UNITS, SYMBOLS ,

AND TERMINOLOG Y
FOR PLAN T PHYSIOLOG Y

This page intentionally left blank

UNITS, SYMBOLS ,
AND TERMINOLOGY
FOR PLAN T PHYSIOLOG Y
A Reference for Presentation of
Research Results in the Plant Sciences
Sponsored b y the International Association fo r Plant Physiology

Frank B. Salisbury , Edito r

Utah State University

New York Oxfor d

Oxford Universit y Press
1996

Oxford Universit y Pres s

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Copyright 199 6 b y Oxford Universit y Press, Inc .

Published b y Oxfor d Universit y Press, Inc .
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Oxford i s a registered trademark of Oxford Universit y Press
All right s reserved. No part o f thi s publication may be reproduced ,
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electronic, mechanical , photocopying , recording , o r otherwise ,
without th e prior permissio n o f Oxford Universit y Press .
Library o f Congress Cataloging-in-Publication Dat a
Units, symbols, an d terminology fo r plant physiology : a reference fo r
presentation o f research result s in the plant sciences / sponsore d b y the
International Association for Plan t Physiolog y ; Frank B . Salisbury, editor .
p. cm .
Includes bibliographical reference s an d index.
ISBN 0-19-509445- X
1. Plan t physiologyTerminology . 2 . BotanyTerminology .
3. Technical writing . I. Salisbury, Fran k B.
II. International Association fo r Plan t Physiology .
QK710.5.U55 199 6
581.1'014dc20 95-5059
3

9 8 7 6 5 4 3 2 1
Printed i n th e Unite d State s o f America
on acid-fre e pape r

CONTENTS
CONTRIBUTORS vii

PREFACE

ix

SECTION L THE BASICS

1. Summary of the International System of Units (SI Units) Salisbury

Quantities and Units 4
Le Systeme International D'Unites (SI) 5
The SI Tables 6
Some Special Considerations 1

2. Rules for Botanical Nomenclature McNeill & Barkworth 2

Documentation 2
Taxonomic Groups (Taxa; singular: Taxon) : Som e Rules of Nomenclature
Form of Scientific Names 2
Special Situation s 2

1
1
22
2
4

3. Statistics Sisson 2
General terms: 2
Measures of Central Tendency 2
Variability 2
Confidence Intervals 2
Test of Hypothesis 3
Regression Analysi s 3
Analysis of Variance 3
Covariance Analysis 3
Nonparametric Test s 4
Miscellaneous 4

7
7
7
8
9
0
1
2
8
0
1

SECTION II: PLANT BIOPHYSICS

43

4. Basic Thermodynamic Quantities Savage 4

Basic Concepts and the Chemical Potential 4
Free Energy and Water Potential 4
Enthalpy 5
Water Potential in the Vapor State 5
Components of Water Potential 5
Water Potential of Aqueous Solutions 5
Theory of the Pressure-Chamber Apparatu s 5

5
5
7
0
0
1
3
3

vi Contents
5. Solutions (Ionic Relations) Dainty 5
Abbreviations Use d as Subscripts an d Superscripts 5
The Tables 5

5
5
6

6. Water Relations Dainty 6

The Tables 6

0
0

7. Energy Transfer Salisbury & Savage 6

Terms, Symbols, an d Units Appropriate in Energy-Transfer Studies 6
Some Equations Used in Heat-Transfer Studies 6

5
5
8

8. Phloem Transport Geiger & van Bel 7

The Tables 7

2
2

9. Electromagnetic Radiation Krizek & Sager 7

The Tables 7

5
5

SECTION III. PLANT BIOCHEMISTRY AND MOLECULAR BIOLOGY 7

10. Plant Biochemistry Black 8

Instructions o n Chemical and Mathematical Usage 8
Abbreviations an d Symbols 8
The Tables 8

1
1
4
6

11. Plant Molecular Biology and Gene Designations Reardon & Price 9
Terminology 9
Gene Designations 10

7
7
5

SECTION IV: PLANT GROWTH AND DEVELOPMENT

109

12. Morphogenesis and the Kinetics of Plant Growth Erickson 11

The Biometry of Growth 11
Shoot and Root Morphogenesis 11

1
1
3

13. Growth Analysis and Yield Components Bugbee 11

14. Plant Movements Haupt 12

Types and Mechanisms o f Movement 12
Control of Movement: Genera l 12
Terms for Induced Movements (Type s of Response) 12
Stimuli 12
Direction o r Sense of Response 12
Terms for Autonomous Movements 12

0
0
1
2
3
4
4

15. Growth Substances Cleland 12

16. Biological Timing Koukkari & Sweeney 12

17. Dormancy, Photoperiodism, and Vernalization Salisbury 13

General Considerations 11
Units for Growth Analysis and Yield Components 11

18. Stress Physiology Fuchigami, Maas, Lyons, Rains, Raison, & Shackel
General Stress-Physiolog y Terms 14

5
6

142
3

Contents vi
Chilling Injury 14
Cold Hardiness 14
Water Stress 15
Salinity Stress 15
APPENDICES: PRESENTING SCIENTIFIC DATA 16
A. Some Suggestions About Scientific Writing Salisbury 16
The Sentence 16
Modifying Words 16
Modifying Phrases and Clauses 17
Verbs 17
Some Further Notes about Punctuation 17
Abbreviations 17
Unnecessary Words 17
Words with Special Problems 17
Some Suggestions about Format and Word Processors 18
Summary 18
B. Standards for Effective Presentations Koning
Slide Presentations 18
Poster Presentations 19

i
4
6
1
4
1
3
4
8
2
5
6
8
8
9
3
5

188
9
5

C. Guidelines for Measuring and Reporting Environmental Parameters

for Plant Experiments in Growth Chambers Sager, Krizek, Tibbitts 202
Purpose and scope 20
2
Introduction 20
3
Definitions 20
3
Instrumentation 20
6
Measurement Technique 20
7
Reporting 20
8
Synoptic Table 21
0
INDEX

217

CONTRIBUTORS
Mary Barkworth, Utah State University, U.S.A.
Clanton C. Black, University of Georgia, U.S.A.
Bruce G. Bugbee, Utah State University, U.S.A.
Robert E. Cleland, University of Washington, U.S.A.
Jack Dainty, University of Toronto, Canada
Ralph O. Erickson, University of Pennsylvania, U.S.A.
Leslie H. Fuchigami, Oregon Stat e University, U.S.A.
Donald R. Geiger, University of Dayton, Ohio, U.S.A.
Wolfgang W. Haupt, Universtat Erlangen-Nurnberg, Germany
Ross E. Koning, Eastern Connecticut University, U.S.A.
Willard L. Koukkari, University of Minnesota, U.S.A.
Donald T. Krizek, U.S. Department of Agriculture, Beltsville, Maryland, U.S.A.
James M. Lyons, University of California, Davis, California, U.S.A.
Eugene V . Maas, U.S. Salinity Laboratory, USDA-ARS, Riverside, California, U.S.A.
John McNeill, Royal Ontario Museum, Toronto, Canada
Carl A. Price, Rutgers University, New Jersey, U.S.A.
William Rains, University of California, Davis, California, U.S.A.
John K. Raison, Macquarie University, North Ryde, Australia (deceased)
Ellen Reardon, Rutgers University, New Jersey, U.S.A.
John C. Sager, Kennedy Space Center, Florida, U.S.A.
Frank B. Salisbury, Utah State University, U.S.A.
Michael J. Savage, University of Natal, Pietermaritzburg, Republic of South Africa.
Kenneth Shackel, University of California, Davis, California, U.S.A.
Donald Sisson, Utah State University, U.S.A.
Beatrice M. Sweeney, University of California a t Santa Barbara, U.S.A. (deceased)
Theodore W. Tibbitts, University of Wisconsin-Madison, U.S.A.
Aart J.E. van Bel, Justus-Liebig Universitat, Giessen, Germany

PREFACE
When one person wishes to communicate some information directly to another person, it
is essential that the two speak the same language; that is, the words and symbols must have the
same meaning for both persons. Suc h a thought provides one motivation for the preparation of
this book, which is designed t o be a reference sourc e for plant physiologists and other plant
scientists who are preparing their research results for publication or other presentation. Th e
primary goal is to provide information about the use of units, symbols, and terminology in the
plant sciences , especiall y plan t physiology. I n addition , w e als o provide som e hint s an d
instructions abou t writing and the preparation of posters and slide presentations for scientific
meetings, including a format for presentation of growth-chamber data.
Section I introduces the basics. It s three chapters consider the use of SI units, rules for
botanical nomenclature, and basic principles of statistics. Section s II, III, and IV present more
detail i n the field s o f plant biophysics, biochemistry , and growth and development. Thes e
sections emphasize SI units whenever that is appropriate, but they also contain many lists of
terms that are used in the plant sciences. Th e appendices contain the hints and instructions for
writing an d for preparing posters an d slide presentations, plus a summary of guidelines for
reporting environmental parameter s for plant experiments in controlled environments. Th e
chapter on biochemistry was modified from The Journal o f Biological Chemistry; i t is included
here as a handy reference. Appendi x C was also prepared for another publication. Al l other
sections were originally prepared for this volume.
Each chapter was first prepared by one or more specialists in the field, and the authors then
sent their chapters to several colleagues. A s a result, the present chapters represent a t least the
beginnings o f a consensu s abou t th e term s an d sometime s symbol s within eac h subfield .
Although th e tim e when al l plan t scientist s agre e o n al l units , symbols, terminology , an d
presentation techniques may be in the distant future (i f it ever arrives), it is hoped that this book
will bring us closer t o such a meeting of the minds. Afte r I had edited the manuscripts sent by
the various authors, the entire book was sent to each author, who often commented about some
chapters beside s hi s o r he r own . Thi s process wa s repeate d severa l time s over a period
exceeding a decade (mostly because the project was set aside several times while other projects

x Preface
were being completed). Durin g this long gestation period, two authors died and several other s
retired! I n spite of the long period from conception t o birth, every chapter includes significant
changes mad e shortly before publication. Th e book presents the most current thinking of its
authors and editor.
The chapters tha t include definitions of terms follow two different approaches : I n some
chapters, terms ar e arranged alphabetically; in others they follow an order in which one term
builds upon the preceding term or terms (a mini-review of the subject). Th e choice of approach
depended upon the author and the subject matter, In the non-alphabetical cases, the number of
terms is rather limited s o that it should be relatively easy to find a term by scanning the lists.
A few references ar e presented, especially where definitions ar e somewhat controversial.
And controversy remains in plant physiology! Pleas e submi t suggestions for future edition s to
me or to the appropriate chapter author .
We have tried to remove inconsistencies an d contradictions althoug h some seem to be inevitable. W e are aware of considerable redundancy, which should make the booklet easier to
use as a reference source. A n editorial inconsistency tha t I have decided to allow concerns the
use of references. Man y show only initials of authors, but when given names were known to me,
I included them. W e have followed a reference style that includes written-out journal names
rather tha n abbreviations and more punctuation than is used in many current journals. Thi s
takes a little more space, but we believe it will make it easier for a reader to use the references.
Several secretaries were involved with the manuscript, but Laura Wheelwright did much
formatting, an d Mary Ann Clark must have spent the equivalent of an intense, full-time year
working on the final formatting of camera-ready copy with much direction from Kirk Jensen,
a Senio r Edito r a t th e Oxfor d University Press. Th e author s an d I wish t o expres s muc h
appreciation to those diligent secretaries; their efforts wer e often "above an d beyond the call of
duty."
F.B. Salisbury
Logan, Utah

APPROXIMATE CONVERSIONS: METRIC

Temperature

Length

Mass

U.S.a
Volume

This char t was prepared by F.B.S. for: Fran k B. Salisbury and Cleon W. Ross. 1969 . Plan t Physiology
First Edition. Wadswort h Publishing Co., Inc., Belmont, California. I t was not used in subsequent edition s
Some letters have been changed to reflect the conventions presented in this book.

GREEK ALPHABET AND ENGLISH EQUIVALENTS

Greek
letter
(roman)

A
B

Greek
letter
(italic)

Greek
name

A
B

Alpha
Beta

a
b

Gamma

g
d
e
z

Delta

E
Z
H

E
Z
H

Epsilon
Zeta

Eta
Theta

I
K

Iota

Kappa
Lambda

M
N

M
N

Mu
Nu
Xi

ks, x

P
r
s
t

Tau

Sigma
Upsilon

Phi
Chi
Psi
Omega

aAt end of word.

th
i
k
1
m
n
o

e(e)

Omicron

Pi
Rho

English
equivalent
(phonetic)

f, ph
ch, kh
ps
o(o)

UNITS, SYMBOLS ,
AND TERMINOLOG Y
FOR PLAN T PHYSIOLOG Y

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I
THE BASICS
This sectio n deals mostl y with constructed scientifi c languages. Ho w do people who
want t o communicat e usuall y achiev e a commo n language ? Mostly , we begi n a s
infants an d jus t us e th e languag e unti l meaning s becom e clear . Bu t ther e ar e
problems wit h thi s approach . Fo r on e thing , people i n different part s o f societydifferent geographica l areas , fo r examplehav e forme d differen t languages .
Furthermore, usag e ofte n produce s language s that lac k logi c an d consistency . A s
scientists, w e would like to communicate effectively with everyone else on the plane t
who migh t share our commo n interests . One solution tha t seem s to be fallin g into
place withou t an y directe d effor t i s th e broadenin g acceptanc e o f Englis h a s th e
language o f scienc e (an d muc h o f commerce , etc.) . A second solutio n fo r scienc e
has involve d a consciou s an d directe d menta l effor t t o create consistenc y an d
uniformity. Group s o f scientists hav e tried t o fin d way s to agree o n ho w to expres s
physical quantities , nomenclatur e o f organisms, and mathematica l symbols (among
other things) . I n thi s section, w e present th e thre e constructe d language s that dea l
with physica l quantities , taxonomi c nomenclature, and statistics:
1. Th e Internationa l Syste m of Unit s for expressin g physical quantities ,
2. Th e adopte d convention s for naming plant material; that is , many of th e
important rule s o f taxonomi c nomenclatur e agree d upo n i n Botanica l
Congresses, an d
3. Statistica l procedure s an d thei r notations ; thes e provid e a measur e o f
significance.
All plan t scientist s wh o wor k with quantitativ e measurements, regardles s o f thei r
specialty withi n th e fiel d o f plant physiology or i n other area s o f botany, need t o b e
conversant wit h thes e two international system s of communication plu s the mean s
of evaluatin g th e reliabilit y of their numerica l data.

1
THE INTERNATIONAL SYSTEM OF UNITS (SI UNITS)1
Frank B . Salisbury
Plants, Soils , an d Biometeorology Departmen t
Utah Stat e University
Logan, Uta h 84322-4820 , U.S.A .
As modern scienc e cam e into being , it depended mor e and more upon the accurat e
measurement o f physica l quantities . Suc h measuremen t require s a syste m o f
standards that i s recognized an d accepted b y all those who would communicate their
measurements t o eac h other . I n respons e t o thi s need , th e metric syste m o f
measurement wa s devised durin g the Frenc h Revolutio n (178 9 t o 1799) . I t was an
attempt to devis e a decima l syste m of measure s tha t woul d simplif y and unif y
calculations. Nearl y a centur y later, recognizin g the nee d t o furthe r improv e th e
system, th e Bureau International des Poids e t Mesures (BIPM ) was se t u p b y th e
Convention du Metre signe d in Paris in 187 5 by seventeen States ; the Convention was
amended i n 1921 . Th e tas k o f th e BIP M i s t o ensur e worldwid e unificatio n o f
physical measurements . I t operates in offices an d laboratories in Sevres, near Paris,
France, unde r supervisio n of the Comite International des Poids e t Mesures (CIPM) ,
which consist s o f 1 8 members, each fro m a different State . Th e CIP M itself come s
under the authorit y of the Conference General e de s Poids e t Mesures (CGPM) , which
consists o f delegates fro m al l the Membe r States (4 6 States i n March, 1991 ) o f th e
Convention du Metre. Th e CGP M meets a t present ever y four years , but th e CIP M
meets ever y year.
By th e mi d twentiet h century , th e metri c syste m wa s bein g widel y use d i n
science, bu t i n many cases, individua l branches of science ha d developed thei r own
specialized unit s an d terms . Fo r example , th e CG S (centimeter.gram.second )
system o f mechanical units , used especially in physics, included such terms a s dyne,
erg, poise , stokes , gauss , oersted, an d maxwel l (all no w considered obsolete) . T o

Earl y version s o f thi s chapte r wer e publishe d as Appendi x A i n Plant Physiology, Fourth
Edition, b y F . B . Salisbur y an d Cleo n W . Ross , publishe d by Wadswort h Publishin g Company ,
Belmont, California , 94002 , U.S.A. , an d i n Journal o f Plant Physiology (Salisbury, 1991). Recen t
study o f th e firs t an d second-leve l authoritie s (describe d in thi s chapter ) has le d no t onl y t o a
somewhat differen t approac h but als o t o som e importan t modification s an d change s i n a fe w units
and th e rule s for thei r use .

4 The

Basics

unify th e metri c syste m further , th e 9t h CGP M i n 194 8 instructe d th e CIP M t o

study an d recommen d th e establishmen t o f a "practical system o f units of measurement suitabl e for adoption b y all signatories to the Metr e Convention " (se e Taylor ,
1991). Thi s conferenc e als o laid dow n a set of principles fo r unit symbols and gav e
a lis t o f unit s with special names . Si x base unit s were established i n 1954 , an d th e
11th CGP M i n 196 0 adopte d th e nam e Le Systeme International d'Unites (English :
International Syste m o f Units) wit h th e internationa l abbreviatio n SI . Th e 11th
CGPM also laid down rules fo r prefixes, derived an d supplementary units, and othe r
matters. Thu s th e S I was born in 1960 , and subsequent meetings have added various
amendments. Th e 14t h CGPM, in 1971, fo r example, added the mole to the original
six base units , making a total o f seven bas e units in the SI , each with its own name
and symbol , which i s th e sam e (wit h slight spellin g differences ) i n al l languages .
The SI is currently by far the best measuremen t system humankind has been abl e t o
develop.
The purpos e o f thi s chapte r i s to presen t th e SI , especially a s it applies t o th e
plant sciences . Th e informatio n presented her e come s fro m variou s sources. I t is
convienent t o thin k o f thre e level s o f authority : Th e first , mos t primar y level, i s
"Le Systeme International d'Unites (SI), 6 e Edition", French an d Englis h texts . Thi s
is th e definitiv e publicatio n issue d i n 199 1 b y the Internationa l Bureau o f Weights
and Measure s (BIPM) . Althoug h thi s publicatio n wa s prepared jointl y with th e
National Physica l Laborator y i n th e Unite d Kingdom , some word s an d practice s
follow Unite d State s rathe r tha n Britis h usage . I n general , this usag e (e.g. , mete r
instead o f metre and lite r instea d of litre) i s closer t o the Europea n usag e than ar e
the Britis h practices . Th e Unite d States translation o f this primary volume is listed
in th e reference s t o thi s chapte r a s Taylo r (1991) ; i t i s virtually identica l t o th e
version publishe d b y the BIP M except fo r a few small matters such as use of the do t
instead o f the comm a a s the decima l marker. On e second leve l source o f authority
is the IS O Standard s Handbook , Thir d Edition , publishe d i n 199 3 b y the Interna tional Organizatio n fo r Standardizatio n (ISO) i n Geneva , Switzerland . I t expand s
upon th e rule s o f th e primar y source , an d thes e expansion s hav e influence d th e
deliberations o f th e CGP M s o tha t som e recommendation s of th e IS O Standard s
Handbook hav e become officia l S I rules. Anothe r second leve l sourc e o f authority
is Special Publication (SP ) 811 of the Nationa l Institute of Standards and Technolo gy (NIST , formerl y the U.S . Nationa l Burea u o f Standards). S P 811 is the "Guid e
for th e us e o f th e Internationa l Syste m o f Units (SI), " prepared b y Taylor (1995) .
The third leve l o f authorit y include s th e man y publications suc h a s thi s on e tha t
attempt to summarize , interpret , and condens e the SI for a give n field . (Se e many
of th e bold-face d entries i n the lis t at the end of this chapter.) Th e rules presente d
here are take n almos t exclusivel y from th e firs t tw o authority levels.
1. QUANTITIE S AND UNITS
In scienc e i n genera l an d th e plan t science s i n particular , we dea l wit h physical
quantities. T o communicate these physical quantities, we use three kinds of symbols:
a symbol for th e physica l quantity, a symbol for a numerical value (i.e., a number),

The International System of Units (SI Units) 5

and a symbo l fo r a unit. Fo r example , i f we want t o communicat e th e lengt h o f
some object , w e can write :
l = 5.6 7 m
If thi s notatio n i s to be a meaningful form o f communication, thos e o f us who want
to communicat e mus t agre e tha t th e symbo l for lengt h is l, that w e will use Arabic
numerals, an d tha t th e mete r (m ) represents a standard unit of length; namely , th e
length o f th e pat h travelle d b y light i n vacuum during a time interval o f 1/29 9 79 2
458 o f a second . O f course , fo r practica l purposes , mos t o f u s wil l trus t th e
manufacturers o f mete r stick s an d othe r measurin g devices , assumin g tha t thos e
manufacturers accuratel y follo w a reliable standard when they create th e measurin g
instruments. Th e variou s nation s hav e bureau s t o insur e thi s accurac y (i n th e
United States , Th e Nationa l Institut e o f Standards an d Technology) , an d a s note d
above, th e ultimat e authorit y for standards goes bac k to the CGP M an d the BIPM .
Remember tha t th e uni t represent s a number . Th e physica l quantit y i s th e
numerical valu e multiplie d b y th e unit . Thus , th e uni t i s subjec t t o algebrai c
manipulations. Fo r example , th e numerica l value can be thought o f as the rati o of
the physica l quantity to th e unit ; i n the abov e example : l/ m = 5.67 . Thi s notatio n
is particularly usefu l i n graph s and i n th e heading s o f columns in tables .
Note that th e symbol for the physical quantity (length in the example) is written
in italic or slanted type (underlined if italic type is not available) , an d th e symbol for
the uni t i s written i n roman (upright ) type. Thi s rul e (listed agai n in Table 4 , #17 )
should b e followe d wit h Greek symbol s as well as those fro m th e Roma n alphabet .
2. LE SYSTEME INTERNATIONAL

D'UNITES (SI)

The SI is a so-called coherent unit system, in which the equation s between numerica l
values hav e exactl y th e sam e for m (includin g the numerica l factors ) a s th e corre sponding equation s betwee n th e quantities . Thi s i s achieved b y defining unit s fo r
the bas e quantitie s (th e base units), and then derivin g further unit s fro m thes e bas e
units based upo n th e equations betwee n th e quantitites. Fo r example, th e equatio n
for spee d (v ) shows speed a s being equal t o th e incrementa l change i n distance (dl )
divided b y the incrementa l chang e i n time (dt) : v = dl/dt. Thu s th e uni t for spee d
is th e mete r pe r second : m/s . Therefore , th e S I include s base units an d derived
units. I n additio n ther e ar e tw o supplementary units, th e radia n (rad , fo r plan e
angle) an d th e steradia n (sr , fo r solid angle) .
The seve n bas e unit s o f th e S I ar e th e meter (length ; metre i s als o used ,
especially i n Britai n an d France) , kilogram (mass), second (time) , ampere (electri c
current), kelvin (thermodynami c temperature) , candela (luminou s intensity) , an d
mole (amoun t o f substance). Thes e units are show n with their symbol s in Table 1 .
Actually, i t i s possible b y combining th e unit s of space (length , area , an d volume )
with those of mass, time, an d temperatur e t o deriv e unit s of any physical quantity .
The basi c uni t o f lengt h i s th e meter (m) , whic h wa s originall y define d a s
equivalent t o th e lengt h of a bar preserve d i n Sevres, France; i n 196 0 i t was defined
as th e lengt h equa l t o 1 650 763.73 wave lengths in vacuum o f the radiatio n corre sponding t o th e transitio n between th e level s 2p10 an d 5d 5 of the krypton-8 6 atom.

6 The

Basics

In 1983 , agai n i n response t o advancin g technology, the mete r was redefined as th e

distance ligh t travel s i n a vacuu m during a tim e interva l o f 1/29 9 79 2 45 8 o f a
second.
Table 1 . Th e Seve n Bas e Units
Quantity

Unit

Symbol

Length (l )

meter

Mass (not weight ) (m a)

kilograma

kg

Time (t )

second

Electric curren t (I )

ampere

Thermodynamic temperatur e (T )

kelvin

K (not K )

Luminous intensit y (I )

candelab

cd

Amount o f substance (n , Q )

mole

mol

For historical reasons, the kilogram is the SI base unit rather than the gram. It is a unit of mass rather than weight.
Although weight is an acceptable synonym for mass, plant scientists should be careful to use mass instead of weight
whenever appropriatewhich is most of the time. (Note that the quantity mass is symbolized with italic m, which is not
to be confused with roman m for meter. See ISO Standards Handbook, 1993.)
As a unit of luminous intensity, the candela was traditionally based on the sensitivity of the human eye; we know of no
application in plant physiology. The lux (lx) is a measure of illuminance based on the candela (1 1x = 1 cd. sr. m-2 );
it has been widely used in plant science but should be avoided.
c

The mole should always be used to report the amount of a pure substance, and in such cases the type of substance must
be specified. To report the amount of a mixture or of an unknown substance, mass must be used.

For historica l reasons , th e gra m is not the S I base unit for mass. Th e kilogram
is th e onl y bas e uni t wit h a prefix . I t i s equa l t o th e mas s o f th e internationa l
prototype o f th e kilogram , mad e o f platinum-iridium , kept a t th e BIP M unde r
conditions specifie d b y the firs t CGP M i n 1889 . Not e tha t weight i s technically a
measure o f th e force produce d b y gravity, whereas the kilogra m is a uni t o f mass.
Mass i s a fundamenta l quantity that doe s no t chang e with the forc e of gravity (for
example, wit h location) . Th e weigh t o f objects , o n th e othe r hand , i s abou t 1
percent les s a t th e equato r tha n a t th e pole s an d i s 82 percent les s o n th e moon .
Thus i t is technically incorrec t t o use the word weight in conjunction with the uni t
kilogram. Th e prope r uni t for weight is the newton . (O n earth, the weight of a 10
kg mass is about 98 newtons.) Although in many technical fields and i n everyday us e
the term "weight* is considered as an acceptable synonym for "mass* plant scientists
should use the term "mass* whenever it is appropriate.
A balanc e balances th e mas s o f a n unknow n object agains t a define d mass;
hence, a balance measure s true mass. Al l balances depend upon an accelerationa l
force fo r thei r function , bu t th e magnitud e of th e accelerationa l forc e doe s no t
affect th e reading . Unfortunately , the magnitud e of accelerational force doe s affec t
the measuremen t o f mas s o n electroni c "balances " becaus e the y ar e reall y scale s

The International System of Units (SI Units) 1

that measur e weight . Thi s i s usually no t a seriou s proble m becaus e th e forc e of
gravity is constant fo r a given location, and electronic balance s an d spring scales are
calibrated with a standard se t o f objects o f known mass.
All object s wit h a mas s als o have a volume and thus displace som e air , which
has a densit y o f 1.20 5 k g m" 3 (standard atmospheri c pressure , dr y air , 2 0 C) . A
correction fo r this volume displacemen t woul d be necessary in some situations (fo r
example, measurin g th e mas s of a heliu m balloon!) , bu t mos t plan t tissue s hav e a
density simila r t o tha t o f water (1,00 0 kg m-3), s o the correctio n i s only about 0. 1
percent.
Note that a quantity of substance ca n be expressed either i n terms of its mass or
the numbe r o f particle s o f whic h i t i s composed : "Th e mole i s th e amoun t o f
substance o f a system tha t contain s a s many elementary entities as there are atom s
in 0.01 2 kilogra m o f carbo n 12 . Whe n th e mol e i s used , th e elementar y entitie s
must b e specifie d an d ma y be atoms, molecules , ions , electrons , othe r particles , o r
specified group s o f suc h particles " (Taylor , 1991) . Plan t physiologist s an d other s
include photon s amon g th e particle s tha t ca n be expressed i n moles, but not e that
the einstei n ( a mol e o f photons) i s not a n S I unit an d shoul d no t b e used . Not e
that 1 mol o f a substance contain s Avogadro's number of particles (no w defined a s
the numbe r o f atoms i n 0.012 k g of carbon 1 2 6.02204 5 x 10 23 particles) .
Following variou s astronomica l definition s of the second (e.g. , 1/8 6 40 0 of th e
mean sola r day), the secon d was defined i n 196 7 a s the duratio n o f 9 19 2 631 770
periods o f the radiatio n correspondin g t o th e transition betwee n th e tw o hyperfine
levels of the groun d state of the cesium-133 atom . Althoug h the minute, hour, day,
week, month, an d year are not officiall y par t of SI, plant physiologists will continu e
to us e them whe n appropriate .
The ampere is defined as that constan t current require d to produce, in vacuum,
a forc e o f 2 x 10- 7 newton s per mete r o f length between two parallel conductor s of
infinite lengt h and 1 meter apart . Becaus e force (the newton) is defined in terms of
length, mass , and tim e (se e Tabl e 2) , current coul d also be defined i n those terms .
The kelvin was define d b y the CGP M i n 196 7 a s th e fractio n 1/273.1 6 o f th e
thermodynamic temperature o f th e tripl e poin t o f water. Tha t sam e CGP M als o
adopted th e nam e kelvin (symbo l K ) t o b e use d instea d o f degree Kelvin (symbo l
K). I n addition t o the physical quantity thermodynamic temperature (symbo l T, unit
K), us e is also mad e o f Celsius temperatur e (symbo l t, unit C) , where t = T - T 0
and T 0 = 273.1 5 K by definition. A n interva l or differenc e o f Celsius temperatur e
can be expressed i n kelvin s a s well as in degrees Celsius .
Luminous intensit y (th e candela) wa s define d i n term s o f th e ligh t intensit y
perceived b y the huma n eye as compared with the intensity of freezing platinum, but
in 197 9 i t was redefined a s monochromatic radiation with a frequency o f 540 x 10 12
hertz and a radiant intensit y of 1/68 3 wat t per steradian . Th e watt (unit for power)
also combine s length , mass , and time. Thus , although the S I recognizes seve n base
units, only the unit s of length, mass, time, temperature, and numbe r (th e mole) ar e
truly basic in that the y are no t derive d from an y other unitsand temperature could
be derive d fro m th e firs t three .

8 The

Basics

Because th e candel a an d it s derivative s wer e base d o n th e sensitivit y o f th e

human eye, and plan t sensitivitie s ma y be very different (dependin g on the pigmen t
involved), the candel a an d it s derivatives (e.g. , the lux ) should not b e used by plant
scientists. Thi s i s true i n spite o f the mor e recen t definitio n based o n monochro matic light . Whil e th e candel a i s o f valu e t o engineer s wh o ar e concerne d wit h
artificial lighting for human beings, other measure s of radiation can be derived fro m
power (th e watt) pe r uni t are a ( W m- 2) or the numbe r (moles) of photons pe r uni t
area time s uni t tim e (usuall y [umol m-2 s-1). Thes e unit s should b e use d b y plant
scientists. I n either case , wav e lengths or frequencie s must be specified.
Table 2 list s th e prefixe s that ar e use d i n th e Internationa l Syste m of Units .
Some third-leve l publication s hav e suggested tha t fou r o f th e prefixe s were "non preferred": centi , deci, hecto, and deka. Althoug h the y were commonly used i n the
metric system, it was suggested that the y should be avoided when it is convenient to
use th e others . Th e first-leve l an d second-leve l sources , however , mak e n o suc h
distinction abou t bein g preferre d o r non-preferred . I n man y cases , usin g thos e
prefixes i s convenient an d lead s t o clarity . I n othe r cases , i t i s logical t o us e onl y
prefixes tha t diffe r b y a factor of 1 000 (103).
Table 2. SI Prefixesa (multiples and submultiples)
Factor

Prefix

da

(10)

deci

hecto

(102)

centi

kilo

(10 )

milli

mega

micro

giga

(106)
9

Prefix

Symbol

deka

tera

peta

(10 )

Symbol

nano

12

pico

1S

femto

P
f

18

(10 )
(10 )

Factor

(10-1)
(10-2)
(10-3)
6

(10- )
(10-9)
(10-12)
(10-15)

exa

(10 )

atto

(10-18)

zetta

(1021)

zepto

yotta

(1024)

yocto

(10-21)
24

(10- )

' Th e firs t syllabl e of every prefix is accented t o assure that the prefix will retai n its identity.

Table 3 show s som e importan t S I derive d unit s with specia l name s tha t ar e
derived fro m th e bas e unit s and ar e o f value to plan t scientists. (Se e Taylor , 1991 ,
for complet e lists. ) Not e tha t th e standar d acceleratio n du e t o gravit y i s a n
experimentally determined unit, and th e unifie d atomi c mass is an arbitrar y unit.

The International System of Units (SI Units) 9

Table 3. Derived Units of Interest to Plant Physiologists
Unit Nam e
Symbol
Quantity (symbol) a
Area (A )

square meter

m2
3

Definition
m.m

Volume (V )

cubic mete r

m.m.m

Speed or velocity (v )

meters pe r secon d

m.s-1

m.s-1

Force (F )

newton

kg.m.s-2

Energy (E) ,
work (W), hea t (Q )

joule

N.m (m 2.kg.s-2)

Power (P )

watt

J.s-1 (m 2.kg.s-3)

Pressure (p )

pascal

Pa

N.m-2 (kg.s-2.m-1)

Hz

cycle s-1

Frequency (v , Gree k nu ) hertz

Electric charg e (Q )

coulomb

A.s

Electric potentia l (V , )

volt

W.A-1 (J.A- 1.s-1; J.C-1)

Electric resistanc e (R )

ohm

ft

V.A-1

Electric capacitance (C )

farad

A.v-1 ( - 1)
c.v.1

Concentration (c )

moles per cubi c

meter

mol.m-3

mol.m-3

Irradiance (energy : E )

watts per squar e

meter

W.m-2

J.s-1.m-2

Irradiance (mole s of
photons)

moles per squar e

meter secon d

mol.m-2.s-1

mol.m- 2.s-1

Spectral irradiance
(moles o f photons )

moles pe r squar e
meter secon d nano meter

mol.m-2 .S-1 .nm-1

mol.m-2.s-1. nm-1

Magnetic field strengt h

(H)

amperes pe r mete r

A.m-1

A.m-1

Activity (o f radioactiv e
source: A )

becquerel

Bq

s-1

Standard acceleratio n
due t o gravity

standard acceleratio n gn
due t o gravity

9.806 6 5 m s-2

Unified atomi c
mass unit

Unified atomi c
mass uni t

1/12 o f mass of 12 C

Electric conductanc e (G ) Siemens

Quantit y symbol s ar e fro m IS O Standard s Handbook , 1993 . Ther e ar e man y varizitions o f suc h symbols .

Technically, velocity i s a vector quantit y requirin g specification o f a magnitud e (speed ) an d a direction, bu t mag nitude i s most importan t i n plan t science .

10 Th

e Basics

Table 4 summarizes th e styl e conventions tha t gover n the us e of S I units an d

that ar e o f interes t t o plan t scientists . Mos t o f thes e rule s ar e fro m th e primary
source o f authority : L e System e International d'Unite s (SI), 6 e Edition (o r th e
American Englis h equivalent: Taylor , 1991), bu t man y rely on the second-level IS O
Standards Handbook an d NIS T Specia l Publicatio n 811 (Taylor, 1995) . A few are
recommendations fro m third-leve l publications ; these ar e noted her e and discussed
further i n the tex t and in relation t o Table s 5 and 6.
Table 4. Summary of SI Style Conventions (Rules)
Names of units and prefixes
1. Uni t name s begi n i n lowercase, excep t a t th e beginnin g of a sentence o r i n titles or heading s in
which al l mai n words ar e capitalized ; tha t is , conventional grammatica l rule s appl y t o names.
Units name d afte r individuals a als o begi n i n lowercase . (Th e "degre e Celsius " migh t appea r
to b e a n exception , bu t "degree " begin s i n lowercas e an d i s modifie d b y "Celsius, " th e nam e
of a n individual . Us e o f "degrees centigrade" i s obsolete.)
2. Appl y only one prefix t o a unit name (e.g. , nm , not mum) . Th e prefi x an d unit name ar e joine d
without a hyphe n o r spac e between . I n thre e cases , th e fina l vowe l o f th e prefi x is dropped :
megohm, kilohm , an d hectare . Prefixe s ar e adde d t o "gram, " no t t o th e bas e uni t "kilo gram." Prefixe d ar e neve r use d b y themselves .
3. I f a compound uni t involving division is spelled out , th e word per i s used (no t a slash or solidus ,
except i n table s i n which space ma y b e limited) . Onl y one per i s permitted i n a writte n uni t
name (se e Rul e 3 0 below).
4. I f a compoun d uni t involvin g multiplicatio n i s spelle d out , th e us e o f a hyphe n i s usually
unnecessary, bu t i t ca n b e use d fo r clarit y (e.g., newto n mete r o r newton-meter) . Th e multi plication (product ) do t ( ) shoul d no t b e used when unit names ar e spelle d out .
5. Plural s of uni t name s ar e forme d b y adding a n "s, " excep t tha t hertz , lux , and Siemen s remain
unchanged, an d henr y become s henries .
6. Name s o f unit s are plura l fo r numerica l values greater tha n 1 , equal t o 0 , or les s tha n -1 . Al l
other value s tak e th e singula r form o f th e uni t name . Examples : 10 0 meters , 1. 1 meters , 0
degrees Celsius , - 4 degrees Celsius , 0.5 meter, -0. 2 degre e Celsius , - 1 degree Celsius , 0. 5 liter.
7. NIS T S P 811 (Taylor , 1995 ) recommend s tha t writte n name s o f units be avoide d mos t o f th e
time; unit symbols shoul d b e used instead . I t is appropriate, however , to use a written name th e
first tim e th e uni t appear s i n a tex t i f it i s felt tha t reader s migh t not b e familia r wit h the unit .
Symbols for units
8. Uni t symbol s shoul d b e though t o f as mathematica l entities : Th e physica l quantit y equals th e
numeral multiplie d b y th e valu e represented b y the uni t symbol . Hence , with fe w exceptions
(see Rule s 7 and 15) , symbol s are use d whe n units are use d i n conjunction with numerals.
9. Writte n symbol s are neve r mad e plura l (that is , by addition o f "s") .
10. A symbo l i s not followe d b y a perio d excep t a t th e en d o f a sentence .
11. Symbol s fo r unit s named afte r individualsa have th e first letter capitalized , but th e nam e o f th e
unit i s written i n lowercas e (se e rul e 1) . Othe r symbol s ar e no t capitalize d excep t tha t th e
second leve l authorities recommen d a capita l L instea d o f a lowe r cas e 1 as th e symbo l fo r th e
liter t o avoi d confusio n with the numera l one (1) . Bot h L and 1 are recognize d b y the primar y
authority a s symbols fo r th e liter . Th e capita l L is recommended here .
12. Symbol s fo r prefixe s greate r tha n kil o ar e capitalized ; kil o and al l others ar e lowercase . I t i s
important t o follo w thi s rule because some letter s for prefixe s ar e th e sam e as some symbols or
another prefix : G fo r gig a an d g for gram ; K fo r kelvi n an d k fo r kilo ; M fo r meg a an d m fo r
milli an d fo r meter ; N fo r newto n an d n fo r nano; an d T fo r ter a an d t fo r metri c ton .
Continued

The International System o f Units (S I Units) 1

Table 4. Summary of SI Style Conventions (Rules) (continued)

13. Us e numerica l superscript s (2 and 3 ) to indicat e squares an d cubes; d o not us e sq., cu. , or c. I t
is also better, when uni t name s ar e writte n out , t o us e the for m "secon d squared " rathe r tha n
"square second" unless volume o r are a ar e being discussed: "squar e meter, " "cubi c meter."
14. Exponent s als o appl y t o th e prefi x attache d t o a unit name; th e multipl e or submultipl e uni t is

treated as a single entity. Thus nm3 is the same as 10-18 m3.

15. Third-leve l sources an d Englis h styl e manual s recommend tha t sentence s shoul d no t begi n with
numerals. Becaus e a unit symbol is always proceeded wit h a number (numeral), a sentence can
never begi n wit h a uni t name o r symbol . Wheneve r possible , a writer shoul d recas t a sentenc e
so it does not begi n with a numeral; if that can't b e done, th e numbe r and unit name shoul d b e
spelled out .
16. Compoun d symbol s forme d b y multiplication may contain a produc t do t ( ) t o indicat e multiplication; internationa l rule s sa y tha t thi s ma y b e replace d wit h a perio d o r a space . I n th e
United States, the produc t do t i s recommended. Compoun d symbol s formed by division can use
a slas h (/) , a horizonta l lin e with units above an d below , or b e indicate d by negative exponents ;
e.g., umol. m .s-1, umol.mol- 1 , etc . I n n o cas e shoul d symbol s b e ru n togethe r (e.g. ,
Wm-2).
17. Becaus e compoun d uni t symbol s ar e mathematica l entities , the y mus t no t includ e nonsymbo l
words o r abbreviations . Thi s i s not tru e of unit names without numerals. Thu s an author mus t
avoid " umol CO2 (mo l o f air)- 1" bu t ca n write: "Dat a ar e presente d a s micromole s o f CO 2
per mol e o f air ( umol.mol- 1)." (Se e discussio n i n the text. )
18. D o no t mi x symbols an d spelled-ou t uni t name s (e.g. , W per squar e meter) , an d never mi x SI
units or thei r accepte d relative s (e.g. , liter, minute, hour , day , plane angle in degrees) wit h units
of another system suc h as the CG S or th e Englis h system (e.g. , mile s per liter , kg ft-3, o r gram s
per ounc e fo r th e quantit y o f fat i n a food) .
19. Th e percen t symbo l (% ) i s an acceptabl e uni t for us e with th e SI : % = 0.01 . Whe n used , a
space i s left betwee n th e symbo l % an d th e numbe r b y which it i s multiplied: X = 25 % = 2 5
x 0.0 1 = 0.25 . Rathe r tha n usin g such terms as "percentage b y volume" (meaningless becaus e
% i s simply a number) , a recommende d approac h i s to presen t dat a a s mL/L, umol/mol , g/kg ,
mol/L, mol/kg , etc . (Taylor , 1995).
20. Uni t symbol s ar e printe d i n roman typ e (uprigh t letters); italic letters (slanted ) ar e reserve d for
quantity symbols , suc h a s A fo r area , m fo r mass , t fo r time , and fo r water potential . Fo r
typewriting or longhand , underlinin g may be use d a s a substitute fo r italics . Accordin g t o thi s
rule, th e Gree k mu , u , whe n use d a s the prefi x symbol fo r micro , should b e printe d i n roma n
type (no t i n italics).
Numerals, often with Symbols
21. A spac e i s lef t betwee n th e las t digi t o f a numera l an d it s uni t symbol . A produc t do t () ,
space, o r slas h (/ ) i s use d betwee n uni t symbol s when mor e tha n on e i s used ; se e rul e 16 .
Exceptions ar e th e degree , minute, and second symbol s for angles or latitude s (e.g., 30 north) .
Note tha t th e degre e Celsiu s (C ) i s a singl e uni t symbo l (n o spac e betwee n an d C ) tha t
should als o b e proceede d b y a space . I t i s incorrect t o us e 1 2 t o 2 5 C (tha t is , to us e
without C) ; correc t form s are: 1 2 C to 2 5 C, (12 to 25) C , or 12-2 5 C .
22. Whe n a quantity is used i n an adjectiva l sense, Englis h rule s of grammar suggest tha t a hyphe n
should b e used betwee n the numera l and the uni t name: a five-hundred-watt lamp . Bu t when
unit symbols ar e used , the hyphe n shoul d b e omitted: a 50 0 W lamp (becaus e th e symbo l i s a
mathematical entity , an d th e hyphe n coul d b e mistake n for a minu s sign).
23. I n th e Unite d States , th e perio d i s used a s the decima l marker although some countrie s (e.g.,
France, Germany , Grea t Britain ) use a comma o r a raised period .
Continued

12 Th

e Basics

Table 4. Summary of SI Style Conventions (Rules) (continued)

24. T o avoi d confusio n (becaus e some countrie s us e a comma a s a decimal marker), a space shoul d
be used instea d o f a comma t o group numerals into three-digit groups; this rule may be followed
to th e righ t a s wel l a s t o th e lef t o f th e decima l marker . Omissio n o f th e spac e i s preferre d
when ther e ar e onl y four digits , unless th e numera l i s in a colum n with others tha t hav e mor e
than fou r digits . (I n spit e o f thi s rule , man y journals tha t consistentl y us e a perio d a s th e
decimal marke r als o us e the comm a t o grou p numeral s int o three-digit groups. )
25. Decima l fraction s are preferre d t o commo n fractions .
26. Decima l value s les s tha n on e hav e a zero to th e lef t o f the decima l (e.g. , 0.2 m).
27. Multiple s an d submultiple s ar e generall y selecte d s o tha t th e numera l coefficien t ha s a valu e
between 0. 1 and 1000. Exception s occu r when the differences between number s being compare d
are extrem e (e.g. , 150 0 m o f 2 m m wire) , an d fo r comparison , especiall y i n tables , simila r
quantities shoul d us e th e sam e unit , even i f the value s fal l outsid e thi s range .
28. Wit h numerals , d o no t substitut e th e produc t do t ( ) fo r a multiplicatio n sign (x) . (E.g. , us e
2 x 2 , not 2.2. )
The denominator
29. Fo r a compound uni t tha t i s a quotient , us e "per" t o for m th e nam e (e.g. , meters pe r second )
and a slas h (/ ; solidus) t o for m th e symbol , with no spac e befor e o r afte r th e slas h (e.g. , m/s) .
Compound unit s may be written wit h negative exponent s (e.g. , m.s-1 o r m s- 1).
30. D o no t us e tw o o r mor e "pers " o r slashe s i n the sam e expressio n becaus e the y ar e ambiguou s
(see Rul e 3) ; negativ e exponent s avoi d thi s problem : J.K-1 .mol- 1 (no t J/K/mol) ; J/K.mo l i s
acceptable because all symbols t o th e righ t of the slas h belong t o th e denominator .
31. Man y third-leve l source s suggest tha t th e denominato r shoul d no t b e a multipl e or submultiple
of a n S I base uni t (e.g.,uN.m- 2 bu t no t N.um-2 ). (Bu t se e discussio n i n the text. )
aIndividuals afte r who m unit s ar e name d include : Antoin e Henr i Becquere l (France , 1852-1908) , Ander s Celsiu s
(Sweden, 1701-1744) , Charle s Augusti n d e Coulom b (France , 1736-1806) , Michae l Farada y (England , 1791-1867) ,
Heinrich Rudol f Hert z (Germany , 1857-1894) , Jame s Prescot t Joul e (England , 1818-1889) , Lor d Willia m Thomso n
Kelvin (Scotland, 1824-1907) , Sir Isaac Newton (England, 1643-1727), Geor g Simon Ohm (Germany, 1787-1854), Blaise
Pascal (France , 1623-1662) , Si r Willia m Siemens (Germany , Great Britain , 1823-1883), Coun t Allessandr o Giusepp e
Antonio Anastasi o Volt a (Italy , 1745-1827) , an d James Watt (Scotland, England, 1736-1819).

The CIP M recognized i n 196 9 tha t users of SI will also wish t o emplo y with i t
certain unit s that ar e no t par t o f it , bu t tha t ar e importan t an d ar e widel y used.
These unit s (along with the unifie d atomi c mass unit and the standard acceleration
due t o gravity) , ar e show n in Table 5 . Not e tha t a goal in settin g up th e International Syste m of Units was to produc e a coherent system , as noted above , a system
in whic h derive d unit s ar e variou s combination s o f th e bas e unit s withou t th e
necessity o f including numerical multiplication factors. Al l of the unit s in Table 5
do requir e th e us e o f suc h factors , an d henc e the y los e th e advantage s o f th e
coherence o f SI units . I t was recommended that thei r us e be restricted t o specia l
cases. I t i s clear, however , that plant scientists will use th e minute , hour, and day
(not to mention the week, month, and year) without hesitation in reporting methods
and results . Th e liter is also a much more convient unit for plant scientists than the
cubic meter, which i s th e officia l S I unit of volume. Thus , we can b e thankfu l fo r
the CIPM' s decision s in 196 9 and fo r Table 5!

The International System o f Units (S I Units) 1

Table 5. Some Units used with the SI but not Officially Part of SIa
Name

Symbol

Value i n SI units

minute

min

1 min = 60 s

hour

1 h = 6 0 min = 3600 s

day

1 d = 2 4 h = 8 6 400 s

degree

1 = ( /180 ) ra d

minute

1' = (1/60) = ( /1 0 800) rad

second

"

1" = (1/60) ' = ( /64 8 000) rad

liter (litre)

L (l)

1 L = 1 dm3 = 10- 3 m 3

metric ton (tonne)

1 t = 10 3 kg

unified atomi c mass unit b

1 u = (1/12 ) of the mass of an atom of

the nuclid e 12 C

standard acceleration g
due t o gravit y c

9.80

6 65 m.s-2

Becaus e thes e units must be multiplied by a factor t o mak e the m equivalen t t o SI units, the y are no t coheren t in the
sense of othe r SI units.

The actua l value of th e unifie d atomi c mass uni t in SI unit s must be determined by experimentation. A t presen t it is
considered
consideredto
to be: u = 1.66 0 540 2(10)
2(
x 10- 27 kg . Th e uncertainty of the last tw o figures, a t the level of one standard
deviation, i s shown i n parentheses.
c

Thi s valu e was confirmed in 191 3 b y the 5t h CGPM . It s symbol , g n, shoul d b e use d instea d o f th e man y symbols
currently used t o indicat e one acceleratio n due t o gravity at the earth's surface (e.g. , g , g, G, G , Xg , etc.).

Table 6 includes som e unit s that were use d with the metric syste m but tha t th e
CIPM recommend s shoul d no t b e use d wit h the SI . A fe w of these units continu e
to b e in wide use among plan t scientists .
3. SOME SPECIAL CONSIDERATIONS
Although mos t o f th e rule s i n Table 4 are explaine d adequatel y in th e table , a few
of the m a s well as some o f the unit s in Tables 5 and 6 are worthy of discussion .
A. Language Conventions with Si-Unit Names and Symbols.2 Writte n ou t
names fo r the unit s follow the rules of grammar (English or other language) , whereas th e uni t symbols shoul d b e though t o f a s mathematica l entitie s b y which th e
preceeding numera l i s multiplied. Fo r example , unit names begin with a lowercase
letter unles s grammatica l rule s cal l fo r uppercas e (i.e. , a t th e beginnin g o f a
sentence an d i n titles) , bu t th e uppe r o r lowercas e o f symbols mus t neve r b e
changed regardles s o f where the y appear. I n English, names ar e often mad e plural

Rule s in Table 4 that bea r on thi s discussio n are : 1, 7, 8, 15, 17, & 22.

14 Th

e Basics

by addition o f s, but S I symbols never are . Number s (usuall y written out ) followe d
by uni t name s use d i n an adjectiva l sens e can be connecte d wit h a hyphe n (e.g. , a
fifty-watt lamp ) bu t th e hyphe n i s not use d wit h symbol s (a 50 W lamp). Becaus e
of the mathematica l natur e o f symbols, it is desirable t o us e them instea d o f names .
Of course the nam e ca n be used th e firs t tim e i t appears if the reader migh t not b e
familiar wit h th e uni t o r it s symbol.
In som e language s i t i s no t uncommo n fo r a numera l t o begi n a sentence ; i n
English thi s should b e avoided, preferabl y by recasting th e sentence, but if necessar y
by writing out th e numera l an d it s unit name .
Table 6. Some Discarded Metric Units
Discarded Metric Unit

Acceptable SI Unit

micron (u )

micrometer ( um )

millimicron (mu )
angstroma (A)

0.1 nanometer (nm)

nanometer (nm)

bara (bar )

0.1 megapascal (MPa);

100 kilopascal (kPa)

calorie (cal)

4.1842 joule (J)

degree centigrad e (C)

degree Celsiu s (C )

hectarea (ha )

10 000 m2 or 0.01 km 2
mole of photons or quant a (mol)

einstein (E )
dalton (Da )

unified atomi c mass unit (u )

(see Tabl e 5)

standard "gravity " (g, g, G, G, xg , etc. )

standard acceleration due to gravity (g n)

molar solution (M)

mol-L-1 (kmol m-3)

molal solution (m )

mol-kg-1

parts per millio n (ppm)

mg.kg-1
{umol.mol-1 (e.g., CO 2 i n air)
(Use k g for mixe d substances and mo l fo r
pure substances and gases. )
1000 mm 3.m-3 (volume ; e.g., liquids)
-1
ug.kg-1
nmol.mol-1
mm3.m-3 (volume ; e.g., liquids)

parts per billio n (ppb )

aIn vie w of existin g practice i n certai n fields , th e CIPM (1978 ) considered tha t these units could b e use d with the SI
temporarily although they should no t be introduced wher e they are no t use d a t present.

Because name s follo w grammatica l rules , i t i s acceptabl e t o us e the m i n

conjunction wit h othe r terms , bu t suc h term s mus t not b e include d wit h S I units :
"Photon flu x wa s measure d a s mole s o f photon s i n th e photosyntheticall y activ e
range (40 0 t o 70 0 nm ) pe r squar e mete r secon d (umol.m-2 .s-1 )." "Dat a ar e
presented a s milligram s of protei n pe r gra m of fres h tissu e (mg/g). " Or : "Protei n

The International System o f Units (S I Units) 1

data ar e presente d o n a fresh-mass basis (mg.kg- 1)." Thi s rule i s often overlooke d
by plan t physiologists , wh o eve n sometime s construc t meaningles s symbol s t o
present thei r data : mg/gf w (meanin g milligrams per gram of fresh weight) or mg (kg
fresh mass)- 1. Th e rul e was discussed b y Downs (1988) and perhap s in othe r thir d
level source s o f which I am not aware , but i t has otherwise been largel y overlooked
by plant physiologists. Th e rule is emphasized, however, by the second-level authorities
(ISO Standard s Handbook , 1993 ; NIS T S P 811 , Taylor , 1995) . Plan t scientist s
should improv e the rigo r of their presentation s b y adhering to this rule .
B. Space Between Numerals and Units and Within Compound Units. Fo r
some unknow n reaso n i t ha s becom e increasingl y commo n t o omi t th e spac e
between a numeral and the unit that follows (e.g. , a 50mL flask) . I n the worst cases,
the spac e o r produc t do t i s omitted betwee n symbol s in a multipl e unit, creatin g
new symbol s tha t hav e n o meanin g (e.g. , Wm- 2, umolm-2S- 1, etc.) . Thi s practic e
breaks Rule s 16 , 19 , and 2 1 i n Tabl e 4 , an d leavin g ou t th e spac e ca n confus e
readers. Us e o f th e produc t do t i s highl y recommende d i n th e Unite d State s
(Taylor, 1995) , but plan t scientists hav e used it only infrequently. It s consistent us e
would remove any ambiguity from multipl e units and would overcome the tendency
to run units together.
C. Italics for Quantity Symbols, Roman for Unit Symbols. Thi s simple practice
is stated i n Rul e 20 , Table 4 , but man y plan t scientists seem t o b e unawar e of it .
Remember tha t unit symbols are printed in roman type (upright letters); italic letters
(slanted) ar e reserve d fo r quantit y symbols, such a s A fo r area , m fo r mass , t fo r
time, and 7 fo r water potential. Fo r typewriting or longhand, underlining may be
used as a substitute for italics. Accordin g to this rule, the Greek mu , u , whe n used
as the prefi x symbol for micro, should be printed in roman (i.e., upright) type whenever possible . Unfortunately , not all word processors allow this. (Not e that Greek ,
Roman, or eve n Cyrilli c alphabets ca n be printed in either roma n or italic type).
D. Only One per or Slash in a Multiple Unit. Thi s is another simple rule that
plant scientist s shoul d appl y more widel y (Rul e 30, Table 4) : D o no t us e tw o or
more "pers " o r slashe s i n th e sam e expressio n becaus e the y ar e ambiguous .
Negative exponent s avoi d thi s problem : J-K-1.mol- 1 (not J/K/mol) ; J/K.mol i s acceptable becaus e al l symbols to th e right of the slash belong to the denominator. I f
this is written out, i t becomes: joule s per kelvi n mole.
E. Only Base Units in Denominators. A s note d i n Rul e 31 , Tabl e 4 , man y
third-level source s sugges t tha t th e denominato r shoul d no t b e a multipl e o r
submultiple o f a n S I bas e uni t (e.g. , uN.m- 2 bu t no t N.um- 2). A s edito r o f
Journal of Plan t Physiolog y during the pas t si x years, I have found thi s rul e t o b e
the mos t difficul t t o enforce . I t goe s against much tradition and sometime s seem s
illogical an d unreasonable . Fo r example , authors hav e long reported amount s or
concentrations o f metabolites, hormones, and other compounds as ug/mg, nmol/mL,
etc. I t ma y seem t o g o against one's intuition to us e the equivalent s of thos e tw o
examples: g/kg , umol/ L (or , using "true" SI units: mmol/m-3 ).
Now I learn tha t th e rul e of onl y base unit s i n denominator s is recommended
solely i n third-leve l sources bu t i s no t a n officia l S I rul e an d i s no t i n th e IS O
Standards Handbook or NIST SP 811 (Taylor, 1995), which are second-level sources
with virtually as much authority as the primary SI publication. I t is not necessar y to

16 Th

e Basics

adhere t o ever y suggestio n pu t fort h i n third-leve l publications . Thus , I will n o

longer tr y to enforc e the rul e in my editing.
Having sai d that , I will nevertheless mak e an argumen t (third level! ) tha t th e
rule (suggestion , at least ) can , in many cases, be quite logical and helpful. Us e of a
single denominator fo r a given quantity by everyone in the fiel d allow s us to thin k
about th e variou s quantitie s withou t th e necessit y o f mentally converting the m t o
our familia r range, differen t perhap s fro m tha t o f th e autho r w e ar e reading . O f
course, t o take advantag e of this rule, those within the field mus t agree t o form new
habits. Fo r example , photosynthesis rate s were previously expressed on the basis of
CO2 uptak e (a s milligrams or moles ) per squar e decimete r o f leaf surface, perhaps
because a square decimete r seeme d t o be an area simila r to the are a o f real leaves .
Most workers now express mos t parameters relatin g t o photosynthesi s on the basis
of a square meter, a s suggested by this rule. Afte r all , few real leaves are exactl y 1.0
dm2 o r exactl y 1. 0 m 2 i n area . (Doe s a banan a lea f approac h 1. 0 m 2?) An d
irradiance i s commonly expressed o n th e basi s of a square meter: umol.m- 2.s-1 o r
W.m-2. Whe n everythin g i s expresse d o n th e sam e basi s (m- 2), comparison s ar e
much easier.
In spit e o f th e tradition s note d above , unit s fo r quantitie s o f mas s can easil y
follow th e rule . Wh y express the amoun t of growth regulator in a tissue sample as
8.5 pg mg- 1 when i t i s just as eas y to writ e 8.5 u g kg- 1? A kilogra m is a relativel y
large amount o f tissue , but i t is easy enough to visualize. Th e researcher probabl y
didn't us e a 1. 0 mg sample of tissue an y more than he or she used a 1. 0 kg sample.
Uniform adherenc e t o thi s rul e would soon familiariz e researcher s wit h its merits.
Nevertheless, applyin g the rul e is optional.
Because a cubi c mete r i s so large , i t ma y seem a littl e les s logica l t o expres s
solution concentration s o n th e basi s o f a cubic meter, which is the S I base uni t of
volume. Nevertheless , man y plant scientists hav e decided to use the cubi c meter as
the base unit for solution concentrations : 1. 0 m3 = 100 0 L; thus a 1.0 mmol/L solution = 1. 0 mol m- 3. Becaus e S I rules allo w use of the liter , however , even thoug h
it is not a n official par t of the system, and because concentrations base d o n the liter
have long been use d by plant physiologist s (and many solutions are made up in liter
quantities), i t is acceptable t o use liters i n most journals that publish papers i n th e
plant sciences . (Se e discussio n belo w o n molarit y and molality ; their traditiona l
units, M and m , should b e phase d ou t o f use by plant scientists.)
It i s no t alway s possible o r desirabl e t o hav e only base unit s in denominators .
For example , spectra l energie s must specif y a narro w wave-lengt h range , th e
nanometer: mol.m-2.S-1.nm- 1 o r W.m-2.nm- 1. (Th e rang e tha t wa s actuall y
measured shoul d always be stated i n the methods section)
In some case s i t ma y be preferabl e t o write out informatio n fo r furthe r clarity .
For example , a stric t edito r tryin g t o enforc e thi s rul e woul d insis t tha t a tem perature gradient of 1 K mm-1 be written as 1000 K m-1. I t would be better t o state:
"A temperature gradien t o f 1 K over a distance of 1 mm was measured."
The recommendatio n o f thi s third-leve l publication : When i t i s logical an d
helpful to do so, use only SI base units in denominators.
F. The Liter: Symbols and Use; Molar and Molal Solutions. Th e lite r (litre in
England, France) i s not a n officia l par t of SI, probably because it is not "coherent. "

Summary of the International System of Units (SI Units) 17

To deriv e i t fro m th e cubi c meter, th e S I base unit , a multiplication factor must be
used ( 1 L = 0.00 1 m 3). Th e 12t h CGP M in 196 4 declared, however, "that the word
'liter' ma y b e employe d a s a specia l nam e fo r th e cubi c decimeter " an d recom mended "tha t th e nam e lite r shoul d no t b e employe d t o giv e th e result s o f hig h
accuaracy volum e measurements. " (Taylor , 1993. ) Thes e statement s effectivel y
defined th e liter as exactly 1 dm3 and the milliliter (mL ) as 1 cm3. Becaus e th e liter
has a convien t siz e an d th e ter m i s traditiona l an d widel y use d b y non-scientists ,
plant scientist s wil l continue t o us e it an d th e milliliter . Eve n th e decilite r migh t
sometimes b e mos t convenient . O f course, we could use dm 3 and cm 3 as easily.
In 1979 , th e CGP M considere d "that , i n orde r t o avoi d th e ris k o f confusion
between the lette r 1 and the number 1 , several countries have adopted th e symbol L
instead o f 1 for th e uni t liter... " I t wa s further decide d "t o adopt th e tw o symbols
1 an d L a s symbol s t o b e use d fo r th e uni t liter , considerin g furthe r tha t i n th e
future onl y one o f thes e tw o symbols should be retained... " NIS T SP 811 (Taylor,
1995) strongl y recommends L as the symbo l for the liter .
The vast majority of papers i n plant physiology express concentrations i n term s
of molarity (symbol M = mol.L- 1) or , especiall y i n th e fiel d o f water relations , o f
molality (symbo l m = mol.kg- 1). Nevertheless , bot h second-leve l authoritie s (IS O
Standards Handboo k an d NIS T SP 811, Taylor , 1995 ) recommen d that th e symbols
for thes e term s b e discontinued . (Althoug h no t state d i n those sources , th e term s
themselves migh t still b e used. ) Th e reaso n i s that thos e uni t symbols (M and m )
are specialize d symbol s that migh t not b e understood b y someone outsid e th e fiel d
(e.g., a physicist), whereas mol.L- 1 an d mol.kg- 1 are simpl e S I units understood b y
anyone familia r wit h SI . Furthermore , m fo r molalit y might be mistake n a s m fo r
meter. Thi s recommendatio n i s confirme d b y a n importan t third-leve l source ,
Quantities, Units an d Symbols i n Physical Chemistry (Mill s e t al. , 1995). Mos t plan t
physiologists will no doubt continue to use the terms molarity an d probably molality,
but i t is recommended tha t th e equivalent SI units be used instead of the traditiona l
symbols. (Bu t see Table 1 in Chapter 10. )
G. The Dalton and the Unified Atomic Mass Unit. Man y biochemists and mos t
(virtually all ) plan t physiologist s us e th e dalton (D a o r D ) a s a uni t o f atomic o r
molecular mass . Th e dalto n has , however, neve r been accepte d b y the CGPM , and
it i s exactly equivalent t o th e unified atomic mass unit (symbol u, Table 5) , which
has bee n considere d an d accepte d b y the CGP M an d is published i n th e first-level
authority. Hence , there seems t o be little excuse beyond tradition to use the dalton.
Most plan t scientist s gav e u p th e einstei n i n favo r o f mol e o f photons . Th e
recommendation i s to begi n t o us e the unifie d atomi c mass unit with its symbol u,
probably with some explanatio n unti l it becomes mor e familiar t o plan t scientists .
H. Equivalent of Gravity at the Earth's Surface. I t is common for biochemist s
and other s t o expres s th e acceleratio n experience d b y a sample being centrifuged as
multiples of the averag e acceleratio n cause d by gravity at th e earth's surface . Ther e
has been almost n o agreement, however , on the symbol that should be used for this
value. O n sees i n various publications: G , g, g, G, g, xg, an d no doubt others. Th e
problem wit h these symbol s is that g is the symbo l for gram, G i s the prefi x symbo l
for giga , and italic s (g ) i s reserve d fo r physica l quantitie s rather tha n units. Bold
facing has no precident in the use of units. Actually , there never shoul d hav e been

18 Th

e Basics

a problem becaus e the CGP M establishe d th e standard acceleration due to gravity i n

1901 and confirmed the value in 1913 . Th e primary and secondary sources show th e
symbol g n. Th e logi c o f thi s symbo l i s tha t th e acceleration o f fre e fall (g ) i s a
physical quantity (hence italics) that can have any value (units: m.s-2), but the
standard acceleration of free fall (indicate d b y the subscript , which is no t i n italics :
gn = 9.80 6 6 5 m s- 2) i s the valu e of particular interest . I t mus t be experimentall y
determined an d i s thus a noncoherent unit , but multiple s of this value can be use d
to describe th e acceleration caused by centrifugation (e.g., sample centrifuged for 20
min at 100 0 g n) or the acceleration experience d i n an orbiting satellite (e.g. , 10- 3 g n).
This symbol , i n context , shoul d b e readil y understoo d b y everyon e withou t an y
special explanation .
I. Other Discarded Metric Units. Tabl e 6 include s a numbe r o f discarde d
metric unit s tha t hav e no t ye t been discussed . On e occasionall y see s micron, but
most of us now use th e micrometer and nanometer. Th e angstrom is seldom use d in
the plan t science s bu t i s still use d in certain fields , whic h is permitted accordin g t o
the footnot e i n Table 6 . Th e bar is still use d in meteorology an d sometime s i n th e
field o f plant wate r relations , bu t it s replacement wit h the coheren t megapascal o r
kilopascal ha s been accepte d b y most plan t physiologists. Th e hectare (ha ) will n o
doubt continu e t o b e use d b y agriculturists instead o f the mor e correc t hm 2 or m 2.
The Calorie (kilocalorie ) i s a part of our moder n dieting culture, but (i n the Unite d
States) s o i s th e Fahrenhei t temperatur e scale ; scientist s us e degree s Celsiu s an d
should also us e joules instea d o f calories. On e often see s parts per million (or billion
or eve n trillion), but i t i s mor e logica l t o us e thei r equivalent s i n unit s o f mass ,
volume, or amoun t of substanc e (e.g. , mg.kg- 1, mmol.kg-1 , mol-L-1). Wit h suc h
units, it i s not necessar y t o specif y th e basis of comparison (i.e., volume, mass, etc.).
IMPORTANT REFERENCES FOR APPLICATION OF S I UNITS
Many of these publication s are no w out o f date an d ar e include d here onl y for historical reference.
The most recen t and most recommende d publication s that have come t o my attention ar e written in
bold face.
American National Metric Council. 1993. ANMC Metric Editorial Guide, Fifth Edition. American National Metric Council, 4330 East/West Highway, Suite 1117, Bethesda, MD 20814.
[Anonymous]. Standar d Practic e fo r Us e o f th e Internationa l System o f Units . AST M E380-89 .
American Societ y for Testing and Materials, 1916 Rac e Street , Philadelphia PA 19103. [No date.]
[Anonymous]. 1992. Guidelines for measuring and reporting environmental parameters for plant
experiments in growth chambers. ASA E Engineering Practice: ASAE EP411.1. American
Society of Agricultural Engineers, 2950 Niles Road, St. Joseph, Michigan 49085-9659. [Thi s
is Appendix C in thi s book.]
[Anonymous]. 1979 . Metri c Unit s of Measure an d Styl e Guide. U . S. Metric Association, 1024 5
Andasol Avenue, Northridg e C A 91103.
[Anonymous]. 1985. Radiation quantities and units. ASAE Engineering Practice: ASAE EP402.
American Society of Agricultural Engineers, 2950 Niles Road, St. Joseph, Michigan 490859659.
[Anonymous]. 1982 . S I Units Require d i n Societ y Manuscripts. Agronom y New s (March-Apri l
1982, p 10-13).
[Anonymous]. 1988. Use of SI (metric) units. ASAE Engineering Practice: ASAE EP285.7.
American Society of Agricultural Engineers, 2950 Niles Road, St. Joseph, Michigan 490859659.

Summary of the International System of Units (SI Units) 19

Boching, P.M. 1983 . Author' s Guide t o Publication in Plant Physiology Journals. Deser t Researc h
Institute Pub . No . 5020. Reno , Nev .
Buxton, D.R., an d D.A. Fuccillo . 1985 . Lette r to the editor. Agronom y Journal 77:512-514. [Thi s
letter include s a summar y of a survey of 97 journals; 7 7 percent eithe r require d o r encourage d
the us e o f S I units.]
Campbell, G.S. , an d Ja n va n Schilfgaarde . 1981 . Us e o f S I unit s i n soi l physics . Journa l o f
Agronomic Education . 10:73-74 .
CBE Styl e Manua l Committee . 1994 . Scientifi c styl e and format : th e CB E manua l for authors ,
editors, and publishers . 6t h edition . Cambridg e Universit y Press, Cambridge , Ne w York. [Se e
also earlie r edition s o f CBE Styl e Manual.]
Downs, Robert J. 1988. Rules for using the International System of Units. HortScience 23: 811812.

Goldman, Davi d T., and R.J. Bell , editors. 1986 . Th e International System of Units (SI). Nationa l
Bureau o f Standards Specia l Publicatio n 330. U . S. Department o f Commerce/National Burea u
of Standards. [Se e Taylo r (1991 ) for th e mos t recent version of this publication.]
Incoll, L.D., S.P . Long, and M.R. Ashmore. 1977 . S I units in publications in plant science. Curren t
Advances i n Plant Sciences 9(4):331-343 . [Thi s article recommended severa l practices that ar e
now in wide use b y plant scientists. I t was a kind o f historical turning point.]
ISO Standards Handbook. 1993 . Quantities and Units. Internationa l Organization for Standardization, Geneve . [Thi s i s th e highl y authorative , second-leve l reference . I t i s availabl e fro m
American Nationa l Standards Institute , 11 West 42n d Street, Ne w York, NY 10036. ]
Mills, Ian, Tomislav Cvitas, Klaus Homann, Nikola Kallay, and Kozo Kuchitsu. 1995. Quantities, Units and Symbols in Physical Chemistry 2nd Edition. Blackwell Scientific Publications,
Oxford, London, Endinburgh, Boston, Palo Alto, & Melbourne.

Monteith, J.L . 1984 . Consistenc y an d convenienc e in th e choic e of unit s fo r agricultura l science .

Experimental Agriculture . 20:105-117.
Petersen, M.S. Decembe r 1990 . Recommendation s for use of SI units in hydraulics. Journa l of th e
Hydraulics Division, Proceedings o f the America n Society of Civi l Engineers 106:HY12.
Savage, M.J . 1979 . Us e o f th e internationa l syste m o f unit s in th e plan t sciences . HortScienc e
14:493-495.
Salisbury, F.B . 1991 . System Internationale: Th e us e o f SI unit s i n plan t physiology . Journa l of
Plant Physiology 139(l):l-7.
Taylor, Barry N., editor. 1991 . Th e International System o f Units (SI). Nationa l Institut e of Standards and Technology Specia l Publicatio n 330 . U.S . Government Printing Office , Washington ,
D.C. [Thi s i s th e Unite d State s editio n o f th e Englis h translation of th e sixt h edition o f "L e
System Internationa l d'Unite s (SI)" , th e definitiv e publicatio n of th e Internationa l Bureau o f
Weights an d Measure s an d thu s the first-leve l authority. Ther e i s also a Britis h versio n wit h
slight differences , a s i n th e spellin g o f "metre, " "litre, " an d "deca. " Th e Unite d State s
version i s fo r sal e b y th e Superintenden t o f Documents , U . S . Governmen t Printin g Office ,
Washington, DC 20402. ]
Taylor, Barr y N . 1995 . Guide for th e Us e of th e International System o f Units (SI). Nationa l
Institute of Standards and Technolog y Special Publication 811. [Alon g with the ISO Standards
Handbook, thi s publicatio n shoul d b e considere d secon d i n authorit y onl y t o "L e System
International d'Unites (SI),* a t leas t for citizens o f the Unite d States. ]
Thien, S.J., an d J.D. Oster . 1981 . Th e international system of units and its particular application in
soil chemistry. Journa l o f Agronomic Educucatio n 10:62-70.
U.S. Metri c Association. 1993 . Guid e t o th e Us e of the Metri c System [SI Version]. U.S . Metric
Assocation, Inc. , 10245 Andaso l Avenue, Northridge, CA 91325-1504 .
Vorst, J.J. , L.W . Schweitzer , and V.L . Lechtenberg . 1981 . Internationa l system o f unit s (SI) :
Application t o crop science. Journa l of Agronomic Educucation 10:70-72 .
Weast, Rober t C. , editor. (199 5 an d ne w editions each year). CR C Handboo k of Chemistry an d
Physics. CR C Press , Boc a Raton , Fla .

