
be regarded as mechanicallyseparableand what part should be regarded
is
difficult
ferences
Difto
as
decide.
belonging to the meat
proper
in the trimming of the piecesmay
for some
of the
account
irregularresults found by recent experimenters.
Extractives.
increases

"

It

the

be

to

appears

muscles,but

extractives

established
of

that

fattening

well as
as
nitrogenous
For example,
causes
a
deposition of fat in and about them.
and Wattenberg's experiments on
in Henneberg, Kern
sheep,
included in Table 65, the composition of the meat
from
the lean
and

from

the very

tissue)computed

fat animal

the muscles

freed
(partially

to the fat-free

state, was

lIbid., Bui. 128 (1908),p. 86.

"

from

connective

358

OF

NUTRITION

Table

71.

Composition

"

ANIMALS

FARM

Meat

Fat-free

of

Thin

of

Sheep

Sheep

Very

Fat

Sheep

Water

79.41

79.02

Insoluble

15.85

15-73

protein

Extractives
Soluble

protein
Non-protein
.

I.29

i-93

2.18

2.17

1.27

1.

Ash

15

extractives

Total

5-25
100.00

It is

computed

the fat animal

Insoluble

that

were

as

the actual

follows

gainsduring the fatteningof

"

38.7 Grms.

protein

Extractives

Protein

Non-protein
Ash

82.0

Grms.

4.2

Grms.

"9.2

Grms.
77.0

Total

Somewhat

similar

results

obtained

were

later

by

Grms.

15.7 Grms.

the

same

Evidently this
experiments on fattening lambs.
increase in the soluble nitrogenous compounds of the muscles
of the factors going to make
is one
gain of
up the observed
proteinby fatteninganimals.
authors

in

fatteningof animals as a
commercial
is a practice based
on
experience,which
process
has shown
that the tenderness and palatability
of the meat
are
is willing
to
materiallyincreased thereby,so that the consumer
pay a higher price for it. It is to this improvement in quality
in the first instance,and only secondarily
to the gain in weight,
446.

that

Object of fattening.
"

the feeder looks

for his

The

profit.
as
composition of the increase in fattening
what
recorded in the foregoingparagraphs serve
to show
are
the principalfactors in this improvement in the qualityof the
meat.
They are, first,the deposition of the intermuscular
The

facts

to the

36"

NUTRITION

Net

storage of energy

in

the feed

that

energy

in

Chapter VIII.
fatteningrations.

The

same

Just

as

is measured

maintenance

prevents,

its net

so

storage of

body

energy

brought

by

be illustrated
may
Kellner upon
a mature

to

of

mixed

basal

second

column

determining

ration

the

fourth

Determination

"

the

This

the net

well

value

for fattening,

followingrespirationexperiment

already

the

to

conception, as

energy

ox, in which

third

extended

meadow

sufficient to

column
energy

hay

computed

contained

gain.

some

cause

the metabolizable
the

added

was

in the

energy
heat production,

observed

fat.

gain of proteinand
72.

by

be

considered

about.

"

of the table shows

rations,the

and

Table

the

by

been

of a feed for
value
energy
which
loss of body energy
it
for fattening
is measured
by

value

the method

two

the net

of determination.

Method

of the

deducted, has
conception may

energy

as

The

been

losses have

various

449.

is not

reason
or
escapes utilization for one
The
of
the
value
net
as
conception
energy
ing
expressavailable after
part of the feed energy which remains

another.

the

"

loss.
accomplished without some
feeding,so in fattening,a considerable

maintenance

portion of

these

for fattening

needs.

its immediate

As

ANIMALS

values

energy

conception.

fatteningmay be
a
possiblefuture
This

FARM

the process
of
Physiologically,
regarded as a storingup by the animal,against
of
scarcity,of feed energy supplied in excess

General

448.

OF

of

Net

Energy

Value

for

Fattening

Energy
Fat
and
Protein
Gained
Body

of

by

Therms

Basal

ration +

Basal

ration

hay

4.24
2.02

Difference
Difference

Each

gain

of

2.22

per lb. of

0.288

hay

pound of hay added to the basal

protein and fat containing 0.288

ration
Therm

resulted

in

of energy.

FATTENING

THE

This

its net

was

Table

with

37

in

MATURE

OF

for

fattening.A comparison
(364),showing the results of a

value

energy

Chapter VIII

of the

361

ANIMALS

value for

maintenance, renders
evident the identityof the method
employed in the two cases,
the only difference being that in one
the comparison between
case

determination

the two

energy

rations is made

in the other

and

net

above

case

in maintenance

the

below

it.

point of

It is evident

feeding,there

is

maintenance

that in

fattening,

considerable

expenditure
the consumption of feed, so that
of energy
consequent upon
is actually stored up in
only part of the metabolizable
energy
the gain by the body.
In the experiment given as an
tration,
illusTherm
of
one
pound of the hay contained
0.714
metabolizable
of which only 0.288 Therm
or
40.7 per
energy,
in
the
recovered
cent
was
gain.
as

450.

The

Relative

values

causes

which

same

of

are

course

of energy

for

result

and

considered

operative to bring about

on

the

heavier

in the formation

would

maintenance

were

addition, it would

In

in

appear
less

fattening.
"

Chapter VIII (367)

the increased expenditure
in

rations of the
that

of fat from
or

more

for

the

fatteninganimal.
chemical
volved
changes in-

proteinsand carbohydrates

evolution

of

heat.

Whatever

expenditure of energy may be thus caused is additional to that

caused
ditions
condirectlyby feed consumption under maintenance
and must
evidentlytend to reduce the net energy value
of the feed by a correspondingamount
; in other words, the net
values of feeding stuffs for fatteningwould
tend to be
energy
lower than those for maintenance.
Such data as are available,
that this difference is a
to indicate
however, do not appear
considerable one in the case of farm animals,and it would appear
that,in the case of cattle at least and presumably in that of other
species,the net energy values of feedingstuffs may be regarded
the same
for fattening
as beingsubstantially
as for maintenance.1
Energy requirements
for fattening
451.

Energy

of

content

gain.

"

Since

the

net

energy

value

as
feeding stuff or ration for fattening,
explained in the
foregoingparagraphs, is that part of its total energy which can
in the increase,it follows that the
be stored up by the animal

of

Compare Armsby

and

Fries,Jour. Agri. Research,

(1915), 435.

362

NUTRITION

of such

ration

OF

FARM

ANIMALS

of net energy
supply an amount
equal to the maintenance
requirement plus the quantity of
contained
in the gain made.
The
latter quantity,however,
energy
be computed approximately from
the data given in
may
" 1 regardingthe chemical composition of the increase in live
of
weight during fattening.Estimating the energy content
protein at 2586 Cals. per pound (5.7Cals. per gram) and that
of fat at 4309 Cals. per pound (9.5Cals. per gram), the energy
of one
in the last column
shown
content
as
pound of increase was
animal

an

must

65 (442).

of Table

the range
apparently questionable results,1
and average
of the remainder
follows.
are
as
what
Although somevariable
indicate
that
the
of
entire
an
on
they
average
fattening period a pound of increase in live weight in cattle,
sheep and swine is equivalent to about 3.25 Therms.

Excluding

Table

"

Energy

per

Pound

Increase

in

Live

Weight
Therms

Maximum

4.002

2.485 Therms

Average

3-245

in
be

can

73.

Minimum

452.
are

two

Influence

The

Therms

just cited
those of an
entire fatteningperiod. There
most
cases
little doubt, however, that the composition of the increase
and its energy
content
vary materiallyas the fattening
of

stage of fattening.
"

results

advances.
This appears
upon

clearlyfrom Henneberg, Kern

sheep. The "fat" animal had been

regarded

was

as

animal

had

weight

occurred.

fat"

animal

and

Wattenberg's

fed for

10

weeks

sults
re-

and

The
according to local standards.
"very fat"
fed for 29 weeks, or until no
further gain in live
As the table shows, the total gain by the "very

fat

been

materiallylower percentages of water, ash

and protein and a higher percentage
of fat, and had
a
10
per cent
parison
higherenergy content than the gain by the "fat
animal,while a combetween
the
the "fat"
the "very fat" animals
shows
and
gain made by the latter during the last 19 weeks of fattening to have
contained

"

Pfeiffer and

Friske's results appear

sisted
exceptional, since the gain apparently conabnormally large extent of water, while the authors themselves
point out
that the gains of dry matter
were
notably less than should have been produced from
the feed consumed.
It would
therefore,that their omission is justified. Soxhseem,
to

an

let's result upon

swine

No.

has

also been

omitted

for

similar

reason.

FATTENING

THE

OF

MATURE

363

ANIMALS

nearly92 per cent of fat and to have had an energy content

also obtained
of 4002
Cals. per pound. The same
tirely
eninvestigators
similar results in the fatteningexperiments on lambs cited in
Chapter XI (458) although naturallythe proportionsof wa,ter and
of the mature
proteinin the increase were
greater thap in the case
sheep.
contained

with thin
earlier stages of fattening,especially
animals, the storage of fat is accompanied by a considerable

During

the

less increase in body protein. As

or
gain of water and by more
to consist
the fattening progresses,
however, the gain comes
to an
increasingextent of fat accompanied by very littleprotein
The
relativelysmall percentage of water.
energy
of a unit of gain in live weight,therefore,
in the later
content
stages of fatteningis materiallygreater than in the earlier
and

will
net energy
Evidently, then, more
be requiredin a fatteningration to produce a pound of increase
in live weight toward the close of the fatteningprocess than at
is entirely
in harmony with the exits beginning,a fact which
perience
of feeders that gains become
increasinglyexpensiveas

stages of the process.

the animals
So far

as

become

fatter.

definite conclusions

scanty data available,it would

of

are

warranted
that

seem

from

the rather

in the earlier stages

fattening a ration supplying (in addition

to

maintenance)

of net energy would be sufficient to support

2.5 Therms
gain of a pound of live weight, while in the later stages the

about
a

requirementmay

rise to 4.0 Therms

Protein
453.

Protein

or

perhaps even

more.

requirementsfor fattening

unnecessary

for

fat

production.

"

It

was

in the
Chapter V (247-249) that body fat,especially
of farm animals,.is derived
case
chieflyfrom the non-nitrogenous
nutrients of the feed,proteinplaying but a subordinate
role in its production, and Kellner
has shown
(769) that the
proportion of the energy of protein which can be stored up by
less than the correspondmature
ing
fatteninganimals is distinctly
So far as
the
nutrients.
for
non-nitrogenous
percentage
simple fat productionis concerned, therefore,it would appear
that a surplus of protein over
that required for maintenance
shown

in

would

be unnecessary

and

possiblydisadvantageouson

account

364

NUTRITION

of its

tendency
(365).
the actual

found

to

of

consumed

in mature

in the

increase

storage of

with, of
material,so
is aware,

proteinas

it has

ever,
(444),how-

in most

variable

been

tion
propor-

epidermaltissue,

or

organs?).

all instances
was

abundant

shown

for maintenance,

So far

that

enous
non-nitrog-

favorable

were

It is to

the rations

necessary
supply of

justmentioned.

for such
the writer

as

the mature

surplusof

animal

has

cells,and in part
nitrogenous extractives

in the internal

yet been

not

of

body

of fat

the conditions

that

rather

growth

number

an

course,

that

and

proteinthan

more

"

of the

fatteninganimals

proteinand

(and

tissue

in

appeared

in part to the

contained

storage of

As

"

remarked, however, that

be

generalmetabolism

small
relatively

protein,due

in the muscular

the

increase.

increase

contain

in part to an
actual
to an

FARM

to stimulate

in

Protein

454.

ANIMALS

OF

fattening
the amount

actuallyrequires
proteinover
any
it
for maintenance, although
can
apparently utilize
necessary
at least to

excess,

in its

some

extent,

to increase

the stock

of

an

protein

body.
requirementof

the

ured
fatteninganimal as measby the observed storage of proteinis relativelysmall,one
pound of increase in live weight containing in round numbers
from 0.02
lb. to 0.08 lb. of protein.

most, the

At

455.

gain

of

Utilization

of feed

proteinby

the

protein.
"

Assuming

fatteninganimal

to

observed

the

represent

real

quirement,
re-

fatteningration must
supply, in addition to the proteinnecessary for maintenance,
after undergoing the various
additional
amount
an
sufficient,
of
of digestionand metabolism, to yield the amount
processes
in the increase of body weight. As
body protein contained
will appear
more
particularlyin considering the subject of
of feed
growth (470,471), little is known
regarding the amount
protein required to yield a unit of body protein. Doubtless
this will differ as between
different individual proteins,depending,
for one
thing,upon the proportionsof the different amino
data are
acids which
they contain,but adequate quantitative
it is evident

that

sufficient

yet unavailable.

as

456.

Protein

in

fatteningrations.

"

In the absence

of definite

of the proteinof
knowledge regarding the availability
the

question of the

amount

of this nutrient

which

the

feed,

should

be

FATTENING

THE

365

ANIMALS

MATURE

OF

suppliedto fatteninganimals may be approached much as was

in Chapfor maintenance
the questionof the amount
ter
necessary
by inquiringwhat
IX, i.e.,

is the least amount

of

digestible
proteinwhich, along with sufficient non-nitrogenousnutrients,
of fattening. If it
rate
has sufficed to support a satisfactory
that of two similar animals or lots of animals receiving
appears
of feed, the one
consuming the smaller amount
equal amounts
of proteingave equallysatisfactory
gains,both as judged by
be concluded
that
the live weight and by the block test,it may
of protein was
at least sufficient,
the smaller amount
although
whether

it cannot

be determined

than

actuallynecessary.

was

it may

not

have

Unquestionably,the proteinrequirements of

been

mature

greater
ing
fatten-

greatlyoverestimated in the past. Wolff's

feedingstandards (791),publishedin 1864, recommended
original
animals

for

been

have

pounds live weight the

digestible
protein:
per thousand

rations
fattening

of

amounts

ing
follow-

"

Cattle

2.5-3.0

lb.

Sheep

3-Q-3-5

lb.

2.7-5.0

lb.

Swine
.

Substantiallythese same
figureshave been repeated more
with a few
from publication
to publication,
less uncritically
or
It is clear,however,
even
exceptions,
up to the present time.
the
based upon
from Wolff's writingsthat his standards were
then prevailing
views of Voit and Pettenkofer
(248)regarding
of animal fat rather than
the importance of protein as a source
actual experimentalresults.
Subsequent investigations,
upon
cattle
the
notably
respirationexperiments of Kellner upon
of
that such large amounts
(p.367),have fullydemonstrated
advantageous for
proteinare neither necessary nor especially

fattening.
that his protein
Indeed, Wolff himself has- demonstrated
l
standard
for sheep was
unnecessarilyhigh. In 1890, he published
in 188 5- 1886 of maize and beans as
the results of a comparison made
feed for fatteningsheep, using two lots of two mature
sheep each.
After a preliminaryfeeding,the followingresults were
obtained in
107 days'feeding:

Sheep.

"

"

Landw.

Jahrb.,19 (1890),823.

366

NUTRITION

Table

74.

Influence

"

OF

of

FARM

ANIMALS

Supply

Protein

Gain

on

by

Mature

i, Fed

Lot

tening
Fat-

Sheep

Lot
on
and

Hay
Beans

and

Kgs.

Weight
Weight

at

beginning

ii8-75

98.61
118.56

18.82

19-95

Gain

Digestible matter

eaten

per

kilograms

1000

live weight
Protein
Total

Lot

Kgs.

99-93

at close

digestible(fat X 2.4)

Fed
Hay
Maize

2,

on

3.26
18.19

s
.

1.

Si

19.20

receivingmaize, produced about the same

gain relatively
the digestible
consumed
Lot
to
matter
as
1, notwithstanding the
of protein supplied. A block test tended
smaller amount
to show
a
the
of
Lot
2.
Subsequent experiments *
slightsuperiorityon
part
on
one
gave confirmatoryresults,barley being compared with beans
The
shows
the
animal each.
table
actual
trients,
following
digestiblenuand
the
total
live
computed per 1000
kilograms
weight,
gain
for each period:
2,

"

Table

75.

"

Influence

of

Protein

Supply

on

Gain

Sheep

Landw.

Jahrb., 25 (1896),175.

by

Mature

tening
Fat-

368

FARM

OF

NUTRITION

ANIMALS

periods having been short in these experiments, the gain in

be satisfactorily
live weight cannot
determined, but on the basis of
of the compositionof the increase
and Gilbert's determinations
Lawes
(442) it may be estimated to have been approximately one pound per
The

day.

Loges
to

reports the results of experiments undertaken

test Kellner's

conclusions,in

which

great gains in weight with mature

of proteinconsumed
absolute amounts
Apparently from 0.75 to 1 lb.
pounds live weight is sufficient to

cattle

as

mature

on

Such

"

record
but

most

are

of

one

as

not

ratio of

stated

10.3 gave

5.7, but the

in the abstract.
1 :

of

digestibleprotein per 1000

the requirements of fully
meet

fattening cattle.

Swine.
are

nutritive

Pomritz

at

show

experiments on
that these

the

fatteningof

swine

as

sheep, need

at

mature

animals, like cattle and

comparatively small surplusof proteinover

for maintenance.
necessary
The
experiments
respiration

by Meissl,Strohmer

and

the

amount

Lorenz

upon

(443) afford a generalillustration of this.

The
followingtable shows the digestible
protein and the metabolizof the feed and the gain of energy
No
able energy
by the animal.
of the high proteinration of Experiment IV over
distinct superiority
the low protein rations of Experiments I and III appears, while the
realized in Experiment II with a moderate
protein
greatest gain was
but
content.
relatively
supply
high energy
the

of fat

sources

Table

77.

"

by

Influence

swine

of

Supply

Protein

on

Gain

by

tening
Fat-

Mature

Swine

Rations
per
100
Live
Weight
Pounds

Digestible
Protein

Metabolizable

Energy

Lb.

I
.

II

IV

16

of

Live
Weight

Animal

III

Gain

140

0.074

70

0.161

Therms

5-n
IO.02

125

0.098

4.10

104

0.410

3-94

Per
Live

100

Lb.

Weight

Energy

Per Therm
Metabolizable

Energy

Therms

Therms

2-54

O.50

5-96
1.48
2.56

O.58
O.36
0.43

Soxhlet,inhis experimentson the same

subject (442),fed two swine
months
old and weighing about 200 pounds each at the beginning
1

Centbl. Agr. Chem., 31

(1902),646.

FATTENING

THE

OF

MATURE

369

ANIMALS

of rice
experiment,a low proteinration consisting
exclusively
for 75 and 82 days, respectively.The protein content
of the ration
and the average
lows
folas
daily gain in live weight per head were
of the

"

Table

78.

Influence

"

Supply

Protein

of

Gain

on

Mature

by

tening
Fat-

Swine

(N X 6.25)

Protein

Digestible

Daily
Live
Initial
Weight

in

Per

Day

Per

and

100

Head

Lb.

II

These

few

results upon
of
possibility considerable
the results

are

Lb.

O.265
O.269

213

mature

swine

Weight
Head

and
Live

per

Weight

Lb.

220

III

Day
Lb.

Gain
Live

are

Lb.

O.I2I

I-

O.I26

1.04

of interest

as

15

showing the

fatteningon low proteinrations. In practice

little
the
since
mercial
comcomparatively
significance
swine is usuallycarried out upon
the immature

of

fatteningof
animal.

The

recorded

experiments show that in the fatteningof

animals
results have been obtained with
mature
as
satisfactory
rations containing0.75 to 1.5 pounds of digestible
protein per
1000
pounds live weight as with those containinga much more
abundant
50 to 100
but with
in

of

supply. Even

these

however, are from

for maintenance,
per cent higherthan is necessary
the exception of a small group
of Kellner's experiments
which
approximatelythe maintenance
requirement

proteinwas

consumed

amounts,

the results fail to show

whether

it is

practicableor advisable to reduce stillfurther the protein content

of fatteningrations. As regards the simple question of
that an amount
of this nutrient
proteinsupply,it appears likely
but littlesuperiorto the maintenance
requirement is all that is
In practice,
absolutelynecessary.
however, the inferior digestibility
of low-proteinrations (723,724) as well as the fact that
such rations are likely
to be less palatablethan those furnishing
liberal supply have to be considered.
The
dition
a more
simple adof

non-nitrogenousnutrients
2

to

maintenance

ration

furnish

might
and

yet

not

economic
in

XII),

of

aspects

connection
the

physiological

it

convert

with

present
aspects

for

material

ample

into

the
the

of

practicable

subject

the

dealing
fattening

meat

more

process.

will

body

ration.

fattening

however,
of

of

production

the

question,

chapter

ANIMALS

FARM

OF

NUTRITION

370

be

production
especially

fat
The

considered

(Chapter
with

the

CHAPTER

XI

GROWTH

"
Cell

457.

microscopic

General

i.

Nature
The

multiplication.
"

cell.

germ

animal

advance

Its

Growth

of

in a single
originates
from
this insignificant

beginning to the size and complexity of maturity is effected

of cells,together with a
by a multiplicationof the number
progressivedifferentiation of function,the whole constituting
of growth. Growth, then, may
the process
be characterized
in an increase of the structural elements
of
brieflyas consisting
in a gain in size
the body, chiefly
by cell multiplication,
resulting
and weight.
The
458.

in

so

increase

Composition of

study of

the

increase.

feed

As

"

with

requirements

factor to be considered

prime

during growth

of

growing animals, a

is the amount

and

made

The
nature
at different ages.
either
be investigated
growth may

of the

gain
made
during
comparative slaughtertests
Of the former

there

by

or
are

means

on

record

growth of pigs for the first 16 days

by Tschirwinsky2 on pigsbetween
months, one by Kern and Wattenberg 3 on
the

on

fatteninganimals,

between

of
a

composition
of the

gain

of
by means
ments.
experirespiration
l
Wilson
study by

after

tigation
inves-

birth,an

the ages of

and

growth of lambs
by Jordan 4 upon the
the

the ages of 6 and 28 months, one

growth of cattle between the ages of 23 and 33 months and one
5
on
by Wellmann
pigs. Data regarding dogs and cats
young
also

are

on

record in

by
investigations

Amer.

Thomas

2 Landw.
Jour. Physiol.,8 (igo3), 197.
Jour. Landw., 28 (1880), 289.
4 Maine
Expt. Sta.,Rpt. 1895, Vol. 2, pp. 36-77.
5 Landw.
Jahrb., 46 (1914), 499.
6 Arch.
(Anat. u.) Physiol., 191 1, p. 9.
7 Arch.
Physiol. (Pfluger),
135 (1910),163.

371

and

Vers.

by Gerhartz.7

Stat.,29 (1883),317-

NUTRITION

372

OF

FARM

ANIMALS

calves
Respiration experiments by Soxhlet1 on three young
of the gain or loss of ash, while
the
included determinations
The
feed being
live weights of the animals
also recorded.
are
of the digestive
exclusivelymilk, the variations in the contents
tract were
probably slightand a computation of the composition
the live weights seems
of the increase based
justified.
upon
The results of both the slaughterand respiration
experiments
contained
in the followingtable,the energy
content
are
being
from
the
fat
and
computed
protein.
Table

1
3
5
6

79.

ier Ber.
Fat- and

"

Composition

Versuchs-Station

of

Wien,

Increase

pp.

or

Live

Weight

101-155.

ash-free dry matter.

from " fat-free body."
Computed
The figuresdiffer slightlyfrom those reported by the author.

in

Growth

By difference.

Two

periods.

GROWTH

and
spiteof irregularities

In

373

gaps

in the

table two

general

in
that the percentage of water
first,
increases with advancing
the gain decreases and that of dry matter
age of the animal, and second, that of the dry matter
gained, an increasingproportionis fat as the animal matures.
clear if the composition of
The latter fact becomes
especially
of the ash-free gain be computed.
the dry matter
and Trowbridge
The result of investigations
by Waters, Mumford
x
are
as
quite in accord
reportedby Henry and Morrison
with the teaching of Table
79, the percentage composition of
the first and the second 500 pounds gainedby young
fattening
steers being as follows :

facts

clearlyshown

are

"

Fat

%
First

lb.

500

Second

lb.

500

459.

48.6
75-

Energy

stored in

of

content

gain.

"

The

of

amount

energy

unit of increase in live

weight shows a fairlyregular

and notable increase as the animal
older, due to the
grows
smaller
and the higher percentage of fat
percentage of water
which

it contains.

per unit in those

animal

of increase in the energy

content
in which no considerable fatteningof the

cases

rate

attempted seems

was

Therms

The

per

results upon

to be

pound and the same

fattening animals

fairlyregularup to about 3.0

thing is also true of most of the

up

to

about

3.5 Therms

per

pound, although the actual energy content per unit,atthe same

animal and the limit is
age is naturallygreater in the fattening
reached

therefore

earlier in life. In both

cases

the limit

seems

to

correspond in a general way with the average energy content of

the gain made
by mature
fatteninganimals as estimated in
Chapter X (451),viz.,about 3.25 Therms per pound. The data,
is much
however, are few and further investigation
to be desired.
Relation
460.

The

dimensions

of

rate

of

of growth to

growth.

"

growing animal
1

Feeds

and

age

If

the

be

platted,there

successive

Feeding, 15th Ed., p.84.

weights
are

or

obtained

NUTRITION

374
what

might

OF

be called the

FARM

of

curves

ANIMALS

of stature.

weight or

These

rise

more
rapidly at first and afterwards
slowly as the animal
the increments
of
approaches maturity. Or in like manner
weight or size observed in successive equal periods (day,week,
be platted,showing at what
month
or
year) may
periodsthe
absolute growth is most
rapid.
It is evident,however, that an increase of a pound in weight
by an animal weighing 500 pounds is relativelymuch less than
increase in a 100-pound animal.
the same
For many
purposes,
of growth to age is afforded
a better expressionof the relation
the
by a computation of the rate of growth, by which is meant
in a given unit of time expressed as a fraction of the
increment

amount

present

instance

the

at

the

just supposed

be in the first case

would

one-hundredth.

one

beginning of
of

rate

time.

growth

in

the second

Thus

in the second

the small

case

in the

weight per day

and

five-hundredth

one

In

that

animal,

in

proportion to its weight, is growing five times as fast as the

larger and may be regarded as showing five times the energy
of growth. An evident advantage of this manner
of expression
is that it permits of a comparison between
animals of very different
weights,as, for example, of sheep with cattle.
461.

Rate

of

growth

different

at

ages.

"

Somewhat

tensive
ex-

and the lower animals, show

observations,both on man
that the rate of growth as just defined diminishes
from
birth
onward, the diminution being more
rapid at first and slower
as
maturity is approached. This subjecthas been discussed in
a

manner
illuminating

most

and
other

others' observations
animals

growth

is

as

well

same

curve

into intrauterine

on
as

expressed by

graduallybecoming
the

by

extends
life.

on

the basis of his

guinea pigs,rabbits,chicks

on

Graphicallythe

man.

descending
and

more

Minot

more

backward

Foster

says:

rate

own

and

of

but
first,
nearly horizontal,while

curve,

steep

without
"It

seems

at

material
as

break

if the

petus
im-

In
growth given at impregnationgraduallydies out."
the early stages of growth, therefore,the anabolic
processes,
which
tend to build up tissue,
while
time goes
as
predominate,
the
the katabolic processes
and
on
over
more
gain more
anabolic until at maturity the two tend to become
substantially
to

balanced.
1

C. S. Minot

Age, Growth

and

Death, Chapter III.

376
Just 1
follpws.
and

OF

NUTRITION

of

lambs

on

have

FARM

been

ANIMALS

included

in the table which

the

in which

made
experiments were
by the method
of comparative slaughtertests, the compositionof the control animals
of the initial protein content
of the
gives an approximate measure
control animal
no
was
tent
body. When
analyzed the initial protein conestimated
from
has been
the live weight.
well as possible
as
the case
in the majority of the experiments it seems
Since this was
the gain of protein per iooo
live weight.
desirable also to compute
Except in the case of very young or very fat animals,the results are
with those computed in the other
likelyto correspond substantially
way, while they have the advantage of being expressed in the manner
usually.adoptedfor formulatingfeedingrequirements.
In those

cases

Table

80.

"

Rate

Gain

of

of

Protein

Gain

Daily
Average

Protein

of

Age
Per

Days

ioo

Body

Per

iooo

Protein

Weight

Live

Cattle

Soxhlet
Soxhlet
Soxhlet

Soxhlet

Fingerling
Soxhlet
De

Vries

De

Vries

Jzn
Jzn

Fingerling
De Vries Jzn
.

Neumann

De

De

Neumann
De

2-945

45

2.419

45

1.272

2.169

47

1.248

2.

5o

0.880

1.500

50

1.082

1.844

54

1.026

57
62

1.320

2.284

o-939

1.611

Landw.

2-755

1.449

Vries Jzn

3.085

1.693

i-795

1.974

32

40

Fingerling

21

2.803

2.276

Vries Jzn

3.024

2.124

Vries Jzn
Vries Jzn

1.722

1.246

Neumann

De

21

1-335

Neumann

Neumann

3-552

37

Neumann

Vries Jzn

3-994

2.076
1.644

38

Neumann

De

2-347
15
18

Vers.

161

63
64
65

0.678

1.209

0-655

1.209

1.020

1.723

68

0.948

1.7*19

69

1.062

1.823

74

0.713

1.

Stat.,69 (1908),393, results of periods3, 5, 7 and

271

9.

GROWTH

377

Daily
Average

Gain

of

Protein

Age
Per

ioo

Body

Per

iooo

Live

Days

Protein

ioo

0.711

1.

150

0.48

0.83

182

0.41

0.76

214

"-33

0.64

297

0.22

0.47

840

0.064

0.089

Weiske

140

0.372

0.651

Weiske

177

0.307

0.499

Weiske

214

0.219

0.360

Weiske

254

0.288

0.449

Just

285

0.233

o.475

Weight

Cattle

Vries Jzn
Fingerling
Fingerling
Fingerling
Fingerling
Jordan
De

192

Sheep

and

Kern

Wattenberg

290

0.272

0.303

Weiske

293

0.179

0.284

Just

3i5

0.182

0.370

Weiske

0.160

0.264

0.180

Weiske

328
360
366

0.360
0.382

Just
Weiske
Weiske

Just

Kern

and

Kern

and

Kern

39o

0.238
0.158

0.315

405

0.178

0.301

436
458

0.033

0.061

0.068

0.074

521

0.087

0.096

745

0.067

0.069

Si

5-553

9.029

7.269

5.621

6.852

5-757

13

4.129

6.675

Zuntz

21

1.840

3-257

Zuntz

26

0-757

1.470

Wattenberg
Wattenberg
and Wattenberg

Swine

Ostertag and
Sanford

and

Zuntz
Lusk
.

Wilson
.

Ostertag and
Ostertag and
Ostertag and
Tschirwinsky
Tschirwinsky

Zuntz

....

114

0.442

0.663

134

0.483

0.740

5-94

7-73

Dog
Thomas

Gerhartz

10

6.44

7.67

Thomas

16

6.71

8-73

54

1.70

2-35

1.82

2-93

6.10

7-57

5-89

7.91

1.60

3-o5

Thomas
Thomas

101

Cat

Thomas

Thomas

18

Thomas

101

378

FARM

OF

NUTRITION

ANIMALS

singleresults obtained in this

but the general teaching of the
be very considerable,
may
way
table is perfectlyclear,viz.,that the rate of growth of protein
tissue,like the increase in size or in weight,whether expressed
live weight, is relatively
per unit of body protein or per iooo
animal
and decreases rapidlyat first and
high in the new-born
more
slowly later,tendingto be asymptotic to the zero line.
Letting the g equal the gain of protein per day per iooo live
pirical
representedby the emweight and a the age in days, a curve
It is obvious

the

that

in

error

equation1

generalaverage of the observed

cattle and
results on
sheep. With swine, the few results appear
of growth during the first three
to indicate a greater rate
This
is shown
months.
clearly in the accompanying graph
(Fig.38) in which the individual results on the different species
is that represhown
are
by the lightlines,while the heavy curve
sented
the
considerable
Of
course
foregoing equation.
by
be expected, and
individual
variations
to
no
are
particular
well
correspondsfairly

with the

attaches
significance

the mathematical

it would

to

this formula

that

seem

broad

express

in

growth

of farm

ruminants

dog and

cat

on

the

in the young
464.

of

Rate

be

may
the

general

indicate

to

of the curve,

used

way
average
at different ages.

seem

of these

form

but

tentativelyto
of protein
rate
The

higher rate

few

results

of

growth

species.

gain of

energy.

"

While

the

rate

of increase

protein,as discussed in the foregoingparagraphs,may be

restricted sense,
of growth in the more
regardedas the measure
of
of the amount
and while it is of importanceas an indication
proteinwhich must be suppliedin the feed, the actual gain in
normal
less production of fat, as is
or
growth includes more
crease
clearlyshown by the data regardingthe compositionof the inin practice
already considered (458). Growth, therefore,
of

involves

storage of energy

gained

but

is difficult to

of
fattening

also in the

draw

young
1

The

an

body not merely in the protein

it
fat
laid
while
accompanying
on,
in the

line between

exact

the

growth

animals.
equation of

rectangular hyperbola.

and

the

GROWTH

379

XI

"
c/i

a
o
o

00
CO
o
I"

If/OOO/

-""rf m

38o

NUTRITION

OF

assumed

be
If it may
in Table

FARM

ANIMALS

in those

that

of the

experiments

corded
re-

considerable
no
tempted
fatteningwas at79 in which
the increase in weight was
approximatelythat due to

normal

growth,the

of energy
contained
in the increase
the daily rate of gain of energy
and
pounds live
per 1000
be computed.
The
the
following table shows
weight may
results

of

such

comparison,

in direct

being computed

81.

Table

amount

"

the

figures per
the

proportionto

Energy

Content

weight.

Daily

of

pounds

1000

Growth

Energy
of

Average
Live
Weight

Average

Animal

Experimenter

Age

Content
Growth

Per

Head

Per 1000
Lb. Live

Weight
Days

Lb.

Cals.

Therms

Thomas

Dog

0.79

58.13

73-35

Thomas

Cat

0.38

14.67

Wilson

Pig

38.47
17.87

Soxhlet

Calf

Gerhartz

Dog

10

Soxhlet

Calf

15

Thomas

Dog

16

i-55

Thomas

Cat

18

0.864

Soxhlet

Calf

21

Wellmann

Thomas

Pig
Pig
Dog
Dog

101

Thomas

Cat

101

Wellmann
.

Thomas

...

Tschirwinsky
Tschirwinsky
Kern
and Wattenberg
Kern
and Wattenberg
.

Lawes

and

Gilbert

Lawes

and

Gilbert

Kern

and

Kern

and

Kern

and

Jordan
Lawes

Wattenberg
Wattenberg
Wattenberg
....

and

Gilbert

Approximate.

"

Average

0.98
138.60

i5i-53

Sheep,Lot

II

Pig
Sheep
Sheep, Lot
Sheep, Lot
Sheep, Lot
Cattle
Cattle

All data

73-58
2634.00
49-31

24.66
5Q-47

3I53-""

22.74

113.40

73-35

39.20
3294.00

45-36
21.74

23

13-52

243-9

18.04

34

17-93

316.5

17-65

54

Pig
Pig
Sheep, Lot

4.12

106.87

3-38

38.02

11.25

5-95

73-Q4

12.28

1.91

27-34

14.32

114

39-51

568.6

14-39

134

34.66

20.36

290

67.02

705.6
328.0

73-42

608.1

290
300

:
l

4.90

8.28

181.88

5041.0

135.58

1185.4

8.74

106.70

712.9

6.68

27.72

II

456
458

521

102.51

434-5

4.24

745

130.07

520.4

4.00

826.10

840
1460

refer to empty

1272.40

1618.0

1.96

6378.0

5.01

weight,exclusive

of hides.

GROWTH

The

of

gain

of energy

381
thus

computed is notably
carnivora
(dogsand cats)during the first two
greater for young
three weeks than that of pigs or calves.
Aside from this,the
or
results on farm animals,although more
less irregular,
or
present
in general the same
pictureas those on the rate of gain of protein,
viz.,a diminishingenergy of growth with advancing age.
The
few instances showing a wide divergencefrom the majority
rate

probably be

may

" 2.

assumed

The

Relative

values

of

questionof

the

to

rapidfattening.

Feed

of

utilization

of the utilization of
the

be due

Utilization

The
465.

to

as

Growth

in

of protein

proteins for growth.

tion
considera-

"

protein in growth necessarilyraises

relative values

of different individual

teins
pro-

in this respect.
As

that

in

pointedout

was

Chapter IX

(398),it appears

probable

the

is essentially
protein requirement for maintenance
amino
acid requirement and
that the relative values
of
an
proteinsfor maintenance
may
prove to depend largelyor wholly
their abilityto supply certain specific" building stones
on
required for the performance of specificfunctions. In the
there is,in addition to this requirement for
growing animal
"

functional

purposes,

demand

for amino

acids

out

of which

In growth, therefore,
be built up.
the
acid requirements may
differ from those for maintenance

body proteinsmay

new

amino

only

not

in

being quantitativelygreater

different.

but in

striking illustration of
of Osborne
and Mendel
by the investigations
of lysinto growth.

tively
being qualita-

this is afforded
1

on

the relation

investigators
they have found that tryptophan
maintenance
Wheat
tains
(399).
gliadin conminute
of
but
While
amount
only a
they
tryptophan
lysin.
have repeatedlysecured maintenance
for long periodson rations containing
the
sole
have
been
unable
to
secure
gliadinas
protein,they
but the simple addition
of lysin enabled
such
with
rations,
growth
The body proteinscontain lysin,
growth to proceed at a normal rate.
ox
muscle, for example, yielding7.6 per cent (50). Evidently this
In

common

is

with other

indispensablefor

Jour. Biol. Chem.,

12

(1912),473;

17

(1913),325;

26

(1916), 293.

382
acid

amino

in order

in the feed

in the

molecules

be

cannot

FARM

OF

NUTRITION

ANIMALS

synthesizedin the body

to permit the construction

tissue,while for maintenance

but

must

of the

be

supplied

new

protein

(399) it appears
that the addition

to

be

adequate
to indispensable. Moreover, they have shown
proteinslike gliadinof other proteinscontaininglysinpermits
that the proportion of the
growth to take place and furthermore
in order
be added
must
to support normal
second protein which
growth is less in proportionas it is richer in lysin.
conclusions
have been strikingly
and Mendel's
confirmed
Osborne
x from
Nollau
and
Kastle
obtained
feeding
by Buckner,
by the results
and
low
mixtures
of
content.
chicks grain
lysin
high
young

that the lack of

It appears
of supportinggrowth,

capable
lysin in a proteinrenders it inalthough it may stillbe adequate
for maintenance
(399),and that the proportion of lysinin those
proteins containing it constitutes a limiting factor for the
of growth which
amount
they can
Tryptophan is
support.
obviously another limitingfactor in this respect, while it must
be regarded as
acids
altogether probable that other amino
become
limitingfactors
belong in the same
category and may
is deficient.
In other words, the amount
if the supply of them
of some
particularamino acid which is available may become
factor which
determines
the rate of growth, just
the minimum
the minimum
mine
as
supply of potassium, for example, may deterThe unsatisfactoryresults
the rate of growth of a crop.
in

obtained
animals

practicewith
be

well

may

maize

due

in

as

the

large part

to

sole
the

feed

for

poverty

young
of the

grain in tryptophan and lysin,it having

of protein they can
that as the sole source
been shown
support
but slow growth (783).
Unfortunately the knowledge available on these pointsis as
sufficient
and affords no
yet chieflyqualitativein character

proteinsof

mixed

foundation

which

on

relative

values

the

of

case

for the

this

of

to

base

proteins in

growth

as

quantitativediscussion

in that

farm

practice. Accordingly, in

of maintenance

consider

of the

it appears
questionsregarding the

sary
neces-

protein
requirement upon the basis of total protein,
largelyirrespective
of its nature.
(Compare Chapter XVII, " 4.)
466.

present

Percentage

animal,

the

to

retention

katabolism
1

Amer.

of

of feed

protein.
"

In

the

mature

protein substantiallykeeps

39 (1915),162.
Jour. Physiol.,

pace

3^4
Table

83.

With

Percentage

"

FARM

OF

NUTRITION

Feed

of

ANIMALS

Protein

Retained

Fingerling

"

relativelysmaller retention is observed.

Thus Neumann
obtained for calves 40 to 70 days old percentages varying
from 38.7 to 48.3, and Tschirwinsky,experimenting on
pigs 100
of
of the
observed
retention
to
to 120
cent
a
old,
days
33.6 per
20.7
digestedprotein. With stillolder animals a yet smaller percentage
retention has been observed, diminishingto nearly zero
with fully
advancing

age,

animals.

mature

467.

Does

not

utilization.

measure

greater percentage retention

of feed

utilization

the

animal

is made

statement

and in that

sense

been

the

basis

customary

proteinis high in
rather rapidlyas

diminishes

and

it has

On

"

the

case

it grows

say that
of the young
to

older.

This

essentiallyfrom a commercial
standpoint
animal
is
the
is capable
it
true.
growing
Only

of feed proteinto increase

using any largeamount
of body protein and the abilityto do this is the more

of

its stock

marked

the animal.

the younger
The

of this

of the

retention

percentage

necessarilyvariable

and

with

the

which

neither
animal

feed

affords

protein,however,

of the efficiency

measure

it into

converts

is

body protein

at different ages.
efficiency
two
important factors to which
viz.,the
especiallycalled by Fingerling,1

of that

nor

The

permits a comparison
comparison is disturbed by
has

attention

of the total amount

influence

of

been

deficient energy

468.

Influence

implied,growth
The

feed

proteinsuppliedand

the effect

supply.

of
is

of

protein supply.

already been
factors.
biological

has

As

"

primarilydependent upon

suppliesmaterial

for

growth

does

but

not

determine

lated
protein as formuin the previous section
(463) represents (so far as the
results are
trustworthy)the capacity of the animal for protein

its maximum

rate.

The

rate

Landw.

Vers.

of increase

of

Stat.,74 (1910), 1.

38S

GROWTH

different ages, but the percentage of the feed protein

this
will depend upon
the relation between
is retained

storage
which

at

of protein actuallysupplied. For

capacity and the amount
example, suppose a calf weighing 100 pounds to be capable of
storingup per day 0.25 pound of protein and to require0.05
If it receives 0.35 pound digestible
pound for maintenance.
amount
proteinin its feed and is able to store up the maximum
this ration,71.4 per cent of the digestible
of 0.25 pound on
protein would be retained,while 28.6 per cent would katabif the feed
in the urine.
But
olize and its nitrogen excrete
of the animal supplied0.45 pound digestible
protein,the gain
still be 0.25
for the animal, but

would

would

only
The
organism is unable

olized.

as

this is the

since

maximum

possible

the percentage of the feed proteinretained

55.6,while 44.6 per cent of it would be katab-

be

pound

pound,

for constructive

and

purposes

Chapter IX (402-404)and

in

shown

to

In

other

suppliedin

the ration

in the

body.

On

of

excess

requiredby

growth and maintenance, the

retained

simplyas

serves

the minimum

over

one-tenth

therefore it is katabolized

words, the greater the

of energy.
of

the added

use

lower

protein
the

will be the

hand,

the other

source

mands
decentage
per-

with rations

protein the percentage retention will increase with

to
the proteinsupply up to the minimum
amount
necessary

deficient in

utilize the

growth capacityof

the animal.

confirmation of the truth of

Fingerling's
experimentsafford striking
the foregoingdeductions from the generallaws of proteinkatabolism.
A calf received daily in one
period 8 kgs. of whole milk with an
addition
of butter
fat and
lactose,while in the succeedingperiod
alone
milk
fed
in
whole
amounts
was
proportionalto the age of the
calf,averaging 11.875 kgs. per day. The results as regardsprotein,
follows :
as
expressedin terms of nitrogen,were
"

Table

84.

Influence

"

of

Protein
of

Supply

Grams

2-5

42.49

25-30

62.97
2

Percentage

Retention

Nitrogen

Digested
Nitrogen
Feed
of

June
June

on

Per

Urinary
Nitrogen

Gain
Calf

Grams

Grams

7.91

28.77

by

Cent
Feed
Protein
Retained
of

34.58

81.4

34.20

54-3

386

NUTRITION

ANIMALS

FARM

OF

sufficient in the first period to

Evidently the proteinsupply was
normal growth. The additional
ensure
supply in the second period,
therefore,had no effect on the gain but simply increased the protein
of energy
for
the added
katabolism, i.e.,
proteinwas used as a source
maintenance
for the production of fat.
or
On the other hand, a supply of proteinnotably insufficient to permit
normal
tention.
gain may yet show a comparatively high percentage rethe same
Thus
calf received in an intermediate
periodonly
milk togetherwith sufficient butter fat and
4 kgs. per day of whole
As compared with the first
lactose to supply the necessary
energy.
periodonly about one-half of the normal gain of protein was secured,
reduced.
yet the percentage retention is but slightly

Table

85.

"

High

Percentage

Retention

of

Nitrogen

cient
Insuffi-

on

Protein

Digested
Nitrogen
Feed
of

Urinary
Nitrogen

Grams

June 2-5
June 13-ii

469.

Grams

Cent
Feed
Retained

Per
by

of

Grams

42.49

7.91

34.58

81.4

19-95

5-5

14.90

74-7

of deficient

Influence

Gain
Calf

supply.

energy

"

But

only

not

in the

of energy
surplusof protein be utilized as a source
but if the energy
manner
supply in the feed
just illustrated,
be diverted from
is inadequate protein may
growth to serve
in the case
fuel material,precisely
of maintenance
as
as
(412),
thus lowering both the observed
tention.
gain and the percentage remay

This

effect is well illustrated

by the followingexperiment by Finthe first two

periodsa limited quantity

gerlingupon a calf receivingin

(10 kgs. per day) of whole milk.
insufficient and
supply became
energy
purposes

so

diminished.

that

the

When,

actual
in

the animal

protein

was

grew

older

diverted

to

the
fuel

the percentage retention both

placed
reperiod,one-half of the milk was

gain

third

As

and

by butter fat, the protein supply being kept at nearly the

level by the addition of egg albumin, the actual gain rose nearly
same
to its original
level and the percentage retention became
even
higher
somewhat
than at first on account
of the
reduced proteinsupply.

387

GROWTH

Table

86.

Influence

"

of

Supply

Energy

Per

Digested
Nitrogen

7-9

Oct.

Meaning

Gain
Calf

cent

Feed
Protein
Retained

by

of

Grams

Grams

Grams

51.84
51-87

12.62

39.22

75-7

19.99

31.84
37.66

61.5
82.3

8.10

45-76

19-27

470.

Urinary
Nitrogen

Feed

or

Oct.

Retention

Nitrogen

of

Sept. 29-Oct.

Percentage

on

of utilization.

The

"

ible
percentage of the digest-

protein of the feed which is retained in the body of the

of the efficiency
growing animal, then, is not in itself a measure
of the animal
organism in converting feed protein into body
protein,since the proportion retained is affected both by the
magnitude of the protein supply in the feed and by the energy
of the

content

ration.

What

then is the

conceptionof

correct

the utilization of the feed

protein?
of protein
As appeared in the previoussection,the amount
which a growing animal can
store
to be a function of
up seems
its age (463),and the attempt was
made
to formulate
mately
approxiin
the capacity for growth
this sense
at different ages.
The
logical
percentage utilization of the feed protein in the physiois
sense, as distinguishedfrom the percentage retention,
the ratio between
the body proteinthus stored up and the least
of feed

amount

which
favorable

proteinin

is necessary

to

of the maintenance

excess

ment
require-

support this growth under

as
conditions,especially

to energy

the most

supply. Suppose,

for

example,that an animal three months old actuallyhas the

capacity,as computed by the formula on page 378, to store
pounds live weight,and
up daily1.23 pounds protein per 1000
that it has been shown
that it can
just reach this capacityon
ducting
a ration supplying 2 pounds of digestible
proteinper day. De(415),there remains 1.5
0.5 pound for maintenance
pounds of protein in the ration out of which is produced 1.23
pounds of body protein. The utilization is therefore 1.23 -fIf, on
2.5 pounds of protein had
1.5

82

per

cent.

the other
to

be

hand,

suppliedin

it

was

found

that

the ration in order

388

bring the gain

to

the

percentage

cent, while

with

be

100

results.

determinations

capacityof

only

1.23
maximum

if the

hand

per cent.
The
writer

is

"

of the

4-

the

2.0

animal,
62

growth
digestibleprotein, the

of

pounds

1.73

Experimental

exact

any

other

ANIMALS

the

to

would

evidentlybe

would
471.

protein up

utilization
the

on

secured

be

of

FARM

OF

NUTRITION

percentage

not

per

could
zation
utili-

of

aware

utilization in the

of protein
amount
justdefined,that is,of the maximum
be produced either from singleproteins
tissue which
can
or from
but interesting
data regardthe mixed proteinsof feedingstuffs,
ing
the utilization of proteinby growing animals
furnished in
are
x
experiments by Fingerling upon calves and by Just 2 on lambs
the influence
in which
of a varying proteinsupply upon
the
sense

nitrogen balance was determined.

Reckoning the maintenance
requirementfor protein at 0.5
results for those
pound per 1000 pounds live weight,Fingerling's
periodsin which the estimated capacity for growth appears to
have

been
Table

fullyutilized
87.

"

were

Computed

follows

as

Utilization

Gain

"

Protein

of

tein
Pro-

of

by

Feed

Protein
Excess
of
Maintenance
in

PER

Calves

Percentage
Utilization

IOOO

Aver-

per

age
Animal

Period

Landw.

Vers.

Age
Days

ObTrue
served
ity
for
Protein

Capac-

IOOO

Crude
Protein

True
Protein

1.94

2.00

34.

O.63

O.62

O.84

O.76

0.85

2.15

Crude
Protein

Gain

Gain

172

O.70

O.67

175

0.63

157

2-44

39^

34-8

33-5
75-o

101,

184

0.66

o-75

O.79

1.04

94.

72.1

211

O.58

0.68

2-59

2.85

26.

23-9

0.53

o.57

I.08

1.29

52.

44.2

237
262

0.48

O.S4

1.

i-37

47-

39-4

309

0.41

0.50

O.31

0.49

161.

102.0

339

0.38

0.44

O.25

0.42

176.

104.8

Prelim-

inary

135

O.87

1.07

2.42

2.81

44-

1-6

187

O.05

0.80

O.74

1.01

Stat.,76 (1912), 1.

13

108.

38.1
1

79.2

Ibid.,69 (1908),393.

GROWTH

389

figuresit appears that in the low protein periods

the estimated capacity of the animals for growth was
lized
fullyutiwith a surplusof digestible
true proteinover
the maintenance
less than that actuallyrecovered
requirement equal to or even
in the growth, while a much
largersupply of protein failed to
secure
any additional growth but simply forced up the protein
In other words, if the estimate for the maintenance
katabolism.
requirement is approximately correct, the utilization of the
have
digestibleprotein in the low protein periods must
proached
ap100
Indeed, in at least two cases it is necessary
per cent.
From

high

these

to

admit

or

that

Table

either

that

the

non-proteinwas
Computed

estimate
used

for maintenance.

Utilization

Substituted

for maintenance

for No.

of

II.

Protein

by

Lambs

is

too

ANIMALS

FARM

OF

NUTRITION

390

direction are contained

Interestingdata pointingin the same
value
of
the nutritive
in the investigationby Just, in which
was
compared with that of protein.
non-protein for lambs
requirement at 0.5 per thousand
Estimating the maintenance
and computing the results of the proteinperiodsas in Fingerthe
that in nearly every
case
ling'sexperiments,it appears
actual gain of proteinwas
only slightlyless than the surplusof
the maintenance
crude proteinabove
requirement,
digestible
it was
while in many
cases
distinctly
greater than the digestible
at
true
protein available. Apparently the non-protein must

to

Neither
to

solve
of

be very

true protein
digestible

been very

their results indicate

proteinin growth,but
high

should

and

the

experimentsupon

lead

General
for

values

for maintenance

growth

is

net

"

conception.

values

caution

to

in the

that

it may

of
interpretation

proteinrequirementsfor growth.

Utilization of energy
472.

if not

high (Table 88).

nor
Just'sinvestigationsare adequate
Fingerling's
tion
the general problem of the maximum
possibleutiliza-

have

must

the utilization of the

growth,while

its

of the animal

to the maintenance

contributed

least have

energy

The

"

values

for growth

conception of
that

net

energy

of net

entirelyanalogous
energy
for
or
fattening. They represent that
to

of the maintenance
suppliedin excess
energy
is able to store
the animal
up in the gain
in mind
It is important to keep this conception clearly

portion of the feed

requirement which
made.

consideringthe utilization of feed in growth and not to

of the young
animal
be misled by the greater economic
efficiency

when

as

producerof

It is

live

familiar

weight increase.

fact

that the

young
when
more

animal

gains in weight

and this has

faster than
mature
relativelymuch
animal
utilizes its
led to the generalimpression that the young
feed more
perfectlythan the older animal, or in other words,
that the net energy value of a feedingstuff for growth is greater
for fattening. It is true that the
than that for maintenance
or
gain in live weight is different in character in the young animal,
therefore
and protein and less fat and
water
containing more
less energy
(458,459) but on the other hand the results recorded
in " 1 show a greater rate of growth as regardsboth protein and
,

ANIMALS

FARM

OF

NUTRITION

392

(366,759), Rubner's
explained in Chapters VIII and XVII
with the energy
ture
expendi"specificdynamic action" is synonymous
if
from
it be subtracted
caused
by the consumption of feed, and
As

is the net

the remainder

the metabolizable

energy
utilization is of

the

course

net

energy

The

energy.

centage
per-

divided by the metabolizable

energy.

be
the percentage utilization would
in this way,
of the followingtable,the second
in the first column

Estimated
shown

as

and

of which

third columns

utilization

the

show

by the writer both with and without

estimated
the fastingkatabolism
as

io

cent

per

computed

as

addition

the data

from

to

mature

on

animals.

Table

89.

Utilization

Estimated

"

Energy

Metabolizable

of

of

Milk

Computed

Writer

by

Fasting
Computed,

Katabolism

Fasting

Using
Rubner's
Factors

Katabolism
in
Same
as
Mature
Animals

Greater
than in

10%

Mature
Animals

Rubner's

84.08

73.10

Soxhlet's

86.18

73-77

75-56

83-99

70.31

75-47

experiments
experiments
Wilson's experiments

it is clear that

While

final conclusions

no

be based

can

upon

been,
figuresobtained as
that the actual experiments
it seems
suggestive,nevertheless,
utilization than
lower
show
with
a
growing animals
average

small

differences

be

would

which

Wilson's

Rubner's

results

ttf the
sense

notion

by

animals.

that
young

these

the

animals

those

Kornauth

and

swine

comparisons afford
of energy

is much

animals.
than

apparently lower

utilization

have

mature

be

grain (757). Certainly

mature

are

results upon

computed from Meissl's and

experiments upon mature
respiration

may

Arche's

between

expectedfrom

Moreover,

these

consuming
little support

in the

higher than

logical
physiothat

by

GROWTH

Experiments

474.

older animals

older

on

those

of Kern

393

animals.

Of

"

experiments upon

and

Wattenberg on lambs and of

Tschirwinsky on pigs (458),permit an approximate computation
of the utilization of the feed energy
during growth and
afford data for some
comparisons, although in neither case
animals employed, the lambs
were
being between
very young
6 and 7 months
old at the beginning of the experimentand the

pigsbetween
the

On

if

9 and

whole, the

results of these

rather lower

anything,a
animals

younger

fail to show

weeks.

10

compared
on
superiority
as

any
of the results of

is true

experiments seem

cate,
indi-

to

percentage utilization by the

with the older.

At

any

the part of the former.

they

rate

The

same
l

experiments by Armsby and Fries upon

old in which the availability
steers
to 27 months
mined
deter10
was
cisive,
While not deby the use of the respirationcalorimeter.
the results seem
lower availability
to indicate a slightly
of
the mixed grainand possiblyof the hay for the younger
animals.
475. Embryonic
Several experimenters,especially
growth.
2
and Bohr
and Hasselbalch,3have
Tangl and his associates
"

determined

the

expended

energy

in

the

development

of

the

in

embryo

oviparous animals, i.e.,in the organizationof the

substances
of the egg into embryonic tissue.
These
gations
investihave shown
that a relativelylarge proportion of the
chemical energy contained
in the egg is evolved as heat during
the process of development, so that the percentage recovered
in the embryo, ranging from 60 to 68 per cent, is distinctly
lower than the utilization of the energy
of milk by suckling
animals

as

computed

in

previous paragraph (473).

over,
More-

they show

that the utilization of the energy of the egg is

notablyless in the earlier than in the later stages of incubation,
low

as

figureas

28

per

cent

having

been

observed

after

10

days'incubation.
The

method

by Tangl
similar

and

eggs

contained

may

Mituch,
from

the

U. S. Dept. Agr., Bur.

Arch.

Skand.

each

same

by the results of

upon

Cals. and

291.38 Cals. chemical

Anim. Indus., Bui. 128 (1911), 51.

93 (1903),327;
98 (1903),490;
Physiol. (Pfliiger),
and

experiments
analysesof

two

three hens' eggs.

From
hen, it was computed that the three

respectively
229.72

(1908),423

be illustrated

104

and 353 ; 14

energy.

(1904),624;

437.

Arch. Physiol.,10 (1900),149

used

(1903),398.

121

and

its contents

sack and

results

the embryo

of incubation

the end

At

ANIMALS

FARM

OF

NUTRITION

394

the energy

90.

Utilization

"

of

Energy

Incubation

in

Eggs
Hen

Originalenergy of eggs
Remaining in yolk sack and
for

contents

in

embryo

used

loss of dry

165.77 Cals.

196.74 Cals.

Cals.

125.66 Cals.

59.87 Cals.

of
substantially

fat.

loss of

No

63.84 Cals.

of the egg

substance

Comparison with the

thus katabolized

that the material

showed

94.64 Cals.

35 to 40 per cent of the energy

recovered but escaped as heat.

matter

from

63-95 Cals.

words,

not

was

VIII

291.38 Cals.

99.24

Percentage recovered

In other

Eggs
Hen

from

Cals.

229.72

productionof embryo

Recovered

yolk

lowing
with the fol-

determined

of each

"

Table

Used

the

separated from

was

consisted

observed.

nitrogenwas

and reptilian
embryos, especially
Experiments on mammalian
by Bohr,2 by Murlin 3 and by Carpenter and Murlin,4appear in
that the
accord
with the foregoingconclusion,since they show
metabolism

of

greater than

that of the mature

the

embryo

per

of

unit

weight is as great or
animal, despitethe fact that the

requirement of the former must be decidedlyless.

of the organThe
growth of the embryo consists essentially
ization
gen
The fact that there is no loss of nitroof protein tissue.
would
indicate
that
chemically the
during incubation
is effected by a cleavage and
resynthesisof protein
process
which
to be a nearly isothermic
(233, 367 d).
process
appears
is
Apparently the organization of the protein into structure
what
calls for the largeexpenditure of energy.

maintenance

476.

Summary.

"

foregoingparagraphs
statements

In

of the

the

case

same

may

experimentalresults mentioned
in the
be briefly
summarized

in the
ing
follow-

"

of

sucklinganimals, while no direct comparisons

animal
the utilization
at different ages are
available,

of the metabolizable
1

Including the

Skand.

3Amer.

The

egg

energy

of milk

for

growth

appears

to

be

membranes.

Arch. Physiol., 10

(1900), 413;
Jour. Physiol.,26 (1910),134.

15

(1904),23.
4

Arch.

Inter. Med., 7 (1911), 184.

GROWTH

would

distinctlyless than
on
case

395

expected from

be

Rubner's

animals.
the utilization of pure nutrients by mature
the utilization appears
to be
of swine, moreover,

than

seem,

that of the metabolizable

The

compared

as

results

in

than
and

477.

draw

and

ones

at

any

rate

fail to

case.

show

relativelylarge

the broader

for the
it

with

in

energy

to

the

more

structure

transformation

embryo.

mature

hypothesis.

formulate

embryonic

of the

of egg substances,
as
relatively
greater in the young

be

final conclusions

permissibleto

sense,

chemical

mere

seems

Provisional
any

older

embryonic growth

on

in
organization,

compared

with

development and indicate a comparatively

utilization of energy.
This large expenditureof
to be required chieflyfor the
development seems

low

rather

less

even

of

energy

greater in the former

that it is any

expenditureof
energy

In the

mals,
anigrain by mature
pated.
although the contrary would naturallyhave been anticiThe results with older animals,while far from conclusive,
if anything,to indicate a lower utilization by younger

animals
show

results

While

"

from

the

it would

be

foregoingdata, it

rash

to

may

be

working hypothesisto the effect

of feed protein (includingthe proteinof
that the conversion
the egg) into tissue requires a considerably greater relative
expenditureof energy than does the conversion of surplusfeed
what might be called the
into fat, the difference representing
work of organization,
of organizedstructure
the formation
i.e.,
and especially
animal
in the young
in the embryo.
It has
been shown
(463) that the rate of growth decreases rapidly
with increasingage.
Accordingly,the work of organizing new
proteintissue,so far as this is measured by the storage of protein,
constitute a steadilydiminishing proportionof the
must
total energy
older
grows

expenditureof
the

increase

the

organism,since

consists

to

as

the animal

diminishing extent

of

increasingextent of fat. The percentage

utilization of the feed energy would therefore,
sis,
upon this hypotheIt would be least immediately after birth
tend to increase.
and after two
would
become
to four months
relativelysmall,
of
the gain. Probably
correspondingto the changing character
by the time an animal has been weaned and is consuming the

proteinand

to

an

normal

feed of its

energy

might be assumed

the percentage
species,
to

be

not

utilization of the feed

much

less than

that

ex-

396

OF

NUTRITION

hibited

proportional

to

energy

values

might

be used

absolute
The

also to

an

the

for

of

rate

to

of course,

mean,

for maintenance

be

the

practically

that
and

at least their relative

measure

values

at any

the

net

fattening
if not

their

growing animals.
clusion
validityof this provisionalcon-

and profitable
field for investigation.
interesting

The

Contrast

would

feeding stuffs

energy
determination

" 3.

animals

of

net

offers

478.

This

it.

and

animal

the mature

by

ANIMALS

FARM

Feed
with

Requirements
fattening.
"

conception of

In

for

the

Growth
case

of

fattening

the feed

as
requirement,particularly
since the extent
of the
artificial,

regards energy, is somewhat

fatteningdepends, within the limits of the animal's capacity,
of feed supplied. Growth, on
the
largely upon the amount
other hand, unless the feed fails to supply the necessary materials
tially
and thus becomes
at a rate substanon
a limitingfactor,goes
obvious
of
determined
by other conditions,the most
which
the species,individualityand
are
age of the animal.
Indeed, it may be said that,within normal limits,the capacity
for growth determines
the feed consumption rather than
the
cause
reverse.
fatteningbut it does not
Heavy feedingmay
of the higher animals, to materially
appear, at least in the case
accelerate
growth, although Eckles * observed the growth of
pared
more
dairy calves to be somewhat
rapid upon heavy as comnance,
with scant
rations.
in mainteIn growth, therefore,
as
ing
bethere is a real requirementto be satisfied,
its measure
the

animal

amount

is

and

character

capable of making

of the increase
under

normal

the young
conditions.
which

results of experi(372) of the interesting

ments
2
under
adverse
Waters
conditions, while
by
growth
upon
3
has been susand Mendel
have
shown
that growth which
Osborne
pended
for a time because of inadequate feed supply may
be resumed
when
this deficiencyis made
good (deferredgrowth). Neither of
these possibilities,
however, invalidates the statement
justmade that
the continued
maintenance
of a normal
rate of growth requiresa
and energy.
definite supply of matter
Mention

Mo.

Soc. Prom.

has been

made

Expt. Sta., Bui. 135, 1915.

Meeting, 1908, p. 71.
Agr. Science, Proc. 29th Annual
3
Amer.
23 (1915), 439;
Jour. Physiol.,40
Jour. Biol. Chem., 18 (1914), 195;
(1916), 16.

GROWTH

Total

479.

increase

in

397

normal

growth

at

different

ages.

"

requirementsof the growing animal as regardsprotein

which such
and energy depend in the first place on the amounts
animal is capableof storingup in normal growth. From
the
an
data regardingthe rate of growth recorded in " i of this chapter,
to
even
though they are somewhat fragmentary, it seems
possible
derive average figuresregardingthe storage of proteinand energy
value as a guide
in growth at different ages which may be of some
in estimatingthe feed requirementsof the growing animal.
it was
that the rate of gam
shown
As regardsprotein,
per
live weight apparentlydoes not vary
widely as between
iooo
cattle,sheep and swine, and an empiricalformula
(463) was
feed

The

given by which

its amount

estimated.

regards energy,

As

for farm

at

animals, but

any

fewer

the

be

approximately

are

available,especially

8i

(464) shows

ing
diminish-

the animal

grows older.
ent
the followingtabulation the daily gain of protein at differof

rate

In

may
data

in Fig.
graphicrepresentation

of the results recorded in Table

39

age

gain of

energy

as

calculated by means
of the formula justmentioned
and the gain of energy estimated from the smoothed graph

ages has

of

Fig. 39.

been

The

of
expression

Table

91.

"

two

togethermay

the normal

Daily

Increase

Age

io

20

30
60
90
120

150
180
210

240

270
300
1

I
2\
1

increase

in

Growth

be taken

in

per

growth at
iooo

approximate

an

as

different ages.

Pounds

Live

Weight

Protein

Energy

Pounds

Therms

days

4-50

24-5

days
days
days
days
days
days
days
days
days
days
days

3-38

21.8

2.70

20.0

I.50

16.O

I.23

13.O

O.96

n-5

0.79

10.0

O.68

9.0

0.59

8-5

0.52

7-5

o.47

7.0

0.42

6-5

year

0.35

5-5

years

O.24

4.0

years

O.I5

2-5

39"

NUTRITION

OF

FARM

ANIMALS

B
o

a
o
o
o

""

"

Therms

per

iooo

pounds live weight

NUTRITION

400

Table

92.

Average

"

Gains

by

Live

ANIMALS

FARM

OF

Growing

Calves

per

Pounds

iooo

Weight

approximate estimates may be made for

the pig from
data reported by Henry.2 At weekly intervals,
up to 70 days old,the computation is based on the live weights
and
in Henry's table. The
gains shown
gains by the older
b. Swine.

animals

Similar

"

estimated

are

larger breeds
assumed

should

that

the

uniform

be

would
old.

The

weigh 250 pounds at one

decrease
rate of gain would

that the results

it

being
mately
approxi-

year,
at

that the

an

reach
energy

the heavier

on

animals

compiled

as

since it may
by heavy
fattening,
mals
for the weights and gains that the anithe figures

from

that

of his statement

basis

obtained

were

computed

the

rate.

It is evident

by Henry

on

weight of

content

it

feedingfor

would

per

about

pound

pounds at about
of gain is estimated

320

10
on

months

the

sumption
as-

uniformly from the average of

about
Cals. obtained
for young
by Wilson and by Wellmann
500
to
pigs (458)up
2485 Cals. as computed in Table 65 of Chapter X
for
Soxhlet's
swine No. 3 at 500 days old.
(442)
1
3

increase

Average of Soxhlet's experiments.

Loc. cit.,
p. 553.

Feeds

and

Feeding, 10th Ed.,

p. 499.

GROWTH

Table

93.

Average

"

Gains

401

Growing

by

Pigs

per

iooo

Pounds

Live

Weight

482.

Estimated

averages.

"

The

foregoing results

have

plotted in Fig. 39 (479) for comparison with those

derived from the computationsof " 1, regardingthe rate of
By drawing smooth curves
through the
storage of energy.

been

results

for

calves

approximate

by

these

appear

for

the
pigs, respectively,

following

of the average
of gain of energy
rate
similar data
No
species have been obtained.

estimates

two

to

and

be

The

available

for

other

speciesof

farm

animals.

to indicate that the results for cattle

graph would seem
for the
well also to sheep,althoughno figures
may apply fairly
latter speciesare
recorded below 300 days old.
Preliminary
assumed
the
basis
of the live weights
by
computations on
Kellner for lambs
at different ages seem
to indicate higher
for animals of the
figuresfor the first six months, especially

mutton

breeds.
2

OF

NUTRITION

402

Table

94.

Estimated

"

Rate

of

Pounds

FARM

Gain
Live

ANIMALS

Energy

of

Day

per

and

iooo

Weight

Cattle
Age

(and

Swine

Sheep?)

days
60 days
90 days
120
days
150 days
180 days
210
days
270 days
365 days
30

Therms

Therms

18.O

20.0

13-5

14-5

11.0

II.O

9.0

9.0

7-5

8.8

6.0

7-5

18 months

5-25

7.0

5.o

6-5

4-75

6.0

4-5

24 months
30 months

4-25

4.0

It cannot

be

claimed

that

the

foregoing computations

are

The
ment
data are
particularlysatisfactory.
scanty, and the eleof personal judgment unavoidably enters, especially
into
the

estimates

of the

value of
energy
there
Nevertheless, while

weight.
divergenciesat

unit

of increase

in live

considerable
very
is after all a certain general

certain

are

points,there
and they may
agreement in the results,
perhaps serve as a first
an
approximation towards
expression of the growth capacity
of farm

animals

in terms

of energy

storage.

It is much

to

be

factor in determining the

regretted that such a fundamental
feed requirements for growing animals
known
is so imperfectly
and the determination
of the amount
and composition of the
increase in growth at different ages, whether
of comparative
by means
calorior
slaughtertests or with the aid of respiration
metric experiments,offers an interesting
field for investigation.
With
the smaller animals, such as pigs,lambs and particularly
fowls,it would appear that such determinations might be made
without
483.

great
Total

attempt
per

ages.

An

energy

show

requirements.

"

The

foregoing figures

approximately the actual storage of energy

pounds of live weight by growing animals at various
adequate ration for such an animal, however, must

to

1000

difficulty.

GROWTH

not

only supplynet

made,

sufficient net
total

the

ages

energy
and from

94

maintenance, the

for

energy

net

Table

from

equalto that contained in the growth

by the foregoingtable, but in addition

energy

indicated

as

403

required by

of the two

animal.

the

the estimated

storage per head

the energy

sum

live

weight

being

Computing
at

different

adding the maintenance

and

requirement computed in proportionto the two-thirds power

of the live weight (347) gives the total energy
requirements
shown

by

Table

b of the

IV

Appendix.

Protein
484.

Minimum

requirements

requirement.

As

"

with

the

energy

of

the

a
feed, the proteinsupply of the growing animal is essentially
certain
factor.
A
deficient
lacking
supply or one
limiting
check
porarily
essential
(465), may
building stones"
growth temor
permanently through simple lack of material,but
that a surplus of protein can
it does
not
materially
appear
"

growth.
of the estimates of the
Granting the approximate accuracy
actual gain of protein in normal
on
growth made
previous
f
eed
of
the
quired
(463, 479),
protein requantity
digestible
pages
in the ration of the growing animal
at any
particular
be conwhat proportionof the latter can
verted
age will depend upon
the percentage
into body proteinand stored up, i.e.,
upon
in the
utilization of the feed protein (470). As was
shown
preceding section, however, this is very imperfectlyknown.
and Just'sresults (471),it be
If, on the basis of Fingerling's
assumed
that the utilization may
approach 100 per cent, then
stimulate

estimated in Table

the amounts
about

of

the rate

0.5 per

for maintenance, would

1000

digestible
protein which must
normal increase of proteintissue.

of

485.

Results

in

experienceseems
proteinthan is
if not
there
the

are

few

(479),with the addition of

91

practice.
"

show

to

that

As
a

suppliedto support

be

be the least amounts

the

of

fact, however,
liberal supply of feed

matter

more

indicated

necessary

tageous
by these estimates is at least advanWhile
in the actual rearingof animals.

on
investigations

determination

of the
1

record

minimum

directed

to
specifically
protein requirements of

In direct proportion to the live weight.

NUTRITION

404

OF

growing animals, there

especially
upon
of live

the increase

are

immature

weight

has

reasonably similar
protein.

ANIMALS

considerable

otherwise
of

FARM

number

of

fatteninganimals, in
been

but

determined

upon

ments,
experiwhich
rations

containingvarying proportions

In the immature

safe to assume
that the
fatteninganimal, it seems
of
is
to the
maintenance)
protein(in excess
appliedsubstantially
of
and
this
that
the
with
growth
growth goes on parallel
support
less independent of it. There appears
or
fatteningprocess but more
evidence that protein specifically
ing,
to be no
stimulates or aids fattenthat conclusions regarding the protein supply drawn
from
so
fatteningexperimentsmay be regarded as applicableto growth without
fattening.
feed

If in such

of feed
experiment, in which the total amounts
consumed
do not
differ widely, it appears
that
the smaller
of protein has been as efficient as the largeras regards
amount
in
to be normal
gain in live weight, and if the gain appears
amount, there is a strong presumption that the lesser amount
of protein was
for
at least sufficient for the needs
of the animal
growth and maintenance, while if a block test shows a normal
character
of increase this presumption is further strengthened.
Obviously, results of this sort cannot be relied on to fix definitely
the lower

an

limit of

proteinsupply,but they may

furnish indications

regardingit.
486.

Experiments

with

cattle.

"

In

the

experiments upon

calves

included in
by Soxhlet, De Vries Jzn and Neumann,
Table 80 (463),
showing the rate of gain of protein,the amounts
of digestibleprotein consumed
as
compared with the actual

gains were

as

follows

Table

95.

"

"

Protein

Consumed

by

Calves

GROWTH

405

the

light of Fingerling'sresults upon

suckling calves
(466),however, there can be little doubt that the proteinsupply
in these experiments was
unnecessarilygreat, especiallywith
In

the older animals.

The

writer

has

elsewhere

weight results bearing upon

fattening cattle which seem
than
at

the

might

several

be deduced

ages.

altered
slightly

form

Those

and

discussed
the
to

indicate

the

experiment by
to a degree the judgment of
to be

as
interpreted

Table

96.

"

The

much

here

are

addition

Protein

tabulated

of

the writer and

as

Requirements

course

the

sents
repre-

figuresare

determinations.
Cattle

of

foregoingestimates correspondin general with

The

in

subsequent

of

summary

indications rather than

Estimated

higher requirements
gains of protein

the actual

results

Schneidewind.2

of immature

protein supply

from

with

of the earlier live

some

the

tein
pro-

requirements for growing cattle as formulated in the WolfTLehmann

and Kellner feeding standards
(790-793) In ments
experi.

upon

two

steers, directed

principally
to

other

questions,

Armsby and Fries observed a normal rate of increase in weight

of digestible
rations containing amounts
protein much
upon
smaller than are called for by current
feedingstandards,although
of the estimated normal gain of proteinfor the corstillin excess
1

U.

S. Dept. of Agr., Bur.

Anim.

Jahrb., 36 (1907), 687.

S. Dept. of Agr., Bur. Anim.

Indus., Bui. 108 (1908), pp. 60"65.

2Landw.
3U.

Indus.,Bui.

128

(1911), pp. 88-90.

406

FARM

OF

NUTRITION

ANIMALS

responding ages. The experimentsdo not show, however,

these amounts
might not have been stillfurther reduced.
Table

97.

Protein

Digestible

"

Armsby

per
and

Pounds

iooo

Live

Weight.

that

"

Fries

x
likewise report the results of unpublished
Henry and Morrison
in which
growing fattening
experiments by Haecker
made
of digestible
steers
satisfactory
gains on amounts
protein

between

intermediate

those

recommended

Kellner

by

for beef

for

dairy breeds.
calves
the other
On
on
hand, Fingerling'sinvestigations
tion
4^-11 months
old, already cited in a discussion of the utilizalower level of
of feed protein (471),indicate that a much
proteinsupply may be adequate to support normal growth.
and

experiments2

The

menthaler

calves from

made

were

four

to

upon
seven

four

months

grade

or

old at

full-blood Simthe

beginning of

The

rations fed
early-maturingstrains.
added
in
consisted of a basis of hay or straw, or both, to which were
varying proportions wheat
gluten,peanut oil and starch with the
varied by varying
of salt. The proteinsupply was
amount
necessary
the amount
of wheat
gluten,the energy values of the rations being
kept as nearly identical as possibleby corresponding changes in the
intended
starch and oil. The experiments were
to test the necessity
for the relatively
of protein called for by the current
large amounts
standards
and also the influence of a deficient energy
supply upon the
of
gain
protein.
and
the trials,

As
the
live
the
1

belonging

from

to

rations,supplyingin
87, the medium
neighborhood of 1.2 pounds of protein per 1000
pounds
of
the demands
weight, were
clearlysufficient to meet
maximum
possibleprotein gain, since an increase of the
appears

Feeds

and

Table

Feeding, 15th Ed.,

p.

670.

Landw.

Vers.

Stat.,76 (1912), 1.

4o8

NUTRITION

Bull and

Emmett

OF

have

FARM

ANIMALS

compiledthe

results of

American
fifty

experiments on
fattening lambs, comprising 5127 animals,
and computed the protein and net energy content
of the rations
consumed.
ing
They divide the animals into four classes accordsubdivide
these classes into groups
to the live weight, and
of digestibleprotein consumed.
A
according to the amount
comparison of these groups shows in general that in each class,
with a liberal supply of feed energy,
the rate of growth
even
increased
the supply of protein increased up to a certain
as
fairlywell-defined
beyond which a further increase
amount,
of proteinhad in general little or no
effect.
The
authors
timate
esthe amounts
of digestible
to ensure
protein necessary
gains by fatteninglambs as follows :
satisfactory
"

Table

Estimated

Protein

Fattening

Lambs

hand, Just'sresults on lambs recorded

88 (471), like those of Fingerlingin calves,point to
lower proteinrequirement.

in Table

On

99.

Requirements

of

the other

488.
93

"

with

Experiments

(481 b),the

swine.

above
distinguished
in
rapidgrowth, especially

swine

is

by its very
The
pig is able to double his
young
and to nearly treble it in two
a week
reached

or

exceeded

of young

by

no

farm

much

is illustrated in Table

As

"

other
the

weight in
weeks, a

animal

with

farm

rupeds
quad-

earlier stages.

little more
rate

the

than

of

growth
ception
possibleex-

fowls.

ingly
growth implies,of course, a correspondlarge storage of protein,a conclusion fullyconfirmed
by
of
the investigations
of Ostertag and Zuntz, of Wilson, and
Sanford
and Lusk, cited in " 1 (463) which
showed
an
average
Such

of

rapid rate

Ills.Expt. Sta.,Bui. 166

(1914).

GROWTH

409

daily gain of from six to nine pounds of protein per thousand

live weight during the first sixteen days after birth.
Plainly,
pigs need a relatively
largesupply of proteinin their feed,
young
but unfortunately no attempts have thus far been reported to
determine
and

the minimum

of feed

especiallyby

older

proteinnecessary

at

different

normal
pigs, simply to ensure
growth. There are on record, however, a considerable number
rations supplying varying amounts
of experimentsin which
of
proteinhave been fed to fatteningpigs and the effects upon the
and upon the rate of increase in live weight
make-up of the carcass
observed.
These experiments have served to demonstrate
in a
the practicaladvantages of a liberal protein
strikingmanner
instances the minimum
quirement
supply and while in many
protein rehave been considerably exceeded,nevertheless,
may
ages

the results

as

whole

perhaps

are

less useful

no

as

guide in

practice.
It is
number

impossibleto include here

of experiments of this sort.
the student

Wolff

in 1876. l

United

States

A
as

may

be

even

an

For

enumeration

of the

large

of earlier investigations

summary

referred to the

considerable

compiled by

number
the

publishedby
summary
of earlier experiments in the

writer

gave

results of the

same

generalnature.
The later experiments upon
be divided into those
this subjectmay
directed more
of the influence upon
to the determination
specifically
and those in which
qualityand chemical composition of the carcass
the increase in live weight was
the principalcriterion.
489.

Effect

of insufficient

Strikingresults

protein upon

the

carcass

of

the

pigs.

"

in young
make-up of the carcass
pigs
have been reported by several investigatorsin experiments in
which
exclusive maize feeding was
compared with the use of
rations
mixed
more
supplying much
protein and ash. The
trials have been popularly spoken of as
Feeding for fat and
In realitythey are
for lean."
quate
a
study of the effect of inadeprotein (and ash?) supply in limiting growth. The
2
first taken
and soon
after by
subject was
up by Sanborn
Henry.3 In general it was found that in the pigsreceivingthe
as

to

"

1
2
3

Ernahrung der landwirtschaftlichen Nutztiere,

Agr'l College, Bui. 10, 14 and 19.
Wis. Expt. Sta.,Rpts 4, 5, 6, 17, 18, 19 and 21.
Mo.

pp.

465-496.

NUTRITION

410

protein (maize)

low

and

organs

in

OF

rations

some

FARM

the

ANIMALS

weights

of certain

cases

of

blood, of

individual

relativelyless than with comparable

high protein (mixed) rations,while on

animals
the

internal

muscles

were

receiving the
other
hand, the

depositsof adiposetissue appearednotablygreater

in the maize-

fed animals.
these

Since

made

were
investigations

fact that

the

mixed

proteins of

it has become
maize

are

nized
well-recoginadequate to support
a

rapid growth (783) and the results reached are to be regarded

being to a considerable
degree the expression of this qualitative

as

deficiency.
As

the

regards

remarked

that in

have

made

purposely made

it clear that

supply

exclusive

experiments it

the difference

instances

most

regards protein was

as

of the

quantitativeaspect

is to

the

between

be

rations

While

the experiments
large.
afford an adefails
quate
to
feeding

maize

of

protein for growing pigs, it does not follow that as

contained
in the contrasting rations
largequantitiesof protein as were
In the later Wisconsin
were
as
experiments especially,
necessary.
the writer has pointed out,1the gain in live weight was
often little
greater

the

on

high proteinthan

Furthermore,
as

feeding,and

while

not

the animals

to

the

weights

in

one

instance

experiments were
did

the low

proteinrations

and

times
some-

less.

even

and

on

afford

not

data

of the various
at

of the
for

compared

were

least the
of

nature

computing

organs
carcasses

more

at

or

the

were

less

close of the

analyzed,the

comparative slaughter
the actual

amount

tensively
ex-

of

tests

protein

to
gained. Moreover, the results upon the carcasses
analyzed 2 seem
indicate that the rations affected the adipose tissue as to its distribution

to
to

rather than as to its total amount.

through the carcass
Finally,the strikingresults as to general thrift,and especiallyas
the growth and strengthof the bones, are probably to be attributed
differences in the ash supply (496) quite as much
as to differences
the proteinsupply.
,

in

the

On

from
valuable
whole, while these investigations
are
the standpoint of practiceas a demonstration
of the ill effects
of a deficient amount
be said
or
quality of protein,it cannot
that this class of experiments affords very definite information
to the actual proteinrequirements of pigs.
as
1

U.

Iowa

S. Dept. Agr., Bur.

Anim.
Indus., Bui. 108 (1908), p. 74.
U. S. Dept. Agr., Bur.
Expt. Sta., Bui. 48 (1900), pp. 373-451;
Indus., Bui. 108, p. 75,.

Anim.

GROWTH

"

490.

Fattening experiments

411

with

Of the

pigs.

"

recent

more

experiments upon the influence of the protein supply upon the

rate of gain of immature
fatteningpigs,four series made by the
l
Halle Experiment Station
at Lauchstadt
and
series of coa
operative
of the German
experiments at a number
experiment
stations under Kellner's generaldirection are of specialinterest.
While
these relate primarily to fattening,the comparative
results with
rations of equal energy
content
should furnish
indications as to the sufficiency
some
of the protein supply,
since the rate
of increase of protein tissue can
hardly be supposed
to differ materially
from that in simple growth without

fattening.
In the Halle

the gradual
to note
experiments it is interesting
loweringof the average protein supply from the high level of
the

first series.
results

content

is

as

The

may

final series
be

obtained

pounds
follows,although the earlier
higherfigures.
per

Table

100.

