THE

AMERICAN

Vol.21,

JOURNAL

No.

11,

or

Printed

Fat,

CLINICAL

November,

1968,
in

NUTRITION
pp.

1291-1301

U.S.A.

Carbohydrate,

Salt,
Further

M.

WALTER

BORTZ,

HE

INCONSISTENT

source

of

plateaus

are

M.D.,3

PAULA

L.

number

be

frequent

1)055i1)ilities.

activity

l)atterns

(2)
time

number

of

trigued

with

stituencv

sil)ie.

time idea
tlailv

(7)

calories

recently

1)erformed

weie

exclusively

in

given

time

amount

all

diets

of

considered

it

observed
carbohydrate

tlifferences
periods

gluconeogenic
or

were

mecimanisms.

providing
From
pital,

time

Division

of

b-

National

I)ivision

Department

Assistant,
of

of

Division

Division

of

of

Research.

(liets

one

were

other

initial

nitro-

a severe
felt

it was
to

study

isitim

Imigimer

study
(7).

was

Time

nearly

nine

sub-

years
lb. at

all were
free from
conmfrom
two
3-hr passes/
confined
to the Metanot

than

diet

All
diet

day.

permitted

time usual

(9)

The

any

physi-

sedentary

this

fed

for

ward

stabilization

were

fat

(to

35%

Follow-

timat was

time subjects

total

studies

to

40

were

made

exclusion

4 g NaCl

calories

reduced
the

high

until

constant.

calories
time

were

percentage
in
was contilmued

containing

weeks,

for-

protein/

carbohydrate.

formula

of 2-3

carbohydrate

Re-

1291

and

usually

In
fat

and
Director,

calories

calories)

120

re-

relatively

Researcim.

g or

duced
to 800 kcal for the rest of time study.
six subjects,
in the first hypocaloric
period,
their
period

Health

80

to maintain
iim a liquid

were

and

Hos-

given

providing

a period

designed

were

remaining

ing

one

was

feedings

of time total

fat

Research,

They

activity

mula

and
to

Formerly

Medicine.

its

with

of the

previous

project;
Aside
subjects
were

the

The

by the

of

time

Ward.

riods,

of

time

MENTAL

of the

weight.

the

lmigh

Lankenau

Institutes

cal

We

activated

Research,

be-

in

activities.

study

Pennsylvania.

AM-08887.
Research
.-ssociate,

search

that

coupled
in some
way to timese
It also
seemed
possible
timat
lmiglmer levels
of dietary
protein

Philadelphia,

Assistant,

of

some

protein,

conserve

of

disease.

bolic

and

between
time fat
were
due
directly
low

Supported
Grant

In

start

week,

carbohy-

g protein/day.

nmeclmanisms

noncarboimydrate,
diet

respon-

absence

that

to

we

(8),

(five mets,
four
women)
were
19-34
all but two weigimed
more
timan 300

the

possible

addition

others

confronted

protocol

to

jects
old;
plicating

and

general

identical

Metabolic

the

salt.

40

content.

The

in-

isocalorically

contained

protein

a tietailed

added

body

wimen

EXPERt

time conbe

or

time

observations

reported

presence

of

In

as imave

content

on

fat

effects.

similar

been

may

interchanged

such

interested,

of

and

varying

iim mvlmiclm dietary

Ward
drate

Imave

that

AND

Smith
salt;

(3-6)

are

physical

time effect

dietary

workers

have

\\Te

simown

of

M.P.H.,4

calorie
restriction.
Timerefore,
we
of interest
to extenti
our earlier

in

influential;

imave

level

of

study

are

offset

gen

most

timere

anti

nmay
capacity

are
fac-

the

However,

Hormonal
(1)

Drenick

cimanging

is

B.S.,

PH.D.5

came

variance

calories

explanation.

otimer
almd

daily

is a

and
of

HOWAT,

HOLMES,

Weight

observed
a number

responsible.
of

loss

dieters.