20 Th

e Basics
CONSULTANTS

Bruce G . Bugbe e Joh

Utah Stat e University Kenned
Logan, Uta h
Louis Sokol * Nat
Boulder, Colorad o Gaithersburg

n Sage r
y Space Center , Florid a
Barry N . Taylor *
. Inst . of Standards & Technolog y
, Maryland

*Dr. Soko l i s president emeritu s o f th e U.S. Metri c Association an d a member of the National Conference o n
Weights an d Measures . H e i s a certified metricatio n specialist Dr . Taylor i s the U. S. representative on th e CGPM .

2
RULES FOR BOTANICAL NOMENCLATURE
John McNeil l
Royal Ontari o Museum
100 Queen's Park
Toronto, M5S 2C6, Canad a
and
Mary E. Barkwort h
Biology Departmen t
Utah Stat e Universit y
Logan, Uta h 84322-5300, U.S.A.
The following discussion provides some recommendations for documenting the plant
material use d i n experimenta l an d othe r studie s an d summarize s th e rule s o f
nomenclature tha t hav e been establishe d a t botanica l congresses hel d ever y five o r
six years for ove r a century (for the mos t recent editio n o f the rules, see Greuter e t
al., 1994) .
1. DOCUMENTATION

It i s imperativ e tha t th e plan t o r funga l materia l use d i n an y experimen t b e

documented. Th e source of the seeds, plants, or cultures used should be cited i n the
publication, eithe r b y indicatin g th e supplie r (e.g. , commercia l source , cultur e
collection) an d includin g any cultivar or strai n identification, or else, in the cas e of
material obtained fro m th e wild, by a statement of the precise geographical location .
In addition , i n comparativ e studies , o r i n thos e i n whic h th e materia l woul d b e
difficult o r impossibl e t o replicat e (e.g. , plant s obtained fro m mos t wil d sources) ,
representative materia l shoul d b e deposite d i n a recognize d herbariu m or cultur e
collection, a s appropriate . Th e herbariu m specimen s should includ e plant s a t
reproductive maturit y plus representative materia l o f any other stage s use d i n th e
study. I f growing the plant s to th e reproductiv e stage is not feasible , the n materia l
from a s matur e a plan t a s possible shoul d b e used . Th e nam e and location o f th e
herbarium or culture collection wher e the specimens have been deposited shoul d be
reported. Thi s can be done concisely by using the internationally accepted abbrevia tions give n in Index herbariorum (Holmgre n e t al. , 1990) , o r i n th e World Directory
for cultur e collection s (Staine s e t al. , 1986) . Th e Curato r o f you r institutiona l
21

22 Th

e Basics

herbarium wil l b e abl e t o provid e advic e o n ho w t o collec t an d preserv e plan t

material fo r depositio n i n th e herbarium . Usefu l advic e ca n als o b e foun d i n
Fosberg & Sachet (1965) , Le e et al.(1982), Radfor d (1986), Savile (1973), and Smith
(1971), or fo r fungi , i n Hawksworth (1974) .
2. TAXONOMIC GROUPS (TAXA; SINGULAR: TAXON)

All plant s ar e assigne d t o species, th e specie s t o genera (sing, genus), an d th e

genera t o families. Althoug h difference s of opinion a s to circumscriptio n o f som e
species, genera , an d familie s exist amon g taxonomists, there i s good agreemen t o n
their limit s fo r mos t flowerin g plants. Withi n some species, infraspecifi c tax a may
be recognized , usuall y a t th e rank s o f subspecies (subsp. ) o r variety (var.). Th e
major rank s above th e famil y (Latin : familia) i n ascending order are: Order (ordo),
Class (classis), Division (divisio) o r Phylum, and Kingdom (regnum). Classification s
at these level s are more controversial, and reference to them is not usually necessary
in physiological researc h publications , unles s the researc h involve s comparison o f a
broad spectru m o f plants. I n further discussio n of nomenclatural practice, w e shall,
therefore, conside r onl y taxa at th e ran k of family an d below. A fulle r accoun t fo r
the genera l biologis t o f th e us e o f scientifi c name s o f plant s i s t o b e foun d i n
Gledhill (1985) .
3. FORM OF SCIENTIFIC NAME S

A. Family names. Famil y names are plura l nouns. The y should be written ou t
in full , wit h th e initia l lette r capitalized , bu t the y ar e usuall y no t italicize d o r
otherwise se t ou t fro m th e res t o f th e tex t i n publication s fro m English-speakin g
countries. Famil y names , apar t fro m nin e exceptions , ar e base d o n th e ste m o f a
generic nam e t o whic h th e suffi x -acea e i s attached . Eigh t o f th e exception s ar e
simple alternatives : Crucifera e [alternativ e base d o n genu s = Brassicaceae] ,
Compositae [ = Asteraceae] , Graminea e [ = Poaceae] , Guttifera e [ = Clusiaceae] ,
Labiatae [ = Lamiaceae] , Leguminosa e [ = Fabaceae ] (bu t se e below), Palma e [ =
Arecaceae], and Umbellifera e [ = Apiaceae] . Th e alternativ e name s ma y be use d
instead o f th e standar d for m bu t nee d no t be . I t is , however , desirabl e t o b e
consistent withi n a paper (i.e. , do not use , fo r example , Poaceae an d Leguminosa e
in the sam e paper) .
The nint h alternativ e name , Papilionaceae , ca n b e use d fo r th e papilionoi d
legumes i f they are regarde d a s a famil y distinc t fro m th e caesalpinioi d an d mimosoid legumes . Th e standar d form s fo r thes e thre e group s o f legumes , i f eac h i s
regarded a s a family , ar e Fabacea e [ = Papilionaceae] , Caesalpiniaceae , an d Mimosaceae. Th e nam e Leguminosa e (Fabacea e i n th e broa d sense ) canno t b e use d if
these three units are treate d a s distinct families .
B. Names of genera. Generic names ar e compose d o f a singl e word . The y
should be italicized, underlined, or se t off in some other wa y from regula r tex t (e.g. ,
written i n roma n i f th e tex t i s italicized) , and hav e th e initia l lette r capitalized .
They ar e singula r nouns, not adjectives . The y should be written out i n ful l unles s
they are use d i n combinatio n with a specifi c epithe t a s the nam e of a species (se e
next item) .

Rules for Botanical Nomenclature 2

C. Names of species. Th e nam e o f a specie s i s a binomial . I t consist s o f th e

name o f th e genu s followe d b y a singl e specific epithet. Th e epithe t (calle d th e
"species name " in zoology ) ma y be hyphenated but i s never tw o separate words. I t
may b e a n adjective , o r a nou n i n apposition , o r i n th e genitive . Th e entir e
binomial shoul d b e italicize d o r se t of f i n som e othe r wa y from th e mai n text .
Underlining i s commonly used when italics ar e no t available .
As noted above , th e initia l lette r o f the generi c nam e must be capitalized. Th e
initial letter of the specific epithet should not be capitalized, although capitalization
is permitted if : (a ) th e epithe t is derived fro m th e nam e of a person (e.g. , Plantago
Tweedyi), (b ) i t i s derived fro m a vernacular nam e (e.g., Dolichos Lablab), o r (c ) i t
was once a generi c nam e (e.g. , Picea Abies). Us e o f lowercase is , however, recom mended i n all cases; i t is never incorrect .
When writin g th e nam e o f a species , bot h th e generi c nam e and th e specifi c
epithet mus t be given . Th e generi c nam e may be abbreviate d to th e initia l lette r
followed b y a period unles s i t i s being used for the firs t tim e in a text or ther e i s a
possibility o f confusion because tw o genera unde r discussion have the sam e initia l
letter. I n th e latte r case , a uniqu e abbreviation consistin g of the initia l lette r an d
one o r mor e others is sometimes use d fo r each o f the generi c names concerned.
D. Names of infraspecific taxa. Th e name of an infraspecific taxo n is a combination o f fou r words , th e generi c name , th e specifi c epithet , th e ter m denotin g
infraspecific rank , an d th e (final ) infraspecifi c epithet . (I t i s possible , althoug h
uncommon, to hav e a hierarchy of infraspecific ranks ; e.g., a subspecies with several
varieties.) Th e ter m denoting rank (e.g., var., subsp.) should be in the same font a s
the bul k of the text , but th e other words are italicize d (or set off in some other way
from th e mai n text), e.g., Stipa nelsonii subsp . nelsonii, Phyllerpa prolifera var. firma,
Trifolium stellatum f. nanum. A s with specific epithets, lowercase should be used for
infraspecific epithet s (bu t see "Name s of Cultivars" below).
E. Citation of Authorities. T o b e accurat e an d complete , i t i s necessar y t o
indicate th e name s o f th e author(s ) wh o firs t validl y publishe d a give n nam e o r
combination. Thi s is usually done by citing them the firs t tim e the plan t or fungu s
name i s use d i n th e text , afte r whic h the nam e ma y be use d withou t citatio n o f
authors. Alternatively , i n paper s treatin g man y specie s fro m a particula r area , a
statement t o th e effec t tha t th e scientifi c nomenclature follows tha t use d i n a well
known Flor a o r Manua l fo r th e are a i s generall y acceptable an d ma y b e mor e
informative. I n addition it i s not necessar y to cit e the author s of genera o r tax a of
higher ran k unles s a pape r specificall y addresse s th e taxonom y of highe r ranks .
Likewise, fo r infraspecific ranks, it is not generall y necessary to cite the authorshi p
of th e specie s name , e.g. , Stipa nelsonii subsp. dorei Barkwort h & Maze. Th e onl y
exception t o thi s i s when th e fina l infraspecifi c epithet i s the sam e a s tha t o f th e
specific epithe t (so-calle d autonyms) . I n such instances , th e autho r of th e specie s
name is given, e.g., Stipa nelsonii Scribner subsp . nelsonii. As scientific names are i n
Latin (o r treate d a s Latin), th e ampersan d (& ) o r th e Lati n word 'et' shoul d b e
used whe n mor e tha n on e autho r i s involved , never th e wor d 'and ' o r it s equivalents in other modern languages. Parenthese s indicate that the taxo n was originall y
named i n anothe r genu s o r a t a differen t ran k but wit h th e sam e (final ) epithet .

24 Th

e Basics

The nam e o f th e perso n wh o firs t validl y published th e combinatio n bein g use d

appears t o the right of the parenthetica l authors . Fo r example, Agropyron cristatum
(L.) Gaertne r show s that th e taxo n was first name d by Linnaeus ("L.") , who coine d
the epithet cristatum for it, but tha t he adopted a different taxonomi c treatment; in
this case he included i t i n another genu s (Triticum). Gaertne r wa s the firs t perso n
to combin e th e epithe t cristatum with th e generi c nam e Agropyron. I n botanica l
nomenclature, unlik e the practice in zoology, the person publishing the combinatio n
being used , a s well as the origina l author(s) , mus t be cited .
If th e name s o f author s ar e connecte d b y 'ex, ' i t mean s tha t th e author(s )
named afte r th e 'ex ' wer e responsibl e fo r valid publicatio n o f th e name , but the y
attributed th e nam e t o th e author(s ) whos e name(s ) preced e th e 'ex. ' Fo r
example, Carex stipata Muhlenb . e x Willd . mean s tha t Willdeno w publishe d th e
combination bu t attributed it to Muhlenberg who, although he had used the epithet ,
had not, i n fact, previously validly published it. I f one wishes to abbreviate the cita tion, on e should retai n the name s of those actuall y publishing the combination; i.e ,
those afte r th e 'ex ' (i n thi s cas e 'Willd.') . Authors ' name s ar e sometime s
connected b y the wor d 'in' . Thi s implie s that th e firs t perso n actuall y named th e
taxon an d provide d th e description , bu t tha t i t was published in a work written by
the author(s ) name d afte r th e 'in; ' e.g. , Viburnum ternatum Rehde r i n Sargen t
means that Rehde r describe d an d named the species, bu t that it was published in a
larger wor k writte n b y Sargent . I n suc h circumstances , th e 'in ' an d th e nam e
following i t are not strictl y part of the author citation and are better omitte d unles s
the plac e of publication i s being cite d (i n this case retainin g only Rehder).
The name s o f author s ca n b e abbreviated , bu t t o avoi d confusion , one shoul d
follow the abbreviation s use d in some standard work (e.g. Hortus Third [Baile y Hor torium, 1976] , the Authors o f Plant Names [Brummit t & Powell, 1992 ] o r a major
Flora of the area) . Abbreviate d name s are followe d b y a period (e.g. , Willd . is an
abbreviation fo r Willdenow in the exampl e above).
In th e case s o f name s o f fungi , ":Pers. " o r ":Fries " afte r th e nam e o f th e
author indicate s tha t suc h name s wer e sanctione d fo r us e b y Persoo n o r Fries ,
respectively, an d hav e a preferred nomenclatural status (Hawksworth, 1984) .
4. SPECIAL SITUATIONS

A. Names of Hybrids. A hybri d between tax a may be referre d t o b y placing a

multiplication sig n x betwee n the names of its two parental taxa; e.g.,Agrostis L. x
Polypogon Desf. , Polypodium vulgare subsp . prionodes Roth x subsp . vulgare. Som e
hybrids hav e bee n give n a nam e of thei r own . Thei r hybri d status i s indicate d b y
placing a multiplicatio n sig n immediatel y before th e name , e.g . xAgropogon P .
Fourn. ( = Agrostis L. x Polypogon Desf.) , Mentha xsmithiana R. A Graha m ( = M .
aquatica L . x M . spicata L. ) I f the mathematica l symbol is not available , a lower
case 'x' should be used (not italicized) and a single space inserted between it and the
name to promot e clarity ; e.g., Mentha x smithiana R.A Graham .
B. Controversial or Unfamiliar Names. I f there is controversy over the nam e of
a taxon , o r i f on e i s usin g th e correc t bu t stil l unfamilia r nam e fo r a taxon , a
familiar alternativ e nam e (synonym ) shoul d b e give n withi n squar e bracket s (o r

Summary o f Rules for Botanical Nomenclature 2

otherwise indicate d parenthetically ) immediatel y after the first mention o f the name ;
e.g., Achnatherum hymenoides (Roeme r & Schultes ) Barkwort h [ = Oryzopsis
hymenoides (Roeme r & Schultes) Ricke r o r Stipa hymenoides Roemer & Schultes] ;
or Elymus lanceolatus (Scribner & J. G . Smith) Goul d [ = Agropyron dasystachyum
(Hooker) Scribne r & J. G. Smith] .
C. Names of Cultivated Plants. Th e name s of cultivated plant s follo w the rule s
of nomenclatur e fo r othe r plant s i n s o fa r a s thes e ar e applicabl e (e.g. , Triticum
aestivum L . for th e commonl y cultivate d species o f wheat), but name s of cultivate d
varieties or race s (terme d "cultivars") ar e subject to additiona l rules. Th e nam e of
a cultivar follow s that of the lowes t botanical rank to which it can be assigned. Fo r
example, cultivar s o f whea t woul d hav e th e cultiva r nam e give n afte r Triticum
aestivum, bu t fo r hybri d te a ros e cultivars , which ar e th e resul t o f extensiv e inter specific hybridization , the cultiva r nam e would follow th e generi c nam e Rosa.
The cultivar nam e is not italicized, but its initial letter is in uppercase. I t should
be pu t betwee n singl e quotatio n marks , e.g. , Taxus baccata 'Variegata' ; unti l
recently i t coul d als o b e precede d b y cv . (fo r cultivar) , e.g. , Taxus baccata cv .
Variegata. Th e grou p o f cultivar s t o whic h it belong s ma y also b e indicated , e.g. ,
Rosa (Hybri d Tea ) 'Peace' .
The name s o f graft-chimera s consis t o f th e name s o f th e components , i n
alphabetical order , connecte d b y th e additio n (plus ) sign : "+ " (e.g. , Cytisus purpureus + Laburnum anagyroides; Syringa xchinensis + S . vulgaris). Fo r furthe r
information o n th e name s o f cultivated plants , see Trehane e t al . (1995).
D. Pleomorphic Fungi. Fung i with differen t phase s i n thei r life-cycl e can hav e
different name s applie d t o thei r various states. Th e fungu s i n all its parts is known
by the nam e of the sexually reproducing stage (teleomorph) , but , where convenient ,
separate name s ca n b e use d fo r th e stage s reproducing b y asexual method s (ana morphs). Anamorp h name s mak e clea r th e phase of the fungu s tha t has been use d
in physiologica l studie s an d s o should b e cited wherever appropriate .
E. Commo n Names . Commo n name s (o r specially formed name s in vernacular
languages; e.g. , English ) ar e permitte d i n mos t journal s o f plan t physiolog y an d
related sciences , but th e scientific name and its author(s) should always be stated in
parentheses immediatel y following th e firs t us e of the common or vernacular name.
REFERENCES
Bailey Hortorium . 1976 . Hortu s third . Macmillan , New York; Collie r Macmillan , London.
1290 p .
Brummitt, R.K. , an d C.E . Powell . 1992 . Author s o f Plant Names . Roya l Botani c Gardens , Kew .
p 732 .
Fosberg, F.R. , an d M.-H . Sachet . 1965 . Manua l for Tropical Herbaria . Internationa l Burea u fo r
Taxonomy an d Nomenclature , Utrecht . p 13 2 (Regnum veg. 39).
Gledhill, D . 1985 . Th e Name s of Plants. Cambridg e University Press, Cambridg e & New York.
Greuter, W. , F.R . Barrie , H.M . Burdet , W.G . Chaloner , V . Demoulin , D.L. Hawksworth , P.M .
Jorgensen, D.H . Nicolson , P.C. Silva , P . Trehane, an d J . McNeill. 1994 . Internationa l Code o f
Botanical Nomenclature (Tokyo Code). Koelt z Scientific Books, Konigstei n Germany. (Regnum
veg. 131) .

26 Th

e Basics

Hawksworth, D.L . 1974 . Mycologist' s Handbook . Commonwealt h Mycological Institute, Kew.

Hawksworth, D.L . 1984 . Recen t change s in the internationa l rule s affectin g th e nomenclatur e of
fungi. Microbiologica l Science s 1:18-21.
Holmgren, P.K. , N.H . Holmgren , an d L.C. Barnett . 1990 . Inde x Herbariorum. Par t 1 . Th e Her baria o f th e World , ed . 8 . Ne w York Botanica l Garden, Bronx , New York. 69 3 p (Regnum
veg. 120) .
Lee, W.L. , B.M . Bell , and J.F. Sutton , editors. 1982 . Guideline s for Acquisition and Management
of Biological Specimens. Associatio n o f Systematic Collections, Lawrence, Kansas.
Radford, A.E . 1986 . Fundamental s of Plant Systematics. Harpe r an d Row , Ne w York. 49 8 p.
Savile, D.B.O. 1973 . Collectio n an d Car e of Botanical Specimens. (Reprin t with addendum) Publ .
1113. Agricultur e Canada , Ottawa .
Smith, C.E . 1971 . Preparin g Herbarium Specimen s o f Vascular Plants. Agricultura l Information
Bulletin 348, Agricultura l Research Service , Unite d State s Departmen t of Agriculture.
Staines, J.F., V.F . McGowan , and V. BD. Skelman. 1986 . Worl d Directory of collections of microorganisms, ed . 3 , 678 p. Worl d Dat a Center , Brisbane .
Trehane, P., C.D. Brickell , B.R. Baum , W.L.A. Hetterscheid , A.C . Leslie , J. McNeill , S.A. Spongberg, and F . Vrugtman, editors. 1995 . Th e Internationa l Code o f Nomenclature for Cultivated Plants 1995. Quarterjack Publishing, Wimborne, U.K. p 175 (Regnum veg. 133).

Consultants
Werner Greute r Noel
Botanischer Garte n Ne
Berlin, German y Bronx

H. Holmgren
w York Botanical Garde n
, New York

David L . Hawkswort h *
CAB-International Mycological Institut e
Kew, England

* Th e author s of this section wis h t o expres s special thanks to Professo r Hawksworth for hi s additions o f material o n funga l nomenclature , which were particularl y helpful.

3
STATISTICS
Donald V . Sisson
Agricultural Experimen t Statio n an d
Department o f Mathematics & Statistic s
Utah Stat e Universit y
Logan, Uta h 84322-481 0
1.

GENERAL TERMS

experimental unit Tha t entit y t o whic h a given treatmen t i s applied. Example s

include a tre e spraye d wit h a give n chemica l o r a petr i dis h containin g see d i n a
particular medium . I n the latte r example, the dish is the experimenta l unit , even if
there are severa l seed s i n th e dish , an d measurement s ar e mad e o n th e individua l
seeds. Th e seed s ar e samples of the experimenta l unit .
experimental error (or MSE) Variabilit y among experimental units that have been
treated alike . Sinc e man y procedures assum e equal variances within the treatments ,
the bes t estimat e of experimental error involve s combinin g o r poolin g th e within treatment variability . Thi s estimat e i s usuall y called th e mean square error, o r
simply the MS E (see pooled variance below) .
replication Th e repeating of the application of a given treatment to more than one
experimental unit . I n the petr i dis h example of the definitio n of experimental unit ,
the seed s ar e no t replication s bu t ar e samples. Thes e sample s ar e sometime s
referred t o a s pseudoreplications.
randomization Th e assignmen t o f treatment s t o experimenta l unit s a t random .
This i s done t o obtai n unbiase d estimates o f the treatmen t effect s an d mean squar e
error. I t remove s persona l bia s or eve n th e appearanc e o f such bias .
local control (often called blocking) A restriction o n the randomizatio n impose d
by th e investigato r i n orde r t o distribut e systemati c variability evenly amon g th e
treatments an d t o reduc e th e unexplaine d variability, or the MSE .

2. MEASURES OF CENTRAL TENDENCY

mean (X ) Th
e arithmeti c averag e o f a set o f values. Thi s i s the mos t efficien t
and common estimate of the "center " of a distribution but it is also affected th e mos t

27

28 Th

e Basics

by extreme value s or outliers . I t i s usually denoted b y

or the su m of all the observations (EX ) divide d b y the number o f observations (n).
median Th e middle observation afte r th e data have been ordered or ranked. I f the
number o f observation s i s a n eve n number , i t i s th e averag e o f th e tw o middl e
numbers afte r ranking . I t i s not affecte d b y outliers.
mode Th e observation tha t occurs with the greatest frequency. I t is not very useful
in smal l samples .
3. VARIABILITY
range (R ) Th

e distanc e betwee n th e larges t an d smalles t observations .

standard deviation (S) Approximatel y th e averag e distanc e fro m th e mea n fo r a

set o f observations. I t i s usually denoted b y

If the dat a ar e normall y distributed, or th e distributio n has the familia r bell-shape d

curve, approximately two-third s o f th e observation s wil l b e withi n on e standar d
deviation o f the mean and approximately 95% will be within two standard deviation s
of the mean .
variance (S2) Th e square of the standard deviation (really just an intermediate ste p
in the calculatio n o f the standar d deviation) ,

coefficient of variation (CV) A measur e o f th e relativ e variabilit y whe n th e

standard deviatio n i s expresse d a s a percentag e o f th e mea n an d th e unit s o
measurement hav e been eliminated .

standard error of the mean (SX) Sinc

e the mea n is itself a variable, it also ha s a
standard deviation . Thi s i s denoted a s

and i s called th e standard error of the mean. Th e standar d error o f the mea n is to
the mea n wha t th e standar d deviation i s to an individua l observation.

Statistics 2

standard error of the difference between two means (Sx1-x2 ) Th e differenc e

between tw o means has a variance that is the su m of the variances of the individua l
means if the two means are independent. Th e standard error of this difference is the
square root o f the variance .

pooled variance I f the assumption of equal variances holds, the information within
groups is pooled t o obtai n

and th e formul a for standard error o f the differenc e betwee n tw o means become s

Note tha t S P i s the sam e a s MS E (mea n square error) an d ca n be expanded t o

accomodate an y number o f groups.
4.

CONFIDENCE INTERVALS

A confidence interval i s a n interva l estimat e constructe d i n suc h a wa y tha t i f a

sampling experiment is repeated a large number of times and an interval constructed
for eac h one , o n the averag e a specified percentage of intervals will contain th e tru e
population value . I f we choose a 95 % confidence level, we usually say that we ar e
95 % confident tha t ou r interva l contain s the tru e population value.
For th e populatio n mean , a confidence interval is found a s follows:
where t is a value from th e table containing Student's t values (in almost all statistics
books) correspondin g t o th e confidence level desired and the degrees of freedom =
n- 1.
For th e populatio n varianc e ( 2 ), a confidence interval is found a s follows :

where th e x 2 (chi-squared ) value s com e fro m a tabl e (foun d i n mos t statistic s

books) correspondin g t o th e appropriat e confidenc e leve l an d wit h degree s o f
freedom = n - 1.

30 Th

e Basics

For a proportion, a confidence interva l i s found a s follows :

where

and Z is the standar d norma l variate, or t with degrees of freedom = .

5.

TEST OF HYPOTHESIS

A hypothesis test i s a procedur e t o determine whethe r a propose d conditio n

(hypothesis) i s reasonabl e o r not :
A. For a Population Mean. Fo r a population mean, u (mu), the conditio n i s
stated a s u 0 where u O is a given value. Fo r example , we could hypothesize that th e
true averag e mas s of a set o f samples was 6.5 g, or u 0 = 6. 5 g. W e us e

The hypothesi s i s rejected i f the calculate d t value exceeds th e value in the t-table ,
with n - 1 degrees o f freedom.
B. Differenc e Between Two Population Means. Fo r the difference between two
population means , th e conditio n i s stated a s
where 8 (delta) i s a given value (usually 0) and

where t ha s n 1 + n 2 - 2 degree s o f freedo m i f th e populatio n variance s ca n b e

assumed t o be equal.
C. Populatio n Variance. Fo r a population variance, the condition i s stated as
where 0

i s a given value, and

22

where X ha s n - 1 degrees o f freedo m an d i s compared to a tabl e o f X values ,

found i n mos t statistic s books.

Statistics 3

D. Two Variances. Fo r tw o variances, th e conditio n i s stated a s

and

where F is compared t o a table of F values found i n most statistics books with n1 -1

and n 2 -1 degrees o f freedom for the numerato r and denominator, respectively.
E. Population Proportion. Fo r a population proportion, (pi)
is stated =
as
, where o i s a give n value, and
O

, the condition

F. Difference Between Two Proportions. Fo r th e differenc e betwee n tw o

proportions, the conditio n i s stated a s 1
-2 = ,
where 6 i s a given value,

and

6.

REGRESSION ANALYSIS

simple linear regression A procedure for relating two continuous variables when
one variabl e (dependen t variable ) i s expresse d a s a linea r functio n o f th e othe r
(independent variable) . A commo n us e i s t o predic t on e variabl e base d o n th e
information provide d by the other . Th e for m o f the equatio n is
where Y

represent s th e predicte d value.

multiple regression A procedur e fo r expressin g on e dependen t variable a s a

function o f two or mor e independen t variables.
least squares techniques On e of the mathematical methods of obtaining estimates
of the terms in a regression equation. Thi s method minimizes the sum of the squares
of the deviatio n o f the observed Y variable (dependent ) from th e value as predicte d
by the regressio n equation .
slope I n th e linea r regressio n equation ,
Y = a + bX
b i s the slop e of the line . I t represents th e predicte d average unit change in Y per
unit chang e i n X . I t i s estimate d (i f th e leas t square s techniqu e is used ) b y th e
formula:

32 Th

e Basics

intercept I

n the linea r regressio n equation ,

Y = a + bX

a i s the Y intercept , o r th e predicte d valu e of Y when X = 0 . Thi s ma y have n o

practical meanin g in man y problems, but i t is still a necessary part of the equation .
It is estimated (i f the leas t square s techniqu e i s used) by the formula:

standard error of estimate Th e squar e roo t o f th e residua l (o r unexplained )

variance i n a regression model . Th e formul a is :

standard error of the slope A measur e o f th e variabilit y of th e slop e o f th e

regression line . I t has the sam e relationship to the slop e as the standard deviation
has to th e origina l variable , X. Th e formula is:

correlation coefficient A measur e o f th e mutua l linear associatio n betwee n tw o

continuous variables . I t is an index as to how closely the actua l points come t o th e
predicted points . Perfec t correlation i s 1 (if the slope is positive) o r -1 (if the slop e
is negative) an d n o correlatio n i s represented b y 0. Th e formul a is :

coefficient of determination Thi s represents the proportion of the variability in Y

(dependent variable ) tha t i s predicted b y X (independen t variable). I t is the squar e
of th e correlatio n coefficient .
7.

ANALYSIS OF VARIANCE

Analysis of variance is a procedure fo r testin g the equalit y of the mean s of tw o o r

more treatments by partitioning the variability into the amount caused by differences
among th e treatmen t mean s an d th e amoun t cause d b y difference s amon g th e
experimental unit s within the treatments .
A. Experimental Designs. Experimenta l designs are th e manne r in which the
treatments ar e assigne d to th e experimental units. Thre e mos t commonly encountered design s are:

Statistics 3

i. Completely Randomized Design (CRD). Th e treatment s ar e assigne d

to experimenta l unit s with no restriction s imposed . Th e linear mode l is
where Yi j represent s an individual experimental unit response ,
p i s the overal l mean ,
ii. (tau ) i s the effec t o f the ith treatment,
Eij (epsilon ) i s the rando m effect associate d wit h the jth experimental
unit assigne d t o the ith treatment .
If w e ar e t o assum e ther e ar e k treatment s wit h n experimenta l unit s i n eac h
group, and le t Y i b e th e averag e of all observations collected fro m experimenta l
units assigned t o i th treatmen t an d Y b e the averag e of all the observations, th e
calculations fo r th e analysis-of-varianc e table are i n Table 1.
Table 1. Analysis-of-variance table for the completely randomized design.
Sources of
Variation

Degrees of

SV

DF

Treatments

k-1

Experimental
Error

k(n-1)

Total

Kn-1

Freedom

Sums of Squares
SS

Mean Square

MS

ii. Randomized Block Design (RBD). Th e experimental units are grouped

or blocked i n such a way that th e variability from bloc k to block is greater tha n
the variabilit y within blocks. Eac h treatmen t occur s once i n each block . Th e
linear mode l is
where Yi j represent s a n individual experimental unit response,
u i s the overal l mea n
ii i s the effec t o f the i * treatmen t
Bj (Beta ) is th e effec t o f th e jth block.
Eij i s th e rando m effect associate d with th e experimenta l unit
assigned t o th e ith treatment and occurrin g in the jth block.

34 Th

e Basics

If w e assum e ther e ar e k treatment s arrange d i n b block s an d le t Y i b e th e

average o f the ith group, Y . b e the average of the jth block, and Y b e the over all average, th e calculation s fo r the analysis-of-varianc e table ar e i n Table 2 .
Table 2. Analysis-of-variance table for the randomized block design.

Sv

DF

Treatments

Blocks

Experimental
Error

Toral

SS

MS

k-1

b-1

(k-1)(b-1)

kb-1

iii. Latin square design. I n the lati n square design the experimenta l units
are groupe d o r blocke d i n tw o dimension s (usuall y designate d a s row s an d
columns) a s oppose d t o on e dimensiona l blockin g i n th e randomize d bloc k
design. Eac h treatment occur s once i n each ro w and once in each column. Th e
number o f treatment s i s equa l t o th e numbe r o f row s an d th e numbe r o f
columns. Th e linea r mode l is
where Yijk represent s a n individual experimental unit response ,
u i s the overal l mean ,
ii i s the effec t o f the ith treatment,
Bj is the effec t o f the jth row,
Yk (gamma ) is the effec t o f the k th column ,
Eijk i s the random effect associate d wit h the experimental uni t in th e
jth ro w and k t h column that was assigned to the i th treatment .
If we let k equal the number of treatments (or rows or columns) an d Y i b e the
average o f the ith treatment, Yj. be th e averag e of the jth row, Y k b e th e aver age of the k th column , the calculation s fo r the analysis-of-varianc e table ar e in
Table 3.

Statistics 3

Table 3. Analysis of variance table for the latin square design.

SV
Treatments

DF

SS

MS

k-1

rows

k -1

columns

k-1

Experimental
Error

(k -1 )(k- 2)

total

k2-1

B. Mean Comparisons. Mea n comparison s ar e procedure s employe d wher e

some o r all of the k treatment s mean s (or averages) are compared in an attempt t o
interpret th e result s of the F-test in an analysi s of variance.
i. Planned comparisons. Planne d comparison s ar e specifi c comparison s
that ar e o f obvious interest, even before th e experimen t is conducted.
a) Factorial experiments. Factoria l experiment s ar e experiment s i n
which th e treatment s consis t o f al l possibl e combination s o f th e differen t
levels o f tw o o r mor e factor s studie d simultaneously . A s a n example ,
consider th e respons e o f a plant to condition s when both temperatur e an d
humidity ar e varied . Le t temperatur e b e factor A wit h a = 3 levels and
humidity b e facto r B with b = 2 levels . Th e resultin g experimen t woul d
have 3 x 2 = 6 treatments where a treatment denote s a particular combination o f temperatur e an d humidity . Assum e that ther e ar e r experimenta l
units in eac h treatment . Tabl e 4 shows the schemati c layout of means.

36 Th

e Basics

Table 4. Means in a two-way factorial experiment.

A = temperatur e
Row Average
B = humidity

Column Averag e
overall
average

Main Effect i s the effec t o f one facto r averaging over the level s o f all of th e
other factors . Thes e ar e teste d i n the analysi s of variance usin g an F tes t
(where th e mea n squar e erro r i s usually th e denominator) , a s indicated i n
Table 5.
Table 5. Partial analysis of variance table for a two-way factorial experimentmain
effects.

SV

df

A main
effect

a-1

B main
effect

b-1

MS

SS

Interaction is the situatio n wher e differences among the levels of one factor, say
factor A , change fro m leve l to leve l o f the secon d factor , say factor B. Th e tes t
for a n interactio n i s also mad e in the analysi s of variance table a s indicated i n
Table 6 .
Table 6. Partial analysis of variance table for a two-way factorial experimentinteraction.
SV

df

AB
in
teraction

(a-b)(b-1)

SS

MS

Statistics 3

Simple effect i s th e effec t o f on e facto r whe n al l othe r factor s ar e hel d

constant. Thes e are not tested directly i n the analysis of variance table. Th e
simple effec t o f B at th e a, leve l woul d be estimated b y Y12. - Y11. . I t
could b e tested by using the concep t o f linear comparisons .
b) Linear comparisons. Linea r comparisons are contrasts between any
2 set s (on e o r mor e mean s i n eac h set ) o f means . Th e simpl e effec t
illustrated abov e is an example, and the tes t would be:

In general, i f the linea r comparison i s of the for m aY 1 bY 2 cY

the variance of the linea r combination i s given by

3,

etc ,

(This assume s tha t th e Y i ' s are independent.)

ii. All possible comparisons.
Fisher's least significant difference test (LSD) . Al l possible differences amon g
the mean s are compare d wit h the LSD value when the

where th e degree s of freedo m fo r the t ar e th e degree s o f freedom associate d

with th e MSE . Thi s tes t ha s a hig h Type I error rat e (whic h also give s a low
Type II error rate. )
Tukey's Test. Th e critica l value is

when q i s a valu e take n fro m a studentize d rang e tabl e (availabl e i n man y

statistical textbooks) . Th e Type I error rate is low (hence the Type II error rat e
is high).
Newman-Keul's Test and Duncan's Test. Test s with intermediate (betwee n th e
LSD and Tukey' s Test ) Type I error rates . Thes e ar e accomplishe d by ranking
the means to be compared an d using different critica l values for different range s
where tw o mean s adjacen t in th e ranking s hav e a range of 2 , one othe r mean
between the m gives a range of 3, etc.

38 Th

e Basics
iii. Orthogonal Polynomials. A
compariso n amon g mean s whe n
regression effects ar e emphasize d and th e objectiv e is to estimate th e for m
of th e response , such as linear, quadratic , cubic, etc. Th e calculations ar e
similar t o linea r combinations , wit h appropriat e weightin g coefficient s
derived fo r eac h ter m i n the polynomial.

C. Variance Components. Variabilit y in a linear mode l is contributed b y two

or mor e effects. I n the mode l

the rando m variability associated wit h the eij effect ca n be designate d a s s e2 .

Likewise, th e variabilit y introduce d b y the treatmen t effects , ti , ca n b e desig nated a s s 2t i f treatments are considere d t o be random. Bot h s 2E an d s2T are
variance component s o f this model .
8.

COVARIANCE ANALYSIS

Covariance Analysis is a combinatio n o f regressio n an d analysi s of varianc e tha t

allows mea n comparison s amon g treatments in the dependen t variable to b e made
after adjustin g for effects o f the independent variable. I n addition, the MSE is based
on deviations fro m a regression mode l rather than deviations from th e mean, hence
the MS E i s usually smaller, and we have a gain i n precision.
As an example of the formulae involved, consider a randomized block design [see
the mode l in equation (29) ] when the amount of nitrogen produced by alfalfa plant s
is measure d unde r differen t moisture-stres s treatments . Eac h experimenta l uni t
consists of 25 seeds. Sinc e germination rates ma y vary, the number germinating may
be used as the independen t variable X. Th e linea r model is

where the ne w term, pXij , i s the effec t o f the germination on that experimental unit.
(See table 6.)

Table 6. Simple linear analysis of covariance table for a randomized block design.
Deviations from Regression
Sv

df

tre atments

k-1

lbocks

experimental

error

treatment plus
experimantal
error

SSx

b-1

(k-1)(b-1)

b(k-1)

(treatment plus
experimental
error)

error
Adj Means
F for testing the equality of adjusted means =

SP

SSy

dF

SS

MS

40 Th

e Basics

9. NONPARAMETRIC TESTS
These are test s tha t mak e a few or n o assumptions regardin g the underlyin g distribution o f th e variable . Th e powe r i s usuall y les s tha n tha t o f a correspondin g
parametric test .
A. Sign Test. A tes t fo r th e media n o f a population . I t classifie s eac h
observation a s being either above (+) o r below (-) the hypothesized median and then
tests to see if the observed proportio n abov e the median , P, differs fro m 0. 5 by using
either standar d binomia l table s o r the norma l approximation to th e binomial :

B. Wilcoxon's Signed-Rank Test. A paired comparison test where the absolut e

differences betwee n pair members ar e ranked, the n reassigned thei r original sign. I f
there is no difference between th e two groups, the expected value of the sums of the
signed ranks should be 0 . A s in many of the non-parametri c tests, a special tabl e is
used t o se e i f the differenc e is significant.
C. Mann-Whitney Two-Sample Test. A test for the equalit y of two populatio n
means where th e dat a for both group s are pooled an d ranked. Eac h rankin g is then
assigned it s accompanyin g grou p identification . Th e su m o f th e grou p wit h th e
smaller sampl e size , R, i s obtained. Th e tes t i s a Z scor e o f the for m

D. Kruskal-Wallis k-Sample Test. A tes t fo r th e equalit y o f the mean s o f k

different samples . I t is the counterpar t of the Analysis of Variance. Al l of the dat a
from th e k group s are ranked a s one combined sample, and the group identification
is then reassigne d t o each ran k value. Th e sums of the ranks, Ri, are then obtained ,
and a chi-squared tes t i s performed as follows :

with k - 1 degrees of freedom.

E. Contingency Tables. Coun t tables where the experimental (or survey) units
are classified according to tw o or more discrete variables in an attempt to determin e
whether th e variable s are relate d o r independent . Ther e ar e man y techniques fo r

Statistics 4

analyzing these table s (categorica l dat a analysis) , but fo r tw o factor s illustrated i n

Table 7 , a chi-squared tes t i s made as follows:

with d f = ( r - l)( c - 1)
or th e expecte d numbe r in the i th ro w and jth column.

where
and c

r = th e numbe r o f rows,
= th e numbe r o f columns.

Table 7. Two-way contingency table.

Germinating

Not Germinating

O11a

O12

Group 1

Rib

R1

Group 2

021

O22

R2

Group 3

O31

O32

R3

C1

C2

nd

Cjc

a Oij represents the number of individuals in the ith row and jth column.
b Ri represents the total of the ith row.
c Cj represents the total of the jth column.
d n is the total sample size.

10.

MISCELLANEOUS

A. Central Limit Theorem. On e o f th e mos t importan t practica l theorem s i n

statistics. I t basicall y says tha t a s the sampl e size increases, th e distributio n o f th e
sample mea n will be norma l with a mean o f u and a standard error of
B. Sample Size. Th e numbe r o f experimental unit s used in each treatmen t o f an
experiment. A n approximatio n t o th e number require d i s
where n i s the require d sampl e size ,
Z i s the standar d norma l variable (1.9 6 i f one i s working with an a-level of
0.05).
s i s the populatio n standar d deviation ,
and D
i s the siz e o f the effec t on e wishe s t o detec t a s a "significant " effect .
Paul N . Hinz
Iowa Stat e University
Ames, Iow a

Consultants
Gary Richardson
Colorado Stat e University
Fort Collins , Colorado

This page intentionally left blank

II
PLANT BIOPHYSICS
The science o f plant physiology relies heavil y upon a variety of biophysical measurements. I t is the goa l of this section t o summarize the symbols, units, and terms tha t
are use d t o expres s th e result s o f thes e measurements . Uniformit y o f expressio n
seems highl y desirable , s o thi s sectio n emphasize s recommende d S I unit s an d
symbols. A t th e sam e time, i t is recognized tha t many plant scientists will continu e
to us e othe r unit s an d symbols , s o acceptabl e alternative s ar e presente d i n a few
cases (e.g. , millimole s pe r lite r a s a n alternativ e t o mole s pe r cubi c meter) .
Definitions o f biophysical term s ar e als o given.
Many physica l parameter s ca n b e considere d a s pairs , wit h on e o f th e pai r
expressing a quantit y an d it s partne r expressin g a potentia l fo r transfe r o f th e
quantity across a barrier. Thus , joules per kilogram are used as units to express th e
quantity of hea t energ y in some substanc e unde r consideration , whil e temperatur e
differences ar e use d t o expres s th e potentia l fo r transfer o f heat fro m on e poin t t o
another (i.e. , from a point o f higher temperature to a point of lower temperature) .
In a thermodynami c system , th e paramete r expressin g th e quantit y is sai d t o b e
extensive (th e valu e i s th e su m o f th e valu e fo r subdivision s of th e system ; e.g. ,
volume), and the parameter expressing the transfer potential is said t o be intensive
(has the sam e value fo r an y subdivision of the system ; e.g., pressure) . Mas s factor s
lend themselve s wel l t o suc h a n analysis . Th e quantit y o f mas s i s expresse d a s
kilograms or moles , whil e the potential s fo r transfe r ar e expresse d i n various ways:
gases pressur
water wate
hydrogen ion s p
in water

e o r partia l pressure (pascals )

r potentia l (pascals , joule s per kilogram )
H unit s

solutes i n water chemica

l potentia l (o r concentratio n o r negativ e loga rithm o f concentration o r activity)
Biophysical measurements i n plant physiology are ultimately dependent upo n th e
concepts develope d i n physics and chemistry. T o a great extent, they depend upo n
thermodynamics; hence , that i s the firs t topi c of this section .

4
BASIC THERMODYNAMIC QUANTITIES
Michael J . Savag e
Department o f Agronomy
University o f Nata l
Pietermaritzburg 320 1
Republic o f South Afric a
The purpos e o f thi s chapte r i s to presen t a simple treatmen t o f some o f th e basi c
thermodynamic concepts involve d in plant physiology, and especially those relatin g
to water potential and its measurement. Th e concepts will be briefly applied to plant
water potentia l an d to water potentia l measuremen t techniques .
1. BASIC CONCEPTS AND THE CHEMICAL POTENTIAL

Purely fro m a n energ y conservatio n standpoint , on e woul d expec t that , fo r a

closed syste m (that is, one with constant mass) , the total (internal) energy change of
the syste m (dE, joules) i s the heat energ y (dQ) added to th e system minus the work
done (dW) by the system :

This expression i s commonly referred to as the first law of thermodynamics. Th e

internal energ y E represent s th e kineti c an d potentia l energie s o f th e molecules ,
atoms, and subatomic particles tha t constitute the (closed) system on a macroscopi c
scale; ther e i s presently no known way to determine E absolutely . Onl y changes in
E, dE , ar e required , however , an d thes e change s ar e normall y determine d b y
experiment. I t is also important to recognize tha t the energy of the system is totally
conserved an d that the energy available for useful work is continually decreasing and
being converted int o energy unavailable for useful work. Indeed , it is this unavailable
energy tha t provide s insigh t int o the concep t o f system entropy .
In simple terms , the chang e of entropy of a system, dS (uni t J K- 1), is the rati o
of th e hea t energ y added t o th e syste m (dQ) to th e thermodynami c temperature T
of the system and, in accordance with thermodynamic principles, is always increasing
for an y rea l system . Thes e tw o statements are ofte n expresse d mathematicall y in
differential for m as :

45

46 Plant

Biophysics

and

Entropy was first introduce d i n classical thermodynamics to provid e a quantitative basi s fo r th e commo n observatio n tha t naturall y occurring processe s hav e a
particular direction . Fo r example , the flow of heat energ y occurs fro m a hotter t o
a coole r region . Equation s 2 an d 3 represen t statement s o f th e second law of
thermodynamics.
Mechanical wor k occur s whe n work done on a system results i n motion . I f F
(with uni t N ) i s the componen t o f the force acting in the directio n o f the displace ment (dl) , th e mechanica l work , dW, equals F.dl. I n this case, d W = F.dl = (F/A)
(A.dl) =
P . d V , where P , whic h i s forc e (F)/are a (A) , i s th e externa l pressur e
exerted o n th e syste m resultin g i n a volume change, dV (= A .dl) .
Combining the first and second law s of thermodynamics (Equations 1 and 2) and
the equatio n d W = P dV describin g th e mechanica l wor k done b y the system , we
have:
This equatio n describe s th e interna l energ y change fo r reversibl e an d purel y
mechanical thermodynami c processes. However , different system s may be subjecte d
to wor k don e b y a numbe r o f force s tha t ma y or ma y not includ e pressur e (i.e. ,
mechanical) wor k forces. Othe r work forces could include work of magnetization,
electrical work, etc. T o allow for the possibility of other work forces involved in th e
closed syste m currentl y considered , w e write:
where dW now represents the total of all other form s of work done by the system on
the surroundings . Thi s work term can be expressed as:

where the jth work term is the product of an intensive parameter Y j an d an extensive
parameter X j (Babcock , 1963 ; Bol t an d Frissel , 1960) . (Se e Chapte r 5 fo r lis t o f
subscripts.)
Including all forms o f work done by the system on the surroundings, the chang e
in interna l energy fo r a closed syste m is expressed by:

In chemical thermodynamics , it is common for the composition of the system to

be varied; i.e. , no t closed . I n such a n ope n system , in addition t o th e variable s of
entropy (S), volume (V), and extensiv e parameter Xj (for given corresponding T, P,
and Yj) , th e compositio n o f th e syste m i s varied . Th e amoun t o f substanc e o f
chemical specie s componen t i , ni (wit h uni t mol) i s used t o describ e th e chemica l
composition o f th e system . Then , choosin g S, V , Xj, and n i a s independent system

Basic Thermodynamic Quantities 4

variables, w e note that th e chang e i n E coul d be due to independen t change s in 5,

V, Xj, an d ni, Th e chang e i n E, fo r example, du e to chang e in 5 only , with V, Xj,
and n i hel d constant, ca n be expressed mathematicall y as:

The term E / S i s called a partial derivative because it expresses th e change

in E with respect t o 5 only.
Hence, dE = (chang e in E wit h respect t o 5 only) + (chang e in E with respec t
to V only ) + (chang
e i n E wit h respec t t o Xj only ) + (chang
e i n E wit h
respect t o ni only).
Hence:

We defin e th e chemical potential ui (J mol- 1) o f th e i th solut e specie s b y th e

partial chang e i n interna l energ y E wit h respect t o n i th e amoun t of substance of
chemical species componen t i, with entropy S, volume V, extensive parameter Xj , an d
other solut e specie s n k (k= i) kep t constant :

From Equatio n 8 and incorporating th e definition o f chemical potential of the

ith solut e species , u i (Equatio n 9) , we get:

2. FREE ENERGY AND WATER POTENTIAL

The Gibbs free energy G (J) i s defined by:

The Gibb s fre e energ y represents th e energy available for usefu l work . Hence ,
the difference in the Gibb s free energ y between two states can be used to predict the
spontaneous directio n fo r a proces s an d indicate s th e usefu l wor k th e transitio n
makes available. Fro m Equatio n 11 , we get:

48 Plant

Biophysics

and hence , from Equatio n 10 ,

so that :

This equation expresse s th e relationship between the Gibbs free energ y and th e
chemical potentia l o f species i. Th e chemical potential of species i roughly indicates
the fre e energ y associated wit h it an d available for performing work. Fo r instance ,
considering th e chemica l potentia l fo r water, u w,

where n k cannot b e nw, then:

where the i' indicate s tha t th e summation cannot includ e th e water component a s
it has already been included . Choosin g T , P, Xj an d n k a s independent variables for
ui, i t can be show n that, where i s a dummy variable in that l . woul d not appea r
if the summatio n were written out :

where

(J mol- 1),

(J mor-1),

and
(J mol- 1).
The quantities Si , Vi , an d Y j ar e partial molar values for entropy, volume, and
intensive paramete r Yj , respectively. Definin g th e i th chemica l species t o b e water ,
we obtai n

Basic Thermodynamic Quantities 4

duw =

d(uw - u

w*

where we define uw* to be the chemica l potentia l o f pure free wate r at a pressure of
101.3 kP a an d a t th e sam e temperatur e a s the wate r wit h chemica l potentia l u w.
Under thes e isothermal conditions, th e temperature difference indicated by dT is zero
so that:

Following integratio n o f Equation 16 , we define:

and
where M w (k g mol- 1) i s the partia l mola r mas s of water, the subscripts m , v , f , an d
n refe r to uni t mass, volume, weight (a force), and amount of substance, respectively ;
and g ( m s- 2) i s the acceleratio n du e t o gravity , V w i s the partia l molar volume of
water, an d m ( J kg- 1), v ( J m- 3, N m- 2 or Pa) , f ( J N-1 or m), and n ( J mol- 1)
refer t o th e specific , volumetric , weight , an d mola r wate r potentials , respectivel y
(Rose, 1979 ; Savage , 1978) .
Water potentia l i s the amount of useful work per unit quantity of water done by
means o f externall y applie d force s i n transferring , reversibly and isothermally , a n
infinitesimal amoun t of water fro m som e standard reference stat e to its position in
the soil, plant , or atmosphere . Th e referenc e stat e i s that of pure free wate r at th e
same temperatur e a s th e wate r i n th e syste m and a t a pressur e o f on e standar d
atmosphere, namely, 101. 3 kP a (adapted fro m Taylo r an d Ashcroft, 1972 , p 153 and
Bolt e t al. , 1975) . Th e S I unit of work is the joul e (J) .

50 Plant

Biophysics

Water potentia l ma y be expressed a s the amoun t o f useful wor k per uni t mass ,
volume, weight, o r amount o f substance (mol ) o f water. Plan t physiologist s us e th e
symbol 7 fo r water potentia l an d usually define it to correspond to a volume basi s
( v ). Som e workers hav e used a mass basis ( m ) and others an amount-of-substance
basis ( n ). I n any system o f units,

where, usin g S I units, p w (k g m-3) i s the densit y of liquid water, where p w = p w(T),
T(C) i s the wate r temperature , an d p w = Mw/Vw .
3. ENTHALPY

The enthalp y (H ) o f a system is defined as :

As in the cas e of the Gibb s free energy , consider th e chang e of the function , i n
this cas e the enthalpy , fro m a n initial equilibriu m stat e t o a fina l equilibriu m state .
Therefore,
dH =

d E + P.dF + V.dP
= dQ + V.dP

where
dE = dQ - P . d F .
Hence, fo r a n isobari c process , d H i s equa l t o dQ , th e hea t energ y amoun t
transferred. I n thermodynamic chemistry where isobaric processes are more important tha n isovolumi c processes , enthalpy i s of greatest use . Fro m th e definitio n of
Gibbs fre e energy , the chang e in enthalpy can be defined via:

4. WATER POTENTIAL IN THE VAPOR STATE

Consider wate r vapo r (whic h ma y be jus t on e componen t o f th e ga s phase )

behaving a s an idea l gas . Then ,
where e (Pa ) i s the partia l water vapo r pressure , an d R (8.314 3 J mol- 1 K- 1) i s th e
Universal ga s constant. W e have, from Equatio n 16,

Basic Thermodynamic Quantities 5

Integrating this equation ove r a vapor pressure range from P = e0 (the saturatio n
vapor pressure ) t o P = e and applying Equation 24:

It ha s bee n assume d tha t isotherma l condition s prevai l durin g the chang e i n

pressure fro m eo to e and that e > 0 kPa.
Substitution o f Equation 1 8 into the equatio n above yields the Kelvi n equation
expressing water potential
=
(Pa ) as a function of fractional relative humidity
v
e/eo:

5. COMPONENTS OF WATER POTENTIAL

Three term s emerg e fro m th e thermodynami c theory a s being component s o f

water potentia l d n ( J mol- 1). I n differentia l form , fro m Equatio n 1 5 applied t o
isothermal conditions , w e have:

It is desirable to partition wate r potential int o components even if there is some

doubt abou t th e partitionin g (Spanner, 1973) . Ignorin g the work term, the secon d
term o f Equation 27 , and writin g V w = d u w / d P an d separating

into a water part an d a non-water part, we have:

52 Plant

Biophysics

The first term, which can be positive or negative, can be written as V w dP an d

expresses th e dependence of the chemica l potentia l o f water, u w, o n pressure P. I n
the olde r literature , thi s ter m i s written as dP but i n mor e moder n literatur e th e
symbol d p i s used. Generally , p i s termed th e pressure potential.
The secon d ter m o f the righ t hand side of Equation 28,

arises from th e contributio n of the dissolved solutes to the chemical potential o f the
water (Dainty, 1976; Slatyer, 1967), commonly referred to as the osmotic component ,
and may be written in traditional notation a s - Vw dpk where p k (Pa ) is referred
to a s th e osmoti c pressur e arisin g fro m th e k th component . I n mor e moder n
literature, th e secon d ter m i s written as V w. d p o r sometime s V W . d S wher e
p (Pa ) i s termed th e osmotic potential and s (Pa ) th e solute potential. Th e old
(or traditional) ter m osmotic pressure, p k, is always positive whereas the more recen t
term osmoti c potentia l p (o r th e solut e potential , s) , i s always negative .
The thir d term o f Equation 28 ,

expresses th e matric component arising from th e solid matter in the system, in which
the chemica l potential o f water is a function o f water content (Dainty , 1976; Slatyer ,
1967), may be written in traditional notation as - Vw . dt wher e - t (Pa ) is referred
to a s the matric potential. Usin g th e notatio n o f the mor e moder n literature , th e
third term of Equation 28 is usually written V w d m wher e m (Pa) is also referred
to as the matric potential. Th e old (or traditional) ter m t is always positive whereas
the more recen t ter m m i s negative.
Integrating Equation 28 and substituting for the various water potential compo nents, we have, with all terms having Pa as their unit:
in traditiona l notation , or , in more modern potential terminology,

Of particular note is the controversy regarding matric potential ( t in Equation

29 and m i n Equatio n 30 ) a s a componen t o f th e total wate r potential . Som e
workers (Passioura, 1980; Salisbur y and Ross, 1991) doubt that matric potential m
can be include d in Equatio n 28 as shown als o in Equations 29 and 30.

Basic Thermodynamic Quantities 5

6. WATER POTENTIAL OF AQUEOUS SOLUTIONS

Applying Equation 27 , valid for isothermal conditions only , under conditions of
constant pressur e an d i n th e absenc e of any work fields , wate r potential becomes a
function o f composition an d concentratio n only . Combinin g Equations 2 6 and 2 7
and considerin g a solution containin g only one solute , sa y NaC1, we get:

where (Pa ) i s related t o vapor pressure (Equatio n 26).

Equation 3 1 form s th e basi s fo r th e us e o f thermocoupl e hygrometers .
Essentially these ar e instruments containing solutes, liquid, and vapor enclosed i n a
sealed cavit y that can be maintained at constant temperature and pressure. Usually ,
the hygromete r measure s th e vapo r pressur e abov e solution , soil , o r plant-tissu e
samples b y the manipulatio n o f energy flow t o an d fro m a thermocoupl e (Savage ,
1982; Savag e and Cass , 1984).
7. THEORY OF THE PRESSURE-CHAMBER APPARATUS
When a transpirin g lea f i s severe d (a t th e petiole) , th e xyle m sa p recedes .
Pressurizing the lea f unti l the water just returns to th e cu t surface give s a measure
of th e hydrostati c pressur e i n th e xyle m (Scholande r e t al , 1965) . Th e pressure chamber apparatu s i s i n fac t analogou s t o th e pressure-membran e (sometime s
referred t o as pressure-plate) apparatus used mainly in soil physics (Passioura, 1980 )
to measur e matri c potential s o f soils an d othe r materials . Th e tw o methods ar e
analogous in that soil matric potential is measured, but in the case of the Scholander
pressure chambe r apparatus , th e matri c potential i n th e apoplas t (o r cel l wall ) i s
measured. Provide d tha t the osmotic or solute potential ( S ) of the apoplastic water
is close t o 0 MPa, the equation:
reduces to

where p is the pressure applied to balance m , the matric potential of the apoplast,
resulting i n a tota l potentia l 7 o f 0 MPa . Hence , th e matri c potential o f th e
apoplast, m , i s equal to - p . I t is usually assumed that the matric potential o f the
apoplast i s equal t o th e tota l wate r potential o f the symplast , s o that th e pressur e
chamber the n measure s the tota l water potential o f the leaf .

ACKNOWLEDGMENTS
This wor k wa s sponsore d b y th e Foundatio n fo r Researc h Development , th e Departmen t o f
Agriculture, an d th e Universit y of Natal South Africa .

54 Plant

Biophysics
REFERENCES

Babcock, K.L. 1963 . Theor y o f the chemica l properties of soil equilibrium. Hilgardi a 34:417-542.
Bolt, G.H . an d M.J . Frissel . 1960 . Thermodynamic s o f soi l water . Netherland s Journa l o f
Agricultural Scienc e 8:57-78 .
Bolt, G.H. , S . Iwata, A.J. Peck , P.AC . Raats , A.A . Rode , G . Vachaud, and A.D. Voronin . 1975 .
Soil physic s terminology. Bulleti n of the Internationa l Soil Science Society 48:26-36.
Dainty, J . 1976 . Wate r relation s o f plan t cells . I n Encyclopedi a o f Plan t Physiology, volume 2 :
Transport i n Plants II: Part A Cells , p 12-35 .
Passioura, J.B . 1980 . Th e meanin g of matric potential. Journa l of Experimental Botany 31:11611169.
Rose, D.A. 1979 . Soi l water: quantities , units, and symbols . Journa l of Soil Science 30:1-15.
Salisbury, F.B . an d C.W . Ross . 1991 . Plan t Physiology , Fourth Edition . Wadswort h Publishing
Company, Belmont , California.
Savage, M.J . 1978 . Wate r potentia l term s and units. Agrochemophysic a 10:5-6 .
Savage, M.J . 1979 . Us e o f th e internationa l system o f unit s in th e plan t sciences . HortScienc e
15:492-495
Savage, M.J. 1982 . Measuremen t o f water potential using thermocouple hygrometers. Unpublishe d
Ph.D. thesis , Universit y of Natal, Pietermaritzburg, South Africa. 16 2 p.
Savage, M.J . an d A . Cass . 1984 . Measuremen t o f wate r potentia l usin g i n situ thermocoupl e
hygrometers. Advance s i n Agronomy 37: 73-126.
Scholander, P.P. , H.T . Hammel , E.D. Bradstreet , an d E.A Hammingsen . 1965 . Sa p pressure i n
vascular plants . Scienc e 148:339-346 .
Slatyer, R.O . 1967 . Plant-Wate r Relationships . Academi c Press, Ne w York.
Spanner, D.C . 1973 . Th e component s o f th e wate r potentia l i n plant s an d soils . Journa l o f
Experimental Botan y 24:816-819.
Taylor, S.A . an d G.L . Ashcroft . 1972 . Physica l Edaphology. W.H . Freema n an d Company , San
Francisco.
CONSULTANTS
Keith L . Bristo w Jac
CSIRO Universit
Townsville, Queensland , Australi a Toronto
Gaylon S . Campbell Georg
Washington Stat e University Wate
Pullman, Washington Pretoria
Alfred Cas s Fran
CSIRO Uta
Glen Osmond , Sout h Australia, Australi a Logan

k Dainty
y of Toronto
, Ontario, Canad a
e C. Green
r Researc h Commissio n
, South Afric a
k B . Salisbury
h Stat e University
, Uta h

5
SOLUTIONS
(IONIC RELATIONS)
Jack Dainty 1
Department o f Botany
University of Toronto
Toronto, Ontario M5S 1A1
Canada
Many event s i n plant s involv e movement s o f substance s a s gase s o r a s solutes ,
molecular o r ionic , dissolve d i n liquids , typicall y water . Th e followin g ar e
recommended symbol s and unit s to be use d in discussion of these movements .
1. ABBREVIATIONS USED AS SUBSCRIPTS AND SUPERSCRIPTS

1 subscrip
in, out subscript
i, o a
co, oc externa
cv, vc
w subscrip
P subscrip
V subscrip
s subscrip
p subscrip
r o r m subscrip
cw subscrip
o, i superscrip

t fo r any component i n a mixture

s o r superscript s denoting the directio n o f a process ; e.g.,
flu x fro m cytoplas m (c) t o vacuol e (v ) or fro m cytoplas m (c) t o
l mediu m (o).
t fo r wate r
t fo r pressur e
t fo r volume
t for solut e
t fo r osmotic potentia l
t for matric potentia l
t for cel l wall
t use d for outside or inside of compartment (e.g., a cell).

A bar over a symbol usually means average; e.g., C=

but i t ca n als o mea n partial molar as in V. .

averag e concentration ,

Current addres s is: Jac k Dainty , Mas Tourriere, F-34270 , Cazevieille , France .
55

56 Plant

Biophysics

2. THE TABLES
Table 1 . Recommende d Unit s fo r Concentration s a, b ( Svmbol C )
solids in solids

mol .kg -1 or mo l mol -1 o r k g kg -1

solids in liquid s

mol-m o r k g m (S I units)
mol-L-1 = M = mola r concentration (no t recommended )
mol- kg -3 = m = mola l concentration (no t recommended) ,
kg -L" (acceptabl e whe n no t a pur e substance ; avoi d mg-mL " ,
etc.)

solids in gases c

mol-m or mol-mol or kg-m-3

liquids in liquids

mol-m o r mol-mo l o r kg- m (S I units)

mol-L" o r L- L o r kg- L (acceptabl e wit h S I units)

liquids in gases c

mol-m or kg-m or mol-mol-1

gases in gases c

mol- mol"1 or mol-m" 3 or m 3- m" 3

Avoid part s pe r million , part s per billion , etc.; L-L" 1
(e.g., /iL-L" 1) is acceptable.

gases in liquids c

Same as gases i n gases.

gases in solids c

mol-mol"1 or mol-kg" 1 or m 3-kg"1

(L-kg"1 is acceptable )

" Thi s table was prepared by F.B. Salisbur y in response to a suggestion o f T.W. Tibbitts .
b
Us e moles for pur e substances ; otherwise , us e kilograms .
c
Whe n volume ( m o r L) is used fo r gases, temperatur e an d pressure mus t be specified .

Table 2. Recommende d Symbol s and Unit s fo r Plant Ionic (Solution ) Relation s

Unit
Parameter
Symbol
concentration

M.
m

mole fractio n
activity
activity coefficien t

i
Xj(orXj)
j)
a

fjJ
v

mol-m"3
mol-L4
(M = molarity ; discouraged)
mol-kg-1
mol-mol"1 (dimensionless)
Same units as correspondin g
concentration
dimensionless; use d when concentratio n
is expressed in mol-m" 3, mol-L" 1, o r
mol-kg ; defined b y a- = f-C- etc .
dimensionless; define d by : a - = v-X-

Continued

Solutions (Ionic Relations) 5

Table 2. Recommended Symbols and Units for Plant Ionic Relations (continued)

Parameter Symbol

Uni

. mo
J-mol

l
"
i
l

partial molar volume

amount of pur e substance n
chemical potential fj.:
electrochemical potentia l J-mo

/-

*i

Note: Chemica l potential for nonelectrolye s or water i s given by:

Ht = n' , + R T I n * j + PF ,
for componen t j. Electrochemica l potentia l has come t o mea n th e chemica l potentia l o f an
ion and i s expressed, for io n j, by:
ftt = $ + tfTlno, + P P, +z,F
In thes e formula e fo r chemica l potentia l an d electrochemica l potential , R i s the ga s constan t
(8.314 J mol" 1 K"1), T i s th e absolut e (kelvin ) temperature , an d F i s th e Farada y constan t
(9.648 x 10 4 C-mol" 1). P i s the pressur e i n Pa, V . th e partia l molar volum e of / i n m 3 -mor 1 ,
Z: i s th e algebrai c valenc y (see entry below) , i p is th e electrica l potentia l i n volts (v), and a - i s
the activit y i n appropriat e units . Th e chemica l and electrochemica l potential s i n th e standar d
states are given by /i,* and ft, * . Fo r nonelectrolyte and ionic solute species, the pressure term, PV j
is usually negligibly small.
electrical potential (o
electrical potential difference

r E)
A# (or AEj

V
V

Note: Electrica l potentia l differenc e i s often symbolize d as V or E; fo r example , th e

membrane potentia l i/r' ' - i/r = A^ is often writte n as V M or E m or V'" or E " .
Strictly, however , th e symbo l E shoul d be reserve d fo r electromotiv e force ; fo r example, th e
Nernst potentia l Ej fo r a n io n (see next entry) . Ej ca n b e considered a n electromotive force .
Nernst potential E

jV

(volt )

Note: Th e symbol s ar e explaine d fo r the chemica l or electrochemica l potential . Th e

superscripts o an d i refer t o th e outsid e an d insid e phases .
algebraic valency z

. dimensionles

Note: Her e use d i n th e sens e o f th e numbe r of electron charges pe r ion . Th e symbols

Z;+ an d 2 " are ofte n use d fo r th e charge s carried b y a catio n and a n anion , respectively .
Continued

58 Plant

Biophysics

Table 2. Recommended Symbols and Units for Plant Ionic Relations (continued)

Symbol

Parameter
mobility

Unit
either: m^s^-V1
or: m-mol'S^-N"1

Note: Th e unit s will depen d o n whether th e drivin g force i s considered a s a voltag e

gradient: m-s'VcV-m" 1), or a s a forc e pe r uni t amount o f ions: m-s'^N-mor 1).
electric current 7
electrical capacitance C
charge Q
electrical resistance R
electrical conductance G
specific electrica l g
conductance

A
F
C
f
S
, g: S-

(amp )
(farad )
(coulomb )
l (ohm)
(siemen )
m

Note: Th e symbol s g, g: can b e considered eithe r a s a slope conductance , <?//<? , or

as a chor d conductance ; e.g. , Ij I (E m - Ej).
transport number t

- dimensionles

Note: Th e transpor t number , t: , is the fractio n o f an electri c current in a solution , or

passing throug h a membrane, fo r instance, carried b y ion/: S t - 1 .
flux 4>
influx <
efflux 0
other (specific ) fluxe

s <

, J mol-m^-s"
jn, <; , J in
out, <0, Joul
oc, 0,,,,, <,, <vc> etc.

Note tha t th e ter m flu x i s used i n transport studie s for amoun t crossing uni t are a pe r
second. Thu s term s suc h a s "rat e o f flux " o r "flu x density " are incorrec t an d shoul d no t
be used .
permeability coefficien t P
diffusion coefficien t D
partition coefficien t K
Michaelis constant

m-s"
irr^-s"
dimensionles

rate constant

A:

velocity (e.g. , of ions ) v

velocity (maximu m rate o f F
transport)
velocity (o f reaction ) v

m-s"
mav mol-m"
mol-s"

1
1

s
mol-mm-3
(or mol-L" 1, molarity, M,
but shoul d b e avoided )
s"'
m3-mol^-s"1, etc .
1

^"1
1

Continued

Solutions (Ionic Relations) 5

Table 2. Recommended Symbols and Units for Plant Ionic Relations (continued)
Parameter
velocity (maximu m o f enzy matically controlle d reaction )
generalized forc e
generalized conductanc e
coefficient
quantity of substance
quantity of isotope
specific activit y

Symbol

Unit
mol-s"1

Vmax

usually J-m^-mol" 1
(or N-mor 1)
mol m"2- s'V'Force"

L
Q

>.

QJ
s

mol
appropriate units;
e.g., th e becquerel , B q (becquerel s ar e
expressed a s s" 1)
appropriate units
e.g., Bq-mol" 1

CONSULTANTS: se e nex t chapte i

6
WATER RELATIONS
Jack Dainty
Department o f Botany
University of Toronto
Toronto, Ontario M55 1A1
Canada
In organism s i n general , bu t particularl y in plants , th e movemen t o f wate r i s of
special importance . It s movemen t b y diffusio n o r bul k flo w follow s th e sam e
thermodynamic principles a s the movemen t of other substances , but it s prevalenc e
in livin g systems has provide d an impetu s for special study and fo r a special se t o f
terms an d measurements . A t present , author s of technical paper s us e one o r th e
other o f th e tw o ways o f expressing the solut e effec t o n th e chemica l potentia l of
water: th e osmotic pressure o r osmotic (solute) potential. Osmotic pressure, which
expresses th e effec t o f solutes as a positive number with dimensions of pressure, i s
the traditiona l approach , but osmotic potential, which considers the solute effect a s
a componen t o f th e wate r potential , expresse s th e effec t a s a negativ e number ,
usually als o wit h dimension s o f pressur e (se e Chapte r 4 on thermodynamics) . I n
spite o f tradition, th e concep t o f potentials seem s most logical fro m th e standpoin t
of thermodynamics, which is the basi s for the universall y accepted concep t o f water
potential. I n th e followin g summar y o f recommended symbols and units , both ap proaches ar e presented , bu t th e us e o f potentials i s strongly encouraged.
1. THE TABLES
The basic recommended terms , symbols, and unit s are summarized in Table 1 . Th e
discussion followin g th e tabl e expand s th e basi c potentia l term s an d give s th e
traditional term s a s well. Tabl e 2 then summarize s the terms , symbols, and unit s
considered i n th e discussion . Tabl e 3 presents othe r term s use d i n discussion s of
plant water relations.

60

Water Relations 6

Table 1. Recommended Terms, Symbols, & Units for Plant Water Relations
Parameter

Symbol

Unit
-1
J.mpl-1

chemical potential of water

Pa

water potential

Pa (Pascal)

components of water
potential: pressure potential
(= hydrostatic pressure),
solute potential, and matric
potential.

Water potentia l i s defined b y

(0.1 MPza = 11 bar; MPa is usually

most approriate but kPA may also
be used; J.mol-1 or J.kg-1 can also
be used.)

where u **w = th e chemical potentia l

(units J.mol-1) of pure water at atmospheric pressure and at the sam e temperatur e
as th e syste m unde r consideration , an d V w = th e partia l mola r volume o f water
equals (1 8 000 mm3.mor-1).
As discussed i n Chapter 4 on thermodynamics, division of the reference chemica l
potential o f wate r (u w - u w*), which ha s unit s o f energ y ( J mol- 1), b y the partia l
molar volum e o f wate r produce s unit s equivalen t t o thos e o f pressure . Thi s i s
illustrated b y the followin g conversions (se e Tabl e 2 in Chapter I of this book):
J.mol-1 = N . m.mol- 1 = m

2.

kg.s- 2.mol-1

Water potentia l i s often writte n as a sum of partial potentials (se e discussio n i n

Chapter 4)
where
P = th e pressur e potentia l (actua l hydrostati c pressure , P ; positiv e o r
negative),
s = the osmotic potential or solute potential (often written p; sometimes,
incorrectly, a s p ; always negative), an d
m = th e matri c potentia l (considere d b y some not t o be a valid componen t
of ; always negative).
The othe r usua l way of writing the equatio n fo r water potential is :
= p - p (- t )
where
and P = water potentia l an d pressure as above,
p = osmotic pressure ( s = -p ; uni t Pa , sometimes osmol.kg-1), and
t = matric potential ( m = -r; unit Pa).

62 Plant

Biophysics

Note tha t osmotic pressure i s numericall y equivalen t t o solute potential bu t

expressed a s a positiv e instea d o f a negativ e value ; ther e ar e als o positiv e an d
negative versions o f matric potential ( *Pm = -T) . Use of osmotic pressure instead of
osmotic potentia l i s traditional an d optional, bu t suc h a usage does no t emphasiz e
that th e paramete r i s one componen t o f water potential . I t i s recommended tha t
osmotic potentia l b e use d i n plant physiology.
Osmotic pressure is , occasionally i n the plan t literatur e an d often in the animal
literature, expresse d i n osmo l kg" 1. A valu e o f X osmol kg" 1 means RT X P a (i n
pressure units) ; tha t is , th e aqueou s solutio n unde r consideratio n ha s th e sam e
osmotic pressur e a s a n ideal solutio n o f molality X mol kg"1.
The matric potential is the (negative ) effec t o f the soli d (and gaseous) phase s o n
the water potential. It s reality is often doubted , particularl y if P and T T are correctl y
interpreted. (Se e J.B. Passioura. 1980 . Th e meanin g of matri c potential. J . of
Exper. Botany 31:1161-1169; F.B. Salisbury and C.W. Ross. 1991 . Plan t Physiology,
Fourth Edition , Wadsworth Pub. Co., Inc; Belmont, California.)
Table 2. Summary of Terms Defined and Discussed in Table 1.

