A BRIEF INTRODUCTION TO PLANT BIOLOG

Thomas L. Rost
Michael G. Barbour
Robert M. Thornton
T Elliot Weier
C. Ralph Stocking
Botany
A Brief Introduction
To Plant Biology
 Botany
A Brief Introduction
To Plant Biology
Thomas L Rost Michael G. Barbour
•

Robert M. Thornton • T. Elliot Weier

C. Ralph Stocking

University of California • Davis, California

John Wiley & Sons

New York • Santa Barbara Chichester
• • Brisbane
Toronto
Copyright © 1979, by John Wiley & Sons, Inc.

All rights reserved. Published simultaneously in Canada.

Reproduction or translation of any part of

thiswork beyond that permitted by Sections
107 and 108 of the 1976 United States Copyright
Act without the permission of the copyright
owner is unlawful. Requests for permission
or further information should be addressed to
the Permissions Department, John Wiley & Sons.

Library of Congress Cataloging in Publication Data:

Main entry under title: Botany.

Based on Botany, An Introduction to Plant
Biology, 5th ed., by T. E. Weir, C. R. Stocking,
and M. G. Barbour.

Includes index.

1 . Botany. I. Rost, Thomas L. II. Weir,

Thomas Elliot, 1903— Botany, An Introduction
To Plant Biology 5th ed.

QK47.B775 581 78-5433

ISBN 0-471-02114-8

Printed in the United States of America

1098765432 1
 preface

Our objective was to provide

shortened version of Botany:
an abridged and
An Introduction
photographs that are large, detailed, and numerous
enough to be true learning aids. (3) A traditional
to Plant Biology, Fifth Edition, by Weier, pedagogic organization written in a clear and direct
Stocking, and Barbour. However, this text is style. (4) An extensive glossary that defines
significantly different from that text and is more than frequently used terms in the text and that shows
a simple abridgement. their etymological derivation.
Like the parent text, this is intended for We take pleasure in acknowledging the following
introductory courses at the university, college, or individuals for their help in the development of the
community college level. Prior courses in biology, manuscript in whole or in part or for providing the
mathematics, or physics are not required, but some materials that are in this book: Alice B. Addicott,
acquaintance with elementary inorganic chemistry is Dorothy Brandon, Dr. Edward Butler, Robin Camp,
helpful for understanding Chapters 2, 5, and 6. Dr. Norma Lang, Jacqueline Lockwood, Walter
Appendix A introduces the basic ideas of chemistry Russell, Lorna R. Thornton, and Dr. John Tucker.
that are needed, and should be valuable for students We also wish to thank the following individuals who
who have no background in chemistry. Much of the reviewed the manuscript: Charles H. Cochise
Field,
parent text has been rewritten and many new College; Jerry Davis, University of Wisconsin, La
illustrations have been added. Topics in the fifth Crosse; David Dallas, Northeastern Oklahoma A& M
edition of Botany by Weier et al. that have been College; and Mary McLanathan, Foothill College. We
extensively revised include: metabolism, absorption are especially grateful to Ted Barkley of Kansas
and transport, photosynthesis, growth, algae, fungi, State University for his meticulous and detailed
and angiosperms. Many detailed new drawings by suggestions on the entire manuscript. Others who
Alice B. Addicott and Jacqueline L. Lockwood provided illustrations are cited at the end of the
accompany these and other chapters. All the book. The many students who have provided
drawings convey important information in a dramatic suggestions for the improvement of the general
manner, but we think that some are original enough botany course taught at the University of California,
in themselves to be contributions to botany. The
Davis, cannot be acknowledged individually, yet they
chapters on bacteria and viruses in the larger text should be aware of our appreciation for them
have been deleted in this adaptation, and much of collectively. We apologize for the unintentional
the material on genetics has been condensed and
omission here of others who have contributed to the
placed in Chapters 3, 7, 10, and Appendix B.
text.
We believe that this text has several features not
shared by any other botany books of comparable
size: (1) A complete, unbiased coverage of botanical Thomas L. Rost
topics with equally detailed sections on anatomy, Michael G. Barbour
cytology, economic botany, ecology, evolution, Robert M. Thornton
morphology, physiology, taxonomy, and a survey of T. Elliot Weier
the plant kingdom. (2) Many original drawings and C. Ralph Stocking
 Digitized by the Internet Archive
in 2011

http://www.archive.org/details/botanybriefintroOOrost
 5 1 1 2

contents

CHAPTER 1 INTRODUCTION 1 DEVELOPMENT OF TISSUES OF THE PRIMARY PLANT

THE SCOPE OF BOTANY 1 BODY 47
ANCESTRY AND CLASSIFICATION OF PLANTS The Primary Meristems 47
Primary Tissues 48
CHAPTER 2 METABOLISM 3
The Primary Vascular Tissues 53
PRINCIPAL MATERIALS 3
SUMMARY OF PRIMARY TISSUES 56
Raw Materials 3
THE DICOTYLEDONOUS STEM 57
Common Metabolic Products 3
Stem Primary Growth 57
ENZYMES AND CATALYSIS 7
Stem Secondary Growth 58
PHASES OF METABOLISM 8
THE MONOCOTYLEDONOUS STEM 64
Photosynthesis, Anabolism, and Catabolism
Primary Growth 64
Respiration 9
Secondary Growth 66
Alcoholic Fermentation 1

STEM MODIFICATIONS 66
THE CONTROL OF METABOLISM 1

Rhizomes 66
The Local Control Within Metabolism 12
Corms 68
The Enzyme Quantities
Control ot 1

Bulbs 68
The Mechanism of Protein Synthesis 12
Tubers 68
The Control of Protein Synthesis 15
SUMMARY 1
Tendrils 68
Cladodes 68
CHAPTER 3 THE PLANT CELL 17 Spines and Thorns 69
CELL STRUCTURE 17 Stolons 69
Technique 18 SUMMARY OF SECONDARY GROWTH 69
Structure and Function 18
The Living Cell 19 Part II Leaves 69
Cell Fine Structure 20 LEAF COMPONENTS 71
Membranes 2C The Blade 71
Ribosomes 20 The Petiole 71
Organelles 23 Simple and Compound Leaves 72
Nucleus 25 Venation 73
Nonprotoplasmic Portions of the Cell 30 ANATOMY OF THE FOLIAGE LEAF 73
SUMMARY 31 Epidermis 73
THE CELL CYCLE 32 Mesophyll 76
Mitosis 32 Vascular System 76
Cytokinesis 36 The Petiole 77
Meiosis 38 LEAF DEVELOPMENT 77
Meiosis — Division
First 42
LEAF SHAPE 79
Meiosis — Second Division 42
Internal Control 79
Recombination Genes of 43
80
Environmental Factors
SUMMARY OF THE CELL CYCLE 43
Leaf Modifications 80
DNA AND ITS REPLICATION 43
LEAF ABSCISSION 81
Replication of DNA 43
SUMMARY— LEAVES 84
CHAPTER 4 THE PLANT BODY 45
Part Stems
I 45 Part III Roots 84
STEM MORPHOLOGY 45 FUNCTIONS OF ROOTS 84
Arrangement of Leaves and Buds 45 TYPES OF ROOT SYSTEMS 85
Position of Buds on a Woody Twig 46 Development of Root Systems 86

VII
 55
27 75 73 1

INTERNAL ANATOMY 88 VARIATIONS IN FLORAL STRUCTURE 125

Root Apex 88 General Description 125
Formation of Primary Tissues 89 Union of Flower Parts 1 28
Secondary Growth in Root 91 Inflorescences 129
SPECIAL ROOTS 93 ANGIOSPERM LIFE CYCLE 130
Mycorrhizae 94 The Androecium 130
Bacterial Nodules 94 The Gynoecium 134
Cont r actile Roots 94 Embryo Sac Development 1 35
SUMMARY 94 Fertilization 1 35

POLLINATION 1 36

Pollinating Vectors 137

CHAPTER 5 THE ABSORPTION AND
SUMMARY 139
TRANSPORT SYSTEMS 96
WATER ABSORPTION AND TRANSPORT 96 Part II The Fruit 140
Water Potential and Water Movement 96 DEVELOPMENT OF THE FRUIT 1 40
The Mechanism of Water Movement in the Plant 99 KINDS OF FRUITS 140
Root Pressure and Guttation 100 Simple Fruits 141
How Roots "Locate" Water 101 Compound Fruits — Formed From Several Ovaries 144
CARBON DIOXIDE ABSORPTION 101 KEY TO FRUITS 146
CONTROL OF CARBON DIOXIDE ABSORPTION AND
Part The Seed
III 146
WATER LOSS 101
SEED DEVELOPMENT 146
MINERAL ION ABSORPTION 103
KINDS OF SEEDS 148
Principal Mineral Elements 103
Common Bean 148
The Absorption of Minerals 105
Grasses 149
OXYGEN ABSORPTION 106
Onion 149
TRANSPORT OF ORGANIC MATERIALS 1 07
DISSEMINATION OF SEEDS AND FRUITS 1 50
Diffusion and Protoplasmic Streaming 107
Agents in Seed and Fruit Dispersal 150
The Mechanism of Phloem Conduction 107
SEED DORMANCY AND GERMINATION 1 50
SUMMARY 109
SUMMARY 154

CHAPTER 6 PHOTOSYNTHESIS 111 CHAPTER 8 THE CONTROL OF GROWTH AND

THE DISCOVERY OF PHOTOSYNTHESIS 1 1 DEVELOPMENT 155
THE CHLOROPLAST— SITE OF ENVIRONMENTAL ADAPTION BY THE YOUNG
PHOTOSYNTHESIS 1 1 PLANT 155
THE THYLAKOID (LIGHT) REACTIONS 1 1 HORMONES AND THE CONTROL OF GROWTH AND
The Light-Capturing System 113 DEVELOPMENT 56 1

Reaction Centers 1 1 Auxins 156

The Transfer of Excitation Energy 1 1
Gibberellins 161

Electron Transport 1 1
Cytokinins 163
Energy Storage by the Thylakoid 1 1
Ethylene 165
Abscisic Acid 1 66
Summary of the Thylakoid Reactions 1 1

C0 2 REDUCTION AND THE FORMATION OF SUGAR The Practical Use of Hormones 167
(DARK REACTIONS) 1 1
LIGHT AND THE CONTROL OF DEVELOPMENT 1 68
EFFICIENCY OF PHOTOSYNTHESIS 120 Seed Germination 168

THE ENVIRONMENT AND PHOTOSYNTHESIS 20 1

Shoot Growth 168

Temperature 1 20 PHOTOPERIODISM AND FLOWERING 168

Light 121
TEMPERATURE AND DEVELOPMENT 1 70

Carbon Dioxide 121

SUMMARY 1 72

Minerals 121
CHAPTER 9 PLANT ECOLOGY 173
SUMMARY 121
COMPONENTS OF THE ENVIRONMENT 173
Moisture 1 74
CHAPTER 7 FLOWERS, FRUITS AND Temperature 1 75
SEEDS 123 Light 175
Part The Flower
I 123 Soil 1 77
FLOWER STRUCTURE 1 23 Fire 180
Carpels and Stamens as Modified Leaves 1 24 Biological Factors 181

Contents

VIII
 5

ECOLOGY AND PLANT POPULATIONS 1 83 CHAPTER 12 THE MYCOTA (FUNGI) 253

ECOLOGY AND PLANT COMMUNITIES 1 84 CLASSIFICATION OF THE FUNGI 253
How May Communities Be Measured? 1 85 Division Mycota (the fungi) 253
Succession 1 86 SUBDIVISION MYXOMYCOTINA 253
VEGETATION TYPES OF THE WORLD 1 87 SUBDIVISION EUMYCOTINA 254
Tundra 1 87 Class Oomycetes 255
Taiga 188 Class Zygomycetes 257
Deciduous Forest 189 Class Ascomycetes 260
Tropical Rain Forest 189 Class Basidiomycetes 265
Savannah and Prairie 190 Class Fungi Imperfecti 270
Scrub 191 THE LICHENS 272
A final note 191 Asexual Reproduction 273
ECOLOGY AND ECOSYSTEM 192 Sexual Reproduction 273
Energy Flow 192 SUMMARY 273
Pollution 192
SUMMARY 193
CHAPTER 13 BRYOPHYTES 274
GENERAL CHARACTERISTICS 274
Economic Importance 275
CHAPTER 10 PLANT TAXONOMY AND Classification 275
EVOLUTION 195 CLASS MUSCI 275
HISTORICAL ASPECTS 195 General Characteristics 275
Pre-Linnaean Period 195 Gametophyte 275
Post-Linnaean Period 196 Sporophyte 276
HOW ARE PLANTS CLASSIFIED? 1 97 Summary of Moss Life History and
Plant Groups 1 98 Characteristics 280
Herbaria and Botanic Gardens 198 CLASS HEPATICAE 280
METHODS OF TAXONOMIC RESEARCH 1 99 General Characteristics and Distribution 280
Traditional (Morphological) 199 Riccia 281
Anatomical 203 Marchantia 285
Biochemical 203 CLASS ANTHOCEROTAE 286
Biological 205
Numerical 206
CHAPTER 14 LOWER VASCULAR
PLANTS 288
EVOLUTION 207
DIVISION PSILOPHYTA 288
The Geologic History of Plants 207
DIVISION LYCOPHYTA 288
Charles Darwin and Evolution 215
Lycopodium 289
The Role of Natural Selection 218
Selaginella 291
How Species Remain Distinct 218
DIVISION SPHENOPHYTA 297
SUMMARY 219
Mature Sporophyte 297
Reproduction 297
CHAPTER 11 ALGAE 222 DIVISION PTEROPHYTA 297
ECONOMIC IMPORTANCE OF ALGAE 224 Mature Sporophyte 302
Ecological Functions 224 Reproduction 302
Economic Uses 227 SUMMARY 305
ALGAL CLASSIFICATION 234
Cyanophyta: the Blue-Green Algae 234
CHAPTER 15 GYMNOSPERMS 310
DIVISION CONIFEROPHYTA 310
Rhodophyta: the Red Algae 234
Classification 310
Euglenophyta 234
Life History of a Conifer 31
Chlorophyta: the Green Algae 237
The Conifer Life Cycle in Perspective
Bacillariophyta: the Diatoms 237
DIVISION CYCADOPHYTA 324
Pyrrhophyta: the Dinoflagellates 237
DIVISION GINKGOPHYTA 324
Phaeophyta: the Brown Algae 237
DIVISION GNETOPHYTA 324
REPRODUCTION 239 SUMMARY 324
Asexual Reproduction: Vegetative 239
Asexual Reproduction: Mitospores 242 CHAPTER 16 ANGIOSPERMS 328
Sexual Reproduction 242 REVIEW OF LIFE CYCLE 328
SUMMARY 252 Male Gametophyte 328

Contents

IX
 Female Gametophyte 328 APPENDIX A BASIC IDEAS IN
Fertilization and Seed Development 328 CHEMISTRY A1
COMPARISON OF ANGIOSPERM LIFE CYCLE WITH
MORE PRIMITIVE PLANTS 330 APPENDIX B GENETICS SUPPLEMENT B1
Protection of the Female Gametophyte 330
Size of Gametophytes 330
APPENDIX C TABLE OF METRIC
Nourishment of the Developing Embryo 330
EQUIVALENTS C1
Fruit 331
EVOLUTION IN THE ANGIOSPERMS 331
The Besseyan System 331
APPENDIX D GLOSSARY D1
SELECTED FAMILIES OF ANGIOSPERMS 332
Magnoliaceae to Lamiaceae 332
APPENDIX E ILLUSTRATION CREDITS E1
Magnoliaceae to Asteraceae 335
Monocotyledonous Line from Magnoliaceae to INDEX OF GENERA 11

Orchidaceae 338
SUMMARY OF ANGIOSPERM EVOLUTION 344 INDEX OF SUBJECTS 17

Contents
 1 CHAPTER

1 introduction

The Scope of Botany

Botany began with tribal lore about edible, medicinal, and, thus, possess the flexibility needed for cooperative
and poisonous From this narrow focus on
plants. movements such as muscle contraction.
familiar leafy plants and mushrooms, curiosity Even though most have walls, there are
plant cells
spread to diverse forms until today more than 550,000 major differences and composition among
in wall structure
kinds, or species, of organisms are identified as part of the organisms of the plant kingdom. In green plants the
the plant kingdom. New species are found continuously strength of the walls results from a network of cellulose
because there are still regions of the world that have not fibers. In the fungi chitin is usually found instead of
been thoroughly explored: the tropics, with their lush rain cellulose, while the bacteria and blue-green algae have
forests; the arctic; and the microscopic worlds of soils, walls with a fishnet structure built from polymers of
oceans, and sediments. another, more complex set of subunits. These major
Perhaps in the earliest days of botany the field could differences in wall structure create a suspicion that the
easily be defined as the study of life forms that are rooted fungi, the bacteria, and the rooted green plants may be
and essentially immobile. But the identification of only remotely related.
additional species and more detailed study have erased The ability to perform photosynthesis is an extremely
any clear boundaries. Thus for example the mosses, or important property of plants (Fig. 1 .5). This process
Bryophytes (Fig. 1.1), have always been considered enables the organism to trap radiant energy from sunlight
"plants" and appropriate subjects for botanists to study. in order to construct organic materials. The foods
But in its early development the moss plant consists of produced by photosynthetic organisms are essential not
green, threadlike filaments that resemble certain species of only for the organisms themselves but also for life forms
aquatic organisms, the algae (Fig. 1 .2). Furthermore, both such as animals (including human beings) that cannot trap
the moss and the filamentous alga have a phase of the life sunlight. However, some of the "plants" discussed in this
cycle in which they produce free-living reproductive cells book do not perform photosynthesis. An example is
(Fig. 1 .3). These cells swim by means of flagella "Indian pipe," a parasitic plant that has roots, stems, and
resembling those of animal sperm cells. Still other algae flowers (Fig. 1 .6). Most bacteria do not photosynthesize;
spend their whole lives as actively swimming, flagellated nor do any of the 200,000 species of fungi (Fig. 1 .7). We
single cells. These discoveries confirm the fact that true have no reason to suspect that the fungi ever had any
natural boundaries between groups of organisms are hard photosynthetic ancestors. It is clear that botanists study
to find. these life forms because they have cell walls and some of

Is there any constant feature that is characteristic of all the life cycle characteristics of the green, photosynthetic
the organisms that botanists study and not of other forms plants.

of life? The answer is, "not quite." But two features — the
presence of cell walls and the ability to perform
photosynthesis — almost serve that purpose and are worth Ancestry and Classification
comment.
special
of Plants
Whenever large, complex forms of life are closely
inspected, they are found to be composed of numerous
microscopic units of living material called cells. In all but Because of the difficulties just described, not all botanists
one kind of organism that botanists study, each cell is agree on the proper way to sort and group the organisms
surrounded by a tough, fibrous cell wall. The walls of included in the Plant Kingdom. Nevertheless, classification
adjacent cells are cemented together, giving the plant as a is both a practical necessity and an important intellectual
whole a rigid shape and preventing individual cells from goal of botanists. In this regard the highest goal is to

moving. The one exception is a small group of organisms organize a true or natural classification of the organisms.
known as slime molds (Fig. 1 .4), which do not have walls The idea of a natural system depends on the belief that

during most of their life cycle. In this feature the slime present-day organisms are related by common ancestry
molds are like animals. Animal cells do not have walls, and that the differences we observe between organisms
 ^,
bacteria*

prokaryotes

blue-green algae*

original
green plants*
ancestor

fungi*
eukaryotes

unicellular
eukaryote
ancestor
animals

Figure 1 .8 Ancestral relationships between modern organisms

as deduced trom protein structure and the structure and function
of cells. The starred groups are all studied in botany.

are a result of extensive changes in heredity (the process common heredity. Two organisms are not likely to have
of organic evolution) over the three billion years that arrived at similarly constructed proteins by chance. The
plants have been on Earth. In a perfect natural proteins built by eukaryotes and prokaryotes are similar
classification organisms would be grouped according to enough to indicate that they arose from a common
the closeness of their ancestry. Of course the construction ancestor, but different enough to suggest that the
of such a classification must depend on indirect evidence, eukaryotes and prokaryotes diverged long before there
since human observers did not trace events earlier than a were any organisms higher than unicells and before there
few thousand years ago. This evidence includes fossils of were any distinctions between plants and animals.
ancient plants as well as observations of similarities and Probably the most significant way to divide up the large
differences between present-day plants (Chapter 10). group of eukaryotes is according to whether or not they
The most fundamental dividing line among life forms have chloroplasts. These are bodies in the cell that
separates the prokaryotes from the eukaryotes (Fig. 1.8). perform photosynthesis. Animal and fungal cells lack
The differences can be seen in the structure of the cells. them. They are present in all the trees and shrubs and
One of several fundamental differences is that prokaryotes nearly all the herbs that make up our familiar landscape.
have their hereditary material (DNA) floating free in the They are also present in a multitude of less familiar forms,
same fluid mass as the rest of the cellular material, includingmany microscopic unicells.
whereas the eukaryotes have their DNA separated from There is evidence to suggest that chloroplasts may be
the rest of the cell contents by a surrounding membranous the descendants of free-living bacteria that entered the
envelope. All the common plants are eukaryotes whereas cells of early eukaryotes, forming a symbiosis that became
the bacteria and the blue-green algae represent the permanent. Comparable symbioses can be seen today
world's only known prokaryotes. The entire animal between bacteria and plants in the root nodules of
kingdom consists of eukaryotes. legumes (Chapter 5), but in these, the partners are still

One of the most recent and promising tools for judging separable. If chloroplasts did arise from symbiosis, there
the hereditary relations between species depends on may have been many points in evolution where various
examining and comparing protein molecules. The species acquired chloroplasts. Alternatively, chloroplasts
hereditary information that is passed from generation to may have originated within a single line of ancient cells,
generation consists largely of instructions for building without symbiosis. These questions remain to be resolved,
protein molecules. Since the number of possible different and they leave us uncertain about how far back we must
proteins is astronomical, a high degree of similarity reach to find ancestral connections between the bacteria,
between the proteins in two organisms indicates a the fungi, and the green plants.

chapter 1 l
Introduction
 A common moss.
Figure 1.1
COLOR PLATE 1

Figure 1 .2 A
filamentous
green alga of the genus
Stigeoclonium.