Estimated

"

from

Protein

trials

show

rations

the different

at

1000

to

seems

whose

weights

seem

Requirements

that

protein

of the animal

to indicate

of

factory
satis-

Fattening

At

weight 77-100 lb
weight 1 10-165 lb
At weight 165-220 lb
At weight over
lb
220

4.0

At

what
some-

Pigs

lb.

3.0 lb.

2.0-2.5
2.0

lb.

lb.

In

initiated by the German

1906 cooperativeexperiments were
of German
AgriculturalCouncil at a number
experiment
stations upon
'the value of potatoes as feed for fatteningpigs
and
the protein supply necessary
especiallyupon
for their
most
complete utilization. The results of experiments upon
this point at eleven stations,
a total of 184 animals, have
upon
been discussed by Kellner 2 under whose
general direction the
3
work
done.
was
According to Kellner
fatteningpigs
young
should receive per 1000
live weight the followingamounts
of protein:
"

Landw.

Jahrb.,28 (1899),947

; 31

(1902),916

; 36

(1907),679

(1910),179.
2

Ber.

Die

Deut.

Landw.

Ernahrung

Rat, Heft

der landw.

(1908).

Nutztiere,5th Ed.,

p.

488.

39, Ergzbd. Ill

OF

NUTRITION

412

Table

The

ioi.

"

Kellner's

of the

outcome

FARM

Standards

ANIMALS

for

Pigs

Fattening

cooperativeexperiments tended

to

firm
con-

that any considerable

standards,indicating
departure
fail
them
will
the
of
from
to meet
requirements
rapid growth in
the best strains of animals
the largest returns.
to secure
or
series perhaps points
On the other hand, the fourth Lauchstadt
in the direction of lower
for the more
standards, especially
animals.
recommends
For
Kellner
mature
breeding animals
smaller amounts
of protein,viz. :
somewhat
these

"

Table

Dietrich

102.

"

recommends

although he too
animals.
to

up

the

Kellner's

recommends

His

figuresfor

age

of about

to 3.3 per thousand

his

are
figures

from
491.

Standards

months,

General

2.0

Ills. Expt.

the

6 months

to

conclusions.

paragraphs that

followed

at 7 to 8 months.

point to

Growth

at

by
For

of age,

Pigs

of

notably largeramounts
less for breedingthan
fatteningpigs are 6.0 to

5.0 to 5.5 up

that

for

of
for

protein,
fattening

7.0 per

1000

tion
gradual reduc-

breedinganimals
ishing
graduallydimin-

maturity.

It is apparent
from
evidence
regarding the
"

Sta.,Circulars 126,

133

and

153.

the

going
fore-

proteinre-

GROWTH

less conflicting.
or
fragmentary and more
side are
investigationslike Fingerling's
sheep (486,487) which appear to show

quirements for growth

the

On

cattle and

on

one

Just'son

413

is

for the animal

that it is possible

to

support what

seems

normal

growth upon a supply of proteinlittlegreater than the

maintenance
actuallystored.
requirement plus the amount
tions
On the other side stand the results of experimentsand observarate

of

the

upon

fatteningof

immature

animals,in which

rations

approximatelyequal as to their content of net energy,

and therefore presumablyequallyeffective for simplefattening,
have produced a greater increase in weight when they contained
much
more
relatively
proteinthan the results of the other class
indicate to be necessary.
Further conof experimentswould
sideration
at

least

of this

that the
For

classes of

two

one

thing, the

while

experimentsare hardlycomparable.
experimentsin which a relatively
high

seemed

fattening experiments.
measured
The effect of the feed was
by the gain in
uncertain
criterion,
weight, which itself is a somewhat

protein supply
live

apparent conflict of evidence,however, shows

considerable

of fat rather
on

than

the

advantageous

share of this increase

of

protein.

all

were

was

due

to

storage

and
Fingerling's

to
contrary, relate distinctly

ments,
Just'sexperigrowth and the
of proteintissue

comparisonsare based on the actual amounts

produced,although it must be admitted that any experimental
would probably tend to make
the excretion of nitrogen
errors
appear too low, and therefore the gain of proteintoo high.
Another

important

difference

experimentslies in the
experimentsthey

nature

between

the

of the rations.

two

In

classes of
olism
the metab-

composed largelyof commercially

pure nutrients such as starch,oil,etc., with only the amount
of roughage necessary
to supply bulk, and
in particular,
the
variations in the proteinsupply were
effected by changes in
of commerciallypure wheat gluten. In the fattenthe amount
ing
experiments,on the contrary, the higher proteincontent of
the rations was
obtained by the use
of the ordinary proteinrich feeding stuffs. What
these experimentsreallyshow is that
a
largerproportionof these feeding stuffs was
advantageous,
but it does not necessarily
follow that this advantage was
due
to the added
protein. For one thing such a modification of
the rations must
have affected the ash supplyto a certain extent.
were

OF

NUTRITION

414

FARM

ANIMALS

stances
however, the influence of those accessory subparticular,
gations
(498,499)which recent investior
growth substances
have shown
to play such an
ing
importantpart in conditionIt seems
not
growth is to be considered.
impossiblethat
in some
such way
have
feeds may
a
stimulating
high-protein
effect upon
the capacityfor growth quite independentlyof their
On the other hand, however, any such stimulating
proteincontent.
effect upon
growth would be absent in experiments made
with pure nutrients added
to a basal ration of hay or straw, and
normal
rate of increase seems
to have been maintained.
yet a fairly
On the whole, one
can
hardly fail of the impressionthat the
estimated
requirements for protein as such in growth have been overutilize its proteinsupply
and that the organism may
In

"

"

made

be

actual
has

than

considerablysmaller

would

feeding standards

economically than the current

indicate ; in other words, that the
more

proteinsupply may
been

supposed

before

growth. Until this impression

is confirmed
extensive investigation,
however, it apby more
pears
to the accepted
the safer course
to adhere
provisionally
standards,and the proteinrequirements for growth as estimated
in Table IV b of the Appendix are based upon
those formulated
by Kellner.
it becomes

Ash
Growth

492.

involves

like the mature

this the formation

requirements

storage

of ash.

growing animal,
for the puringredients
poses
The

"

in

requires mineral
Chapter V (268-272),but

of

new

one,

enumerated

tissue and

the storage of ash

involves

in

limitingfactor

in addition

of
especially

ingredientswhich

must

to

the skeleton
be

derived

clearlyby the data recorded in

" i (458) regardingthe composition of the increase,its ash
1.42 per
case, ranging from
content, aside from one exceptional
the

from

cent

feed.

to 6.18 per

493.

Rate

of

This

cent.

storage

are

elements
are

in

growth.

"

Data

regardingthe

rate

in

growth are not very numerous

largelyconfined to experimentson the two important
tions
and
calcium
phosphorus. The principalinvestiga4
those of Soxhlet,1
Neumann,2 Lehmann,3 and Weiske

of storage of mineral
and

is shown

ier Ber. Versuch-Station

Landw.

Vers.

Stat., 1

elements

Wien, pp.
(1859), 68.

101-155.

2
4

Jour. Landw.,
Jour. Landw.,

41
21

(1893), 343(1873), 130.

416

NUTRITION

fact shown

in

of ash

may
direction.

in the

same

of

fluctuations

to

nutrients

organic

draw

to

Total

494.

total
may

of

weights

during growth.

ash

each

of
ash

the

old

in the
would

be

Table

104.

as

"

from

entire

of the

those

much

serve
largerre-

bodies

follows

Total

at
:

notion

bodies

in
to

the

yses
Gilbert's anal-

farm

of

of

during growth

and

Lawes

old be assumed

animals

them.

the

Thus

if

be for cattle 400

sheep 50 kilograms(no pounds)

pig 50 kilograms,then, applying the
store sheep and store pig respectively,
for

the half-fat ox,

of mineral
amounts

daily retention

average

to

Some

"

ingredientcontained

analysesof
the total

points

far less sensitive

assimilated

elements

by computing

six months'

animal

mature

organism appears
its daily supply of ash than
because
it has relativelya

weight at one year

kilograms(880pounds) and
for

less storage

or

The

the live

and

in the

even

of mineral

the

more

upon.

secured

of

occur

retention

amounts

be

ANIMALS

(435) that

IX

Chapter

elements

FARM

OF

elements
head

per

birth in the

in the

bodies

and

(includingthe stock
of the sheep and
case

the
tained
con-

pig)

"

Retention

of

Ash

Ingredients

During

Growth

GROWTH

417

In order to compare
the figuresthus obtained with the results
recorded
in the previous paragraph for the retention
during

short

periods,it is necessary to eliminate the influence of varying

The
retention
weight by computing the results per 1000.
of calcium and phosphorus as thus computed agrees very well
with that found in the balance experiments with the exception
of Forbes' high result for phosphorus with the pig. On
the
other hand the computed retention of the alkalies is strikingly
less,a fact for which no obvious reason
appears.
Table

105.

Average

"

Daily

Retention

per

Live

iooo

Weight

Pig
6

Potassium

0.018

Sodium

0.009

Calcium

0.084

Magnesium

0.004

Phosphorus
Chlorin

On

O.051
0.006

0.003

the whole it appears that our knowledge of the ash

of growing animals, that is,of the actual amounts

up in normal
495.

ing
DurFirst
Months

growth, is

Availabilityof
formulate

ments
requirestored

quitefragmentaryand unsatisfactory.
ash

ingredients

of

feed.

"

If it is difficult

from

existingdata any trustworthyestimates

of the ash requirements of growing animals, it is even
more
to

difficult to

make

of

amount

the feed in
extent
are

to

which

statements

the results recorded

in the

to formulate
trustworthy,it is possible

Thus, Weiske's
to

definite

regardingthe total
be suppliedin
element which must
any particular
order to meet
those requirements,although to the
any

35

mgrs.

of

results

previousparagraphs
the minimum
supply.

sheep show a retention of from

phosphorus per kilogram of live weight.
on

20

If

these

figuresrepresent the normal requirements,it is evident

that a ration containingless than this amount
would
not supply
enough phosphorus for normal
growth. What
surplusabove
is necessary
in the feed would depend on the prothis amount
portion
of the feed
2

phosphorus which

is

capableof

solution in

4i8
the

elimination

the

on

element

sufficient

amply

suppliedin

still more

and

digestivetract,

one

ration

ANIMALS

FARM

OF

NUTRITION

the effect of other

upon

of

phosphorus.

to

meet

might

be

the

An

of this

amount

demand

normal

quite inadequate

ments
ele-

when

in another

of

different character.

path of excretion for

ash elements
(164, 199), a computation of the digestibility
some
in the ordinary sense, by comparing
of these elements
in feeds and feces,gives an
the amounts
entirelyfalse idea of
shown
in Chapter IX
their availability.
Moreover, it was
lost
elements
are
(429-433) that the rate at which mineral
the qualitative
from
the body depends to a large degree upon
compositionof the ash of the feed, variations in the supply of
sometimes
element
affectingmateriallythe gain or loss of
one
it was
another.
In particular
pointed out that the proportion
intestines

the

Since

basic

and

of acid

are

the

and

elements

normal

to

less

degree

ratio

the

of

have
striking effects of this sort.
potassium to sodium
may
dition
For
example, in Weiser's experiments on pigs (493),the ad-

of 5 grams

of

calcium

carbonate

to

ration

of

iooo

of maize
not
only changed a loss of calcium into a gain
grams
the phosphorus balance,
effect on
but also produced the same

phosphorus supply which was previouslyinsufficient

able to support a material gain.
the body was
maintain
that

so

to

Table

io6.

Ash

"

Balance

or

Swine

with

and

Calcium

without

Carbonate

Phosphorus

Calcium

Maize

alone

In feed
In feces

1.

In urine

1.0686
1.2682

and

1298

2.2570

0.8134

0.1384

Balance

Maize

2.6731

0.1996

0.3973

1.2682

3.0704

3.0704

CaCO%

In feed
In feces

In urine
.

2.8167

2.1950

Balance
2.1950

1.2602

1.6960

0.0766

0.2714

0.8582

0-8493

2.1950

2.8167

2.8167

GROWTH

It has

419

been

maintained, principallyon the basis of Soxhlet's

of the ash
experiments on calves (493), that the availability
ingredientsof milk, and particularlyof its calcium and phosphorus,
is especiallyhigh. The
in
retained
the
percentages

body

of the five trials were

the average

on

Total

"

ash

53-o

Potassium

20.7

Sodium

29.1

Calcium

97.0

Magnesium

30-S

Phosphorus

72.5

Chlorin

Neumann's

3.8

experimentswith

that the

however, render it evident

of calcium

phosphorus

and

in the

elements
elements

It

body.

somewhat

one-half

great

as

milk

skim

experiments on

milk

Calcium

used

Phosphorus

animals

therefore excreted

and

the calcium

on

mixed

the excess,
in the feces.

and

Weiske's

rations showed
and

milk

Period

calcium

in Neumann's
is that

the

44-5
42.7

the

be
1

Landw.

%
%

%
41.8%
44-1

phosphorus in

the urine

experiments (493) with older calves

percentage

of
availability

the

fullyas great as that observed for

experiments,and here too the natural
demand

for these

elements

rather than any lower availability

per se, is the
assimilation.
It is well established that the
may

Period

obviously required less of these elements

and

Lehmann's

in Neumann's

was

47-8%
.Si.i%

older

for these

alone.

Period

The

retention

scarcely more
experiments,viz.,in the

in Soxhlet's

as

(493),

hardly be supposed that these

can

experiments, yet the percentage retention

than

high

large demand

less assimilable in the skim

are

of the

cause

the

was

older calves

in

cause

the

phorus
phosskim
clusion
con-

body,

of the less

phates
inorganicphos1
quite completely assimilated,and Fingerling
Vers. Stat.,79-80

(1913),847;

86

(1915),75.

NUTRITION

420

has

shown

the

OF

FARM

ANIMALS

thing to be true
phosphorus compounds, as well as of
feeding stuffs,while in case
of

should

two

which

496.

doubt

some

contain

elements

the
of

would
of

Effects

three

times

normally be

deficiency
make

regarding the

of

the
stored

ash.

any definite
ash supply of

in the

ability
avail-

serious

fact that such

the feed

it

seems

quantitativestatements
growing animals there is
an
as

insufficient

supply.

already indicated,

both

directlyand
generallyconnotes
deficiency

consequences

These

body.
while

But

"

a
particular,
deficiencyof calcium,

have

centrated
con-

quantitiesof mineral

necessary
evidence
of the evil effects of

abundant

may
the

of

phosphorus
roughages * an

observed.
was
per cent
Kellner's conclusion
that

on

or

scarcelypossibleto

In

varietyof organic

approximately 50

throw

facts

of

same

of

account

on
an

acid ash

(431).
2

Kellner

dogs

and

cites

experimentsby.Roloffand by Voit, in which

in calcium

pigs receivingfeed poor

showed

and

deficient

young

growth

the skeleton

developed severe
pathologicalsymptoms,
showing
3
in
lected
ash
Forbes
has colingredients (Rachitis)
deficiency
of which
of experiments on this subjectin some
a largenumber
marked
effects on
the composition of the bones
observed
while
were

notable

in others

these effects

not

were

distinct.

very

In

still more

recent

experiments by Weiser
pigs, a diet deficient in calcium
upon
the growth and produced a skeleton containing an excess
and organic matter
and deficient in ash.
water
Contrary to the

(428) the

of Aron

deficient in calcium

bone

and

ash

of
sults
re-

calcium-poor rations was

of alkalies,
excess
especially

the

on

contained

stricted
re-

an

sodium.

Of farm

animals, pigs are

partlybecause

their

growth

they often receive

are

apt

to

be

low

rations

results in the
and

Burnett
1
2
3
4
6

has

Biochem.

is

grain rations which

that
(431). Henry 5 has shown

almost

in calcium

supplementing such

likelyto suffer in this way,

cause
relativelyrapid and partlybe-

most

exclusive

with

production of

confirmed

these

calcium
heavier
results.

phosphate
and

or

bonate
car-

stronger bones,

Hart

Ztschr., 37 (1911), 266.

Ernahrung der landw. Nutztiere, 6th Ed., p. 177.
Ohio Expt. Sta., Tech. Bui. 5, pp. 384-390.
Biochem.
Ztschr., 66 (1914), 95.
Wis. Expt. Sta., 6th Rpt., pp. 6-41 ; Bui. 25.
Neb. Expt. Sta.,Buls. 94 and 107 and 23d Rpt,

and

McCol-

GROWTH

lum

that confined

found

421

pigs on

ration of maize

alone and

failed to grow, while the addition of an

of salts enabled nearly normal
growth to be

drinking distilled water

artificialmixture
made.

sheep a deficiencyof calcium is not

roughages are usuallyrich in this

cattle and

With

feared,since
and roots, however, are rather low
Straw
those
certain by-product feeds,especially
meal, distiller's grains,etc., which have
to be

with

element.

and

so

are

gluten feed and

been subject to extraction

like

water.

of

Forms

497.

in calcium

usually

phosphorus.

questionis
inorganicphosphorus
discussed

much

"

organic and
compounds. It was stated in Chapter V (258),that the animal
body is apparently able to synthesize its organic phosphorus
2
has
given
compounds from inorganicphosphorus. Forbes
His
a
very complete review of the literature of this subject.
general conclusion is that it has not been proven that a supply
of organic phosphorus is essential,
although he regards the
for
all the purposes
proof that inorganicphosphorus can serve
needs phosphorus as being incomplete. As
which any animal
of the two, the facts alreadynoted
regards the relative efficiency
in Chapter IX
garding
(437,438) and in the following paragraphs,rethe importance of accessory
substances,in particular
the so-called growth substances,in nutrition strongly suggest
has
of organic phosphorus which
that the apparent superiority
that of the relative value

been

observed

in

of such

presence

of

compounds

and

the

accompanying

substances

the latter

to

not

been

have

experimentsmay

some

as

due

to

the

phorus
organicphos-

such.

Accessorysubstances
498.

Relation

of

fats

to

growth.

Chapter V (265),in consideringthe

that it had
minimum

apparently been
of

amount
"

lipoids)was

fat

(or

at

necessary

however, have led

shown

to

It

"

functions

in

of the nutrients,

that the presence of a certain

stances
subleast of ether-soluble
for

growth.

different

Later

gations,
investi-

of
interpretation

Jour. Biol. Chem., 19 (1914),373.

Expt. Sta.,Tech. Bui. 5, pp. 318

Ohio

mentioned

was

to

365-

these

OF

NUTRITION

422

ANIMALS

FARM

the

lated
technique of experimentation with isohas been
of Rohmann,
nutrients
developed by the work
and
Mendel
and
McCollum, Osborne
others, it has become
evident
that it is not the lipoidsas such but some
substance
or
earlier results.

As

associated

substances

for

while

all fats

McCollum
of

mixtures
for

it has

and

Osborne

considerable

time,

reachingperhaps "
cessation of growth
the

but

fat,olive oil,almond

decline in

of

animal

an

by

effect

have

75

or

no

tinued
con-

means

growth.

on

found

fat may

no

about

added

the

to

liver oil

hand, purifiedbutter-fat,cod
the ration

to

that

normally

grow

days,

100

fed

rats

and

after

or less abrupt
weight,there is a more
decline
in
stantially
weight. Subby a speedy
when
certain forms of fat (lard,
beef

ensues

oil)are

that

favorable

and Mendel

after

followed

result

same

essential

are

shown

purifiednutrients containing
of the mature

to

been

capableof exertingthis

are

Both

which

hand, growth has been maintained,

least,on a practicallyfat-free diet,

at

other hand

the

on

them

one

time

considerable

the

On

growth.

with

which

weight this decline

has

is

or

ration,but if,on
certain

ceased

other

to

fats be added
and

grow

and

promptly stopped

the other

begun

to

practically

normal

These results indicate the existence of two

growth resumed.
of fats,one
of which
aids growth while the other does not.
groups
Evidently,therefore,the growth supportingproperty does not reside
in the

glyceridsthemselves

499.

substances.

Growth

McCollum

in

rejectsFunk's

specific vitamins

numerous

soluble

to

is associated

A,

water-soluble
with
fats

abundant

fats.
thus

other

The

the

"

in water

leaves

than

and

other,called
ciated
asso-

from

all table
vegebut
sufficie
insmall
in
to

appears

be

relatively

of

these

McCollum

Jour. Biol. Chem., 23 (1915), 181 and

and 361.
Jour. Biol. Chem., 19 (1914), 373.

(1916),333

is absent

It is present

grainsbut

calls fat-

apparentlynever

plants.
specific
growth
growth is,however, apparent

factors

Hart

and

he

the

fats,while

fat-soluble A

in the

influence

by

tions,
investiga-

existence
hypothesis
and
distinguishedonly two
substances) both of which are
of

certain

far examined.

in the

markedly

with

B, is soluble

amounts

of later

essential

That

the basis

(orclasses of
which
growth. One, lipoid-soluble,

substances

growth

substances.

accompanying

some

On

"

"

of

41

but

231

who

; 25

found

(1916), 105

substances
from

recent

may
periments
ex-

that the freedom

; Amer.

Jour. Physiol.,

XII

CHAPTER

PRODUCTION1

MEAT

"

flesh of

the

the

on

into

"

By

Meat

of

"

animal

an

and

the internal

greater

adiposetissue (" fat ") and

or

fat,and
501.

is

skeleton

other

offal

cally
separablemechaniof

which,
complex composition.

of connective

of fat has

less accumulation

the

and

general

lean meat, both

of somewhat

tissue in which

place and

taken

is

of

reserve

its characteristic

Proportion

of lean meat

of fat and

in

lean

to fat tissue in the

carcass.

is

carcass

body to a minimum
production of large amounts

while

proteins.
The proportion
able,
naturallyquitevari"

ing
chieflyon the feedreducing the store of fat
the
heavy feedingmay cause

the age
upon
of the animal, insufficient nutrition
in the

the

are
ingredients

depending somewhat

of it.

but

Thus

and

Lawes

proportion of fat in the carcasses

analyzed by them (97) to vary from 15.3
Jordan observed a range of 18.80 to 24.62 per

found

in

material.
non-nitrogenous, energy-yielding
in the narrower
The
lean meat, or
meat
sense
(86),
tissue along with more
less
or
primarily of muscular

a
essentially

consists

hide

organs,

generalsense

are
latter,
adiposetissue (94) consists

The

is understood

from
distinguished

as

in this

Meat

Production
"

meat

especiallythe

but

hand

one

the other.

on

Nature

Definitions.

500.
way

i.

the

of the
to
cent

Gilbert
animals

ten

48.3

per

in steers

cent.

2-2

of 10.39 and
Tschirwinsky reports the extremes
of 1.31
found
a minimum
40.92 per cent in pigs,while Wilson
2
in new-born
pigs. Atwater
gives the followingas
per cent
the average
fatness :
compositionof a side of beef of medium
years

old.

"

The

discussions

in this

chapter follow, to

by the writer in U. S. Dept. Agr., Bur. Anim.

2 U.
S. Dept. Agr., Office Expt. Stas., Bui.
424

considerable

extent, those

Indus., Bui. 108 (1908).

21

(1895),p.

35.

presented

PRODUCTION

MEAT

Table

107.

"

Average

425

Composition
Medium

of

Side

Beef

of

of

Fatness

Water

54-77

Protein

17.20

Fat

27.07

Ash

0.96

100.00

Lean

of meat,
by the

cuts

is indicated
l

Emmett

dry

cent

per

samples

been

removed.

in the

fresh

The

matter.

may

contain

average

The

of beef round

minimum

substance,or
of the

much

less fat than

Grindley and

Thus

foregoingstatement.

analyzedseven

visible fat had

3.19

however,

from

figurefor fat

12.29

per

analyses

seven

which

the
was

cent

of the

was

as

lows
fol-

"

Table

108.

"

Average

Composition

of

Visible

Fat

with

Seven

Samples

of

Beef

Removed

In

Water

Round

Water-free
Substance

Ash
.

Protein
Extractives
Fat
.

which
he
carefullyprepared lean meat
used as representing
substantially
proteinfeed,and which had
freed from all visible fat,0.91 per cent
been most
painstakingly
of ether extract in the fresh substance,equal to 3.77 per cent of
the dry matter.
The term
meat
commonly suggests to the mind the muscular
with
almost
tissue of the animal, and has become
synonymous
Voit

found

in

the

Ills. Expt. Sta., Bui. 162.

426

OF

NUTRITION

ANIMALS

FARM

proteindiet. It is evident,however, that the commercial

involve
the productionof considerably
growing of meat
may
fat than proteinand that,in so far as this fat is actually
more
is far from
being the distinctively
consumed, meat
protein
a

which

food
"

nutritive

of which

valuable

fat,although a

so-called

meat

it is

the distinctive ingredient

are

produced and eaten,

nutrient,is

to

certain

while
sidiary
sub-

extent

accidental.
in

Corresponding

general
meat, viz.,
distinct
tissue and adipose tissue,two
physiological
involved in meat
are
production,viz.,growth and

Processes

502.

proteinsof

the

for the sake

and

the

"

that

equally true

the

Thus

side of beef,calculated
of the average
in the
On the other hand, however, it
is about
i : 3.5.

ratio

usual manner,
is

ordinarilyconsidered.

it is.

the

to

way
muscular
processes

involved.

main

two

"

of

constituents

commercial

fattening.
Growth.
unfit

to

the

not

The

"

serve

utilize

from

feed.
or

less

be concluded
a

whole

birth

at

be

the

in the

marked

is

usually regarded as
this
were
Moreover, even

food.

human

it would

highestdegree
assimilative

uneconomic

powers
tissue

of

the

(meat)
production of body
involves
Consequently the production of meat
This may,
for specialreasons,
growth in all cases.
early,as in the productionof lamb or veal, but as

animal

young

more

as

case

fail to

to

animal

for

the world's

the

commercial

meat

supply

is derived

from

approaching maturity. This growth of animals

of an
from birth to approximate maturity consists essentially
increase in the protein tissues (457), the rate of which
is essentially
animals

at least

determined
animal

and

can

at

by
most

the
be

of
individuality
stimulated by an
slightly

nature

but

and

the

creased
in-

proteinsupply (403,484).
Fattening. Fattening,on the contrary, is a process which,
is largelyunder
the control of the
in a given animal
at least,
the quantity of feed
feeder.
it is dependent on
Substantially
"

requirementsfor
and growth, and there is lackingany definite proof
maintenance
that the actual storage of energy in the form of gain for a given
of excess
feed is seriouslyaffected either by the age or
amount
consumed

the

by

the

animal

in

excess

of

the

individualityof the animal.

Fattening, therefore,may
the greater demand
take place at any age, although of course

PRODUCTION

MEAT

for

for material

growth

in the young

427
animal

tends to reduce

the

proportion of the feed available for fattening.

The
prime object of fattening(446) is an improvement in
the quality of the meat
by the depositionof fat between the
fibers of the meat, and to some
extent
tractives
by increasingthe exof the meat
itself. The large depositsof fat about
the internal organs
to

are

certain

and

extent

the skin

under

The

waste.

incidental

are

subcutaneous

to

this and

fat affords

quality of the meat, and of course

is of some
the adipose tissue of the carcass
value,but these fat
ity
depositslargelyrepresent the pricepaid for the improved qualof the meat
It is not impossiblethat the traditions
proper.
of the market
the process of fatteningto be pushed
cause
may
a

index

convenient

beyond

what

to

the

is necessary.

improvement in quality may be, and to a considerable

extent
is, secured by a comparatively short period of high
feedingafter growth has been nearly completed. It is obvious,
the prohowever, that no sharp line can be drawn between
cesses
of growth and fattening.A calf or yearling may
be
fattened while growing,and a two-year-oldsteer will continue
The
to grow
to some
two
extent
while being fattened.
cesses
proThis

shade

into each

other

and

considerations

economic

will

decide whether

by

ously
or less simultanethey shall be carried on more
singleproducer or at different times by two different

individuals.

brief,meat
production may be defined as a combination
of growth and fattening,
which may
be either simultaneous
or
successive,but the production of protein tissue is the primary
of fat,although adding
objectin view, while the accumulation
In

to

the nutritive value

or

less a

is

to

of the meat,
palatability

is more

of the present

chapter
of growth and
consider the applicationof the principles

of food
503.

Factors

mechanism

then, as

precedingchaptersto

this branch

of

meat
meat

production.

From

"

producing animal
of which

the

may

the

economic

be looked

material

upon

contained

by
uct
feedingstuffs is converted into the finished prodhuman
production,
consumption. Regarding meat
and quality of
a
manufacturing process, the amount

in the various
for

in the two

purpose

production.

point of view, the

a

The

secondarymatter.

discussed
fattening

as

and to the

means

raw

428

NUTRITION

three factors

is plainly
dependent upon

productionobtained

the

ANIMALS

FARM

OF

and
of the mechanism
the efficiency
first,
; second, the amount
material
supplied; third, the conditions
quality of the raw

under

"

which

The

2.

the

Of

the mechanism

Animal

as

operated.

Factor

Production

Meat

in

the animal

just mentioned,

factors

three

is

may

fairly

of the
prime importance. The success
to
the capacity of his animals
feeder depends primarilyupon
of raw
into a
materials
convert
profitablylarge amounts
finished product of high quality.

be said

be the

to

of

one

Early maturity
504.

the

upon

of

Definition

maturity.

importance

Strictlyspeaking,a
its growth
one
i.e.,
reached

in meat

more

or

less distinct

is one

animal

the increase

in which

rightlylaid
though
alproduction,
is

which
of

has

senses.

completed

protein tissue

has

of two animals
In this sense, that one
limit first is the earlier maturing.
this natural
limit.

its natural
reaches

which

in two

mature

"

stress

earlymaturity

of

is used

the term

Much

"

limit of size,
substantiallythe same
this conception of early maturity is, of course,
synonymous
with a greater absolute rate of proteingrowth (460),while if the
latter be expressedrelativelyto the weight of the animal, as
of size.
in previous pages, the same
thing is true regardless
The
term
early maturity,however, is used also in a quite
of the animal rather
to the conformation
different sense, referring
has
than to completed growth. Thus, if a steer at 22 months
and
reached sufficient size to
attained the typical beef form
viously,
Obof the market, he is said to be mature.
the demands
meet
With

this does

but

This

commmercial

not

rather

size of

and

carcass

to

reach

impliesa greater rate

he

of

completed his growth,

able
growth to furnish market-

has

sufficient

conception

note, however, that

certain

that

mean

he has made

simply that
meat.

reach

which

animals

maturity,

in

It is
physiological.
element.
involves
a
physiological

than
it
as

well

this at

of

words,
important

other

as
an

conformation

certain

early age

growth,whether

almost

measured

is
to

manded,
is de-

necessarily

physiologi-

PRODUCTION

MEAT

429

cally by increase of protein tissue or practically

by gain in
weight.
the term maturity is used, however, the
In whichever
sense
reduces itself to the question of rate of growth. The
matter
ing
greater the initialimpulse to growth, the sooner, other thingsbeequal,will the animal complete his growth, while if the rate
of growth can
be made
tion
rapid the desired accumulasufficiently
be reached before
of meat, and consequent weight, may
maturity. In other words, the rate of growth may
physiological
be looked upon as expressingthe capacityof the machine, since,
determined
stated in Chapter XI, it is substantially
as
was
factors and is apparently littleaffected by the feed
by biological
supply,provided only that the latter is adequate.
505.

Economic

significance.
"

material

that there is any

between
economy

There

no

seems

difference

reason

to

logical
regardsphysiorapid growth and slow growth ; that
that the storingup of certain
to suppose
is,there is no reason
of protein and energy
in the body of an animal in one
amounts
month
requiresany greater or any less total feed supply, in
suppose

addition
the

same

to the maintenance
amounts

feedingmay
(722). In

in the

diminish

the

as

the

storingup of
months'
two
time, except as heavy
of the ration
percentage digestibility
requirement,than

that if a gain of one

words, it may be assumed
tion
pound in live weight contains 2500 Calories of energy, the rathe mainof net energy
above
tenance
must
supply that amount
the
requirement within the time required to make
gain, whether that time be one day or three.
the economic
From
point of view, however, there is a very
important difference which explainsthe stress laid upon early
It is plain that, other
maturity in meat-producing animals.
thingsbeing equal,the animal which inherits the greater initial
impulse to growth, and in which that impulse dies out the more
slowly,will reach either physiological
maturity or a given size
and

other

weight sooner

It makes

than

the

one

in which

that

impulse is

less.

however, to the producer

difference,
of beef cattle,
for example, whether
a calf weighing 100
pounds
at birth has the capacity to reach
a
pounds at
weigh of 1200
he requiresthree years to do it. This
two years old,or whether
is not, however, because there is any material difference in the
the
of feed which the animal requiresto manufacture
amount
a

very

material

NUTRITION

430

pounds

noo

in

comes

of

from

expenditurefor

well

the

has

the

ANIMALS

difference

regardsfeed cost
maintenance, since each pound
as

pounds, must be maintained

until maturity. The
animal, then,
of growth and
which
in
matures

originalioo
on

higher rate

costs

two

the

as

it is laid

the time

which

FARM

The

of increase.

the

gain,as

OF

owner

notably less expenditurefor

years
the
than

one

maturing

savingin

cost

of attendance

in three

feed

years, to say nothing of the

in interest on the investment.

and

Age
Influence

506.

of

cost

on

gain is made more

as
compared with
younger
and in
in growth proper

the
the

undisputed
cheaply by the

more

older animal.
commercial

an

This is true both

fatteningof partly

animals.1

mature

On

It is

"

rapidlyand

that

fact

production.

hand, it was

the other

shown

in

(472-476)that
capacityof the young

Chapter XI

experimentalevidence that the

animal
for making a more
rapid gain is due to any greater
in the conversion
of surplus digestible
physiological
economy
there is

no

(720)
of the young
that the digestive
animal is not materially
power
animal.
As regards protein,
different from
that of the mature
that the loss of nitrogenousmaterial in the
the indications are
of feed proteininto body protein is not
actual conversion
dinarily
ormaterial

tissue,while

into

great and

animal, while

no

greater in the younger

507.

Causes

it

regardsenergy

as

also

established

been

greater in the old than

was

in the young
that the probabilities

shown

that its utilization

in favor of the view

are

than

is

it has

is less rather

in the older animal.

than

of greater economy.

"

More

or

less confusion

of

thought has resulted from this apparent conflict of evidence,

son
while feedingexperiments like those cited by Henry and Morrihave

been

the greater
This

made

the

and
digestive

confusion

has

arisen

of unwarranted

basis

assimilative
to

powers

inferences

largedegree through failure

Compare
568-572.

Henry

superiorityof the
and

Morrison, Feeds

and

younger

to

animals.

of young

and
commercial
distinguishbetween
physiological
of the
and it is important to secure
a clear conception
of the commercial

as

to

economy
elements

animal.

Feeding, 15th Ed.,

pp.

431-434,

512,

NUTRITION

432

Table

109.

"

Table

Relation

iio.

FARM

OF

of

"

Feed

Weight

ANIMALS

of

Pigs

Consumption

to

by

Lambs

Digestible
Eaten
Live

Organic
Matter
Live
Kgs.
50
Weight

per

Weight
In

Consumed

Feed

In

Proportion
Weight

to

Kgs.

Period

Period

II
.

Period

III

Period

IV
.

Period V
Period

VI

20.5

I059
1029

28.9
32.6

Grams

25-5

35-0

757

35-3

755

38.0

710

Period

VIII

40.5

681

Period

IX
.

39-o

690

57-5

549

as

Grams

787

870

VII

But

Surface

850

Period

Period X

Proportion
to

575

the maintenance

requirement is approximately proportional

to the body surface.
Consequently the feed consumption
the animal
does
older
with
crease
the innot
keep pace
grows
in

its maintenance

requirement, so

that

constantly

diminishing proportionof its feed is available for productive

For
of
example, in Periods I and X respectively
purposes.
Weiske's

experiment it

may

be

computed

that the metaboliz-

PRODUCTION

MEAT

able

of

energy

the

rations

433
and

consumed

requirements per day and

maintenance

Table

hi.

In ration

Required

1568
for maintenance

Available for gain

In Period

ration

available for

was

cent

were

of

Feed

Period

Cals.

2209

807 Cals.

1613 Cals.

761 Cals.

596 Cals.

of the metabolizable

growth as compared

with

of the

energy

only

27 per

X.

regardsthe

increase may

"

Cals.

of the
brief,then, the undisputedsuperiority

In
as

I,48.5 per

in Period

cent

approximate

Period

Energy

Metabolizable

head

Availability

Diminishing

"

the

of feed

amount

be

requiredto produce a

reasonably ascribed

animal

young

unit of

"

concentrated
First,to the fact that his feed is often of a more
and
matter
nature, containinga greater proportionof digestible
perhaps causing a smaller expenditureof energy in connection
with its digestionand assimilation.
Second, to the fact that the gain of live weightin the young
and especially
animal contains a less percentage of dry matter
the

than

represents the storage of less energy

therefore

of fat and

increase in the older animal.

same

consumption of the animal, especially

feeds
used
for
the
more
simple
generally
bulky
upon
increase
the
maintenance
quirement,
reas
rapidly as
growth, may not
the
feed is
that an
so
increasingproportion of
and is unavailable to produce
requiredsimply for maintenance
Third, that the

total

feed

increase.
Production

511.

of the

gain made

even

at different ages

factor in

is a material
is of

of lean meat.

"

The

difference in the

which, as

determiningthe

cost

nature

has
of

justbeen shown,
gain in live weight,

greater importance in another aspect of the

matter.

Chapter XI (460-463),the capacity for growth

for increase of proteintissue,
is especially
in the stricter sense, i.e.,
animal
characteristic of the young
and decreases rapidlyas he
be materially
that it can
grows older,while it does not appear
As

in

shown

OF

NUTRITION

434

FARM

ANIMALS

proteinsupply in the feed. It is of the highest

economic
importance, therefore,to utilize to the full the
abilityof the young animal to lay on proteintissue. In the early
abundant
supply
stages of growth,he is able to utilize a relatively
of feed proteinwhich, if given to an older animal, would
largely
and
be lost so far as growth is
undergo protein katabolism
time the total feed per head
quired
reconcerned, while at the same
stimulated

by

the

is smaller.

for maintenance
to

stint

protein supply

the

of

The

the

of

feeder

afford

cannot

animal, while the

young
be completed or approach

growth can
demands, the
nearly enough to completion to satisfymarket
more
economically will it be conducted.
conclusions
The
regarding the rate of increase of protein
tissue considered in Chapter XI
chiefly
are, however, derived
of the gain or loss of total nitrogenous
from
determinations
besides
the edible portion,the proteinof the
matter, including,
ternal
skin,hair,hoofs, horns and other epidermal tissue,of the inIt is important, therefore,
and of the skeleton.
organs
to inquireinto the rate of increase of the edible portion of the
earlier the

process

carcass.

Table

112.

"

Gain

of

Fat-free

Lean

Meat

and

Total

Protein

by

Lambs

Gain
Aver-

age
Character
Ration
of

Lot

and

per

Week
Head

Age
of

Ani-

mals
Kilo-

Days

Growing
Growing
Fattening
Fattening
Fattening

290

gramsKilograms
O.I14

O.056

521

"053

.031

745

.042

.029

290

.130

.077

.040

.026

458

that the increase of edible meat

fairlyassumed
will be in a generalway
proportionalto the increase of total protein,
be considerable
it is equally clear that there may
departures from
the average.
Unfortunately,however, the data upon this point are
While

it may

be

MEAT

scanty, owing

the laborious

to

About

PRODUCTION

the

only

and

435
of such

expensive nature
available

results

those

are

of

ments.
experi-

Kern

and

Jordan on steers, that is,these are

which
the only ones
permit of a comparison of the rate of growth of
in successive
similar animals
periods. Both these experiments have
been outlined in Chapter XI
(458) Table 112 shows the gain of fatlean
in
free
Kern
and Wattenberg's experiments as compared
meat
with the estimated
refers
gains of total protein. The term "meat"
of
"butcher's
freed
from
sinews
the
and
the
meat
only to
pieces,"
connective
tissue by passage
coarser
through a meat
grinder. The
in general,parallelto that of
production of fat-free lean meat
was,
total protein,diminishing with
advancing maturity. Apparently,
the
of
of
total
rate
however,
gain
proteindiminished less rapidlythan
Wattenberg

lambs

on

and

those

of

that

of edible meat.

Jordan's experiments include a comparison of the weights and

chemical
composition of two pairs of animals at the end of twentymonths'
and seventeen
months'
seven
feedingrespectively.The protein
of the lean meat
after mechanical
separationfrom the fat tissue,
and the total body protein,were
follows :
as
"

Table

So

far

as

"

conclusions

it would
results,
parallelto that
with

113.

Gain

can

of

be

Protein

safelydrawn

that the rate

appear
of total protein and

of

Nos.

has

and

been

were

somewhat

computed

to the

from

gain of

like the

being somewhat
age, the diminution
than in the latter. At all ages
case

content

Cattle

by

more

the

lean

latter

these few
meat

runs

diminishes

rapidin

the

mer
for-

storage of total

The
protein
lighter animals than Nos. 2 and 3.
live weight of the heavier animal in each case.

436

NUTRITION

OF

FARM

ANIMALS

exactly,the increase in the fat-free body)

creasing
considerablyexceeds the gain of lean meat
proper and with into become
maturity this difference seems
relatively

protein(or,more

greater.
512.

Best

the

narrower

the

immature

for

age

"

of increase

sense

While

fattening.

animal,

of

proteintissue is confined to
improvement of its quality by the

the

fatteningprocess is an essential part of

Fattening,however, may be
any

in the life of the animal.

time

in

is to be

hands

the

production in

meat

of

the

commercial

meat

effected at

practically

Assuming that

same

an

from

owner

duction.
pro-

animal

birth until

fattening
slaughter,at what stage should the distinctively
from
be
growth
begun?
process as distinguished
It is evident that the beginningof the fattening
process may
be delayed too long. To take the extreme
be
case, it would
obviously uneconomical
and

to add

then

fatteningperiod.

to full maturity

animal

an

While

there is

no

reason

of feed

the amount

productionof a

actually expended in the

unit of fat would be materiallygreater than if the

fatteningwere

conducted

to

that

first to grow

suppose

during the

latter part of the

growing

period,the expenditurefor maintenance, care, interest,etc.,

added to the cost of production. On
would be simply so much
the other hand, heavy fattening
rations,
largeamounts
containing
if they do not interfere with
of non-nitrogenous
nutrients,even
the growth of young
animals, are uneconomical, tendingeither
to

overload

the meat

with

fat

or

toward

the accumulation

of

cheap internal fat,and making the animal ripefor the butcher

has been properly
before his capacityfor producing lean meat
utilized.

limited market

of
exists,

course,

for fat lambs

and

productionof the world's meat supply it is

important to utilize the capacityfor growth up to a point at
least approaching maturity. Too
early fatteningtends to
produce an animal which, even if not of inferior quality,must
be maintained
in a fat condition until the growth of lean meat
veals ; but

has had

for the

an

beginningof

opportunity
the

to

overtake

fatteningshould

be

that
so

of fat.

timed

Plainly the

that

it will be

to be
completed by the time the rate of gain of lean meat ceases
profitableunder the existingmarket conditions.
The periodin the life of the animal
at which
fatteningshould
begin,then, will depend upon its inherited capacityfor growth,

MEAT

PRODUCTION

437

If this is
its rate of growth as defined on previous pages.
i.e.,
rapid,as, for example, in the improved breeds of swine especially
less extent
be
with cattle and sheep, it may
and to a somewhat
practicableto begin the fatteningalmost from birth,the innate
tendency to growth assuring sufficient size and weight by the
To
this
time good marketable
condition is attained.
secure
result,however, it is necessary to use rations containinglarge
and such
of easily digestiblefeed in a small bulk
amounts
rations are
necessarilycomparatively expensive. Moreover,
expenditure of feed
growth as well as fatteningrequires an
siderable
(473-476)a not inconenergy, and as appeared in Chapter XI
ever,
The capacityof an animal to consume
one.
feed,howanimal
is put on
is limited and when
a
relativelyyoung
full feed,the more
growth he makes the less feed will remain for
fattening. This corresponds with the experience of practical
feeders that mature
animals will reach a higher condition in a
given time than the young ones.
Under
conditions,as a rule,only the best
present economic
grade of animals having to a high degree the quality of early
maturity can be profitablyhandled in the way just indicated.
With

inferior in this respect, the

usuallyis a period of growth upon

animals

more

economical

cedure
pro-

comparatively cheap

roughage,followed
short period of intensive fattening,
beginning,
by a relatively
lost.
however, before the capacity for growth has been entirely
of supporting
The
lies,of course, in the possibility
economy
cheap feeds during
growth and maintenance
relatively
upon
considerable

rations,consisting

to a

longer time necessary in the

will depend to a large extent upon
the

The

stuffs.
be about
For

the

actual
same

the individual

who

case

of

of inferior animals

the relative

costs

and

of

fatteningitself is

of the

feed cost
in either

extent

feeding
likelyto

case.

raises and

fattens his

own

animals,

then, it would appear to be economical, so far as the feed cost

is concerned, to use as early maturing animals as possibleand
for market
to fit them
to push them
at as early an
as
so
age
of.
as
they are capable
When, however, as is notably the case in beef production,the

rearingof
in

to

animals

and

their

fatteningfor market are

considerations
important economic

hands, other
modify this conclusion.

In

this

case

the

in

ferent
dif-

enter

business

of

438
the

NUTRITION

feeder

OF

ANIMALS

FARM

the quality of the

substantiallyto enhance
able
depends to a considermeat, and the profitof the transaction
the difference between
extent
buying and selling
upon
prices,and includes a large element of speculation. While it
which
is true that the animal
still retains more
less capacity
or
for growth will make
the cheaper gains, nevertheless,
if the
market
price of such animals is relatively
high compared with
that of more
animals
it may
be more
mature
profitableas a
business
proposition to feed older animals, even
though the

feed cost

is

pound

per

of

gain is higher.
Condition

513.

It is generally
admitted

Decreasing gains in fattening.

"

that in the

of the

fatteninganimal the rate

of gain in live weight decreases as the fatteningprogresses until
a limit is reached
beyond which the increase,if obtained at all,
is slow and
are
costly. Several causes
responsiblefor
very
this :

case

nearly mature

"

First,the maintenance
requirementof the animal increases
with its gain in weight (393). The
capacity of the digestive
however, undergoes no correspondingincrease,and
organs,
of excess
feed is correspondingly
reduced
consequently the amount
and its proportion in the ration made
less,so that the
total feed requirement per unit of gain will be greater.
Second, the appetiteof well-fattened animals not infrequently
diminishes,resultingin a lessened consumption of feed. This
of excess
again has a double effect,
diminishingthe total amount
feed to total feed.
feed available and reducing the ratio of excess
the close of the
Third, a unit gain in live weight toward
fatteningperiodrepresents a largerstorage of energy than the
same
gain at the beginning(452).
514.

Effect

is natural
state

on

that

of fatness

of
economy
for a lot of

economy
"

the
"

of

gain.
"

condition

should

have

For

"

all these

of the animal
marked

effect

"

on

reasons

that

it

is,its

the rate

and

gain. Georgeson
reports the following results
of
3-year-oldgrade Shorthorn steers, the number
in the table meaning in each case
the number
from

days stated
the beginningof

the
1

feeding:

Kansas

"

Expt. Sta., Bui.

34,

p. 95.

OF

NUTRITION

440

ANIMALS

FARM

gain during the later stages of feedingas

found by other experimenters.
There

is

actual

the

not

sufficient evidence

no

utilization

percentage
of

the advance

with

both

breeds

recognizedmeat
singleanimals

animals

differences

marked

characterize

they yieldfor

reasons

and
to

in

which

particularthere

physiological
causes,

have

has

the

differences between

exist between

for these differences

of meat

such

individual
the feed

sumed,
con-

always been

not

been

common

tendency

unwarranted
and
some
power,
these pointshave gained currency.

Digestive

another

over

difference

to a

although those

who

digestionwhat

is

there

in the extent
make

to which

this assertion

properly
differences

are

animals, but

animals

The

"

as

more

they

clusions
con-

breed
superiorityof one
regardsfeeding capacity is often

power.

the feed is

often
"

or
cribed
as-

digested,
by

understand

utilization."

doubtedl
Un-

between
digestivepower
of manifestlyabthe case
normal
found
to be comparatively

in

except in
been

have

termed

to

difference in digestive

as

assimilative

on

different

the individual

and

the

or

animal

qualitieswhich

return

clearlyseen,
assign them

515.

Individuality

regardsthe

but

or

feed diminishes

excess

important factors in the economy

generally recognized. It is a fact of

that

as

of the

whether

are

production is
observation

and

inherited

those

show

to

generally

fattening.

Breed
That

record

on

been

had

of the

for the marked

slightand quite insufficient to account

in production (718,719). Neither

ences
differ-

any evidence
that the improved breeds of meat-producing animals
possess
the ordinaryunimproved
in this respect over
any superiority
is there

animals.
An

illustration of the latter fact is afforded

Armsby
the digestivepowers

Fries,1in which

and

of

no

material

pure-bred beef

in

difference
animal

experiments by
observed

was

and

"

scrub"

in
at

the approximate ages of one, two and three years.

The same
ments
experiof
losses
material
differences
in
also failed to show
the
any
energy

in urine

energy

which
1

was

and

methane,

so

that

the

percentage

metabolizable,especiallywhen

U: S. Dept. of Agr., Bur.

Anim.

of

the

computed

Indus., Bui. 128 (191 1).

on

feed
the

MEAT

PRODUCTION

441

energy of the
animals.
The

the same
for the two
digestedmatter, was substantially
for
the
the
of
and for
figures
digestibility
dry matter
the percentage of the digestible
each
metabolizable,
energy
being the
of
two
follows
were
as
:
periods,
average
"

Table

116.

Digestibility

"

Assimilative

516.

power.

designatebroadly

the

digested nutrients

of the

producer able

meat

kind

given

more

to

Pure-bred

by

This

"

abilityof
feed

form

from

tissue than

new

term

the

into
a
can

and

Scrub

body
unit

be

may

organism

to

tissue.
of

Steers

used

to

the

convert

Is the

good

digested feed of

the inferior animal

In

words, is the net energy value of the feed affected by the

As yet there has been scarcely
individualityof the animal?
scientific investigationbearing upon
this question, but
any

other

such

evidence

of material
Such

of the

does

indicate

not

the

existence

differences in this respect.

of Kellner's

(449) as
animals

is available

as

show
energy

of the

were

determinations

made

upon

of net

similar

values for fattening

energy
feeding stuffs with different

generallygood agreement
feed,although it does
experiments whether or not
a

of the

as

regards the utilization

not

the

appear
animals

from
used

the

counts
ac-

differed

materiallyin type.
The experimentsby Armsby and Fries,justreferred to, were
directed
of this question. They failed
to the investigations
more
specifically
to demonstrate
any decided advantage on the side of the pure-bred

FARM

OF

NUTRITION

442

ANIMALS

utilization

supplied in
of the maintenance
concerned, the slight
excess
requirement was
difference
observed, especiallyin the earlier years, being perhaps
for by the greater tendency of the pure-bred steer
accounted
to
lay on fat.
of breed
In the aggregate a considerable number
tests of cattle,
experiment stations,
sheep and swine have been made by the American
animal

far

so

as

the results of
the

show

to

individual

of which

some

have

of feed

quantity

cases

as

whole

breed

one

been

of the

energy

by Henry x so as
unit of gain. While
in
less,
to be found, neverthe-

summarized

consumed

per

fluctuations

considerable

the results
of

the percentage

are

certainlyfail to indicate
another

over

marked

any

in this respect, and

later

periority
su-

ments
experi-

to
we
given materiallydifferent results. When
come
consider the other possiblefactors,such as differences in live weight,
in maintenance
requirement, in total feed consumed, etc., we must
clear evidence
of any
results give no
conclude that the recorded

have

not

specificindividual
involved

processes

breed

or

differences

in the conversion

equally true, of course,

that

they

the

in

of feed into
fail to

prove

actual

physiological
tissue,although it is
the

of such

absence

differences.

It
a

than to
that it is necessary
to look elsewhere
capacity of the
greater digestiveand assimilative

clear

seems

supposed

typicalmeat-producing
superiorityover

Thus

requirement.

pounds

4.72 Therms
live weight

various

factors

pointed

out

and

figuresof

muscular
called
found

the

were

of

one

activityof

the

his

nomic
eco-

individual.
specialized
It

"

shown

was

inconsiderable

not

differences

individuals

as

in the

of cattle the

case

in

regards

the

tenance
main-

extreme

of net energy
7.43 Therms
per 1000
for thin animals.
Of the
observed

affectingthe

that

rest, and

that

different

exist between

may

less

explanation of

an

requirement.

(376,391)

VIII

Chapter

the

maintenance

The

517.

for

animal

maintenance

the

most

animal

requirement, it was
important is the degree of
when

even

in the

state

of

so-

requirement
decidedly lower maintenance
pared
by Armsby and Fries for a pure-bred beef steer as comdue
with
scrub was
there interpreted
to
a
as
probably

more

the

and
disposition

nervous

greater restlessness

of

the

latter.
It is
rations

clear,however,
the

which

one

Feeds

and

that
has

of two
the

animals

lower

Feeding, 10th Ed.,

receivingidentical

maintenance
pp.

328 and

511.

requirement

PRODUCTION

MEAT

443

largersurplus for growth or fatteningand, other

the greater increase per unit of
things being equal, will make
lower maintenance
To what
extent
total feed.
a
requirement
remains
is characteristic of high-bred meat-producing animals
If it appears
to be a generalfact,it would
to be determined.
the part of the latter.
on
go far toward explainingany superiority
of
It is not impossible,
also,that differences in the amount
muscular
important part in fattening
activitymay play a more
hitherto reported. In
than in the experiments on maintenance
able
the latter,the experimental conditions necessitated considerof motion, while under the conrestriction of the freedom
ditions
the

will have

of

perhaps be

wider

practicea

scope may
the individualityof the animal in this
Feed

518.

consumption.
is
superiority

individual

element

of

important

abilityof

the

to

respect.

Another

"

afforded

animal

an

to

consume

of feed.
Of two
animals
otherwise
regularly large amounts
day
similar,it is clear that the one which is able to consume
after day the heavier ration is the better meat
producer. It is
not
however, that the heavier feeder makes a
always realized,
profitableuse of his feed because, as pointed
relativelymore
in Chapter VIII (360),assuming the maintenance
out
ment
require-

the productivepart of the ration forms a

be the same,
of the large eater.
larger part of the total ration in the case
to

Consequently, since
makes

the

There

but

breeds

doubtless

are

in this

animals
as

gain, not

paid for, this

because

simply because

he

he

is able

marked

differences

between

respect. Whether

whole

in view

doubtful

to

better

be

animal

utilizes his
to

consume

of it.

more

of

economical

more

feed

excess

all the feed must

possess any
of the results

individual

the

specific
meat-producing
in
this
advantage
respect appears
record regardingthe feed cost
on

Apparently the quality is one

rected,
which
the attention of breeders has not been speciallydiwhich, it would seem, might well repay
yet it is one

gain with

different breeds.

attention.
519.

that

Type
the

factor

in

breeds

as

and

conformation.

conformation

of

"

meat

It is

animal

fact
well-recognized
is

very

important

determining his sellingprice. The improved meat

to live weight,
a rule show
a higherratio of dressed

better distribution of fat in the finished carcass,

somewhat

OF

NUTRITION

444

largerproportionof
of meat.
They are
an

animal

ANIMALS

the

higherpriced cuts

all

important

of meat, but

renders

FARM

there

is

factors

as

higher quality

in the

evidence

no

efficient

more

any

and

that

economic
their

duction
pro-

possession

of feed into

converter

meat.

economic

The

importance of a rapid
of the consequent early maturity has been
rate of growth and
considered
in previousparagraphs (504,505).
of common
It is a matter
experiencethat there exist marked
Early maturity.

520.

"

individuals

differences between

of the

weight finallyattained by the

the

speciesboth

same

animal

mature

and

as

as

to

the

to

It is natural to interpret
this
growth at the same
age.
differences
in the
fact as indicatingcorrespondingindividual
of proteintissue,but the writer is
of growth, especially
rate
of any recorded
on
not aware
experimentsbearingspecifically
that the qualityof early maturity is
this point. It is true
popularlyattributed to the meat breeds, but as regards cattle
rate

at

of

least

Henry

has

shown

that the beef breeds

such

as

the data

that
show

fail to prove

hand

at

greater

gain in live
dairybreeds,

of

rate

weight or a greater weight at maturitythan do the

although it is likewise true that other elements than simply
into the economic
the weight enter
conception of maturity.
If it is correct
to

in

variations
a

breeder

noted

above

rate

of

gests
sug-

the

the feeder.

" 3.
521.

differences

growth of protein tissue,it

of much
interest both
to
investigation

the

field for
and

the individual

ascribe

to

Feeding

Feeding

as

Meat

for

related

to

Production

individuality.
"

The

facts considered

capacityof the animal

of vegetableproducts into
for the conversion
as
a mechanism
meat.
They (and other less important ones) determine the
degree to which, from the commercial
standpoint,the animal
is able to utilize the feed given it. Favorable
modifications
of any
of these factors are of advantage because
they enable a
largerand more
profitable
productionto be secured.
in the

previous section relate

in

Feeding stands
purpose

is to

somewhat

supply the
1

Feeds

and

material

to

the

different
upon

Feeding, 10th Ed.,

relation,in

which
p. 329.

the

that

mechanism

its

MEAT

PRODUCTION

445

It is of

prime importance to the feeder that his animals

the largestpossibleproductive capacity,
shall have
but while
which the animal can
the maximum
produce is determined by
its breed and individual characteristics and cannot
be materially
the amount
affected by feeding,
which it actuallydoes produce
must
in any given case
of material
depend upon the amount
suppliedto it in its feed. Production may be limited by a
deficient feed supply,although it cannot
be forced above a certain
works.

maximum

by increasingthe ration.

Feed

522.

supply

requirements.
of

Since feed is to be looked

"

upon

material for the animal

mechanism, it is clear
that the kind and amount
required will depend primarilyupon
the capacityof the animal.
The young
animal,with his marked
of the specific
more
capacity for growth, will requirerelatively
materials for growth, viz.,protein and ash, than will the older
animal.
The early maturing animal, with his greater rate of
total feed per day than the one
growth,will requiremore
turing
maa

as

more

and

raw

slowly. The
utilize largetotal
in

larger amounts

order

animal

with

capacity to

of feed must

amounts

that

the

his

advantage

be

sume
con-

given these

in this respect

may

be

time

according to circumstances the requirements for the production

of fat. The feed requirements for these two
purposes

have

but
already been considered in the two previouschapters,
be convenientlyrecapitulated
here with more
particular

fullyutilized.
As alreadypointed out, meat
of
production is a combination
growth and fattening,the latter process being superimposed
The feed requirements of the meat-producing
the former.
upon
include in the firstplacethe requirements for
animal, therefore,
normal
growth, to which are added during a longeror shorter

may

emphasis

on

economic

Protein
523.

Relation

that it is
of meat

relations.

requirementsfor

to age.

"

It

only during growth

in the

narrower

was

meat

shown

production
in

Chapters X and XI
considerable production

that any
i.e.,of muscular
sense,

tissue,takes

place,and
young
and more

likewise that the energy of growth is greatest in the

animal and diminishes,at first rapidlyand then more

slowly,until physiological
maturity,when

but

slight

446

OF

NUTRITION

FARM

ANIMALS

proteinand stillless of meat proper can be

secured.
Evidentlythe questionof the necessary proteinsupply
in the rations of meat-producinganimals is of specialimportance
the
of
during
earlystages
growth.
of the total

increase

524.

Minimum

protein supply

animal, then, in
be

growth must
in addition

to

order

suppliedin

for

to

growth.

utilize

The

"

fullyhis

his feed at each

ducing
meat-pro-

capacityfor

stage of that growth,

his maintenance

requirement, with at least as

he is capable of storing up in his

much

protein as
digestible
growth. Whether
any greater quantity than this is necessary
gree
deor
advantageous is,as has been shown
(491),still to some
unsettled question. Some
with
an
experiments,
especially
cattle and sheep,indicate that any considerable surplusis unnecessary
for normal
the
other
hand, feeding
growth, while,on
with ruminants
cate
indiextent
experimentswith pigsand to some
that amounts
of those thus computed
considerablyin excess
at least greater gains of live weight.
assure
525. Protein
requirements in fattening. While growth
and fattening
be regarded physiologically
distinct proas
may
cesses,
it is economicallyimportantin the practice
of meat
duction
prothat they should go on
less simultaneously.
more
or
The
animal
the very young
is not
growth of even
simply a
but normally includes more
less
or
production of protein tissue,
fat production,while in proportionas one
has to deal with
earlymaturing animals it is desirable to begin the fattening
proper at a comparativelyearlystage of growth (512)
There
for regardingthe actual fattening
to be no
reason
appears
different in the growing and
as
being essentially
process
"

in

the mature

animal.

that in the latter

required

for

case

It has
no

maintenance

reason

combined

material

shown, however
excess

is necessary.

supply of buildingmaterials
no

been

is

of
So

(453,456)

proteinover
far

as

the

there
concerned,therefore,

that
mere
seems

that the actual proteinrequirementfor

suppose
growth and fatteningis any greater than that for
to

normal

growth without fattening.The conclusions regarding

the proteinrequirements for growth recorded
in Chapter XI
(482),therefore,may be regarded as applicablealso to young
animals
that are being fattened,especially
rived
desince they were
in part from results on immature
fatteninganimals, and

from

this

point of

view

the increased

feed

supply requiredfor

448

OF

NUTRITION

ANIMALS

FARM

proteinthan is called for by the estimates

digestible
supplymore
in Appendix Table IV b. When
pensive
exproteinfeeds are especially
smaller proportionof proteinmight doubtless
even
an
to economic

be used

advantage,

even

without
digestibility,
supply, especiallyin the case
maturity.

loss of

some

of

Specific effects

527.

animals

of

of the

approaching
must

be

experimentsupon

the

stuffs.

feeding

also of the fact that in most

taken

though at the expense of

unduly curtailingthe protein

"

Account

protein requirement thus far reported the variation in

proteinsupply was effected by varying the proportionsof
in the

concentrates

for maize.

meal

however,

such

not

are

we

to

not

may

of the ration but may

serve
stimulate the growth process or
While

tain
cer-

cottonseed
ration,such, e.g.,as substituting
As was
suggestedin Chapter XI (491),

substitution

the

suppose
nutrients

only affect

the ash

substances

to introduce

ance
bal-

which

perhaps the fatteningprocess.

that

such

substances

can

take

(738),they might enable the

place of actual
or
they might,
fullyutilized,
proteinin the ration to be more
the
for more
by stimulating fattening
process, create an appetite
the

feed.
At

rate

any

it

seems

that the addition

of certain

oil

rations

meals,

them

to

to

the

feed

consume

generalexperienceof stockmen
feeds rich in protein,
the
especially
fatteninganimals tends to induce
freelyand thus (518)to yieldmore

to be the

of

more

profitable
gains.
Energy requirements
for
528.

Combined

growth

and

meat

production

fattening.
"

An

attempt

was

in

made

Chapter XI (480-483)to estimate approximatelythe

values requiredat different ages for normal growth
net energy
without material fattening.To the extent to which
the fattening
time, these requirements
process is to be carried on at the same
must
evidentlybe increased by amounts
equal to the
additional
desired

in

by

net

energy

stored

up

in the increase

of

sue
adipose tis-

expected. Subject to the limitations indicated

previous paragraphs, this additional energy may be supplied
the

or

addition
to the

of either

terials
nitrogenousor non-nitrogenousma-

growth ration.

MEAT

PRODUCTION

449

Chapter X that a pound of increase in

live weight in the mature
fatteninganimal is equivalentto about
of energy.
If it is allowable to apply this average
3.25 Therms
to the fatteningof younger
animals, this would be equivalent
that
to sayingthat for each pound of increase in weight above
It

due

to

in

estimated

was

of net energy
3.25 Therms
to the requirementsfor growth as estimated in

growth proper,

be added

about

should

Chapter

(480-483). The energy requirement of the meat animal,

will obviouslydepend on its capacity to produce gain
therefore,

XI

of flesh or fat and

the extent

capacity,and

and
specific

no

to which

it is desired to utilizethis

invariable

requirementscan

be

formulated.

If, for the reasons

given in
proteinin the
previousparagraphs,the proportionof digestible
ration is kept above a certain limit,the questionof the amount
of net energy to be suppliedresolves itselfinto the question of
total amount
of feed to be given and this
the most
profitable
529.

Total

amount

of feed.

"

depends upon a varietyof conditions.

It has alreadybeen pointedout (512)that only with animals
having a rapidrate of growth and maturing earlyis it advisable
to begin intensive
feedingbefore a fair degree of maturity is
reached.
With
ordinary animals the major portionof their
growth may be more
cheaplysupportedupon pasture and the
small amounts
of concentrates,
ordinary roughages with relatively
since the growth process
be materiallyhastened
cannot
by heavy feeding. When, however, the time for beginning
of expensiveconthe fattening
centrates
process involvingthe use
is reached
(533),whether this be earlyor late,it is
in order to reduce
important to hasten it as much as practicable
the cost of maintenance,attendance,etc.,and the question
of feed becomes an important one.
of the most
amount
profitable
530.
Heavy feeding profitable. That comparativelyheavy
feedingof fatteninganimals is economicallyadvantageous is
of practical
and is evident from
shown by the experience
feeders,
the fact,to which attention has alreadybeen called several times,
that a less proportionof the heavy ration is requiredfor the
of the animal.
maintenance
Were
this the only factor involved,
of
it would follow mathematicallythat the greater the amount
"

the greater would be the growth per unit of feed

therefore that the appetiteof the fattening
animal should

feed consumed
and

NUTRITION

450
be

stimulated

the

OF

FARM

ANIMALS

ever,
possible. In fact,howother considerations
in to modify this conclusion.
come
531.
Influence
on
tent
digestibility. Overfeeding to the exand throwing the animal
of causingdigestivedisturbances
to

greatest

extent

"

"

"

off feed

is of

and

rather

feeder

the

than
from

aside

But

turbance
avoided, since the resultingdis-

subsequent lessened consumption

advantage from the increased amount

the

outweigh any
may
It is the regularuniform
animal

be

to

course

with

one

this

that is

danger, it

consumed

results

The

increases.

well

seems

of mixed
digestibility
decreases
productivefeeding,

in

rations,such
more

or

record

on

less

of the
efficiency

Influence

532.

that

as
digestibility

in the net

decrease
to be
to

somewhat

this effect

quantity
respect (722)
of the nitude
magtend to diminish

just noted

the

the

that when

values.

value

is,of

"

Such

equivalentto

course,

ration

become

of the

ration

is flooded

organism

matter

less

with

There

energy
efficient in

of

As
as

The

the

appears

yet

in

seems

net

energy
reportedhave

rangingfrom

energy

of

appears

actually
producing

in other words

"

the resorbed

products

and

stimulated
are
processes
the digestedmatter
escapes

the katabolic
of digestion,
share

(764),the
Chapter XVII
hardly sufficient to warrant

that in addition

and

is increased

of

decrease

of the rations.

the
digestibility,

on

the amount

gain as

rations.

energy

energy

be

the

as

in this

however,
general impression,

from

resorbed

net

on

that

would

as

for any quantitative

estimate
but it is evident that it must
of this effect,
the

profitable

established

scarcelysufficient

are

eaten.

capriciousappetite.

the percentage
used

be the

to
likely

feed

of

evidence

on

a
as

this

larger
heat.

"

point

positivestatements.

feedingstuffs which have thus far been

been
obtained chieflyin experimentson rations
ing
submaintenance
to only moderately heavy fatten-

rations,and

values

the

of

results show

no

distinct

indication

of

of feed.
On the other hand,
amounts
increasing
that
render it quite conceivable
considerations
physiological
and so increasing
metabolism
the effect of the feed in stimulating
be relatively
the heat production(365)may
greater on a high
than on a low nutritive plane.
less falling
off in the nutritive effect of
or
Apparently more
be anticipated,
is increased must
ration as its amount
a fattening
of
of decreasing digestibility
whether
or
account
on

decrease

with

PRODUCTION

MEAT

451

utilization of the

combination
a
or
digestiblematter
this diminution, within the limits of the
Whether
of the two.
animal's capacity to consume
feed, is sufficient to offset the
economic
advantage of such increased consumption remains to
be shown, although Morgen * reports experiments on sheep in
which
gains in
very heavy rations, actually produced smaller
live weight than lighterones.
Finallyit should be remembered
that it is the actual gain of chemical energy by the animal which
relation to the feed enis believed to bear a tolerablyconstant
ergy.
It has been repeatedlypointed out that the gain in live
of energy
weight is a very uncertain indication of the amount
It is quite conceivable
that the largergain to be
stored up.
lessened

expectedon
dry

matter

the

lighterration,and
not

may

be

contain

the heavier ration may

less protein and more
or
that

less water

fat than

consequentlythe

proportionalto

the increase

533.

or

of

Proportion

considerations

wiped

to

In

heavier

feeding

out.
to

concentrates

apply

produced on
increase in weight
that case,
recognized by the

higher quality of the gain were

advantage attached
market, the economic
be diminished

more

that

in feed.

unless the

would

and

in

the

roughage.

"

first instance

The
to

going
fore-

varying

feedingstuffs. In the case of

be effected
herbivora,however, heavy feeding must
necessarily
of roots
to
or
by increasingthe proportion of concentrates
roughage, the higher cost of the former per unit of net energy
than offset by the economic
being more
advantage incident to
amounts

of the

the

much

an

addition

mixture

same

largeramount
of

of

which

concentrates

can

When

be consumed.

contains

such

large proportion of

carbohydrates (as in the case of maize or roots)it would appear

of the rations would suffer to a certain
(724)that the digestibility
of the ration,while
extent
owing to the low protein content

rapid passage of the

feed through the digestive
tract of heavilygrained ruminants
which
has not,
and suggests a decrease in total digestibility
however, been experimentallyconfirmed.
It is clear from the foregoingthat under
534. Standards.
or
nearly mature
fatteninganimals,
ordinaryconditions mature
fattened in the United
such as the cattle ordinarily
States,
should be fed as heavilyand pushed as rapidlyas the capacity
common

observation

indicates

more

"

Futterung unci Schlachtergebnisse,

pp.

22

and

33.

conclusion

reached

was

Such

an

of

intensive

be

compared with roughages,it is economy

extent.
largestpracticable
these conditions

in
significance
at least

afford

basis

it is evident

feedingstandard

the amount

as

for

the

mony.
quitein hareffected only by a

unless the latter

are

to

as

Under

This

permit.

investigation
appear
feeding can

and

of concentrates

use

feeder will

that
so
feeders,
long ago by practical

experienceand

results of

ANIMALS

FARM

skill of the

the

and

of the animals

free

OF

NUTRITION

452

very
use

expensive

them

that there is very

in the

ordinarysense,

to the

little
so

far

of feed is concerned.

it is true,
It may,
preliminarycomputation of the amount

feeding,if this is of any importance,

is
induce
the
but in actual feedingthe problem
to
animals,
of the art of the skilled feeder,to consume
large
by means
of feed without
amounts
injuryto their appetitesor digestive
of
capacity,and this is largelya questionof the individuality
lot. The one
the animal
or
thing to be kept in mind is to see
the
that the supplyof proteinin the ration is sufficient to ensure
normal
a
relatively
growth of proteintissue,since this causes
rapidincrease in weight.
definite reFor younger
more
quirements
fatteninganimals, somewhat
indicated in a
might be formulated in the manner
previousparagraph (528)on the basis of the requirementsfor
fatteningand for growth as estimated in ChaptersX and XL
l
The
of American
compilationby Bull and Emmett
ments
experilambs
referred
in
cluded
XI
to
on
(487)infattening
Chapter
data regardingthe computed net energy content
of the
rations.
They conclude that the production of satisfactory
of digestible
gains requiredthe followingamounts
proteinand
net energy
pounds live weight.
per iooo
of feed

Table

requiredfor

117.

"

season's

Requirements

of

Fattening

Lambs

Weight

Ills. Expt.

Sta.,Bui. 166 (1914).

per

iooo

Lb.

Live

MEAT

PRODUCTION

453

similar

No

of farm animals have

compilationsfor other species
valuable
material in the
reported,although much

yet been

the

of
publications

"
While

Influence

4.

the

awaits

experimentstations

capacityof

the animal

as

supply of feed sufficient in quantityand

that

the

capacityare

under

the conditions
influence

producer and a
qualityto fullyutilize
meat

great factors in

two

which

discussion.

Conditions

External

of

such

the animal

production,yet

meat

is

kept are

without

not

the results obtained.

on

Temperature
of

Teachings

535.

animal

from

"

feed, it

the

warm
fairly

in sheds
amount

that
better
a

this

of this

feeding in

number

the writer

same

siderable
recentlya coning
has been reportedshowpracticeactuallygives

and

more

comparisons have

leave

doubt

no

as

to

results of
been

the

which

Critical

536.

(354),there

is

marized
sum-

validity
the principles

in harmony with
conclusion,while it is entirely
governing the influence of external temperature

metabolism

upon

in

previouschapters.
in Chapter
shown
As was
temperature.
certain approximate temperature, called
a
were

for

time,however,
being successfully

the

quarters. The

warm

of such
l

of feed

emphasis has been

importance of providing

feed lots and

experimentalwork
supposedly uneconomic

considerable

by

open

of

than

returns

oxidation

rived
de-

that cold

conclude

were
cattle,in particular,

and

of materials

considerable

the economic
upon
quarters for live stock. At
of

to

wasteful

past

great numbers
fattened

natural

seemed

surroundingswould lead to
simple heat production,and
laid in the

temperature of the

by the katabolism

is maintained

body

Since the

practice.

discussed
"

VII
the

critical temperature, at which

outflow of heat
the minimum
justbalances the necessary heat productionresultingfrom the
internal work

and

below

which

more

or

less oxidation

of tissue

required to maintain the normal temperature of the body.

Furthermore, it has been shown
(395-397)that the digestion

is

and

assimilation
1

U. S. Dept.

of feed

and

Agr., Bur. Anim.

its conversion

into tissue result in

Indus., Bui. 108 (1908),pp. 79-86.

the

OF

NUTRITION

454

ruminants, and

in the

of heat
with

this

lower the critical temperature as compared

In other words, there is
that of the fastinganimal.
for each

and

heat

the

which

the effect of

that

heat, especially
internal production

to
virtually

is

animal

for each

of

relatively
large amounts

of

evolution

ANIMALS

FARM

ration

becomes

produced

gotten rid of by radiation

and

certain

above

temperature

in part

an

excretum,

to be

evaporation.

of heat productionover
likelythat a certain excess
the body temperature
is
that absolutelyrequiredto maintain
advantageous, both as promoting the comfort of the animal and
of temporary fluctuations
a margin in case
as affording
especially
On the other hand, both our
of temperature.
own
sations
personalsenof practical
stock feeders show that
and the observations
both
an
affecting
high temperature is debilitating,
unnecessarily
then, it is desirable to
appetiteand generalhealth. In practice,
keep the thermal surroundings of the animal within the range
somewhat
above
above
the critical point but not
indicated
much
to affect the appetiteand thrift. It is evident
as
so
so
that the limits of this range may
vary widely with the kind of
It appears

"

animal

and

537.
the

the

with

of ration.

Amount

of the

amount

The

"

ration.

influence

requirementsfor protectionfrom

cold

of this factor upon

is clearlyindicated

by what has already been said. The heavier the ration,other

heat will be evolved
thingsbeing equal, the more
during its
conversion

digestionand
feed thus

have

their

at

into

tissue.

disposala large amount

and

naturallycan thrive under

detrimental
might be seriously
relativelylightrations.
a

ox
fattening

Table

118.

Metabolizable

Energy

the

gave

Thus

energy

stored

as

conditions
to
one

Heat

538.

of heat

Excess

over

Age

like animals

and

of

in

surplusheat

experimentson

Fattening

5"92o
20,740

Cals.
Cals.

15,060 Cals.

....

5, 680

maintenance

of different

full

26,600 Cals.

of ration

gain

weight

on

"

Production

Energy evolved as heat

Computed maintenance
requirement
Excess

animals

of exposure
which
growing animals on

young,
of Kellner's

followingresults :

Excess

"

Mature

Cals.

37-7%
of

animals.

sizes,under

"

like

The

internal

work

conditions,appears

of
to

456

NUTRITION

OF

FARM

ANIMALS

equal,then, it will clearlybe desirable to have the water

approximate as nearly as possiblea continuous
while if the
by having it constantlyaccessible,
watered only at intervals the temperature of the water
are
need

be

to

rather

higherthan

in the other

sumption
consumption
con-

stock
may

case.

Shelter
and from wind may
be of quite
protectionfrom rain or snow
as much
importance as protectionfrom low temperatures simply.
is
541. Precipitation. An
important factor in the case
the amount
ing
of precipitation
(rainor snow) to be expecteddurthe feeding period. In cold weather
the low
temperature
A

"

of the water
of

which

loss of heat

affected

the

by

than

and

seem

as

has

once

of the animal's

thickness

the coat

out

of the

some

would

be

coated

the heavier

thoroughly

the
nature

of the

climate

shelter the animals

is
relatively,

the animal

or

that

as

is
regardshumidity and precipitation

from

the
that

storms

temperature

in its

in many

localities

be

efficient as

may

as

bearing
roof to
a

tight

dentally,
inciadvantage of the roof, already mentioned
of a dry bed, thus
is that it providesthe possibility
penditure
only adding to the comfort of the stock but avoiding exing
meltof energy in warming up or evaporating
water
or

barn.

snow

542.

One

"

ice.

or

Wind.

"

All

are

familiar

with

compared with a stillone

large part of the protectivevalue

windy day
A

it

observed, are largelyindependentof

thermometer, and it is clear that the

quite as important a factor as

and
the question of shelter,
on

not

when

will be

of the

indications

animal

Still greater,

wet.

on
falling
requiredto melt the snow
which
it is compelled to lie.

it
effects,

portant
im-

more

is the

the heat
upon
These

Far

coat.

quired
expenditure of heat reafter it is wet, and this,as it would
cate,
to indiexperiments with sheep seem

greater with

become

cause

be regarded as practicallyan addition

may
of both losses being
to the cold air,the extent

this,however,

dry

to

is the

which

due

to that

the skin of animals

to

penetrates

the coat

of

as

an

animal

fibers of the material.

resides

Wind

the greater

severityof

of the

temperature.

same

or
clothingof man
the
in the air entangled between
tends to replacethis air with fresh,

of the

PRODUCTION

MEAT

greatlyreduces the protectiveeffect.

thus

cold air and

have

therefore,
may
feeding.
Insolation.

543.

457

effects of the weather

The

"

distinct economic

A windbreak,

in stock

value

are

appreciably

by the exposure of stock to direct sunlight. Aside from

direct effect of the light
inconsiderable amount
as such,a not

modified
any

of heat is imparted to the

this is likely
to be

weather

the

hot months
its heat
work

is

reverse

the sun's rays.

During cold
distinct advantage,but during the

body by

production below

that

from
resulting

cannot

duce
re-

its internal

and assimilation of its feed,it may

riously
sedigestion
heat
tax its powers
to disposeof the additional
imparted
direct sunlight. In this case
shelter of some
sort may

and

the

by

the

be

required for opposite reasons

cold months.
and

the animal

Since

true.

For

to the wind

exposure

the animal

similar

to

to
a

reasons

may

be of

those

obtainingduringthe

supply of cool,fresh water

great advantage in helping

get rid of its surplusheat.

Other conditions
644.
^

Exercise.

The

"

well-known

fact that muscular

tion
exer-

of tissue

is

accomplishedat the expense of the katabolism

and ultimately,
at the expense of the feed,would seem
therefore,
of the meat-producat firstthought to indicate that the activity
ing
should

animal
case

of the

element

be restricted

as

much

as

practicable. In the

growing animal, however, another

enters

exercise tends

into the case,

to stimulate the

very

important

the fact that moderate

namely,
growth of the muscular

system,

Since this is
words, the productionof lean meat.
of
the essential objectsought, a normal and reasonable amount

or, in other

muscular
allowed

activityon
and

growing animal should be

exercise
though the muscular
feed.
more
Accordingly,young

the part of the

encouraged, even
involves the consumption of
stock should be given the freedom
of the pasture or range
to
time care
while at the same
as
great an extent as practicable,
should be taken to supply abundant feed containinga sufficient
supply of protein in order that enough material may be present
to supply the demand
for growth stimulated by the exercise.
In the case
of breedingstock, especially,
most
a
important
consideration is that of the health and stamina of the animal,

458

NUTRITION

which

can

above

principles

the

on

of

other

while

ensure

it

then,
reduce

to

should

"

be

of

the

close

is

at

545.

rapid

to

and

is
of

have

this

the

the

ters
quar-

in

may

the

case

similar

For
of

animals

they

important

supply

be

ing
fatten-

the

that

so

portion,
pro-

should

and

nature.

water

In

comfortable

possible,

tein
pro-

health

feed.

keep

to

bedding,

timid

stage,
of

endeavor

end

particularly

its

of

fattening

the

for

water

the

requires

as

the

well

as

of

utilization

of

consumption

physiological
for

be

never

"

involving

feed,

reason,

should

It

supply.

of

this

taken

as

more

sons
rea-

animals

hand.

production,

amount

For

desirable

Water

amounts

exercise

This
account

on

reached,

the

plentiful

for

growth

of

is

with

stock.

sufficient

assimilation

To

of

fattening

little
maintain

to

undisturbed

as

the

comparatively

and

classes

allowed.

reached

necessary

The

opportunity

an

frequently

possible.

as

down."

lie

sheep
it

of

all

to

way

have

stage

provided,

kept

and

eat

only

this

quiet

be

is

is

overconfinement.

given

which

amount

as

be

appetite

as

the

animal

should

is

normal

general

there

hand,

tissue,
to

animals

ANIMALS

through

than

exercise

case

the

in

should

hogs

FARM

suffer

to

apply

and

movement

In

fail

hardly

particular,

In

OF

the

that

relatively

large

corresponding
of

purposes

the

mentioned

previously

one

forgotten

animal.

(540)
,

it

is

desirable

possible,
too

cold

that

stock

at

all

times

to

be

consumed

and

should
that

freely

have
the

by

water

the

ready

access

supplied
animals.

to

should

water,
not

if
be

CHAPTER

MILK

"
546.

The

i.

XIII

PRODUCTION

Physiology
of

Components

milk.