Loss

in proportion
calorie
weigimt

Following

time

to

diet.
This
loss became
control

pe-

were
placed
on 24-day
diets
60%
protein;
time remaining
up of fat or carbolmydrate
to
of

and

time

other.

3 g KC1

1mm seven

were

used;

of

time

in

time

Downloaded from www.ajcn.org by on December 8, 2007

could

time

weigimt

to

frequently
Obviously

discouragiimg.

tors

of

RATE

confusion

Weight

Studies12

WILLIAM

and

1292

Bortz

et al.

6TE

CHO
Diel

4400

Calories

800
Ssb1DR

663

FAT

169

PRO

80

52
30
80

20
0

0
54

120
0

80

80

80

Coories

445

Diet
gin

435

800

3300

CHO
FAT
PRO

469
32

54

54
80

80

80

SC

304

425
294
415
284
405

274

395

385

264

375
254
365

244

355
0

20

30

40

50

60

70

80

90

20

30

40

50

60

70

80

90

Oars

Downloaded from www.ajcn.org by on December 8, 2007

Subj

Days

N S

800

3300

Calories

.ASCC

V
CHO
Diet

FAT

469
1 3

PRO

80

4000

Calories
20

120

54
80

0
80

80

800

CHO 560
FAT
56
PRO
80

120
0
80

54
80

0
54
80

120
0
80

340
450

330

440

430

320
-5
A

420

310

410
300
400

290

390

380

280
0

20

30

40

50

60

70

80

Dajs

FIGS.

otimer
and

studies

no

chenmical

100

per

total

dients

of

casein,6

sucrose,

Artificial
protein

loss

potassium

was

diets

these

sodium

and

feedimmg.

diet

180

mg
basic

purified
corn

on

800-kcal

added,

showed

The

were

cornstarclm,

content.
before

Every
analyzed

The
the

6 days
for

6 Supplied
tories,

mg

weight

ap-

and

Evansville,

free
as

subjects

were

blood

fatty
Casec
Indiana

acids
by

Mead

was

butter.

Johnson

and

were

(13),
ammonia

milled.

check
riodic

or

(12).
made

and

sodium
mmitrogen

Urinary

dioxide
quotients

KC1.

Correspolmding
measured

(14),

levels

and
(17)

urine
for

urinary

potassium

of urea

(15).

mmitrogen

were

was

creatinine

on completeness
measurements

carbon
piratory

weighed

NaCI

cholesterol

collections

In five experiments,

collected

(10),

(1 1), and

added

nitrogen

first feeding.
venous

without

ingre-

powdered

oil,

diets

potas-

flavorings
were
added
to improve
The
diets were analyzed
chemically
for

taste.

daily

or
of

daily

the

Daily

sodium

analysis

proximriately
siunm

1-4.

Days

also

(16)
deter-

nmeasured

as

of collection
(18).
of oxygen
consumption

production
determined

were
made
(19).

and

Peand
res-

and
RESULTS

triglycerides
Labora-

Figures
terations

1-4

in

slmow

dietary

weigimt

response

carbolmydrate

to

anti

al-

fat

Fat,
in

time

absence

firmation
paper,

of

rate

of
the
initial

after

of

weight

witim time
of calories.
Figures

was

essentially

5-1 1 show
of

Whereas

time results
timere was
following

boimydrate
loss

the

and

80-g

of

protein

time

study,

study

anti

(Figs.

except

that

In
ceiving
Time

are

supplements

on
fat

those,

weight

fat

generally

initial

It

as

previously
self-limited,

all

comes

offset

homeostatic
nutritional
sine-wave
sodium
dietary

time

by

shown.

object

periods

that

be

Figures

19-22

in

were
an

mechanisms
One
can
extending

data,
changes.

which
beChanges

the

subjects

to

added.

amazing

to

bear
sodium.