Parameter
water potential
pressure potential (or
pressure)
solute or osmotic potentia l
osmotic pressure

Unit

Symbol
y (usuall y negative, ca n be
positive)

Pa
J-kg 4
Pa

o r P (ca n be + o r - )

J-kg"1
Pa

*,
( V^', always negative)

J-kg"1
Pa

7T

J-kg-1

(S", = ir ;

TT i s always positive)
matric potential

*m

Pa

J-kg"1

Crr;7m = -r>

Tm i s always negative;
T is always positive)
Table 3. Other Terms Used in Discussions of Plant Water Relations.
Parameter

Symbol

volume flux

Jv

solute flux
solute permeability

***,
P, or o>.

Unit
m-s"1

mol-m^-s"1
m-s"1

mol-m^-s'^Pa'1

Continued

Water Relations 6

Table 3. Other Terms Used in Discussions of Plant Water Relations (continued)

Parameter Symbol

Uni

Note: u s i s defined b y the equation : flu x = u> sRT&Cs an d is thus given by <a sRT = P s.
hydraulic conductanc e L

m-s"

-Pa'1

Note: Thi s refer s usuall y to th e cel l membran e an d i s ofte n incorrectl y calle d hydraulic
conductivity. I t i s no t normalize d t o uni t thicknes s o f th e barrier . Th e ter m hydrauli c
conductivity is correctly used when, for example, referring to the conductivity to water of the
cell wall material; th e sam e symbo l is usually used, bu t th e unit s are m 2-s"'-Pa"1.
hydraulic resistanc e R

( = l/L p-A, wher e A = area ) Pa-s-m"

Note: Refer s t o cells , tissues, organs , or entir e plants.

diffusivity D

-*'1

Note: Denote s the "speed " a t whic h changes in water potential propagate within tissue s
and incorporate s cel l and wall conductance s to water and thei r storag e capacities .
diffusional permeabilit y fo r P
water
reflection coefficien t a

j m-s'
dimensionles

Note: Fo r a membran e tha t i s leaky toward th e solut e (i.e. , no t semipermeable) , volum e

flux, J v, i s given, not b y Jv = L p(bP - ATT) , bu t b y / v = LJhP - aAir). Th e reflectio n
coefficient, a , expresse s th e rati o betwee n th e apparen t osmoti c pressur e an d th e
thermodynamic osmotic pressure ; a , i s always less than one, an d usuall y 0 < a < 1 .
volume modulu s o f elasticity e
(for a cell) e

P
i s defined as V(dPldV)

a
3

non-osmotic volume b

osmotic coefficient c/

> dimensionles

viscosity 7

7 N-s-m"

(often give n as % or m - m )

extensibility (o f cell wall) m

rn^N^-s"

kinematic viscosity rjj

- m-s"

thickness of unstirred laye r 8

surface tensio n T

N-m"

64 Plant

Biophysics
CONSULTANTS FO R TH E CHAPTER S B Y JACK DAINTY

Mary A . Bisso n
SUNY - Buffal o
Buffalo, Ne w Yor k

John A Milbur n
The Universit y of New England
Armidale, Australia

Julian Collin s
University o f Liverpool
Liverpool, Englan d

Michael G . Pitman
CSIRO
Dickson, Australia

John Cra m
University o f Newcastl e
Newcastle, Englan d

Ronald J . Pool e
McGill Universit y
Montreal, Quebec , Canad a

Robert F . Davis (deceased )

Rutgers Universit y
Newark, Ne w Jersey

Leonora Reinhol d
Hebrew Universit y of Jerusale m
Jerusalem, Israe l

Dieter Jeschk e
Estenfeld, German y

Roger M . Spanswick
Cornell Universit y
Ithaca, New York

Betty L . Kleppe r
USDA-ARS
Pendleton, Orego n
William J. Luca s
University of California
Davis, California
Enid A . C . MacRobbie
Botany Schoo l
Cambridge, Englan d
E. Marr e
Universita' degl i di Milano Studii
Milano, Italy

Ernst Steudl e
Universitat Bayreuth
Bayreuth, German y
Michel Thellie r
Faculte de s Sciences de Roue n
Mont-Saint-Aignan, France
Alan Walke r
University of Sydney
Sydney, Australia

7
ENERGY TRANSFER
Frank B . Salisbury
Plants, Soils , and Biometeorolog y Departmen t
Utah Stat e University
Logan, Uta h 84322-482 0 U.S.A .
Michael J . Savag e
Department o f Agronomy
University o f Natal
Pietermaritzburg 320 1
Republic o f South Afric a
An excellent exampl e of plant biophysics is the applicatio n o f physical principles t o
understand th e energ y exchang e betwee n a plan t an d it s environment . Here , we
present a summar y table o f terms, symbols, and unit s tha t ar e appropriat e fo r thi s
endeavor followe d by some equation s tha t ar e ofte n used.
1. TERMS, SYMBOLS, AND UNITS APPROPRIATE IN ENERGY-TRANSFER
STUDIES
The symbol s i n Table 1 are mostl y arrange d alphabetically , wit h Roman letter s
first, the n Gree k letter s (bu t som e parameter s hav e alternativ e Roma n o r Gree k
symbols).
Table 1. Terms, Symbols, and Units.
Parameter
Symbol

Unit

absorption coefficient, like reflectio n

coefficient, nee d no t hav e A (wavelength) specified

unitless

absorptance o r absorptivit y

unitless (J-J" 1)

heat energ y storage

J-kg'1, W-m' 2

boundary layer , superscript

bl

conduction, superscrip t

65 Continued

66 Plant

Biophysics

Table 1. Terms, Symbols, and Units (continued)

Parameter
volumetric heat capacity
(at constant volume)
specific heat capacity of dry air (at
constant pressure)

Symbol
c

Unit
J-m^'-C1
or J-nr 3 -K 4
J-kg^'-C 1
or J-kg^-K' 1

m2-s4

emissivity or emittance in infrared

region, for example

eIR and EIR

unitless

water evaporation site, superscript

radiant energy, kinetic energy

diffusion coefficient of species j

Sleaf

nvs"1

coefficient, convective transfer coefficient, heat energy transfer coefficient

heat energy convection

hc

W-nr^-C'1
or W-m^-K'1

Planck's constant

=6.626 x ID'34 J-s-photon 4

a quantum of radiant energy

hv

sensible heat energy transfer

W-m' 2

leaf conductance

heat energy, subscript

infrared

h
IR (near infrared:
800 to 3000 nm)
(far infrared:
3000 to 70 000 nm)

unitless

thermal conductivity coefficient of

region,/

K>

W-nr 2 -C 4
orW-m- 2 -K 4

eddy diffusion coefficient of gaseous

species

metabolic heat energy

foliar absorption coefficient

m2-s4
W-m'2, J-kg 4

photosynthetic irradiance
(photosynthetically active radiation)

PAR

W-m- 2

photosynthetic photon flux (photosynthetically active radiation, photon

basis)

PPF

mol-m'2-s4
(moles of photons per square
meter second)
(usually /imol-m -s )
(also: mol-m^-d' 1 )

partial pressure of gaseous species j

Pa
(usually kPa)
Continued

Energy Transfer 6

Table 1 . Terms , Symbols , and Unit s (continued)

Parameter
vapor pressure, leaf and air

Symbol
e

l>ea

Unit
Pa

net irradiance

W-rrT2

Universal gas constant

8.3143 J-mol^-K' 1
8.3143 m 3 -Pa-mor 1 -K- 1
8.3143X1Q-6 m 3 -MPa-mor 1 -K- 1
(1.987 cal-mol^-K4)
(0.083143 L-bar-mol^-K- 1 )

reflection coefficient

unitless

reflectivity (A indicates wavelength

reflected)

r^ or r(A)

unitless

resistance for gaseous diffusion for

species j

s-m"1

s-m

boundary-layer resistance (a for aerodynamic)

diffusive resistance within a leaf
relative humidity

r or r

s-m" 1

RH

percent (%)

W-m"2

solar irradiance; i.e., global

irradiance

turbulent air, superscript

ta

transpiration, superscript

temperature

K(C)

kinetic energy per amount of

substance

J-mol" 1

ultraviolet

specific latent heat of vaporization;

tranpiration or condensation; specific
latent heat of fusion
velocity, wind speed
water, water vapor, subscripts
distance

UV(UV-A 320 to
400 ran)
(UV-B 280 to
320 nm)
(UV-C <280nm)
KorL

J-kg 4 , W-m' 2

m-s" 1

w, wv

x, or 8 (delta)

Continued

68 Plant

Biophysics

Table 1. Terms , Symbols, an d Unit s (continued )

Parameter

Unit

Symbol

Pa-K'1
Typical value is 66.6
Pa-K'1
(at 20C and 100 kPa:
sea level)

psychrometric constant

Y (gamma)

thickness of air boundary layer

Sbl (delta)

difference or change in the quantity

that follows

A (delta)

emissivity or emittance in infrared

region (as example)

6jR (epsilon)

unitless (J-J" 1 )

H. (lambda)

nm

*max (lambda)

nm

wavelength of radiation
wavelength corresponding to the
maximum absorption coefficient in
an absorption band or to the
maximum photon (or energy)
emission in an emission spectrum
frequency of electromagnetic
radiation
density of dry (unsaturated) air
Stefan-Boltzmann constant

m (usually mm)

v (nu)

s'1, Hz (hertz)

P (rho)

kg-m- 3

a (sigma) or S (delta)

=5.673 x 10'8 W-m^-IC 4

2. SOME EQUATIONS USED IN HEAT-TRANSFER STUDIES

A. The Energy Balance Equation for a Leaf Surface (all values can be expressed
as watts per square meter: W-m"2):

Q + H + V + B + M +A = 0
where
Q=
ne t irradianc e (positiv e i f leaf i s radiating less energy tha n th e radian t
energy absorbed fro m it s surroundings),
H=
sensibl e heat flux transfer (includes conduction and convection; negative
if lea f loses more hea t energ y than it gains),
V=
laten t hea t flux ; th e transpiratio n ter m (negativ e when water i s vaporizing; positive whe n condensing o r freezing) ,
B=
storag e flux (positive whe n leaf temperatur e is increasing),
M=
metabolis m an d othe r factor s (positiv e when heat i s produced), an d
A=
advecte d hea t flu x fro m lea f t o ai r (positiv e for advectio n fro m ai r t o
leaf; advectio n i s the horizonta l flow of airi.e. , wind).
At constan t lea f temperatur e an d ignorin g metabolism an d advectio n (whic h
could b e important but i s difficult t o measure) : Q + H + V = 0.

Energy Transfer 6

B. Radiant Energy Flux Absorbed by a Leaf Surface (Q abs; W.m-2):

Qabs = eQPAR + e'Qth

where
eQPAR = tota l absorbed irradianc e i n the PAR region (W.m- 2),
and

e'Qth =tota l absorbed (thermal ) irradianc e outsid e PA R region (W.m-2) ,

e and e ' = lea f emissivitie s i n the two spectral regions .

C. Radiant Energy Flux from a Leaf (or any) Surface (Qe; W.m-2):
Qe = e s T4
where
Qe = Radiant energ y flux (W.m- 2),
e = emissivit y (abou t 0.9 8 fo r leaves a t growin g temperatures) ,
a = Stefan-Boltzman n constan t (5.67 3 x 10- 8 W.m- 2-K-4), and
T = absolut e temperatur e o f the lea f (K )
This Stefan-Boltzmann Law is applied i n th e nex t equation .
D. Net Irradiance at a Leaf Surface (Q; W.m-2):
Energy flux emitted by a leaf (Stefan-Boltzmann law) is subtracted fro m the absorbe d
radiant energ y flux (Qabs) :
Q = Qabs -

eIRsT4

where
Q = energy flux (W.m- 2)
Qabs

= absorbe d energ y flux (W.m-2), and

eIR = emissivit y or absorptivit y of the lea f fo r long-wave (thermal) radiation; typicall y about 0.9 5 fo r livin g leave s a t norma l temperature s
(same a s e' above) .
Often, th e abov e equatio n i s written (se e Monteit h an d Unsworth, 1990) :
Q = Is - rI, + Lenv - emsT4
where
2
IS = th e sola r irradianc e inciden t a t th e lea f surface (W.m- ),
r = th e lea f surfac e reflectio n coefficien t (decima l fraction) , and
Lenv = th e environmental longwav e irradiance inciden t at th e leaf surfac e
(W.m-2).
E. Sensible Energy Flux Transfer by Convection at a Leaf Surface (H; W.m-2):

70 Plant

Biophysics

where
Ta =

ai r temperature ( K or C) ,

Tl =

lea f temperatur e (K or C) ,

AT = T

a-

Tb

cp =

specifi c hea t capacit y o f dr y (unsaturated ) ai r ( = 100 0 J.kg- 1.K-1)

at constant pressure ,

p=

densit y o f dry air (1.20 5 kg.m- 3 at 20 C an d 10 0 kPa),

ra=

boundary-laye r resistanc e (s.m- 1), and

ga =

boundary-laye r conductanc e (m.s- 1).

The convective transfer coefficient (h c; W.m-2.K- 1), als o calle d th e heat transfer
coefficient (proportiona l t o the reciprocal of the boundary layer resistance), ma y be
used t o calculat e sensible energy transfer H (W.m- 2):

F. Latent Energy Flux of Water Vapor at a Leaf Surface (V; W.m-2), the
Transpiration Term:

where
et = vapor pressur e i n the leaf ; i.e., withi n the substomata l cavit y (Pa) ,
ea = vapo r pressur e o f the air (Pa) ,
ra = boundary laye r resistanc e (i n air) (s.m- 1),
rl = diffusiv e resistanc e withi n the lea f (s.m- 1),
r=

psychrometric constan t (typicall y 66.6 Pa.K- 1), and

gl and g a = leaf an d boundary-laye r conductivitie s (m.s- 1), respectively .

Energy Transfer 7

REFERENCES
Campbell, Gaylo n S . 1977 . A n Introductio n to Environmenta l Biophysics. Springer-Verlag , New
York, Heidelberg , Berlin . 15 9 p.
Gates, Davi d M . an d L a Verne E . Papian . 1971 . Atla s o f Energ y Budget s o f Plan t Leaves .
Academic Press , Londo n an d Ne w York. 27 9 p.
Gates, Davi d M . 1968 . Transpiration an d lea f temperature. Annua l Review of Plant Physiology
19:211-238.
Larcher, Walter . 1995 . Physiologica l Plant Ecology , Thir d Edition . Springer-Verlag , Berlin ,
Heidelberg, New York. (Translate d b y Joy Wieser) 50 6 p.
Monteith, J.L. and M.H. Unsworth. 1990 . Principle s of Environmental Physics. Edwar d Arnold :
London, 29 1 p.
Nobel, Park S. 1983 . Biophysica l Plant Physiology and Ecology. W.H . Freeman an d Company , San
Francisco. 60 8 p. [Th e symbols and unit s used i n this chapter were modified fro m thos e i n this
text book. ]
Raschke, Klaus . 1960 . Hea t transfe r between th e plan t and th e environment . Annua l Review of
Plant Physiolog y 11:111-126.
CONSULTANTS
Donald T . Krizek
USDA Agricultura l Research Servic e
Beltsville, Maryland
John C . Sager
John F. Kennedy Space Cente r
Kennedy Space Center , Florid a

8
PHLOEM TRANSPORT
Donald R . Geige r
Department o f Biology
University o f Dayto n
Dayton, Ohi o 45469-232 0 U.S.A.
Aart J.E . va n Bel
Botanisches Institu t 1
Justus-Liebig Universita t
Senckenbergstrasse 17
D-35390 Giessen, Germany
In this chapte r term s ar e defined and SI units are presented whe n appropriate .
Table 1. Some Terms and Units Used in the Study of Phloem Transport.
Term
Description of Concept
Units
BASIC AND DESCRIPTIVE TERMS:
photoassimilates Organi c compounds produced b y photosynthetic carbon fixation .
translocation Lon

g distanc e transpor t o f solute s throug h sieve tube s o r othe r

structures specialized for longitudinal transport.

allocation a Flo
(partitioning) ica

w o f photoassimilates int o various compartments or biochem l pathway s within sourc e an d sin k regions . I n sourc e organs ,
carbon i s allocate d t o variou s use s includin g export . I n a sin k
organ, carbon enter s into compartments or i s used fo r synthesis ,
storage, or energ y metabolism.

partitioning a Distributio
(allocation)
pressure flow Th
hypothesis em

n o f translocated photoassimilate s among sinks.

e theor y o f osmotically-drive n pressur e flow withi n th e phlo ; pressur e build s u p i n th e siev e element-companio n cel l
complexes i n th e sourc e a s water move s into these sieve elemen t
members i n respons e t o hig h solut e concentration s therein ;
pressures within the siev e elemen t member s ar e les s i n th e sin k
regions a s solute s exi t fro m th e phloem . Firs t propose d b y E .
Munch i n 1930 .

Continued

Phloem Transport

73

Table 1. Some Terms and Units Used in the Study of Phloem Transport (continued)
Term

Description of Concept

pressure flow A

s applie d t o phloe m transport , th e mas s flow o f wate r an d

solutes along a pressur e gradient .

Units

osmotically Flo
w tha t arise s fro m negativ e osmoti c potentia l withi n siev e
generated flow tube s (phloe m translocation ) o r xyle m vessel s (roo t pressure ,
root exudation , guttation) .
mass flow Flo

w o f solute alon g with solvent .

sieve-element/ Cellula
companion-cell branche
complex physiologica

r comple x interconnecte d b y specialize d unilaterall y

d plasmodesmata ; th e comple x act s a s a n integrate d
l uni t i n photoassimilate transport .

apoplast Th

e interconnecting cel l walls and water-fille d xylem elements i n

a plan t throug h whic h wate r an d dissolve d solute s ca n mov e
freely. I n a sense , th e "dead " par t o f a plan t but , fro m a func tional standpoint , excludin g the suberin-fille d Casparian strips.

symplast Th

e interconnected , throug h plasmodesmata , protoplast s o f a

plant. Som e author s woul d exclud e th e centra l vacuoles . I n a
sense, the "living " part o f a plant.

QUANTITATIVE OR
translocation
profile

MEASUREMENT TERMS
A plo t o f solut e concentratio n versu s distance (spatia l profile) or
solute concentratio n a t a particula r locatio n versu s tim e (tem poral profile) .

bidirectional Simultaneou
s transpor t o f solute s i n opposit e direction s i n th e
transport sam
e fil e o f siev e elements . Th e proces s ha s no t bee n demon strated i n th e sens e o f th e definition . Bidirectiona l transpor t
may occu r unde r som e circumstances , fo r example , in a fil e o f
cells as a resul t o f cytoplasmic streaming.
source A

regio n i n which net flux of solut e int o elements i s sufficient t o

cause ne t expor t fro m them .

sink A

regio n i n whic h there i s ne t efflu x o f solute s an d wate r fro m

sieve elements , resultin g in import int o them .

phloem loading Th

e proces s b y which product s o f carbo n assimilatio n enter int o

the sieve-element/companion-cel l complex . Th e proces s ma y be:
a) apoplasti c a s whe n solute s ar e transporte d acros s th e plasm a
membrane of the sieve-element/companion-cel l complex , b)
symplastic a s whe n solute s ar e transporte d fro m surroundin g
mesophyll int o th e siev e elemen t companion cel l comple x
through symplasti c connections , c ) symplasti c an d apoplasti c
when bot h pathway s operate i n parallel or alternatively.

phloem Th
unloading solute

e processe s tha t brin g abou t th e ne t transpor t o f wate r an d

s fro m siev e element s i n sin k organs int o th e surrounding
sink tissues . Passag e ou t o f th e siev e elements may b e symplastic
or apoplasti c or both , depending on th e natur e of the sink .
Continued

74 Plant

Biophysics

Table 1. Some Terms and Units Used in the Study of Phloem Transport (continued)
Term Descriptio
n o f Concept
Units
FLOW TERMS:
phloem export Rat
rate sourc

e o f phloe m translocatio n o f a specifie d solut e ou t o f a

e organ. A basi s fo r comparison suc h as per leaf , per area ,
or pe r plan t shoul d b e specified.

mg-s'1
or
mol-s"-1

phloem import Rat

rate organ

e o f entr y o f a specifie d solut e throug h phloe m int o a sin k

. Th e rat e shoul d b e expresse d o n a suitable basi s suc h as
per specifi c sin k organ , o r pe r fres h o r dr y mas s o f th e sin k
organ.

mg-s"1
or
mol-s"1

phloem mass Flu

flux sectiona

x o f a specifie d solut e throug h a uni t of sieve elemen t cross - mg-m -2- -s 1

l area.

translocation Linea
speed (velocity ) siev

r distanc e travele d pe r uni t time b y the solutio n i n a file of

e element s o r b y a concentration fron t i n a phloe m bundle .

m-s-1

osmotic-potential Th e differenc e i n osmoti c potentia l i n a fil e o f siev e element s

gradient ove
r a specifie d distance o f phloem path .

MPa-nT1

t flux of water enterin g siev e elements .

m'-m-V1

phloem pressur e Th e differenc e in turgor pressur e i n a file of sieve elements ove r

gradient a
specifie d distance.

MPa-m 4

volume flo

w Ne

" Unfortunatel y th e tw o terms ar e use d i n opposite ways by different authors. Car e must be take n t o determine which
way a give n autho r chooses to appl y thes e terms .

CONSULTANTS
Susan Dunfor d Coli
University of Cincinnat i Universit
Cincinnati, Ohi o Gle
Walter Eschric h Joh
GOttingen, German y Universit
Donald B . Fishe r
Washington Stat e University Pete
Pullman, Washingto n Th
R.M. Gifford
CSIRO
Canberra, ACT , Australi a Joh

Lim C . Ho Newcastle
Institute o f Horticultur e
Littlehampton, Wes t Sussex , Englan d Fran

n F. Jenner
y of Adelaide
n Osmond , Sout h Australia, Australia
n A. Milburn
y of New England
Armidale, NSW , Australia
r E . H . Minchin
e Horticulture and Foo d Researc h
Institute o f New Zealand Ltd .
Lower Hutt, New Zealand
n W . Patrick
University of Newcastle
, NSW, Australia
k B . Salisbury
Utah Stat e University
Logan, Utah

9
ELECTROMAGNETIC RADIATION
Donald T . Krizek
Climate Stres s Laborator y
U.S. Departmen t o f Agriculture, AR S
Beltsville, Marylan d 20705-235 0 U.S.A .
John C . Sager
Biomedical Operation s an d Researc h Offic e (MD-RES )
John F . Kennedy Space Cente r
Kennedy Space Center , Florid a 32899-000 1 U.S.A .
An accurate description of the radiation environment used in controlled-environment
and othe r studie s is fundamental to plan t science. Sinc e formal approva l o f the S I
(Systgme International d'Unites) i n 196 0 b y th e Conference Generate de s Poids e t
Mesures, ther e ha s bee n increasin g interes t amon g plan t scientist s i n tryin g t o
standardize terminology used in describing electromagnetic radiation , which includes
the huma n visuall y evaluate d wavelength s calle d light . Th e quantitie s use d t o
describe and evaluate light (e.g., the candela, lumen, lux), however, are not applicable
to plan t physiology . Th e followin g list o f terms , symbols , and unit s is base d o n
recommendations give n i n th e CI E (Commission Internationale d e I'fcclairage)
International Lightin g Vocabular y publishe d i n 198 7 an d i n othe r reference s
attached.
Table 1. Terms, Symbols, and Units Basic to Studies of Radiation".
Quantity
Symbol

Units

RADIATION " INCIDENT ON A FLAT SURFACE (2 DIMENSIONAL):

radiant energy *

Qe

radiant exposure

He

J-m" 2

energy flux (irradiance) E

spectral energy flux * E
(spectral irradiance)

W.m-2
e^

number of photons N
(number of quanta)
Avogadro's number
(mole) of photons

W-m~

dimensionles
Q

-nm" 1
s
mol

75 Continued

76 Plant

Biophysics

Table 1. Terms, Symbols, and Units Basic to Studies of Radiation (continued)

Quantity

Symbol

Units

mol-m-2
mol.m-2.s-1

photon exposure H
pho
spectral photon flux c E

. mol-

m -s^-nm"

RADIATION ARRIVING AT A POINT (3 DIMENSIONAL):

The quantities , symbols, an d unit s defined fo r radiatio n inciden t on a fla t surfac e (two
dimensional) also apply to radiation incident on a point (three dimensional). However , th e
term "fluence" has been defined as the amount of radiation incident on a spherically shaped
receiver. Specifically , fluence i s the integra l of flu x at a point over al l directions about th e
point. I n norma l conditions , where radiatio n come s fro m al l directions, on e mus t us e a
spherical sensor t o measur e fluence.
MATERIAL OR REACTION RESPONSE TO RADIATION:
absorptance, absorption a
factor (ratio of absorbed
to incident radiation)

dimensionles

absorbance A

dimensionles

reflectance (ratio of p
reflected to incident
radiation)

dimensionles

transmittance T

dimensionles

" The term intensity should not be used to describe radiation falling on a surface or a point Intensity (symbol f) refers
to the source', e.g., the sun or a lamp.
The sam e symbo l i s use d fo r th e correspondin g energy (e ) o r photo n (p ) quantit y wit h th e subscrip t use d wher e
confusion migh t occur.
c
Spectral data shoul d b e shown wit h th e abscissa as a wavelength scal e with lo w values t o the left Discret e responses,
such a s actio n o r emissio n spectr a shoul d b e give n i n term s o f photons .

Table 2. Terms Used in Studies on Special Aspects of Photobiology (Some Relate

to the Quantum Theory of Light)
Quantity Symbol

Unit

PHOTOSYNTHESIS:
frequency

&1, Hz

wave number

m'1

wavelength

m (visible spectrum: nm)

fluorescence

F
F

initial
maximum
variable
terminal

dimensionless

* max
F

= F

max ' F<f>

Continued

Electromagnetic Radiation 7

Table 2. Terms Used in Studies on Special Aspects of Photobiology (continued)

Quantity Symbol
Unit
s
quantum yield a
(ratio of effect to
number of photons)

dimensionles

half-peak band-width ^1/2

photosynthetically active radiation PA
(Integral over pho
active wavelengths, 400 to 700 (e.g.
nm) (Se

nm

R Specif

y wavelengt h interval the

first time used
, 40 0 to 70 0 nm)
e nex t two entries for units)

photosynthetic irradiance

PI W-m"

photosyntheti

PPF

mol-m-2.s-1
(usually jumol-m"2^"1)
(sometimes mol- m -d" 1)

PHOTOTROPISM: Al l terminology is referenced t o th e basi c definitions and unit s of electro magnetic radiatio n
PHYTOCHROME (phy):
total phytochrome Plot

= Pf r + Pr dimensionles

far-red-absorbing form Pf

r dimensionles

red-absorbing form P

r dimensionles

fraction of phytochrome present in 4

Pfr form with respect to Ptot at
photoequilibrium

> dimensionles

difference in absorbance at two h

different wavelengths

A dimensionles

change in difference in absorbance A

after irradiation with a second
actinic source

A/1*2 dimensionles
^

" For a detailed description of current phytochrome nomenclature, the reader is referred to Quail et al. (1994)

REFERENCES
ASAE Engineering Practice : ASA E EP285.7. 1988 . Us e o f SI (Metric) Units . America n Societ y
of Agricultural Engineer s (ASAE) , 2950 Nile s Road, St. Joseph, Michiga n 49085-9659.
ASAE Engineering Practice: ASAE EP402. 1990 . Radiatio n quantitie s and units. America n Society
of Agricultural Engineer s (ASAE) , 2950 Niles Road, St. Joseph, Michiga n 49085-9659.
ASAE Engineering Practice : ASAE EP411.2. 1992 . Guideline s for measuring and reporting environmental parameter s fo r plan t experiment s in growth chambers . America n Societ y o f Agricultural
Engineers (ASAE), 2950 Niles Road St. Joseph, Michigan 49085-9659. (See appendix C.)
American Societ y fo r Horticultura l Science Working Group o n Growt h Chamber s and Controlled
Environments. 1980 . Guideline s for measuring and reportin g the environment for plan t studies.
HortScience 15(6):719-720 .

78 Plant

Biophysics

Commission International e d e l'Eclairage . 1987 . Internationa l Lighting Vocabulary. CI E Publ. No.

17.4. Geneve , Suisse .
Downs, R.J . 1988 . Rule s fo r using the International Systems of Units. HortScienc e 23(5):811-812 .
Holmes, M.G. , W.H . Klei n an d J.C . Sager . 1985 . Photons , flux , an d som e ligh t o n philology .
HortScience 20(1):29-31 .
Krizek, D.T. 1982 . Guideline s fo r measuring and reporting environmental conditions i n controlledenvironment studies . Physiologi a Plantaru m 56:231-235.
Krizek, D.T. an d J.C. McFarlane . 1983 . Controlled-environmen t guidelines . HortScienc e 18(5):662 664 and Erratu m 19(1):17 .
McCree, K.J. 1972 . Th e action spectrum, absorbance , an d quantum yield of photosynthesis i n crop
plants. Agricultura l Meteorolog y 9:191-216 .
McFarlane, J.C . 1981 . Measuremen t an d reportin g guideline s fo r plan t growt h chambe r
environments. Plan t Scienc e Bulleti n 27(2):9-ll.
Mitchell, C.A . an d H.C . Dostal . 1977 . Light intensity , footcandles an d lu x are obsolet e terms .
HortScience 12:437-438 .
Monteith, J.L . 1984 . Consistenc y an d convenienc e i n th e choic e of units for agricultura l science.
Experimental Agricultur e 20(2):105-117 .
NBS Technica l Not e 910-2. 1978 . Self-Stud y Manua l on Optica l Radiatio n Measurements , Par t
1Concepts. Unite d State s Governmen t Printin g Office, Washington , DC.
North Centra l Regiona l 10 1 Committe e o n Growt h Chambe r Use . 1984 . Qualit y assuranc e
procedures for accurac y in environmental monitoring-Draft proposal . Biotronic s 13:43-46 .
Quail, P.H., W.R . Briggs , J. Chory, R.P . Hangarter , N.P . Harberd, R.E . Kendrick , M. Koorneef, B.
Parks, R.A . Sharrock , E . Schafer , W.F . Thompso n an d G.C . Whitelam . 1994 . Spotligh t o n
phytochrome nomenclature . Plan t Cel l 6:468-471.
Salisbury, F.B. an d C.W. Ross . 1991 . Plan t Physiology, Fourth Edition . Wadswort h Publishin g Co.,
Belmont, California . Appendi x B : Radiant Energy : Som e Definitions, pp. 494-501.
Shibles, R.M . 1976 . Committe e Report : Terminology pertaining to photosynthesis. Cro p Scienc e
16:437-439.
Shropshire, Jr. , W . an d H . Mohr , editors . 1983 . Photomorphogenesis . Encyclopedi a o f Plan t
Physiology, V. 16 A and 16B . Springer-Verlag , New York.
Smith, H. an d M.G . Holmes , editors . 1984 . Technique s in Photomorphogenesis. Academi c Press ,
New York .
Spomer, L.A . 1980 . Guideline s fo r measurin g and reportin g environmenta l factor s i n controlle d
environment facilities . Communication s in Soil Science an d Plan t Analysis 11(12):1203-1208.
Spomer, L.A . 1981 . Guideline s fo r measurin g an d reportin g environmenta l factor s i n growt h
chambers. Agronom y Journa l 73(2):376-378 .
Thimijan, R.W . an d R.D . Heins . 1983 . Photometric , radiometric , an d quantu m ligh t unit s o f
measure: a review o f procedures fo r interconversion. HortScienc e 18(6):818-821 .
CONSULTANTS
Steven J . Brit z
U.S. Departmen t o f Agriculture, ARS
Beltsville, Maryland

J. Michae l Robinson
U.S. Departmen t o f Agriculture, ARS
Beltsville, Maryland

Gerald F . Deitzer
University o f Maryland
College Park , Maryland

Walter Shropshire , Jr .
Omega Laborator y
Timonium, Maryland

Elisabeth Gant t
University of Maryland
College Park , Maryland

Ambler Thompso n
U.S. Department o f Commerc e
Gaithersburg, Maryland

III
PLANT BIOCHEMISTRY AND MOLECULAR BIOLOGY
Much of plant physiology and plant science in general is plant biochemistry. Fo r th e
most part , traditiona l plan t biochemistry is the same as general biochemistry , but a
few specia l feature s appea r i n th e table s o f Chapter 10 . Durin g recent years , plant
physiologists have become deeply involved in understanding the biochemistry of plant
genetics, a field tha t i s often calle d molecular biology, the topi c of Chapter 11 .

10
PLANT BIOCHEMISTRY
Clanton C . Black, Jr .
Biochemistry an d Molecula r Biolog y Departmen t
Life Science s Building
University o f Georgi a
Athens, Georgia 3060 2 U.S.A .
The followin g discussio n an d table s hav e bee n extracte d fro m th e Instructions to
Authors, normall y published annuall y in a January issue of Th e Journal o f Biological
Chemistry (use d by permission), modified somewhat to more closely conform with SI
notation an d with special referenc e t o th e plan t sciences .
1. INSTRUCTIONS ON CHEMICAL AND MATHEMATICAL USAGE
A. General. I n preparing a manuscript fo r publication, make references i n th e
text to simple chemical compounds by the use of formulas when these can be printed
in single horizontal lines o f type. D o no t us e two-dimensional formulas in running
text. Cente r chemica l equations , structura l formula , an d mathematica l formula s
between successive lines of text. Prepare such structural formulas and mathematical
equations in a form suitable for direct photographic reproduction and include them
on a duplicate sheet at the end of the paper. Similarly , long sequences o f amin o
acids o r nucleotide s usuall y reproduce bette r an d will be fre e fro m printers ' error s
if they ar e printe d with a laser printer , drawn in ink, or typewritte n by the author .
(Boldfaced print is best.)
B. Ionic Charge should b e designated a s a numbered superscript followin g the
chemical symbol ; e.g., Mg 2+, S 2-. Th e notation Mg(II ) is also acceptable .
C. Optically Active Isomers. Name s o f chira l compound s whos e absolut e
configuration i s known may be differentiated by the prefixe d R - and S- (see IUPA C
(1970) J . Org. Chem. 35, 2849-2867) . Whe n th e compound s ca n b e correlate d
sterically with glyceraldehyde, serine , or another standard accepte d for a specialized
class o f compound, SMAL L CAPITAL LETTERS D-, L-, and D L may be use d for chira l
compounds an d thei r racemates . Wher e the directio n o f optical rotatio n i s all that
can be specified, (+)-, (-)-, and ()- or dextro, laevo, and "optically inactive" are used,
but i n suc h instances the condition s of measurement must be specified .

81

82 Plant

Biochemistry an d Molecular Biology

D. Isotopically Labeled Compounds. Th e followin g guideline s conform to th e

recommendations adopted b y the IUB Committee of Editors of Biochemical Journals
(CEBJ). Fo r mor e detailed instruction s consul t the IUPAC-CNO C Recommendations o n Isotopicall y Modife d Compounds (1978) Eur. J. Biochem. 86, 9-25.
For mos t biochemica l usage , an isotopicall y labele d compound i s indicated by
placing the symbo l for th e isotop e introduce d i n square brackets directly attached
to the front of the name (word) or formula as in [ 14C]urea, [a- 14C]leucine, L-[methyl 14
C]methionine, [ 3H]CH4. I f the specific position of the labeling is known, it should
be indicated a t leas t th e firs t tim e the compoun d is mentioned o r i n the Material s
and Method s section ; thereafter , th e les s specifi c notatio n ca n b e used . Th e
following rule s gover n mos t situations .
The isotopi c prefi x precede s tha t par t o f th e nam e t o whic h i t refers , a s i n
sodium [14C]formate, iodo[14C]aceticacid, 2-acetamido-7-[ 131I]iodofluorene, fructose,
l,6-[l-32P]bisphosphate, B-hydroxyl[ 14C] aspartate , 17B-[ 3H]estradiol, E . coli
[3H]DNA. Term s suc h a s 131 I-labeled albumi n shoul d no t b e contracte d t o
[131I]albumin, sinc e nativ e albumi n doe s no t contai n iodine; however, 131 I-albumin
and [ 131I]iodoalbumin ar e both acceptable .
The symbo l indicatin g th e configuratio n shoul d preced e th e symbo l fo r th e
isotope; e.g, D-[ 14C]glucose; L-[l- 14C]leucine; (JR)-[14C]ethanol.
The sam e rule s appl y when the labele d compoun d is designated b y a standard
abbreviation or symbol, other tha n the atomic symbol; e.g., [a- P]AT P or [ 32P]CMP
(not CM 32P).
When isotope s o f mor e tha n on e elemen t ar e introduced , thei r symbol s ar e
arranged i n alphabetica l order , fo r exampl e [3- 14C,2,3-3H,15N]serine. Onl y th e
symbols 2 H an d 3 H shoul d b e use d for deuterium and tritium , respectively. Whe n
more than one positio n in a substance is labeled by means of the same isotope an d
the positions ar e not indicated, the number of labeled atoms is added as a right-hand
subscript, as in [14C2]glycoli c acid. Th e symbo l U indicates uniform an d G general
labeling; e.g., [U- 14C]glucose (wher e the I4 C i s uniformaly distribute d among all six
positions) an d [G- 14C]glucose (wher e the 14 C is distributed among all si x positions,
but no t necessaril y uniformly); i n the latte r case it is often sufficien t t o write simply
[14C]glucose.
When known, the positions of isotopic labeling are indicated by Arabic numerals,
Greek letters, or italicized prefixes (a s appropriate) placed within the square brackets
and befor e th e symbo l of th e elemen t concerned , t o whic h they are attache d by a
hyphen; example s ar e [l- 2H]ethanol, [l- 14C]alanine, L-[2- 14C]leucine (o r L-[a 14
C]leucine), [carboxy- 14C]leucine, [Me- 14C]isoleucine, [2,3- 14C]maleicanhydride,
[6,7-14C]xanthopterin, [3,4- 13C,35S]methionine, L-[methyl- 14C]methionine, [1- 14C,2"C]acetaldehyde.
The form s 14 CO2, 32 PO4, 32 Pi ar e acceptabl e rathe r tha n th e mor e formall y
correct1 [14C]O2 or [ 14C]CO2, [32P]Pi, etc. However , the square brackets are not t o
According to the IUPAC Recommendations (reference above), a distinction is made between isotopically substituted
compounds (carrier-free material) where square brackets are not used (e.g. 14CO2, Na125I, CH3-C2H2-OH, (14C) carbon
dioxide, sodiu m (125I ) iodide , (2-3 H2) propanol ) and isotopicall y labeled compound s where square brackets are used ,
either in the formula or in front of the name or formula (e.g . [14C]O2, [3H]CH3I, Na[125I], C[ 2H]3CH2O[2H] and other
examples give n abov e an d i n th e IUPAC Recommendations).

Plant Biochemistry 8

be use d whe n th e isotopi c symbo l i s attache d t o a wor d tha t i s no t a specifi c

chemical name , abbreviation , o r symbo l (e.g., 131 I-labeled, 3 H-ligands, 14 C-steroids,
14
C-amino-acids). Not e tha t th e abbreviatio n fo r Curi e i s Ci . The SI unit is the
bequerel (Bq). 1 Bq = 1 disintegration pe r second o r 60 disintegrations pe r minute .
1 Ci = 3 7 x 10 9 disintegrations pe r second = 3 7 GBq. 100 0 dpm = 0.4 5 nCi = 16. 7
Bq. Th e S I unit i s preferred.
E. Spectrophotometric Data. Author s reportin g spectrophotometric dat a must
indicate th e relatio n betwee n th e symbols used. Althoug h a number of alternatives
exist, i t i s recommended tha t author s follo w th e symbol s and terminolog y adopted
by IUPAC (1970) Pure Appl. Chem. 21, 1. Beer' s law may be stated as: A = -Iog 10T
= elc , wher e A i s th e absorbance ; T , th e transmittanc e ( = I /Io); e , th e molar
absorbance coefficient ; c, the mola r concentration o f the absorbing substances; an d
l, the lengt h o f the optica l pat h in centimeters. Unde r thes e condition s e ha s th e
dimensions L-mol1-.cm- 1 (not cm 2 mol- 1). Th e ter m absorbance i s preferre d t o
optical density.
If Beer's law is not followe d by a particular substance in solution, this should be
explicitly stated; eve n in such cases th e substance ma y be characterized by reporting
the absorbanc e a t a specifie d concentration . Whe n spectrophotometri c measure ments are mad e with the us e of a radiant energy source that is not confine d strictly
(as in a line spectrum) t o th e wavelengt h or frequenc y specified , the exac t value of
e wil l be somewhat ambiguous ; report th e spectral characteristic s of the source .
F. Molecular Weight and Mass. Ther e ar e tw o equivalen t expression s tha t
should be distinquished: molecular weight (Mr, relative molecular mass) is the rati o
of th e mas s of a molecule t o 1/1 2 o f the mas s of carbon 12 . Henc e i t i s dimensionless. Molecular mass (symbo l m) i n contrast i s not a ratio and can be expressed i n
daltons (symbo l Da) or in atomic mass units (symbol u). Th e molecular mass is th e
mass o f on e molecul e o f a substance ; i t i s thu s th e mola r mas s (M ) divide d b y
Avogadro's number . Th e dalto n i s defined as 1/1 2 o f the mas s of carbon 12 .
It i s correct t o sa y either "th e molecula r mas s of X i s 10,00 0 daltons " (or "1 0
kDa") o r "th e relativ e molecula r mas s (molecula r weight ) Mr = 10,000), " bu t i t i s
not correc t t o expres s M r i n daltons. On e ca n use expressions such as "the 1 0 kDa
peptide" an d "th e mas s o f a ribosom e i s 2. 6 x 10 7 daltons" (o r "2 6 MDa"), eve n
for a n entity that i s not a definable molecule. Avoi d the use of k as a shorthand for
1000 o r fo r kDa (kilodalton) .
When presenting estimate s o f relative molecula r mass from ge l electrophoresi s
data, be sure to includ e the scale use d to estimate the molecular mass as one of the
ordinates o n th e figure, not just the locatio n o f the various standards used.
G. Equilibrium and Velocity Constants. Dissociatio n constants , association constants, an d Michaeli s constants shoul d ordinaril y be written i n terms of concentra tion; the unit s should alway s be clearly indicated at the point where the equilibrium
constant i s defined an d wher e its value is given.
Values o f rat e constant s shoul d b e similarl y specified , first-orde r velocit y
constants bein g generall y given as s- 1. Second-orde r rat e constant s ar e ordinarily
given i n M-1.s- 1 (bette r SI : L.mol-1.s- 1).

84 Plant

Biochemistry an d Molecular Biology

H. Composition of Solutions and Buffers. Th e composition of all solutions an d

buffers shoul d b e specifie d i n sufficien t detai l t o defin e th e concentratio n o f eac h
species. Fo r ordinar y buffers, suc h as 0.1 mol.L-1 sodiu m acetate, p H 5.0 , it will be
assumed that the molarity refers to the total concentration o f the various species that
buffer a t th e indicated pH an d that the concentration o f the counterio n i s sufficien t
to neutraliz e th e charg e of the ionize d buffe r species . Th e composition of mixtures
should b e indicate d b y th e us e o f a diagona l (/ ) o r a colon . Example s are :
chloroform:ether (9:1) ; 1-butanol/acetic acid/water (75:20:5). Hyphen s or dashes are
not acceptabl e fo r thi s purpose . Giv e th e complet e unabbreviate d nam e an d th e
source (o r a reference tha t give s the complet e composition ) for all culture media.
2. ABBREVIATIONS AND SYMBOLS
All abbreviations used in a text, except those specifically indicated below (see
Tables 1 to 10), should be defined in a single footnote, inserted at the beginning of
the paper or immediately after the first such abbreviation. Abbreviations used only
in a table or figure may be defined in the legend.
Abbreviations are hindrances to readers in fields othe r than that of the author,
to abstractors, an d to scientists i n foreign countries. Therefore , their use should be
restricted t o a minimum . O n th e othe r hand , i t i s sometimes convenien t t o us e
abbreviations o r symbol s fo r th e name s o f chemica l substances , particularl y i n
equations, tables, o r figures . A limited use of abbreviations and symbols of specified
meaning is therefore accepted . However , clarit y is more important than brevity.
For som e o f th e mos t importan t biochemica l reagents , coenzymes , etc. , shor t
abbreviations are universally employed; e.g., ATP, NAD, RNA. Th e creation of new
abbreviations o f thi s kin d shoul d b e restricte d t o a n absolut e minimum . The
following abbreviations are obsolete; do not use them: TCA, PCA, DTE, DOC,
DMSO (us e Me2SO), SAM (use AdoMet).
Do not abbreviate pyridoxal, pyridoxamine, deoxypyridoxine, thiamine, cocarboxylase, pantothenate, folate , pteroylglutamate, trichloroacetic acid , perchloric acid , the
tricarboxylic acid cycle, and member s thereof. Most trivial names are sufficiently
short that they do not need further shortening.
Names of enzymes are usually not to be abbreviated except in terms of substrates
for whic h accepted abbreviation s exist (exceptions are ATPase, DNase , and RNase) .
Authors shoul d us e th e Recommende d (trivial ) Nam e give n b y th e IUPAC/TUB
Committee o n Enzyme Nomenclature in "Enzyme Nomenclature Recommendations
(1984)" (1984 , Academi c Press) . Excep t fo r ver y commo n enzymes, the reactio n
catalyzed shoul d als o be included .
Class names, such a s fatty acids, protein, etc. , or short terms (poly, furan, folate ,
etc.) ar e no t t o b e abbreviate d eve n whe n a n associate d ter m i s abbreviate d o r
symbolized (e.g. , poly(X), not PX ; H4folate, not THF) .
Symbols o r abbreviation s othe r tha n thos e liste d i n th e IUPAC-TU B Recommendations shoul d b e used onl y in those situations where an objective case may be
made for necessity; none should be used when pronouns and similar short terms may
replace a long word or phrase . The y should always be defined i n each paper. Suc h
ad hoc abbreviations and symbols should not conflict with recommended symbols and

Plant Biochemistry 8

should b e introduce d onl y whe n repeate d us e i s required . If , i n exceptiona l

circumstances, symbol s o r abbreviation s ar e use d i n a n abstract , the y shoul d b e
defined i n th e abstrac t a s well as in th e bod y of the paper .
When other abbreviations for chemical compounds ar e needed, the maximum use
should b e mad e o f standar d chemica l symbol s (C , H , O , N , P , S , Na , Cl , etc.) ,
numerical multiples (subscripts 2 and 3, not di or D or T etc., as in Me 2SO, Me 3Si-),
and o f trivia l name s an d thei r symbol s (e.g. , folate , P, Me , Pr , Bu , Ph , Ac ) (se e
Tables 3 t o 10) . Thes e symbol s ma y be combine d t o represen t mor e comple x
symbols, suc h a s Tos-Arg-OMe , i n whic h th e basi c structur e (arginine ) remain s
recognizable.
One o f the area s o f biochemistry for which special symbols ar e essentia l i s that
of macromolecules. Ther e are three main series of symbols for the monomeric units,
those fo r amin o acids , monosacharides , an d mononucleosides , o f which the amin o
acid series is the oldest .
The monomeri c unit s o f protein s ar e generall y designate d b y three-lette r
symbols: a capita l followe d by two lowe r cas e letters . Th e abbreviation s shoul d
ordinarily no t b e use d fo r th e fre e monomer s i n th e tex t o f papers . A standar d
treatment ha s been devise d fo r th e thre e group s of macromolecules, built up fro m
these units . Fo r th e amino acid residues in polypeptides, the residue with the fre e
a-amino grou p (i f one i s present) i s place d a t th e lef t o f th e sequenc e a s written .
Where th e sequenc e o f residues i s known, the symbol s are writte n i n order lef t t o
right and joined by short lines (dashes, hyphens). Wher e the sequence is not known,
the symbol s are separate d b y commas, enclosed i n parentheses. Example : Ala-Gly(Met,Pro)-Lys mean s that th e sequenc e o f methionine and proline is unknown.
For th e polysaccarides, symbol s for th e sugar s ar e joine d b y short dashe s o r
arrows to indicate th e link s between units . Th e position an d nature of the links are
shown b y numerals and th e anomeri c symbol s a an d B . Fo r example :
Maltose Glcpa1-4Gl
c or Glcpa 1 4Gl c
Lactose GalpBl-4Gl
c o r GalpB 1 4Gl c
The arro w point s awa y fro m th e hemiaceta l link . If the das h is used, it i s assumed
that th e hemiaceta l lin k is to the lef t o f it. Whe n it is necessary to indicate furanose
or pyranose , th e italicize d (underline d i f italics are no t available ) letter f,or p afte r
the saccarid e symbo l may be used ; e.g. , Ribf for ribofuranose.
Macromolecules compose d o f repeating sequence s ma y be represente d b y th e
prefix 'poly ' o r the subscrip t n , bot h indicatin g 'polymer of.' Th e symbol s for th e
monomeric unit s o f th e sequenc e ar e enclose d i n parentheses . Thus , poly(lys ) or
(Lys)n i s polylysine ; poly(Ala-Lys ) o r (Ala-Lys) n i s a linear polyme r consistin g o f
alanine and lysine in regular alternating sequence, and poly(Ala, Lys) is the irregular
(random) copolymer o f equal amount s of these amino acids. Betwee n poly and th e
parenthesis ther e is no intervenin g spac e o r hyphen . Th e n ma y be replace d b y a
definite number , a n averag e (e.g. , 10) , o r a rang e (e.g. , 8 t o 12) , a s appropriate .
'Oligo' may replac e 'poly ' for shor t chains . See als o the legen d to Tabl e 8
regarding nucleoside an d mucleotid e symbols.
GeneticsA. guid e t o nomenclatur e i n bacteria l genetic s ma y b e foun d i n
Demerec, M. , Adelberg, E.A. , Clark , A.J., an d Hartman , P.E. (1966 ) Genetics 54,

86 Plant

Biochemistry an d Molecular Biology

61-76. Not e that the genotypes are italicized (underline d i f italics are not available) ;
phenotypes ar e not . (Se e als o Chapte r 1 1 in this volume.)
Genetic designations fo r various bacteria, bacteriophages, animal viruses, algae,
and other materials ar e listed i n "Genetic Maps " (Editor Stephe n J. O'Brien). Thi s
publication ca n b e purchase d fro m Col d Sprin g Harbo r Laboratory , Fulfillmen t
Department. P.O . Bo x 100 MM, Cold Spring Harbor, NY 11724. Th e nomenclatur e
of variou s bacteri a i s liste d i n Bergey , Manual o f Determinative Bacteriology, 8t h
Edition, Waverl y Press, Baltimore , M D 21202. Th e nomenclatur e o f transposabl e
elements i n prokaryotes can be found i n A. Campbell et al. (1979) Gene 5, 197-206 ,
or Szybalsk i and Szybalsk i (1979) Gene 7, 217-270 .
3. THE TABLES

Table 1. Abbreviations of Units of Measurement and of Physical and Chemical

Quantities. Thes e abbreviations ma y be used without definition. The y are no t followed by periods. Th e same form i s used in the plural. Se e Chapter 1 and Sectio n
II for more information about most of these and other units. Som e are not SI units.
Name
Units of Concentrationa
molar (moles/liter )
millimolar (millimoles/liter)
micromolar (micromoles/liter )
nanomolar (nanomoles/liter )
picomolar (picomoles/liter )
Other Units
becquerelc
curie
dalton
unified atomi c mass unit (thi s is the S I equivalent o f the dalton ; its use i s preferred; se e
Chapter 1 )
equivalent (shoul d be avoided )
counts per minute
revolutions per minute
cycles per secon d (hertz )
calorie
kilocalorie
swedberg (10- 13 s)

Unit/Symbol
Mb, mol.L- 1 (preferred )
mM (rathe r tha n 10- 3 M) ,
mmol.L-1 (preferred )
uM (rathe r than 10- 6 M) ,
[umol.L-1 (preferred )
nM (no t muM) ,
nmol.L-1 (preferred)
pM (no t uu M),
pmol.L-1 (preferred)
Bq

Ci (no t SI )

Da (no t SI )
u
eq (no t SI )
cpm (no t SI , but acceptable )
rpm (no t SI , but acceptable )
Hz
cal (not SI , use J)
kcal (no t SI, use kJ )
S (not SI )

Continued

Plant Biochemistry 8

Table 1. Abbreviations of Units of Measurement (continued)

Name

Unit/Symbol

Physical and Chemical Quantities

absorbance A
equilibrium constant K
Michaelis constant K
relative molecular mass M
retardation facto r R
average acceleration of gravity at earth's g
surfaced (us e to repor t centrifugation , etc. )
specific rotatio n [a]
sedimentation coefficien t s
sedimentation coefficien t i n water a t 2 0 C ,
extrapolated t o zero concentration
diffusion coefficien t D

m
r
f
n

t
s20,w

Thennodynamic Termse

Gibbs energy change (formerly F ) G

entropy change S
enthalpy change

Term s such as milligram percent (mg% ) should not be used. Mas s concentrations should be given as g/kg, mass/volume
concentrations as g/L, etc. Th e lite r (preferre d symbo l L) i s accepted fo r us e with th e SI (see Chapte r 1 in thi s book).

Th e letter M is not an abbreviation for mole (mol); it is reserved fo r molar. Us e mM for 10- 3 M and uM for 10- 6 M .
Avoid designatin g concentration s a s umo l pe r mL . Th e designatio n should , i n thi s case , properl y b e m M (i.e. , milli molar). Maintai n consistency in the use of units in situations where they are to be compared (e.g. , d o not juxtapose 10
M and 10-5 M). A s discussed i n Chapter 1 , second-level discussion s o f the SI state that molar and M should be replaced
by ti e mor e readil y understandabl e (t o nonchemists ) mol-L-1 , mmol-L-1 , etc. Thi s i s supported b y physical chemist s
although, a s indicate d i n thi s table , Th e Journal o f Biological Chemistry continue s t o accept th e ter m molar (M).
c

1 becquerel = 1 dps or 60 dpm. 1 Ci = 3.7 x 1010 B q (37 GBq). Becquere l i s the preferred ter m in the International
System o f Units .

d In the SI, the symbol g (note italics) stands for the acceleration caused by gravity at any location (e.g., on the moon
well as earth). Th e subscript n (not italics ) i n the symbol g n indicates that the symbol stands for the average acceleratio n
caused b y gravity at th e earth's surface (9.80665 m s-2); tha t is, gn i s a unit (Se e Chapte r 1.)
e

Fo r thermodynami c term s se e th e Recommendations o f the Interunio n Commissio n o n Biothermodynamic s (J. Biol.

Chem. 251, 6879-6886 . 1976 )

Table 2. Abbreviations for Semisystematic or Trivial Names. Thos e abbreviations

preceded b y an asteris k ma y be use d withou t definition .
Abbreviation
AMP , ADP, and ATPa Adenosin
* cAMP , cGMP etc. Cycli
CMP-NeuAc Cytidin
* Co

A (o r CoASH ) Coenzym

Name
e 5' -mono, di-, and triphosphate s
c AMP (adenosin e 3':5'-monophosphate), etc.
e monophosph o /V-acetylneuraminic acid
eA
Continued

88 Plant

Biochemistry an d Molecular Biology

Table 2. Abbreviations for Semisystematic or Trivial Names (continued)

Name
Abbreviation
CoASAc
Cm-cellulose
CMP, CDP , an d CTP a
DEAE-cellulose
DNA
DOPA or Dop a
dTMP, dTDP, and dTTPa
EDTA
EGTA
FAD and FADH 2
FMN
GDP-Fuc
GDP-Man
GMP, GDP , an d GTPa
GSH an d GSS G
Hb, HbCO, HbO 2, metHb
Hepes or HEPE S
IMP, IDP, an d ITPa
Me2SO
NAD, NAD+ , an d NAD H
NADP, NADP + , and NADP H
Pi
PPj

RNA
SDS
TEAE-cellulose
TMP, TDP , an d TIPa

Tris
UDP-Gal
UDP-GalNAc
UDP-Glc
UDP-GlcNAc
UDP-GlcUA
UDP-Xyl
UMP, UDP , an d UTPa
a

Acetyl coenzym e A

O-(Carboxymethyl)cellulose

Cytidine 5 ' -mono- , di-, and triphosphates

O-(Diethylaminoethyl)-cellulose
Deoxyribonucleic aci d or deoxyribonucleat e
3,4-Dihydroxphenylalanine
Thymidine 5'-mono- , di, and triphosphate s
Ethylenediaminetetraacetate
[Ethylenebis(oxyethylenenitrilo)]tetraacetic aci d
Flavin-adenine dinucleoiid e an d it s full y reduce d for m
Riboflavin 5'-phosphat e
Guanosine diphosphofucos e
Guanosine diphosphomannos e
Guanosine 5'-mono- , di, and triphosphate s
Glutathione an d it s disulfide for m
Hemoglobin, carbo n monoxid e hemoglobin, oxyhemo globin, methemoglobi n
4-(2-Hydroxyethyl)-l-piperazineethanesulfonic acid
Inosine 5'-mono- , di- , and triphosphate s
Dimethyl sulfoxid e
Nicotinamide-adenine dinucleotide and it s oxidized and
reduced form s
Nicotinamide-adenine dinucleotide phosphate an d it s
oxidized an d reduce d form s
Inorganic phosphat e
Inorganic pyrophosphat e
Ribonucleic aci d or ribonucleat e
Sodium dodecy l sulfat e
O-(Triethylaminoethyl)cellulose
Ribosylthymine 5 ' -mono, di- , and triphosphate s
Tris(hydroxymethyl)aminomethane
Uridine diphosphogalactos e
Uridine diphosph o N-acetylgalactosamin e
Uridine diphosphoglucos e
Uridine diphosph o N-acetylglucosamine
Uridine diphosphoglucuroni c acid
Uridine diphosphoxylos e
Uridine 5'-mono , di- , and triphosphate s

Th e d prefix ma y be use d t o represent the corresponding deoxyribonucleoside phosphates ; e.g., dADP . Th e various
isomers o f adenosin e monophosphat e ma y be written 2'-AMP- , 3'-AMP , o r 5'-AM P (i n cas e o f possibl e ambiguity) .
A simila r procedur e ma y be applied t o other nucleosid e o r deoxyribonucleosid e monophosphates .

Plant Biochemistry 8

Table 3. Miscellaneous Symbols. Mos t of these abbreviations ma y be used withou t

definition. Som e (e.g. , Q, K ) should b e define d th e firs t tim e the y are used .
Symbol

Name

Symbol

Name

Ferredoxin
Menaquinone
Plastoquinone
Phosphoric acid residu e
Phylloquinone
Pteroic acid (pteroyl- )
Pteroylglutamic acida
PyridoxylTocopherol

Fd
MK
Q
P- o r - P
K
Pte
PteGlu
PxyT

TQ
Tocopherolquinone
Ubiquinone
Q
CD
Circular dichroism
ORD
Optical rotar y disperson
NMR
Nuclear magneti c resonance
ESR
Electron spi n resonanc e
EPR
Electron paramagneti c resonance
IR spectra
Infrared spectr a
UV
Ultraviolet

Folate and folyl- are not abbreviated.

Table 4. Symbols for Amino Acids. Th e symbol s precede d b y an asteris k ma y b e

used withou t definition . Th e us e o f the one-lette r abbreviation s (i n parentheses )
should be restricted to comparisons of long sequences in tables, lists, or figures , and
for suc h special use a s tagging three-dimensiona l model s o f proteins. The y shoul d
not b e use d i n paper s wher e th e single-lette r syste m fo r nucleosid e sequence s i s
employed, a s i n repeatin g codons . Di(-amin o acids ) ar e liste d i n appendi x B of
Nomenclature of -Amin o Acids, CBN (1975) Biochemistry 14, 449-462 .
Name
Alanine
3-Aminopropionic aci d
Arginine
Asparagine
Aspartic acid
Aspartic acid or asparagin e
4-Carboxyglutamic acid
Cysteine
Glutamic acid
Glutamine
Glutamic acid or glutamine
Glycine
Half-cystine
Histidine
Homocysteine
Homoserine

Symbol
Ala (A )
BAla
Arg (R )
Asn (N )
Asp (D )
Asx (B )
Gla
Cys (C )
Glu (E )
Gln (Q )
Glx (Z )
Gly (G )
Cys
His (H )
Hey
Hse

Name

Symbol

Homoserine lacton e
Hse>
Hydroxylysine
Hyl
Hydroxyproline
Hyp
Isoleucine
He
(I)Leucine
Leu (L )
Lysine
Lys (K )
Methionine
Met (M )
Ornithine
Orn
Phenylalanine
Phe (F )
Proline
Pro (P )
5-Pyrrolidone-2-carboxylic acid <Gl
u
(pyroglutamic acid; 5-oxoproline )
Serine
Ser (S )
Threonine
Thr (T )
Tryptophan
Trp (W )
Tyrosine
Tyr (Y )
Valine
Val (V )

90 Plant

Biochemistry an d Molecular Biology

Table 5. Symbols for Carbohydrates and Organic Acids. Thos e symbol s preceded
by a n asteris k ma y be use d withou t definition . Pyranos e and furanos e forms ar e
designated where necessar y by the suffixe s p an d /.
Carbohydrate
Simple sugars
Arabinose
Fructose
Fucose
Galactose
Glucose
Mannose
Rhamnose
Ribose
Xylose
Derivatives of various sugars
N-Acetylglucosamine
Glucosamine
2-Deoxyglucosea
Glucuronic acid
Reductive Pentos e Phosphate Cycle
6 Phosphogluconi c Acid
Fructose-l,6-bisphosphate
Fructose-6-phosphate
Xylulose-5 -phosphate
Sedoheptulose-7-phosphate
Sedoheptulose 1,7-bisphosphat e
3-Phosphoglycerate
Glyceraldehyde-3-phosphate
Dihydroxyacetone phosphate
Erythrose-4-phosphate
Fructose-2,6-bisphosphate
Glucose-1,6-bisphosphate
Sialic Acid
Organic Acids of the Tricarboxylic Acid cycle
Oxalacetic Acid
Citric Acid
a-Ketoglutaric Acid , 2-oxoglutorat e
Succinic Acid
Malic Acid
Fumeric Acid
Isocitric Acid
Pyruvic Acid
Phosphoenolpyruvic Acid
Cis-Aconitate

Symbol

Ara
Fru
Fuc
Gal
Glc
Man
Rha

Rib
Xyl

GlcNAc
GlcN
dGlc
GlcA
RPPC
6PGL
Fru-l,6-BP or FBP or F1,6-P 2
Fpu-6-P o r F6 P
Xylu-5-P or X5P
Sedoh-7-P o r S7P
Sedoh-l,7-BP o r S1,7-P 2
3-PGA
GAP
DHAP
E-4 P or Ery-4-P
Fru2,6-Bp o r F2,6-P 2
Glul,6-Bp o r G1,6-P 2
Sia

OAA
CIT
a-KG, 2-O G

SUC

MAL
FUM
ISOCIT
PYR
PEP
cACN

aThe prefix 'd' indicates a 2-deoxysugar. Isomers may be designated similarly with a positional numerial; e.g., 3-aThe prefix '
deoxyglucose; 3-dGlc .
Note: I n case s where the distinction between N-acetyl and O-acety l i s important, NeuNA c or NeuOA c ar e acceptabl e
with definition . Likewise , NeuNGc and NeuOG c ar e acceptabl e fo r th e glycolyl analogs.

Plant Biochemistry 9

Table 6. Symbols for Pyrimidine and Purine Bases. Thes e symbol s shoul d b e
defined excep t thos e marked wit h an asterisk .
Base
Adenine *
'a base' Bas
Cytosine *
Guanine *
Hypoxanthine Hy
6-Mercaptopurine (thiohypoxanthine ) Sh
Orotate Or
'a purine' Pu
'a pyrimidine'
Thymine *
Uracil *
Xanthine *

Symbol
Ad
Cy
Gu

Th
Ur
Xa

e
e
t
a
p
y
o
r
Pyr
y
a
n

Table 7. Symbols for Nucleosides and Nucleotides. Symbol s preceded b y an asteris k

may be used withou t definition. Tw o systems are recognized, one using three-lette r
symbols for the common nucleosides an d a capital italic P for the phosphoric residue,
the othe r usin g single capita l letter s fo r th e commo n nucleosides and a lowe r case
p fo r th e phosphori c residue . Th e three-lette r symbol s should b e use d whenever
chemical changes involvin g nucleosides o r nucleotides are discussed. Th e one-lette r
symbols ar e intende d fo r th e nucleosid e residue s i n sequences or partia l sequence s
only; i n thes e the y shoul d alway s be connecte d b y hyphens (for internal phospho diester 3 ' -5' linkages) , an d the terminal phosphoric residu e should be indicated by
the letter p. Codon s ma y be indicated i n the tex t without hyphens or th e termina l
p's. Th e 2'-deoxyribonucleoside s ar e indicate d b y the prefi x 'd' . Fo r incompletel y
specified base s i n nucleotid e sequences , see (1986 ) J. Biol. Chem. 261 , 13-17 .
Nucleoside
Adenosine
Bromouridine
Cytidine
Dihydrouridine
Guanosine
Inosine
6-Mercaptopurine ribonucleosid e (6-thioinosine )
'a nucleoside '
Orotidine
Pseudouridine
'a purin e nucleoside '
'a pyrimidin e nucleoside '

Symbol
Three-letter One-letter
* Ad o
BrUrd
* Cy d
* Gu o
* In o
Sno
Nuc
Ord
*r d
Puo
Pyd

*A
B
*C
D or h U
*G
*I
M or M P
N
O
or Qa
*
R
Y

Continued

92 Plant

Biochemistry and Molecular Biology

Table 7. Symbols for Nucleosides and Nucleotides (continued)

Symbol
Three-letter One-lette r

Nucleoside

Nir

Ribosylnicotinamide
Ribosylthimine

* Th d

Thiouridine

Srd

* dTh d
* Ur d
* Xa o

Thymidine (2'-deoxyribosylthymine )
Undine
Xanthosine
Phosphoric residu e

-P

Sor sU

*d T
*U
*X
p or _ b

a For computer work

For interna l phosphodieste r bonds, use a hyphen .

Table 8. Symbols for Specific Preparations of Nucleic Acids. Thes e symbols may be
used without definition .
Name
Symbol
Complementary DNA , RN A cDNA
Heterogeneous nuclear RNA hnRN
Messenger RN A mRN
Mitochondrial DNA, RN A mtDNA
Nuclear DNA , RN A nDNA
Ribosomal RN A rRN
Transfer RN A tRN
Chloroplast plasti d or DNA , RN A ctDNA

, cRNA
A
A
, mtRNA
, nRNA
A
A
, pDNA , ctRNA, pRN A

Table 9, Buffers. Th e Buffe r name s ma y be use d without definition.

Buffer name
Systematic Description
ACES 2-[(2-amino-2-oxoethyl)amino]ethanesulfoni
c acid
ADA [(carba18moylmethyl)imino]diaceti
c acid
BES 2-[bis(2-hydroxyethyl)amino)ethanesulfoni
c aci d
Bicine N,N-bis(2-hydroxyethyl)glycin
e
BisTris 2-[bis(2-hydroxyethyl)amino]-2-(hydroxymethyl)-propane-l,3-dio
l
CAPS 3-(cyclohexyIamino)propanesulfoni
c acid
CHAPS 3-[(3-cholamidopropyl)dimethylammonio]-l-propanesulfoni
c aci d
CHAPSO 3-[(3-cholamidopropyl)dimethylammonio]-2-hydroxy-l-propanesulfonat
CHES [2-(N-cyclohexylamino)-ethanesulfoni
c acid ]
CDTA 1,2-cyclohexylenedinitrilotetraaceti
c aci d
EDTA ethylenediaminetetraaceti
c acid

Continued

Plant Biochemistry 9

Table 9. Buffers (continued)

Buffer name
EGTA
EPPS
HEPES
HEPPS
MES
MOPS
PIPES
TAPS
TEMED
TES
Tricine
Tris

Systematic Description
[ethylenebis(oxyethylenenitrilo)]tetraacetic aci d
Acceptable abbreviation fo r HEPPS (use HEPP S definition) .
4-(2-hydroxyethyl)-l-piperazineethanesulfonic acid
4-(2-hydroxyethyl)-l-piperazinepropanesulfonic aci d
4-morpholineethanesulfonic aci d
4-morpholinepropanesulfonic aci d
1,4-piperazinediethanesulfonic aci d
3-{[2-hydroxy-l,l-bis(hydroxyraethyl)ethyl]amino}-l-propanesulfonic aci d
N,N,N',N'-tetramethylethyIenediamine
2-{ [2-hydroxy-l ,1 -bis(hydroxymethyl)ethyl]amino}ethanesulfonic aci d
N-[2-hydroxy-l ,1 -bis(hydroxymethyl)ethyl]glycine
2-amino-2-hydroxymethylpropane-l,3-diol

Table 10. Tentative Rules and Recommendations of International Scientific

Unions
Group/Title
General
Abbreviations an d symbol s fo r chemica l names of
special interes t i n biological chemistry.
Abbreviations an d symbols : a compilation (1976).
Citation o f bibliographi c references i n biochemical
journals.
Biothermodynamics
Recommendations for the measurement and presentation of biochemical equilibriu m data.
Recommendations for the presentatio n of thermody namic and related dat a i n biology (1985 )
Labeled compounds
Isotopically modifie d compounds b
Stereochemistry
Fundamental stereochemistry c
Enzymes
Enzyme nomenclature . Recommendation s (1984 )
The nomenclatur e o f multiple forms o f enzymes
Catalytic activity
Units o f enzyme activit y (1978)
Symbolism an d terminolog y i n enzyme kinetics
Ami no acids, peptides, and proteins
Nomenclature an d symbolis m for amin o acids and
peptides. Recommendation s (1983)

May be found ina

(1966) J. Biol. Chem. 241 , 527-53 3
(1977) Eur. J. Biochem. 74 , 1-6
(1973) J. Biol. Chem. 248 , 7279-728 0
(1976) /. Biol. Chem. 251 , 6879-688 5
(1985) Eur. J. Biochem. 153 , 429-43 4
(1978) Eur. J. Biochem. 86 , 9-2 5
(1979) Eur. J. Biochem. 102 , 315-31 6
(1970) J. Org. Chem. 25 , 2849-2867
(1984) Academic Pres s
(1977) J. Biol. Chem. 252 , 5939-594 1
(1979) Eur. J. Biochem. 97 , 319-320
(1982) Eur. J. Biochem. 128 , 281-29 1
(1985) J. Biol. Chem. 260 , 14-4 2

Continued

94 Plant

Biochemistry an d Molecular Biology

Table 10. Tentative Rules and Recommendations (continued)

Group/Title

May be found ina

Abbreviations an d symbol s fo r the descriptio n o f th e

conformation o f polypeptide chains
Nomenclature o f iron-sulfur proteins
Corrections
Nomenclature o f peptide hormone s
Nomenclature of human immunoglobulin s
Nomenclature o f glycoproteins, glycopeptides , an d
peptidoglycans. Recommendation s (1985 )
Nomenclature o f electron-transfer proteins . Recom mendations (1989 )
Carbohydrates
Tentative rules for carbohydrat e nomenclature . Par t
1. (1969)
Corrections
Conformational nomenclatur e fo r five- and six-membered ring forms of monosaccharides an d thei r
derivatives (1980 )
Nomenclature o f unsaturated monosaccharide s (1980 )
Nomenclature o f branched-chain monosaccharide s
(1980)
Abbreviated terminolog y of oligosaccharide chain s
(1980)
Polysaccharide nomenclature (1980 )
Symbols for specifyin g th e conformatio n of polysaccharide chains
Lipids
The nomenclatur e of lipids. Recommendation s
(1976)
Nucleotidcs and nucleic acids
Abbreviations an d symbol s for nuclei c acids, polynucleotides and their constituent s
Corrections
Abbreviations an d symbol s for th e descriptio n of
conformations o f polynucleotide chains
Nomenclature fo r incompletel y specified base s in
nucleic acid sequences . Recommendation s (1984)
Phosphorus
Nomenclature o f phosphorus-containing compounds
of biochemica l importance. Recommendation s
(1976)
Steroids
The nomenclatur e o f steroids. Revise d tentativ e
rulesd
Amendments (1971 ) and correction s

(1970) J. Biol. Chem. 245, 6489-6497

(1979) Eur. J. Biochem. 93, 427-43 0
(1979) Eur. J. Biochem. 102 , 31 5
(1975) J. Biol. Chem. 250, 3215-3216
(1974) Eur. J. Biochem. 45, 5-6
(1987) J. Biol. Chem. 262,13-18
(1991) Eur. J. Biochem. 200, 599-61 1
(1972) J. Bio. Chem. 247, 613-634
(1972) Eur. J. Biochem. 25, 4
(1980) Eur. J. Biochem. 111, 295-298
(1981) Eur. J. Biochem. 119 , 1- 3
(1981) Eur. J. Biochem. 119 , 5- 8
(1982) J. Biol. Chem. 257, 3347-3351
(1982) J. Biol Chem. 257, 3352-335 4
(1983) Eur. J. Biochem. 131 , 5- 7
(1977) Lipids 12 , 455-46 8

(1970) J. Biol. Chem. 245, 5171-5176

(1971) J. Biol. Chem. 246, 489 4
(1983) Eur. J. Biochem. 131 , 9-1 5
(1986) J. Biol Chem. 261, 13-1 7
(1977) Proc. Natl. Acad. Sci. U.S.A.
74, 2222-223 0

(1969) Biochemistry 8 , 2227-224 2

(1971) Biochemistry 10 , 4994-499 5
Continued

Plant Biochemistry

95

Table 10. Tentative Rules and Recommendations (continued)

Group/Title

May be found ina

Quinones
Nomenclature o f quinones with isoprenoid sid e chains (1975 ) Eur. J. Biochem. 53 , 15-1 8
Carotenoids
Tentative rule s for the nomenclature of carotenoids (
Revisione (1975
Cyclitols
The nomenclature o f cyclitols. Recommendation s (
(1973)
Folic acid
Nomenclature and symbols for folic aci d and related (
compounds
Corrinoids
Nomenclature o f corrinoids (

) J. BioL chem - 247 > 2633-264 3

) Biochemistry 14 , 180 3

1972

) Eur. J. Biochem. 57, 1- 7

1975

) Eur. J. Biochem. 2 , 5-6

1967

1974

) Biochemistry 13 , 1555-1560

Retinoids
Nomenclature o f retinoid s (1983
Tetrapyrroles
The nomenclatur e o f tetrapyrrole s
Tocopherols
Nomenclature o f tocopherols an d related compound s (
(1981)
Miscellaneous (vitamins)
Trivial name s o f miscellaneous compound s o f impor- (
tance i n biochemistry

) J BioL Chem. 258, 5329-533 3

(1980) Eur. J. Biochem. 108 , 1-3 0
) Eu.r. J. Biochem. 123 , 473-47 5

1982

1966 J. BioL chem

Vitamin B6
Nomenclature fo r vitamin B6 and related compound s (

1973

Vitamin D
Nomenclature of vitamin D (1981) (

1982

- 241 , 298 7-2994

) Eur. J . Biochem. 40 , 325-327

) Eur. J . Biochem. 124 , 223-22 7

Mos t of these documents have also been publishe d in other journals, e.g., Biochemistry, Biochem. J., Eur. J.
Biochem., Boichim. Biophys. Acta, ARch. Biochem. Biophys.
The secon d edition o f a Compendium of thes e documents is available from Portlan d Press Inc. , Ashgate Publishing
Co., Ol d Pos t Rd. , Brookfield , VT 05036-9704; o r Portlan d Press, Ltd., P.O. Bo x 32, Commerce Way, Colchester
CO2 8HP , Essex , U.K . Th e Pric e is 18.00/U.S. $36.00. Postag e is 2.00/U.S. $2.50 . A 15 % discount is
allowable on order s fo r 1 0 copies or mor e to a single address. Paymen t must accompan y th e order.
b
Th e fina l versio n ma y be foun d i n (1979) Pure Appl Chem. 51, 353-380.
c
Th e fina l versio n may be found i n (1976) Pure Appl Chem. 45 , 11-30.
d
e

Th e definitive rule s for nomenclatur e of steroids may be found i n (1972 ) Pure Appi Chem. 31, 285-322.
Th e definitive rule s ma y be found i n (1975) Pure Appl Chem. 41 , 407-431.