Figure 1 .3 A stage of
later
development the alga
in
Stigeoclonium. The small free
green cells are swimming
reproductive cells.

jiX*l*f
Figure 1 .4 Plasmodium of
the slime mold Physarum
COLOR PLATE 1

polycephalum. (continued)

Figure 1 5 A species of
Mimulus, a green flowering
plant.

Figure 1 6 The
nonphotosynthetic flowering
plant Monotropa uniflora.

Figure 1.7 The fruiting body

(basidiocarp) of Amanita
muscaria, a true fungus.
 CHAPTER

2 metabolism

The life cannot be understood without

of the plant
1
Common Metabolic Products
discussing its chemistry.
Since muscular
movements and nervous responses are lacking, the There is a noticeable difference between the simple raw
visible signs of life in most plants are limited to slow materials that the plant takes in, such as C0 2 H 2 0,
, and
changes such as the growth of organs. But if we consider mineral elements, and the complex final products of
the units of matter called molecules, of which plants are +
metabolism. For example, the compound NAD which we ,

composed, the plant body proves to be a place of +

will meet again later, has the formula C 21 H 28 14 N 7 P 2 and
incessant, complex activity. This chemical activity is is formed from raw materials by a complex series of
known collectively as metabolism. occurs throughout It
reactions.
the plant body, at all stages of life except the state known Nearly all the molecules formed in metabolism contain
as dormancy. (We shall meet the subject of dormancy carbon and hydrogen; other elements may or may not be
later in discussing seeds and buds.) With its thousands of present. Such molecules are called organic compounds
different chemical reactions, metabolism makes the plant because they are rarely found in nature except as the
body millions of times more active chemically than the products of metabolic activity by living organisms. By
surrounding nonliving environment. contrast, the simple molecules that plants take in as raw
materials are termed inorganic compounds.
Organic compounds are produced in such great variety
Principal Materials that only a few types can be discussed here. But there are
several classes of organic compounds that have universal
importance and can be used as the basis of an
Raw Materials introduction to metabolism.
Carbohydrates are composed of the elements C, H,
The raw materials from the environment and
plant extracts
and O; they have the general formula (CH 2 0) n where n
converts them into a great variety of more complex
may be any number. The simplest carbohydrates are the
products.
sugars, which are important both as sources of energy
Water is the substance that plants take up in the
and as building units in the construction of many other
greatest quantity. The plant acts as a wick between the
kinds of compounds.
moist soil and dry air, about 90% of the water that enters
Let us consider briefly the structure of a sugar
the plant is later evaporated away. The remaining 10%
molecule. In one group of simple sugars, the hexoses,
remains in the plant, where it provides bulk, serves as a
each molecule contains six carbon atoms and has the
medium for storing and transporting dissolved materials,
general formula C 6 H 12 6 For example, glucose has the
.

and contributes atoms to the metabolic system. Water is

directly consumed or produced in many reactions. Having
straight-chain structure shown in Fig. 2.1 A The CHO —
end is an aldehyde group. The sugar molecule may occur
the formula H 2 0, water is a major source of the elements
in two forms. When not in solution the carbon atoms form
hydrogen (H) and oxygen (O).
a straight chain. When the sugar is in solution, four or five
Carbon dioxide (C0 2 ) is taken up by the land plant from
of the carbon atoms (depending on the kind of sugar) and
the air. This compound is the plant's chief source of
an oxygen atom form a closed ring (Fig. 2.1 Q. In the
carbon (G) and oxygen.
straight-chain form, note the difference in the end carbon
Mineral elements are also taken up by the plant from the
atom of glucose and fructose. As noted, the —CHO of
environment. These are discussed extensively in Chapter
glucose an aldehyde. The C=0 of fructose (Fig. 2.1fi)
is
5. They include nitrogen (N), phosphorus (P), sulfur (S),
characterizes a ketone.
and several other elements. They are usually taken up
The —OH and —
H groups of the sugar molecule may
from the where they occur as ions in solution with
soil,
be arranged in different positions in the ring without
water. Some of them (e.g., magnesium and potassium) are
changing the relative numbers of carbon, hydrogen, and
present as single charged atoms, whereas others occur as
oxygen in the formula. In fact, a shifting of these atoms, as
ionic compounds with oxygen and hydrogen (e.g., N0 3 ~,
~ -2 in the examples of glucose and fructose, results in sugars
NH 4 + S0 4 2 and HP0 4 ).
, ,

with different chemical properties. Sixteen different

hexoses are possible, but only a few occur naturally in
1
Readers who have not previously studied chemistry may find it

useful to read the Appendix before pursuing this chapter. Molecules such as these sugars with the same
plants.
 water molecule. For example, the disaccharide produced
from glucose and fructose is the commonest of all sugars,
H
V CH 2 OH
sucrose:

H— C— OH CH.OH
C=0
1

HO— C —
1

HO— C—
H— C— OH H— C— OH
1

H— C— OH
I
H — C — OH
CH 2 OH CH 2 OH

06 H 12°6 06 H 1206
H OH OH H

A B
Sucrose can easily be split into fructose and glucose. One
Straight-Chain Structures
water molecule consumed in the process. Reactions
is

such as this, where water splits another compound, are

called hydrolysis reactions. They are important steps in
CH ? OH breaking down stored food molecules. Sucrose, our
1
*

c -0 common table sugar, is very important as a mobile food

H /\
\/ H
\H storage compound most plants.
in

p Three or more monosaccharide molecules may join to

IV OH form tri-, tetra-, or polysaccharides. The latter are
-1/ OH
composed of the union of many simple sugar molecules.
H
1

OH As with the formation of a disaccharide a water molecule

is given off for each pair of simple sugar molecules united.

Polysaccharides are not generally soluble in water, nor are

HOH 2 C ^^ ^o.
\ OH
they sweet. Starch and cellulose are the two most
abundant polysaccharides in plants. Each is composed of
0H/| a long chain of glucose molecules. In starch, the chain
j,r CH 2 0H may be coiled because of the way the glucose units are
1 1
linked together and some chains are branched, while in
OH H cellulose the chains are unbranched and more or less
straight. Cellulose (Fig. 2.2) is a major structural material
Ring Structures
in the plant, while starch is a reserve, water-insoluble
Figure 2.1 Glucose and fructose, two hexose sugars, have the
same formula but different structures. Both can exist as a straight food.
chain or ring. A and C, glucose; B and D, fructose. The union of relatively simple molecules, like sugar, into
long-chain gigantic molecules composed of the repetition
of simple units is a common chemical process known as
chemical composition but a different internal arrangement
polymerization. We shall meet it again in our discussion
are called isomers.
of proteins and nucleic acids.
The most common hexoses in plants are glucose and
Lipids form a very diverse collection of compounds; the
fructose. Sugars with three carbon atoms are called
chief similarity among all of them is a tendency to be
trioses; with five and seven carbon atoms, pentoses and
insoluble in water (that is, molecules of lipids do not
heptoses. The pentose sugar ribose is a constituent of
readily mingle with water molecules).
the giant molecules that carry hereditary information (the
Cutin and suberin are two waxy lipids that often coat
nucleic acids). Ribose is also part of several energy-
the surfaces of plant organs and serve to limit water loss.
carrying molecules such as ATP (Fig. 2.8).
Some waxes (e.g., carnauba) are widely used in
plant
Simple sugars are called monosaccharides. The union
furniture and automobile waxing compounds.
of two of these molecules produces a disaccharide and a
Fats are simple and abundant lipids that are composed
of fatty acids united with the three-carbon alcohol,
glycerol. A fatty acid is a molecule that has an acidic
CH 2 0H H OH CH 2 0H
I I
group at one end; the rest of the molecule is a long
u c— carbon chain to which little other than hydrogen is
\ /I °\
c
r ^/c
0H
i\ /
H
V
H
\
c
/H attached. In lauric acid (Fig. 2.3) the chain is composed
H / \ / \H / \ / \0H of 1 2 carbon atoms.
-0 \'_ -c H
Figure 2.3 shows the structure of glycerol and fatty
I

H OH CH 2 0H H OH acids, and the way in which they react to produce a fat.

Cellulose Three molecules of water are produced in the process.
The fat molecule itself is nonpolar and does not mingle
Figure 2.2 Structural formula of glucose units in cellulose readily with water. This means that fat molecules tend to

chapter 2 I Metabolism
 C

A Glycerol + 3 fatty acids 3 water + fat

HOHHHHHHHHHHH
H— — — — — — — — — — — — — — —
C OH + HO C C C C C C C C C C C C H —
H-C OH+R' HHHHHHHHHHHH
— I I I I I I I I I I I I |

H— C — OH + R"
I

H
H

H — C — lauric acid
I

3H 2
— C — fatty acid
I

H R'

H — C — fatty acid
I

R"
I

Figure 2.3 Formation of a fat by condensing fatty acids with

glycerol. A, in words; 6, structural formulas. The fattyacid
shown in detail is lauric acid.

accumulate in droplets. They are rich in energy and their side chain. The side chain is not shown in detail here

makes them good food storage compounds.

insolubility because from one amino acid to another. The 20
it differs
Phospholipids are similar to fats, but here one of the kinds of amino acids differ from one another according to
fatty acids is replaced by a phosphoryl group: the side chains they possess. The exception to the picture
shown above amino acid proline, in which the R
is the
group bends over and attaches to the N of the amino
group. Several amino acids are shown in Fig 2.4.
Proteins are polymers built by attaching amino acids
together, end to end (Fig. 2.5). The bond is made between
OH the amino group of one molecule and the carboxyl group
of another. A molecule of water is released in making this
One end of the phosphoryl group binds to the glycerol bond.
unit, while the other end of the phosphoryl may be The bond between the amino acids is termed a peptide
attached to any of several different organic groups. The bond and therefore proteins are sometimes called
phospholipids are unusual in that they are water-soluble at polypeptides. Some proteins have as few as 16 amino
one end (the phosphoryl part, which is polar) while acids while others may contain hundreds. An average
insoluble at the other end (the nonpolar fatty acid tails). protein might contain 150 to 200 amino acids. Proteins
These molecules tend to line up at the boundary of any also differ in the kinds of amino acids they contain and in

water mass in which they happen to be immersed; the theirsequence along the chain. With these differences an
insoluble parts jut out of the water. Phospholipids are immense variety of proteins is possible. No single
essential to the structure of membranes in the plant cell organism can manufacture more than a small number of
(Chapter 3), and their importance is a result of their the possibilities.
semisoluble property. It is impossible to exaggerate the importance of proteins.
Amino acids are important as the molecular units from They serve structural roles, storage roles, and regulatory
which proteins are built. Some of the amino acids also roles; in addition they are the agents (enzymes) that
serve as carriers for temporarily storing nitrogenous govern chemical reactions in metabolism Their ability to

groups. There are 20 common kinds of amino acids. With perform such a variety of functions is a result of their
one exception they are alike in their basic structure: structural variety. Each kind of protein, with its unique
amino acid sequence, performs just one function. Proteins
C can perform hundreds of different functions because there
are hundreds of different proteins in the organism.
//
H ?N C — The precise shape of the protein determines its function.
The shape is determined by the sequence of amino acids.
OH But that sequence alone is not the only factor responsible
for the definite shape of the protein. Since rotation is
carboxyl group
possible around some of the bonds in the protein polymer,
the protein has a potential for assuming many different
The carboxyl group readily donates its hydrogen nucleus patterns of coiling or folding. Of these possibilities, there is

to water; hence acts as an acid. Thus the amino and

it usually only one pattern of folding that is associated with a
carboxyl groups give the compound its name, amino acid. biological function (Fig. 2.6). An improperly folded protein
The symbol R signifies a special group of atoms called a has no function except as a store of amino acids.

Principal Materials
 H f

H
i
NH 2 1 L__°_ _J
^n i

Side chain Amino acid

H— c— V
1

I
\
NH 2 0H
Glycine

—A
Hi H

H — — C C
I

C—
\OH
—
I

VV\ C C C Hi

Tryptophane
NH- ri

H
LrJ' H I_Rj'"
H
I

NIC
I H II I

H
A — N— C— —
I

C
H
i
H
i
H
i

C
H
i

-C—
C^ C

// I I

OH H h
N H H H NH- mr
Arginine
l_l

H
(
V 1st 2nd 3rd
H— S — — C— C
I

amino acid amino acid amino acid

I

H NH- OH
Figure 2.5 A polypeptide chain
is a part of a protein molecule.

Cysteine
The backbone molecule is formed by many amino
of the protein
acids joined by the union of the amino group (NH ) of one amino
2
acid to the acid group (COOH) of another amino acid by the
removal of a water molecule. The R groups represent side chains
Figure 2.4 Some amino acids. of the different amino acids.

In the living organism each protein ordinarily maintains occur regularly along the polymer; the folding brings if

the pattern of folding that has functional value. Several them close together, they can form a hydrogen bond
forces contribute to the stability of the folding. Most (dotted line):
important, the surrounding water forms a cage, with polar
water molecules attracted to one another by hydrogen
bonds. Some of the side chains of the protein molecule
/
C = H — N.

are also polar and can interact with the surrounding water,
ionizing or forming hydrogen bonds. But other side chains Many such hydrogen bonds form within the folds of a
are nonpolar. These cannot break through the web of typical protein, often giving rise to coiled or helical foldings
forces that unites the surrounding water mass. Therefore in part of the protein chain. In addition, attractions and
the protein folds in a shape that places the nonpolar side repulsions can occur between ionized side chains. Strong
chains out of contact with water. Most proteins assume a electron-sharing bonds known as disulfide bridges can
globular or rounded shape with the interior occupied by also occur to firmly cross-link two folds of the protein
nonpolar parts and the surface, in contact with water, when the two side chains of the amino acid cysteine (Fig.

carrying the parts that are attracted to water. 2.4) come together.
Dehydration (removal of water) allows the protein to Most of the forces that maintain the folding of proteins
unfold. This is one reason why an abundance of water is are relatively weak. For this reason the folding, hence the
vital to the normal operation of the organism. function,can be disrupted by such factors as elevated
The folding of the protein molecule may also be temperatures (which leave the amino acid sequence intact
influenced by interactions between the side chains. For but tangle or denature the protein); changes in acidity;
instance, the groups and the addition of specific small molecules that may bind
H to the protein and modify the balance of internal forces.
The environmental sensitivity of proteins goes far toward
-C— and -N- explaining the physical limits of life.

chapter 2 I Metabolism
Figure 2.6 Proposed folded configuration of a molecule of
cytochrome c, a plant protein.

Enzymes and Catalysis variety of reactions, involving different combinations of

materials and products. The development of an organized
raw

plant body with a shape determined by heredity implies a

Enzymes are vital in two ways. absence the
First, in their careful selection of the chemical reactions that occur and
molecules in the body react so slowly thatwould be life when they occur. Enzymes are selective in speeding
nonexistent at the pace at which we know it, even at the reactions. Each kind of enzyme affects only a narrow
slow pace of plant life. Enzymes speed reactions. Second, range of reactions.
the molecules of life are capable of an almost infinite The property of selectivity or specificity mentioned

Enzymes and Catalysis

above is widely attributed to a "lock and key" relationship
between the enzyme and the molecules (substrates) that
react with it. Enzymes are protein molecules. The tolding
of the protein includes a furrow or pocket (the active site)
which has a shape complementary to the shape of the
substrate molecule. This allows the enzyme and substrate
to bind together; in contrast molecules with other shapes
cannot bind to that particular enzyme.
The enzyme functions as a broker or catalyst, opening
an avenue for rapid reaction but giving no energy for it.

Reactions are driven by the kinetic energy of molecular

collisions and sometimes by light energy that the
molecules absorb. Without catalysts, biological molecules
rarely react because their existing structure is maintained
by internal forces too strong to be overcome by the forces
that are generated in most molecular collisions. Enzymes
often serve to weaken the stabilizing forces within the
molecule so that the energy of collisions may be enough
to bring about the reaction. The enzyme may act by (a)
warping the substrate molecule; (b) temporarily reacting Figure 2.7 Phases of metabolism. Anabolism produces complex,
high-energy products from simpler raw materials. Catabolism
with the molecule so that its internal organization is
breaks down fuels to form compounds such as ATP and NADH
altered; or (c) changing the electrochemical environment, that are needed for anabolism. Photosynthesis completes the
cycle.
which affects the pattern of electronic movements in the
molecule. Sometimes an enzyme may also speed a
The second phase of metabolism is termed anabolism.
reaction by holding two substrate molecules together
This heading refers to pathways that build more complex
longer and with more exact orientation than would have
molecules from simpler ones. Photosynthesis, which
been possible in free solution. This gives the molecules
makes sugar from carbon dioxide and water, could have
more opportunities to collide effectively.
been put in this category but was discussed separately for
special emphasis. The construction of proteins from amino
acids is an example of an anabolic process. Anabolism is
Phases of Metabolism the basis of all developmental processes and reproductive

events.