In

"

Production

Milk

of

addition

to

milk

water,

of the four great groups

of nutrients,
representatives
fats,carbohydratesand ash.
viz.,proteins,
stance
The principal
Proteins.
protein of milk is casein,a sub(55). This
belongingto the group of phosphoproteins
protein is peculiarto milk, not being found elsewhere in the
body.

contains

"

In addition

to

casein,milk contains also

lact-albumin

Their presence
in small amounts.
paraglobulin
of
the casein by means
by precipitating
of peptones, possiblydue
the filtrate. Traces
also found in milk.
of a proteolytic
are
enzym,

be demonstrated

may

acid and
to

and

heating

the presence
has

been

1.79 per cent to 4.23 per cent and

of the other proteins
from 0.25 per cent to 1.44 per cent.

that

According
observed

Fats.

to

Konig, the casein

to

vary

Fats

"

varying

from

average

to

stated

in

in

and

as

the diameter

simplya

condensation

of these

generalway to range from

a
singlecubic centimeter of
into

runs

formerlydescribed

milk

ules
microscopicglobin
loidal
colthe
suspension
of

the

millions.

surrounded

by

protein nature, but the supposed membrane

as

to

cow's

In

their number

globuleswere
of

be

millimeter

0.01

milk

held in

greatlyin size and

globulesmay

0.0016

in the form

in milk

occur

solution of casein.
fat

of milk

content

of the

proteinof

The

fat

membrane

is

the

now

garded
re-

milk, due

surface tension.
Milk

of
fats,is a mixture of a number
As compared with body fats,the
triglycerids.

fat,like other animal

simplefats
fat of milk

or

rich
relatively
melting point.

is

low
relatively

consequentlyhas a
from
is especially
distinguished

in olein and
It
459

460

OF

NUTRITION

molecular

fatty acids of low

Chapter I (30), where
of

important

proportion
already noted in

weight, as
the principalconstituents

list of

considerable

"

of these so-called

given. The presence

acids which can
(i.e.,
an

of

the presence

body fat,however, by

ANIMALS

FARM

be distilled in
for

means

of

current

detection

the

volatile

of

is
"

fattyacids

steam) affords

adulterations

of

butter.

The

the other

cent

varies

widely. For the cow a

1.67 per cent is reportedby Konig. Six per cent,
hand, is a high figure,althoughoccasionally
7 per

of

minimum
on

of fat in milk

percentage

is reached.

Babcock
for

observed

that

states

givingas

cow

much

as

is the

cent

9 per

pounds

15

mum
maxi-

of milk

daily.
and

fat

milk

The

also

varying

lecithins and

cholesterins

of

coloringmatter, derived, as
shown, chieflyfrom the carotin of

amounts

Eckles

and

Palmer

of

carries traces

have

the feed.
Milk

Carbohydrates.
"

in

contains

solution

disaccharid

itself,
namely, lactose,or milk sugar (13). In
distinction from fat, the percentage of lactose in fresh milk
shows
comparativelysmall variations,averaging about 5 per
in cow's milk.
The
cent
souring of milk is brought about by a
peculiar

to

of the

fermentation

milk

sugar

by

which

its molecule

is

split

of lactic acid.

into four molecules

tioned
organicingredientsof milk should also be menin appreciable
in the
acid,which occurs
quantities
citric

the

Among
form

of calcium

Ash.

about

citrate.
total

The

"

cent

0.7 per

mineral

according to Van

the ash
Qualitatively,
in all animal

matter

of milk

substances.

Its

in

milk

cow's

averages

Slyke.2

contains the

ingredientsfound
quantitative composition,however, as
same

hand, and with that of

the tissues on
the other, shows some
interestingrelations.
Bunge 3
gives the followingfiguresfor the composition of the ash of the serum
compared

with

the blood

of cattle blood
added
of

Lawes

and
and

serum,

of the

Gilbert's

ash

on

the

of cow's

one

milk.

figuresfor the ash

these have

To
of

calf for the sake

comparison.
Jour. Biol. Chem., 17 (1914),191-264.
Jordan, The Feeding of Animals, 1908, p. 305.
3Ztschr. Biol.,10 (1874),301 ; 12 (1876),191.

1
2

been

MILK

Table

119.

Composition

Percentage

"

461

PRODUCTION

Serum
Cattle
Blood

Ash

of

of

Cow's

K20

22.1

55-i

5 -40

1.6

MgO

0.6

2.6

0.1

0.04

CI

20.0

47.1

P2O5

2.20

0-53
0.12

24.8

40.37

tions
animals,having a shorter period of growth, the relal
for
the rabbit Bunge
more
striking. Thus,
reports

smaller

are

the

43-95

21.3

3-4

With

3.82

13-9

CaO

Fe203

OF

3-2

Na20

Bodv
Calf

Milk

even

followingresults.
Table

120.

"

Composition

Percentage

Serum
Rabbit
Blood

of

Ash

Body
of

of

Days
Old
Rabbit

14

3-2

10.8

54-7

6.0

CaO

1.4

35-0

MgO

0.6

2.2

0.0

0.2

K20
Na20

Fe203
CI

P205

It appears

that while

sodium

and

47-8

4.9

3-o

41.9

chlorin

are

the

predominant

ingredientsof the blood serum, these elements are present in milk in

phorus
relativelysmall proportions,while potassium, calcium and phosin
milk
of
the
t
he
ash
latter,
closelyresembling
predominate
that of the body of the same
species.
547.

figuresas
cow's milk
1
2
3

see

cites the
following
Wing
showing approximately the average composition of
3
accordingto various authorities.

Average

composition.

"

fiirPhysiologie,II, 188.
by Sellheim in Nagel's Handbuch
its Products, 1897, p. 17.
Milk
For data regarding the composition of the milk of other speciesthan
Schaefer's Text Book of Physiology,Vol. I, p. 125.

Quoted

and

cattle,

462

NUTRITION

Table

121.

OF

ANIMALS

Composition

Average

"

FARM

American

Casein

3-40

3-30
3.00

3.02

3-40
0-45

O.70

4.88

4-55

4.60

4.80

0.71

o.75

o.75

"-75

100.00

Milk

548.

in

two

glands.

number,

body, although in

The

"

one

100.00

milk

animals

many

or

or

two

has

The

compound

in

structure

milk

40.

Manual

(Armsby,

gland.

of milk

Lobule

of

great number

Feeding.)

of Cattle

in

Fig. 40,

in groups
of
outlet
shown
as

common

alveoli
which

lead

the "active

are

passages
cells but

These

lined with

are

of

ducts

3 to 5
in the

cluster.

the

different

unite to form

alveoli

sort

and

alveoli,

are

olus
alve-

of

an

inch

in diameter

lobules

form

nective
con-

and

vessels,nerves

to

having

figure. Internally,the
cells (Fig.41),
epithelial
The

.ducts

or

also lined with

which

largerones,

Each

layer of

outer

singlelayerof
milk.
agents in secreting

leadingfrom

to

schematically
correspondingto the single

berries of the grape

consists of an

united

Its

shown

are

carrying capillaryblood
lymphatics. These alveoli are about -^
are

sheep

consists

It
grapes.
of acini or

tissue

and

and

gland.
roughly compared

be
may
of
bunch

three of which
"

horse

tubulo-acinous

of

outlets

lobes, in the cow

the hog from ten to
gland is classified

in

that

the

two

three,and

or

into

having separate

Thus

gland

line of the

is subdivided

gland

lobes

more

fourteen.
as

glands,properly speaking,
each

teats.

each

99.60

IOO.OO

side of the median

each

on

two

Fig.

87-75

3-25

"-53

Ash

are

87-75
2.80

Albumin

Sugar

mann)

French

(Cornevin)

87.60

3-69

(Fleisch-

(Oliver)

87.17

Fat

Milk

German

English

(Babcock)

Water

Cow's

of

as

do

not

shown

to the teat, emptying first into

finally
the base of the
a cavitylyingnear
cistern,"

in

epithelial
produce milk.

the so-called
teat.

which

Fig.42,
In

"

milk

compound

464

FARM

OF

NUTRITION

ANIMALS

their full'development only in the

reach

of

formation

the

of the alveoli and

time, a rapid growth

that

At

new

ones

pass
That

substances
stimulating

into the blood

other

fluid found

of the

in

Starling,the formation
so

reach

the fetus which

the milk

glands.

cooperate, however, is shown

apparently well-established

the

by

least

at

may

and

perhaps
to this

stimulus

(Hormones) in

of the mother

causes

the

occurs,

growth being,according to Baylissand

of certain

stages of pregnancy.

later

fact that

the

the

moval
regularrevirginanimal,

glands of the
mechanical
lead to the formation
even
stimulation,may
considerable
in
instances even
quantitiesof milk, in some

or

of

the male.
550.

The

secretion

secretion and
is

not

clearlyshown

of milk.
mere

That

"

formation

milk

filtration of material

from

is
the

true

blood

the facts

position
alreadystated regarding the commilk.
As
all
of
the principal
was
pointedout,
organic
of the milk are
ingredients
peculiarto it. Casein and lactose
in the animal body, and while the prinnot found elsewhere
are
cipal
milk
in
also found
the body fat,their
are
simple fats of
different in the milk fat and the latter is specially
are
proportions
characterized
by the presence of glyceridsof the lower acids
series.
of the aliphatic
marked
more
Furthermore, even
titative
quan-

by

differences exist between

milk

and

those

of the blood

the mineral
From

serum.

elements

all these

of the

facts,it is

gland is a producing or secreting

organ and
of the milk are largely
manufactured
ingredients

clear that the milk

that

the solid

in it out

of materials

derived

from

the blood.

propounded by Virchow
found
wide acceptance.
According to this theory,milk production
of a physiological
consists essentially
fattydegeneration
of the epithelial
cells of the alveoli. The microscopeshows that
the cells of the activelysecreting
gland are largerthan those
in the restinggland and more
less filled with fat globules,
or
It was
the side toward the cavityof the alveolus.
on
especially
held that while this process went
the cell divided,forming
on
and that finally
the cell next to the cavityliquefied,
two or more,
settingfree the fat globules which it contained and, perhaps
A

theory

of milk

the

Milk

production was

tissue.

addition

of

with

secretion

more

thus

first

less water, constituted

regarded as a form of the
or

the milk.

growth

of

465

PRODUCTION

MILK

however, has generallyfailed to

Subsequent investigation,
show
satisfactoryevidence of cell division. A modification
of Virchow's theorystillheld is that while there is no cell division,
the outer portionof the protoplasm is sloughed off and dissolved,
forming the milk, and is again renewed by the growth of new
protoplasm. The weight of opinion,however, regards milk
served
productionas a true secretion,entirelyanalogous to that obin other
a

glands.

cells,but

of

breaking down

It is not

believed that there is normally

that

secreted materials into the alveolus

cells of other
well

as

to

as

This is held

the other

analogous to that
cellsof
by the epithelial

material

The

as
precisely
to apply to

of the

secretion of milk

continuous

the

the

The

is in

process

the obvious
intestine,

in the active udder

is

move.

more

or

less

the

possiblycontain
of milk produced in a singlemilkingby a reasonamount
ably
tion
productivecow, and it is well recognizedthat a rapidsecreof milk occurs
duringsucklingor milking. In other words,
milk gland,like other glands,reacts to a specific
stimulus.

551.
source

secreting
fat globules

the materials

however, that the cavities of the udder

the

do the

product accumulating in the cavities

of the gland. Fleischmann
long ago showed,

process,

passages

their

resorptionof digested

the small

being the direction in which

difference

latter extrude

ingredientsof milk.

ways

many

and

glands.

the

Sources

of

ingredients of

of the material

feed,the

milk

immediatelyfrom
brings about

extensive

its
the

milk.

While

"

in the milk

contained

gland draws

cannot

supply of

is of

material

blood,while

at

the

transformations

chemical

the ultimate
the

course

for milk
same

in

duction
protime it

the

stances
sub-

supplied. Probably all the ingredientsof the

milk should be regarded as products of the chemical activity
of the epithelial
cellsof the glands,althoughthe extent
to which
the original
material is modified varies.
and globulin
552. Origin of milk
proteins. The albumin
of milk are quitesimilar to the corresponding
substances in the
blood.
The casein,on
the other hand, is radicallydifferent.
In the firstplace,it is,as alreadystated,a conjugatedprotein
the
containingsome
phosphorus-bearingradicle. Whether
latter is derived exclusively
from the organicphosphorus compounds
of the feed has not
been demonstrated, although it
phosphorus compounds (phosappears probablethat inorganic
thus

"

466

OF

NUTRITION

be utilized

phates)may

of the

sources

as

ANIMALS

FARM

phosphorus of the

milk

(257,258, 497).
gation
production of casein,however, is not simply a conjuThe
of a simple protein with
a
phosphorus group.

The

of casein

constitution

is

different

markedly

from

of

that

the

of the muscles,as is shown

or
by
proteinsof the blood serum
the proportionsof its various
cleavage products as given in
Chapter I (50)so that if casein is formed from the proteinof the
,

blood

considerable

tissue,a

or

of

reconstruction

cules
mole-

their

hand, if the casein of the milk

cells of the udder, in the manner
epithelial
suggested in Chapter V (226,227), from the simplercleavage
for the milk gland.
productsin the blood, the process is specific
is necessary.
is built up in the

553.

Origin

the other

On

of milk

the
249) in discussing

fats.

It

"

of

sources

stated

was

body

in

fat that

Chapter V (247latter

although the

be derived
show
in part from the fat of the feed and
some
may
of its characteristics,
nevertheless,the production of fat must

be

of the fat cells,

to the activity
essentially
to a simple deposition.
first place,it has been demonstrated
by the researches

regarded

and

not

In the

of

and

Jordan

formed

from

In

others
the

that

milk

or

with

one

determined.

which

feed,a

could

from

possiblybe
the

of the results

Table

1
2

very

N.

poor

After

considerable

fat

carbohydratesof

Jordan'sl experiments

ration
milk

due

as

cows

balance

fed

in fat and

was

carbohydrates.

left
The

the

the

for

accounted

as

body fat

either with

production

maximum

an

amounts

protein and
which
could only have
followingtable gives a

by

"

Production

of

the

Fat

by

ordinary

of fat in the

"

122.

be

may

the feed.

were

deducting

well

as

Cows

Y. (Geneva) Expt. Sta., Buls. 132 and 197.

Digested protein of feed less gain of protein by the animal.

fat

of

fat of the
been

duced
pro-

summary

467

PRODUCTION

MILK

of fat from
perhaps be pointed out that the formation
have occurred
carbohydratesin these experiments may not necessarily
that the main
in the milk gland itself. It is entirelyconceivable
have
taken
place elsewhere
portion of the synthesisof the fat may
and that the fat or its precursors
were
simply transferred to the milk
gland.
It should

of
Second, it has also been shown by a considerable number
experimentsthat,as in the case of body fat,the fat of the feed
sensiblyaffect the propertiesof the milk fat. Not only
may
have
changes in the melting point,iodin number, and other
of
properties

fat been

butter

found

follow in

to

generalway

in the feed

fat,but characteristic ingredients

changes
of foreignfats given in the feed have been detected in the milk.
While it is not necessary
to conclude, and is indeed
unlikely,
is
it
that the feed fat
as
were
simply transferred,
mechanically,
the other hand, that relatively
to the milk, it is clear,on
large
able to pass through the
are
fragments of the fat molecule
it evident
These
facts render
epithelialcells into the milk.
of milk fat. Not only so, but experithat feed fat is a source
ments
later (613),
to be mentioned
by Morgen and his associates,
similar

to

seem

show

that

animals
of milk
idea

The

certain

least)conduces

at

extent

in it of the lower

is

acids

of the

or,

breaking up

of the carbon

on

be
may
simplercarbon

aliphaticseries,which
fat from

arise during the partial

may
in the feed fat which probably

hand,

chain

into milk

precedesits transformation
a

is

synthesisof

the other

compounds,

As

efficient production

most

to a
produced synthetically
perhaps supported also by the presence

steps in the

intermediate

to the

fat.

it may
general conclusion,therefore,

be stated that the

fat of milk may

have its origineither in the fat or
it may
of the feed,or in both.
Whether
from

protein has not been demonstrated

but reasoningby analogy with the formation
be regardedas at least very probable.
must
554.

Origin

of lactose.

"

The

yieldingupon

hydration dextrose

its derivatives

are

and

it is also

bivorous
(inher-

fat.

that the fat of milk

considerable

of fat in the feed

amount

abundant

constant

lactose of milk
and

in the carbohydrates

also be produced

experimentally,
body fat, it

of

is a disaccharid

galactose.

Dextrose

in the feed of herbivorous

of
ingredient

the blood.

On

or

animals
the other

468

OF

NUTRITION

FARM

ANIMALS

contains
ordinary feed of herbivora
drates
carbohyis
latter
transformed
the
apparently
yieldinggalactose,
and
has not
into glycogen quite promptly
at any
rate
been
found
in the blood,while animals
receivingfeed containingno
The
galactose (carnivora,e.g.)produce lactose in their milk.
to be that the galactosehalf of the lactose
probabilityseems
in the milk
is manufactured
gland from the dextrose of the

hand,

the

while

blood.
of

Sources

555.

ash.

ingredientsof the milk,

phosphorus, are, of course, derived
The

"

ash

including its sulphur and

ultimatelyfrom the correspondingingredientsof the feed. In
rations containing
liberal milk production on ordinary winter
from
vestigat
inof organic nutrients,however, it appears
a sufficiency
of calcium,
by Forbes 1 that considerable amounts
magnesium and phosphorus may be drawn from the relatively
large store contained in the body, presumably to be replaced
in later stages of lactation.
556.

milk

that

of

Character

milk

production is

production.
a

of

form

tissue

it is essentially
true
incorrect anatomically,

compositionof
the feed

on

comparing
and

milk

supply

of the

in that

comparison its

milk, it

growth
so

far

This

in

is evident

sugar should
Taking Babcock's

is

probably

the chemical
it makes

clearlyshown

is

by

of
organicmatter
weight of growing animals.

protein to fat
increase

as

statement

which

the demands

concerned.

milk

of fat.

amount

are

the ratio of

the solids of

In

product and

the

the

While

"

in the

to make

that in order
be

reduced

to

fair

the

figures(547)as

equivalent
representing

vided
compositionof milk, the 4.88 per cent of sugar diby 2.25 is equivalentto 2.17 per cent of fat,which added

the average

per cent of fat present as such makes

equivalentof 5.86 per cent, while if milk sugar were
the

to

3.69

by

fat the

total

this

organicmatter

would

amount

parts of

total fat

thus
to

placed
re-

9.41

would

organic
contain 37.73 per cent
proteinand 62.27 per cent of fat.
Comparing these figureswith those given in Chapter XI (458)
that
for the composition of the increase in growth, it appears
the proportionof protein to fat is greater than that computed

per

cent.

On

basis, 100

matter

of

for young
The

animals

computed

except in the earliest stages of growth.

energy
1

Ohio

content

of average

milk

Expt. Sta., Bui. 295 (1916).

solids is 2620

469

PRODUCTION

MILK

of
the energy
content
animals, but less than

Cals. per pound. This is greater than

the dry matter
gained by very young

general way,
be said that milk solids correspondin proportion
then, it may
of proteinand in energy value per pound to the gainsmade
by
that

growing animals
557.

months

of

Rate

pounds per

of

dry

neighborhoodof

production of milk

solids.

three months

beef calf three

"

old.

growth of approximately
day, containingperhaps three-fourths of a pound
with

matter

very

moderate

day would

contain

The

in the

when

be assumed

old may

1.5

stages of growth. In

in later

computed

an

make

to

about

of about

content

energy

yield of

Cals.

2200

pounds of average milk per

lent
pounds of total solids equiva-

15

1.92

words, considerably
than twice as great a productionwould be effected by the
more
relativelysmall bulk of the secretingcells in the udder as by
to

5030

other

In

of energy.

Cals.

is further

it
body of the calf. When
is
that the product of the dairy cow
economic
value
as
a
producer of human
this point Jordan says : 1
On
is
milk
pounds of average
year
per
the

whole

edible,her

all

food

"

considered

cow

not

great
vious.
ob-

becomes

yielding6000
regarded as an
annual
tion
produc-

however, the
amount
less than
of not
an
780 pounds of milk-solids,
in the body of a cow
least double the dry matter
weighing

at

pounds.

900

This

animal.

unusual

When

we

means,

consider

that this manufacture

of

new

only duringa singleyear, but through

tensive
the entire adult life of the animal, we
begin to realize how exthe food supply. Still more
the demands
are
upon
yieldingannually over
strikingis the case of high-gradecows
the performand when we remember
half a ton of milk solids,
ance
of Clothilde,whose
26,000 pounds of milk produced in

material is carried

year

on

not

certainlycontained

than

more

2500

pounds

of solid

wonderful powers
regard the cow as possessing
of transmutation.
Her
capacity for the rapid and economical
food of the highestqualityis not equaled
production of human

matter,

by

any

558.

from

we

must

other animal."
Factors

of milk

production.

productionin one
latter,broadly speaking, an
meat

animal

is what

is
1

very

The

Milk

essential

increase

sought, and

"

while

productiondiffers
particular.In the

in the whole
the

Feeding of Animals, 1908,

body of the

product may
p.

308.

vary

in

NUTRITION

470

the feed

quality,all

market

requirement
In

milk

OF

FARM

consumed

is available

production,on

ANIMALS

the

in

for

of the

excess

the

of

production

contrary, what

tenance
main-

is desired

gain.
is the

singleset of glands. An increase in weight in

the mature
is not
dairy cow
sought. At best it represents a
than the one
in view, while
diversion of feed to other purposes
the activityof the
considerable fattening tends to check
any
milk glands. In feeding for milk
production, therefore,it is
tenance
to consider not
only the surplusfeed above the mainnecessary
requirement but the factors affectingthe distribution
milk productionon
of that surplus between
hand
the one
and
The
art of feedingfor
growth or fatteningon the other hand.
milk consists in stimulatingthe milk productionto the greatest
economicallypossibleextent and in supplying.the feed material
secretion

of

for

necessary

this

production,while

avoiding,in

the

mature

animal, any material increase of body tissue.

The
factors governing milk production are
essentiallythe
of animal
mal,
in other branches
same
as
production,viz.,the aniand the feed supply.
the environment
In milk production,however, the relative importance of the
first and

second

productionfor
the distribution

is greater than

conditions
the

that

reason

of the

they

forms

of

materially influence

may

feed between

excess

in other

milk

productionand

tissue increase.

" 2.

The
The

559.

Animal

as

Factor

in

Milk

Production

prime factor in successful dairyproductionis the

secretingtissue
which
rate of activity
and of yieldinga secretion of good quality,the most
scrupulous
and the most
abundant
care
feedingwill inevitablyfail to yield
animal.

latter possesses
abundant
is capable of being stimulated
to a normal
Unless

the

returns.
satisfactory

Individuality
560.

Includes
may

is

simply

an

breed
be

said

differences.
to

aggregate of

related

individuals.

It

discuss

problems of

breeds

include
more

or

is outside
and

"

that of

breed, since

less similar
the

of

influence

The

scope

and
of

dividual
in-

breed

genetically

this work

breeding,and this branch

to

of the

NUTRITION

472

three

of these

poorest animal
animal.
breed

was

the

between

has been

of the

some
investigations,

differences

shown

by

same

the

statistical

numerous

have

strikingof which

most

of the

averages
the
exist among

greater differences

even

of the country

cows

common

individuals

That

breeds.

for the three

differences between

greater than

much

were

picked herds, however, the production of the

only from 50 to*6o per cent of that of the best
the

Moreover,

remarkably good considering

In
to a public test.

are

incident
necessarily

conditions

the unfavorable
each

of these records

the lowest

Even

ANIMALS

FARM

OF

been

lected
col-

by Eckles.1
562.

Influence
that

the
(i.e.,

on

utilization

the

of

of milk

amount

of

economy

in the udder

feed

the

solids of

from

feeding.
a

in

it is

While

the

narrower

likely
un-

sense

factured
given composition manu-

of nutritive

equal amounts

supplied)is materially affected by

"

the

substances

of
individuality

the

is
economic
sense
animal, the feed utilization in the broader
this factor.
be conIt must
stantly
largely dependent upon
very
borne
in mind
that, as already stated (558),efficiency
in milk productionis in largepart a question of the distribution
of maintenance.
Some
of the feed suppliedin excess
animals,
able to
are
by virtue of individual or inherited peculiarities,
of excess
feed into milk without
ing
stortransform large amounts
up any considerable portion of it in the form of body tissue.
if well fed
tend to remain
Such animals
spare in body and
of milk.
They are the typicaldairy
produce large amounts
wellthe
animals.
Other
a
individuals,on
contrary, have
marked
tendency in the opposite direction,viz.,toward the
fed heavily, they utilize
production of body tissue. When
little or
feed chieflyin this direction and show
the additional

tendency toward an increase in milk production. These

The
two
are
types, of course,
typicalmeat-producinganimals.
shade into each other by imperceptible
gradations.

no

The

important bearing

dairy
Here

animals
it may

will

simply

individuals

paragraphs

abilityto
into

be

which

noted

has

milk

been

due

to

large amounts
rather than

Dairy Cattle

and

into

Milk

facts

the

upon

nutrition

of

(606-610).
of certhat the superiority
tain
illustrated in the preceding
considered

further

is doubtless

consume

be

of these

later

considerable
of feed

body

and

extent

convert

tissue.

Production, 191

1, pp.

18-126.

to

the

the
plus
sur-

PRODUCTION

MILK

Influence

563.

on

473

of lactation.

course

That

"

individuality

determining the rate at which

milk yield falls off with advancing lactation is shown
in a subsequent paragraph (568).
specifically
important part

plays an

Influence

564.

composition of milk.

on

that

observation

common

in

cows

more

of

matter

regards the

as

vary

It is

"

the

richness

of cream
butter
or
milk, that is,as regards the amount
be obtained from a given weight of milk.
Various
which can
breed tests at experiment stations have served to define more
the comand breed upon
position
exactly the influence of individuality
of their

The

of milk.

and

illustrate this influence

to

at

two

in the

results

The

average

in

from

test, and

the average
results
of the milk.
fat content

Table

124.

Average

"

each

proportion

to

are

Composition

ranging from

time

24

intended

are

ferent
difcompare
of the results obtained

table shows

covering periods of

years.

primarilyto

not

the averages
three different experiment stations
The

breeds.

in tests

in Table

results cited

station

the

for several breeds

eightmonths to
are
given weight

number

arranged in

of

Milk

of

of

under

cows

the order

Different

of the

Breeds

Num-

ber

Casein
and

Ash

Albu-

min

Holderness

Ayrshire
Shorthorn
Devon

Guernsey
Jersey

maximum

224;

50

Maine
New

Solids

0.68

3-14

5-n

346

12.39

12.29
12.82

12.45

3
2

3.22

4.89

3-5i

IS*

S-i3

3-27

4.80

3-64
3-65

o.73

0.76

3-74

5-32

4-54

14.36

0.75

3.78

4.94

4.96

1443

o.75

3-92

4.98

5-25

14.90

evidentlythe

OF

Cows
Aver-

aged

0.67
0.67

is

Fat

than

tose

Fat

Total

%
.

Holstein-Friesian

Lac-

most

variable

ingredientof milk,

its

in these averages
by more
exceeding its minimum
Along with the increase of fat there is
per cent.
Jersey Expt. Sta.,Rpt. 1890, pp. 223Expt. Sta.,Rpt. 1890, p. 29; New
York
(Geneva) Expt. Sta., Rpt. 1891, pp. 94-104.

also

of the

increase

an

these

but

and

25 per cent
increase of 12

an

per

changes, the

Influence

565.

of the

nature

differences

by computing
with
total solids,

Table

125.

Average

"

not

composition

on

clearer
free

of
range
increase

the

solids,

of the

the

and

differences

regularlywith

lactose

other

constant

most

total

comparatively small

ingredients.The
of the milk.
ingredient

of the

the

fat,the totals
cent, respectively. There is also
the proportion of ash, while the

extreme

it does

cent, while

per

per
in

cent

of

that

than

contrary, shows

the

lactose, on

protein and

total

21

ANIMALS

FARM

smaller
relatively

are

being

11

OF

NUTRITION

474

milk

of

is

regular
ir-

being
the

crease
in-

evidentlythe

solids.

The

"

real

composition,however, is rendered
percentage composition of the water-

in

in Table

the results shown

Composition

Milk

of

125

Solids

"

Different

of

Breeds

Num-

ber

Casein
Ash

"

AND

OF

Fat

Lactose

Albumin

Holderness
.

Holstein-Friesian

Ayrshire

Devon

Guernsey
Jersey
.

5-49

25-44
26.20

41.24

7-93

39-79

28.56

40.01

28.39

5-86

26.37
26.27

38.55

29.32

5-29

26.04

37.06

31.61

5.20

26.20

34-23

34-37

5-o3

26.31

33-42

35-24

5-23

Shorthorn

aged

5-45

Cows
Aver-

foregoingtable,it appears that the percentages of

ash and of total proteinin the milk solids are very constant,
the averages
from
but very slightly
the singlefiguresdiffering
essential difference in
of 5.36 and
26.10, respectively.The
the composition of the milk solids lies in the proportionof
lactose
to fat, the former
decreasingas the latter increases,
taken togetheris pracwhile the total percentage of the two
tically
the
from
cent
less
than 0.7 per
average
constant, varying
From

of

the

68.53 per

cent.

In other

words, it appears

from

these

figures

producing milk rich in fat the secretingcells form

more
fat,while, as
relativelyless lactose and correspondingly
that

in

cows

PRODUCTION

MILK

Table

475

by

shows, this difference is accompanied

124

tively
rela-

smaller secretion of water, so that the percentage of total

solids in the resultingmilk is greater.

Cooke,1 in 1890, drew

over

analyses of

2400

of the

Haecker2

United

has

the
milk

States

study of

tions
reported by the experiment starecently
up to that date, and more

the
substantially

reached

from

conclusion

same

analysesof 544 individual samples

result

same

from

of milk

from

the Minnesota

Station herd.
566.

be

of composition
Variability

that

noted

the

in

foregoing conclusions

the average composition of the milk

for comparativelylong periods. The
of the

animal.

same

drawn

are

of the

It should

"

from

individuals

same

composition of the milk

does vary quite
frequently

however, may and

its average
widelyfrom one milkingto another without affecting
composition as computed from analysesof a number of milkings.
in the case
of the fat because
This has been observed
especially
same

cow,

far

have

determinations

more

been

of

made

this

stituent
con-

other,the fat being both the most valuable

easily determinable ingredient.Variations as

than of any
and

the

great
the

as

most

same

3 per

cent

per

cow

cent

are

not

in the fat content

not

very

differences of

and

uncommon

there is

Whether

rare.

of successive

as
proportion of lactose,
the previous paragraph,does not

that these variations

are

due

and

even

tion
correlated varia-

in the averages

in the

in

milkingsof

appear.

largely

to

compared
It is presumed

external

influences but

factors of environment
specific
has been traced with certainty,
although Spier3 believes them
It is evident that correct
to be due to incompletemilking(575)
comparisons of the yieldsof different animals, or of the same
be made
animal at different times,can
only on the basis of the
of days.
average yieldand composition for a number
in the composition of milk from
The extent of this variability
one
milkingto another appears to be an individual peculiarity,
the milk of some
uniform in daily composition
cows
being much more
An
than that of others.
example of this
interesting
has been reported by Farrington.4
no

definite connection

with

any

1
2
3
4

Expt. Sta.,Rpt. 1890, pp. 97-100.

Expt. Sta.,Bui. 140 (1914),p. 51.
Jour. Highland and Agr. Soc, 1909, p. 287.
Vt.

Minn.

Ills.Expt. Sta.,Bui.

17

(1891),p.

9.

476

ANIMALS

FARM

OF

NUTRITION

Stage of lactation
production

Milk

567.
has

of nature

state

of the young

shortlybefore,while

or

becomes

Milk

"

tion
produc-

of the offspring. In a
objectthe nourishment
it is a periodic
function,beginning at the birth

its

as

periodic function.

dependent

less

as

the

on

the young
animal gradually
mother
it diminishes
in

has greatly proAlthough man

longed
that the
the period of milk production of the cow,
so
short and in some
she goes dry is relatively
time during which
milk production
instances is eliminated
nevertheless,
altogether,
character and undergoes marked
stillretains its periodic
changes

intensityand

during the

of

progress

Influence

568.

of

finallyceases.

yield

on

the

produced, but

of milk
at

different stages in the

at

the

does

of the
case

fallingoff

slow

rapiddecrease
of farrow

going dry for

the milk

cows,

comparatively slow
The

greatlyfrom
animal

569.

to

cow

and

by

cow

observed

any

Influence

toward

the end

570.

Bearing
the

of lactation

continue

for

much

may

from

to show

longer time.
be called,
however,

vary

to

year

general the
fat tend to increase,
especially
the quantity of milk falls off

when

of

milk.

In

"

changes in quantity,these variations in

often
are
scarcely
irregularand sometimes
until the rapid fallingoff in quantity sets in

of the lactation.

the end

changes in

more

the

all

at

cally
typi-

conditions.

of lactation

rapidly. Like
composition are
manifest

by

is

calvingapproaches,all but
In the
longer or shorter time.

composition

on

there

in

year with the same

often not
plained
irregularities
readilyex-

and

marked

show

the birth
weeks

two

followed

months

amount

next

they

as

amount

rule,the

immediatelyafter

percentages of total solids and of

toward

As

productionmay

decrease

of lactation

curves

of the

effect

differ widely
may
in different animals

and

lactation.

for several

the time

as

exceptionalcows
a

animal

same

of the

increase for one

but shows
or
an
young,
of the cow.
Following this maximum,

case

evident

most

of decrease

rate

its maximum

reach

not

The

"

gradual decrease

periodin
corresponding

of milk

the

milk.

of

is the

lactation

advancing

lactation.

on

experimental methods.

yieldand composition of

must

be

taken

account

"

milk

The

unavoidable

with

the advance

of in all

experiments

on

MILK

production,and

milk

PRODUCTION

render

477

the

interpretationof their results

peculiarlydifficult. It is obvious, e.g., that if a change of the
ration of a cow
is accompanied by a decrease in her milk yield
part at least of the decrease
and

not

to

be due to the progress of lactation

may
On
the change of feed.
the other hand an

of the milk

increase

yield in

later

period of the experiment

be partly offset by the natural
shrinkage in milk. In
may
brief the later periods of an experiment are at a disadvantage
compared with the earlier periods.
methods
for eliminatingor attempting to eliminate this
Two
influence of lactation have been used, viz.,the period system
and

the group
The

system.
the

In

duced
period system, as introby Wolff, Kiihn and others of the earlier experimenters,
the animal receives an identical ration in two or more
periods
well removed
from each other in point of time
usually the
and from the results of these periodsthe
firstand last periods
gredient
daily rate of decrease in the yield of milk and its inaverage
is calculated.
On the assumption that had the same
ration or treatment
been continued
unchanged this rate would
have
been
uniform
be
throughout the experiment, it may
mediate
computed what yields would have been secured in the interperiods. A comparison of these computed yieldswith
those actually observed
is taken as the measure
of the effect
of the change in feed or other conditions.
The
of
accuracy
571.

period system.

"

"

"

this method

depends
that

the

of

course

the

on

yields would

have

of the

correctness

decreased

at

sumption
as-

uniform

rate.

572.

The

introduced

each

other

groups

relative

with

animal

animal

receive

production

his

to
or

of the group

use

with

group.
same

group

animal

or

seeks

system,
group

of

on

with

to

the
other
an-

long preliminaryperiod

ration

is determined.

was

Copenhagen

period system

an

In

the

system

their determinations

itself. The

compare

the

in

successors

(702). The

units

hand, attempts
check

The

"

in connection

Station

so-called feed

compare

both

system.

by Fjord and

Experiment
the

group

or

treatment

and

their

One

of the groups
is then
the same
continued
while with the other group
treatment
on
the factor to be tested is introduced.
Finally,in a concluding

period,both

groups

are

again treated

as

in the initial

'

period.

478

OF

NUTRITION

of the

combination

two

receives

the other

uniform

ANIMALS

also be used, one

may
rations in successive periods,

systems

being fed varying

of animals

group
while

FARM

ration

throughoutthe

entire

experiment.
A

of the methods

discussion

complete

very
of

of

eliminatingthe

fluence
in-

advancing lactation in the interpretationof the results of

experiments on milk production is to be found in a recent article by

Morgen.1

" 3.

Influence

The

The

word

which

all those

affect

may

to be

is here

environment

to summarize

term

Environment

oe

milk

important of

more

which

used

looselyas

influences

external

The

production.

particularlysensitive

external

to

tion
Produc-

convenient

other

than

dairy cow
in the

feed

appears
of the

conditions,some

considered

are

Milk

on

following
paragraphs.

Milking
is but

Milking

an

of the

sucklingof the
in securing the milk is likely
efficiency

imperfectimitation

and naturallyits
young,
to be affected by a variety of circumstances.
573.

Frequency

of the udder
at

of

milking.

"

As

already stated,the cavities

hold

heavy milkers cannot

milkings
milking. Between

one

in

considerable

accumulation

there

of matter

all the milk

produced
be

evidently may

in the

alveoli

and

canals

which

the effect of diminishing the secreting

to have
appears
activityof the epithelialcells through what might be crudely

called
effect

"

back

of

secretion

pressure." Suckling

relievingthis

"

or
"

pressure

milking
and

would

have

the

perhaps rendering

time it seems
to act as a
easy, while at the same
direct stimulus to secretion.
At any rate it is a fact that more
frequentmilking tends to increase the yield of milk, especially
in the

more

of

and in the earlier stages of lactation.

good cows
The
effect of frequent milking is strikinglyillustrated in the
off in the
The abrupt falling
followingexperiments by Kaull.2
milk yield when
the milking was
made
pervery frequent may
"

case

Landw.

Cited by Kellner, Die

Vers.

Stat.,77 (1912), 351.

Ernahrung der landw.

Nutztiere, 6th Ed.,

p. 521.

480

OF

NUTRITION

FARM

ANIMALS

it is more
concentrated
than that yielded
fat,i.e.,
The
differences in the composition
after the longer interval.
of the night'sand morning's milk, which
have been the subject
of so much
this basis,the
discussion,appear explicable
upon
the morning's and night's
interval between
milking beingusually
the night'sand morning's.
less than that between
solids and

be

the

to

be greater than

percentage

of

solids-not-fat

fact of the greater richness

The

little.
"

is well

known.

caused

as

by

in the

contained

small
well

If successive

"

found

portions will
while

milking.

milking be analyzed, the percentage

same

pings

of

Completeness

575.

the

of

This

udder.

The

the entire anatomy

as

untenable.

of the

earlier ones,

is held

comparatively
so-called
strip"

at

was

time

one

cream

fact,however,

of the

difference

The

in the

risingof the

of milk

amount

of fat in the later

varies

difference

actual

an

portions of

the milk

on

that

plained
ex-

but

paratively
com-

in the milk

udder, renders

is probably due

to a

cistern,as
explanation
partialretention
and canals,they
this

stagnationof the fat globulesin the alveoli

out
being afterward washed
by the portions of milk secreted
during the latter part of the milking. Incomplete milking
not only fails to get this fat,thus lowering the qualityof the
milk
that
(compare 566), but it appears
actually obtained
or

of

secretion
is

of

retention

the

the

the

practicable.The
the

the

it

as

be

to

seems

influence

been

much

as

exercise
small
extent.

to
to

upon

the

United

"

various

methods

with

of

manipulating
probably due

most

to

cows

chines
ma-

remove

hand.

exercise,
fatigue

exertion
a

the

milked

necessary

of muscular

In

check

Hegelund method, are

Similarly,in the use of milking

exertion

discussed

basis.
been

the

of

to

milking, therefore,it
out
as
completely as

or
by
portions,
strippings,

Muscular
The

be

tends

alveoli

all forms

cow

this influence.

largelyto

the

advantages of

udder, such

the last

in
In

milk.

that

important

fat

milk

upon

secretion has

comparatively slender
States

the

mental
experi-

questionhas usually

of motion
and
of allowing freedom
desirability
while
in Europe, especially
dairy cows,
among

the

farmers, cows

are

used

for draft

to

not

inconsiderable

Feed

576.

of

discussion

(391)to

of exercise.

cost

the very

Attention

"

maintenance

the

marked

481

PRODUCTION

MILK

in

requirement

effect of muscular

called in the

was

Chapter

VIII

exertion in increasing

of body fat or of the non-nitrogekatabolism,especially

nous
the
feed.
It
has been frequentlyargued
suppliedby
ingredients

the

this fact that the amount

from

should be restricted

cows

indeed

have

as

reasoning that since the

feed into milk
exertion

was

any

possible. Not

as

confine

to

as

their

to

few

dairy
men
dairy-

entirely,

cows

objectof their business is

diversion

to

convert

the support of muscular

and
This, however, is a very narrow

waste.

much

far

so

gone

of exercise allowed

of it to

inadequate view of the subject. Most authorities on dairying

of exercise for dairy cows
amount
ficial.
beneas
regard a moderate
* in
Thus
Martiny
1871 cites five authorities on this
point and expresses the opinion that exercise and moderate
work
increase rather than decrease the yield of milk, while
has

work

severe

an

unfavorable

effect upon

both

the

yieldand

quality. Similar opinionsare expressedlater by C. F. Miiller,

These earlier
Fleischmann, Kirchner,Konig and Von Klenze.
data

of the

are

rather than of actual

empiricalobservations

experiments.
"

influence

those of
the

less

or

more

Morgen's investigations. Of actual experimentsupon

577.

the

of

nature

of

Morgen

muscular
2

at

exertion

milk

upon

the Hohenheim

production,

Experiment Station

are

under
made
convincingbecause they were
strictly
comparable conditions and especiallybecause the relative
of work
amounts
performed in the different periods were
most

determined.
The

Simmenthal

two

being used

for draft.

employed

cows

The

work

was

done

the sweep power dynamometer used

upon work productionby the horse
of work

the

accustomed

were

at

slow walk

to

upon

Wolff in his

by
experiments
(386a, 670, 779),the amount
performed being regulatedin part by the resistance of

dynamometer

in part

required,so that

of hours of work
by the number
approximatelysingle,double and quadruple

work

The

done.

was

and

ration

unchanged throughout the

of

1 1
1
3

fed,which
trials. The

periods,approximating two

Die

Milch, Part I, pp. 345-435Eleven

to twenty-six days.
2

liberal one, was

experiment consisted

was

weeks
2

Landw.

each
Vers.

of alternate rest
Stat.,51 (1899), 117.

482

NUTRITION

FARM

OF

ANIMALS

periods,beginning with a rest period, so that each

work periodwas
precededand followed by a rest period.
of work performed caused some
The rather moderate
amount
of milk produced, the effect tending to
decrease in the volume
work

and

be

littlegreater in the

periods in

which

most

work

done.

was

chieflya decrease in the amount

decrease,however, was
in the yieldof
of water
secreted,although a slightdiminution
served.
total milk solids,
ranging from 10 to 85 grams per day, was obIn other words, the effect of the work
to render
was

The

the

somewhat

milk

effect,however,

concentrated.

more

was

upon

the

The

notable

most

yieldof fat, which

showed

an

in every case
but two.
This increase was
pensated
comfor by a decrease of the fat-free solids,
so that analyses

actual increase

showed

of the milk

higher percentage

of fat and

while the percentage of fat-free solids remained

in

obtained

cases

some

solids,

changed.
unpractically

Quite similar results,although

by less rigorousmethods, have been

Confirmatory results.

578.

of total

"

Backhaus,3 Torssell 4 and Dolreportedby Dornic,1Stillich,2

gich.5
Observations
by Sturtevant,6Henkel 7 and Hills 8 upon the
effect of fatigueon the yieldand compositionof milk are also
in accord with the results of experimentsupon
work
and exercise
of
milk
the quantity
in showing a tendency to reduce
and

the

at

yieldof

actual

from

Aside

time

same

by

the

cows

of their milk

both

the percentage and

the

fat.

clear that

appears

to increase

and

questionof
a

considerable

without

with

an

the

effects of

amount

overexertion,it

of work

any serious diminution

actual increase in the

may

be performed

of the volume

yieldof fat,its

in Morgen's
lightestwork
ingredient.The
experimentswas roughlyequivalentto hauling a load of a ton
much
more
if miles over a smooth level road. This is certainly
labor than the ordinarycow
will perform when
turned loose in
a comfortable
yard or paddock.
most

valuable

2
Jahresber. Agr. Chem., 39 (1897),529.
Ztg., 25 (1896), 331.
Expt. Sta. Rec, 10 (1899), 85.
Agr. Chem., 28 (1899), 492;
4
5
Jahresber. Tier Chem., 33 (1904),382.
Expt. Sta. Rec, 12 (1901), 381.
6N. Y. (Geneva) Expt. Sta.,Rpt. 1882, p. 25.
7 Landw.
Vers. Stat.,46 (1896), 329.
8 Vt.
Expt. Sta.,Rpts. 1894, p. 162, 1898, p. 367 and 1899, p. 309.
1

Milch

Centbl.

483

PRODUCTION

MILK

for all muscular

that the energy

It is still true, of course,
exertion is ultimately supplied by the feed.
the

just mentioned
be

required for this

feed

extra

the instance

In

purpose

may
contained in

the basis of the data

approximatelyestimated,on

matter
Chapter XIV, at two-thirds of a pound of digestible
per
of
of
The
about
maize.
to
feed
a
eight-tenths
pound
day, equal
of
the
exercise
taken
turned out in the
cost
ordinarily
by cows
and
be far outweighed by the tonic
yard must be insignificant
effects of fresh air,sunshine and freedom
their health and
on
some
general condition,while in the case of heavily fed cows
be
of
exercise may
ency
possibly
advantage in diminishing the tendThe
to fatten.
for
question of turning out dairy cows
exercise,then, virtuallyreduces itself to the question whether
of the labor involved is repaid by the effect upon
the cost

the health

of the animals.

Temperature. Shelter
579.

the

Air

temperature.

relations between

The

"

general principlesregarding

external

heat

temperature,

production
(350-356),

supply, already discussed in Chapters VII

VIII
(395-397)and XII (535-543),apply also to the dairycow.
in full flow of milk is
Like the beef steer, the well-fed dairy cow
of feed above
her maintenance
ration
consuming a largeexcess
and is producing a correspondingly
of heat.
largeamount
feed

and

example, in
heat production of
For

the estimated

and
on

ration

maintenance

Table

127.

experiment reportedby Jordan 1 the computed

two
cows
(disregarding
slightchanges in weight)
would
have been produced
of heat which
amount
an

"

were

as

Estimated

Heat

follows

775

Computed
Estimated

heat

production
heat production
.

maintenance

No.

1200

18.67 Therms

10.10

heat

The

Cow

10

Lb.

Therms

production was greater in one

in the other by 54 per cent than the estimated
than
which is a considerablygreater excess
Kellner's fatteningsteers.
The

Cows

of

No.

12

Lb.

21.10

Therms

13-7o

Therms

on

ration
.

"

Production
Cow

Weight

Feeding of Animals, The

Macmillan

case

by 85

amount

that

Co., New

on

per

cent

and

maintenance,

computed (537) for

York, 1908,

p. 310.

484

FARM

OF

NUTRITION

ANIMALS

of

maintenance

body temperature goes, then,

paratively
might not be subjected to comreason
no
why a cow
appears
without
low
causing any increased
temperatures
katabolism
the
for the sake of heat production solely. That
factors of size and weight,humidity of air,and the amount
same
far

So

and

as

mere

of ration

character

question is
580.

in the

as

case

of the steer

obvious.

Shelter,

etc.

The

"

question of

shelter

does

with the cow

The
and with the steer.
principle
wind, insolation and temperature
precipitation,
the same
the cow
water
are
on
as
on
qualitatively
same

apply in

render

which

reasons

differ

not

influence

in

the

into the

enter

of

of

drinking
and

the steer

shelter desirable in the

one

caze

the other.
The

foregoingfacts,however, are
scarcelysufficient to justifythe conclusion that a dairy cow
be treated in this respect like a beef steer.
In making a
may
quantitative application of these facts in practice,certain
modifying factors requireconsideration.
Relative body surface. Even
casual
the most
comparison
with the beef steer is sufficient to show
that
of the dairy cow
in form and to raise the supposition
that
they differ materially
the ratio of body surface to weight may
vary considerably in
581.

Modifying factors.

"

"

the
of

two

surface

body

that

The

types.

the

spare

of

writer is not

cows

but

can

angular form
surface

more
relatively

one

than

of any
hardlyavoid

aware

of the

measurements

the

impression

typicaldairy cow

the compact,

exposes

form

rounded

of the

beef animal.
Condition.

"

Outdoor

winter

largelywith fatteninganimals

experiments have been

considerable coveringof

feeding of
and

conducted.

it is with
With

practiced

cattle is
them

such

that most

an

animal

fat is

usuallyacquired before the onset

of extreme
cold weather, while the typical
dairycow devotes her
feed to milk production and carries very little body fat. There
are

definite data

no

but
that

doubtless
it

Skin
thinner

it is

might
and
than

have

hair.

"

that

as

to

the

protectivevalue

of

fat

covering

of heat and it would

seem
poor conductor
considerable
influence in reducing radiation.

The
of the

skin

of the
and

dairy cow

is

reputed to

be

therefore be a better
may
The coat of hair of the cow, too, is apt to be
radiator of heat.
shorter and lighterthan that of the steer, whether
a result
as
steer

MILK

PRODUCTION

485

breeding or of continuous shelter and warm

quarters, and is
that extent
a poorer
protectionagainstloss of heat.

of
to

all these reasons,

dairy cow may well be

it is clear that the loss of heat from

For

like

weight under
the

former

The

limit of external

katabolism

direction

consideration
and

is caused

of

in connection

with

the

is that
dairy cows
direction of production.

the

and

steer

that

be

of

sequently
con-

below

temperature

may

production.

shelter for

upon

conditions

the

higher for the

for the latter.

than

582.

external

same

minimum

additional

which

the

rapid than that from

more

the

of

Another

"

important

question of temperature
their possible influence

Stress

laid

was

at

the outset

of this discussion of the factors of milk

production (558)upon
beef production and milk production

the essential difference between

due
of

the fact that in the latter it is

to

singlegland and

simply the

tion
secre-

general increase of the whole

ever,
body which is desired. The activityof the milk gland, howis much
sensitive to external influences than, for
more
fore,
example, that of adipose tissue. It is quiteconceivable,therethat a degree of cold or exposure which, from the standpoint
of heat production merely, might not
requireany additional
a

katabolism

maintain

not

the

less
body temperature, might neverthethe formation of milk, especially
if the cow
jected
subwere
to it suddenly. In such
it would be anticipated,
a case
either that feed previouslyused for milk productionwould be
stored up as body fat or else,if the cow
continued to eat the
same
amount, would lead to a stimulation of the generalbody
katabolism and so to an
unnecessary increase in heat production.
In other words, exposure
to cold might conceivablyneither
to

check

increase the feed

consumption nor diminish the total utilization

of surplus feed but might, nevertheless,
be a disadvantage because
it diverted the current
of productive activities from the
formation
583.

of milk

Results

in

is available

of temperature for
Plumb

to

other and

practice.
"

undesired

Only

of

production.
dence
experimentalevi-

meager

regarding the practicableor desirable limits

dairycows.

compared the feed consumption

lots of purchased cows,

about one
hour per day
1

forms

one
on

of which
sunny

was

days

and

turned

while

Ind. Expt. Sta.,Bui. 47 (1893),pp.

the

yield of

milk
out

into

other

89-96.

was

the

two

yard

turned

486

NUTRITION

shelter from

some

The

exposed

lot ate

the sheltered lot and

comparabilityof

the

Brooks

stable kept
which

550 F.

at

and

lots of
a

its percentage of fat

2

Jordan

and

more

milk

more

ad

No

with

libitum.

hay

than

proof of

heated
artificially

the temperature of
produced in the warm

one,
was

lower.

was

recorded

an

but

less

milk.

between

cows

feed

somewhat

cooler,unheated

Rather

reported.

Richards

consumed

cows

more

two

the weather

regard to

grain but
produced 161.1 pounds
the two
lots is given.

exchanged

is not

stable but

much

ANIMALS

FARM

every day without

The
the wind.

eighthours

for

out

OF

the milk

yieldof

number

of

cows

periodsin which the stable temperature

at about
450 and 550 F. More
maintained, respectively,
was
of
four and more
milk was
butter fat in
out
produced in three cases
of
four
at the higher temperature.
two
out
cases
3
Spier reports experimentsat four farms, on a total of 88 animals
and ventilation to milk yield.
the relation of stable temperature
upon
upon

He

in alternate

feed

uniform

calls attention

to

the fact that both

these

the average
the average
consumed

of

warmer

results for

there is

128.

Influence

"

of
on

record

Ventilation
Milk

of actual

Mass.

Wis.

feed

consumed

weights of the animals.

and

Stable

Temperature

Production

(Hatch) Expt. Sta.,Rpt. 1895,

Expt. Sta., 21st Rpt., 1903-1904,
Jour, of Highland and Agr. Soc, 1909,

no

of the live

during the several periodsnor

Table

involved

in

followingtable shows
periods as compared with
preceding and following periods and likewise
rations
the entire experiments. The average

stated but

are

are

The

upon the influence of shelter.

in two
the results obtained
speciallycold

experiments

factors

p. 39.
p. 143.
pp.

255-306.

488

NUTRITION

the whole

On

OF

agree with
indicate that the

deductions

the
need

for

been

overemphasized,but

limits

within

Much

doubtless
of

treatment

have

likelyto

insufficient

are

effect

more

to it

accustomed

and

coat

such

on

animals

the weather

as

low

the

has
the

yield.
previous

carry

temperature

grows

and

affect

not

than

on

establish

to

does

temperature

winter

are

data
physiological
quarters for dairy cows

depends,as Spierpointsout, upon

the cows.
Warmly stabled animals

than

rather

experiments as

from

warm

stable

which

ANIMALS

be said that such

it may

record

FARM

seems

those

on

mer
sum-

ually
grad-

colder.

kept in the stable most of the time the question

of temperature is of special
interest in its relation to ventilation.
stable must
in most
be warmed
a cow
cases
Practically,
simply by the heat derived from the animals themselves and
of more
be obtained
a high temperature can
or
only by means
Where

cows

are

less restricted ventilation.

perfectventilation,with

more

vigorof

health and

" 4.

the

584.

Meaning

of

identical with

(470).

discussed
the milk

milk
the

is the

conceptionof
here

of

ratio

liberal

is substantially

growth already

protein contained in
protein which is required

favorable

Surplus protein katabolized.

production requiresa

considered

tion
the utiliza-

the

of feed

the most

conditions.
it is evident

While

"

supplyof proteinin

the

actuallysecreted in the milk is determined

of the animal, precisely
as
by the individuality

amount

storage of protein in the

the form

increase

of milk

it appears

While

at

the

Production

of its utilization in

to the least amount

possibleto

used,

of protein
The

"

Milk

in

productionas
that

It

produce it under
585.

Feed

utilization

milk

be

can

its beneficial effects upon

of

of utilization.

proteinin

temperatures

animals,is possible.

Utilization

The

The

to

If low

of the

case

will the

growing animal.

amount

of

that

the

ration,

marily
priis the

It is not

proteinsecreted

the supply of proteinin

by increasing
be true

that

of
activity

in

the feed.

the milk

glands
can
proteinsupply
by an
(599) it is nevertheless true that the animal producesan amount
of milk determined
essentially
by its capacity and any surplus
be

to

stimulated

somewhat

abundant

of

protein over

that

necessary

for this purpose

is katabolized

MILK

is the

justas

for that
a

This

with

case

matter

to

supply of

surplussuppliedto

mature

material

489

PRODUCTION

animal.
and

not

Feed
cause

animal, or

young

tially
protein is substanof production.

is

illustrated in experiments by Jordan x in which

strikingly
the protein supply of two cows, beginning with a liberal amount,
was
one-half and then graduallyincreased
to about
graduallydiminished
again to the originalquantity. The followingtable shows the average
nitrogenbalances of Cow No. 12 of the second series of experiments,
the dailyresults being grouped into periodsas indicated.
Table

130.

"

Average

Daily

Nitrogen

Balance

of

Cows

Gain
Body

by

Grams

Jan. 30-Feb.
Feb. 6-Feb.
16-Feb.

Feb.

26-Mar.

Mar.

8-Mar.
1

Mar.

17-9

16.3
2.2

0.3

18

6.9

26

8-Mar.

16

Feb.

Mar.

28

12.2

-f- 10.6

8-

Apr. 7
Apr. 7-Apr. 14

9-4

in
yieldsdecreased in quite a normal way with the advance
like that of total milk solids,
ing,
diminishthe yieldof protein,
lactation,
the
about
while the percentage of proteinin the latter remained
of
the
middle
there
rations
On the low protein
seems
periods
same.
the
of
milk
o
ff
in
have
been
amount
to
some
proteinproduced
falling
have
been
in comparison with what
might
expected on an unchanged
in
is
small
but
the
difference
two
or
one
ration,
periodsin which
except
Aside
from
the protein supply reached the lowest limit.
this,the
of digestible
protein
principaleffect of the variations in the amount
diminish
the
of
excreted
increase
to
amount
or
was
nitrogen
supplied
and fell with the
in the urine, which, as the table clearlyshows, rose
supply of nitrogenin the food.
The

586.

Estimates

of

utilization

of

protein.
"

In

attempting
feed protein

regarding the utilization of

for the productionof milk protein,then, it is evidentlynecessary
of protein in the ration,since such an excess
to avoid an
excess
to

reach

conclusions

N. Y.

(Geneva) Expt. Sta., Buls.

132

(1897) and

197

(1901).

is

OF

NUTRITION

490

FARM

subject to rapid katabolism, so

ANIMALS

that

high proteinrations will

show a low apparent utilization of the proteinfor

necessarily
milk justas they do for growth (468)
On the other hand, too
small a supply of proteinmay
the tissue proteins of the
cause
of milk proteinso
body to be mobilized and utilized as a source
that a direct comparison of feed proteinand milk proteinwould
give too high a result. To determine the utilization of feed
it is necessary, while maintaininga sufficient
protein,therefore,
of the ration
supply, to reduce the protein content
energy
as
nearlyas possibleto that which is just sufficient to prevent
loss of body proteinand
then to compare
the feed protein
a
minus the maintenance
requirement with the milk protein.
of
Such
an
experiment obviously requires a determination
the nitrogenbalance of the animal, and
few of the
relatively
reported investigationson milk production include such a
determination, while in none
yet reportedhas the sufficiency
of the energy
of respiraby means
supply been demonstrated
tion
There
inconsiderable
experiments.
are, however, a not
number
of experimentson
record in which the live weights of
.

the
of

been

have

animals

well

protein but
digestible

milk

plus those

estimated

maintained

and

little greater than

to be necessary

in which
those

found

for maintenance

been
of

amounts

adequate for the productionof at least moderate

milk without drawing on the body protein.

in the
have

amounts

Naturally an exact balance of the income and outgo of nitrogen

will rarelybe secured.
In most
it is necessary
the
to compare
cases
feed protein with the algebraicsum
of the milk proteinand the gain
loss of body protein, the comparison being more
or
nearly correct as
the latter factor becomes

Table
of

131
number

shows
of

smaller.

the

low

computed utilization of the protein

protein rations,the daily maintenance
protein being estimated as 0.6 pound per

requirement of crude
1000
pounds live weight in direct proportion to the latter. It
includes the experiments by Jordan upon
of milk
the sources
fat,the results of one of which as regardsproteinhave justbeen
cited,an experiment by Hayward x the results of which
as
the
tensive
regards
nitrogen balance are still unpublished, the exexperiments upon the minimum
protein requirements
1

Penna.

Expt. Sta.,Rpt.

1901-1902,

pp.

314

to

396.

MILK

Table

131.

"

Utilization

PRODUCTION

of

Protein

491

in

Milk

Production

FARM

OF

NUTRITION

492

ANIMALS

Laboratory for Agricultural

Research
in Copenhagen l and unpublishedrespiration
ments
experiby Kellner.2 The experiments of Hart and Humphrey
of

dairy

carried

cows

in the next

mentioned

on

the

at

paragraph,when

computed

in the

same

high percentage utilization of the digested

protein,although the gains and losses of body proteinare
disturb
the
to
as
comparison.
relativelyso considerable
also show

way,

Haecker's

low

proteinrations

in

as

1902-3-4-5,
to afford another

noted

on

example of the
subsequent page (602),seem
high utilization of feed protein.
While too much
weight should not be attached to the results
since they include
of comparisonslike the foregoing,especially
estimate
of the protein requirement
less uncertain
a
more
or
to indicate beyond
for maintenance, they nevertheless seem
reasonable

doubt

that

low

on

feeding stuffs may

without any very largeloss.
least

some

587.

of

values

Relative

proteinrations
be

converted

proteins for

the
into

milk

proteinof at
milk protein

production.
"

The

ing
precedingchapters(400,465) regardfor maintenance
and
the relative values of different proteins
for productionrender it altogether
probablethat they also differ
of milk protein. No
in value as sources
experiments on this
but Hart
and Humphrey 3 in
point have as yet been reported,
have compared the mixed
series of experimentson
two
cows
proteinsof maize, wheat, gluten feed, oil meal and distillers'
using maize
grains with proteins prepared from milk (784),
and silageas roughage. They found the average
stover
centage
perconsiderations advanced

in

recovered in the
nitrogen which was
gain (or minus the loss)of the body protein

of the resorbed
milk
to

yieldplus the

be
Skim

milk

powder

59

Casein
Maize

Wheat
Gluten

feed

Oil meal

Distillers'grains
1

Denmark-Beretning

for landokonomiske

"

59%
40%
36%
45%
61%
60%

Kgl. Veterinear of Landbohojskoles Laboratorium

Translated
6ode, 1906, and 63de, 1907, Kobenhavn.

fra den

Forsog.

Rationale
du Betail.
by Mallevre, Society de l'Alimentation
Compte Rendu
et i2eme
ueme
Congres.
2 Die
Ernahrung der landw. Nutztiere, 6th Ed., 1912, p. 551.
3
Jour. Biol. Chem., 21 (1915), 239; 26 (1916), 457.

de

MILK

the

If

493

probable requirement for protein maintenance

from

deducted

PRODUCTION

the

total

nitrogen,the

resorbed

be

utilization of

remaining protein,calculated as in the experiments of the

ing
previousparagraph, was notablyhigher,approachingor reachthe

100

cent

per

in several

instances.

differences observed

The

tuations
largelydue, however,to flucbody protein,the formation of

were

in the

milk

gain or loss of
protein being quite uniform

from

period, and
this fact seems
the results of somewhat
to render
questionable
relevance as regards the specialquestionof comparative values
of milk protein, although they do show
marked
as
sources
period

to

differences in total efficiency.

The
588.

utilizationof energy

for milk
The
values
net
production.
energy
value of a feedingstuff or ration for milk production is
Net

energy
identical

"

in

conception with

that

for

fattening(448) or

for

It is that part of the feed

growth (472)already considered.
of the maintenance
requirement which
energy suppliedin excess
is recovered in the product. For example, if a cow
produces
of
four
lb.
milk, containing(604) 336
per day 20
per cent
Cals. of energy per pound, the total of 6720 Cals. would
be the
dition
be supplied in the ration in advalue which must
net energy
to that requiredfor maintenance.
As pointed out in Chapter VIII
(371),it cannot be assumed
that the net energy values for maintenance,fattening
or growth
apply to milk production,but the values for the latter purpose
be determined by direct experiment. As yet, very scanty
must
data are
available on this point, the only results yet reported

being three
589.

contained

Complete

nitrogen, carbon
on

in

brief

by

preliminary
paper
l

Kellner
balances.
energy
and energy balances,
determined

oxen,

of three

cows

Kellner.

reports

"

as

in his

the
periments
ex-

rations and

receivingmixed

in their milk yield. By the method

varying considerably
in Chapter XVII
it is estimated that the
(768-772),

scribed
denet

energy values of the rations and the percentage utilization of

their metabolizable energy for fattening
would be :
"

5terInternat. Kongress

fur

Milchwirtschaft,
191

1.

Estimating
from
the

their

maintenance

weights on
of

oxen

ANIMALS

requirements of the animals

maintenance

the

live

FARM

OF

NUTRITION

494

results on
of his average
(381),Kellner obtains the following
basis

the

showing a considerablyhigher utilization for

energy
milk productionthan that computed for fattening.
balances

Table

132.

"

Energy

li Kilogram starch value

Balances

of

2.356 Therms

Dairy

net

Cows

energy.

496
*

has

farm

of

nutrition

likewise

animals

determined

partialenergy balances of
for an entire year on rations just sufficient to
ten milking cows
their live weight. In these experiments the percentage
maintain
of the rations is computed for eightof the cows
on
digestibility
Eckles

the

basis

of results

animals,while
was

were

dried

digestiontrials

five of

on

the

metabolizable

of the

energy

rations

assumption of

Table

gain

no

loss

or

by

135.

"

Percentage

Utilization

3.7

puting
com-

the

body, the followingvalues

productionare obtained.

the

percentage utilization in milk

Milk

of

Energy

Metabolizable

in

Production

Cow

No.

206

Cow

No.

304

Cow

No.

400

63.36 %
67.60%
66.90%

Cow

No.

43

5J-36%

Cow

No.

62

Cow

No.

Cow

No.

27

72.82%
60.24%
62.89%

Cow

No.

63

5o-35%

Average

61.94%

Haecker,2 in discussingthe
with

at

(753)and
organicmatter
per kilogram of digestible
the results,
exactly as in Jordan's experiments,on

for the

the

requirement of all but one of the

live weight experiments after the cows
results reportedin Chapter VIII
(381).

in

off,with

Estimating the
Therms

in

the maintenance

determined

cows

obtained

the

results of extensive

experiments

dairyherd
Experiment Station,has
nutrients
of the feed and the solids of
compared the digestible
the milk by reducing both to their carbohydrate equivalent.3
Subtractingthe estimated maintenance
requirementfrom the
total carbohydrateequivalent (" nutriment
") of the feed, he
finds
cent

that of the remainder

was

years

recovered

in the

being 54.65 per cent,

in

were

of the Minnesota

lower

generalmaintained.

utilization of energy

Jordan'sand
1

Mo.

Minn.

The

Eckles'

from

milk,
while

the

This

seems

than

that

experiments.

Expt. Sta., Research

Bui. 7.
Expt. Sta., Bui. 140 (1914),p.
fat of the feed

to 66.22
50.25 per cent
per
for nine
the general average

It

the product added

to the carbohydrates and
called " nutriment," are, of course,
approximately
and

the metabolizable

weights

to indicate

of the

cows

decidedly
computed in Kellner's,
be noted, however,
must
a

45.

is multiplied by the factor

and

of the milk

live

energy

2.2

and

that of the milk

protein. The
proportional

of the feed

sums,
to the

by 2.25

which
energy

respectively.

are
tent
con-

MILK

PRODUCTION

497

of the rations in Haecker's

digestibility
experiments
from
estimated
was
figureswhich, according to Eckles'
average
in milk, although
results (722),are probably too high for cows

that the

the other hand

on

also
591.

Net

"

values

for

comparison

by the

for the maintenance

estimate

quirement
re-

high.

seems

energy

fattening.
obtained

Haecker's

probably greater

of Kellner's
l

author

same

milk

and

the utilization of metabolizable

results

for

(589)with those
and

by Armsby

in either

energy

than

Fries

for

maintenance,

to indicate
fattening seems
clearlythat the net
for milk production are
values
distinctly
higher than
energy
those for the latter purposes, although no direct comparisons
the same
on
feedingstuff or ration can be made;
Both Jordan's and Eckles' results tend to confirm this conclusion,
which
is further strengthened by the fact, to which
Eckles calls attention,that with one
exceptionthe actual energy
of the milk in his experimentswas
content
greater than the net

growth

value

energy

by

or

the

available in the ration

of Kellner's

use

Unfortunately, no
singlefeeding stuffs
been

The

Cause

or

of

higher

net

energy

apparently higher net

compared

as

in

shown

in

values

energy

with tissue

the difference

to

As

the net energy

values
upon
nutrients for milk productionhave

results

impossibleat present
quantitativecomparisons.

592.

computed

as

factors.

that it is

so
reported,

exact

"

producingit

for milk

and

be

compositionof
the

yet
any

production.

values for milk

production may

the

make

to

of

tion
produc-

cribed
plausiblyas-

the

products.
crease
of the in-

Chapters
XI,
consists
in
fattening
chieflyof fat (441-443)and even
the case of growth fat makes up a considerable proportion
of it
(458)except in extreme
youth. In average milk, on the comtrary,proteinand milk sugar constitute two-thirds of the total
organicmatter and carry over one-half of the total energy.
X

organic matter

in

It

not

seems

protein into

milk

milk

may

sugar,

of energy

as
or

1
2

improbable that

the

conversion

of

digestible
protein,or of digestiblecarbohydratesinto
involve a comparativelysmall expenditure

hydrates
compared with the synthesisof fat from carboprotein. If such be the case, the organicmatter
i.
Sta., 53 (iqoo),
Jour. Agr. Research, 3 (1915), 435;

Landw.

Vers.

7 (1916), 379.

498

NUTRITION

of the

milk

energy
would

of the

the increase

matter

could

593.

would

retain

of

ANIMALS

of the

larger percentage

from

matter
digestible

tissue

body

which

it

which

formed

was

that

chemical
than

digestible

same

produce.

Computation
it be

Let

"

FARM

OF

of

equivalent

assumed

carbohydratesof the feed

compounds of milk

values

for fattening.

net

energy

the

digestibleprotein and

that
be
may
without

converted
loss and

into
that

the

sponding
corre-

the

expenditure
carbohydrates
is the same
that observed
as
by Kellner (769)for the production
of body fat.
Then
each gram
of protein or carbohydrates in
the milk would
requirethe supply in the feed of one gram of
digestibleprotein or carbohydrates respectively,while each
of energy

in the

of milk

gram

production of milk

fat if manufactured

fat from

from

carbohydrateswould

requireabout 3.9 grams of the latter.

The
of energy
in milk
recovered
corresponding amounts
production and in fatteningrespectivelywould, according to
the

foregoingassumptions,be

Table

136.

Computed

"

Energy
in

Supplied

1
1

Feed

in

follows

protein

gram

carbohydrates
carbohydrates

gram

gram

Recovered

"

Milk

in

Production

and

Fattening

Produced

gram

3.9 grams

as

Energy
Recovered
Milk
in

Milk

in

ing
l

Cals.

5.7

Cals.

2.24

carbohydrates

4.1

Cals.

2.37 Cals.

fat

9.23

Cals.

9.23

protein

gram

Energy
Recovered
Fattenin

Cals.

approximatelythe amount
be requiredfor the
would
of net energy
for fatteningwhich
of milk of known
production of a given amount
composition.
Thus
four
Table
cent
to
milk, according
144 (604),
average
per
of
contains
drates
carbohy3.08 per cent of protein,4.85 per cent
this

On

and

contained
the

amount

the

same

basis,it is easy

4.0

in

of
feed

per

cent

pound

to

compute

The

of fat.

of

such

milk

actual
would

have
which
would
energy
if used for fatteningwould

Cals.

of

net

the

Conversely, an amount
as
computed from
energy
1

Kellner's

amount

be
been
have

of

energy

336 Cals.,while
recovered
been

from

only

252

containing 252 Cals. of

results of fattening experi-

feed

factors.

MILK

ments

would

energy

in four

PRODUCTION

suffice to

shown

in the

Table

137.

support

the

The

milk.

cent

per

499

storage of 336 Cals. of

method

of

computation is

following table.
Recovered

Energy

"

Four

in

Cent

Per

Milk

and

in

Fattening

Energy

Protein

Carbohydrates
Fat
...

Total

per

100

Total

per

pound

is afforded

(589).

In

4.1

3-08
4.85

9.23

4.00

2.24

3.08

19.8 Cals.

2.37

4.85

9.23

4.00

36.9 Cals.

Cals.

6.9 Cals.
1

1-5 Cals.

36.9 Cals.
Cals.

55.3

Cals.

Cals.

252

Confirmation

"

Energy
covered
ReFattening

17.5 Cals.

336

of

this

hypothesis

by the results of Kellner's respirationexperiments

putes
substantiallythe way just outlined, Kellner comthe actual chemical

by his cow
only 10.367 Therms
a

in

74.2

results.

Equivalent

in

that while

that

5.7

grams

Kellner's

594.

Recovered
Milk

ration

was

of

13.907
net

energy

support the observed

and

of the milk

solids produced

Therms, this was

equivalentto
value for fattening,
and therefore,

supplying this

for maintenance

energy

amount

body gain

in

excess

should

of that

quired
re-

be sufficient to

milk

for
production. For the three cows
which
results are reported,the requirements for net energy
as
thus computed compared with the estimated net energy values

of the rations
Table

138.

"

were

Net

as

Energy

follows

Values

"

Fattening

for

in

Kellner's

periments
Ex-

Cow

Therms

Total

in ration

Required

11.403

for maintenance

4.806

6-597
Required

.928

for

body gain
Available for milk production
Computed requirement for milk

5-669

tion
produc,

6.079

The

of net

FARM

ANIMALS

actuallyavailable for milk production

correspondquite closelywith the amounts
computed
requiredaccording to the foregoingassumptions, and
amounts

be

to

OF

'NUTRITION

500

Kellner

states

number

of

that

energy

this

also

was

the

in

case

considerable

his

unpublished experiments, although in others,

especiallythose in which a surplusof feed was given, the agreement
far from being so good, the difference in one
was
case

reaching24

per cent.

Quite in harmony
paragraphs is
"

in milk

when

the

is the

equal

generalconclusions
by

statement

Eckles

of the

going
fore-

that in his

periments
ex-

of energy in the feed produced more

energy
the per cent of fat was
low than when
it was
high.
efficient when
of feed is more
used
given amount
medium

production of

production of

low

to

fat is

other

in fat.

greater

tax

constituents

from
It appears
the animal
upon
of

the

milk

this
than

carrying

value."

energy

"
595.

the

the

therm

Apparently a
to produce milk
that

with

5. Feeding

Feeding

Milk

for

secondary

factor.

Production
"

As

has

already been

a
urged, the feedingof a milking animal is in a certain sense
of successful
secondary factor in dairying. The possibilities
milk
the capacity of the
production depend primarilyupon
the maintenance
animals as milk producers and upon
of such
environment
will give free play to this capacity. Feed,
an
as
the other hand, while equallynecessary, is after all essentially
on

the

supply of

works

and

raw

cannot

material

upon

which

the

animal

mechanism

greatlystimulate production,though

it may

limit it for lack of material.

The

thing is substantially
true, of course, of all forms
of productivefeeding,
in the feeding
the case
but it is especially
for the reason
of dairy animals
alreadynoted (558),that it is
the product of a single
gland and not a generalincrease of body
tissue which
is desired.
in
Improper rations,therefore,may
this case
if sufficient
not
only limit the total productionbut, even
in quantity,may
if deficient in quality deflect production
from milk to fattening,
or possiblyto greater muscular
activity,
and

same

thus fail to utilize fullythe

1

milk-producingcapacityof

Loc. cit.,
p. 137.

the

MILK

PRODUCTION

501

a
Feeding, therefore,while in a sense
secondary
factor is nevertheless an important one.
Feed
596.
of
Regarded solely as a source
requirements.
of milk, the daily ration must, of
material for the formation
contain
of protein and ash and
an
adequate amount
course,
a
quantity of non-nitrogenous nutrients sufficient to furnish
material for the manufacture
of the non-nitrogenous ingredients
of the milk, while it must
also supply enough energy for
activities of the body, includingmaintenance
the physiological
and the energy
expended in the processes of milk formation.
be taken into conIn addition to this,however, there must
sideration

animals.

"

the
of substances

of
possibility
which

the presence
absence in the feed
or
have
a
specificeffect on the milk

may

by stimulatingor depressingits action as a whole

the character of its action and so
or
by affectingqualitatively
the compositionof the milk.
There is,of course, a possibility
of such specific
of production,but it is
effects in other forms
in milk production for evident reasons
obvious
most
and has

gland,either

been

studied

most

in that connection.

Protein
Milk

597.

milk

requirements
for milk production

rich

in

productionis,of

The

of
physiological
purpose
to support the growth of the young.
(462),is the progrowth, however
duction

protein.
"

course,

essential feature

of

The

protein tissue,which, in the sucklinganimal,

is relatively
rapid,and in order to support this growth the milk
less proportional
to
contain proteinin amount
must
more
or
milk
is decidedly
the rate of growth of the species. Cow's
protein in character,the ratio of protein to non-nitrogenous
ingredientscorrespondingroughly with that of the increase
made
old (556) Moreover, in the
by an animal three months
of the cow, man
has been able to increase greatlythe natural
case
crease
milk-producingcapacity,with, of course, a correspondinginof

new

in the total amount

moderate

over

0.6

yieldsof champion
598.

Minimum

protein formed.

Even

the

tion
pounds of milk of average composipound of protein,while the extraordinary

dailyyieldof

contains

of milk

cows

20

contain

several

protein requirement.

growth, it is evident that the least

"

amount

times

this amount.

Just as
of

in the

case

of

protein
digestible

OF

NUTRITION

502
which

is the

animal

requirements of the milk-producing

of the body
quantity requiredfor the maintenance
the

possiblymeet

can

ANIMALS

FARM

in the milk
of proteincontained
proteinplus the actual amount
yielded. For example,if a iooo-pound cow is to produce daily
containing3.2 per cent of total protein,
25 pounds of milk

digestibleform
in the milk plus
at least the 0.8 pound of proteincontained
the approximate 0.6 pound presumably required for body
maintenance, or a total of approximately 1.4 pounds. A less
evidently result either in a falling
supply than this must
it is evident

milk

off in the
milk

in

yieldor

in

contain

must

body proteininto

of

conversion

protein.
much

How

by

an

utilization

"4

ration

her

that

of

this

this minimum

than

more

adequate

ration will

amount

depend

the

upon

plied
sup-

percentage

already discussed

protein in the sense

chapter (584-586),i.e.,upon
feed

of the

be

must

in

proportionof

the

it

tion
capableof conversion into milk protein. Thus in the illustrajustemployed, if 80 per cent of the surplusfeed protein
be utilized the proteinrequirement would be 1 .0 pound for
can
milk productionplus 0.6 pound for maintenance, or 1.6 pound
with
is parallel
that of the
instead of 1.4 pound. The
case
proteinrequirement for growth discussed in Chapter XI (484491) and in both instances the experimentaldata available are
insufficient for a final conclusion,although the probabilities
of a high percentage utilization
to indicate the possibility
appear
under
599.

favorable
Protein

conditions.
as

stimulus

the

to

milk

glands.

"

The

going
fore-

subject. In
of milk protein
that the amount
them it has been tacitly
assumed
fixed.
It
manufactured
by the milk glands is substantially
well established,
seems
however, that in addition to furnishing
the nitrogenous
material
for the manufacture
of milk protein,
considerations

matter

do

of the feed may

glands,causinga
of all the milk

more

solids.

not, however, exhaust

act

to

active
In

extent

some

secretion

other

full
to
to

not

words, it

greater or less surplus of proteinover

calculations

as

the

stimulus

to the

only of proteinbut
would

the amount

appear

that

indicated

by

foregoingis necessary if it is desired to take

of the milk-producing capacity of the animal or

like the

advantage

delay as much as possiblethe

advancing lactation.

natural

shrinkage in milk due

OF

NUTRITION

5"4
Table

139.

Influence

"

Protein

of

ANIMALS

FARM

(Results per day

Supply
and

on

Production

Milk

head)

siderable
proteinrations were able to support a conmilk productionwithout
causingthe body proteinto
liberal supply
be drawn
nevertheless,a more
materially,
upon
of digestible
accompanied by a distinctly
greater
proteinwas
productionof both total milk solids and milk protein.

Although

the low

extensive
investigationsof
gen's investigations. The
l
milk
production by sheep
Morgen and his associates
upon
of trials in which an exchange between
include a large number
M

or

"

comparativelypure proteinon
the

on

afford

other

was

factors.

in the

one

rations.

hand
The

and

starch

or

oil

results,therefore,

regarding the influence of the protein

distinguishedfrom the possibleeffects of associated
the ration of the low protein
In nearly all instances

valuable

supply as

made

the

data

protein
surplusof digestible
above
the total of milk protein plus maintenance
protein. In
the followingtable,computed by the writer,the experiments

period

Landw.

contained

Vers.

considerable

Stat.,61 (1904),1

; 62

(1905),251

; 64

(1906), 93;

66

(1907), 63.

MILK

have

been

PRODUCTION

grouped accordingto

505

the amount

of this

surplus,and

the average
percentage increase in the yield of milk solids and
of milk fat which resulted from an increase of the feed protein
has been
Table

computed

140.

On

for each

Influence

"

of

group.
Protein

Supply

Milk

on

Production

the

to furnish conwhole, Morgen's investigations

seem
clusive
evidence of a stimulating
milk
effect of protein on
duction.
proEven
when the proteinsupply alreadylargely
exceeded
minimum
demand, a further addition was in most instances

the

followed

by

usuallyin

that

distinct increase in the

of milk

fat.

It should

yieldof
be

milk

solids and

said,however, that

respectableminority of the individual experiments failed to

show this effect. Of the nineteen singletrials in which protein
substituted for fat, eleven showed
increased yield of
was
an
milk
1

solids and

six

Digestibleprotein

an

minus

increased

yieldof

milk

requirements for maintenance

fat.
and

for

Out

of the

growth of wool.

506

OF

NUTRITION

ANIMALS

FARM

substituted for carbohydrates,

thirty-onetrials in which protein was
twenty-sixshowed an increased yieldof milk solids
and twenty-one an increased yieldof milk fat. In thirteen out
of additional
of the entire fiftyexperiments,therefore,the presence
increase in the milk solids,
an
proteinfailed to cause
while in twenty-threetrials it failed to produce an increase of
milk fat.
Effect

600.

feeds.
protein-rich

of

In addition

"

to

tions
investiga-

paragraph,in which the effect

of an interchangeof practically
was
studied,a
pure nutrients
of experimentsare
record in which an
considerable number
on
of a ration in digestible
enrichment
proteinhas been effected
by an interchangeof feeding stuffs,as for example, by the
like those noted

of cottonseed

substitution
In
with

all these
one

in the last

two

or

meal

for maize

experimentsthe low proteinrations contained,

proteinabove
exceptions,a surplusof digestible

the milk

proteinplus the estimated

increase

of

of milk
from

maintenance, yet
followed

by
the
digested,

digestible
protein was
unit of organicmatter

per

cent

per

meal.

to

39

per

cent.

further

largeryield

increase ing
rangin the experiments

As

what
previousparagraph,the results appear somethe
showing no consistent relation between
capricious,
of proteinsuppliedand the relative increase of milk production

described

excess

in the

secured.
be

It should

low-proteinrations
of maize
extent
protein,which has
inferior nutritive value (783).
proteinsof

601.

the

Protein

fed

in

American

experimentsthe

of these

that in many

added

consisted

to

since been

shown

practice. On
"

considerable

the

to be

of

basis of

recommended
a standard
experimentsand observations,Wolff
for dairy feeding callingfor 2.5 pounds of digestible
protein
dailyper 1000 pounds of live weight. Although later modified
versally
unifor many
by Lehmann, this standard was
years almost
doubted
accepted on Wolff's authority,supported by the uninstances the addition of protein-rich
fact that in many
feedingstuffs to ordinaryfarm rations materiallyincreased the
milk
to indicate
yield. Later observations,however, seem
in practice
that while proteinis important the amount
necessary
has

been

somewhat
1

Die

overestimated.

Ernahrung

der landw.

Nutztiere, 1876,

p.

548.

PRODUCTION

MILK

Woll

extensive

an

regards the
successful dairymen were

United

the

practicein

make

first to

the

was

507

States

as

study of dairy
protein supply,

using rations
called for by Wolff's
was
the rations reportedas compared

rindingthat very many

supplyingmateriallyless proteinthan
The

standard.

as

follows

Digestible

Matter

with Wolff's standard

Table

of all

average

141.

was

"

"

Rations

Dairy

in

Woll's

Dry matter
Digestiblematter
Protein

24.51

Fat

Woll

ratio

points out that this

that

be

not

guide for practice,and

protein than the Wolff
profitable.
Somewhat
in

represent

production,expresses the
and althoughit does not
feeding experience,
that less proteinwould not be sufficient or

either
would

more

while it does not

average,

of milk
investigation

results of American
demonstrate

Lb.

i:5-9

scientific

any

0.74

Nutritive

Lb.