96 Plant

Biochemistry an d Molecular Biology

REFERENCE

[Anonymous]. 1994 . Instruction s t o authors . Th e Journal of Biological Chemistry 269(1):777785. 199 4 by The America n Societ y fo r Biochemistr y and Molecula r Biology, Inc.
CONSULTANTS
William H . Campbel l Dougla
Michigan Tech . Universit y Universit
Houghton, Michiga n Columbia

s D. Randal l
y of Missouri
, Missouri

Jan Mierny k Gregor

USDA, ARS Universit
Peoria, Illinoi s Athens

y Schmid t
y of Georgi a
, Georgi a

Jack Preis s
Michigan Stat e Universit y
East Lansing , Michigan

11
PLANT MOLECULAR BIOLOGY and GENE DESIGNATIONS
Ellen M. Reardon an d Carl A Pric e
Waksman Institut e
Rutgers Universit y
Piscataway, N J 08855-0759
1. TERMINOLOGY

The word s define d i n thi s sectio n represen t terminolog y commo n t o molecula r

biology integrate d wit h certai n phrase s usefu l i n biochemistry , microbiology , an d
genetics. A few terms are defined within other definitions; these are also printe d in
boldfaced type . Word s i n italics ar e themselve s define d elsewhere althoug h italic s
may also be use d fo r scientifi c names and eve n fo r emphases.
2D gel Two-dimensiona l gel ; an electrophoretic techniqu e based on runnin g a gel
under one circumstance, e.g., a pH gradient, rotating the gel 90, and rerunning the
gel under differen t conditions .
35 S promoter A

strong promoter from cauliflowe r mosaic virus (CaMV).

amplified-fragment-length polymorphism (AFLP) A

mapping.
allele A

techniqu e use d i n genome

n alternat e form o f a gene at a specific locus.

anticodon A

nucleotide triplet complementar y to a codon.

ballistic transformation Th
host.

e us e of a particl e gu n t o inser t foreig n DNA int o a

base, kilobase (b, kb) Adenin e (A) , guanine (G), cytosine (C) , thymine (T), an d
uracil (U) . Base s ar e elements of DNA and RNA Th e lengths of DNA and RN A
sequences are measured i n bases (100 0 b = 1 kb).
branch point Th e poin t i n th e replicatio n o f a nucleotide chai n wher e ne w
nucleotides are added .
cauliflower mosaic virus (CaMV ) A
centromere Regio
attaches.

DNA virus that infects plants.

n o f th e chromosome to whic h th e mitoti c or meioti c spindle

97

98 Plant

Biochemistry an d Molecular Biology

chloroamphenicol transacetylase (CAT ) A

as a reporter gene o r selectable marker.
chromatin Th

bacterial enzyme encoded by cat; serves

e material o f chromosomes, consisting o f DNA and histone proteins .

chromosome A
self-replicatin g modul e o f a genome consistin g o f DN A an d
proteins. Note : nuclea r chromosome s ar e physicall y very differen t fro m chromo somes o f organelles and prokaryotes. Vira l chromosomes ma y be of DNA or RNA .
circular A pofynucleotide chain , usuall y of DNA, in the for m o f a circle; 3 ' - and
5' -end s ma y be hydroge n bonde d o r joined covalentl y (see closed circular).
cis Locate

d o n th e sam e stran d o f DNA .

closed circular A
coding sequence A

covalently close d circl e o f DN A

set o f codons that encod e a protein .

codon A ihree-nucleotide segment o f RNA specifyin g a n amin o aci d o r transla tional stop signal .
colony A

grou p o f cells, normall y on a plate, derive d fro m a single cell.

colony hybridization A metho d fo r detectin g th e presenc e o f a specifi c sequence

of DN A amon g bacterial colonie s o n a plate .
compatibility Th e coexistence o f two genetic systems in the same cell or organism;
e.g., phage with a host bacteriu m or organelles wit h a nucleus.
complementary DNA (cDNA) A
reverse transcriptase .

single-stranded DN A cop y of an RN A mad e by

complementation Th e rescue of function to produce a wild-type phenotype by two

separate mutant s within the sam e cell; distinguishable from recombinatio n i n which
a wild-type gene supplement s a mutation.
constitutive Gene

s that ar e alway s expressed; housekeepin g genes .

construct A plasmid i n whic h geneti c o r structura l elements, suc h a s gene s o r

restriction sites , ar e introduce d through artificia l means.
controlling elements Sequence s tha t contro l th e expressio n o f genes b y inserting
into o r retractin g fro m them ; transposable elements; transposons.
cosmid Aplasmid
plasmid DNA .
cryo- Ver

includin g a lambda phage cos site that permits packaging of the

y cold conditions , a s in liquid nitrogen .

dalton, kilodalton (Da, kDa, kD) A uni t o f measuremen t o f molecula r mass ;

kD=1000 daltons, wher e 1 dalton = 1.66 1 x 10- 24 g; equivalent t o th e unified atomic
mass unit (u) , which i s an S I unit defined as 1/1 2 o f the mas s of an ato m o f 12 C.
deoxyribonucleic acid (DNA) Th e macromolecul e in which genetic information is
stored; th e primar y genetic materia l of all cells.
domain A three-dimensional arrangement of amino acids with specific catalytic or
binding properties .

Plant Molecular Biology an d Gene Designations 9

downstream

Proximal; in the 3' direction.

ectopic gen e A gene, usuall y on e transforme d int o anothe r species , tha t i s

expressed i n other tha n it s normal locatio n o r stag e of development.
editing Th e addition, removal, or substitution of nucleotides to a DNA or RNA to
generate or restor e correct coding .
electrophoresis Th e movemen t of a mixture of macromolecules into a semi-solid,
porous mediu m i n respons e t o a n electri c fiel d t o determin e it s mass/charg e rati o
(size) o r t o separat e i t fro m othe r components . Thi s techniqu e i s used t o separat e
both protein s an d nucleic acids. ( A liquid medium can also be used, but toda y such
applications ar e onl y for special purposes. )
electroporation Th
foreign DN A

e openin g o f pore s i n cell s b y electric shoc k s o a s t o inser t

enhancer A DN A sequenc e ofte n bu t no t alway s dista l t o th e promoter tha t

increases promote r activity , and therefore, transcription.
exon Th

e codin g portio n o f an RN A transcript o f a split gene .

expressed sequenc e ta g (EST ) cDNA s for protein gene s that are expressed unde r
selected conditions . ES T ofte n refer s t o a partia l sequenc e o f th e cDN A use d t o
identify site s i n the genom e tha t encode a specific gen e product.
expression, gen e expressio n Th e accumulatio n of specific gen e product s usually
under define d environmenta l conditions .
gel A semi-soli d mediu m int o whic h protein s o r nuclei c acid s ar e subjecte d t o
electrophoresis, to determin e thei r size or to separate them from othe r components .
gene Althoug h i t i s impossibl e t o "define " a gen e t o th e mutua l satisfaction of
biochemists, molecula r biologists , and geneticists, in this context a concise meanin g
might b e th e sequenc e of DNA tha t encodes a gene product (an RN A o r protein) ,
including al l upstream an d downstream sequences involve d in th e expression o f th e
gene.
gene famil y A se t o f gene s whos e sequence s diffe r onl y slightl y tha t encod e
identical products ; within a single species, a multigene family.
gene taggin g Th e additio n o f a marker t o a gene; often b y the introductio n o f a
transposon o r insertio n element .
gene transfer Transformation;
th e insertion of foreign DN A into a host conferring
a trai t no t previousl y inherant i n that organism.
genome Su
virus.

m o f al l geneti c informatio n of a n organism , nucleus , organelle , o r

jS-glucuronidase (GUS ) Bacteria l enzyme encode d by the uidA gene ; a selectable

marker whose product can be directl y visualized in man y tissues.
heat shock A n upwar d temperature shift inducin g stress that results in a quantitative or qualitativ e alteration in gene expression.

100 Plant

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heterologous Perceive d a s different ; ofte n use d i n referenc e t o DN A t o mea n

having dissimilar sequences.
histone A smal l number o f highly conserved protein s tha t comple x with nuclear
DNA to for m chromatin.
homologous Perceive d a s having a common ancestor; ofte n use d i n referenc e t o
DNA to mea n havin g similar sequences .
hybrid duplex A double-stranded molecul e compose d of complementary DNA or
RNA strands , o r singl e strand s o f complementar y DNA o r RN A fro m differen t
species.
hybridization Th
hybrid duplex.

e matin g o f complementary DN A an d RN A strand s t o for m a

hydropathy plot A
macromolecule.

ma p depictin g th e hydrophili c and hydrophobi c domains of a

immunodetection Th

e identificatio n of a specific polypeptide by use of antibodies.

induction Increase d expressio n o f a gene i n respons e t o a n externa l factor ; e.g. ,

expression o f the gen e encoding nitrat e reductas e in response t o nitrate ; see repression.
insertion Introductio n of a nucleotide or nucleotide chain into RNA or DNA; may
occur naturally , as by insertion element, transposon, o r T-DNA.
insertional mutagenesis Chang e o f geneti c informatio n b y introductio n o f a
nucleotide chai n int o a gene; gen e is usually rendered inactive .
intron Non-codin
ing.

g portion o f an RNA transcript tha t is removed during process-

iso-electric focusing (IEF) Separatio

their isoelectric points .

n o f proteins i n a gradient gel to th e p H of

lariat A n intermediate structur e forme d i n an RNA molecule during the excision

of certai n kind s of introns.
lawn Th

e confluenc e of bacterial colonie s o n a Petri plate.

library Transforme d bacteria , cosmids , YACs , etc. , whos e inserts represen t th e

entire genome (genomi c library) , or transcripts (cDNA library ) o f a n organis m expressed unde r define d conditions.
ligation Th

e joining of insert and plasmid DNA, usually by the action of T4 ligase.

locus Sit e o n a chromosome at whic h a specifi c gene, or allele o f tha t gene , i s

located. I n molecular term s a locus can be defined t o a single nucleotide, wherea s
the precisio n o f a locus determined b y segregation analysis is limited to abou t 2 0 to
2000 kb or 1 centimorgan (se e morgan).
megaplasmid A plasmid o f 10 0 k b o r more ; e.g. , ni f plasmids , which encod e
enzymes o f nitroge n fixation .

Plant Molecular Biology an d Gene Designations 10

missense DNA Geneti c error resulting in a gene product with the wrong sequence.
morgan (M ) A uni t tha t expresse s th e relativ e distanc e betwee n gene s o n a
chromosome; one M equals a crossover value of 100 %; 1 centimorgan is equivalent
to 2 0 to 2,00 0 k b in higher plants. Name d in honor o f Thomas Hun t Morgan .
multigene family A set o f genes within a species tha t encode simila r or identica l
products.
mutation An y change i n the sequenc e o f DNA in a genome.
nonsense DNA Geneti c error tha t result s i n premature termination of transcription int o RNA .
northern blot Th e hybridizatio n on a membrane of a specific radiolabele d probe
with RNA transferre d from a n electrophoretic separation . Note : northern blot is not
capitalized; see Southern blot.
nucleoid Regio
concentrated.

n o f a prokaryote , plastid , o r mitochondrio n wher e DN A i s

nucleotide Th e basic subunit of DNA and RNA, composed o f a base (A, T or U ,

G, C), a sugar (deoxyribose or ribose), an d a phosphate .
nucleus Th
matin.

e portio n o f the cell , often membrane bound, that contain s the chro -

palindrome Inverte d repeat s o f DN A suc h tha t th e sequenc e i s the sam e when

read forwar d o r backward ; e.g., CAGTTGAC , o r i n the English language, "Madam,
I'm Adam. "
phage A

virus that infect s bacteria .

phenotype Th e displa y of characteristics exhibited by an organis m resulting fro m

the expressio n o f its genome in the existin g environment.
plaque Clearin
bacteriophage.

g o f bacterial lawn caused by a virus infection initiated by a single

plaque-forming unit (PFU ) A quantitativ e measur e o f th e numbe r o f viruse s

descended fro m a single colony require d t o clear a n area o f a bacterial lawn.
plasmid DN A that can replicate independently within a bacterial cell; engineere d
plasmids serv e as cloning vectors fo r the insertio n of foreign DN A into a host.
plating Growin g bacterial colonies , transforme d or otherwise , on a Petri plate .
ploidy Th e number s of copies o f a genome found i n a given species; haploid has
one copy ; diploid, two; etc .
polyadenylation Th e post-transcriptional addition of adenylates to the 3' - end of
a mRN A molecule ; poly(A) + an d poly(A) " refe r t o th e presenc e o r absenc e o f
multiple A residues.
polymerase Th

e enzym e by which DNA and RNA ar e replicated fro m a template.

102 Plant

Biochemistry and Molecular Biology

polymerase chain reaction (PCR ) A reactio n i n whic h a templat e o f DN A i s

elongated b y cycling of temperature s i n a reactio n chambe r with primer(s) an d al l
possible nucleotides i n excess .
polynucleotide A

chain o f nucleotides; DN A or RNA .

post-transcriptional regulation Regulatio n o f gene expression afte r th e synthesi s

of th e primar y transcript, a s at th e leve l o f processing or degradatio n o f transcripts.
post-translational regulation Contro l of gene expression afte r translation, as at th e
level o f processing o r degradatio n o f proteins .
premature termination Terminatio n o f transcription befor e the en d o f the coding
sequence resultin g i n an incomplet e transcript .
primary transcript A n RN A synthesized directly fro m it s DNA template prio r t o
processing, polyadenylation , o r editing .
primer Short , single-strande d nucleotid e segmen t tha t initiate s th e replicatio n o f
DNA fro m a template .
probe A DNA, RNA, or protein use d to identify o r isolate a specific target DNA ,
RNA, o r protein .
promoter Th e region o f DNA that binds RNA polymerase II to initiate transcription; promoters ma y be constitutive o r "turned on" in response to a variety of signals.
See signal transduction.
protein structure: primary (1 structure) Th

e amino-acid sequence of a protein .

protein structure: secondary (2 structure) Th e loca l conformation or foldin g of

the protein' s backbone, a s into helice s or pleated sheets .
protein structure: tertiary (3 structure) Th
a protein .

e three-dimensiona l organization of

protein structure: quarternary (4 structure) Th e organization of some protein s

from multipl e polypeptid e subunits . Homopolymeric proteins ar e compose d o f
identical subunits; heteropolymeric proteins ar e compose d o f dissimilar subunits.
pseudogenes A gen e rendere d non-functiona l b y additio n o r deletio n t o it s
structure; probabl y related t o duplicate d genes .
pulsed-field electrophoresis Th e separatio n o f genomic, or very large DNAs i n a
semi-solid mediu m b y electric pulses rather tha n constan t current .
random-amplifled-polymorphic DNA (RAPD) Th e amplificatio n of sequence s o f
DNA distribute d randoml y throughou t a genom e b y PC R technolog y t o analys e
genetic structur e o r relatednes s o f populations .
reading frame A
polypeptide.

specific mode of reading codon triplet s t o produc e a functional

receptor Recipien t o f a signal, such as a hormone , protein, light, etc., tha t binds
to a specific site an d initiate s a reaction. Se e signal transduction.

Plant Molecular Biology an d Gene Designations 10

recombinant A n organism whose genetic makeup is altered by the stable insertion

of a foreign DNA .
regulation Contro

l o f gene expression.

reporter gene Th e codin g regio n o f a surrogat e gen e whos e produc t i s easil y

observed, eg. , CAT , GUS , LacZ , fuse d t o th e promote r o f a nativ e gen e whos e
product i s difficul t t o detect . Demonstratio n o f th e surrogat e gen e analyze s th e
native gene: its function, localization, an d responses to developmental an d environ mental signals.
repressor Negativ e regulatio n o f a gen e i n respons e t o a n externa l factor ; e.g. ,
decreased expression o f the gene encoding nitrate reductase in response t o ammonium; see induction.
restriction enzyme A n enzyme , usuall y of bacteria l origin , tha t cut s DN A a t
specific base pairs, producing restriction fragments.
restriction-fragment-length polymorphism (RFLP) Characterizatio
by comparison o f sizes o f selected restriction fragments.

n of a genome

Ri plasmid Plasmi d o f Agrobacterium rhizogenes; plasmi d confer s "hairy-root "

pathogenicity to Agrobacterium species.
ribonucleic acid (RNA) A polynucleotide in which the sugar is ribose rather tha n
deoxyribose; th e molecule o f which messenger-, ribosomal-, and transfer-RNAs, and
some viral genomes ar e composed .
RNA editing Th e substitution , additio n o r remova l o f base s i n a transcript t o
produce a "correct" transcript .
scoreable marker A

trait tha t ca n be identifie d in a bacterial lawn .

second messenger A small molecule within a cell that relays a chemical messag e
from outsid e the cell , a s cyclic AMP.
selectable marker A trait whose presence or absence enables one to grow colonies
or organisms with or without a specific essential component; e.g., antibiotic-resistan t
mutants in the presence o f that antibiotic, or in the absence of an otherwise essential
nutrient.
selection Identificatio

n o f a specific trait usefu l i n singling out a transformant.

selective amplification of microsatellite polymorphic loci (SAMPL). A

nique use d i n genome mapping .

PCR tech -

sequence A specific string of bases in a protein o r polynucleotide . A s a verb, t o

sequence means t o determin e th e sequence s o f amino acids in a protein o r bases i n
a polynucleotide .
signal sequence, signal peptide A portion , usuall y th e N-terminus , of a protei n
that is recognized and removed co-translationally by the endoplasmic reticulum (ER)
prior t o transpor t o f the protei n int o the lume n of the ER ; se e transit peptide.
signal transduction Th e proces s b y which th e perceptio n o f a n externa l signa l
induces a result: th e expression of a gene or set of genes; e.g., the steps between th e

104 Plant

Biochemistry an d Molecular Biology

absorption o f ligh t b y phytochrom e an d th e synthesi s o f protein s o f th e light harvesting complex .

sodium dodecyl sulfate (SDS ) A detergen t use d t o separat e polymeric , o r
apoproteins into thei r monomeric subunits . Se e protein structure: quarternary.
Southern blot Transfe r o f DNA tha t ha s been separate d b y electrophoresis to a
membrane i n order to probe it by hybridization; after E.M . Southern. I n recognitio n
of Southern's contribution , subsequent researcher s name d RNA or protein blots after
directions also . Se e northern blot, western blot.
spliceosome A comple x o f macromolecules tha t remove s introns and joins exons
during RNA processing .
splicing Th

e joining of exons into a mature mRNA .

gene interrupte d b y an intron, or non-codin g sequence .

split gene A

stop codon A codon tha t cause s terminatio n o f transcription; specificall y UAA ,

UAG, and UG A
subtraction library A library of cDNAs containing only those transcripts expresse d
under a defined condition afte r eliminatio n of transcripts from differin g conditions .
suppression Th
sequences.

e phenotypi c correction o f a mutation without changing any gene

T-region of Ti plasmid Th
genome.
telomere Sequence
replication.

e portion of a Tiplasmid that is transferred into the host

s at both termin i of each eukaryotic chromosome that facilitate

temperature-sensitive mutation On e i n which a wild-type phenotype is expressed

at the permissive temperatur e and a mutant phenotype at a non-permissive (usually
higher) temperature .
template Th e stran d o f DN A o r RN A tha t serve s a s th e mode l fo r it s ow n
replication o r transcription .
Ti plasmid Plasmi d o f Agrobacterium tumefacien s tha t confer s pathogenicit y i n
crown-gall infection .
trans O

n th e opposit e strand o f DNA; see cis.

transcript A

n RN A tha t i s copied (transcribed) from a gene .

transcription Generatio
transduction Introductio
transformation Th
of a n organism .

n o f an RN A cop y of a DNA sequence .

n o f foreig n DN A int o a genome by a phage vector.

e introduction of a gene previously not inherent in the genome

transient gene expression Expressio

integrated int o a genome .

n of a foreign or ectopic gene that has not been

Plant Molecular Biology an d Gene Designations 10

transit peptide Th e N-terminus of a protein tha t is recognized post-translationall y

and (usually ) removed durin g transport o f a protein acros s a membrane; se e signal
peptide.
translation Formatio
amRNA

n o f a protein o n a ribosome accordin g to the instructions in

transon Exons encode d separatel y fro m on e another , with other gene s betwee n
them o r on opposite strands.
transposition Th

e insertio n o r excisio n o f a transposon t o o r fro m a genome .

transposon A transposable elemen t usually flanked by inverted repeat sequences;

see controlling elements.
twintron A

n intron within another intron .

upstream Dista
downstream.
vector Plasmid;

l or 5' t o the promoter; opposite th e direction of transcription; see

means of introducing foreign DNA int o a host .

western blot Immunodetectio n o f proteins blotted t o a membrane. Note : western

blot i s not capitalized ; se e Southern blot.
yeast artificial chromosome (YAC) A construct including centromere, telomeres, an
origin o f replication an d multipl e cloning sites fo r the insertio n o f foreign DNA .
2. GENE DESIGNATIONS

Any discussion o f gene nomenclature mus t be prefixed with the subject to which th e
terminology i s to be applied: specifically , the traditiona l genes of genetics and plant
breeding, contraste d wit h genes tha t hav e been clone d an d sequenced .
A. Nomenclatures of traditional genetics. A specifi c locus o n a chromosom e
that i s associate d wit h a uniqu e phenotyp e wil l be referre d t o her e a s a geneti c
system base d on segregational analysis. Segregationa l nomenclature s fo r individua l
plant specie s ar e distinct , an d ther e i s n o systemati c effor t t o assig n commo n
designations t o simila r gene s i n multipl e plants. Mutant s with similar phenotypes
may have no similarit y at th e molecula r level; they can only be distinguished on th e
basis of their loci on chromosomes. Thre e maize mutants with shortened internodes ,
for example , migh t b e calle d dwarf1 , dwarf2 , an d dwarf3 . Year s late r i t migh t b e
discovered tha t dwarfl i s due t o a lesion i n the biosynthesi s of giberellin, dwarf2 t o
a defec t i n a recepto r fo r gibberellin , an d dwarf 3 du e t o som e totall y unrelate d
process. A dwarf mutant in arabidopsis might be due to the same lesion in giberellin
biosynthesis, bu t geneticist s ca n not affor d t o wait until the biochemical functio n of
the gene product has been determined before designations ar e assigned t o the genes.
B. Nomenclature of sequenced plant genes. Th e Commissio n on Plan t Gen e
Nomenclature (CPGN) was founded i n 1991 under the auspices of the International
Society for Plant Molecular Biology. Th e goal of the CPGN is to unify nomenclature
across th e plan t kingdom : a gen e encodin g nitrat e reductas e i n tomat o o r i n
arabidopsis or i n pe a would have the same name. Th e establishmen t of plant-wide

106 Plant

Biochemistry an d Molecular Biology

designations, a uniqu e developmen t i n biology, i s aided b y a lon g tradition amon g

plant scientist s o f sharing ideas regardles s o f the specie s unde r study.
Once a gene ha s been isolate d an d sequenced, w e can make comparisons with
genes fro m othe r organisms . A common nomenclatur e fo r sequenced plan t gene s
began i n 198 3 wit h a proposa l fro m Hallic k an d Bottomle y fo r designation s fo r
chloroplast genes , no w univerally adopted. I n 198 8 Lonsdal e an d Leaver propose d
a uniform terminology for mitochondrial genes. CPG N extended the basic principles
to al l sequenced plan t gene s (CPGN , 1994) .
C. Gene families. Th e guidin g principle o f the CPG N nomenclature i s that all
genes throughout the plant kingdom that encode the same product are members of the
same gene family, (cf . Fig. 1) . Th e same product means that if the sequence s o f two
products are identical , the y are obviously the same. Bu t small differences in coding
sequences commonl y occur within and among species, as in RbcS genes, such that th e
products-the small subunits of ribulose-bisphosphate carboxylase-may be very similar
but ar e no t identical . Thes e slightl y different smal l subunits are nonetheles s per ceived a s being "the same" and a s belonging to a single, plant-wide family o f genes .
There ar e man y instance s wher e gene s ar e classifie d furthe r o n th e basi s o f
sequence; e.g., four distinguishabl e sequence s encod e functionall y identical larg e
subunits o f ADPglucose synthase , and fou r distinguishabl e sequences encod e func tionally simila r superoxid e dismutases. Fo r gene s encodin g both set s o f proteins ,
coding sequences withi n a set of genes are 80 to 90 %, but may be only 50 % similar
among sets, both within and among species.
The requiremen t fo r simila r codin g sequence s i s no t absolute . Protein s dis playing activitie s o f 1-aminocyclopropane-l-carboxylat e oxidase , fo r example , ar e
encoded b y gene s wit h a variet y o f codin g sequence s tha t d o no t fal l int o an y
discernible natura l groupings . I t seem s reasonabl e i n suc h case s t o assig n thos e
genes, a t leas t fo r the present , t o a single family .
D. Gene families and multigene families. Multipl e gene s tha t belon g t o th e
same plant-wide gene famil y within a single species comprise a multigene family. Thi s
is consistent wit h conventional usage . Differen t member s of multigene families ar e
distinguished by member numbers, based generall y on th e chronolog y of isolation .
E. Gene designations. A specifi c gen e in an individua l species o f plant is full y
defined b y four fields :
Membership i n a plant-wide gene famil y
Species o f plant
Genome (nuclear , chloroplastic , mitochondrial, viral)
Membership in a multigene family
The gen e symbo l identifying memberhi p in a plant-wid e gene famil y i s usuall y
three letters followed by a number, such as Adh1 or Gln2, but mor e letters ma y be
used a s needed. Th e letter s can be followed by a capital letter instea d of a number
where dictated b y prior usag e or b y homology with bacterial genes, such as RbcS o r
TubB.
Nuclear-encoded gene s alway s star t wit h a capita l letter, a s i n th e example s
above, while genes encoded i n chloroplasts or mitochondri a start with a lower-case

Plant Molecular Biology an d Gene Designations 10

letter, such a s rbcL or cox2. Thus , th e gen e encoding the smal l subunit of ribulosebisphosphate carboxylas e i s RbcS i n higher plant s but rbcS i n chromophytic algae.
The fou r field s denotin g a gen e shoul d b e state d explicity . A n abbreviate d
format ca n als o b e use d provide d i t i s defined i n advance. Th e thir d membe r o f a
multigene famil y in A. thaliana, encoding th e light-harvesting complex type I LHCII,
for exampl e (cf . Fig. 1) , could b e represente d a s Lhcb1;At;3 .
F. Public Databases of Plant Genes. Detaile d informatio n o n gene s i n a
number o f plan t specie s ar e availabl e throug h a maste r database , PGD, whic h i s
maintained b y th e Nationa l Agricultura l Librar y o f th e USDA . Th e specie s
represented i n PGD includ e A. thaliana, maize and other grains , an d soybean, an d
are bein g incremente d o n a n almos t dail y basis . PG D als o contain s th e CPG N
listings o f sequenced plan t genes . Th e electroni c addres s o f PGD is:
http://probe.naiusda.gov:8300.html

Fig. 1. Designations for plant-wide gene families. Th e CPG N classifies sequenced plan t genes into
families based primarily on the functio n o f the gene product . I n this example of the light-harvesting
complex typ e I LHCII, every gen e i n the plan t kingdom encoding this protei n i s a membe r o f thi s
gene famil y an d bear s th e gen e symbo l Lhcbl. Th e ISPMB number i s a consecutivel y applie d
identification numbe r use d i n the management o f the CPG N databases. Gene product numbers ar e
part o f a numerical system being developed b y the CPG N for the classification o f related familie s of
genes analogous t o Enzyme Commission numbers. Specifi c gene s in individual species of plants ar e
identified b y a name or mnemonic o f their plant-wide gene family, the genus and species of the plant,
and member numbers, representative o f thei r occurrence in multigen e families .

108 Plant

Biochemistry an d Molecular Biology

References

CPGN. 1994 . Nomenclatur e o f Sequenced Plan t Genes . Plan t Mol. Biol . Reptr . 1 2 Supplement:
S1-S109.
Hallick, R.B. 1989 . Proposal s for the naming of chloroplast genes. II . Updat e to the nomenclature
of gene s fo r thylakoi d membran e polypeptides . Plan t Mol. Biol. Reptr. 7:266-275 .
Hallick, R.B., an d W . Bottomley. 1983 . Proposal s for the namin g of chloroplast genes . Plan t Mol.
Biol. Reptr . 1:38-43 .
Jansson, S. , E. Pichersky , R. Bassi , B.R . Green , M. Ikeuchi, A. Melis , D.J. Simpson , M . Spangfort,
L.A. Staehelin , an d J.P . Thornber . 1992 . A nomenclatur e fo r th e gene s encodin g th e
chlorophyll a/b-bindin g protein s o f higher plants. Plan t Mol. Biol. Reptr . 10:242-253.
Lonsdale, D.M. an d C.J . Leaver. 1988 . Mitochondria l gene nomenclature. Plan t Mol. Biol. Reptr .
6(2):14-21.
We ar e indebte d t o th e authoritie s wh o compris e ou r mor e tha n sixt y working group s fo r th e
development o f CPGN's commo n nomenclatur e for sequenced plan t genes. Withou t their expertis e
and continuing input, we would b e unable to make databases o f approved designation s for sequence d
plant gene s availabl e to th e scientifi c public, and without charge.

Consultants for Section 1

Sewell P . Champ e Eri
Waksman Institut e Coo
Rutgers Universit y Rutger
Piscataway, Ne w Jersey Ne
Frank B. Salisbury Juli
Utah Stat e Universit y E.I
Logan, Uta h Experimenta

c La m
k Colleg e
s University
w Brunswick, New Jerse y
e Vogel
. D u Pont d e Nemours & Co., Inc.
l Station
Wilmington, Delawar e

IV
PLANT GROWTH AND DEVELOPMENT
The physica l an d chemica l processe s i n plant s tha t for m th e basi s fo r th e units ,
symbols, an d term s presente d i n th e previou s section s al l tak e plac e i n th e highly
complex machiner y tha t w e recogniz e a s a livin g organism: th e roots , stems , an d
leaves wit h their xyle m an d phloem, parenchym a and pith, and all the othe r tissue s
made up of cells that in turn are highly complex machines with their cytosol, nucleus,
and sundry organelles. Thos e physical and chemica l functions are targets fo r much
study i n plan t physiology , bu t th e trul y astoundin g thin g abou t livin g organism s
including plants i s that the y are sel f generating. Al l that machiner y comes fro m a
single cell , the zygote . Discoverin g ho w this happens i s the goa l o f studies i n th e
subfields o f plant growt h and development. Thi s section attempt s to assembl e th e
terms, symbols, and unit s of measurement used by researchers i n those fields.
Most growt h an d developmen t i s influenced , sometimes strongly , b y changin g
environmental factors . Thi s is obvious in the case of the tropisms and the induction
of reproductiv e growth , but i t ma y be a little les s obvious in some other case s such
as responses to salinity or chilling stress .
This section s begin s wit h terms an d unit s that describ e th e change s i n siz e an d
complexity tha t w e recogniz e a s growt h an d development , continue s wit h plan t
movements an d reproduction , an d the n present s a larg e chapte r wit h subsection s
based o n plan t response s to various stress factors.

12
MORPHOGENESIS AND THE KINETICS OF PLANT GROWTH
Ralph O . Erickson1
Department o f Biolog y
University of Pennsylvani a
Philadelphia, Pennsylvani a 1910 4 U.S.A .
This chapte r deal s wit h terms used i n studying the kinetic s of growth. A few terms
are defined within other definitions; these are also printed in boldfaced type. Word s
in italics ar e themselve s define d elsewhere ; italic s ar e als o use d fo r symbol s tha t
represent physica l quantities (a s in the res t o f this book) .
1. THE BIOMETRY OF GROWTH.
absolute growth rate Rat e o f chang e o f x (size ) wit h respec t t o t (time) , dt/dt ;
dimensions o f x t- 1 (e.g. , m m d- 1). Whe n th e rat e i s approximately constant fo r a
period o f time, growth i s then terme d linear.
allometry Simpl e allometr y obtain s when the relativ e growt h rates o f two measured attribute s o f a n orga n o r organism , y an d x, ar e i n a constan t proportion .
Formulated a s y = ax k, log y = log a + klogx, wher e a i s a constan t an d k i s th e
allometric coefficient; dimensionless.
anisotropic Havin

g differen t propertie s alon g different directions .

anisotropic growth Magnitud

e of growth vector differ s i n different directions .

autocatalytic growth function (or curve)

function. Th e transformation ,

a symmetrica l sigmoi d
is linea r an d ca n b e use d t o

evaluate a an d k graphicall y or b y least square s fo r given data o n x vs. t.

cell production rate Loca

l rate o f formation of new cell walls.

convective rate of change Rat e of change associated with movement of a particle

to a new location i n the growt h field .
deposition rate Loca l net rate of production or import of a metabolite;
where p i s density, t is time, and u is growth velocity.
1Current address is: Ralp h O . Erickson , 192 0 Dog Kennel Road , Media , PA 1906 3
111

112 Plant

Growth and Development

determinate growth Obtain s whe n the approximate maximum size of an organ o r

organism i s determined, presumedl y genetically, a s in the growt h of a leaf. Growt h
curve i s usually sigmoid.
exponential growth function (or curve) Assumin g that the relativ e rate of change
ofx wit h t, r=dlnr/dt, i s constant, x=aert, or lnx=lna+rt. Plottin g Inx against t gives
a straigh t line , fro m whic h a and r may be estimated .
growth Increas e in a measured attribute , x, of an organ or organism, as a function
of time, t ; x = f(t).
growth field Representatio n of the spatial distribution of a developmental variable .
growth velocity v x; rate of displacement o f a material particle locate d at x.
indeterminate growth Obtain s whe n growth is potentially not limite d (excep t by
external factors) , as is often th e cas e with apical growth of a root o r shoot .
kinematics Th e stud y o f motion and shape change, apar t fro m consideration s of
mass and force .
local derivative Rat e of change associated wit h a spatial location; e.g. , the rat e of
change o f the positio n 3 mm from th e ti p of the root .
logistic function Sam e a s autocatafytic growth Junction.
material derivative Rat e o f chang e associate d wit h a real , o r material , plan t
element.
monomoleculargrowth function (or curve) x = a(l - e' kt); represent s increase of
a measure d attribute , x, at a decreasin g rate , to a maximu m value , a. The linea r
transformation, ln(a - x) = In a - kt, allow s graphical or least-square s estimation o f
a an d k fro m dat a o n x vs. t.
relative elementa l growth rat e (RELEL) Fo r growth along an axis, x,
for growt h of a surface, referred to coordinates , x, y, the growt h
RELEL
tensor i s used; REGR

dimension, t-1.

relative growth rat e Rat e of change relative t o x with respect t o time and proporMay b e symbolize d r; dimension , t- 1. Whe n r i s
tional t o
approximately constan t to r a period o f time, growth is then terme d exponential .
Richards growth functio n (o r curve ) x
= a(l be-kt)1/(1-m; take s a variety of
asymmetrical sigmoi d forms . Th e transformation , I n ((x)1-m + 1) = In b - kt , i s
a

linear fo r appropriate choic e o f sign, a and m, an d can be used to evaluate b and k ,

from give n dat a o n x vs. t. A computer solutio n i s advisable.
sigmoid growt h curv e A plo t o f a measured attribute, x, agains t time, t, showing
an early phase of acceleration, an d later a deceleration phase, approaching a limiting
value.

Plant Growth and Morphogenesis 11

2. SHOOT AND ROOT MORPHOGENESIS

apical cell Th e singl e initia l presen t i n th e apica l meriste m o f som e root s an d
shoots, as in many lower vascular plants. Divide s to form a segment cell merophyte,
and a successor apical cell; this process usuall y occurring periodically.
decussate O f leaves on a stem, arranged in pairs at each node, each at right angles
to th e nex t pair abov e o r below . Symbol , 2(1,1).
development Chang
differentiation Increas

e fro m on e patter n o f growth to another .

e i n histological complexity.

divergence angle Inphyllotaxis, th e difference i n angular position of two successive

leaves o n a stem, primordia a t the shoo t apex , scales o n a cone, etc .
Fibonacci sequence Th e integer sequence, 1 , 1, 2, 3, 5, 8, 13, . . ., in which each term
(after th e firs t two ) i s th e su m o f th e precedin g tw o terms . Ther e ar e relate d
sequences, suc h as the Lucas sequence, 1 , 3, 4, 7, 11, . . .(See phyllotaxis).
generative helix or spiral Th e sequence of leaves on a stem, or of primordia at th e
shoot apica l meristem , numbered i n order o f distance along the axis , or i n order of
radial distanc e fro m th e cente r o f the meristem . Ther e ma y be one or more spirals
in k-jugate phyllotaxis.
growth zone Tha t portio n o f a root , shoot, o r othe r structur e in which cells ar e
formed an d enlarge .
initial (1 ) A meristemati c cell tha t divides to for m a new initial cell, plu s a cell
that divide s furthe r t o ad d cells t o th e plan t body; (2) a meristemati c element tha t
differentiates int o a mature specialized element , as a metaxylem initial.
intrusive growth Tha t type of growth in which the growing cell penetrates between
existing cells and in which new areas of contact ar e formed betwee n the penetratin g
and neighborin g cell s (cf . symplastic growth).
leaf plastochron index (LPI) Indicate s th e developmenta l age of a leaf; fo r a leaf
with seria l numbe r i , LPI = P I - i . Whe n th e lengt h o f a lea f i , Li, equal s th e
reference length , Lr, the n LPI = 0.
meristem Tha t portio n o f a growt h zone i n whic h cel l formatio n (cell division )
occurs, accompanie d b y cell enlargement.
parastichy Inphyllotaxis, a helical or spira l rank of leaves or scale s along a stem,
or aroun d th e cente r o f a shoot apica l meristem; e.g., a 3-parastichy passes through
every third leaf along th e stem .
phyllotaxis Th e arrangement of leaves on a shoot, of primordia at the shoot apical
meristem, o f scale s o n a pine cone , etc. ; formulate d b y citin g a pai r o f oppose d
parastichies connecting neighborin g leaves that differ i n serial number by integers, m
and n; symbol (m,n). Thes e integers are often consecutiv e terms of the Fibonacci sequence, as (3,5). Wher e whorls of k leaves occur at each node, the symbol is k(m,n).

114 Plant

Growth and Development

plastochron (1 ) The period of time between the commencement of two successive,

repetitive processes, a s between th e initiation s o f two successive leaf primordia ;
(2) by extension, th e tim e perio d betwee n correspondin g stage s o f development o f
two successiv e leaves .
plastochron ratio (a) (1 ) At the shoot apex, a = rn/rn+l, where rn is the radial distance fro m th e ape x to the cente r o f a leaf primordium n , and rn+1, i s the distanc e
to th e nex t younger primordium ; (2 ) more generally, the rati o o f a measured attri bute of two successive leave s o f a growing shoot, as, for leaf length (L), a = L n/Ln+l.
plastochron index (PI) Indicate

s comparativ e developmenta l ag e of a shoot :

, where L i s leaf length, Lr i s reference leaf length (e.g. ,

L, = 1 0 mm), and n is serial number of the leaf just longer than Lr. Assume s leaves
are growin g exponentially with equal relative rates, and plastochron is constant.
relative plastochron growth rate Relative growth rate o f leaves or lea f primordia ,
regarding time as measured in plastochrons; p = ma, where a is the plastochron ratio;
dimension, t- 1, where t is measured i n plastochrons.
steady growth A s i n a root , growt h in which the patter n of cell productio n an d
enlargment i s invarient with time.
symplastic growth Adjacen t cel l walls do not alter thei r position relative t o each
other, and no new areas of contact are formed; this leads to expansion with minimum
shear i n the cros s section .
CONSULTANTS
Peter W . Barlo w
University o f Bristo l
Bristol, Englan d

R.F. Lyndo n
University of Edinburg h
Edinburgh, Unite d Kingdo m

P.W. Ganda r
Plant Physiolog y Division , DSI R
Palmerston North , Ne w Zealan d

T. Sach s
The Hebre w Universit y of Jerusale m
Jerusalem, Israe l

Paul B . Gree n
Stanford Universit y
Stanford, Californi a

Frank B . Salisbury
Utah Stat e University
Logan, Uta h

Zygmunt Hejnowic z
Silesian Universit y
Katowice, Polan d

Wendy Kuhn Silk

University of California
Davis, California

Aristid Lindenmaye r
University of Utrecht
Utrecht, Netherland s
(deceased 1989 )

R.F. Williams
Australian Nationa l University
Canberra City , Australia

13
GROWTH ANALYSIS AND YIELD COMPONENTS
Bruce G. Bugbe e
Plants, Soils and Biometeorolog y Departmen t
Utah Stat e Universit y
Logan, Uta h 84322-482 0
1. GENERAL CONSIDERATIONS

Plant growt h analysi s values an d yiel d component s ca n easil y and convenientl y b e

expressed wit h S I units, an d yet S I units are ofte n incorrectly used t o repor t thes e
measurements. Lik e many units reported in the literature, the reasons are historical.
A majo r reference boo k o n th e subjec t of plant growth analysis (Hunt, 1982 ) use d
metric and not S I units. However , a recent updat e of this book (Hunt , 1990) i s now
consistent with SI rules, except for the use of non-base units in the denominator (mg,
weeks), which could b e avoided (se e Chapter 1) . Th e widespread use of the hectar e
as a unit of area ha s caused thi s unit to remai n allowable in many applied agricultural journals, but its use creates a communication barrier between basic and applied
researchers tha t could be resolve d i f both groups used th e S I unit for area: m2.
The mos t problematic growth analysis parameters are those with a unit of time
in the denominator . Unlik e measurements that integrate plant physiological func tions ove r a shor t tim e interva l (seconds) , growt h analysis measurements typically
integrate the results of many physiological functions over days or weeks. A measured
carbon exchang e rat e would be expressed with SI units as mol m- 2.s-1 with positive
values durin g the ligh t perio d and negativ e value s durin g the dar k period . The
integrated result o f the diurnal carbon flux, th e crop growth rate, is typically expressed a s kg.m-2.d- 1. Thi s uni t i s no t strictl y allowabl e according t o som e author s
because o f th e us e o f day s i n th e denominator , bu t th e correc t unit , kg.m-2.s-1,
does no t indicat e th e tim e o f da y when th e integratio n occurred. Despit e thes e
problems, th e recommende d tim e interval is day for crop growt h rate. Th e us e of
days in the denominator is also important when expressing integrated daily radiation
flux (se e Chapte r 9). Hun t (1990 ) argues that some biological processes need t o be
integrated ove r lon g interval s an d thu s use s week s i n th e denominato r o f som e
growth analysi s units . Th e us e o f week s a s a uni t o f tim e i s no t recommended .
Unlike a 24- h interval, a 7-d interval ha s no special significanc e to a plant. Whe n
measurements are mad e at weekly intervals, the results should be reported as a daily
average. Thi s avoid s the confusio n of an additiona l unit o f time.
115

116 Plant

Growth and Development

Specific leaf area , a common growt h analysis measurement, ha s historically been

reported a s cm 2 g- 1. Th e us e o f thi s uni t resulte d i n a rang e o f number s fo r th e
parameter between abou t 5 0 and 300 cm2 g-1, but the gram is not an SI base unit and
its us e shoul d b e avoide d i n th e denominator . Th e bas e uni t i s the kilogram , kg.
Presumably in an effort t o utiliz e SI units, specific leaf area ha s been reporte d a s m2
g-1, whic h result s i n a very small rang e o f values fo r th e paramete r (betwee n 0.00 5
and 0.030). Th e preferre d unit , m 2 kg- 1, results i n a range of values between 5 and
30; it shoul d therefor e b e used t o expres s specifi c leaf area .
2. UNITS FOR GROWTH ANALYSIS AND YIELD COMPONENTS
The following are some examples of preferred units for reporting growth analysis
measurements an d yield components . Remembe r tha t denominator s shoul d b e SI
base units, wit h th e exceptio n o f time wher e da y is accepted. Numerator s wit h o r
without prefixe s ca n b e use d s o tha t th e rang e o f values wil l be betwee n 0. 1 and
1000. Th e followin g example s sho w th e bas e unit s i n bot h numerator s an d
denominators, a s is customary when showing SI units.
Table 1. Terms and Units for Plant Growth Parameters
relative growth rate
or specific growth rate

kg.kg-1.d-1 (g i s appropriat e fo r th e numerator ; g.g-1.d- 1 i s

the uni t most ofte n used , bu t i t i s not recommende d becaus e i t
uses g, a non-Si bas e uni t i n the denominator )

net assimilation rate

kg.m-2.d-1 ( g is appropriate fo r th e numerator) .

leaf area ratio

m2.kg-1 (m
2.

lea f area pe r k g total dr y biomass) .

lea f are a pe r k g leaf dry mass).

specific leaf area

m kg- (m

specific leaf mass

(the invers e of specific lea f area, ofte n calle d

kg.m -2
specific lea f weight).

leaf mass ratio

kg.kg-1 (k

crop growth rate

kg.m-2.d-1 ( g is appropriate for th e numerator ; it places th e

typical rang e of values between 1 and 100) .

seasonal crop yield

kg.m-2

harvest index

The ratio of harvested or edible (usable) biomass to total biomass

or to above-ground biomass. Th e convention used must be specified. A s a ratio , harves t inde x i s dimensionless ; i t ca n b e
expressed a s a decima l fraction o r a s a percen t (b y multiplyin g
the fractio n b y 100) .

nutrient concentration in
plant tissue
(N, P, K, etc.)

mol kg- 1. Thi s is typically expressed a s a percent, g.kg- 1, or a s

ppm. g.kg- 1 i s a correc t Si-unit , bu t inorgani c element s ar e
pure substances, and thei r concentratio n is thus better expresse d
as mole s rathe r tha n a s mass . Expresse d o n a mas s basi s i t
appears that N an d K are a t simila r level s in plant tissue; i n fact ,
the numbe r of N atoms is about 3 times higher tha n the number
of K atoms. Th e physiologica l importanc e o f nutrient s i s determined b y their number , not thei r mass.

g lea f mas s per k g total dr y biomass).

Growth Analysis and Yield Components 111

Other compound units are use d in plant growth analysis and yield. Adherenc e
to SI rules should be made with the exception of the use of days in the denominator
for som e compoun d units .
Note agai n tha t th e bes t uni t of area i n the denominato r is the squar e meter.
If th e magnitud e of values i s above 999 or belo w 0.1, th e valu e of the uni t in th e
numerator shoul d b e change d b y adding or deletin g the appropriat e prefix . Fo r
example, 0.0 3 kg-m^-d" 1 i s incorrect ; 3 0 g-m^-d" 1 i s correct . (Thi s rul e ca n b e
broken i f th e rang e o f values being compared i s greater tha n 1 0 000 because th e
comparison amon g values is easier i f the sam e prefix i s used throughout.)
Table 2. Typical Ranges of Values for Plant Growth Parameters.
Unit

Typical
Value

Trend over Time

g-kg-'-d- 1

10 to 400

decrease

net assimilation rate

or unit leaf rate

g-m^-d-1

1 to 30

decrease

leaf area ratio

m2-kg-1

10 to 60

decrease

specific leaf area

m^kg-1

5 to 30

decrease

leaf mass ratio

(leaf weight ratio)

kg-kg'1

0.4 to 0.8

decrease

g-m^-d'1

1 to 40

rapid increase
then gradual
decrease

/Mnol-m^-s"1 c

1 to 40

decrease

Parameter
relative growth rate or
specific growth rate (RGR)

crop growth rate

single leaf photosynthetic rate

(CO2 exchange)
leaf area index

-2
leaf ground

0.01 to 10

large increase
(sigmoid curve)

" Th e uni t g- g d i s a commonly used unit for RGR, but it is not recommende d because the unit of
grams is not a base uni t an d shoul d no t b e used i n th e denominator. Th e uni t ( d ) is also used fo r RGR be cause the gram s cancel ou t Thi s is confusing, however , because RGR represent s g new growth per g of existing
biomass.
Leaf weigh t ratio is th e commonl y use d ter m bu t k g is a unit o f mass , s o weight i s incorrect
c

Us e moles for pur e substances , suc h a s (imo l o f CC^; us e kilograms or grams for mixed substances , such as
grams o f biomass (ne t assimilation rat e o r cro p growth rate , g-m - d ) .

Table 3. Growt h analysis quantities derived from plan t mass and leaf area
Symbol

Instantaneous
Value

Absolute Growth Rate

AGR

dm/dt

Relative Growth Rate

XGR

\im - dm/dt

Leaf Area Ratio

LAR

Specific Leaf Area

Individual Plant

Derived Quantity

u!

Formula for Mean Value over Time

Interval (t rt,)

Dimensions

Units

m -t' 1

kg-d' 1

Qnm2-\n m,) I(t2 -/,)*= NAR -LAR

m-m"' -r'

kg-kg-'-d' 1

LA/m

[(LAI/m, ) + (L^/m^/2 =

A -m-'

m2-kg

SLA

LA/Ln

[(.Lu/L^ + duOL^W

A -m' 1

m2-kg

Leaf Mass Ratio

1MR

LJm

[(Z,n/m;) + (Z.^/m,)]/2

m -m'1

kg - k g1

Specific Leaf Mass

SIM

L./LA

\(Lml/LAI) + (L

m-A-'

kg m' 2

Net Assimilation Rate

NAR

( 1 I LA ) - dm/dt

(nh - m,V(f, - 1,) (In LA2 - In LA1)I(LA2 - L A1)

m-A-' -f

Percent Dry Mass

PDM

(mD/mF) 10 0

Crop Growth Rate

CGR

1/GA dm/dt

Leaf Area Index

LAI

LA/GA

(m2 -m,) I{t2-t1)" = Am/ At

SLA -LMR

M/La)V2

kg m'2 d' 1

[(mDI/mn) + (m D2/mr2)}/2

% = 0.01

(m2- m,)l(t 2 - 1,) 1/GA = NAR LAI

m-A-' T '

kg m-2 d' 1

(LAI+LM)/2-l/GA

A -A' 1

m2 m- 2

LA = Leaf Area, L m - Lea f mass, G A = Ground Area , / = time, A = Area, m = mass = Weight, m DORf = Dry or Fresh mass. Not e tha t m = meter, but m = mass (became roman typ e
is used for units but italic type i s used for physical quantities ; see Chapter 1) .
a
b

The most frequently used form of each equation i s boldfaced .

In m 2 - In m, = In (m 2/m,).

Growth Analysis an d Yield Components 11

REFERENCES
Causton, Davi d and Jil l Venus . 1981 . Th e Biometr y o f Plant Growth . Edwar d Arnold , London .
Hunt, R . 1978 . Plan t Growt h Analysis . Edwar d Arnold , London .
Hunt, R. 1982 . Plan t Growt h Curves : Th e Functional Approach t o Plant Growth Analysis . Univ .
Park Press. Baltimore .
Hunt, R. 1990 . Basi c Growth Analysis . Unwi n Hyman Ltd., London , UK ; and Winchester, MA.
CONSULTANTS
Ray Wheele r Charle
NASA Kenned y Space Cente r Agricultur
Florida Kentville
Carl Rosen
University of Minnesot a
St. Paul, Minnesot a

s F. Forne y
e Canada
, Nova Scotia, Canad a

14
PLANT MOVEMENT S
Wolfgang Haup t

Institut fii r Botani k un d Pharmazeutisch e Biologie

der Universita t Erlangen-Nilrnber g
Staudtstrasse 5 , D-91058 Erlangen GERMAN Y
This chapte r deal s wit h terms use d i n th e stud y o f plant movements. A fe w terms
are defined within other definitions ; these are also printed in boldfaced type. Word s
in italics are themselve s define d elsewhere i n this chapter .
1. TYPES AND MECHANISMS OF MOVEMENT
ciliary movements Bendin g of cilia or flagella of eukaryotic cells, caused by sliding
of microtubula r doublet s alon g eac h other , usuall y resulting i n locomotion . Th e
microtubules slid e because o f a tubulin-dynei n interaction.
flagellar movements Rotatio n o f flagell a i n bacteria , drive n b y proton-motiv e
force, resultin g in locomotion .
running (running phase) Locomotio n o f peritrichous bacteria (having a uniform
distribution o f flagell a ove r th e bod y surface; movement more o r les s straigh t forward), usually caused b y counterclockwise rotation o f flagella .
tumbling (tumbling phase) Irregula r movement o f peritrichous bacteri a withou t
locomotion, usuall y caused b y clockwise rotation o f flagella. Th e running phase i s
interrupted b y a tumbling phas e eithe r autonomously in more o r les s regula r inter vals, o r a s a respons e t o a phobic stimulus. Usually , the directio n o f movement i s
randomly different befor e and afte r a tumbling phase.
cytoplasmic streaming Movemen t of cytoplasm (sometimes with nuclei) within the
cell a s th e resul t o f actin-myosi n interaction; differen t type s are distinguishe d according to th e regularit y of velocity in time and space with which the cell organelle s
are displaced. (Not e tha t th e ter m velocity comprises both speed an d direction! )
growth movement Irreversibl e curvatur e caused by differential growth ; i.e., plasti c
extension o f cell walls, either differentiall y i n a single cell on opposite flanks, or, in
a multicellula r organ , by differential cell growth on opposite flanks .
Note: A growt h curvature can apparently be reversed , but i n fact thi s is caused
by an opposite curvatur e located clos e t o th e firs t one .

120

Plant Movements 12

bending an d bulgin g Differentia l growth can caus e a curvatur e in eithe r o f two

ways: (1 ) In an organ with a subapical growing zone, curvature is oriented awa y from
the growth-stimulated flank (o r toward the growth-inhibited flank) : bendin g (some times also bowing). (2 ) In an organ with an apical growth center, shifting of maximal
growth ou t o f the cente r result s i n outgrowth in that direction; hence , curvatur e is
oriented towar d th e growth-stimulate d side : bulging .
torsion Growt

h movemen t involvin g twisting of the organ .

turgor movement Reversibl e curvatur e caused by asymmetric change in length o r

volume, eithe r in a single cel l b y asymmetric elastic extension/relaxatio n o f the cell
wall upon uptak e or loss of water, or in a multicellular organ by differential swelling/
shrinking of cells o n opposite flanks.
cohesion movemen t A chang e in cel l shap e resulting in curvature and caused by
a los s o f water beyon d th e ful l relaxatio n o f the extende d elasti c wall. Becaus e of
cohesion o f water in the vacuole, its adhesion t o the surrounding cytoplasm, and the
firm attachment o f the latter t o the wall, this loss of water results in an elastic inward
deformation o f the wal l (negative turgor) . Thi s slow movement can be followe d by
a sudden return of the cell to its original volume and shape if the elastic deformation
becomes stronge r tha n the cohesion, thu s allowing water-vapor bubbles to be formed
in th e vacuole . A classica l example : annulu s cells o f the fer n sporangium .
hygroscopic movemen t (als o swelling, shrinking ) Reversibl e curvatur e caused by
water uptak e into/wate r loss ou t o f intermicellary spaces o f the cel l wall ; does no t
need livin g structures onc e a suitable wal l texture ha s been developed .
2. CONTROL OF MOVEMENT: GENERAL
autonomous (endogenous) Self-generate d movement, not depending on an external stimulus; the controllin g facto r is sometimes calle d internal stimulus.
induced (exogenous ) A n externa l stimulus determine s tha t a movemen t (o r a
change i n movement ) start s or ho w it i s executed.
stimulus (signal, input signal) A physical or chemical factor that, when interacting
with th e organis m o r cell , ca n elici t a movemen t o r modif y a n alread y existin g
movement. It s quantity is characterized b y its intensity and its duration. I t can act
in a scalar o r i n a vectorial way . A stimulus regulates the movemen t bu t doe s no t
provide th e energ y for it . Thus , i f such a factor als o provides energ y for th e move ment, on e ca n debat e whethe r i t shoul d b e calle d a stimulu s (e.g. , photokinesis in
cyanobacteria, desmids , etc.) .
receptor A structur e o r substanc e i n cell s o r organisms , with which a stimulus
interacts (stimulus perception).
perception Interactio
processes.

n betwee n stimulus and receptor tha t starts th e transduction

susception Sometime s th e ter m perception i s restricte d t o thos e processe s tha t

involve livin g structures . I n thi s case , th e ver y first , pur e physica l effec t o f th e

122 Plant

Growth and Development

stimulus i s calle d susception; e.g. , displacemen t (sedimentation ) o f a statolith-lik e

structure, or excitatio n o f a pigment molecule. Perception i n the stric t sense , then ,
would be the actio n o f the statolith o r th e excited pigment molecule on the respec tive sensitive cel l structur e (e.g. , a membrane).
transduction (stimulus transduction, signal transduction, transduction chain)
Biochemical o r biophysica l event s tha t follo w perception, finall y resultin g i n th e
response; transductio n usuall y includes amplificatio n processes an d a tim e la g between perceptio n an d response.
signal transmission Par t o f th e transduction processe s bridgin g the locatio n o f
perception an d tha t o f respons e i f they do no t occu r a t th e sam e sites (structures ,
compartments, cells , organs) .
latency (latency time) Tim e interva l betwee n star t o f stimulus and response . It s
precise definitio n depend s o n th e respons e paramete r tha t i s measure d (e.g. , i n
phototropism th e degre e of curvature measured unde r th e microscop e o r wit h th e
naked eye ; a differential membran e potential).
reciprocity (reciprocity law; in photoresponses: Bunsen-Roscoe law) Th e respons e
depends quantitativel y neithe r o n th e intensit y o f th e stimulus alone, no r o n it s
duration, but o n the produc t of both parameters. I f reciprocity holds, this is usually
taken as indicating a lack of rate-limiting processes in the transduction chai n o r i n
recovery processes o f the receptor. Accordingly , reciprocity is restricted t o a certain
range of intensity and duration of the stimulus . (I t often doe s no t hol d for very low
intensities o r very short durations. )
dose Tha t fractio n of the applied stimulus (product of intensity and duration) that
reaches th e receptor.
adaptation Relaxatio n o f response wit h persistent o r repetitiv e stimulation ; it is
sometimes difficul t t o distinguis h whethe r th e stimulus change s th e receptor o r
transduction syste m suc h tha t sensitivit y o r responsivit y i s reduce d (sensory o r
response adaptation).
tonic effect Chang e i n sensitivity or responsivit y to a stimulus caused b y external
or internal conditions .
3. TERMS FOR INDUCED MOVEMENTS (TYPES OF RESPONSE)
taxis (tactic orientation; formerly topotaxis) Movemen t of motile organisms, cells
or organelles, th e prevailin g direction of which is determined by the directio n o f th e
stimulus. Strictl y speaking, taxis is not alway s a single response, but ma y be a result
of a series of responses (e.g. , chemotaxi s of bacteria).
phobic response (formerly phobism or phobotaxis) Transien t chang e of movement
of motil e organisms induce d b y a change in magnitude of a stimulus. Th e effectiv e
change ca n b e a n increas e o r decreas e o f the magnitude ; that is , a step-up o r stepdown o f th e stimulus . Afte r th e previou s movement is resumed, its directio n may
have changed, bu t wit h n o relatio n t o th e directio n of the stimulus . I n inhomoge-

Plant Movements 12

neous fields , repeated phobi c response s ca n result i n patterns o f distribution o f th e

organisms; e.g. , accumulatio n i n a "light trap".
kinesis Th e spee d o f movemen t o f motil e organism s o r cell s depends , unde r
steady-state condition s o f inpu t an d output , o n th e magnitud e (an d sometime s
probably als o direction) o f a stimulus.
dinesis Applie s whe n th e spee d o f cytoplasmic streaming or th e fractio n o f cytoplasm involve d i n streamin g i s controlled b y a stimulus.
tropism (tropistic movement) Curvatur e of organs or cells, the direction o f which
is determined b y the directio n o f the stimulus. Tropism s ar e usuall y growth move ments; solar tracking, however, i s an example of a phototropism that in some plant s
is caused b y turgor changes .
nasty (nastic movement) Curvatur e o f organ s o r cells , th e directio n o f which is
determined b y morphologica l o r physiologica l organization ; i t i s induce d b y a
stimulus bu t independen t o f it s direction . Note : th e term s epinasty, hyponasty,
nyctinasty, cyclonasty ar e use d t o describ e autonomous curvatures; thus , they do no t
deal wit h true nasties .
growth response Transien t chang e i n growt h rat e induce d b y a chang e i n
magnitude o f a n externa l factor . Th e ter m i s sometimes als o use d t o denot e th e
dependence of steady-state growt h rate on a stimulus. Mainl y used when light is the
stimulus; then it is traditionally and inconsistently called light growth response rather
than photo growth response.
strophism Torsion

induce d by an externa l factor.

4. STIMULI. Note : stimul i ar e indicate d b y prefixes attache d t o -taxis , -nasty ,

-tropism, etc. :
photo- (formerly: helio-) Induction
polaro- Orientatio

by or orientatio n wit h respect t o light.

n wit h respect t o th e plan e of polarization o f radiation.

scoto- Induction b y darkness.

Note: Strickl y speaking , darknes s canno t b e a stimulus but denote s th e absenc e
of ligh t as a stimulus .
gravi- (formerly: geo-) Orientatio n wit h respec t t o a n acceleration , especiall y
gravity, but als o centrifugal acceleration .
agravi- (formerly: ageo-) Indicate
accelerational stimulus.

s tha t the orga n does no t respon d t o th e

thigmo- (formerly: hap to-) Induction by or orientation with respect t o mechanica l

contact wit h a solid body .
Note: Thigmotropism may be use d as a general term when a distinction cannot
be mad e between tropistic an d nastic response i n tendrils.
seismo- Induction by vibration (e.g., by wind action, shaking, usually also by sudden
contact wit h a solid body or a fluid) . I t should not b e use d to replac e traumata-.

124 Plant

Growth an d Development

thermo- Induction
temperature.

b y temperatur e o r orientatio n wit h respec t t o a gradien t i n

chemo- Induction by a specific substance o r orientation wit h respect to a gradien t

of suc h a substance .
hygro- (sometimes less correctly: hydro-) Induction
or orientation with respect t o a gradient o f either.

b y fluid water or water vapor,

rheo- Induction

by or orientatio n wit h respect t o streamin g water or air .

aero- Induction

by or orientatio n with respect t o air , usually its oxygen content.

traumato- Induction

by or orientatio n wit h respect t o wounding.

electro-, galvano- Induction

current.
magneto- Induction

by or orientation wit h respect t o an electrical fiel d o r

by or orientatio n wit h respect t o a magnetic field.

avoidance response Movemen

t awa y fro m a nearby solid barrier .

step-up, step-down Denote s in which direction th e magnitude of the stimulus has

to b e change d i n order t o induce a phobic response o r a growth response.
5. DIRECTION OR SENSE OF RESPONSE
positive I n scalar stimulation : Wit h increase o f stimulus the spee d o f movemen t
increases an d vic e vers a (sometime s als o "direct") . I n vectorial stimulation : th e
movement ha s a prevailin g component towar d the sourc e o f a stimulus.
negative I n scalar stimulation : Wit h increase o f stimulus the spee d o f movement
decreases an d vice versa (sometime s als o "inverse") . I n vectorial stimulation: Th e
movement ha s a prevailing component awa y fro m th e stimulu s source.
ortho- Orientatio

n paralle l o r antiparalle l t o th e direction o f a vector stimulus.

plagio- Orientatio n a t som e determine d o r constan t angl e between 0 an d 180

to the stimulus source .
dia- (not transversal) Orientatio n a t righ t angle to a stimulus source .
Note: Fo r a ful l classificatio n o f a movement, one should combin e orientation ,
stimulus, and typ e of response; e.g. , positiv e phototaxis , plagio-gravitropism.
one-instant mechanism (spatial sensing of direction) Th e direction o f stimulus is
sensed a t on e instan t b y comparing the arrivin g stimulus at tw o (or more ) receptor
sites.
two-instant mechanism (temporal sensing of direction) Th e direction of a stimulus
is sensed a t one receptor site only, comparing the arriving stimulus at two instants i n
time; the arriving stimulus is modulated, e.g., by rotation of the cell or organism during locomotion .
6. TERMS FOR AUTONOMOUS MOVEMENTS. Note : Th e term s of this section
containing nasty i n th e wor d ar e no t nastie s accordin g to th e definition ; however,

Plant Movements 12

they are occasionall y use d i n a purely descriptive way when the controlling facto r is
not known .
nutation Autonomous growth movement ove r extende d periods , i.e. , curvatur e of
organs (o r chang e o f curvature) , cause d b y differential flan k growth , which is no t
induced b y an externa l stimulus. (Th e ter m should not b e used to denote nastic or
tropistic movements, which are induced movements. )
epinasty Growt h curvature in a morphologically downward sense, caused by faster
growth rat e o f the uppe r sid e (e.g. , adaxial side o f leaves).
hyponasty Growt h curvature in a morphologically upward sense, cause d by faster
growth o f the lowe r sid e (e.g. , abaxial side of leaves) .
nyctinasty A diurnal periodic upwar d and downward movement, usually of leave s
or petioles , mainl y controlled b y the physiologica l cloc k but , in addition , synchro nized by an external factor. Sometime s not restricted to autonomous movements, but
used als o fo r movement s tha t ar e induced b y rhythmic light-dark changes , o r onl y
those that ar e induce d b y light-dark transition in the sens e o f scotonasty.
circumnutation(formerly: cyclonasty) Periodi c change of growth curvature, the ti p
of the orga n ideall y moving around a circle or cone; movemen t occurs as the regio n
of highes t growt h rat e rotate s aroun d th e organ . Circumnutatio n i s no t alway s
autonomous but ca n be the result of tropistic stimulation s with extended after-effects .
autotropism A tendency of an organ to grow straight and to straighten a curvature
induced b y a tropistic stimulus.
CONSULTANTS
Donat-Peter Hade r Andrea
Universitat Erlangen-Nurnber g Universita
Erlangen, German y Bonn
Anders Johnsso n Hemmin
University o f Trondheim Universit
Dragvoll, Norwa y Goteborg
Francesco Lend Masamits
C.N.R., Istitut o d i Bioflsica Toky
Pisa, Ital y Tokyo

s Siever s
t Bonn
, German y
g I. Virgin
y of Goteborg
, Swede n
u Wada
o Metropolita n Universit y
, Japa n

Hans Machemer Gottfrie

Universitat o f Bochum Universita
Bochum, German y Giessen

d Wagner
t Giesse n
, German y

Wilhelm Nultsch Manfre

Universitat Marbur g Universita
Marburg, German y Karlsruhe

d H. Weisensee l
t Karlsruh e
, German y

Peter Schopfe r
Universitat Freibur g
Freiburg, German y

15
GROWTH SUBSTANCES
Robert E . Clelan d
Department o f Botany
University of Washington
Box 355325
Seattle, WA 98195-532 5 U.S.A.
Terms use d t o describe plant growth substances have been used in widely divergent
ways. A s a result, there has been little attempt to standardize the definitions. Th e
definitions presented here are based, as far as possible, on the most common usage
at present. Som e terms, such as abscisic acid and ethylene, are not defined here, as
they refer to a single compound .
A few terms ar e define d within other definitions ; these ar e printe d i n boldfaced
type. Word s in italics are themselve s define d elsewhere.
antiauxin A compoun d tha t antagonize s th e biologica l actio n o f a n auxin, and
whose inhibition kinetics are strictly competitive. Fo r example, p-chlorophenoxyisobutyric acid is an antiauxin because it shows competitive inhibitor kinetics, but 2,3 ,
5-triiodobenzoic aci d (TIBA ) an d naphthylphthalami c aci d (NPA) , whic h ar e
inhibitors of polar auxi n transport and show non-competitive inhibitor kinetics, ar e
not antiauxins.
auxin A compound that has a spectrum of biological activities similar to, but no t
necessarily identical with those of indoleacetic acid. Thi s includes the ability to: (1 )
induce cell elongation in isolated coleoptile or stem sections, (2 ) induce cell division
in callus tissues in the presence o f a cytokinin, (3) promote lateral root formation at
the cu t surfac e o f stems , (4 ) induc e parthenocarpi c tomat o frui t growth , an d (5 )
induce ethylene formation.
auxin antagonist A compound that antagonizes the biological action of an auxin.
The inhibitio n kinetic s ca n be eithe r competitiv e o r non-competitive . TIB A an d
NPA can be auxin antagonists, even though they are not antiauxins.
bound auxin A molecule i n which an auxin i s bound to another compoun d (e.g.,
sugar, amin o acid , o r macromolecule ) vi a a covalen t bond . Sometime s calle d a
conjugated auxin.

126

Growth Substances 12

cytokinin A compound tha t has a spectrum of biological activities similar to those

of trans-zeatin . Thi s include s th e abilit y to : (1 ) induce cel l divisio n i n callus cell s
in the presenc e of an auxin, (2) promote bud or root formation from callu s culture s
when i n appropriat e molar ratio s t o auxin , (3) delay senescence of leaves , an d (4 )
promote expansio n o f dico t cotyledons . Th e ter m cytokini n is often restricte d t o
compounds tha t contai n a n adenine ring structure; other compounds with cytokinin
activity are calle d cytokinin-like.
florigen A compound o r group of compounds, produced in leaves under inductive
conditions, tha t i s transported i n th e phloe m t o buds and cause their developmen t
to chang e fro m vegetativ e t o floral . T o b e considere d a florigen , a compound(s )
must b e effectiv e o n a wide range o f species, an d o n short-day , long-day, and day neutral plants . Fo r thi s reason , gibberellins ar e not considere d t o b e florigens. A s
yet, the natur e o f florigen is unknown.
gibberellin A compound containin g the ent-kaurene ring structure. I f it is active
in a higher plant , it will have a spectrum of biological activities similar to thos e of
gibberellic acid (GA 3). Thi s includes the abilit y to: (1 ) promote extension i n dwarf
genotypes o f pea , corn , an d rice , (2 ) induc e d e nov o synthesis o f hig h p I form s o f
a-amylase i n barley aleurone cells , and (3 ) induce or promote flowering in selected
long-day plants when under short-day conditions. Gibberellin-Iike is used for all substances tha t hav e GA-lik e biologica l activity, but whose chemical structure has no t
been defined .
hormone-binding protein A protein that binds the hormone in a saturable, specific
manner. (See phytohormone.)
hormone receptor A hormone-binding protein that is shown to have a physiological
effect afte r bindin g of the hormone .
hormone sensitivity A n ill-defined , ambiguous term. I n general, it refer s t o th e
amount o f respons e (i.e. , responsivity) t o a give n amoun t o f hormone . Th e
sensitivity will depen d upon : (1 ) the concentratio n o f hormone fro m bot h endoge nous an d exogenou s source s tha t is present a t th e site of action, (2) the concentra tion of hormone receptors, (3) the affinit y o f the receptors fo r the hormone, and (4 )
the coupling of the hormone-receptor t o the final observed response. I t is not usefu l
to tal k abou t hormon e sensitivit y o r a chang e i n sensitivit y unles s th e ter m i s
carefully qualifie d as to th e typ e of sensitivity that is being considered .
phytohormone (plant hormone) A compound, produced i n th e plant, that a t low
concentrations ( 1 umol.L-1 or less) modulates o r regulates some aspects o f the bio chemistry o r physiolog y of cell s distan t fro m it s sit e o f synthesis . I n man y cases,
hormones ma y als o influenc e th e cell s i n whic h they are produced . Example s of
phytohormones are : auxins, gibberellins, cytokinins, abscisic acid , ethylene .
plant growth regulator A compound that when applied at lo w concentrations ( 1
utmol-L-1 or less ) modifie s th e growth or development of the plant. A plant growth
regulator ca n b e eithe r a n endogenou s compoun d (e.g . indoleaceti c acid ) o r a
synthetic compound (e.g.a-naphthaleneacetic acid), but the term is primarily applied
to synthetic compounds.