The action of each enzyme is limited to the performance The third phase of metabolism is known as catabolism.
of only one simple catalytic task. A single reaction Catabolic pathways extract energy from fuel (food)
between two substrates usually involves changes in only molecules, for use in powering anabolism and for driving

about two bonds. Therefore the construction of a complex the transport of materials from one location to another.
substance such as a protein from simple materials Other catabolic pathways can dismantle all the kinds of
requires an extensive series of reactions with almost every molecules that are built during anabolism (though a given
step catalyzed by a different enzyme. A series of such plant may lack some of these pathways). Proteins, lipids,

reactions is termed a metabolic pathway. Most of the and carbohydrates as well as less common compounds
reactions in a pathway require two substrates, one that can serve in the plant as fuels because of these pathways.
was made in the previous step of the pathway, and the Anabolism and catabolism are linked by several kinds of
other made by a separate pathway. Thus metabolism is molecules that act as carriers of energy and building
actually a web or network of intersecting and branching materials. The most prominent and universal of these are

pathways. The enzymes determine the pathways that are ATP (adenosine triphosphate) and the pyridine
+ +
available, thereby shaping metabolism as a whole. nucleotides (NADP and NAD ).
ATP is produced from ADP (adenosine diphosphate)
and phosphate ions (Fig. 2.8). Energy is required to
Photosynthesis, Anabolism, and couple the extra phosphate group onto ADP; molecules
Catabolism with three phosphate groups joined in series are unstable,

The whole of metabolism is far too complex to discuss

Adenosine Phosphates
here. However, we can distinguish three general phases of NH,
metabolism (Fig. 2.7). One phase is photosynthesis. This I

is a complex system of pathways by which green plants

energy to build sugar molecules. It is the primary
HN \ OH OH OH
use light I

way in which energy is brought into the living system to

HC.
C CH, O
1

P O
1

P —O ~ P OH
II II II

power chemical constructions, growth, and repair. Its o o O

occurrence in plants is especially important to man Adeni Ribose >, P
2
'3

because we cannot use light energy ourselves but must

Figure 2.8 The structure of ATP (adenosine triphosphate). ADP
derive food (i.e., energy-rich molecules such as sugars) has only two phosphate groups and AMP has only one. AMP is
from other organisms and ultimately from plants. an example of a nucleotide.

chapter 2 I Metabolism

8
 2.7). Thus we can characterize ADP as a phosphate
ATP + H-OH ADP-H P-OH
carrier as well as an energy carrier, which travels back
and between catabolism and anabolism. These
forth
ATP + H-OR ADP-H + P-OR relationships help to explain why phosphorus is an
Figure 2.9 Two reactions of ATP. The first is hydrolysis; the essential element in the life of the plant.
energy stored in ATP is released as heat. In the second, an Just as ADP picks up phosphate and (in the form of
organic compound replaces water and accepts phosphate from
ATP, storing part of the energy and becoming more reactive as a ATP) carries it to anabolic pathways, so also NAD + picks
result. R can be any of a wide array of organic groups. up electrons and hydrogen ions at several points in
catabolism. Carbon dioxide, the starting material from
which plants build organic materials, contains no
reactive, and rich in energy. If molecules of ATP and water hydrogen, but all the complex functional molecules of
are put contact and supplied with a suitable catalyst,
in
metabolism are rich in hydrogen. Carriers such as NAD +
hydrolysis will spontaneously occur and heat will be get hydrogen from the fuel molecules (e.g., sugars) that
released, resulting in a final equilibrium that very strongly
are being broken down in catabolism. The hydrogen is
favors ADP and inorganic phosphate (Fig. 2.9). The role of
then supplied to anabolic pathways.
catabolism is to drive this hydrolysis reaction in reverse,
using energy from fuel molecules. By means of suitable
enzymes, the anabolic system allows ATP to react with Respiration
organic molecules but not with water (Fig. 2.9). With a
portion of the ATP molecule attached, the organic product The principal events in catabolism form a series of
inherits some of the reactivity (energy) of the ATP and can reactions known as respiration. Dozens of reactions are
participate more readily in reactions that yield complex involved in this process, using many enzymes. All of these
products. Note that the organic reactant in Fig. 2.9 is reactions cannot be presented in detail here, but we can

exchanging an H atom for a phosphate-containing group. outline some of their most important aspects in general
Evidently, attachment to phosphorus-containing groups terms. Three distinct phases in the respiration system can
tends to make organic compounds more reactive and be distinguished.
energy-rich. In most reaction pathways the phosphate- The first phase is known as glycolysis (Fig. 2 10). This
containing group is eventually released, to be recycled is a series of some 1 1 reactions in which molecules of the

back to catabolism and built into new ATP molecules (Fig. sugar glucose are trimmed and modified for entry into the

glucose

sugar
1st
phosphorylation
ATP

ADP •<-

(
(P)

>
glucose 6— phosphate )

f^
c fructose

(P)-^C-
6— phosphate
J
2nd sugar
phosphorylation
ADP -<-

~f fructose 1, 6— diphosphate j
sugar cleavage

pyruvic acid
formation NAD
c
(20
glyceraldehyde 3— phospha
D G hydroxyacetone phosphate
J
reduction
of nucleotide oxidation of
NADH <- Ur
^ triose

2ADP (P)

phosphorylation
of ADP
2ATP

pyruvic acid

Figure 2.10 Steps in the process of glycolysis

Phases of Metabolism
subsequent phase of respiration. The effect of glycolysis and electrons from each three-carbon unit; and at two
may be compared to trimming logs to a size that will fit points in the process the three-carbon compounds give up
into wood stove.
a phosphate to ADP, making ATP
Though glucose is rich in energy, it is too stable to start Glycolysis is important to the plant partly because some
the process of breakdown without help. Thus at two points of its intermediate products are useful in anabolism Only
in glycolysis the sugar molecules are reacted with ATP so about 17% of the energy of glucose is transferred to
that the sugar acquires a reactive phosphate group. This energy-carrying compounds. A energy (3%) is lost as
little

process is termed phosphorylation. Glucose-phosphate heat. And almost 80% of the energy still lies trapped in the
undergoes a series of internal rearrangements, acquires final product, pyruvate.
another phosphate from ATP, and is split into 2 three- The second phase of respiration, termed either the
carbon compounds (sugar cleavage). Another reaction citric acid cycle or the Krebs cycle (after its discoverer,
makes these three-carbon compounds identical. Another Sir Hans Krebs), breaks pyruvate down to carbon dioxide
long series of internal reorganizations then converts each (Fig. 2.1 1). In this process the hydrogen that was present

three-carbon unit into a molecule of the final product of in the pyruvate molecules is transferred to hydrogen-

glycolysis, pyruvic acid or pyruvate. In the course of carriers, chiefly NAD + .Water molecules also enter,
+
these changes a molecule of NAD picks up hydrogen donating oxygen to help form C0 2 and giving up their

entrance of
carbon into acetyl-CoA
the acid
cycle }
oxaloacetate

citric

acid cycle
\
cis-aconitate

\
isocitrate
C° 2
-^K
a-ketoglutarate

rADP+PM

ATP

final stages
of respiration

electron
transport
T T t Y t
and
->- coenzyme Q

oxidative
phosphorylation Y Y Y Y
cytochrome b
ATP
available
(ADP 4- pP) -> ATP
to do
Y YY
work in eel
cytochrome c

cytochrome a

(adp + Pi\- ATP

2H+ + '/2
2 -i-H 2

Figure 2.1 1 The relation between the citric acid cycle, electron
transport, and oxidative phosphorylation

chapter 2 I Metabolism

10
 C

+
hydrogen to NAD There are two points in the process
.
very high affinity for electrons. Its position as the final
where organic intermediates pick up phosphate from the electron acceptor explains why oxygen is necessary for
surrounding solution and transfer it to ADP, making ATP. most life.

Pyruvic acid enters this phase of metabolism by In the electron transport process, two or three
attaching to a complex of enzymes. One of the three molecules of ATP are made from ADP and phosphate for
carbon atoms of pyruvate is cleaved off and released as a each pair of electrons that travel through the chain. Since
molecule of C0 2 The two-carbon remainder, still bound to
.
many charged hydrogen were formed during
carriers
the enzyme, is an acetyl group: earlier phases and each donates a pair of
of respiration
electrons to electron transport, this last phase of
respiration is the place where the greatest abundance of
ATP is produced: about 32 molecules produced for every
H — acetyl group
glucose molecule consumed in respiration. In comparison,
glycolysis and the citric acid cycle each produces only
H
two molecules of ATP per glucose molecule consumed.

The acetyl group is transferred to a mobile carrier Overall, the complete breakdown of glucose yields six
compound that is called coenzyme A. The combination is C0 2 and six H 2 molecules, absorbing six 2 molecules
highly reactive, and coenzyme A can transfer its acetyl and trapping about 40% of the energy of glucose in the
group to several different compounds. Thus in anabolism form of some 36 molecules of ATP. The rest of the energy
acetyl-coenzyme A donates two-carbon units, which help is released as heat during the various steps of respiration.
to build more complex molecules. But
its most frequent

role is groups into the citric acid cycle.

to feed acetyl
The cycle begins when an enzyme transfers the acetyl Alcoholic Fermentation
group from coenzyme A to a four-carbon acid called
Most higher plants are not able to live for long periods of
oxaloacetate. The product, with six-carbon atoms is citric
time in the absence of 2 Without 2 electron transport
The
.
,

acid. rest of the citric acid cycle consists of a series

stops and ATP production is much reduced. In contrast,
of reactions that gradually remove all the atoms that were
some fruits, notably apples, may live and produce C0 2 for
originally brought in as the acetyl group: the carbon and
long periods even stored in an atmosphere with little
if
oxygen are released as C0 2 the H is transferred to
.
2
;

Yeast may live and reproduce with very little 2 producing

carriers. The final reaction produces another oxaloacetate
,

large amounts of C0 2 and ethyl alcohol as wastes.

molecule. This makes the whole process appear as a
But yeasts have only a limited tolerance for alcohol.
cycle, though also represents an open-ended flow of
it

When the alcohol concentration in the environment

materials from pyruvate to other products.
reaches about 12%, the yeast cells become inactive. This
Among the intermediates of the citric acid cycle are
is why natural wines do not have alcohol content above
several compounds that are starting materials for
12%. Yeast actually grows faster with an 2 supply, but
pathways phase of catabolism therefore
of anabolism. This
thenit does not produce alcohol. Only a few bacteria
is a material interface between catabolism and anabolism.
grow better without 2 present.
It is commonly described as a "hub" of metabolism. Since
In higher plants that are deprived of 2 , the pyruvic acid
pathways ending with lipids and proteins meet here, this
formed during glycolysis is converted into carbon dioxide
cycle is the location where the breakdown of lipids and
and ethyl alcohol. This also happens in yeast. This
proteins feed into respiration.
process, called alcoholic fermentation, occurs in two
The principal result of glycolysis and the citric acid
steps. First, an enzyme separates a C0 2 molecule from the
cycle is the transfer of hydrogen from sugar to carriers
pyruvic acid. This leaves a two-carbon compound,
such as NAD + The loaded hydrogen carriers are
acetaldehyde. Next, the NADH that was formed in
.

extremely reactive (rich in energy) and may donate H to

glycolysis donates its H to the acetaldehyde, forming
many molecules that are being built during anabolism. But
alcohol. NAD + is released to take part in glycolysis again
most of the H is consumed in the third phase of
Thus, during growth without 2 , the energy trapped in
respiration, the electron transport system.
NADH used to form alcohol. For each glucose molecule
is
The electron transport system acts as an ATP generator,
fermented to alcohol a net of only two ATP molecules are
which harvests the energy of reactive hydrogen by letting
formed. This is less than 3% of the energy available in
it combine with oxygen. The system consists of catalysts
glucose. About 6% of this energy is lost as heat, while
that are grouped into many short chains. Each chain
almost 84% is still locked in the alcohol and is unavailable
draws H from the loaded hydrogen carriers. The hydrogen
to the plant In addition, the alcohol itself may be toxic.
nuclei are released early in the chain: they are picked up
+ Fermentation is not an efficient way to extract food energy,
by water molecules to form H 3 The electrons remain .

compared to complete respiration.

and pass along the chain of catalysts as if they were on a
bucket brigade. Several of the catalysts in the electron
transport chains have iron ions that are converted to the
Fe
+2
form when they take up an electron, and then return
+3
The Control of Metabolism
to the Fe form as they pass the electron on to the next
catalyst in the chain.The final catalyst is an enzyme that
+
combines its electrons with H (from the surrounding In preceding sections we have outlined the basic structure

water) and 2 to
form water molecules. Oxygen has a of metabolism. We now turn to the control systems that

The Control of Metabolism

11
balance the reaction rates within the pathways and that The Control of Enzyme Quantities
direct the pattern of productions.
The rate of activity along a pathway may also be
controlled by adding or subtracting enzyme molecules.
Local Control Within Metabolism Indeed, whole phases of metabolism may be completely
turned on or off by this means. More than any other
Since metabolism comprises many pathways, its control is factor, the timed and controlled production of enzymes

also complex. But several kinds of controls occur guides the transformation
of a seed into an adult plant. It

repeatedly at different places in the metabolic system and, has been mentioned before that an almost unlimited
taken together, they constitute the total control system. variety of different proteins could theoretically be made,
Compartmentation is one important aspect of control: whereas real plants actually build only a few hundreds or
the enzymes that govern various pathways are often thousands. The ability to build a particular set of proteins
confined in organelles so that they are separated from the is a property that the plant has acquired from its

enzymes of other pathways by differentially permeable ancestors. Biologists currently believe that the hereditary
membranes. This confinement permits the membrane to control of protein synthesis — both the kinds proteins of

exert control over the flow of molecules between and the timing of their production — form the essential
pathways. Compartmentation is further discussed in basis of all heredity. Therefore the subject of protein

Chapter 3. (includingenzyme) synthesis and its control have primary

Regulatory enzymes are found at many points in the importance in any study of life.
network of metabolism, particularly where pathways To control enzyme quantity, systems are needed for
branch off. Regulatory enzymes differ from other enzymes removing old enzyme molecules as well as building new
in can be controlled by
that their rate of catalytic activity ones. Plants often contain relatively nonspecific protein-
special small molecules known as modulators. Modulators degrading enzymes known as proteases. Proteases break
act by attaching to the enzyme at a particular location down proteins more or less indiscriminately, so that
known as the allosteric site. The attachment changes the accessible proteins must be continually replaced if the
shape of the enzyme and its ability to work with its normal quantity present is unchanged.
to remain

substrate. Depending on the enzyme and the modulator, The control of enzyme synthesis is complex. The control
the attachment may either speed or slow the enzyme's system needs (1) a set of recipes, which spell out the
action. A single regulatory enzyme may have sites to amino acid sequences of each type of protein that the
attach as many as a dozen different modulators. This gives plant can build; (2) a reading system that can translate the

the regulatory enzyme a fine degree of sensitivity to recipes into protein molecules; and (3) a control system
conditions in the surrounding plant body. that instructs the reading apparatus on the specific recipes

Often the final product of a pathway acts as a modulator to use at each moment.

to inhibit a regulatory enzyme at the start of the pathway.

This relationship is calledfeedback inhibition. It provides
a supply-and-demand control of production: if the product The Mechanism of Protein Synthesis
is formed faster than it is used, its concentration rises and
the regulatory enzymes governing the pathway are The recipes for proteins are technically known as genes.
inhibited. The attachment between modulator and enzyme Recipes are stored in molecules of deoxyribonucleic acid
is weak and reversible, so a later demand for more (DNA). Like proteins, DNA is a polymer. Its subunits are
product, signaled by a drop in concentration of the called nucleotides (Fig. 2.8). Each nucleotide of DNA
modulator, releases the enzymes from inhibition. consists of a molecule of the sugar deoxyribose to which
To understand the operation of regulatory enzymes is it is attached a phosphate group and a complex organic

important to realize thatmany copies of each kind of group known as a base. In the polymer, the phosphate of
enzyme are present in the system, and that some enzyme one nucleotide is linked to the sugar of the next. Thus the
molecules may be free of modulators at the same time that sugar and phosphate groups comprise a "backbone" of
other molecules of the same enzyme are bound and the polymer with the bases jutting out to the side. There
inhibited (or promoted). This makes the rate of the are four kinds of bases in DNA: adenine, guanine,
reaction smoothly adjustable in proportion to the levels of cytosine, and thymine (Fig. 2.12). These are often
modulator concentrations. symbolized as A, G, C, and T. Just as an English sentence
Regulatory enzymes balance the rate of catabolism carries information in its sequence
so also the of letters,
against the needs of anabolism. The anabolic processes DNA polymer carries information about proteins in its
consume ATP, converting it to ADP and phosphate ions. It sequence of bases.
has been found that one of the enzymes in glycolysis is The structure of DNA is a little more complex because
regulatory, with ATP as an inhibitory modulator and ADP the DNA polymers combine in pairs, wound together as a
as a promotory modulator. If the rate of anabolism double helix (Fig. 2.13). This pairing is maintained by
increases, the cell soon contains less ATP and more ADP. hydrogen bonds that occur between the bases of the two
This change increases the rate at which the regulatory polymers. For such a helix to be stable, the two DNA
enzyme in glycolysis operates. The result is that glycolysis polymers must have a complementary sequence of
runs faster, feeding the other phases so that respiration as bases. The required relationship is that wherever one
a whole is accelerated. ATP is therefore produced at a polymer has an A, the other must have a T; and wherever
higher rate. one has a G, the other must have a C. Thus for a polymer

chapter 2 I Metabolism

12
 H N H

II 1 II H

H— fT X— C— t /

1 II 1 II II
-G-
1 1

-^ C \ /C v H

H H H
Uracil Cytosine Thymine

T- -A-

A- -

Sr -G--C-
II

-C--G-
N^^C-\ H—
' C
W C ^|/
C—
^ V N /C
T- -A-

Adenine Guanine
C- -G-
A- -T-

Figure 2.12 The bases in DNA and RNA. T- -A-

with the sequence ACCTG, the complementary polymer

would have the sequence TGGAC. The reason tor this :--g-
requirement is that the bases ditter in size and shape; only
the A-T and C-G pairs have the proper match of size and
shape to fit comfortably within the helically folded Figure 2.13 Schematic diagram of the double helix in a portion
arrangement. Note that when the two polymers are of a DNA molecule. Alternating sugar, S (deoxyribose); and
phosphate, P, groups make up the backbones of the strands.
combined in the double helix, the important base
Attached to each sugar unit is one of the purine or pyrimidine
sequence is hidden in the interior of the helix. In this form bases, adenine, A; thymine, T; guanine, G; cytosine, C. The
the genes cannot be read and the information in the DNA strands of the helix are held together through hydrogen bonds
between the base pairs: adenine to thymine and guanine to
is not available for protein synthesis. To permit reading of
cytosine. A, one strand is displaced along the axis of the helix tor
the recipes, the DNA helix must unwind. Possibly the the convenience of diagramming; B, a diagram of the parallel
primary importance of the helical arrangement strands in the helix.
is that it

protects the DNA

and prevents untimely reading.
The reading system is a two-phase process, consisting enzyme reaches the end of the DNA segment that was to
of transcription followed by translation. be read, the RNA molecule is released and the enzyme
Transcription is the transfer of information from DNA to may detach and return for another act of transcription.
a similar substance, RNA (ribonucleic acid). RNA is also The RNA molecule that results from this act of
built of nucleotides, but its sugar is ribose instead of transcription may now be read by the protein-synthesizing
deoxyribose; and in place of the base thymine, RNA machinery, as instructions for building a protein. Because
carries the base uridine. Otherwise the two kinds of of its intermediary is termed
role, this kind of RNA
polymer are similar. Transcription works in the following messenger RNA or mRNA. The
an intermediary
use of
way (Fig. 2.14). Under the influence of an enzyme called information carrier presumably allows the organism to
RNA polymerase, the DNA double helix uncoils at the control its information more effectively; when no longer
region containing the recipe of the protein to be needed, the mR M(\ molecule may be destroyed, leaving
manufactured. The enzyme attaches to one of the two the original DNA intact for possible later transcription.

DNA polymers. Free subunits of RNA, which have been Translation is a more complex process than
built by the metabolic system, attach to the bases of the transcription. Here the mRNA polymer is "read" by a
DNA polymer according to the base pairing rules: where complex a ribosome, with a protein molecule
unit called

the DNA has an A, an RNA subunit bearing the base uracil as the product. The ribosome consists of some 50
(U) attaches by hydrogen bonds; where DNA has C, a different protein molecules together with three large
subunit with G will attach; and so on. The enzyme moves segments of a special type of RNA known as ribosomal
along the DNA polymer and connects the RNA subunits to RNA. (Ribosomal RNA is produced by transcription from
one another so that an RNA polymer results. This RNA special regions of DNA by the same process that yields
polymer has a base sequence complementary to the DNA mRNA. However, ribosomal RNA is embedded in the
molecule, with the substitution of U for T. When the ribosome where it presumably serves a structural role.)

The Control of Metabolism

13
 free RNA subunits

Figure 2.14 Transcription. A segment of DNA

about to be read has uncoiled from the helix. The
segment may contain genes for one or several proteins. The RNA polymerase enzyme, shown as a
circle, has built part of an RNA molecule and is waiting for the next free subunit (which should carry
the base guanine) to arrive.

Ribosomes need help to read the information carried by

mRNA. Specifically, there is required a set of enzymes amino
called amino acid enzymes and a set of RNA
activating acid

molecules of a third type, known as transfer RNA ( f RNA).