2.15 Lb.
13.27 Lb.

Carbohydrates
...

age
Aver-

similar

89 2-1 893 with

advantageous, it does afford a safe

indicates that rations containing less
standard

observations
additional

the

calls for

probably more

are

2
reported by Phelps

were

feature

in

that

several

stances
in-

subsequentlymodified at the suggestion

ration
the
and
the
of the experimenter
new
yield on
determined.
a supply of 1.9 to 2.5 pounds
Phelps recommends
digestible
proteinper head according to the productivenessof
the cow, the amount
to be based on the yieldof milk rather than
the rations fed

on

the live

economical
602.

were

weight, and

believes

productionthan

Experiments
observations

on

and

those

herds.

"

such

rations

will

Haecker

tensive
reported exprotein supply

has

the

(1892) and 38 (1894).

(Storrs)Expt. Sta.,Rpt. 1897, pp. 17-66.
Expt. Sta., Buls. 71, 79, and 140.

Wis. Expt. Sta., Buls. 33

Conn.

Minn.

more

containingless protein.

experiments on

give

5o8
of the

dairy herd

ANIMALS

FARM

OF

NUTRITION

Station, leading to the

of the Minnesota

standard

that the Wolff -Lehmann

calls for

clusion
con-

unnecessarily

of protein.
largeamounts
regarded as normal and
During nine years, yieldswhich were
either on the basis of total amounts
produced or
satisfactory,
secured on rations containing,
of feed consumed
per unit of milk, were
2
with the exception of the year
1 895-1896, about

pounds live weight.1

also made
During three of these years, comparisons were
of cows
between
a group
receivingabout 2 pounds of digestible
proteinper day and 1000 pounds live weight and one receiving
rations
about
1.5 pounds. In the earlier years the low protein
appeared as efficient as the higher ones, but toward the end of
deficient vitality,
showed
the three years the low proteingroup
apparentlyindicatinga lack of protein.
In all nine years, the (estimated)digestible
proteinin the
the
surplus over
high protein rations supplieda considerable
nance
Estimating the mainteprotein of milk plus maintenance.
requirement of protein at 0.7 per 1000, Haecker makes
the followingcomparisons: 2

pounds

of

proteinper
digestible

1000

"

Table

Bui.

142.

140,

"

Protein

p. 43.

Supply

of

Dairy

Herd

Ibid,, p.

54.

PRODUCTION

MILK

The

of the milk

proteincontent

509

from

the

low

protein groups

approximate estimate indicates that it

less than the surplus of feed protein
could not have been much
over
maintenance, thus furnishingfurther instances of an apparently
high percentage utilization of feed protein (586).
the indications
that such very low protein rations
While
are
clear that a surplus of 40 or 50 per
were
inadequate,it seems
of available protein over
in the milk was
that contained
cent
ample to support normal production.
the
Woll 1 has reporteda nine-year series of observations
on
dairy herd of the Wisconsin Station,the time being divided into
three periods of three years each, during the first and third of

is not

reported,but

which

the rations had

second

three years

consumed

pounds

an

it

was

nutritive ratio of 1:7, while during the

The
estimated
1 : 6.
digestibleprotein

day by

per

over
weighing slightly

cows

was

Average
Average

Table

143.

"

of
of

and

periodsA
period B

Surplus

1.76 pounds.
1.97

Available

of

pounds.

Protein

Available
Protein

Period

A,

Low

protein

"
.

Average

C, Low

protein

"
.

Average

f
Period

B, High protein

Average

in

Herd

Milk
Protein

Lb.

Lb.

I.03

0.72
0.80

1.41
1.

Period

1000

18

0.69

Rations

Surplus
Available
Protein

Per

Cent

43

76

_7"
63

1. 21

0.74

0.82

0.58

41

1. 11

o-73

52

1.07

0.76

41

1. 00

0.69

45

1-54
I.18

0.70

120

"-59

100

1.22

0-63

94

i-3i

0.64

105

Wis. Expt. Sta., Research

Bui. 13.

of

OF

NUTRITION

510

results for the entire year and

the whole
showed
a somewhat
on

likewise

The

economical

average

The

the

of the

milk

3.38 per

cent

Summary.

603.

individual

"

both

cows

smaller
is not

decidedly more
supply of protein.
mating
estibut
stated,

allowing 0.6 pound per 1000 for

approximate surplus of the available

requirement)

maintenance
winter

rations

was

yieldand

to

as

shown

as

of the nine years.

In view of the great differences

in each

143

rations

and

proteinmaintenance, the
protein minus
protein (digestible
the milk production in the
over
in Table

for the winter

greater and

production on

proteincontent

it at

ANIMALS

FARM

between

compositionof milk, it

is

figurecan express the proteinrequirement for

milk productionper day and head, but that it must
vary with

clear that

one

no

and

the amount

character

produced.

of the milk

ble
fairlywell established (586)that the digestibe
converted
rations
feed proteinof ordinary mixed
may

It appears

into

be

to

proteinwithout

milk
a

moderate

rate

any very great loss and

be
of milk productionmay

that

quently
conse-

maintained,

furnishinga comparativelysmall
the milk
protein plus the
protein over
surplus of digestible
least for

at

requirementfor
On
pure

the other

on
was

maintenance.

hand,

proteinsand

however

(599,600),both experimentswith

in which

those

an

increase

in the

tent
proteinconfeeds
protein-rich
of excess
protein

by the use of
to indicate clearlya stimulatinginfluence
milk production,
although in the majority of cases the effect
pated,
not very large. Contrary to what
might have been anticiproteinof the ration
however, an increase in the digestible
of rations

seem

rations

time, on

has

secured

been

mals
quiteas effective with anireceiving
alreadyon a high plane of proteinnutrition,i.e.,
the minimum
requirement as with those
a
large surplus over
This
level of protein supply.
much
lower
a
on
appears
with
especialclearness in Morgen's experiments on sheep.

appears

The

to have

results

been

therefore

on

the whole

fail to

indicate

the limits

within

which

stimulatingeffect is manifest or to establish any quantitative

the surplusproteinsuppliedand the additional
relation between
for any estimate
milk yielded. They afford no basis,therefore,
of milk productionby means
of the extent to which a stimulation
under any given
of excess
proteinwill be economically profitable
this

conditions.

NUTRITION

512
If there

FARM

ANIMALS

available definite

were

of

values

energy

OF

feeding stuffs

knowledge regarding the net

for milk production, the foregoing

figuresfor the total energy of the milk would

serve
basis for estimating the net energy
as
a
supply requiredfor
the production of a given yield of milk
for any
of the
one
ten grades,25 pounds of 4 per cent milk, for example, requiring
8400 Cals. of net energy in the feed.
336 X 25
=

605.

Equivalent

values

energy

of determinations

of the net

for

In

fattening.
"

values

energy

of

the

sence
ab-

feedingstuffs

production (588)it is impossibleto make direct use, in

the manner
justindicated,of the foregoingdata regardingthe
of milk.
Pending such determinations,however,
energy content
it appears
possibleto estimate the net energy requirements in
the feed of dairy cows
in another
way, viz.,by computing from
the compositionof the milk, in the manner
alreadydescribed
for milk

(593),the

of

amount

fatteningwhich
of milk
yield.

unit

a
requirement
certain assumptions,that
on

to

is

equivalentin

Thus

it

was

energy

estimated,

of feed energy required

for the productionof one pound of average 4 per cent milk would,
have produced a gain of only 252 Cals.
if appliedto fattening,
in

of net

ration
energy

pound

this way
of

ingly,
Accordactuallypresent in the milk.
in excess
of maintenance, 252 Cals.
containing,
for fattening
would
have been adequate to produce

place of the 336

a

of

Cals.

milk

the amount

pound

one

the amount

In
energy.
requiredfor the production

of net

energy
of each
of the

of milk

grades included

previous table may be computed.

By this device of reducingthe total
to the

of net

of

Cals.

containing 336

content

energy
energy for

in

the

of the milk

fattening,it appears
possibleto utilize the net energy values of feeds obtained by
and
Kellner
others
in maintenance
or
fattening experiments
for milk.
Such a method
a basis for computing rations
as
is,of

equivalentamount

and
provisional,

course,

the basis for it at present is somewhat

the best one

slender,but it seems
now
for computing rations,however,
use
to

take

well-fed
some

which

into account
cows

the

the fact shown

of the
digestibility

lower
than
the
5 per cent
used in computing net
are

available.
it appears

by

Eckles

energy
the figuresfor the equivalent energy
for

necessary
(722)that

also

with

the average
digestioncoefficients

rations

average

In its actual

is

on

Accordingly,
fatteningas computed
values.

MILK

PRODUCTION

513

grades of milk have been increased by 5 per cent,

giving the followingresults,which may be used provisionally
of the Appendix
from
the figuresof Table
VII
to compute
the rations requiredfor the production of milk of different
for the several

grades.
Table

606.

145.

Equivalent

"

Concurrent

Energy

fattening.
"

Values

Were

for

Fattening

all the

surplusfeed above
milk
production, it

requirement applied to
to compute the amount
would be a comparativelysimple matter
of feed energy requiredin a dailyration. Thus, if a cow
ing
weigh1000
pounds were
capable of producing 25 pounds of 4.5
per cent milk daily,the net energy requiredin her ration would
the

be

maintenance

computed

as

follows

"

production25 lb. of milk "291

For

milk

For

maintenance

Cals.

7.275 Therms
6.000 Therms
13.275

has

Attention

been

called several

times, however,

Therms
to

the

milking animal at least two forms of production

are
possible,
viz.,milk and increase of body tissue (fattening),
only the former of which is usually desired. To these may
perhaps be added, as a third form of production,a possible
stimulation of the incidental muscular
activityof the animal
by heavy feeding (609). Evidentlyif conditions are such that

fact that in the

Including5
2

per

cent

allowance

for difference in

digestibility.

OF

NUTRITION

514

is diverted

part of the feed energy

ration

must

would

be necessary

supply

both

established

other

that

such

forms

may

these other purposes,

to

plane of nutrition.

"

It appears

experience and

common

the
than
duction.
pro-

to be well

by direct experiment

of energy
from milk productionto
in fact take place before the maximum
capacity

diversion

glands is

milk

of the

by

ANIMALS

energy
per pound of milk
utilized for milk
latter were

net

more

if all the

of

Influence

607.

FARM

reached.

moderate

On

rations,the

net

satisfyingthe maintenance
requirement,may
milk
utilized
for
be
entirely
production. As the
apparently
does not continue
feed is increased,however, the animal
to
after

energy,

available

utilize all the

the limit of its

with

then

for milk

productionup to
suddenlybegin to utilize any

the heavier rations

On

higher plane

the concentration

the organincreases,
ism
and
at
a point varying
nutrition,

in the

digestednutrients

reaches

energy

capacityand

surplusfor fattening.
of the

net

of

fluids

body

this greater concentration

different individuals

of available

speak,robs
for milk production.
milk glandsof feed intended
of
608.
of individuality. The
Influence
individuality
material

fatteningto begin,which,

causes

so

to

"

is

animal
cows

important

most

having

so-called beef

breeds, this point

at

which

as
fattening,

energy

the

With

in this connection.

tendency toward

inherited

an

factor

the

in the

begins

be

to

productionand fatteningmay be reached

on
comparativelylightrations. Such animals can be brought
to their maximum
milk-producingcapacity only at the
up
of a considerable
expenditureof feed for concurrent
expense
for dairy purposes.
fatteningand are likelyto be unprofitable
nance
On lightrations,givinga moderate
yield of milk, the maintedivided

between

milk

requirement
cost, while

with

constitutes

heavier

too

largea proportion of

feedingproduction is

largelyto fattening.
the typical dairy animal,
With
but a slighttendency to fatten,the
towards

the amount

of the milk

glands,or

on

the other

the feed

directed

too

hand, having

be increased well
may
capacity
requiredto support the maximum
that
in exceptional
to
even
cases
point,
up
feed

material
diversion to fattening. Such
any
if of large milk-producing
capacity,are the
animals, especially

without

causing

dairy animals
profitable

so

far

as

the cost of feed is concerned.

MILK

relations between

The

demonstrated

in

such

of

and

those

as

Hess

Carlyle3
4

Haecker
609.

it

and

Weld

and

of Waters

Pennsylvania station,those by Woll and

Wisconsin
station,and especiallythose by

the Minnesota

Stimulation

from

beyond

diversion

station.

of katabolism.

obtained

is increased

part by

Waters, Caldwell

the

at

returns

feed

the

at

at

515

ing
supply,milk productionand fattenthe foregoing paragraphs have
been clearly
number
of investigations
a
on
dairy feeding,

in

outlined

PRODUCTION

the feed of the

certain

maximum

net

while

dairycow
may
milk

from
energy
to be true also that heavier

seems

of

But

"

the
as

ing
diminishits amount

be

explained in
tening,
production to fat-

feedingmay

cause

to be oxidized,
largerproportionof the digestedorganicmatter
either as the result of greater muscular
activityor by a direct
stimulation of the katabolic processes.
This is especially
evident
in breed
tests in which
heavy rations have been consumed.
Striking illustrations of it are afforded by the results of the
Purchase
tests of dairy breeds at the Louisiana
Exposition in
5
and by the extensive comparisons
1904 as computed by Haecker
of German
breeds reportedby Hansen.6
a

610.

Diminishing

returns

from

feed.

"

It is evident

from

the

of a very
foregoingthat,with the possibleexceptionof cows
be
pronounced dairy type, the maximum
yield of milk can
secured only at the expense
of a simultaneous
production of
less body fat and perhaps also of a stimulation
of the
more
or
of the body. Consequently,beyond the
katabolic processes
point at which this fatteningor stimulation begins, the milk
productionper unit of net energy in the feed must
necessarily
be a diminishing
of
one, and it is clear that the determination
the net energy requirements for milk productionis to a considerable
economic

problem.
Milk
will be produced at the least feed (energy)cost per
pound when the ration is so adjusted as to produce as great a
extent

yieldof
1

Penna.

Wis.

an

milk

as

is

possiblewithout

causingfattening.7If

the

2
Expt. Sta.,Rpt. 1893, p. 24-36.
Ibid.,Rpt. 1895, p. 24-55.
Expt. Sta., 17th Rpt. (1900), p. 37-61.
4 Minn.
5
Minn.
Expt. Sta., Buls. 79 and 140.
Expt. Sta., Bui. 106, p. 158.
6 Landw.
Jahrb., 35 (1906), Ergzbd. IV, 147-236; 37 (1908), Ergzbd. Ill, 2362er Ber. vom
Dikopshof (1911), 210, 430.
410;
7
It may
be presumed
that the stimulatingeffect upon
the katabolism
occurs
chieflyin heavy feeding which causes
fatteningalso.

516

FARM

OF

NUTRITION

point the milk yieldwill

tend to fall off while the maintenance
requirement remains
The maintenance, therefore,
will consume
constant.
practically
total feed energy
so
that, exactlyas
a largerpercentage of the
the net
in growth or in fattening,while
requirement
energy

supply is

energy

decreased

of

for the formation

total

the

net

milk

and

below

ANIMALS

this

unit of milk

remains

stant,
approximatelycon-

both maintenance
to support
necessary
production increases relatively
per unit of
energy

product.
hand,

the other

On

katabolism, it

stimulate

to

or

if the feed is increased

is clear

as

to

that

will be

of milk

unit

so

tening
fat-

cause

the

energy
creased
apparentlyin-

produced
reasons
alreadyexplained. Such an increase in
for the
be economicallyjustifiable
the feed cost, however, may
of any form of intensive production.
reasons
as in the case
same
be doubted
milk
commercial
In average
production,it may
the rations should be made
whether
heavy enough to cause
any
requirements per
for the

fattening,and

considerable

far

so

mated
this is the case, the esti-

as

values
per unit of milk
energy
the basis for computing rations.

net
serve

as

in Table

145 may
If,however, feed is

ing
relatively
cheap and dairyproducts high in price,the diminishstill be profitable
due to heavier
returns
feeding may
up
of
milk
unit
certain
to a
point even
though more
per
energy
be supplied in order to support concurrent
must
fattening,
the fact that

while

more

less of

or

the

fat stored in the

body

early

be utilized for the support of milk productionin the

stages of the next lactation is also to be considered.

may

for milk production

requirement

Fat
611.

Is fat essential?

It

"

was

noted

in

the
discussing

tions
func-

with the requirements

(265)and also in connection
growth (498,499) that the presence in the feed

of the nutrients
for
of certain

fats

be essential

growth.

to

analogous

to

That

the

milk

body

in

whole

fat

as

Since

growth

the fats of the feed exert

productionas

associated with

of substances

or

any

or

on

well

as

largeamounts

appears
productionis in many

milk

it is of

them

interest

to

to
spects
re-

inquirewhether

either on milk
effect,
specific
the productionof milk fat.
body fat may be manufactured in
such

from

other

nutrients

has

been

shown

MILK

beyond questionby

PRODUCTION

517

the

experiments of Voit,Kuhn and Fleischer,

M.
Fleisher,Wolff and especiallyby those of Jordan,1 while
the latter investigatordemonstrated
that milk fat can
be
from
formed
carbohydrates (553). Jordan's experiments on
of Morgen 2 on sheep and goats,
well as the later ones
as
cows,
likewise show
that relatively
of milk
be
large amounts
may
rations

produced on
from

or

which

the

made

of

feedingstuffs very

poor in fat
been extracted.

larger part of the fat has

It is

scarcelyfeasible

such

animals

milk

productionwith

and

up

to

prepare
the writer is not

absolutelyfat-free
aware

such

rations for

of any experiments on
it is clear that at most

rations,but
of fat can
but very small amounts
be regarded as indispensable.
of fat to rations.
612. Addition
Experiments in which the
fat content
of ordinary rations has been increased,either by
the direct addition of fat in one
form or another or by the substitution
of fat for carbohydrates,
have given very contradictory
"

results.
been

An

increased

percentage of fat in the milk

has

frequentlyobserved, sometimes accompanied by an

increase in the actual yieldof fat and sometimes
not, while in
other cases
the results have been entirely
negative. In many
instances the experiments are complicated by the fact that the
fat was
simply added to a basal ration, thus increasingthe
of feed.3 The
total amount
most
recent
are
investigations
those undertaken
a
common
plan under the auspices of
upon
the German
AgriculturalCouncil at ten German
experiment
very

stations

with, in all,196

reportedby
The

the results of which

cows,

have

been

Kellner.4

increase in the fat of the rations

effected

substituby the tion

of rice feed 5 for rye meal
and starch,so that fat replaced an
of carbohydrates. The
cluded
equivalent amount
results,therefore,inif such there
"specific"effects of these two feedingstuffs,
any
Per
were
1000
(618).
pounds live weight, the fat-poorrations contained
f
at
and
the
to
fat-rich
0.25
0.50 pound digestible
0.47 to 1.10
pounds.
1

N.

Landw.

3
4
5

nor

was

Y.

(Geneva) Expt. Sta., Buls. 132 (1897) and 197 (1901).

Vers. Stat.,61 (1904), 1 ; 62 (1905),251 ; 64 (1906), 93.
Compare Kellner, Die Ernahrung der landw. Nutztiere, 6th Edition, pp. 564-566.
Reichsamt

des Innern
to Hansen

Berichte

rice feed

According
although this effect did
in those of Fingerling (613).

liber
has

not

the

appear

Landwirtschaft,Heft

and

2.

specificeffect of depressing the fat

manifest

in most

of these

duction,
pro-

experiments

5i8

in the percentage

associated

ANIMALS

regarding the fat content

of milk yielded,it
the total amount

according to
increase

FARM

with

Table

in the total

decrease

146.

Effect

"

of

yieldand

Increasing

At

Experiments

Fat

Bonn

"

Pommritz
Kiel

Breslau

+
"

Triesdorf

Weihenstephan
Lauchstadt

Darmstadt

Jena

crease
In-

Average

It

147.

"

average

of fat for

Influence

clear

seems

conditions

expected

%
0.2
%
3-1 %
2.3 %
7.1 %
2.1%
0.5

from

two

of the

5.o%

0.3%
2.6%

of

from
of

Individuality

the

3-7%

as

Effects

foregoingresults

4-8%
1.0%
10.8 %

animals

on

0.6%

carbohydrateswas

3.3%

7-9

however,
experiments the

for the individual

decrease

the substitution

Table

OR

9-8%
6.6%

differences in cows,

example, in

For
or

2.5

crease
In-

Decrease
(" )
Content
Fat
in
Milk
of

-2.7%

Striking individual
increase

Percentage

(+)

Yield

0.5

versa.

Rations

of

-6.7%

vice

(+) OR
(" ) of

Decrease
Milk

that
appears
in general
was

of fat in the milk

Percentage

Danzig

of the milk

the results

Grouping
an

OF

NUTRITION

served.
ob-

were

of

range

consequent
follows

of

that

Fat

on

"

Increase

under

the

practiceno material advantage can be

increasingthe digestiblefat of dairy rations

OF

NUTRITION

520

FARM

ANIMALS

of fat, may
favorable
a
by-products containinga minimum
pated
the yieldof milk and its percentage of fat be anticieffect upon
from an increase in the supply of digestible
fat.
614.

Influence

afford any

of the

None

"

periments
ex-

milk

supplyupon

regardingthe

data

exact

energy.

of the fat

the influence

on

of

utilization

on

production
gain or loss of

concurrent

of the gaseous excreta were

made.
the observed
It is impossible,
whether
therefore,to determine
since
tissue,

determinations

no

brought about by a stimulation of milk

of fattening,
i.e.,
by modifying the
expense

effect of the feed fat

production

the

at

the energy of the feed was

or whether,
utilized,
in the milk gland was
its influence,
the metabolism
ally
actu-

direction
under

in which

effected

of the ash

estimate

mineral

for milk production

requirements
data

no
Practically

615.

economically.

more

Ash

The

are

elements

on

record

the

milk.

in the milk

that

which

dairycow

It is true

"

trustworthy

can

be based.

the outgo of
without special

that

be determined

may

figures are available

This, however, is but a single

reasonably accurate

which

element

upon

requirementsof the

outgo in

and
difficulty
from

was

it may
be estimated.
in the problem. It became

evident

ash

requirements for maintenance

and those for growth in Chapter

in

in

consideringthe

Chapter

IX

(421-436)

(492-497)that neither the

of the mineral
actual availability
elements
of feeding stuffs
the influence of the amount
and qualityof the ash supply
nor
the losses in feces and
gated
investiurine has been
sufficiently
upon
to permit any
conclusions as to the influence
satisfactory
XI

of these factors.
*

Kellner

has, however, computed the approximate requirements

for calcium

phosphorus from the outgo in the milk.

Accepting Henneberg's estimate of 71.4 grams of calcium and
of phosphorus per 1000
21.8 grams
kilogramslive weight for the
maintenance
average

and

requirements, he
found

amounts

assumption

in the

that

only

adds

to

milk

upon

one-third

ash

is available.

for

per

day by

cow,

Computed
thousand
pound
1

Ernahrung

landw.

these
the

his results

Nutztiere, 6th Ed.,

are

p. 595.

as

tionable
ques-

the

pounds

20

the

times

somewhat

one-half

to

yieldof

three

feed

of milk

follows

"

MILK

Table

616.

148.

The

that
the

Estimated

"

supply
will

dairy rations

PRODUCTION

Requirements

feed.

in the

"

the

usuallymeet

only in exceptionalcases
calcium supply.

Forbes

52

for

Kellner

Milk

states

Production

that

ordinary

requirementsjuststated

will it be necessary

to

and

supplement

has

shown, however, that rations fullyadequate, so

far as organicnutrients are concerned,to support a considerably
ments
greater milk productionthan that on which Kellner's requirebased

permit very material losses

of mineral ingredients,
of calcium, magnesium and
especially
reported for six
phosphorus. Complete ash balances are
are

animals
live
With

on

may

nevertheless

three different

weightsof

the

cows

rations,all of which
resulted in

and

maintained

the

gains of body protein.

live weight of about 935 pounds and an average

average
dailymilk yield of about 36 pounds (16.38Kgs.),the calcium

and

an

phosphorus requirementsas
method

were

and

the actual

computed

according

suppliedin

amounts

the

to

rations

"

Table

149.

"

Calcium

Ohio

and

Phosphorus

Expt. Sta., Bui.

for

275

Milk

(1916).

ner's
Kell-

Production

NUTRITION

522
Whether

shrinkage of

made

usually

cows

up

lactation,and whether
of the factors in the natural
the body is one
production, as suggested by Forbes, are

of
milk

investigation.

evident,however,

information

much

fresh

again in
this depletionof the

for future

It is

ANIMALS

relatively
large losses by
copiousmilk production and are

reserves

matters

FARM

such

accompany
the later stages of
mineral

OF

that

of the

none

regardingthe

real ash

foregoingdata afford
requirementsof dairy

cows.

Feed
617.
"

odor

or

taste
to

to milk

in

feedingstuff
of

which

small

or

proteinor

do not

production

flavoring substances

By

"

presence

its content

substances

are

whose
of

stimulus

substances.

Flavoring
those

meant

as

ration
energy.

yieldmatter

is

improves the
while not adding materially
In other words, they
to the body
or
energy

amounts

ordinary sense, but which may nevertheless affect the

course
or
rapidityof metabolism.
That
the flavor or aroma
of feedingstuffs is not an insignificant
element
in determiningtheir commercial
value, not only
is well established
for milk productionbut for other purposes,
by practicalexperience. This superiorityis doubtless due
more
largelyto the fact that a palatablefeed is consumed
freely
of
milk
in
the
flavor.
In
than one
case
production,
lacking

in the

however, it

limits,

various

to the

milk
In

that, within certain rather narrow

appears
flavoringmaterials may act as a direct stimulus

of fat.
gland,causing a greater yieldof milk and especially
Morgen's experiments on milk productioncited on previous
extensive
of rations consistinglargelyof
made
use
was

pages
almost

flavorless

materials.

be

impossibleto secure
rations supplying equal
to

made

up

of

normal

With

such

rations

it

was

yieldsequal to those obtained

of protein and
amounts
energy

feeds.

The

addition

to

these

found
from

but

flavorless

as
fennel,anise or hay
rations,however, of such substances
the introduction of malt sprouts, caused
a distinct
or
distillate,
increase in the milk yield,so that, with rations containing a
secured.1
of fat,almost
or
sufficiency
quitenormal results were
in increasing
observed
the fat
Moreover, a distinct effect was
production and the percentage of fat in the milk.
1

Landw.

Vers.

Stat.,61 (1904),1.

MILK

PRODUCTION

523

*
Subsequent experimentsby Fingerling
fullyconfirmed these
results.
The addition
to the flavorless rations,or
to damaged
the impregnationof
fennel,or even
hay, of salt,hay distillate,
rations with the odor of the latter substances,caused a marked

in the

increase

yieldof

milk

and

in its content

of fat

well

as

as

in the percentage of fat in the milk

normal
rations were
without
to

while similar additions

solids,
effect. Fingerling's
experiments
likewise show clearly,
ing
however, that this effect of flavorcan
materials,while of much
interest,
physiological
rarely

of much

be

economic
of the

the claims

etc.,

It

also shown

was

powders,

that

certain

cake
feeding stuffs (malt sprouts, palmnut cake, cocoa
beet molasses)when added to a ration of damaged hay and
nutrients increased the milk and fat yieldsto about the
extent
to

as

some

(549)does
618.

not

production of

pure
same

the hormones

Specific effects

feeds.

of

the

general.

feedingstuffs

The

may
the quality of the milk

"

The

production of

appparentlyby their influence

naturallyto a consideration of
of feeds in

of milk

production

appear.

certain feeds stimulate

that

and

these effects are due

flavoringwith fennel. Whether
form of nerve
to
stimulus,either general or specific,
or

increased

an

to

support

no

feeds,milk

condimental

numerous

largelyadvertised.

so

they lend

importance and

the

on

the

fact

just noted

milk

and

that

of milk

fat,

flavor of

so-called

"

rations,leads
"
effects
specific

belief has

promote
to

an

long been held in practice

milk production and improve

extent

of

fullyexplainedby the
which they supply.
energy
not

of
or
digestiblematter
On the other hand, there has been no general agreement as to
what particular
feeding stuffs possess this power, and scientific
have been led to question the existence of such
investigators
the compositionof milk.
A discussion
effects,
particularly
upon
of the literature of the subjectup to 1903 by Lemmermann
amounts

and

Linkh

affords

strikinginstances

of the

tween
bediscrepancies

different

experiments. The effects of such feeds as

palmnut meal, cocoa
meal, and cottonseed meal, for example,
able
are
reported by different experimentersas favorable,unfavoror

Vers. Stat.,62 (1905),11;

(1911), 163.
Landw.
Jahrb., 33 (1903), 564.

^andw.
373;
2

indifferent.

74

64

(1906),357;

67

(1907),253;

71

(1909),

OF

NUTRITION

524
Defective
much

ANIMALS

experiments is doubtless

of

planning

FARM

of this confusion.

In

the

instances

many

responsiblefor
experimentershave

lightbasal ration,as in the

familiar experiments by G. Kiihn
on
palmnut meal so frequently
referred to.
one
feeding stuff for another,
Others, while substituting
of digestible
that the total amount
have failed to show
matter
plied
supfor instance,
extensive investigations,
was
unchanged. In some
oil meals and similar feeds have been interchangedin amounts
ing
supplyother
ingredients.
equal quantitiesof protein without regard to
be
conditions
could
Under
concordant
results
not
such
expected,and
simply

added

the

feed

be tested

to

to

is
and Linkh that the evidence
agree with Lemmermann
inconclusive,while their own
specific
experiments,although indicating
one

but

can

effects for various

dairy herds

trials with
and

S torch

unit

"

and

carried

by

convincing.

the extensive

by Fjord, Friis

in Denmark

out

feeding

afford the basis for the so-called

which

Scandinavian

or

these trials

In

is afforded

negativeevidence

Similar

scarcelymore

are
feedingstuffs,

"

feed

comparing rations (702).

puted
refeedingstuffs,includingmany
of

system

varietyof

effects on milk production,

were
compared
specific
ordinary farm grains and failed to exert any material
to have

with

influence

on

the milk

secretion

other than

the variations in the

explainedby

what

may

be

the total

and

protein content

plausibly

experimentalmethod.
effect on the production
specific

of the rations incident

nutrients

to

the

indications of a
particular,
of milk fat are lacking.
More
positive results have been reached,however, in two
recent investigations,
viz.,in a series of investigations
by Hansen
at the AgriculturalAcademy
Bonn-Poppelsdorfand in a series
of cooperative experiments on
under the
palmnut meal made
auspicesof the German
AgriculturalCouncil.

In

Hansen's

619.

nine

series

experiments.
on

63

cows,

the various

feedingstuffs

comparison

ration

in this respect.
nutrients
net

energy

in the

Jour. Landw.,
Landw.
; 40

for others
Care

was

extendingover
be

tested

which

taken

experiments
2

5 years,

appeared to
to

keep

in which

substituted

were

the

be

total

cluded
in-

in

indifferent

digestible

after the first 3 years, the estimated

rations,or
values
(starchvalues),unchanged.

Ill, 171

to

Hansen's

"

22 (1874),178.
Jahrb., 35 (1906), 125 ; 35 Ergzbd. Bd. IV, 327
(191 1),Ergzbd. Bd. I, 129.

; 37

(1908),Ergzbd. Bd.

PRODUCTION

MILK

feeding stuffs

distinct effects of certain

results show

The

525

on

apparently quite independentof the

yieldwhich were
nutrients or of energy values, or of the protein
supply of digestible
the milk

supply, and which

the

consistent when

were

repeated.
1
three
Hansen
distinguishes

experiments

were

feedingstuffs.
(apparently
of the first group,
including maizena
Those
gluten feed),maize and oats, increase the quantityof milk but
depress the percentage of fat,so that the total yieldof fat is not
of the second
including
group,
materially changed. Those
residues,maize distillers'grains,and to a
palmnut meal, cocoa
of these

groups

"

"

less

the

total

yield of

crease
legumes, inmateriallyaffectingthe
and

meal

cottonseed

degree linseed and

fat without

the

that the percentage of fat in the milk is

of the third group,
cake,
including poppy

quantity of milk, so
Those

increased.
"

false flax

diminish

"

the

cake, rice feed and

do not

fat but

yieldof

to

cake,
degree sesame
sensiblyaffect the quantity
a

less

milk, so that the percentage of fat is decreased.

3
the same
plan,
on
substantially
In a subsequent investigation
has compared the effects of palmnut cake containing
Hansen
of

respectively
5.55
He

amounts.

the

and

12.42

concludes

per

that

proportionof palmnut

the

effect
specific

in the ration

cake

in the cake.

of fat contained

fed in different

fat when

cent

and

increases with
with

the percentage

finds that to

He

secure

about 2 pounds per 1000

results in practice,
pounds
significant
live weight of fat-rich cake and 2 J to 3 pounds of the poorer
ferent
have difDifferent individual animals
grades are necessary.
to the effects of palmnut cake
degrees of susceptibility
but the result

can

be obtained

if sufficient is fed.

principalcriticism to be made of Hansen's experiments

short
so
usually 7 days
is that the experimental
periodswere
It is not
preliminaryand 7 days for the experiment proper.
unusual
an
experiencein dairy feeding experiments to see a
change of rations followed by a temporary stimulation of the
rally
and the questionnatumilk productionwhich is not sustained,
to be
the
arises whether
specific effects which seem
The

"

"

"

demonstrated
for

in the first week

or

two

would

have

continued

longer time.
1

Loc. cit.,Bd.

Landw.

40,

Jahrb.,

187-188.
(1914), 30-

pp.

47

Camelina Sativa.

Cooperative experiments.

620.

the

under

ANIMALS

FARM

OF

NUTRITION

526

The

"

ments
cooperative experi-

the German

auspicesof

AgriculturalCouncil

made
palmnut cake or meal and were
different institutions with,
plan at seven
according to a common
The
in all,1 86 cows.
experimental periodscovered about one
month
regarded as a
each, of which the first 5 to 7 days were
between
4 pounds
preliminaryfeeding. The comparison was
of

the influence

relate to

of

palmnut

of

meal

meal

of maize

mixture

protein and

values

energy

sulted
lactation,the substitution of palmnut meal reonly in each of the 7 experiments as a whole, but with

of

the advance
not

nearly all the individual

The

production.

The

increased.

quantity of

total

150.

milk

effects of the

of fat in the milk

meal

palmnut

Effects

"

Palmnut

of

Meal

Milk

on

Increase
Daily
(+)
Decrease
(" ) in
Yield

Milk

Percentage

Bonn
.

Griefswald

Friesdorf

to

smaller

the effects of the

It

amounts.

immediatelyfollow
reachingits maximum
1

was

the

3-24

48

3-25

2-97

O.28

22

3-i7

3-05

0.09

64

3-5i

3-17

O.04

15

3-68

0.06

25

3.85
3-78

13

4.21

4-05

Berichte

change

uber

milkers

than

that

of feed but
course

seemed

meal

palmnut

of

one

Landwirtschaft,Heft

in

On Check
Ration

O.58

also observed

in the

Palmnut
Meal

good

were

On

3-58

-f-O.02

that

Fat

62

general,cows

of

"

Weihenstephan

In

as

O.29

"

Hamburg
Jena

were

Milk

Milk Fat
Grams

Kgs.

was

Production

or

Danzig

tially
substan-

yieldedwas

the percentage

average

of the fat

distinct increase

followingtable.

in the

Table

in

cows

unaffected,so that
shown

weight and an amount

and peanut meal
supplyingequal
(computed). After correctingfor
live

pounds

1000

per

3-5i

more

those

ceptible
sus-

yielding

the effect did not

developed gradually,
two
or
weeks, and
21

and

23.

528
the

NUTRITION

624.

Influence

centered

percentage

on

this substance.

around

increase
specific

of Kuhn

the work

an

increase

in the

The

well

as

conservative

to

has

percentage
percentage

content,

down

in 1868

fat is

discussion

view

up by Jordan in 1908.1
much
discussed and much

point was thus summed

This question has been
from

the

in the

fat.

in the

Since

"

increase

in the water

selves
them-

of the evidence.

An

result from

a decrease
solids,
i.e.,

of total

contented

of fat in milk.

ingredient of milk,

of fat in the milk may

from

conservative,have
the conflicting
nature

pointingout

valuable
specially

the

"

ANIMALS

milk,while others,more
with

this

FARM

OF

as
on

gated
investi-

the present

day.

experimentshave been conducted for long periods and

pared
periodsin which very moderate rations have been comwith very largeones, highly nitrogenousfoods with those

Many
short

protein content, dry with green or succulent materials,

class with one
and grains of the same
another, and, in a great
consistent
no
majority of cases, the verdict has been that
of

low

'

exist between

relation

to
appears
the ration and the

the

character

quantity or

of

has
writer
The
composition of the milk.'
examined
the results of nearly all the important experiments
of this character of which he could find a record,and in but few

could

cases

in the

decrease
relation

he

discover

there

that

proportionof

to variations

in the ration.

appeared in the milk

change
in the ration,but

milk

in most

was

solids which
In

some

immediately

instances

bore

cases

after

of this kind

or

logical

temporary

violent
there

change

was

return

by

causes

to

the

not

under

his

normal

very
small

product. In a
manent
proportionof experiments,the milk appeared to sustain a perThe
modification.
extensive
weight of
though not
that the qualityof milk
testimony bears out the statement
mined
be changed at will by the farmer, but is largelydetercannot
a

soon

animal's

increase

material

control,such

as

breed

and

viduality
indi-

the latter,
although feedingand treatment, especially

have

or

more

less influence

upon

the

character

of the

milk

secreted."
position
feeding stuffs upon the comof milk is associated with the question of the so-called
specific effects of feeding stuffs (618). As was pointed out

Much

"

of the

allegedeffect

of

"

The

Feeding of Animals, 5th Edition, The

PP- 3i7-3i8.

Macmillan

Co., New

York, 1908,

MILK

PRODUCTION

that question,both
considering

in
of

of

many

the

529

the

planningand

execution

older

defective and their

experiments were
be regarded as inconclusive. The
recent
more
and of the German
of Hansen
cil,
AgriculturalCoun-

results must

experiments
the other
on

hand, as

well

the

as

of Morgen
investigations

their associates upon

the influence of feed fat
of condiments upon
afford numerous
milk production,
parently
ap-

Fingerlmgand
and

unquestionable instances of
the fat content

upon

effect of the ration

an

of the milk.

example, the experiments of the German

Agricultural
Council on palmnut meal
(620)showed an average increase in
the percentage of fat rangingin the different experiments from
For

to

cent

per

cent, while individual

per

showed

cows

even

strikingdifferences. Morgen's results on the specific

effect of feed fat when added to fat-poor
rations (613)and likewise
results regardingthe influence of condiments
Fingerling's
effect
more
(617) afford even
strikingexamples of the same

more

Apparentlyit

be admitted

must

the fat content

of milk

may

that this effect appears

of the ration.

and

625.

Influence

such

conditions

some

be

affected by
distinctly

to

be associated with the fat supplv

of

percentage

on

from

it appears

that, under

fat in

of these latter

solids.

the

feeding

more
Further-

"

experiments as

cluded
in-

determinations of the total solids of the milk that the

increase

i.e., that the

a
essentially

was

proportionof fat

This

milk

"

fat content

in

to

one-sided

marked

other solids in the milk

was

creased.
in-

results

was

of fat to the feed.

also obtained

were

Similar
in

althoughmuch less
experiments
Fingerling's

the influence of condiments.

upon
In Hansen's experiments,
too, the influence on
of the milk
increase

was

due, in

the

decrease of the

or

the
solids,

being

over

while

Lmdsey1

cocoa

increase

notably the case in the majorityof experiments

in
of
the
which
fat
in the
on
fat-poorrations,
proportion
solids was
increased by from 12.5 per cent to 23.5 per cent

by the addition

mrik

"

increase

5 per

cent

has

majorityof instances,
largelyto
percentage

decrease

of fat contained

over

the

Hansen's

meal.

an

in the

comparison rations
in. fullyone-third of the experiments
or

confirmed

the fat content

Mass.

Expt. Sta.,Bui.

155.

results

as

regards

NUTRITION

530

It
fat

the

in

the

of

626.

this

be

those

been

market

milk

is

most

essentially

altogether
of

selection
such
on

of
and

feeding
might

on

suited

to

easily

and

at

that

earlier

and

less
real

well

be

meet

the

the

is

be

made

the
to

meal.

is

of

one

subject

those

of

of

possible
the

by

depressed

of

the

ing
dairyThe

intelligent
the

time

short

in

has

As

particular

by

secondary

increase

is

matter

of

same

it

due

producer.

secured

If

observed

it may

milk

from

are

successful

demands

the

one.

milk

of

influence

At

true

(564).

in

any

feed

remains

still

animals

certainly

while

been

similar

palmnut

as

permanently

has

stuffs,

animal

not,
lished
estab-

importance

factor

prime

must

apparently

composition

in

factor.

stress

It

commercial

while

negligible
to

hand,

investigations
Council

as

other

the

that

feed

extent

the

of

secondary

of

the

selection,

indicate

to

seem

the

facts

the

differences

insisted,

best

and

breeding

in

are

individual

capacity

of

quality

not

which

and

the

centage
per-

presumed

fact, represent

considered.

just

differences

repeatedly
is

its

temporarily

much

Too

"

physiological

evidence

sources

breed

of

and

fat

be

in

point

matter

practice.

the

on

major

different
to

in

laid

the

by

as

of

may

of

proportion

sort.

Significance

however,

it

this

on

yield

influenced,

and

feed,

did,

experiments
of

that

the

the

that

total

be

may

obtained

results

conclusive
effect

of

facts

the

as

milk,

nature

ANIMALS

FARM

foregoing
well

as

fresh

the

the

least, by

the

solids,

milk
in

some

from

clear

seems

OF

fat

the

by
yield

unsuitable

considerable

large
German

scale

is

results

factor

experiments,
an

feed

it

is

suitable
to

any
or

if,

choice

importance
cooperative
Agricultural

CHAPTER
WORK

Prime

627.

the
feed in
of

prime

PRODUCTION

of

purpose

XIV

feed.

excess

for which

purpose

of its maintenance

excess

the external

either natural

domesticated

work

mechanical

work

Aside from

"

animal

mature

It is true

that

more

material

reserve

which

be

may

the

feed may
be
that a fattening

than is requiredfor this purpose

and
of the animal may
result. The
latter,however, is
of

in

enforced

or

consumed

layingaside

consumes

requirement is the production

tivities,
required for its diverse ac-

in the wild animal

animal.

tion,
reproduc-

simply a

utilized later

and, however

be regarded as
important economically,may
of the
incidental. Any considerable fattening
physiologically
work
animal is not only a diversion of energy from the main
of the

purpose

carried
with

the

by

feedingbut

of the United

i.

or

The

weight

it may

extensive

to

be

interfere

respiration.

mules

are

the working animals

substantially
ence
refer-

these animals.

The

Physiology
Nature

628.

extra

an

discussion will have

States,the following

chieflyto

"

if too

animal,while

heart action and

Since horses

constitutes

muscles.

"

of

of

Work

muscular

Mechanical

Production

work
work

is

performed by

an

(84,85),of which there are two

kinds called,respectively,
and smooth, or
striped,or striated,
of the microscopic
muscles from the appearance
non-striated,
fibers of which they are composed. The skeletal muscles, by
of which external work is performed, are striated muscles.
means
animal

They

by

are

means

of its muscles

also called

voluntarymuscles
53*

because

they are

inner-

OF

NUTRITION

532

the

from

vated

and

cerebrospinalsystem

under

are

of the internal organs

muscles

The

of the will.

the

trol
con-

chiefly

are

being the conspicuousexception,

nervated
to a very limited
degreesubjectto the will,being infrom
the sympatheticnervous
system. In the study
with the
production,therefore,we have to do chiefly
muscles,the

non-striated
and

ANIMALS

FARM

are

of work

heart

phenonena of striated voluntarymuscles.

contraction
The physiologyof the muscle and of muscular
is a very complex subject and wide differences of opinionexist
aspects of it. All that is attempted here is to
regardingmany
for a proper
outline such generalfeatures as seem
necessary
comprehension of its relations to nutrition.
Contraction.

629.

livinganimal

the

When

"

is

usuallya

nerve

is,it tends

it contracts, that

muscle

in
stimulus, which
stimulus,is applied to a

suitable

to

shorter

grow

thicker.

and

shorteningand thickeningof
the muscle is built up.
A single
the individual
fibers of which
stimulus,such, for example, as that caused by the making or
as
breaking of an electric circuit,givesrise to what is known
If such a stimulus is
contraction or twitch.
a simple muscular
repeatedwith sufficient frequency it produces a series of simple
in a state of contraccontractions
which fuse together,resulting
tion
which
continues,subjectto the effects of fatigue,as long
This

as

change

is

brought

about

by

acts.

This

form

the stimulus

received

the

ordinarycontractions
system,

nervous

of muscle

shorteningof
or

be done
in the

When

the

that

are

but

seem

momentary,

the

muscle

for

the

necessarily
imply

Contraction

the

with

in connection

in

ology
physiactual

an

either be isotonic

may

contracting overcomes

example, in raisinga weight, the

is said to be isotonic.

in the

of

When, on the other hand, the

the' muscle are fixed,evidentlyno work

mechanical

sense

the muscle

but

exerts
physiological
sense, i.e.,

is called
630.

the

In

not, however,

the muscle.

points of attachment
can

has

livinganimal
muscles, brought about by

used

as

resistance,as,

contraction

contraction

in their character.

does

isometric.

constant

the

those

contraction

term

of muscular

tetanus."

of

even

tetanic
essentially
The

"

of

name

Chemical

either

an

pull.

still contracts

Such

traction
con-

isometric contraction.

changes in
isotonic

or

contraction.

"

In

isometric,there

muscular

occurs

traction,
con-

rapid

PRODUCTION

WORK

in the

contained

of materials

katabolism

by the circulation together with

it

of their chemical

oxidation,yieldingchieflycarbon

to

the details of the process,

or

katabolism

dioxid

the views

of muscular

general features

muscle

brought to

corresponding

This

energy.

an

Certain

533

of

mation
transfor-

is in effect

and

water, but

as

differ.
physiologists

katabolism

fairlywell

are

is
accompaniment of contraction
oxidation
but a rapid,almost
an
not
explosive,breaking down of a
substances
substance
or
tion
present in the muscle, causing the producdioxid.
of carbon
It has been
shown, according to Zuntz and
Nevertheless,it
Loewy, that the muscle contains no free oxygen.
contracts
instantaneouslywhen
stimulated,while the effects upon
lated
the blood supply follow later,circulation and respiration
being stimumore,
Furtherby the carbon dioxid and other products formed.
it has been
shown
that, under certain conditions at least,a
muscle
and give off carbon
dioxid
in the
continue
to contract
may
made

First

out.

entire absence

immediate

the

of oxygen.

or
activityof the muscle, there is established more
of
with
the
blood
less distinctly
increased
a state
equilibrium
supply,
taken
the
and
carbon
dioxid
muscle
being
given off,
oxygen
up by
of the
number
of
the
dextrose
to
a
while,according
experimenters,
blood also disappearsduring its passage
Other
through the muscle.
products of muscular
katabolism, notably lactic acid and potassium
the
so-called
tend
to
mono-phosphate
fatigue products
accumulate
in the muscle and diminish
and finally
suspend its ability
to respond to a stimulus.
Fatigue of the muscles usually results
lack
of these substances
and not
from
from a gradual accumulation

With

continued

"

"

of material

631.

which

to be

Energy
takes

katabolized.

transformations.

place in

conversion

muscular

of chemical

"

The

katabolism

contraction
energy

of matter

impliesan

into kinetic

energy.

lent
equivaThe

in the two forms of heat

finally
energy thus transformed
appears
and
visible motion
the two
(work) though the ratio between
may

widely

vary

under

different conditions.

As

regards the

intermediate

stages of this process, relativelylittle certain

be said that
knowledge is yet available. Broadly, it may
there

siders
possiblegeneral views. The first of these conthat the potentialenergy
is
of the material katabolized
first converted
into heat, and that subsequentlya portion of
are

this heat

two

is converted

into mechanical

motion.

The

second

NUTRITION

534

generalview considers
produced,a portionof
the

other.

it has

been

not

that heat

differences to

In other words, the

efficiency.
as
being a heat engine.

on

work

of the

conceive

of sufficient temperature

broadest

and

simultaneously

are

existence

is not

less

muscle

in

general
view, stated
prevailing

The

garded
re-

quent
outline,is that in the chemical
changes conseis
in
heat
and
stimulus,energy
part liberated as

expended in producingor maintainingtension

it is as if by
fibers. To use a simple illustration,
the elasticity
of a cord supportinga weight were
increased.

that

no

for its observed

account

muscle

in

The

be lifted for

cord

certain

the muscles

is,when

are

up does mechanical
this increased
tension is also

would

contract

distance.
free to

some

process
denly
sud-

and

the

weight

contraction,

shorten,the increased
In

isometric

converted
finally

in

to be

In isotonic

work.

set

in

of the muscle

part

would

portion

authoritythan Englemann, but nevertheless

In pargenerallyaccepted by physiologists.
ticular,

it is difficult to

the

ANIMALS

the energy taking one

form and a
former
view has been
supported by

The

distinguishedan

FARM

OF

sion
ten-

contraction

heat, as for

into

contraction appliedto simply

example in the case of muscular
sustaininga weight. In this case no work in the mechanical
is done, but energy
is expended in what
has sometimes
sense
been

work

called
"

632.

"

static work."

is the muscular
Tonus.

"

In

assumed
that
tacitly
is absolutelyrelaxed.

in

effort
the

familiar

foregoingparagraphs

before and
Such

of

illustration

"

static

required in standing.
after

is not

of rest, so-called,
there

state

tension

is

contraction

normally
a

it has

greater

the
or

been

the muscle
case.

less

Even

degree of

of the

muscles, especially
during the wakening hours,
tonic contraction.
In
known
other words, the
tonus
as
or
the stretch,as is shown
on
livingmuscle is slightly
by the fact
that

it gapes

with

the bone

open

when

cut

or

shortens

when

its connections

tension,like the much greater

is maintained, in part at least,
set up in active contraction,
one
in the muscle, which
taking
by a continual katabolism
respires,
and giving off carbon dioxid.
In other
words, the
up oxygen
"
resting muscle is in a state of slightisometric contraction
and is doing " static work."
just
According to the principles
in such a muscle finally
enunciated,all the energy transformed
takes the form
of heat, so that, as indicated in Chapter VII
"

are

severed.

This

536

NUTRITION

and

rapidlyremoved

are

fact,it

In

the

excessive
more

is

blood

during

Since the heart

is

supply of

that

contains

is

oxygen
during work which
less carbon

sured.
en-

is not

dioxid

and

rest.

muscular

ANIMALS

abundant

an

usually true

venous

than

oxygen

FARM

OF

it is obvious

organ,

that this

in the

circulatory
activitymust add materiallyto its
metabolism.
In the performance of work, therefore,there is
and energy, not only for the work
of
an
expenditure of matter
increase

muscles,but likewise for the additional work of

and
in their experiments upon
Zuntz
Hagemann

the skeletal

the

heart.

the

justmentioned

horse

katabolism
cent

due

of the

634.

the

to

that

compute
of

work

total katabolism

the

heart

work

amounts

the

3.8 per

to

of the

body.
greater activityof

The

Respiration.

"

during moderate

muscular
exertion
would
upon
efficient aeration
provisionmade for more
increased

activityof

circulation

futile

be

consequent

the

not

were

in the

of the blood

the

respiration.Under
the stimulus of the carbon
dioxid and other katabolic
products
of muscular
which
the blood,the respiratory
enter
ments
moveactivity
increased in frequencyor depth or both, as described
are
in Chapter IV (194),thus making possiblea more
rapidgaseous

lungs through an

exchange

between

action

usually so

is

contains

smaller

the

blood

and

efficient that

proportion

of

the

air in the

the

expiredair

carbon

dioxid

lungs. This
during work
it does

than

the total

during rest, notwithstandingthe fact

that

eliminated

like
respiration,

is

much

is maintained
case

as

by

greater metabolism

Effectof

Since

muscular

in the latter that

a

As

greater.

work

the function

of
activity

lation,
circu-

in the former

action,it is true

greater

quantity

sitates
neces-

for that purpose.

upon

proteinkatabolism

already indicated,knowledge

of the

details of muscular

katabolism

is still meager.
The
student of nutrition,
however,
is less directly
interested in these details than he is in knowing
the

aggregate effect of the

of matter

and

performance of work
by the body
energy

conditions,since it is this latter which must

the feed supply. Much
effort has therefore
studies of the influence of muscular

exertion

be

upon
under
made

been
upon

the

penditure
ex-

varying
good by

devoted
the kind

to

and

WORK

of material

amount

will be convenient
the

upon

of

PRODUCTION

broken

537

in the

down

body during work.

consider the effects of muscular

to

protein katabolism

and

second

work, first

the

upon

It

katab'olis

non-nitrogenousmaterial.

work

for

the

"

for the work

itselfwas

done.
have

to

seem

of which

This

Liebig'sview, although

was

based

been

for

natural
un-

it does

experimental

performed at the expense

the muscles, causing an
in

was

of

nitrogenousby-productsand an increased
protein in the feed,while the carbohydratesand

excretion

demand

actual

any

upon

He

creased

not

the subthat
stance
suppose
and yielded the energy

consumed

taught that work

katabolism of protein in

results.
of

the

to
early physiologists

of the muscle

not

Since

muscles, by means
is performed, consist largely
of protein,it was
Early views.

635.

fats of the feed

regarded as simplyheat

were

fat

and

producing

materials.
636.

Analogy

with

engine.

between
(274^276)

VI

of itselflead
does not

do work

fuel

body

of this view.

truth

its own

by burningup

material,and

analogy drawn in Chapter

an
engine,however, might

and

question the

to

one

the

The

"

if it is well

An

engine

ing
substance but by burnthe

constructed

due

wear

to

the work

imposed upon it is comparativelyslight. It might

be reasonablyexpected, therefore,
that the machinery of the
animal body would prove to be at least as perfectly
constructed
artificialmachine

an

as

the energy
the materials
the

indeed
Carl

and

least

at

of fuel material

into work

enteringinto its
under

case

Voit, whose

normal

equallycapable of

been

was

destroying

That

structure.

own

conditions

results have

without

ing
convert-

such

first shown

fully confirmed

by

is

by
later

investigators.
Voit's
and

liberal meat

work

diet.

foUowing tables,while

two

of

on

'

on

The
the

dog alternately
resting
treadmill both when
fastingand
a

upon

results
second

are

shown

in

the firstof the

results
the average
and
Voit 2
Pettenkofer

contains

later series of similar experiments

by

man.

uber
y^ersuchunSen

Muskelbewegungen
Die

was
investigation1

doing considerable

upon

A/r

first

Ernahrung
2Ztschr.

f.

auf

den

der landw.

den

Einfluss

Stoffwechsel.

des

Kochsalzes, des Wassers,

i860.

Nutzthiere,pp. 386-388
Biol.,2 (1866),478.

Summarized

by E.

v.

und

Wolff

der

in

538
Table

151.

Number

Effect

"

II

III

1500

Table

152.

Effect

"

of

Work

Grams

258

186

14.3

\Work

872

518

16.6

[Rest

123

145

II.Q

I Work

527

186

12.3

[Rest

125

143

10.9

fRest

182

1060

109.8

I Work

657

I330

117.

140

1081

109.9

/Work
\Rest

1500

Urea
Excreted

fRest

[Rest
IV

Dog

Grams

Grams

of

Urine
Excreted

Water
Drunk

Grams

Katabolism

Protein

on

Meat
Eaten

ment
Experi-

of

Work

of

ANIMALS

FARM

OF

NUTRITION

on

412
03

164

114.1
6

no.

1040

Katabolism

Protein

of

Man

Urea

Ex-

creted

Grams

Fasting

26.5

Rest
Work

25.0

Average diet
33-6
36.8

Rest
Work
.

In

while there was

of the man,
a
great increase in the
of carbon
dioxid
and water
practically
excreted, there was

the

amount

case

in the

increase

no

meat

diet

attributes

only,
to

excretory

there

was

637.

in every

deficiencyof

direct effect of muscular

Influence

of

nitrogen. With
case

the

dog fastingor
which

small increase

non-nitrogenousnutrients

on

and

not

Voit
to

the

exertion.

non-nitrogenous nutrients.

"

While

Voit's

animal

quite in harmony with present conceptionsof the

of energy,
converter
they
a
organism as essentially

aroused

considerable

results

seem

controversy

as

to

criticism at the time

the

source

of muscular

and

led to

energy.

an

extended
The

effect

WORK

of work

the

upon

539

protein katabolism
varied

the most

under

PRODUCTION

gated
repeatedly investi-

was

conditions

with

results which

their face to be conflicting.Some

upon
increase in the excretion of nitrogenduring or
marked

observers

such

no
work, while in other investigations

The

key

these

to

conflictingresults

effect
to

seems

peared
ap-

found

following
apparent.

was

have

been

first

1879 in experiments upon the work

of feed was
horse.1 He found that so long as the total amount
performed were
ample, variations in the quantity of work
the proteinkatabolism.
without effect upon
If,however, the
by Kellner

discovered

work

increased

was

to

in

sufficient

amount

an

to

cause

falling

weight of the animal, thus indicatingthat the energy

the excretion of nitrogen in the urine
insufficient,
supply was
found that if either
increased promptly. Furthermore, it was
added
to a ration which
was
just
carbohydratesor fat were
off in the

sufficient to enable

the animal

work, the demands

upon

increased

without

to

perform

the animal

causing

any

could

increase

of
given amount
be correspondingly

in the

olism.
protein katab-

be illustrated by the followingsummary

may
experimentin which the addition to the ration consisted
This

an

in which

starch and

of revolutions

in the number

of work

the amount

of the

153.

"

Effect

of

Starch

on

Protein

Katabolism

Horse

Landw.

of

performed is expressed
dynamometer
sweep
power

used.

Table

of

Jahrb.,8 (1879),701

; 9

(1880), 651.

of

Working

effect of

The

small

even

how

determine

ANIMALS

of work

excess

of the horse

nitrogen excretion

attempted to

FARM

OF

NUTRITION

540

was

increasingthe

sharp that

so

much

in

work

could

Kellner

be

even

performed

at

of a given weight of starch or fat by increasing

the expense
the
the animal
demand
it just failed to
upon
up to the point where
in the

increase

an

cause

nitrogenexcretion

and

fall in live

weight.
number

considerable

is the

nutrients

which

katabolism

sometimes

of
that in the presence
of carbohydrateseven

that

chief

experimentshave fully
of non-nitroga deficiency
enous
of the increased protein

recent

more

conclusion

Kellner's

confirmed

of

cause

during work

occurs

sufficient amount

work

severe

and

have

shown

of fats and

especially
performed without

be

can

work
tinued
conIndeed, moderate
increasingthe nitrogenexcretion.
of days has in some
for a number
been accompanied
cases
by a gain of nitrogen,a fact apparentlyquitein accord with the
ercise.
common
experiencethat the muscles are strengthened by exIt is clear that the body normally uses
non-nitrogenous
of the energy expended in muscular
materials as the source
work,
nal
exactlyas it does in the case of the energy requiredfor its inter-

activities.

Only

when

the

is inadequate does it resort

materials
supplyof non-nitrogenous
to

the katabolism

of

of energy
for external work, precisely
as
exclusive
fastingor on
proteinfeedingas a source
source

638.

work

Gaseous

is its very
and

katabolism

effect of work

marked

effect in

upon

In

"

strikingcontrast

the excretion

of

nitrogen

the consumption of
increasing

of carbon

dioxid

and

water.

This

oxygen
increase

experienceand too well

than an illustration.
to requiremore
scientifically
obvious

fact of such

for

matter
of non-nitrogenous

increased.

exchange

the excretion

is too

the

upon

the minimal

with

of energy

(339,407).

internal work

Effectof

protein as a
it does during

an

from

common

increase

was

shown

in the researches

lished
estabThe

of Lavoisier

by the earlier experimentersin this field,such

Smith in 1859.
as
Scharlingin 1843, Hirn in 1857 and especially
The
investigationsof Pettenkofer and Voit in 1866, however,
and

confirmed

appear

to have

methods.

been

Their

the first to be executed

results

regardingthe

accordingto

modern

influence of work

upon

WORK

PRODUCTION

541

proteinkatabolism have already been cited (636) but may

with those obtained with the aid of
be repeatedin connection
the respiration
apparatus.
the

Table

154.

639.

"

Influence

Effects

Work

of

Gaseous

on

Exchange

Man

of

mon
Experiment confirms the comobservation that the increased pulmonary exchange consequent
exertion
muscular
begins almost immediately,
upon
reaches its maximum
in a very
short time and
disappears
are

promptly when
absorptionof

immediate.

the work

"

This

ceases.

is

of which
considerable
no
oxygen,
to be stored up in the body in the free state.

of

the excretion

of carbon

dioxid,more

be held in solution in the blood

slightlag in

some

In

or

lymph

case

less of this gas can

and there is consequently

of this prompt
to the
adjustment of the respiration
of work, determinations
of the pulmonary exchange by
of the forms

one

of work

of

respirationapparatus

the katabolism.

upon

The

it possible
to compare
the gaseous
work with that of the same
animal
very

carbon

sharply the

dioxid

work.

excretion

use

in

described

studyingthe

of this method

effects

renders

exchange during periodsof

at

rest

and

thus

to

additional
caused

in

mine
deter-

consumption
oxygen
amount
by a measured

and
of

The

ease

short

pears
ap-

In the

its excretion.

useful
are
Chapter VI (297-299)
especially

the

amount

view

amount
some

and

of the

true
especially

of the apparatus required,

comparative simplicity
with which the respiratory
changes can be followed in

periods,and

the fact that both

oxygen

and

carbon

dioxid

be

can

for

OF

NUTRITION

542

work

investigations
upon

of this method

use

production.
material

non-nitrogenous

normal

under

Since

of

Nature

ANIMALS

led to the extensive

determined, have

640.

FARM

muscular

conditions

katabolized.

exertion

does

not

"

crease
in-

protein katabolism, it follows that the substances

be substantially
must
oxidized for the performance of work
of
either carbohydratesor fats. If the former, each volume
dioxid given off will correspondto an equal volume
carbon
of
the

taken

oxygen
be i.o.

On

up ; that is,the respiratoryquotient (296) will

the other hand, if the material oxidized consists

quotient will be approximately

solelyof fat, the respiratory
0.7, while if both are beingconsumed, it will have an intermediate
value.
Moreover, it is comparativelysimple to calculate from
the respiratory
quotientthe proportionsin which the two are
being katabolized.
proportions of fat

Investigationsof this sort show that the

and
for the percarbohydrates katabolized
formance
within
wide
both
limits,
may
vary
groups

of work

being readilyavailable
Sources
641.

Proteins

of

of energy.

sources

for

energy

matter.

sumption
Liebig's as-

"

in
protein katabolism
contraction
muscular
the source
impliedthat the proteinswere
for many
of the energy manifested,and this view prevailed
years.

Voit,in i860,

When

increase

an

of

work

muscular

non-nitrogenous

vs.

(635) of

as

the

(636)that muscular

showed

exertion

is not

any material increase in the protein

seemed
unavoidable
that non-nitrogenous

necessarily
accompanied by
katabolism, the inference
materials
This

were

the

main

of

sources

muscular

energy.

radical to be at once
too
conclusion,however, was
in opposition to Liebig'sauthorityand numerous

cepted
ac-

genious,
in-

but not

alwaysconvincing,
hypotheseswere advanced to
explainthe observed phenomena on the assumption that the
of the energy expended.
proteinswere, nevertheless,the source
642.

Fick

however,

and

Wislicenus'

experiment.

"

The

first

attempt,

comparison of the work performed

quantitative
with the energy
available from the proteinkatabolized during
licenus
its performance was
the famous
experimentof Fick and Wisx

at

in 1866.
1

These

observers

Naturf.
Vrtljschr.

made

an

ascent

Gesell. Zurich, 10, 317.

of the Faul-

tained
quantitiesof fat and glycogencon-

the small

from

Aside

work.

ANIMALS

FARM

OF

NUTRITION

544

the energy for work under these conditions

derived only from the proteins
or their cleavage

in the meat

could have

been

products.

These

to

largeextent

results show
a

as

of muscular

source

be used

clearlythat proteinmay
but

energy,

it is

theless
never-

particularly
is,as already
with farm animals, the
energy
of the feed.
stated,through the non-nitrogenous ingredients
It is by no means
however, that a certain amount
impossible,
in a muscular
tion.
contracbe necessary
of proteinkatabolism
may
of the protoplasm of
Such a contraction is a function
fibers and it is conceivable that a portion of the
the muscle
of the proteinsand nucleoarisingfrom the katabolism
energy
appearingas heat in the
proteinsof the muscle and ordinarily
off,so to speak,to aid in producing
restingmuscle may be switched
In other words, it is possiblethat a
the contraction.
in order
be necessary
certain level of proteinmetabolism
may
that

true

under

ordinary conditions,and

supply of

main

the

maintain

to

the

of

potentialenergy

Such

would, of

fact

the amount

of

the matter

"

2.

non-nitrogenous materials
have

course,

The

While

Body

an

Efficiency

the

of

important bearing upon

the energy

expended

Body

as

at present

Motor

results

is immediate

substance

work.1

into

proteinrequiredfor a working animal, but

belongsin the realm of speculation.

General
645.

transforming

for

conditions

favorable

most

in work

of

source

productionis

energy.

of

course

"

rived
de-

source,
ultimatelyfrom the feed consumed, its immediate
of body substance, and
stated in "i (630),is the katabolism
as
of labor in the
amount
animal may
an
perform a considerable
It will aid
fastingstate at the expense of stored-up material.

in the

discussion

of the

of the animal
as
efficiency
the
with which
efficiency
to inquirewhat
i.e.,

material
1

katabolized
Compare Armsby,

complicatedquestionof the
to consider first the
a prime motor
body utilizes this stored-up energy,

somewhat

percentage of the total energy

for work

production is

Principlesof

Animal

Nutrition,

of the

recovered
pp.

207-209.

body
in the

WORK

PRODUCTION

545

done, deferringto the followingsection a study of the

of the animal as a converter
of feed energy into useful
efficiency
work and of the feed requirementsof work animals.

work

Mechanical

646.

regardedas

efficiency of muscle.

machine

mechanical

into

in somewhat
the
efficiency
engine or an electric motor.

the

muscle
does

is recovered

lifts a

0.2344 X
in the muscle

-f-

0.2344

Much

one,

0.4688

ten

therefore,speak"of its
as

of

of that

grams
of

if

Thus

done.

steam

If the increased katabolism

to be

shown

were

muscle

50 per cent, that

would be recovered

is,50

per cent

has

devoted

been

machine.

io"4

would

be

work.

study of
complicated

the

to

subjectis

The

of the total

mechanical

as

caused

0.4688

the

as

isolated

an

through one centimeter, it

work, equivalent (308) to

of
efficiency

experimentalwork

singlemuscle
and

sense

by its contraction

mobilized

energy

the

weight of

calories, the

gram

same

in the work

centimeters
gram
io-4 gram calories.

10

may,

energy

in this sense
is meant
By efficiency
the total energy mobilized during a contraction

proportionof
which

one

be

may

of chemical

for the conversion

and

work

muscle

"

unanimityof

nism
views upon it,especially
to the mechaas
been reached.
contraction,has by no means

of muscular
As

of the muscle as a converter

of energy,
regardsthe efficiency
fact is perfectly
well established,
however, one
viz.,that it
varies within quitewide limits,dependingespecially
upon the

load

as

related

of

shortening.

647.

the

to

Mechanical

capacityof

efficiencyof

the muscle

the

body

as

of energy
mobilized in each muscle
performanceof a certain form of work were
amount

and

the degree

upon

whole.

"

concerned

known,

If the
in the

it is

ceivable
con-

that,assuming each muscle to act with its maximum

theoretical efficiency
an
efficiency,
might be computed
average
for the whole

of muscles.

group

The

conditions

for the

imum
max-

of a muscle, however, seldom or never

obtain
efficiency
in the working animal, and
consequently this hypothetical
is not
attained.
Of its many
serve
efficiency
muscles, some
largelyor wholly to maintain the relative positionsof the
different parts of the body, i.e.,
their contractions are isometric
(629) and consequentlyhave an efficiencyapproachingzero.
Others
the

contract

maximum.
2

to

Some

varying extent

and

muscles, owing

to

under

loads less than

their anatomical

re-

546
lations,work
vary with
of labor

which

the

by

of

notably

at

animal

advantage than others,while

secondary activities,
(633,634),
respiratory
organs
various

their share of energy

consume

the

to

sets

up
and
circulatory

the

work.

of the

performance
of these

Some

nature

of these

extent

and

severityof

the

ogous
analloss of energy
are
of a heat engine,
the cylinder
of

sources

comparable with

are

the

the

and

work,

rectly
di-

contribute

and yet do not

losses from

to the radiation

while others

with

varies

secondaryactivities

ANIMALS

group of muscles is called into action will

of the work.
Moreover, the performance

nature
an

FARM

less mechanical

to which

the extent

the

OF

NUTRITION

the internal resistances of the

engineitself.
afford

adequate

no

fore,
muscle, thereof
of estimating the efficiency

of the
efficiency

of the

Determinations

means

isolated

rect
body as a whole and the latter must be determined by diis made
by causing
experiment. Such a determination
of work under conamount
ditions
the animal to perform a measured
either directly
which also permit the measurement,
as
of indirect calorimetrydescribed in
heat or by the methods
Chapter VI, of the total body energy metabolized.
and Cathcart 1 upon
in experimentsby Benedict
Thus
a man
through the
ridinga bicycleergometer, the subjectbreathed
universal
respirationapparatus
mouthpiece of a Benedict
of which the oxygen
bon
consumption and the car(298),by means
the

data

could

elimination

dioxid

in the

metabolized

of energy

the amount

these

From

determined.

be

body

was

puted
com-

of mechanical
work done
compared with the amount
measured
as
by the ergometer. For example, in one of these
piratory
tests the energy
output per minute as computed from the resand

exchange
done
1.02

per
-7-

minute

6.32

recovered

as

equivalentto

was
1

6.32 Cals.,while the mechanical

was

6. 1

per

useful

In other

words,

of the

total energy
output
the remainder
taking the form

cent

work,

Cals.

1.02

work

was

of

heat.
648.
the
of

Gross

net

efficiency.
"

Comparisons

like that of

precedingparagraph give what is called the gross efficiency

the body, i.e.,
they show what proportionof the total energy

metabolized
It is
1

and

during

analogousto

Muscular

Work;

work

the

is recovered

of
efficiency

Carnegie Inst,

an

in the

useful work

engineas computed

of Washington,

Publication

No.

done.

from

187 (1913)-

PRODUCTION

WORK

comparison of

the

But

the brake

horse

body katabolizes

547
with

power

"

requirementfor

the support of its internal

quired
rerespects to the energy

subjectof Benedict
experimentproduced during rest (lyingon a

Cathcart's

couch)

1.09

sumption.
con-

and

matter

(341,342) analogous in some

to run
an
engine without load.

and

steam

liberates energy
for
the performanceof external work,
it
i.e.,

other purposes
than
has a maintenance
work

the

The

If this maintenance
Cals. of heat per minute.
quirement
rebe subtracted
from the total energy
output during

there is left 5.23 Cals.,as the additional energy

requiredfor the performanceof the 1.02 Cals. of measured
work

work.

in this way

Computed

of
efficiency

an

1.02

output
nal
exter-

5.23

-?-

19.5 per cent results. This has been called the net efficiency.
It shows
the utilization of that portion of the energy
output
=

which
the

is

in the

expended

production of external work

forms

of internal work

requiredfor

physiological
processes
as

distinct from

the various

included in the maintenance

requirement.
arises in
difficulty

In

in this way
computing the net efficiency
Thus
in an
decidingupon the proper deduction to be made.
subtract from
the
experiment like that just cited,one may
total energy
output of the body during work, not only the

expenditure for maintenance

during rest but likewise
energy
the ergometer and causing it to rotate
that caused by sitting
on
without

load,and the remainder

metabolized

for

the

more

when

at

energy
rest.

requiredthe
Cals.

per

minute

work, from

performance
being 6.32 Cals.
that the
minute

per

The

be
of

total output of energy

work, it was determined
Cals.

may

same

when

regarded as the

the
per

useful
minute

energy

work.

The

during the

subjectmetabolized
1.13
riding without load than

load in the work

added

fore,
experiment,there(1.09-f-1.13) 4.10
expenditureof 6.32
for the performance of
Cals.
of
1.02
=

"

of 24.9 per cent may

be
efficiency
in experiments with the work horse one
Similarly,
subtract the energy
expended during horizontal locomotion
which

an

of that metabolized
with
1

the useful work

For

compare

Reach,

duringrest

Biochem.

and

Cathcart's

Ztschr., 14

compare

may
stead
in-

the remainder

done.1

discussion of the various

Benedict

and

puted.
com-

base

lines for the computation of efficiency,

publication already cited, pp. n 2-136, and

Jahrb., 37 (1908), 1053.
(1908),430; Landw.

also

548

NUTRITION

This

of

method

the engineer,and
is accustomed
a

FARM

estimate

usuallyemployed by
any
criticized
it
has
severely. The engineer

the losses due

of brake

power.
horse power

comparison
exists,however,

of the

animate

is available.

It is

for

and

resistance of

indicated horse

determining the indicated

if indeed

motor,

conception,and only

the internal

to

horse power

method

No

ANIMALS

computation is unlike

Schreber

to

engineby

an

OF

it

the method

of

of any

permits

comparison

responding
cor-

lined
justout-

if the

mated
engineer had no indicator and estithe efficiency
of his engine by deducting from the total steam
the engine empty
and compared
consumption that required to run
the

remainder

the

animal, however,
the

If,on

with

basis

as

the external
like that

work

The

done.

internal

work

of

engine, is largelymechanical.
in locomotion
computation of the efficiency

of Zuntz's

of

the

that this internal work

is performed
be assumed
(662),it may
with approximately the same
the external work, then
as
efficiency
the net efficiency
of the animal
will be somewhat
analogous to the
of the steam
in the cylinderof the engine.
efficiency

649.

while

Gross

the

efficiencyvariable.

efficiency
may

net

under

considerable

will vary

subjecthad
13.7

cent

per

Total energy

observed

if

Thus
as

gross

of work

much

(load)to

done

Benedict

and

with
per minute
would
been
have
efficiency

instead of 16.1 per

work

Cal. -5- 0.195

maintenance

1.09

Cals.

Total

3.71

Cals.

0.51

Cals.

in work

work,

0.51

"

done

efficiency
to the

point at

which

of the

has

for

efficiency
beginsto be affected,
increase with increasing
load as Benedict
This
experimentallyto be the case.
requirement

Arch.

the

upon
is identical with

that

in connection

with

of energy
already been called

material

cent

the net

maintenance

of the utilization
attention

2-62 Cals.

13.7 per

the gross efficiency

will
and
Cathcart
show
influence

only

expended per minute.

For

Up

the

cent.

mechanical

Gross

tenance
main-

Cathcart's

For

Recovered

that

stant
conregarded as substantially
ciency
variety of conditions,the gross effi-

done, only half

his
efficiency,

net

same

be

be

with' the ratio

requirements.

It should

"

products. If

the

159
Physiol. (Pfluger),

useful

work

(1914),276,

tion
computato

which

zation
the utili-

performed

WORK

be reduced
the gross
650.

to

of
efficiency

work

the

animal

example

549
in horizontal

also becomes

course

locomotion,

zero.

The

figuresfor either gross or

the efficiency
for the time during which the
Since,however, it is not practicableto stop

Efficiency per

show
efficiency
is being done.

net

for

as

zero,

PRODUCTION

machine

day.

"

the

when

demand

for work

ceases,

the

the degree to which the

for the entire 24 hours, i.e.,
efficiency
body energy is utilized in practice,will evidently vary with
of hours work done per day. Thus if Benedict and
the number
8 hours per day at
Cathcart's subject had been able to work
his gross efficiency
the same
rate as in the experimentjustcited,
for the 24 hours

would

been

have

as

follows

"

Energy expended
480 minutes

work

960 minutes

rest

Cals.

"6.32
@

1.09

Cals.

Cals.

3034

1046 Cals.

4080 Cals.
Work

done

480 minutes

Cals.

1.02

Efficiency
per day

Cals.

490
12.01

per

cent

hand, if he had worked only one hour per day,

it may
be presumed that both the net and gross efficiency
of the
work
would
have
been
production during the hour of work
the same
but the efficiency
for the 24 hours would
substantially
have been much
lower,viz.,
On

the other

Energy expended
60 minutes

1380 minutes

work
rest

@
@

6.32 Cals.
1.09

Cals.

379

1504

Cals.
Cals.

1883 Cals.
Work

done
60 minutes

work

1.02

Cals.

Efficiency
per day
In

discussions

61 Cals.
3.24

per

cent

of the

of a man
animal as a
an
or
efficiency
in comparisons with artificial motors,
motor, and particularly
it is essential to distinguish
whether the net, or the gross
clearly
is meant
and likewise to base the comparisons upon
efficiency

the

Since

apparently less
affected than the gross efficiency
by variations in the intensity
of the work, it appears
and duration
to be the most
logical
performance

per

day.

the

net

is

comparison in the case of

convenient
in practice.

the most
651.

the animal

on

inclined

an

of ascent

work

into work

separately.The
addition

other

of

which

being

as

course

work, such

as

it

tally
horizon-

performs

locomotion, work

for
efficiency

is of

same

of

forms

the

and

well

draft,either

work

The

plane.

therefore be subdivided

draft and

without

or

as

quadruped performs work

"

locomotion, with

of

or

total work.

of

Analysis

means

may

ANIMALS

FARM

of

method

by

OF

NUTRITION

550

each

form

of

puted
com-

of man,
but in
true
turning a crank with

(stationarybicycle)or liftinga
of analyzing
weight directlymay be performed. The method
worked
of a quadruped has been
the work
out
especially
by
hands

the

with

or

the

feet

and
conveniently illustrated from Zuntz
Hagemann's investigationson the work horse.1 The methods
of indirect calorimetrywere
used, carbon dioxid production
Zuntz

and

be

may

consumption being determined with the Zuntz apparatus

calculated.
and
the
(279)
correspondingenergy output
done
The
work
a
was
specialtread-powerlocated in the
upon
air,and during the rest experimentsthe animal likewise
open
The inclination of the platform of
stood in the tread power.
the power could be varied,and it could also be driven by a steam
engine,so that by settingit horizontal the work performed by
and

oxygen

the animal

reduced

was

traveled

was

to

measured

that of locomotion

by

revolution

alone.
counter

The

and

tance
disin the

draft the animal

pulledagainst a dynamometer.
The
apparatus used is illustrated in Chapter VI, Fig. 33 (313).
This is an
locomotion.
652. Horizontal
important factor
in work
siderable
production, since it requires the expenditureof conof the body at each step
energy in successive liftings
thus
and
the overcoming of internal resistances.
The energy
chanical
expended does not ultimatelyproduce any work in the meThe work of locomotion,
as heat.
sense, but all appears
cident
innot useful work
therefore,is in a sense
althoughnecessarily

experimentson

"

the

to

If the
the

performance of

tread

total energy

power

be

output

set

by

be

the

work.

horizontal
the

subjectwill

called,by analogy with

expenditure in locomotion.

the gross

from
during rest (standing)

the

iLandw.

Jahrb.,18 (1889),1;

and

driven
measure

motor,

what

may

(648),the gross
efficiency

Subtractingthe
23

by

total output
(1894),125;

energy

output

during locomotion

27, Ergzbd. Ill, (1898).

OF

NUTRITION

552

performance of

be

to be

0.004395

0.107041

follows

as

Since

"

of ascent, the two

"

0.4035

cals.

1.0795

cals-

of

and

be

can

computed

"

0.3746 cals.

the

that

values

is

kilogram meter

one

of work

formulated

equationsthe

these

ANIMALS

kilogram meter

one

followingequations may

From

FARM

6.5856 cals.

equivalentto

efficiencyin

net

cals.,it

2.344
work
of

the

ascent

lows
folwas

the net efficiency

In effect,
in
35.73 per cent.
2.344 -1- 6.5856
is computed by deductingfrom the total energy
of ascent
work
and
for horizontal
expended for maintenance
output the amounts
=

ured
comparing the remainder with the measresults given in the previousparaThe
graph
computed
expended in locomotion were

and

locomotion

of ascent.

work

for the

energy
this scheme.

according to

of draft.

Work

654.

by

the average
of maintenance

On

excess

traveled

meter

The

similar method.

horizontal.
in

"

in
efficiency

The

draft

was

puted
com-

tread

set nearly
was
power
of sixteen trials the total energy output
per kilogram live weight and per

of ascent
1.5021 cals.,the work
0.0051 15
and the work of draft 0.153127
kilogrammeters.
the energy
expended in the performance of 1

was

kilogrammeters
Lettingz equal
kilogram meter
be formulated

net

of work

of

draft,the followingequation

may

"

y +

0.005115

0.153127

1.

5021

cals.

Substitutingaverage values for x and y, the value of

of 32.84 per cent.
equivalentto a net efficiency
7.143 cals.,
Correction

655.

it was

found

that

speed.

for

the

"

In

experimentsmade

expenditureof

The

or

draft,it is

method
of which
1

to
necessary
of doing so is a method

need

not

be gone

Compare Armsby, Principles of

is

walk

increased
energy per meter
is illustrated by the averages

materiallyas the speed increased,as

alreadycited (652)and is shown
paragraph (663). In computing
ascent

at

more

the

take
of

fullyin a
of
efficiency
account

ing
succeedwork

of this fact.

approximation, the

into here.1
Animal

of

Nutrition,pp. 507-508.

tails
de-

PRODUCTION

WORK

Summary.

656.

separatedin

be

Table

155.

the

Net

"

from

the

the

efficiencyin

net

done

the total work

into which

of work

forms

summary

is apparent

As
body of the horse as a motor.
foregoingexplanations,the table shows

the

the various

of
regarding the efficiency

results

Hagemann's

and

of Zuntz

following table contains

The

"

553

can

justdescribed (651,654).

manner

Efficiency

of

Horse

the

Forms

Different

in

of

Work

Work

at

Walk

Net
Effi-

Expenditure
Energy

Net

kgm. work

without
of 2.91 miles
Speed of 3.36 miles
Speed of 3.66 miles
with
The same
Speed of 3.36 miles

load:
per hr.
per hr.
per hr.
load on back
per hr.

Relative

Trot

Net
Effi-

Expenditure
Energy

ciency

of

Kgm.

cals.

Kgm.

6.8508
6.9787

2.9116
2.9660

34-3

7-36471

3-1300

6.502

2.7634

36.2

75190

3.1960

31-3

7.4240

4-393"

22.7

10.0780

10.3360

of Chauveau's

into

dextrose,with
its energy,

31-96'

33-7

3-iSSo1
4.28202

3i-7

'

23-4!

0.3256

0.5478 "

0.3666

0.3929

0.3914

utilization

view

0.6007 '

of

fats

and

theory 4 that fat

the

elimination

before it

can

as
serve

carbohydrates.
first be

must

heat

of

for the

In

"

converted

considerable

as
directly

it becomes
physiological
processes,
relative extent
the energy
to inquireto what
carbohydratesis utilized in muscular work.
energy

Slow

cals.