All

time

nitrogen

whose

quotient

stabilization

diet

diet,

was
of

was

035;

and

all

0.85;

on

the

bal-

no

demon-

(22),
(23).

are

and
the

we

critical

body
wahormonal

composition

reported
of

and

the

results

of

obese

subjects

in

carbohydrate

were

switched

firm

of our

those

tlmis study

cltmsion
weight

earlier

did

that
loss

report

not

affect

was due
capacity

bohydrate.

et

Katz

carbohydrate

diet

fasting.

regardless

and

bring

experiments
protein
content
the

prior

con-

any difference
in time rate
between
fat and
carbohydrate-

containing
diets
and fluid-retaining

in

ni-

body

shown

of

(20),
(21),

fat-carbohydrate
interclmange
to a total of 22. The higher

drate

urinary

are

witimin
an 800-kcal
formula
data
of the present
study
con-

caloevi-

of

diet

a group

and

despite
marked
was
particularly

analyses

in

fat

protein

high

It imas been

of

and

(7)

that

stay

study.

to serum
lipids
electrolyte
balance

near

to

carbohydrate
events

isocaioricaliy
diet.
The

of

periods.
definitive

of intense

which

to

diet

subjects

capacity

nitrogen
equilibrium
rie restriction.
This
during

high

clmoiesterol

respiratory

modifications

studies

effect.
curve,

represent

those

minerals

serum

of dietary
of metabolic

Recently

as other

Body
weigimt
cimanges
seem
to be related
time urinary
sodium
fluctuations.

More

time

roles
a host

levels
weiglmt

em-

observed,

in urinary
potassium
appear
rougim
correlation
with
urinary

dent

on

importance
ter and

loss,
Conperiods

are

a crude

are
diet

should

changes

through

strateti

average

The
in

fat

period.

imowever,

sodi tim-sensitive
overcome
the

ances

the

time carbohydrate
diet, 0.73.

nitrogen,
subjects
re-

carbohydrate

retention.

these

fatty

the

are

by sodium

mixed-diet
early

sodium

imagine

present
to

sodium,
in the

of time high

pimasized,
that

the

in

parts

the

simow

in

free

on

low.

subjects

our

higher

DISCUSSION

12-18
the
balances

time

versely,

in

comparable

mineral
first

iim

subject

time fluctuations

characterized

as

ni-

studies.

show

triglycerides

degree.

Figures
potassium

and

been

time results

4-6)

of a lesser

are

imad

higimer

al.

was
the

Our

as

stutiies,

of

time

dietary

carbohydrate

sodium

retention.

to the
sodiumof dietary
car-

(24)

have

effective

reversal

of

salt

however,

level

of

reported

as

carbohy-

loss

due

indicate

protein

appears
This

of

confirms

to
that

intake

to

mediate
an

earlier

Downloaded from www.ajcn.org by on December 8, 2007

earlier

CC

subjects

nitrogen
time initial

was

diet

values

formulas

The

Subject

five

diet,
althougim
the latter
difference
is
statistically
significant
(Table
mm). On

both

diet.

There

lower

extremely

fat

in

during
time fat
periods
carbohydrate
periods
(see
12-22),
particularly
in the

formula

lipid
and

fat
not

rapid

of

period.

Serum
acids

made

all,
total
urinary
highest
during

trogen
excretion
than
during
the
bar graphs,
Figs.

to

1293

were

In
was

i).

hypocaloric

subjects

period

content

linear

similar

reinstitution

Loss

source

a period
of weight
stabiintroduction
of time car-

upon

Weight

(Table
excretion

to the

seven

and
trogen

con-

potassium.

grossly

of

formula

weigimt

the

and

are

six

In

response

sodium

in

Salt,

previous
loss, the

of

weight

added

above,

studied
lization

sodium.
of our
weight

independent

presence
those

added
results
rapid

loss

and

Carbohydrate,

Bortz

1294

et al.