To understand how these components cooperate let us
consider the nature of the "language" or "code" in which
DNA and RNA carry information.
In English, a word consists of a group of letters that
may be anywhere from one to perhaps 20 letters in length.
The RNA language also has words, technically known as
codons. All the codons have the same length, however.
Each consists of three bases along the RNA molecule.
The mRNA molecule acts as a string of three-base
codons. Most of these codons represent the names of
particular amino acids. With three bases per codon and
four kinds of bases (A, C, G and U), there are 64 possible
different codons. This is more than enough to name all the
20 kinds of amino acids. Some of the extras codons serve
as punctuation, instructing the ribosome where to start
and stop along the RNA message. Most of the extras are
synonyms, however, for amino acids that have already
been named by other codons. For example the amino acid
isoleucine is represented by six different codons.
For each codon that names an amino acid, there is a
unique, matching kind of transfer RNA (fRNA). There are
61 kinds of fRNA. The fRNA's are all alike in that they are
small compared to mRNA: each contains about 80
nucleotides each, whereas mRNA may contain thousands.
mRNA
I I I I I I I I I I

Also, all the fRNA's fold into cloverleaf shapes (Fig. 2.15)
that are stabilized by the same kind of base pairing used codon
in the DNA Each fRNA molecule can carry
double helix. Figure 2.15 The structure of a fRNA molecule.
its appropriate amino acid at one end. The other end of
the fRNA molecule is specialized to bind with the
appropriate mRNA codon. This binding to mRNA is 2.16).The enzyme uses energy from ATP to join the
achieved by means of a set of three bases, forming an amino acids to the fRNA molecules. Once joined, the
anticodon, on the fRNA molecule. The anticodon is amino acids are much more reactive than free amino
complementary to the codon. Thus a fRNA with the acids.
anticodon would bind to the mRNA codon GUC.
CAG The next step in building proteins is for the "loaded"
The fRNA cannot pick up amino acids by itself. To do fRNA's to attach to the right mRNA codons. The ribosome
this, amino acid activating enzymes are needed. There seems to serve as a kind of "workbench" on which the
are 20 of these enzymes, one for each kind of amino acid. reading of mRNA is completed (Fig. 2.17). The ribosome
Each enzyme has an active site that is shaped to accept attaches to the mRNA molecule at a "start" codon, and
only the correct amino acid and fRNA combination (Fig. attaches to a loaded fRNA that matches this codon. The

chapter 2 I Metabolism

14
 amino acid

amino acid

anticodon

Figure 2.1 6 Diagram showing one manner in which fRNA could select a specific amino acid from
an amino acid pool. Nofe that ATP and an enzyme are involved. The binding sites on the enzyme are
specific for a certain amino acid and for the molecule of fRNA bearing the anticodon for that amino
acid.

ribosome shifts its grip on the mRNA, moving to the next produce mRNA depends on the presence or absence of a
codon. Eventually another loaded fRNA arrives, matching repressor protein that blocks the way. The repressor
this codon. The ribosome detaches the first amino acid binds to the DNA at a site near the promoter called the
from its fRNA and attaches it by a peptide bond to the operator site. Repressors are analogous to the locks on
newly arrived amino acid. The ribosome shifts its grip to filing cabinets. They are similar to regulatory enzymes in

the next codon and ready to receive another loaded

is that their shapes can be changed when special small
fRNA. This cycle of steps is repeated again and again with signal molecules are attached to them. Depending on the
one amino acid after another being added to the growing particular repressor and signal molecule, attachment may
protein until the ribosome reaches a stop codon. At this either improve or decrease the repressor's ability to bind
point the ribosome releases both the mRNA and the with DNA. Therefore the reading of genes is controlled by
completed protein. The ribosome, the mRNA, and the supplying or removing the signal molecules. Such
fRNA may be reused many times to produce more protein molecules may be products of local metabolism, they may
molecules. enter from the environment, or they may migrate from a
distant point of origin elsewhere in the organism. These
possibilities may give a basis for keying the production of
The Control of Protein Synthesis enzymes to prevailing conditions, as well as coordinating
development in various parts of the organism.
The system that controls the reading of the genes is not
The control of gene reading by repressors and signal
well understood in plants. However, we have some
molecules known as the operon mechanism. The extent
is
knowledge ot the controls in bacteria and these may give
to which this mechanism operates in higher plants is
us an idea of the possible systems.
uncertain, and additional modes of control undoubtedly
Control requires accuracy in selecting the right gene to
remain to be discovered.
be read at the right time. In bacteria, the task of locating
the right gene falls to the RNA polymerase enzyme. The
Summary
DNA molecule contains many genes arranged in a series.
The points where transcription may begin are marked by 1. The plant body is built and governed by a complex
special regions of DNA known as promoter sites. These system of reactions called metabolism.
sites presumably have special base sequences that permit 2. The system builds organic compounds
plant metabolic
the enzyme to bind to the intact DNA helix. from carbon dioxide, water, and minerals absorbed
Whether an enzyme that has found a promoter site may from the environment.

Summary

15
 Figure 2 17 Steps in translation In A, the ribosome binds an
mRNA molecule with two codons in working positions A partially
completed protein is bound to fRNA 1 and another fRNA with its ,

amino acid has just arrived at the second working site. B, as the
next step, the ribosome shifts the protein from fRNA 1 to the
newly arrived amino acid fRNA 1 is now free to depart. C, the
polypeptide chain
ribosome has moved along the mRNA molecule a distance of one
codon, so that /RNA 2 is now in the left-hand working site A
third fRNA (3) with its amino acid has bound to the next codon.
Step B can now take place again. This cycle of events occurs
each time a new amino acid is added to the protein.

The catalysts are special proteins called enzymes.

5. Each enzyme catalyzes only a specific kind of
reaction, so that the course of metabolism is

determined by the selection of enzymes on hand.

anticodon
6. Sequences of reactions are called metabolic pathways.
The pathways branch and join as a network.
7. The three main phases of metabolism are
tpcpcp
g jffl t? pg g gg gg gg a p g g p
!

J i i i i ii i i
"p photosynthesis, anabolism, and catabolism. Anabolic
ribosome ./ I_ mRNA pathways build complex molecules; catabolic
pathways break down fuels to produce building
codon
materials and provide energy for anabolism and
transport processes.
8. Energy and hydrogen are carried from catabolism to
anabolic reactions by recyclable carrier compounds
such as ATP and NADH.
9. The principal events in catabolism comprise
respiration, which is the oxidation of fuels such as
sugar, fats, and amino acids to produce ATP.
10. Respiration consists of three blocks of reactions:
glycolysis, the citric acid cycle, and the electron
transport system. Many intermediate products formed
in the first two phases are used as building blocks in

anabolism.
11. In the absence of 2 , respiration cannot be completed,
but glycolysis may continue with the terminal product,
pyruvic acid, being converted to ethyl alcohol instead
of feeding into the citric acid cycle. This is the process
of alcoholic fermentation. It yields much less ATP than
respiration.
12. The choice of metabolic pathways and their rates are
controlled by mechanisms that include separation of
enzymes in organelles, the action of regulatory
enzymes that change their rate of action on signal,
and the production and destruction of enzyme
molecules. Some of these controls use feedback
signals to pace reactions.
13. Protein (enzyme) synthesis uses the primary
hereditary information of the organism as recipes for
building particular proteins.
14. The hereditary information is permanently stored in the
base sequences of DNA molecules. The first step in

protein synthesis, termed transcription, consists in

building disposable molecules of mRNA that have base

sequences matching that of DNA.
15. Protein synthesis is completed by ribosomes (units

built of RNA and protein) that assemble amino acids

into proteins, using mRNA molecules as templates to

determine the proper amino acid sequence. This

process, termed translation, also involves many
enzymes, fRNA molecules, and energy sources.
3. The most abundant compounds formed by plants are 16. The control of protein synthesis by the operon
carbohydrates, lipids, amino acids, proteins, and mechanism, found mainly in bacteria, is based on
nucleic acids. blocking the process of transcription through the
4. Nearly all reactions in metabolism require catalysis. reversible attachment of proteins to the DNA molecule.

chapter 2 I Metabolism

16
 3 CHAPTER

3 the plant cell

the
Galen,Asia of the great Greek doctors,
last who lived enclosed air." He estimated that a cubic inch of cork
Minor during
in the second century A.D., would contain about 1259 million such cells.
thought that all organs, such as spleen, brain, Because of the prominence of cell walls in plant tissue,
and kidney of animals, leaves, stems and roots of plants, the cell was soon considered the unit of structure and of
were a "sensible element, of similar parts all through, life in plants. However, during these early years, zoologists

simple and uncompounded." Others before him had considered the tissues as the true centers of life in the
thought that animal tissues were simple coagulated animal body. There were supposed to be 21 different
"juices" seeping through the walls of the intestine. No one change within themselves, depending upon
tissues, able to
dreamed that these tissues and their plant counterparts the organ which they were located. In these tissues life
in

had an astonishing and complicated structure, a structure was thought to reside. However, numerous observations
that could not be seen until the invention of a microscope, upon protozoa and animal tissues led to a gradual
which Zacharias Jansen, a spectacle maker of Holland, accumulation of evidence that cells also existed in

accomplished in 1590. This instrument was later improved animals. 1838 the German zoologist Theodor Schwann
In

upon by Robert Hooke, an Englishman, who was not only and the botanist Matthias Jacob Schleiden collaborated in
interested in optics but was also an architect and an a paper entitled, Microscope Investigations on the
experimenter with flying machines. Hooke, who lived from Similarity of the Structure and Growth in Animals and
1636 to 1703, examined all sorts of natural objects with Plants.
his improved microscope. Among these were thin slices of This paper established on a firm basis the theory that
cork (the dead outer bark of an oak). Figure 3.1 shows the cell is a basic unit of structure in both plants and
cork tissue as Hooke saw it under his microscope. This animals. However, another 30 years of research were
illustration was published in 1664 in an article entitled necessary for the general acceptance of such a new idea:

Micrographia, or Some Physiological Descriptions of that organisms are composed of cells, and that cells
all

Minute Bodies Made by Magnifying Glasses. The term cell are indeed the basic units or building blocks of life.
was first used by Hooke to denote in cork the "little boxes This chapter is divided into three parts. The first

or cells distinct from one another . . . that perfectly examines plant cell structure, the second summarizes cell

division and the cell cycle, and the third reviews the
process of DNA replication.

Cell Structure

There are two large groups of organisms that differ in

basic cellular structure. Bacteria and blue-green algae

(perhaps more correctly, blue-green bacteria, Chapter 11)
constitute one group called prokaryotes. Prokaryotic cells

are considered to be primitive and are characterized by

their lack ofa membrane-enclosed nucleus and of
membrane-bound organelles. The second group
comprises all other plants and animals, the eukaryotes.
Eukaryotic cells separate many of the metabolic processes
of the cell into discrete membrane-bound, protoplasmic
particles called organelles. Each organelle has a
characteristic structure and function wherever occurs. it

The organelles are the nuclei (Fig. 3.4), the mitochondria

(Fig. 3.7), the endoplasmic reticulum (Fig. 3.11), the

dictyosomes (Fig. 3.15), the microbodies (Fig. 3.18), and

the plastids (Figs. 3.12 and 3.14). Although the ribosome
is a small particle rather than a membrane-bound
Figure 3.1 Cork tissue as Robert Hooke observed it under his
microscope. organelle, does play an important role in protein
it

17
Table 3 1

Some Dimensions

inch cm mm fim nm A

1 inch 1 2.54 25.4 25.4 x 10 3 254 X 10

6
25.4 x 10 7
4
1 cm 0.393 1 10 10 10
7
10
8

1 mm 0.393 x 10"
1
10" 1
1 10
3
10
6
10
7

4
1 /xm 0.393 x 10" 10" 4 10~ 3 1 10
3
10
4

7
1 nm 0.393 x 10" 10" 7 10" 6 10" 3 1 10
8 10~ 8
1 A 0.393 x 10~ 10" 7 10~ 4 10" 1
1

The symbol for micrometer (/xm) now replaces the micron (/x); nanometer(nm) replaces millimicron (m/x).

synthesis and is included in this list. The organelles are electrons very much as they are examined with a light
bathed ground cytoplasm through which diffusion of
in beam. However, electrons are not visible to the eye, and
materials occurs, in which certain phases of metabolism some device, such as activation of phosphors on a
take place, and in which the activities of the organelles fluorescent screen, must be used to enable us to see the
are coordinated. beam of elections. There are two types of electron
Eukaryotic plant and animal cells differ in several basic microscopes: transmission electron microscopes and
ways: (a) Plant cells specialized to carry out scanning electron microscopes. In the first type, the beam
photosynthesis are provided with one to several passes through the specimen as in the compound
organelles, the chloroplast (Fig. 3.12), in which reside microscope. In the second, the beam is reflected as with
most of the enzymes, energy compounds, and the stereodissecting microscope.
intermediate substances required for photosynthesis, (b) For transmission electron microscopy, small pieces of
With few exceptions, plant cells are provided with a cell tissue are killed in glutaraldehyde and osmium tetroxide,
wall composed and secondarily deposited
of cellulose then dehydrated in a graded series of alcohols and
materials that gives greater rigidity and lessened mobility embedded in a plastic that must be held at 70°C for 24
to plants, (c) Most plant cells are provided with a large' hours to harden it properly. Sections of tissue and plastic,

central aqueous vacuole (Fig. 3.6). about 70 nm thick (0.07 /xm), are stained with a heavy-
As you recall, Robert Hooke calculated that there would metal stain such as lead citrate, which binds to

be 1259 million cells in a cubic inch of cork. This figure is membranes. The tissues are then observed with the
based on his estimate of about 1 100 cork cells along a electron microscope in a high vacuum. A beam of
line 1 inch long. One thousand cells along 1 inch would electrons is passed through the sections, and an image is
give us about 40 cells for the length of 1 mm; therefore a formed on a fluorescent screen. Magnetic fields bend the
single cork cell is '/40 mm in diameter. It now becomes beam of electrons just as glass bends light rays. Thus, the
convenient to use a smaller unit of measurement, the lenses in the electron microscope are electromagnets. The
micrometer (micron) (/xm). There are 1000 /xm in 1 mm, dark masses seen on the screen of the transmission
and a cell '/40 mm in diameter would be 25 /xm in diameter. electron microscope will not be images of organic matter
This is about correct for a cork cell, although many plant but shadows of the high-atomic-weight metals, osmium
cells are larger. Table 3.1 lists other units of measure from the fixative and lead from the stain, which have
commonly used. become preferentially bound to some components of the
protoplasm.
For viewing with the scanning electron microscope, an
Technique object is mounted on a small metal plate and placed in a
vacuum in the path of a beam of electrons. Living tissue
Living cells may be observed for many hours on the light may be examined directly; however, rapidly desiccates it

microscope by maintaining them in a proper medium. in the vacuum and does not provide a good reflecting

Because the parts of the protoplast have an index of surface for the electrons. This means that plant parts to be
refraction close to that of water (i.e., the protoplasm bends studied with the scanning microscope must usually be
light rays to the same degree that water does), many killed, dehydrated carefully, with a final drying in liquid

details of cell structure are not apparent in living cells. C0 2 and then shadowed or coated with a thin layer of
,

More detail can be observed by killing cells in various gold. Thus prepared, the tissue has a normal appearance,
mixtures of alcohol, acetic acid, chromic acid, does not deteriorate further, and has a better reflecting
formaldehyde, or osmium tetroxide. The latter two surface in the electron beam.
compounds are because when they
of particular interest,
kill cells they do not coagulate the proteins. The killed
Structure and Function
tissue is dehydrated, embedded in paraffin, and cut into
sections from 5 to 20 /xm thick. When this tissue is Cells may be considered by referring to their
mounted on a glass slide and properly stained, much (a) structure, (b) activity, or (c) chemical organization. It is

detail becomes apparent. necessary to know the interrelationships of all three levels
Plants or plant tissues may be examined with a beam of to understand the cell as a whole. Within a cell, certain

chapter 3 I The Plant Cell

18
activities are confined to definite structures or organelles. solutes, and sometimes water-soluble pigments. The
Photosynthesis, for example, takes place in chloroplasts, interfaces of the cytoplasm-vacuole and the
while respiration is confined to mitochondria. cytoplasm-cell wall are characterized by special
Cells are dynamic living units that are maintained in cytoplasmic membranes that exercise some control over
balance with their surroundings only through the the passage of substances into and out of the cytoplasm.
expenditure of energy. The maintenance of a steady living These membranes are not visible with the light microscope
state through the expenditure of energy is known as but, because of their activity, their existence has long
homeostasis. Disruption of the source of energy results in been recognized. The membrane separating cytoplasm
the death of the cell. and vacuole is known as the tonoplast; that bounding the
Cells are continually renewing their substance. To keep outer surface of the cytoplasm is the plasmalemma. The
pace with the increasing number of cells, the total mass of cytoplasm and plasmalemma may be removed, leaving the
protoplasm must also increase. Cells also accumulate tonoplast as a sac enclosing the vacuole. A single
materials to be stored and used in their own metabolism. nucleus (Figs. 3.2, 3.38) is embedded in the cytoplasm.
For instance, leaf cells synthesize sugar from simpler In the living cell it generally appears structureless,
substances, and store it in large amounts as starch. although with careful focusing, one to four small denser
The products, or accumulations, of protoplasmic activity bodies, the nucleoli (nucleolus, singular) can be seen
(water, sugar, cellulose, hormones, and similar items) are (Fig. 3.2). There is also an envelope (membrane) around
found in the dynamic cooperating unit of protoplasm, the nucleus.
called the protoplast (Figs. 3.2, 3.3A,B). The protoplast The cytoplasm in a leaf cell, especially when warmed by
secretes about itself a cell wall which frequently later the light of a microscope, will show active streaming, in
receives a deposition of secondary products. All of the which the organelles are carried. In time the velocity
protoplast, plus the cell wall, make up the plant cell. reaches 5 to 10 mm per min. This rapid flowing of the
cytoplasm is a dramatic indication of its plasticity, and

The Living Cell

•L <-
In a such as that in the aquatic plant Elodea,
living leaf,

the chloroplasts are the most prominent organelles. They

are embedded in the cytoplasm, which is confined to the
periphery of the cell, or to thin strands crossing the clear
central vacuole (Figs. 3.2, 3.3A.B). The central vacuole
contains water with dissolved salts, various organic

V \

w*-J&
K

cytoplasm

.
•
/
cell wall

chloroplasts
fa >

B
Figure 3.3 Leaf cells from the aquatic plant Elodea. A, living
untreated cells; note the distribution of chloroplasts around sides,
Figure 3.2 Diagram of a living cell from an Elodea leaf. When top, and bottom of the cell, with a vacuole occupying the rest of
viewed on the light microscope, organelles in living cells, other the cell. B, in cells recently killed with formaldehyde, the
than chloroplasts, lack contrast. The nucleus and nucleolus are protoplast may withdraw slightly from the cell wall, chloroplasts
generally visible. Mitochrondria are seen only with perfect lighting may form irregular clumps, and the nucleus frequently becomes
conditions, x700. visible.

Cell Structure

19
should be recalled when studying microscope slides and bounded externally by the plasmalemma (Figs. 3.4, 3.5,
electron micrographs. 3.7)and internally by the tonoplast (Figs. 3.4, 3.4, 3.6).
The plasmalemma separates the protoplast from the
external cell wall or other nonliving systems. Within the

Cell Fine Structure protoplast, the tonoplast separates the protoplasm from
the vacuole. In any organized living system membranes
A thin slice of a cell from a young leaf, as it appears in the are needed to separate high-energy from low-energy
transmission electron microscope, is shown in Fig. 3.4. regions.
Cells at this stage of development are not conducting Note that each of these membranes consists of a light
photosynthesis at their potential capacity. These young line sandwiched between two dark lines. Such membranes
cells tend tohave relatively dense protoplasm, small are known as unit membranes (Fig. 3.7). They have been
vacuoles, and a large nucleus. The cells shown in Fig. 3.4 thought to be composed of a central bimolecular layer of
have several small chloroplasts, mitochondria, lipid sandwiched between two monomolecular layers of
microbodies, dictyosomes, endoplasmic reticulum, and fully extended protein (Figs. 3.7, 3.8A.B).
many small vesicles and ribosomes. This abundance of Two other membrane models have been recently
organelles indicates a high level of metabolic activity. —
proposed the subunit model (Fig. 3.8C), and the fluid
Compare the "texture" of these cells to that of the young mosaic model (Fig. 3.8D). The two models are in some
two figures,
root cell in Fig. 3.5. Both of the cells in these ways similar, both consisting of globular protein molecules
from a young and a young root, are highly active. The
leaf partially or completely embedded in a lipid bilayer. The
primary difference between them is that the root cell has presence of protein actually within the membrane and not
colorless starch-storing plastids (amyloplasts) while the just on its surface helps to explain certain aspects of
leaf cellcontains green chloroplasts. Now examine Fig. membrane permeability (Chapter 5).