'

portionof

10.7% grade
18.1% grade
For 1 kgm. work of ascent, with load:
iS-8% grade
For 1 kgm. work of draft:
1.5% grade
8.5% grade
Locomotion
per meter
per kg. mass

657.

at

of ascent, without

load

Speed

Net

ciency

of

For

Work

of

source

of much

terest
in-

of fats and

and

Hagemann's extensive investigations,

particularly
those upon the horse,there were
very considerable variations
'The
the proportions of fat and carbohydrateskatabolized.

In Zuntz
in
in

individual

trials in

the

which

Singleexperiment.

Independent of speed.
Compare Armsby, Principlesof Animal

Two

same

kind

experiments.

of work
Work

Nutrition,pp.

was

per-

probably excessive.

153-154

and 399-405.

in many

also show

formed

FARM

OF

NUTRITION

554

this,however,

similar variations.

cases

the

ANIMALS

percentage of energy

standing
Notwithutilized did

tion
materiallyin these instances and there is no indicavary
be expectedaccordingto
of any such differences as would

not

theory.
question has also been

Chauveau's
The

in

associates

his

and

investigateddirectlyby
experiments on dogs and on man.

experiments,the

these

feed

consisted

Zuntz
In

largelyas possibleof

as

carbohydrates or fat,respec(protein,
tively),
its
that
metabolism
the
at
was
so
largely
body
expense.
method
of investigationwas
the same
as
substantially

the nutrient

The

be

to

has

that which

tested

metabolism

energy
Table

Proteins

156.

described.

The

kilogram and

meter

Nutrients

for

just been
per

Comparison

"

of

final results for the

traveled

Work

were

"

Production

only

Chiefly fat
Chieflyfat (body freed from carbohydrates by

phloridzin)
Much

sugar

with

Much

sugar

and

The

proteins
little proteins

differences

are

quitesmall,while,as
for energy

2.6 cals. represent the demand

pointsout,

Zuntz

per unit of work

if

when

it should,accordingto Chauveau's
carbohydratesare the source
clusively
theory,rise to about 3.68 cals. when the energy is derived exfrom

Later

and

conclusion.
and

fat.
elaborate

more

Atwater

Benedict

and

and

Cathcart

experimentson

man

led to the

Benedict,1 Benedict
3

also

report

and

same

Milner,2

experiments upon

which, while not regarded as conclusive,indicate a possible

of
fats
all
but
with
not at
one
slightinferiority
comparable
that demanded
by Chauveau's
theory. On the whole, then,

men

U.

S. Dept. of

U.

S. Dept. of

Muscular

Work

Agric, Office Expt. Stas.,Bui. 136 (1903), 182.

Agric, Office Expt. Stas.,Bui. 175 (1907), 234.
;

Carnegie Institution

of Washington,

Publ. No.

187 (1913),145-

PRODUCTION

WORK

the

conclusion

if any
difference exists
of fats and of carbohydratesit

that

warranted

seems

in the utilization of the energy

is too small

be of much

to

practicalsignificance.

Conditions

efficiency
affecting
the
from
appears
of the animal
efficiency

Efficiency varies.

658.

(656),the

summary

As

"

net

is comparatively
high in the

motor

exceedingin
broad

instances

most

555

of the

case

about

that with this animal

generalway

body as a
horse,considerably
be said in

It may

cent.

30 per

foregoing

of the

one-third

(i.e.,
of maintenance
or
plus locomotion)is
It is also evident,
in the mechanical
work
done.
recovered
however, that the organism has no one fixed degreeof efficiency
wide range
but that the latter may
through a somewhat
vary

for
energy metabolized
of maintenance
in excess

under

form
specific

exertion

of muscular

different conditions.

grade, with

the

of

ascent

moderate

grade

cited
a

up

vestigat
yet in-

of work

Of all the forms

draft.

without

or

The

"

done

work

largelyto

experimentsthus far
by walking horizontallyor

done.

of work

Forms

659.

refer

or, in other

words,

which
one
body by the legs,appears to
of over
is performed most
36
economically,a net efficiency
This
for man.
per cent being reported both for the horse and
percentage, however, decreases considerablyas the angle of
is increased.
For horizontal locomotion,
ascent
as alreadynoted
of 35 per cent.
Draft up a
(652),Zuntz computes an efficiency
in the
less efficiently
slightgrade was
performed somewhat
of the horse, the percentage being approximately 31,
case
done with an efficiency
while draft up an 8| per cent grade was
be the

of the
the lifting

of less than

Other

23 per

forms

cent.

of work

to

appear

be

performed

Thus
degree of efficiency.
experiments on
done by turning a crank with the
work was

decidedlylower
done

on

Benedict

efficiencies than

treadmill.
and

Cathcart

The
on

same

man,

those
was

true

in which

the work

the

work

was

Loc.

cil.,
p.

125.

was

done

figurescomputed
stationarybicycle,the maximum
of
from
net efficiency 6 subjects
ranging
20.4 to 25.2 per
1

the

experimentsof
:

less

shown

have

hands

in the

in which

man

in which

with

upon
for the
cent.

556

difference

The

Species.
"

of the human

and

body

differences in the

to

is done

ascent

finds that
the

660.

the work

of

traveled

show

example,Horse
expended

locomotion

utilized energy

but,
back

the

to

energy

locomotion

per

0.53 to

kilogramlive weight.
Hagemann's
experiments

than

differences between

in conformation.

given load

did Horse

on

No.

For

his back

with

Horse

III.

No.

to

slightlyless
a

extent

than

less
considerably

extent

hand,

like No.

XIII,

No.

ing
Ill in ascenddraft

in horizontal

carried

economicallythan No. III.

possiblebearingof these facts upon

breeding opens

661.

Training

unfamiliar

up.

and

any unaccustomed
than it is later.

an

load

his

on

This

motions

of work
is due

more

exercise

of

of

group

is much

in part to

of

speculation.
experiencethat

familiar

fatiguingat first
the fact that in making
more

of muscles
called
are
groups
accessory
skill has been
later when
more
necessary

activitythan are
acquired. The experienceof
example

It is

fatigue.

form

questionsof heredity

field
interesting
"

into

cellent
bicycleis an exthis.
In the second
place,however, simple
crease
of muscles in a particular
to inseems
way
a

learner

on

the

their average
mechanical
efficiency.
This effect may
be illustrated by the results
of

former

horse,viz.,about

differences

carried

XIII

the

about

more

The
and

in horizontal

and

Zuntz

due

that

Klein

to

the other

on

but

of

and

grade

work

than
Horse
No.
Ill in horizontal
energy
less in trotting.No.
walk
but
II likewise

more

at

does

horse

individual
interesting

No.

expenditure of

performed, since the

distance and

meter
"

animals, presumably

II

than

Individuality.
the horse

efficiency
is evidently
due largely

in both cases.
equal efficiency
is performed by the ox with
ascent

calories,
per

0.55 gram

less

of work

kind

the

about

with

unit of distance

between

that of the horse

as
efficiency
by the
expends much more
energy

animal

ANIMALS

just noted

net

same

upon

FARM

OF

NUTRITION

experiments by
the

Gruber

carbon

dioxid

hill climbingthe amounts

Series I
Hill
Hill

upon

himself

excreted

excreted

in

he

which

termined
dein

Thus

during work.

in twenty

series

of two

minutes

were

"

climbingwithout
climbingafter 1

^entbl.

practice
days'practice

40.98 grams
....

Physiol.,26 (1912),722.

32.22

grams

WORK

Series II

PRODUCTION

557

Hill

climbing without practice

Hill climbing after 14 days'practice
That

the less

of accessory

38-83 grams
.

muscles

31.00

is not

grams

the

only cause
is indicated by experiments
of this increase in efficiency
upon
shown
that
w
hich
have
the gradual strengthening
convalescents,
of the muscles
results in a more
economical
performance of
their work, largelyindependent of any
specialtrainingfor a
use

kind of work.
particular
Conversely,fatiguehas

been

shown

by

materially increase the relative amount

unit of work.
the
Schnyder 1 summarizes

of metabolism

to

that it is
which
which

the

work

determines

the

not

have

to

seems

the relation of muscular

662.

Intensity of

itself but

Work.

results

quired
re-

sion
conclu-

regarding

metabolism.

to

It has

"

effort

metabolism,

anticipatedHill's
tension

per
in the ment
state-

matter

the muscular

of

amount

observers

numerous

already been

shown

(649)

that the gross efficiency

of the body tends to increase with the
of units of work formed
intensityof work, i.e.,with the number
perin a unit of time, for the reason
that the proportion
of the total energy
expended which is devoted to useful work
increases.
On the other hand, common
observation
tends to
show

that this

be true

only within limits,and that excessive

is performed uneconomically.
intensityof the work may be increased by increasing

work

The

can

either the

speed, the load moved, or

be anticipated,
therefore,that

would

of these factors would

one

663.

Influence

locomotion
evident.

of

speed.

involves
The

travelingthe

one

In

the

horizontal

case

or

with

increased

Ztschr. Biol.,33

at

energy

per

vastly
a

efficiency.

net

and

to

(1896),289.

more

about

than

results

Hagemann's

increased

an

kilogram weight and

speed of

energy
rate.

(652) show

walk

is

energy

travelinga mile

moderate

speed, while locomotion

distinct increase up
1

horse, Zuntz

locomotion

expenditure of

at

It

increase of any

the track athlete

distance

same

of the

of ascent.

great speed in horizontal

top speed obviouslymetabolizes

at

undue

diminished

That

"

an

angle

largelyincreased expenditure of

horse

race

result in

the

meter

at

trot

7^

miles

net

distance

showed
an

on

no

hour.

Jour. Physiol.(London), 42 (1911), 1.

558
Much

study has
The

man.

Brezina

of the

influence
be

to

seems

of

the net

to be true

was

done

the

at

results

in

tial
exponen-

work

of ascent

that

the expenditure in
upon
available.
With
are
man,
found
power
distance in walking

experimentson

meter

of

tread

speed

just indicated.
been

found

velocities

at

Since

this

that both

the net

and

efficiency
the

speed.
efficiency

gross

be
In

much

siderably
con-

found

was

nearlyindependent of

high speeds,increasingvery

of

According to
increase of speed in

gait.
"

(656)an

of

gait from

in the net

energy

moderate

limit

up

to

on

the contrary, Benedict

Such

certain

tions
speed only slightvaria-

in
efficiency

horse

walk

to

expended

weight and distance,although an

in

the

as

an

crease
in-

regarded as
of the

none

being
portant
im-

an

rapidly

more

speed.

increase

than

it became

their

Influence

at

that

beyond

locomotion,it follows that the

Cathcart1

on

change

marked

experiments cited was there any air resistance,the work

In actual practice this is
a
stationary apparatus.

mann's

by

that

bicycleergometer, which might

draft, decreased as the speed increased.

of

664.

the
3

also have

must

factor
than

found

minute)

net

than

the maximum

and

done

on

clearlya

unit of locomotion

per
2

per

also of horizontal

Benedict

various

shows

kilogram and
independent
substantially

of ascent

form

per

the

upon

Reichel

expenditure

grade

in work

meters

upon
less marked

and

below

and

speed

results

No

Brezina, Kolmer

in work

energy
Reichel

while below
velocity,

much

locomotion.

Murschhauser

shown.

were

The

horizontal locomotion
in
upon
literature of the subject as
marized
sum-

expended

speed (about 80

function

up

and

ANIMALS

expenditure of

increases.

maximum

FARM

extensive

Benedict

in the net

speed

been

somewhat

by

that

OF

NUTRITION

causes

and

no

by

extent

is lifted

at

motions,

such

brought

into

are

further

doubtless

step and

swinging
play.
as

Carnegie Institution

Biochem.

Biochem.

Carnegie Institution

the
the

Washington, Publication
(1914),170.

of

notable
unit of

trottingspeed
With

man,

tion
find that locomo-

economicallyin running

able
brought about to a considerheight to which the body
degree to which extraneous
in rapid walking, are
arms

of

Ztschr., 63
Ztschr., 65 (1014), 16 and

per

increase.

Murschhauser

differences in the

each

involves

in the

increase

Hage-

the horse obtained

for locomotion

given speed is performed more

walking.
differences

trot

and

Zuntz

No.

231

No.

187 (1913),138.

(1915),pp.

12-28.

35.

Washington, Publication

560

FARM

OF

NUTRITION

ANIMALS

Net

Grade

Efficiency

io%
20%

39%
31%
27%

30%

"
is the

As
needs

available

Requirements

Feed

in

case

be

to

3.

feedingfor

supplied with

form

with

and

proteinsand

ash

The
667.

the

Functions

Work

for

the

other purposes,

adequate

an

certain

working animal
of energy

amount

forms
specific

of

in

matter, particularly

ingredients.

requirementsof

of protein.

As

"

matter

shown

in

Chapter IX

(418),

per

nance
dailyprotein requirementof the horse for simple mainteis apparentlyabout
the same
that of other farm
mals,
anias
crude protein
0.6 pound of digestible
viz.,approximately,
1000
pounds live weight, although the experimentaldata

are

rather

scanty.

"

of this

pended
chapter(641-643)that the energy exin muscular
work
is practically
derived from the katabolism of carbohydratesand fats,the protein katabolism
being
of
unaffected by work
so
long as an ample supply
enous
non-nitrog-

It

in

appeared

nutrients

is available.

One

might

at

first

thought be

inclined to

conclude,therefore,that the simpleaddition of nonration would

suffice to
nitrogenous material to a maintenance
enable it to support a correspondingamount
of work production
and that a maintenance
ration of digestible
protein would also
be a sufficient supply for the working horse.
Such a conclusion
It is quite conceivable
would, however, be premature.
that

in order

to

maintain

the muscle

as

an

efficient instrument

for

converting chemical energy into mechanical work, a higher

be necessary
than is required
plane of protein metabolism
may
it in nitrogen equilibrium when
to support
doing no work,
while the possibility,
for example, of a favorable influence of
abundant
an
protein supply on the blood circulation and on
the

nervous

system

should

the demand
especially,
high level. Since the

introduced

into

the

of
blood

body,

not

be

the

animal

is the
an

overlooked.
for

vehicle

adequate

In

oxygen
by which
stock

of

fast work
reaches
oxygen

blood,

is
or

561

PRODUCTION

WORK

be necessary and
of haemoglobin, would
particularly
to assist in securing this.
liberal supply of protein seems
more

conjecturesshould
role
not
insignificant

any

of these

may
work

play

without

evidence

any

If

the proteins
to be true
prove
in the production of muscular

of the

fact

appearing

in the total

nitrogenexcretion.
specificinvestigations
regarding the minimum
proteinrequirement of the work horse
been
to have
seem
made, but the extensive experiments of
and
others referred to on
Wolff, Grandeau, Muntz
previous
668.

The

protein requirement.

afford numerous
pages
results were
obtained

althoughin

instances
from

none

"

in which

rations

of them

No

was

entirely
satisfactory
comparativelylow in protein,
the

supply reduced

to

the

tensive
requirement. Similarly,in Langworthy's exx
in
of
rations
fed
amples
expractice,numerous
compilation
of low protein rations are
to be found.
fact,it would be difficultto compound from ordinaryfeed-

maintenance

In

stuffs a ration sufficient to support any considerable amount

of
work
without
protein than is presumably
introducingmore
Such
required for simple maintenance.
being the case, the
principalpoint to be taken into consideration is the effect of a
reduced proteinsupply upon the digestibility
of the ration (723normal
725). Any ration carryingsufficient protein to ensure
furnish ample protein for work
doubtless
digestion would
production in all ordinary cases, with the possibleexceptionof
work at high speed. A nutritive ratio,computed in the usual
(709),of 1 : 10 or 1 : 12 would unquestionablyensure
ample
way
for
and
for
slow work,
protein
probably
moderatelyrapid work
of man,
is well known, experienceor traalso. In the case
dition
as
have led to the general employment of high protein
rations by athletes. On the other hand, however, Chittenden 2
has shown
that the protein supply of athletes and soldiers,
as
of sedentaryoccupations,may
well as that of men
be reduced

below

much

the usual level without

these

however,the
experiments,

than

the amounts

which

for the maintenance

or

in
Even
efficiency.
proteinsupplywas much higher

loss of

experiments have shown

of nitrogenequilibriumin man
recent

doingonlylightwork.
1
2

U. S.

Dept. Agri.,Office Expt. Sta.,Bui. 125 (1903).

Physiological
Economy in Nutrition,1904.

to

fice
suf-

at rest

562

requirements.

Ash

669.

As

"

ANIMALS

FARM

OF

NUTRITION

in

pointed out

ters,
previouschap-

requirementsof an animal deserve greater

than they generallyreceive,while in the case

the ash

growth, at least,the

So

is necessary.

in the ration

however,
of these

far

sufficient data

no

the

as

working horse
available

seem

of

substances

of certain accessory

presence

sideration
con-

for

is

cerned,
con-

cussion
dis-

topics.

The

energy

requirement

preceding
of
between
the efficiency
made
section certain comparisonswere
and that of artificialengines.
the animal body as a prime motor
but includes
The animal body, however, is not only a prime motor
Economic,

670.

or

also the furnace

over-all

efficiency.

In

the

in which

the fuel is burned

and

"

complete power plant,such

example, rather than an engine.

in this
for

Just

respect

the energy

as

of the

fuel of

as

locomotive

resembles

locomotive,
is

subjectto

incompletecombustion and to radiation of

the cylinders,
reaches
so
heat before the steam
portionsof the
of the feed escape in the excreta
or
are
expended in the
energy
various processes incident to the formation in the body of those
katabolism
substances whose
yieldsthe energy for a muscular
certain losses due

to

contraction.

expendituresare largelyunavoidable,
they constitute part of the energy requirementsof the work
of
animal, and from the economic point of view the efficiency
the animal is measured
by a comparison of the total feed energy
ciency,
effidone.
This might be called the economic
with the work
of a steam
plant as
comparable to the over-all efficiency
computed by a comparison of coal consumption with the brake
obtained.
horse
Any such comparisons, of course,
power
Since

must

animal
basis
Few

these

take

losses and

account

of

the

maintenance

requirement of

the

the
therefore be made
on
doing no work and must
of the 24-hour output of work
(650).
ciency
effiof the economic
satisfactorydirect determinations
of the relation
animals in the foregoing sense, i.e.,
of work
of the work done to the feed (ortotal feed energy)required
when

for its performance,have

yet been

reported.

WORK

The

extensive

heim

by

have

been

the
on
investigations

Kellner

continued

and

referred

Chapter
requirement,were
requirementsfor work.

horse initiated at Hohen-

work

under

to in

Wolff's

and which
direction,
(386 a) in their bearing upon
intended
primarilyto determine
Unfortunately,however, as there

VIII

the maintenance

the energy
noted, the measurements

563

PRODUCTION

of the work

done

in the earlier

experiments

In the comparatively
subsequentlydiscovered to be inaccurate.
few later experiments of 1891-94, various
mixed
rations were
fed.
the total energy
consumed
could
While, therefore,
per unit of work
be computed, only few and uncertain
data
for individual feeding
stuffs can
be deduced
and the results are therefore of small general
value for the particular
phase of the subjectunder discussion here.
were

671.

Net

values

energy

for

work

production.

"

of the energy
requirement of the work animal
be approached in a somewhat
different way.
The

expended

energy

(630,631),is derived
The
is to

tion
ques-

may,

ever,
how-

in work

production,as alreadystated
stances.
primarilyfrom the katabolism of body sub-

function

replacein

The

of the feed

so

far

as

energy

is concerned

the

body the energy thus expended. The net

value
of a feeding stuff for work
production,then, is
energy
measured
of body energy
which it can
thus reby the amount
place.
is precisely
The case
as
parallelto that of maintenance
discussed in Chapter VIII
(370). The net energy value of a
feedingstuff for the latter purpose is measured by the extent to
which it prevents loss of body energy
of internal
as
a
consequence
work, while the
is measured
a

loss of

working

by
body

the

net

for the former

purpose

to which

it prevents or makes
good
to external work.
Conversely,the

extent

due

energy

value

energy

animal

requiresin addition to maintenance

a
supply
of net energy in its ration equal to the amount
of body energy
katabolized for work production.
In view

of this close

in maintenance
warranted
two

and

in work

that the net

are
purposes
with a steer

between
similarity

the functions

of feed

production,the assumption seems

energy

values of

feedingstuffs for

these

the same.
in an
Thus
substantially
iment
experalreadydescribed (364) it was found that one
,

pound

of

timothy hay contributed

of the animal.

If the

Cals. of external

work

net

same

and

502 Cals. to the maintenance

animal had been requiredto do 167

had

performed it with the same

age
averviz. about one- third,
he would have
as the horse,
efficiency
,

564

NUTRITION

katabolized

hay would

it would
been

have

Cals.
containing 167 4- "
502
that one
be anticipated
pound of timothy
sufficient to replacethis energy in the body.
=

the performance of
Similarly,

would

horse

to

the feed necessary

supply about
In

brief,the

in the

manner

Cals. of external

1000

by a
body

3000 Cals. of
would
have
to support this work
available

Cals. of net

3000

work

of about

the mobilization

cause

and

energy,

ANIMALS

substance

body

and

of energy

FARM

OF

energy.

values for maintenance, determined

energy
and tabulated in the
described in Chapter XVII
net

Appendix, may be regardedas also net energy values for work

production,and the energy requirements of the work animal
values.
of these net energy
be expressedin terms
may
It is plain,in the lightof
672. Net
requirements.
energy
of net energy required
that the amount
the foregoing
discussion,
by an animal for work production may be regarded as equal to
in the performance of the work.
the body energy metabolized
in " 2, it is possible
to estimate
the data contained
From
proximat
ap"

how

much

be

must

perform a

known

in

energy

mobilized

of its maintenance

excess

in the

horse, e.g.,

of

work

of mechanical

amount

of

body

quirement
re-

to

kind.
specific

having a draft of 100 pounds

miles on a level road would do 10,560 foot tons of mechanical
20
Table
Calories.
(651) shows
work, equivalent to 3421
155

Thus

the

net

horse

in

hauling a

load

the horse

of
efficiency

Accordingly,the animal would

the performance of this work
and

his feed

in addition

must

the

to

in draft to be about
have
4321

to mobilize
-5-

0.313

31.3 per cent.

in his body for

10,929 Calories,
of net energy

supply this amount

requirementsfor locomotion,maintenance
therefore

other purposes.
of
The total expenditure

or

body energy during the performance

horse,as appears from " 2, includes substantially

four factors in varying proportions,
viz.,the expenditurefor
(ordescent)
maintenance, for horizontal locomotion,for ascent
the
of
estimate
for draft.
A fairlyaccurate
net
and
energy
requiredto do a certain pieceof work may therefore be obtained
by computing the requirement for each of these factors separately
and
recorded
for
from
the data
net
efficiency
already
of work

by

the

adding the results.

For example, let
pounds

hauls

load

it be
of

supposed that

2000

horse

pounds, having

weighing 1100
horizontal

draft

WORK

565

PRODUCTION

pounds, 15 miles per day, including5 miles up a 1 per

The mechanical
cent
grade, at a speed of 3J miles per hour.
the weight of the animal plus
work
performed consists of lifting
the load 264 feet and in overcoming a draft resistance of 100
fore
pounds through 79,200 feet. The total mechanical work, thereof

100

is

follows

as

Table
Draft

158.

Ascent

"

Example

"

5280

100

15

The

of

amount

in the

body

the mechanical

to

the

818,400 foot pounds

0.265 Therms

8,738,400 foot pounds

2.830 Therms

of

in

Table

in locomotion

the horse must

which

energy

work

percentages

2.565 Therms

of this

performance

Done

Total

Work

of

foot pounds

7,920,000

264

3100

Computation

of

metabolize

daily task will be that corresponding

draft and of ascent, computed from
pended
ex155, together with the energy
the

accordingto

table and the energy

same

The total requirement of

requirement for maintenance.
will be as follows : 1
per day, therefore,

net

ergy
en-

"

Table

159.

Example

"

Computation

of

For

draft

For

ascent

2.565 -f0.265

For

locomotion

0.262

-r-

Net

of

that

0.343

0.773

Therms

3.930

Therms

15

4.356 Therms

be

of

rations.

are

it is evident

horse

if the net

of feed stuffs
but

as

we

(775-778)

For

data

regarding the
(760,773, 774),but
JThe

computation

energy

the ox,

net

energy

in this

animal's
could

of 5 for all forms

efficiency

be

case

body

on

correspondingration
of the feedingstuffs
the

of the

case

very
in work

as

way

requirement

net

of

may
to be

horse,the

energy

values

been

mined
directlydeterlations
complicated calcusatisfactory
contrary,fairly

values of feed stuffs

somewhat

of work.

such

some

States,such

the

Therms

energy

possess have not

results of somewhat

of the
efficiency

values

now

the

are

net

that the

of the United

animal

work
principal

the

in
Unfortunately,

known.

in

Having

"

just illustrated determined

computed

used

17.254

Calculation

the work

8.195 Therms

Total

673.

Requirement

0.313

(385)

For maintenance

Energy

are

available

few determinations

of the

production have yet been

simplifiedby

assuming

uniform

net

566

NUTRITION

the indications

although
reported,
different from

FARM

OF

ANIMALS

(659)that

are

it is not

widely

of the horse.

that

Using the net energy values for the horse obtained by Zuntz and
Hagemann's method of computation and contained in Table VIII
of the Appendix, rations may
readilybe computed for this animal
in the same
pending
as for any
other,their accuracy degeneralmanner
of the net energy values used.
Thus,
upon the accuracy
in the case
just supposed,the requirement of net energy was
the figuresof the table it is easy to comFrom
pute
17.254 Therms.
the
ration
would
the
that
meet
following
requirement.
Table

160.

Example

"

Ration

of

for

Work
Net

10

lb. meadow

10

lb. oats

hay

Energy

3.270
8.820

4.1 lb. maize

Therms

5.164 Therms
17.254

The

Therms

in practicelies in
principaldifficulty

of the amount

of

of work

at

work

more

possibleto make
performed which

done.

farm

With

computation

any
would

the determination

animals

it
less irregular
intervals,

or

of

Therms

the

doing

riety
va-

hardly

seems

mechanical

work

be

trustworthyor which would justify

The
of the ration of the farm
the time consumed.
sufficiency
be judged of by the live weight and conhorse will ordinarily
dition
of the animal, and the principal
of tables of net energy
use
values will be as an aid in securingthe necessary
feed energy
at the cheapestrate per unit.
of horses or mules
On the other hand, where a largenumber
used

are

would
work

for the

degreeof

and

in

compute

It appears
lead to considerable

consideringthe

(392),a surplus of
animal

to

Feeding
estimates

be

formulated, as

has

fairlyreliable
not

it

of the

estimates

unlikelythat

reasonable

such

tations
compu-

since,as was pointed

requirements of the horse

economy,

the

medium,
light,
been done by

horse.

and

to

stimulate

this

expenditureof

an

for

conditions

requirements with

unnecessary
activities.

standards
for

uniform

especially
apt

seems

and

muscular

under

maintenance

feed

restlessness

in minor
674.

feed

accuracy.

might
out

to

of work

make

possible to

seem

done

kind

same

"

More

heavy

various

work

writers.

or

ergy
en-

less
may

bitrary
ar-

also

568

efficiency

gross
from

case

work

and

per

OF

NUTRITION

to

the

of

day,

i.e.,

will

it

hours

stopped

for

computed

work,
of

up

on

modern

with

keep

and

Therms

45.230

motor

work

was

basis,

being

are

done

would

to

for

be

24

hours

6.26

if

portion

be,

as

or

as

of

to

the

cent,

an

internal

although
the

about

hours

or

the

of

about

ditions
con-

to

necessary

were

if

and

ment,
require-

practice,
it

idle

machinery

reduced

per

the

is

animal

might

during

continuously

run

only

be

actual

In

for

the

bodily

of

tenance
main-

to

maintenance

be

much

work

engine

an

efficiency

very

steam

his

would

locomotive.

of

were

as

animal

supposed,

24-hour

over-all

this

animal
head

imagine

vary

intensity
the

useful

case

necessary

the

steam

an

full

energy

we

his

of

the

hours,

18

only
feed

total

the

If

day.

the

of

In

will

percentage

hours

ratio

the

as

remaining

with

him

charge

that

the

of

number

varies.

per

this
with

increase

the

vary

requirement
worked

will

with

ANIMALS

however,

body,
It

case.

decrease

FARM

time.

bustion
com-

actual

"

PART
IV

THE
FEED
SUPPLY

XV

CHAPTER

Part

of

Sources

676.

the

III

animals

and

STUFFS

FEEDING

THE

feeding stuffs.

the several

In

"

requirements of different classes of farm

for different forms of production have been considered.
feed

primitive forms of animal

times
the pastoralhusbandry of ancient
the

In

chaptersof

husbandry, such

more

feeding

States, these requirementswere

United

of the western

the range

or

as

met

by

ing
products of the soil. Increaspopulation and rising land values, however, inevitably
tensive
intend to the displacement of pastoralagricultureby more
forms in which
a much
greater variety of feedingstuffs
is available for domestic
animals.
Forage crops are grown

the

for

consumption

use

of the

in the winter

natural

and

supplement the

to

pasture ; grainis produced in

human

consumption and

of the effective demand

excess

utilized

deficiencies of the

as

stock feed ;

for

as
finally,

this

surplus of grain decreases with the growing requirement for

human
food, a great variety of residues from the preparation
of the crude products of the farm for man's
use, the by-product
available to the stockman.
feeds,becomes
It does

consider

either the

the

of

the present work to

with the
connected

of agronomy
the technical
stuffs or
which

brief consideration

principalclasses
to

province of

problems

production of feeding
manufacturing processes
feeds,but

the

fall within

not

yield the
of the

feeding stuffs

discussion

of the

details of the

various

by-product
of the
generalproperties
desirable

seems

as

an

duction
intro-

principlesdetermining their

nutritive values
677.

Classification.

"

The

three main

classes of

feedingstuffs,
the coarse
fodders,

already stated in Chapter II (111),are

or
roughages,consistingof the vegetativeorgans of plants,the
and tubers,and the concentrates, the latter comprising
roots
both the grainsand similar farm products and the by-products
as

of

divers

industries.

The

members
57i

of

these

three

classes of

OF

NUTRITION

572

FARM

ANIMALS

feeding stuffs may be variouslygrouped for different purposes,

but
the following scheme, although not
strictlyconsistent,
the purpose

serve

may

of this discussion.

of feedingstuffs
Classification
Roughages,

or

fodders.

coarse

Dried
Grasses

Legumes
Straws

Fresh
Grasses

Legumes

Silage
and

Roots

tubers

Concentrates
Farm

products
Cereal grains
Leguminous grains
Oil seeds

Dairy products
By-products
By-products of milling
industries
By-products of fermentation
By-products of oil extraction
By-products of starch and glucosemanufacture
By-products of sugar manufacture
By-products of the packing house
classes of
of these various
following characterization
feedingstuffs is reproduced without material change from an
The

earlier article

"

by

the writer.1

i.

Roughages,

or

Coarse

Fodders

terized
characroughages are
large percentage of crude
chemicallyby a relatively
of the plant. They usually
fiber,which forms the framework
this ingredo not contain very much
dient
protein,although in some
shows
a
fairlyhigh percentage. The proportionof crude
678.

General

characters.

Bailey'sCyclopedia of

American

"

The

Agriculture,1908, Vol. Ill, pp. 58-92.

THE

fat is small and

free extract,

FEEDING

STUFFS

includes much

along with

more

besides

573

true

fat.

less starch and

or

The

sugar, includes

great variety of less familiar carbohydratesand

whose

nutritive

value

nitrogen-

of other

stances
sub-

is

problematical. By far the

is supplied
largerproportion of the roughages in common
use
by two classes of plants, the grasses (Gramineas),including
maize, and the legumes (Leguminosae) Furthermore, crops
"

belonging

to

both

these classes

may

be

used

for fodder

when

but

lowed
partiallymature
(hay, maize forage),or they may be alto ripen, the grain may
be removed, and the residue
(straw,stover)used for feeding.

The

largershare of the hay crop and

of the pasturage of the United States is suppliedby plantsknown
in a restricted and popular sense
as
grasses, such as timothy,
red- top. To these must
be added, as a most
blue-grass,
tant
impor679.

The

grasses.

of

source

"

forage

in

the

United

States,maize,

Indian

or

which

botanicallyis a grass, although not commonly so

called. The foragesuppliedby these plants has a very wide
of nutritive value, depending on a varietyof conditions.
range
Chief among
these is the stage of maturity at which
the crop
corn,

is utilized.
thin

In

growing
young,
consist of nearly pure

and

filled with

vegetation the cell walls are

while the cells are
cellulose,
protoplasm whose chief ingredientsare proteins.

active

Hence, forage cut

percentage of crude

at

this stage shows

fiber and

low
relatively

high percentage of proteins.

rich in protein and
Young and tender pasture grass, relatively
low in crude fiber,
in value,
even
approach the concentrates
may
illustrated by the followingcomparison of the dry matter
as
of
a

sample

of young

Table

162.

pasture grass with that of average

"

Comparison

of

Pasture

Grass

and

oats

Oats

"

NUTRITION

574
the

As

plant
and

more

the

At

ANIMALS

cell walls

the

matures,

thicker

grow

impregnated with tough, woody

more

time,

same

FARM

OF

soluble

more

carbohydrates, as

being produced, while

and

come
be-

material.

starch

and

the

to occupy
protoplasm comes
but a small part of the cell. The
fullymature
forage,
in
i
s
crude
rich
fiber
of
resistant
contains
a tough,
therefore,
sort,
much
carbohydratematerial in generaland tends to be poor in
proteins. For example, three samples of meadow-grass, cut
the followingcomposition,reduced
at different dates, had
to a
are

sugar,

uniform

percentage of

Table

163.

water

Composition

"

of

"

Hay

Cut

at

Different

Dates

June 26
(over-ripe)
Water

15.0

Ash

6.2

Crude

protein

Crude

fiber

7.2

32.4

...

Nitrogen-free extract
Ether

extract

36.9
2-3

IOO.

Accompanying
In
digestibility.
resistant

this
the

the action

change in compositiongoes
first place, the crude
fiber
of

the

decrease

becomes

in

more

digestiveorgans.

Furthermore,
the less soluble crude fiber seems
to have
a
tendency to protect
the contents
of the cells from
At
digestion.
rate,
any
of the protein,and, to a less dethe percentage digestibility
gree,
that of the other ingredientsalso suffers. The percentage
to

of the several ingredients

of the above
digestibility
found
to be as follows
grass, omittingthe ash, was
Table

164.

Percentage

"

Digestibility

of

Hay

Cut

samples of
:

at

"

Different

Dates

26

June
%
Crude

protein

Crude

fiber

55-5
61.

...

Nitrogen-freeextract

55-7

Ether

43-3

extract

576

NUTRITION

the other

On

hand, of

FARM

the

course,

diminishes
(stover)
plant grows older.

leaves

alone

the

as

OF

681.

of

Proportions

ANIMALS

of the
digestibility
as

vegetative

in the

of other grasses

case

organs.

stalks and

The

"

composition

and

of the grasses is also materially

affected by the
digestibility
The
influence of
proportionsof the various vegetativeorgans.
the largeproportionof seed in the maize planthas alreadybeen
mentioned.
In general,
the leaves of the grasses, and of other
tender
and
contain
less crude
forage plantsas well,are more
fiber and
more
proteinsthan the stems.
Leafy species and
varieties therefore tend
to have
a
higher feedingvalue than
those which
consist more
largelyof stems, and any influences,
such as thickness of planting,
manuring, season, and the like,

affectingthe

relative

proportion of leaves,tend

the value of the crop.

is to make

from

the

The

it, extremely

9.8 per
The

cent.

are

grasses

result of all these

made
grass, or of the hay or silage
variable.
American
analyses of timothy

follows

as

166.

"

total

"

Protein

and

Fiber

in

Grasses

Various

Total
Protein

Per

Kentucky
Meadow

....

blue-grass
fescue

Orchard-grass
Maize forage x

Crude
Fiber

Cent

Per

these variations

in

differences

in

Cent

5.9-10.4

24.0-31.8

5-3-12-9

17.7-26.8

4.5-11.8

20.8-31.9
28.9-38.3

6.6-10.4

Oats

That

factors

proteinranging from 3.8 per cent

cent and fiber varying from
22.2
per cent to 38.5 per
corresponding variations in hay from a few other

Table

Red-top

affect

to

composition of

hay, for example,show

to

combined

also

2.7-

6.9

7-5-24-7

5-2-

9-5

23.1-30.9

responding
compositionare accompanied by corhas
been
pointed
digestibility already
out.
Moreover, the percentage of crude fiber in roughage
index of the relative expendito be a fairly
accurate
appears

Entire

plant,usually containingconsiderablymore

water

than

hay.

STUFFS

FEEDING

THE

577

of energy in digestion
(770). Not only does coarse, woody
matter, but what it does contain
forage contain less digestible
ture

rived
animal, pound for pound, than that defrom forageof a better quality.
the clovers,
682.
The
alfalfa,
legumes
peas, beans,vetches,
of forage second only to
constitute a source
and the like
the grasses in importance, while their value
as
renovating
a
peculiarpositionin agriculture.Broadly
crops gives them
speaking,leguminous forage may be said to differ from that
main
tions
of the grasses in two
points. First,under like condiit is notably richer in proteinsthan the latter. Second,
is less valuable

to

the

"

"

there is

difference between

marked

more

the

physicalproperties

legumes, the rather coarse

to the leaves with
stems
turity.
advancing maincreasingrelatively
mature
legumes is therefore likely
Hay from somewhat
of crude fiber than
to be bulky, to have
a higher percentage
the same
to be less digestible.For
grass hay, and relatively
it is more
reason
subject to mechanical losses in curing,which
of the

stems

and the leaves in the

quality. For all these reasons, the composition

of
and
digestibility leguminous forage show an even
greater range than those of the grasses, and the importanceof
In brief,the influences
marked.
timely cutting is still more
of the grasses
which
affect the compositionand
digestibility
but
the same
affect those of the legumes in substantially
way
its

likewise lower

to

an

greater extent.

even

vegetativeorgans of the plant

after the removal
of the ripe or nearlyripeseeds.
Since the
uble
ripeningof the seed consists largelyin the transfer to it of sol683.

consists of the

Straw

materials
straw
extent

from

the leaves

and

stems, it follows that the

materials in
will be poor in digestible
and the degree to
of seed formation

proportionto
which

the

the seeds

ripen. Furthermore, those parts of the plantmost distant from

found to be most
the seed are
completelyexhausted of food
The straw of the common
small grains is relatively
material.
very

in

poor

proteinsand fat,while still containingnot

amounts

of

digestible
carbohydratesand

siderable
incon-

stances.
related sub-

tough, woody character,however, as indicated by

its high percentage of crude fiber,
large
pointsto a relatively
expenditureof energy in its digestion,and its real nutritive
Its

value

is therefore
2

low.

Wheat

and

rye

straw

stand

at

the

578
foot

NUTRITION

the

of

OF

list,while

Sheep
the upper
parts. The

are

and

FARM

ANIMALS

and

uable.
valare
more
barley straw
ing
especially
adapted to utilize straw, consumvaluable
the coarser
more
parts and rejecting
oat

(stover) constitutes

of maize

straw

valuable

less woody than that of the small

feedingstuff. It is relatively
and is more
high degree of digestibility,
grains,has a relatively
its complete conTo
secure
sumption,
palatablethan ordinary straw.

however, it

is necessary
to cut
the additional
questionedwhether

been
cut

is worth

fodder

legumes is

molds

"
Contain

Roots, Tubers

2.

much

the

water.

coarse

convenience

cucurbita.

other

on

from

8o

other.

fiber in their

low, and

hand

one

and

them

With

They

are

In
the

the

centrated
con-

cluded
be in-

may

crops

(ofwhich

less water,

potatoes

and
their

(beets,

proper

like)the percentage of

tubers

to

root

erties
prop-

the

fruits,notably pumpkins
characterized
by
especially

A second
66 to 82 per c^nt.
of these feedingstuffs is the low

also

the

constitute

certain

The
95.
chief representative)
contain
vary

of

straw

tubers

and

the

on

the

largeproportionof water.
turnips,
carrots, mangels and
may

in the

Fruits

and

Roots

"

fodders

feeding stuffs
for

eaten

feeding stuffs,
differing
markedly in their

distinct class of
from

cutting. The

it has

proteinthan that of the cereals and lower

On
the
correspondinglyhigher digestibility.
is usuallycoarse
and liable to
and unpalatable,
and other fungi.

fiber,with
other hand, it

684.

material

it,and

richer in

in

contain

of

labor

the

shred

or

water

are

the

the

being aprange
proximat
acteristic
charequallymarked
percentage of crude

Their percentage of crude proteinis

dry matter.
ferior
a
largeshare of it consists of non-proteinof in-

nutritive value.
685.

source

of

carbohydrates.

of the
crops consists largely
In the tubers starch is the

beets,especially
sugar
and

this substance

nutritive

has

more

"

The

dry

matter

of these

readilysoluble carbohydrates.

predominant carbohydrate,while in
beets,cane
sugar occupies this position,
been

shown

to have

lower
distinctly

value, for ruminants at least,than starch. In other

root
pectin
crops, the carbohydrates consist largelyof gums,
and
other
the
substances,
pentose carcompounds, including

FEEDING

THE

bohydrates, whose
There

are
or

more

STUFFS

nutritive

exact

579

value

is

still uncertain.

in fruits,
also present in roots, and
particularly
nutritive value is low.
In
less organic acids whose
of their succulent

consequence

and

tender

nature, tubers,and

be
and may
especially
roots, have a high degree of digestibility
presumed to require little energy for their digestion. They
of carbohydratematerial,even
therefore a valuable source
are
inferior
ingredients are of somewhat
of tubers is more
In general,the dry matter
value.
valuable
than that of roots.
On the other hand, the dietetic effects of
of labor
roots are especially
prized,but the considerable amount

though

of

some

requiredfor

their

their cultivation

"
686.

Comparison

3.
with

tends

restrict their

to

use.

Concentrates

The

roughage and

with

roots.

"

The

centrated
con-

"

feedingstuffs,or

plies,
imconcentrates,"as their name
of nutriment
those which contain a large amount
in
are
small weight and bulk.
a
They stand in contrast, on the one
is accompanied
hand, with roughage, in which the real nutriment
tible
by a largeproportion of woody fiber and other indigesmatter

which

adds

the

to

weight and

bulk

without

rially
mate-

increasingthe nutritive value. On the other hand, they

excel the roots and tubers because, while the dry matter
of the
latter is very valuable, it is largelydiluted,so to speak, with
The

water.

concentrates

are

therefore

the main

reliance for

the

The
rapid, intensive production of meat, milk or work.
be subdivided
into farm
concentrates
products and the
may

by-productfeedingstuffs.
Farm
687.

The
recent

and

indeed

cereal

grains.

products
"

times,the main
are

still in many

The

'

grains were,

reliance of
sections

until comparatively

of concentrates,
of the United
States.
users

Corn, oats, barley,rye, peas, beans, rice and at times even

These
wheat, are feedingstuffs whose value needs no advocate.
for the use
seeds contain, stored away
of the young
plantlet,
proteins,fats and carbohydratesof the most valuable character
and

"

the highesttype
representing

of

vegetablefood."

Their

580

OF

NUTRITION

is

FARM

ANIMALS

the form

chieflyin

of true

proteinsof
recognizednutritive value, their carbohydrates are largely
starch,and their ether extract
chieflytrue fat. Being closely
related to the nutrition of the young
plant,the composition of
smaller variations
seed shows much
than
the properlymatured
fodders.
The
that of the coarse
degree of maturity of the
seed,however, materiallyaffects its compositionand in much
nitrogenous matter

the

as

way

same

earlystages

that

of the

fodders.

coarse

'

the

In

of seed

from

ripeningof

it does

dantly
formation, the proteinand ash flow abunthe vegetativeorgans to the seed, while later the

the seed is largely

an

which
therefore,
Any influences,
of the seed, such as drought

produce a seed
Light,shriveled

richer in

accumulation
check
or

of

carbohydrates.

the normal

development

lodging of the grain,tend

proteinand

poorer

to

in

carbohydrates.
high in protein.

grain,therefore,tends to be
of unripe seeds differ to a considerable
Moreover, the ingredients
The
from those of ripe seeds.
extent
ample,
nitrogen,for exis to a largerextent
in the form of non-proteinrather
and the carbohydrates
in the form of sugars
than true protein,
are
rather than starch,as in the ripegrain.
of one
sort or another
688.

and

Composition

of cereals.
digestibility

"

cereal

The

by a medium
percentage of protein
(8 to 14 per cent),chieflycomposed of true protein,a rather
low percentage of fat (1.5to 6 per cent) and a high percentage
of carbohydrates,
largelystarch. Their ash is small in amount
and in it potassium and phosphorus acid are prominent,while
Maize
contains
rather less protein
but littlecalcium is found.
than the other cereal grains,
with correspondingly
high percentages
grains are

characterized

of starch

and

of fat.

While

it has

been

shown

that the

notablyincreased by selection and

affected
effects of the latter have not yet sensibly
the character of the commercial
The naked grains(maize,
crop.
a comparatively
high percentage digestibility,
rye, wheat) show
and both in this respect and as regards their compositionexhibit
less variation
than the hulled grains (oats,barley). In
valueless
the latter,the variable proportion of the relatively
hulls to the kernel causes
both compositionand digestibility
to
of
the extremes
shown
vary greatly. Oats, for example, have

proteincontent
breeding,the

6 and

of

corn

can

be

17 per cent proteinand 3 to 7 per cent

in compositionand value.
resemble straw

of fat.

The

hulls

They therefore

in-

THE

proportionof

the

crease

581

FEEDING

STUFFS

crude

fiber in the

grain,and

spondingly
corre-

and nutritive value.

digestibility
ing
The place of the cereal grainsin feedUses
of cereals.
689.
practice is clearlyindicated by the foregoing statements.
They enable the feeder to introduce into his rations,without
of easily
unduly increasingtheir bulk or weight, large amounts
and
highly nutritious ingredients.Of themselves,
digestible
they contain a fair proportionof proteinfor many
purposes,
not
but
of
for
mature
are
capable
animals,
they
especially
offsettinga deficiencyof proteinin the other ingredientsof
the ration,nor do they supply enough of this ingredientto meet
diminish

its
"

of
fullythe demands
productivedairycow.

rapidlygrowing animal

the

or

the

highly

leguminousgrainsshare the
and like
of the naked cereal grains,
generalphysicalproperties
them
contain feed materials (proteins,
carbohydrates,fats)of
the highestgrade. They are especially
trast
characterized,in conwith the cereal grains,by their relatively
high percentage
of protein,ranging according to American
analyses from 20
of them, as the soybean and the lupine,
Some
to 42 per cent.
690.

grains.

Leguminous

"

The

of fat,but
also carry notable amounts
richer in this substance
than
not
are

than

richer in ash
acid and

lime.

concentrated

rations in

they are

the

more

common

ones

cereals.

They are
regardsphosphoric

cereals,notably as
Their digestibility
is generallyhigh. Like the
the

valuable

cereals,they are
a

the

as

sources

of total

feed
digestible

form, but unlike these they serve

protein. Aside

the most

from

certain

available materials

technical

in

also to enrich

by-products,

for this purpose,

and the
both for this purpose
and

leguminous feeding crops,

for their effects on the soil,deserves careful consideration.
oil seeds, such as flax,cotton
and
The
691. Oil seeds.
culture of

"

as
directly
feedingstuffs because
value.
seeds contain a high perof their commercial
These
centage
in
of
of protein,while
the carbohydrates
placeof much
of the cereals and legumes a largepercentage of oil is found.
Flaxseed
contains
considerable quantity of so-called
a
cilage,"
muwhich swells up with water
and has a
to a slimy mass
is
the digestive organs.
Cottonseed
very soothingeffect on
fed to cattle to some
extent, usuallyeither boiled or roasted,but

rape,

are

commonly

not

used

"

is

regarded

as

dangerous for growing swine.

582

ANIMALS

FARM

OF

NUTRITION

By-products
Nature.

692.

by-productfeedingstuffs

The

"

by which the
processes
preparedfor man's use, either as food
of technical

are

the residues

products

of the soil

for other

or

purposes.

ing
technical processes are : the millmore
important
of cereal foods ; the manufacture
of grains; the manufacture
of starch and gluof alcoholic liquors
cose
; the manufacture
of these

The

of

By-products

693.

wheat,
stuffs.

grain for millingand

The
grindingproper.
mixture

according to

Milling residues,particularly

milling.
"

of the

familiar

ing
by-productfeedsecured in cleaningthe
screenings
the bran and
middlingssecured in the
screeningsare an exceedinglyvariable
sides
bethe qualityof the grain, containing,
grains,a great varietyof weed
seeds,

the most
among
They include the

are

lightand broken
fragmentsof straw, sand and earth,as well as
While some
fungi,and dirt of all sorts.
feedingvalue, the possibledanger to
animals, and

through

the

the infestation

of

of oils.

the extraction

; and

of sugar

manufacture

; the

of

are

the health

fields with

the

great caution

demand

manure,

of

ous
spores of numerof these have undoubted

in

as
screenings

to

is

be

to

weed

the

seed
use

feed.

bran

of the

of

dition
Its ad-

middlings
regarded as an
or

adulteration.
Bran

"

The

bran of wheat

of
essentially
the seed-coats of the grain,
the layerof so-called gluten
cells immediately beneath
them, and a proportionof
the inner,flourypart of the
grain varying with the perfection
of the milling. The
seed-coats of the graincontain
or

Fig. 43.

"

Partial

section

(Bailey'sCyclopedia
i,
coat.

of wheat

American

grain.

ture.)
Agricul-

Seed pod. 2, Outer seed coat.

3, Inner seed
4, Gluten cells. 5, Starch cells. (Jordan.)

richer

the

of

in

proteinsthan

the

the
as
therefore,
flour,does it become

inner

bran
at

once

is

rye

consists

most

while

the

part of the
more

of its crude

fiber,

gluten

cells

kernel.

In

are

portion,
profrom
perfectly
separated

richer in

proteinand

in crude

584

OF

NUTRITION

which

becomes

rancid

rather

FARM

ANIMALS

easilyand

often

renders

the

terial
ma-

be
unpalatable. It is asserted that this ranciditycan
the
bran or polishas soon
as produced.
preventedby kiln-drying
Uses

of

regard

to

milling by-products.
the

than

has

millingby-products largelyas

it is true

While

There

"

whole

sufficient to

that

the bran

wheat

or

enable

tendency

of

sources

protein.

and

other

them

been

middlingsare richer in tein

procereal grains,the difference is
offset to

marked
degree
any
the deficiencies of other ingredientsof the ration in this respect.
of digestible
They are to be regarded primarilyas sources
not

to

the
whole, with a tendency to increase somewhat
with the good
proportion of protein in the ration.
Familiarity
its comparative safetyas
qualitiesof wheat bran in particular,
its good dietetic effect have
a feed in inexperiencedhands, and
tended
to an
it rules
When
exaggerated idea of its feed value.
high in price it is usuallypossibleto substitute other feeding
stuffs for it,partially
or
wholly,which will furnish both protein
matter

as

and

energy

cheaply.

more

Buckwheat

contrary, often

cheap
a

the

middlings,on
of

source

ration

furnish

protein for

otherwise

deficient

in it.
694.

Breakfast

ture
manufac-

the

In

"

dues.
food resi-

of the great varietyof

so-called cereals,
fast
breakor

foods,now
a

of

the

on

considerable

quantity

by-products
the

In
most

oatmeal,

lates.
accumu-

of the

case

of

common

the

ket,
mar-

these,

residue

of
chiefly

sists
con-

the hulls of

the oats

togetherwith some
the lighter
grains.

of

Oat
Fig. 44-

"

Partial

section

(Bailey's Cyclopedia of

of

American

oat

grain,
Agricul-

ture"-)
o, Hull,

cells.

Seed

(Jordan.)

coat.

4,

Gluten

cells. 5. Starch

"

hulls

The

have
themselves^

scarcely
than

feeding value

more

the
i,

hulls.

straw,
11

resemble

"

in

which

they
"

,"

composition,

proportion of lightoats

the

while

raise the value.

to

in the

but

market

hulls

Oat

585

STUFFS

FEEDING

THE

is

sufficient materially

not

rarelyoffered

are

usually disposed of

are

in

such

as

of

one

two

First,they are made the basis of various proprietary

feeds,cheap by-productsof various sorts being added, usually
of the protein-rich
by-products shortly
includinga small amount
ways.

and

by
are

advertisingtestimonials.
worthless,it is evident that the oat

means

no

offered under

are

various

While

abundant

with
no

feeds

These

be described.

to

of the addition

because

valuable

more

names

they

are

hulls themselves
to

them

of other

ing,
ultimatelypays the cost of mixand advertising. The second
to which
use
transportation
oat-hulls are
pecially
put is the adulteration of the mixed feeds, esand oat feeds, which
are
freely offered on the
corn
considerable
market.
Since it is difficult to recognize even
a

materials,while the

of this sort, such

adulteration

only

from

consumer

of known

manufacturers

guarantee

as

mixed

feeds should
or
integrity

be

purchased

under

factory
satis-

purity.

to

of pearled
by-product of the manufacture
barley,is similar in feedingvalue to oat hulls.
of hominy from
Hominy feed. In the manufacture
corn,
the hull,the germ and the more
starchyparts of the kernel are
rejectedand constitute hominy feed, or hominy chop, which

Barley feed, a

"

is similar

the whole

to

except that
has

compositionand digestibility,

percentage of fat is greater. Consequently it

its

higher feedingvalue,although the fat is likely

rancid on long keeping and thus lower its quality.

somewhat

become

to

kernel in

695.

of

By-products
of

the

fermentation

industries.

"

The

in the
liquorsconsists essentially
of the starch of grainsor potatoes into sugar and
conversion
of this sugar by means
of yeast.
the subsequent fermentation
be consumed
The resultingliquor may
directly(beer,ale) or

manufacture

distilled,
yieldingthe

be

it may

alcoholic

The

"

are

of

of

the

one-third

about
from

roots

malt, by

growth

the

first step in the process is the preparation

moistened
to
allowing
barley
germinate.

The

sprouts.
of

distilled

alcohol.

liquorsor commercial
Malt

concentrated

more

sprouts is stopped by drying when

inch

long, and

these

grain, constitute malt

barley,they

have

the

they

dried sprouts, separated

sprouts.

Being young

generalpropertiesof

all young

586

NUTRITION

FARM

OF

ANIMALS

plantgrowth,containinga high percentage

of

much
nitrogen,

of

non-protein,and a low percentage of crude fiber.

Brewers7
ing
grains. The next step in the process is the mashof the ground malt and other grain with warm
In
water.
this process,
the diastase of the sprouted barley acts on
the
starch of the grain,transforming it into sugar.
In the manufacture
of beer or ale,the resultingliquidis drawn
mented
off and ferbrewers'
as
leaving a residue known
separately,
grains,
is used extensivelyas a dairy feed.
which
In the fresh state it
is valuable,but is subjectto the disadvantageof fermentingor
souring very readily,and tending in this state to injure the
it in the form

of

"

Somewhat
recently,economical
quality of the milk.
cesses
profor dryingit have been perfected,
and the dried brewers'
grainsconstitute a valuable feed which can be shipped like
other

any

dried feed.

Distillers7 grains.

alcohol,the liquidis
alcohol

fermented

with

in contact

the

grainsand

tillers'
disas
off,leavinga residue known
wetter
slop. This residue is much
grainsor distillery
less
than brewers'
but
is
grains,
subjectto fermentation,since
the sugar has been more
Large quantities
completelyremoved.
of it are now
put on the market in the dried form, both under
the

its
no

then

preparation of distilled liquoror

the

In

"

distilled

of which
contain
trade names,
some
It constitutes
suggestionof the real nature of the material.
and

name

own

valuable

of stock

source

regarded
quality.

are

as

various

feed.

the poorest and

The

grainsproduced

those from

maize

as

from

rye

of the best

the starch of
objectis to convert
the grain as completely as possibleinto sugar and then into
This results in increasing
the percentage of all the other
alcohol.
ingredientsin the residues.
They contain accordinglya high
In

all these processes

the

percentage of protein with also

of crude

fiber than

somewhat

greater percentage

the

ordinarygrains. They serve, therefore,

whole but also to
not
matter
as
a
only to supply digestible
of protein in the ration.
correct
a deficiency
696.

By-products
oils from

called
value.

oil cake

Of

or

of oil extraction.

various
oil

in

meal,
the

The

extraction

seeds
oil-bearing

these,cottonseed

extensivelyused

"

some

and

United

of which

leaves
have

linseed

meal

States

and

are

are

of

mercial
com-

by-products,
high feeding
the

only ones

typicalof

the

FEEDING

THE

seeds of cotton

The

others.

and flaxseed about

is extracted

pressure,
meal.
The

At

"

fat,the

by the

or

oil

The

of

use

fat and

8 to 9 per

of fat.

cent

therefore

very

being ground

cottonseed,which

have

of the

true

Linseed meal.

pressure and
removed

at

by

low

fiber.

Cottonseed

tough,

black

meal

hulls

feeding value.

of

This

is

the

is

pecially
es-

cottonseed
grades of commercial
lower pricethan the standard grade.

the flaxseed both

oilis extracted from

naphtha, the

of

means

latter

resultingoil-meal and

being

pletely
com-

recovered

from

the

"

"
of extraction removes
the fat
process
"
"
old process
of pressure, and the
the
in fat and contains
is somewhat
poorer

new
again. The
completelythan

resultinglinseed meal

proteinthan

more

the

very

Linseed

"

the

inferior

sold

are

from

is made

44

little crude

contain

meal

only by

cottonseed

to

per cent of crude proteinand

It should be nearlyfree from the hulls
to

40

extensivelywith

adulterated

somewhat

cake

highest grade of cottonseed

contains

meal, which

oil is extracted

cotton

present

hulled seed and

more

35 per cent
variable.
somewhat

proteinand

by pressure
residue still containing some

resultinghard

the

use

of each

protein.

Cottonseed meal.

by

cent

30 per

the seeds either

from

solvents,leaving a
rich in

fat and

are

contains about

per cent

22

rich in both

flax

percentage, however, being

latter

and

and

cottonseed

Hulled

protein.

587

STUFFS

of extraction

by

years, however,
meal
new-process

is

old-processmeal.

the

The

for

cess
pro-

in recent
pressure has been so far perfected
that the difference between
and
the old-process

less than formerly. The protein

distinctly
meal appears to be slightly
less digestible
of the new-process
than that of the old-process
meal, which tends still further to
oil meals.

Other
from

the

seeds

palm

and

the

not

appear
the United
The

697.

the

to

as

have

States

two.

also manufactured

peanut, the

common

commercially
soybean,the oil

resultingoil cakes or meals are

feeding stuffs in European countries but do
The

yet found

as

to

also be

mentioned

classed
of

glucoseare

access

considerable

any

meal

By-products
and

are

palm.

cocoa

corn-germ

feeds may

Oils

"

of

extensivelyused

Starch

the

the difference between

reduce

as

starch

made

an

to

of

the feed market

extent.

in connection

with the

gluten

oil-meal.
and

glucose manufacture.

in the United

States

"

chieflyfrom

588

NUTRITION

The

maize.

starch

is

being

with acid.

of the starch

ANIMALS

grinding and

coarse

carried

off in

the

suspension and

is manufactured

Glucose

to settle out.

FARM

separatedby

starch

of water, the

use

OF

lowed
al-

further treatment

by

In the

the parts of the kernel which

preparationof the starch,

rejectedare the hull,the germ
and the more
glutinouspart
of the interior of the grain

are

from

which

the starch

cannot

be

completelyseparated.
Corn
The
{maize)bran.
hulls are comparativelylow
"

in

protein and

contain

fiber.

siderable
con-

sold

When

called
separatelythey are
com
position
bran,althoughthe comof commercial

indicates

Fig. 45.

(Bailey's Cyclopedia of American

4,

Outer

layer of skin.

Gluten cell. 5. Starch

germ

"

contains

ture.)
Agricul-

of
1,

ture
admix-

some

of the germs.
Germ
meal.
The

kernel.

Partial section of maize

"

ples
sam-

about

30 per
has
a

oil, which

Inner layer of skin.

cells. (Jordan.)
2,

value

cent
mercial
com-

is secured

and

by pressingthe
residue
per

constitutes

cent

of

oil,and

which

meal,

germ
in the

The
germs.
about
still contains
7

neighborhoodof

1 1

of crude

cent

per

protein.

glutinousresidue of the kernel

constitutes
gluten meal, containing,in general,30 to 40 per
of crude protein with a comparativelylow percentage of
cent
fat and fiber. Some factories mix the glutenmeal and the hulls,
Gluten

and

and

meal

feed.

The

under

the

"

sell the mixture

approximately24
crude

fiber and

germs

are

sold

cent

per

or
are

dry.
made

gluten feed,which

of

of crude

of fat.

cent

protein,6

Sometimes

the

"

and

under

diverse

of

hulls and
"

"

or

names

sold

tains
con-

cent

per

the

sugar feed
In fact,various mixtures

togetherunder

feed,"either wet
main
products

starch

of the three

commercial

invariably
glucose products should
purchased on a guarantee as regardscompositionand purity.
698.
to
Sugar has come
By-products of sugar manufacture.

names.

be

6 per

name

These

various

"

be manufactured

from

sugar-beetsto

considerable

extent

in

THE

United

the
from

FEEDING

States, while
is

cane

sugar

589

STUFFS

in certain

regionsthe

important industry.

an

Sugar-beetpulp. The sugar is extracted

in what is known
of water
cut beets by means

from

"

process.
known

as

residue

beet

pulp, which
to

90

about

to

85

to

87

is

commonly

those of roots

be

may
Its
pressing.

by

cent

and

the

the fresh

In

of water, which

cent

per

finely

beets minus

essentially
sugar

95 per

to

similar

are
properties

is

the

the diffusion

of the other soluble substances.

some

it contains

state

as

this constitutes what

from

The

and

sugar

manufacture

duced
re-

general

it occupiesmuch

the

Its digestible
matter
consists chiefly
place in the ration.
of pectinsand gums,
of carbohydrates belongingto the group
inferior to the sugar of the beets but, according to
somewhat
recent
investigation,
fullyas valuable as the digestiblematter
same

of

The

mangels.

but

beet

wet

be

may

portation,
pulp is too heavy to bear long transpreservedin the neighborhood of the

however, dried
dried beet pulp, containing not more
is now,

factoryby ensiling. It
market

as

of

on

the

5 to

10

put

than

about equally
pulp is relatively
valuable with the wet pulp,especially
if soaked in water, as it
should be before feeding.
cent

per

Molasses.
beets

sugar

molasses.

water.

or

of true

with

sugar

This

dried

the further manufacture

In

"

50 per
cent

The

and

of sugar either from

there remains, as a final residue,the

cane,

contains

to

20

of sugar,

cent

protein,and

other substances
of

25 per

cent

scarcelymore

8 to

per

10

of doubtful

of

cent

of water, approximately
than one-half per

non-protein,
along

nutritive value.

It is

tially
essen-

easily soluble

carbohydrates, principally
Beet
laxative
molasses, in particular,has a marked
sugar.
action,commonly ascribed to the potassium salts present in it
but perhaps due quite as much
For this reason,
to the sugar.
is required to accustom
animals to it graduallyand not to
care
overfeed with it. Its laxative qualities
said to be valuable
are
a

source

used in small

when

for horses in

amounts

preventingattacks

of

colic.
Molasses
is an
the

"

inconvenient
so-called

These
A

feeds.

Owing
material

molasses

consist of molasses

large number

to
to

feeds

its

molasses
physicalproperties,
handle.
To avoid this difficulty,
have

dried

of concentrated

down

been
on

put
some

on

the

market.

suitable material.

feedingstuffs

have

been

used

this

for

and

purpose,

proportion

of

yields

value

especial

%,

80

small

of

amount

of

rendering

blood
of

fat,

rich

in

is

tankage
be

is

while

bought

other

the

ash

in

on

vary
a

which

of

are

poultry.
of

which

digestible.

the

it

contains

contains

It

and

it

method

bones

of

than

protein

its

composition

centage
per-

it

renders

calcium
of

the

considerable

contains

especially

after

scrap

less

contains

in

left

residue

much

which

guarantee.

house

packing

ash.

hand,

widely

value

slaughtering

modern

contains

the

of

the

made.

are

The

trimmings

from

with

are

"

protein,
is

ingredients,

to

they

and

of

bone

obvious

feeds

by-products

swine

little

scraps,

the

these

the

which

Tankage
on

likely

of
but

meat

but,

As

always

the

in

essentially

house.

packing

dried

all
and

consists

Tankage

rich

together

which

nitrogenous
of

dried

house.

scale

feeding

fat

of

packing

large

practically

All

out

especially

is

been

pulp.

highly

the

in

blood

Dried

the

of

ANIMALS

also

the

on

number

FARM

materials
of

animals

meat

over

the

to

By-products

699.

has

it
molasses

forming

pulp,

beet
in

OF

NUTRITION

590

and

tively
rela-

phorus.
phos-

manufacture,
and

should

ANIMALS

FARM

system had the advantage of simplicity.Experiencehad

The

fairlydefinite idea of the quantityof hay required

of production. It was
only necessary to
given amount

afforded
for

OF

NUTRITION

592

hay values what weights of the available

of
feeding stuffs would produce equal effects. The simplicity
due
the calculations,
especiallyto the fact that the relative
value of a feed was
expressed by a singlefixed number, led to
from

compute

the

"

rapidadoption of the system.

assigned and
hay value was
table

beautiful

each

To
in

feedingstuff a

short

time

nite
defi-

had

one

the most
mation
inforexact
gave
of the most
diverse feedingmaterials

which

constructed

regardingthe value
in comparison with hay. Anything which
appeared in any way
its place in the table and
each new
suited for feedingfound
feeding stuff which the progress of agronomy
provided,directly
likewise quickly incorporated. It went
or
was
so
indirectly,
far that even
the salt suppliedto the animals was
computed in
*
hay values."
in part on the results of pracThaer
himself based his figures
tical
carried
subsequentinvestigators
experiments. Numerous
direct comparisons of feedingstuffs on an extensive scale
out
formulated.
and not one
but several tables of hay values were
Unfortunately,these tables differed widely from each other,
of them givingtwo or three times as great a hay value as
some
another

to the

grain or

and

roots

for

of the feed

content

into account
out

hay, was

limitations led to various

estimatingthe hay

that

the

limited
un-

feeds,as for instance

impossible. Such discrepancies
of the

modifications

methods

of

Boussingaultregarded the protein

factor,while Nathusius took
principal

values.
as

the

also the content

somewhat

also

evident

was

of different classes of

substitution
of

It

feed.

same

elaborate

fiber and

of crude
method

in

an

Wolff

attempt

to

worked

retain the

The
for a feed.
reckoning with a singlenumber
of this,however, graduallycame
to be recognized,
impossibility
and the hay values have now
only a historical interest.
convenience

701.

hay

Practical

values

relative

of

has

feeding trials.
become

nutritive values

comparisonsof

obsolete
of

the results

Settegast, Die

Die

"

But

while

the idea

of

system

of

determining the

feedingstuffs on the basis of direct

has survived in
obtained in practice

Fiitterungslehre,
1879, p. 4.
1861,
Fiitterungslehre,

landwirtschaftliche

the

pp. 455-456.

FEEDING

OF

VALUES

RELATIVE

STUFFS

593

in
vigor. A very considerable share of the investigations
stock feedingduringthe last two decades,especially
perhapsin
the United
States,has consisted of experimentsintended to

full

the effects of the substitution of

determine
in

feed for another

one

ration.

trial has an important

Undoubtedly the so-called practical
part to play in the development of a sound theoryof feedingas
well

garded,
aspects of the subject. Re-

in relation to the economic

as

however, simplyas

mination
deterquantitative
feedingstuffs it is subjectto
for the

means

of the relative values of

the
precisely

generalled
is

to

true

as

uncertainties

the old

as

tempt
at-

hay values, and in this respect has in

results.
concordant
more
or
satisfactory
scarcely
in the later as in the earlier experimentsthat
bination
a
feeding stuff may vary widely with the com-

determine

to

It

limitations and

same

the effect of

it is fed and

in which

the conditions

under

which

it is used.
702.

Feed

units.

An

"

fundamental
within

conception of
and
restricted field,

is found

in

thus

relative value

expressingthe
and

interesting
attempt to revive
hay values in a modified form

the so-called feed

his associates in Denmark

of

to

the
and

advantage of
singlenumber,

retain the

feed

by

unit system devised by Fjord

used also in
and extensively

Sweden.1
feed unit system, like that of hay values,is essentially
a
system of empiricalequivalentsaccordingto which feeding
stuffs may
replaceeach other. Instead of hay, the basis of
The

weight of grain (corn,barley,wheat or

rye or a mixture of grains). This is called a feed unit and the
of other feeds requiredto equal the feed unit have
amounts
in very extensive cooperative
been determined
feedingexperiments
with
by the group system (572)with swine and especially
The experimentsthemselves have been executed
dairy cows.
results for
The
with every
precautionto ensure
accuracy.
dairy cows, as revised by Woll for American
feedingstuffs,
and the Danish values for swine and for the horse are givenby
comparison is

and

Henry
1

For

more

unit

Morrison

as

follows

complete discussion of the feed unit system

Expt. Sta.,Circular No. 37.

Feeds and Feeding, 15th Edition,p. 127.
2Q

"

compare

consin
Woll; Wis-

NUTRITION

594

Table

167.

Amount

"

of

OF

FARM

Different
Feed

ANIMALS

Feeds
Unit

Required

to

One

Equal

Feed

Required
Unit
Equal
i

to

Feed

Average

FOR

DAIRY

Range

COWS

Concentrates

Corn, wheat, rye, barley, hominy feed, dried

brewers' grains,wheat
middlings, oat shorts,
in roots
molasses
beet
dry matter
pulp,
peas,

1.0

Cottonseed

0.8

Oil

meal

meal, dried distillers'grains,gluten feed,soybeans

0.9

Wheat

bran, oats, dried beet pulp, barley feed,

malt

sprouts

I.I

Alfalfa meal, alfalfa molasses

and

Hay

feeds

1.2

straw

Alfalfa hay, clover hay

2.0

i-5-

3-0

Mixed

hay, oat hay, oat and pea hay, barley and

pea hay, red-tophay
Timothy hay, prairiehay, sorghum hay
Corn stover, stalks or fodder,marsh hay, cut straw
Soilingcrops, silageand other succulent feeds
.

Green

alfalfa

Green

corn,

sorghum, clover,peas

and

oats,

nery
can-

refuse
Alfalfa
Corn

silage
silage,
pea

vine silage

brewers' grains
Potatoes, skim milk,buttermilk

Wet

Sugar beets
Carrots

Rutabagas
Field beets,green rape
Sugar beet leaves and tops, whey
Turnips,mangels, fresh beet pulp
The value of pasture is generallyplacedat 8
the average,
units per day, on
varying
kind and

and

The
those

10

with

condition

values for pigs and

for dairy

in Wisconsin

to

cows

Circular,No.

horses

the values
37.

are
as

those
revised

given

in the

by Woll

Danish

for American

valuation

table

feeding stuffs

RELATIVE

Table

VALUES

Amount

167.
"

feed

and

Logical basis

unit values

FEEDING

Different

of

Feed

703.

OF

Unit

of feed

have

Feeds

STUFFS

Required

595
Equal

to

One

{Continued)

unit system.

The

"

Scandinavian

broad

experimental basis. The results

of the experiments have
been
reasonably consistent
in general the feed unit values correspond well with the

relative

net

values

energy

except that
rich feeds.

they

ascribe

discussed
somewhat

in

the

followingchapter
to protein-

higher values

Nevertheless,the logicalbasis of the system has the

defect that is inherent

Chapter V (263),feed

has two

incommensurable.

are

value of

in all such

feed

with its value

as

as

distinct functions
It is as

of

source

As

systems.

shown

was

and

impossibleto

same

in

these functions
combine

the

proteinor other structural material

of energy, and to express the result in

it is to compare
the relative values of food

source

singlenumber, as
and water
A protein-richfeed like cottonto a starvingman.
seed
meal, for example, will necessarily
produce a greater effect
a

when
to

added

an

deficient in

matter

deficient in

abundance

values, under
fed in mixed

protein than when

of that

the
protein precisely

of fact the feed units

"

when

ration

containing

one

material
As

to

added

ingredient;with

reverse

would

be true.

only claimed to be equivalent

ordinaryconditions of feedingthese animals,
are

rations that would

contain

over

certain

596

OF

NUTRITION

of

minimum

pointedout,

x
protein."
digestible
"

different farms

and

crops

priceof purchased

the

and

Henry

similar

region where

small

As

system has been

feed unit

The

ANIMALS

FARM

Morrison

evolved

have

in

paratively
com-

the
on
grown
feeds does not vary
are

widely throughout the district."

The

writer

the

feed

unit

and

units

energy
those who would

to

system

values.

net

agree with
in this country

able

is not

feed

of

Comparison

704.

with

"

introduce

its wide

variety

of the feed
The applicability
feeding stuffs and conditions.
the presence
units, as just pointed out, is conditioned
upon
As thus limited,however,
of sufficient protein in the rations.

of

theypractically
attempt

to

the relative values

measure

as

sources

the use of the net energy values

of energy, and for this purpose
cally
to be considered in the next
chapteris justas simplearithmetiand

equallyaccurate,

First,the

energy
based
on

to their

lead

units

it is far inferior in

of

causes

extent

hand, aim

expectedfrom

of the

of energy

body

maize
per hundred

fed

as

part of
of

which

it

this

can

the

They

show

what

to

which

than

product which may

i.e.,the
feedingstuff
of

"

contribute
of

new

an

one

ration,would
would

to

the maintenance

age
Thus, if avervalue of 85 Therms

tissue.

energy

that

of fat and

amount

85 Therms,

to discover

attempt

no

worse

or

the amount

means

maintenance
an

comparisonsof

tablish
others,but esfeed and product. Energy values,on

buildingup
example,has

meal, for
pounds,

with

weight of

unit

accuracy,
Second, the feed

direct

on

differences.

to show

to the

or

of the animal
value

to the

pedagogic value.

is better

relation between

the other

amount

or

increase in numerical

some

observed

the

feedingstuff

one
no

be

further

different materials

the results with

the

their very
which may
investigation

purely relative values,based

are

not

and
discovery of new
of comparison. The feed unit,on the other
is concerned,
dead end so far as investigation

leadingto nothingbeyond
while

advantages.
values.
empirical

investigationsand
physiological

great improvement

constitutes

immense

rational and

are

to stimulate

stillbetter methods

hand,

values

net

They are
tend
imperfections

while it has two

hundred
conserve

pounds
in the

proteinhaving
otherwise

be

an

burned

of

it,
body

energy
up

to

that, if
support the vital activities. Furthermore, it means
added to the maintenance
ration,the maize will furnish ma1

Woll, loc.

cit. p. 13.

FEEDING

OF

VALUES

RELATIVE

STUFFS

597

produce a quantity of milk or of meat having

Still further,the investigations
value of 85 Therms.

terial sufficient to
an

energy

by which

these facts

also show

established

are

pounds of maize
gross energy of 100
unused in the various excreta, while about

meal, about 50 escape

expended in the various processes
52 are
consumption and assimilation of the feed.
of

of the losses suffered

the nature

product

expected. Such

be

to

value and

expressedin
"

of

are

as

data

connected

as

different
entirely

an

the final amount

these

as

the

words, they

In other

well

with

have

pendent
inde-

an

from

nature

those

the feed units.

Based

Values

Relative

2.

of the

out

Therms

approximately187

show

that

Composition

on

and

Digestibility
705.

Chemical

composition.

"

system of hay values, attempts

Sprengel and
chemical

others

to

by Davy, Einhof,

made

were

indeed

earlier

the

rise of the

the

feeding stuffs

compare

analyses,and

before

Even

the basis of

on

hay

values

were

for
comparisons (700). The methods
of feeding stuffs were
the chemical analysis
graduallyimproved,
although they stillremain quiteimperfect,but along with this
in

based

part

improvement

problem
a

the

came

is at bottom

Physiological functions
the nutrition

on

basal facts
fats and

the

of

and
physiological

nutrients.

not

In

"

particular

investigationsof Bischoff and

of carnivora

served

to

establish those

carbohydrates,
regarding the functions of proteins,

ash in nutrition

by

question.

teachingsof Liebig and

Voit

recognitionof the fact that the

clearer

of relative values

chemical
706.

such

on

later

which. have

and
investigations

have

confirmed

been
been

outlined

and
in

tended
ex-

ter
Chap-

the earliest to suggest the

of these principles
to comparisons of feeding stuffs
application
Haubner

V.

and

to

have been

feedingof farm animals,while to Grouven 2 belongs the

of having first formulated
the requirements of animals

the

credit
and

appears

feeding stuffs in terms of the different classes

the total
based on
His tables,however, were

the values of

of nutrients.
nutrients

found
1
2

by

chemical

analysisand

were

comparatively

i860.
Gesetze der Ernahrung des Fleischfressers,
Vortrage iiber Agriculturchemie,1858.

598
soon

the

ANIMALS

FARM

OF

NUTRITION

data based

accurate
replacedby more
nutrients.
digestible

on

determinations

of

investigations. It is to
Henneberg's and Stohmann's
1
the fundamental
investigationsof Henneberg and Stohmann
at the Weende
Gottingen, that we
Experiment Station,near
indebted for the inauguration of a system of comparing the
are
with little material
has endured
values of feeding stuffs which
the
change up to the present time. These investigatorswere
in studying the nutrition of herbivfirst to apply systematically
for other
the physiological
ora
already demonstrated
principles
707.

"

and

classes of animals

determinations

their

base

to

the
upon
Their
earlier

of the

body.
of feedingstuffs
digestibility
experimentsdeal chieflywith
of investigation,
Later
and
rations.
a
comprehensive scheme
outgo

well

as

as

the

upon

income

the

of the

includingdeterminations
and

begun

Table

168.

but

of

Computation

Clover

Chemical
Ash
Protein

Digestible

Nutrients

Maize

Hay

Meal

15-03

13-73

5-49

125
8.80

10.24

1.36

Non-protein

0.25

1.89

28.61

fiber

Nitrogen-free extract
Ether

of

composition

Water

Crude

laid out2

was

completed.

never

Example

"

excreta,

gaseous

36.98

70.44

364

2.29

extract

100.00

Percentage digestibility
Protein

Non-protein
Crude

fiber

Nitrogen-free extract
Ether

18.40%
66.43%
100.00%
32.40%
97-75%
95-74%)

46.48%
53-19%
100.00%
50.27%
68.94%
65.02%

Ash

extract

Digestiblenutrients
5.49X0.4648=

Ash
Protein

10.24

1.36 X

Non-protein
Crude

Ether

28.61

fiber

Nitrogen-free extract
extract

1. 000

0.5027

0.6894

X 0.6502

36.98 X
2.29

0.5319

2.55%
5.45%
1.36%
14-38%
25.49%
1.49%

Beitrage zur Begrundung einer rationellen Futterung

and 1864.
aNeue
Beitrage, etc., 1870.
1

1.25

0.1840

8.80

0.6643

0.25

1.000

1.89 X

0.3240

0.9775

3.64 X

0.9574

70.44

0.23%
5-85%
0.25%
0.61%
68*.85%
3-48%

der Wiederkauer,

i860

6oo

OF

NUTRITION

Table

170.

ANIMALS

Nutrients

Digestible

"

FARM

Reduced

Carbohydrate

to

Equivalent

Clover
Hay

Maize
Meal

5-85%

Digestibleprotein
Digestible carbohydrates equivalent

to

enous
non-nitrog77-54%

nutrients

Total

709.

nutrients

The

50.03

nutritive

ratio.

By the method

"

83-39%

just illustrated

is expressed
matter
digestible
which correspondto the two functions of the
by two numbers
nutrients alreadydescribed
(263) The digestible
proteinshows
the structural
contribute towards
the feedingstuff can
what
needs of the body, while the carbohydrateequivalentof the
non-nitrogenous nutrients shows what portionof the
digestible
of fat or of energy.
nutrients can serve
digestible
only as a source
The ratio between
these two quantities
gives a useful indication
of feedingstuffs is suited
stuff or mixture
as to whether
a feeding
for forms
of productionlike growth or milk production,which
requirea considerable supply of protein,or whether it is better
adapted for those which, like work or fattening,make special
the content

of

in

feeding stuff

demands

for fuel material.

This

so-called

"

nutritive

ratio

"

nutrient ratio)is obtained by a simpleproportion. Thus

(better,
in the two
instances just given,it is computed as follows,the
the nutritive ratio :
second half of the proportionconstituting
"

710.

For

clover

hay,

5.45

For

maize

meal

5.85 :

Significance of results.

44.58

77.54

"

1 :

1 :

Under

8.2
13.3

the stimulus

of Hen-

neberg and Stohmann's

pioneer work and under the leadership
of feeding stuffs
of Wolff, investigationof the digestibility
was
activelytaken up in Germany and later in the United
labor exStates and other countries,and as the result of much
pended
complete knowledge
during the last fiftyyears a fairly
nutrients supand proportionsof the digestible
of the amounts
plied
lated.
by most of the ordinary feeding stuffs has been accumuExtensive

tables

of averages

have

been

publishedby

RELATIVE

VALUES

OF

FEEDING

601

STUFFS

various

of the agricultural
authors,includingmany
experiment
for the feeder to learn what
stations,and it is an easy matter
and
amounts
kinds of digestiblenutrients
given feed or
any
ration
In

will

view

that

the

should

supply.
of the

understood.

the

feed

tables it is

As

units.

is inferior to

these

In

crudely,to
feedingstuffs in

that

methods

the

compare

the animal

which

of

matter

mere

comparison of feeding stuffs

nutrients
digestible
on

of such

use

of the results
significance

exact

be

extensive

actual

body.

the

on

based

the

important
they embody
logicalconception,
basis

hay

on

of their
values

or

ever
attempt is made, howeffects

produced by

determination

of

the

tibility,
diges-

the contrary, affords no direct information whatever

to the nutritive
effect of the materials digested. It is not
on

as

weigh the animal in a digestiontrial. The

comparison of feeding stuffs on this basis is between what they
contain and not between
what they accomplish.
nutrients have been of great
Nevertheless,tables of digestible
value in promoting more
rational and profitable
feeding,but it
even

is

to

necessary

evident
becoming increasingly

the

truth.

The

essential

comparison outlined
units of energy
a

basis of the

introduction

accurate

to

this volume

to

their

seek

methods

to

than

"

as

its content

" 3.

newer

is not

that
but

and

in

of matter

and

on

the

in

body

so

in

the

lumina
ilthe

These

1870.
elaborate

available to the earlier

were

at

were

Stohmann
"

Beitrage
determine, by more

Conditions

they employ
stitute
they con-

that

discussed

were

Neue

but part of
methods
of

express

the

basis of

the actual effect of the feed

well

of

and which
early methods
a
manner
by Henneberg

methods

newer

feature

they

comparison
the logical
conceptions which

as

return

in

that

and

ers,
experiment-

of the animal

as

energy.

Affecting

Digestibility

only is the current method

of estimating
the relative values of feedingstuffs,
described in
as
the previoussection,
based largely
of the mathe digestibility
terials
on
in question,but the latter is also a most
important fac711.

Digestibilityvariable.

das Ziel und

Uber

stationen

auszufuhrenden

die Methode

"

der

Not

auf

den

landwirtschaftlichen

thier-physiologischen
Untersuchungen.