128 Plant

Growth and Development

polar transport Transpor t tha t i s dependen t o n metabolism , mor e rapi d tha n

diffusion, an d predominantly unidirectional. Th e term has primarily been restricted
to th e movemen t o f hormones , especiall y auxins. Transpor t throug h th e phloem ,
where a numbe r o f compound s ar e movin g unidirectionall y together , i s no t
considered t o b e polar transport .

CONSULTANTS
Robert Bandurski
Michigan Stat e University
East Lansing , Michigan

Richard P . Pharis
University o f Calgary
Calgary, Alberta, Canad a

Peter J. Davie s
Cornell Universit y
Ithaca, Ne w Yor k

Bernard O . Phinney
University o f California
Los Angeles, California

Michael L . Evan s
Ohio Stat e Universit y
Columbus, Ohi o

Frank B . Salisbury
Utah Stat e Universit y
Logan, Uta h

Richard D . Fir n
University o f Yor k
York, Unite d Kingdo m

Andreas Siever s
University o f Bonn
Bonn, German y

Arthur W . Galsto n
Yale Universit y
New Haven, Connecticu t

Lincoln Tai z
University o f California
Santa Cruz , Californi a

Russell L . Jones
University o f California
Berkeley, California

Kenneth Thimann
The Quadrangle
Haverford, Pennsylvania

A. Car l Leopold
Cornell Universit y
Ithaca, Ne w York

Anthony J. Trewava s
University o f Edinburgh
Edinburgh, Scotland

Hans Moh r
Universitat Freibur g
Freiburg, German y

16
BIOLOGICAL TIMING
Willard L . Koukkari
Department o f Plant Biology , University of Minnesota
St. Paul, Minnesota 5510 8 U.S.A .
Beatrice M . Sweeney 1
Department o f Biological Sciences , University of California
Santa Barbara , Californi a 9310 6 U.S. A
This chapter deal s with terms used in the study of biological timing. A few terms are
defined withi n other definitions; these are also printed in boldfaced type. Word s in
italics are themselve s define d elsewhere .
acrophase La g o f th e maximu m (peak) o f a mathematica l curve (e.g. , cosine )
versus a reference ; th e phase angle o f the cres t o f the fitte d mode l in relatio n t o a
reference tim e point .
aliasing Misrepresentatio n o f a frequency to b e lowe r (o r perio d t o b e longer )
because intervals between consecutivel y spaced samples were too long . (Sometime s
called folding effect.)
amplitude Paramete r o f a rhythm , which fo r a mathematica l (e.g. , sinusoidal )
curve, is half the range from th e peak to the trough. I t is sometimes (e.g., by certain
astronomers an d biologists ) use d for the entir e rang e from pea k to trough.
annual Yearly

biological clock A biologica l variabl e showin g rhythmicit y (especiall y circadian

rhythms) an d implyin g a mechanis m tha t impart s time information ; se e als o ref erences t o photoperiodis m i n Chapter 17.
biological cycle Sequenc e o f events i n an organism that repeat i n the sam e orde r
and at the sam e interval through time. [Whe n used in the context of the lif e history
(life cycle ) o f an organism , the interva l through time may vary.] On e cycl e may be
represented a s a circle (360) .

1Deceased.
129

130 Plant

Growth and Development

biological rhythm A change i n a biological variabl e that recur s (repeats ) wit h a

specifiable frequenc y an d pattern . Generall y viewed a s meeting th e criteri a o f an
endogenous rhythm.
chronogram Plo t o f a measure d variabl e o n th e ordinat e ( Y axis ) agains t th e
measure o f time on the abscissa ( X axis) and used to illustrate the state of a variable
over time .
circadian pacemaker A postulated master ocillator that coordinates the period and
phase o f circadian rhythms.
circadian rhythm Biological rhythm havin g a perio d o f abou t 24 h (circa, about ;
dies, day ) an d satisfyin g othe r criteria , suc h a s havin g a labil e phase tha t ca n b e
shifted b y external environmental synchronizers an d continuin g for mor e tha n on e
cycle (free-running) i n th e absenc e o f externa l environmenta l synchronizer s (e.g. ,
under LL o r DD).
circadian time (CT ) Tim e tha t spans the circadia n period . Th e choice o f when a
cycle start s i s arbitrary , especially unde r free-running conditions. Som e biologist s
designate 0 0 h as the en d of the dark span (dawn) of an environmental LD cycle , or
the phas e correspondin g t o thi s poin t i n constan t condition s (subjective dawn).
Others selec t 0 0 h as the middl e of the dark span or the phase corresponding to this
point i n constant conditions .
circannual rhythm Biologica l (endogenous) rhyth m having a period of about (circa)
a year . Sometime s use d whe n endogenieit y has no t bee n demonstrate d o r i s no t
known.
circaseptan rhythm Biologica
(circa) seve n days.

l (endogenous) rhyth m havin g a perio d o f abou t

circatrigintan rhythm Biologica l (endogenous) rhyth m havin g a perio d o f abou t

(circa) 30 days. (Sometime s called circalunar rhythm.)
clock time Tim e provide d b y a cloc k o r watc h (nonbiologica l system) ; also se e
biological clock and circadian time.
cosinor analysis Dat a fitte d by cosine(s) of given period(s) (e.g., 2 4 h and 7 d) by
a least-squares procedure an d used to estimate rhythm parameters (e.g. , acrophase(s),
amplitude(s), an d MESOR).
cosinor (polar) display Summar y of rhythm parameters (amplitude an d acrophase
of each harmonic) estimated b y cosinor analysis an d displayed on pola r coordinate s
so that th e phas e angl e of the maximu m (acrophase) o f each cosine curve included
in the mode l is shown by the angular direction of a vector and the amplitude of each
component i s represented b y the length of the vector. Th e 95% confidence limits of
both for each harmonic are shown as an ellipse around the tip of the vector. Some times th e erro r ellipse is plotted to th e ri m for purposes o f comparing acrophases ;
in suc h cases, th e lengt h of the vecto r i s not proportiona l to th e amplitude.
damping A decreas e i n th e amplitude o f an oscillatio n (rhythm ) ove r time ; very
common i n plant s as they mature and develop.

Biological Timing 13
dark break Interruptio
DD Abbreviatio

n o f the ligh t spa n o r LL wit h a dark span (o r pulse).

n fo r continuous darkness .

desynehronization A chang e i n th e phas e relationshi p betwee n tw o o r mor e

rhythms in an organism (internal) ; o r between th e rhythms of the organism and th e
environmental cycle s (external) . Ther e ma y be desynchronization i n phase and/o r
frequency.
diel Th e 2 4 h day; 24 h period; rarely , if ever, use d when discussing plant rhythms
but sometime s use d i n animal research .
diurnal Ter m use d i n reference t o either a daily cycle or th e light spa n o f a 24 h
day (e.g. , diurna l animal s compare d t o nocturna l animals) . Dependin g upo n th e
context, ofte n bes t replaced b y other terms (e.g. , daily , circadian, light span, etc.) .
endogenous rhythm A rhythm that persists (free-runs) under constant environmental condition s fo r mor e tha n on e complet e cycle . T o qualif y a s a biological rhythm,
the oscillations must be shown to repeat wit h approximately the same period i n the
absence of environmental cycle s o r synchronizers .
entrainment Couplin g o f th e period o f on e rhyth m t o tha t o f anothe r cycl e o f
about th e sam e length ; fo r example , the settin g o f a circadian rhythm to exactl y 24
h by an LD environmenta l cycle . Thi s usuall y involves both period and phase.
free-running rhythm Se
frequency Th

e endogenous rhythm.

e numbe r o f cycles in a unit tim e o r I/period .

frequency demultiplication Synchronizatio n t o a lon g period by a cycl e that i s a

submultiple o f that period . (Th e convers e ma y also be observed, i n which case i t is
referred t o a s frequency multiplication.)
h Abbreviatio

n ofte n use d fo r hour(s) . (Se e Tabl e 5 , Chapter 1. )

harmonic Ter m use d t o describ e feature s of a periodi c curve . A nonsinusoida l

periodic function can be mathematically expresse d as the sum of cosine curve s with
period ta u ( ) , tau/2 , tau/3 , ...etc . Ta u i s called th e fundamental period, an d th e
cosine wit h perio d ta u i s th e fundamental term or first harmonic. Cosine s wit h
periods tau/2 , tau/3...etc . ar e calle d th e second, third, ... etc. harmonics. A curv e
consisting onl y of a fundamental ter m i s purely sinusoidal with period tau . [Note :
Tau i s sometimes use d a s an abbreviatio n fo r period.]
high frequency oscillations Ultradian oscillations o f biologica l variable s havin g
periods less tha n 3 0 minutes. Se e ultradian rhythms.
hour-glass timer A mechanis m capabl e o f timing only a singl e tim e perio d an d
likened t o a n hour-glas s (b y contrast t o a pendulum timer); thus , not a rhythm.
infradian rhythm Biological rhythm having a period appreciably longer tha n 24 h;
usually periods longe r tha n 2 8 h; therefore, a circannual rhythm i s infradian .
LD Abbreviatio n for a light:dark cycle; for example, LD 15: 9 would indicate 24 h
cycle(s) i n which a 1 5 h ligh t span alternates wit h a 9 h dark span.

132 Plant

Growth and Development

light break Interruptio

n o f the dar k span or DD with a light span (o r pulse) .

LL Abbreviatio n fo r continuou s light . Preferably , the energ y levels and spectra l

characteristics should remai n constant .
masking Alteratio n o f rhyth m parameter s o r characteristic s b y externa l (e.g. ,
environmental) conditions . Maskin g may be responsibl e fo r causing the amplitude
of a circadian rhythm to increase , decrease , o r b e unexpressed ; o r i t ma y change a
sinusoidal curve to a non sinusoida l on e o r visa versa.
MESOR A n acrony m (midline estimating statistic of rhythm) use d i n cosinor
analysis t o indicat e th e mea n of the mode l fitte d t o th e data.
oscillator Se

e biological clock.

pendulum timer A n oscillating mechanis m capable of timing rhythms and likene d

to th e pendulu m of a mechanical cloc k (b y contrast t o a n hour-glass timer).
period Th e tim e require d t o complet e on e cycle . Biological rhythms ca n b e
classified accordin g t o thei r free-runnin g period s (e.g. , ultradian, circadian, and
infradian).
phase Measur e o f timin g o f an y instantaneou s stat e withi n a cycl e (e.g. , peak ,
trough) o f a rhythmic variable versus a reference (either interna l or external) .
phase angle difference Angula r difference between the phase of one cycle and that
of another.
phase response curve A graphic representation o f the extent of phase shifts cause d
by treatments (perturbations ) o f short duration relative to one cycle and positione d
at differen t part s o f th e cycl e (a pulse experiment). Phas e shift s ar e ofte n plotte d
against the rhyth m stage (e.g . circadian time) when the treatmen t wa s administered.
phase shift A change in th e phase o f one cycle relative to tha t o f the origina l o r
previous cycle . Sometime s referred to as rephasing. I f the phase is advanced in time
relative t o the reference cycle , the phase shift i s positive (+) an d involves the earlier
occurrence o f events within a cycle; if the phase is delayed, the phase shift i s negative
(-) an d involve s th e late r occurrenc e o f events within a cycle.
photoperiod Lengt h o f the ligh t span in a 24 h LD cycle . Commonl y used in th e
literature, althoug h light span (or photofraction) woul d b e preferabl e becaus e
photoperiod ha s been used to refer to the light span (as defined here ) or to the whole
daily cycle (comprisin g bot h th e ligh t an d dark spans).
rhythm splitting Th e subdivisio n of rhythmic processes int o two or mor e groups
with similar or different periods , sometimes observed in organisms under free-runnin g
conditions.
self-sustained oscillations Se

e endogenous rhythm.

semidian rhythm Biological rhythm wit h a perio d o f abou t 1 2 h . (Th e term ,

circasemidian, als o appear s in th e literatur e to indicat e a period o f 1 2 2 hours.)

Biological Timing 13

sinusoidal rhythm A rhyth m that, when th e observe d measurement s ar e plotte d

as a functio n of time, exhibit s a curve that approximate s a sine curve.
skeleton photoperiod A synchronizing light:dark cycle with short ligh t spans marking the beginnin g an d th e en d o f the usua l light span o f a 24 h cycle. Fo r example ,
the skeleto n photoperiod o f a usua l L D 10:1 4 cycl e coul d includ e th e followin g
sequence: 3 0 min ligh t span , 9 h dar k span, 30 min light span, and 1 4 h dark span .
subjective day Unde r constan t condition s (e.g. , L L or DD) , i t i s the spa n corre sponding t o th e ligh t spa n o f a previously used 2 4 h LD environmenta l cycle .
subjective night Unde r constan t condition s (e.g. , L L or DD), it is the span corre sponding t o th e dar k spa n o f a previously use d 2 4 h LD environmenta l cycle.
synchronizer A n environmenta l signa l tha t ca n rephase (o r entrain) a rhythm .
Also called th e Zeitgeber. Synchronizer ma y be preferred since it does not convey th e
impression o f "givin g th e time " (a s Zeitgebe r does) , bu t merel y t o se t th e phas e
and/or period of an endogenous rhythm.
transient cycle Cycl e o f an abnorma l lengt h occurrin g immediatel y after a phase
shift (o r stimulus) for synchronization t o a new cycle. Mor e tha n one transien t cycle
may be observe d before th e usua l period i s reestablished .
ultradian rhythm Biological rhythm havin g a perio d appreciabl y les s tha n 2 4 h
(usually less than 20 h). Ultradia n oscillation s wit h periods les s than 30 minutes ar e
called high frequency oscillations.
Zeitgeber Se

e synchronizer.

CONSULTANTS
Ruth Satte r (deceased ) Bernar
University o f Connecticu t Universite
Storrs, Connecticu t Besango
Germaine Cornelisse n Guillaum e Fran
University o f Minnesota Uta
Minneapolis, Minnesot a Logan
Ola M . Heide Theres
Agricultural Universit y o f Norway Universit
AS-NLH, Norwa y Bruxelles

d Millet
d e Franche-Comt e
n Cedex , Franc e
k B . Salisbury
h Stat e Universit y
, Uta h
e Vanden Driessch e
e Libr e d e Bruxelle s
, Belgium

17
DORMANCY, PHOTOPERIODISM, AND VERNALIZATION
Frank B . Salisbury
Department o f Plants, Soils , an d Biometeorolog y
Utah Stat e Universit y
Logan, U T 84322-482 0 U.S.A .
This chapter deals with terms used in the study of vernalization, photoperiodism, an d
dormancy. A few terms ar e defined within other definitions ; thes e ar e also printe d
in boldfaced type. Word s in italics are themselves defined elsewhere althoug h italic s
may also be use d fo r scientific names .
abortion Arres t o f developmen t o f a structure . I n som e species , especiall y i n
bulbous plants , the terms flower abscission and blasting are frequently used for flower
abortion. Abscission is the abortion o f a structure that shrivels, dries up, and rapidly
sheds; blastin g i s the abortion o f a structure tha t shrivels, dries up, but usuall y does
not shed . I n certain studies , especially with roses, blindnes s i s used for early abortion o f the flower .
absolute response Se e qualitative response.
after-ripening Use d b y some author s with reference to an y change tha t goe s o n
within a dormant see d o r bud during the breaking of dormancy. Othe r authors have
used the term in a more restricted sense , limiting it to maturation changes that occur
in th e embry o durin g storage. Th e firs t us e is preferable .
allelopathic substances Organi c chemicals that are produced by one plant and that
harm anothe r plant , sometimes by inhibiting germination.
ambiphotoperiodic plants Plant s tha t respon d (e.g. , flower ) onl y whe n give n
photoperiods tha t ar e shorte r tha n som e daylengt h o r longe r tha n som e longe r
daylength (e.g. , shorte r tha n 1 4 h or longe r than 1 8 h); opposite t o intermediate-day
plants. Thi s respons e i s rare, but i t was reported i n Madia elegans (Lewi s and Went ,
1945) an d i n Setaria verticiltata (Matho n an d Stroun , 1960) .
annual A plant wit h a life cycl e fro m see d t o see d tha t is completed i n only one
growing season (o r on e year).
anthesin Se e florigen , whic h Chailakhya n (1968 ) suggest s i s a combinatio n o f
gibberellin an d anthesin .
134

Biological Timing 13

anthesis Th e tim e o f coming into ful l bloo m (e.g. , i n grasses, th e tim e when th e
anthers ar e extende d fro m th e flowe r an d pollen i s released) .
antiflorigen A

transmissibl e stimulu s that maintain s the vegetative state.

apical meristem (apex) A

meristem borne at th e ti p of a vegetative plan t stem .

autonomously-inductive plant (self-inductive) Flowerin g occurs more-or-less independently of day length (a s in day-neutral plants) and more-or-less independentl y of
any othe r specia l environmenta l treatment . Tha t is , the respons e occur s unde r a
variety of constant environmental conditions .
axillary meristem Meristems i n the angle (axil) formed by the leaf petiole and th e
stem; potentiall y capabl e of forming a branch.
biennial A plan t tha t live s two growing seasons an d flowers and die s durin g the
second season . Typically , biennial s gro w vegetatively during the firs t season , ar e
induced t o flower by the lo w winter temperatur e experience d betwee n th e seasons ,
and flowe r an d di e th e secon d season ; tha t is , they hav e a n absolut e vernalization
requirement. Ofte n thei r flowerin g i s als o promote d b y o r require s lon g days .
(Many wild biennials ma y sometimes be to o smal l after th e firs t seaso n t o becom e
vernalized, i n which case the y might live for mor e than two seasons, althoug h they
flower only once before dying.)
bolting Rapi d elongatio n o f a flowering stem fro m a vegetative rosette, often i n
response t o vernalization o r lon g days.
caulescent plant A
to a rosette plant.

plant with leaves distributed along an elongated stem; opposite

critical daylength (critical day, critical photoperiod) I n plant s with an absolut e

daylength requirement , th e daylengt h or photoperio d tha t mus t b e exceede d t o
initiate long-day responses (e.g., flowering of long-day plants) or to inhibit short-day
responses. (Som e author s hav e define d critica l daylengt h as th e daylengt h tha t
produces th e smallest detectable response , or even 50% flowering, but those defini tions shoul d not be used.)
critical nightlength I n plants with an absolute daylength requirement, the night length o r darkperio d tha t mus t b e exceede d t o initiat e short-da y responses (e.g. ,
flowering of short-da y plants o r formatio n of potato tubers ) o r t o inhibi t long-day
responses. (Som e author s hav e defined critical nightlength as the nightlength that
produces th e smalles t detectabl e response , o r eve n 50 % flowering , bu t thos e
definitions shoul d no t b e used.)
daylength (or day length) Sometime s (a s in these definitions) written as one word
in th e sens e o f the photoperio d i n a natural 24-hour cycle of light and darkness , as
this might influence plant growth or development. (Otherwise , in English the term
is correctly written a s tw o words: da y length.)
day-neutral plants (DNP ) Plant s that do not require a specific daylength treatment
for flower initiation or othe r photoperiodicall y controlled response.

136 Plant

Growth and Development

determinate Wit h reference t o an organ such as a leaf, flower, or fruit tha t grows
to a certain size and then stops growing; stems and roots, because the y are produce d
by apical meristems, may continue t o gro w indefinitely, and are thu s indeterminate.
developmental arrest A limitatio n o n see d developmen t tha t prevent s a viabl e
embryo from germinatin g during growth of the seed. (Th e term could be applied t o
other structures a s well.)
devernalization Reversa l of the promotion of flowering induced by exposure to low
temperatures (i.e. , by vernalization) b y an immediate exposure to hig h temperature s
(e.g., 30 C). I f a period o f time elapses a t neutral temperatures between vernalization an d th e high-temperatur e treatment , devernalizatio n usuall y fails. (I n som e
perennial plants, Chrysanthemum, fo r example, prolonged exposure to low irradiance
or shor t days also reverse s th e effect s o f vernalization. )
donor I n graftin g experiments , th e graf t partne r tha t i s assumed t o provid e th e
stimulus (promotiv e o r inhibitory ) to th e receptor.
dormancy Th e condition o f a seed or other plant organ when it fails to germinat e
or gro w because i t ha s not bee n provide d with some special set of conditions (e.g. ,
a perio d o f lo w temperature , suitabl e wavelength s o f light , a treatmen t tha t wil l
scarify th e seedcoa t o r leac h ou t inhibitors ) althoug h i t ha s bee n provide d wit h
moisture, oxygen , and temperature conditions that are suitable for germination and
growth after th e specia l requirements have been met. On e special condition can be
sufficient tim e for the embry o to mature.
Dormancy a s define d her e ha s bee n calle d endogenous o r innate dormancy or
endodormancy (Lang et al., 1986) a s contrasted to imposed dormancy, which prevails
if a n essentia l facto r (e.g. , H 2O o r O 2) i s lacking . Impose d dormanc y i s calle d
quiescence here. Seed-coat-imposed dormancy and embryo dormancy have also bee n
distinguished.
Pomologists hav e use d rest i n th e sens e o f dormancy a s define d her e (Samish ,
1954). Becaus e this use is rather specialized , it would be well to avoid the term rest.
evocation Earl y responses o f receptor tissu e following environmental triggering or
other form o f induction; usually related t o the flowering responses tha t occur at th e
shoot ape x afte r arriva l of flower stimuli and prio r t o flower differentiation (floral
initiation); defined by Evans (1969) to distinguish from induction, which occurs in the
leaf.
facultative response Se

e quantitative response.

florigen (floral stimulus) A postulate d flowering hormone or chemica l stimulus

believed t o aris e i n th e leave s o f certai n plant s i n respons e t o a n appropriat e
environmental treatmen t (suc h a s lon g o r shor t days ) and i s translocate d vi a th e
phloem t o th e bu d apices wher e it causes evocation and flower initiation. Florige n
may als o aris e autonomousl y i n th e leave s o f day-neutral plants o r b e transmitte d
from a n induce d t o a noninduce d plant through a graf t union .
fractional induction Induction cause d by one or more cycles of inducing conditions
(e.g., shor t days ) interspersed wit h one or mor e cycle s of noninducin g conditions
(e.g., long days).

Dormancy, Photoperiodism, an d Vernalization 13

germination Th e su m o f th e physica l and chemica l processe s within a see d tha t

lead t o visible penetratio n o f the see d coa t b y the radicle (embryoni c root).
grafting Th e process of combining two separate plants or plant parts (as two stems
or a bud an d a stem) with the intention tha t they will unite and grow. Th e process
may be carrie d ou t i n man y ways. Se e stock, scion, donor, receptor.
hard seed Seed s that ar e kep t dorman t by hard, impermeable seed coats ; ca n be
induced t o germinate by physical, chemical, or microbial abrasion/decomposition of
the seed coat.
impaction A treatment in which seeds are vigorously shaken to dislodge a cork-like
filling (th e strophiolar plug) i n th e strophiolar cleft o f th e seedcoat , allowin g
penetration o f water an d oxyge n and leading to germination.
indeterminate Th e conditio n o f an apical meriste m of the shoot o r root that ha s
the potentia l t o gro w indefinitely . Vegetativ e meristems are indeterminate . Con trasted to a determinate meristem, which produces a structure such as a leaf or flowe r
and the n cease s to exis t as a meristem .
induced state Conditio

n o f a plant that ha s been induced.

induction A phenomenon i n which some response (e.g., flowering) can be caused

(induced) in an organis m by some treatment (e.g., an environmental condition such
as cold or short days), and the response continues , or typically first appears, after th e
treatment ha s been discontinued . Sometimes , inductio n appears to occu r autonomously i n th e absenc e o f an y obviou s treatmen t (i.e. , unde r a variet y of constan t
environmental conditions) . Inductio n precedes evocation an d initiation.
inhibitor Substanc e preventing germination, growth, flowering, or other responses ;
that is , it need s t o be leached out , oxidized, or otherwis e metabolize d (e.g. , broken
down or bound) to permit th e response. (Ca n also be applied externally to inhibit.)
initiation, flower Th e beginnin g o f flora l differentiatio n (i.e., a morphologica l
change) tha t follow s induction an d evocation.
intermediate-day plant (IDP) A plant that fails to respond photoperiodically when
days are eithe r to o shor t o r to o long; usually with referenc e to flower formation.
juvenility Stat e o f a usuall y youn g plan t tha t i s incapabl e o f flowerin g unde r
otherwise suitabl e conditions ; ofte n associate d wit h a variet y o f morphologica l
features. (Se e ripeness-to-flower. ) Som e matur e wood y plants (i.e. , trees ) retai n
juvenile feature s i n thei r lowe r parts , a phenomeno n sometime s calle d secondary
juvenility. Terminatio n o f juvenility (achieving maturity) ma y differ fro m achieving
ripeness-to-flower i n tha t plant s tha t reac h maturit y may flower (and/or lose thei r
juvenile morphological features ) whether conditions change or not, whereas a plant
that ha s reached ripeness-to - flower typically requires some special treatmen t (e.g. ,
long days, short days , low temperatures) t o actuall y flower.
long-day plant (LDP) A plan t that flowers or otherwis e responds when th e day s
are longer than some minimum length (depending on the species) and the nights are
shorter tha n some maximu m length; opposite in response to short-day plant. Long day plants typically respond best t o continuou s light.

138 Plant

Growth and Development

long-short-day plant (LSDP) A plan t tha t require s a sequenc e o f long day s

followed b y short day s for som e photoperiodi c response , usuall y flowering .
maturity Th

e stag e reache d b y a plant when juvenility i s terminated.

meristem A tissue or group of tissues capable of cell division, enlargement (ofte n

elongation), and differentiation t o produce determinate or indeterminate structures of
the root/shoo t system . Apical o r axillary meristems of angiosperm s ma y becom e
flowers. Vegetativ e meristem s ar e indeterminate, capable i n principl e o f growing
indefinitely t o produc e stem s and roots .
minimum leaf number Th e minimu m number o f leave s tha t mus t be produce d
before flora l initiation occur s (in plants that produce the first flowe r a t the terminal
bud) o r tha t mus t b e produce d befor e a plan t achieve s th e conditio n know n as
ripeness-to-flower (o r ripeness-to-respond), i n which conditio n i t ca n become induced
in response t o a photoperiodic o r other environmenta l treatment.
monocarpic species Plant s tha t flowe r an d produc e seed onl y once an d the n die .
They ma y be annuals, biennials, o r perennials. Contraste d wit h pofycarpic species.
night interruption (night break or light break) Wit h reference t o inhibitio n o r
promotion o f some photoperiodic respons e b y interrupting the dar k period wit h an
interval o f light . Dependin g upo n species , th e ligh t brea k wil l b e mor e o r les s
effective dependin g upo n whe n i t i s give n an d upo n th e irradianc e leve l an d th e
spectral quality . Long-day plants typicall y requir e a muc h longe r perio d o f
interruption at a higher irradiance leve l tha n do short-day plants. I f the ligh t brea k
is effective , i t act s a s a long-da y treatmen t (promotin g a long-da y response o r
inhibiting a short-day response). Respons e to a night interruption is an excellent test
for a true photoperiodi c response . (Se e photoperiodism.)
nightlength (night length) Sometime s written as one word in the special sens e of
the dar k perio d i n a natura l 24-hou r cycl e o f ligh t an d darkness , a s thi s migh t
influence plant growth or development. (Otherwise , in English the term is correctly
written a s two words: nigh t length. )
null-response technique A n approach used in studies on photoperiodic phenome na and on other photobiological response s i n which irradiance levels of two opposing
wavelengths (i.e. , red and far-red i n most applications) ar e balanced to produce th e
same plan t respons e a s woul d b e produce d b y darknes s o r b y som e whit e ligh t
source.
perennials Plant s that live for an indeterminate number of growing seasons. Mos t
perennials arepofycarpic an d flower once each year when they are sufficiently mature,
but som e are monocarpic (flowe r onl y once afte r severa l years and then die) .
phase change I n th e contex t o f juvenility, th e transition fro m th e juvenile to th e
mature condition, permittin g sexual reproduction.
photoperiodism Th e respons e o f organisms to th e relativ e length s of day and/or
night. Mos t response s involv e changes in growth and development (e.g., flowering,
tuber formation , and dormancy) . (Adjectiv e form i s photoperiodic.)

Dormancy, Photoperiodism, an d Vernalization 13

phytochrome Plan t pigment consisting of two interconvertible form s absorbing in

the re d (c.a . 660-nm ) o r far-re d (c.a . 730-nm ) region s o f light . Phytochrom e i s
directly involve d i n man y photomorphogeni c reaction s (e.g. , photoperiodism ,
dormancy breaking, deetiolation , chlorophyll formation, etc.). (Se e Chapte r 9. )
polycarpic species Plant s tha t flowe r fo r mor e tha n on e seaso n (ofte n man y
seasons); al l polycarpi c species are perennials. Contraste d with monocarpic species.
prechilling (stratification) Th e treatmen t i n whic h imbibe d (moist ) seed s o r
dormant plant s ar e subjecte d t o col d temperature s (usuall y a fe w degrees abov e
freezing an d preferabl y fluctuating ) fo r som e interva l o f tim e wit h th e goa l o f
breaking dormancy an d promoting active growth; not t o b e confused wit h vernalization, which i s a promotion o f reproductive growt h by cold treatment .
precocious Ter m applied to switching of developmental pathway s prior to natural
maturity, a s in flowerin g induce d chemically or germination o f immature seed.
qualitative response (absolute response, obligatory response) A plant response,
usually flowering, that absolutely depends on some daylength, temperature, or othe r
environmental stimulus . I f the plant does not experienc e the required stimulus , the
response does not occur (e.g., the plant remains vegetative). Opposit e to quantitative
response.
quantitative response (facultative response) A plant response tha t is changed in
number o r developmenta l rat e (e.g. , more flowers or tubers ) by a treatment suc h as
a particular da y length o r an exposure to a period of low temperatures. Opposit e t o
a qualitative or absolute response. I n the absence of the treatment, the response stil l
occurs bu t a t a much slower rat e or fewe r organ s are produced.
quiescence Applie d t o a viable, nondormant seed or bud that fails t o germinate or
grow only because i t ha s not bee n provide d with suitable temperature, oxygen, and
moisture conditions. Ha s also been called imposed dormancy or ecodormancy (Lang
et al. , 1986) . Se e dormancy.
recalcitrant seeds Seed s that have a very limited storage period that usually cannot
be extended b y dry and cold conditions . The y are generally large and frequently ar e
tropical species ; the y germinate at onc e afte r ripening .
receptor I n graftin g experiments , th e graf t partne r that is assumed to receiv e th e
stimulus (promotiv e o r inhibitory ) from th e donor.
rejuvenation Reversio n (usuall y onl y partial ) o f matur e wood y plant s t o a
secondary juvenile phase by such treatments as pruning, grafting, and treatment with
gibberellins.
rest Use d by pomologists (Samish , 1954) and others in the same sense as dormancy
as defined here. Rest has als o been use d in the sens e o f quiescence as defined here .
Hence, the ter m i s ambiguous and would best b e avoided.
revernalization Effectiv

e chillin g treatment of plants that have been devernalized.

ripeness-to-flower (Bluhreife, ripeness-to-respond) A little understood condition

of a plant that is reached a t a certain age when it is capable of becoming induced t o
flower in respons e t o a photoperiodic o r othe r (e.g. , temperature ) treatment . Th e

140 Plant

Growth and Development

response unde r discussio n i s usually that o f flowering . Se e juvenility, maturity, an d

phase change.
rosette plant A plant with leaves comin g from a greatly shortened ste m at ground
level; typica l o f man y biennial s durin g th e firs t yea r (e.g. , beets) ; opposit e t o a
caulescent plant.
scarification Breakin g o f th e seedcoa t barrie r b y mechanical treatmen t suc h a s
abrasion b y san d o r grave l o r microbia l action ; allow s penetratio n o f wate r an d
oxygen an d thu s allows germination .
scion Th

e detached plan t par t tha t is grafted t o a stock.

short-day plant (SDP) A plant that flowers or otherwise respond s when the days
are shorter than some maximu m length (dependin g on the species) and/o r when the
nights are longe r tha n some minimum length; opposite t o long-day plant.
short-long-day plant (SLDP) A plant tha t respond s photoperiodically (flowerin g
is usually the response ) to a sequence o f short day s followed by long days.
stock Th
grafted.

e roo t o r roote d plan t t o whic h a detache d plan t par t (th e scion) i s

stratification Se

e prechilling.

summer dormancy Dormanc y of buds established during long days of summer or

early autumn, usually in preparation fo r winter conditions.
thermoperiodism Growth , development, or behavioral responses o f organisms to
alternating da y and night temperatures. I n earlier researc h o n storage of bulbs, this
term referred t o the requiremen t fo r different temperature s applied durin g different
developmental stages . T o avoid confusion, this second usage should now be avoided.
vernalin A substanc e postulate d t o aris e i n respons e t o vernalization an d t o
promote reproductiv e growth . It s existence i s seriously doubted.
vernalization Th e induction o f flowerin g i n a plant , mois t seed , o r developin g
embryo o n th e mothe r plan t throug h exposur e t o lo w temperatures (usuall y a few
degrees abov e freezing) . I n man y (perhap s most? ) cases , vernalizatio n lead s t o
ripeness-to-flower, afte r which some other treatment is required to produce flowering.
(For example , th e biennia l strain of Hyoscyamus niger ha s an absolut e requiremen t
for lon g days following vernalization.)
viable Applie d t o th e conditio n o f a seed tha t i s alive and capable of germination
when provided with suitable environmenta l conditions; these conditions may include
treatments t o brea k dormancy.
winter annuals Plant s tha t germinat e i n lat e summe r or autumn , spending th e
winter a s seedlings , an d flowerin g an d fruitin g durin g th e nex t growin g season .
Flowering ma y be promote d b y exposure to th e lo w temperatures of winter.

Dormancy, Photoperiodism, and Vernalization 14

REFERENCES
Chailakhyan, Mikhai l Kb . 1968 . Interna l factor s o f plan t flowering . Annua l Review s o f Plan t
Physiology 19:1-36 .
Evans, Lloy d T . 1969 . Th e natur e o f flower induction . In : L.T . Evans , editor . Th e Inductio n of
Flowering, Th e Macmilla n Company o f Australia, Sout h Melbourne , p 457-480.
Lang, Greg , Rebecca Darnell, Jack Early, and George Martin. 1986 . Repl y to letter. HortScienc e
21(2):186.
Lewis, Harla n an d Frits W. Went. 1945 . Plan t growth under controlled conditions . IV . Respons e
of Californi a annuals t o photoperio d an d temperature . Amer . J . Bot . 32:1-12.
Mathon an d Stroun . 1960 . Thir d Internationa l Congres s o f Photobiology, Elsevier , Copenhagen .
p 384-386.
Samish, R.M . 1954 . Dormanc y i n woody plants . Ann . Rev . o f Plan t Physiol . 5:183-204 .
CONSULTANTS
These terms with preliminary definitions were published in the Flowering Newsletter, which was edited
and issue d b y Abraham H . Halev y (no w b y Georges Bernier) . Th e followin g scientists responde d
with comment s tha t strongl y influenced the fina l definitio n o f terms as presented here .
Suresh C . Bhargav a
Indian Agricultural Research Institut e
New Delhi, Indi a

Moshe Negbi
The Hebre w Universit y of Jerusale m
Rehovot, Israe l

Georges Bernie r
University o f Lieg e
Liege, Belgiu m

E. H. Robert s
University o f Readin g
Reading, Englan d

Charles F . Clelan d
U.S. Dept . of Agriculture
Washington, D.C .

Kenneth C . Sanderso n
Auburn University,
Auburn, Alabam a

Abraham H . Halev y
The Hebre w Universit y o f Jerusale m
Rehovot, Israe l

Max Saure
Diplom-Agraringenieur
DorfstraBe 1 7
Moisburg, German y

Wolfgang Haup t
Universitat Erlangen-Nurnber g
Erlangen, German y
Jean-Marie Kine t
University o f Lieg e
Li6ge, Belgium
Rodney W . King
CSIRO
Black Mountain , Canberra , ACT , Australi a
Donald T . Krize k
USDA/ARS
Beltsville, Marylan d
Wim d e Mun k
Bulb Research Cente r
Lisse, Netherland s
Klaus Napp-Zin n
Botanisches Institut der Universita t Kol n
(Cologne), German y

Walter W . Schwabe
University o f Londo n
Wye, England
Atsushi Takimoto
Kyoto University
Kyoto, Japa n
Kenneth Thimann
University of California
Santa Cruz , California
Daphne Vince-Prue
Goring-on-Thames
Reading, Englan d
Jan A . D. Zeevaar t
Michigan Stat e University
East Lansing , Michigan

18
STRESS PHYSIOLOGY
Leslie H. Fuchigam i
Department o f Horticulture
Oregon State University
Corvallis, Orego n 9733 1
Eugene V . Maas
U.S. Salinit y Laboratory, USD A ARS
Riverside, Californi a 9250 7
James M . Lyons
Department o f Vegetable Crop s
University of California, Davis Campus
Davis, California 95616
D. William Rains
Department o f Agronomy and Range Science
University of California, Davis Campus
Davis, California 95616
John K . Raison (deceased )
Plant Physisolog y Unit
CSIRO Divisio n o f Food Researc h & School o f Biological Science s
Macquarie University
North Ryde, 2113, N.S.W . Australia
Kenneth A . Shackel
Department o f Pomology
University of California , Davis Campus
Davis, Californi a 95616-8683
The fiel d o f stress physiolog y is not onl y of considerable theoretica l importance ; i t
is highly significant to agriculture. I n practice, researchers ten d to specialize within
at leas t fou r subfields : chillin g injury, col d stress , water stress, an d salinit y stress.
Yet, thes e subfield s hav e several basi c terms in common. Thus , this chapter begins
with the term s commo n to stud y of all plan t stresses an d is then divide d into fou r
sections representin g th e fou r subfields . Som e bold-fac e terms are als o define d in
the contex t o f othe r definitions ; words i n italics ar e define d elsewher e o r ar e
botanical names . Se e Chapter 6 for terms, units, and symbols used to describe plant
water relations (e.g., water potential and osmotic, matric, and pressure potentials, etc.),
142

Stress Physiology 14

1. GENERAL STRESS-PHYSIOLOGY TERMS

acclimation Adjustmen t of an organism to changes in external environments; thes e
anatomical o r physiological change s ar e beneficial and increase the organism's resis tance o r tolerance to subsequen t environmenta l stress .
adaptation Anatomica l o r physiologica l characteristic s o f a n organism , usuall y
genetically fixed , tha t enabl e i t t o liv e in a given environment.
avoidance A n acclimation or adaptation tha t reduces th e intensit y of stress at th e
cellular level .
calorie Th e unit of energy required t o raise the temperature of one gram of water
by one degre e centigrade . Equa l t o 4.1842 joules (exactly) , the preferred SI unit.
conditioning Exposur e t o temperature s slightl y abov e th e critical temperature
chilling rang e fo r various period s o r other treatments tha t ca n limit the magnitude
or affec t o r dela y th e onse t o f th e primary and/o r secondary events leadin g t o th e
development o f visible symptoms.
critical temperature Th e lowest ambien t temperature a t which the whole or part s
of a living organism ca n endure fo r 30 min without injury. Th e critical (threshold)
temperature ma y vary with the species, tissue , stage of growth, etc. Som e species ar e
damaged b y cool temperature s abov e th e freezin g poin t (chilling-sensitive); other s
only by subfreezing temperatures . (Critica l temperatur e i s not t o be confused with
the chemica l definitio n o f critical temperature , whic h is the temperatur e o f vaporization o f a liquid. )
dehardening Synonymou s with deacclimation. The loss in plant tissues and organs
of resistance to variou s stresses.
hardening Conditioning or acclimation of an organism to a particular stress, which
results i n increased resistance t o tha t stres s an d sometimes t o other stresses. Wit h
relation to low-temperature stresses, hardening is a term used to describe physiological event s tha t lea d t o a lowerin g o f th e critical temperature fo r letha l injur y o f
chilling-insensitive plant s expose d t o freezin g temperatures . Th e ter m hardening
should not be used in relation t o chilling; the more appropriate ter m is conditioning.
photo oxidation Oxidatio

n o f a substance cause d b y the absorptio n o f a photon .

primary event(s ) Th e primar y cellula r sensor(s ) o r trigger(s ) tha t initiate(s ) a

series of secondary events leading eventuall y to th e visibl e symptoms of such strains
as chitting injury. Th e primar y event(s) mus t occur a t th e critica l stress leve l (e.g. ,
critical temperature) fo r th e specie s o r tissue , mus t be rapid, and in th e shor t term ,
reversible.
regrowth Th e ultimate viability test to determine survival following a stress. Plant s
or cells are grow n or quiete d fo r a given period o f time following a stress and the n
evaluated fo r eithe r roo t and shoo t regeneratio n o r increas e i n mass.
repair Th e proces s b y which th e stress-induce d injur y o f a plan t i s partiall y or
completely reverse d followin g remova l of the stress .

144 Plant

Growth and Development

secondary event(s ) Th e metaboli c an d cellula r change s followin g (directl y o r

indirectly) as consequences o f the primary event and that lead to the visible symptoms
of stress-induced injury . Th e secondary events are both time- and stress-dependent .
In th e shor t ter m th e change s induce d ar e reversibl e i f th e stres s i s removed .
However, i f the stres s is maintained, th e tissu e become s unabl e to recover .
strain Th e observe d (deleterious ) biologica l change s tha t occu r i n respons e t o
stress. (Th e term s stress and strain ca n be used in a manner analogous t o thei r us e
in physics: stress is the forc e applied to an object, as a metal bar; strain is the actua l
change i n shape o f th e object , a s bending of the bar. )
stress An y environmental condition tha t is capable of causing a biologically injurious chang e (strain). Sinc e plant s ar e autotrophs , an y chang e tha t directl y o r
indirectly reduces plant growth (biomass accumulation) must be considered biolog ically injurious, even i f there ar e beneficial consequences fo r othe r aspect s of plan t
function. Complet e description s o f a give n stress (o r strain ) shoul d includ e it s
magnitude, duration, and rate of development.
symptoms of injury Visibl e manifestation s of th e secondary events tha t reflec t
injury t o th e cell s and tissue s cause d by chilling.
tolerance Abilit y o f a n organis m o r it s cell s o r othe r part s t o surviv e the ful l
impact o f a stress. Se e also avoidance.
viability Th e stat e of living , growing, or developing . Th e viability test estimate s
the relativ e o r absolut e surviva l o f a n organism. Fo r example , commo n cold hardiness viability tests includ e visual browning, conductivity, regrowth, vital stains
such a s 2,3,5-tripheny l tetrazoliu m chloride, florescei n diacetate, Evan' s blue , an d
neutral red .
2. CHILLING INJURY1
ameliorate T o provid e a treatment o r set of conditions tha t reduce the impac t of
a chilling treatment b y alterin g th e tim e cours e o f sympto m development . Thus ,
amelioration i s confined to describin g changes in the tolerance of the plant tissu e t o
the impose d chillin g stress.
chilling Th e ac t o f exposin g plan t materia l t o a non-freezin g low temperature.
This exposur e ma y or ma y not b e beneficial to th e plant .
chilling injury A descriptiv e ter m fo r th e physiologica l injury t o man y plants ,
particularly those warm-season species (e.g. , crops) of tropical or sub-tropical origin,
when the y ar e expose d t o low , but non-freezin g temperatures .
chilling-insensitive Thos e plants that typically continue to grow and develop, albeit
slowly, and ca n complete thei r lif e cycl e when continuously exposed to chilling tem peratures. Thes e plants ar e primaril y cool seaso n specie s o f temperate origin .

Original authors of this section were J.K. Raison an d J.M. Lyons.

Stress Physiology 14

r change s tha t correct th e advers e effect s o f th e secondary

chilling repair Cellula

events.

chilling reversal Reversa l o f the primary event (s), which would be rapid and direct
and shu t of f further stimulatio n o f the secondar y events.
chilling-sensitive plants Thos e plants that are injured by exposure to temperatures
below about 1 0 C to 1 5 C, but above freezing. (Th e warm-season crops of tropical
or sub-tropica l origin s hav e receive d th e mos t study. ) Al l stage s o f growt h an d
development o f the entire plan t (excep t perhap s the dry seed) ar e susceptible. Thi s
susceptibility limit s th e seaso n o f growth, geographic distribution, an d postharves t
storage conditions of these plants. (Harveste d plan t parts, especially fruit s o f some
temperate plants , notably apples, pears, cranberries, asparagus, and potatoes, exhibit
chilling damage durin g storage whe n exposed fo r extended periods a t temperature s
very close to freezing , i.e., aroun d 2 C to 3 C . However , thes e temperature s do
not limi t growt h or geographi c distributio n o f thes e specie s a s exhibited b y warmseason crops.)
chilling temperature An y temperatur e belo w th e critical temperature, bu t abov e
freezing, tha t cause s injury .
chilling tolerance Th e abilit y of chilling-sensitive plants o r plan t parts , t o endur e
the metabolic dysfunctio n and/or harmful consequences tha t result from exposur e to
chilling temperature s an d t o surviv e if th e abus e is not sustaine d beyond a certai n
lethal point . Chillin g toleranc e i s use d t o describ e thi s differentia l respons e t o a
chilling stress and should be confined to describing differences i n the time and course
of th e developmen t o f chilling injury symptoms .
chilling treatment Th e proces s o f exposur e t o a chilling temperature fo r a tim e
period sufficient to caus e injury .
conditioning Se

e definition i n Section 1 of this chapter .

critical temperature Se

e definition i n Section 1 of this chapter.

dysfunction A n impaire d functioning of plant tissues in response to chilling. Th e

impairment i s reversible i f the tissu e i s returned t o a nonchilling temperature afte r
a period of exposure. Howeve r thi s dysfunction become s irreversibl e after a longer
period of time a t th e chillin g temperature.
hardening Se

e definition i n Section 1 of this chapter.

intermittent warming Interruptio n of a chilling exposure with brief warm periods

before th e critica l tim e is exceeded an d injur y occurs . Chilling-sensitiv e tissue can
be kept fo r extende d period s a t chilling temperatures i f th e critica l tim e i s no t
exceeded befor e intermittent warming and sufficient tim e is spent at the warmer temperature fo r th e tissu e t o recove r o r repai r prio r t o returnin g t o th e chillin g
temperature.
primary event(s) Se
secondary event(s ) Se

e definition i n Sectio n 1 of this chapter.

e definition i n Sectio n 1 of this chapter.

146 Plant

Growth and Development

sensitivity to chilling Th e ter m use d t o distinguis h relativ e sensitivit y (amon g

chilling-sensitive plant species) o n th e basi s of their critical temperature an d critica l
time t o develo p chilling injury.
symptoms of injury Se

e definition in Sectio n 1 of this chapter .

3. COLD HARDINESS2
anaerobic stress A
stress imposed on an organism as a result of the absence of the
free oxyge n of air. Se e also ice encasement and flooding.
bacteria nucleation inhibitors Chemical
nucleation by ice-nucleation bacteria.
calorie Se

s othe r tha n bactericide s tha t inhibi t ic e

e definition i n Section 1 of this chapter .

chilling requirement Low-temperatur e requiremen t t o overcom e dormanc y i n

seeds and buds . (Se e dormanc y in Chapte r 17 ; also calle d endo-dormancy)
cold hardiness Wit h referenc e t o th e exten t tha t plants ca n survive temperature s
below freezing . Ca n b e quantitativel y expressed a s the critical temperature.
cold injury Injur y incurred by biological materia l due to temperatures below 0 C .
The ter m i s often use d interchangeabl y with winter injury, freezing injury an d frost
injury.
cold protection Method s of guarding against injury from temperatures below 0 C.
The ter m i s often use d interchangeabl y wit h frost protection and generally refers to
protection o f blossoms i n th e spring .
cold shock Impositio n o f a brief, non-freezing temperatur e resulting in a strain t o
the organism ; ma y or ma y not induc e furthe r acclimation .
convection Th e mass movement of heated liquid or gas. Whe n used in discussions
of col d hardiness , th e ter m generall y refers t o mas s movement of heated air .
critical temperature Se

e definition in Section 1 of this chapter.

deep supercooling Abilit y of organisms to supercool a t a temperature below that

of intracellular freezin g of water t o a s low as the homogeneou s nucleatio n poin t o f
pure water, approximately -40 C. Thi s mechanism to avoid freezing exists in tissue
such a s xylem ray parenchyma an d dorman t flower buds . Refe r to supercooling.
degree growth stage model (G S Model) Numerica l syste m fo r quantifyin g th e
annual physiological growt h stages of buds of temperate plants . Th e annual cycle is
divided int o 360-degree growth stages an d fiv e majo r poin t event s (0 G S and 360
GS = onse t sprin g growth ; 90 G S = maturit y induction poin t whe n plan t firs t
becomes responsiv e t o photoperiod ; 18 0 GS = vegetativ e maturity and the onse t
of dormancy (se e definitio n in Chapter 17) ; 270 GS = maximu m rest ; 315 GS end o f rest whe n chillin g requirement i s satisfied).

Original autho r o f thi s section was Leslie H . Fuchigam i

Stress Physiology 14
dehardening Se

e definition i n Section 1 of this chapter .

differential thermal analysis (DTA) Metho d o f determinin g th e exothermi c

temperature differenc e between a reference an d a sample, usually biological, bein g
frozen, o r th e differentia l endothermi c temperatur e o f a sample being thawed.
electrical conductivity Th e curren t tha t will flow fro m on e fac e o f a uni t cub e of
a given substance t o the opposite face when a unit potential difference is maintained
between thes e faces. A technique use d t o determin e electrolyte leakage (membran e
integrity) followin g a stress.
electrical impedance Th e application of alternating electrical current to pre-frozen
or froze n plant tissue s to predict damag e or measur e injury , respectively .
electrolyte leakage A technique use d t o determin e cel l o r tissu e viabilit y by estimating membran e integrity . Followin g a stress, the tissu e i s shake n i n a give n
quantity of water fo r a predetermined period , and the initia l electrical conductivity o f
the effusat e i s determined. Th e tissue is then killed, either in liquid nitroge n or by
heat, shaken fo r a given time, and the fina l conductivity determined. Viabilit y or th e
extent of cellular o r tissue damage is estimated by the percentage loss of electrolytes .
electrometric method (and cell freezing) Measurin g the electrolyt e leakag e fro m
tissue caused b y membrane damag e from freezing . Variou s electrometri c method s
are used ; refer s t o electrical conductivity, electrical impedance, conductivity, an d
electrolyte leakage.
endotherms Hea

t consume d durin g a thawing event.

evaporative cooling Coolin g cause d by the vaporizatio n of a liquid. I n th e even t

of cold hardiness , i t would be the reductio n in temperature of biological matte r du e
to th e loss of water t o th e atmosphere .
exotherm Hea

t los t durin g a freezing event.

extracellular freezing Th

e crystallizatio n of apoplastic water.

extraorgan freezing Mechanis m o f freezin g tolerance i n plan t organ s b y wate r

translocation fro m supercoole d tissue s o r organ s t o nucleatio n center s i n adjacent
tissues (extratissue freezing) or outsid e the organ s (extraorgan freezing).
flooding Se

e definitio n in Section 4 of this chapter.

freeze avoidance Lac k o f water crystallization i n tissues at subzer o temperature s

caused by the absence o f either intrinsi c or extrinsic nucleators. Th e tissue supercools
and thereb y escape s injur y cause d b y ice formation.
freeze dehydration Los s o f symplastic water caused b y the vapor-pressur e defici t
created b y ice in the extracellula r spaces. Th e degree of dehydration is a function of
temperature; a s the temperature decreases , freeze-dehydratio n increase s resultin g in
a concentratio n o f the cel l contents .
freeze desiccation Se
freeze tolerance Th
apoplastic spaces .

e freez e dehydration.
e ability of a cell or tissue to tolerate the presence of ice in th e

148 Plant

Growth and Development

freezing injury Injur

y o f biological materia l cause d b y sub-zero temperature .

freezing point Th e maximu m temperatur e attaine d b y th e releas e o f hea t

(exotherm) followin g the initiatio n o f ice crystal formation. Th e temperature o f ice
crystal initiation ma y vary, depending o n th e presence o f nucleating agents , but th e
freezing-point temperatur e i s constant fo r an y given solution .
freezing-point depression Th e lowerin g o f th e freezing-poin t temperatur e o f a
solution b y the presenc e o f osmotically activ e compounds .
frost A deposit of one o f several forms of ice crystals as a result o f sublimation of
water vapor o n the eart h o r earth-borne object s colde r tha n 0 C :
Advection Frost: Occur s fro m th e movemen t o f larg e col d ai r masse s int o a n
area fo r several day s resulting in severe low temperatures an d often accompanied by
strong winds.
Blackfrost: A dry freeze that occurs when the dewpoint is low, preventing watervapor crystallizatio n o n object s an d resultin g in the interna l freezing o f vegetation.
Hoarfrost or White Frost: A deposit of interlocking ice crystals formed by direct
sublimation o n objects .
Radiation Frost: Occur s o n calm , clea r night s whe n ther e i s unimpede d
radiation fro m th e earth resultin g in strong temperature inversions . Usuall y occurs
in th e earl y mornin g hours an d i s characterize d b y relatively mild subfreezin g air
temperatures.
frost hardening Se e acclimation. Thi s proces s ma y be divided into thre e phases ,
based o n th e environmenta l stimulu s and the typ e of changes occurring .
1st Stage: Triggere d b y shor t days , whic h stimulat e th e productio n o f a
translocatable hardiness-promoting factor, predominantl y an active metabolic process.
2nd Stage: Triggere d b y low temperature, often a mil d frost . Bot h metaboli c
and physica l changes ar e involved .
3rd Stage: Foun d i n hardy woody species tha t have been expose d to prolonge d
freezing temperatures , resultin g i n physical alterations .
frost heaving Partia l o r complet e upliftin g o f a surfac e cause d b y ice expansion
and resulting i n exposure an d injur y t o plants .
frost plasmolysis Contractio n o f the dea d protoplast fro m th e cel l wall following
a lethal stress. Primarily due to the inability of the cell to reabsorb and maintain turgor followin g a lethal stress. (See freeze dehydration.)
frost protection Method s of guarding against injury from temperature below 0 C.
Generally refer s to protectio n o f blossoms in th e spring.
hardiness promoter(s) Naturall y occurring substance(s) synthesized in plants that
induce(s) freezin g resistance. Se e also: translocatable hardiness promoter(s) .
heterogenous ice nucleation Nucleatio n tha t i s catalyze d b y a soli d o r liqui d
substrate tha t allow s groups of adsorbed water molecules t o assum e configurations
that ar e able t o promot e furthe r condensation . Thes e nucleator s enhanc e th e
probability that a cluster of water molecules of critical dimensions can form , which
results i n crystallization.

Stress Physiology 14

hexagonal (H11) phase Th e H1l phas e is basically a liquid crystal in a hydrocarbon

matrix penetrated b y hexagonally packed aqueou s channel s towar d which the pol e
groups of the lipid are oriented. Lipid s in the H1l phas e provide a permeability barrier between interna l an d external environments.
homeohydric plant (or homoiohydric) Se

e Section 4 of this chapter.

homogenous nucleation temperature Th e temperatur e a t which a water nucleu s

forms tha t can be recognized by other water molecules as a structure resembling ice.
Such cluster s o f molecule s aris e spontaneousl y b y rando m fluctuations . Th e
probability tha t suc h a cluste r ca n serv e a s a n effectiv e nucleu s fo r crystallization
depends o n it s siz e an d it s lifetime , which are a functio n o f temperature . Wate r
spontaneously nucleate s a t -38. 1 C . Th e approximat e homogenou s nucleatio n
temperature, T n, fo r typica l plant solutions is given by the equation : A Tn = -( 2 A
Tm + 38.1 C) , where A Tm i s the melting point depression for the solution i n C .
ice deletion mutants Mutant s of ice-nucleating bacteria that do not ac t as ice nucleators. (Se e ic e nucleation active bacteria.)

ice encasement Th e partia l or complete coverin g of an organism by ice resulting

in anaerobic stress.
ice nucleation active bacteria (INA) Bacteri a capabl e of causing ice formation at
sub-zero temperature s clos e to 0 C .
intercellular freezing Se

e extracellular freezing .

intrinsic ice nucleator Nucleator s o f ice formin g within plan t tissue s an d organ s
resulting in crystallization a t relativel y warm subzer o temperatures.

inversion layer A meteorologica l phenomeno n i n whic h temperatur e rise s wit h

increased elevation instea d o f falling (o r decreases les s than the adiabatic lapse rate
would predict) . Ai r doe s no t ris e by convection throug h an inversion layer .
killing temperature Th e temperatur e a t whic h an organis m cannot recove r fro m
the stress as measured b y a viability testoften estimate d with the LT 50.
LT50 Th e temperature a t which 50 % of the population survives an artificial temperature treatment .
lamellar phase Orientatio n o f th e phospholipid s wit h thei r polar-hea d group s
exposed t o a n aqueou s environmen t an d th e acy l sid e chai n oriente d towar d th e
center o f the phospholipi d bilayer.
low temperature exotherm (LTE) Exotherm o r exotherms tha t appea r a t a lower
temperature tha n the large exotherm that represents freezin g o f extracellular water.
The low temperature exotherms represent a small percentage of the total water. Lo w
temperature exother m (LTE) pe r s e refers t o th e exotherm that occurs at approxi mately -37 C t o -4 0 C, the nucleatin g temperature o f homogeneous water.
low temperature injury Ambiguou s term used for injur y occurrin g from temper atures above freezing (chilling injury) t o temperatures as low as -196 C . A better
term migh t be freezing injury, th e temperatur e at which th e organis m no longer has
tolerance.

150 Plant

Growth and Development

melting point depression Th e decrease in the temperature neede d t o melt a solid

(e.g., ice) du e to th e additio n o f solutes .
membrane fluidity A property of membranes determined by fatty-acid chain length,
saturation level , an d sterol components .
membrane permeability Th e degree t o which a membrane will allow a solvent o r
solute to penetrate.
minimum survival temperature Temperatur e abov e which cells eithe r tolerat e or
avoid freezin g and belo w which they are killed .
phase transition temperature Lipid s in membranes exist in one of several possibl e
ordered structure s in which the individual lipid molecules have more or less motional
freedom. I n a functional membrane, the lipids need to exist in a fluid state to allow
rotational freedo m an d t o b e abl e t o diffus e i n th e plan e o f th e membrane . Th e
temperature a t whic h th e lipid s adop t a hexagonall y parked structure , thu s losin g
their diffusiona l freedoms , an d ente r a ge l phase i s known as the phas e transitio n
temperature.
photo oxidation Se e definition in Section 1 of this chapter .
plasmolysis Th e withdrawal of water fro m a plant cel l causin g the protoplas t t o
contract awa y from th e cel l wall (whic h may shrink elastically i f the cel l ha d bee n
under turgor) .
poikilohydric plant Se e definition i n Section 4 of this chapter .
poikilotherm A n organism whose temperature varies with that of its environment.
radiation frost Hea t radiate d fro m surfac e o f soil , trees , an d an y othe r soli d
particles t o reduc e th e temperatur e lo w enough t o caus e fros t conditions . Soli d
particles los e hea t mor e rapidl y tha n air , reducin g th e surfac e temperature .
Eventually th e ai r at lo w levels become s coo l an d heav y compared t o ai r a t highe r
elevations. Occur s mor e readil y o n cloudles s night s whe n ther e i s no barrie r fo r
retaining th e hea t (i.e. , a barrier tha t radiate s th e heat back to earth) .
regrowth Se e definition in Section 1 of this chapter .
repair Se e definition i n Section 1 of this chapter .
snow mold A diseas e o f cereal s cause d b y th e fungu s Calonectria graminicola.
Characterized b y abundant white mycelium and found beneath prolonged snow cover.
A similar disease cause d b y the genera Typhula, Sclerotium, or Fusarium, which are
particularly prevalen t i n turf-grasses.
sunscald A winter injury phenomenon i n which an otherwise hard y woody plant
is partially thawe d o n its sunward side; when the sun disappears, the thawe d tissues
experience rapi d intracellula r freezin g leading t o injur y an d death .
supercooling Als o called undercooling or subcooling. Coolin g of a substance (i.e. ,
water) belo w th e temperatur e a t which a change of state (i.e. , liqui d to ice ) woul d
ordinarily tak e plac e withou t suc h a chang e o f stat e occurring . Coolin g o f water
below it s freezin g point withou t freezin g takin g place; thi s result s in a metastabl e
state. (Se e als o homogenous nucleation temperature an d deep supercooling.)

Stress Physiology 15

T50 or TK50 Se e LT50.

thawing Th e process of melting of ice formed within or surrounding the organism .
Ice formed within the organis m ma y be extracellular or intracellula r in nature .
theory of minimum artical cell volume Merryma n proposed tha t cells canno t re cover from a greater volume decreas e than 40 % to 50 % of the unfrozen volume .
thermoavoidance A n organism that escapes high temperature (e.g., by transpiration
and coolin g o f leaves, movemen t o f leaves in response t o hig h temperature).
thermophile A n organis m that thrive s at hig h temperature.
thermophilic Havin g th e abilit y to surviv e or eve n thriv e at temperature s abov e
those considered t o b e lethal fo r most organisms.
thermophily Th e tolerance o f certain organism s to temperature s between 3 0 C
and 10 0 C .
thermostability Th e ability of proteins, enzymes, membranes, etc. to maintain their
integrity with increasing temperature .
thermotolerance Th e ability of an organism or its cells or other part s (tissues and
organs) t o surviv e the ful l impac t of a temperature stress (i.e. , se e freeze tolerance).
Tmax Temperatur
Tmin Th

e a t whic h 100 % of the plant s are killed .

e minimu m temperature tha t results i n injury .

translocatable hardiness promoter(s) Sam e as hardiness promoter(s). Substance(s )

produced i n respons e t o a short photoperio d tha t i s perceived b y phytochrome in
leaves an d translocate d i n the phloe m t o other plan t parts.
viability Se

e definition i n Section 1 of this chapter .

visual browning A method o f determining injury t o plant tissues by observing th e

presence o r absenc e of oxidative phenolic browning.
winter chilling Temperature s adequat e fo r th e chilling requirement t o brea k
dormancy. (Se e Chapte r 17. )
4. WATER STRESS3
Despite the larg e body of knowledge concerning the role o f water in the physiology
of plants, there is a lack of unanimity among plant physiologists regarding the nature
and physiologica l basi s o f plan t response s t o water-limite d conditions . Fo r thi s
reason, the definitions recommended below are mainly descriptive and not intende d
to imply specific underlying physiological mechanisms for, or attribute adaptive value
to, th e term s defined . Th e basi c approac h t o stress/strai n follow s tha t o f Levit t
(1980).
antitranspirant A
to plants .

substance that reduces transpirational water loss when applied

Original autho r o f thi s section was Kenneth A. Shackel .

152 Plant

Growth and Development

bound water Th e quantit y of water i n a tissue tha t i s strongly held b y adsorptiv e

(matric) force s an d i s difficult t o remov e b y tissue drying . Sinc e thi s quantit y will
depend o n the drying conditions, th e utility of a distinction betwee n boun d an d free
water i s questionable; hence , thi s term i s not recommended .
cavitation Spontaneou s occurrence of a vapor phas e i n a liqui d unde r tension .
Thought t o occu r i n the xyle m of plants under moderat e t o sever e water stress an d
to resul t i n a reduce d capacit y of the xylem t o conduct water.
critical period A perio d durin g crop developmen t i n which water stress will have
a mor e profoun d effect o n harvestabl e yiel d than th e sam e stress a t othe r periods .
Can be associated wit h stress-related reduction s i n early crop growth, which will be
compounded ove r time , or, for crops grown for their reproductiv e structures , direc t
stress-related effect s o n reproductiv e development .
dehydration (desiccation) tolerance Abilit y t o continu e a plan t functio n (i.e. ,
reduce strain) despite reductions i n plant water potential ( ) .
dehydration (desiccation) avoidance Abilit y to preven t reductions i n plant water
potential ( ; i.e., reduce stress) despite reductions in environmental water availability.
drought A condition i n which the availabilit y of soil water on a meso- o r macroenvironmental scal e i s generally insufficient t o suppl y the maxima l requirements of
otherwise well-adapte d plants .
drought avoidance Th e abilit y of a plant to survive or yield under drought conditions by increasing it s water suppl y (e.g., by deep rooting) relative to other plant s in
the sam e environment . Doe s no t impl y desiccation avoidance.
drought escape Completio n o f the entir e lif e cycle , o r critica l portion s thereof ,
during drought-free period s i n an otherwise drought-dominated environment .
drought resistance Th e ability of a plant to survive or yield under drought conditions relative t o other plant s in the same environment. Drought tolerance, avoidance,
and escape ar e type s of drought resistance .
drought stres s Th e degre e t o whic h soil-wate r availabilit y fail s t o mee t th e
maximal requirement s o f otherwis e well-adapted plant s ove r a meso - o r macro environmental scale .
drought tolerance Th e abilit y of a plant to survive or yield under drought condi tions despit e th e lac k o f availabl e soi l wate r t o mee t it s maxima l needs. Plan t
survival ma y be based o n reduction s i n water need (e.g. , stomata l closure) ; hence ,
drought toleranc e does no t impl y desiccation tolerance.
evaporative demand Th e capacit y o f th e aeria l environmen t t o caus e wate r
evaporation fro m a give n object , usuall y a plan t o r plan t canopy . Evaporativ e
demand i s influence d b y man y environmenta l factor s includin g radiatio n load ,
temperature, vapor pressure deficit (VPD), an d win d spee d i n additio n t o plan t o r
plant-canopy characteristics .
flooding Th e partia l o r complet e covering of a n organis m b y water resulting in
anaerobic stress.

Stress Physiology 15
hardening Se

e definition i n Section 1 of this chapter .

homoiohydric Plant s tha t maintai n a relativel y constan t leve l o f hydratio n

compared wit h large changes tha t occu r i n the water potential o f their environmen t
(e.g., vascula r plants) .
hydrophyte A plan t adapte d t o environmenta l condition s o f partia l o r ful l
submersion i n fre e water .
mesophyte A
availability.

plan t adapte d t o environmenta l condition s o f moderat e wate r

osmoregulation (osmotic adjustment) Partia l o r complet e maintenanc e o f cel l

turgor ove r a rang e o f tota l water potential s b y regulated change s i n cel l osmoti c
potential. Thi s process i s well documented i n the turgo r an d volume maintenanc e
of certai n alga l specie s when expose d t o change s i n th e osmoti c potentia l o f thei r
aqueous environment . Th e adaptiv e importanc e o f thi s process i n vascula r land
plants i s controversial .
paraheliotropic Orientin g parallel t o the incident solar radiation. Paraheliotropi c
leaf movement s i n plant s unde r water stress reduce s intercepte d radiatio n an d ca n
reduce leaf temperatur e an d leaf transpiration .
phraeatophyte A
plan t wit h the abilit y to thriv e in a dry environment, i n which
high evaporativ e demand s ar e me t b y havin g root s dee p enoug h t o reac h a
permanent wate r tabl e (se e drought avoidance).
poikilohydric Plant s that equilibrate with the water potential of their environment
and, for lan d plants, ar e capabl e o f assuming a wide range of hydration states.
Relative Water Content (RWC) Th
e wate r conten t o f a tissu e expresse d a s a
percent of the wate r content o f the sam e tissue at ful l hydratio n (=
0).
SPAC Th e concept o f transpirational water flow through aSoil-Plant-Atmosphere
Continuum, usuall y i n term s o f a n Ohms-La w analog y with A T a s th e potentia l
gradient, transpiratio n a s th e flux , an d th e hydrauli c resistance betwee n an y two
points o f the pathwa y as the rati o gradient/flux.
succulents Plant s havin g thick leave s with a high tissue-water content. Typicall y
xerophytes tha t hav e a thic k cuticle, lo w transpiration rates , an d th e hig h water use
efficiency (WUE) associated wit h CAM metabolism .
vapor pressure deficit (VPD) Th e difference between the actua l vapor pressure of
water i n th e ai r an d it s saturate d (i.e. , 10 0 % relativ e humidity ) vapo r pressure ,
expressed i n pascals : Pa . I t i s als o logica l t o us e th e ter m vapor density deficit
although thi s i s seldo m done . (Vapor density is expressed i n gram s of wate r pe r
cubic meter o f air).
water deficit An y value of tissue tha t is below the highest value exhibited by that
tissue i n it s mos t hydrated natural state. Doe s no t impl y water stress.
water stres s Ca n refe r t o a n environmenta l condition of either exces s or lac k of
water, but th e ter m i s usually used t o indicat e a lack of water. Wate r stres s a s a n
environmental conditio n (sometime s referre d t o a s drought stress) shoul d b e

154 Plant

Growth and Development

expressed a s th e amoun t o f water availabl e to a plant compare d wit h it s maximal

requirements. Wate r stress as a physiological conditio n ma y be expressed as plantwater potential ( ) , but the association betwee n an d water-stress-related injur y is
controversial. Se e stress i n Section 1 of this chapter .
water status Th e genera l conditio n o f a plant relative to any limitations impose d
by a lac k of water .
water use efficiency (WUE) A
comparative measure of plant productivity per unit
water used, in which the appropriate units for productivity and water use will depend
on the objectiv e o f the comparison . Agricultura l WUE can be defined for a growing
season either as yield per unit irrigation water applied or biomass produced pe r unit
of transpiration, wherea s instantaneou s physiological WUE can be defined as moles
of carbon dioxid e fixe d b y photosynthesis pe r mol e of water transpired .
xeromorphy Morphologica l characteristics , suc h as small , tough leaves , a heavy
cuticle, etc. , that ar e generally important for adaptation to dry environmental conditions.
xerophiiy Th e combined adaptiv e morphological and physiological characteristic s
of plant s able to gro w under dr y environmental conditions.
xerophyte A
availability.

plant adapted to environmental conditions of severely restricted water

5. SALINITY STRESS4
acclimation Se

e definition in Section 1 of this chapter .

biosalinity Interpla

y of saline habitat s and th e organism s living within them.

cation exchange capacity (CEC) Th e total quantity of cations that a soil or othe r
material can adsorb a t a specific pH, usuall y expressed a s centimoles (millimole s is
preferred SI ) of a specifi c cation pe r kilogra m of exchanger.
chloride salinity Salinity

i n which chlorid e is the dominan t anion.

clay dispersion Th e separatio n o f cla y int o individua l component particle s re sulting from th e presence of monovalent cations. I n saline soils, sodium is frequently
responsible for th e dispersio n o f clay. Soil s with dispersed cla y are dens e and hav e
a greatl y reduced permeability .
compatible solute A n organic solute that accumulates inside cells without causing
severe metabolic disruption and may aid osmoregulation; for example, glycine-betaine,
proline, glycerol.
cyclic salt Sal t derive d fro m th e se a or salt lakes that is deposited o n plant s and
soils fro m win d or rainfall.

Original authors for thi s section were D.W. Rains and E.V . Maa s who wish to express special
thanks t o Richar d H . Niema n for hi s thorough review of and suggestion s for thi s set o f definitions .

Stress Physiology 15

dehydration Los s of water fro m plan t tissue s i n excess of water uptake .

desalination Th e removal of salt from soil or water by physical or chemical means .
electrical conductivity (EC) A measur e o f th e abilit y of a solutio n t o conduc t
electrical current ; i t is correlated wit h the concentration o f the ions in the solution .
EC i s used t o expres s th e concentratio n o f salt in saline soil solutions and i s given
as decisiemens pe r mete r (dS/m ; millisiemens pe r mete r i s preferred SI) .
euhalophyte (true halophyte) A plant whose growth is maximum a t a soil wate r
salinity of about 1 0 g total dissolve d solids pe r lite r ( S 0. 8 MPa; EC 2 0 dS/m)
and decreases at eithe r highe r o r lowe r salinities .
exchangeable sodium percentage (ESP) Th e percentage o f the total exchangeabl e
cations tha t i s sodium. Soil s wit h high ESP ca n be difficult soil s t o reclaim .
flocculation Th e clumpin g o f cla y particle s int o aggregate s a s a resul t o f th e
neutralization o f charged surface s of the clay .
glycophyte A plant tha t grows optimally in nonsaline habitats .
halomorphic soil A suborder o f the intrazonal soil order, consisting of saline and
sodic soils forme d unde r imperfect drainage in arid regions and including the grea t
soil group s Solonchak or Salin e soils , Solonetz soils, and Soloth soils .
halophyte A plan t tha t grow s and complete s it s lif e cycl e in saline habitats .
hardening Se e definition in Section 1 of this chapter .
ionic effect A toxi c or nutritiona l effec t o f specific ions on a plant.
leaching requirement (LR) Th e amoun t o f water i n exces s o f the plan t require ment tha t i s neede d t o remov e sal t fro m th e roo t zon e i n orde r t o preven t soi l
salinity fro m exceedin g a specified value.
mangroves Grou p o f woody plants (about 1 1 genera) tha t grows in saline aquati c
habitats (e.g. , marine estuarie s an d swamps ) and is capable o f tolerating sea water.
Common gener a includ e Avicennia and Rhizophora.
mesophyte A plant adapte d t o a moderately moist habitat.
miohalophyte A plant that exhibits maximum growth in nonsaline soils but steadily
decreasing growth with increasin g salinity; such plants tolerate highe r salinit y than
gtycophytes.
oligohalophyte A plant adapte d t o habitat s of low salinity ( s 0.0 4 t o 0. 4 MPa
at fiel d capacity ; EC 1 to 1 0 dS/m).
osmoregulation Change s i n cell osmotic potential tha t tend to maintai n turgor .
osmotic adjustment Change s in concentration o f certain cell solutes (se e compatible solute) in response to changes in cell water potential. Thes e changes contribut e
to osmoregulation. (Se e als o osmoregulation i n Section 4 of this chapter.)
osmotic effector Se e osmoticum.
osmotic shock Osmotic stress cause d b y a sudde n and drasti c change in externa l
osmotic potential.