3.6, which was taken from a mature leaf of alfalfa. In

comparison to the young leaf and root cells these mature
cells are less "busy" in terms of their organelle Ribosomes
complement; they also contain a large dominant central
vacuole. Note that the nucleus and other organelles are The cytoplasm, after glutaraldehyde and osmium fixation,

pressed against the cell wall. The chloroplasts are also is moderately electron transparent and has a soft gray
quite large and numerous. granular appearance. In it always appears at least one
type of densely stained granule, roughly angular and from
17 to 20 nm in diameter (Figs. 3.4, 3.7). The enzyme
Membranes RNAase specifically degrades RNA. Treating tissue with
RNAase results in the disappearance of these granules,
Perhaps the most notable thing about cells at this with no change taking place in the other organelles. This
magnification is their compartmentalization into isgood evidence that RNA is present in these particles,
membrane-bound organelles. The protoplasm itself is the ribosomes. Ribosomes are present in plastids and

Figure 3.4 Immature cells from apex of sunflower (Hellanthus

annuus). Note the small vacuoles (v), large nucleus (n) and other
organelles —
mitochondria (m), chloroplasts (c) and dictyosomes
(d), X6416.

chapter 3 I The Plant Cell

20
Figure 3.5 Immature cells from young root tip of Gladiolus sp.
Notice the similarities to the immature leaf cells in Fig. 3.4; small
vacuoles (v), amyloplasts (a), mitochondria (m), and dictyosomes
(d) are apparent, x 4583.

Figure 3.6 Mature cells alfalfa (Medicago sativa) leaf.

from an
The vacuoles (v) are larger in these old cells. Note also that
much
the nucleus (n) is proportionally smaller than in immature cells.
Chloroplasts (c) and mitochondria (m) are also abundant, X4191

Cell Structure

21
 (6Y6Y6Y6Y0Y0;

(ofoYotbibto)

222222222222222222

655655555555556555
WXMMAAWAAA/W
Figure 3.7 Enlarged view of mitochondria (m) in a corn root cell
(Zea mays). The nucleus (n), a dictyosome (d), and numerous
ribosomes (r) are also visible, x 25,333.

70= 10A 40 = 5 A

Figure 3.8 Membrane models. A, B, unit membrane; C, subunit

model; D, fluid mosaic model.

proteins

chapter 3 |
The Plant Cell

22
 .

Figure 3.9 Electron microscope photograph ot an immature

wheat (Triticum aestivum) leaf Numerous organelles are present.
Note especially the coiled polyribosomes (arrows), x6231

Organelles

Endoplasmic Reticulum
In cross section, the endoplasmic reticulum is represented
by profiles of two parallel membranes separated by a
narrow light space about 4 nm wide (Figs. 3.9, 3.10,
3.11). These profiles are actually cross sections through
extensive, flattened vesicles. Note that these membranes
form a closed system; their ends are never open to the
ground cytoplasm. The endoplasmic reticulum frequently
has ribosomes pressed to its outer, or cytoplasmic,
surface. This is known as rough endoplasmic reticulum.
Smooth endoplasmic reticulum lacks ribosomes and is
net as common as rough endoplasmic reticulum (Fig.
3.10). There is considerable evidence suggesting that
protein synthesis may be associated with the rough
endoplasmic reticulum. For example, endoplasmic
reticulum is most highly developed in cells active in
protein synthesis. It is poorly developed in mature leaf
cells (Fig. 3.6). The endoplasmic reticulum is associated

Figure 3.10 Endoplasmic reticulum in an apical cell of the alga with plasmodesmata, or cytoplasmic connections from
Chara. Both rough and smooth endoplasmic reticulum are present cell to cell across cell walls (Figs. 3.4, 3.7, 3.9).
and are continuous with each other. A microbody is also visible.
x 1 s.tjoo.
Mitochondria

The mitochondrial envelope separates internal

mitochondria, as well as in They appear
the cytoplasm. to mitochondrial space from the cytoplasm (Figs. 3.4, 3.7).
be lacking in nuclei. Some micrographs show them The mitochondrial envelope is double and each
attached to the cytoplasmic side of the nuclear envelope. component is a typical unit membrane. The outer
If they do occur within the nucleus, they are probably component forms a continuous barrier around the
confined to the nucleolus Ribosomes are
(Fig. 3.5). mitochondrion. The inner membrane is thrown into folds
frequently associated with thecytoplasmic-membrane called cristae (Figs. 3.7, 3.1 1). Internal to the cristae is a
system, the endoplasmic reticulum (Figs. 3.9, 3.10). Under fluid called the mitochondrial matrix. Cristae in plant
certain circumstances they appear in groups, frequently as mitochondria frequently are sparse, irregularly arranged,
helical aggregations. These arrays are designated as and not always seemingly connected with the inner
polyribosomes or simply polysomes (Fig. 3.9). membrane of the envelope.

Cell Structure

23
 ER

r ' ' > .•-•/.•

Figure 3.1 1 Meiocytes of Selaginella to show the endoplasmic
reticulum (ER) and other membranes surrounding cellular
organelles, x 16,250.

Plastids thylakoids. Some of these occur in cylindrical stacks

(Fig. 3.12) called grana. The grana are connected at
The chloroplasts (Fig. 3.12) are bounded by an envelope irregular intervals by membranes called frets.
of two membranes, just as are the mitochondria. The
plastids have a granular background material, the stroma. Kinds of Plastids. Plastids respond to environmental
Particles having the properties of cytoplasmic ribosomes conditions by reversibly changing their structure, contents,
may be seen in the stroma. Clear areas are also present, and function. Dark-grown seedlings, for example, are long
and in favorable sections, such clear areas contain an and spindly and contain plastids with an elaborate
array of slender fibrils that represent chloroplastic DNA. crystallinelike structure usually called a prolamellar body
Also present are closed membranous sacs called (Fig. 3.13). In nongreen embryonic tissue and in roots are

Figure 3.12 Chloroplast from alfalfa (Medicago sativa) leaf. The

parts of the chloroplast, the grana (g), frets (f), stroma (s), stored
starch (st) and surrounding envelope (e) are clearly shown,
x 16,696.

chapter 3 l
The Plant Cell

24
 approximately 28 nm in diameter (Fig. 3.17). Their function
is not exactly understood, but connected to certain
it is

cellular events. The spindle apparatus, for example, is

composed of bundles of microtubules and evidence
indicates that they are involved in chromosome
movements found
(Figs. 3.27, 3.28). Microtubules are also
in bands around cells and somehow regulate
that spiral
the pattern of secondary and primary cell wall formation
(Fig. 3.30). They also occur in the cilia, or flagella, of

motile cells of all eukaryotes and may, therefore, be

involved in motility.

Microbodies
Preparations of different kinds of plant and animal tissues
show a variety of spherical organelles that vary from about
mitochondrial size to a size very much smaller. These
organelles can be distinguished morphologically from
mitochondria, since they have a single outer membrane
rather than a double membrane envelope that is

characteristic of the mitochondria. Cisternae and cristae

are absent from microbodies, and the central area
Figure 3.13 An etioplast from a yellow leaf of a dark-grown dense under the electron
frequently appears rather
bean seedling (Phaseolus vulgaris), x 20,000.
microscope and may contain a variety of crystals (Fig.
3.184).
Considerable research is being carried out to determine
the exact chemical and physiological significance of these
found small plastids that will later develop into normal organelles. Evidence is accumulating that many different
plastids; these are called proplastids. Other colorless
enzymes may be found in single-membrane-bound
plastids, leucoplasts, are found in the epidermal cells of
spherical organelles in both plant and animal cells. To
leaves, in onion, in storage tissue of apples, and in other
some extent, the types of enzymes found depend on the
white tissues. Some of these plastids store large amounts
tissue and kind of cells appears
being studied. Thus, it

of starch and may be called amyloplasts (Figs. 3.12,

that there is not just one kind
microbody but that the
of
3.14).
term "microbody" as defined above includes a fairly large
As fruitsripen on trees and leaves prepare to fall in the
number of different kinds of organelles that contain
end of summer, they change from green to red, orange, or different enzymes and perform different functions in the
yellow. This results from the destruction of chlorophyll,
cell.
accompanied by an accumulation of yellow or red
Beginning students can appreciate some of the
pigments known as the carotenoids. The plastids with a terminology when they realize that
confusion in
dominance of red and yellow pigments are chromoplasts. microbodies isolated from leaves and containing a
A chromoplast from ripe tomato is shown in Fig. 3.14
complex of oxidative enzymes have been called
(right).
peroxisomes (Fig. 3.18/A); microbodies isolated from
castor bean seeds and containing other oxidative enzymes
Dictyosomes
have been called glyoxysomes (Fig. 3.18S).

Dictyosomes are composed of from 5 to 15 circular

flattened vesicles, or cisternae, aligned in stacks (Fig. Nucleus
3.15). In cross sections, many small vesicles appear at the
margins of the cisternae. However, face views reveal that Electron micrographs of nuclei show few details not

the margins of each cisterna form a coarse net with true already recognized by the light microscopist (Figs. 3.4,

vesicles only at the extreme outer regions (Figs. 3.15, 3.5). There are one or more nucleoli and some denser
3/f 6). The peripheral vesicles, in most cases, contain areas embedded in a granular nucleoplasm. The denser
areas may represent sections of chromosomes, but such a
granules of secretory material elaborated in the
relationship has not yet been definitely demonstrated. High
dictyosome cisternae, possibly in collaboration with the
magnification sometimes suggests fibrils in sectioned
endoplasmic reticulum. There is considerable evidence
nuclei.
that the precursors of cell wall material are synthesized in
The nuclear envelope is double and is provided with
the dictyosomes. The stack of cisternae be appears to
pores (Figs. 3.4, 3.6). Nuclear pores are evident in special
polarized; that is, there is a forming face and a concave
disappearing face that is somehow used up in the
preparations of frozen nuclei, when fractured and viewed
formation of the vesicles (Fig. 3.16). through the electron microscope (Fig. 3.19). These pores
appear as distinct holes, and although there is some
Microtubules evidence that rather large molecules can pass through
them, not all investigators agree that they do serve as
Microtubules are hollow tubes of indefinite length and are passageways.

Cell Structure

25
 chloroplast

proplastid

Figure 3.14 Diagram to show the interconversions of plastids;

bottom center (proplastid), center (etioplast), top center
(chloroplast), left (leucoplast) and right (chromoplast).

chapter 3 I The Plant Cell

26
 Figure 3.15 Four dictyosomes in a cell from the root tip of a
water plant (Hydrocharis). One dictyosome has been sectioned at
right angles to its cisternae, and another one almost parallel to its
cisternae. Note the peripheral net around this second dictyosome
The two other dictyosomes have been sectioned obliquely to the
cisternae and some material appears to be passing from them to
the vacuole, x 1 5,000.

endoplasmic reticulum

ribosomes

Figure 3.16 Diagram to show membrane flow from ER through a

dictyosome to the plasmalemma.

Cell Structure

27
Figure 3.17 Microtubules (mt) are shown in a parallel array near
the cell wall, x 62,480.

X >
L '*

|
«T.-:. .:
j^^^
^4

v
> -
.

Figure 3.18 A
electronmicrograph of a type of microbody, the
peroxisome (p), which is found in leaf cells, x 54,500. B, A
second type of microbody, the glyoxysome (G), is found in seeds
and is associated with lipid (L) digestion during seed germination,
x 20,71 4.

chapter 3 | The Plant Cell

28
Figure 3.19 Freeze-etch preparation of an onion root tip cell,
showing the organelles as they appear after being frozen, rather
than chemically fixed. The similarity between the chemical fixation
image and the frozen image is striking, x 10,000.

Cell Structure

29
Nonprotoplasmic Portions of the Cell

Vacuoles
Vacuoles are definitely a part ot the protoplast, but are
involved in passive, or active, mechanisms to remove
organic or inorganic materials trom the protoplasm.
In many young cells the cytoplasm occupies much of
the space in the cell; small vacuoles are, however, present
(Fig. 3.20A). As the cell grows larger, the small vacuoles
within the cytoplasm increase in size, coalesce, and
become fewer in number (Figs. 3.206.C). Finally, when
the cell has attained its mature size, only a few large
vacuoles, or even only one, may remain (Fig. 3.20D). The
protoplasmic contents (nucleus and cytoplasm) of the cell

liecompressed against the cell wall. The nucleus may

occupy a position near the center of the cell, where it is

connected with the cytoplasm around the cell wall by

strands of cytoplasm (Fig. 3.20D).
Vacuoles do not contain pure water, but rather a dilute
solution of many substances. This aqueous solution in the
cell is termed cell sap. Among the substances dissolved
in the cell sap are (a) atmospheric gases, including
nitrogen, oxygen, and carbon dioxide; (b) inorganic salts,
such as nitrates, sulfates, phosphates, and chlorides of
potassium, sodium, calcium, iron, and magnesium; (c)
organic acids, such as oxalic, citric, malic, tartaric; (d)
salts of organic acids; (e) sugars, such as grape sugar
(glucose) and cane sugar (sucrose); (f) water-soluble
proteins, alkaloids, and such as
certain pigments,
anthocyanin. Generally, the sap is slightly acid. The
cell

concentration of cell sap varies from cell to cell, and it

may vary in the same cell during the course of the cell's
life.

The most common pigments of the vacuolar sap are the

anthocyanins. These pigments are responsible for the
red, purple, or blue of the petals of many flowers or of
other parts of plants. The yellow coloration of poppy
flowers is due to yellow vacuolar anthocyaninlike pigments
called anthoxanthins. The red color of roots and leaves of
garden beets is due to another vacuolar pigment called
betacyanin.

The Cell Wall

The protoplast is surrounded by a plasmalemma or

cytoplasmic membrane. Outside this membrane, and
surrounding the entire protoplast, is a rigid wall

synthesized by the protoplast that it encloses. Walls of

adjacent cells are cemented together by an intercellular Figure 3.20 Stages in the growth of a cell. Progressively older
cells shown from A through D, X2000.
substance, the middle lamella (Fig. 3.21 A), which is

composed of pectin (polymers of various sugars,

especially rich in partly oxidized galactose) and certain
other substances. The first wall formed by the protoplast is cellulose impregnated with other substances. Some of
the primary wall, made of cellulose and other these substances, notably lignin, slows decay; others, like

carbohydrates. When the cell ages, the protoplast may suberin and cutin (Figs. 3.21 B,Q, are waxy and protect
deposit more wall material on the primary wall. Thus, a leaves and stems against water loss. In addition, certain
secondary wall (Fig. 3.21 D,E) is formed, and the other materials may enter into the composition of the cell
completely mature cell wall many may have a thickness wall — gums, tannins, minerals, pigments, proteins, fats,

times greater than the primary wall. some tissues, the In and should be emphasized that in mature hard
oils. It

secondary wall is stratified and composed of several tissues, such as wood, lignin may be deposited not only in
layers. In others, the cells do not lay down any secondary the secondary wall but also in the primary wall and middle
wall material. The secondary wall may be of cellulose or of lamella.

chapter 3 l
The Plant Cell

30
 cellulose

middle lamella

Figure 3.21 Development of cell walls during primary growth

while stem is elongating. A, cell with cellulose walls. B, cutin may
be laid down on the outside of an epidermal cell. C, in another
location, suberin may be deposited within the cellulose primary Figure 3.22 Types of inorganic crystals found in the vacuoles of
cell wall to form a cork cell. Lignin is deposited within the primary
and in the cell walls of older nonliving cells. A,
living cells
wall and between the primary cell wall and the protoplast. D, raphides; B, a cluster of crystals; C. a single crystal, x2000.
during rapid elongation, it is laid down as rings or as a spiral
band. E, when elongation stops (except for pits), the lignin forms
a complete secondary wall around the cell.

Although the walls of cells vary considerably in protoplasm if it attains a high concentration in the cell. By
composition in different species, and from one part to its union with calcium, the soluble oxalic acid is converted
another in the same individual plant, cellulose constitutes into the highly insoluble calcium oxalate, which will not
the greatest percentage of the material of which most cell injure the protoplasm. In addition to calcium oxalate,
walls are made. It is synthesized by the protoplast and crystals of calcium sulfate, calcium carbonate, or protein
deposited across the plasmalemma by some mechanism. are sometimes found.
Living cells are interconnected with each other through
cytoplasmic channels called plasmodesmata (Fig. 3.7).
The plasmalemma lines these small channels, and
Summary
endoplasmic reticulum can often be observed within them. 1. A cell may be divided into protoplast and cell wall.
2. Protoplasm is divided into an array of particles,
Inorganic Crystals
membranes, and organelles.
Cells with crystals are found in almost all plants and in 3. The plasmalemma is the membrane separating the
many different plant tissues. Crystals form within vacuoles protoplast from the cell wall, and the tonoplast is the
and vary in chemical composition and in form (Fig. 3.22). membrane separating the vacuole from the cytoplasm.
The most common crystals are of calcium oxalate; it is 4. At least three major theories have been proposed to
generally held that they are an excretory product of the explain membrane structure. The unit membrane
protoplast formed by the union of calcium and oxalic acid. model consists of a bilayer of protein with a middle
This acid is soluble in cell sap and is toxic to the layer of lipid. The subunit and fluid-mosaic models

Summary

31
 propose that globular proteins are embedded in a lipid photosynthetic processes, and respiratory activity all

bilayer. share a common cytoplasm. These are prokaryotic

5. Ribosomes are small particules having a high RNA cells.

content. Polyribosomes are aggregates of ribosomes

that are involved in protein synthesis.
6. The endoplasmic reticulum consists of an extensive
array of flattened vesicles. It may or may not have The Cell Cycle
ribosomes associated with the outer surface of the
vesicles and appears to be involved in protein
synthesis.
Cells don't simply divide, but instead must progress
through a sequence of four precise steps known as the
7. Mitochondria are small organelles about 0.5 jum in
cell cycle (Fig. 3.23). G1 refers to the period preceding
width and from 1 to 3 jum length. They are bounded
in

by a double membrane envelope. The inner

DNA synthesis. During G1 the cell accumulates the
chemical energy and proteins needed before DNA
component invaginates to form cristae, which are
synthesis can occur. For example, if an inhibitor of protein
surrounded by a homogeneous matrix. Respiration is
synthesis were added to a cell in G1 , that cell would be
localized in the mitochondria.
8. Chloroplasts are approximately 5 x 10 /xm in size.
also inhibited from entering DNA synthesis, which is the

They are bounded by a double membrane envelope. second step (S) in the cell cycle. G2 is a third step; it

precedes nuclear division (M). During G2 energy is stored

The aggregate to form
internal chloroplast lamellae
cylindrical grana, which are connected by intergranal
that enables chromosome movement to take place. In
addition, proteins such as those used to construct the
lamellae (frets). The photochemical reactions of
microtubules, which make the spindle fibers, are
photosynthesis take place on the membranes; the
accumulated. Inhibition of any of these events will mean
enzymatic reactions of photosynthesis are located in
that cells will be prevented from dividing.
the stroma.
may contain
Each plant species has a characteristic average amount
9. In addition to chloroplasts, cells
of DNA per nucleus. Pea (Pisum sativum) plants, for
leucoplasts, amyloplasts, proplastids, and _9
example, have 7.9 pg (1 picogram = 10 g) DNA per
chromoplasts.
nucleus; longpod bean (Vicia faba) plants have 24.3 pg
10. Dictyosomes are stacks of from 3 to 10 flattened
per nucleus. Cytologists (scientists who study cells) have
cisternae.Each cisterna is surrounded by a peripheral
net. Dictyosomes appear to be involved in the
established the generalization that the more DNA a cell

has in its nucleus, the longer it will take for that cell to
synthesis of various cellular products.
progress through the cell cycle. As an example, pea cells
11. Microtubules are of indefinite length; they are about 28
take 14 hours to progress once through the cell cycle,
nm in diameter, and have an internal core about 8 nm
in diameter.They are known to be involved in and longpod beans take approximately 18 hours. Each
stage of the cell cycle generally takes a characteristic
chromosome movements, to possibly regulate the
proportion of the total cell cycle time. G1 usually is the
pattern of cell wall fibril deposition, and to somehow
most variable in length; mitosis is usually the shortest
control cell motility.
period. Table 3.2 gives the durations of different stages of
12. Microbodies are all bounded by a single membrane.
the cell cycle in three plants and total cycle time (CT).
They are variable in size and in morphology. They are

closely associated with various types of intracellular M, the last of the four stages of the cell cycle, refers to

enzyme activities. nuclear division, the actual separation of chromosome

13. The nucleus is bounded by a double membrane replicates to form derivative nuclei. This division may be
envelope provided with pores. The nucleoplasm
appears to be characterized by a closely packed array
of unit fibers about 22.5 nm in diameter and of Table 3.2
indefinite length. Cells may
and even differentiate
live Cell Cycle Durations
for a short time without a nucleus; however a nucleus
is required for the continued life of a cell and for cell DNA
division. Amount CT G1 G2 M
14. Vacuoles contain aqueous solutions within the (pg) (hours)
protoplast, and are separated from the cytoplasm by
the tonoplast. Inorganic crystals, when present, are Hellanthus annus 6.3 7.8 1.2 4.5 1.5 0.6
located in vacuoles. (sunflower)
15. The which bounds the protoplast, is formed
cell wall,
Pisum sativum 7.9 14 5 4.5 3 1.2
embedded in an amorphous matrix.
of cellulose fibrils
(pea)
Other compounds (suberin, pectin, cutin, and lignin)
may be present. Plasmodesmata are cytoplasmic Vicia faba 24.3 18 4 9 3.5 1.9
connections between cells through the primary cell (longpod bean)
wall.