Versuchs-

602

NUTRITION

OF

ANIMALS

FARM

determining the actual production values discussed

(742),the excretion
chapter,since,as there shown

in

tor
next

the

constituted

feces

greatest,

the

although not

in the
in the

only,loss

of

by the feed.
of a feeding stuff or of its several
The percentage digestibility
constituents,however, has not a fixed and invariable value,
of a chemical
compound, but may be
analogous to the solubility
less by a varietyof conditions,although to a
affected more
or
the
than is frequentlysupposed. This arises from
less extent
in Chapter III (155) that portions of ingredients
fact noted
tract
capable per se of solution and resorptionin the digestive
and reappear
in
actuallyescape digestionfor various reasons
in
the feces.
Any conditions which influence the digestibility
this way, however, necessarilyaffect the value of the feeding
method
stuff by whichever
determined, and the more
tant
imporchemical

suffered

energy

be

of them

may
conditions

in this connection.

convenientlyconsidered

affect,or which are supposed to affect,

the potentially
the degreeof completeness with which
digestible
of a feeding stuff are
vided
actuallydigested may be diingredients
into those relating
itself and those relating
to the animal
The

to

which

the feed.

Conditions
712.

which

Variation

factors

various

of the
found

be

must

to

at

different

borne

in

mind

the animal

times.
in

"

An

studying

important fact,
the

influences

of

digestion,is that the percentage digestibility

individual
has been
feeding stuff by the same

upon

same

vary

relatingto

more

or

less at different times.

especiallyby G. Kiihn.1 In experiments

of meadow
the digestibility
hay by cattle the variations in the
upon
of the dry matter, which
is the one
ject
least subpercentage digestibility
o.6 to 2.1, averaging 1.3, and the digestibility
to error, ranged from
variations.
for
of the organic matter
about
the same
That
showed
the nitrogen-free
extract
averaged 1.8,while in the case of the crude
fiber,proteinand ether extract it reached 3.3. These variations were
of experiment.
the possibleerrors
shown
to be materiallylargerthan
served
obsmaller, variations were
Similar, although relativelysomewhat
rations of hay and bran.
Kiihn points out that
Moreover,
on
the larger
differences were
found in those cases
in which
the maximum
This

has

been

shown

Landw.

Vers.

Stat.,29 (1883),129,

147

and

153.

OF

VALUES

RELATIVE

603

STUFFS

FEEDING

made.
No
could be traced
connection
singletrials were
and the condition of the animals.
the variations in digestibility
between
L
similar
The writer
observed
difference in two experimentsupon
a
one
sheep with clover hay while the other sheep of the pair showed
the feces
In later experiments 2 in which
difference.
no
significant
of three steers were
quantitativelycollected daily for periods of 56
that the digestibility
it was
shown
and 27 days on identical rations,
3
and nitrogencomputed from overlappingtenof the air dry matter
ment
day periods,varied at times from the average for the whole experiexperimental error.
by amounts
greater than the estimated
4
Hall
Emmett
likewise
and
observed distinct
Mumford, Grindley,
with
obtained
in
coefficients
the
cattle in sucfluctuations
cessive
digestion
of
from
to
two
weekly periodsfollowingpreliminaryperiods
of

number

four weeks.

dry feed and the writer

foregoingexperiments were
upon
is inclined to attribute them, to a considerable degreeat least,
to irregular
voidingof the feces.
All the

713.

of the digestive
differences in the anatomy
of different speciesmight naturallybe expected

Species.
organs

The

"

result in differences in the extent

to

speciesis digested.
of the feed whose

This

which

the feed of these

of those ingredients
especially
is due to the action of
digestibility

is true

so-called

organized ferments

to

less
or
which, therefore,will be more
dependent upon the opportunitieswhich the digestivetract
affords for the stagnationof the feed and so for the activity
of these organisms.
direct comparisons of
Few
714.
Species of ruminants.
the digestibility
of the same
feedingstuff by different species
and

"

of ruminants

are

alimentarycanal
find

to

record.

on

in these

comparatively

identical

view

species,one

small

feeding stuffs

In

are

of the
would

of the
similarity
naturallyexpect

differences in the extent

to

digested.

way

In

general

which

this

expectationis borne out by the average results of a largenumber

of recorded
digestionexperiments upon feeds bearingthe same
although not of identical composition. Thus Wolff,5
name,
in 1874,compared the results of about 40 German
experiments
1

Amer.

Penna.

The

of
4

Jour, of Science, 28 (1885),368.

Expt. Sta.,Bui. 42 (1898),pp. 129-141.
daily excretion of dry matter
not
determined
was

small

error

due

to

Ills.Expt. Sta.,Bui. 172

lack of exact

(1914).

and

there

is

bility
possi-

uniformity in the air drying.

BLandw.

Fiitterungslehre.

604

of nine

comparisons

that

considered
of ruminants
and

the

be

Such

made

similar

obtained

others

been

generally
one
species

with

without

the favorite
is

as

it has

material

error

imal.
experimentalan-

available,however, leads

to

of this conclusion.

modification

some

this sort

been

evidence

direct

have

experiments.

applied to

goat has

or

Hall

digestioncoefficients

may

sheep

American

ANIMALS

and

comparisonsof

the basis of

On

FARM

sheep and Jordan

cattle and

on

OF

NUTRITION

Comparisons of the digestivepowers of cattle and sheep for

identical feedingstuffs have been reportedby Frear,2the Mississippi
Station,3Bartlett,4Kellner,5Tangl and Weiser,6Zuntz,7
Voltz.8

and

experimentalresults,while

The

when

consistent,seem,

and

extensive

not

taken

in connection

togethe
al-

not

with

the

the conclusion
general comparisonspreviouslymade, to warrant
ence
that as regardsthe better grades of roughages the differbetween
cattle and sheep is not marked
in digestivepower
results,
and, with the exception of Zuntz's rather remarkable
be the case
the same
to
would
as
seem
regards concentrates.
of the coarser
On the other hand, it would appear that in the case
forms of forage a distinct difference exists in
and less digestible

favor of cattle.

is inclined

Kellner

greater percentage of

the

intestine of cattle

action of

extensive

The

715.

as

horse

Wolff9

The

most

in Hohenheim

in the

water

this difference to
of the

contents

compared with sheep,which

the organized ferments.
with

compared

comparisons have been

feedingstuffs by horses
animals.

ascribe

to

made
and

"

number

more

of

of identical
digestibility
by sheep as representingruminant

extensive
from

ruminants.

favors

lower

of

the

trials of

1877

to

1884

this sort
but

were

Tangl

and

made

by

Weiser10

of several
samples of
compared the digestibility
hay by horses and by sheep or cattle and Langworthy11has
compiled the results of a largenumber of digestionexperiments
also

have

U.

S. Dept. Agr., Office Expt. Stas.,Bui. 77

(1900),90.

3
Eighth Rpt. (1895), 79.
Expt. Sta.,Rep. 1890, 58.
Bui.
'Maine
5Landw.
Vers. Stat., 63 (1906), 313.
(1904).
no
Sta.,
Expt.
6Landw.
35
(1906), 205.
Jahrb.,
7 Jahrb. Ver.
in Deutschland, XII
(1912), 324.
Spiritus Fabrikanten
8Landw.
Jahrb., 45 (1913), 422.
9
Grundlagen fiir die rationelle Futterung des Pferdes, 1886.
10Landw.
Jahrb., 35 (1905), 159.
11 U. S.
Dept. Agr., Office Expt. Stas.,Bui. 125, p. 44.
2

Penna.

Table

171.

Digestibility

"

STUFFS

FEEDING

OF

VALUES

RELATIVE

Sheep

by

by

and

605

Horses

Digestibility

Percentage
Number
of

NitroCrude
Dry
gen-Ether
Crude
Organic ProExments Mattract
free
ter Matter teinFiber Extract
Experi-

ROUGHAGE

Wheal

straw

Sheep

45

Horse

20

23

57

59

Meadow

hay

"

Horse

hay

"

.......

hay

Meadow

"

54

68

62

46
23

47

47

57

39

56

56

59

57

56

62

51

47

48

57

36

55

24

62

64

65

63

65

54

42

57

20

average

Sheep
Horse

37
18

inferior

Sheep
Meadow

27

59

superior

Sheep

10

5o

5i

62

Sheep

65

76

73

80

76

65

Horse

54

62

69

57

66

13

55

56

50

61

56

37

63

29

Horse
Dried

Red

pasture grass

clover hay

5i

5i

56
56

58

59

7i

45

66

4i

58

58

73

40

70

14

70

7i
68

80

30

76

83

66

21

74

7i

87

90

87

79

9i

84

85

87

86

65

93

13

Sheep

90

89

66

93

75

Horse

80

83

Sheep
Horse

Alfalfahay
Sheep

12

Horse
Oats

Sheep

13

Horse
Beans

Sheep
Horse
Peas

77

89

97

78

78

94

5i

5i

27

79

62

9i

85
63

Lupins

Sheep
Horse

7i

72

Maize

Sheep
Horse

90

Results regarded by Wolff

as

9i

78

100

of questionable accuracy.

94

606

both

on

OF

NUTRITION

horses

rilminants.

contained

are

In

and

general,the

of ruminants

FARM

ANIMALS

The

results

parisons
com-

in Table

171.
have
shown
comparisons

over

of Wolff's

horses

in the

distinct

superiority
roughages,especially
and
tract)
exnitrogen-free
a

digestionof

(crude fiber
regards those ingredients
so-called digestionis wholly or in part a fermentawhose
tion.
in the better grades of foragethe crude fiber was
Even
the whole
on
by horses than by
considerablyless digestible
ruminants, although three of Tangl's experimentsare exceptions,
while less difference appears
as
regards the nitrogen-free
and scarcely
extract
any as regards the crude protein. On the
other hand, little difference was
observed
in most
in the
cases
of the total organicmatter
and nitrogen-free
extract
digestibility
as

of

In

concentrates.

fiber
the

the

the

latter

of
digestibility

the

crude

also

low but in view of its small amount

and
relatively
uncertaintyin the results little significance
consequent
was

attaches

this difference.

to

notably lower figuresfor the

The

of the ether extract by horses

digestibility
from
a
larger excretion of ether-soluble
in the feces of these animals
in

rather

than

arise in all probability

excretory
from

any

products

real difference

digestibility.

Comparisons of
the digestibility
of identical feeds by swine and by sheep have
been reported by Honcamp,
the feedand Milliner,1
ing
Neumann
stuffs being wheat, rye and the by-products of their milling.
Although the results upon the individual animals of the same
age
speciesfluctuated somewhat, as is not unusual (718),the averresults showed
the part of either
material superiority
on
no
species.
716.

Swine

Owing

to

compared

the small

with

ruminants.

of

percentage

crude

"

fiber contained

in the

ingredientare naturallyquite variable

and of no especialsignificance.Aside from this,there seems
to have
of the
been a slightsuperiority
the part of the swine in the case
on
rye products (with the exceptionof the germ) while with the wheat
with the coarser
the case, especially
was
milling
products the reverse
to have
products. The swine seem
digested the crude proteinfully
well
the
all
the experiments.
as
as
sheep in

feeds, the results upon

this

Bretsch, Losche
Fingerling,
to
especially
1

Landw.

Vers.

test

and

the relative

Stat.,81 (1913),205.

Arndt,2

in

signed
experiments de-

digestive
powers

of

sheep and

2Ibid., 83 (1913),181.

6o8

of

OF

NUTRITION

ether-soluble

more

also

or

the

higherthan

even

of

fiber

in

some

of crude

concentrates

cases,

but

are

in others

for ruminants.

obtained

those

of the small percentage

view

crude

swine

with

materiallylower

equal to

excretory products in the feces of these

figuresfor

The

animals.

ANIMALS

FARM

In

fiber in the concentrates

correspondingrange of possibleerror, however, the

results on
this point are of little significance.The crude fiber
Crude
of roughage is but imperfectly
digestedby swine.
tein
prowould appear, on the whole, to be rather more
completely
swine
than by ruminants, possiblyindicatingthe
digestedby
of more
nitrogenousexcretory products in the feces
presence
and

the

of the latter.
717.

of

collectingthe

feces

of

determinations

few

who
Bartlett,1

made.

Owing
fowls separatelyfrom

with

compared

Fowls

has

swine.

upon

in all recorded
Table

173.

number

"

Digestibility

the

the

difficulty

paratively
urine,com-

have

of such

been

ments,
experi-

digestioncoefficients

average

experimentsup

to

animals

these

reporteda

gives the followingas the

obtained

"

to

by

1910.
Fowls

Ether
Extract

Bran, wheat

Beef scrap
Beef (leanmeat)
.

Barley

Buckwheat

Maize,
Maize,

37.00

95.00

86.30

67.86

89.22

whole

88.11

cracked

Maize
Clover

87.60
87.60

India wheat

35-50

83.80

85.71

Millet
Oats
Peas

87.89
80.01

Wheat

53-oo
22.60

Rye
Potatoes

Maine

Expt. Sta., Bui. 184,

1910.

RELATIVE

VALUES

FEEDING

OF

609

STUFFS

to be relatively
difficultof digestion
fiber appears
by
this ingredientwere
able
varifowls and the results obtained
upon

Crude

apparentlycapricious. Aside

and
of

results with

for swine

those

from

this,a comparison
quite a close general

shows

classes of animals.
the two
agreement between
differences
718. Individuality. In addition to the specific
"

justconsidered,differences
of the
digestibility
same
species. To

such

also

to

seem

In their

in the

or

this may
be due to abnormalities,
chronic diseases of the digestive

animals

distinct individual

differences

occur.

compilationof
Jordan

evidence

extent

some

in normal

but

organs,

observed

feeding stuff by individuals of the

same

defective teeth

as

likewise been

have

and

the results of American

1

Hall

of such differences in

unable

were

experiments,
digestion

find conclusive

to

and

digestivepower

inclined

are

the apparent variations which were

observed
at different times already considered
to the variability

largely

to attribute

experiments by G. Kiihn,2however,

The

the discussion of the latter

of

is true

grade Shorthorn

steers

for five years.

Carmichael, Newlin

observed

under

to

afford indubitabl

differences in cattle and

experiments by the writer

were

cited in

were

also

seem
possibility,

instances of individual
same

which

(712).

observation

differences
significant

in which

the

three

at different times

wise
Grindley4 have likethe digestive
of
powers

and
in

individual

pigs. On the other hand, Christensen and Simpson 5

three series of digestiontrials on alfalfa hay for two successive
years, usingfour range steers each year, and failed to find

made

any

consistent individual

The

differences.

existence of time variations in

it somewhat

(712)renders
digestibility

difficult to decide whether

an

observed

difference

in the

digestionof the same

feedingstuff by two animals is really
whether it is in a sense
dental.
accior
expressionof individuality
A comparison based on a singledigestion
trial as ordinarily

an

made
results
over

1
2

3
4

is liable to

be

and
misleading,

to

correct

secure

of trials or
trial extending
a
requires either a number
is ordinarily
a longerperiod than
employed. On the

U. S. Dept. Agr., Office Expt. Stas.,Bui. 77

Landw.
Vers. Stat.,29 (1883), 129, 147, 153.

Expt. Sta., Bui. 42 (1898),124.

5 New
Science,July 2, 1915, p. 38.

(1900),88.

Penna.

Mexico

Expt. Sta.,Bui.

91

(1914).

610

NUTRITION

whole, however, the

the same
speciesmay

OF

FARM

conclusion

ANIMALS

that
justified

seems

differ to

animals

of

in their

digestive
that these individual differences are probably less
power
than appear
to be indicated by the results of singledigestion
much
trials and are certainly
in any degree
too small
to account
extent

some

but

for the economic

in

observed
three

Breed.

of

differences

Wolff

as

by

and

the differences

Even

digestiontrials rarelyexceed

usuallymateriallyless than this.

The
selves
foregoingfacts are sufficient of themimprobable the existence of any considerable

"

been confirmed

and

cent

render

to

breed

the results of short

four per

or

719.

differences in animals.

are

regards digestion,and

this conclusion

the

of

experimentsof Haubner
Armsby and Fries.3 The

taken

togetherfail to indicate any

digestivepower of different breeds

and

or

Hofmeister,1

recorded

material

has

data

difference in the

between

pure-bred and

scrub animals.
There
which

exists
show

somewhat

generalimpression that

animals

themselves

superior as producers of meat, milk,

of individual
or
etc., whether as the result of breed, heredity,
that superiority,
in part at least,to a superior
variation,owe
digestive
; that is,it is supposed that the improved breeds
power
of farm
breed
of

animals
are

able

to

feed than

and

can

the

extract

superiorindividual
more

nutriment

inferior animals.

The

animals

from

reasons

within

given weight
for the

doubted
un-

economic

of some
individuals over
others
superiority
have
in previouschapters. So far as differences
been considered
in digestive
are
concerned,however, the experimental
power
evidence
gives littlesupport to the popularimpression.
of the
720. Age.
Comparisons of the digestivepower
animals
made
same
(lambs) at different ages were
by Wolff 4
"

in

1871-72 which

six

and

fourteen

led to the conclusion

months

the

the ages of
of the
digestibility

that between

percentage

is
this conclusion
unchanged and
practically
taken
confirmed
by the results of an experimentby Weiske 5 underprimarilyfor another purpose.
the effect of the performance
721. Work.
on
Investigations
of work
of rations have
the digestibility
naturallybeen
upon

feed

remained

"

3
4

2 Landw.
Jahrb., 1 (1872),533.
Stat.,12 (1869), 8.
U. S. Dept. Agr., Bur. Anim.
Indus., Bui. 128.
5
Landw.
Ibid.,9 (1880), 205.
Jahrb.,2 (1873), 221.

Landw.

Vers.

RELATIVE

made
been

the

upon

VALUES

horse.

and

FEEDING

Experiments
and

reportedby Wolff

Grandeau

OF

LeClerc

6ll

STUFFS

this

upon

subject have

his associates at Hohenheim

and

by

Paris.

at

Wolff's1

animal,a draft horse

experimentswere
upon a single
weighing about 550 kgs. (1200pounds). The ration remained
in all the periodsand was
the same
insufficient to maintain
the
The
weight of the animal in the periods of heavier work.
work was
that of draft,done at a slow walk (about 1.9 miles
per
per
to

hour) with in most

instances

draft of 60

kgs.,the total work

day (notincludingthat of locomotion)ranging from 475,000
these
Under
1,800,000 kilogram meters.
conditions,no

effect

the

on

of the
digestibility

mixed

rations

employed

was

observed.
Grandeau
several

and

LeClerc's

different horses

experiments
both

at

upon

made
investigations2were
upon
of the Paris Cab
and included
Company
work
and others upon
motion
simple loco-

walk

and

together with

trot

rest

periments.
ex-

The

plan of the experimentsdiffered from that of Wolff's

in some
important particulars. The animals used were
lighter
(about 400 kgs. as compared with 550 kgs.)and apparentlyof
active temperament
indicated by their more
a more
as
rapid
walk, the velocityof which varied from 2.6 to 3.0 miles per
hour.
The work, which was
that of draft,was
done on a dynamometer
similar to Wolff's.

The

draft

was

about

half of that

of work 3 was
experiments and the total amount
less,ranging in most cases from 400,000 to 600,000
considerably
in one experimentof
kilogrammeters per day, with a maximum
in all the
was
785,000. Its amount
approximatelythe same
experimentsin each series and was not greater at a trot than
at a walk.4
Finally,correspondingto the main purpose of the
to study the feed requirements of cab
experiment,which was
horses,the rations in the periodsof work or of walking exercise
of Wolff's

heavier

than

in those in the

periods of rest, the increase

beingone-tenth in the experiments on locomotion and, in most
work.
The
cases, one-half in the experiments on
proportions
were

Landw.

L'alimentation

Not

Jahrb., 8, Ergzbd. I (1879),73 ; 16, Ergzbd. Ill (1887), 53-71.

du cheval de trait; Berger-Levrault et cie,1882-89.
that
of locomotion.
including

The

total work

in the former

case

of the greater expenditureof energy

was,

of course,

in trottingas

somewhat

compared

account
greater on
with walking (664).

6l2

NUTRITION

of the different
the

feedingstuffs

except in

same,

OF

FARM

in the

few

very

rations,however, remained

cases.

whole, and despite some

on
singleingredients,Grandeau
with Wolff's in showing that work
On

ANIMALS

the

in
irregularities
and

LeClerc's
at

even

the

results

results

somewhat

agree

rapid

affect the digestibility

of rations.
On
materially
the other hand, they show a distinct decrease of the percentage
the work
in the periodsin which
done at a
was
digestibility
that this effect can
be ascribed to the
It scarcely
trot.
seems
walk

does

work

as

not

such,since

experimentsand

the measured
not

was

horizontal

mere

greater

at

at

less than

was

than

at a

trot,in

some

trot

locomotion

produced the

to have

least,seems

amount

walk.

over,
More-

instances

which
effect,

same

in Wolff's

at

apparently

gait. It is true that the rations were

heavier
in the work
periods and that this (722)may possibly
is apparent why
have
but no reason
affected the digestibility,
it should have produced a greater effect in the trotting
periods
is due

to

the difference in

in the

than
The

walking periods.

influence of work
1

and
by Tangl
after 36 hours
hours

by
of

later and
and

has

Scheunert.2

A
the

fasting and
the

analyzed.

animal

of

contents

On

investigatedin a different way

of oats
fed
was
weighed amount

also been

the

killed from

was

and

stomach

the

to

one

small

five

intestines

of the

crude
none
assumption
of
the
was
portion
alimentary tract,
the results show
the
of
the
that work
feed from the
delays
passage
stomach to the intestines,
especially
during the first one or two hours.
the gastricjuicepenetrates the largermass
As a consequence,
of the
feed more
of it is neutralized
slowly and more
by the saliva,so that
the stage of starch digestionis prolonged and that of proteindigestion
shortened, the result being that more
carbohydratesand less protein
are
digested. In the later stages of digestionthe differences tend
the intestinal
to equalizethemselves, while
the effect of work
upon

weighed

fiber of the oats

digested in this

found

digestionwas

that

to

be small.

Scheunert

was
digestibility

As

considerablyincreased by
noted, however, the results cover
only

of digestion. The
and

one

themselves.

question of
1

Arch.

Arch.

the

Only

method

of

results show
actual

comparison
very

computes
the

performance

the

first five

is

upon

Physiol. (Pfliiger)
; 65 (1896), 545.
Physiol. (Pfliiger)
; 109 (1005), 145;

or

confessedlyan

considerable

digestionexperiments

the total effect of work

that the total

of work.
six hours

mate
approxi-

variations

suffice to

among

decide the

digestion.

Landw.

805.
Jabrb.,34 (1905),

VALUES

RELATIVE

OF

Conditions
722.

rations
the

by.

of

Quantity

regard

the

feed.

relatingto
"

Current

of
digestibility

consumed.

amounts

FEEDING

As

the

613

STUFFS

feed

methods

of

computing

stuffs

unaffected
as
feeding
regards exclusive feeding

of exnumber
periments
roughage, the results of a considerable
by various investigatorsappear to justifythis view.
mixed
fewer exconcentrates
rations of roughage and
With
periments
been
have
made, but the results indicate a distinct

with

the

ration

when
percentage digestibility

the

in

decrease

is increased

considerably above

the

that

of

amount

requiredfor

maintenance.

on
earlyexperimentsof Henneberg and Stohmann
in which
of clover and
cattle include several cases
varying amounts
the digestibility
fed and in which
meadow
was
substantially
hay were
and
the
found
unaffected
to be
same
was
by the quantity consumed,
of
Hohenheim
in
number
true by Wolff at
a
experiments on sheep.1
2
the
Later
same
on
investigator
sheep
comparative experiments by
confirmed
his earlier results,
have
also those of
and horses have
as
while
Tangl and Weiser 3 upon sheep fed alfalfa hay or alfalfa silage,
4
distinct effect of the quantity
likewise found
and Girard
Miintz
no
of alfalfa hay by the horse.
In
the digestibility
consumed
upon
5
each of two steers
subthree experimentsby Armsby and Fries
on
a
ration of timothy hay was
maintenance
slightlybetter digestedthan
five
of
ration in
out
cases
a maintenance
six,but the differences were
small, amounting to from 1.0 to 2.7 per cent on the dry matter.
Later unpublished experiments have given similar results. In earlier
authors on different amounts
of clover hay,
experiments 6 by the same
observed.
differences were
no
practically
Kellner 7 obtained
Mixed
the followingresults in four
rations.
ration consisting of
of a mixed
periods in which varying amounts
meadow
hay, dried molasses beet pulp, rye bran and cottonseed meal

Roughage.

"

The

"

fed

were

and

to cattle.

Compare

Wolff, Die

Ernahrung

der

landwirtschaftlichen

Nutztiere, pp.

63

64.
Landw.
Vers. Stat.,21 (1878),19.
Jahrb., 1 (1872), 533;
and
3Landw.
Vers. Stat.,74 (1911), 277
282.
4 Centbl.
Agr. Chem., 27 (1898), 756.
5U. S. Dept. of Agri., Bur. Anim.
Indus., Bui. 128 (1911),p. 27.
6U. S. Dept. of Agri.,Bur. Anim.
Indus., Bui. 74 (1905),pp. 12-13;
2Landw.

101
7

(1908),pp.

11-13.

Ernahrung

landw.

Nutztiere, 6th Ed.,

p. 49.

Bui. No.

6 14
Table

174.

Effect

"

FARM

OF

NUTRITION

Amount

of

ANIMALS

Mixed

of

Ration

Consumed

on

Digestibility

of mixed
rations sufficient
digestibility
thirteen
months
and
for maximum
milk production,
later,when the
ration of the same
of
mixture
cows
were
dry, that of a maintenance
of the ration
feedingstuffs. About thirtyper cent of the dry matter
from hay, thirty-six
derived
was
per cent from silageand thirty-four
per cent from grain.

Eckles

Table

175.

Mumford,
of four

the

determined

Digestibility

"

Grindley,Hall

different

mixed

Mixed

of

and

determined

Emmett3

rations

cattle

of

Rations

and

hay

than

Cows

by

the

grainby

digestibility
pairsof

four

maintenance
receivingrespectively
slightly
One animal of
one-third feed,two-thirds feed and full feed.
lower digestivepower
than the
fed pair showed
a
distinctly
more

all the

periods

two-thirds
animal

feed

in this

the

same

case

are

shown

true

was

in the last two

Expt. Sta., Research

Approximate.
Mo.

and

of

periods.

one

The

in the table in

ration,

animal

the fullother

receiving the
the other

results upon

parenthesis:

"

Bui. 4.
3

Ills.Expt. Sta.,Bui. 172

in

(1914).

6i6

would

One

NUTRITION

OF

naturallybe

inclined to ascribe the lower

of heavier

rations

rapid passage

more

the

through

lessened

consequent
It would

in

the

been

on

resorbable

feces.

Some

indicated

in

ANIMALS

their greater bulk and

the digestivetract, and
of

extent

that

seem

and
digestible
appear

to

FARM

the

bacterial

liberal rations

may
of the

thus
reasons

in part to

tions.
fermenta-

material

as

tially
poten-

digestionand

escape
for

discussingthe feces

digestibility
relatively

this have
a

ready
al-

feed residue

of undigested
(155). The presence of considerable amounts
grains or fragments of grain in the feces of heavilyfed animals
is readilydemonstrated
but
by washing out the finer portions,
actual
the extent
of this
digestionexperiments to determine
loss have not yet been reported.
723.
of carbohydrates.
It has been established by
Excess
numerous
experiments that an undue
proportionof carbohydrates
in a ration tends to reduce its digestibility,
especially
by
ruminants.
The effect is most
distinct when
bohydrates
carpure digestible
added
also when
to a ration,but is manifest
are
large
of
amounts
feedingstuffs rich in carbohydratesare introduced.
An
example of the former is afforded by an experiment by
1
divided
into three periods,
in
It was
G. Kuhn
two
on
oxen.
the first of which
the animals received a dailyration of 9 kgs.
of hay to which, in the second
and third periods,
2 kgs. and
3.5
adcfed.
of
starch
the
were
Assuming
kgs. starch,respectively,
of
been
the
to have
followingamounts
completelydigested
the several ingredientswere
computed to have been digested
from the hay by Ox V.
"

Table

178.

Nutrients

"

Digested

from

Hay,

with

and

without

Starch

Ether
Ex-

Period

tract

Grams

No

II

kgs. of starch

41

III

3.5 kgs. of starch

38

starch

Landw.

No

Vers.

starch

40

Stat.,44 (1894),470-472.

could

be detected

in
microscopically

the feces.

617

STUFFS

FEEDING

OF

VALUES

RELATIVE

Periods II and III

in

the feces
Exnressed in another way,
which ,
of hay ingredients
the followingamonnts

coined

ac-

digestible
escaped

be regarded
must
the results of Period I,
of the addition of starch
which under the influence
as

cordme
but

to

digestion.
Table

179.

Escaping

Hav

or

Nutrients

DtoEsxiBLE

Feces

in

NlTROORGANIC
Matter

Dry
Matter

Period

^f
Pro-

CRTOE
Fiber

free

Kgs. starch
starch
3.5 Kgs.

II

III

232

217

44

635

632

15"

example of
foregoingis a typical

The

IGrams

I Grams

Grams

Grams

tract

Extract

Grams

"THER
Ex.

GJW-

tein

Grams

76
180

299

the results of numerous

on'
siir
^erimtnts
^"t,hay
added
cellulose
magnitude
mted
Other thingsbeing equal,
^"^*X
usuallyincreased,
total
in which

ruminants

substances

and

to

ana

to

been

have

even

"-

the

of the

rations.

in

as

effect has

has

amount

Its

added.
tityof carbohydrates

vara

con

Jerably
in different f*^""*""%?Z"
"

uniformly the

been

has

of
depression

coefficientsare
of

lowered by the escape

theseTn
stances

or

of

less of the added

to

Wolff,1swine

appear
of carbohydratesthan are
Grandeau and Alekan

and

ruminants
"

by

or

crease

of

be

so

as

^s""

that

well

as

the

same

such

:=

terial
ma-

adecan

^"""jU"

six experimentswith

Stat., 19 (1876),273.
Chem., 49 (1906),350;
A
gr.
Jahresber.

iLandw.

few

nkewi*
apparently

clearlyfollowed

carbohydrates. Thus
"

as

in

may

some

to tins influence

ve

indicate

to

seem

in

r lations
althoughthe quantitative
digestibility,

always

not

that

""*""""

to

^Feeingst^rich
carbohydrates,
Isstarch
sugar,

MgJ

carbohydrate(starch)

less sensiU

be much

diges ion

potential

itself escaped digestion.According

ha

course

be noted, however,

It should

more

tne

constitutes

since, of
digestibility,"

in the feces.

matter

This

same.

beet

molasses

'

Vers.

Ann.

Sci. Agron., 1904,

T
I, 30,

33c

by

6i8

OF

NUTRITION

Lehmann
like

and

by Kellner

later trials
of the crude

the

substance,

depressionin

basal

only effect

this

the

ration,but

was

the

on

gestibil
di-

in two

digestibility

fiber.

questionhas

Wolff in

that

showed

carbohydrates, caused
pure
of all the ingredientsof a

the

The

Kellner

by

one

ANIMALS

FARM

been

regard to the

of these materials

however, by
especially
investigated,

feedingof

tubers

and

ing
Heavy feed-

roots.

is

generallystated,on the strengthof his

experiments,to result in a pronounced decrease in the digestibility
of the remainder
of the ration,although,as Wolff himself
conclusive.
pointsout, the evidence is by no means
In

Wolff's

quantitiesof
had

extensive
roots

been

potatoes

or

were

added

to the ration

of

increasing amounts

180.

Computed

"

the
as

is
the

to

seems

compute

the

assumption

case

(161)and

show
Wolff

has

series of

any decrease
carried out the

as

Sugar

as

the

feces contained

the

see

sugar

beets

"

method

of

Hay

of

with

and

whether

comparison, however,

in the

roots

of
digestibility
to

the

the

first instance

hay, just as

on

in the

the coefficients thus obtained

proportionof
this

roots

fed is increased.

manner

for his entire

singletrials.

Landw.

Jahrb., 25 Erzgbd. II (1896), 117.

Stat.,53 (1900), 199.
3Ernahrung landw. Nutztiere, 5th Ed., pp.
4 Landw.
lahrb., 8 Ergzbd. I (1879),123.
2Landw.

that

Beets

computation in
experiments,numbering in all no
1

that

Digestibility

of the
digestibility

of unaltered

of concentrates

found

was

increased,the

follows

reasonable

more

it

assumption

Percentage
without

What

was

and

undigested nutrients. For example, in one

hay and sugar beets,the percentage digestibility

experiment with meadow

of the hay, computed on
was
were
completelydigestible,
Table

experiments on sheep 4 increasing

fed along with hay whose digestibility

previouslydetermined,

of roots

amount

series of

Vers.

158-175.

The

aver-

RELATIVE

VALUES

OF

FEEDING

619

STUFFS

results for total

and for nitrogen-free

extract
organicmatter
are
uniform
in
series
of
each
there
was
what
somefairly
experiments,although
the
individual
variation
in
and
fail
to
more
trials,
give anyof a diminished
with
the
decided indication
digestibility
increasing
The computed digestibility
of roots consumed.
of the crude
amounts
less variable,
but on the whole
or
a decreased
proteinis more
tibility
digesof this ingredientas the proportionof roots
to hay increased
to be plainlyshown.
seems
age

is,of

It

in which

roots

of the fecal

added

are

matter

determine
to

is derived

in

digestion
experiment

basal ration what

from

the

roots

and

tion
propor-

what

from

remaining ingredients,and such results as those of Wolff

be interpretedeither as showing a fairlyconstant
tibility
diges-

the
may

of both

the

on

or,

impossibleto

course,

showing

hay.

To

reasonable
the

hay and the roots (asidefrom crude protein),

of the roots, as
assumption of complete digestibility
of the
progressive depressionin the digestibility

the

the

writer,the former

althoughit should

course,

experimentsthe
increased

was

by
consumed,

appears

an

absolute
amount

amount

on

the whole

be added
of crude

the

that in

more

some

of

fiber in the feces

greater than that contained

in the

thus

demonstratinga depressionof the digestibility

of this constituent of the hay. Probably the truth
lies between
the two
views.
It is unlikelythat roots
tirely
enare
digestibleand, on the other hand, it is probable that a
the digestibi
extent
largeproportion of them may diminish to some
roots

of other

It is
with them.
feeding stuffs consumed
to be remembered, however, that roots
contain not altogether
inconsiderable quantitiesof crude proteinwhich, as shown
in a
following paragraph (727),tends to offset the effects of their

carbohydrates.
725.

Cause

of

diminished

of protein.
digestibility
"

tion
Atten-

alreadybeen called (163-167)to the influence of the

tibility
excretory productscontained in the feces on the apparent digesof the nutrients and especially
of protein. According
to Kellner's and
Pfeiffer's results,the digestionof each 100
material,
of dry matter, whether proteinor nitrogen-free
grams
results in the excretion in the feces of approximately0.4 gram
of nitrogenin the form of these excretory products. If,then,
to a basal ration,the nitrogenous
a kilogram of dry starch be added
excretory products in the feces are increased by aphas

620

OF

NUTRITION

proximately25 grams,
is digestedfrom
digestionmay

have

not

ANIMALS

that

apparently25

basal

ration,while

so

the

FARM

been

affected.

less of programs
tein
in reality
the true
in Kiihn's

Thus

iment
exper-

hay and starch (723) the nitrogenous excretory

of the
products corresponding to the 1646 grams dry matter
consumed, would be approximately 40 grams,
2 kgs. of starch
with

while

the

actuallyfound

excess

was

44

the

grams,

difference

being insignificant.
The

is

agreement

instance

and

in

been

cited

was

so

close

as

the addition

experiments on

protein determined.
seems
justifiedthat at

influence

the

of

bohydrates
car-

digestibility

true

Nevertheless, the
least the

in this

general

largerpart

of the

of

bohydrates
carbohydratesand of feeding stuffs rich in carof the proteinof the
the apparent digestibility
on
to the fact that,when
added to a basal ration,they

feed is due
increase

the

the

other

the

true

nitrogenous excretory products in the feces. On

be remembered, that while
hand, however, it must
be lowered, it is, as
not
digestibility
already
may

(167),the apparent

pointed out
of

have

always

means

the

conclusion

the

no

of the

none

which

(166) of

by

real

advantage

its feed.

derived
the

Whether

in the

feces

by

which
digestibility
the

increased

animal

from

excretion

measures

the
of

digestion
nitrogenous

after

carbohydratefeedingbe due to an
to both
bined,
comor
apparent or a real depression of digestibility,
it is none
the less a loss of protein from the body.
In general the depression in the percentage digestibility
of
matter

the

protein is greater
As

justifythe

the
*

Kellner

gredient
the basal ration is in this inpoorer
has pointed out, however, this does

frequentlymade that the magnitude

of the depressionis dependent upon
the nutritive ratio of the
The
difference
is purely a mathematical
feed.
crease
A deone.
of the digestibility
of the protein by 50 grams,
for example,
is relatively
greater in a basal ration of oat
very much
crude
straw, containingonly 140 grams of apparently digestible
protein than in a basal ration of meadow
hay containing 430
not

grams
The

of

statement

digestibleprotein.

fact that

apparent

the

addition

of
digestibility

readily explicablefrom
1

Landw.

the
this
Vers.

of

protein tends to decrease the

protein of a basal ration is also
Pfeiffer's experpoint of view.
Stat.,44 (1894),344.

VALUES

RELATIVE

OF

FEEDING

621

STUFFS

(162)showed that the increase in the nitrogenous excretory

whether
the
productsin the feces was about the same
consisted of carbohydrates
added digestible
matter
or of protein.
Consequently,the addition of proteinto a ration would tend
of the protein just as
to diminish the apparent digestibility

iments

the addition

would

carbohydrates.

of

when given in
non-proteins,
especially
vegetablematerial and roots, likewise increase
The

of the feces,but

the

tent
nitrogenconreview of the literature of the subjectl

of the proteins,
the increase consists,
case
that,as
least in largepart, of metabolic productsand does not indicate
of the protein,
although
digestibility

decrease in the true

any
it does, of course, decrease
726.

diminished

of

Cause

available

the amount

of
digestibility

to

the

organism.

carbohydrates.

"

of the non-nitrogenous ingredients

depressionof digestibility

The

of the feed of ruminants

different cause, viz.,to
in the rumen,
and

more

entirely

an

the fact that these effects are

cesses
pro-

observed

insoluble carbohydrates
of
comparatively

less of the

or

the feed

to

lends strong support to this view.

stated (128-132)that the disappearance

already been

It has

due

in the fermentation

modification

be

to

appears

this class of animals

chieflyon
of

of green

in the

shows
at

the form

during its passage through

the

alimentary tract

is

due,

in ruminants, to a bacterial fermentation, occurparticularly

ring
in the first stomach
and yielding
principally
chieflycarbon

dioxid,methane
shown
and
the

and

that when
are

sugar,

organic acids.

the

soluble

more

introduced

of

methane.

It

can

of the
is due

like
carbohydrates,

into the ration

organismsand undergo
amounts

Furthermore, it has been

the

the

scarcelybe

same

they

are

attacked by

fermentation, yielding
responding
cor-

characteristic
doubted

that

product,
gaseous
gestibil
the decreased di-

less soluble
to

starch

carbohydratesunder these
partialdiversion of the activityof

cumstance
cir-

the

resultingfrom the action of

the saliva on the starch or to the sugar directlyadded, since
these substances
are
readily attacked than
presumably more

organisms to

ferment

cellulose and
The

the like.

action of

effect of

an

excess

explainedas
"

the maltose

Compare

nitrogenous substances in counteractingthis

of readilysoluble carbohydratesis plausibly
due to its supplying more
nitrogenous food
U. S. Dept. of Agr.,Bur.

Anim.

Indus., Bui.

139-

62

OF

NUTRITION

ANIMALS

FARM

and
organisms and so stimulatingtheir multiplication
and the fact that readilysoluble nitrogenousmaterials
activity,
salts seem
acids or ammonium
like amino
to be particularly
action
of
The
effective is quite in harmony with this view.
nitrogenous materials in stopping the excretion of undigested
starch,on this view, would be explainedas due to an increase
of the proportionfermented, leavingless to be acted on by the
juicesof the intestines.
digestive
for the

of

addition

of

Effect

727.

protein.
"

It

shown

was

in

the

bohydrates
containinga largeportion of cardeficient in protein,i.e.,
therefore relatively
and
those having a wide nutritive ratio,are
likelyto show an im-s
Correctingthis
especially
by ruminants.
paired digestibility,

paragraph that

last

rations

by increasingthe proteincontent

condition

increase

of such rations

tends,

their

digestibility.
of the effect of
by
investigators
the addition of nearlypure protein(wheat glutenwith 78 per
fish meal with 96 per cent of crude
of crude protein,
cent
or
proteinin the organic matter)to a basal ration. In general,
of the
such an addition has had littleeffect on the digestibility
nants
proteinof the basal ration,but in several experimentson rumiof crude fiber,
increased digestibility
cases
an
and, in some
with
of the nitrogen-free
extract, has been observed,especially,
in protein. In other
basal
rations poor
instances,however,
in protein was
when
the deficiency
less marked,
particularly
this effect has been either slightor entirelyabsent and the
is true of such experimentson
swine
have
been thus
as
same
the
record
in which
also on
far reported. Experiments are
addition of feedingstuffs rich in protein,such as oil cake or
of the crude
increased
the digestibility
legumes,has distinctly
be

would

as

expected,to

Trials have

of

fiber
has

basal

Effect

ration
excretion

stopped an

728.

made

been

of

as

and
of

others

"

of

in which

undigested starch

non-protein.

of ruminants
extracts

several

The

addition

digestible
non-proteinin

rule tends

to

diminish

such

an

addition

in the feces.
to

the basal

the form

the apparent

tion
ra-

of

plant
digestibility

to increase the excretion

proteinof the basal ration,i.e.,
of nitrogenin the feces,
while the simplerforms of non-protein,
such
as
salts,have not usuallyproasparaginor ammonium
duced

of the

this effect.1
1

Compare

U.

S. Dept.

Agr., Bur. Anim.

Indus., Bui.

139

(1911),pp. 14-28.

624

FARM

OF

NUTRITION

ANIMALS

order that the green foragewas.

Armsby
similar experiment upon
a
subsequentlymade

same

and
a

Caldwell

stantiall
by sub-

cow

plan, using mixed grasses cut while still

and
to pasture grass, and
corresponding substantially
young
has reported comparisons of the same
three
sort
on
Morgen
the

same

The

sheep.
in

Table

results of the four

average

While

181.

comparisons are shown

to
experiments are
open

earlier

the

the whole
the conclusion apon
particulars,
pears
of forage is not
warranted
that the digestibility
very
and
diminished
removal
of
that
water
materially
by the simple
the lower value of ordinarydry roughage as compared with green
forage is largelydue to differences in maturity and composition.
of coarse
ders
fodThe digestibility
730.
Cutting of roughage.
is not increased
ficult
by cutting,and, indeed, it would be difin

criticism

some

"

conceive

to

in either

since

that

how

the feed

case

the same
practically
experiments by Kellner

extent.

to

chaff

and

was

carried

Table

182 shows

straw

cut

731.

182.

Grinding
to

his

into inch

Table

in

of

This
the

which

the extent

of

have

such

effect,

an

during mastication
in
shown
strikingly
preparationof straw

is

grindingit to

fine meal.

comparisonbetween wheat straw and barley

and a half lengthsor finely
ground.

grain.

"

solvents,their

Cut

The

and

outer

Finely

coats

Ground

of seeds

are

When

Landw.

sistant
re-

being to protect the seeds from

in large
whole grain is fed,especially

purpose

Expt. Sta.,Rpt. 1888, p. 60; AgriculturalScience, 3, 295.

Vers. Stat.,75 (1911), 321.
Ernahrung landw. Nutztiere, 6th Ed. p. 266.

iPenna.

to

Digestibility

"

external influences.

could
process
is comminuted

VALUES

RELATIVE

FEEDING

OF

STUFFS

625

animals with

imperfectteeth,
and, protectedby the
more
or
unacted
tract relatively
outer
coats, passes through the digestive
intact grainsof corn, oats, etc., still
Such apparently
upon.
of
are
a familiar sightin the droppings
capableof germination,
amounts

greedy feeders

to

or

to

mastication

less of it escapes

heavily fed animals.

visible losses,however,

Such

are

not

confined

of whole

grainbut, although less obvious, extend

crushed

grain as

well.

to the
to

feeding

cracked

or

If, for example, the feces of full-fed

grainbe washed out, a

considerable amount
be recovered,
of fragments of grainmay
the amount
sequent
dependingupon the total quantityfed and the conrapiditywith which it passes through the digestive

cattle

tract.

receivingcracked

corn

or

other

Moreover, it is evident that the mechanical

separation

necessarily
imperfect.Not only may the sieve
but it is likeother things than fragments of grain,
wise
that any undigestedfragments of the latter which

by washing
hold back
clear

is

through and
be lost,so that fine meal or well-masticated grainmight suffer
a
greater loss through incompletedigestionthan would be
indicated by such tests.
While it is to be supposedthat smaller
fragments will undergo more
rapid solution in the digestive
of passage
tract than largerones, it is evident that the rapidity
are

smaller than

through the

the meshes

organs

is

an

of the sieve will pass

important factor and that

small bits may,

under some
while on the other
complete digestion,
whole

grainmight

be

almost

the results
Qualitatively,
'

well

as

even

paratively
com-

circumstances, escape

hand, with lightfeeding,

digestedas

when

ground.

reached

by washing out the feces are

readilybe misleadingas regards

but they may

of great interest,
the actual advantage of grinding.

the relative digestifew investigations

Surprisingly
bility
upon
of ground and unground grainhave been reported. Jordan
and Hall,1in their compilation
of American
experiments
digestion
comparisonswith horses
up to 1900, present two
and two with swine,all of which show the ground grainto be
than the unground,the difference with respect
more
digestible
to the dry matter
rangingfrom 3.3 to 14 per cent.
Gay,2in experimentsupon oats with a horse weighingabout
1
2

U. S. Dept. Agr., Office Expt. Stas.,Bui. 77, P- 97Centbl. Agr. Chem., 25 (1896),
729.

626

OF

NUTRITION

ANIMALS

FARM

kilograms(750 lb.)and receivingper day 3 kilogramsof

the following
results :
and
2 kilogramsof hay, obtained

340
oats

"

Table

183.

Digestibility

Percentage

"

of

Oats

by

Horse

Ether
Extract

Whole

40.90

....

Crushed
.

59-46

54.78

Ground

Gain

by crushing

18.56

Gain

by grinding

13-88

While

results

the

just cited

are

or

more

less

variable,and

of the nitrogen-free
digestibility
in Gay's experiments seem
extract
peculiar,the results as a
whole
by swine and
clearlyshow an increased digestibility
that the
horses as a result of grinding,while they also show
less perhaps than
difference is apparently not
great
very
would have been expected.
Gay also reports the followingresults of similar experiments
a
sheep weighing 81 kilograms and eating500 grams of
upon
of alfalfa hay:
oats and
750 grams

while the small differences in the

"

"

Table

With

184.

the

"

Percentage

exceptionof

it is difficult

to

Digestibility

the

determine

ether

no

Oats

extract, whose

by

Sheep

digestibility

accurately(165),the percentage

is practically
identical
digestibility
as

of

in the

three

cases.

So

far

singleexperiment goes, therefore,it indicates that there is

advantage in grinding oats for sheep. Experiments upon

RELATIVE

it does
whole

not

other

more

lacking,but

are

should

ruminant

627

STUFFS

feeding stuffs

surprisingthat
completely than

appear

grain

with

and

other ruminants

FEEDING

OF

VALUES

non-ruminant.

digest
As

whole, the results upon the influence of grindingon digestibility

and in particularthey afford no
formatio
inare
comparatively meager
to the

as

of whole
digestibility

the relative

in the amount

effect of variations

grain and

of

coarse

fed upon
fine
or

meal.
732.

Acids.

extensive

The

"

of

use

question of the influence of acids

feeding stuffs.
the

silagelends interest to
the digestibility
of
on

of meadow
hay with and
digestibility
acid (0.75
without the addition of sulphuric
per cent SO3) by one
sheep,using two periodson each ration,and obtained almost
crease
absolutelyidentical results,with the exceptionof a slightinin digestibility
of the ash and ether extract of the acidified
hay. Kellner 2 added a much largerproportionof lactic acid
(2.67per cent)to a ration of hay and maize fed to a sheep and
likewise observed practically
effect on the digestibility.
no
of organicacids as are ordinarily
Apparently,then, such amounts
in silageand other feeds are without effect
consumed
on
digestionin the case of ruminants and this conclusion is to
a certain extent
supportedby the generalresults of experiments
which have shown
that ensiled forageis fullyas digestible
as
of acid conthe same
The amounts
material carefully
dried.
sumed
under normal
pared
conditions are after all not largeas comized
and neutralwith the quantitiesproduced in the rumen
l

Weiske

compared

by the
stimulate
and
for

saliva.

be

be

Condiments.

for the various

to

excessive

amounts

of

acids

may

produce scouring is doubtless true,

such as the horse,
presumed that other species,

example,may

able

That

and
peristalsis

it may

733.

the

increase

so

sensitive

more
"

One

to

of the

acids than ruminants.

exaggerated claims

proprietarycondimental feeds
of
materiallythe digestibility

is that

made

they

are

rations to which

slightestscientific basis for this claim

exists. All experimentersagree that they are without influence
for exin this respect. Recent
ample,
investigationsby Fingerling,3
in which fennel,anise,fenugreekand malt sprouts were

they are

added.

Jour. Landw.,

Landw.

Not

the

2
Ernahrung
(1885),21.
Vers. Stat.,62 (1905),41-57.

33

landw. Nutztiere, 6th Ed., p. 56.

628

both

added

FARM

OF

NUTRITION

ordinaryfeeds

to

ANIMALS

and

flavorless materials

to

ration made

showed

effect upon

no

of abnormally

up

the percentage

digestibility.
Water

734.

writers

drinking.

the

on

by
particularly

feeding tends

after
feed

his animals

Even

far from
sort

of

of farm

drinking on

been

that

asserted

gastricjuiceand

the

numerous

feeder has

digestion,
drinking

to wash

been

the

advised

to

would

be

warranted, since

the

stomach,

serves
being the sole organ of digestion,
largelyas a
reservoir
the
and
extensive intestines
(119),
preliminary
animals
afford ample opportunityfor the digestion
of

substances

any

by

feedingrather than after feeding.

supposed facts true, it is questionablewhether

drawn

conclusions

and

laid

before

the

were

It has

dilute the

to

of the stomach

out

water

the

horse.

been

of water

supposed effect
the

has

Stress

"

of

matter

dilution

which

may

fact,however,

occurs

has

as

action in the stomach.

As
escape
such washing out
no
or
degree of

been

has

As

supposed.

already been

semi-solid rather
of the stomach
are
(131),the contents
than
much
less
liquidand, as shown
by their stratification,
mixing of them takes placethan is sometimes imagined. Scheul has
shown
that in the horse the largerpart of the water
nert
stated

drunk

well

is

filled with
in

retained,but
stomach
water

has

and

feed.
no

contents

which

did

exceed
the

enters

the

In

case

material

no

stomach

and

quite promptly discharged into

is especially
the case
when
the

This

intestine.

of the

along the walls

passes
and
contents

contrary

the

about

effect in the

the

of

small
is

stomach

is

water

more

total- dilution
io

duodenum

case

its

around

the

entire
the

Moreover,

per cent.
is rather

rapidlyresorbed
of feed into the
transportation

largeintestine.
In

of

view

these facts it is

few
a

trials which
have
digestion
decrease
in digestibility
as
a
Gabriel and

Weiske

difference

in the

hay, whether
organic matter
1

Arch.

Landw.

the
was:

been

surprisingto
show

made

find that the

no

evidence

of

drinking after eating.

result of

cant
experimentson two sheep found no signifiof a ration of oats and
percentage digestibility
2

the water

before

not

in

was

given

animals.

The

before

or

after

percentage

feeding or kept

of the
digestibility

"

144 (1912), 411

Physiol. (Pfliiger),
Vers. Stat.,45 (1895),311.

; 151

stantly
con-

(1913),396.

629

STUFFS

FEEDING

OF

VALUES

RELATIVE

II

Sheep

Water

Watered

before

Watered

after

Tangl,1
found

in

feeding,

nearly
and

water

tended

Jahresber.

to

on

before

when

in

this

found

that

in

the

secretion

decrease
animals

Chem.,

to

seem

on

28

the

(1899),

be

in

effects

661.

dog

general
of

water

Expt.

was

restricted

gastric

of

supply

of

with

Rec,

going
fore-

those
in

drinking
Sta.

results

The

juice.

accord

or

less

also

the

are

water

during

digestibility

of

of

watered

connection
the

horses,

consumption
were

corresponding

Suggestive
who

the

they

different

four

upon

drinking,

than

the

that

studies

Agr.

experiments

less

case.

extensive

of

was

and

Lambert,2

results
Hawk's

watered
and

every

Foster

number

when

irregular

after
in

feeding
feeding

that

was

animals

the

before

constantly

25

of

man.

(1911),

16.

CHAPTER

PRODUCTION

THE

"

General
the

By

"

chapter,is

weight of

the

FEEDING

OF

STUFFS

Considerations

the

relative values

the actual

meant

of

production values

from
distinguished

stuffs,as
the last

VALUES

i.

Definition.

735.

XVII

feeding
in

discussed

effect produced by

unit

in

substance

maintaining an animal or in supporting

the processes of growth and fattening
of
milk
work
or
or
production. That such production values will also express
relative values scarcely
needs mention.
Even
the
at their best, comparisons based
on
digestible
"

nutrients,"such
have

become

only the

as

have

familiar

relative and

It is true

that

to

been

in vogue
for many
years and
show
all students of the subject,can

to

the absolute

not

the

extent

which

to

values

of

it may

be

feedingstuffs.
assumed

that

the

nutrients as determined
digestible
by analysesand digestion
experiments actuallyconsist of proteins,carbohydratesand

fats,their

furnish
may
value of the material consumed.
amount

useful clue to

Even

the

nutritive

then,however, it affords

expected,while in
of most
the case
as
appeared in Chapters II and
feedingstuffs,
material has been but very
of the digested
III,the actual nature
incompletelyinvestigated.Neither the chemistryof feeding
quantitativemeasure

no

stuffs

nor

the

of the

results to be

of their various

behavior

constituents

in metabolism

is

the basis for any

well known
to serve
as
sufficiently
ter
trustworthyestimate of their actual nutritive effect. The latbe determined
can
only by a direct trial with the animal,
and during the past two decades considerable progress has been

made
736.

the
a

in this direction.
Determination

production value

unit of

of

production values.

is the effect

the feed under

produced upon

consideration.
630

"

The

By

definition

the animal

by

generalmethods

632

to in Part

referred

III

evident that,
renderingit increasingly

are

quite aside

from

of

the

nature

and

intensityof

energy,

the

course

its value

it appears
substances
may

growth,
while, on

supply of

the

the

feed

more

or

as

ticular
par-

yet ill-defined

observations
to

less toxic character

the

on

clusive
ex-

indicate that similar

follow

may

the

regardedas
consumption of feedingstuffs ordinarily
definite a
It is important,therefore,
to secure
as
of these new
facts
as
possibleof the significance
relation to the older conceptionsof productionvalues.

sive
exces-

ful.
healthtion
concep-

in their

Significance of "accessory ingredients." It is clear

738.
that

and

in
limitingfactor, particularly
diseases,
development of specific

products(499)seem

of wheat

use

In

processes.

of certain

absence

material

profoundly influence

metabolic

the

constitute

structural

may

lead to the
may
the other hand, McCollum's

or

effects of

as

of

that

ANIMALS

FARM

OF

NUTRITION

the

"

"

ingredients (usingthis simply as

accessory

summary
indicated
in the

for the

term

various

venient
con-

classes of substances

paragraph) influence the nutritive value

of a feedingstuff in an
different fashion than does
essentially
the quantity of available ash, protein,and
which
it
energy
of productionwhich the
supplies. The latter limits the amount
feedingstuff can support ; the presence or absence of the former
the extent to which
this potential
determine
value is actually
may
in
realized. Thus
Chapter XI, experimentsby Osborne and
and by Hart
and McCollum
Mendel
described
(498,499) were
that a mixture
of pure nutrients may
which show
be prepared
which shall contain an abundant
supply of complete,proteins,
of ash and of energy but upon which young
animals (rats)
fail to
grow,

last

while the addition

of substances

associated

support normal

In

of minute

mixture

certain

with

growth.

"

such

to

amounts

fats enables the rations to

aspects of the matter,

some

these

"

accessory ingredients might be crudelycompared with the

lubricants of a machine, which
furnish neither
of themselves

material,but

nor

power

consumption of
conduce
A

The
so

to

the

fuel to

which
be

enable

more

production of

lack of lubricants in the

power

used
efficiently
largeroutput.

case

raw

material

would

and

from

the

therefore

ably
justsupposed might conceiv-

affect the output of a machine

in one
undue
friction might slow down
the
that less

derived

pass

or

both

of two

ways.

as

whole

machine

through

it in

given time,

PRODUCTION

THE

VALUES

FEEDING

OF

633

STUFFS

certain more
delicate parts of the
might affect specifically
and cause
of the machine
machine and so reduce the efficiency
it

or

it

to

yieldless finished productper

which

In

determined.

been

in the

two

ways

affects the nutrition of

"

have

of these

unit of

of

case

"

deficiencyin

animal does not

an

It would

stances
sub-

accessory

to

appear

however, that,
probable,

seem

animal, for example,a deficient dietary

young

material consumed.

raw

stop the whole group of anabolic processes

involved in growth.1 The organs would thus be rendered incapable
to slow down

acts

of

convertinga normal dailyamount

and

substances
would

or

decrease
corresponding

follow.
presumably

that such feed

eaten
actually

was

as

requirementmight
have

In such

be

just as

in feed

in

excess

efficient in
as

words, it is conceivable that lack of

In other
"

in

may,

sense,

considered. In

accessory

regulated
by
has

producing gam
m

the

"

The

normal

not

stances
sub-

accessory

rather than

the

writer has failed to

this aspect of the matter

has been
the

substances,"the feed consumption has

of the
appetite
been reported.

the

even

and

ration.

all reported
investigations
practically
upon

"

influence of

ration.

consumption

of the maintenance

affect the economic

of the
efficiency
physiological
in which
.note any experiments

body

into

it is quite conceivable

case

value per unit

great a production

as

been

of feed

animal

and

in

many

stances
in-

undoubted

ot
importance of the accessory ingredients
writers,to a tendency
feedingstuffs has led,on the part of some
which as yet appears hardly justified
the signifito minimize
cance
values in the older sense.
The subject
of the production
and the field too broad to warrant
is too new
clusions,
dogmatic con-

The

but

it stillremains

feeding stuff is

that the

prime

supply structural material

to

function

and

in this respectare
potentialities

the

body, and

its

productionvalues.

its

true

That

the

results attained

energy

for

in
expressed

by its use
long been

practiceare affected by other considerations has

for
recognized. Thus, if a feeding stuff is unpalatable
and is not
eaten
reason
freely,the portion consumed

in

of

some

may

show
a
high nutritive effect per unit and yet the use of the
feed be inadvisable. The presence of toxic substances might
Naturally such
specificmetabolisms
1

metabolisms

an

effect might be brought about by a retardation of certain

which the whole growth process depended and the specific

upon

affectedmight differin different cases.

634

NUTRITION

prevent the
and

yet

tolerance

of

use

the

OF

ANIMALS

feed in sufficient amounts

nutritive

might

FARM

effect of the

feed

to

within

be

profitable

the

limits of

be considerable.

Production

of balance
values,then, if determined
by means
experimentsmade under normal conditions,are to be regarded
as
of
showing the potentialvalues of feedingstuffs as sources
and energy, i.e.,
their worth as constituents of a ration
matter

which

contains

sufficient amounts

"

are

The

study of

"

important group
of feedingstuffs
potentialities
that in the future

already outlined
for the various

"

the

the normal

ensure

"

of factors

energy

the
influencing

for

be

must

ash, protein
of

purposes

Production

Values
Energy

extent

which

to

to

(or amino

the

It is possible

the

ments
require-

acids) and

the requirementsfor
feeding,

necessary
organs

Regards

as

apparentlyvery

added

"

efficient functioning
of the cells and

2.

in the broadest

actuallyutilized.

are

there

substances

accessory

"

gredient
inaccessory
of metabolism.
course

substances
accessory
has revealed an additional and

of the term

sense

to

necessary

"

of whatever

to

the

secure

of the

Energy

most

body.

"

Net

Values

Recapitulation. The consideration of the processes of

nutrition in Part II, and in particular
the study of metabolism
of the balance
and
in Chapters V and VI, has
of nutrition
739.

"

body is primarilya transformer of

the most
and
chemical
that quantitatively
important
energy
In the several
function of the feed is to supply this energy.
chaptersof Part III the conceptionof net energy values was
developed and the requirementsfor net energy by different
cussed.
disspeciesof farm animals and for different purposes were
shown

that

It

the

animal

is apparent

from

those

discussions

that

the

net

productionvalue of
defined in the preceding
a
as
feedingstuff
section.
It appears
desirable at this point,therefore,to recapitulate
of the
the general facts regarding the energetics
in previous chapters, and
animal body which
contained
are
the production
in greater detail their bearingupon
to consider
of a certain amount
values of feedingstuffs,
at the expense
even
of repetition.
energy

value

is

only another name

as
regardsenergy

for the

PRODUCTION

THE

740.

Gross

contained

energy.

FEEDING

supply of an
energy
chemical
and
the
energy,

as

635

STUFFS

The

"

in its feed

OF

animal

is

maximum

furnish for the vital activities

can
any substance
in the body is measured
its oxidation
bustion.
by its heat of comwhich

amount

by

VALUES

This

called its gross energy

(315) to avoid
the implication that it represents the total amount
of energy
with

associated
741.

has

been

the

Losses

of

substance.

energy.

that this maximum

case

the feed escapes

general classes.

rarely,if

proportionof

unutilized.

These

ever,
happens, howpractically
every

ever,

effect is realized.

smaller

larger or

It

"

In

the chemical

losses of energy

energy
are

of

of two

First,a portionof the chemical energy of the feed fails to be

transformed
at all,leaving the body as chemical
in the
energy
and in the combustible
visible excreta
from gastric
gases arising
and

intestinal fermentations.

Second, another portion of the chemical energy of the feed is

indeed transformed, but at ordinarytemperatures virtually
results
merely in a superfluousheat production. It is true that
the metabolism
consequent upon feed consumption is not only
be regarded as a necessary
unavoidable
but may
expenditure of
with digestion
energy for the support of the activities connected
and assimilation.
Nevertheless, from the standpoint of the
net gain or loss by the organism this portion of the feed energy,
which ultimately takes the form of heat and escapes from the
body, must be regarded as a loss.

The
742.

in the

of

derived

not

the

none

ration

With

Chemical

"

undigestedfeed

constitutes

must,

chemical

of

on

escapes in the feces

which
they contain and
While

the latter

the feed

consumed,

they carry.

immediatelyfrom

loss of

less,be

the energy

herbivorous

energy

energy

residues

excretory products which

portion is
but

in feces.

Losses

in the

both

losses

incompletelykatabolized matter, it
included in estimatingthe net effect
balance of the body.

animals,

the

excretion

in

the

feces

stitutes
con-

the greatest loss of chemical energy, and the one

which
varies most
different feedingstuffs or different species
as between
of

animals,as is apparent from

the results recorded

in Table

187

636

NUTRITION

is

(749). This
much

contain

the

With
the
The

FARM

ANIMALS

true of the energy

especially
indigestible
matter, but even

materials

because

OF

loss

swine

former

through

it is

animals

"3

of the

with

feces is

the

the

The

herbivora
on

trates.
concen-

the

upon
alreadybeen

urine

gestible
di-

more

siderable.
conrelatively

conditions

has
digestibility,
previouschapter.

feces,i.e.,
upon

roughages,which

less than with

relatively
are
usuallylargelyfed

influence of various

the

of

losses in

discussed

in

is

the vehicle
especially
from
the body of the end products of protein
for the removal
is the most
familiar and frequently
katabolism, of which urea
abundant.
Numerous
other nitrogenoussubstances,
the most
the purinsand,
however, are contained in the urine,particularly
in herbivorous
animals, hippuric acid.
Moreover, as stated
in Chapter V (224,225), the urine of herbivora
in particular
contain
considerable
quantitiesof non-nitrogrelatively
enous
may
of
which
little
excretory products regardingthe nature
is known.
All these substances
carry off a portionof the gross
743.

in

Losses

urine.

"

of the feed as unused

chemical
of
the amount
energy
energy,
the loss being measured
That
by their heats of combustion.
the extent
the

of these losses cannot

of the urine

nitrogencontent

The

loss of chemical

satisfactorily
computed from
has alreadybeen pointedout.

be

in the

energy

urine, as

from

appears

relativelysmall percentage of the

total loss.
be expected,it is quite variable,being
As would
higher as the feed supply is richer in protein,and lower with
substances
where the loss in the feces is
indigestible
relatively
larger.
744.
Fermentation
The
losses.
products,chiefly
gaseous
gestive
of
fermentation
of
the
the
methane,
carbohydratesin the diof ruminants, carry off contract of herbivora,especially
siderable
Table

187,

constitutes

"

of

amounts

chemical

unused

methane, for example,having

energy,

one

of combustion

heat

gram

of

of 13.344

Calories.
The
methane

Armsby

extent

to

fermentation
and

Fries

greater
the

same

which

the

be
to
appears
observed
that
have

light than

on

carbohydratesare

on

medium

likewise observed

authors
1

Jour. Agr'l Research,

a
3

attacked

somewhat

with

cattle

by

the

variable.
tinctly
it is dis-

heavy rations and

singleinstance of indior

(1915),445.

viAud

difference in this

Zuntz

and

in
instances
reportedstriking
m
particular
fermentation
methane

associates" have

Ms

theextent

whkh

Recently

animals

respect between

637

STUFFS

FEEDING

OF

VALUES

PRODUCTION

THE

of the

Tasteen
markedly affectedby
especially
by the order

in which

the feeds

and

the rations

make-up of

the

consumed, while

were

the practical
stress upon
"
laid much
his associates have
differences
results. No such marked
importance of these
but the range
Fries' experiments
observed in Armsby and
It is perhaps too ear y
not so wide.
eedhig stuffs used was
but they should
of Zuntz's results,
udge of the full significance
and

V61U

were

Tf
to

C
It

held by not
of

and

as
digestion

of

coefficientas
digestion

the other

On

few,

notion,

the

to correct

serve

hand, however,

it

"?^!%""*
process
perfectly
definite

sort

constant.

of chemma

quitepossibe

seems

to

over-

Simate
the effect of such

Joule

net

energy
to

they appear

variations in the digestiveprocess

On the whole
values of feeding stuffs.

of far less

be

other factors to

than
significance

%rpSceentagerof

is lost in the ferthe gross energy which

Table 188 (749)
appears from
the case of feedingstutts
in

"JntLgases',
as

""""

greatest
It is naturally

large

^iSri-l^te^espeda^toea-l
verv

neimental

S,

the

While

ex

and

losses in feces
determination of the energy
simple
relatively
easy task, requiring
comparatively

annhances

the

determinationof the fermentation

teTthe of
sfta
use

the

piratTon
apparatus.
to
it is possible

ever

losses.

fermentation

Computation of

745

of accuracy
degr'e

somewhat

of such

apparatus,how

an

losses with

compute the fermentation

the results of the

methane
experiment
The

neces-

and costlyrescomplicated

In the absence

from

losses

fair

ordinarydigestion

fermentation attacks chieflyor

of cattle
Xly the carbohydrates(135,140) and the of methane
duced
prothat the
shown
particular,
it has
of total
proportionalthe
is in general
digestd
fcrudeand nitrogen-free
Shydrates
in

amount

been

extract

fiber

with

experiments
Kellner3 in forty-four
the average

car

amount

to

rations found

methane

excretion

cattle
to

Landw.

7SHS0
765 ; Landw. Vers. Stat.,
Jahrb.,44 (1913),
(1913),3*5Jahrb.,44 (1913).68S; 45

Landw.

Vers. Stat.,S3

.Landw.

in

case

415(1900),

on

mixed

be 4.2 grams
(."), 78..

638

NUTRITION

for

100

estimated
in

methane
of

addition

Fries

and
for

for concentrates.

4.7 grams

for the smaller ruminants

In

basis.

With

the

age
unpublished experiments,Kellner's l averLater experimentsby Armsby
to 4.3 grams.
given slightly
higheraverages, viz.,4.8 grams

roughages and

it may

that

on

later

have

computed

were

increased

was

ANIMALS

of total carbohydratesdigested,
and the
grams
results for ruminants
recorded
in Table
188 for the

each

losses

FARM

OF

been

have

that the average

safelyassumed
applicableto these speciesalso.
the horse the principal
seat of the
and

(128). Since

ccecum

of the feed

similar

sults
re-

reportedbut probably

be

is the colon

No

for cattle is substantially

fermentation

methane

the

hydrates
soluble carbo-

more

ing
or entirely
digestedbefore reachlargely
these organs,
methane
is much
less copiouslyproduced
than in the case
of ruminants
and may
be regardedas derived
of crude fiber.
chieflyfrom the fermentation
In respiration
rations
of oats, hay
experimentson mixed
and

the

Lehmann,

straw,
result

of

of

gram
more

of methane

fiber and

hydrogen

recent

and

Zuntz

eight rather

total excretion
crude

are

discordant
of 4.73

in addition

per

observed

experimentsan
grams

per

100

as

average
grams

gested
di-

excretion of 0.203
digestedcrude fiber. In

an

average

100

grams
experiments,Von der

Hagemann

Heide, Steuber

and

Zuntz,4

obtained
using a Regnault-Reiset respiration
apparatus (298),
for the methane
excretion
digestedcrude fiber
per 100
grams
on
9.06 grams
hay and 2.28 grams on straw
pulp. Using
the average
of these rather
mentatio
discordant
experiments,the ferlosses in the case
of the horse may
be approximately
of crude
fiber digested.
computed from the amount
Swine with their simpleralimentarycanal suffer but small
losses from fermentation in the digestive
tract.
5
and
found
of
the amounts
Reinhardt
Fingerling,Kohler
combustible
their

form

Klein

obtained

gases excreted
of Pettenkofer

in three
the

small

too

to

apparatus.

experiments with

followingresults

be
Von

determined
der

Heide

with
and

Regnault-Reisetapparatus

"

Ernahrung landw. Nutztiere, 6th Ed.,

Jour. Agr'l Research, 3 (1915),p. 450.
4
Biochem.
Ztschr.,73 (1916),161.
6Biochem.
Ztschr., 55 (1913),195.

p. 94.
3

Landw.

Landw.

Jahrb., 23 (1894),125.
Stat.,84 (1914),197-

Vers.

640

NUTRITION

tract
digestive

the

OF

in the

or

ANIMALS

FARM

of metabolism

course

in the tissues.

(323),this convertible portionof the

feed energy has been given various
gators,
names
by different investisuch as "physiological
heat value," fuel value," available
energy,"etc., but followinga suggestionmade earlier by
the writer it is here designatedas
metabolizable
energy."
stated

As

in

VI

Chapter

"

"

"

Method

747.

of

determining.

is apparent

As

"

from

the

of the metabolizable
foregoingparagraphs,the direct determination
of a feedingstuff or ration requires
the measurement
energy
of the

and

of the

outlined

and

amounts

solid,liquidand

in

heats

of the feed

of combustion

excreta
by the methods
gaseous
These
quantitiesbeing known,

a
Chapter VI.
Thus
the
simple subtraction gives the metabolizable
energy.
results of the experimentused as an illustration in Chapter VI
detailed form, were
more
as follows :
(322),
put in a somewhat
"

Table

187.

Example

"

Determination

of

Heat
of
Combustion
Dry
or
Matter
per
Gram

Daily feed
hay

Linseed

meal

Daily

Energy

Energy
Feed
of

of

Excreta

Cals.

Cals.

Timothy

Energy

Metabolizable

of

4,556

Cals.

27,727

1,811

5,iii

excreta

Feces

4,831
0,23c1

Urine
Methane

14,243
1,210

1,896

13,344

Metabolizable

By difference

energy
.

29,538

748.

Correction

for

experiment the animal

protein and therefore
energy, viz.,
In
In

loss

gain or
gained
stored

Per

protein.
"

In

the

foregoing

66.6 grams
of fat and
15.2 grams
up in its body equivalent amounts

protein,5.7 Cals.
fat,
9.5 Cals.
1

of

12,189
29,538

66.6

15.2

gram

380 Cals.
144

fresh urine.

Cals.

of
of

PRODUCTION

VALUES

Cals. of energy

contained

THE

The

144

actuallytransformed

not

into other

transformation had

such

641

STUFFS

fat,however, although

in the

capableof
requiredit,

forms

of energy, were
of the organism

demands

the

FEEDING

OF

and therefore constitute part of the metabolizable energy of the feed.

With the 380 Cals. contained in the proteinstored up, however, the
would

their energy

been katabolized,part of
grams
bolic
escaped in the resultingnitrogeneousmeta-

Had

is different.

case

these

have

66.6

products. According to Rubner

derived from

each gram of urinary nitrogen

is equivalent
Cals. of chemical energy.
to 745

lean meat

katabolism

The

would
of the 66.6 grams of protein,
therefore,
chemical energy of the urine by 83 Cals, while

the

Cals. would

have

transformed.

been

This

When

respiration
apparatus

combustible
from

the

outlined

an

of

case

with

as

loss of protein

amount

to
possible

the

olizable
estimate the metabfrom
has

of the

visible excreta.

the

been

additional labor

The

requiredis so small that it is to be hoped that

digestion
experimentsit may be undertaken whenever

thus

that

in future

possible

in this way

the

extensive data may

be secured remore
garding
metabolizable energy
of feeding stuffs. While

results do

not

show

(750),they constitute

the

values
production

of the rations

important step toward

an

their

more

determination.

exact

Experimental

749.

limited

number

with swine in which

Results.

of

from

the

There

the losses of energy

been

considerablylargernumber

estimated

"

are

experimentswith

have
respectively

methane
a

ready
al-

manner

ordinary digestion
experimentto which

of combustion

such

mated
esti-

the collection of the urine and determinations of the heats

added

and

of the
be

may

considerable degree of accuracy

measured

for the determination

carbohydratesin
digestible
that it is

only 297

evidentlybe subtracted.

their
available,

not

so
(745),

energy

results of

is

gases

In

creased
in-

of 83 Cals. must

amount

correction to the urinaryenergy

consequentlybe added as a
in computing the metabolizable energy.
from the body a similar correction must

have

on

what
some-

cattle and

in the

determined

record

few

feces,urine and

while
directly,

the losses of methane

have

in

been

digestible
carbohydrates(crudefiber plus

extract)in the manner

nitrogen-free
sults
just described. The rerecorded in Table 188,1which
of these experimentsare
shows
1

the percentages of the gross energy

This table is not

to be

claimed to be

fairlycomplete.
2

an

exhaustive

which

were

compilationof data,

carried
but

is believed

642

NUTRITION

several

off in the

which

OF

FARM

the percentages
and, by difference,

excreta

metabolizable.

were

The

organic matter
digestible
it forms a
subsequently,

of

gram
appear

of metabolizable

of it

energy

metabolizable
is also

added, since,as

convenient
when

energy

basis

for the

per
will
putation
com-

direct determinations

available.

not

are

ANIMALS

Table

188.

"

Apparent

Metabolizable

Energy

3a
a s
"

Author
So
"
"

o
Cals.
Cattle

Roughages
hay
Meadow
hay
Timothy hay
Red clover hay
red
Mixed
timothy and
clover hay
Alfalfa hay
Hay from irrigatedmeadows
Ensiled hay

Tangl,
Armsby
Armsby

et al.

Armsby
Armsby
Tangl,
Tangl,

and

Fries

3-390

and

Fries

3-605
3.600
3.698

Oat

Kellner

3-740
3-3IO

Meadow

Kellner

....

straw

Wheat

straw

Kellner

Straw

pulp

Kellner

Maize

stover

Armsby

3-5oi
3-437

and Fries

3.487

and Fries

3.486

et al.
et al.

3-640
and Fries

3-45Q

Average

3-529

Concentrates

Maize

meal

Wheat

bran

Hominy

chop
grains No.
grains No.

Mixed
Mixed

i
2

Millet
meal

Palmnut

Distillers'

slop

molasses

Beet

molasses

Distillers'

residue

Fries

4-075

Fries

3-9io

Fries

3-879
3-787
4.849

536i

Kellner

3-473

4.519

et al.

Kellner

4.287
3-603

Kellner

4-792

Starch

gluten

2.703

et al.

Tangl,
Tangl,

Wheat

3-954

from

beet molasses

3-797

Fries

Voltz, et al.
Voltz, et al.

Pumpkins

grapes

Fries

(from

potatoes)
Beet

Armsby and
Armsby and
Armsby and
Armsby and
Armsby and
Tangl, et al.
Voltz, et al.

Average

4.078

Estimated.

Not

corrected

to N.

equilibrium.

VALUES

PRODUCTION

THE

Table

FEEDING

STUFFS

643

(Continued)

Energy

Metabolizable

Apparent

ii

OF

Author

Sheep

Roughages
hay
hay

Meadow
Meadow

....

....

from

peat meadows
alkali soil

from

same,

Hay from
Hay
Hay

irrigated

Alpine hay

TangI, el al.
Voltz, el al.
Tangl, el al.
Tangl, el al.
Tangl, el al.
Tangl, et al.

Average

Tangl, et al.

Alfalfa hay

Average
Dried potato vines
Same

fruit

with

Voltz, et al.
Voltz, et al.
Voltz, et al.

....

Hay and dried potato vines

ensiled potato
and
Hay

Voltz, et al.
Voltz, et al.

vines

Wheat

straw

Average
Concentrates
Oats

Tangl, et

Millet

Tangl,
Tangl,

Corn-and-cob meal
Palmnut

Distilleryslop from
molasses

et al.

Voltz, et al.
Voltz, et al.

meal

Lentils

Beet

al.

el al.

toes
pota-

Voltz, et al.
Voltz, et al.

....

23-2

18.6

Average
Horses

Meadow

Roughages
hay
....

peat meadow
alkali soil

Hay from
Hay from
Hay from same, irrigated
Alpine hay
Sour meadow hay
Silagefrom same
.

Tangl, et al.
Tangl, el al.
Tangl, et al.
Tangl, et al.
Tangl, et al.
Tangl, et al.

Tangl, et

19

0.8

59-3

1.61
2.0

50.4

al.

55-1

66.1

50.1

1.61

66.4

1.5

70.0

1.61

"

39-4
29.4

45-4
43-

44.6
28.4
24.7

3-707

3-854
3-803
3-741
3.915

3.607
3-352
3-712

Average
Concentrates
Oats

Distillery residue
grapes

Tangl,

et al.

41.4

Tangl,

et al.

66.9

3.7

0.21

54-7

4-493

from

and beet molasses

0.8

30.9

4-76i
4.627

Average

Estimated.

Not

corrected to N.

equilibrium.

644
Table

88

(Continued)

Energy

Metabolizable

Apparent

"

ANIMALS

FARM

OF

NUTRITION

Losses

Percentage

Author
In
Urine

In
Feces

In
Meth-

ane

O
M

w
hi

w
u

o
n

P*

"**
,

"

Oats,
Oats,
Oats,
Oats,

hay
hay
hay
hay
Oats, hay
Oats, hay

Horses

Rations

Mixed
and

straw

Zuntz

and

Hagemann

4-474

and

straw

Zuntz

and

Hagemann

3-236

and

straw

Zuntz

and

Hagemann

3-403

and

straw

Zuntz

and

and

straw

Zuntz

and

straw

Zuntz

Average

Hagemann
and Hagemann
and Hagemann

3-803
3.980
5-Q52

3-991

....

Swine
Concentrates

Tangl,
Tangl,

Millet

Pumpkins
Barley and

gluten

Starch

pulp

Straw

Sugar
oil

Peanut

dried yeast
Same

palm oil

dried

Computed
Computed
Whole

potatoes
for oil
dried

milk

milk

Skim

milk

and

saccharified

and

raw

and

al.

4.908
4.076

al.

3-952

al.

3-75o

al.

8-997

V. d. Heide

and Klein

4-237

V. d. Heide

and Klein

4.442

V. d. Heide

and

4.160

V. d. Heide

and Klein

V. d. Heide

and Klein

Klein

9-552

toes
pota-

starch
Skim

5.629

al.

milk

Skim

4.521

al.

...

for

al.

and

Barley, dried potatoes

Same

3-460

Fingerling,et
Fingerling,et
Fingerling,et
Fingerling,et
Fingerling,et
Fingerling,et
Fingerling,et

meal

Wheat

4-335

et al.

little flesh

meal
Flesh

et al.

starch

4.111

9.8

4-7

Wellmann

3-4

11.3

4-519

Wellmann

3-8

10.5

3-825

Wellmann

3-5

7-5

fat
.

5-467

Wellmann

5-994

Averages
Flesh

meal

and

wheat
5. 269

gluten
Whole

5-467

milk
....

Peanut
Other

oil
.

8-997

rations

4-055

Geese
Maize
Millet

Tangl,
Tangl,

et al.

23-9

3-753

et al.

43-8

2.723

Tangl,

et al.

53-6

2.323

Ducks
Millet

Estimated.

VALUES

PRODUCTION

THE

Significance of

750.

olizable energy,

metabolizable

the

645

STUFFS

energy.

metab-

By

"

simply the
body, with no implication

in the

transformation

proportionof

the

to

as

FEEDING

already explained, is

as

capable of

energy

OF

meant

transformed

thus

energy

utilized

by the organism. The heat evolved

during the methane fermentation,for example,constitutes part
of the metabolizable
thus defined,although it does
as
energy

which

not

be

can

into the tissue metabolism.

enter

metabolizable

The

of

energy

its productionvalue, since it takes

the

classes of losses

two

of the feed energy

estimation

is thus

its chemical

rejectedunused.

of the metabolizable

important step in
value as regardsenergy, and constitutes an
since
simple determination of digestibility,
and

Real

751.

energy
in the

manner

of

so

regardsenergy
on
one

the
of

obtain

as

thus

energy

advance
it takes

the

over

account

of

of those in the feces.

energy.

by
of
digestibility
urine

the

The

"

olizable
metab-

the basal
were

respectively.By
272.3
here used, the algebraic
of these
sum
the

is subtracted

from

its metabolizable

computed

the

energy.

expresses

the

available for conversion

net

The

time

amounts

of

putation
com-

is virtually
from

metabolizable

body

and

the

of the latter

energy

energy

increase in the amount

in the

the

by 1865.9

method

losses of energy
gross

ried
car-

ration that

increased

the

molasses

amount

Cals.

and

to

as

of energy
135.8 Cals.,while at the same

the
regarded as representing
molasses

more

or

determination

digestive
processes.
Kellner's experimentsbeet

lost in the feces and

Cals. and

one-half

fixing

of all the influences which

basal ration diminished

depressed the

amounts

in

factor

feedingstuff as determined experimentally

illustrated in a precedingparagraph (747)is the

off in the methane

it

is subject.

feedingstuff exerts
For example, in
to

well

as

apparent metabolizable

aggregate effect as

added

methane

of

only one

feedingstuff is,
ascertainingits production
energy

therefore,an

the losses in urine and

ure
meas-

energy

The
of

not

of

account

Obviously,however, it is an essential
from one-fourth to
value,since frequently

that

or

which

to

does

feedingstuff

may

of

be called

the apparent metabolizable

energy.
On the other hand, the results for the metabolizable
of the

nutrients
digestible
corrections

for these

recorded

in the next

energy
clude
paragraph in-

secondaryeffects. They

aim

to show

646

NUTRITION

the actual

OF

FARM

ANIMALS

of metabolizable

amounts

energy
of its
digestedportionsof the feed irrespective

i.e.,to

"

give a

its real metabolizable

express

more

in the

contained

Such

energy.

energy is better
of questionsof feeding.1The

adapted for

use

already discussed in Chapter III (167)between

apparent digestibility.
of metabolizable

Computation

While,

"

metabolizable

in the

absence
of

the parent
apin a discussion

distinction is similar

that

nutrients.

energy

while
feedingstuff regarded by itself,

metabolizable

752.

figures

of metabolizable

idea of the store

accurate

suppliedby the
secondaryeffects

energy

of

from

to

real and

digestible

respiration
apparatus,

feedingstuff or ration may be

estimated with a fair degree of accuracy
lined
by the method outin previousparagraphs,not every experimenter
is equipped
the

to determine

energy

the heats

of combustion

of the feed and

the visible

of computing them is availmethod

able.
satisfactory
been made to compute
Various attempts have accordingly
of feedingstuffs from chemical data.
the metabolizable
energy
and by Atsuch method
One
is that employed by Rubner
of the food of
for estimatingthe metabolizable
water
energy
and of carnivora as described in Chapter VI (324),their
man
factors for protein,
carbohydratesand fat beingapplieddirectly
nutrients of feedingstuffs,
and several tables
to the digestible
of energy values as thus computed have been published. Later
however, showed that the results thus obtained
investigations,
much
of herbivorous
too high in the case
pecially
animals, eswere
cite but a singleinstance,experiTo
of ruminants.
ments
in Table
cattle by the writer
on
gave the results shown
189 for metabolizable energy as compared with those computed
earlier
by the use of Rubner's factors,and Kellner's somewhat
results 3 led to the same
generalconclusion.
for this discrepancy.The
There
two
reasons
are
principal
of the carbohydratesin the
first is the extensive fermentation
digestivetract of ruminants, leadingto a relatively
largerloss

excreta, and

no

in the

of energy
is the
more

fact that

the

gases excreted.
urine of herbivora

material (224)than
non-nitrogenous
carnivora.

or

combustible

Compare

Penna.