156 Plant

Growth and Development

osmotic stres s Externa l osmoti c potentia l below or abov e cel l osmoti c potential
sufficient t o cause strain (e.g., reduced yield). Osmoti c stress almost always refers to
external osmoti c potentials belo w those in the tissues, but osmotic potentials abov e
those o f th e tissu e (i.e. , les s negative ) ma y have som e deleteriou s effect s suc h a s
causing certain fruit s (e.g. , cherries ) t o split.
osmoticum A solut e tha t decrease s osmoti c potential s o f cel l solution s a s it s
concentration increase s (i.e. , any solute).
physiological drought Plan t water deficit (i.e. , low tissue water potential) cause d
by salinity, lo w soil wate r potential, o r othe r stress factors.
polyol A

compound containin g many alcoho l groups.

reclamation Wit h reference to saline soils, the process of removing excess sal t t o
a leve l tha t permit s production o f plants without significant advers e effects .
saline adaptation Geneti c modification of individuals in a population that increases their abilit y to surviv e excess salt.
saline adjustment Physiologica l and biochemical changes of individual plants that
increase thei r ability to surviv e excess salt .
saline-sodic soil A

soil tha t i s both saline an d sodic.

saline soil A soi l tha t ha s a n electrica l conductivit y in a saturated-paste extrac t

greater tha n 4 dS/m. Th e solubl e salt conten t o f such soils is sufficient t o interfer e
with the growth of many plant species. Sensitiv e crop plants are affected a t half this
salinity and highl y tolerant one s a t abou t twice this salinity.
saline stress Externa l salt concentrations sufficiently high to reduce plant growth.
Injury ma y resul t fro m osmotic stress, io n toxicitie s (se e ionic effect), and/o r
nutritional imbalance .
salinity Presenc e of inorganic ions in solution. Th e term often i s used to refer t o
ion levels hig h enough t o cause osmotic stress. Th e predominant ions include Na +,
Ca2+, Mg 2+, Cl- , SO42-, and HCO 3'. Thes e ions may have specific io n effects a s well
as osmoti c effects . Boro n an d othe r toxi c substances ma y be presen t bu t d o no t
contribute significantl y to salinity.
salination Th

e proces s o f accumulation of soluble salts in the soil .

salinity threshold Th e maximum salinity a plant can tolerate without a reductio n

in growth below tha t which occurs unde r simila r but nonsalin e conditions .
salt-affected soil A soil containing excessive concentrations of soluble salts and/or
exchangeable sodium.
salt balance A steady-state concentration o f salt determined by a balance between
the influ x o f salt an d th e efflu x o f salt fro m th e system . Thi s balanc e i s frequently
disturbed b y inappropriate managemen t o f water and/o r soil s o r a chang e i n th e
climatic conditions .
salt glands Specialize d cells or group s of cells on o r nea r the surfac e o f leaves or
stems tha t secrete salt an d hel p control levels of salt in plant cells.

Stress Physiology 15

salt resistance Th e abilit y o f plant s t o tak e som e actio n o r emplo y som e

mechanism t o avoi d detrimenta l effect s o f salinity. Th e mechanis m o f resistanc e
should b e specifie d i f it i s known.
salt tolerance Th e abilit y of plants to cope with salinity.
saturated soil paste A particular mixture of soil and water. A t saturation, the soi l
paste glisten s as it reflect s light , flows slightly when the container i s tipped, an d th e
paste slide s freel y an d cleanl y from a spatula .
saturation extract Th
tent.

e solution extracte d from a soil a t its saturation water con -

saturation percentage Th e water content of a saturated soil paste, expressed o n a

dry-mass percentage basis .
sodic soil A soil that has an exchangeable sodium percentage (ESP) greate r tha n 15.
The exchangeabl e sodiu m of such soils often adversely affects soi l structure and may
be deleterious t o plan t growth.
sodicity Presenc

e of excess sodiu m in soils wit h a pH rang e of 8.5 to 10.0 .

sodium adsorption ratio (SAR) A relatio n betwee n solubl e sodiu m and solubl e
divalent cation s tha t ca n be use d t o predic t exchangeable sodium percentage of soi l
equilibrated wit h a give n solution. I t i s defined as follows:

where concentration s ar e expresse d i n millimoles per liter .

succulence 1 . A morphological conditio n denoting thick, fleshy plant organs with
a hig h water content. 2 . Juiciness ; measure d as the water content pe r uni t surface
area o f plant tissue . (Se e definitio n i n Section 4 of this chapter. )
sulfate salinity Salinity
xerophyte Se

i n which sulfate is the majo r anion.

e definitio n in Sectio n 4 of this chapter.

REFERENCES

[Anonymous]. 1984 . Glossar y o f soil science terms. Soi l Sci. Soc. Amer., 67 7 South Segoe Road ,
Madison, Wisconsin , p 38.
Burke, M.J. , L.V . Gusta , H.A , Quamm e an d C.J . Weiser . 1976 . Freezin g an d injur y i n plants.
Annual Review o f Plant Physiology 27:507-528 .
Chapman, V.J . 1960 . Sal t Marshe s an d Sal t Desert s o f th e World . Interscienc e Pub., Inc . New
York, p 392 .
Franks, F. 1981 . Biophysic s and biochemistry of low temperatures and freezing. In : G.J. Morri s an d
A. Clark , editors . Effect s o f Lo w Temperature s o n Biologica l Membranes . Academi c Press ,
London, p 3-19 .
Levitt, J . 1980 . Response s o f plant s t o environmenta l stresses. Vo l 1 , Chilling , freezin g an d
temperature stresses . Academi c Press, Ne w York.

158 Plant

Growth and Development

Levitt, J. 1980 . Response s of plants t o environmenta l stresses, Vol 2, Water, radiation , salt, and
other stresses. Academi c Press , Ne w York.
Li, P.H. an d A Sakai , editors. 1978 . Plan t Cold Hardiness and Freezing Stress . Vo l 1. Academi c
Press, Ne w York.
Li, P.H . an d A Saka i 1982 . Plan t Cold Hardines s and Freezin g Stress. Vo l 2. Academi c Press ,
New York.
Little, R.J . an d C.E . Jones . 1980 . A dictionary of botany. Va n Nostran d Reinhold, Ne w York.
Luyet, B.J . 1968 . Th e formatio n o f ic e an d th e physica l behavior o f th e ic e phas e i n aqueou s
solutions and i n biological systems. In : J. Hawthorne and E.J. Rolfe , editors. Lo w Temperatur e
Biology of Foodstuffs. Pergamo n Press , Ne w York, p 53-77 .
Lyons, J.M. an d R.W . Breidenbach . 1987 . Chillin g injury . In : J . Weichman , editor. Postharves t
Physiology of Vegetables. Marce l Dekker, Inc. , New York, p 305-326.
Mazur, P. 1969 . Freezin g injur y i n plants. Annua l Review of Plant Physiology 20:419-448.
Raison, J.K. an d J.M. Lyons . 1986 . Chillin g injury: a plea for uniform terminology . Plant , Cell, and
Env. 9:685.
Sakai, A. and W. Larcher. 1978 . Fros t survival of plant responses and adaptation to freezing stress.
Springer-Verlag, Ne w York.
Salisbury, F.B. , an d C.W. Ross. 1992 . Plan t Physiology , Fourth edition. Wadswort h Publishin g Co. ,
Belmont, California.
Saltveit, M.E. , Jr . an d L.L . Morris . 1990 . Overvie w on chillin g injur y o f horticultura l crops. In :
C.Y. Wang, editor. Chillin g Injury o f Horticultural Crops. CR C Press, Inc., Boca Raton, Florida .
p 3-15 .
Sinclair, T.R . an d M.M . Ludlow . 1985 . Wh o taugh t plant s thermodynamics ? Th e unfulfille d
potential o f plant water potential . Aust . J . Plan t Physiol. 12:213-217.
Staples, R.C. an d G.H. Toenniessen , editors . 1984 . Salinit y Tolerance i n Plants - Stategies for Crop
Improvement. Joh n Wile y & Sons, New York.
Turner, N.C . 1979 . Drough t resistanc e an d adaptatio n to water stres s i n crop plants . In : Stres s
physiology i n cro p plants . H.W . Mussel l and R.C . Staples , editors . Wile y (Interscience), Ne w
York.
United States Salinity Laboratory Staff . 1954 . Diagnosi s and Improvement of Saline and Alkali Soils.
U. S. Dept. Agr. Handboo k No . 60. p 160 .
Weiser, C.J . 1970 . Col d resistanc e an d injur y i n woody plants. Science . 69:1269-1278 .
Weiser, C.J . 1982 . Plan t Cold Hardines s and Freezing Stress. Vo l 2. Academi c Press, New York.
Consultants
John S . Boyer Ton
University o f Delaware Orego
Lewes, Delawar e Corvallis

y H.H. Che n
n Stat e University
, Oregon

Kent J . Bradfor d Emanue

University of Californi a Universit
Davis, California Davis

l Epstein (retired )
y of Californi a
, Californi a

William J. Bramlag e Lelan

University of Massachusetts USD
Amherst, Massachusett s Riverside

d E. Francoi s
A Agricultural Research Servic e
, Californi a

R. Willia m Breidenbac h Catherin

University of Californi a USD
Davis, Californi a Riverside

e M. Grieve
A Agricultura l Research Service ,
, California

Michael J. Burk e Laurenc

Oregon Stat e Universit y Universit
Corvallis, Orego n Saskatoon

e V. Gusta
y o f Saskatchewan
, Saskatchewan, Canada

Stress Physiology 15
Anthony E . Hal l
University o f California
Riverside, Californi a
Robert L. Jefferie s
University o f Toronto
Toronto, Ontario , Canad a

Mikal E . Saltvei t
University of California
Davis, California

Delmer O . Ketchi e
Washington State Universit y
Wenatchee, Washingto n

Richard C . Staple s
Cornell Universit y
Ithaca, Ne w York

Mark A . Matthew s
University of Californi a
Davis, Californi a

Donald L . Suare z
USDA Agricultural Researc h Service
Riverside, Californi a

Richard H . Niema n (retired )

USDA Agricultural Researc h Servic e
Riverside, Californi a
James W. O'Leary
University o f Arizon a
Tucson, Arizon a

Charles G . Suhayda
USDA-Agricultural Research Servic e
Pasadena, California

Robert E . Paull
University o f Hawaii a t Mano a
Honolulu, Hawai i
Robert W . Pearc y
University of California
Davis, Californi a
Albert C . Purvis
University o f Georgi a
Coastal Plai n Experiment Statio n
Tifton, Georgi a
Harvey A . Quamm e
Research Statio n
Summerland, BC , Canad a
James D. Rhoades
USDA Agricultural Research Servic e
Riverside, Californi a
Frank B . Salisbury
Utah State Universit y
Logan, Uta h

Michael C . Shannon
USDA Agricultura l Research Servic e
Riverside, Californi a

Karen Tanin o
University of Saskatchewa n
Saskatoon, Saskatchewan , Canada
Ralph Weimber g (retired)
USDA Agricultura l Research Servic e
Riverside, California
Chien Y i Wang
U.S.D.A. Beltsvill e Agricultura l Researc h
Center, Maryland
Conrad J . Weise r
Oregon Stat e Universit y
Corvallis, Orego n
Clyde Wilso n
USDA Agricultura l Research Servic e
Riverside, California
Michael Wisniewski
USDA Agricultural Researc h Service
Kearneysville, Wes t Virginia
Anne F . Wron a
University of California
Holtville, California

APPENDICES
PRESENTING SCIENTIFIC DATA
Anyone ca n d o experiments , bu t i f th e result s ar e t o qualif y a s science , mor e i s
required tha n special techniques, logical thought processes, and elaborate equipment .
It i s not th e whit e laborator y coat s an d foul-smellin g chemicals , o r eve n th e fiel d
stations o r telescope s o r orbitin g satellites , tha t qualif y a work as science. T o b e
science, the results o f one's investigations must be presented t o one's colleagues fo r
their evaluatio n an d perhap s fo r the m t o us e in their ow n attempts t o expan d th e
limits o f huma n understanding . Dat a mus t b e publishe d o r otherwis e presented .
"Publish o r perish " may be thought of by some as an unfair deman d placed o n goo d
teachers, but without publication (presentation) , a n investigation does not qualif y as
science. Indeed , unles s th e result s o f one' s effort s ar e communicate d t o one' s
colleagues, th e resources (ofte n ta x funds) use d to produce thos e result s ar e wasted.
Presentation of data requires the use of language. Typically , results are described
in a scientific paper, s o Appendix A is a discussion of writing. Th e discussio n doe s
not emphasiz e th e usua l forma t o f a scientifi c paper (Introduction , Method s an d
Materials, Results , Discussion) ; suc h a patter n i s wel l know n t o scientists , an d
specific detail s can be seen in the journals t o which authors inten d t o submit thei r
manuscripts. I t is imperative tha t authors consult such sources before preparing and
submitting manuscripts . Th e discussio n her e present s som e principle s o f Englis h
grammar and style. Althoug h spac e limitation s eliminat e a complete analysi s of the
English language , a n attempt i s made to cover th e basic s in a somewhat logica l way
and to emphasize a few points that often ar e not appreciated. Scientist s al l over th e
world mus t now prepare som e o f their manuscript s in English even when English is
not thei r nativ e language . Althoug h goo d cop y editor s kno w th e rule s discusse d
here, many scientist author s apparentl y do not , an d fe w journals do an y extensive
copy editing. Hence , it is easy to find example s in the curren t scientific literature of
the problems describe d i n Appendix A .
Data are als o communicate d at scientific meetings . Traditionally , this has bee n
an oral presentation illustrate d b y slides. Durin g recent years , such slide talk s have
included man y tex t slide s a s wel l a s th e traditiona l photographs , dat a graphs ,
diagrams, and othe r figures . Compute r program s have made it possible to prepar e
beautiful an d elaborat e slides . Nevertheless , suc h talk s ar e sometime s poorl y
presented an d poorl y received . Member s o f the audienc e ma y have to struggl e t o
understand what is being discussed. I n many cases, communicatio n could be better
achieved i f a few simple rules were followed durin g preparation and presentatio n of
161

162 Appendices
the slides . Appendix B discusses many of these rules. Man y scientific societies no w
emphasize poster s ove r ora l presentations , an d again, many posters ar e difficul t t o
assimilate i n th e availabl e time. Thus , Appendi x B also discusse s some rules an d
suggestions tha t can improve communication between a poster presenter an d his or
her audience. W e recognize that writing styles, slide talks, and poster presentation s
represent the personal expressions of their authors. B y the same token, Appendices
A an d B represent persona l viewpoints. W e hope thi s wil l no t detrac t fro m thei r
potential value .
Plant scientists ofte n depen d upon growth chambers in their research , but ther e
are many kinds of chambers and many ways that the experimental conditions ca n be
reported. Thus , Appendix C presents guideline s fo r measurin g and reportin g th e
environmental parameter s o f growth-chambe r experiments. Th e guideline s wer e
formulated by a special committee of the American Society of Agricultural Engineers
(ASAE). Th e guidelines are essentially an application of the principles presented in
Section II . The y have been edite d slightl y to conform , a s far a s possible, t o othe r
recommendations i n this book.

A
SOME SUGGESTIONS ABOUT SCIENTIFIC WRITING
Frank B . Salisbury
Plant Scienc e Departmen t
Utah Stat e Universit y
Logan, Utah 84322-482 0 U.S.A .
As a budding young scientist a t th e university , I found m y English classes t o be very
distasteful. Ther e seeme d t o b e n o logicn o sciencet o the language , onl y a n
assemblage o f arbitrary rules. The n I lived for two years in Switzerland and learne d
German, a highl y logical language . Gradually , through analog y with the Germa n
grammar I was enjoying s o much, I began t o se e som e glimmering s of logic in my
native tongue. Later , a s I became involve d in text-book writing, the cop y editors at
Wadsworth Publishing Company piqued my curiosity by the changes they made in my
manuscripts. Wh y should which sometimes be changed to that, for example? I began
to browse the style manuals and rule books, findin g a little more logic in the English
language, ofte n burie d i n th e assemblag e o f arbitrary rules.
For th e pas t fe w years, I hav e bee n editin g submitte d manuscript s fro m th e
Americas an d Pacifi c Rim countrie s fo r the Journal of Plant Physiology . O f cours e
there ar e special problem s face d b y authors whose native tongue is not English , and
part of the discussio n her e has them in mind. Furthermore , not al l my fellow nativ e
English speaker s se e th e structur e o f Englis h a s I do. Rule s o f punctuation , fo r
example, base d o n th e logi c o f the languag e as I came t o understan d it an d a s th e
manuals describe it, seem to be breaking down. Usag e continually changes the rules.
What follow s summarizes my personal vie w of how the logi c of English can b e
expressed wit h th e hel p of punctuation and a suitable choice of words. A s fa r as I
know, my approach t o th e topi c i s unique although I can back up ever y rule with
references t o the much more extensive manuals, and my discussion has been checked
by three grammarians. I hope tha t you will approve of this approach and apply its
recommendations. Tim e will tell ho w much usage will change the rule s and henc e
my approachand perhaps mak e th e language even more arbitrary and less logical !
Clearly, th e bes t communicatio n require s a n agreemen t o n languag e convention s
between bot h reade r an d author . A prope r comprehensio n o f suitabl e writin g
conventions i s essential, just as acceptance an d understanding are required for proper
use of the othe r units , symbols, and term s presented in this book. T o this end, th e
following i s offered .
163

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Presenting Scientific Data

1. THE SENTENCE
While i t may be impossible t o define a sentence i n a broad sense that will cover
all example s (Pinckert , 1986) , a sentenc e i n technica l writin g is seldom difficul t t o
recognize. Indeed , grammarian s recogniz e onl y six basic sentenc e structure s (se e
box). A complet e sentenc e contain s a t leas t on e subjec t ( a noun) wit h it s verb
(Plants grow. Jesus wept.), and most of the time it also includes an object of the verb
along with modifying adjectives, adverbs, various phrases tha t act as modifiers, and
often a conjunction tha t join s one word with another, one phras e with another , o r
one claus e wit h another. Prepositions ma y be placed i n front o f nouns t o show the
relationship o f the nou n to othe r word s in the sentence, an d pronouns (which, in a
sense, are reall y nouns) ma y be used t o substitute for other nouns . Sometimes , bu t
not ofte n i n technical writing , an interjection may be added although i t ha s no rea l
relationship t o anything in the sentence . (A h ha, we discovered that plants grow.) A
sentence may include any of the eight parts of speech (writte n in bold face above), bu t
the key to recognizing a complete sentenc e is to recognize the subject and the verb.
The verb i s needed fo r the predicate (what is being said about th e subject, including
the verb wit h or without objects , complements , o r modifiers) . I f either th e subjec t
or the verb is missing, the resul t is a sentence fragment (or an incomplete sentence)
rather tha n a complete sentence .
Writing elementary sentence s seldo m cause s any difficulty fo r a scientific writer,
but problem s sometime s aris e whe n a sentenc e contain s mor e tha n on e subjec t
and/or mor e tha n one verb. Shoul d tw o ideas, each with a subject and predicate, b e
included i n on e sentence , o r shoul d the y be separate d int o tw o sentences ? Ho w
should the relationship betwee n these ideas be formulated and expressed? Thes e are
decisions tha t a n author mus t make in the attempt to best communicate what he or
she wants to say. I t is essential t o kno w the availabl e options i f one i s to mak e th e
best decisions. Th e relationship betwee n tw o ideas can be expressed at several levels ,
and thes e ar e indicate d b y various system s of punctuation.
A. Closely Related; Subject or Verb is Shared. Man y authors , includin g
technical writers , ti e togethe r (coordinate ) tw o subject-predicat e idea s wit h a
coordinating conjunction and omit the subject before the second verb , knowing that
the reader wil l refer to the original subject to understand the second predicate . Th e
sentence just presente d provide s a n example . Th e subjec t is authors (modified by
several words) , th e firs t ver b i s tie, an d th e secon d ver b i s omit, whic h als o ha s
authors a s it s subject . Th e idea s ar e tie d togethe r b y the conjunctio n and, which
coordinates th e tw o ideas . Coordinating conjunctions includ e and, but, or, nor, for,
and so. Not e tha t th e firs t clause ( a group o f words with expressed o r understoo d
subject an d predicate ideas ) i s independent because it has both a subject and a verb,
while the secon d clause i s dependent on th e first . Th e secon d claus e i s dependen t
because i t lack s a subject o f its own but depend s o n th e independen t claus e fo r it s
subject (a s i n thi s sentence , followin g but). Som e dependen t clause s lac k a ver b
instead o f a subject althoug h thi s i s less common . Th e questio n concerns ho w th e
relationship between th e two ideas in such sentences should be communicated to th e
reader b y punctuation. Sinc e bot h idea s shar e a commo n subject, the y are closel y
related, an d logi c suggest s tha t n o comm a (n o paus e i f spoke n aloud ) i s neede d

Some Suggestions about Scientific Writing 16

THE SIX SENTENCE STRUCTURE S I N ENGLISH

The English sentence has six basic patterns. N o matter how complicated a sentence is, it can be broken dow n into one or more of these
designs:
1.

subject verb

2.

subject verb + modifier s

adjective adverb
prepositional phrase

3.

subject verb

4.

subjective complemen t (eithe r a n adjective ,

noun, o r pronoun)
subject verb object

5.

subject verb object

indirect objec t

6.

subject I verb

objective complemen t object

Examples;
1. Jesus wept.
2. The dirty clothes are probably in the hamper.
3. I t i s cold. I t tastes sour. This is he. (Th e complemen t i s either
an adjectiv e o r a nominative-case nou n or pronoun. )
4. W e found him. Sh e measured the plant. (Th e object is always in
the objectiv e case , a matter o f concern onl y with pronouns. )
5. H e gave her the ring. (Th e indirect object, her, defines the recipi ent o f the actio n o f a transitive verb.)
6. Th e sight turned hi s hair grey. Th e speech made everyone angry.
We elected him president (Th e objectiv e complement tell s what
happened t o th e object a s a result o f the actio n o f the verb. )
This material was supplied b y Moyle Q . Rice .

before th e conjunction . Th e majorit y o f style manuals and rul e book s on Englis h

writing agree (althoug h som e suggest that but is an exception an d should always be
preceded b y a comma).
It i s commo n practic e i n moder n writing , however , to inser t th e unneede d
comma. Becaus e usage rathe r tha n logi c actually dictate s ho w w e shoul d write,

166 Appendices:

Presenting Scientific Data

adding th e unneede d comm a ma y now be so common tha t i t ca n hardly be called

incorrect. I t i s usage tha t teache s a reader t o understan d the sam e convention s a s
the author. Nevertheless , a reader seldom if ever fails to understand such a sentence
with n o comm a just because th e comm a i s omitted . Therefore , i t i s logica l t o
recommend, i n spite of increasing usag e to the contrary, that the comma between a
dependent an d a n independen t claus e connecte d wit h a conjunctio n b e omitted .
What i f the tw o clauses becom e s o long and complex that th e autho r strongl y feels
a nee d fo r a pause an d want s t o inser t a comma to sugges t it ? The n th e comm a
should probabl y b e added , bu t i t migh t be bette r t o brea k th e sentenc e int o tw o
sentences or to go to one of the next levels by repeating the subject and making both
clauses independent ; that is , avoid creating th e problem .
Once the concepts are understood, punctuation should put onto paper what the
author want s th e reade r t o feel . A comm a mean s a pause , an d suc h a paus e i s
usually not needed if the second clause is dependent. Fo r example, one can read this
sentence aloud an d ca n note that n o paus e is needed befor e the and. Th e subjec t
(one) o f the first claus e carries over to the second, dependent clause. (Th e two verbs
are ca n read an d ca n note; the secon d ca n could be omitted.)
If you want to paus e for the sake of emphasis, the appropriate punctuation is a
dash (called a n em dash), which was invented just to indicate a long pause that shift s
the emphasi s to th e en d of the sentence . A dash () i s longer than a hyphen (-) or
made on a typewriter with two hyphens (--); when used like thisit is usually not se t
apart with spaces. I t is easy to overuse the dash-and thus to weaken its effect (lik e
that). Furthermore , dashe s ar e seldom use d in scientific writing. A s a punctuation
mark, the das h convey s a certain emotion , an d emotion mus t generally be avoide d
in technical writing. Som e grammarians suggest that the dash should never be used .
B. Closely Related Independent Clauses Connected by a Coordinating Conjunction. Ofte n two more-or-less equal ideas, each expressed with both subject and verb,
are tie d togethe r (coordinated ) wit h a coordinating conjunction. I n this case, bot h
logic an d th e styl e manuals (especiall y i n America ) sugges t th e us e o f a comm a
before th e connectin g conjunction , and usag e agai n seems t o b e goin g toward n o
comma. Nevertheless , th e recommendatio n i s to us e the comma . I used i t i n th e
sentence about styl e manuals and usage , and her e i t i s in thi s sentenc e a s well. I
doubt tha t anyon e will be upset. Th e rule books generall y say that short , indepen dent clause s following a conjunction do not need the comma before the conjunction
and tha t make s some sense. (Di d I need a comma before that last and ? Probabl y
not.) Thus , ther e i s som e leewa y i n th e us e o f a comm a befor e a coordinatin g
conjunction connectin g independen t clauses , but a s a general rule, that comma can
be helpful. I t should b e use d mor e frequently than it is.
There are times when it is needed for clear understanding. Without it, a reade r
may hav e t o bac k up , star t again , and realiz e onl y on th e secon d readin g tha t th e
noun followin g th e conjunctio n wa s th e subjec t o f a secon d independen t claus e
instead of the secon d object of the firs t clause : W e measured the auxin and gibberellin
was present but was not measured. I t would be wel l t o recas t tha t sentence , bu t a
comma afte r auxin would help i t a s it stands.

Some Suggestions About Scientific Writing 16

C. Closely Related Sentences (no Conjunction). Sometime s an author wants to

show that tw o sentences ar e closel y related; the y may not be related closel y enoug h
to connect the m with a conjunction, however. I n such a case, one has the option of
simply writing two sentences, eac h terminate d with it s own period, or on e ma y put
a semicolon between them , as in the previous sentence. O f course, there ar e othe r
possibilities. Woul d the opening sentence hav e been better this way: Sometimes an
author wants to show that two sentences are closely related, but they may not be related
closely enough to connect them with a conjunction. Tha t is, those two ideas could have
been coordinate d wit h the conjunctio n but.
There ar e tw o kind s o f error s tha t appea r i n thi s situation . Bot h shoul d b e
avoided. Th e firs t i s the comma fault (o r comma splice) in which the tw o complet e
sentences are connected wit h a comma but no conjunction. Thi s is considered a fatal
error by all editors, it must be avoided under all circumstances. ( I hope you caught
it just now. )
The secon d erro r i s being supporte d b y some limited usage; indeed, ther e is a
tendency for scientific authors to feel tha t they are being especially "scientific" when
they use however instead o f but: An author may want to show that two sentences are
closely related, however they may no t b e related closel y enough to connect them with a
conjunction. Word s lik e however, rather, nevertheless, thus, indeed, an d other s ar e
connectives an d no t conjunctions . The y ar e als o calle d conjunctive adverbs; they
both modif y an d connec t a t th e sam e time . The y are punctuate d as interjections .
Thus, i f the autho r fel t compelle d t o us e however i n tha t sentence , i t shoul d hav e
been punctuate d lik e this: A n author may want to show that two sentences are closely
related; however, they may not be related closely enough to connect them with a conjunction. Tha t is, the second par t of the compound sentence (afte r the semicolon) shoul d
be punctuated in the sam e manner as when a conjunctive adverb begins a sentence:
However, they may not be related . . . . I n general, however, an author should realize that
overuse of such connectives sounds stuffy, affected , professorialan d scientific, if that
is what usage has taught us. Nevertheless , writing can often b e greatly improved by
eliminating such words. Surely , a reader deserve s th e credit for being able to see the
connection between ideas without such unneeded "help." (Were connectives overused
in thi s paragraph ? Probably. )
D. Les s Related Sentences. Thes e ar e separate d with periods. O f course they
are no t unrelated , whic h i s wh y the y ar e combine d togethe r i n a paragraph .
Separation wit h period s i s th e usua l an d appropriat e wa y o f writing , bu t i t i s
important to have the other levels at one's fingertips in case they better communicat e
an author' s ideas . Bu t avoid th e comm a fault !
E. Two Ideas are Connected with a Subordinating Conjunction. W e have been
discussing situations i n which two ideas ar e about equal to each other an d are tie d
together i n various ways (comma, semicolon) or not at all. I n another situation, two
clauses o f unequal importance are tie d together in the sam e sentence b y a subordinating conjunction. Ther e ar e man y kinds of subordinating conjunctions including
relative pronouns (that, which, who) and conjunction s of tim e (after, since, before,
when, while, as, and until), place (where, wherever), purpos e (so that, in order that),
comparison (than, as, a s if , but if , as though, whereas), condition (if, unless), conces -

168 Appendices:

Presenting Scientific Data

sion (although, though, even though), and caus e (because, since, and th e weake r for
and as). An y of these and other s ca n be used to show how the subordinat e claus e
relates t o th e mai n clause . Changin g tw o coordinat e clause s connecte d wit h a
conjunction t o a mai n claus e wit h it s subordinat e claus e ca n ofte n clarif y an d
generally improv e th e writin g (Pinckert, 1986) . Th e easy way is to us e coordinat e
clauses. A better way, although on e tha t require s som e menta l effort, i s to clarif y
relationships b y forming subordinate clauses. S o that it does not sound like an after thought, i t ofte n help s t o pu t th e subordinat e claus e firs t (a s i n thi s sentence) .
Consider thes e tw o sentences: Galleys should b e returned t o the editor after they have
been carefull y read. After th e galleys have been carefully read, they should b e returned
to the editor.
The rul e fo r punctuatin g between a mai n clause an d it s subordinat e clause i s
simple: I f the subordinat e claus e comes first , i t i s followed b y a comma; if it comes
after th e main clause, it is not. ( . . .it is not i f it comes after th e main clause.) Becaus e
a subordinate claus e shoul d never stand alone (formin g a sentence fragment rathe r
than a complet e sentence) , i t mus t alway s occu r i n th e sam e sentence a s it s main
clause. Thi s means that it should not be separated fro m a preceding main clause by
a comm a o r semicolon. Tha t i s the rule , althoug h it sometimes seems appropriat e
to ad d th e comm a fo r emphasi s (a s was done here). I n the mos t informal writing,
an author ca n add a dash to provide strong emphasis on th e subordinate phrase o r
clausealthough dashe s ten d t o b e overuse d b y writers who ar e insecur e i n thei r
knowledge o f punctuation. Whe n in doubt , add a dash! O f course thi s should be
avoided.
There is a complication. Som e subordinating conjunctions (e.g., although) can
be used a s conjunctive adverbs . I n such cases, punctuation should follo w th e rule s
described abov e fo r such conjunctiv e adverbs.
F. Beginning a Sentence with a Coordinating Conjunction. A t leas t on e
important questio n remains : I s it correc t t o begi n a sentenc e with a coordinating
conjunction (and, but, etc.)? Becaus e coordinating conjunctions are normall y used
to splic e together tw o clauses, they cannot come at the beginning of the sentence if
both clause s ar e presen t (a s th e subordinatin g conjunction because di d i n thi s
sentence). Bu t what abou t a sentenc e tha t begin s with a conjunction but contain s
only one clause ? Sometimes , for emphasis, an author may want to arbitrarily make
two sentence s ou t o f a compoun d sentenc e tha t ha s a coordinatin g conjunction
between tw o independent clauses . Doin g so ties th e sentenc e tha t begin s with th e
coordinating conjunction to the thought of the previous sentence mor e closely than
would be th e cas e without the conjunction . I t i s acceptable t o begin a sentence with
a conjunction. An d sometimes it can provide impact. I n thi s case, th e coordinatin g
conjunction act s more lik e a conjunctive adverb. Bu t like many devices of this type,
it ca n be overdone . I t should be use d with care.
2. MODIFYING WORDS
For th e mos t part , the us e o f modifyin g word s is relatively eas y in th e Englis h
language, but a fe w problems arise.

Some Suggestions About Scientific Writing 16

A. Adjectives. Adjective s ar e words that modify (explain ) nouns and pronouns .

Because the y ar e no t decline d accordin g t o gende r an d cas e a s i n man y othe r
languages, thei r us e i n Englis h i s quit e simple . Th e fe w problems tha t appea r i n
technical writin g concer n us e o f th e hyphe n i n compoun d adjectives , us e o f th e
comma when mor e tha n on e adjective modifies a noun, and distinguishing between
adjectives an d adverbs . (Som e language s don't us e articles, which are special adjec tives tha t limi t or give definiteness t o the things represented b y the noun. A an d an
are indefinit e articles ; th e i s th e definit e article: A beaker says ther e i s onl y on e
beaker bu t doesn' t specif y whic h one; th e beaker an d the beakers refe r t o a specifi c
beaker o r beaker s tha t ma y have already been mentioned. )
A compound adjective consists o f two or more words that act as a single idea t o
modify a noun . Th e compoun d adjectiv e i s usuall y formed b y placin g a hyphe n
between th e tw o or mor e word s ( a four-year-old tree).
The tw o words migh t bot h b e adjectives, on e migh t be an adjectiv e an d on e a
noun, or both might be nouns, but the combination acts as an adjective: a cold-water
shower. Eve n adverb s ca n b e par t o f compoun d adjectives . T o qualif y a s a com pound adjective , th e word s mus t ac t togethe r i n a specia l wa y rather tha n eac h
modifying th e nou n independently . Far-red light provide s a good example . W e ar e
not talkin g about a distant, re d light ; instead, we are discussin g light in th e far-red
part o f th e spectrum . Usually , when tw o or mor e adjective s precede a noun, eac h
modifies th e nou n independently: a bright, red, metal-halide lamp. Thes e are calle d
coordinate adjectives, and i n the example, one i s a compound adjective that consist s
of tw o noun s (metal an d halide) actin g togethe r a s a n adjectiv e that describe s th e
lamp.
It i s sometimes difficul t t o kno w whether a comma should b e use d betwee n a
series of coordinate adjectives . Curren t usag e often tends to eliminate commas, bu t
in technica l writin g i t help s t o us e them . I f there ar e mor e tha n tw o coordinat e
adjectives, comma s become mandatory; if there ar e onl y two, a helpfu l devic e i s t o
see whether th e insertio n o f and i s acceptable. I f it is, the tw o adjectives ar e quit e
independent o f each other , an d th e comm a should be added. I f the and sounds ou t
of place , the tw o adjectives ar e relate d t o eac h othe r almost as they would be in a
compound adjective; th e comma should be omitted. I f a bright and green light doesn' t
sound right , leav e ou t th e comma : a bright green light. I f a ho t an d re d light i s
acceptable, us e the comma : a hot, red tight. Clearly , the distinctions can be fine, an d
decisions mus t be lef t t o th e author , bu t th e author mus t be aware of the accepte d
conventions.
Two words in a compound modifie r might be an adjective and a noun, with th e
adjective an d th e nou n actin g together t o modif y anothe r noun : a short-day plant.
This doe s not sa y that th e plan t i s short o r tha t i t is a day plant but tha t th e plan t
responds t o short days. Not e tha t the combination short days b y itself does no t for m
a compound adjectiv e but i s simply a case of short acting as an ordinary adjective t o
modify th e wor d day. Th e hyphe n i s appropriate onl y whe n the combinatio n acts
together as an adjective to modify another noun (which is sometimes implied but no t
stated: irradiate th e seeds with far-red).

170 Appendices:

Presenting Scientific Data

It is appropriate in scientific writing to combine a number (an adjective) with a

unit (a noun) t o form a compound modifier when the number and unit are written out:
a four-hour night, a ten-milliliter aliquot, a five-gram sample, etc. (But , note : four
hours, te n milliliters, fiv e grams, etc.) A s discusse d i n Chapte r 1 , however, whe n
numerals and unit symbols are used, the hyphen should be omitted. Thi s i s because
the uni t symbo l shoul d b e though t o f a s a mathematica l symbo l tha t stand s fo r a
physical quantity . Thus , a 0. 5 kg sample i s th e mas s o f th e standar d kilogra m i n
Sevres, France , multiplie d b y 0.5 . Us e o f th e hyphe n destroy s thi s relationship .
When numerals an d symbols are spelled out , the rules of grammar apply; when thei r
symbols ar e used , mathematica l rule s apply .
Sometimes tw o or eve n thre e nouns ac t together a s an adjective. Logic would
suggest tha t the y shoul d b e connecte d wit h a hyphen , but usag e ofte n make s th e
hyphen see m ou t o f place: cell-wall materials or cell wall materials'? Mos t author s
would us e the secon d form , and most editors woul d probably change th e firs t t o th e
second form . Bu t stan d u p fo r logi c i f you fee l s o inclined, eve n i f your creation is
likely to b e changed b y an editor.
Adjectives ca n b e th e presen t o r pas t participle s o f verbs, i n which for m the y
often appear i n compound adjectives, especially in scientific writing: a light-controlled
switch (past participle); a light-controlling switch (present participle). Not e tha t th e
meanings ar e differen t i n these two cases. I n the first , ligh t controls th e switch ; in
the second , th e switc h control s th e light . Althoug h thes e form s ar e commo n i n
technical writing, the comparable longer constructions ar e often easier to understand:
a switch that light controls; a switch that controls the light.
To summariz e som e recommendation s fo r th e us e o f hyphen s i n compoun d
modifiers: D o no t us e wit h a combinatio n o f number s an d unit s tha t modifie s a
noun ( a 100 W lamp). Us e with nearly all other compound modifiers; this will make
it easier for most reader s an d offen d onl y those who have decided tha t punctuatio n
should be eliminated jus t for the sake of eliminating punctuation. Avoi d adding th e
hyphen between som e compoun d modifier s that ar e ofte n see n withou t i t (the high
school reunion), usually consisting o f two nouns rather than an adjective and a noun.
Do no t us e a hyphen betwee n a n adver b and an adjectiv e if the adver b ends i n -ly .
Otherwise, compoun d adjective s tha t include adverbs are often hyphenated : a welldone steak (bu t a steak that wa s well done), a well-known expert, a loose-fitting
garment (bu t a loosely fittin g coat), a well-guarded secret (bu t a carefully guarded
secret). Al l compoun d number s ar e hyphenate d (sixty-seven samples and there were
only sixty-seven), bu t chemica l term s neve r are : hydrogen sulfide gas.
Some word s hav e prefixe s or suffixe s tha t ar e alway s hyphenated: self-rule, ex husband. Other s ar e combine d wit h th e wor d an d neve r hyphenated : overdose,
underground, coworker. Whe n i n doubt , consult a dictionary.
B. Adverbs. A n adverb can modify verbs, adverbs, and adjectives; it is a versatile
part o f speech. I f a modifie r is modifying anythin g but a noun or pronoun , i t i s an
adverb. Adverb s tell where, when, how, or how much. Man y adverbs end i n -ly, but
many do not, an d not al l words ending in -ly are adverbs. Mino r trouble arises when
adverbs and adjective s are confused , bu t thi s occurs most often i n speech an d no t i n
technical writing. W e naturally write: Th e good experiment wa s done well. Not : Th e

Some Suggestions About Scientific Writing 171

well experiment wa s done good. Well i s the adver b that modifies the verb was done,
while good i s the adjectiv e tha t modifie s the nou n experiment,
Note tha t sense-impression verbs ar e followe d b y th e adjectiva l rathe r tha n
adverbial forms : I t tastes good. I t smells bad. H e looks sick. I feel ba d (rather tha n
I feel badly) . I feel good. ( I feel well means that I feel healthy rather than sick if well
modifies I-o r tha t I have a good sens e of touch if well modifie s feel.)
A word o f warning about adverbs : Man y writers seem t o have the notio n tha t
adverbs ar e ver y elegant, bu t ofte n the y are inherentl y vague. Ho w elegant i s very
elegant? Ho w vagu e i s inherently vague ? W e ca n ofte n tighte n ou r writin g by
eliminating very, too, greatly, really, actually, extremely, quite, rather, slightly, fairly ,
somewhat, t o a certain extent, and ver y man y others! O f cours e thes e word s ar e
sometimes useful .
C. Prepositions and the Objective Case. Englis h is relatively easy for foreigners
to lear n (t o begi n with , a t least ) becaus e nouns , a s wel l a s adjectives , ar e no t
inflected accordin g to case and gender. Thi s means that there are few problems with
the us e o f prepositions, whic h ar e word s tha t ar e place d i n fron t o f noun s an d
pronouns t o sho w the relationshi p o f the nou n to other words in the sentence : fil l
to the calibratio n mark , for a good reason , behind th e scutellum , with care, between
the lines , etc . Noun s followin g preposition s ar e alway s i n th e objective case, bu t
since English noun s have the same form in the subjective and the objective cases, we
seldom give the matter any thought. Tha t is probably why we may have trouble when
we use Englis h pronoun s tha t d o have a special for m i n th e objectiv e case. Ther e
are onl y si x of the m i n moder n English : me , him, her, us, them, an d whom (plu s
whomever and th e obsolet e thee). 1
We seldo m hav e troubl e whe n th e nou n follow s th e prepositio n directly , bu t
many Americans (wh o seem to have little sense of case because it is such a small part
of the language ) say between you and I instead of the correct between you and me, an d
it i s not uncommo n t o hea r with we girls instea d o f with us girls. Direc t o r indirec t
objects o f verb s ar e als o i n th e objectiv e case : H e gave her the unit, an d it s probe
contacted him an d me. I n technical writing, we use fewer personal pronouns anyway,
but w e must be careful when we do use them.
D. Personal Pronouns. A s a matte r o f fact , w e shoul d probabl y us e mor e
personal pronoun s i n our scientifi c writing. I t is highly artificial t o put ou r writing
always i n th e thir d perso n b y saying the author did thi s or tha t instea d o f simply
saying / o r w e did it. Man y modern editor s ar e no w insisting on the firs t perso n
instead o f the out-of-date third person. Fo r one thing, an author should be willing
to tak e responsibilit y fo r his or he r experimenta l results and philosophica l sugges tions by speaking i n th e firs t perso n instea d of hidin g behind some almos t anonymous author who seems t o be doin g the writing. Sayin g the author doesn't mak e a
paper an y more objective .

The subjective form s are /, he, she, we, they, who, an d thou; possessiv e forms are my, mine, his,
her, hers, its, our, ours, their, theirs, whose, and thine; it and you ar e th e sam e in the subjective and th e
objective cases .

172 Appendices:

Presenting Scientific Data

3. MODIFYING PHRASES AND CLAUSES

A. Restrictive and Nonrestrictive Phrases and Clauses. Problem s encountered
with phrase s ar e mostl y concerned wit h how they should b e punctuated. Ofte n th e
author must decide how to punctuate, depending on the exact meaning that he or she
wants t o convey . Th e ke y to makin g th e decisio n correctl y i s t o understan d th e
concept o f th e restrictiv e versu s th e nonrestrictiv e phras e o r clause . A restrictive
phrase o r claus e contain s informatio n that i s essential t o understandin g some par t
of th e sentenc e no t include d i n th e phras e o r clause ; a nonrestrictive phras e o r
clause add s informatio n tha t ma y well b e importan t bu t tha t i s no t essentia l t o
understand th e part s o f the sentenc e no t include d in the phrase o r clause .
These two possibilities can often be illustrated with the same sentence, whic h is
why th e autho r mus t decide whic h one expresse s hi s or he r intention . M y son, the
doctor, sent me a letter. Th e doctor i s a nonrestrictiv e phrase tha t suggest s that th e
speaker has onl y one son , who happen s t o b e a doctor. M y son the doctor sent me
a letter. I n this case, the speake r ha s more tha n one son, but th e restrictiv e phras e
the doctor tell s whic h so n sen t th e letter . Th e differenc e i s th e punctuation .
Nonrestrictive words , phrases, or clause s are se t off by commas; restrictive one s ar e
not. Her e i s another example : Th e girl standing in the garden waved t o him. Yo u
can identif y th e gir l only by knowing that sh e was standing in th e garden . Th e girl,
standing in the garden, waved t o him. I t happens that the girl who waved was standing
in the garden , a bit of interesting bu t nonessentia l information.
In correc t application , th e us e o f the relative pronouns which o r that depend s
upon whether the claus e tha t is introduced i s restrictive or nonrestrictive . Which i s
the nonrestrictiv e relativ e pronoun ; that i s restrictive . Th e automobile that wa s
speeding wa s completely destroyed. T o kno w which automobil e wa s destroyed , w e
must know that it was speeding. Th e automobile, which was speeding, wa s completely
destroyed. Th e importan t thing that the sentence tell s us is that the automobile-no
question whic h one-was destroyed; a s it happens , it was speeding. I n our reading ,
we gain the sens e of whether a phrase or clause is nonrestrictive or restrictive b y the
use or omissio n o f the comm a or commas . Whe n ther e ar e relativ e pronouns , th e
use of which or that fortifies tha t sens e o f restrictiveness o r nonrestrictiveness .
In scientific writing, most phrases or clauses introduced by relative pronouns are
restrictive; suc h phrase s shoul d b e introduce d b y that without a preceding comma .
When a n occasional nonrestrictive phras e or clause is used, it should be introduce d
by which, proceeded b y a comma. T o improve one's writing, an author should engage
in a "whic h hunt," replacing restrictive whichs with thats.
This seem s lik e a simpl e enoug h rul e t o follow , an d i n scientifi c writing th e
concept o f restrictive o r nonrestrictiv e can often conve y important information that
an autho r migh t want th e reade r t o comprehend . Judgin g by published technica l
articles, however , man y authors an d editor s ar e unawar e of th e rule . I t i s onl y
consistently followe d in the best-edited , non-technica l magazines or books. Appar ently man y writers hav e th e mistake n idea tha t which i s more forma l tha n that an d
thus shoul d be use d i n th e mos t formal o r technica l writing. Th e us e of thes e two
relative pronoun s ha s littl e o r nothin g t o d o wit h formality ; i t i s al l a matte r o f
restrictiveness o r nonrestrictiveness . Th e recommendation is to follo w th e rul e but

Some Suggestions About Scientific Writing 17

not t o get too upse t when others fai l t o follow it. Perhaps , as the years go by, it will
be adhered t o by more an d more author s an d editors .
That an d wh o provid e anothe r pai r o f relativ e pronouns . Again , th e rul e i s
simple, and agai n it is often broken. Wh o should be used with reference t o people;
that i s used fo r everythin g else. A claus e beginning with who (o r whom if it i s th e
object of a sentence or a preposition) ca n be restrictive or nonrestrictive; us e of the
comma tell s when it i s nonrestrictive .
Nonrestrictive introductory phrases o r phrases a t the end of a sentence should
be set of f from th e rest of the sentenc e wit h a comma in formal writing. Usag e has
led to the omission of this comma when the phrases are short, but more frequent use
of a comma to set off introductory or final phrases would lead to clearer writing. A s
reading experience demonstrates, th e sligh t break or pause indicated b y the comma
often contribute s t o eas e of understanding, as in this sentence.
The discussion o f how to punctuate nonrestrictive phrases o r clauses brings up
a logica l rul e tha t i s often violate d b y modern writers: Put a pair of commas, or
none, between subject and verb, or verb and object, or subjective complement. T o
say it anothe r way : A subjec t an d it s ver b shoul d neve r b e separate d b y a singl e
comma (unless the comma occurs between coordinate adjectives). I f subject and verb
are separate d b y a nonrestrictiv e phras e or clause, there mus t be two commas that
surround th e phras e o r clause . Th e rul e is broken, especiall y i n technica l writing,
because the subject may be modified with so many words and phrases that the author
feels the reader will run out of breath by the time he or she gets to the verb; the rest
afforded b y a comma seems t o b e in order. Bu t this comma will be very distracting
to a reader who is really paying attention. Th e reader i s anticipating the action, th e
verb, an d i s confuse d b y bein g tol d t o paus e jus t befor e gettin g ther e whe n n o
nonrestrictive (parenthetical ) materia l justifie s th e pause . Th e sample that ha d a
large, green leaf attached to the brown stem with an expanded petiole base but virtually
no thickened cuticle or acute lobe was chosen for th e herbarium. Some author s might
be tempte d t o pu t a comm a befor e th e was. O f course , th e sentenc e woul d b e
improved b y recasting i t a s two sentences .
B. Parenthetical Phrases or Clauses. Nonrestrictiv e phrases or clauses (as just
described) ar e parenthetical, which is to say that they contribute important informa tion but are not essential t o understand the rest of the sentence or to its grammatical
structure. I f the sentence is correctly constructed, the parenthetical phrase or clause
can be removed (alon g with the punctuation that sets it apart, the parentheses i n this
case), and what remains will still be a grammatically correct sentence . Parenthetica l
phrases ca n be punctuated i n four ways: wit h commas (as we have been discussing) ,
with parentheses2 (round brackets), with brackets (square brackets), and with dashes.
The choic e belong s t o th e author , bu t th e choic e ca n conve y an author' s feelin g
about th e parenthetica l material . I f commas ar e used , th e informatio n i s closel y
related t o th e sentence , almost restrictive , we might say.
2
In Britain and other United Kingdo m countries, the term parentheses is generally applied simply
to portions of text that are parenthetica l while bracket is a generic term for all parenthetical symbols:
round brackets ( ) , square brackets [ ] , curly brackets{ } , and angle d brackets< > .

174 Appendices:

Presenting Scientific Data

If parentheses (round brackets) are used , the materia l is more of a side issue .
It is important (or it would not be included at all), but it is not as closely related t o
the res t o f th e sentenc e a s would be th e cas e whe n commas are used . Use d to o
often, parentheses ca n be distracting, always confronting the reader with extraneous
information tha t may seem beside th e point. Thi s feeling can often be changed just
by changin g som e parenthese s t o commas . I f th e informatio n reall y i s almos t
unrelated t o th e res t o f th e sentence , however , i t shoul d be in parentheses , an d a
long sentence can sometimes b e made easier t o understan d by placing some of th e
material i n parentheses. I n a sense , thi s removes it fro m th e sentenc e althoug h it
might remai n exactly where it was. (Th e material might also be moved somewhere
else.)
Brackets [square brackets] are ofte n reserve d i n forma l writing for comment s
inserted by an editor. Th e editor can be the author if he or she is quoting someone
else but needs to insert an explanation or comment in the quoted material. Bracket s
can also be used as parentheses within parentheses: Evans and an assistant (Gillespie,
who made her own study of a flightless bird [the kiwi] in Australia) spent several
difficult months in the field.
Dashes are much less formal and should seldom if ever be used in technical writing-unless the autho r feels justified i n adding the stron g emphasis provided by use
of the dash or dashes-and if the author is confident that the editor will not remov e
the dashes ! I f comma s ar e include d i n a phras e se t of f with dashes, th e dashe s
become essential . (Th e exclamatio n poin t i s als o seldo m use d i n technica l writ ing-for th e sam e reasons dashe s are seldom used!! [ I tend t o overus e both!] )
Two point s abou t parenthetica l materia l nee d t o b e noted : First , sinc e al l
parenthetical material is by definition nonrestrictive, parenthetical phrases (regardless
of how they are punctuated) that are introduced by a relative pronoun should always
use which (o r wh o o r whom) instea d o f that. Second , sinc e i t shoul d alway s b e
possible t o remov e a parenthetical phras e or clause without affecting th e structur e
of wha t i s left , a n autho r mus t neve r us e doubl e commas , (lik e this) , aroun d
parentheses o r brackets .
Where i s the perio d place d i n relatio n t o material in parentheses o r brackets ?
If th e materia l i n parenthese s come s a t th e en d o f a sentenc e an d i s itsel f a n
incomplete sentenc e (sentenc e fragment) , th e perio d i s place d outsid e o f th e
parentheses (lik e this) . I f the materia l i n th e parenthese s come s a t th e en d o f a
sentence but b y itself forms a complete sentence, then such a parenthetical sentence
should b e se t withi n its ow n parenthesis . (I n suc h a case , th e firs t lette r o f th e
parenthetical sentenc e should be capitalized, and a period shoul d be placed a t th e
end o f th e sentenc e an d befor e th e las t parenthesis , lik e this. ) Ther e i s n o rul e
saying tha t parenthetica l materia l must be included in some other sentence ; it can
and often should stand on it s own, as in the example . Ther e i s also no rule saying
that parenthetical material cannot form a complete sentence within a sentence (it can
be distracting , as here, so i t i s well to avoi d the practic e when possible), bu t i f th e
complete-sentence, parenthetica l material can be placed after it s "parent sentence,"
it migh t just as well be give n a lif e o f it s own, cut of f from it s parent . Whe n it i s
included within another sentence, i t is not punctuate d as an independent sentence .

Some Suggestions About Scientific Writing 17

4. VERB S

A. Plural an d singula r verbs . Plura l verbs must be use d with plural subjects ,
singular verbs with singular subjects. Tha t is , a verb must agree with its subject i n
person an d number .
A subject consisting of two or more singular nouns connected with and is plural:
One nou n an d anothe r nou n mak e a plura l subject . I f tw o singula r noun s ar e
connected by or, the subjec t is singular (but plural if the nou n closest t o th e verb is
plural).
A singula r subject followe d b y a modifyin g prepositiona l o r othe r phras e tha t
contains plura l nouns or more than one singular noun is nevertheless singular (as in
this sentence and the on e that begin s the previous paragraph).
Some nouns taken from language s other than English form thei r plurals in ways
that ar e no t alway s familiar; watch for these (datum an d data, medium an d media,
etc.; se e Section 8) .
B. Ver b tense. Ver b tens e should be consistent. I t is usually logical to use th e
past tens e i n describin g methods , materials , an d result s i n a scientifi c paper: W e
found that applied LA A strongly promoted elongation o f intact pea plants. Th e
experiments were done in the past, and it is conceivable that they might give different
results i f repeate d (i f al l determinin g condition s ar e no t know n or understood) .
Hence, th e hones t wa y to describ e the m i s to us e the pas t tense . Avoi d changin g
tense i n th e middl e o f a descriptio n o f method s o r results , usuall y i n a singl e
paragraph. Publishe d result s ma y be described wit h the presen t tense : Yang e t al.
(1993) showed that a continuous supply of auxin enhances stem elongation in intact
plants.
C. Participles. Englis h form s a present participle by adding -ing to the infinitive
of th e verb . Thi s i s combined wit h som e for m o f th e ver b to b e to emphasiz e a n
action tha t i s occurring (or tha t was occurring or has been occurring or will be occurring). Thi s i s such a n importan t par t o f the Englis h languag e that nativ e speakers
virtually neve r use it incorrectly , but it is ofte n difficul t for writer s whos e nativ e
tongue i s not English . Th e tendenc y is to us e this verb form to o often , when i t is
not needed . Eve n nativ e speakers ca n frequently tighte n their writing by changing
to th e simpl e form s (i t occurs, occurred, has occurred, will occur, for example) .
A special problem i s the dangling participle, whic h is a participle that cannot b e
connected immediatel y an d unmistakabl y with th e word(s ) t o whic h i t refers .
Because the antecedent of the verb is often left to the reader's imagination, sentences
with dangling participles ca n often b e quite ludicrous: Coming into the greenhouse,
the large skunk cabbage gave off a n overwhelming stench. (Wh o entere d th e green house? Th e writer or th e skunk cabbage?)
English usuall y forms a past participle b y adding -ed to the verb and combining
it with another auxiliar y verb, usuall y a for m o f to have. Thi s verb for m indicate s
that a n actio n wa s begun i n th e pas t relativ e t o th e tim e being referre d t o bu t i s
completed i n that time being referred to, which can be the present, the past, or even
the future : I have measured. Sh e ha s measured. H e ha d measured. Yo u will have
measured. Th e pas t participl e can usuall y b e replaced wit h a simple past o r futur e
form: / measured. Sh e measured. H e measured. Yo u will measure. Nevertheless , it

176 Appendices:

Presenting Scientific Data

is sometimes appropriate t o us e the past-participl e form . Thi s for m usuall y causes

no trouble for native speakers unless the past participle is irregular or differs from
the simple past , also formed by adding -ed. Example s o f irregular form s include: /
give it. I gave it. I have given it. I do it. I did it. I have done it. She comes home.
She came home. Sh e has come home. H e goes. H e went. H e has gone. Consul t a
dictionary i f in doubt abou t th e pas t participle .
D. Passive voice. Author s o f journa l article s ofte n us e th e passive voice by
combining th e pas t participl e o f a transitiv e verb with any form o f the ver b t o be:
The water potential was measured. Thi s constructio n omit s th e perpetrato r o f th e
action, th e on e wh o did th e measurin g in th e example . Th e first-perso n persona l
pronouns ar e avoide d s o the writing may seem mor e objective o r "scientific. " (Di d
you catc h th e passiv e voic e i n tha t sentence? ) Actually , us e o f th e passiv e voic e
allows one t o avoi d responsibilit y fo r one's actions, to divorce th e worker fro m hi s
or he r work . Tr y t o us e th e active voice: Instea d o f saying, the leaves were treated
(passive voice), say we treated th e leaves (activ e voice) .

5. SOME FURTHER NOTES ABOUT PUNCTUATION

The discussio n s o far has followed my "logical" approach t o the structur e o f th e

English languag e with its simple-to-compound sentences , restrictiv e an d nonrestric tive modifier s (word s an d phrases) , an d specia l ver b forms . Thi s discussio n ha s
placed man y items of punctuation int o tha t "logical " context, but severa l important
although ofte n arbitrar y rule s di d no t fi t int o tha t evolvin g but somewha t limited
discussion. Thu s th e followin g brie f outlin e (based o n Pinckert , 1986 ) summarizes
the importan t rule s o f punctuation ; item s alread y discusse d i n som e detai l ar e
presented i n small type :

A. Avoi d sentence fragments (incomplete sentences terminated with a period), and do not use
a comma when a perio d (o r semicolon) shoul d be used; that is, avoid the comm a fault .

B. En d question s wit h a question mark, but d o not us e a question mar k with

indirectly quoted questions: Sh e asked wh o Bob was.
C. Reserv e th e exclamation point for tru e exclamation s o r command s (Drop
dead!), which are seldo m use d i n technical writing ! Whe n the exclamatio n point is
used fo r emphasis , a reade r soo n get s tire d o f suc h exclamator y writing! It s us e
suggests sentence failure! Avoi d using the exclamation point just for emphasis! (Bu t
its use in the last sentenc e is valid because that is a direct command, an imperative.)
D. Use th e semicolon:
betwee n independen t clause s not connecte d with a coordinatin g conjunction .

t o separat e comple x items in series whe n each item itself consists of a serie s
of items : These ar e important plant hormones: auxin, typically a stem growth
promoter; the gibberellins, also promoters of stem growth; cytokinins, stimulators
of cell division; and ethylene and abscisic acid, sometimes called stress hormones.
E. Use commas:
- betwee n item s in a series, including before the and that precedes the last item :
The equipment included a camera, portrait lens, an d flash attachment. Thi s
serial comma, as i t i s called , i s ofte n omitte d b y moder n writers . Mos t
manuals still recommend its use.

Some Suggestions About Scientific Writing 177

- afte r man y adverbs, phrases , an d clauses tha t introduc e a sentence an d are
followed by a voice pause : T o avoid precipitation, it is essential to stir continually.
t o enclos e nonrestrictiv e phrases and clauses.
befor e a coordinating conjunction introducing an independen t clause.

t o se t of f direct addres s and othe r parenthetica l interrupters . "Carol, will you

fix breakfast?"
- t o introduce o r interrupt shor t quotations . "Not," sh e said, "on your life."
t o preven t misreading . "Let's talk about Prof. Jones, and good scientists." Suc h
style commas can be inserted t o emphasize an adverb (She snored, heavily.) o r
to giv e extra emphasis t o modifier s of equal importance .

F. Use th e dash t o indicat e an extende d paus e an d giv e emphasisbu t seldo m i f at al l i n

technical writing.
G. Use commas, parentheses, dashes, and brackets to se t of f parenthetical material.

H. Use ellipses ( a serie s o f thre e periods ) t o sho w tha t somethin g ha s bee n

omitted fro m a quotation. I f the ellipses come at th e end of the sentence, add th e
fourth period. Us e brackets [square brackets] to show that something was inserted by an editor.
I. Whe n use d a s a mar k of punctuatio n within a sentence , a colon mean s as
follows, a n example follows, here i s th e explanation o r list, or here i s what he o r sh e
said. Whateve r follow s a colo n refer s back to wha t immediately preceded it . I t is
logical t o us e a capital lette r afte r a colon i f a complete sentenc e follows , but many
editors wil l chang e th e capita l lette r t o lowe r case . I f a lis t follow s a colon , th e
words ar e no t capitalize d unles s the y ar e prope r noun s o r mus t otherwis e b e
capitalized. Th e colon i s also use d in several other ways in scientific writing as ofte n
noted i n this boo k (e.g. , t o sho w the proportion s o f components i n a mixture used
in chromatography : water:aceti c acid:butano l 5:1:3).
J. The apostrophe i s use d t o for m contraction s (don't worry) an d possessive s
(Joule's book; originally a contraction o f Joule his book). I f the possessiv e i s a plural
ending i n s o r a z sound , th e apostroph e come s afte r th e s ; some manual s suggest
adding another s , but becaus e ther e is so much disagreement, thi s can be a matter
of persona l choice . On e suggestio n i s tha t th e s should b e adde d i f i t i s easil y
pronounced: Descartes's essays (becaus e neithe r s i s pronounced i n Descartes).
Usually the extr a s is not adde d in technical writing. Indeed , contractions ar e use d
less ofte n i n technica l writin g than in less forma l writing.
K. Quotation marks are use d around direct quotations but not aroun d indirect
quotations. I f the quotation extend s over more than one paragraph, quotation marks
begin each paragrap h but end only the last paragraph. Quotatio n marks are used to
set apart a word or phrase that is used in some sense other than the usually accepted
one, bu t thi s i s easil y overdon e an d shoul d be avoide d a s muc h a s possible . Pu t
commas an d period s insid e closin g quotatio n marks ; pu t semicolon s an d colon s
outside. Othe r punctuatio n (questions marks and exclamation points) should be put
inside the closin g quotatio n mark s only when the punctuation is actually part of the
matter being quoted: H e asked: "Should th e catalyst b e added?" I s it really, a s written,
"larger than the primary leaf?
L. Hyphens are used :
i

n compoun d adjectives.

178 Appendices:

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-t

o form compound nouns , although it is often acceptable t o combine the two

nouns into one word without the hyphen: mailman instead of mail-man, for
example.
t o divid e word s a t th e en d o f th e line , but rule s fo r hyphenatio n are to o
involved fo r summary here. Whe n in doubt, check in a good dictionaryo r
trust you r word processor' s hyphenation feature.
M. The slash is used in several technical applications, as noted elsewhere i n this
booklet (e.g. , i n fractions : 1/20) . I t i s als o sometime s use d i n th e expressio n
. ..and/or. . . in scientific writing. Som e editors do not like this and will suggest . . .or . .
both.
N. Underlining i s use d i n plac e o f italics i n type-writte n o r hand-writte n
material. Ther e are instance s i n scientific writing where italics or underlinin g must
be used , as i n scientifi c names o f organism s (se e Chapte r 2). Itali c typ e has bee n
used appropriatel y in thi s appendi x to se t of f examples. Italic s can also b e used fo r
emphasis, but thi s is discouraged i n technical writing. Foreig n words are underline d
or italicized , especiall y if they are unfamiliar .
6. ABBREVIATIONS
Science speaks its own language with specialized words that we must all learn in
our respectiv e fields . Perhap s tha t i s why we always see m t o wan t t o inven t eve n
more term s b y constructin g abbreviations . O f cours e man y abbreviation s o r
acronyms are recognized; the y are already a part of the language of plant physiology:
ATP, DNA , IAA, 2,4-D , NADP, SDP, and many more. Journal s often publis h list s
of suc h accepte d abbreviation s an d expec t author s t o us e them . (Not e table s i n
Chapter 10. ) Bu t i t i s an impositio n t o expec t a reade r t o lear n a handfu l o f new
abbreviations in orde r t o rea d a paper. A few abbreviations (fou r o r five? ) ma y be
justified to avoid the constant repetition o f some complex terms or phrases, but most
authors will achieve a more sympatheti c audienc e i f new abbreviations are kep t t o
a bare minimum . I t i s also helpful i f the newl y introduced abbreviations are easil y
distinguished from eac h other. Consider , for example, a series o f treatments with or
without auxin, light or dark, at morning, noon, or night: ALM, ALN, ALNi, NALN,
NALN, NALNi, ADM, ADN, ADNi, NADM, NADN, an d NADNi. Pitt y the reader !
7. UNNECESSAR Y WORDS
We hav e a grea t tendenc y t o expan d ou r writin g b y using words tha t ar e no t
needed or tha t hav e shorter an d more concise counterpart s (Heiche l et. al. , 1990).
1. Som e words can simpl y be droppe d (e.g. , simply i n thi s case) :
prior histor y (all histor y is prior)
careful study , careful examinatio n (how else would you d o it? )
very (thi s word only contributes something in certain negative constructions:
It isn't very effective.)
it i s shown that (seldo m needed)
it i s a fact that (seldo m needed)
it is emphasized that (seldo m needed)
it is known that (seldo m needed)

Some Suggestions About Scientific Writing 17

2. Som e word s ca n be replaced b y more descriptive an d concise terms :

Instead of:
Use:
in th e absenc e of withou
t
higher i n comparison t o mor
e than
was found t o b e wa
s
in the even t tha t i
f
small number o f fe
w
was variable varie
d
additional added
, more, or othe r
approximately abou
t
at th e presen t tim e no
w
at tha t poin t i n time a
t tha t time
establish sho
w
identify find
, name , or sho w
in a timely manner promptl
y
necessitate caus
e or nee d
appears t o be seem
s (als o overused , ofte n t o avoi d
making a factual statement: I t seems to
be raining. Better : I t i s raining.)
8. WORDS WITH SPECIAL PROBLEMS
Some words are use d incorrectly so often i n scientific writing that some reader s
may never lear n to use them correctly when they become authors. Som e of these ar e
homonyms, which are pair s o r groups of words with identical or similar sounds bu t
different spelling s an d meanings . Th e followin g list include s som e o f the proble m
words includin g a fe w homonyms. Hopefully , th e lis t include s many of th e word s
that pos e special problem s fo r speakers whos e native tongue i s not English . I f you
are i n doub t abou t th e correc t meanin g an d spellin g of a word, check with a goo d
dictionary o r i n a manual that list s such words (e.g., CBE Style Manual Committee ,
1994; d e Mell o Vianna , 1977 ; Strun k and White, 1979) .
accept, except Accept mean s t o receiv e o r admit , to regar d a s right or true , or t o
bear u p under . Except mean s t o leave out , exclude, or excuse .
affect, effect Eac h ca n be used either as a verb or as a noun, but mos t of the time ,
affect i s used as a verb (t o influenc e or caus e a change in) an d effec t i s used as a
noun ( a resul t o r consequenc e o f an action). A s a noun, affect i s used onl y as a
technical ter m i n psychology. Effect i s used as a verb in the sens e o f to cause (t o
bring about or make). Technica l writer s like to soun d scientifi c by using effect a s
a verb, bu t i t often lead s t o confusion , and i t i s easy to sa y cause instead. Eff
as a verb sounds affected . Furthermore , affect i s a rather weak verb. I t i s better
to tel l how somethin g i s affected: increased o r decreased, for example .
as Whe n a s is a preposition, meanin g in the role , capacity, or functio n o f (see like,
as below) , i t i s alway s followe d b y a nou n o r pronou n i n th e objectiv e case .
Otherwise, th e cas e ca n depen d o n th e sens e tha t i s desired : Yo u need he r as
much as I (i.e. , as much as I need her). Yo u need her as much as me (i.e. , as much
as you need me). Ambiguit y can result when as i s used as a conjunction instea d

180 Appendices:

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of since o r because: Sh e did not hear the bell as she was on th e terrace. Di d sh e
fail t o hea r th e bel l because sh e wa s on th e terrac e o r while sh e wa s o n th e
terrace?
as . ..as, s o . . . as I n positive comparisons , as...as is the construction tha t is used: a
tough as nails. I n negative comparisons, eithe r as . . a.s or so . . .
as can be used: ...no
as (o r so) skilled a s his technician. Th e firs t as should not b e omitted in positiv e
comparisons. (Don' t say : Th e sound wa s clear a s a bell. Say : ...as clear a s a
bell.)
can, may Ca n i s used t o indicat e abilit y to d o something ; may, to ask , grant , or
deny permission t o d o it . Thi s distinctio n should be followed in formal writing.
datum, data Traditionally , especially i n technical writing, datum ha s bee n consid ered singula r ( a fac t o r singl e ite m o f information ; a singl e number ) an d data
plural, bu t popula r usag e ha s almos t eliminate d th e singula r datum fro m th e
language, an d data i s almos t universall y used a s a singula r noun ( a collective :
information organized for analysis). A few of us continue to say . . . thesedata are .
but a s our generatio n die s off, data wil l no doubt be used only as a singular noun.
(I fin d thi s regrettable! )
due to Thi s expression i s often overused in technical writing. I t is correct whe n it
is used a s a predicat e adjectiv e that follow s som e for m o f th e ver b to b e an d i n
the sens e o f caused b y or attributable to: Th e broken centrifuge wa s largely du e t o
faulty maintenance. Thi s coul d b e replace d b y ...was caused largely by... I t i s
somewhat les s correct , although commonly used, i n th e sens e o f because of , o n
account of , owing to, o r through: Th e centrifuge failed du e t o faulty maintenance.
In forma l writing , it would be better t o say . . .because o f faultyits, it's It s is the possesiv e for m o f the pronoun it, but used in this case without th e
apostrophe: ...the graph with it s curves... It's i s a contraction o f i t is an d i t has:
It's no t new; it's been done before. (Overus e of the contraction s sometimes makes
the writing seem to o informal. )
et al. Thi s is properly use d in bibliographies t o mea n and others. Not e th e perio d
after th e secon d element . Becaus e i t i s Latin , som e editor s insis t tha t i t b e
italicized (o r underlined) .
information I n som e language s othe r tha n English , th e comparabl e ter m fo r
information ma y be a plural (e.g., French, Spanish, Russian) or may be used a s a
plural (e.g. , German) . I t i s never correct i n scientific English to us e information
as a plural: informations.
lay, lie La y (t o put , place , o r prepare ) alway s take s a direct objec t (lay it down);
that is, lay is a transitive verb. Li e (t o recline or be situated) never does; that is,
lie is an intransitive verb. Bu t th e pas t tense and past participle of lay is laid, and
the pas t tens e o f li e i s lay, the pas t participl e i s lain. Thi s certainl y leads t o
confusion. Si t an d set are equall y troublesome.
like, as Like an d a s are correctl y use d as prepositions expressing different senses .
In this case, like indicates resemblance to the object mentioned: H e looks like his
brother. I t ca n always be replaced b y similar to. A s indicate s a role, capacity , or
function: H e serves as Department Head. (On e could say serves like a Department
Head, referrin g to someone wh o is not a Department Head bu t serve s like one. )

Some Suggestions About Scientific Writing 18

It is less desirable t o us e like as a conjunction t o introduc e a clause, althoug h i n

formal writing this can be done i f the clause is an elliptical one in which the verb
is not expressed : ...looks like a good day for science. Suc h a clause is not accept able in forma l writing if th e ver b i s expressed: ...looks like i t will b e a good day.
In such a case, like must be replace d b y as, as if, o r as though: ...looks as if i t will
be a good day. Fea r o f using like incorrectly ma y tempt a writer to us e as when
like, used a s a preposition, is called for : Sh e acted like (not as) an idiot.
may, might Originally , might was the pas t tens e o f may, bu t no w both verb s ar e
used a s subjunctives expressin g possibilit y o r permissio n i n presen t an d futur e
time. The y diffe r i n intensity rathe r tha n in time. I n the sense of either possibili ty or permission , ma y i s stronger tha n might: H e may go makes a stronge r cas e
for hi s havin g permissio n t o g o o r th e likelihoo d tha t h e wil l go than doe s h e
might go. (Se e can , may ; can is used t o expres s abilit y to d o something. )
medium, media Lik e datum/data, medium i s the singula r form o f media, but media
is often use d as a singular collectiv e t o refer to the means of mass communicatio n
taken a s a whole. I n careful writing, media as a subject should alway s be treate d
as a plural , an d eac h individua l mean s o f mas s communicatio n shoul d b e
expressed a s th e singula r medium: Television i s an influential medium. Together,
television, radio, newspapers, an d periodicals make up the media. Scientifi c writers
should b e carefu l t o distinguis h betwee n thes e tw o forms of the nou n when they
are use d i n their origina l sens e of a surrounding or pervading substance i n which
bodies exist or move: th e environment. A n object ca n only be surrounded by one
medium, bu t differen t object s ca n be in different media. Bacteriologist s and plant
pathologists ofte n incorrectl y us e media t o refe r t o a singl e sterilize d nutritiv e
substance fo r cultivatin g bacteri a o r fungi . Medium i s also use d i n othe r sense s
including tha t o f an intervenin g thin g through which a forc e act s o r a n effec t i s
produced, o r a perso n throug h who m communication s com e fro m th e dead .
Medium a s an adjectiv e als o refer s to somethin g intermediate, a middle state o r
degree.
nor No r is a conjunction used to express continuin g negation; often it is paired with
neither: He was neither for the idea nor against it. She had no experience as a
physiologist nor did the subject interest her. Th e word or can be used instead o f no r
when th e elements are within a single independent clause , and it is clear tha t th e
negative sens e carrie s ove r t o th e elemen t tha t i s introduced: H e was not for o r
against the idea. She had no experience or interest in physiology.
percent, percentage Percent (no longe r writte n per cent) i s specific and follow s a
number or numeral: 1 8 percent. Percentage is nonspecific and should not b e used
with a number : A small percentage died o n day three. (Us e a spac e betwee n a
numeral an d th e percen t sign : 1 8 %. Se e Chapter 1.)
precede, proceed Precede mean s t o com e befor e i n time , order , o r ran k o r t o
introduce. Proceed mean s t o g o forward o r onward , to undertak e a n action .
principal, principle Principal ha s severa l meaning s bot h a s a nou n an d a s a n
adjective. A s a n adjective , it mean s leadin g or chief . A s a noun , it mean s th e
leader or person i n charge or the capital or main body of an estate o r a sum owed

182 Appendices:

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as a debt: Th e principal o f th e school is my pal Principle i s used only as a nou n

meaning a basic truth, rul e o f human conduct, o r fundamenta l law.
proved, proven Bot h ar e pas t participle s o f th e ver b t o prove, bu t proved i s
preferred: H e ha d proved his point. Proven i s more commo n when use d a s a n
attributive adjectiv e befor e a noun: a proven record. I t is also the for m use d in
the phrase not proven. (Becaus e o f variabilities cause d by chance, and because of
possible alternativ e explanations , scientists mus t be wary of saying that something
is proven. W e often striv e to disprove a hypothesis.)
respectively Th e word means singl y in the orde r designated . I n scientific writing,
it i s often quit e clea r tha t th e dat a presente d i n one brie f lis t ar e give n i n th e
same orde r a s th e name s o r othe r informatio n i n anothe r list , an d th e wor d
respectively insult s the intelligenc e o f the reader : Plants in the dark and in the light
were etiolated an d green, respectively. Wit h few exceptions, i t i s best t o writ e th e
sentence so that respectively i s not needed : Proline concentrations were 0.5 molL-1
for sample one, 1.32 mol.L-1 for sample two, and 3.56 mol.L-1 for sample three.
Don't write : Proline concentrations were 0.5, 1.32, and 3.56 mol.L-1 for samples
1, 2 , an d 3 , respectively. Th e secon d wa y ma y be shorter , bu t i t i s inherentl y
confusing.
shall, will I n the mos t forma l writing, use of these words was governed by a series
of comple x rule s tha t woul d b e understoo d b y few modern readers . I n moder n
usage, both are used to indicate futurit y a s well as determination, compulsion , o r
obligation, bu t will is becoming more common in construction of the future tens e
(opposite o f the ol d rule) : Tomorrow, we will set up the experiment. Becaus e this
can b e ambiguous , i t i s a goo d ide a t o us e som e othe r constructio n whe n
determination i s the sense to be expressed: Tomorrow, we must (have to , certainly
will) se t u p th e experiment. Shall is sometime s use d t o emphasiz e resolve : W e
shall overcome. Bu t shall is slowly disappearing fro m th e language .
since, becaus e Since is often used in the sens e of because, which is still a good use .
Some editors , however , migh t insis t tha t it s us e b e restricte d t o th e followin g
sense: ...the time that has elapsed since some event.
so Whe n the conjunction so introduces a clause that gives the purpose of, or reason
for, a n actio n state d earlier , it i s usually followed by that and no t precede d b y a
comma: W e changed the solutions so that toxins would not build up . I n that usage,
it is a subordinating conjunction. S o is used alone as a coordinating conjunction
when th e claus e tha t i t introduce s state s a resul t or consequenc e o f something
preceding: W e had t o finish, so w e worked late.
so-called Thi s hyphenate d adjectiv e precede s a nou n tha t i s no t enclose d i n
quotation marks : Th e dark period wa s interrupted b y a so-called night-break.
Sometimes i t i s used sarcastically : ...a so-called leader.
then, than Mos t commonly , then means at that time or refers to order o r position :
First the seed germinates, then the seedling develops. Than i s used in comparisons:
Cultivar A is more drought hardy than cultivar B.
that, which , who , who m Se e th e discussio n i n Sectio n 3. A o n Restrictive an d
Nonrestrictive Phrases and Clauses. That (o r wh o o r whom) shoul d be use d t o
introduce restrictive phrases and clauses. Which, who, or whom (especially which),

Some Suggestions About Scientific Writing 18

preceded b y a comma , are use d t o introduc e nonrestrictive phrases an d clauses.