16. The type of cell just summarized is highly Data from J. Van't Hot. 1974. The duration of chromosomal DNA
synthesis, the mitotic cycle and meiosis in higher plants. In
compartmentalized. It is known as a eukaryotic cell.
Handbook of Genetics, Vol. II, R. C. King (Ed.). Plenum, New
More primitive cells are not compartmentalized; DNA, York.

chapter 3 l
The Plant Cell

32
 cell

differentiation

Figure 3.23 Diagram to show the stages of the cell cycle: G1

(pre-DNA synthesis phase), S (DNA synthesis phase), G2 (post-
DNA synthesis phase), and M (mitosis).

either by mitosis, which two genetically identical

results in physiological activities including absorption, conduction,
derivative nuclei, or by meiosis, which results in the reproduction, photosynthesis, and support.
formation of four nuclei that have a reduced chromosome The formation of new tissue cannot take place without
complement. Each body cell contains two identical sets of cell division. During the period of division, both nucleus
homologous chromosomes. One set is originally and cytoplasm divide. It is customary to designate the
contributed by each parent; hence a cell before division is period of nuclear division as mitosis and to divide it into
called a diploid cell and is said to have a 2n chromosome four phases: (a) prophase, (b) metaphase, (c) anaphase,
number. A cell derived by meiosis, however, has a and (d) telophase. The division of the cytoplasm is known
reduced chromosome complement and is said to be as cytokinesis. Mitosis gives rise to derivative nuclei
haploid. Haploid cells have a 1n chromosome having identical gene complements. Cytokinesis gives rise

complement. Whether it is mitosis or meiosis that occurs to two new parcels of cytoplasm that are similar, but
depends on the cell type or tissue (group of cells) that is probably never identical The period of
(Fig. 3.24).
dividing. Each type of cell division will now be examined in preparation for division known as interphase and
is

detail. consists of the stages G1 S, and G2. The sequence

, of

events taking place during interphase is equal in

importance, in the complete process of the cell cycle, to

Mitosis the events that take place during mitosis and cytokinesis.

Prophase
Cell division is apparently required because a single
nucleus with a full complement of genes can control only The DNA strands in the interphase nucleus are long,
a small amount of cytoplasm. Increase in size and slender, and seem tangled. The onset of mitosis is
complexity of the plant body requires cell division. The heralded by the presence of definite chromatin threads
resulting increase in the number of cells makes possible (Figs. 3.24A, 3.25A). These threads gradually shorten and
the specialization of tissues for the different structural and thicken and become easier to see (Figs. 3.246, Q. They

The Cell Cycle

33
 nuclear membrane

chromosome romosome chromosome chromosome

Prophase

half-
chromosomes

. cell

plate

kinetochore

40 MM
Metaphase Early anaphase Late anaphase Early telophase

Figure 3.24 Mitosis, diagrammatic representation; A to D, stages

in prophase. Two pairs of chromosomes are represented; the
"green" pair have a median kinetochore, while in the "gray" pair
the kinetochore is close to one end. The chromosomes shorten
and thicken, and chromatids become apparent, E, metaphase; F,
early anaphase; G, late anaphase; H, telophase.

stain more heavily with certain dyes. For this reason they microtubules (Fig. 3.26). These structures, the
are called colored (chromo) bodies (soma), or microtubules, or spindle fibers, plus any adherent
chromosomes. Each chromosome consists of an nucleoplasm is called the mitotic spindle.
individual strand of DNA. The nucleolus slowly decreases The chromosomes are now visibly composed of two
in size and finally disappears during this stage (Figs. closely associated halves, each half being known as a
3.24A to D). chromatid such as corn, which
(Figs. 3.24E.F). In plants,
becomes apparent that at late prophase each
It have been each chromosome can be
intensively studied,
chromosome is composed not of one but of two threads recognized and numbered. There are 20 chromosomes in
coiled about each other (Fig. 3.24Q. The nuclear corn, but only 10 different types that can be distinguished
membrane disappears toward the end of prophase. by their size and form. There are thus two chromosomes
of each type. The 20 chromosomes of corn may be
Metaphase arranged in 10 pairs. Maps have been prepared of corn
Forces active within the cell now arrange the chromosomes showing the relative positions of the genes
chromosomes, or at least a specialized portion of each along them. Since the chromosomes split longitudinally,

chromosome (the kinetochore or centromere), in the each gene is replicated and each chromatid contains a full

equatorial plane of the cell (Figs. 3.24E, 3.25C.D). Spindle set of genes.

fibers are attached to the kinetochores. As the

nucleoplasm elongates, the chromosomes, or at least the Anaphase
kinetochores, are moved to the equatorial plane of the cell
(Figs. 3.24E, 3.25D, 3.26). Spindle fibers extend from the The chromosomes do not remain long in the equatorial
chromosomes to the opposite poles of the cells. Other plane. The chromatids soon separate from each other and
fibers apparently reach to the poles but are not attached move to opposite poles of the cell. This period of
to the chromosomes. The electron microscope separation of chromatids is anaphase (Figs. 3.24F,G,

demonstrates that these spindle fibers are bundles of 3.25D.E).

chapter 3 I
The Plant Cell

34
 Figure 3.25 Mitosis as followed in
living cells of the endosperm of seeds of
the blood lily (Haemanthus katherinae).
(Photographs taken with Nomarski
optics.) A, early prophase; the nuclear
membrane is still present. Note the clear

W£
zone of cytoplasm surrounding the
nucleus. B, the nuclear envelope has
disappeared, and the clear zone still
surrounds the chromosomes. A few
short spindle fibers are already present
C, full metaphase; the coiled chromatids
are distinct. The two chromatids moving
to the upper pole have separated and a
few spindle fibers may be seen. D, early
anaphase; the chromatids are moving to

if % opposite poles of the cell. E,

midanaphase; spindle fibers are in
evidence between the chromosomes
and the poles of the cell. F, telophase;
the chromatids have aggregated at the
opposite poles of the cell. A nuclear
envelope has not yet formed. Time
intervals after A, 6, 14 min; C, 64 min;
D, 74 min; E, 93 min; F, 107 min,
x 2000. G, telophase; the chromosomes
are tightly clustered at opposite poles of
the cell, and the cell plate has started to
form; x4000, H, late telophase; the cell
plate is almost continuous, and fibrils
may be seen extending poleward a short
distance from the plate; X4000
Telophase Cytokinesis
When the divided chromosomes have reached the
In the great majority of cases, the division of the nucleus
opposite poles of the cell, they group together and the
is followed by the division of the cytoplasm. Light
nuclear membrane and become
the nucleolus again
microscopy has demonstrated that a cell plate forms at the
apparent. This period known as telophase (Figs. 3.24/-/,
is
equatorial plane of the cell at right angles to the spindle
3.25Fto H). Cells in telophase will now begin to recycle fibers (Figs. 3.25G,H, 3.28). From electron microscopy we
through the cell cycle (G1-S-G2) (Fig. 3.23). know that the spindle fibers are groups of microtubules. At
the equatorial region, the spindle fibers are surrounded by
Polarity of Cell Division
an amorphous material (Fig. 3.28A). Vesicles appear in

If the spindles in a meristematic tissue were oriented at this region and eventually fuse to form the first barrier
random, an irregular mass of tissue would result. This dividing the derivative protoplasts (Fig. 3.286). This
does occur when a single cell or small group of cells from structure, formed known as the middle
of pectin, is

a carrot root in tissue culture produces an irregular mass lamella. The new formed by the deposition of
cell wall is

of cells called a callus (Fig. 3.27A). For orderly growth cellulose by each protoplast on its side of the middle
there must be a precise orientation of the mitotic spindles. lamella. The resulting structure is the primary cell wall.

A polarity is established so that, in general, the axes of The formation of the primary wall and, subsequently, the
spindles are parallel with each other and with the axis of secondary wall poses an interesting problem of genetic
the shoot or root (Fig. 3. 276). When the spindle axes are control. The cellulose is laid down outside the protoplast,
oriented parallel to the root-shoot axis, the divisions are apparently resulting from the polymerization of many sugar
called anticlinal. When the axes are perpendicular to the molecules to form long, unbranched molecules of

root-shoot axis, the divisions are called periclinal (Fig. cellulose.The molecules are organized into a crystalline
3.276). array by hydrogen bonds to form long unbranched fibrils
Polarity seems to be inherent in cells and in tissues of that are visible in the electron microscope.
which they are a part. Changes in polarity may be induced After removal of the amorphous materials from the wall,
by hormones. Occasionally, the orientation to be assumed the array of fibrils may be studied with the electron
by the spindle can be detected in plants at the cellular microscope. When first deposited by the protoplast, the
level before metaphase. In some instances, the cellular fibrils are in parallel array and form a band around the
organelles will pass largely to one end of the interphase protoplast (Figs. 3.29A 3.30). The mechanism bringing
cell. According to some observations microtubules form a about this precise arrangement is not understood.
circular band at the equatorial plane of the cell during However, the microtubules in the cytoplasm are also
interphase, just before the onset of mitosis. These present in a similar parallel array. It is logical to postulate

microtubules may be involved in establishing the location that the cytoplasmic microtubules may be involved in the
of cytokinesis, but we do not yet know how the placement orientation of the cellulose fibrils.

of the microtubules is controlled. As the cell lengthens, the deposition of fibrils by the

Figure 3.26 Electron micrograph of thin section of metaphase

chromosomes in the endosperm of Haemanthus katherinae,
showing the aligned chromosomes with microtubules (spindle
fibers) attached to the kinetochore. Magnification x 40,000.

chapter 3 l
The Plant Cell

36
Figure 3.27 Influence of polarity on cell divisions and
development of form. A, nonpolarized cell divisions result in an
irregular mass of cells; B, cell division parallel with the axis of the
root (gray arrow and cells) results in growth in length; cell
division parallel with the circumference of the root (dark green
arrow and cells) increases the circumference of the root; cell
division at right angles to the circumference of the root (light
green arrows and cells) increases the diameter of the root.

'

\ l;H

1.;

Figure 3.28 The formation of the cell plate in the endosperm of

Haemanthus katherinae. A, early stage in plate formation; the
microtubules are clustered into groups that are associated with a
denser material, apparently in the plane of the future cell plate,
x 26,000. S, late stage; the microtubules are fewer in number and
are not clustered; vesicles are present and apparently fusing to
form a continuous separation phase between the sister cells,
x 26,000.

The Cell Cycle

37
Figure 3.29 Diagram to show deposition and change in protoplast continues, with no change in the orientation of
orientation of the cellulose fibrils in an elongating primary cell
wall. A, B, and C, represent increasing age and length of the
the newly deposited As elongation continues, the
fibrils.

same cell. The cellulose fibrils are first deposited parallel to the fibrils, now further removed from the protoplast, assume a
circumference of the protoplast. They are then pulled out of this position more in line with the axis of the elongating cell.
orientation as the cell wall elongates. (In the actual wall, the
different stages of fibril orientation would grade into each other.)
A section through a primary cell wall after completion of
Green represents the earliest fibrils deposited, and light gray elongation shows the oldest cellulose fibrils to be parallel
represents those most recently deposited.
to the long axis of the cell; the most recently deposited
cellulose fibrils encircle the cell at right angles to the long
axis (Fig. 3. 29 A to 3.29Q. Intermediate fibrils have
intermediate positions.

microtubules ^ ,
^ j*
Meiosis

Meiosis is a type of nuclear division that is important

•
r i
during the life cycle of plants, because it involves a
reduction in the chromosome number by one half. This
process is significant in sexual life cycles because it offers
a mechanism for genetic exchange and recombination
during reproduction. The role of meiosis in plant life cycles
is discussed further in Chapter 7 and in the chapters
surveying the plant kingdom.
The chromosomes during meiosis are
division stages of
similar to the division stages of chromosomes undergoing
mitosis. Meiosis begins with the chromosomes present as
long, slender uncoiled DNA strands (Figs. 3.31 A, 3.32A).
They proceed to shorten and thicken and split into
. ^
chromatids as in mitosis (Fig. 3.32C) The chromatids
Figure 3.30 Microtubules in the root tip of Arabidopsis thaliana.
The microtubules are oriented parallel to the circumference of the
separate at anaphase. In meiosis (but not in mitosis), there
cell wall and thus parallel to the cellulose fibrils, x 50,000. is a pairing of homologous chromosomes so that four

chapter 3 l
The Plant Cell

38
 *1

** N
f 1 \

i
*• M t&9
Figure 3.31 Meiosis in lily anther. Photomicrographs showing:
A, early prophase Note paired threads. B, late prophase
I. I, each
body represents two paired chromosomes; note chiasmata. C,
late prophase Paired chromosomes; two chiasmata present.
I.

D, metaphase Note spindle fibers extending from chromosomes.

I.

E, anaphase F, telophase
I. G, prophase II. H, metaphase
I. II. /,

anaphase J, telophase
II. II.

The Cell Cycle

39
 > r "\ r

J V. J k
paired homologous
chromosomes

crossover
crossover crossover

r | s t

crossover crossover crossover crossover

Figure 3.32 Diagram of meiosis, two pairs of homologous

chromosomes are shown. A, early prophase with unpaired
chromosomes. B, the homologous chromosomes have paired. C,
the chromatids have formed. Those of a single chromosome are
sister chromatids. Here and in the following diagrams they are
represented as paired parallel lines (this is not the normal
situation). D, markers on the chromosome, future location of
crossover indicated by arrows. E, metaphase Nonsister
I.

chromatids of homologous chromosomes have broken and, at the

crossover region, have exchanged pieces with each other. F,
early anaphase Kinetochores separate; sister chromatids are
I.

still associated at the kinetochore but, because of the crossover,

nonsister chromatids are associated distally to the kinetochore. G,

early telophase The chromosomes start to regroup into a
I.

nucleus; note that there is only one member of a homologous pair

represented. H, interphase. Two sister chromatids attached at the
kinetochores. /,early anaphase II. The chromosomes have moved
to the metaphase plate and the kinetochores have split. Here
II

chromatids are moving to opposite poles of the cell. Note the

disposition of the markers. J, early telophase II. K, four
meiospores are formed. L, meiospores that would arise if the
crossover had occurred distally to the third marker or in the short
arm of the chromosome; the three markers in each chromosome
would have remained linked in their original order.

chapter 3 The Plant Cell

40
r

{ ^ j

The Cell Cycle

41
chromatids, two derived from each ot the pairing Mm, the relationship between these three pairs would not
homologous chromosomes, are associated at mid- change: M, N, and O would remain linked together as
prophase (Figs. 3.328.C). This introduces a complication, would m, n, and o on the chromatids of the original
because the chromatids of paired homologous chromosomes. However, in Fig. 3. 32E chiasmata are
chromosomes are able to exchange partners with each shown between Nn and Oo on the green chromosome
other by tangling their chromosome arms and exchanging and between Rr and Ss on the gray chromosome.
segments (Fig. 3.32E). Such exchanges between Thus, a third step in the prophase of meiosis is (c) the I

chromatids, derived from opposite members of a formation of chiasmata resulting from the breaking and
homologous pair of chromosomes (nonsister chromatids) rejoining of chromatids from homologous chromosomes. It

(Fig. 3.32F), may later be expressed as visible changes in is normal for some chiasmata to form; however, some

the offspring. chromatids produce no chiasmata.

In order for meiosis to take place, there are two As in mitosis, the nucleolus disappears. The nuclear
sequential divisions that result in four cells from a single membrane also disappears during the later stages of
initial cell. However, DNA replication occurs only during prophase I (Figs. 3.318, 3.32C).
the interphase preceding the first division. Each of the four
has only one complete set of
resulting cells, therefore,
Metaphase I

chromosomes. Thus, during meiosis the two sets of At metaphase I (Figs. 3.32C,D), the kinetochores of the
homologous chromosomes found in a diploid parent are paired homologous chromosomes pass to the equator of
reduced to a single set of chromosomes, and the cell or the cell.
plant bearing this single set is said to be haploid.
Since two divisions are required for the completion of Anaphase I

meiosis, it is common
two divisions by the
to designate the
In anaphase I, the kinetochores of whole chromosomes
numerals and and the phases as prophase
I II I,
separate from each other and, with their associated
metaphase anaphase telophase prophase
I, I, I, II,
chromosomes, move to opposite poles of the cell.
metaphase anaphase II, and telophase II.
II,
However, because of chiasmata formation, whole,
complete chromosomes are not separated from each
other, for on the side of the chiasma away from the
Meiosis I
— Firs
.—
t Divisio n
, II !
kinetochore, sister chromatids will separate just as in

mitosis (Fig. 3.32F). Anaphase I of meiosis is thus the

Prophase I
separation of two chromatids. Since the chromatids have
become variously modified, the separation involves both a
During prophase I of meiosis (Figs. 3.31 A, 3.32A,B,C) the separation of whole chromosomes (at the kinetochores)
chromatin contracts to form chromosomes, each with two and a separation of sister chromatids as in mitosis (across
chromatids; in this respect the process resembles mitosis. the chiasma, Fig. 3.32F).
The process is complicated, however, because before the
chromatids become apparent, the homologous Telophase I

chromosomes pair.
Following anaphase I, the chromatids group together at
In so doing, they approach and coil about each other opposite poles of the cell (Figs. 3.31 F, 3.32G) and
(Figs. 3.31 8, 3.32B). The two chromatids of a
immediately prepare for the second meiotic division. Each
chromosome are visible only after the pairing of the
telophase chromosome consists, as usual, of two
homologous chromosomes is well-advanced (Fig. 3.32Q. chromatids. In this sense, the 1n, or haploid number of
The resulting figure is composed of two paired chromosomes, is present. However, must be it

homologous chromosomes, with four chromatids remembered that some of these chromosomes consist of
(Figs. 3.318, 3.32Q. While they are paired, chromatids
two chromatids derived from the same parent (sister
frequently break at several points and rejoin in such a way
chromatids), and others consist of two chromatids derived
that a given reconstituted chromatid may be composed of
from each of the two parents (nonsister chromatids).
parts of four chromatids (Fig. 3.32Q. This breaking and
rejoining of the chromatids is called crossing over, and
the crossover formed by the chromatids involved
interchange known as a chiasma (chiasmata, plural)
is
in the Meiosis — Second Division
(Fig. 3.32E). Chiasmata may occur at any point along the There now occurs a second meiotic division that separates
paired chromosomes. Figure 3.32D represents two pairs of these rearranged chromatids. During prophase (Figs. II

homologous chromosomes, each with three pairs of genes 3.31 G, 3.32H), chromosomes again form, each with two
to serve as markers; Mm, Nn, and Oo on the green composite chromatids. The kinetochores approach the
homologs and Fir, Ss, and Tt on the gray homologs. The equatorial plate, forming metaphase (Figs. 3.31 H, 3.321). II

symbols Mm, Nn, etc. are used to show the hypothetical As in mitosis, the kinetochores now split and separate. In
positions, on homologous chromosomes, of gene loci. anaphase single chromatids move to opposite poles of
II,