0
0

03

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to

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Fat,

Carbohydrate,

Salt,

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Weight

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1295

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al.
et
Bortz
1296

is

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Ii
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Fat,

Carboimydrate,

Salt,

and

Weight

Loss

1297

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&

S.tt DR
NITROGEN
80gm

BALANCE

psiem

NOIS7C/ci-KC/

3300

Calories

Stab

gm

19

Days

SubtCC

NITROGEN
80gm

BALANCE

pem

ADA6C/o,

KC/

Calories

4000

Stab.

CHO

High

fat

High CHO

-I

Sulo:TE

BALANCE

10

20

30

40

50

20

80gmpem

44004-

800

21

70

80

90

100

hO

-+3700

NS

Si4

Days

60

Days

NITROGEN
80

BALANCE

gm prolem
KC/

No4loOo-

V#{176}#{176}

Catones

4i

--

-6:
4

N2_
qm
C
F
-2

j
20

1OC.lO

30

..
403.50.i60

FIGS.

the

19-22.

equilibriunm

Effect

of

between

dietary
nitrogen

fat

and
intake

carbohydrate
and

7O

38O01l39O.

Days

22

urinary

interchange
excretion.
1298

on

urinary

nitrogen.

The

x-axis

represents

COO

Downloaded from www.ajcn.org by on December 8, 2007

-4

Calories

High

-I
flr

NITROGEN

tot

High

Fat,
report

of

specificity
In sum:

Bloom

(21)

Carbohydrate,
that

of carbohydrate
essentially,
over

indicated

and

our

excretion
the figure
derives
gram
80-,

earlier

values
635

and

take
not

on

relate

timat

by

during

all

nitrogen

al.

N5

Fat

in

day

in

their
periods

to

the

different

7.4

141.0

of

the

8.2

blood

the

dietary

are

the

number

by

each

III

excretion

intake

of 24-

value.

and

as

related

fat-carboimydrate

interchange

40-g

N,
hr

monia

N, g/24
hr

Coeatinine
g/24

N,
hr

Un-

protein

diet

Urinary
g/24

N,

studies

periods

80-g

account-

periods

protein

Fat

ed for N,
g/24 hr

diet

and

periods

protein

Fat

6.3

8.5

6.3

6.2

12 . 5

13.9

12 .6

11 .3

18.9

19.1

18.9

18.5

studies

periods

Carbohydrate

(80-g

diet

studies

periods

Carbohydrate

periods

inData

Stabilization

11.1

8.88

Mixed

17.2

12.93

Fat

14.0

9.66

1 .65

0.71

1 .98

Carbohy-

12.6

8.20

0.96

0.58

2.86

0.82

0.71

0.69

1 .06

0.74

2.47

represent

CC,

and
(120-g

during

the

tiespite
subjects

pro-

gen

Stabilization

16.1112.16

Mixed

19. 7215

Fat

l9.0714.92
l8.53l4.64

Carbohy-

.06

represent
for

the

averages
respective

1-2
in

balance.

lates

on
to

time

1.19

0.69

2.78

of

1 .25

0.51

2.39

g/day

for

humans

1 .34

0.54

2.01

stores

are

extremely

of
dietary

values
periods.

of urine

brain

which

cannot

sustain

During

our

received

no

fat

is

diet

said

any
fat-diet
dietary

our

negative

nitro-

of

this

ob-

particularly

glucose
(26).

inthat

restriction

slight
significance

the

2.21

fecal

it is apparent

calorie

obligate

0.59

urinary

collections

additional

only

Time

all

6-day
periods.

g/day,

marked

were

1.15

drate
data

the

servation,

Diet

deitary

an

of

of

1 to

in

appropriate

loss

PP

averages

obtained

Acknowledging

GS,

intake)

the

values

trogen

drate

The

118.4
18.4

errors.

nitrogen

protein

nitrogen

collections

ml

136.4
11.8

during

represented

urinary

averages

6 days

TABLE
Total

take)

tein

99.4

parentheses

their

Diet

Subjects

the

standard

in

dietary

been

(25)

and

l20-g

protein

(14)

0.09
0.68

0.06

every

Numbers

Fat
Am-

Urea
g/24

hr

NW

Cholesterol,
mg/100

aol

N Intake,
g/24 hr

Total

VB

0.92

represent

obtained

periods

ap-

difference

data

samples

Carbohydrate

Subjects

(12)

Carbohydrate

periods

N, g/24

Triglycerides,
mg/100

onEq/

excretion

Urinary

levels

senmistarvation.