Who o r whom (dependin g o n case ) shoul d b e use d whe n th e referenc e i s t o
people. Which i s alway s used whe n i t i s precede d b y that a s a demonstrativ e
pronoun: W e often long for that which is impossible.
thus, thusly Thus is always the choice. Bot h are adverbs, and thusly never need s t o
be used .
toward, towards Eithe r form i s acceptable, bu t toward i s most common in moder n
writing (especiall y i n the Unite d States) .
unique Somethin g unique is in a class by itself withou t an equal. A s an adjective,
it is an absolute. Therefor e expression s o f comparison shoul d be avoided: most
unique, rather unique, etc. Bu t some modifiers are acceptable an d logical: nearly
unique, more (o r most) nearly unique. Th e same i s true o f such othe r absolutes
as perfect or dead.
were Were ca n b e use d a s the pas t subjunctiv e mood o f the ver b to be to expres s
conditions tha t are clearly hypothetical or contrary to fact: I f i t were only possible.
Such statement s ma y express a wish: / wish that the data were complete. Ofte n
they ar e precede d b y if: I f ou r budget were only larger. Informally , was i s often
used instea d o f were, and sometime s were i s incorrectl y use d instea d o f wa s i n
indirect questions o r when the conditional statemen t is not really contrary to fact:
...if the report was true, changes would be necessary. They asked if he was agreeable.
9. SOME SUGGESTIONS ABOUT FORMAT AND WORD PROCESSORS

Most journals have printed instructions with many details on suitable format fo r
manuscripts. Th e CBE Style Manual Committee (1994) als o ha s a n extensiv e
discussion of format. Author s mus t carefully study such instructions before writing
and certainl y befor e submitting a manuscript to a particular journal. I f the forma t
does not follow that of the journal, reviewers may assume (sometimes correctly) that
the manuscript was submitted to another journal and rejected. A t the very least, an
incorrect forma t tells the editor that the authors were too careless to bother checking
such details before submitting a manuscriptand thus may be careless in carrying out
the scientific study that is being described. Incorrec t formatting also fails to consider
the editors and reviewers who must read th e manuscript .
A fe w suggestions, mostl y my own personal preferences , com e t o mind : Manu scripts submitte d fo r publicatio n shoul d always b e doubl e spaced ; thi s include s
captions, footnotes , quotations , everything. Th e double spacing is to help reviewers,
copy editors, an d compositors edi t an d otherwise mark the text . (Becaus e it is now
so eas y t o reproduc e an d eve n t o rewor k a manuscript , it i s quit e i n orde r fo r
reviewers t o writ e directl y o n th e manuscript. ) Title s shoul d b e bot h brie f an d
descriptive. Becaus e th e titl e i s usually the firs t thin g a potentia l reade r sees , it s
importance can' t b e overestimated. Nex t in importance is the abstract , which must
also b e as brief as possible (s o as not t o discourage a potential reader) whil e at th e
same tim e conveyin g all th e ke y point s includin g reason s fo r th e stud y an d th e
important conclusions . Al l plan t materia l must b e accuratel y name d both i n th e
abstract an d i n th e section s describin g method s an d material s (see Chapte r 2) .
Figures an d table s conve y th e actua l dat a produce d b y th e stud y tha t i s bein g

184 Appendices:

Presenting Scientific Data

reported, s o i t i s extremel y importan t t o mak e the m a s complet e an d eas y t o

understand a s possible. (Se e th e followin g section fo r many tip s tha t ca n appl y t o
published paper s a s wel l a s t o ora l an d poste r presentations. ) I t i s a commo n
practice t o us e different symbol s and lines fo r different treatment s shown in figure s
and the n t o defin e thes e symbol s an d line s i n th e caption . Sometime s thi s i s
necessary because there may not be room on the figure to label the curves, but if the
curves ca n b e labelle d o n th e figure , i t become s muc h easie r fo r a reade r t o
understand wha t i s bein g presented . Wh y shoul d a n autho r pla y game s wit h
potential readers? Wh y should a reader have to learn a code to understand a figure?
In an y case , th e captio n shoul d presen t enoug h informatio n t o mak e th e figur e
understandable withou t having to rea d the text. Suc h details as dates an d statistica l
treatments shoul d als o be included.
The preparatio n o f manuscript s ha s bee n greatl y aide d b y word processing
programs tha t reduc e th e tim e require d t o creat e an d especially t o revis e a manuscript and, most important, make it possible virtually to eliminate typographica l and
other errors. Th e spell checker is especially valuable, and grammar checkers can also
help, especiall y thos e authors whose native language is not English. Suc h programs
are now used by almost everyon e world-wide, and many journals accept manuscripts
in electroni c for m o n a disc . Nevertheless , ther e ar e a fe w minor problem s o r
irritations tha t resul t fro m th e us e of word processing programs.
There is a tendency to put too much faith i n the word processor an d not to proof
the fina l manuscrip t before i t is submitted. Perhap s the mos t common errors aris e
from editin g the manuscrip t on the compute r screen. On e makes a change but may
forget t o remov e al l the materia l being replaced, for example. I t is still important t o
carefully proof the final document.
The word processor usuall y provides a capability not previously enjoyed by most
authors: th e abilit y t o justif y th e righ t margi n (i.e. , alig n it, a s i n thi s book) . I t
seems tha t fe w author s ar e abl e t o resis t th e temptatio n t o us e thi s capabilit y
although i t sometime s produce s som e of those mino r irritations. Althoug h we are
used t o reading type-set materia l with justified righ t margins, there are two reasons
why "amateur" justification of the righ t margin is not alway s a good idea :
First, man y authors ar e reluctan t t o hyphenat e (and it i s somewhat easier fo r a
typesetter at the printin g press to work from a manuscript that has no hyphenation).
The result with a justified righ t margin is that sometimes an exceptionally long word
will not quit e fit at th e en d o f the lin e and is automatically moved (wrapped) by the
word processor t o th e nex t line, leaving a large space that must be divided betwee n
the words that remain i n the line . Thi s produces a line with a few words separate d
by large spaces , lik e these , whic h can be distracting for a reader, wh o shoul d
be abl e t o concentrat e o n th e conten t o f the paper . Wha t does righ t justification
gain fo r the edito r an d th e reviewers ? I s understanding the pape r aided in any way
by a justified right margin?
Second, some printers (especially dot-matrix) are not capable of dividing the space
left a t th e en d o f a lin e int o smal l fraction s o f a millimete r an d proportionin g it
evenly among all the spaces between words and letters in the line. Instead , they work
only with whole spaces (columns) , and thi s means that space s between some words

Some Suggestions About Scientific Writing 18

are ofte n a t leas t on e spac e large r tha n space s betwee n othe r words . Thi s uneve n
spacing occurs especiall y wit h non-proportional font s (e.g. , Courier). Som e reader s
also fin d thi s distracting .
If an author must justify th e right margin of a manuscript (and it can make a good
initial impression) , grea t effor t shoul d b e expende d t o hyphenat e correctl y (a s
professional typesetter s have always done), an d only the bes t o f printers capable of
microjustification shoul d b e use d fo r th e fina l product . Otherwise , right-margin
justification gains nothing while it provides an added irritation that may not produc e
the desire d receptiv e attitude in a reviewer.
10. SUMMARY
A. The sentence.
1. Tw o ideas in a sentence connected with a coordinating conjunction may share
a commo n subjec t (o r sometime s a verb), in which case the y should not b e
separated with a comma or other punctuation: Th e sentence presents one idea
and adds another to fortify the first.
2. Independen t clause s connecte d b y a coordinatin g conjunctio n shoul d b e
separated b y a comma before the conjunction : Th e first clause has a subject
and verb, and the second clause also has both subject and verb.
3. Closel y related sentence s no t connected with a conjunction may be separated
by a semicolon; thi s ties th e idea s together i n a special way (as here). Sep arating suc h sentence s wit h a comm a i s calle d a comma fault o r comma
splice, th e habi t must be avoided. (Di d you notice th e example? )
4. Les s relate d sentences are separated by periods. Thes e two sentences provide
an example .
5. Whe n tw o ideas ar e relate d t o each other wit h a subordinating conjunction,
they shoul d b e separate d b y a comm a i f th e subordinat e phras e o r claus e
comes firs t i n th e sentenc e (when i s th e subordinatin g conjunction i n thi s
case); otherwise, n o comm a is needed.
6. An d i t i s acceptable t o begin a sentence wit h a coordinating conjunction (as
here; use d mor e a s a conjunctive adverb) althoug h this practice shoul d no t
be overdone .
B. Modifying words.
1. Compoun d adjective s ar e forme d b y connecting with a hyphen : tw o adjectives, an adjective and a noun, or tw o nouns: near-ultraviolet radiation, a tenwatt lamp, cell-wall structure. (Bu t omit the hyphe n with numerals followed
by unit symbols: 10 0 W lamp.)
2. I n technica l writing , i t i s importan t t o us e adverbia l form s whe n a verb ,
adverb, o r adjectiv e i s being modified: a n unusually concentrated solution.
3. Noun s used as direct o r indirect objects or followin g preposition s ar e always
in th e objectiv e case , which in English is only evident when expressed b y th e
personal pronoun s me, him, her, us, them, and whom (o r whomever).
4. Althoug h some editors migh t disagree, authors would do well to use personal
pronouns in writing technical articles for the scientific literature: W e homogenized th e tissue i n a buffer solution. Fo r on e thing , thi s avoid s us e o f th e

186 Appendices:

C.

D.

E.
F.
G.
H.

Presenting Scientific Data

passive voice : Th e tissue was homogenized. . . . (Passiv e voice i s appropriate

for Method s an d Material s sections. )
Modifying phrases and clauses.
1. A restrictive phras e or clause contains information that is essential t o understand som e par t o f th e sentenc e no t include d i n th e phras e o r clause ; a
nonrestrictive phras e o r claus e add s informatio n tha t i s no t essentia l t o
understand th e part s of the sentence no t include d in the phras e or clause .
Nonrestrictive phrase s or clauses ar e se t apart by commas, but restrictiv e
phrases o r clause s ar e not : Th e hypocotyl sections i n auxin solution curved
down. Th e hypocotyl sections, in auxin solution, curved down. (Th e autho r
determines th e restrictiveness. )
The relative pronoun that is used to introduce some restrictive phrases o r
clauses; it is not precede d b y a comma: This was the sample that we examined.
The relativ e pronou n which i s use d t o introduc e som e nonrestrictiv e
phrases o r clauses ; i t i s precede d b y a comma : W e examined this sample,
which [incidently ] nearly escaped us .
Introductory o r fina l nonrestrictiv e phrases o r clause s ar e se t of f with a
comma: I n preparation for th e experiment, w e collected th e necessary glassware.
The relative pronoun who (or whom) is used with reference to people; that
(or which) is used fo r everythin g else: Th e teacher wh o drew th e diagram. . . .
The plant that grew. . ..
Put tw o commas ( a pair), or none, between subject and verb, or verb and
object o r subjec t complement : Th e hypocotyl sections, in gibberellin solution,
curved upward. Not : Th e hypocotyl sections i n gibberellin solution, curved
upward.
2. Parenthetica l phrase s o r clauses can be surrounded by commas, parentheses,
brackets, or dashes. Th e choice i s up to the author. I f a phrase in parentheses comes a t the en d of a sentence, th e perio d goe s outside the parentheses .
(Complete sentences initiated with the firs t wor d capitalized an d terminated
with a period ca n also be include d in parentheses, lik e this.)
Verbs.
1. Verb s mus t agree i n number (singular or plural ) with their subjects.
2. B e consisten t i n ver b tenses ; us e th e pas t tens e t o describ e method s an d
results.
3. Us e the presen t an d past participle s correctly .
4. Wheneve r possibl e an d appropriate , us e th e activ e instea d o f th e passiv e
voice.
Further notes on punctuation. Thi s section contain s some rules not containe d
in the previou s discussion , which also often concer n punctuation . Becaus e th e
notes are alread y in a summary form, the y are no t repeate d i n this summary.
Abbreviations. Avoi d using too man y ne w abbreviations.
Unnecessary words. Tighte n you r writing b y dropping unnecessary words an d
phrases an d b y using simple, concise form s wheneve r possible.
Words with Special Problems. Chec k the lis t here or us e a good dictionary to
use these words correctly.

Some Suggestions About Scientific Writing 18

I. Suggestions about format and word processors. Chec k instruction s to author s

published by the journal t o which a manuscript i s to be submitted. Watc h fo r
errors tha t ten d t o appea r whe n on e use s a wor d processor ; proo f th e fina l
printed manuscrip t carefully. Us e a high quality printer and avoid justifying th e
right margi n unles s yo u us e a suitabl e proportiona l fon t an d us e prope r
hyphenation a t th e en d of lines tha t require it.
REFERENCES
Anonymous. 1993 . Th e Chicag o Manua l of Style, Fourteenth Edition . Th e Universit y of Chicago
Press, Chicag o and London ,
de Mello Vianna, Fernando. 1977 . Th e Written Word. Houghto n Mifflin Co. , Dictionary Division,
Two Park Street , Boston , M A 0210 7
CBE Styl e Manual Committee . 1994 . Scientifi c styl e an d format : th e CB E manua l for authors ,
editors, an d publishers . 6t h edition . Cambridg e University Press, Cambridge , New York. [Se e
also earlier edition s of CB E Styl e Manual.]
Heichel, G.H. , D.E . Kissel , C.W . Stuber , G.A . Peterson , J.L . Hatfield , R.G . Hoeft , R.J . Wagenet ,
T.J. Logan , W.A . Anderson , an d W.R . Luellen . 1990 . Becom e a Mor e Successfu l Author .
Journal o f the Soi l Science Societ y of America. 54[Sept-Oc t '90](5):iv-vii .
Pinckert, Robert C . 1986 . Pinckert' s Practica l Grammar. Writer' s Diges t Books , 993 3 Alliance
Road, Cincinnati , Ohio 4524 2
Strunk, William , Jr . an d E.B . White . 1979 . Th e Element s o f Style . Th e Macmilla n Company,
Toronto. Man y writers say this i s still th e bes t guid e fo r goo d writing .

CONSULTANTS
Ross E. Konin g Shirlen
Eastern Connecticu t Stat e Universit y Uta
Willimantic, Connecticu t Logan

e M. Pope, Emeritus Professo r

h Stat e University
, Utah

Andrea L . Peterso n Moyl

Utah Stat e Universit y Uta
Logan, Uta h Logan

e Q. Rice, Emeritu s Professo r

h Stat e University
, Utah

B
STANDARDS FOR EFFECTIVE PRESENTATIONS
Ross E . Koning
Biology Departmen t
Eastern Connecticu t Stat e University
Willimantic, CT 06226-229 5 U.S.A .
As scientists working in a rapidly advancing discipline, we must communicate effec tively a t regional , national , an d internationa l meetings . Whil e publication i s th e
permanent record o f research progress , we rely heavily upon meetings presentation s
to communicate our most recent ideas and results. Ther e are two common forms of
meetings presentation : th e oral report an d th e poster. Thi s appendi x is primarily
designed to assist with the preparation o f slides for the oral report, but much of the
information applie s equally well to artwor k prepared fo r posters .
To communicat e effectively , ora l presentation s mus t b e designe d t o optimall y
deliver ideas and finding s within a timed interval (usually less than 15 minutes). T o
achieve this, the artwork prepared for an oral report must be quite different fro m th e
artwork prepared fo r publication. Slide s cannot have the finenes s o f detail nor th e
complexity required of publication graphic s (for saving precious space in journals).
Since an oral report lack s the luxury of long explanations and detailed study, each
slide (o r viewgraph ) mus t hav e a simpl e format , mus t b e fre e o f nonessentia l
information, mus t b e readil y understood, an d mus t hav e a single , clea r purpose .
Each slide must be visible, legible, attractive, and integrated with the other slides and
the oral presentation. I f properly designed, your graphics should catch the attentio n
of the audience , reinforce your spoken ideas, and make your communication easier,
faster, an d mor e exciting.
Presentation plannin g software for computers may assist researchers i n designing
effective graphics , but becaus e th e powe r t o desig n is still th e prerogativ e o f th e
scientist, som e guidelines are needed t o us e this software t o best advantage. Man y
examples of poorly designed slides generated by computer are evident at professional
meetings. Thus , the guideline s presented here should be followed b y those who use
computer graphic s facilities as well as those usin g older methods.
Clearly, slides must not detract fro m th e presentation, so a key word for thinking
about graphic s i s simplify! Yo u d o no t wan t you r audienc e t o wallo w i n th e
particulars o f your approac h bu t t o understan d your result s an d t o com e t o your
conclusions (literally to see what you mean). I t is important to keep methodological
188

Standards for Effective Presentations 18

and statistica l detail s fro m cloudin g th e finding s o f your research . A n intereste d

party ca n ask questions about method s an d analysis after you r presentation o r i n a
private conference . You r finding s mus t stand abov e your process .
The followin g sections present som e o f my ideas abou t presentations .
1. SLIDE PRESENTATIONS
A, Planning your slides. T o kee p th e audienc e listenin g an d interested , yo u
should organize and plan your presentation carefull y before any artwork is attempted .
The process is essentially the construction of a storyboard. Again , computer software
has proliferate d t o assist busines s worker s and (t o a lesser extent ) scientist s i n this
process. First , compos e you r mai n messag e i n 2 0 word s o r less . Thi s shoul d
constitute you r title. Assembl e a sequence o f similar short message s for each piec e
of evidence leading to your conclusion. Desig n a graphic that will communicate each
element o f th e evidenc e a t a glance. Thi s migh t be a graph , a photograph , o r a
simple phrase. Allo w only one idea pe r graphic! Th e audience must both loo k an d
listen, so it is critical to keep the slides and the spoken word simple and coordinated .
To hel p organiz e you r presentation , a slid e o f you r question s o r mai n point s
might be projected nea r th e start of the presentation. A s you progress throug h your
presentation, th e outline slide could b e show n again with a topi c highlighte d i n a
contrasting colo r t o conclud e th e correspondin g sectio n o r t o introduc e th e nex t
point i n your presentation. A t th e en d of the presentation, th e lis t of questions o r
main point s migh t be shown agai n with answers to reinforc e your summary.
For complicated figures , use the build-up routine. Suppos e you wish to show the
difference betwee n tw o curves sharing a common abscissa ( x axis). I n the firs t slide ,
the ordinat e an d absciss a ar e describe d an d th e firs t curv e is shown. I n the secon d
slide, th e additiona l ordinat e i s adde d an d th e secon d curv e i s shown. (Th e firs t
curve and its ordinate ma y remain o r may be drawn with thinner lines, or relegate d
to a different colo r i n this slide.) I n the third slide, the area between the two curves
is hatched t o emphasize th e differences. I n this way, three slides are used to presen t
your findings. Now , to driv e the tren d home firmly , follo w u p with a text slide o f a
phrase boldl y proclaimin g th e tren d (light stimulates shoot growth). Thi s fina l tex t
graphic coupled wit h the build-up method makes your point clearly and memorably.
Instead o f dryl y describing on e grap h with tw o lines o n a singl e slid e ove r a two minute period, yo u will spend perhap s 3 0 seconds o n each of four slides , each slid e
having a clear purpose . I t helps your audience understand you better, an d you will
have hel d thei r attentio n t o your presentation .
Not al l graphic s i n your presentation nee d t o hol d information ; plain-color (no
information) slides draw attention t o you and to important conclusions that you will
simply state. Thi s incredibly valuable and effective techniqu e is seldom used but will
really make your audience liste n t o what you are saying. Sinc e the previou s slide is
replaced b y plain color o n the screen , th e audience is forced to hal t its examinatio n
and t o liste n t o your interpretation .
Your planne d presentatio n shoul d hav e enough slides t o preven t boredo m fo r
your audience. Yo u shoul d plan to chang e slides at 3-5 slides per minut e (no mor e
than 30 seconds spen t o n each slide). Limitin g each slide to one idea should assure

190 Appendices:

Presenting Scientftc Data

that you have enough slides . I f you do not have enough slides planned to accommo date this change rate, then you probably hav e planned slide s that present more than
one idea! Thes e need t o be simplified. Walkin g into a 15-minute presentatio n wit h
five complicate d slide s assure s boredo m fo r your audienc e an d disrespec t fo r you .
This is particularly tru e for weary audiences afte r a few days of a national meeting .
On th e othe r hand , i f yo u alread y hav e 5 0 slide s fo r a 15-minut e presentation ,
adequate rehearsa l wil l hel p yo u decide i f you cannot presen t al l the slide s (ideas )
you may have planned. If , during rehearsal of your planned presentation, yo u cannot
remember a particular poin t yo u want to make , you need anothe r graphi c elemen t
that wil l remind yo u of what was important an d will help driv e this point hom e t o
your audienc e a s well.
Examine your presentation plan to be sure you have included title slides, question
slides, evidence slides, and conclusion slides. Chec k carefull y t o not e whethe r you
need duplicat e slide s fo r graphic s tha t ar e t o b e show n mor e tha n onc e i n you r
presentation. Th e audienc e ha s no patience fo r you to give verbal instructions t o a
projectionist t o tr y to locat e a previou s slid e i n a slid e tra y and the n t o retur n t o
another specifi c slide t o continu e you r presentation . Never turn back! Th e story board mus t be unidirectional !
B. Preparing the artwork. Artwor k mus t be designe d t o a 2-height-by-3-width
ratio, which convert s directl y t o th e 2 4 x 36-mm forma t o f the standar d slide . I t is
best t o kee p th e artwor k awa y fro m th e edge s o f th e fram e an d t o hav e centra l
weight to the figure . Colo r and lettering weight can be used to emphasize. Realiz e
that wester n audience s wil l view the slid e from uppe r lef t t o lower right, so items in
the uppe r righ t and lower left corner s o f the diagram are of less importance and may
go unnoticed i n a complicated slide .
Legibility i s a most-importan t criterion . Make it bold! Us e larg e fonts ! I f
your artwork outpu t i s on 8. 5 x 11-inch or on A4 paper, tape it to a wall and stan d
three meter s away . I f you ar e usin g a compute r scree n t o prepar e artwork , again
move thre e meter s fro m th e scree n t o chec k fo r legibility . A t thi s distance , you r
graphic ha s th e sam e visua l size a s a projectio n scree n viewe d fro m th e bac k o f a
lecture room . Ca n yo u stil l rea d everything ? I s the poin t o f th e graphi c clearl y
observed fro m thi s distance? I f not, then the lettering fonts , symbols, line thickness ,
or other elements o f the artwor k must be made larger or bolder .
As a fina l chec k fo r legibility , hold th e graphi c that ha s been converte d t o a 2 x
2-inch slid e 4 0 cm fro m you r nose. Chec k t o b e sur e you can read everythin g and
that th e poin t o f th e graphi c i s clearl y observed fro m thi s distance . I f not , the n
either th e letterin g o n th e artwor k must be proportionally large r o r the slid e mus t
be retake n t o mor e nearl y fill th e frame .
Legibility i s assured i f a fe w simple rule s ar e applie d t o al l graphic s use d i n a n
oral presentation . A Koda k publicatio n (Koda k #2 , 1986 ) i s a goo d technica l
discussion o f legibilit y parameter s an d give s excellen t tip s fo r makin g your slide s
readable.
Text. Eac h textua l graphic should be limite d to a fe w (less tha n 10 ) words. A
text slid e shoul d b e though t of a s a n idea-gram . I t make s your poin t dramatically
and remind s yo u o f wha t t o sa y withou t havin g note s t o handl e (o r shuffle!) .

Standards for Effective Presentations 19

Complete sentence s are seldo m legibl e o n a screen, so you should never hav e much
to rea d o n a slide . I f you read a slid e t o th e audience , yo u will bor e them ; i f th e
audience read s a slid e while you discus s it , the y are no t listenin g carefully t o you .
Helvetica o r other sans-seri f typefaces (suc h a s thi s one) ar e much more readabl e
in graphics tha n serif-fonts such as Times Roma n (lik e this) . Th e font siz e must be
tall enoug h s o tha t onl y 1 2 lines would fil l th e slid e area fro m to p t o bottom , an d
wide enoug h s o tha t onl y 36 characters woul d fill th e slid e are a fro m lef t t o right .
The fonts must be bold; the thickness of the lines used to form each character shoul d
be betwee n 1/1 0 t o 1/ 5 of th e heigh t o f th e character . I f these siz e guidelines ar e
followed strictly , then everyone, eve n those in the back of the conference room , will
be able to see what you intend t o show them. Thes e legibility rules appl y equally to
lettering on tables, graphs, an d other artwork.
Tables. Prepar e a slide showing onfy th e portion of the tabl e tha t you intend t o
discuss; leav e ou t unuse d data . Excessiv e ra w dat a an d particularl y unprocesse d
statistical table s ar e roadblock s betwee n yo u an d your audience ; you r audienc e i s
likely t o b e mor e intereste d i n plan t physiolog y than i n statistica l analysis . I f you
plan t o focu s onl y o n a fe w cell s i n you r table , the n reformulat e th e tabl e fo r
projection. Reserv e the complet e tabl e as a photocopy for private discussions with
interested conferees . Alternatively , reserv e th e complet e table as an informational
slide afte r you r fina l slid e a s potentiall y usefu l i n answerin g a questio n fro m th e
audience. Moreover , i f you wish to reac h a large r audience, a handou t migh t b e
desirable. However , i f you do have a handout, pass it out onl y when the tal k is over,
so the audienc e will be listening and not readin g during your presentation. B e sure
your name, address , an d phone numbe r are printed on the handout so a member of
the audienc e ma y contact yo u later fo r discussion.
In any case, table s for your oral presentation shoul d have no more than fiv e rows
or columns t o sta y within the limits of legibility and sensibility. I t is better t o divide
a large table into smaller portions o n several slides than to present an illegible smear
of tiny digits for several minutes . I t is important to note tha t graphs (line, bar, pie)
are usuall y far mor e illustrativ e an d memorabl e than number s in tables .
Line graphs should be bold an d legible. Man y computer graphics programs draw
axis and plotted line s to o finel y b y default an d need t o be modifie d fo r making bold
projection slides . Ther e should be no more than eight marked ticks on any axis, and
all lettering shoul d conform to the legibilit y rules for text (above). Again , computer
programs usuall y default t o smal l font s suitable fo r publicatio n but unsuitabl e fo r
projection. Ideally , you should show only one or tw o lines or curve s on each slide ,
but you may build up to a multiple-curve figure by revealing a new line on successiv e
slides.
Bar graphs should hav e no more than eight bars and, if stacked bars are used, the
bars ma y be (shoul d be ) reveale d sequentiall y or i n group s by color o r textur e fo r
contrast.
Pie charts shoul d hav e fewe r tha n 1 0 slices . Colo r o r textur e contras t an d
"pulled-out" slices ca n b e employe d fo r emphasis . Sequentia l emphasis in a slide
series ma y better communicat e your findings.

192 Appendices:

Presenting Scientfic Data

Colors can be used t o great advantage , but th e overuse o f colors i s only distracting. Th e ease of applying color i n computer-generated graphic s tempt s researcher s
to us e too man y colors. Wheneve r possible, associat e a color wit h a particular typ e
of information. Th e color s mus t contrast wel l in order t o b e distinguished. Ligh t
colors on a dark background are very legible. Dar k colors on a medium background
are seldo m appreciated . Extrem e contrast i s needed fo r partially-darkene d lectur e
halls, bu t i n fully-darkene d room s extrem e contras t fatigue s th e visio n o f you r
audience. Hig h contras t blac k and white positives and negatives will be more easily
appreciated i f dye d t o a paste l color . Thi s take s th e "edge " of f o f th e excessiv e
contrast. Thi s i s particularly important if you intend to mix black-and-white artwork
with typical color slide s or othe r continuous-tone artwork .
Do no t underestimat e th e specia l connotation s o r emotiona l impac t o f certai n
colors (Xerox , 1985) :
Red: stop
, danger , fire , anger , warmth, passion, excitemen t
Blue: police
, navy, sea, serenity , sky, fidelity, water, coolnes s
Green: go , growth , trees , country , spring , restfulness , youth , freshness ,
money (USA )
Yellow: caution , sunlight , cheerfulness, heat, light, life
White: hospitals , sterility , purity , innocence, peace , calm
Gray: somberness
, dignity , quietness, age, wisdom, gravity
C. Making the slides. A variety of techniques can be employed to conver t welldesigned legibl e ar t wor k t o effectiv e slides . Scientist s wit h acces s t o compute r
graphics equipment can generate slides directly from their computers. Severa l highly
sophisticated program s are now on the market to create beautiful, multicolored slides
(e.g., Harvard Graphics , Power Point , Slid e Write, WordPerfect Presentations, etc.) .
It is possible t o photograph thes e directly from th e screen although resolution i s not
as good a s when th e compute r fil e fo r the slid e i s sent t o a fil m recorde r designe d
to expos e th e fil m accordin g t o th e file . Suc h recorder s cos t a t leas t a thousan d
dollars (althoug h price s hav e been droppin g sinc e the y first cam e o n th e market) .
If yo u don' t hav e th e recorder , yo u ca n generat e th e presentatio n graphic s o n
diskettes tha t ar e the n sen t t o agencie s t o b e converte d to slide s (a t cost s around
$5.00 pe r slide. )
If th e compute r facilitie s are no t availabl e to you , or i f you haven't tim e t o g o
through the sometimes involve d process of using computer programs to produce th e
slides, yo u can use th e tried-and-tru e photographi c processes tha t everyone ha d t o
use before th e compute r revolution . Basically , these processe s involv e puttin g th e
graphs, tables , words , o r othe r material s o n pape r o r othe r suitabl e mediu m an d
photographing th e result s t o mak e slides . Suc h slide s ca n b e draw n b y hand o r
produced wit h suitabl e softwar e on a computer attache d t o a high-qualit y printer.
Often, th e Universit y or othe r organizatio n ha s a photography laboratory tha t will
make th e slide s fro m th e ar t work . Som e workers lik e to o r mus t go through th e
entire proces s themselves , usin g technique s describe d i n th e appendi x t o thi s
appendix.
One quic k and simpl e approach is to photograp h the ar t work , on a goo d copy
stand, usin g color slid e fil m (direc t positive). Thi s give s fairl y satisfactor y result s

Standards for Effective Presentations 19

even wit h black-and-white copy, providing that you remember to over expose b y 1/2 to
one full f stop s o tha t th e backgroun d wil l appea r brigh t instea d o f gre y whe n
projected. O f course , i f you us e color film , yo u can add colo r t o th e ar t wor k o r
even us e color filters when makin g the slides .
H. Showing the slides. Ther e is no replacemen t fo r adequate rehearsal o f your
slide presentation . I t wil l show errors i n you r planning, errors i n logic , error s i n
legibility, an d certainl y hel p you fee l mor e confiden t when you actually deliver th e
oral report at the conference. O f course, this rehearsal must begin sufficiently befor e
the conferenc e s o that error s ca n be corrected. Yo u should begi n t o prepare you r
oral report a s soon a s possible.
During rehearsa l b e sur e you r slide s ar e i n mount s tha t wil l functio n i n th e
standard Koda k Carousel 14 0 tray. Thi n plastic mounts jam less frequently tha n any
other type. Th e mount s should b e numbered so that if a stack of slides is dropped,
order ca n be quickly restored. Yo u should place a dot or an x in the proper corne r
of th e moun t s o tha t i t will be oriente d properl y at th e conference . Th e standard
convention fo r placement o f this mark is to hol d the moun t so that th e slide can be
properly read and to mark the lower left corne r of the mount. Th e projectionist will
then reorien t th e moun t so the mark is in the upper right corner. Thi s inversio n of
the slid e will assure prope r orientation o n th e screen. B e sure to check this orientation o f your mark during rehearsal!
When yo u trave l t o th e conference , carr y your slides i n a carry-o n bag to avoid
loss wit h your checke d baggage . I f your rehearsa l ha s bee n adequate , yo u shoul d
arrive, slide s i n hand , a t th e conferenc e confiden t tha t you r presentatio n wil l b e
organized, legible , an d understandable.
At th e sessio n fo r your presentation , arriv e before the beginnin g of the sessio n
to presen t you r (marke d an d ordered ) slide s t o th e projectionis t an d familiarize
yourself with the roo m an d its facilities. Brin g photocopies and maybe an overhea d
transparency o f eac h graphi c just i n cas e th e slid e projecto r fails . I f the projecto r
lamp burns out, you r 15-minut e time slot i s too shor t t o have the lamp replaced i n
time for you to finis h wit h slides .
When i t i s your turn to speak , remember never read a prepared speech. Ther e is
no more boring method o f delivery, and your audience unconsciously wonders if you
did the work you are presenting. "I f you did the work, then why must you read about
it?" Yo u shoul d b e abl e t o discus s the artwor k without a written text. Moreover ,
many auditori a ar e no t equippe d wit h readin g lights . Le t you r artwor k be your
notes. Never read a slide t o th e audience! Th e onl y thin g worse tha n projecte d
sentences i s projected paragraphs . You r audience thinks, "Why give an oral presen tation; jus t publis h th e work! " Grammaticall y correct sentence s wit h complicate d
logic and suitably condensed fo r journal publication are incomprehensible in an oral
presentation; us e simple, direct, conversationa l English .
Be careful in your use of a pointer, especiall y a light pointer (e.g. , a laser pointe r
that project s a smal l red dot). I t i s very distracting to a n audienc e when the ligh t
from the pointe r dance s or wave s around excessivel y on the scree n and especiall y
when it dances all over the room while you are talking but not pointing. Hol d it still
or mov e i t slowl y to emphasiz e what yo u want , an d turn i t of f when i t i s not being
used!

194 Appendices:

Presenting Scientfic Data

I. Special language problems. Yo u fac e a n especiall y difficul t challeng e i f you

must giv e your presentatio n i n a language tha t i s not your nativ e tongue . Englis h
has become the international languag e for scientific presentations, an d most scientist s
everywhere no w rea d an d ofte n spea k English . Man y scientist s whos e nativ e
language i s not English , however , d o not fee l confiden t enoug h i n the languag e t o
give an oral presentation withou t a written text, which breaks the rule just presente d
to never read a prepared speech. I n spite o f this rule, it is common a t internationa l
meetings t o hea r presentation s tha t ar e rea d i n Englis h b y non-English-speakin g
scientists. Th e fac t tha t th e talk i s read instea d o f being presented spontaneousl y is
already enough to lose the interest of many if not most in the audience, bu t often the
speaker i s not highl y proficient in English, so that his or her unfamilia r accen t add s
still another solid barrie r betwee n th e speaker an d the audience. Ther e are severa l
options availabl e t o you if you must present a talk in a language in which you are no t
highly competent. Th e following two options depen d on the status of your presentation:
i. You arrive at the meeting with a conventional manuscript, perhaps one that
has not even been edited by an English-speaking editor. Yo u shoul d strongl y
consider askin g a fello w scientis t whos e nativ e languag e i s Englis h (o r th e
language o f th e conference ) t o rea d th e pape r fo r you. I f you do , thi s perso n
should have time to study the manuscript before presenting it , making grammatical and stylistic suggestions tha t will improve understanding. Th e reader shoul d
practice reading th e manuscrip t and be admonished to read slowly with frequent
pauses, eve n insertin g spontaneou s remark s tha t hel p t o clarif y wha t i s bein g
presented. Remember , however , tha t no t al l scientist s whos e nativ e tongu e i s
English ar e skillful reader s wh o can read smoothl y without backing up to correc t
small errors. I f there is time, the frien d migh t even study the manuscript until he
or she has become s o familiar with its contents tha t he or she can use your slides
to presen t th e materia l withou t readin g it . Remember , yo u will b e ther e t o
interrupt th e perso n wh o i s givin g your talk ; you ca n correc t an y mistake s o r
misconceptionsand yo u wil l b e th e on e wh o answer s question s durin g th e
discussion period. B y having someone familiar with the language read your paper,
you will almost certainl y communicat e with the audienc e better tha n if you read
the pape r yourself.
ii. You have time before going to the meeting to follow the suggestions that are
presented in this section but still feel that you must read the paper. Remembe r
that a n ora l presentatio n differ s i n significan t ways fro m a publishe d technica l
paper. Man y of thos e who mus t present thei r tal k in a foreign language simply
read a manuscrip t tha t ha s bee n prepare d exactl y a s i f i t wer e goin g t o b e
publishedas often it will be. Suc h a format i s especially difficult fo r an audienc e
to follow. Instea d o f using this rather standar d procedure, outlin e your talk as a
series o f logically related, simpl e ideas, and prepare at least one slid e for each of
those ideas , a s discusse d above . The n write , probably in your ow n languag e t o
begin wit h s o tha t idea s will flo w freely , a shor t paragrap h that describe s eac h
slide. Havin g done this, you can then either practice the English version until you
are proficientpossibl y s o proficien t tha t yo u won't hav e to rea d i t afte r allo r

Standards for Effective Presentations

19

you can persuade a n English-speaking fello w scientist to read th e paragraphs for

you. Yo u migh t eve n hav e tim e t o sen d you r slide-relate d manuscrip t t o th e
friend befor e th e meetin g s o that h e or she can practice reading it. I n any case,
preparation base d upo n idea s an d slide s rathe r tha n the usua l technical-manu script forma t will lead t o muc h better communicatio n with your audience .
2. POSTER PRESENTATIONS
Part o f th e cultur e o f moder n plan t scienc e i s tha t mos t presentation s a t
professional meeting s are now in the form of posters. Th e poster session consist s of
two distinct parts: th e physical poster an d informal discussions of the research wor k
presented o n the poster. Researcher s prepare th e physical poster at home but must
also b e prepared fo r the discussion s to tak e place at th e meeting.
A. The Physical Poster. Eac h society has specifications for their poster sessions ,
and these are usually found in the brochures calling for abstracts to be submitted for
the meeting. Th e specifications provide the scientist with dimensions and orientation
for th e display boards. Th e entire poster mus t be prepared with th e size and orientation i n mind.
A four-foot-squar e poster (commo n i n th e U.S. ; a littl e ove r a squar e meter )
mounted wit h the to p o r center nea r eye-leve l i s simple to prepare. Th e title goe s
at th e top , an d graphi c element s place d virtuall y anywhere on th e boar d wil l b e
visible at a glance. I n general, a Western viewe r will examine a poster fro m lef t t o
right an d fro m to p t o bottom , s o th e element s ar e usuall y arrange d t o match .
Contorted path s t o follo w th e element s should be avoided.
If th e specification s ar e fo r a 4x8-foo t rectangl e (o r comparabl e metri c dimensions), the preparations mus t be more careful. I f the orientation is horizontal at eye
level, th e autho r mus t prepar e th e poste r wit h the ide a tha t th e viewe r will work
across th e poste r fro m left-to-right . Moreover , wit h th e genera l left-to-righ t
movement o f session participant s throug h the poste r displays , it will be difficul t fo r
discussants t o hav e to mov e back to th e lef t fo r a second ro w of graphic elements .
Some meeting s avoi d thi s movemen t proble m wit h 4x8-foo t board s standin g
vertically. Th e reade r wil l work down a poste r wit h thi s orientation , bu t graphi c
elements place d abov e 6-fee t fro m th e floo r o r belo w waist-height are difficul t t o
observe. Thes e area s shoul d b e restricte d t o ancillar y element s suc h a s larg e
photographs no t needin g muc h close examination . Th e essentia l element s o f such
a poste r shoul d b e placed i n a 4x4-foot square centered a t ey e level.
Bold, sans-serif fonts (like this) are preferred for all text. Fanc y or calligraphic
fonts shoul d be avoide d because o f inherent poo r legibility. Th e title of the poste r
should be lettered a t 90-point o r larger type. Th e title should state the major point
or findin g o f th e researc h clearl y and i n a s fe w words a s possible . I n genera l i t
should match the titl e found i n the published abstract and program booklet fo r th e
meeting. Nearby , the author(s ) names and universit y or researc h organizatio n and
location shoul d appea r i n moderate-size d typ e (perhap s 60-point) . Th e majo r
headings o f th e poste r design should also be in 60-point type. Th e bulk of any text
elements should be i n 30-point type or larger.

196 Appendices:

Presenting Scientfic Data

The poste r i s no t a "journa l article o n a board, " and lon g passages o f tex t ar e
completely inappropriat e fo r a poster session . Instead , the text elements shoul d be
largely idea-grams that lea d th e viewe r through the othe r graphi c elements to th e
conclusions draw n from th e research . Th e sequence of presentation ma y be similar
to a journal article, however .
A cop y o f the poste r abstrac t migh t be the firs t graphi c element afte r th e title .
The abstract puts the entire research body into a concise paragraph that a viewer can
read to determine whether she or he wishes to continue to examine the poster o r go
on to something else . Thi s migh t be the longest tex t element o n the poster .
A brie f introductio n presentin g th e backgroun d o f th e researc h an d perhap s
introducing the particula r organis m studied might come next . Thi s should b e very
brief and cover onl y the essentials . Thre e o r fou r sentence s woul d be a good guideline. A photograph coul d b e appropriate here .
Unless th e researc h detail s th e developmen t o f a ne w procedure, a n effectiv e
poster migh t simply include a flow chart rather than a long text explaining methods.
In many cases a photograph or drawing communicates what would take many words
to explain in text .
Most o f th e graphi c elements o n a poste r presen t th e researc h results . Thes e
graphic element s consis t o f photographs , graphs , tables , autoradiograms , etc .
Effective poster s frequentl y hav e simpl e figur e caption s o r title s declarin g th e
interpretation draw n fro m eac h accompanyin g graphi c element . Eac h graphi c
element shoul d b e larg e an d bold . Th e boundin g rectangle for graph s and chart s
might be 2 0 x 30 cm. Th e minimu m type size should be 30-poin t for al l lettering.
Graph symbols should approach 6 mm in width, and connecting lines should be 2 to
4 m m wide . Th e graphi c element s ca n b e slightl y mor e comple x tha n thos e
presented i n slide s fo r ora l presentations , bu t n o mor e tha n thre e line s shoul d
normally appea r o n a grap h (unles s th e figur e present s a famil y o f closely relate d
curves), and bar charts should be limited to fewer tha n 10 bars. Eac h element should
have a brief explanatory caption, bu t lon g passages are best avoided .
There i s n o discussio n sectio n o n a poster . Neve r wast e preciou s spac e o n
discussion tex t elements . Th e purpos e of a poste r sessio n i s to personall y discuss
your researc h wit h intereste d viewers . Thi s interactio n betwee n scientist s i s th e
beauty of the poster session . Th e verbal discussions cover the details of the researc h
and suggestions for improvements, etc. You r poster should provide the evidence and
support for the verbal discussion. O f course, the main conclusions will appear in the
abstract. Thi s i s important because som e viewers will stud y your poste r when you
are no t availabl e for discussion.
The las t graphi c elemen t i n th e poste r sequenc e should b e a summar y o f th e
research findings . A bulleted lis t i s sometimes effective; a concludin g mechanistic
model diagra m might be more memorable .
Very fe w viewers ar e prepare d t o writ e down a lis t o f references t o tak e hom e
with them , s o a literature-cite d sectio n i s generall y a wast e o f poste r space .
Established scientist s wil l kno w th e fundamenta l literature , an d a n intereste d
newcomer will write to you after th e meetin g for references an d reprints. Havin g a
business card with you migh t b e a hand y and considerate alternative.

Standards for Effective Presentations 19

Some scientist s brin g summarizin g handouts wit h the m t o giv e t o intereste d

parties. Whil e thes e coul d b e a s formal a s an articl e reprint , the y are generall y a
"miniature" poster wit h a few literature references. I t is difficult t o judge how many
copies o f a handout t o brin g with you.
B. Setting up the poster. Ther e ar e man y style s o f poste r graphi c elements .
Careful us e o f colo r i s effective ; excessiv e us e o f colo r i s distracting . To o man y
graphic element s ar e intimidatin g an d stifl e discussion . Man y peopl e brin g th e
individual poster elements an d their captions, etc. on individual cards or sheets, often
on 8.5x11 or A4 sheets. Thes e hav e to be mounted on the display boards individually; spacing and alignment must be adjusted at the meeting. Thi s takes time that you
might choose to spen d i n oral session s o r otherwise. A better pla n is to attach th e
poster element s ont o large r cardboard s wit h spacin g and alignmen t determined a t
home. Thu s th e assembly of the poster a t the meeting involves only putting a small
number of cardboards together . O f course, th e smaller sheets ma y be easier t o pack
in your briefcas e o r suitcase .
C. The poster discussion. A t the prescribe d time , the author(s ) are t o stand a t
the poste r an d discuss th e researc h wit h interested viewers. Thi s is the opportunity
to shar e ideas , t o commen t o n technique s an d interpretations , t o improv e th e
science, an d sometimes t o make new friends. Th e author should not approach eac h
passer-by an d launch int o a dee p presentation . Instead , the autho r let s th e viewer
initiate a discussion . Th e viewe r usuall y is allowe d t o lea d th e discussio n i n a
particular direction . O n th e othe r hand , occasionally a viewer will simply ask th e
author to "present " th e research. The n th e author is free to launch into a discussion
in his/he r ow n direction. Man y viewers will simply want to examine the poste r an d
draw their ow n unspoken conclusion s withou t discussion.

Appendix to Appendix A
Photographic Techniques for Creating Slides
For thos e wh o lac k compute r photographi c facilitie s o r budget s fo r outsid e
consultants, Kodak' s pamphle t # 3 (1987 ) an d it s addition s (Koda k #4 , 198 2 an d
Kodak #5 , 1982 ) ar e helpfu l a s a n overvie w of th e proces s o f makin g your own
lecture slide s with simple and relatively inexpensive photographic techniques. The y
present som e o f th e option s availabl e t o mak e reasonable slide s fo r presentation .
Another Koda k pamphlet (Koda k #6,1985) is an excellent how-to booklet for several methods o f producing text slides. A small poster (Koda k #7,1987) review s film s
(and thei r availabilit y an d processing) , thei r uses , an d technique s fo r particula r
applications (text , charts, line drawings, prints, electron micrographs, chromatograms,
electrophoresis gels , autoradiograms, gross specimens , and lab scenes) .
Your artwor k wil l likel y fal l int o tw o categories : continuou s ton e an d hig h
contrast.
Continuous tone artwork consists of images containing various shades of gray or
various colors. Thes e ar e best rendered int o slides by photography with continuous
tone black-and-whit e (e.g., Koda k Rapid Proces s Cop y Film o r reverse-processe d
Technical Pa n Film ) o r colo r slid e fil m (e.g. , Ektachrom e or Kodachrome).

198 Appendices:

Presenting Scientfic Data

High contrast artwork consists of images containing only black and white or vastly
different shade s o f color. Thes e are bes t rendere d int o slide s b y photography with
high-contrast black-and-whit e film s [e.g. , Dektol-processed Kodalith , Technical Pan ,
or Precisio n Lin e (LPD4 ) films ] o r high-contras t colo r fil m [Vericolo r Slid e Fil m
(SO-279)]. Kodalith , Technica l Pan , and Vericolo r film s produc e a "negative" slide
of th e artwork . Th e dar k lines o n th e artwor k are clear o n the slid e an d the whit e
background of the artwork will be intensely black (black-and-white films) or intensely
colored (colo r determine d b y filte r selectio n wit h th e Vericolo r film) . Koda k
Precision Lin e fil m produce s a positive imag e (intensely blac k line s o n clea r back ground). A s mentioned previously , high-contrast black-and-clea r slides fatigu e the
vision o f your audience i n fully darkene d room s and should be dyed so that the clea r
areas tak e o n a light paste l shad e t o reduce contrast .
D. Methods for continuous tone slides. Koda k Technical Pa n Fil m 241 5 i s a n
incredibly fine-graine d negativ e fil m tha t ca n b e processe d t o variou s degree s o f
contrast (Koda k #8 , 1982) . I t therefor e make s a universa l fil m fo r al l sort s o f
applications. Wit h a POTA developer [1. 5 g l-phenyl-3-pyrazolidinone (i.e., Phenodone), 3 0 g sodium sulfit e pe r lite r distille d wate r use d fo r 1 5 min a t 2 0 C wit h
agitation], norma l contras t blac k and white negative s ca n be mad e fo r printin g o n
photographic paper . Norma l contrast blac k and white slides can be made by reversal
processing a s describe d i n th e bo x a t th e en d o f thi s appendix . Extremel y high contrast negative s fo r reversed-tex t slide s o r publicatio n print s ca n b e mad e b y
exposing th e fil m a t AS A 20 0 (1/3 0 s a t f/1 1 usin g 4 15 0 W photofloods ) an d
developing th e fil m i n undilute d Dekto l fo r 2 min a t 2 0 C wit h continuous slo w
agitation. Whil e th e reversa l processin g coul d b e use d t o mak e high-contrast ,
normal-text slides , LPD 4 i s a more convenien t alternative .
As note d above , colo r transparenc y film s (Ektachrome , Kodachrome , an d
equivalent) ca n be used t o make slides from artwork . Colo r reproductio n can be an
advantage t o distinguis h portion s o f pie charts , bars i n histograms, etc. However ,
standard blac k an d white artwor k loses some contras t with these films an d appear s
dark gra y o n ver y ligh t gra y background . Moreover , an y corrections an d surfac e
irregularities i n the artwor k will be visible in the fina l slid e because o f the low-con trast colo r rendition .
Color transparency film s ar e very useful fo r showing the plan t used, the method s
and equipmen t employed , an d your colleagues fo r the study, but tables , graphs , and
line drawing s from you r data ar e bette r presente d i n slides o f higher contrast .
E. Methods for high contrast slides. Th e best quality high-contrast negatives ar e
prepared fro m Kodalit h (o r similar ) graphi c art s film s processe d i n an y o f th e
Kodalith or similar (undilute d Dektol) developers. Thes e materials give an intensely
black background with very clear line-images . Th e film i s insensitive t o red safelight
and, therefore, als o to red or other faint-color guidelines on your artwork. Sinc e th e
contrast o f the fil m i s so high, corrections t o artwork made by clean erasure , white out ink , an d clea n correctio n tap e ar e invisible . An y undesirable mark s tha t d o
appear ca n be blotte d ou t o n th e negativ e with a n opaque ink (e.g. , fro m a perma nent-black marking pen) or special opaquing material available in photo stores. Th e
negative ca n b e printe d t o mak e publication print s or i t ca n be mounted i n a slide

Standards for Effective Presentations 19

mount for projection a s a reversed-text slide . A s a slide, the extreme contrast tempts
one t o dy e the fil m prio r t o mounting . Thi s is especially tru e when projection i s to
be in a small, completel y darkene d room . However , i n large auditoria wit h incom pletely drape d window s or "hous e lights" , the extrem e contras t i s highly desirable .
The very popular blu e backgroun d slides ar e made with SO-279 Vericolo r Slid e
Film (Koda k Pamphle t E-24) . Thi s i s exposed fo r 6 seconds a t f/1 6 through a n O
(range) "G " filter t o artwor k illuminate d by 2 x 500 W Photofloods held 1 m away
from an d a t a 45 angl e t o th e artwork . Th e fil m i s processed fo r 5 min at 3 5 C
in Unicolor K 2 Chemistry (or equivalent C-41 processing). I f you do not want to do
color photographi c processing , yo u ca n tak e th e expose d fil m t o a "One-Hou r
Photofinishing" company and ask for negatives only. Thes e ca n then be mounted i n
Pakon (o r equivalent thin-plastic ) slide mounts. (Th e company might do the mounting.)
In a pinch , you ma y substitute Kodacolo r I I (o r equivalent ) for SO-279 , expos e
it after metering a t the manufacturer-recommende d ASA, and have it processed fo r
negatives only . Th e background colors will be weaker and the printe d areas wil l be
slightly orange .
LPD4 Kodak Precision Lin e Film is a direct-to-positive film for making black-line
slides fro m black-lin e artwork. Th e fil m i s exposed fo r 1 0 s at f/9. 5 using 2 x 500 W
Photofloods a t 45 an d 8 0 cm from cop y center. I t is developed 1 min in undiluted
Dektol at 20 C with slow continuous agitation. Fix , wash, and dry as any other film .
The brilliant clea r backgroun d and crisp black lines of these slides mak e them suitable fo r us e i n a larg e auditorium , fo r us e i n wea k projectors , an d fo r us e i n
inadequately darkene d rooms . Thes e slides are excellent i n any projection environment. A disadvantage i s observed whe n these slide s are projected i n sequence wit h
color slide s o r other less-brilliant slides . Th e contrast ca n be painfully excessive ! I f
this is anticipated, th e backgroun d can be dyed to reduce contrast. I t is undesirable
to reduc e contras t ver y much, so very dilute solution s o f water-soluble dyes should
be used to obtain weak staining of the protein emulsio n (Frost, T.M . and P.A. Jones ,
1982).
F. Adding color to black-and-white slides. Negative s and slide s from black-andwhite film s ma y be dye d i n dilut e solution s o f water-soluble dyes. I sugges t one percent o r more-dilut e solution s of Tartrazine Yellow, Acid Orang e II , or Naptho l
Green. Addin g glacial aceti c aci d t o mak e the dy e solution 0. 5 % aceti c aci d will
improve th e uniformit y of th e staining . A brie f rins e i n wate r afte r stainin g will
prevent formatio n of opaque dy e crystals on th e fil m (Homer , J.A . an d C . Pennington, 1974) .
Individual line s o f typ e ca n b e emphasized , particularl y o n black-and-whit e
reversed-text negatives , by highlighting them with water-soluble ink from fel t markers
(e.g., Vis a Vis ) designed for overhea d projection . I f the marks are mad e on th e
shiny (backing ) sid e o f th e film , the y ca n b e washe d of f easil y an d reapplie d a s
needed.
G. Recovering from disasters. Farmers Reducer i s an amazin g treatmen t that
can eliminat e the exces s silve r in th e region s of a black-and-white slide or negative
intended t o b e clear . A n overexpose d o r overdevelope d negativ e slid e ca n b e

200 Appendices:

Presenting Scientfic Data

corrected by cutting silver from the film. Farmer s Reducer consists of two solutions:
Part A (37. 5 g potassium ferricyanid e / 500 mL water) and Part B (240 g/L sodium
thiosulfate). Thes e hav e a reasonabl y lon g shelf-life , bu t mixture s o f thes e tw o
solutions are effectiv e fo r les s than 3 0 min and, therefore , mus t be prepare d onl y
immediately before use . Th e standar d mixtur e is 1 part A with 4 parts B and 30
parts water. Th e slide, negative , or print is agitated i n the solution fo r as long as it
takes t o remov e th e unwante d silver . I f 30 mi n elaps e befor e completion , fres h
solution mus t b e prepare d an d th e proces s mus t b e repeated . Afte r sufficien t
clearing, was h th e fil m i n fiv e volume s o f water , dry , an d mount . Thi s simpl e
treatment ca n save you fro m havin g to repea t th e whol e exposure an d processin g
routine. It s availability also eliminates rationalizin g the use of a single substandard
slide fro m a roll of otherwise goo d frames .
There ar e intensifier s tha t migh t be helpfu l fo r correctin g underexposur e an d
underdevelopment, but I have not personally tried them. I guess I tend to err on the
other sid e i n an attemp t t o hav e very high contrast an d the deepes t possibl e black
areas. Th e intensifiers can increase th e contrast o f thin, continuous ton e negatives
but ma y be les s usefu l wit h slides .

Reversal Processing for Technical Pan Film

Use the followin g steps i n the dark:

5 minutes 7 0 C developer (D-1 9 undilute d stock solution)
5 volume water rinse
3 minute bleac h ( 6 g K2Cr2O7 + 7. 5 mL Sulfuric Acid pe r 500
mL water)
5 volume water rins e
2 minutes clearing (5 0 g sodium sulfite per 500 mL water)
Use the followin g step s i n the light:
5 volume water rins e
30 seconds per side of reel reexposure with 100 W bulb ca. 30 cm
1 minute Developer (D-19 a s above)

5 volume water rinse

Standards for Effective Presentations 20

REFERENCES
Frost, T.M. an d PA . Jones . 1982 . Do-it-yoursel f blac k and whit e slides. Bull . Eco . Soc . Amer .
63:16-17.
Homer, J. A, and C. Pennington. 1974 . A simple and rapid method o f adding color t o photographi c
projection materials . Southeas t Electro n Microscop y Societ y Abstracts .
Kodak #1 . 1975 . Koda k Publicatio n S-30, Plannin g and Producing Slide Programs .
Kodak #2 . 1986 . Koda k Publicatio n S-24, Legibility : Artwor k t o Screen .
Kodak #3 . 1987 . Koda k Publicatio n M3-106, Makin g lecture slides.
Kodak #4 . 1982 . Koda k Publicatio n M3-515, Makin g lecture slides: Workshee t #1 .
Kodak #5 . 1982 . Koda k Publicatio n M3-516, Making lectur e slides: Worksheet #2 .
Kodak #6 . 1985 . Koda k Publication S-26, Reverse-tex t slides .
Kodak #7 . 1987 . Koda k Publicatio n P-15, Koda k film s fo r lecture slides.
Kodak #8 . 1982 . Koda k Publicatio n P-255, Kodak Technical Pan Film 2415 .
Kodak #9 . 1984 . Koda k Publicatio n F-5, Koda k Professiona l Black and White Films.
Kodak #10 . 1983 . Koda k Publicatio n G-73, Koda k Precisio n Line Films.
Xerox. 1985 . Communicat e effectivel y wit h slides. Reorde r Number 610P153110. Xero x Repro duction Centers , Xero x Square , Rochester , N Y 14644 . (Ver y nic e bookle t put s muc h of thi s
appendix down i n an outlin e with color examples. ]
CONSULTANT
Frank B . Salisbury
Utah Stat e University
Logan, Uta h

c
GUIDELINES FOR MEASURING AND REPORTING
ENVIRONMENTAL PARAMETERS FOR PLANT
EXPERIMENTS IN GROWTH CHAMBERS
Developed b y the American Society of Agricultural Engineers Environment of Plant
Structures Committee ; approve d b y th e ASA E Structure s an d Environmen t
Standards Committee ; adopte d b y ASAE , Marc h 1982 . Revise d Marc h 1986 ;
reconfirmed Decembe r 1989 ; revise d Februar y 1992 3.
Submitted by:
John C . Sager
NASA, Joh n F . Kennedy Space Cente r
KSC, FL, 32899-0001 U.S.A.
Donald T . Krizek
USDA Climat e Stress Laborator y
U. S. Department o f Agriculture, ARS ,
Beltsville, M D 20705-2350 U.S.A .
Theodore W . Tibbitts
Department o f Horticultur e
University of Madiso n
Madison, WI 53706-1590 U.S.A .
SECTION 1: PURPOSE AND SCOPE
1.1 Th e purpos e o f thi s Engineerin g Practic e i s t o se t fort h guideline s fo r th e
measurement of environmental parameters that characterize the aerial and root
environment i n a plant growth chamber.
1.2 Thi s Engineering Practice establishes criteria that will promote a common basis
for environmenta l measurements for the research community and the commercial plant producer .
1.3 Thi s Engineering Practic e promote s uniformit y an d accuracy in reporting data
and result s i n the cours e o f conducting plant experiments.

Submitted Jul y 16 , 1993 ; include s a fe w recen t modifications . Thi s i s ASAE Engineerin g

Practice: ASA E EP 411. 2
202

Guidelines for Reporting Environmental Parameters 20

SECTION 2: INTRODUCTION
2.1 Th e aeria l environmen t i s characterize d b y th e followin g parameters : ai r
temperature, atmospheri c compositio n includin g moisture an d carbon dioxid e
concentration, ai r velocity, radiation, and the edge effects o f wall/floor on thes e
parameters.
2.2 Th e roo t environmen t i s characterized b y the followin g parameters : mediu m
composition and quantity, nutrient concentrations, wate r content, temperature ,
pH, electrica l conductivity, an d oxygen concentration .
2.3 Measurin g an d reportin g thes e variou s parameter s wil l b e covere d i n th e
sections that follow . Th e definitions of the parameters indicat e th e symbol and
units i n th e forma t (symbol , units) . Measurement s shoul d b e mad e tha t
accurately represen t th e mea n an d range o f the environmenta l parameter s t o
which th e plant s ar e expose d durin g the experimenta l period , t o indicat e th e
temporal variations , both cyclic and transient, and the spatial variations over th e
separate plants in th e chamber .
2.4 Th e definitions , measuremen t techniques , an d reportin g procedure s provid e
criteria an d promot e uniformit y i n measurin g an d reportin g environmenta l
parameters, bu t these guidelines should not be used to select the environmenta l
parameters applicabl e t o a particula r experiment . Othe r parameter s ma y be
applicable t o a particular experiment or special environments such as elemental
concentration i n hydroponi c solutions , pollutan t concentratio n i n ai r qualit y
research, and spectra l qualit y ratios i n photobiology .
2.5 Whe n measurement s ar e made , th e chambe r shoul d b e operatin g wit h con tainers an d plant s locate d i n th e chamber . Provisio n shoul d be mad e t o tak e
all measurements wit h minimum disturbance t o th e operating environment .
SECTION 3: DEFINITIONS
3.1 Radiation: The emission and propagation of electromagnetic waves or particles
through spac e o r matter .
3.1.1 Radiant energy (Q e, J): Th e transfe r o f energy of radiation .
3.1.2 Energy flow rate ( e , W) : Th e rat e o f flo w o f energy , a fundamental
radiometric unit ; also called radiant power.
3.1.3 Spectral distribution: A functional or graphic expression o f the relatio n
between th e spectra l energ y flux, spectra l photo n flux , o r fluence rate pe r uni t
wavelength, an d wavelength .
3.1.4 Spectral energy flow rate ( e y , W-nm" 1): Th e radian t energ y flow rate
per uni t wavelength interva l a t wavelengt h Y
3.1.5 Energy flux (E e, W.m- 2): Th e radian t energ y flow rate pe r uni t plan e
(flat) surfac e area ; als o calle d irradiance .
3.1.6 Spectral energy flux (E e,y , W.m-2.nm- 1): Th e radian t energ y flo w rat e
per uni t plane surfac e per uni t wavelength interval a t wavelength Y.

204 Appendices:

Presenting Scientific Data

3.1.7 Energy fluence (F e, J.m- 2): Th e radian t energy dose tim e integra l pe r
unit spherica l area .
3.1.8 Spectral energy fluence (Fei, J.m-2.nm- 1): Th e energy fluence per uni t
wavelength interva l a t wavelength y
3.1.9 Energy fluence rate (F et, W.m- 2): Th e radiant energ y fluence per uni t
time. Th e sam e a s radiant energ y flux (irradiance) fo r norma l inciden t (perpendicular) radiatio n o n a plane surface.
3.1.10 Spectral energy fluence rate (F e,t,y, W.m-2.nm- 1): Th e radian t energ y
fluence rate per uni t wavelengt h interva l a t wavelength y
3.1.11 Photon (uni t = q ; i.e., one photon): A quantum (the smallest, discret e
particle) o f electromagneti c energ y with an energ y of hc/A. ( h = Planck' s constant; c = speed o f light; y = wavelength). It s energy is expressed in joules (J).
3.1.12 Photon flow rate ( p , q.s- 1 or mol-s- 1): Th e rat e o f flow of photons.
3.1.13 Photon flux (E p, q-m-2.s-1 or mol-m-2.s- 1): Th e photo n flo w rat e pe r
unit plane surface area; sometimes als o called photon flux density to emphasize
the unit area .
3.1.14 Spectral photon flux (Ep,y, q-m-2.s-1.nm- 1 o r mol-m-2.s-1.nm-1) :
The photo n flux per uni t wavelength interval at wavelength A.
3.1.15 Photon fluence (F p, q-m"2 or mol-m" 2): Th e photo n flow rate pe r uni t
spherical area .
3.1.16 Photon fluence rate (F pt, q.m-2.s- 1 o r mol-m- 2.s-1): Th e photo n
fluence per uni t time . Th e same as photon flux for normal inciden t radiation.
3.1.17 Spectral photon fluence rate (F ptli, q-m-2.s-1.nm- 1 o r mol-m- 2-s4
nm-1): Th e photon fluenc e rat e pe r unit wavelength interval at wavelength y
3.1.18 Light: Visuall y evaluated radiant energy, with wavelengths approximately ranging between 38 0 and 78 0 nm, based on sensitivit y of the huma n eye.
3.1.19 Illuminance (Ev . 1x): The luminous flux (light incident pe r uni t area) .
NOTE: (a) Radiation instrument s that measure illuminance are not recommended. The y shoul d onl y b e use d alon g wit h recommende d radiatio n
instruments for historical comparison , (b) Conversion factor s from illumi nance t o radiation ar e spectrally sensitiv e an d thus unique for each specified
source.
3.1.20 Photosynthetically active radiation (PAR, q.m- 2.s-1, mol-q-m- 2. s 4 ,
or W-m- 2): Th e radiatio n i n the wavelength rang e of 400-700 nm . Measure d
as th e photosyntheti c photo n flu x (PPF), i n quanta.m- 2-s-1, or mol.m- 2-s-1
or photosyntheti c irradianc e (PI) in W-m- 2 for th e specifie d waveband , y1-y2
(400-700 nm).
3.1.21 Photomorphogenic radiation (q.m- 2.s-1, mol.m- 2.s-1, o r W-m- 2):
The radiatio n wit h wavelength s approximatel y ranging between 300-80 0 n m
contributing t o photomorphogeni c response s (i.e. , phototropism , flowering ,

Guidelines for Reporting Environmental Parameters 20

reproduction, elongation , dormancy ) i n relatio n t o th e relativ e quantu m

efficiency o f th e spectra l qualit y of th e radiatio n i n severa l discret e spectra l
regions. Measure d a s the photo n flux in average quant a .m-2.s-1, or i n energy
flux in W-m" 2 for th e specifie d waveband, y1-y2.
NOTE: Th e specfi c response s t o photomorphogeni c radiatio n mus t b e
biologically quantifie d and carefull y measured fo r eac h response spectrum
(action spectrum).
3.2 Temperature: Th e therma l state o f matter wit h referenc e t o it s tendenc y t o
transfer heat. A measure o f the mean molelcular kineti c energy of that matter .
3.2.1 Temperature, dry bulb (T, C) : Th e temperature of a gas or mixtur e of
gases indicate d b y a n accurat e thermomete r protecte d fro m o r correcte d fo r
radiation effects .
3.2.2 Temperature, wet-bulb (Tw, C) : Wet-bul b temperature i s the tempera ture indicate d b y a wet-bul b sensor o f a psychromete r constructe d an d use d
according t o instructions .
3.2.3 Temperature, dewpoint (Td, C) : Th e temperatur e o f a n ai r mas s a t
which th e condensatio n o f water vapo r begin s a s th e temperatur e o f th e ai r
mass i s reduced . Also , th e temperatur e correspondin g t o saturatio n vapo r
pressure (10 0 % relative humidity ) for a given air mass at constan t pressure .
3.3 Atmospheric moisture: Th e water vapor component of the mixture of gases of
the atmosphere .
3.3.1 Water vapor density (P r, g.m-3 or Pa) : Th e rati o o f th e mas s of water
vapor t o a given volume o f air, als o calle d absolut e humidity . I t ma y also b e
measured a s partial pressure . (Wate r vapo r pressure).
3.3.2 Relative humidity (Hr, percent): Th e ratio of the mol e fraction o f water
vapor present i n the air to the mole fraction of water vapor present in saturated
air at the same temperature and barometric pressure. I t approximates the ratio
of the partial pressure o r density of the water vapor in the air to the saturation
pressure or densit y of water vapor at th e same temperature .
3.3.3. Water vapor deficit (e d, Pa): Th e differenc e betwee n saturatio n wate r
vapor pressur e a t ambien t temperatur e an d actual vapor pressur e a t ambien t
temperature.
3.4 Air velocity (V, m-s-1): Th e tim e rate of air motion along a directional vector .
3.5 Carbon dioxide concentration ([CO 2], umol-mol-1 or Pa) : Th e carbon dioxid e
component o f th e mixtur e of gases o f th e atmosphere . Curren t expressio n o f
units of equivalent gas concentration ar e iimol-mol- 1, parts per million (ppm),
or u-L-L- 1, bu t the y d o no t expres s standar d temperature an d pressure , STP,
correction. Us e o f partia l pressure , Pa , i s preferre d i n nonstandar d atmospheres.
3.6 Watering (volume, L): Th e additio n of water to th e substrat e specified a s t o
the source , th e times , the amount , and the distribution method.

206 Appendices:

Presenting Scientific Data

3.7 Substrate: Th e medi a comprisin g th e roo t environmen t specifie d a s t o type ,

amendments, an d its dimensions (containe r size) .
3.8 Nutrition: Th e organic an d inorganic nutrient salt s necessary fo r plant growt h
and development . Formul a and/o r macr o an d micr o nutrient s ar e specifie d
within th e substrat e a s mol-m- 3 or within liquid solution a s mol-L-1.
3.9 Hydrogen ion concentration (p H units) : Th e hydroge n io n concentratio n
measured i n th e substrat e o r liqui d media over a range of 0 to 1 4 pH units .
3.10 Electrical conductivity (y c, mS.rn- 1): Th e electrica l conductivit y withi n th e
solid o r liqui d medium .
3.11 Accuracy : Th e exten t t o whic h the reading s o f a measuremen t approac h th e
true values o f a single measure d quantity .
3.12 Precision: Th e abilit y of the instrumen t t o consistently reproduc e a value of a
measured quantity .
SECTION 4: INSTRUMENTATION
4.1 Radiation. Sensor s shoul d b e cosin e correcte d an d constructed o f material of
known stability, known response curve , and lo w temperature sensitivity . Suc h
relationships shoul d b e specifie d an d availabl e fo r eac h sensor . B y definition
fluence measurement s can only be taken with spherical sensor s an d cannot b e
derived fro m measurement s take n wit h any plane-surface sensors. Th e sensi tivity an d linearit y ove r th e spectra l respons e an d irradianc e rang e shoul d b e
specified by calibration o r direct transfer from a calibrated instrument . Spectra l
measurements shoul d be made with a bandwidth of 20 nm or less in the 300-800
nm waveband.
4.2 Temperature. Sensor s should b e shielded with reflective material and aspirated
(> 3 m.s- 1) fo r air measurements .
4.3 Atmospheric moisture. Measuremen t shoul d b e mad e b y infrare d analyzer ,
dewpoint sensor , or psychromete r (shielde d an d aspirated at > 3 m.s- 1).
4.4 Air velocity. Senso r should hav e a range of 0.1 to 5. 0 m.s- 1.
4.5 Carbon dioxide. Measuremen t shoul d b e made by an infrare d analyze r with a
range o f 0 to 100 0 umol-mol- 1 o r greater .
4.6 Hydrogen ion concentration. Senso r shoul d hav e a rang e o f 3. 0 t o 10. 0 p H
units.
4.7 Electrical conductivity. Senso r shoul d have a range of 1 to 10- 2 mS.m-1 (1-100
milliohms resistance) .
4.8 Expected instrument precision and measurement accuracy. Tabl e 1 gives these
percentages, whic h indicate full scal e precisio n or accuracy . Furthe r definition
of thes e requirements ca n be foun d i n reference 27.