Each of these letters refers to a gene on one homologous the cell (Figs. 3.31/, 3.32 J) and are reconstituted in
chromosome; for instance, M is an allele for the telophase into nuclei (Fig. 3.32K"). Each nucleus formed
II

corresponding gene m on the other homolog. The two in this way contains one of the four chromatids derived

together, Mm, are called an allelic pair. chiasmata If from the pairing of homologous chromosomes in prophase
occurred between Oo and the kinetochore, or to the left of (Fig.3.32L).

chapter 3 I
The Plant Cell

42
 Walls develop about each new nucleus and associated 8. The stages of mitosis are prophase, metaphase,
cytoplasm, thus forming cells with the "In or haploid anaphase, and telophase.
number chromosomes. Since the chromatids within
of 9. The spindle, which somehow directs chromosome
each of these four cells
have been variously modified by movement, is constructed from many microtubules.
crossing over, each of the four cells may be genetically 10. The middle lamella is the first layer to separate the two
different. In what specific way do they differ and of what derivative protoplasts.
importance is this variance in the life cycle of plants 9 11. Vesicles collect around the microtubules in the
equatorial plane of the cell. They fuse to form the
middle lamella. Cellulose formed by the two daughter
Recombination of Genes protoplasts and deposited on the middle lamella forms
the primary wall.
The distribution of genes during meiosis depends on the 12. Meiosis involves two divisions with only one period of
separation of the chromatids, and on the location and the DNA replication.

amount of interchange taking place between the 13. Meiosis is different from mitosis in that during
chromatids. If, for instance, as in Fig. 3.32D, the prophase I homologous chromosomes join
pairs of
interchange took place to the left of Mm, then M, N, and O together. After both divisions each new haploid cell
would move with the kinetochore at anaphase and the I, contains one half the number of chromosomes of its
two telophase nuclei would contain the markers M, N, O
I
parent cell.

and m, n, o; these nuclei would remain linked together so 14. Meiosis is important because its mechanisms for

that only two types of meiospores M, N, O and m, n, o recombination of genes allow variation in the genes of
(Fig. 3.32L) would result. The same would happen if the resulting progeny.
exchange occurred to the left of Rr. T, S, and R would
remain linked, as would s, and r. The number of different
t,

kinds of meiospores to be derived from one to several DNA and its Replication
linked allelic pairs is two or four, depending on whether
crossing over has occurred between the alleles. Without Deoxyribose nucleic acid, DNA, is the relatively simple
crossing over, two of the meiospores would be similar; molecule in which, it appears, is stored all the information

with crossing over, all four meiospores would be different.

needed for the development of the several hundred
The proportion of meiospores having crossover thousand plant species, past and present.
chromosomes in a large number of meiotic divisions will There may be variations in the type and distribution of
depend on the number and location of interchanges. the nitrogen bases in DNA from different species, but
there is no reason to suppose that there has ever been
Now, consider the distribution of the genes, Oo and Tt,
which are on nonhomologous chromosomes. Each of the any fundamentally different kind of carrier of genetic
four meiospores are different in the example shown; they
information than the DNA known today.

are OT, Ot, oT, and ot.

Enzymes regulate metabolic reactions in the cell. Each
Combining segregation of chromatids during meiosis different kind of enzyme is a different protein and regulates
a different reactionin the metabolism, growth, and
with chromatid interchanges, or crossing over, brings
about variation of the genetic constitution of the resulting
development of the cell. Since each kind of protein has its
meiospores or gametes. In either case, gametes of
own exact sequence of amino acids, the way in which the
cell regulates the sequence of amino acid in proteins is
different chromosomal makeup will ultimately fuse to
very important (Chapter 2). We now know that the basis of
produce a new diploid plant that may be quite genetically
genetic information is the sequence of bases that each
different from the parent. Thus, meiosis greatly enhances
the possibility for variation, and results in genetic
part of the DNA molecule contains. Precise duplication of

recombination from generation to generation.

DNA insures that each new cell contains the genetic
information needed for all of its potential functions.

Summary of the Cell Cycle

Replication of DNA
1. The cell cycle consists of G1-S-G2-M. G1 and G2
are important, because it is during this time that cells We shall nowconsider the manner in which DNA itseif is

. 'prepare for DNA synthesis (S) and nuclear division replicated. From each double helix (Fig. 2.13) two new
(M). double helices must form, without at the same time,
2. Cell division increases the numbers of cells. disturbing the precise ordering of the base pairs (Fig.
3. All cells have the potentiality to divide but are normally 2.12). The dividing double helix may be several million
blocked from doing so. nanometers long and, since a single base pair occupies
4. Nuclear division is called mitosis; cytoplasmic division the space of only a few nanometers, there are many base
is called cytokinesis. pairs along a double helix.
5. The derivative nuclei are genetically identical. Precise information telling us the exact time and location
6. The derivative parcels of cytoplasm may be unlike of the synthesis of new DNA may be obtained by using

each other, but are usually similar. radioactive precursors of nucleic acid. A commonly used
7. In general, mitotic spindles are oriented parallel to substance of this type is radioactive thymidine. It seems to
each other and to the axis of the shoot. A definite, enter cells and to be directly incorporated into nuclei of
predetermined polarity produces this orientation. dividing cells of both plants and animals. Root tips of

Summary of the Cell Cycle

43
Tradescantia (a common house plant) grown in a culture
solution containing radioactive thymidine (a torm of the
DNA base thymine) show that interphase may be divided
into three periods (the cell cycle). During the first stages
the nucleus increases in volume, but there seems to be no
incorporation of radioactive thymidine; this is G1. Later,
radioactive material is incorporated into newly synthesized
DNA (s). A period follows in which the nucleus maintains
a constant size without any further incorporation of labeled
thymidine (G2). Refer back to Table 3.2 to reexamine the
time it takes for a cell to transverse each stage of the cell
cycle.
But these observations do not tell us the manner of
replication of the double-stranded DNA helix. It is possible
that a whole new double-stranded helix might be
replicated, thus totally conserving the old double helix
Or the coiled double strands could somehow
intact.

separate and a new strand could be formed by each old

strand. This is semiconservative; half of the new helix
contains old DNA. Or the replication could take place at
random throughout the DNA molecule.
Careful experiments conducted by Dr. J. Herbert Taylor
when he was at Columbia University, demonstrated that
DNA replication was, in fact, semiconservative. The
mechanism for this replication can be explained in the
following way.
Assume that a double helix is present in early
interphase at the onset of replication. This helix may
untwist, separating the two strands (Fig. 3.33). Each
strand now serves as a template for the formation of a
new strand. Thus, following replication with radioactive
thymidine available, each chromatid will contain a strand
bearing the radioactive material. In a subsequent division
in the absence of radioactive thymidine, the radioactive
old
strand will serve as a template for a new nonradioactive
strand as well as the old nonradioactive strand.

Figure 3.33 Schematic diagram showing how one "old" double-

stranded molecule of DNA could be replicated into two new
double-stranded molecules of DNA. Each new molecule has one thymine
ne — '
*— adenine *— guanine >— cytosir
old and one newly synthesized strand.

chapter 3 I
The Plant Cell

44
 —

CHAPTER

4 the plant body

The vegetative plant body consists of three organs

stems, leaves and roots. In this chapter we will
the distribution of internal cells and tissues (anatomy).
Stems are elongated organs that form the axis on which
examine their external form (morphology) and the lateral appendages, leaves and buds, are attached
internal structure (anatomy). One
serious problem in The place on a stem where leaves and buds are attached
presenting this material photographs and drawings
is that is called a node, the portion between nodes is an

are two-dimensional, and words are often inadequate to internode. Some plants have erect stems, others have
explain the pattern and symmetry of plant parts. So, while horizontal, creeping stems (Fig. 4.35), and some have
reading these sections, you should visualize the plant's stems that do not elongate except at flowering time.
internal structure in three-dimensions. Try touse your (These are rosette plants, Fig. 4.1 C.) All stems whether
"mind's eye" as a kind of X-ray vision to see the length, short or long, horizontal or erect, are distinguishable as
width, and breadth of cells and tissues and their stems by the presence of nodes and internodes.
interconnections. It will become apparent, as you study
plant structure, that all organs are integrated functionally
and structurally. This means that the entire plant acts as Arrangement of Leaves and Buds
one unit, and is not simply an aggregate of isolated parts.
A three-year-old twig of walnut in winter condition is

shown in Fig. 4.2. The the twig generally bears a

PART ONE tip of

large terminal leaf bud. At regular intervals along the

stem, other buds may be seen; they are called lateral
Stems buds. Note that below the base of each lateral bud there
is a scar that was made when a leaf fell from the twig; this

Stems provide mechanical support for leaves in erect is a leaf scar. Vascular bundle scars (Fig. 4.2) may be
seen within each leaf scar; strands of food- and water-
plants, and are an axis for attached leaves in horizontal
conducting tissues passing from the stem into the leaf
plants. Flowers and fruits are also produced in positions
were broken when the leaf fell, leaving these scars. Buds
on stems that allow for pollination and seed dispersal.
and leaves are usually borne in this relationship to each
Stems provide a pathway for the conduction of water
other; buds form in the angle made by the stem and the
and mineral nutrients from roots to leaves, and for transfer
leaf stalk. This angle is termed the leaf axil, and
of foods, hormones, and other metabolites from one part
consequently these buds may also be called axillary buds.
of the plant to another.
Protecting the young immature leaves and cells within
The normal life span of plant cells is from 1 to 3 years.
the bud is a series of overlapping scales, bud scales.
Water and mineral salts in dilute solution move in dead
They are usually shed when the bud develops into a new
cells, but this movement depends upon the activity of
shoot, and they also leave scars, bud-scale scars. The
living cells in leaves and roots that are generally less than
part of a stem or twig between sets of terminal-bud-scale
three years old. Stems in herbaceous perennials (Fig.
scars is generally formed during one growing season. For
4.1 A) and in 2000-year-old redwoods, or bristlecone pines
instance, growth made by the twig this year is set off from
(Fig. 4.1 B), annually provide new living tissue for normal
growth made last year by means of a ring or girdle or
metabolism of the plant. Other stems are modified for the
storage of plant products.
terminal-bud-scp'3 scars (Fig. 4.2). When scales of a
terminal bud fall off in spring, they leave a number of
Stems thus have four major functions: (a) support; (b)
closely crowded scars that form a distinct ring.
conduction; (c) the production of new living tissue; and (d)
Examination of twigs several years old shows that growth
storage.
in length may vary from year to year, as revealed by the
different spacings between the terminal-bud-scale scars.
There is no increase or decrease in the length of any
Stem Morphology portion of a stem after that portion one year old. is

The slightly raised areas on the bark are lenticels. They

are composed of cells that fit loosely together, with air
The organs of plants can usually be distinguished from spaces between, which permit passage of gases inward
one another by their external shape (morphology) or by and outward.

45
Figure 4.1 A, bristlecone pine (Pinus aristata); gymnosperms
and woody angiosperms achieve longevity through secondary
growth, which annually results in a cylinder of new tissue around
the trunk. The older tissues die. S, Iris; many other seed plants
attain longevity by the continued growth of new primary tissue at
the apex of the shoot. Older parts of the stem die. New leaves
form at the apex, roots arise in nodes behind the apex, and older
portions of rhizome die. C, Echeveria sp. rosette plant with
shortened internodes. Note the elongated internodes of the floral
branches to the left and right.

Position of Buds on a Woody Twig

In is just one leaf bud and one leaf
the walnut twig there accessory buds. Walnut (Juglans sp.) may also have
at each node. This arrangement of buds and leaves on the more than one bud in the leaf axil (Fig. 4.3/A).
stem is spoken of as alternate (Figs. 4.3A AAA.C.E). is It Not infrequently, buds may arise on the plant at places
the most common type of bud and leaf arrangement. Ash, other than leaf axils. They may appear on stems, roots, or
maple, lilac, and many other plants have two leaves even leaves, and give rise to new shoots. Such buds are
opposite each other at each node, and a bud in the axil of called adventitious buds. Their formation may be
each leaf (Figs. 4.38, 4.4D.F). This arrangement of leaves stimulated by injury, such as occurs in pruning.
and buds is spoken of as opposite. When three or more Dormant or latent buds arise in a regular fashion in the
leaves and buds occur at each node, as in Catalpa, leaf leaf axil, but their development is usually inhibited by the
arrangement and bud arrangement are said to be whorled dominance of the terminal bud. This mechanism, apical
(Fig. 4.3G). dominance, is fully discussed in Chapter 8.
Some plants have several buds in or near the leaf axil. From the above discussion is seen that buds may be
it

For example, apricot (Prunus sp.) often has a group of classified by their arrangement on the stem, which may be
three buds in the leaf axil: a central bud, which develops (a) alternate, (b) opposite, or (c) whorled; by their position
into a side branch, and two lateral ones, which are called on the stem, which may by (a) terminal, (b) lateral

chapter 4 |
The Plant Body

46
 —

terminal bud (axillary), (c) accessory, or (d) adventitious; and by the

nature of the organs into which they develop, which may
be (a) leaf, (b) flower, or (c) mixed.
As a rule, the terminal bud of a stem is the most active
and grows more vigorously than any of the axillary buds.
Usually, the lowest lateral buds on a year's growth of the
shoot remain dormant and do not develop into branches. If
the terminal bud is removed, however, as may be done in
pruning, lateral buds, otherwise dormant, may become
active.

Development of Tissues of
the Primary Plant Body
The Primary Meristems
Plant organs (leaves, stems, roots, and flower parts) are
obviously different from each other morphologically (i.e..

based on their external form). But, if we examine their

internal anatomy is apparent that
it all organs are
composed of similar structural units — cells and tissues
that are dissimilarly arranged. Each cell (cell type) is

modified to make it ideally suited to perform one or more

specific functions. Tissues are organizations of cells of
one or more types that have a common origin and a
common collective function.
Cell types and tissues develop by a process called
differentiation. A differentiating cell progresses through a
series of steps that result in the cell becoming mature and
functional. Each different cell type "experiences" slightly

different differentiation steps. The formation of new cells,

and the initiation of differentiation in plants takes place in

specific regions calledmeristems. Meristems can also be

categorized. The apical meristems occur in the shoot
and root tips. Apical meristems are the source of all other
meristems. They form the three primary meristematic
tissues (or primary meristems): protoderm, ground
meristem, and procambium. These three primary
two years meristems will differentiate into the three primary tissues:
ago epidermis, ground tissues ("pith and cortex") and
vascular tissues (xylem and phloem). Now we will
consider in detail each of the primary meristems and
Figure 4.2 Three-year-old twig of walnut (Juglans regia), x 1
tissues in the plant body.

Protoderm

This is the outermost layer of cells (Figs. 4.5A.D, 4.6). It

develops into epidermis — the special primary tissue that

covers and protects all underlying primary tissues. The
epidermis prevents excessive water loss and yet allows for
the exchange of gases necessary for respiration and
photosynthesis.

Ground Meristem
The ground meristem comprises the greater portion of
meristematic tissue of the shoot tip (Figs. A.5B.D, 4.6). Its

cells are relatively large, thin-walled, and isodiametric. The

regions forming from the ground meristem are (a) pith, in

the very center of the stem, and (b) cortex, in a cylinder

just beneath the epidermis and surrounding the vascular

Development of T issues of the Primary Plant Body

47
Figure 4.3 Twigs showing three methods ot bud and leaf
arrangement. The position of leaves is shown by the leaf bases
and scars. A, alternate, walnut (Juglans regia); B, opposite, lilac
(Syringa vulgaris); C, whorled, Catalpa, x '/2 -

tissue. A note of clarification must be added here. Primary Tissues

Traditionally speaking, the pith and cortex are not tissues,
but instead are stem and root regions that are composed Primary tissues stem are differentiated from the
of the
of the ground tissues, parenchyma, sclerenchyma, and three primary meristematic tissues —
protoderm, ground
collenchyma. These will be discussed in the following meristem, and procambium —
and that these three are
section. derived from the apical meristem of the shoot tip. In
woody plants, we must look for primary tissues of the stem
a very short distance behind the stem tip. Even before the
Procambium end of the first season's growth, differentiation of these
primary tissues from primary meristematic tissues is

These cells usually appear first as strands among ground completed, and secondary tissues may be formed in

meristem cells (Fig. 4.6). In cross section (Fig. 4.5C.D) abundance. Whereas primary tissues are derived from
strands appear as isolated groups of cells arranged in a primary meristematic tissues of the shoot, secondary
circle.Sometimes a continuous procambium cylinder is tissues are the result of production of new cells by the
formed. As seen in a transverse section of a single vascular cambium and by the cork cambium. The
procambium strand, procambium cells are smaller than origins and nature of these two types of cambia are
those of the surrounding ground meristem; in longitudinal discussed later.

section, they are much longer, and some of them may be

pointed at the ends. Procambium cells give rise to primary
The Epidermis
vascular tissues (Figs. 4.5E.F, 4.7). These primary tissues The epidermis is usually a single superficial layer of cells
carry out several functions and are divided rather rigidly that covers all other primary tissues, protecting them from
into two groups according to these functions. Food drying out and, to some extent, from mechanical injury. It

synthesized by photosynthesis is conducted in the is the limiting layer of cells between the plant and its

phloem. Water and mineral salts are conducted in xylem. environment. In surface views (Fig. 4.7, 4.8), epidermal

chapter 4 The Plant Body

48
 leaf bud

leof scar

bundle scar

leaf bud
leaf scar

bundle scars

Figure 4.4 Leaf scars and bud arrangement of different species

of woodyplants. A, walnut (Juglans regia), x2; B, catalpa
(Catalpa bignonioides), x2'/2 C, tree of heaven (Ailanthus
;

altissma) x 2; D, box elder (Acer negundo), x 2; E European

,

plane (Platanus acerifolia), x3; F, buckeye (Aesculus californica),

x2.

Development of Tissues of the Primary Plant Body

49
 Figure 4.5 A to F: diagrams showing the pattern ot vascular
development and the positions ot the three primary meristems.
apical Vj.K
meristem

leaf primordium

protoderm

procambium

ground meristem

•A?*- bud primordium

Figure 4.6 Longitudinal section of the shoot apex of the bean

(Phaseolus vulgaris), x 90.

chapter 4 |
The Plan t Body

50
 epidermis

procambium

Figure 4.7 Cross sectional view of clover (Trifolium sp.) stem

showing the primary vascular tissues, xylem and phloem. The
epidermis, and the stem regions, the cortex and pith are also
illustrated, x339.

called guard cells. Between each pair of guard cells is a

epidermis
small opening, or pore, through which gases enter and
guard cells
leave the underlying stem tissues. Two guard cells plus
the pore make one stoma (pi. stomata) (Fig. 4.8A). Guard
cells differ from other epidermal cells by their crescent
shape and the fact that they contain chloroplasts. Stomata
are common in the epidermis of leaves, floral structures,
and fruits as well as of stems. Stomata are discussed more
thoroughly in Part II of this chapter.
Epidermal appendages, such as hairs, may occur on
young stems (Fig. 4.44).These structures will also be
discussed in the section on leaves.

The Cortex
This complex region, derived from ground meristem, forms
beneath the epidermis as a cylindrical zone that extends
inward to the primary phloem (Figs. 4.7 A,B, 4.15). The
following tissues or cell types may be found within it:

Figure 4.8 Epidermis; cross section of alfalfa (Medicago sativa) parenchyma, collenchyma, sclerenchyma, and
stem showing epidermal cells, guard cells, and cortex with a secretory tissue.
substomatal chamber, x400.