Landw.

The

The

is the

Nutrition,pp.

son
rea-

carries off much

case

results of direct determinations

Armsby, Principlesof Animal

Expt. Sta., Bui. 71, p. 7.
Vers. Stat., 53 (1904), 440-449.

second

291-293

with
on

and

man

swine
333-335.

648

NUTRITION

Table

190.

OF

Metabolizable

"

FARM

ANIMALS

Energy

Digestible

of

Nutrients

per

Gram

Cattle

Protein

(N

From

6.25):
wheat
gluten
wheat gluten (N

From

beet molasses

From

mixed

From

Cals.

From

mixed

and
From

Cals.

Cals.

4-958
3-984

ration

Swine

4.894

5.7)

X
.

grain

Horse

4.083

of

oats, hay

3.228

straw

meadow

From

hay
timothy hay

From

straw

1.272

3-"57(?)
(?)

Fat:
From

peanut oil

8.821
...

From

hay (etherextract)
Carbohydrates :
Starch, Kellner's experiments
Starch, Kiihn's experiments
Nitrogen-freeextract (assumed)
of straw
Crude fiber,
pulp
of hay fed alone
Crude fiber,
Crude fiber,of hay added to basal

8.322
3-763
3.648

4.185

3.606

3-3H

ration
Crude

3.606

Crude

fiber,of oat straw

fiber,of wheat straw
fiber,of mixed ration

753.

Computation

Crude

3.001

3-523

metabolizable

from

energy

digestible

simpleand direct method of computation

the
total
be
based
on
ible
digesthowever,
employed,
may,
of
the
ration.
As already pointed out,
organic matter

organic

matter.

"

of

3-437

the

differences shown

the

gross

energy

due

are

chemical
and

energy

methane

more

in Table

of different

chieflyto

188 between

feedingstuffs

the
which

differences in the

carried off in the

feces,while

percentages of
are

lizable
metabo-

proportionof

the

the losses in urine

lizable
Accordingly,the metaboorganicmatter necessarily
energy
per unit of digestible
exhibits much
smaller
variations than
that per unit of dry
a
matter, and in fact shows
strikingdegree of uniformity.
Selectingthose averages which appear most trustworthy,the
results may

are

far

more

be summarized

uniform.

as

follows

"

Table

VALUES

PRODUCTION

THE

191.

Energy

Metabolizable

"

OF

FEEDING

per

Kilogram

649

STUFFS

Digestible

ganic
Or-

Matter

Mean

Therms

Roughage
Cattle

3-53

Sheep

3-56

Horse

3-7i
Concentrates

Cattle

4.04

Sheep

3.85

Horse

4.62

Swine

4.40

mixed

rations

Table

192.

when

degree of uniformityappears

similar

compared,as

are

"

Organic

Matter

in

on
:

"

Digestible

Rations

Mixed

Number
Single
Trials

of

Kilogram

per

results
shows

following
summary

the
Energy

Metabolizable

the

Minimum

Maximum

Therms

Therms

Mean

Therms

Cattle

Kellner and

Kdhler

Armsby and
Voltz, et al
Tangl, et al

Fries

All

3-72

3-48

3.60

26

3-89

3-5i

3-73

4.12

3-76

3-98

_8

4-H

3-"4

3^5

76

4.12

3-48

3-67

16

4-30

3-5"

3-79

19

4-Q5

3-i5

3-79

35

4-30

3-i5

3-79

4-47

3-24

3-99

4-3i

4-19

4-25

4-43

3-48

4.06

experiments

38

Sheep
Tangl, et al
Voltz, et al
All

experiments

....

Horse

Lehmann,

Zuntz

and

Hage

mann

Tangl,
All

et al

experiments

14

....

Excluding feeds containing

much

oil.

ANIMALS

FARM

OF

NUTRITION

650

the latter,
Feeding stuffs,rich in proteinand fat,especially
naturallygive higher values,as is illustrated by the following

Table

193.

from

taken

likewise
results,

Table
Energy

Metabolizable

"

188.

Organic

Kilogram

per

Matter

meal

Palmnut

4.85 Therms
3.98 Therms

Cattle

Sheep
Wheat

Digestible

gluten
4.79 Therms

Cattle
Flesh meal

5.63 Therms

Swine

of

can

unit for

the

of

venience,
practicalconit is believed
that for the approximate computation
of ordinary feeding stuffs or rathe metabolizable
tions
energy
is known
of digestibleorganic matter
whose
content
or
be estimated,the following
factors may be used,at least for

the

Taking

pound

ruminants, without
Table

194.

as

serious

error

"

Energy

Metabolizable

"

reasons

Pound

per

Digestible

ganic
Or-

Matter

Horses

Therms

Roughage
Mixed

1.683

rations

roughage and

"

1.810

concentrates

Concentrates
Grains

similar material

and

less than

With
With

more

Oil meals

digestiblefat
fat
5 per cent digestible
materials high in protein
5 per cent

and

Losses

It
energy

of

losses of
losses
energy
second

in

stated

was

of
a

2.096

is

heat

production

subject to
into

discussed

class of losses has

two

classes of

the

gross

losses,
viz.,

in the excreta
and
energy
heat.
The
losses of chemical

chemical

conversion

been

previousparagraph (741) that

feeding stuff

through

in

energy

untransformed

have

than

now

in the
to

precedingparagraphs. The

be considered.

754.

feed

of

Influence
from

is evident

"

of

has become

OF

to increase

of the effect varies within

of the

and

of

wide

rather

nitude
mag-

limits

according
and physicalproperties
less difference of

or

his associates have

and

As

"

physiology. The

is still more

Zuntz

causes.

metabolism.

on

the chemical

feed, while there

to its

opinionas
"

animal

speciesof

the

to

651

STUFFS

(365),the fact that the

the heat production of an

Chapter VIII

commonplace

FEEDING

consumption

feed tends

consumption of
animal

VALUES

PRODUCTION

THE

called

"

creased
digestion and have attributed it largelyto inand
and glandularactivityof the digestive
muscular
in this field have
Numerous
investigations
excretory organs.

it

of

work

made

been

carnivora

on

usuallyvery largeexcept when much protein

these
recent
The more
experimentson
species
work
shown
that the mechanical
of the digestive

is consumed.

have

to

organs

is but
"

the increase of

in which

man,

on

is not

the metabolism

appear

or

small

is not

factor and

fortunate

that

With

one.

the

"

term

herbivora

of digestion

work

and

especially

hand, the total effect on the heat

more
marked, and the
production is quantitativelymuch
with

ruminants,

mechanical

the other

factor is of greater

Results

755.

on

(364,449) the

on

cattle.

As

"

significance.
illustrated in

earlier chapter

an

effect of feed

consumption upon the metabolism

of ruminants
be determined
by comparing two periodsin
may
of the same
which different amounts
feedingstuff or ration are
of heat production on
the heavier
consumed, the increment
ration being compared with the additional amount
of feed consumed.
The experimentsthus far reportedhave been almost
cattle,the principalones
exclusively
being the pioneer
upon
l
of
of Kellner and
and the later ones
Kohler
investigations
Armsby and Fries.2
Most

of Kellner

maintenance
but

rations.

computed
indicated

manner

few

Only

from

data

from
in

and

Kohler's

The

heat

the

experiments were
productionwas not

balance

Chapter

of their results

regarding a few
Kellner's book, the increments

VI

have
of

of

carbon

and

made
measured

on

super-

directly,

nitrogen

in

(329),i.e.,by indirect calorimetry.

as
yet been published in full,but
the

other

of heat

experiments

production may

contained
be

in

computed.

Landw.
Vers. Stat., 47 (1896), 275; 50 (1898), 245; 53 (1900), 1-474.
Ernahrung der landwirtschaftlichen
Nutztiere, 6th Ed., Berlin, 191 2.
2
Jour. Agr. Research, 3 (1915),435.
1

the

Die

652

NUTRITION

supermaintenance rations
calorimeter.
respiration

from

derived

Table

the two

195.

and

summarized

elsewhere

As

FARM

ANIMALS

Fries' experiments included

and

Armsby

OF

series

the heat

measured

was

of

Heat

Production

and

directlywith

by the writer,the average

of experimentsare as follows

Increment

"

submaintenance

both

by

results
:

"

Cattle

Roughage
Timothy

hay
Red clover hay
Red
clover hay
Mixed
hay
Alfalfa hay
"Grass
hay"
Meadow
hay
Rowen

Maize

stover

Barley
Oat

straw

straw

Wheat

straw

Straw

pulp

Maize

meal

Concentrates

feed

Hominy

bran

Wheat

Grain

mixture

No.

Grain

mixture

No.

:
.

meal

Cottonseed

....

meal

Linseed

meal

Palmnut

meal

Peanut

molasses

Beet

Starch
oil

Peanut
Wheat

In

Kellner
2

gluten

this and
and

Wheat

following tables,A"F

signifies
Armsby

and

Fries

and

K"K

Kohler.

bran, 14.28

meal, 42.86 per

3 Corn
meal, 60

per

cent;

corn

meal, 42.86

per

cent;

old

process

linseed

cent.

10

per

cent.

per

cent

crushed

oats, 30

per

cent

; old

process

linseed

meal,

Results

756.

In

"

FEEDING

respiration
experiments
Wattenberg, reported by

and

of

varying amounts
Henneberg and Pfeiffer,1
basal

of

ration
from

of flesh meal

conglutinor

of

in the form

barley meal.

hay and

the recorded

shows

the
Table

amount

196.

lost

Increment

of

as

writer2

has

basal

ration

difference between

tein
proto

puted
com-

metab-

and
the

the
two

heat.

Heat

Production

by

Sheep

notably lower than those obtained by Kellner

for wheat
gluten fed to cattle,although in the three middle
periodsthey are higher than those found with that speciesfor
other concentrates, but there are several points of uncertainty
in the experimentalresults and the method
of computation is
On the whole, pending further investigation,
an
approximate one.
it appears
probablethat the results obtained with cattle
without very serious error, be regarded as applicableto
may,
other speciesof ruminants.
757.
Results
swine.
The data regardingthe increment
on
of heat productionconsequent
the consumption of feed by
on
The

results

"

of energy

added

experimentsthe

the

to

nearly pure
were

The

results of these

of the additions
olizable energy
of the resultinggain. The
energy

653

STUFFS

series of

Kern

sheep by

two

upon

sheep.

on

OF

VALUES

PRODUCTION

THE

are

"

swine, although

abundant

than

those for

sheep, are still

rather
Respiration experiments made
meager.
by Meissl,
and Lorenz3 in a study of the sources
Strohmer
of animal fat,
1

more

Jour. Landw., 38 (1890), 215.

Biol.,22 (1886), 63.

3Ztschr.

Principlesof Animal

Nutrition, pp. 463-465.

654

NUTRITION

and

cockle

be made

may

due

hardt3

feed

to

der

of Von

and

Kornauth

by

Heide

and

toward

of

Neither

of

Arche

FARM

ANIMALS

the

upon

nutritive

value

of

the basis of estimates

of the energy expenditure

the later investigations

consumption, while
and Klein,2of Fingerling,
Kohler
and Reindirected more
Wellmann,4 were
specifically

study of

OF

the energy

relations.

the two

first mentioned
included any
investigations
the same
but by substantially
method
determinations,
as that
energy
appliedin the previousparagraph to experiments with sheep,assuming
maintenance
an
requirement, the heat increment per
average
be computed.
unit of feed may
Von der Heide and Klein, in Zuntz's laboratory,
have measured with
of
of
the
the aid
a respirationapparatus
Regnault-Reisettype (298)
of three swine on
the metabolism
than
basal ration slightly
a
more
sufficient for maintenance
of barleymeal, dried potatoes
and consisting
and

dried yeast, and

also the effect of the addition

of dried potatoes and

was

two

to

this basal

ration

palm oil. The energy of the feed and excreta

mals
determined.
Estimating the fastingkatabolism of the three anibe computed as in the
from the body surface,the results may
crements
previous experiments. A computation from the total heat inabove

the

in live

of

basal ration

without
(i.e.,

correction for the differences

higher results.
Kohler and Reinhardt, in experimentson two growing
Fingerling,
swine about
old, added approximately pure nutrients
eight months
(starch,peanut oil,straw pulp, wheat gluten,flesh meal and sugar)
of ground barleywith a littleflesh meal.
to a basal ration consisting
The animals gained steadilyin weight. By a comparison of the first
and
last periods,on
the
the basal
ration,the authors compute
of body surface to
meter
fasting katabolism per square
average
have been
1044.67 Cals.,which
fairlywell with the average
agrees
(377), viz., 1089 Cals. per square
computed in Chapter VIII
meter.
Taking the average of the first and last periodsas the basal
ration,in order to eliminate the effects of the increase in live weight,
and subtractingit from the results of the intermediate periods,the
in proportion to the surface of
fastingkatabolism being estimated
weight) gives somewhat

the animal, the heat

computed
while

by

the small

in the

manner

increment

due

illustrated

per gram
1

Landw.

Vers.

Landw.

Vers.

dry

matter

in the

Stat.,40 (1892),177.
Stat.,84 (1914),149.

nutrients

in the

in the first and

of flesh meal included in the

of

added

for starch

correctingthe results obtained

amount

the

to

? Biochem.
4

Landw.

be

followingtable,
last periodsfor

ration,the

barleymay

may

energy

diture
expen-

likewise be estimated.
Ztschr., 55 (1913), iQS-

Jahrb.,46 (1914),499.

656

NUTRITION

the

in heat

of the total increment

cent

contrary, caused
of swine

FARM

OF

production.

fullyas great
in that

ANIMALS

an

Straw

pulp,on

increase in the heat production

of cattle.

Fingerlingexplainsthis
that the straw pulp was
rather
fermented
the supposition
upon
than digested. He failed,however, to find any corresponding
excretion of methane
(745),and Von der Heide, Steuber and
x have
small evolution
of comobserved only a relatively
Zuntz
bustible
as

this

from

gases

material

in

of the

case

horse.

The

regardsoil and proteinare not readilyexplicable

since,accordingto Kellner,they are not subjectto the methane
differences

as

fermentation.
Table

198.

Increment

"

of

Heat

Production

by

Swine

Heat

In-

cremen

per

Experimenter

100

LB.

DRY

Matter
Eaten

Therms

Grains

Rice

Barley
Barley
Dried

Meissl,et

al.

al.

Meissl, et

et
Fingerling,

V. d. Heide

potatoes

Flesh meal

al.

and

Klein

et al.
Fingerling,

Mixed

rations

Rice, flesh meal and whey

Cockle, barley and maize
Rape cake, barley and maize
.

Meissl,et al.

41.

Kornauth

and

Arche

Kornauth

and

Arche

Skim

milk

and

flour

Wellmann

24.4

27.9

60.9

Single nutrients
Starch

et
Fingerling,

al.

31-93

Cane

Fingerling,et
Fingerling,et
Fingerling,et
Fingerling,et

al.

47.22

al.

60.56

al.

Si-67

Straw
Wheat
Peanut

Palm

sugar

pulp
gluten
oil

V. d. Heide

oil

Biochem.

Ztschr.,74 (1916),161.

al.

and

3Q-3S

Klein

105.92

758.

VALUES

PRODUCTION

THE

Experiments

the

on

OF

horse.

FEEDING

No

"

657

STUFFS

experimentson

this

reportedin which the energy expendituredue

mined.
deterto the consumption of a singlefeeding stuff has been
the only data available are those derived
Practically
of Zuntz
and Hagemann, the
from the extensive investigations
results of which
regardingthe fastingkatabolism have been
animal

have

considered

been

in

(385). On

Chapter VIII

the basis of their

periments
ex-

expenditureand the net

of the
value from the composition and
digestibility
energy
with that employed
identical in principle
ration by a method
in the experimentson
cattle alreadydescribed.
The
ments
experitute
conducted
so
were
however, as to constidifferently,
and
distinct method
be more
a
they may
practically
considered in connection with the computation of
conveniently
the

they compute

net

values discussed in

energy
759.

Results

(365,366)

carnivora.

Mention

"

in

of the fact that

subsequentparagraphs (775-778)
was

carnivora,as

made
well

in

as

Chapter VIII

in herbivora

and

consumption of feed stimulates the heat production,

called by Rubner
the specific
increase having been
dynamic
the

omnivora,
the

on

energy

action.
of Lusk

It is evident

that

experiments

virtuallydeterminations

were

species. While having

no

direct

like those

of net

of Rubner

and

energy values for these

the question of the tive
nutri-

bearing on
farm animals, these

data have been

feedingstuffs for
sirable
deextensivelyquoted in related physiological
writingsand it seems
include
here.
Rubner's
made
them
later
to
experimentswere
the
above
critical
at about
330 C, or considerably
temperature for
of the
the dog, a fact which
is of importance in the interpretation
results (395-397).
in which
A balance
experiment with a respirationcalorimeter
nearlyenough fat was fed to supply the requirementfor energy gave

values of

Table

199.

Increment

"

of

Heat

Production

by

Dog

on

Fat

Diet

658

NUTRITION

OF

kilogram live weight the results

per

stated in

in substance

which

are

in Table

shown

different

somewhat

form

ANIMALS

FARM

that used

from

identical with

remarkable
figuresappear somewhat
is
with
that the comparison
virtually
body
that the

means

for maintenance

valuable

of feed

energy
as

are

by Rubner

but

his.
in view

These

it

which

199,

fat.

fat is

of

the

fact

preted,
Literallyinter-

only 87

per

cent

as

of mobilized
the energy
body fat plus a
true, it implies a larger expenditure of

protein. If this be
in the digestionof fat or a greater stimulatingeffect of the
energy
cell activitythan now
resorbed fat upon
seems
probable, since the
katabolism
of resorbed feed fat can
hardly differ greatlyfrom that of
body fat. Rubner 's figureis the result of a single experiment and
unfortunatelyit enters into the computation of all the other results.
little

It is
much

heat

lower

in which

an

additional

heat

for fat.

increment

of olive oil

emulsion

interest,therefore,that Lusk

of much

matter

elimination

be

to

In two

calorimetric

given

was

0.92

per

to

and

cent

found

experiments
he

dog

has

found

1.49

the

per cent
of energy

of the energy of the oil,so that on the average

98.8 per cent
of the fat was
available for maintenance, a much
higher figurethan

Rubner's.
Both

Rubner

find the

and Lusk

effect to be

marked

most

an
by protein. In two other experimentsby Rubner
maintain
fed.
the dog was
meat
nearly sufficient to
a

small

the feed per

of

amount

fat, the

average

In

of lean

amount

The

metabolizable

kilogram live weight being as follows

produced
meat

tained
con-

energy

of

"

protein 56.70 Cals.

In fat

4.95

Cals.

61.65 Cals.

Using the data afforded by

due to the proteinmay
Table

200.

"

Increment

of

the
be

Heat

ment
experiment on fat,the heat increcomputed as follows :
"

Production

Jour. Biol. Chem.,

13

by

(1912),38.

Dog

on

Meat

Diet

those

his method

Lusk's

should
"

Table

201.

that

"

of the feed

As

portion of the
after

net

in

Stated

It
following table.
figuresare not applicable
of herbivora.

Energy

there have

been
due

those

results from

gross

Available

considered

the

the increased

to

the

its

of

energy

deductingthese
value, or

energy

Rubner's

erroneous.

the

consumption of feed.
in the introductoryparagraphs of this section,

pointed out

that

although

way,

values

and

excreta

substantially

for Dogs

energy

previousparagraphs

in the
losses of energy
heat productionwhich

as

659

STUFFS

same

Metabolizable

of

Net
the

in

these

Results

Average

In

the

computation

Percentage

"

in

contained

nutrients
digestible

FEEDING

computed

of
are

averages

OF

report results agreeing

clearlyunderstood

be

to the

when

of Rubner

they regard
and

Lusk

and

Williams, Riche
with

VALUES

PRODUCTION

THE

feedingstuff

mains
re-

classes of losses constitutes its

two

production value

different
slightly

which

the net

way,

as

energy

regards

energy.

value is

equal

the metabolizable

minus
the increment
of heat proenergy
duction.
It differs from
the relative value, based
the
gestible
dion

to

nutrients

or

the

metabolizable

of all the losses of energy

760.

Net

values

energy

foregoingdiscussions
and
than

net

energy

for any

other

for cattle.

that

values

to which

are

the

data

much

speciesof

farm

in

count
taking acthe feed is subject.

energy,

It is apparent

"

from

regarding losses of
more

abundant

animals.

the

energy

for cattle

Combining

the

66o

OF

NUTRITION

ANIMALS

FARM

195 (755)regardingthe losses due

production with those regarding the losses

of Table

data
heat

in the

energy

(749)gives the
Table

202.

results

Net

"

contained
Values

Energy
Per

in

recorded

excreta

Hundred

of

Pounds

another
in

the

form

of

increased
chemical

in Table

188

followingtables,1the
Stuffs

Feeding

of Dry

to

for

Ruminants

Matter

Net
Energy
Values

Therms

Roughage
Timothy hay
Red
clover hay
clover hay
Red
Mixed
hay
Alfalfa hay
"Grass
hay"
Meadow
hay

48.63

Rowen

33-88

46.64

36.79
43-37

34-io

Maize

stover

Barley
Oat

36.43
41.69
40.23

33-89
26.03

straw

straw

Wheat

straw

Straw

pulp

11.07

89-54

Concentrates
Maize

meal
.

92.58

feed

Hominy

97.84

bran

Wheat

60.51

Grain

mixture

No.

78.79

Grain

mixture

No.

83.82
84.79
82.93

meal

Cottonseed

Linseed

meal

Palmnut
Peanut

meal

78.89

meal

81.56

Beet

molasses

Starch
Peanut

Wheat

82.10

81.89

oil

161. 71

gluten

Penna.

Expt. Sta., Bui.

68.45

142.

Estimated

from

digestible
organic matter.

THE

VALUES

PRODUCTION

OF

FEEDING

first showing the losses of energy and the

and the second
ioo
pounds of dry matter
of
various

the

the

gross

losses and

of

energy

the net

(745,749, 756), it

the

energy

appears

energy values per

the percentage distributio

net

feeding stuffs between

values.

As

probable that

be used also for other classes of ruminants

may

66l

STUFFS

dicated
alreadyin-

these values

without

serious

error.

Table

203.

Distribution

"

of

Energy

of

Feed

in

Ruminants

Net
Energy
Value

Roughage

Timothy hay

24

....

Clover

hay
Mixed
hay
Alfalfa hay
Meadow
hay
Maize
Oat

23
21

17

stover

....

21

....

20

straw

Wheat

!3

straw

Extracted

....

straw

48

"
.

Concentrates
Maize

meal

Hominy
Wheat

46

feed

46

....

bran

Grain

29

mixture

No.

mixture

Grain

No.

37

40

Beet

molasses

49

....

Starch
Peanut
Wheat

761.

form

43
oil

38

gluten

Net

27

....

energy

values

for

swine.

"

the results of the determinations

Combining
of the heat

in the

same

increment

consumption of feed by swine (757) which are

recorded
in Table
198 and such data regardingthe losses of
contained in Table 188
as are
energy in feces,urine and methane
(749)yieldsthe net energy values shown in Table 204 :
caused

by

the

"

662

NUTRITION

Table

204.

Net

"

OF

Values

Energy
Per

Stuffs

Feeding

of

Pounds

.100

ANIMALS

FARM

for

Swine

Matter

Dry

Net
Energy
Values

Experimenters

Therms

Grains

Barley

Meissl, et al.
Meissl, et al.
Fingerling,et

Dried

V. d. Heide

Rice

Barley
potatoes

158.7
121.9
al,

120.5

and

Klein

Flesh meal

....

Rice, flesh meal

whey
Cockle, barley

Meissl,et
and

al.

149.6

and

Kornauth

Arche

milk

flour

and

138.3

Kornauth

cake, barley
maize

Skim

190.2

and

maize

and

al,

Rations

Mixed

Rape,

1OI.3

et
Fingerling,

and

Arche

139.2

Wellmann

I35-0

SingleNutrients
Starch
Cane

sugar

Straw

pulp

Wheat

gluten

oil

Peanut
Palm

oil

Fingerling,et

al.

151.

et
Fingerling,

al.

119.0

Fingerling,et
Fingerling,et
Fingerling,et

al.

92.O

V. d. Heide

al.

152.3

al.

386.7

and

Klein

762.

results

of

of

Comparison

3I4.7

roughage and

in

recorded

concentrates.

The

"

age
aver-

crease
foregoingtables for the total inin metabolism
resultingfrom the consumption of a unit
work
of digestion
i.e.,for the so-called
dry matter
the

"

"

"

in the widest

we

sense

are

"

scarcelyin

accord

Unconsciously misled

by

have

to think

been
1

accustomed

Omitting

one

very

an

with

common

unfortunate
of the

restless animal.

more

ceptions.
con-

nology,
termicoarse

664

NUTRITION

in

effect of maize

marked

and

Hagemann's

effect of maize
also made

This

result

is of

observations

metabolism

of the

on

activitywas

stimulating

horse, which

were

increase in the

no

Grain

reported.

of maize

interest in view
the

on

the
upon
the standinganimal, although

muscular

minor

ANIMALS

the metabolism,
noticeably
increasing

during standing.
especially
of Zuntz

FARM

OF

mixture

No.

i,

likewise showed

meal,
a similar
containing43 per cent
No.
although with grain mixture
effect,
2, containing60 per
less marked, possiblyon
much
of maize, it was
cent
account
lower

of the
per

protein(12.5as

with

compared

17.5

cent).
Influence

764.

of

the

of feed

of amount

the

that
and

of

content

consumed.

foregoingparagraphs it

both

the

increment

losses of chemical

has

In

"

been

the

cussions
dis-

sumed
tacitlyas-

in the excreta

energy

of heat

production consequent upon feed

consumption are proportionalto the quantityof feed ingested,
that the net energy values per unit of feed are substantially
i.e.,
unaffected by the amount
consumed
or
trition
by the plane of nuof the animal.

This
rations
the
one.

Part

not

seems
are

to accord

less
relatively

fat animal

with

the

effective than

utilizes its feed

less

It became

III, of

generalbelief

that

lighterones

than
efficiently

clear,however, in the
the feed requirements for

course

of the

various

forms

heavy

and

that

the

thin

study,in
of production,

varietyof factors are influential in determining

the actual outcome
of feedingoperationsand that diminishing
from
returns
sarily
heavy or long continued feeding do not necesof
of the feed as a source
imply a diminishingefficiency
On the other hand, however, surprisingly
body material or energy.
voted
little specific
to have been deinvestigation
appears
that

to

this fundamental

question.

ence
might influthe net energy
value of a feedingstuff either by affecting
the extent
chemical
is lost in the excreta
to which
(i.e.,
energy
the metabolizable
the magnitude of the
energy)or by affecting
losses due to increased heat production.

Obviously,differences

in the

amount

consumed

That
in mixed
rations the
Influenceon metabolizable energy.
suffer more
less on heavy feeding has already
or
digestibility
may
been shown
in Chapter XVI
(722),notably in Eckles' and Armsby and
"

experimentsin which
rations varying in amount

should be noted

that

maintenance

moderate

or

the other

On

of 8 to

decreases

hand, however,

it

was

found

the

within

the

unmistakable
in the

body

in mind

by

the

that

unit

weight

in the

digestivetract
exchange
energy

of feed.

amount

fermentation

extensive

more

as

in the latter

experiments
distinctly
greater on

were

in the amount

of feed

sumed,
con-

experiments,failed to show any

actuallyliberated
quantity of energy

the

considerable

used.

were

of these

limits

effect upon
from

10

production rations

losses of energy
in the methane
the lighterrations so that the differences
that

665

STUFFS

observed
per cent were
by from 42 to 186 per cent, although it
in Armsby and Fries' later experimentsonly sub-

Fries'
on

FEEDING

OF

VALUES

PRODUCTION

THE

heat

Moreover,

of the additional
of the

of fermentation

does

borne

secured

energy

lighterration
and

be

it must

is liberated

not

enter

into

body tissues,so that the difference in the

net nutritive effect is likelyto be less than that in the metabolizable
far such a compensation would
How
as
ordinarilydefined.
energy
the

occur

in

more

liberal

of the

feeding is difficult at present to

production. It is believed by

say.

however,
Influenceon heat
some,
the
metabolizable
that, aside from differences in digestibility,
etc.,
derived
from
the
feed
is
utilized on
less efficiently
actually
energy
than
rations
and
fat
than
thin
on
animals, i.e.,
light
heavy
by
by
less
that a unit of metabolizable
supplied yields
product.
energy
creased
far
the inThis does not
So
as
exactlyprobable,a priori.
appear
heat production is due to mechanical
work
of digestion,
it
that it would
be substantiallyproportionalto the
would
appear
of dry matter
amount
consumed, except possiblyon extremelyheavy
rations.
So far as it is due to a stimulation
of the body metabolism
by the digestive products resorbed
(367 e) it would appear more
in
with
accordance
the general laws of mass
action, it
likelythat,
be
function
of
the
would
a
diminishing
quantity present. Certain
Grafe and Miiller,
authors, especially
have, it is true, reportedexperiments
"

which
on

to

appear
It has

are

heavy

claimed

to demonstrate

rations

of

the writer

so-called "luxus

carbohydrates,but

tion"
consump-

their results

scarcely

entirelyconclusive.

heat
in Chapter X (450) that any
already been shown
production arisingfrom a synthesisof body substance, such as that
of fat from carbohydrates,for example, and which might be supposed
of the feed energy
to result in a decreased
on
supermainefficiency
tenance
as
compared with submaintenance
rations,is apparently not
of
sufficient in amount
values
affect
the net energy
to materially
as
1
the writer has elsewhere
discussed
feedingstuffs. As regards cattle,
the results of Kiihn's and Kellner's
respirationexperiments in their
bearing on this question, reaching the conclusion that their general
1

Principlesof Animal

Nutrition,pp. 466-471.

666

OF

NUTRITION

ANIMALS

FARM

hypothesisthat the proportion

utilized is substantially
of the metabolizable
independent of
energy
not
the quantity of feed, provided that the changes in the latter are
in
of the fermentations
so
great as to materiallymodify the course
the digestivetract.
species1
Armsby and Fries' results on the same
tendency

tend

confirm

to

of

be in favor

to

seems

in the

of the latter

amount

the whole

On
the

most

set

by

the

seem
probabilities

of feed

capacityof

feed

rather

over

wide

Influence

765.

of

range

as

the

be

that the limit to

in herbivora

at

least,is

similate
alimentarycanal to digestand asby the capacityof the organism to

it

to

the

by

organs

of resorb-

values may be
strictly
constant, at least nearly so

to be the case,

regarded as being, if

to

energy,

transmitted

If this proves

tion.

the

than

utilize the material

of feed

production per unit

heat

distinct evidence

no

increased.

was

efficient use
the

afforded

conclusions,since they

these

increase

an

the

of

not

the net energy

feeding.

of age,

and

breed

individuality.
"

That

ferences
dif-

"

"

individualitymay exist between

with which
animals as regards the efficiency
they utilize the
of their feed and consequentlyas regardsthe net energy
energy
values of the latter does not
probable a
particularly
appear
priori. Such data on these pointsas are available have been
referred to in ChaptersXI and XII, the generalconclusion being
due

to age,

breed

reached

that the evidence

of any

marked

growth

of very

766.

Influence

or

is insufficient to establish the existence

differences of this sort

young

except, perhaps,in the

animals.

of kind

of

production.

It will not

"

have

caped
es-

foregoingdata regardingnet energy values

whether
relate entirely
to the productionof body tissue,
directly
in work
in maintenance
in growth or fattening
or
or
indirectly
production. While it is perhaps unlikelythat the values for
the discussions in
these various
are
identical,
stricly
purposes
notice

that the

Chapters VIII, X, XI,

XIV

and

in the

present chapter

seem

probable that the differences are not of sufficient

of these net enwith the use
ergy
magnitude to interfere seriously
values for the computation of rations in practice.
ever,
of animal husbandry, howAs regardsan important branch
in Chapter XIII,
shown
viz.,milk production,as was
scarcelyany accurate data regardingnet energy values are yet
to

render

it

Jour. Agr. Research,

(1915),472-476

and

Fig.

2.

THE

PRODUCTION

VALUES

OF

FEEDING

667

STUFFS

available,although it appears
probable that they are higher
than the corresponding values for tissue production. A tentative
method

but

of

the present net

utilizing
the requirements of dairycows
was
definite experimental
data are much

" 3.
767.

Computation

The

of

Net

It is apparent

energy
there
to

puting
com-

proposed (605),

be desired.

Energy
from

in

values

Values

the

foregoingparagraphs
that the number
of actual experimentaldeterminations
of net energy
values as yet recorded is comparativelysmall
and that it can
hardly be increased very rapidly,while it is
method
to apply the laborious
of respiration
obviouslyimpracticable
and calorimeter experimentsto all the great number
of feedingstuffs now
in use.
Determinations
of the metabolizable energy, in which at least the energy of the feedingstuffs
and

Importance.

of the visible

energy

has been

excreta

while there

numerous,

of

"

are

determined, are

record the results of

on

digestion experiments in which

made.

were

statistical data

It is

rather

more

great

ber
num-

determinations

no

highly important that

the

of
of

mass

thus

in tables of
accumulated, and summarized
the compositionand digestibility
of feedingstuffs,
should not

be

incontinentlythrown
point of view has revealed

contrary, it should
connection

with

overboard
more

be utilized

the

as

simply because a newer

its deficiencies. On the
clearly
to the fullest extent
in
possible,

yet rather meager

results of the

experimentalmethods, for computing

values of such feedingstuffs as have not yet
to direct investigation.
recent

the

net

been

more

energy

subjected

nutrients
Computationfrom digestible
768.

attempt
feed

as

Kellner's
to

make

the

source

metabolism
amount

of

investigations.
"

To

Kellner

is due

the

first

of the conceptionof
practical
application
of energy
to the body. In 1880, in his

the

vestigat
in-

the relations between

muscular
activityand
upon
in the horse (637) he determined
the additional

work, which

result of the addition

expressedhis

to

the

animal

was

able to

his rations of starch

results in terms

and

perform
of fat.

as

He

of the percentage of the energy

668

of the starch

called attention

of combustion

years

later,after

able

769.

of

point
Kellner

of

values

Energy

work

nutrients,and Zuntz and his

begun their investigations
upon

from

cattle

with
investigations

Sixteen

of

sociates
as-

the

Kellner
energy,
the extensive
undertake

standpointof

the

subject and

the

to

return

to

and

of the

determinations

feedingstuffs.
publishedhis fundamental

had

of the horse

metabolism
was

Rubner

useful work

as

and

of nutrients

replacement values
(385,651-656) had

the

recovered

was

of
desirability

the

to

heats

on

which

fat

or

ANIMALS

FARM

OF

NUTRITION

frequentlycited

previouspages.
Taking as his
of feeding stuffs,

on

digestiblenutrients.

"

nutrients
departure the digestible
sought first to determine the net energy

of the

values

protein,carbohydratesand fats for cattle by adding

digestible
in as pure form as possible
to a basal ration in
these substances
the manner
already described (449). The results with cattle
obtained in this way on starch,straw pulp,sugar, wheat gluten
and oil are included in Table 202 (760) In that table,however,
these materials are regarded in the lightof feedingstuffs and
.

losses and

the energy
as

Kellner
as

which

include

and

whole

computed
whole

but

determinations

resorbed
Table

from

205.

"

values
energy
all its effects.

net

the net

of

Energy

his

ever,
howpurpose,
stances
values,not of these subFor

protein,carbohydratesand
showed
to have
digestibility
the followingresults.1

fat

of the

them, with
Net

energy

relate to the substance

Values

of

been

Nutrients

Digestible

for

Cattle

Per
Per
of

100

Dry

Pounds
Matter

of

Cent
Metab-

olizable

Energy

Therms

Protein
Starch and
Cane

sugar

Ether
of

crude fiber

IOI.6

45-8

107. 1
8l.2

64.1

204.I

S4-i

227.3

56.8

258.6

64.6

51.6

extract

roughage

of grains and their by-products

of feeding stuffs containing over

....

5 per

fat

xLandw.
95-159-

Vers.

Stat., 53

(1900),1-474;

cent

Ernahrung landw.

Nutztiere,6th Ed.,

PRODUCTION

THE

VALUES

obtained

those

dogs, and

on

by

nutritive

analogous

are

(759) in

Lusk

differences between

the

differences in the

the

and

by Rubner

669

STUFFS

in the last column

Obviously the percentages

to

FEEDING

OF

the

two

ments
experi-

sets

phasize
em-

of the

processes

two

species.
Kellner

then

proceededto
of these factors to the ordinary feeding
test the applicability
stuffs of cattle. With a certain number, notably the oil meals,1
of his factors
the net energy
values as computed by the use
of protein,carbohydrates and
from the amounts
fat actually
digestedshowed a close agreement with those found in direct
experiments with the respiration
apparatus. The digestible
for crude

Correction

770.

nutrients of these materials

pure

fiber.

"

of full value

were

as

compared with

starch,gluten or oil.

Table

206.

Net

"

Energy
Per

100

Values
Pounds

of

Dry

Oil

Meals

for

Cattle

Matter

Difference

meal

Cottonseed
Peanut

meal

Palmnut
Linseed

On

meal

meal

0.2

2.0

were

ranging from

30

2.1

strikingcontrast with the oil meals is

roughages,whose net energy values as directly
much
lower than those computed, the deficit

hand,

by the

determined

with the

"

the other

afforded

1.8

per

cent

to

80

per

cent

and

being greatest

materials.2
Kellner found
digestible
this deficit to be more
nearly proportionalto the crude fiber
than to any other ingredientof the feeding stuffs,ranging
from 46.3 Therms
to 76.7 Therms
per 100 pounds of total fiber.
from the computed
the average
of 61.7 Therms
By subtracting
which
obtained
net
agreed well
values, results were
energy
coarsest

and

with

those

hays

but stillshowed

as

follows

secured

least

experimentsin the case

for the
considerable discrepancies

in

direct

"

Loc. cit.,p. 160.

Ibid., p. 162.

of the
straws,

670

NUTRITION

Table

207.

Net

"

OF

Values

Energy
Per

Pounds

100

ANIMALS

FARM

of

Dry

Roughages

Cattle

for

Matter

Differ-

ence

Per

Wheat

straw

Sample
Sample
Oat

Cent

straw

Barley

straw

Meadow

38.0

24.1

19.5

hay

Sample
Sample
Clover

Rowen

i-3

2.5

hay
hay"

"Grass

4.1
7-o

6.0

finer materials

For

chaff,presumably requiringa less

mastication,31.8 Therms
pounds of total
per 100

expenditurefor
fiber

crude
cent

43-4

or

deducted.

was

of crude

more

scale

was

For

fiber the

green
same

foragecontaining16
deduction

was

for that

dry forageand
fiber,the

like

same

for

as

containing 4 per cent

these
chaff,while between

This

used.

correction

for crude

or

made

for

less of crude

limits

fiber

as

per

was

sliding
applied

only to roughage.
Kellner
to the

ascribed

mechanical

this apparent effect of crude fiber

and
work requiredfor its mastication

largely
portation
trans-

through the alimentarycanal,but in part also to the

fermentations
it is subject; in other words, he ascribed
to which
of digestion."In reality,
work
it to the so-called
however,
the crude fiber can
be regardedonly as a convenient
empirical
"

measure

of the

oil meals

and

ner's
energy

own

differences between

roughage.

less with

far from

roughages
work

factor in it.

than

such

as

the

alreadybeen shown from Kellfrom

others
(762)that the loss of
beinggreater, is on the whole rather

It has

experimentsand

in this way,

concentrates

with

concentrates

and

that

the

chanical
me-

digestive
organs is probably a rather small
less net energy value,not
Roughages have relatively
of the

672
it.

from

bran

of wheat
*

Allen

by

Table

example,the

For

209.

"

net

values of alfalfa

energy

hay and
given
digestibility

compositionand
computed as follows :

having

would

ANIMALS

FARM

OF

NUTRITION

be

the

"

Computation

of

Net

According

Values

Energy

per

Pounds

100

Kellner

to

AlfalfaHay
Digestible
protein
carbohydrates
Digestible
e
Digestiblether extract

10.58

10.75 Therms

1.071

39.98.Therms

2.041

1.016

X
X

37.33

1.38

2.82 Therms
53.55

Total crude fiber

Net

energy

0.617

25.00

15.43 Therms

38.12 Therms

value
Value
(Relative

Bran

Wheat

protein
Digestible
Digestiblecarbohydrates
Digestibleether

%)

77

12.01

1.016

=12.

41.23

1.07

2.273

2.87

extract

20

energy

Starch

772.

62.88

value

values.

values, as

energy

obtained.

were

"

In

results

Kellner's
from

is evident

"-52 Therms

to

in

realitynet
which

by

avoid

Therms

48.42 Therms

were

method

the

order, however,

0.77

Therms

44.16 Therms

62.88
Net

Therms

the

use

they
of

required to express the net energy values

large numbers
of
and
also to avoid the introduction
rations in Calories,.
terms, he converted
called

"

starch

values."

them

for

starch

The

into what

use
practical

value

the

of
familiar
un-

he

feedingstuff
starch (assumed to
of

of pure
defined as the amount
briefly
value.
net
perfectlydigested)which has the same
energy
Thus, Kellner's table gives the starch value of maize meal as
or 81.5 pounds per 100
pounds.
81.5kilogramsper 100 kilograms,
has a
One
pound of starch,accordingto Kellner's results (769),
The starch value of 81.5 given
value of 107 1 Cals.
net energy
value
is equivalentto a net energy
for maize meal, therefore,
of 1071 X 81.5
87,286 Cals.,or 87.29 Therms, per 100 pounds
and converselythe starch values of the alfalfa hay and wheat
spectively.
bran of the previousparagraph would be 35.59 and 45.21, re-

may
be

be

1
2

U. S. Dept. Agr., Farmers' Bulletin

In
other words, Kellner's starch

values per

100

lb.

22

(Rev.), iooi,

values

pp. 8-9.

multiplied by

1.071

net

energy

THE

PRODUCTION

VALUES

FEEDING

OF

Kellner's starch values

magnitude

those

as

and

yieldnumbers
alreadyfamiliar in

avoid

unfamiliar

have

It appears

use.

to

They accomplish these

whole conception on which the

it

confusion

of

thought in connection

some

the writer to have

to

to

in

been

an

believes the

use

that

there

are

with

their

unfortunate

cession
con-

advantages

based

on

energy,

is readily
avoided
the Therm

(308).

expressed in Therms
pounds are of
per 100
order of magnitude as the familiar figuresfor digestibl
if 100
the
nutrients,and even
kilograms be made

same

basis of calculation
reasons,

volume

perts,
ex-

values

energy

these

for

even

manifest

franklyand boldlyto a system

while the objection
to the use
of largenumbers
by the employment of a largerunit of energy,
the

erature
lit-

recent

of energy in terms
of
quantities
feeder can
intelligent
readilylearn
his computation of rations,as not a

going over

Net

in

express

units of energy in
few have alreadydone, and
to

nutrients
digestible

that

attempt
He

matter.

tables of

order of

same

instances
striking
is not always easy,

shown

avoid

to

of the

units.

ends,however, by ignoringthe
system is built up, while

673

STUFFS

they

values

energy

expressedin

are

inconveniently
large. For
feedingstuffs in the present
per 100 pounds.

not

are

of

Therms

Computationfrom digestible
organicmatter
773.

from

Independent
the

of chemical

composition.

"

It is

of Kellner's somewhat
foregoingdescription

method

that

it is

based
essentially

on

the

apparent

cated
compli-

digestible

fats of the

older relative values

protein,carbohydrates and
while it involves in its execution certain more
or
(705-710),
less empirical
corrections which are at bottom
simply methods
of applying the average
results on
net
typicalfeedingstuffs
to other materials.
Armsby and Fries * have proposed a
which seeks to attain the same
method
end more
and
directly
tions
simply,relatingthe energy content and the necessary deducto

the total

dry

matter

or

total

matter
digestible

of the

feedingstuff independentlyof its chemical composition.

The energy
of a feedingstuff is just as definite a
content
or
fats,and
quantityas its content of protein,carbohydrates,
1

Jour.Agr. Research,

(1915),486.

674

NUTRITION

ANIMALS

FARM

OF

entirelypossibleto trace the distribution of that energy

in the body quiteindependently
of any knowledge of the chemical
compositionof the material. Not only so, but it is believed
values there are
that in discussingenergy
distinct advantages
and perhaps also as regards accuracy,
in
as
regards simplicity,
garding
cuttingloose as far as possiblefrom the conventional data rechemical compositionand digestion
coefficients and in
with quantities
of energy.
dealingdirectly
it is

This

is

statement

comparisons based
The
for

no

the

problems
any lightthat
relations

energy

is that in whatever

methods

on

them

approach

only
phase
energeticaspects

to

by

of them.

their

that

and

complex

too

one

aspects, to

of

as

are

stigmatize

to

either valueless

be thrown

can

touch

degree

their chemical

from

methods

of nutrition

understood

be

to

means

chemical

to refuse

us

and

on

by

extent

we

can

or

fluous.
super-

difficult

too

any method,
The
point
be

separated
two

possess

pendent
inde-

them.

As

already pointed out,

of a feedingstuff is equal to
the net energy
value
its methe heat production caused
tabolizable energy
minus
by its
consumption. It has been shown (753)that the metabolizable
of a feedingstuff,when
determined
not
directly,
may
energy
be computed approximatelyfrom the total digestible
organic
factor. If from
this
matter
by multiplyingby a proper
result there
be subtracted
the energy
expenditure due to
feed consumption, either as
mated
estias
or
directlydetermined
774.

Method

from

of

that

the net

computation.

"

of

energy

similar

feeds, the

value.

Thus

remainder

in the

same

stuffs

is approximately

feeding
gestibl
pound of dion
previous
two

justused to illustrate Kellner's method each

organicmatter, accordingto the averages
of metabolizable
would
contain i.6o Therms
(753-755),
pages
in the hay and
in the bran; the average
1.77 Therms
energy
losses of energy in heat production per pound of dry matter
would

be for the

Therm, and
follows

as

as

case,

the

hay

0.5303

Therm

computation of

and

for

the net energy

the

bran

0.5339
values would be

*
"

digestible
protein,carbohydrates and fats enter into the calculation simply
of obtaining the total digestible
organic matter
means
when, as is usually the
the computathis is not reported separately. If the latter is the case, then
tion
is,as stated above, independent of the chemical composition.

The

THE

Table

PRODUCTION

210.

VALUES

Computation

"

Armsby

to

Alfalfa

dry matter
Digestible

Total

Protein
.

Carbohydrates
Fats

Wheat

lb.

Bran

88.5 lb.

lb.

37-33

Pounds

100

12.01

lb.

41.23

lb.

1.38 lb.

2.87 lb.

lb.

56.11 lb.

digestibleorganic

Total

matter

Loss

energy

value

energy

production

in heat

The

49.29

....

Metabolizable

Net

per

Fries

and

10.58 lb.

Values

Hay

91.6

675

STUFFS

FEEDING

Energy

Net

of

According

OF

49.49

1.60

78.86 Therms
48.58 Therms

56.11 X 1.77
88.50 X 0.5339

99.31

Therms

47.25

Therms

30.28 Therms

52.06 Therms

computation is

of

method

same

91.60 X 0.5303

of

applicableto

course

data at hand
speciesthan cattle,so far as the meager
permit. The results of such computations, based upon the
of American
feeding
composition and digestibility
average
stuffs,are contained in the tables of the Appendix.
other

Computation of
Zuntz

775.

by

and

Zuntz

(758)is

illustrated for cattle.

energy.

"

for

the horse

The

"

method

ployed
em-

for

computing net energy

similar to that just
substantially

nutrients and
digestible
expendituredue to
energy
Metabolizable

method.

metabolizable

The

the

776.

values

energy

Hagemann's
and
Hagemann

values for the horse

from

net

from

energy

is estimated

it is subtracted

the

consumption

the

of the feed.

the results of five

From

puted
com-

digestion

in
rations of oats, hay and
straw
1888 and
different proportions made
at intervals between
1891, they
of the total digestiblenutrients
the metabolizable
compute
energy
and

metabolism

experiments

on

the digestedfat multipliedby 2.4)to

(including

average

3.96 Cals.

digestibleorganic
corresponding to 3.99 Cals. per gram
writer
in
Table
the
188
matter
as
(749). In the
computed by
nutrients were
not determined
experiments,the digestible
respiration
estimated by combining the results of the same
directlybut were

per

gram,

digestionand metabolism
experiments in various ratios according
consumed.
to the proportion of oats, hay and straw
777.
of heat
Increment
Experiments upon
production.
man,
made
by Magnus-Levy in Zuntz's laboratory,had previouslyshown

five

"

Landw.

Jahrb., 27 (1898);Ergzbd. Ill, 211-236, 276-279, 418.

676

NUTRITION

food

cent

of the metabolizable
assume

energy

effect than

estimated

by

means

that

consumption of more
being added for cutaneous
is ascribed
be

to

mastication

feed

of

Cals. per gram.

is likewise related

of
per gram
of total crude

feed

the

by

brief,Zuntz

"

horse

be

to

estimated

energy,

at

equal to
the

obtain

energy

the net

the

to

of

sumption
con-

cent

9 per

rate

nutrients,plus 2.65 Cals.

digestible
fiber present, and by subtraction of

of

3.96 Cals.

for each

gram
these amounts
value.

energy

computation may be convenientlyillustrated

the data given by Langworthy * for timothy hay. Zuntz
100
factors, recalculated
Hagemann's
pounds for
per
method

of

convenience, become
and

expended
energy
its
crude
fiber content,
to

net

compute

the metabolizable

and

is

in the

In
value.
energy
the heat production due

of

Computation

metabolizable

from

its amount

and

hay

The
total heat increment
0.565 Cals. per gram.
crude fiber,
at 2.65 Cals. per gram.
therefore,is estimated

of

The

ference
respiration.This dif-

fiber of the

at

Hagemann

from

tion
correc-

The

2.086

being estimated

the

intestinal

and

intervals

rations,a small

less diverse

or

crude

the

to

of the

determined

was

apparatus in short periodsat various

after the

and

Hage-

nutrients
digestible

respiratory exchange, which

the

from

of the Zuntz

778.

eaten.

9 per

and

marked
hay produced a much more
augmenting the heat production of the horse

found

grain in

did

as

per gram

by about
Zuntz

of the horse.

addition,it was

computed

of the food

that this result is applicableto the

of the feed
In

ANIMALS

FARM

consumption increased the total metabolism

that

mann

OF

for crude

of the heat
Table

fiber

for

Therms.

1.202

productiondue

211.

"

Computation

On

the

to

Net

or
for

1.796 Therms
energy
this basis the calculation

metabolizable

the

be

would

hay

Value

Energy

as

follows

per

100

"

Pounds

Horse

Digestible nutrients
Protein
Crude

1.25 lb.
lb.

fiber

12.39

Nitrogen-free extract
Fat (1.18 X 2.4)

21.29

37-72

Total

crude

fiber

Metabolizable
Increase
9 per

lb.

2.83 lb.
lb.

29.00 lb.

1796 Therms

energy

37.72

cent

Additional

of metabolizable
for crude

energy

fiber
.

67.75

Therms

0.09

1.202

Therms

29.00

6.10 Therms

34.86 Therms

Total
Net

67.75 Therms

of metabolism

energy

40.96 Therms
value
1

U. S.

26.79 Therms

Dept. Agr.,Office

of

Expt. Stas.,Bui.

125, p. 14.

THE

from

is evident

As

VALUES

PRODUCTION

the methods

believe it to be

which

of considerable

criticism,the

estimate,based

upon

for the

from

assumption

very
during the

considerable

is

relatively
greater
the first to point out
of

with

of

of

feed, as
roughage

that

values

the

his

on

practice.

second,

of 9 per
and more

for 24 hours

be

experimentson

have

Hagemann

increase

digestionof

estimating

two

comparativelyshort

and

the

computation has been the subject

principalpoints being, first their

that the metabolism

the results of

were

strictly
exact, but

nutrients,and
digestible

Qualitatively,Zuntz
the

the factors

of

the results of

effect of the
the

which

by

677

close approximation
sufficiently

Hagemann's method

and

STUFFS

computationsof rations in

to base

Zuntz

FEEDING

of calculation is not

reached, this method

authors

OF

with
be

stuffs.

in

the fact that

than

may

cent
pecially,
esputed
com-

experiments.
respiration
demonstrated
clearly

metabolism

energy
well as

this effect must

feeding

man,

the

horse

this increase

grain,and

they

into

account

taken

There

be

may

were

in

difference

of their figuresand certainly

quantitativeaccuracy
direct methods, involvingfewer assumpinvestigationsby more
tions
and complex calculations,
are
greatlyto be desired,but until
those
continue
such results are obtained,we may
to use provisionally

opinion as

reached
1

to

in the

779.

the

just described.

manner

Wolff's

of

method

the

computation.

horse

made

"

His

extensive

Hohenheim

tions
investiga-

in

working
1877 to 1894
and antedating the investigationsthus far mentioned, led Wolff to
of estimating the relative net
still simpler approximate method
a
upon

at

values of feeds for the horse.

energy
It was
shown, on the average of a considerable
that the

nutrients
digestible

from

roughage

both for work

from

production and for maintenance

grain. Wolff found, however, that if

were

omitted
and

also that

this

less efficient

were

those derived

were

the

the

He
for maintenance.
necessary
the crude fiber in the rations of the horse

valueless and

fiber

crude
digestible

fiber-free nutrients
that

sons,
compari-

the fiber-free
comparisons,the ratio between
and
work
comparatively uniform
performed was
of
assumption yieldeduniform results for the amount

from

nutrients

than
the

of

number

that

the

therefore

remaining

apparently

was

digestiblenutrients

cluded
con-

might

be

garded
re-

equal value whether derived from grainor from roughage.

Expressed in the light of our present conceptions, this is practically
to the
equivalentto saying that the net energy value is proportional
as

amount

Wolff

of

of fiber-free nutrients.
is careful

to

say

that

the

crude
digestible

and virtually
valueless,
regards the
convenient
a
empiricalmeasure

amount

of the

fiber is

of crude

fiber

apparently
as

ing
furnish-

difference in the value

of

678

NUTRITION

OF

FARM

nutrients of roughage
digestible

the

ANIMALS

compared with those of grain.

case
explained in part by the rather
limited variety of feeding stuffs employed in the experiments. The
addition
of
hay with, in some
roughage was meadow
cases, a small
usually oats, partiallyreplaced in some
straw, while the grain was
instances by other feeds.
Whether
the same
relation between
fiberdone would
hold in widely different rations
free nutrients and work
such

That

is

not

apparent.
Wolff's results

only. They do not show the actual

rations but only that it was
proportional

energy in the
fiber-free nutrients.

the

to

relative

are

of net

amount

differ

The

the

considerablyfrom

energy

net

Hagemann's method,
of 9

metabolizable

cent

per

assumes

of the

value of

uniform

method
of

gives the
less than

Wolff's method
exceed

" 4.

crude

and

Zuntz

Production

Differences

per

Values

in

by

Zuntz

and

the deduction

as

values

proteins.
"

if it has

negative value
Values

cent.

Hagemann's

Relative
780.

fiber

the fiber-free nutrients

from

latter would

and
it
second, because
energy,
for crude fiber,
while Zuntz and Hagemann's

zero

55

of the

content

as
computed
energy
it does not include

first because

to

as

is doubtless

is the

gestibil
di-

according

computed
would

therefore

siderably
con-

figures.

Protein

Regards

of proteins
As

appears

from

the

sions
discus-

precedingsection,the productionvalues of feeding

gree
destuffs as regardsenergy may alreadybe formulated with some
is much
needed.
of accuracy, althoughfurther investigation
Concerning the production values as regards protein,the
the protein of
situation
is far less satisfactory.
For
years
a
singlechemical
feedingstuffs has been treated as if it were
known
substance
the different proteins
to exist in feeding
; i.e.,
stuffs have been assumed
to have
equal nutritive
substantially
of the

values.
and

The

more

physiologyof

investigationsinto the chemistry

proteins,however, have resulted in an

recent

the

As has
change in the point of view.
in previous chapters (340,398, 465, 552),it

entire

been
is the

fullyshown
constituent

which
proteinsare splitin digestion
the materials
of which body proteinis constructed,and
out
are
the processes
quire
of maintenance, growth or milk
productionrefor their support, not
proteins as such, but certain
acids as cannot
and proportions
of such of the amino
amounts
amino

acids into which

the

680

OF

NUTRITION

FARM

ANIMALS

tained
vegetableproteinsderived from seeds is known, those conin roughagesand in roots have not yet been investigated,
although a beginninghas been made l in determiningthe proportions
acids which
of the different groups of amino
are
yielded
ing
(crudeprotein)of various feedby the total nitrogenousmatter

of

stuffs.
tion
appeared in previouschapters,such informathe proteinrequirementsof farm
is available respecting
as
animals has been derived from experimentsin which only the
considered
without
total proteinsuppliedwas
reference to its
kind.
no
Practically
knowledge is available as to the amino acid
requirementsof the various farm animals for different purposes.
the productionvalues of the various single
were
Third, even
proteinsknown, it would not be possibleto estimate from them
the productionvalues of the mixed
proteinsof feedingstuffs,
since a deficiencyin one
proteinmight be compensated by a
higherprosurplusin another and the mixture show a much
duction
value than either of its ingredients
separately. Thus,
which lacks lysin,
as
alreadynoted, the value of wheat gliadin,
is practically
for growth, while as part of a mixture
with
zero
other proteinssupplyinglysinit may
have a high value,the
replacementof 25 per cent of it by lactalbumin,for example,
renderingthe mixture fullyadequate to support normal growth.
Each
particularmixture of proteinswould have its own
duction
proof the
value, which might differ widely from the mean

Second,

values

as

has

for the individual

constituents.

differences in proteinsare doubtless of much

qualitative
in progress
and
the researches
can
significance,
hardly fail
rational method
of valuation than
ultimatelyto lead to a more
in use, but
that now
as
yet they do not afford an adequate
basis for expressingthe values of feedingstuffs in generalas
The

of

sources

the purposes
to adhere
necessary

protein.

it stillseems

For

of the stock

fore,
feeder,there-

to the older method

which

digestible
proteinof a feeding stuff as expressing
tually
approximatelyits productionvalue in this respect, thus virrations the protein
assuming that in ordinary mixed
each
balance
will largely
deficiencies of the different ingredients

regards

the

other, and

this method

has

been

followed

Grindley,Joseph and Slater; Jour. Amer. Chem.

2762: Nollau; Jour. Biol. Chem., 21 (1915),611.
1

in the
Soc,

37

tables

of the

(1915), 1778

and

THE

VALUES

PRODUCTION

should

FEEDING

OF

68

STUFFS

be

done, however, with a distinct

consciousness
of the inadequacy of the method
and with the
hope that it may ultimatelybe replacedby one having a more
This

Appendix.

scientific basis.

Meanwhile, notice should be taken of the results of several

the mixed
recent
proteinsof a few feeding
investigations
upon
those of the cereal grains.
stuffs,
particularly
783.

of maize

value

Low

The

proteins.
"

demonstration

of

of the zein of maize for either maintenance

insufficiency
or
growth (781) has tended not unnaturally to produce the
impression that this important feeding stuff is relatively
valueless as a source
of protein. Zein,however, is not the only
proteinof maize.
According to Osborne and Mendel 1 the mixed
the

proteinsof

maize

made

are

follows

approximatelyas

up

Zein

"

%
31%
22%
6 %

41

Maize

glutelin

Globulins, albumins
Insoluble

and

proteoses

in alkali

100%
Glutelin
the

yieldsall the
is probably true

same

amino

acids

of the other

which

zein

proteinsof

lacks

maize.

and
dently
Evi-

the results of
to be valueless

as

experiments on zein do not show maize

of protein,although they do indicate
source

low value and this conclusion

relatively
and Mendel
the experimentsof Osborne
on
pigs. On the other hand, however,

were

able

to

obtain

only maize
Osborne
mixture

of

manufacture

glutelinin

has
on

been
rats

Hart

confirmed
and

and

by

of Waters

McCollum

normal

growth of pigs on rations supplying

proteinbut supplemented by salts.
a

investigatedthe nutritive value of the

proteinscontained in the "corn gluten" produced in the
of starch from maize
and consisting
chieflyof zein and
the proportion of approximately 100
In such a
to 44.
and

Mendel

have

mixture, the deficiencies of the zein are to a greater or less extent

compensated for by the glutelin,and the mixed
proteinsnot only
able to support rather
but were
proved adequate for maintenance
of casein,
of lactalbumin
addition
slow growth. The
or
to them

however, rendered

them

much

more

efficient and

induced

normal

growth.
'Jour. Biol. Chem.,

18

(1914),1.

Ibid., 19 (1914),373-

682

NUTRITION

Waters
manner

In

each

maize.

FARM

OF

ANIMALS

in

experiments on growing pigs has shown in a striking

of Osborne
and
the practicalsignificance
Mendel's
results.
of the four trials reported,one
lot of animals received only
gredient
The others were
given maize with the addition of ash in-

either by direct additions of salts or in the form of the

called protein-free
milk, while still others received an addition

complete proteins,as
from

cases

blood

in

receivingonly maize
same

due

not

was

was

when

true

was

to

ash

lack

of

possible,derived in some
The
others from milk.
growth of the lots
either very slow or practically
and the
zero
was
added, showing that the failure to grow

nearly

and

so-

of

ash-free

mineral

as

When,

matter.

however,

plete
com-

added
normal
to the maize, steady and
proteinswere
growth
and
of
the
showed
a
carcasses
responding
corplace
comparative analyses
of
the
total
animals.
The
sults
restorage
body protein by
live
follows
to
as
:
weights were as

took

"

Table

212.

Influence

"

Nature

of

of

of

Protein

Supply

on

Growth

Pigs

Daily
Gain

Lb.

trial

Second

Maize

alone

Maize

and

ash

Maize

and

blood

O.21

0.19

albumin
.

Maize,

blood

albumin

Third

and

1.00

ash

1. n

trial

Maize

alone

Maize

and

ash

Maize

and

milk
protein-free

Maize

and

milk

protein

Fourth

trial

o.
o.

Maize

alone

Maize

and

ash

Maize

and

milk
protein-free

Maize

and

milk

Maize

and

casein

0.06

1.50

o-37
0.32

albumin

0.51
.

1.05

1-34

Fifth trial
Maize

alone

Maize

and

milk

0.25
ash
.

Maize

and

tryptophan

Maize

and

milk

Maize

and

casein

0.13
0.22

albumin

Proc.

Approximate.

Soc. Prom.

1.

19

1. 01

Agr. Sci. (1014),p.

The

exact

initial

7.

weights are

not

given in

the report cited.

THE

784.

VALUES

PRODUCTION

Values

of other

cereal

FEEDING

OF

683

STUFFS

at the
Investigations

proteins.
"

Experiment Station led to the conclusion that not

only the proteinsof maize but the unbalanced proteinsof other
inferior to milk proteinsas sources
cereal grains are distinctly
of proteinfor growth and milk production.
Wisconsin

Hart, Humphrey
alfalfa proteins for
to

20

much

growing

per

of

cent

McCollum

reports

proteinsof

milk

nitrogen balance

the

in

of

retention

of

mately
approxi-

nitrogen as compared with

milk proteins in later experiments

for

maize,

no

did it exceed

case

in
utilizing

growth,

that

so

wheat

pigs in which the

young
oats, of casein and of skim

and

The

compared.

were

in the different trials but

that

and

maize

the

series of trials on

effects of the

varied

comparisons of maize

institution.

same

on

in two

heifers observed

higher figuresobtained

the

at

24

Morrison

and

the author
the

presents

protein supply
for believing

reasons

the animal

amount

the results

not

are

was

affected

capable
in

the

Chapter XI (468) by surplus protein being

katabolized.
On the higher protein rations,from 20 to 34 per cent
of the resorbed
retained
in the body of the animal,
nitrogen was
while,contrary to what would naturallybe expected, the percentage
retention was
decidedly lower on rations supplying less protein.
The milk proteins,on the other hand, showed
a decidedlyhigher percentage
casein
and
for
skim
milk proretention,
viz.,for
51 per cent
teins
discussed

manner

66 per
Hart and

in

cent.
3

Humphrey

have

compared

the

proteins of maize,wheat,

of protein
gluten feed, oil meal, distillers'grains and milk as sources
for milking cows
(587). Unfortunately,the effects were
chieflyon
the body protein,so that the only comparison possible is between
the algebraicsums
of body protein and milk protein. Computed in
this way,
for three animals
the average
percentage efficiency
was,
for milk proteins,59, for maize 40, for wheat
36, for gluten feed 45,
for oil meal 61 and for distillers'grains60.

785.

their

Alfalfa

proteins.

comparisons

(784),found
the

as
none

of

maize

the

and

alfalfa

and

Morrison

proteins just

in

tioned
men-

did the total

total

nitrogenof

of these Wisconsin

requirement of the animals

1

Hart, Humphrey

nitrogen of alfalfa to show about

percentage retention in both growth and milk production

same

In

"

maize.

experiments is the maintenance

taken

Jour. Biol. Chem., 13 (1912),133.

26 (1916),457Ibid.,21 (1915),239;

into account
2

in

Ibid., 19 (1914),323.

computing

684

NUTRITION

OF

ANIMALS

FARM

of the protein. If this be done, using

percentage efficiency

the

approximate data contained in Chapter

percentages of the proteinssuppliedin excess
the

the
(415-417),

IX

of maintenance

in every case.
increased
distinctly
be concluded, therefore,that the low percentages
It cannot
computed by the Wisconsin investigatorsshow that only these

which

be

would

retained

were

proportionsof the cereal proteinsare capable of

ever,
into body proteins. On the other hand, howtransformation
correction
would
result in making the
such a conjectural
the
different proteins appear
relative differences between
of calculation used.
by the method
greater than those shown
small

rather

the relative values of the mixed

upon
of feedingstuffs have come
to the writer's notice.
other

No

studies

Value

ofnon-protein
conclusion

previousparagraph (782)the
for the present the only available

In

that

of

values

is the

feeding stuffs

which

they

contain.

the

"

of

protein
digestible
tein
pro-

applicationof

proteinor

this method

the basis of

whether

decide

to
necessary
"
crude
shall be the

"

the

reached

was

of the

measure

total amount

In

itjbecomes

teins
pro-

true

"

son
compari-

protein as

termined
de-

(104-107)
; in other
shall be assignedto the

methods

by existingconventional
words, to decide what value, if

any,

-protein.

non

786.

the fact that

"

the

non-proteinmake
of

matter

in

with

Following

investigations.

Early

substances

under

the

share

of

much
feeding stuffs,

numerous

of

their

determine

efforts to
that

considerable

up

grouped

the

true

collective term
the

labor

nutritive

proteins.

recognitionof

the

These

nitrogenous

has

been

value

as

pended
expared
com-

investigations

While
by the writer elsewhere.1
the generaltendency
much
diversityof opinionhas prevailed,
of questionable
value.
the non-protein
has been
to consider
as
in particular,regarded
Kellner,the leadingGerman
authority,
been

have

it

as

valueless.

787.

New

chemistry
1

summarized

viewpoint.
of the

of Animal
Principles

Bui. 139

(1911).

"

With

proteinsand

advancing knowledge
of the

Nutrition, pp. 52-58

chemical

; U. S.

of the

mechanism

Dept. Agr.,Bur. Anim.

of
Indus.,

THE

VALUES

PRODUCTION

protein nutrition,however, it

OF

has

FEEDING

become

685

STUFFS

evident
increasingly

of little real significance

of these earlier results are
that many
and that the questionof the nutritive value of non-proteinmust

approached from a different standpoint. It has become

pletely
evident, for example, that attempts to replace proteinscomby a singleamino acid or even by two or three of them
must
quires
necessarilyfail,since the formation of body protein rebe

the

of all its constituent

presence

proper

proportions. For

amino

acid

to

ration

the

can

the

reason

be effective

ration happen
particular

that

same

only if

addition
the

of

an

proteins of

deficient in that

to be

in

buildingstones

one

stituent.
con-

of the non-protein
Furthermore, experimentswith ingredients
which do not form part of the proteinmolecule are of questionable
significance.For example,asparagin,which has been a
favorite subject of investigationfor reasons
of convenience,is
found among
the cleavageproducts of the proteinsbut benot
longs
the class of acid amides.

to

contribute
the

to the

formation

of

So far

as

proteinonly

it could

after conversion

into

asparticacid (47) and it has not yet been shown

over,
body can undo the amide linkageof nitrogen. Morecludes,
appeared in Chapter I (60-67),the non-protein in-

related

that the
as

in

variety

addition

to

acid

amides

of

like

asparagin, a great
but remotely reare
lated

nitrogenoussubstances which
chemicallyto the proteinsand whose

at

appears,

nutritive

value

is

best doubtful.
It would

stuff

as

the

appear
source

that the value of the

of

body protein must

non-proteinof a feeding
be determined
by precisely

the value of
thing which is believed to measure
individual protein or of the mixed
an
proteinsof feedingstuffs,
viz.,the kinds and proportionsof amino acids which it can
isting
yield,since there is no evident reason
why an amino acid exready formed in a feedingstuff should differ in value from
the same
substance splitoff from proteinin the process of digestion
If this be admitted, however, the distinction made
in
same

proteinand non-proteinin feedingstuffs

years between
becomes rather meaningless. If the value of each is measured
by
recent

its amino

acid content, then what is needed to fix the production

values of feedingstuffs as regards proteinis a knowledge of

the kinds

and

amounts

of these

compounds

which

the

feeding

686

stuff

as

of whether
first

are

the

they

exist in

produced in

Indirect

788.

of

case

the

soluble,as it were
digestivetract of
of

utilization

consideration

the

of the

was

by herbivora.

value

digestive
processes of these
stated in Chapter III (141)that

nitrogenouscompounds
and

which

non-protein,

play

large

so

animals.

the presence
in the feed tends to stimulate

of
activity

In

"

factor

another

of the

organisms

part in the
It

the animal.

non-protein

to the ferment

viz.,its relation
a

furnish,irrespective
predigested,form or

of ruminants,
herbivora, especially

into

enters

ANIMALS

its crude protein)can

(i.e.,

whole

FARM

OF

NUTRITION

of soluble

the multiplicati

these

organisms, thus bringingabout

increase in the excretion
of methane
and in the proportion
an
of carbohydrates
apparentlydigested. It was likewise indicated
of non-proteinin the
that the proteinproduced at the expense
of the microorganisms might serve
of
as
a source
multiplication
proteinto the body and thus bring about an indirect utilization
of the non-protein. Much
experimentalevidence supporting
the extensive
this view is on record,particularly
investigations
which
of Morgen and his associates,
where
have been discussed elsel
general conclusions regarding
by the writer. Three
the behavior
in the body were
of non-protein
drawn, viz. :
of non-proteininto proteinappears
In ruminants, a conversion
effected by the microorganisms of the digestivetract.
to be
"

The

of this conversion

extent

in the

to be relatively
greater
appears
salts and asparaginthan in that of

of ammonium

case

the

non-protein of vegetable extracts.

The proteinthus formed
from non-protein
seems

subsequently. The apparent formation

observed
investigators
by some
appears
in the metabolic
action
specific

of

the

digested
indigestible
protein

of

to be

in the

products contained

to be

due

to

increase

an

feces,'caused by

vegetable extracts

upon

the

the

digestive

tract.

By

of its conversion

means

in the feed of ruminants

and
for

also

growth,

as

in rations

used

in these

deficient

U.

the

experiments were

substitution of the former

1

S. Dept.

may

of

source

however,
Quantitatively,

bacterial

into

serve

protein for
in protein.
various
much

Anim.

protein
non-

tenance
indirectlyfor mainmilk, and probably
of

forms

inferior to

for the latter caused

Agr., Bur.

protein,the

Indus., Bui.

a
139

non-protein
proteinand a

marked
(1911).

falling

688

involved

thus

higher

somewhat
of
since

than

difference

of

simply
in

if
It

deficiency
worst

make

to

computation.

the

at

the

tends

the

FARM

OF

NUTRITION

the

ANIMALS

protein

crude

the

protein

is, therefore,

protein

may
increase

tends

to

latter

respect

an

limit

on

seldom

of

the

the

the

considerable.

basis

safe

while

production
cost

rations

the

made

were

error

the

is

of

content

side,
a

ration,

plus
sur-

and

XVIII

CHAPTER
COMPUTATION

THE

"
Origin.

790.

hay values described in Chapter XVI

of comparing the
methods
place to new

As

"

Standards

Feeding

i.

RATIONS

OF

the

(700)gradually gave
cal
values of feedingstuffs based upon improved methods
of chemianalysisand upon investigationsinto the generallaws of
followed to express the nutritive
an
nutrition,
attempt naturally
instead of in terms
requirements of animals in a similar manner
Thus
of gross weight of feed or of hay values.
originatedthe
for different speciesof farm animals which
feeding standards
later came
to be popularlyregarded more
less in the lightof
or
or
prescriptions
recipesfor

791.

Early

standards.

"

successful

feeding.

earliest

The

suggestion along this

l
originatedwith Haubner
in 1857, amplifiedthe suggestion,but
Lingethal,1

line

to

seems

1858

have

the first

was

formulate

to

about

1840.

Grouven2in

the requirements
specifically
*

farm

of

in terms
of dry matter,
animals, expressingthem
total protein,
total fat (etherextract),and
carbohydrates
in
material
soluble
acids
and alkalies).In other words,
(total
.

"

the crude

Wolff

nutrients

took the

the basis of Grouven's

were

"

standards.

step in advance

by making the digestible

nutrients as determined
of Henneberg and
by the methods
Stohmann
(707-710)the basis for comparisonsof feedingstuffs
and
for expressingfeed requirements. His feedingstandards
first published in 1864 in Mentzel
were
and von
Lengerke's
Landwirtschaf
tlicher Kalender
and were
also incorporated
in his
widely read book, Die landwirtschaftliche Futterungslehre,in
1874.
of

These

next

standards

attempted

to

formulate

protein,carbohydratesand
digestible
1

Quoted by Grouven
sel,1863, p. 327.
2

fats which

should

be

Kritische Darstellung aller Futterungs-Versuche. Kas-

Vortrage iiber Agricultur-Chemie,1858.

2

the amounts

68q

690
in

contained

for

rations

in order to secure
purposes
conditions.
Thus, the
average

various

satisfactory results under

Wolff standard for dairy cows
Standard

Feeding

was

Cows

Milk

for

ANIMALS

FARM

OF

NUTRITION

"

Day

per

and

Pounds

iooo

Live

Weight

Total dry matter

24

Digestibleprotein
Digestiblefat
Digestiblecarbohydrates
.

0.4

that

means

of

mixture

any

pounds
pounds
pound

pounds

12.5

ratio

Nutritive

This

2.4

5-4

suitable

feeding stuffs

digest 2.5 pounds of protein and

pounds of non-nitrogenous nutrients per day will constitute
suitable ration and produce a good flow of milk.
which

from

can

cow

The

Wolff

standards

few

years

later

and

others, and

and

found

with

wide

into

introduced

were

the

United

13
a

States

through the writings of Johnson, Atwater

by the writer's translation of Wolff's book,1
of stock

students

among

currency

feedingand

popular writers.
of the

Modifications

792.

standards.

Wolff

modifications
of investigation
progress
standards formulated
fifty
years ago was

1864

1896 Wolff's

to

Mentzel

and

standards

were

Lengerke'sKalender

von

with

That

"

should

be

the
in

made

expected. From
published annually in
unchanged.
practically
be

to

under
the charge of
continued
1897 to 1906 they were
additions and modifications,
some
Lehmann, who introduced
the principal
ones
being the subdivision of the standard for dairy
From

milk

accordingto

cows

and

milk

or

wool

breeds

793.

in the

the well known

constitute

These

yieldand

Kellner's

standards.

"

the distinction between

standards

for

growing

the

animals.

standards.

Wolff -Lehmann
Both

meat

Wolff

and

the

Wolff-

expressedin terms
standards,as already noted, were
nutrients.
of the so-called digestible
Kellner, in 1905, in the
Nutzfirst edition of his Ernahrung der landwirtschaftlichen
of starch
tiere,proposed the system of calculation by means

Lehmann

values
and

formulated
to

has

(772)which
this

new

since

table of

been

associated

feedingstandards

method.
1

Manual

of Cattle Feeding, 1880.

with

his name,

ing
expressedaccord-

In

differ

respect Kellner's standards

one

and

ones

constitute

691

RATIONS

OF

COMPUTATION

THE

from
radically

ceding
prethe earlier

While

notable advance.

standards, like the earlier tables of feedingstuffs,assume

to

protein,carbohydrates and fats from different sources

of substantiallyequal nutritive value, Kellner's figures

be

take

gestibl
di-

of those

account

in the values

differences

of nutrients

as

have
been revealed by recent investigations
which
energy
and
the needs of animals in this respect in
express

of

sources

fact,although not in form, net energy values. In

addition, his standards
regard only the true protein as of
the very high requirementsof
value and they reduce somewhat
for protein as postulatedby early authors.
fattening animals
the plan
In other respects, however, they are
on
substantially
of the Wolff -Lehmann
standards, i.e.,they are in form prescription
for
rations
for
different
or
recipes
purposes.
what

in

are,

"
794.
was

Limitations
to

necessary

of

as

Requirements

feeding standards.

the outset

From

"

it

guard against misconceptionsarisingfrom the

definite form

very

Feed

2.

which

the

were
feeding standards
sented.
pretended
inTheir authors insisted from the first that they were
adhered
generalguides and not as fixed rules to be rigidly

But

to.

in

the human

mind

craves

recipe and there has

persistenttendency to substitute for the study of the

of nutrition a series of exercises in appliedarithmetic.
principles
the
have interpreted
Others again,perhaps misled by the name,
of
demand
as
feeding standards
representinga physiological
been

the animal

"

sort

at, but concerningwhich

feeding, to be aimed

ideal in

of moral

concessions

have

to human

to be made

and
of circumstances.
the pressure
fallibility
The difficulty
or
inherent,more
less,in all forms of feeding
in the earlier ones, is that they fail
standards,but especially
of the fact that the feedingof farm
to take sufficient account

animals

buy

is

some

an

economic

average
as

the

norm

problem.

for his

raw

material

of his

own

When

high pricesprevailed he might

a

factoryor

maximum

amount

not

garded
might be reof the capacity
business,irrespective

of

amount

would

manufacturer

of the market

of

raw

which

find

material

finished

product.
dle
it profitableto han-

for the

and

so

to

reduce

the

692

NUTRITION

FARM

OF

ANIMALS

at the risk of some

costs, even
percentage of his overhead
in the manufacturingprocess.
of efficiency
In the contrary
he might find it necessary
below his
to run
considerably

capacityin order

to tide

over

bad

loss
case,
imum
max-

In somewhat

season.

for the stock feeder to adapt his

similar fashion it is necessary
rations to the economic
conditions under which he works.
While
the animal

be handled

like

in

machine

nevertheless,
factory,
has appeared in previouschapters,
it shows
as
a large
in its requirements both quantitatively
and
degreeof flexibility
fixed standard
is either physiologically
No single
qualitatively.
or
economicallynecessary for productivefeeding.

795.

cannot

feeder's

The

problem.

As

"

the

feeder

looks

at

his

animals, the problem which they present is a threefold one.

furnish them
with sufficient repairmaterial
First,he must
and energy
he must
to keep the body machinery running,i.e.,
ration.
The
supply a maintenance
requirementsfor this
variations,have
although subject to individual

worked

with

out

problem

is

Second,

some

of accuracy

this part of his

and

relatively
simple.

in addition

his animals

degree

pose,
purbeen

to

maintenance

with the amounts

of matter

for the

the meat,

them

his task

productionof
to yield. Here

ration,he

milk
is much

of energy
work
which

and
or

supply

must

necessary

he desires

less

simple.
has been emphasized in
previouschapters,that the producing capacityof the animal
is the prime factor in the problem. No argument
is necessary
that a cow
to show
quires
producing thirtypounds of milk dailyreIt is evident

in the first place,as

greater addition

to

her

maintenance

ration than

does

having a capacityof only fifteen pounds, or that a steer

which can gain three pounds dailyneeds more
surplusfeed than
one
capableof making only one pound of gain. Good business
that the better animal be givenfeed sufficient
demands
economy
in amount
and
kind to permit its producing capacity to be
fullyutilized,thus reducingthe relative cost of maintenance.
On the other hand, it would
be an
obvious waste
to give a
one

mediocre

producer a ration adequate

times the production of which
it is capable.
Third, the feeder,like the manufacturer, must
to

those

or

poor

market
of animal

conditions.