Guidelines for Reporting Environmental Parameters 20

Table 1. Expected Instrument Precision and Measurement Accuracy

Measurement
Parameter Instrumen
t accurac
y of
precision readin
g
Radiation
Flux
Spectral flux
Temperature

1 %
1%

10%
5%

0. 1 C

0. 2 C
0. 2 C

Relative humidit y
Dewpoint temperatur e
Water vapo r densit y
Air velocity
Carbon dioxid e

2%
0. 1 C

0. 1 g.m'3
2%
1 %

5 %
0. 5 C
0. 1 g.m-3
5 %
3 %

H+ concentratio n
Electrical conductivit y
Salt concentratio n

0. 1 pH

0. 1 pH

5%

5%

Air

Soil or liqui d

Atmospheric moistur e

PH

0. 1 C

SECTION 5s MEASUREMENT TECHNIQUE

5.1 Photon and energy flux. Measurement s shoul d b e take n ove r th e to p o f th e
plant canop y t o obtai n th e average , maximum , and minimum readings, an d a t
least a t th e star t an d en d o f each stud y and biweekl y if studies exten d beyon d
14 days.
5.2 Spectral photon or energy flux. A measurement should be taken at th e center
of the growin g area, at leas t a t th e star t an d end of each study.
5.3 Air temperature. Measurement s shoul d be made at the top of the plant canopy
at leas t daily , 1 h o r mor e afte r eac h ligh t an d dark perio d begins , t o obtai n
average, maximum , an d minimu m data . Continuou s measurement s ar e
recommended.
5.4 Soil and liquid temperatures. Measurement s shoul d be made at th e cente r of
the containers in the growing area, obtaining average, maximum, and minimum
readings a t th e middl e of the light and dark periods at th e start o f the experi ment. Continuou s measurement s during the entire study are recommended .
5.5 Atmospheric moisture. Measurement s should be made at the top o f the plan t
canopy in the center o f the growing area daily, 1 h or more after eac h light and
dark period. Continuou s measurement s are recommended .
5.6 Air velocity. Measurement s shoul d be taken at th e to p of the plan t canopy, at
the star t an d en d o f th e studies . Obtai n average , maximum , an d minimu m
readings over th e plants . I f instantaneous devices are utilized , 10 consecutive
readings should be take n a t eac h locatio n an d averaged.

208 Appendices:

Presenting Scientific Data

5.7 Carbon dioxide. Measurement s shoul d be taken at the top of the plant canopy
continuously durin g th e perio d o f th e study . A time-sharin g techniqu e tha t
provides a periodi c measuremen t (a t leas t hourly ) i n eac h chambe r ca n b e
utilized.
5.8 Watering. Th e quantity of water added t o each container o r average per plan t
at eac h waterin g shoul d b e measured . Soi l moistur e shoul d b e measure d t o
provide th e rang e betwee n waterings .
5.9 Nutrition. Measuremen t o f nutrients added to a volume of medium or concen tration o f nutrients added in liquid culture should be obtained at each addition .
5.10 Hydrogen ion concentration. Th e pH o f th e liqui d solution s i n a nutrien t
culture system should be monitored dail y and before each pH adjustment. Th e
pH o f the solutio n extracte d fro m soli d medi a should be measured at th e star t
and en d o f studies an d befor e and afte r eac h pH adjustment .
5.11 Electrical conductivity. Conductivit y o f th e liqui d solution s i n a nutrien t
culture syste m shoul d b e monitore d dail y durin g th e cours e o f eac h study .
Conductivity of the solution extracted fro m soli d media should be measured a t
the star t an d en d o f each study.
SECTION 6: REPORTING
6.1 Photon or energy flux. Report th e average and range over the containers a t the
start o f th e study , and th e decreas e o r fluctuation s fro m th e averag e ove r th e
course of the study. Th e source of radiation and the measuring instrument/sensor should be reported. Illuminanc e should not be reported excep t fo r historical compariso n i n conjunction with other radiatio n measurements .
6.2 Spectral photon or energy flux. Report the spectral distribution (graphical) and
the integra l (photo n o r energ y flux ) a t th e star t o f the study . Th e sourc e o f
radiation an d th e measurin g instruments should be reported.
6.3 Air temperature. Repor t th e averag e dail y readings with extreme s ove r th e
growing area for the light and dark periods with the range of variations over th e
course of the study.
6.4 Soil and liquid temperatures. Repor t th e averag e readings a t th e star t o f th e
study fo r th e ligh t and dar k periods .
6.5 Atmospheric moisture. Repor t the dail y average moisture level fo r both light
and dark periods with the rang e over th e cours e o f the study.
6.6 Air velocity. Repor t the average and range over containers at the start and end
of th e study .
6.7

Carbon dioxide. Repor t th e mea n of hourly average concentrations an d range

of averag e readings over the perio d of the study .

6.8 Watering. Repor t th e frequenc y o f watering , source , an d amoun t o f water

added dail y t o eac h container , and/o r th e rang e i n soi l moistur e conten t
between waterings.

Guidelines for Reporting Environmental Parameters 20

6.9 Substrate. Repor t th e type of soil and amendments, or components of soilless

substrate, and container dimensions.
6.10 Nutrition. Repor t the nutrient s added t o solid media . Repor t th e concentra tion of nutrients in liquid additions and in liquid culture solution along with the
amount and frequenc y of all additions.
6.11 Hydrogen ion concentration. Repor t th e mode and range in p H over the perio d
of th e study.
6.12 Electrical conductivity. Report th e average and range in conductivity over th e
period o f the study.

SECTION 7: SYNOPTIC TABLE

7.1 Tabl e 2 is a synoptic table o f the materia l presente d i n th e previou s section .
Table 2. Guidelines for Measuring and Reporting Environmental Parameters for Plant Experiments in Growth Chambers*
Measurements
Parameter Units

Radiation
Photon flux
y1 - y 2 nm , with cosin e
correction o r

umol.m-2.s-1
(y1 -y2 nm)
or

Energy flux (Irradiance) ,

y1 - y 2 n m with cosine
correction

W.m-2
(y1 -y2 nm )

Where to take Whe

At to p o f plant canopy.
Obtain maximu m and minimum over plan t growing
area.

n to take Wha
Minimum measurements :
at star t and finis h o f each
study and biweekl y if studies extend beyond 1 4 d.

t t o report
Average ( extremes ) ove r
containers a t start o f study.
Percent decreas e o r fluctu ation fro m averag e ove r
the cours e of the study.
Source o f radiation and
instrument/sensor.

Continued

Spectral photon flux

y1 - y2 nm, i n < 20 n m
bandwidths with cosin e
correction o r

umol.m-2.s-1.nm-1
(y1 - y 2 nm)
or

Spectral energy flux

(Spectral irradiance)
y1 - y 2 nm , in <10 nm
bandwidths with cosin e
correction

W.m-2.nm-1
(y1 - y2 nm)

Photosynthetic photon
flux, PPF,C y400 - y700 nm
with cosine correction o r

umol.m-2.s-1
(y400- y70 0 nm )
or

Photosynthetic irradiance,
PI,C y400 - y700 nm with
cosine correctio n

W.m-2

At to p o f plant in center of
growing area.

Minimum measurement :
at star t an d end o f each
study.

Spectral distributio n of
radiation with integral (y 1
- y 2) a t star t o f study.
Source of radiation an d
instrument/sensor.

At to p of plant canopy.
Obtain maximu m and minimum over plan t growing
area.

Minimum measurement:
at star t an d finis h o f each
study and biweekl y if studies extend beyond 1 4 d.

Average ( extremes ) ove r

containers a t star t o f study.
Percent decreas e or fluctu ation fro m averag e ove r
the cours e of the study .
Source of radiation an d
instrument/sensor.

(y400- y70 0 nm )

Temperature
Air
Shielded an d aspirate d
(> 3 m.s-1) device

At to p of plant canopy.
Obtain maximu m and minimum over plan t growing
area.

Minimum measurement :
measure once daily during
each ligh t an d dark perio d
at leas t 1 h after ligh t
change. Desirable : contin uous measurement .

Average o f once daily readings (o r hourl y average values) fo r th e ligh t an d dar k

periods o f the stud y with
extremes for th e variation
over th e growin g area.

Temperature
Soil and liqui d

In cente r o f container .
Obtain maximum and minimum ove r plan t growing
area.

Minimum: measur e a t th e
middle o f the ligh t and
dark periods a t the star t of
the study . Desirable : con tinuous measurement .

Light an d dark perio d

readings at th e star t o f th e
study (o r hourl y average
values of 2 4 h i f taken).

Continued

Table 2. Guidelines for Measuring and Reporting Environmental Parameters (continued)

Measurements
Parameter Units
Atmospheric Moistur e
Relative humidit y (RH )
with aspirated psychrometer, dewpoint hygrometer ,
or IRG A or

a
% RH , dewpoin t temperature, or g-m"3
or

Where to take Whe

n to take Wha

t to report

At to p o f plant canopy in
center o f plant growin g
area.

Minimum: onc e during

each ligh t an d dark perio d
at least 1 h after light
changes. Desirable :
continuous measurement.

Average o f daily readings

for bot h ligh t and dark
periods, with range of daily
variation during studies.

Vapor deficit , VP D or vapo r

difference

kPa o r g-m- 3

Air Velocity

m.s-1

At to p o f plant canopy.
Obtain maximu m an d
minimum reading s over
growing area .

At star t an d end o f studies .

Take 1 0 successive readings
at each locatio n and age .

Average readin g and range

over containers a t start and
end o f th e study .

Carbon Dioxide
Mole fractio n

nmol-mol

At to p o f plan t canopy

Partial pressure
Concentration

Pa
raol-m

Minimum: hourl y measurements. Desirable : continuous measurements.

Mean o f hourly average

concentrations an d range
of average concentration s
over th e perio d o f the
study.

Watering

liter (L )

Substrate

At time s of water addi tions.

Frequency o f watering.
Amount of water adde d
and/or rang e i n soil moisture content betwee n waterings.

At beginnin g of studies.

Type of soil and amendments. Component s o f

soilless substrate. Wate r
retention capacity . Con tainer dimensions .
Continued

Nutrition

Soil media
mol-m-3 o r mol-kg-

At time s of nutrient additions.

Nutrients added t o solid

media. Concentratio n o f
nutrients in liquid additions
and solution culture .
Amount an d frequenc y o f
solution additio n and re newal.

Liquid cultur e
mol-L-1

pH

pH unit s

In saturate d media, extract

from media o r in solution
of liquid culture.

Start an d end of studies in

solid media. Dail y in liquid
culture. Befor e each pH
adjustment.

Mode an d range durin g

studies.

Electrical conductivit y

mS-m-ld (millisie mens per meter )

In saturated media, extrac t

from medi a or i n solution
of liquid .

Start and end of studies in

solid media . Dail y in liquid
culture.

Average an d range durin g

studies.

"USDA Nort h Central Regiona l (NCR 101) Committee on Controlled Environment Technology and Use , June 1978; Revise d b y ASAE Environment of Plant Structure s Committee , Oc t
1978; Revise d by NCR 10 1 Committee, March 1993. Publishe d in par t in th e following references: 1 , 17, 18, 22, 27, 28, 34, and 37 .
a
r

Repor t in other subdivisions of indicated unit s if mor e convenient

The energ y flu x (irradiance ) is also commonly reported i n J.m-2.s-1 (equals W.m-2) .

Referre d t o as photosynthetically active radiation (PAR) fo r general usage .

mS.m- 1 = 1 0 umho-cm- 1.

214 Presenting

Scientific Data
BIBLIOGRAPHY

American Societ y fo r Horticultura l Scienc e Working Group o n Growt h Chamber s an d Controlle d

Environments. 1980 . Guideline s fo r measurin g and reportin g th e environmen t fo r plant
studies. HortScienc e 15(6):719-720.
ASAE. 1988 . ASA E Engineering Practice: ASA E EP285.7, Use of SI (Metric) Units. Ameri can Societ y o f Agricultural Engineers , St. Joseph, M I 49085.
ASAE. 1985 . ASA E Engineering Practice : ASA E EP402, Radiation Quantitie s an d Units .
American Societ y o f Agricultural Engineers , St . Joseph, MI 49085.
Bell, C.J . an d D.A . Rose . 1981 . Ligh t measurement an d th e terminolog y of flow. Plant , Cell an d
Environment 4:89-96 .
Bickford, E.D . an d S . Dunn. 1972 . Lightin g for Plan t Growth. Th e Ken t Stat e University Press,
Kent, Ohio .
Biggs, W.W. an d M.C . Hansen . 1979 . Instrumentatio n for biologica l an d environmenta l sciences .
LI-COR Inc . Lincoln , Nebraska .
C.I.E. 1970 . Internationa l Lightin g Vocabulary. Publ . No. 17 . Commission Internationale de
I'Eclairage, Paris , France.
Downs, RJ . 1975 . Controlle d Environment s fo r Plant Research. Columbi a Universit y Press,
New York, NY .
Downs, R.J . 1988 . Rule s fo r usin g the internationa l systems o f units. HortScienc e 12(5):811-812 .
Geist, J . an d E . Zalewski . 1973 . Chines e nomenclatur e fo r radiometry. Applie d Optic s 12:435 436.
Holmes, M.G. an d L . Fukshansky . 1979 . Phytochrom e photoequilibriu m in green leave s unde r
polychromatic radiation : a theoretica l approach . Plant , Cel l and Environmen t 2:59-65.
Holmes, M.G. , W.H . Klei n and J.C . Sager . 1985 . Photons , flux , an d som e ligh t o n philology.
HortScience 20(1):29-31 .
IES Lightin g Handbook. 1981 . Fift h edition . Illuminatin g Engineering Societ y o f North Ameri ca, New York, NY .
Incoll, L.D., S.P . Lon g and M.R . Ashmore . 1977 . S I units in publications in plant science.
Current Advance s i n Plant Scienc e 9(4):331-343.
Kerr, J.P. , G.W . Thurtel l an d C.B . Tanner . 1967 . A n integrating pyranometer fo r climatological
observer station s an d mesoscal e networks . Journa l of Applied Meterolog y 6:688-694 .
Kozlowski, T.T. , editor . 196 8 an d 1976 . Wate r Deficit s an d Plan t Growth . Vol . 1 , Development ,
Control an d Measurements . Vol . 4 , Soil Wate r Measurement , Plan t Responses and Breeding
for Drough t Resistance . Academi c Press , Ne w York, NY .
Krizek, D.T . 1982 . Guideline s fo r measurin g and reportin g environmenta l conditions i n controlled-environment studies . Physiol . Plant. 56:231-235.
Krizek, D.T . an d J.C . McFarlane . 1983 . Controlled-environmen t guidelines . HortScienc e
18(5):662-664 and Erratu m 19(1):17 .
Langhans, R.W. , editor . 1978 . A Growt h Chambe r Manual . Cornel l University Press, Ithaca ,
New York.
LI-COR. 1982 . Radiatio n measurement s an d instrumentation . Publ . No. 8208-LM. LI-COR ,
Lincoln, Nebraska .
McCree, KJ . 1972 . Tes t o f current definition s o f photosynthetically active radiatio n agains t leaf
photosynthesis data . Agricultura l Meteorology 10:443-453 .
McFarlane, J.C . 1981 . Measuremen t an d reportin g guidelines for plan t growt h chamber environ ments. Plan t Science Bulleti n 27(2):9-ll.
Mohr, H . an d E . Schafer . 1979 . Gues t EditorialUnifor m terminolog y for radiation : A critical
comment. Photochemistr y and Photobiolog y 29:1061-1062.
Monteith, J.L. 1984 . Consistenc y and convenienc e in the choic e of unit s fo r agricultura l science .
Experimental Agriculture 20(2):105-117.

Guidelines for Reporting Environmental Parameters 21

NBS Technical Note 910-2. 1978 . Self-Stud y Manual on Optica l Radiation Measurements , Par t
1Concepts. Unite d State s Government Printin g Office, Washington , DC.
Norris, K.H . 1968 . Evaluatio n of visible radiation for plant growth. Annua l Review of Plant
Physiology 19:490-499 .
North Centra l Regiona l 10 1 Committee o n Growth Chambe r Use . 1986 . Qualit y assuranc e
procedures for environmenta l contro l an d monitorin g in plant growth facilities. Biotronic s
15:81-84.
Percival Manufacturin g Co. 1981 . Guidelines : Measuring and Reportin g Environmen t fo r Plant
Studies. Availabl e as a plastic card.
Rosenberg, N.J . 1974 . Microclimate : Th e Biologica l Environment. Joh n Wile y & Sons, New
York, NY.
Rupert, C.S . an d R . Latarjet . 1978 . Towar d a nomenclature and dosimetric scheme applicable to
all radiations . Photochemistr y an d Photobiology 28:3-5 .
Salisbury, F.B. an d C.W . Ross. 1991 . Plan t Physiology. Fourt h edition . Wadswort h Publishin g
Co., Belmont , California.
Sestak, Z., J. Catsk y and P.G . Jarvis , editors. 1971 . Plan t Photosynthetic Production, Manual of
Methods. Junk , The Hague, Netherlands .
Slatyer, R.O. 1967 . Plant-Wate r Relationships. Academi c Press, New York, NY.
Spomer, L.A 1980 . Guideline s for measurin g and reporting environmenta l factors in controlle d
environment facilities . Commun . Soil Science an d Plan t Analysis 11(12):1203-1208.
Spomer, L.A . 1981 . Guideline s for measuring and reportin g environmenta l factors in growth
chambers. Agronom y Journal 73(2):376-378 .
Thimijan, R.W . an d R.D . Heins . 1983 . Photometric , radiometric, and quantu m light unit s of
measure: A review of procedures fo r interconversion . HortScienc e 18(6):818-821 .
Tibbitts, T.W . an d T.T . Kozlowski , editors. 1979 . Controlle d Environmen t Guidelines for Plant
Research. Academi c Press , Ne w York, NY.
Tooming, K.G. 1977 . Sola r Radiatio n an d Yiel d Formation (Solnechnay a radiatsiya i formirovanio urozhaya) Gidrometeoizdat , Leningrad .
Zelitch, I . 1971 . Photosynthesis , Photorespiratio n an d Plan t Productivity. Academi c Press , Ne w
York, NY .

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INDEX
Air, 7
temperature, 207-208
velocity, 205-208 , 21 2
Alanine, 89
Algebraic valency, 57
Aliasing, 12 9
Allele, 97
Allelopathic substances, 13 4
Allocation, 72
Allohydroxylysine, 89
Allometry, 11 1
Ambiphotoperiodic plants, 13 4
Ameliorate, 144
Amino acid residues, 85
Amino acids, 93
Amount of pure substance (mol), 5-6, 57
AMP, 87
Ampere (A), 5-7
Ampersand, 23
Amplified-fragment-length polymorphism , 97
Amplitude, 129
Anaerobic stress, 14 6
Analysis of variance, 32
Anamorphs, 25
And, 164, 168
Angstrom (obsolete), 14 , 18
Anisotropic growth, 11 1
Annual, 129, 134
Antecedent, 17 5
Anthesin, 13 4
Anthesis, 13 5
Antiauxin, 126
Anticodon, 97
Antiflorigen, 13 5
Antitranspirant, 151
Apex, 13 5
Apiaceae, 22
Apical:
cell, 113
meristem, 13 5
Apostrophe, 177 , 180
Arabic numerals, 5
Arabinose, 90
Arbitrary rules, 16 3
Area, 5, 9
Arecaceae, 22
Arginine, 89
As, 179, 18 1

A (symbol for amp), 6

Abbreviations, 178 , 186
Abortion, 13 4
Abscissa, 18 9
Abscission, 13 4
Absolute, 18 3
growth rate, 111, 118
response, 134, 139
Absorbance, 76-77 , 83, 87
Absorptance, 6 5
Absorption:
coefficient, 6 5
factor, 76
Absorptivity, 6 5
Abstract, 18 3
Acceleration:
caused by gravity, 1 7
of free fall, 1 8
Accelerational force , 6
Accept, 17 9
Acclimation, 143 , 154
Accuracy, 206
ACES, 92
Acronyms, 17 8
Acrophase, 12 9
Active, 18 6
voice, 17 6
Activity, 56
coefficient, 5 6
of radioactive source , 9
ADA, 92
Adaptation, 122 , 143
Addition (plus) sign, 25
Adenine, 91
Adenosine, 91
Adjectival sense , 11 , 14
Adjective(s), 23 , 164-165, 169-170, 181 , 185
ADP, 87
ADPglucose synthase , 10 6
Advection frost , 14 8
Adverb(s), 164-165 , 169, 171-170, 177 , 185
Adverbial forms, 18 5
Aero-, 12 4
Affect, 17 9
AFLP, 97
After-ripening, 13 4
Ageo-, 12 3
Agravi-, 12 3

217

218 Index
Asparagine, 8 9
Aspartic acid, 89
Asteraceae, 22
Articles, 16 9
Artwork, 188, 190, 192-193
Atmospheric moisture, 205-208, 21 2
Atomic mass unit (u), 83
Atoms, 7
ATP, 8 7
Atto (prefix = a), 8
Attributive adjective, 18 2
Auditorium, 19 9
Author(s), 23-24
Authorities, 2 3
Autocatalytic growt h function, 111
Autonomous, 121 , 124
Autonomously-inductive plant, 135
Autonyms, 23
Autotropism, 12 5
Auxin, 126
antagonist, 126
Average, 55
Avoidance, 14 3
response, 12 4
Avogadro's number, 7
(mole) of photons, 7 5
Axillary meristem, 13 5

Black-and-white artwork, 192

Blackfrost, 14 8
Blasting, 134
Blindness, 134
Blocking, 27
Blocks (statistics), 3 9
Blue background slides, 199
BlOhreife, 13 9
Boldface, 17 , 164
Bolting, 135
Botanical congresses, 21
Bound:
auxin, 126
water, 15 2
Boundary layer, 65
resistance, 6 7
Bowing, 12 1
Brackets, 173 , 174, 177, 186
Branch point, 97
Brassicaceae, 2 2
Bromouridine, 91
Build-up routine, 189
Bulging, 121
Bunsen-Roscoe law, 12 2
Bureau International des Poids et Mesures, 3
Business card, 196
But, 164-165, 167-168

Bacteria nucleation inhibitors, 14 6

Balance (to determine mass), 6
Ballistic transformation, 97
Bar (bar, obsolete), 14 , 18
Bar:
charts, 19 6
graphs, 19 1
'Base, a,'91
Base, kilobase (b, kb), 97
Base unit(s) (of the SI), 4-6, 12 , 15
in denominators, 15-1 6
Basic:
patterns, 16 5
sentence structures, 16 4
Basis of sequence, 10 6
Because, 180
Becquerel (Bq), 9, 12, 86
Bending, 121
and bulging, 12 1
BES, 92
Bibliographies, 18 0
Bicine, 92
Bidirectional transport , 7 3
Biennial, 13 5
Binomial, 23
Biological:
clock, 12 9
cycle, 12 9
rhythm, 130
Biosalinity, 15 4
Biothermodynamics, 93
BIPM, 3-6
BisTris, 92

C (symbol for coulomb), 9

Caesalpiniaceae, 22
Calorie (obsolete), 14 , 18, 86, 143, 146
cAMP, cGMP etc., 87
CaMV, 97
Can, 180
Candela (cd), 5-8

Capitalized, 10, 22-23

CAPS, 92
Caption, 184
Carbohydrates, 94
Carbon dioxide, 206, 208, 212
concentration, 205
4-Carboxyglutamic acid, 89
Carotenoids, 9 5
Case, 171
CAT, 98
Catalytic activity, 93
Cation exchange capacity (CEC), 15 4
Caulescent plant, 135
Cauliflower mosaic virus, 97
Cause, 179
Cavitation, 152
cDNA, 98
CDP, 88
CDTA, 92
Cell:
freezing, 14 7
production rate, 111
Celsius (C), 7, 10 , 12, 14
Centi (prefix = c; nonpreferred), 8
Centimeter, 17
centimetergranvsecond (cgs), 3

Index 21
Central Limi t Theorem, 41
Centrifugation, 1 8
Centromere, 9 7
Cesium-133 atom, 7
CGPM, 3-6, 13, 17
COS, 3, 11
CHAPS, 92
CHAPSO, 9 2
Charge, 5 8
Chemical potential, 5 7
of water, 6 1
Chemical terms, 170
Chemo-, 12 4
CHES, 92
Chilling, 14 4
injury, 144 , 149
-insensitive, 144
repair, 145
requirement, 146 , 151
reversal, 145
-sensitive, 14 5
temperature, 14 5
tolerance, 14 5
treatment, 14 5
Chloride salinity, 15 4
Chloroamphenicol transacetylase, 98
Chloroplasts (plastid) DNA, RNA, 92, 106
Chromatin, 98
Chromatography, 17 7
Chromosome, 9 8
Chronogram, 13 0
Chronology o f isolation, 10 6
Ciliary movements, 12 0
CIPM, 3, 4, 12-1 3
Circadian:
pacemaker, 13 0
rhythm, 130
time, 130
Circalunar rhythm, 130
Circannual rhythm, 130
Circasemidian, 13 2
Circaseptan rhythm , 130
Circatrigintan rhythm , 130
Circular, 98
dichroism, 8 9
Circumnutation, 12 5
Cis, 98
Cis-Aconitate, 9 0
Citric Acid, 90
Class (classis), 22
Clause(s), 164 , 168, 172-173, 18 5
Clay dispersion, 15 4
Clock time, 13 0
Closed circular, 98
Clusiaceae, 22
Cm-cellulose, 88
CMP-NeuAc, 87
CMP, 88
CO2 exchange, 117
CoA(orCoASH), 87
CoASAc, 88

Codon, 98
Coding sequence(s), 98, 106
Coefficient, 6 6
convective transfer, 66
heat energy transfer, 66
of determination, 32
of variation, 28
Coherent unit system, 5, 12-13, 16
Cohesion movement, 121
Cold:
hardiness, 14 6
injury, 14 6
protection, 14 6
shock, 14 6
Collective, 180
Colon(s), 17 7
Colony, 98
hybridization, 98
Color slide film, 192 , 197
Colors, 192
Comite International des Poids et Mesures, 3
Comma(s), 11, 164-169, 172-174, 176-177 , 185 186
to group numerals, 12
Comma fault (or splice), 167, 176, 185
Commission on Plant Gene Nomenclature, 105
Common Names, 25
Communicate, 164
Communication, 161-162, 188
Compatible solute, 15 4
Compatibility, 98
Complementary:
RNA, 92
DNA, 92, 98
Complementation, 98
Completely randomized design, 33
Components of water potential, 51
Composition of solutions and buffers, 8 4
Compound:
adjective(s), 169-170 , 177 , 185
nouns, 178
numbers, 170
symbols, 11
units, 12 , 15
Computer graphics, 19 2
Concentration(s), 9, 56, 212
of metabolites, 15
Conclusion, 189
Conditioning, 143, 145
Conduction, 65
Conference Generate des Poids et Mesures, 3
Confidence interval, 29
for a mean, 29
for a proportion, 30
for a variance, 29
Conjugated auxin , 12 6
Conjunction, 164-168 , 179, 181, 185
Conjunctions of time, 167
Conjunctive adverb(s), 167-168, 185
Connectives, 16 7
Constitutive, 98

220 Index
Construct, 98
Contingency tables , 40
Continuous tone :
artwork, 197
slides, 198
Contractions), 177, 18 0
Contrary to fact, 18 3
Control, 12 1
Controlling elements, 9 8
Controversial o r unfamiliar names, 24
Convection, 14 6
Convective:
rate of change, 111
transfer coefficient, 70
Convention d u Metre, 3
Coordinate, 16 4
adjectives, 169 , 17 3
clauses, 16 8
Coordinated, 166-16 7
Coordinating conjunction , 164 , 166 , 168 , 176 , 18 5
Copy:
editors, 161, 16 3
stand, 192
Correlation coefficient, 32
Corrinoids, 9 5
Cosinor (polar) display, 130
analysis, 130
Cosmid, 98
Coulomb (C), 9 , 12
Counts per minute, 86
Covariance analysis , 3 8
CPGN, 106-10 7
Critical:
daylength, 15 2
nightlength l5 2
period 15 2
photoperiod, 135
temperature, 143 , 145-14 6
Crop growth rate, 116-11 8
Cruciferae, 2 2
Cryo-, 98
CT, 13 0
CTP, 8 8
Cubed, 1 1
Cubic:
decimeter, 1 7
meter, 12 , 16
Cultivar(s), 21, 25
Cultivated Plants , 2 5
Culture collection, 21
Curie, 8 6
Current, 7
cv (ab. for cultivar), 2 5
Cycles per second (hertz), 86
Cyclic salt, 154
Cyclitols, 95
Cyclonasty, 12 5
Cysteine, 89
Cytidine, 91
Cytokinin, 12 7
-like, 127

Cytoplasmic streaming , 12 0
Cytosine, 9 1
d (symbol fo r day), 1 3
DD (continuous darkness), 13 1
Dalton (Da or D, obsolete), 14 , 17, 83, 86, 98
Damping, 13 0
Dangling participle, 175
Dark break, 131
Dash(es), 166 , 173-174, 177 , 186
Data, 180 , 19 1
Dawn, 130
Day (d), 7, 12-1 3
-neutral plants, 135
Daylength or day length, 135
Deacclimation, 14 3
Dead, 183
DEAE-cellulose, 88
Deci (prefix - d; nonpreferred), 8
Decimal:
fractions, 1 2
marker, 4, 11
system, 3
values less than one, 12
Decimeter, 16-1 7
Decussate, 11 3
Definite article, 16 9
Degree(s):
angles or latitudes, 11 , 13
centigrade (obsolete), 10, 14
Celsius (C), 10 , 18 , 11 , 14
kelvin (obsolete; use kelvin without degree), 7
Degree growth stage model (GS Model), 14 6
Deep supercooling, 146
Dehardening, 143 , 14 7
Dehydration (desiccation), 15 5
avoidance, 15 2
tolerance, 15 2
Deka (prefix = da; nonpreferred), 8
Demonstrative pronoun, 183
Denominator (in SI units), 12 , 15
Density, 7
of dry (unsaturated) air, 68
2-Deoxyglucose, 90
Deoxyribonucleic acid, 98
2'-Deoxyribosylthymine, 92
Dependent, 16 4
clause, 16 6
Deposition rate, 11 1
Derived units 4, 5, 8, 12
Desalination, 155
Desynchronization, 13 1
Determinate, 13 6
growth, 112
Development, 113
Developmental arrest, 136
Devernalization, 13 6
Dia-(not transversal), 124
2,4-Diaminobutyric acid, 89
Diel, 13 1
Difference:

Index 22
between two population means, 30
between two proportions, 3 1
or change in the quantity that follows, 68
Differential thermal analysis, 14 7
Differentiation, 11 3
Diffusion coefficient : 58 , 87
of species/, 66
Diffusive resistanc e within a leaf, 67
Diflusivity, 63
Dihydrouridine, 91
Dihydroxyacetone phosphate, 9 0
Dinesis, 123
Diploid, 10 1
Direct:
address, 177
quotations, 17 7
Direction, 122-12 4
Diurnal, 131
Discarded metric units, 14
Discussion section on a poster, 196
Distance, 67
Divergence angle, 113
Divisio, 22
Division, 10-11, 22
DNA, 88, 98
DNPorDnp, 88 , 135
Documentation, 21
Donor, 13 6
Domain, 98
Dormancy, 13 6
Dose, 12 2
Double:
commas, 17 4
spaced, 18 3
Downstream, 99
Drought (including: avoidance , escape, resistance,
stress, and tolerance), 15 2
DTA, 147
dTDP, dTMP, dTTP, 88
Due to, 180
Dyed, 192 , 198
Dyes, 19 9
Dyne, 3
Dysfunction, 14 5
EC (symbol for electrical conductivity), 155
Ectopic gene, 99
Eddy diflusion coefficient o f gaseous species, 66
Editing, 99
EDTA, 88, 92
Effect, 17 9
EGTA, 88, 93
Einstein (obsolete), 7, 14 , 17
Electrical:
capacitance, 9, 58
charge 9
conductance, 9, 58
conductivity (EC), 147 , 155, 206, 208-209, 213
current 5-6, 5 8
impedance, 14 7
potential (difference), 9 , 57

resistance, 58
Electro-, 124
Electrochemical potential, 57
Electrolyte leakage, 147
Electrometric method, 147
Electron:
paramagnetic resonance, 89
spin resonance, 89
Electronic "balances", 6
Electrons, 7
Electrophoresis, 99
Electroporation, 99
Ellipses, 17 7
Embryo dormancy, 136
Emissivity in infrared region, 6 8
Emphasis, 176-178
Encoding, 106
Endodormancy, 136
Endogenous, 121 , 136
rhythm, 130-131
Endotherms, 147
Energy, 9
balance equation, 68
flow rate, 203
fluence, 204
fluence rate, 204
flux, 75, 203, 210
English, 194 , 195
classes, 16 3
system, 11
English-speaking:
editor, 194
fellow scientist, 195
Enhancer, 99
Enthalpy, 50
change, 87
Entrain, 133
Entrainment, 131
Entropy change, 87
Enzyme commission numbers, 107
Enzymes, 93
Epinasty, 125
Epithet, 23-24
EPPS, 93
Equilibrium:
constant, 87
velocity constants, 83
Equivalent (obsolete), 86
Erg (obsolete), 3
Erythrose-4-phosphate, 90
ESP, 155
EST, 99
et, 23
etal., 180
Euhalophyte, 155
Evaporative:
cooling, 14 7
demand, 15 2
Evocation, 13 6
ex, 24
exa, 8

222

Index

Except, 17 9
Exchangeable sodium percentage, 155
Exclamation: point(s), 174, 176-177
Exclamatory writing, 176
Exogenous, 12 1
Exon, 99
Exotherm, 14 7
Expected instrument precision and measurement accuracy, 206
Experimental:
designs, 32
error, 27, 39
unit, 27
Exponential growth function, 112
Exponents, 11
Expressed sequence tag, 99
Expression, 9 9
Extensibility, 63
Extracellular, extraorgan, and extratissue freezing,
147
F (symbol for farad), 9
Fabaceae, 2 2
Factorial experiments, 35
Facultative response, 136 , 139
FAD and FADH2, 88
Fahrenheit temperature scale, 18
Family (familia), 2 2
names, 22
Family, gene, 106
Farad (F), 9
Faraday, 12
Farmers reducer, 19 9
Felt markers, 19 9
Femto (prefix = f), 8
Ferredoxin, 89
Fibonacci sequence, 11 3
Figures, 18 3
Film(s), 198-19 9
recorder, 192
First harmonic; second, third, etc. harmonics, 131
First law of thermodynamics, 4 5
First person, 17 1
personal pronouns, 17 6
Flagellar movements, 120
Flocculation, 15 5
Flooding, 147, 152
Flora, 23
Floral stimulus, 136
Florigen, 127, 136
Flower initiation, 13 6
Fluorescence (including: initial , maximum, variable,
terminal), 76
Flux, 58
FMN, 88
Folding effect, 12 9
Foliar absorption coefficient, 6 6
Folic acid, 95
Fonts, 191, 195
For, 16 4
Force, 6-7, 9, 46

Foreign words, 178

Formality, 172
Format, 183, 187
Fractional induction, 136
Free-running rhythm, 131
Free energy, 47
Freeze (including: avoidance, dehydration, desiccation, tolerance), 147
Freezing:
extracellular, extraorgan, and extratissue, 147
injury, 14 8
point, 148
point depression, 148
French Revolution, 3
Frequency, 8-9, 76, 131
multiplication and demultiplication, 131
of electromagnetic radiation, 68
Fries, 24
Frost, 148
hardening, stages of, 148
heaving, 148
injury, 14 6
plasmolysis, 148
protection, 148
Fructose, 90
Fructose-6-phosphate, 90
Fructose-2, 6-bisphosphate, 90
Fructose 1 , 6-bisphosphate, 90
Fucose, 90
Fumeric Acid, 90
Fundamental:
period, 131
term, 131
Fungi, 22, 24-25
Future tense, 182
Futurity, 182
gn (symbol for standard acceleration due to gravity),
9, 13-14, 18
Galactose, 90
Galvano-, 124
Gases in gases, liquids, and solids, 56
Gauss, 3
GDP, GDP-Man, GDP-Fuc, 88
Gel, 99
2-dimensional, 97
Gene, 99
designations, 105-106
encoding, 107
expression, 99
family(ies), 99, 106
product numbers, 107
symbol, 106
tagging, 99
transfer, 99
Generalized:
conductance coefficient, 5 9
force, 5 9
Generative helix or spiral, 113
Generic name, 23
Genitive, 23

Index 22
Genome, 99, 106
Genus, 22-23
Geo-, 12 3
Geographical location, 21
German, 16 3
Germination, 13 7
Gibberellin, 12 7
-like, 12 7
Gibbs energy change (formerly F) , 8 7
Gibbs free energy (G), 47
Giga (prefix = G), 8, 10, 17
Global irradiance , 67
Glucosamine, 90
Glucose-1, 6-bisphosphate , 9 0
Glucose, 90
Glucuronic acid, 90
p-Glucuronidase, 9 9
Glutamic acid, 89
Glutamine, 89
Glyceraldehyde-3-phosphate, 9 0
Glycine, 8 9
Glycophyte, 15 5
GMP, 8 8
Graft-chimeras, 2 5
Grafting, 13 7
Gram, 6, 10 , 17
Gramineae, 2 2
Grammar:
English, 164-18 7
as applied to SI units, 1314 ;
checkers, 18 4
Graphic(s), 188-190
elements (e.g., on poster), 196-197
Graphs, 191
Gravi-, 12 3
Gravity, 6-7, 17
Greek mu, 11 , 15
Greek symbols, 5
Growth, 11 2
analysis quantities, 118
field, 11 2
movement, 120
response, 12 3
velocity, 11 2
zone, 113
GSH and GSSG, 8 8
GTP, 88
Guanine, 91
Guanosine, 91
GUS, 99
Guttiferae, 22
h (symbol for hour), 13, 131
Half-cystine, 89
Half-peak band-width, 77
Halomorphic soil, 155
Halophyte, 15 5
Handout(s), 191 , 197
Haploid, 10 1
Hapto-, 123
Hard seed, 13 7

Hardening, 143, 145, 153, 155

Hardiness promoter, 148
Harmonic, 131
Harvest index, 116
Hb, HbCO, HbO2, metHb, 88
Heat, 9
shock, 99
Heat energy:
convection, 66
storage, 65
subscript, 66
Heat transfer:
coefficient, 7 0
studies, 68
Hectare, 10 , 14, 18
Hecto (prefix = h; nonpreferred), 8
Helio-, 123
Helium balloon, 7
Henry, 10
Hepes or HEPES, 88, 93
HEPPS, 93
Herbarium: 21-2 2
specimens, 21
Hertz, 9-10, 12
Heterogenous:
ice nucleation, 148
nuclear RNA, 92
Heterologous, 100
Heteropolymeric proteins, 10 2
Hexagonal (H,l) phase, 149
High contrast artwork, 198
High frequency oscillations, 131
Histidine, 89
Histone, 100
Hoarfrost, 14 8
Homeohydric plant, 149
Homocysteine, 89
Homogenous nucleation temperature, 14 9
Homoiohydric, 149, 153
Homologous, 100
Homology, 106
Homonyms, 179
Homopolymeric proteins, 102
Homoserine, 89
lactone, 89
Hormone, 127
binding protein, receptor, sensitivity, 12 7
Hour-glass timer, 131
Hour (h), 7, 12-13
However, 167
Hybridization, 100
Hybrid duplex, 10 0
Hybrids, 24
Hydraulic: conductance and resistance, 6 3
Hydro-, 124
Hydrogen ion concentration, 206, 208-209
Hydropathy plot, 100
Hydrophyte, 15 3
Hydrostatic pressure, 61
Hydroxylysine, 89
Hydroxyproline, 89

224 Index
Hygro-, 12 4
Hygroscopic movement , 12 1
Hyphen(s), 10 , 11, 14, 166, 170 , 177 , 18 5
Hyphenation, 178 , 184 , 18 7
Hyponasty, 12 5
Hypothesis test, 30
difference between two means, 30
difference betwee n two proportions, 31
mean, 30
proportion, 31
two variances, 31
variance, 3 0
Hypoxanthine, 9 1
Hz (symbol for here), 9

Ice:
deletion mutants, 14 9
encasement, 14 9
nucleating bacteria, 14 9
nucleation active bacteria, 14 9
nucleators, 14 9
IDP, 88, 137
IEF, 10 0
Illuminance, 204
Immunodetection, 10 0
IMP, 88
Impaction, 13 7
Imperative, 17 6
Imposed dormancy, 13 6
In (in botanical nomenclature) , 2 4
INA, 149
Incomplete sentence, 16 4
Indefinite articles, 16 9
Independent, 164
clause(s), 166 , 17 6
Indeterminate, 13 7
growth, 112
Index herbariorum, 2 1
Indirect object, 16 5
Induced, 121-122, 137
state, 137
Induction, 100 , 13 7
Infinitive, 17 5
Information, 18 0
Informational slide, 19 1
Infradian rhythm, 13 1
Infrared, 6 6
spectra, 89
Infraspecific taxa , 22-23
Inhibitor, 13 7
Initial, 11 3
Initiation, flower, 137
Innate dormancy, 13 6
Inosine, 91
Input signal, 12 1
Insertion, 10 0
Insertional mutagenesis, 10 0
Intensifies, 200
Interaction, 36
Intercellular freezing, 14 9
Intercept, 32

Interjections), 164 , 16 7
Intermediate-day plant, 137
Intermittent warming, 145
Internal stimulus, 121
International Bureau of Weights and Measures, 4
International Organization for Standardization, 4
International Cod e of Nomenclature fo r Cultivated
Plants, 25-26
International language, 19 4
International Scientific Unions, 93
Intransitive verb, 180
Intrinsic ice nucleator, 149
Introductory or final nonrestrictive phrases, 18 6
Introductory phrases, 17 3
Intron, 100
Intrusive growth, 113
Ionic:
charge, 81
effect, 155 ;
(solution) relations, 56
Ions, 7
Inversion layer, 149
Irradiance, 9, 75, 210
Irregular forms, 176
Iso-electric focusing, 100
ISO Standards Handbook, 4, 15 , 17
Isocitric acid, 90
Isoleucine, 89
Isotopically labeled compounds, 82
ISPMB number, 107
Italic type, 5, 11, 15, 17, 22-23, 25, 17 8
ITP, 88
Its, it's, 180
Joule (J), 9, 12 , 14, 18
Justify the right margin, 184, 18 7
Juvenility, 13 7
Kelvin (K), 5-7, 10 , 12
a-Ketoglutaric Acid, 2-oxoglutorate, 9 0
Killing temperature, 14 9
Kilo (prefix = k), 8 , 10
Kilocalorie (kcal or Cal, obsolete), 8 6
Kilogram (kg), 5, 6, 10
Kilohm (kCi), 10
Kilopascal (kPa), 14 , 18
Kinematic viscosity, 63
Kinematics, 112
Kinesis, 12 3
Kinetic energy, 66
per amount of substance, 67
Kingdom, 22
K-jugate phyllotaxis, 11 3
Kodak pamphlet, 19 7
Kruskal-Wallis k-Sample Test, 40
Krypton-86 atom, 5
L (symbol for liter), 10, 13 , 17
Labeled compounds, 93
Labiatae, 2 2
Lamellar phase, 149

Index 22
Lamiaceae, 2 2
Language conventions with Si-unit names and symbols, 13
Language problems, 19 4
Languages, 1
Lariat, 100
Latency, 12 2
time, 122
Latent energy flux, 70
Latin, 23
square design, 34
Lawn, 100
Lay, lie, 180
LD (long day), 13 1
LDP (long-day plant), 13 7
Le Systeme International cTUnites (SI), 6c Edition, 4
Leaching requirement, 155
Leaf:
area index, 117-118
area ratio, 116-11 8
conductance, 66
mass ratio, 116-118
plastochron index , 113
weight ratio, 11 7
Least squares techniques, 31
Legibility, 190-19 1
Leguminosae, 22
Length, 5, 7
Lettering, 190-19 1
Leucine, 89
Levels of authority, 4
Library, 100
Ligation, 10 0
Light, 204
break, 132, 138
growth response, 12 3
span, 13 2
trap, 12 3
Light-harvesting complex type I, 107
Like, as, 180
Line graphs, 19 1
Linear, 111
model, 33-34, 38
comparisons, 37
Lipids, 94
Liquids:
in gases, 5 6
in liquids, 56
List 177
Liter (L), 4, 10, 12-13, 16
Litre (British, French spelling), 4, 13 , 16
LL (continuous light), 132
Local:
control, 27
derivative, 11 2
Locomotion, 12 0
Locus, 100, 105
Logic in the English language, 163
Logistic function, 112
Long-day plant (LDP), 137
Long-short-day plant (LSDP), 138

Low temperature:
exotherm, 149
injury, 14 9
Lowercase, 9-10, 13 , 23
LPI, 11 3
LR, 155
LT50, 149
LTE, 14 9
Luminous intensity, 5-7
Lux, 8, 10
Lysine, 89
m (symbol for meter), 6, 14 , 17
M (symbol for morgan), 10 1
Magnetic field strength, 9
Main effect, 3 6
Malic acid, 90
Mangroves, 155
Mann-Whitney Two-Sample Test, 40
Mannose, 90
Magneto-, 124
Marking pen, 198
Mass, 5-7, 16
communication, 181
flow, 73
Masking, 132
Material derivative, 112
Mathematical entities, 11
Matric potential, 61, 62
Maturity, 138
Maxwell, 3
May, 180-181
MejSO, 88
Mean, 27
square error, 27
Mean Comparisons, 35
Duncan's test, 37
Fisher's least significant differenc e test, 3 7
Newman-Keul's test, 37
Tukey's test, 37
Meetings presentations, 18 8
Melting point depression, 150
Median, 28
Medium, media, 181
Mega (prefix = M), 8 , 10
Megapascal (MPa), 14
Megaplasmid, 10 0
Megohm (M ) , 10
Member numbers, 106
Membership in a:
plant-wide gene family, 10 6
multigene family, 10 6
Membrane:
fluidity, 15 0
permeability, 15 0
Menaquinone, 8
6-Mercaptopurine, 9 1
ribonucleoside, 91
Meristem, 113, 138
Merophyte, 11 3
MES, 93

226 Index
Mesophyte, 153 , 15 5
MESOR, 13 2
Messenger RNA , 92
Metabolic heat energy, 66
Metastable state, 150
Methionine, 89
Methods and results, 186

Meter (m), 4-5, 9-1 0

Metre (British, French spelling), 4- 5

Metre Convention, 4
Metric:
ton, 10 , 13
metric system, 3
Michaelis constant, 58 , 87
Micro (prefix = u), 8, 11, 15
Micrometer (um) , 14 , 18
Micromolar (obsolete ; use micromoles/liter), 8 6
Micron (obsolete; use micrometer), 14 , 18
Midline estimating statisti c of rhythm, 132
Might, 18 1
Milli (prefix = m), 8, 10
Millimicron (obsolete), 14
Millimolar (millimoles/liter) , 8 6
Mimosaceae, 2 2
Minimum:
leaf number, 13 8
survival temperature, 15 0
Minute (min), 7, 11-1 3
Miohalophyte, 15 5
MissenseDNA, 10 1
Mitochondria, 10 6
Mitochondrial DNA , RNA, 92
Mnemonic, 10 7
Mobility, 58
Mode, 28
Modifier, 17 0
Modifying:
phrases, 172
words, 18 5
mol (symbol for mole), 6, 14
Molality, 1 7
Molar and molal solutions (obsolete), 14, 16
Molar (obsolete; us e moles/liter), 8 6
Molarity, 1 7
Mole, 4- 7
of photons, 14 , 17
fraction, 56, 212
Molecules, 7
Molecular:
mass, 8 3
weight, 83
Monocarpic species, 13 8
Monochromatic light , 8
Monomolecular growth function, 112
Month, 7, 12
MOPS, 93
Morgan (M), 10 1
Movement, 12 0
mechanisms of 120
MSE, 27
Multigene:

families, 10 6
family, 10 1
Multiple:
regression, 3 1
unit, 15
Multiplication, 10-1 1
sign, 24
factors, 1 2
(product) dot, 10
Mutation, 101
N (symbol for newton), 9
JV-Acetylglucosamine, 90
NADandNAD+, 88
NADH, 88
NADP, NADP+, and NADPH, 88
Names, 13,1 4
of units, 1 0
Nano (prefix = n), 8, 10
Nanometer (nm), 14 , 18
Nanomolar (obsolete), 8 6
Nastic movement, 123
Nasty, 123
National Agricultural Library of the USDA, 107
National Physical Laboratory, 4
National Institute of Standards an d Technology, 4- 5
National Bureau of Standards, 4
Negative, 12 4
exponents, 11-12, 15
Neither, 181
Nernst potential, 57
Net assimilation rate, 116-118
Net irradiance, 67 , 69
Nevertheless, 167
Newton (N), 6-7, 9-10 , 1 2
Nifplasmids, 10 0
Night break or interruption, 138
Nightlength, 13 8
NIST, 4
SP 811, 15 , 17
Nomenclature of:
sequenced plant genes, 105
traditional genetics, 10 5
Non-osmotic volume, 63
Nonrestrictive, 173 , 17 4
phrase, 17 2
phrase or clause, 186
phrases and clauses, 177 , 18 3
phrases or clauses, 18 6
Nonsense DNA, 101
Nor, 164, 18 1
Norleucine, 89
Northern blot, 101
Noun(s),22, 164-165 , 169-171, 181, 185
in apposition, 23
Nuclear DNA and RNA, 92
Nuclear magnetic resonance, 89
Nuclear-encoded genes, 106
Nucleic acids, 94
Nucleoid, 101
'Nucleoside, a,'91

Index
Nucleotide(s), 94 , 101
Nucleus, 10 1
Nuclide 12C , 13
Null-response technique , 13 8
Numbers), 4, 14
Number of photons, 75
Numeral(s), 11
to begin a sentence, 1 4
Numerical value, 4
Nutation, 12 5
Nutrient concentration i n plant tissue, 116
Nutrition, 206 , 208-209, 213
Nyctinasty, 12 5
Objective:
case, 171, 185
complement, 16 5
Object(s), 164-165 , 185-186
Obligatory response, 13 9
Oersted (obsolete) , 3
Ohm( ) , 9, 12
Oligohalophyte, 15 5
One-instant mechanism, 12 4
Opaquing material, 19 8
Optical:
density, 8 3
rotary disperson , 8 9
Optically activ e isomers , 8 1
Or, 164, 181
Oral:
presentation, 191 , 194
report, 18 8
Order (ordo), 22
Ordinate, 18 9
Orotate, 9 1
Orotidine, 9 1
Ortho-, 12 4
Orthogonal polynomials , 38
Oscillator, 13 2
Osmoregulation, 15 5
Osmotic:
adjustment, 153 , 155
coefficient, 6 3
effector, 15 5
potential, 61-6 2
potential gradient, 7 4
pressure, 61-6 2
shock, 155
stress, 156
Osmotically generate d flow, 73
Osmoticum, 15 6
Outline slide, 189
Ounce, 1 1
Overhead transparency , 19 3
Oxalacetic acid , 90
Pa (symbol for pascal), 9
Palindrome, 10 1
Palmae, 22
Papilionaceae, 2 2
PAR, 204, 213

Paraheliotropic, 15 3
Parastichy, 11 3
Parentheses, 23 , 173-174, 177 , 186
Parenthetical:
interrupters, 17 7
material, 17 7
phrases, 173, 186
Partial:
molar, 55
molar volume, 57
pressure, 212
pressure of gaseous species j, 66
Participles, 170 , 175, 186
Partition coefficient, 5 8
Partitioning, 7 2
Parts of speech, eight, 164
Parts per:
billion, 1 4
million, 14 , 18
Pascal (Pa), 9, 12
Passive voice, 176, 186
Past:
participle, 170, 175
tense, 17 5
Pause, 166, 177
PCR, 10 2
Pendulum timer, 132
Peptides, 93
Per, 10, 12 , 15, 24
Percent, 18 1
dry mass, 11 8
symbol (%), 11
Percentage, 18 1
by volume, 11
Perception, 12 1
Perennials, 13 8
Period(s), 132, 167, 174, 176-177, 185-186
Perfect, 18 3
Permeability coefficient, 5 8
Personal pronouns, 171, 185
Peta (prefix = P), 8
PFU, 10 1
PGD, 107
pH, 213
Phage, 101
Phase, 13 2
angle difference, 13 2
change, 138
response curve, 132
shift, 13 2
transition temperature, 150
Phenotype, 101, 105
Phenylalanine, 89
Phloem:
export rate, 73
import rate, 74
loading, 73
mass flux, 74
pressure gradient, 74
unloading, 7 3
Phobic response, 122

227

228 Index
Phobism, 12 2
Phobotaxis, 12 2
Phosphoenolpyruvic acid , 90
3-Phosphoglycerate, 9 0
6-Phosphogluconic acid , 90
Phosphoric acid residue, 89 , 92
Phosphorus, 94
Photo, 12 3
growth response, 123
oxidation, 143 , 15 0
Photoassimilate, 7 2
Photofraction, 13 2
Photograph, 19 6
Photographic techniques for creating slides, 197
Photomorphogenic radiation , 20 4
Photon(s), 7, 204
and energy flux, 207
exposure, 76
flow rate, 204
fluence, 204
fluence rate, 204
flux, 76, 204, 210
or energy flux, 208
Photoperiod, 13 2
Photoperiodic, 13 8
Photoperiodism, 13 8
Photosynthesis rates, 16
Photosynthetic irradiance, (PI), 66 , 77, 211
Photosynthetic photon flux, (PPF), 66 , 77, 211
Photosynthetically active radiation (PAR), 66, 77, 204
Photosynthetically active radiation, photon basis, 66
Phototropism, 77, 123
Phraeatophyte, 15 3
Phrase, 164 , 17 3
Phylloquinone, 89
Phyllotaxis, 11 3
Phylum, 22
Physical quantity, 3-4, 10 , 18
Physiological drought, 15 6
Phytochrome, 13 9
far-red absorbing form, red-absorbing form, and
total, 77
Phytohormone, 12 7
Pi; 88
PI, 11 4
Pico (prefix = p), 8
Picomolar (pM; use pmol/L), 86
Pie charts, 191
PIPES, 93
Plagio-, 124
Plain-color (no information) slides, 18 9
Planck's constant, 6 6
Planned comparisons, 3 5
Plant:
growth regulator, 12 7
hormone, 12 7
or fungal material, 21
tissues, 7
Plant-wide family, 106
Plaque forming unit, 101
Plasmid, 101

Plasmolysis, 150
Plastochron:
index, 114
ratio, 114
Plastoquinone, 89
Plating, 101
Platinum, 7
Pleomorphic Fungi, 25
Ploidy, 101
Plural, 13
subject(s), 17 5
verbs, 17 5
Plurals of unit names, 10
Poaceae, 22
Poikilohydric, 153
plant, 150
Poikilotherm, 150
Pointer, 193
Poise, 3
Polar transport, 128
Polaro-, 123
Poly(A)- and Poly(A)+, 101
Polyadenylation, 101
Polycarpic species, 13 9
Polymerase, 101
chain reaction, 102
Polynucleotide, 10 2
Polyol, 156
Polysaccarides, 85
Pooled variance, 29
Population:
mean, 30
proportion, 31
variance, 30
Positive, 124
Possessives, 17 7
Poster, 188 , 19 7
abstract, 196
discussion, 19 7
presentations, 19 5
Post-translational regulation, 10 2
Post-transcriptional regulation, 102
Power (watt = W), 9
ppb, 14
PPj
PPi, 88
88

ppm, 14
Precede, 181
Prechilling, 139
Precision, 206
Precocious, 139
Predicate, 164
Preferred S I unit, 8, 17
Prefixes, 4, 8, 10, 170
Premature termination, 10 2
Prepared speech, 193
Present:
participle, 170, 175
tense, 175
Presentation(s) 188-190
Presenting scientific data, 16 1
Pressure, 9; 62

Index 22
chamber apparatus (theory of), 53
flow, 73
flow hypothesis, 72
potential, 61-6 2
Preposition(s), 164, 171, 179-180, 18 5
Prepositional phrase, 165
Primary:
event, 143 , 145
transcript, 10 2
Primer, 10 2
Principal, principle, 18 1
Printer(s), 184, 187
Probe, 102
Proceed, 18 1
Product, 10 6
Product dot, 11, 15
Projection screen, 19 0
Projectionist, 19 3
Proline, 89
Pronoun(s), 164-165, 169-17 1
Promoter, 10 2
Proof, 18 7
the final manuscript, 18 4
Proportional font(s), 185, 187
Protein structure :
primary, 10 2
quarternary, 10 2
secondary, 10 2
tertiary, 10 2
Proteins, 93
Proved, proven, 18 2
Proximal, 99
Pseudogenes, 10 2
Pseudoreplications, 2 7
Pseudouridine, 91
Psychrometric constant, 6 8
Pteroic acid (pteroyl-), 8 9
Pteroylglutamic acid, 89
Public databases o f plant genes, 10 7
Publication, 188
Publish or perish, 16 1
Published results, 17 5
Pulsed-field electrophoresis, 10 2
Punctuation, 163 , 164, 166, 168, 170, 172, 176
'Purine, a,'91
'Purine nucleoside, a, ' 91
Pyridoxyl-, 89
'Pyrimidine, a,' 91
'Pyrimidine nucleoside, a,' 91
Pyroglutamic acid ; 5-oxoproline, 89
5-Pyrrolidone-2-carboxylic acid , 89
Pyruvic acid, 90
Qualitative response, 139
Quantitative response, 139
Quantity of:
isotope, 59
substance, 7, 59
Quantum:
of radiant energy (a quantum), 66
yield, 77

Question mark(s), 176-177

Quiescence, 139
Quinones, 95
Quotation marks, 25, 177
Quotient, 1 2
Races, 25
Radian, 5
Radiant energy, 66, 75, 203
flux, 6 9
Radiant exposure, 75
Radiation, 203, 206, 210
frost, 148 , 150
Raised period, 11
Random-amplified-polymorphic DNA, 102
Randomization, 27
Randomized Block Design, 33
Range, 28
RAPD, 102
Rate constant, 5 8
Rather, 167
Reading frame, 102
Recalcitrant seeds, 139
Receptor, 102, 121, 139
Reciprocity law in photoresponses:
Bunsen-Roscoe law, 122
Reclamation, 156
Recombinant, 103
Reductive Pentose Phosphate Cycle, 90
Reflectance, 76
Reflection coefficient , 63 , 65, 67
Reflectivity, 6 7
Regnum, 22
REGR, 112
Regrowth, 143, 150
Regulation, 103
Rehearsal for presentation, 19 3
Rejuvenation, 13 9
Relative:
elemental growth rate, 112
growth rate (r), 112, 116, 117, 118
humidity, 67, 205
molecular mass, 87
plastochron growth rate, 114
pronoun(s), 167, 172, 186
water content, 153
RELEL, 11 2
Repair, 143, 150
Rephasing, 132-13 3
Replication, 27
Reporter gene, 103
Represser, 103
Research results, 196
Resistance for gaseous diffusion fo r species/, 67
Respectively, 182
Response adaptation, 122
Rest, 13 9
Rest endo-dormancy, 14 6
Restriction:
enzyme, 10 3
fragments, 10 3

230 Index
-fragment-length polymorphism , 10 3
Restrictive, 17 3
phrase, 172
phrase(s) or clause(s), 186
phrases and clauses, 18 2
Retardation factor, 8 7
Retinoids, 9 5
Revernalization, 13 9
Reversal processing for technical pa n film, 200
Revolutions pe r minute, 8 6
RFLP, 103
Rhamnose, 90
Rheo-, 12 4
Rhythm splitting, 13 2
Riplasmid, 103
Ribonucleic acid, 10 3
Ribose, 9 0
Ribosomal RNA, 92
Ribosylnicotinamide, 9 2
Ribosylthimine, 9 2
Richards growt h function, 112
Right margin, 187
Ripeness-to-flower, 13 9
RNA, 88, 103
editing, 10 3
Roman, 1 5
alphabet, 5
(upright) type, 5, 11
Rosette plant, 14 0
Round brackets, 17 3
Rules for botanical nomenclature, 21
Running, 120
phase, 120
RWC, 15 3
s (symbol for second), 6
S (symbol fo r Siemens), 9
Salination, 15 6
Saline: adaptation , adjustment, sodi c soil, and stress,
156
Salinity, 15 6
threshold, 15 6
SAMPL, 10 3
Sample size, 4 1
Salt:
-affected soil , 15 6
balance, 15 6
glands, 15 6
resistance, 157
tolerance, 157
SAR, 15 7
Sarcosine, 89
Saturated soi l paste, 157
Saturation:
extract, 15 7
percentage, 15 7
Scales, 6
Scarification, 14 0
Science, 16 1
Scientific:
paper, 161

writing, 16 3
Scion, 14 0
Scoreable marker, 10 3
Scoto-, 12 3
Scotonasty, 12 5
SDP (short-day plant), 14 0
SDS, 88 , 104
Seasonal cro p yield, 116
Second (SI unit, s), 5-7, 1 3
Second:
law of thermodynamics, 46
messenger, 10 3
symbols, 1 1
level sources, 1 5
Secondary:
event, 144-14 5
juvenile phase, 13 9
juvenility, 137
Sedimentation coefficient, 8 7
Sedoheptuose-7-phosphate, 9 0
Sedoheptuose 1,7-bisphosphate , 90
Seed-coat-imposed dormancy, 13 6
Segregational:
analysis, 105
nomenclatures, 10 5
Seismo-, 12 3
Selectable marker, 103
Selection, 10 3
Selective amplification of microsatellites polymorphi c
loci, 103
Self-inductive, 13 5
Self-sustained oscillations, 13 2
Semicolon(s), 167-168 , 176-177, 185
Semidian rhythm, 132
Sense-impression verbs, 17 1
Sensible:
energy flux transfer, 69
heat energy transfer, 66
Sensitivity:
of the human eye, 8
to chilling, 146
to growth regulators, 12 7
Sensory, 12 2
Sentence(s), 164 , 167 , 18 5
Sentence fragment, 164, 168 , 17 4
Sentences should not begin with numerals, 11
Sequence, 10 3
Serial comma, 176
Series, 17 6
Serine, 8 9
Sevres, France, 3, 5
Shall, 182
Short-day plant (SDP), 140
Short-long-day plant (SLOP), 14 0
Sialic Acid, 90
Siemens (S), 9-10, 1 2
Sieve-element/companion-cell complex, 73
Sigmoid growth curve, 112
Sign test, 40
Signal, 12 1
peptide, 10 3

Index 23
sequence, 10 3
transduction, 103 , 122
transmission, 122
Simple:
effect, 3 7
linear regression, 31
Since, 18 0
Since, because, 18 2
Single:
family, 10 6
leaf photosynthetic rate , 117
Singular: nouns , subjects), and verbs, 175
Sink, 73
Sinusoidal:
curve, 13 2
rhythm, 13 3
Sit, 18 0
Skeleton photoperiod , 133
Slash, 11-12 , 15, 178
Slide(s), 188, 190-192, 197-20 0
Slide presentations, 18 9
SLDP (short-long-day plant) , 14 0
Slope, 31
Snow mold, 150
So, 164, 182
...as, 180
-called, 18 2
Sodicsoil, 15 7
Sodicity, 15 7
Sodium:
adsorption ratio, 157
dodecyl sulfate, 104
Software, 188-18 9
Soil and liquid temperatures, 207-208
Soil-plant-atmosphere continuum , 153
Solar:
irradiance, 67
tracking, 123
Solids: in gases, liquids, and solids, 56
Solidus, 1 2
Solonchak soils, 15 5
Solonetz soils, 15 5
Soloth soils, 15 5
Solute:
flux, 62
potential, 61 , 62
permeability, 6 2
Solution concentrations, 1 6
Source, 7 3
Southern blot, 10 4
SP 811 (NIST publication) , 4
SPAC, 15 3
Space, 5
between numerals and units, 15
to group numerals, 1 2
Spatial sensing of direction, 12 4
Special Publication (SP) 811, 4
Species, 22-23
of plant, 10 6
Specific:
activity, 59

electrical conductance, 5 8
epithet, 23
growth rate, 116-11 7
heat capacity of dry air, 66
latent heat of fusion, 6 7
latent heat of vaporization, 6 7
leaf area, 116-118
leaf mass, 116, 118
rotation, 8 7
Spectral:
distribution, 20 3
energies, 1 6
energy flow rate, 203
energy fluence, 204
energy fluence rate, 204
energy flux, 75, 203, 211
irradiance, 9, 75, 211
photon fluence rate, 20 4
photon flux, 76, 204, 211
photon or energy flux, 207-208
Spectrophotometric data, 83
Spell checker, 18 4
Spelled-out unit names, 11
Spliceosome, 10 4
Splicing, 10 4
Split gene, 10 4
35 S promoter, 97
Square:
brackets, 24 , 173-174, 177
meter, 1 6
Squares, 11
Standard:
unit, 5
"gravity", 1 4
acceleration due to gravity, 8-9 , 12-14 , 17
deviation, 28
Standard error of:
estimate, 32
difference betwee n two means, 29
mean, 28
slope 32
Standards, 3
Statistical tables, 191
Steady growth, 11 4
Stefan-Boltzman:
Constant, 6 8
Law, 69
Step-up, step-down, 12 4
Steradian, 5
Stereochemistry, 93
Steroids, 94
Stimuli, 12 3
Stimulus, 121, 12 3
perception, 12 1
transduction, 122
Stock, 140
Stokes, 3
Stopcodon, 10 4
Strain, 21, 14 4
Stratification, 139-14 0
Stress, 14 4

232 Index
Strophiolar clef t or plug, 137
Strophism, 123
Style:

commas, 17 7
conventions, 1 0
manuals, 163 , 165
Subcooling, 15 0
Subject, 164-165, 173, 175, 185-186
complement, 186
Subjective:
complement, 165
and objective cases, 171
Subjective day or night, 13 3
Subjunctive mood, 18 3
Subordinate:
clause, 16 8
phrase, 185
Subordinating conjunction, 167, 185
Subspecies, 22
Substrate, 206, 209, 212
Subtraction library, 104
Successor apical cell, 113
Succinic acid, 90
Succulence, 157
Succulents, 15 3
Suffixes, 17 0
Sutfate salinity, 157
Summer dormancy, 14 0
Sunscald, 15 0
Supercooling, 15 0
Superoxide dismutases, 106
Superscripts (an d SI units), 11
Supplementary units, 5
Suppression, 10 4
Surface tension, 63
Susception, 12 1
Swedberg (not SI), 86
Symbol(s) 13-14, 17
Symbols for the monomeric units, 85
Symplastic growth, 11 4
Symptoms of injury, 144 , 146
Synchronizer, 13 3
Synonym, 24
t (symbol for metric ton), 13
T50, 151
T-region of Ti plasmid, 104
Tables, 183, 191
Tactic orientation, 12 2
TAPS, 93
Tau, 131
Taxis, 122
Taxon, 22, 24
Taxonomic groups, 2 2
TDP, 88
TEAE-cellulose, 88

Technical writing, 172-173, 176

Teleomorph, 2 5
Telomere, 10 4
TEMED, 93
Temperature, 5 , 7, 67, 205-206, 211

dewpoint, 205

dry bulb, 205

gradient, 1 6
-sensitive mutation , 10 4

wet-bulb, 205
Template, 104
Temporal sensing of direction, 124
Tense, 175 , 186
Tera (prefix = T), 8, 10
TES, 93
Test of hypothesis, 30
Tetrapyrroles, 95
Text, 190
slide, 189-190
Thawing, 151
Than, 18 2
That, 172-174, 182, 186

The International Syste m of Units (SI Units), 3-2 0

Then, 18 2
Theory of minimum artical cell volume, 15 1
Thermal conductivit y coefficient o f region j, 66
Thermo-, 12 4
Thermoavoidance, 15 1
Thermocouple hygrometers, 5 3
Thermodynamic temperature, 5- 7
Thermoperiodism, 14 0
Thermophile, thermophilic , and thermophily, 15 1
Thermostability an d thermotolerance, 15 1
Thickness of:
air boundary layer, 6 8
unstirred layer, 63
Thigmo-, 12 3
Thiohypoxanthine, 9 1
6-Thioinosine, 9 1
Thiouridine, 92
Third person, 17 1
Threonine, 89
Thus, 16 7
Thus, thusly, 18 3
Thymidine, 9 2
Thymine, 91
Ti plasmid, 10 4
Time, 5, 7
Times earth's gravity (g n), 87
Tissue sample, 16
Title(s), 183, 18 9
TK50,151
TMP, 88
Tmax and Tmin, 151
Tocopherol, 89
Tocopherolquinone, 89
Tocopherols, 9 5
Tolerance, 144
Tonic effect, 12 2
Tonne, 1 3
Topotaxis, 12 2
Torsion, 12 1
Toward, towards, 18 3
Tranpiration or condensation, 67
Trans, 104
Transcribed, 10 4

Index 23
Transcript and transcription, 10 4
Transduction, 104
chain, 12 2
Transfer RNA, 92
Transformation, 10 4
Transient:
cycle, 133
gene expression, 10 4
Transit piptide, 10 5
Transitive verb, 176, 180
Translation, 10 5
Translocatable hardines s promotor, 151
Translocation, 72
profile, 7 3
speed, 74
Transmittance, 76
Transon, 105
Transport number, 58
Transposable elements , 98
Transposition, 10 5
Transposon, 10 5
Transversal, 12 4
Traumato-, 12 4
Treatments, 3 9
Tricine, 93
Triple point of water, 7
Tris, 88, 93
Tropism, 12 3
Tropistic movement, 12 3
True halophyte, 155
Tryptophan, 89
TTP, 8 8
Tumbling, 120
phase, 12 0
Turbulent air, 67
Turgor movement, 121
Twintron, 10 5
Two-instant mechanism, 12 4
Two variances, 3 1
Type size, 196
Typefaces, 19 1
Tyrosine, 89
u (symbol for unified atomic mass unit), 9, 13-14, 17
Ubiquinone, 89
UDP, UDP-Glc, UDP-GalNAc, UDP-Gal, UDPGlcUA, UDP-GlcNAc, and UDP-Xyl, 88
Ultradian:
oscillations, 13 1
rhythm, 133
Ultraviolet, 67, 89
Umbelliferae, 2 2
UMP, 88
Undercooling, 15 0
Underlining, 11 , 15, 22-23, 178
Unfamiliar names, 24
Unified atomi c mass unit (u), 8-9, 12-14 , 17
Unique, 183
Unit, 5
leaf rate, 117
name(s), 10-11 , 14

symbol(s), 10-1 1
Units:
used with the SI but not officially part of SI, 13
of concentration, 86
Universal gas constant, 6 7
Unnecessary words, 178, 186
Uppercase, 13 , 25
Upstream, 105
Uracil, 91
Undine, 92
Usage, 1, 163, 165-167
UTP, 88
V(symbol for volume, electric potential), 9
V(symbol for volt), 9
Valine, 89
Vapor density deficit, 15 3
Vapor pressure:
leaf and air, 67
deficit, 15 3
Variance, 28
components, 38
Varieties, 22, 25
Vector, 10 5
Velocity, 58
wind speed, 67
Verb(s), 164-165, 170, 173, 175, 185-186
tense, 175
Vernacular name, 25
Vernalin, 140
Vernalization, 140
Viability, 144, 151
test, 14 4
Viable, 140
Viewgraph, 188
Viscosity, 63
Visual browning, 151
Vitamin:
B6, 95
D, 95
Volt (V), 9
Volta, 12
Volume, 5, 7, 9, 16
flux, 62
modulus of elasticity, 63
Volumetric heat capacity, 66
VPD, 15 3
W (symbol for watt), 9
Water:
deficit, 15 3
evaporation site, 66
potential 11 , 47, 62
potential in the vapor state, 50
potential of aqueous solutions, 53
status, 154
stress, 15 3
use efficiency, 15 4
vapor, 67
Water vapor, 67
deficit, 20 5

234 Index
density, 20 5 Wor
Watering, 205, 208 , 21 2 Worl
Watt (W), 7 , 9, 12 Writin
Wave lengths, 5 , 8, 68, 76 WUE,
Wave number, 76
Week, 7, 1 2 Xanthine
Weight, 6 Xanthosine
Were, 18 3 Xeromorphy
Western blot, 105 Xerophily
Which, 172 , 174 , 182 , 18 6 Xerophyte
Which hunt, 17 2 Xylose
White frost, 14 8 Xylulose-5-phosphate
Who, 173 , 182 , 18 6
Who, whom, 173 , 18 2 YAC
Wilcoxon's Signe d Rank Test, 40 Year
Will, 18 2 Yeas
Winter: Yoct
annuals, 14 0 Yott
chilling, 15 1
injury, 14 6 Zeitgeber
Wish, 18 3 Zept
Word processing, 184 , 18 7 Zett
Words with special problems, 17 9

k (W), 9
d Directory, 2 1
g conventions, 16 3
154
, 91
, 92
, 154
, 154
, 154 , 15 7
, 90
,9 0
, 105
, 7, 12
t artificial chromosome, 10 5
o (prefix = y), 8
a (prefix = Y), 8
, 13 3
o (prefix = z), 8
a (prefix = Z), 8