TABLE
Urinary

the

inter-

lipid
FFA,

The

carbo-

previously

et

on

______________dietary

serum

liter

uses
anti

balance

has

Keys

experiments

fat-carbohydrate
on

Diet

respectively.

but

Tlmis

demonstrated
classic

higher,

striking.

dietary

these

intake,
one
1 g nitrogen

nitrogen

the

of

change

present

equilibrium.
During
the fat-diet
there
was a slightly
more
negative
balance
on the lower
protein
inthan

Effect

II

to
Body

limited

prolonged
periods,
carbohydrate

re-

requirement

be

about

200

carbohydrate
anti

obviously

glucose
time
at

need.
subjects
all;

yet

Downloaded from www.ajcn.org by on December 8, 2007

proaches
periods
nitrogen

1299

TABLE

ex-

the

studies,

then,

periods,

Loss

12.6, and 18.9 as time

intake
for the 40-,

protein

It is apparent,

Weight

time

nitrogen

To

to protein
g protein/

120-g

hytirate

is

one.

the figures
6.3,
per day nitrogen

and

in this
regard.
a long
period
of

weight
loss, a calorie
is a calorie.
Table
iii
illustrates
urimmary
cretion
figures
derived
from
study

Salt,

Bortz

1300
their
by

nitrogen
only

tary

excretion

a small

exceeded

amount.

conditions,

intake

Under

therefore,

these

one

must

die-

periods.
the 80-g

con-

distinctly

dude
either
that brain
and other
glucosedependent
organs
do not require
as much
glucose
as proposed,
or that
glucose
is be

ing
formed
dated.
Cahill

by processes
et al. (27)

as
have

yet unelucishown
that

potential
gluconeogenic
substrates
timan protein
(i.e., glycerol,
pyruvate),
erated
during
calorie
privation,
provide
sufficient
substrate
carbon
supposed
et

glucose

al.

(28)

duction

have

reported

content
for
this

glucose

from

gives
process

production

glucose-14C

Wrenshall
that

as calculated

other
gencannot
for the

glucose

the

pro-

urinary

ni-

erroneously
low
when
compared
as

estimated

turnover

data.

the

suggest

time possibility
of glucose
formation
from
another
major
source,
which
by implication is body
fat. Alternatively,
Owen
et al.
(29) have
documented
ketone
bodies
by human
vation.
A study
in vitro
on brains
demonstrate
bodies
ter

the
oxidation
of
brain
during
starin our laboratory

of starved
rats,
preferential
over

a period
Whatever

glucose

by

of ketosis
the

however,
use
of
time brain,

did not
ketone
even

reconciliation

of

nitrogen.

Under

time

mi-

conditions

total
shown

starvation
Owen
et
that
ammonia
is the

nous

excretory

sents
tions

a byproduct
that
yield

skeletons
resents

product.

creased
organic
ing
starvation

specimens)
the

was

higher

sodium
part
of
implicated
vated

less

in the

higher

stud-

protein

by

marked

in

diets.

in-

quantitative
random
urine
subjects

on

Whether

bear
a
unknown.

cause
The

these

and
effect
increased

excretion
noted
during
the early
the fat-tiiet
period
has also
been
(32) as a cation
cover
for ele-

organic

acids.
SUMMARY

Eleven
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were
perfornmed
on nine
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Fat and carboimydrate
were
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in an 800-kcal
constant-protein

diet.

loss

No

was

difference

noted

apart

to alterations

in

dynamics.
ably

low

on

in
from

rate

and

fluid-balance
was

regimens.

those
a lower

weight

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all

of

that

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Nitrogen

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Ammonia

1.

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