As

for two

or

adapt his practice

pricesof feedingstuffs

products rise,he

will tend

three

to

feed

fall and

more

in-

OF

COMPUTATION

THE

but here he
tensively,
The
dairy cow
of this.

the

encounters

law

of

affords,perhaps, the
increase in the

An

693

RATIONS

turns.
diminishingre-

lustrati
strikingil-

most

quantityof

her feed above

increase in milk
ration may
be expected to cause
an
time an increasing
secretion but at the same
proportion of the
a

moderate

feed

extra

will be

diverted

fattening(606-610). Similarly,

to

erate
proteinsupply appears adequate to support modulus
milk productionbut largeramounts
to act as a stimseem
of the milk glands and to increase the yield
to the activity
but presumably at a diminishing
of milk (603),
The dairyrate.
man's
problem is to utilize these stimulatingeffects up to the
point at which the increase in yieldis offset by the added cost
the solution
of the ration,and
of this problem requires experience
in which littleaid can
and good judgment and is one
be afforded by feedingstandards.
of dairy feeding appliesin
is so emphaticallytrue
What
greater or less degreeto all forms of animal production. Even
a

rather small

there

though

may

be

decrease

no

in

of

utilization

the

the

feed in the strict

stimulation

diminished
digestibility,
physiological
sense,
incidental bodily activity,
or
position
changing com-

of

of

increase

tend

make

to

rations

heavy

effective than

less
protein rations relatively

or

high

moderate

more

ones.

the feeder needs

What

is not
a

knowledge

various
a

so

much

conditions

of

is available

the
to

to

kind

and

he

capacityof

of

standards,as

requiredunder
of a unit of product
example, or a pound of
"

weight,for

the extent

knowing

can,

group

of feed

for the manufacture

in live

ligently
this situation intel-

meet

to

his

mation
this infor-

which

animals, proportion

thus go far toward

each and

securing

efficient production. It appears

desirable,therefore,
different point of view from that which
a somewhat

most
assume

has

the

to

standard,or

given quality. To

the feed

fixed

of the amount

pound of increase

milk

in order

largelyprevailedin
of

the past and

substitute for the

to

the modified

feeding standards

conceptionof

ception
con-

feed

requirements.
796.

Feed

the first to
the feed

requirements.

apply this

idea

requirements for
1

Minn.

Haecker

"

to

appears

Expt. Sta.,Bui.

have

been

productionand to formulate
productionof a pound of milk

milk

the

to

79

(1903),pp.

104-107.

694

NUTRITION

ANIMALS

FARM

OF

modified

by his subsequent experiments,

of requirements * has
become
this statement
known
the
as
Haecker
a
standard, although, strictlyspeaking, it is not
in the older sense.
The
writer 2 subsequently pubstandard
lished
a

As

grades.

of different

tentative

statement

pound

of milk

per

of the

proteinand
containingfour per

ments
requireenergy
of fat and
cent

illustrated the

computation of rations on this basis,without,

however, attempting similar estimates for other grades of
milk.
and
Eckles 5
Later
Woll
Humphrey,3 Savage 4 and
have
adopted various forms of the same
conception. Henry
6

Morrison

and

have

included

Haecker's

standards

in their modified

requirements for

Wolff-Lehmann

milk

production and
also similar data, based
on
unpublished results by the same
.experimenter,for growing fattening steers, and have also
somewhat

widened
and
On
and

introduced

range
minimum

of

different

several

in

chapters of
of

consideration
of

terms

therefore

are

in

open

relative values

the

to

the

of

trients
nu-

sources.

Requirements

797.

and

ignoring differences

from

figures.
foregoingrequirements
Eckles',are expressedin

all of the

digestiblenutrients

criticism

poses
pur-

maximum

hand, however,
standards,with the exceptionof
of

for other

standards

and

the other

terms

The

of the

the

Part

feed

the

of

terms

III

protein and
devoted

were

requirements of

digestible
proteinand

of net

energy.

primarilyto
farm

"

animals

in

In the case
energy.
be regarded as to a

of

tain
cermaintenance, these requirements may
degree fixed and capable of computation upon the basis
of the size of the animal, being related either to its weight or

its

to

body

surface.

contrary, the

(in excess

of

therefore made

In the

product. The results

togetherin Tables I- VI

of these

convenience

of the

head

1
8
3
6

estimates

for normal

Minn.

of

productivefeeding,on the
obvious
method
of comparison is that of feed
maintenance) with yield,and an attempt was
to estimate the feed requirementsper unit of
case

of the

growth

at

estimates

have

been

brought

Appendix, which include also for

total requirements per day and

different

weights and

ages.

Expt. Sta., Bui. 140, p. 56.

Bui. 346 (1909),pp. 19-25.
Dept. Agr., Farmers'
4 N.
Wis. Expt. Sta., Research
Bui. 13.
Y. (Cornell) Expt. Sta., Bui. 323.
6 Feeds
Mo.
Bui. 7.
and Feeding, 15th Ed., p. 669.
Expt. Sta., Research
U.

S.

696

NUTRITION

in

reduced

latter
at

are

times

the

ANIMALS

with

may,

safety,be

siderably
con-

instances.

some

tables

the

Furthermore,
of

FARM

quite possiblethat they

it is

and

OF

of the

Appendix

include

mates
esti-

no

ash

because
the
requirements. This is not
unimportant, for it is not improbablethat they may
be a controlling
factor,but simply because study

in this field has

progressedfar enough

not

permit

of their

emphasize here

certain

to

formulation.
while it has seemed

But

defects

of

desirable to

feedingrequirementsas formulated, as a precaution

their
uncritical
to
are
means
no
against
by
use, they
valuable
be rejectedas worthless but are capable of affording
feeder.
aid to the intelligent
By their use he can get a general
and can
idea of the feed requirementsof his animals
compute
rations which will approximatelysupply the requisite
amounts
His ability
of proteinand energy.
feeder will be shown,
as
a
the conditions which will modify
in his power
to estimate
first,
the feed requirementsof his particular
his
animals and cause
the

feeds to vary
which he can

and
second, in the skill with
average,
the dailyresults and modify his feeding
interpret

in accordance
799.

Dry

ration

from

the

with

them.

matter.

contains

variable,has

also be

must

its limits.

indication
matter

amount

of feed which

total volume

dry

The

"

shown

an

In

of

taken

dry

into

which

matter

the
The

consideration.

although rather
requires,
venient
computing rations the most conanimal

of the bulk

of the feeds

is the

of Tables

in the first column

percentage of
and

VII, VIII

Appendix. In very general terms it may be said

that
should be given from 20 to 30
a
1000-pound ruminant
pounds of dry matter
per day, 25 pounds being perhaps a
of the

IX

fair average,

while

for the

horse

smaller

amounts

will

be

appropriate.
An

examination

stuffs contain
to their

in

of the tables shows

much

dry matter than

where
heavy feeding,

more

do
the

that concentrated

proteinand
the

ing
feed-

proportion
Evidently,then,

energy

in

forage crops.
is to give the animal all the
purpose

possible,the ration should consist as largelyas practicable

of concentrated
because
feedingstuffs,
only in that way can the
creasing
of nutriment
be obtained
without unduly inrequired amount
the bulk of the ration. In lightfeeding,on the contrary,
feed

OF

COMPUTATION

THE

697

RATIONS

roughage may
predominate,because it is usually relatively
of feed in a bulk
cheaper and can supply the required amount
which

the' animal

"

can

consume.

Method

3.

the

examplesgiven on

The

following
pages
of

illustrations of the method

as

and

not

model

as

Computation

of

rations.

intended

are

using the tables

of

Limitations

of the

simply

Appendix

forbid

space
who

the

of examples, but the reader

the
multiplication
grasps
will have
serious difficulty
in applying it to his
method
no
will be acquired with surprising
own
conditions,while facility
rapidity by practice. It will be observed that the form of

tables

these

methods

the

and

of

computation do

differ

not

materiallyfrom those which have been used for many

years in
"
though
computing rations on the basis of
digestiblenutrients,"althe significance
of some
of the figuresis different. It
that the digestible
be added
proteinin the tables is true
may
include
the non-protein.
protein that is, it does not
"

Consequently
in

smaller

the

than

percentages,

rations

feed

average

condition
meal.

and-cob

are

Such

should

therefore make

"

pounds

1000

of

bunch

head

per

be fattened

to

energy
the
therefore,

on

"

animals
it is

value

grow

make

the

clover

and

gain of

sufficiently

computed

and

(Table I) for

to

should

dailygain
To

the average
weight of
the maintenance
use

1250

corn-

weigh
not

quire
re-

3.5 Therms
of

pounds,

this must

be

as

the

be 7 Therms.
requirement,which will increase

accurate

than

in better

hay

400

requirementwould
For

to

pounds. They
2 pounds per day.
gain of 1 pound live

average

in the feed ; for

heavier.

"

feeders

and

for market

ready

an

the maintenance

added

somewhat

are

gain of
It may
be estimated (Table III) that a
weight by animals of this grade will requireabout
of net

mated
esti-

amounts

cattle,if of good grade, should

pounds each when

over
200
days to

1400

the

as

succeeding pages,

on

required.

old,averaging

years

well

as

in the older tables.

Total

800.

the

pounds, viz.,7

pounds
requirement

1200

Therms.

This

of this section are

The contents
reproduced by permission of the Honorable
of Agriculture,
Secretary of Agriculture, from Bulletin No. 459 of the U. S. Department
prepared by the writer.
1

6g8

NUTRITION

total net

the

makes

hay,

it is

of Table

VII

of net

ANIMALS

on
requirement per day 14 Therms
of the whole feedingperiod.
that 2 pounds of grain will be fed for each pound
from
the figuresin the last column
easy to compute

the average
If we
assume
of

FARM

OF

the
as

energy,

energy

of feed

amount

follows

requiredto supply 14

Therms

"

Therms

In

100

In

200

pounds of average clover hay

pounds of corn-and-cob meal
pounds of feed
pound of feed

In 300
In

To

supply

of total

14

Therms

38.68
151.60

190.28

634

requires14

feed,consistingof

pounds of
of hay and
This, of

pounds

course,

feedingperiod.

hay

pounds

and

14.72

numbers, 7J pounds

of meal.

represents the average

the

At

22.08

of clover

7.36 pounds
meal, or, in round

corn-and-cob
15

0.634

4-

feed

beginning the

for

ration

will

the whole

naturallybe

lighterand consist to a largerextent of hay, while the amount

of feed,and especially
the proportionof grain,will be gradually
the end of the feeding,the animals
increased until,toward
are
consuming all the grain they will take,with only enough hay to
insure

the

too, the
in the

of the

course

assumed

was

that

decided

and
upon
required. Let us

and

whether

and
corn

digestion. Naturally,
proper
is given will usuallybe varied

feeding.

Improvement

801.
it

bulk
necessary
form
in which
the

of

ration.

the

the

feeding stuffs

attention
now

In

"

take

was

directed
the

up
of the

to

foregoingexample
be

used
the

simply to

question from

had

been

quantity

the other

end

ration may
not
study
yield some
suggestionof possibleimprovement.
meal,
According to Table VII, clover hay and corn-and-cob
contain
in 100
respectively,
pounds :
see

"

Total
Dry
Matter

Pounds

Clover

hay

Corn-and-cob

meal

Digestible
Protein

Pounds

Net
Energy
Value

Therms

87.1

4.9

38.68

89.6

5-7

75.80

THE

The

COMPUTATION

of clover

7J pounds

contain

OF

in

hay

699

RATIONS

the

will

ration

evidently

"

87.1

0.075

4.9

0.075

0.075

38.68

6.53 pounds of dry matter.

protein.
"-37 pound of digestible

of net

Therms

2-9"

preciselysimilar computation
gives the followingresults :

value.

energy

for the

corn-and-cob

meal

"

89.6

0.15

5.7

0.15

75.8

Adding

X0.15

of

0.85 pound
11.37

these amounts,

of

pounds

13.44

Therms

dry

matter.

digestibleprotein.

of net

energy.

find that the total ration contains

we

Net
Energy
Value

Therms

Clover

hay, 7! pounds
meal, 15 pounds
.

2.90

Corn-and-cob

n-37

Total

The

14.27

quantity

estimated
fixed

of

estimated

but

of such
fall below

IV

that which

is evident

VII

that

of

the

ration

is less than

were

that

beef cattle of this age

might produce fair gains,

advantage
the gain would

growth and

the

by

fail to take full

of the capacity

likely

most

increase in the tein

prothe ration more
efficient.

anticipated.An

was

we

the ration in this respect,it

into it some
feed much
richer

introduce

must

either of those

small amount

change

composing
we

or

meal,

of

one

in the ration.

old-processlinseed

meal,

needed

it is evident that what

only a

marked

the

like the above

feeds,like glutenfeed
that

of feed

amounts

expectedto make
change in
any marked

protein than

Table

that

observe, however, that

We

be

make

To

be

to

cattle for

might

in

basis.

digestible
protein in

in Table

corresponds with

course,
the
because

necessary,

weight. A ration
it probably would

and

of

energy,

that

on

upon

amount

be

to

of

meal

ration will foot up

for
as

need

is

the oil

it.
one

On

consulting

of the

uct
by-prod-

meals, etc., and

of these will be needed

Thus, if we
2

pounds

follows

"

substitute

to
2

also
effect

pounds

of the corn-and-cob

NUTRITION

700

OF

ANIMALS

FARM

Net
Energy
Value

Therms

hay, i\ pounds
meal, 13 pounds
Old-processlinseed meal, 2 pounds
Clover

2.90

....

Corn-and-cob

9-85

1.78

Total

14-53

comparatively small additional expense we are

able to improve the ration materiallyby adding the lacking
protein,and there is littledoubt that the improved ration would
produce a more
rapid gain and, under ordinaryconditions,a
the total gain
more
one
as
well,either by increasing
profitable
or
shorteningthe feeding period.
Thus

802.

at

Computing

available for

ration

from

given feeding stuffs.

"

There

dairyherd field-cured corn forage(including

the ears),clover hay, corn
meal, wheat bran and glutenfeed.
Table VII shows
if of good average
that these feeding stuffs,
quality,will furnish in 100 pounds :
are

"

Net
Energy
Value

Therms

Corn

forage
hay

43-94

Clover
Corn

38.68

meal

85.20

Wheat

bran

53-"o

Gluten

meal

84.15

The

cows

previousyears
4

per

850 pounds

average
an

of

cent

requirement

of 20
average
fat.
According

of

approximately:

animals

such

per head

pounds
to

per

and

of milk

Table

day

produced in
day testing

have

I,
and

per
the maintenance
head

would

"

Digestibleprotein

0.43

Net

5.40 Therms

energy

pound

be

COMPUTATION

THE

For

productionof

the

20

OF

pounds of

needed, according to Table

be

RATIONS

4 per cent

milk there would

"

Digestibleprotein (0.05 X 20)

(0.27 X 20)
energy

pound

1.0

5.4 Therms

Net

requirements per day and head

total feed

The

701

are

therefore

Digestible
Protein

Pounds

Net
Energy
Value

Therms

For

maintenance

0.43

5 -4o

For

milk

I. OO

5 -4o

production

Total

1-43

problem, then, is to find a

stuffs which will yield these

mixture

The

and

of energy,

and

which

shall have

of

amounts

coarse

fodders.

digestibleprotein

suitable bulk.

ration is to fix upon the

It is usuallydesirable to use
as

first step in the construction

The

IO.80

of the available feeding

of

amounts

of

since they are

large a proportion of these as possible,
usually
of feed than
cheaper sources
grain. On the other hand, the
of them

amount

which

animal

an

can

the individual animals, and

upon
only be told by trial,but we should

depends
can

from

12

to 14

pounds

of

dry

matter

is limited.

consume

Much

the proper amount

probably aim to get

in the form

of

coarse

fodder.

foragebeing a cheap feedingstuff,we shall naturallyuse

this freely,
with probably some
hay for variety. By a little
find that 10 pounds of corn forageand 6 pounds of clover
we
trial,
of
and the amounts
hay will give us 13.4 pounds of dry matter
Corn

digestible
protein and of

energy

shown

below

"

this

To

The

requirement.
will take, at
trial. We

to

the

pounds of

sufficient grainto

to add

have

we

amount

proper
a

venture,

bring the

we

pounds

Adding this

bran.

wheat

ANIMALS

FARM

OF

NUTRITION

702

meal

corn

the ration

to

up

ascertain

must

of

ration

have

we

by
and
:

"

Net
Energy
Value
Therms

forage,10 pounds
Clover hay, 6 pounds
Corn meal, 4 pounds
Wheat
bran, 2 pounds
Corn

Total

1.06
18

II.

low in

the

is

requirement as
regardsenergy,

ample as
digestible
protein. The
that

shows

more

feed

may

rather

low

added

be

to

computed,
erably
consid-

but

figurefor dry
the

if

ration

it evident that what

energy makes
feed,but feed of a different composition,

the total for net

but
desirable,

is not

is needed

totals with

these

the ration

rind that

matter

2.32
3-41

....

Comparing
we

4-39

more

Gluten
proteinalong with rather less energy.
this requirement,and substituting2 pounds of
meal answers
meal
it for 2 pounds of corn
gives a ration which, while still a
with the
trifle high in energy,
agrees as closelyas necessary
computed requirements. Thus :

supplyingmore

"

Net
Energy
Value
Therms

forage,
pounds
Clover hay, 6 pounds
Corn meal, 2 pounds
Corn

4-39

10

Wheat

bran,

Gluten

meal,

Total

This
whole

2
2

1.70

pounds

1.06

pounds

1.68
11.

....

ration

herd,
hardly need

correspondswith
since
be

and
milkers

2.32

it is based

said

that

it should

of
capacities

gettingmore

and

the average
its average
on

the

the

be

requirementof the
performance. It

modified

individual

ones
lighter

15

to
cows,

less.

suit the
the

quirements
re-

heavy

Let
a

us

ANIMALS

followingfeedingstuffs
:
pricesnamed
the

suppose
at the

dairyman

FARM

OF

NUTRITION

7"4

(40 cents

Corn

meal

per

offeedsper

ton

bushel)

$25
25

Wheat

bran

21

Wheat

middlings (flour)

24

brewers'

Dried

grains

Gluten

meal

Cotton

seed meal

23

27

(prime)

30

Old-processlinseed meal

supply of

The

animal

each

to

"

Prices
Oats

available

are

day

per

feed

coarse

32

33

farm

the

on

pounds

of

is sufficient to furnish

silageand

1000
pounds
hay ; the cows
average
to produce per day about
24 pounds

each

and

of milk

pounds
be

may

testing4.5

of clover

expected
per cent

fat.
the
compute, in precisely

first step is to

The

previous example, the

per day as follows :

estimated

same

way

requirements of

in the

as

these

cows

"

Digestible
Protein

Pounds

maintenance

For
For

pounds of

24

Protein
Net

24

energy

Total

The

24

milk

according

to

of
Table

6.96
i-75

silageand
VII, the

digestible
protein,and

net

hay available will furnish,

of dry matter,
followingamounts

energy

value

"

Pounds

Clover

silage,32 pounds
hay, 8 pounds

Total

12.96

clover

Total
Dry
Matter

Corn

6.00

1.25

0.29

requirement

amount

Therms

0.052
X

0.50

Net
Energy
Value

8.42

6.97
iS-39

Digestible
Protein

Pound

0.19

Net
Energy
Value
Therms

5-09

"39

3-09

.58

8.18

COMPUTATION

THE

OF

RATIONS

705

questionnow is what feedingstuffs is it most economical

to buy (or to refrain from
sellingif in stock) to complete the
ration. The first step in deciding this questionis to compare
The

various

the

feeds
of

unit

of energy
value at

sources

as

net

energy

and

see

lowest

the

computationgives the followingresults :

which

nishes
fur-

one

price. This

"

Cost
of
Therm
Net
Energy
Value

Cents

I.85

Oats
Corn

meal
.

bran

Wheat
Wheat

I.98
1.60

middlings
brewers' grains

Dried

Gluten

meal

Cottonseed

2.15
1.60

1.67

meal
meal

Old-processlinseed

1.86

Evidently,if it were
the

I.41

"

animals,we

should

simply a questionof supplyingenergy to

a unit
corn
use
meal, since that supplies

lower price than any

of energy at a much
stuffs. If it were
thought desirable to add

of the other

varietyto

the

feeding
ration,

middlings would obviouslybe our next choice.

It is evident,however, without
going through the labor of
computation,that while corn meal and wheat middlings may
be used in the ration,neither will supplyenough proteinif used
exclusively.Of the available feeding stuffs which are rich
wheat

in

proteinand
of this

and

It is not

pounds
to

of

meal

table shows

:
z

"

not

meal

next.

comes

great,

difficult to

coarse

very

is

corn

the

therefore

may

and

fodder

closelyto

we

shall

to

serve

is

ingredient,
glutenmeal

cottonseed
the two

one.

which

balance

the cheapest,
relatively
the

While

difference

naturallytry

determine

3^ pounds

ficiency
the de-

by
of

few

the

cheaper

trials that

gluten meal,

tween
be-

2"

in addition

sponding
available,will give a ration correthe following
the requirements,
as

706

NUTRITION

FARM

OF

ANIMALS

Net
Energy
Value

Therms

silage,32 pounds
hay, 8 pounds
Corn meal, 2" pounds
Gluten feed,3! pounds
Corn

5-09

Clover

3-09

Total

2.13

2-95

13.26

....

ration shows

This

close

as

could

proteinrequirement as
The
surplusof energy.
indicates that

ration.

computed
desired,but contains a slight

fodder

coarse

more

might

have

been

ter
mat-

used had

ration.
very economical
however, is nearly as cheap as a
a

of
source
meal,
more
as
glutenmeal, while it contains considerably
tein.
proit may
It seems
worth while, therefore,to see whether
the necessary
possibleto secure
cheaplyby
proteinmore

Cottonseed

be

the

comparativelylow figurefor dry

priceand apparentlyhave

not

be

with

agreement

available,with the probable effect of cheapening the

lowest in
have
As it is,we
used the feeds relatively

it been

energy

an

using a

smaller

meal.

Three

exactlythe
Making

pounds

same

this

placeof the gluten

cottonseed
meal will supply almost
of proteinas 3 J pounds of glutenmeal.

of the former

amount

of

amount

substitution,the

ration

feed in

stands

as

follows

"

Net
Energy
Value

Therms

silage,
32 pounds

Corn
Clover

5-09

3-09

2.13

hay, pounds
meal, 2\ pounds
Cottonseed
meal, 3 pounds
8

Corn

2.70

Total

13.01

This
than
is
as

trifle cheaper. The

computed requirementseven better

that it
a simplecomparison shows
grain portionof the two rations costs

ration agrees with the

the previous one, while

follows:

"

THE

RATIONS

OF

COMPUTATION

707

Second
Ration

meal

Corn

meal

Gluten

meal

Cottonseed
Total

It thus appears

that the ration made

meal

expensivecottonseed

more

is

with

up

the somewhat

actuallythe cheaper.

The

fed for 200

small, yet for 30 cows
to $30. Such a difference is only likely
days it would amount
in this instance,
assumed
to be found, however, when, as was
be had at a relatively
cheap
feed very high in proteincan
some

difference,to

is

sure,

general,it

In

rate.

be

may

be said that when

differences in the cost

marked

of

Therm

there

no

very
value in
of energy
the ration,that one
are

feedingstuffs constitutingthe bulk of

at the
feeds which suppliesenergy
of the various high-protein
be used, although it is always
lowest cost should ordinarily
wise to check up this point,as in the example justgiven.
While in the foregoing
804. The
compounding of rations.
examples an exact dailyration is computed, it would, of course,
to weigh out
in most
separately
cases
be utterlyimpracticable
the

"

each

day's ration

for

feeds at intervals would

each

animal.

often

Individual

yieldvaluable

weighings of

information

and

be undertaken, but for the ordinary routine

might profitably
be used.
must
of feeding,simplermethods

be advantageously
the grain feed may
practicable,
in the desired proportionsin as large quanmixed in advance
tities
the
the storage capacityavailable and
tion
as
proper preservaof the materials will permit. Where facilitiesare available,
be
of grainrequiredfor all the animals may
the whole amount
ditional
adfor each feeding,without much
weighed out daily,or even
the individual
In distributingthe grain to
labor.
animals, regard,of course, should be paid to their productive
ration as
The
capacity and their individual peculiarities.
When

The skill of the feeder is

animal.
is for the average
to the individual.
in adaptingit in qualityand in amount

computed
shown

708

weighings

individual

Doubtless

useful

be

would

intervals,

at

control

as

ANIMALS

FARM

OF

NUTRITION

the

on

already

as

gested,
sugof

accuracy

the

distribution.

problem

simple.

fodder,

control
all

In

of

made

or

less

seek

in

actual

the
of

the

would

conducted
of

hundreds
be

will

play

animals

are

Finally,

animals.

rations
feeder
cannot

in

are

it

absurd

be
should

guides

wisely

using

the

the

place

of

take

fed,

being

be
and

not

resources

experience

to

or

go

feed

the

scale

important

an

would

which

sought

of
The

to.

amount

it

is

it

foolish,

be

to

in

the

They

recipes.

and

his

or

profitably
two

three

or

these

may

command,

good

feeding
scores

that

remembered

at

the

may
of

in

saving

Where

case

to

expense

the

exactness

more

therefore,

which

part.
an

best

at

be

can

more

than

upon

necessary.

that

estimate
but

would

apportioning

and

weighing

practicable,

is

sense

best

realizing

in

exactness

latter

the

It

if

weighed.

desirable.

very

requirements,

average

cut

or

comparatively

readily

not

common

expresses

silage

weighing,

matters

approximation.

an

extreme

the
is

ration

computed

The

is

contrary,

difficult

more

still

is

matter

is

however,

occasional

an

similar

and

these

the

feeding,

the

upon

usually

When,

the

on

here

even

bulk.

its

trucks,

in

Long

Nevertheless,
a

of

handled

is

is

fodder

coarse

account

on

fodder

as

of

weighing

The

judgment.

puted
com-

aid
but

the

they

APPENDIX

APPENDIX

712
III.

Table
.

Requirements

"

Growth

All

"

Species

for
"

in

Fattening
Addition

with
to

no

Considerable

Maintenance

the

Requirement

Table

IV.

Requirements

"

for

Growth

with

no

Considerable

Fattening

a.

Per

Pound

Weight, in Addition
Requirement

of Increase

in

Live

Estimated

from

Estimated

from

Estimated

proteincontent

protein content
experiments on

to the Maintenance

of increase.

fattening (456).

of increase.

APPENDIX

b. Per

Day

and

Head, Including Maintenance

(1) Cattle

(2) Sheep

(3) Swine

Based

on

Kellner's standards.

713

APPENDIX

7i4
V.

Table

Add

pound

of milk

Table

VI.

for each

of the several grades.

Requirements

"

Production

Milk

for

requirement the following amounts

maintenance

the

to

Requirements

"

Per

Work

for

Production

Digestible
Protein

Net

Half work
One-fourth

8 hrs. per
4 hrs. per

"

"

work

"

AVERAGE

DRY

ENERGY

VALUES

day
day

hrs. per

day

2.0

18.2

1.4

11.

AND

PROTEIN

STUFFS

FEEDING

PER

7.6

1.0

DIGESTIBLE

MATTER,
OF

Energy2

Therms

Pounds

Full work

(674)

Horse

the

Live Weight

Pounds

1000

by

100

NET

POUNDS

able
recentlypublisheda very valuanalysesof feedingstuffs and of
compilationof American
the results of American
experiments,and on this basis
digestion
nutrients in a great
of digestible
have
calculated the content
varietyof feeding stuffs.
With the permissionof these authors and with the cooperation
of Assistant
Professor Fred Silver Putney, of The Pennsylvania
the writer has computed from their tables the net
State College,

Henry and

Kellner's

Feeds

and

Morrison

have

standards.

Feeding, 15th Edition,

pp.

To

be

computed from

633-666.

Table

VIII.

APPENDIX

values

energy

of the

important feedingstuffs in the manner

Chapter XVII
(773,774) with the results regarding

in

described

715

more

reportedin

ruminants

Station

Bulletin No.

142

and in Bulletin No.

Experiment
Department of Agriculture.Those

of the
459

vania
Pennsyl-

of the U. S.

tions
results,with a few addihere reproduced and the computaand
are
corrections,
tion
well
basis
the
also
been
has
as
extended, as
now
meager
available will permit,to the data regardingswine suppliedby

Henry

tables.

Morrison's

and

in part from

the

for
figures

The

and

source

same

the horse

are

rived
de-

in part from Zuntz

values being
energy

the net
Hagemann's investigations,
gators
computed according to the method proposed by those investi(775-778).The tables show primarilythe net energy
values for maintenance
or
good reason
fattening.There seems
for believing,
however, that they may be taken without serious
and

represent also the

to

error

net

energy

values

for

growth and

productionand at least the relative values for milk

production.
Henry and Morrison's tables include only the crude protein
of non-proteinhas been estimated
(N X 6.25). The amount
from the crude proteinby the writers on the basis of Kellner's
for work

averages.
VII.

Table

"

Values

per

100

Pounds

for

Ruminants

Net
Energy
Value

Dried
and

Hay
Brome
Corn

grass,

Roughage

fodderfrom

40.83

smooth

(maize) fodder

(earsincluded, medium

dry)
Corn

(maize)

dry)

Therms

cereals

stover

43-94

(earsremoved, medium

Kafir fodder,high in water

Kafir stover, high in water

Millet,Hungarian
Mixed
timothy and clover
Oat hay

31.62
34.28
27.65
46.96

40.85
32.25

APPENDIX

716

VII.

Table

Values

"

per

Pounds

ioo

for

(Continued)

Ruminants

Digestible
Net
Dry
Matter

Dried

Hay
Orchard
Prairie
Red

grass

hay

top

fodder, computed

Sorghum

to

80

cent

Timothy,
Timothy,

before

per

analyses

Timothy, early

bloom
to

full bloom

Timothy, late bloom to early seed

Timothy, nearlyripe
Hay

and

True
Protein

Pounds

Pounds

Pounds

88.4

4-7

3-3

44-93

93-5

4.0

2.9

40.42

90.2

4.6

3-9

51.22

80.0

Therms

dry

matter

all

Crude
Protein

Roughage

fodderfrom cereals

and

Energy
Value

2-5

i-5

32.20

88.4

3-0

2.2

43.02

92.8

4-7

2.9

43-52

87.2

3-6

2-5

47.40

85-1

2.4

1.8

37-54

87.5

2.2

1.8

38.59

91.4

10.6

7-i

34-23

93-8

154

fodderfrom legumes

Alfalfa,all analyses
Alfalfa,before bloom
Alfalfa,in bloom
Alfalfa,in seed
Clover, alsike
Clover, crimson
Clover, red, all analyses
Clover, red, before bloom
Clover, red, in bloom
Clover, red, after bloom
white
Clover, sweet
all
analyses
Cowpeas,

92.5

.......

.....

Soybeans

10.5

6.7

36-23
32.33

6.2

32.23

7-9

5-3

34-42

89.4

9-7

6.9

87.1

7.6

4.9

36.21
38.68

89.6

8-5

87.7

89.6

Cowpeas, before bloom

Cowpeas, in bloom to early pod

10.3

11.6

5-4

42.17

86.1

8.1

5-3

39.12

77-9

6.8

4-5

34-51

6.7

38.98

91.4

10.9

90.3

i3-i

92.2

17.8

89.4

12.6

91.4

11.

9.2

37-59

12.8

33-54

9-5

39-n
44-03

Straws

Barley
Buckwheat
Oat
.

Rice

Rye
Wheat

0.6

36.61

85.8

0.9

90.1

4.2

3-2

4-55

88.5

1.0

0.8

92.5

0.9

0.4

34.81
23.63

0.7

0.5

17-59

0.7

o-3

7.22

'.'."'.'.'.
92.9

91.6

APPENDIX

Table

VII.

"

Values

per

100

717

Pounds

for

Rumtnants

{Continued)

Digestible
Net

Dry

Energy

Matter

Fresh

Green

Blue

fodder
before

heading
Kentucky,
headed
out
Kentucky,
grass,
Blue grass, Kentucky, after bloom
Buckwheat, Japanese
Cabbage
Cabbage, waste outer leaves
Corn
(maize fodder, dent, all analyses
Corn
(maize fodder, dent, in tassel
Corn
(maize fodder, dent, in milk
Corn
(maize fodder, dent, dough to glazing
Corn
(maize fodder, dent, kernels glazed
Corn
(maize fodder, dent, kernels ripe
Corn
all analyses
(maize fodder, flint,
Corn (maize fodder, flint,
in tassel
Corn
in milk
(maize fodder, flint,
Corn
kernels glazed
(maize fodder, flint,
kernels ripe
Corn (maize fodder, flint,
Corn
(maize fodder, sweet, before milk stage
Corn
(maize fodder, sweet, roasting ears or
grass,

....

later

(maize) fodder, sweet,

Millet,Hungarian
Corn

ears

fodder

Orchard

Value

Pounds

Pounds

Pounds

Therms

23.2

2-3

23.8
36.4
43-6
36.6
8.9

Roughage

Blue

Oat

True
Protein

cereals,etc.

Green

Barley

Crude
Protein

grass

removed

2.0

14.08

3-7
2.8

2.8

14.82

2.2

17.77

1.9

1.6

21.01

2.2

i-5

17.78

1.9

i-3

8.87

14.1

i-7

1.1

23.1

1.0

0.8

14.9

1.1

0.8

9-52

19.9

1.0

0.8

13.64

25.1
26.2

i-3

1.0

17-35

1.1

0.8

16.74

34-8

i-5

1.1

22.48

20.7

1.0

0.8

10.6,

0.9

0.7

15.0

0.9

0.7

10.39

21.0

1.0

0.8

13-49

27.9

1.2

0.9

17.84

10.0

O.8

0.6

7.82

20.3

1.2

0.9

13.38

21.5

0.8

14.26

1.9

1.1

17.24

26.1

2-3

2.0

14.06

29.2

i-7

1.1

15.81

16.7

2.6

1-7

13-07

21.3

2.1

1.4

15-99

24.9

0.7

0.4

15-37

1.1

18.36
18.89
26.36
18.75

24.2

32.1

i-3

0.8

46.4

i-5

1.0

Wheat

27.4

2.8

1.9

Green

Clover, crimson

6.89

I.O

1.8

Alfalfa,before bloom
Alfalfa,in bloom
Alfalfa,after bloom
Clover, alsike

13-53

27.6

Rape
Rye fodder
Sweet
sorghum fodder
Timothy, before bloom
Timothy, in bloom
Timothy, in seed
fodder

7-05

14.60

legumes
.

19.9

3-5

1.9

9.20

25-9

3-3

1.8

11.50

29.8

2.1

i-3

11.

24-3

2-7

1-5

17.4

2-3

1.6

14.56
10.83

10

718

APPENDIX

VII.

Table

Values

"

per

ioo

Pounds

for

Ruminants

{Continued)

Digestible
Net
Dry
Matter

Green

Fresh

Green

Energy
Crude
Protein

True
Protein

Value

Pounds

Pounds

Pounds

Therms

26.2

2-7

1-7

15-87

2-7

1.8

16.74
17.30
10.42

Roughage

legumes

Clover, red, all analyses

Clover, red, in bloom
Clover, red, rowen

27-5
344

3-3

2.2

16.3

2-3

1-7

16.6

2.9

2.1

23.6

3-2

2.4

12-53

20.8

3-o

2-3

10.44

24.2

3-i

2-5

12.70

18.1

3-5

2.4

n-95

26.3

1.1

O.6

15.90

21.0

1.0

O.4

11.96

25-3

1.2

O.6

14.27

19.6

0.5

0.3

8.98

27.8

i-3

O.8

7.26-

Cowpeas

22.0

1.8

I.I

11.05

Soybeans
Sugar beet pulp

27.1

2.6

1-5

n-59

10.0

0.8

0.5

Apples

18.2

0.4

O.I

Beets, common
Beets, sugar

13.0

0.9

O.I

16.4

1.2

0.4

Carrots

11.

Cowpeas
Peas, Canada

field

Soybeans, all analyses

Soybeans, in bloom
Soybeans, in seed
Vetch, hairy

9.78

Silage
Corn
Corn
Corn
Corn

(maize),well-matured, recent
(maize),immature
(maize),from
(maize), from

frosted

analyses

ears

field-cured stover

Clover

Roots, Tubers,

and

Mangels

9-32

Fruits

9.4

15-92

7.84
11.20

0.9

0.5

9.21

0.8

O.I

5-68

21.2

1.1

O.I

18.27

Potato

flakes

87.9

3-6

0.4

72.68

Potato

flour

89.4

1.4

0.1

80.09

1.1

0.6

6.05

Potatoes

Pumpkins,

field

8-3

1.0

0.3

8.46

1.0

0.4

6.16

90.7

9.0

8.3

Buckwheat

87.9

8.1

7.2

59-73

Corn

89-5

7.5

7.0

Rutabagas

10.9

Turnips

9-5
Grains
Cereal

grains

Barley

Corn

(maize),dent
(maize),flint

Corn

(maize) and

cob

meal

89.94

87.8

7-7

7.2

85-5"
84.00

89.6

6.1

5-7

75.80

APPENDIX

720
VII.

Table

Values

"

per

Pounds

ioo

By-Products
industries

Fermentation
Distillers' grains,wet

....

Malt
Malt

sprouts

Milling
Buckwheat

bran

Buckwheat

hulls

Buckwheat

middlings

....

feed

Hominy
Red
dog flour
Rice bran, high grade

....

Rice meal
Rice

polish

Rye

bran

Wheat

bran

Wheat

middlings, flour
middlings, standard

Wheat

....

Oil extraction

meal, low in fat

meal, high in fat

Cocoanut
Cocoanut

,.

hulls

Cottonseed

meal, choice
Cottonseed
meal, prime
oil meal, maize
Germ
Linseed meal, new
process
Linseed meal, old process
Cottonseed

....

cake

Palmnut
Peanut

cake from

hulled nuts

cake, hulls included

Soybean meal, fat extracted
Peanut

Sunflower

seed

cake
.....

Starch

Gluten

feed

Gluten

meal

manufacture

Starch feed,dry

feed,wet
Sugar manufacture
Molasses, beet
or black strap
Molasses, cane

Starch

for

Ruminants

(Continued)

APPENDIX

VII.

Table

Values

"

per

100

721

Pounds

for

(Continued)

Ruminants

Net
Energy
Value

By-Products
Therms

Sugar manufacture
Molasses

beet

pulp
pulp,dried

76.28
75-87

9-32

8.99

....

Sugar beet
Sugar beet pulp, ensiled
Sugar beet pulp,wet
Packing house
.

Dried

68.12

blood

Tankage
Over

protein
cent protein
cent protein
per cent protein

60 per cent

55-60

per

45-55
Below

per
45

VIII.

Table

"

93-Q4

83-58
72.96
54.16

Values

per

100

Pounds

for

the

Horse

Net
Energy
Values

Therms

48.82

Alfalfa hay

clover

hay
Timothy hay
Red

Wheat

straw

Beans

39-94

26.64

"

20.90
109.40

....

Corn

(maize),dent

112.80

Corn

(maize),meal

132.70

Oats

93-44

Peas

105.20
101.60

Linseed
Carrots
Potatoes

cake

16.60
....

3 A

35-7Q

APPENDIX

722

IX.

Table

"

Values

per

ioo

Pounds

for

Swine

Net
Energy
Values

Therms

Grains

106.08

Barley
Corn

(maize),

Corn

(maize)

meal

Corn

(maize)

and

118.82'

dent

120.25
cob

meal
.

Pea

meal

rough

Rice,

110.98
123.68

Rye
seed

Sorghum

100.59

JWheat

108.85

Milling
Red

103.30

122.43

products

flour

dog

107.02

Wheat

bran

Wheat

middlings,

74.95

standard
.

Oil

meal,

Linseed

old

meals

process

110.85
.

meal

Soybean

103.73

108.42

Sundries
Dried

blood

Tankage,
Potatoes

Skim

116.89
over

60

per

cent

protein

109.39

24.69
milk

14.74

APPENDIX

")
"

"S"

itMlOOM

to

"

723

t^

00
-"*

000

00

N*w

"n

"

t!-

t^Tj"u

"

c5

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APPENDIX

INDEX
80
Abomasum,
Accessory substances,348,
significanceof,632

Alfalfa proteins,
value of, 683
Alkali ratio of ash, 342

421

Alkaloids,37
Amids, 37

Acid:

hippuric,163
hippuric,synthesisof, 163
uric,synthesisof, 171
Acidity in ash, significance
of, 341
Acidosis,336
Acids

in plants,38
acids,37
from simple proteins,
28, 29
in plants, 38
occurrence
relative values for growth, 381
required for maintenance,314
Ammonia, formation of,in katabolism
proteins,165
of feed,influence on
Amount
occurrence

Amino

excretion of,338
influence

digestibility,
627
neutralization of, 337
nucleic,34
anabolism, 168
autogenesis,169
cleavages, 170
deaminization, 171
katabolism, 170
metabolism, 168
169
synthesis,
on

effects of temperature,
metabolizable
milk
net

production, 515
values, 664
energy

Amylase, 78, 86
Amylopsin, 78, 86
Anabolism, 145
of fats,171
nucleic acids,168
phosphorus, 180

of,in digestion,
40, 158
of, 159

Adipose tissue,
58, 424
compositionof, 59

160
simpleproteins,

Age:
best for

449

664

of methane, 665

in feeding stuffs,
40
metabolism

energy,

454

production,269

organic,40
formation

production, 443,

meat

Animal

fattening,
436

factor in meat

production,428
production, 470
composition of gain, 431
as prime motor,
192
cost of production of meat, 430
Araban, 15
610
digestibility,
Arabinose, 9
effects of temperature, 454
Arteries,126
for
requirements
nance,
mainteAscent, work of, 551
energy
307
efficiencyof body in, 551
feed consumption, 431
Ash, s
net energy
acid and basic,340
values,666
alkali ratio of, 342
production of lean meat, 433
relation of growth to, 373
balance, 216
relation of protein requirements to,
maintenance
of,339, 344
body, proportionof in bone, 48
445
in offal,56
Albuminoids,S3
Albumins, 33
bone, compositionof, 48
influence

as

milk

on

725

of

INDEX

726
Ash,

in

of feed, 332

content

correction of deficiencies in,346

of, in feedingstuffs,67

determination

digestionof, 101
effects of deficiencyof, 420
6
ingredients,

availabilityof, 417
balancing of, in ration,343
deficiencies in, 339
of, 118, 333, 343
digestibility
excretion of, 141
functions of, 187

skeleton

significanceof results of, 241

Barley feed, 585
Bases, organic, 37
Bile, 86
Blood, 123
coagulation of, 125
red, 124
corpuscles,
white, 124
of, 127
course

Body:

plant,567
comparison with power
compositionof entire,61

of, 178
of,338

reserve

as

fat- and

losses of, 334

of, 334
of milk, 460
sources
of,468
outgo of, in milk, 520
proportionof, in animal, 5
in feeding stuffs,5
rate of storage of, in growth,
requirements for growth, 414
maintenance,

substances

physicalregulationof,262

significanceof acidityin, 341

in dairy rations,521
total
retention of, during
:

Autogenesisof

441

on,

individualityon,

Balance

growth,

Bone, 47
ash composition of, 48
composition of, 47
proportionof body ash in,48
proteinin, 48
Bones

416
of breed

441

nucleic acids, 169

Breakfast

food

Breathing

residues,584

example of, 206

216

assimilative power,

matter,

expenditure,194

and

digestivepower,

202

nitrogen, 202
determination

of, 203

example of, 203

nutrition,192, 201
includes

water,

216

energy,

216

experiments,

comparison

with

200

metabolism
.

practicalexperiments,244

441

440,

610

early maturity, 441

feed consumption, 443
maintenance
requirements, 442
meat
production, 440
values, 666
net energy
Brewers'
grains,586
By-products, 582
gations,
investiof, 582
nature
of fermentation
industries,585
-

241

of, 137

composition of milk, 473

composition of milk solids,474

example of, 240

income

338
ingredients,

of ash

reserve

of, 134
regulation of rhythm
Breed, influence on

carbon, 205

Balance

as

Bran, 582
rice,583
rye, 582
wheat, 582

mechanics

of ash, 216

energy,

regulation of,

chemical

263

supply

influence

for work,

544

"

332

Assimilative power

of energy

sources

temperature,

414

production,520
production,562

work

ash-free,65

fat-free,64
efficiencyof. (See Efficiency)
expenditure by, 192
schematic, 195

causes

milk

"

plasma, 125
platelets,
124

indispensable,
332
metabolism

experiments, continued
investigations,
agricultural
243

Balance

continued

"

milling,582

INDEX

728
Computation,

Dextrins,

continued

"

Concentrates,

72, 579

comparison with roots, 579

apparent,

by

roughage, 662
of digestibility
of, 115
proportionof, to roughage, 451, 696

determination

for,671

relative values

of,

influence

Condiments,

bility,
digesti-

on

627
Condition, influence of,
of gain, 438
on
economy
meat
production,438
rate of gain in fattening,438
601,
affecting digestibility,
Conditions
602, 613
Conditions, external, influence

on

milk

relation

Conformation,

of, to

meat

duction,
pro-

443

of energy,

Conservation

219

Contraction, muscular, 532

chemical changes in, 532
energy

Corn

transformations

in,533

bran, 588
meal, 587

Cottonseed

264, 453
by feed consumption, 308
Crude
fiber,13
compositionof digested, 120

Critical temperature,

of

net

energy

values

for,

of, in feeding stuffs,71

determination

heat
of, on
horse, 676

influence

production

of

proportionoS pentosans in, 71

Cutting

excretion, outgo of chemical

in, 231
energy
of roughage, influence
of, on
624
digestibility,

Cytoplasm,

42

Dairy rations
addition

influence on,
of acids,627

addition of protein,622
610

breed, 610
condiments, 627
conditions relatingto the animal,602
feed, 613
cutting of roughage, 624
drinking, 628
drying, 623
of carbohydrates, 616
excess
feeds rich in carbohydrates, 617
grindingof grain, 624
heavy feeding, 450
individuality,609

roots, 618

tubers, 618
water
drinking, 628

work,

cereal grains,580
determination
concentrates,

of nucleic

acids, 171
simple proteins,165
reversible,166
in ash

of,346

effects of, 420

of

of, 115

maturity

on,

574

forage, influence
on,

of,

621

maize

proteinin, 506

correction

610

laboratory determination of, 116

of ash ingredients,
118, 333, 343
carbohydrates,diminished, cause

ether extract, 119

influence
grasses,

ash in, 521

Deficiencies

ruminants,

604
species of ruminants, 603
swine compared with ruminants, 606
conditions
affecting,601
definition of, in
determination of, in,
114
influence of excretory products, 118

of fat to, 517

Deaminization

with

603
species,

669

Cutaneous

compared

non-protein,622
proteinsupply,447
quantity of feed, 613

lowered

correction

120

horse

age,

of,

production,453
production,478

meat

14

Dextrose, 8
(SeeNutrients)
Digestible nutrients.
Digestibility,in, 601

of, 697
of total feed required, 697
method

of maturity

575

nitrogenous substances, 119

of,619
protein,diminished, cause
601
variable,
variation of, at different times, 002
Digestion, 77
changes in, 101
chemistry of,89

INDEX

Digestion,

"

conversion

continued
colloids

of

729
of feeding, influence

Economy
into

loids
crystal-

in, 102

Efficiencyof body,

experiments,example of, 114

as

of, 112
time requiredfor, 113
of protein cleavage in, 98
extent

compared

economic, 562

intestinal,
87

gross

methods

molecular
of ash,

in, 103
simplification

motor,

and

544

544

with

conditions

in work

of individuality

472

on,

power

plant,567

affecting,555
net, 546

of ascent, 551

draft,552

101

carbohydrates, 89
cellulose,
89
disaccharids,95
electrolytes,
101
fats,88, 95
hemicelluloses,92
96, 100
non-proteins,
nucleic acids,99
pentosans, 91

phosphorus,
83, 88, 95
proteins,
by erepsin,
98
pepsin,96
101

trypsin,97
starch, 79, 88, 92
in intestines,
94
stomach, 93

sulphur,101
of, 77
general plan of, 77

organs

solution of nutrients in, 10 1

uniformity of nutritive material, 103
work of, 277
differences between feedingstuffs,
663
with
roughage compared
trates,
concen662

influence of breed, 440

individuality,440
from
feed in milk
Diminishing returns
production, 515
Disaccharids, 10
digestionof, 95
general propertiesof, 1 1
Distillers' grains,586
Draft, work of, 552
efficiencyof body in,552
Dried blood, 590
Drinking, influence of, on digestibility,

Digestivepower,

628

Dry matter, 3
of body, composition of fat- and ashfree,65
requirementsof,695
Drying, influence of, on digestibility,
623
Duodenum,
84

influence on,
of fatigue,556
forms

of work, 555

gait,558
grade, 559

individuality,
556
intensityof work,
load, 559

557

speed, 552, 557

training,556
mechanical, 545
over-all,562
per day, 549
variable,548, 555
Efficiencyof muscle, 545
Electrolytes,digestionof, 101
values
Embryo, net energy
of, 393
Emulsification
Emulsion

of fats in

for growth

digestion,95

of fats,19

Energy :
available,233
balance of, 216
example of, 240
in milk production, 493, 495
chemical, 218
outgo of, 229, 635
conservation of, 219
definition of, 216.
expenditurein,
internal work, measure
of, 256
feed consumption, 275
significanceof, 277
locomotion, 550
influence of speed on, 552, 557
for work, body substance
source
as
544

proteinas
forms

source

of, 542

of, 217
gross, 227, 635
income
of, 226
katabolism of, in fasting,
256
constancy of, 256
kinetic,218
measurement
of, 325

of,

INDEX

73Q
Energy,

continued

"

Energy,

continued

"

sheep, 292
swine, 285

outgo of, 235

losses of, 229,

235, 635
chemical, 299, 635
in feces,230, 635
fermentation, 230, 636
computation of, 627

heat

relation of temperature
milk

of

net

of, for work, 542

of, on retention of
protein, 386
total, not measured
by heat of combustion,
sources

energy

values

with, 271
computation of,
from

223

transformations
in muscular

nutrients,646
digestible

utilization of,
in growth, 390
milk production,493

for the horse, 675

generalconceptionof, 231
on,

of amount

of, 218
contraction, 533

units, 220

organic matter, 648

factors for, 234

influence

511

production,562, 564

supply, influence

urine, 231, 636

metabolizable, 231, 639

comparison

production,

work

production,235, 650

to, 308

production,448

meat

work

of feed, 664

production,544
(See Net

Energy Values)
milk
of, on

of

values, net.
Environment,

in combustible

production,478
reactions reversible,150
Enzym
Enzyms as agents in metabolism, 148
digestive,78
extracellular,
148
intracellular,
149
in the body, 150

of determining, 640

method

nutrients,648
digestible
feeding stuffs,642
real and apparent, 645
of, 645
significance
for,
233
synonyms
net.
(See Net Energy)
outgo of, 229, 235, 635, 650
gases,

230,

636

influence

feces,230, 635

Epithelium,105

heat

Erepsin, 79, 87, 98

action of, on proteins,
98
Esophagus, 79

production,235, 650

urine, 231, 636

productionvalues as regards, 634

of,318
proteinas source
rate of gain of, in growth, 378
requirements
for fattening,361

growth, 399
of cattle,
399
sheep,401

Ether

of ash

of age,

manner

140

water, 142
:

feed cost of,481

307

fattening,306
plane of nutrition,305
stage of fattening,362
temperature,

ingredients,
141

dioxid, 139
nitrogenous products,
Exercise

on

carbon

swine, 400
maintenance, 267
factors affecting,304
influence

extract

digested, 122
of, 119
digestibility
of feeding stuffs,70
Excretion, 123, 139
functions of kidneys in, 140

304

of stating,283

methods

of determining, 281

modified

conceptionof, 284

of, on meat production,457

production,480

influence
milk

yield of milk fat,482

Expenditure, balance of income and, 194
in horizontal
of energy
locomotion,
55o

Extractives, percentage of, in lean meat,

of cattle,288

357

farm

animals, 280, 303

fowls, 301

horses,295

Farm

animals, composition of bodies of,

62

INDEX

731
Fats,

Fasting :
katabolism

in, 251
functions of proteinin, 255
katabolism, 249
computation of, 282
conditions affecting,258
expended in, 251, 257
energy
energy

influence

reactions of, 18

digestionof, 88,

95

oils and, 19

distinction between

emulsification of, in digestion,

95
emulsion of, 19
formation
from

on,

fat, 252
external temperature, 262, 265
muscular activity,261
previousfeeding, 253
size of animal, 258
standing and lying, 262
substances katabolized,in, 249

of carbohydrates from, 178

carbohydrates, 155,

protein,168,

of body

174

173

functions of, 186

of, 18
hydrolysis
katabolism

of, 176
melting pointsof, 19
metabolism of, 171
molecular

of, 17

structure

native, 19

proteinkatabolism

in, 251
normally small, 251
variable, 251
of in
and lean, proportions

Fat

continued

"

chemical

of,1

occurrence

of milk, 459
carcass,

oxidation of, at /3carbon

atom,

177

physicalpropertiesof, 18

424

relation of, to growth, 421

Fat:

addition of, to dairy rations,517

96
of, in digestion,
saponification

animal,

effects of feeds associated with,

specific

of, 173

sources

body, influence

of

on

ism,
fasting katabol-

proportionof in offal,56
computation of gain or loss of, 205

compositionof

crude, determination of, in feeding

stuffs,70
gain or loss of, 205
manufacture
of, 172
minimum
of, for milk production,519
mobilization of reserve, 177
of feed, resynthesis
of, 171
of milk, influence

of exercise

on

527

Fattening, 350
best age for,436

252

yield

increase

during growth, 448

content of gain in, 361
energy
requirementsfor,361, 448
energy
equivalentenergy values for,572
gain of proteinin, 354, 364
influence of condition on, 438
compositionof lean meat, 356
on

529

net

production,protein

for,

object of,358, 427

animals, 35"
of mature

nance,
requirements for mainte-

energy

363

proportionof, in

meat, 425
relative utilization of carbohydrates

and, for work production,553

requirements of, for milk production,

352,

concurrent, in milk production,

513
contrast with growth, 396

of,482
originof,466
percentage of, in lean meat, 356
milk, influence of feed on, 528
milk solids,influence of feed on,
unnecessary

in, 350,

353,354-364

306
for,360

values

energy

pigs,proteinrequirementsof, 41 1
proteinrequirementsfor,363, 446
rations,proteinin,364
requirements,350, 359, 361, 363,

44",

5i6
storage of, 172
Fatigue, influence of,

of, on energy
362
utilization of proteinin,364

stage of, influence

on

efficiency of

body, 556
milk production,
482
Fats, 16
anabolism of, 171
animal, elementary composition of, 21
chemical changes in resorptionof, 108

Fatty acids, 17
Feces, 105,
as

109

excretory

product, 109

feed residue, 109

compositionof, 11

quirements,
re-

INDEX

732
Feces,

"

classification of, 571

in, 635
in, 230

composition of, 66
determination

Feed:
as

stimulus to milk

production,522

crude

of,
production, 651
the
horse, 675
by
metabolism, 651

influence

on

heat

influence

on,

of age,

431

breed,

443

individuality,
443
significanceof

energy

fat in, 70, 71

protein in, 68
nitrogen-freeextract in,71
non-protein in, 69
protein in, 67
true protein in,67
,
water
in, 67
direct comparisons of, 591
ether extract of, 70
metabolizable
of, 639, 642
energy
production values of, 630, 634, 678
relative values of, 591, 597
rich in carbohydrates, influence of, on
crude

expended in, 275

increases heat production, 273

on

of

ash in, 67

consumption,
energy

continued

Feeding stuffs,

continued

"

losses of energy
outgo of energy

expenditure in,

277

digestibility,
617
e
ffects
of,448
specific
associated with fats,527
milk production, 523
on
of, 571
sources
sundry ingredients of, 39
digestibility,
practical,592
Feeding trials,
Fermentation
industries,by-products of,

from, in milk
diminishing returns
production, 515
dual function of, 183
influence of, on
composition of milk,
527

quantity of,

influence

of,

on

613
requirements, 691, 693, 694
for growth, 396
maintenance, 280, 313

production,445

meat

milk

production,

500

supply,569
aspects of, 631

two

surplus,disposal of, 350

of, for meat

total amount

449

unit system,

S85
Fermentation, losses of chemical
energy
in,636, 639
computation of, 637
Flavoring substances, 41
milk production,522
influence of, on
Fluids, digestive,78
production, Forms of work, influence of, on efficiency
of body, 555
Fowls

logicalbasis of, 595

comparison of,

with

net

with

energy

for meat

production,451
horse,566

Kellner's,690
limitations of, 691
originof, 689
Wolff's,689
modifications
of, 690
Feeding stuffs,571
ingredientsof, 632
accessory
significanceof, 632
choice of, 703
classes of, 72

requirements for

maintenance

of, 301

production, 488

Feeding as related to individuality,

444
Feeding standards, 689
early,689

swine,

608

energy

values, 596
utilization of, in milk

the

by, compared
digestibility

units, 593

Fruits, 579
Fuel value, 283
Functions

of ash ingredients,187, 190

carbohydrates, 186
fats,186
feed, dual, 183
non-nitrogenous nutrients,187
nutrients,182
597
physiological,

proteins,185
water, 190
Gain

in

fattening,
361

energy

content

of,

733

INDEX

Gain

growth,

in

energy

content

of,

373
dition,
of, in fattening,influence of con438
Gait, influence of, on efficiencyof body,

558
Galactans, 14
Galactolipins,23
Galactose, 9
Gaseous exchange increased by work, 540

through the skin, 139

Gastric juice,82
Gelatinoids,33
Germ
meal, 588

in, 371
composition of, 371
involves storage of ash, 414
of, 375
measure
minimum
of protein for,446
nature
of, 371
values for,390
net energy
of cattle,energy
requirements for, 399
protein requirements for,404
requirements for,401
sheep, energy
protein requirementsfor, 407
swine, effect of insufficient protein
on,

409

requirementsfor,400
protein requirementsfor,408
energy

parotid,79
salivary,79
sublingual,79
submaxillary, 79
Globulins, 33
Glucose
manufacture,

protein requirements for,403

results in practice,403
rate of, 373
at different ages,

by-products of,

587
Glucosids, 10
nitrogenous, 37
Glutelins,33
Gluten feed, 588
meal, 588
Glycogen, 14
computation of gain or loss of, 207
of body, 61
content
conversion of, to dextrose in the liver,
153

formation

gain or
muscle,

of, in liver,153

loss of, 205

of gain of energy

374

in, 378

protein in, 375

storage of ash in, 414
relation of fats to, 421
relation of, to age, 373
relative values of amino

acids for,381

proteins for,381
retention of ash during, 416
retention of protein in, 382
influence

of energy

supply on, 386

protein supply on, 384

substances, 41, 348, 422
total increase in, at different
utilization of energy
in, 390

ages,

397

proteinin, 384, 387, 388

storage, 61
influence

rate

feed in, 381

154

Gums,

Glycoproteins,35
Grade,

continued

"

increase

rate

Glands

Growth,

of,

on

15

efficiency of

Haemoglobin, 135
body, 559
Haemoglobins, 35
bility
Grain, influence of grinding on digestiof, 624
Hay values, 591
Grasses, 573
Heart, 125
measurement
influence of maturity on
of, 221
Heat
composition
energy,
unique, 220
of, 573
of combustion, 223, 228
digestibility
of, 574
Grinding of grain, influence of, on digestibility, outgo of, 235
of increase in, 275
production,causes
624
increased by feed consumption, 273
Gross energy,
635
of feed, 665
influence on, of amount
Group system, 477
crude
fiber,
Growth, 371
676
ash requirements for,414
feed consumption, 651
with fattening,396
contrast
by the horse, 675
centrates,
roughage compared with conrequirements for,399
energy
662
fattening during, 448
losses of energy
feed requirements for,396
in, 650
^

INDEX

734
feeding,influence of,

Heavy

on

450
net

continued
bility, Individuality.
digestimeat
production, 440
"

milk

values, 450

energy

production,514

values, 666
yield of milk, 471
Ingredients of milk, sources
of, 465
Initial and final states, law of, 223
ciency
Intensity of work, influence of, on effiof body, 557
Intercellular substance, 46
Horse :
computation of net energy values for, Intestine,large,85
small, 84
675, 676, 677
nants,
rumiwith
by, compared
Inulin, 14
digestibility
Invertases, 79, 78
604
Investigation,methods
of, 194
requirements for maintenance
energy
of details of metabolism, 194
of, 295
Ionic concentration, maintenance
of, 188
feeding standards
for,566
influence of feed consumption on heat
of, 181
Iron, metabolism
production by, 675
of, 6
occurrence
metabolizable
shelter
for,675
Isolation,
from, 457
energy
for
nance,
mainteprotein requirements of,
in
profitable

production,449

meat

net

energy

Hemicelluloses, 13
digestion of, 92
Hexosans, 12
Hexoses, 8
Hominy feed, 585

Jejunum, 84

329

influence

Humidity,

of,

temperature,

effects

on

of

Juice,intestinal,87
pancreatic, 86

455

Hydrolysisof simple proteins, 164

Hydrogen,

losses of energy

in, 639

Ileum, 84
of a ration, computation
Improvement
of, 698
Income, balance of expenditureand, 194
of energy,
Increase

influence

226

of age

on,

431

composition of,

350,

352,

353,

364
of, 352

content

energy

proteinin, 364

substances

total at different ages,

course

397

of
power,

work on, 540

nucleic acids, 170

441

of lactation,473

phosphorus, 180

digestibility,
609
digestivepower, 441
of feeding,472
economy
efficiencyof body, 556
feed

proteins,162

requirements,

of ammonia

in, 165
nitrogenous end products of, 162
two
stages of, 164
formation

consumption, 443

maintenance

in,249

intermediary, 157
in fasting,251
energy
constancy of, 255
fats,176
non-nitrogenous matter, influence of

Individuality:
feeding as related to, 444
assimilative

katabolized

of carbohydrates, 156

growth,
composition of, 371
content
of, 373
energy

on

262, 265
activity,261
previousfeeding, 253
size of animal, 258
standing and lying, 261
of proteinvariable, 251
temperature,

muscular

in fattening,

354,

fat, 252

external

composition of, 198

influence

on,

of body

influence

Katabolism, 145
computation of per unit of surface,258
to standard
weights, 250
fasting,249
conditions affecting,258
computation of, 282

sulphur,179
442

products of incomplete,230

INDEX

736
Maltase, 79, 87
Maltose, 11
Manifolds, 80
Mannose, 9
Matter

"

of, 202

balance

dry, 3
requirements of,
for fattening,363
growth, 403, 414
meat
production, 445
milk production, 501, 520
work production, 560
Maturity :
definition of, 428
early,428
economic
significanceof, 429
of breed

influence

continued
Metabolism,
general scheme of, 182
influence on, of feed consumption, 651
investigations,comparison of, with
balance experiments, 241
of details of, 194
of ash ingredients,178
calcium, 181
carbohydrates, 152
fats,171
iron, 181
magnesium, 181
nucleic acids, 168
nucleoproteins, 168
organic acids, 159

on,

158
carbohydrates,
pentose
157

pentosans,

phosphorus, 180

444

influence

of,
on
composition of grasses, 573
maize forage, 575
of grasses, 574
digestibility
maize forage, 575
Meat, definition of, 424
fat-free,composition of, 52
proportion of fat in, 425
Meat
production, 424
animal as factor in, 428
combined
growth and fatteningin,448
requirements for,448
energy
factors of, 427
feed requirements for,445
feeding for,444
feeding standards for,451
in, 449
heavy feeding profitable
influence

on,

of age,

430

453

of, 424

in, 426
protein requirements for,445
involved

relation of conformation

to, 443

type to, 443

total amount

of feed for,449

Metabolism, 144
a gradual process,
analytic,146
definition of, 144
enzyms

as

Metabolizable
(See Energy)
energy.
Metaproteins, 35
of, 636
Methane, heat of combustion

influence of amount

of

feed

of, 665
losses of energy
in, 637, 639
production of, in digestion,90,
of investigation,194
Methods
Middlings, buckwheat, 583
wheat, 583

on

duction
pro-

94

Milk:

ash, 460
of, 468
composition of,461
carbohydrates, 460
origin of, 467
of, 459
components
composition of, 461
sources

temperature,

processes

sulphur,179
oxidative,146

average

condition,438
drinking water, 455
exercise,457
external conditions,453
shelter,456
nature

potassium, 181
proteins, 160
sodium, 181

147

agents in, 148

generalconceptionof, 144

influence

on,
of breed, 473

completeness of milking, 480

feed, 527
frequency of milking, 479
stage of lactation,476
animal, 475
variabilityin same
content
of, 511
energy
fat, influence of exercise on yield of,
482
fats,459
originof,466
glands, 462
development of,463

INDEX

continued
Milk,
protein as stimulus to,

Milk

"

influence

of feed

on

in, 528
proteins,459
origin of, 465
secretion of,464
solids, composition
breed
influence

on,

502

percentage

of fat

of, influence
percentage

of production of, 469

of ingredients of, 465

sources

yield of, influence on

of completeness of milking, 480
frequency of milking, 478
individuality,471
stage of lactation,476
Milk
a

production,459
periodicfunction, 476

animal

as

factor in, 470

requirements for, 520

of,468
concurrent
fattening in, 513
diminishing returns from feed in, 515
balances in, 493, 495
energy
requirements for,511
energy
factors of, 469
fat requirements for,516
character

as

stimulus

to, 522

feeding a secondary factor in, 500

feeding for, 500
influence

on,

of environment, 478

exercise,480
fatigue,482
flavoring substances, 522
frequency of milking, 478

individuality,
514
plane of nutrition,514
protein-richfeeds, 506

proteinsupply,

504,

507

shelter,484
stage of lactation,476
483
modifying factors,484
minimum
of fat for, 519
protein for, 501
values for,493, 497
net energy
equivalent fatteningvalues, 498
outgo of ash in, 520
physiology of, 459
protein as stimulus to, 502
requirements for, 501
relative values of proteinsfor, 492
temperature,

"

of

continued

on,

523

of katabolism

in, 515
supply of ash in, 521
utilization of,
in, 493
energy
feed in, 488
protein in,488
estimate of, 489, 491
Milking, completeness of, influence of on
composition, 480
yield,480
frequency of, influence of on composition,
479

yield,478
Milling, by-products of, 582
of, 584
uses
Mineral

ash

feed

of

on

fat in, 529

rate

production,

requirements for, 501

specificeffects of feed
stimulation

474

of feed

737

matter, 5

Molasses, 589
Molasses
feeds,589
Molecular
simplificationin digestion,103
Monosaccharids, 8
composition of, 8
Motion, tissues of, 50
Motor, efficiencyof body as, 544
Mouth, 79
Muscle
extractives,37
fat-free,composition of, 52
mechanical
efficiencyof, 545
Muscles, 50, 531
composition of, 51
structure
of, 50
Muscular
work, nature of, 531
Net

energy

below

critical temperature,

310

Net

values, 271, 278, 634, 659

energy
comparison of feed units with, 596
with metabolizable.energy,
272
computation of, 667, 673, 677
for the horse, 675, 676, 677
from digestiblenutrients,667
organic matter, 673
tion,
independent of chemical composi-

673
importance of, 667
correction of, for crude
determination
of, 272
for cattle,659

fiber,669

different purposes,

279

fattening,360
growth, 390
of embryo, 393
older animals,

393

INDEX

738
energy

"

"

net

production,494, 497
equivalentfattening values,498

ruminants, 660
swine, 661
work

production,563
on,

of age,

666
of feed, 664

amount

breed, 666
heavy feeding, 450
individuality,666
of

kind
method
of

nutrients, 668
digestible

example of,203
Nitrogen factors,69

Nitrogen-freeextract :
compositionof digested,.121
of, 72
of, in feeding stuffs,71
Nitrogen, free,not excreted, 202
Nitrogenous products, excretion of, 140
constituents

determination

matter

influence of work
katabolized

on

in work, nature

of, 540

originof, 160
Non-proteins, 36
determination
of, in feeding stuffs,69
digestion of, 96, 100
generalpropertiesof, 36
of,36
groups
indirect utilization of, 622
622
influence of, on digestibility,
factors
for, 70
nitrogen
occurrence
of, 36
value of, 324, 684
for maintenance, 324
Nucleoproteins, 34
metabolism
of, 168
Nucleus
of cells,
42
Nucleic acids,34
digestionof, 99
t

Nutrients

digestible,599

computation of, 598

of metabolizable

energy

from, 646

as

measure

ratio,600

Oat

hulls,584
Offal,composition of, 55
proportion of body ash in, 56
fat in, 56
proteinin, 56
Oil extraction,by-products of, 586
Oil meals, 587
seeds, 581
fats and,
Oils,distinction between

19

ethereal,40
80

Organic acids,production of

of, 542

of urine, 159

216

energy,

effect,live weight
of, 196

total,195
Nutritive

Omasum,

katabolism

from, 667
of, 648

values of, 668

energy
significanceof, 600
functions of, 182
mutual
replacement of, 270
non-nitrogenous,
effect of deficiencyof, 320, 324
surplus of, 321, 324
functions of, 187
functions of, 597
physiological
solution of, in digestion,101
Nutrition, balance of, 192, 201
Nutritive

and
ing,
fattenrelative,for maintenance
361
of, 189, 335
Neutrality, maintenance
Nitrogen balance, 202
determination of, 203

energy

net

includes

production,666

of determination, 271

Non-nitrogenous

values

energy

metabolizable

milk

influence

continued

Nutrients,

continued
values,
suckling animals, 391
maintenance, 271

Net

in digestion,

90

Organic matter, 4
able
digestible,computation of metabolizfrom, 648
energy
values from, 673
net energy
subdivision
of, 4
maintenance
of, 188
Osmotic
pressure,
335

Outgo of chemical
in cutaneous

energy,

229

excretion, 231

feces,230
urine, 231
heat, 235
kinetic energy,

work,

235

235

efficiencyof body, 562

Oxygen :
absorption of,by blood, 13s
through skin, 139
consumption of, in metabolism, 146
determination
of, 208
supply of, 132
Oxyhemoglobin, 136
Over-all

INDEX

Packing house, by-productsof, 590

Pancreas, 86
Parotid glands, 79
Passage of feed from stomach, 83
Paunch, 80
Pectins, 15
Pentosans, 15
digestionof, 91
fermentation of,91
metabolism of, 158
proportionof, in crude
Pentoses, 9
metabolism of, 158
Pepsin,79, 82, 96

Peptids,30,
Peptones,
Period

"

continued

determination

of,630
of feeding stuffs,630 ,634, 678
Prolamins, 33
Proteans, 35
Proteases, 79, 86, 87
:

of, influence of, on

bility,
digesti-

622

of energy,
318, 542
to milk glands, 502
body, fluctuations of, 319
as

fiber,71

35

35

system, 477

source

stimulus

proportion of, in bone, 48

offal,56
of diminished
digestibilityof,
cause
619
cleavage, extent of, in digestion,98
computation of gain or loss of, 204
consumed
by calves,404
lambs, 407
crude, determination
of, in feeding
stuffs,68
digestibility
of, 119
functions of,in fasting,255

production,543, 560

work

gain of,in fattening,354

loss of, 202
or
in dairy rations,506
fatteningrations,364
increase in fattening,364
of rations,
influence of,on digestibility
33i

insufficient,effect of,
swine, 409
katabolism,

on

growth

of

dependent on supply,322
in fasting,251

305

milk production,514

Plasma, blood,

125

Polypeptids,31
n
Polysaccharids,
chemical structure

terminology of, 12
Potassium, metabolism
of, 6
occurrence

of, 181

of body with,
S67
of, compared with body, 567
efficiency
Practical feeding trials,
592
Precipitation,shelter from, 456
animal

Production

as, 192

values,
regards protein,678
634
energy,
definition of,630

as

3B

variable,251
normally small, 251
in work, influence of non-nitrogenous
nutrients on, 538
influence on, of feed supply,316
,

of, 1

plant,comparison

motor,

values,

addition

respiration
apparatus, 212
Phosphatids, 22, 23
Phospholipins, 22
Phosphoproteins, 35
Phosphorus :
anabolism
of, 180
digestionof, 101
forms of, 7, 180, 421
inorganic,value of, 421
katabolism
of, 180
metabolism
of, 180
occurrence
of, 7
Pigs:
requirementsof, 400
energy
feeding standards for,412
gains by, in growth, 401
proteinrequirements of, 411
Plane of nutrition,influence of,on energy
requirements for maintenance,

Prime

Production

Protein

Pettenkofer

Power

739

work, 536
of,
for growth, 446
maintenance, 316,

minimum

milk

323

production,501

nitrogen factors for,69

nutrition,plane of, 324
of feed, storage of, 319
optimum of,for maintenance, 323,
minimum
of, 254
physiological
values
as
regards,
678
production
in
increase
of
growth,
of,
rate
375
retention of, in growth, 382

330

INDEX

74o
Protein,

"

continued

influence

of energy

on,

Proteins,

protein supply, 384

requirements,
computation of, to unit weight, 325
for fattening,363, 416, 446
growth, 403
of cattle,404
sheep, 407
swine, 408
results in practice,403
maintenance, 313, 323
nature
of, 313
meat
production, 445
milk production, 501
work production, 561
of cattle,326, 367, 404, 501
horses, 329, 561
sheep, 327, 365, 407
swine, 329, 368, 408
relation of, to age, 445
rich feeds, influence of, on
milk production,
506
supply, influence of, on digestibility,

originof, 465
physical propertiesof, 25
putrefaction of, 99
relative values of, 678
for growth, 381
maintenance, 315
milk production, 492
simple, 25, 26
anabolism
of, 160
classification of, 32
cleavage products of, 28
composition of, 26
deaminization
of, 165
reversible,166
formation
of,
ammonia

in katabolism
of, 165
carbohydrates from, 167
fat from, 168
hydrolysisof, 28, 164
katabolism
of, 162
metabolism
of, 160
olism
nitrogenous end products of katabof, 162
non-nitrogenous residue of, 163
proportions of cleavage products in,

"

447
_

milk

production, 504, 507

retention in growth, 384
surplus,katabolized, 317, 488
of, in feeding
true, determination
stuffs,68
for fat production, 363
unnecessary
utilization of, in fattening,364
growth, 384, 387, 388
milk production,488, 491
limited,318

Proteins,24
alfalfa,values of,683
cereal,values of, 683
chemical
changes in resorptionof, 107
coagulated, 35
coagulation of,26
conjugated, 25, 34
derived, 25, 35
primary, 35
secondary, 35
digestion of, 83, 88, 95
by erepsin,98
"pepsin,96

trypsin,97
formation

of fat

functions

of, 185

continued

"

supply,386

31

of, 27
synthesis of, 30
from digestiveproducts, 160
two
stages in katabolism of, 164
unbalanced, 679
Proteoses, 35
Protoplasm, 42
composition of, 44
Ptyalin, 78, 79
conditions of action of, 92
Pulmonary
exchange, investigationof,
structure

214

Putrefaction

of

proteins,99

Pylorus,83
Quantity of feed, influence

incomplete,679

tibility,
diges-

Raffinose, n
of growth, 373

Rate

at different ages,

from, 173

of, on

613
Quotient,respiratory,207

Rations

374

compounding of, 707

maize, low value of, 681

nomenclature
of, 24

computation of, 565, 689, 697

from given feeding stuffs,700
improvement of, 698

of milk, 459

for work

production,calculation of,565

INDEX

Rectum, 85
Regnault-Reiset

respirationapparatus,

209

values

Relative

feeding stuffs, 591,

of

597

Requirements :
for fattening,350, 359, 361, 363
growth, 396, 399, 403, 414
maintenance, 269, 280, 313, 332
production, 444, 445, 448
meat
milk production, 500, 501, 511, 520
work production, 560, 562
of ash,
for growth, 414
maintenance, 332

production,520
production,562

milk
work

dry matter, 696

for fattening,361
energy
growth, 399
maintenance, 280, 303
meat
production, 448
milk production, 511
work
production, 562, 564
feed, 691, 693, 694
fat for milk production, 516
363
proteinfor fattening,
growth, 403
maintenance, 313, 323
production, 445
meat
milk production, 501
work
production, 561
Residue, non-nitrogenous, of simple proteins,
163
Resorption,
chemical
mechanism

105

changes in, 107

of, 106

Respiration,123,
apparatus,

132

influence of work

polish,583

456
precipitation,

457

wind, 456
of, on

209

on,

536

regulation of, 137

Respiratoryquotient, 207
Reticulum, 80
Reversible reactions,150, 166
of metabolic
reactions,152,
Reversibility
9

604
swine, 606
requirements for growth, 401
energy
maintenance, 292
influence of feed consumption on heat
production by, 653
pared
maintenance
requirements of, comwith cattle,294
proteinrequirements for growth, 407

production, 456
production,484
fasting
Size of animal, influence of, on
katabolism, 258
of ash ingredients,
Skeleton as reserve
influence

of tissues,136

Rice bran, 583

Saliva, 79
action of, in stomach, 73
starch, 92
on
Saponificationof fats in digestion,95
Schematic
body, 195
Scleroproteins,34
Sheep :
by, compared with horse,
digestibility

sun,

208

iS3

696
in,
proportion of vegetative organs
576
Rumen, 80
Ruminants, digestibilityby, compared
with horses,604
swine, 606
speciesof, 603
values for,660
net energy
Rumination, 81
Rye bran, 582

Shelter from

in, 106

Pettenkofer, 212
Regnault-Reiset,
calorimeters,236

Rhamnose,

Roots, 73, 579

618
influence of, on digestibility,
Roughage, 72, 572
comparison of, with concentrates, 662
general character of, 572
influence of cutting on digestibility
of,
624
proportion of, to concentrates, 451,

maintenance, 456

paths of, 107

role of osmosis

741

meat

milk

338
exchange through, 139
Skin, gaseous
Slaughter tests, comparative, 199, 351
of, 181
Sodium, metabolism
6
of,
occurrence
Solution of nutrients in digestion,101
603
Species,influence of,on digestibility,
of ruminants, digestibility
by, 603
dynamic action, 275
Specific

INDEX

742
effects
Specific
associated
on

on

fats,527

nucleic acids,169

production,523

milk

influence

Speed,

Synthesis
of hippuricacid,163

of feeds,448

with

of,

on

160

body, 552, 557

expenditure in locomotion,

of, 161

seat

energy

Standing

digestive products,

simple proteinsfrom

efficiency of

uric acid, 171

552, 557
and
lying, influence

of,

on

in the

Syntheticprocesses

body, 146

fastingkatabolism, 261
Starch, 13
digestionof, 79, 88, 92
in intestines,
94
stomach, 93
fermentation
of, in digestion,94
manufacture, by-products of, 587
values, 672
Steapsin, 79, 86
Stomach, 79
of hog, 81
horse, 81
ruminants, 80
sheep, 80
of feed from, 83
passage
Straw, 577
values for
Suckling animals, net energy
growth of, 391
Sucrase, 79, 87
Sucrose, 10
Sugar beet pulp, 589
Sugar manufacture, by-products of, 588
Sulphur,
digestionof, 101
katabolism
of, 179
metabolism
of, 179
occurrence
of, 7
Sun, shelter from, 457
Sundry ingredients of animals,

chemical
regulation of, 263
physical regulation of, 262
critical,
264, 453
lowered
by feed consumption, 308
effects of extremes
of, 266
external,influence of age, on effects of,

body,

454

humidity, on
live weight

effects of, 454

fastingkatabolism,

influence of, on

262, 265

requirements for

energy

nance,
mainte-

304

production,453

meat

milk

of

production, 483
modifying factors,484
drinking water, influence

of,

on

production,455
requirements for

meat

Tissues

animal,

39

45

classification of, 45

per

unit

connective, 49
elastic,49

epidermal,57
composition of, 57
of, 57

functions

digestibility
by, compared

on

on

effects of, 455

effects of, 454

maintenance, 308
Tendon, 49
Tissue, adipose, 58
compositionof, 59

of, 258
Surplus feed, disposalof, 350
with

fowls,

608

ruminants, 606
effect of insufficient proteinon

growth

of, 409
requirements for growth, 400
maintenance, 285
influence of feed consumption on heat
production by, 653
values for,661
net energy
protein requirements for growth, 408
maintenance, 329
energy

of ration

amount

relation of, to energy

plants,40
Surface, computation of,
computation of katabolism

Swine

Tankage, 590
Temperature,

of alimentation, 54
chemical
compositionof,55

motion, 50
58
supporting,46

reserve,

Tonus,

534

Total

feed

required, computation

of,

697
Training, influence
body, 556

of,

on

efficiencyof

Triglycerids,
17, 19
elementary compositionof,20

*HE

following pages

books

by

the

same

contain
author

or

advertisements
on

kindred

of

subjects

of Feeding
Farm
Principles
By

SLEETER

Associate in Animal

Animals

BULL

Nutrition, University of Illinois

$1.75
illustrated,

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in stock

general,elementary
feeding.
underlying the art of feeding animals have been
will not only be suitable
that the book
presentedin such a manner
for collegecourses
in general feeding,but will also
for use
text
a
as
who
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be valuable to the farmer
education
in
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agriculture.
The

course

scientific facts

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author

The

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presented rather
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binations
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discussion

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and
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the author
Chemical Society,Washington, D.C.
Journal American
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The

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By

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AgriculturalExperiment

volume

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general principlesof bio-

practicalside of feedinganimals
relate specifically
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general reader will also find the

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Not
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Scientific
Feedingof Animals

The

Translation

Authorized

WILLIAM

KELLNER

0.

Professor

By

by

B.Sc,

GOODWIN,

Ph.D.

AgriculturalChemistry, and Head of the Chemical Department,

South-Eastern Agricultural
College (Universityof London), Wye, Kent.

Lecturer

on

$1.75

Cloth,i2mo,

An

English translation

authorized

of Dr.

generallaws

feeding of animals

underlie the

which

scientific foundations

work

explainsin simple language

It

Kellner.

O.

of the valuable

which

upon

the

the
the

and

of
principles

animal

valuable

books

Animals.

The

nutrition rest.

"I wish

that it is

to say

English language

in the

is

author

He

planned

Feeding

on

in

extremely lucid

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covers

such

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of the most

one

concise

expressionand

his field in

will

Farm

give

the

that

manner

reader

most

is well

excellent

knowledge of the generalprinciplesof feeding."

"

Kellner's

"Dr.

in this
to

be

subjectis

needed,
have

to

P.

Ohio

S. Plumb,

Charles

standing as
too high for
I feel

and

done

sure

service in

any

State
student
words

and

investigator

of commendation

that the translator

rendering this

work

"

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and

lisher
pub-

available
Henry

Breedingof Animals

The

By
Professor of Animal

B.

F.

Husbandry,

of the College of Agriculture, and

Dean

of the Experiment

deals first with

text-book

methods
The

animal

notable

achievements

in modern

how

of the

the constitution
to

heritanc
in-

ment
develop-

of the most

one

through man's

be

been

representedin

thus transmitted

and

germplasm

the

qualitieshave
to

come

breeding.

from

parent

is a subjectof great scientific and practical

interest.
offspring,

The

principleswhich

scientific

all be

breeding may

development. The
which

of animal

of the cell and

described and
The

The

breeding.

practice of

animal

inheritance,reproduction and

those scientificprinemphasizes particularly

ciples

text

the basis of

recognizedas

are

the

govern

classified under

well established
been sufficiently

of

is

man

valuable

qualitieshave

these

to

agriculture.How,

these highly specializedand

efforts,

acquired,and

animal

resultingin

qualitiesuseful
highly specialized

of

questionsof

peculiar to

are

of the domestic

improvement

pages, $1.75

animals, but emphasizes the principles,

practiceswhich

and

illustrated,
304

the fundamental

plants and

to

common

Director

Station of the University of Missouri.

i2tno,

This

M.S.

MUMFORD,

to

afford

physicalbasis

of

heredityand which have

real basis for the

practice

heredityin the germplasm

changes resultingin transmission

significant

the

are

illustrated.

to the practice
physiologyof reproductionand its applications

breeding is the subjectof

of

art

important chapter. The

development which

heredity and

of the breeder's

an

are

discussed.

The

constitute

the real basis

practicalquestionsof

and
sterility,
crossbreeding,
grading,fertility,
in the

lightof

the

most

modern

sex

development

of

science.

THE
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