Parenchyma. The principal tissue of the cortex is

parenchyma (Figs. 4.7, 4.9G,/) consists of isodiametric
It

cells with thin walls, made mostly of cellulose, and with

cells are elongated in the direction of the stem's length; in protoplasts that remain alive for a long time. We may
transverse section, they are usually isodiametric. Its speak either of a parenchyma tissue, where many
protoplasm forms a thin layer that lines cell cavities and parenchyma cells are found together, or of individual
normally retains its living properties for a long period. parenchyma cells.
The outer tangential wall of cells exposed to air is Parenchyma tissue is characterized by the presence of
usually thicker than the other walls, and its surface layer intercellular air spaces, which vary greatly in size; in some
is usually coated with a waxy substance called cutin. This parenchyma tissues they are difficult to find, while in

superficial layer is the cuticle, which is quite impermeable others they are very apparent (Fig. 4.7). Because
to water and gases. There may be cracks or other parenchyma cells retain active protoplasts, they function in
imperfections in the cuticle, however, and water vapor the storage of water and food, in photosynthesis, and
may pass out of the plant at those points. The inner walls, sometimes in secretion. When the parenchyma cells
parallel to the stem surface, are thinnest, and radial walls, contain chloroplasts, they are collectively referred to as
at right angles to the surface, often taper in thickness chlorenchyma. Parenchyma cells can be reprogrammed
toward the inner wall (Fig. 4.8). to differentiate into different cell types. One example would
Young stems usually possess specialized epidermal cells be the changes parenchyma cells undergo in response to

Development of Tissues of the Primary Plant Body

51
 cytoplasm

Figure 4.9 Cell types and tissues. A and B, fibers in longitudinal

view; C, fibers in cross section; D, collenchyma in cross section;
E, collenchyma in lontitudinal view; F, sclereid; G, parenchyma;
H, stone cells; /, woody parenchyma.

chapter 4 |
The Plant B ody

52
being wounded. In this case, the wounded cells on the
surface simply die, and the inner cells divide and torm a
layer that is quite similar to the bark on the outside of
stems.
Parenchyma cells are not confined only to the cortex
and pith of the stem, but also occur in practically all other
types of tissue and in other organs of the plant.

Collenchyma. The outermost cells of the cortex of

young stems, lying just beneath the epidermis, often
known as collenchyma. This tissue
constitute a tissue
may form a complete cylinder, or may occur in separate it

strands. Collenchyma cells are elongated, often contain

Figure 4.10 Transfer cells (A) around vascular bundle in leaf of
chloroplasts, and are living at maturity. The walls of
Armeria Corsica, x 1887.
collenchyma cells are composed of alternating layers of
pectin and cellulose. In the most common type of
collenchyma, the cell walls are thickened at the corners
(Figs. 4.9D.E). These thickenings are quite flexible and will
Transfer Cells. Frequently, cells located in positions of
stretch without resistance, in much the same way as active solute transfer will show an irregular extension of
heated plastic. Collenchyma is, therefore, an ideal the cell wall into the protoplast. Since the plasmalemma
strengthening tissue because it strengthens and at the follows the contour of the wall, its surface is greatly
same time allows normal tissue growth. Collenchyma extended. Mitochondria appear to aggregate adjacent to
serves as a strengthening tissue in young expanding these areas of increased membrane surface. The
stems and also in the petioles of leaves. morphological picture thus presented (Fig. 4.10) suggests
an adaptation that facilitates transport from one cell to

another, or from the interior to the exterior of the plant.

Sclerenchyma. The main functions of sclerenchyma
These cells have been called transfer cells.
cells are support and, in many cases, protection. Their
shape and the thickness and toughness of their walls The Pith
contribute to the ability of these cells to support and
protect the stem. Thickness and toughness of walls are The pith makes up the central core of many stems. It is
increased by deposition, within the original cellulose wall, composed mostly of parenchyma cells that store food
of a substance known as lignin. Lignin is made by the products such as starch. Sclerenchyma cells (especially
protoplast and deposited on the wall. When this secondary sclereids) can also be found in the pith. The cellular

wall is completely formed, the protoplast usually dies. region between vascular bundles is sometimes called a
pith ray (Fig. 4.5F).
There are two types of sclerenchyma cells: (a) sclereids
and (b) fibers (Figs. 4.9AS.F.H).
Of various types of sclereids, the most common are
stone cells. Other types of sclereids are branched,
resembling very irregular stars (Fig. 4.9F). Some sclereids The Primary Vascular Tissues
are derived from parenchyma cells by a pronounced
The term "vascular" pertains to tissues that conduct
thickening of cell walls; others arise from separate
various substances in liquid form.
meristematic cells.
In vascular plants, water and different water-soluble
Sclereids occur not only in the cortex of stems but also
inorganic salts from soil, as well as food substances, are
in the hard shells of fruits, seed coats, and bark, in pith of
conducted throughout the plant in well-defined vascular
some stems, and in certain leaves.
tissues.
Fibers are elongated, strengthening cells that are thick-
In a young stem, very near the tip (Figs. 4.5, 4.6),
walled, and usually pointed at the ends (Figs. 4.9AS).
vascular tissues occur as separate bundles, primary
Their walls may or may not be lignified. The walls may
vascular bundles. Each primary vascular bundle is
become so thick that the cell cavity, the lumen, almost
from a procambium strand and consists
differentiated of
disappears. Simple pits form in their thick walls (Fig.
primary xylem and primary phloem. If secondary growth is
4.9/A)- Fibers are sometimes very elastic and can be
to occur, a thin band of meristematic tissue destined to
stretched to a great degree without losing their ability to
become vascular cambium remains between primary
return to their original length. Protoplasts of fibers often
xylem and primary phloem (Fig. 4.17).
disappear as the cells attain maturity.
The Primary Phloem
Secretory Cells. Secretory cells are parenchymalike and Phloem in angiosperms may possess several types of
contain dense protoplasm. They secrete various cells: sieve-tube members, companion cells, fibers,
substances, such as resinous materials (Fig. 4.27) and parenchyma. A sieve tube is a vertical
sclereids, and
nectar in many flowers. Many epidermal hairs are row each cell is known as a sieve-tube
of elongated cells;
secretory cells (Fig. 4.44C). member. These are the conducting elements of phloem

Development of Tissues of the Primary Plant Body

53
(Fig. 4.1 1) that translocate sugars, produced in the leaves companion cells have a normal protoplast with a full
by photosynthesis, to other plant parts (see Chapter 5). complement of organelles (Fig. 4.1 1), these cells possibly
Among angiosperms, a sieve-tube member and a regulate the metabolic activity of the sieve-tube members
companion cell are sister cells; that is, they originate by that have no nucleus. Plasmodesmata connect the
division from the same procambial cell. Young sieve protoplasts of companion cells and sieve-tube members.
elements have the usual complement of organelles: A characteristic structural feature of mature sieve-tube
nucleus, plastids, mitochondria, and dictyosomes. As the members is the sieve plate. It may occur in the end or
element matures, its protoplast becomes greatly modified. side walls (Fig. 4.1 1). The end wall between two adjacent
Its nucleus is thought to disintegrate. Plastids lose most of sieve-tube members is thickened and strands of cytoplasm
their internal membranes, but usually retain starch. pass through pores adjoining them. With the exception of
The mitochondria become small. The cytoplasm, much a few trees such as palms, where they live longer, sieve-
reduced in amount, becomes reduced to a thin peripheral tube members live and function about 1 to 3 years. New
layer. The central part of the cell is occupied by a mass of ones are formed annually.
strands or tubules. This mass may be seen with the light In many studies on the structure of mature sieve-tube

microscope, and has been called slime. Since is now it members, a carbohydrate known as callose is seen
known to be a protein, is more correctly referred to as
it around the margins of pores in the sieve plate (Fig. 4.12).
P-protein (Fig. 4.11). At maturity one or more companion In some instances, protein may also collect at the sieve
cells lie adjacent to each sieve-tube member. Since plate. Obviously, this would block the pores of the sieve

sieve plate sieve— tube members

phloem
parenchyma
cells

phloem parenchyma

companion ce

sieve -tube plastids

parenchyma plastid

Figure 4.1 1 Phloem tissue from the stem of tobacco (Nicotiana),

X400.

chapter 4 |
The Plant Body

54
plate and obstruct movement ot food materials. It has been not alive The tracheid wallmay not be the same thickness
demonstrated in other tissues that callose forms very throughout All the wall may be thickened except for
rapidly in response to wounding. numerous small, circular, or oval areas called pits (Fig.
In gymnosperms there are sieve cells rather than sieve- 4.13Q. There are two types of pits in xylem cells: simple
tube members; these have tapered end walls without sieve pits and bordered pits. Simple pits as shown in Figs.
plates and they do not connect to form sieve tubes. 4.9/4 and occur in fibers, sclereids, and in parenchyma
/,

cells when they have secondary walls. Pits form opposite

The Primary Xylem each other in the secondary walls of adjacent cells. A pit-

pair is not a hole in the wall, since the primary wall and
The conducting cells that occur in primary xylem of the middle lamella of the two communicating cells remain
vascular plants are tracheids and vessel elements. These These primary layers, however, are penetrated by
intact.

cells conduct water and mineral salts. Associated with plasmodesmata while the cells are living. The type of pit
them may be fibers (xylem fibers) and parenchyma known as a bordered pit (Figs. 4.13A.E) occurs in

(xylem parenchyma). tracheids, vessel elements, and some xylem fibers. This
A tracheid is a single elongated cell more or less type of pit is more structurally complex. It consists of an
pointed at its ends (Fig. 4.13). Functioning tracheids are expanded border of the secondary cell wall that extends

**v
-

- ><^!IBM

P— protein body

sieve-tube men-.ber

pore

sieve plate

sieve-tube member

Figure 4.12 A, longitudinal section of three sieve-tubemembers

and one companion The nucleus and various organelles are
cell.
present in the central companion cell. Most organelles are
missing in the left sieve-tube member, but are present in the one
on the right, Cucurbita maxima, x 8300. B, micrograph of sieve
plate between two sieve-tube members from Cucurbita maxima,
X4600.

Development of Tissues of t he Primary Plant B ody

55
secondary wall

border

torus

primary wa

Figure 4.13 Tracheids and vessel elements from secondary

wood. A, tracheid from spring wood of white pine (Pinus); B, tip
of tracheid from wood of oak (Ouercus); C, tip of vessel element
from wood of Magnolia; D, tip of vessel element from wood of
basswood (Tilia); E, diagram of bordered pit.

over a small pit chamber. Within the chamber is a tracheids, and most xylem fibers. They function in the
diaphragmlike primary cell wall. In gymnosperms, the storage of water and foods, which, as we have learned, is

center portion, the torus, is thickened and impregnated one of the principal functions of parenchyma wherever it

with a waxy material. The torus apparently acts as a valve occurs in the plant. Parenchyma may also conduct
that plugs the pit opening during times of drought materials for short distances.
(Chapter 5). Xylem fibers are similar to the fibers described
A vessel element is a single cell with oblique, pointed, elsewhere.
or transverse ends. A vessel is a series of vessel elements
differentiating end to end, with perforated end walls.
Vessels are often several centimeters long, and in some
Summary of Primary Tissues
vines and trees they may be many meters in length. 1. Buds are characteristic of woody stems. Bud scales
Before the protoplasts disappear, vessel walls become enclose and protect rudimentary leaves surrounding an
thickened, forming a secondary wall (Fig. 4.14); the apex. The apex may be either a vegetative shoot or a
thickening material is laid down on primary walls in floral apex. A apex terminates growth. Woody
floral
various patterns so that in some places the secondary plants each year require new vegetative growth, which
walls are thick and in others thin. The material deposited is normally accompanied by flowers.

is cellulose; later, the layers of cellulose become lignified. 2. Primary growth brings about the elongation of stems
The end walls of the vessel elements also dissolves before and establishes the basic pattern of cells and primary
the protoplasts disappear. Thus, the deposition of tissues characteristic of the particular stem and upon
thickening material forming the secondary walls and the which the functioning and future growth of the stem
dissolution of end walls are functions of living cells. After depend.
these events have taken place, the protoplast dies. 3. Primary meristematic tissues are apical meristem,
The secondary walls of vessels in angiosperm stems are protoderm, ground meristem, and procambium.
deposited in several different patterns (Figs. 4.14A8): 4. The primary plant body is composed of (a) the
annular, scalariform, reticulate, and pitted. The ends of epidermis, which may be differentiated into three cell
the vessel elements (Figs. 4.1 3AD), are generally on a types: epidermal, guard cells, and epidermal hairs; (b)
slant and, although open, they may have bars of wall cortex,which is composed of collenchyma,
material across them. The shape of vessel elements may sclerenchyma (fibers andand parenchyma;
sclereids),
indicate evolutionary relationships among plants (Chapter (c) vascular tissue, composed
xylem (fibers, of

10). Vessel elements do not occur in small veins in leaves, tracheids, vessel elements, and parenchyma) and
and are lacking most gymnosperms and in the lower
in phloem (fibers, sieve-tube members, companion cells,
vascular plants. In these forms, tracheids occur in and parenchyma); (d) pith, composed largely of
elongating regions, and they may have annular and spiral parenchyma and sometimes accompanied by sclereids;
types of secondary walls. (e) pith rays, composed of parenchyma cells.
Xylem parenchyma cells outlive vessel elements, 5. The functions of these cells and tissues are as follows.

chapter 4 |
The Plant Body

56
 (-parenchyma cells

annular
r reticulate
-- sp r pitted

Figure 4.14 Types of vessels that occur in primary xylem in an

elongating branch. A, annular vessels were formed first, and are
therefore the oldest and most stretched. The pitted vessel formed
last is the youngest. Elongation has stopped, so the pitted vessels
will not be stretched. B, photomicrograph of vessels during
primary growth in cleared node of Alternanthera, x 30.

Parenchyma is used for the storage of water and food which will continue to differentiate into the cortex and pith,

and the conduction of materials for short distances. respectively.

Collenchyma, sclereids, fibers, and tracheids are the Between these two tissues, near the apex, is a ring of
strengthening or mechanical tissue elements. Tracheids residual meristem cells that retain the cellular
and vessels (series of vessel elements) conduct water characteristics of the apical meristem (Fig. 4.5Q. This ring
and mineral salts. Sieve tubes (series of sieve-tube will become a cylinder of discrete procambium strands
members) conduct foods. that later differentiate into primary xylem and phloem
Collenchyma and sclerenchyma may occur in patches (vascular bundles). Each bundle is separated from others
or completely surround the stem just underneath the by regions of parenchyma cells (Figs. 4.5D.E, 4.15). The
epidermis. Sclerenchyma is frequently associated with formation of vascular bundles from the residual meristem
vascular bundles. is apparently in response to leaf development. The
mechanism is not exactly understood, but it may involve
the production of some substance by the leaf primordia
(immature leaves on the shoot tip) and its transport
The Dicotyledonous Stem downward to the residual meristem ring. When the
substance reaches a certain critical concentration, it

Stem Primary Growth apparently induces the residual meristem cells to form
bundles of procambium and then xylem and phloem. Each
The tip of the stem consists of small immature leaves bundle is called a leaf trace and, as it matures, will lead

enclosing a dome-shaped apical meristem. The apical from the stem into a leaf, connecting it to the axis of the

meristem is composed of dividing cells, arranged in stem (Fig. 4.47). The vascular system of the stem actually
various ways, that give rise to the leaves, buds, and the consists, for themost part, of interconnected leaf traces.
primary meristematic tissues (Figs. 4.6, 4.5). The apical Other traces, however, connect to buds (bud traces), and
meristem is said to be indeterminate, that is, if conditions some traces end at the apical meristem without
were apex could grow continuously. We know,
ideal the connecting to either a leaf or a bud.
however, that this doesn't happen in nature either The role that leaves play in vascular differentiation and
because environmental factors are limiting or because the in the pattern of vascular bundles has been determined by
onset of flowering usually causes vegetative growth to elegant experiments. In one of these experiments, the
stop. In addition, each plant species is genetically leaves and leaf primordia were removed around the apex
programmed to develop within a certain size/age range. As the stem elongated the new leaf primordia
(Fig. 4.1 6/A).

Beneath the apical meristem are the regions of the three which formed were destroyed. Anatomical examination of
primary meristematic tissues —
protoderm, ground the stem which differentiated during the weeks of the
meristem, and procambium. The ground meristem starts to experiment revealed that the "vascular tissue" remained
differentiate first. These tissues form a cylinder near the as an unbroken cylinder (Figs. 4.1 68, Q. The cells making
outside of the stem and a core in the inside (Fig. 4.5Q, up the cylinder did not differentiate completely, and

The Dicotyledonous Stem

57
 collenchyma epidermis fibers

kTOSg;
f.y
i
'&&
phloem

-
•
_

"\ i
z& pith

//
Figure 4.15 Young alfalfa (Medicago sativa) stem showing
primary tissues Notice the vascular bundles that are composed of
xylem and phloem, the epidermis, and the cortex and pith
regions. The regions between vascular bundles consist of only
parenchyma cells, x 14.

vascular bundles did not form. If leaves were later (Lycopersicon), sunflower (Helianthus), and alfalfa

permitted to develop, the newly formed vascular tissue did (Medicago), also increase in diameter. This lateral
have bundles and did differentiate into xylem and phloem. thickening involves the activation of a secondary meristem,
This experiment demonstrated that leaves are needed for the vascular cambium (Fig. 4.17, 4.18). The vascular
vascular differentiation and for the formation of procambial cambium is composed of two parts the fascicular —
strands and vascular bundles. cambium, which forms from within the vascular bundles,
and the interfascicular cambium, which originates from
parenchyma cells that lie between vascular bundles (Figs.
Stem Secondary Growth 4.1 7A,D).
Recall that thefirst stage of development of vascular

Initiation tissues stems was the formation of a cylinder of residual

in

meristem. Next, the newly formed leaves acted on this

During primary growth, stems increase in length. Perennial
cylinder to induce portions of it (vascular bundles) to
stems and some annual stems such as those of tomato
become procambium strands and then vascular bundles of
xylem and phloem. After development of the primary xylem
and phloem, in some plants a portion of the procambium
remains undifferentiated. At some time during the plant's
life cycle, very early in the case of woody plants and later

in the case of herbaceous plants that develop secondary

growth, this residual procambium reinitiates divisions to

become the fascicular cambium (Figs. 4.17 A,B).
Simultaneously, parenchyma cells adjacent to the
fascicular cambium, but between vascular bundles, are
also stimulated to divide, to become the interfascicular
cambium. Together, both components form the vascular
cambium that will make secondary phloem to the outside
and secondary xylem to the inside of the stem (Fig. 4.19).

Organization

Close examination of a woody stem reveals that the cells

that make up wood are actually oriented in two ways
(Figs. 4.20A8). Some cells are elongated parallel with the
axis of the stem,making the axial system. The axial
system is composed of vessel members, tracheids, fibers,
Figure 4.16 Diagram of an experiment to show the effect of
and parenchyma. In the phloem, sieve-tube members,
removing leaves on vascular development in the stem of Geum
sp. A, top view of shoot tip, dotted lines indicate cuts to remove fibers, companion cells, and parenchyma make up the
young leaves and leaf primordia that are not yet visible. 6, "pith axial system. The source of these cells in the vascular
plug" that forms several days after removing leaf primordia. C,
cross section of pith plug to show the "provascular" tissue that
cambium are fusiform initials. A second system, the ray
forms in a complete circle or cylinder when leaves are removed. system, is composed of cells oriented at right angles to

chapt er 4 |
The Plant Bo dy

58
 -residual procambium
-pith ray

primary phloem
secondary phloem
cambium
secondary xylem
primary xylem

Figure 4.17 Formation of a complete cylinder of vascular cambium. A, at completion of primary

growth, some meristematic cells remain between primary xylem and primary phloem. (Residual or
"leftover" procambium shown in dark green.) Parenchyma cells appear in pith rays between vascular
bundles (light green). 6, the residual procambium becomes reactivated to form the fascicular
cambium; some of the parenchyma cells of the pith ray become meristematic to form the
interfascicular cambium (dark green). Together, the fascicular cambium and interfascicular cambium
= vascular cambium. C, parenchyma cells of pith between the meristematic cells of the bundles have
returned to a meristematic state, forming a cylinder of cambium (dark green). D, complete cylinder of
secondary xylem and secondary phloem (light gray) is formed by the vascular cambium. Parenchyma
cells are shown in light green, vascular cambium in dark green, and secondary tissues in light gray.

cortex

secondary xylem

Figure 4.18 Older alfalfa (Medicago saliva) stem showing secondary growth. The vascular cambium
is a continuous cylinder that forms progeny cells to the inside, the secondary xylem, and to the

outside the secondary phloem. The primary xylem, pith, cortex, and epidermis are also shown,
X25.
 the axis of the stem (Figs. A.2QA.B). The rays that make
up this system develop from ray initials in the vascular
cambium. Rays are living channels through which
nutrients and water move laterally in stems with secondary
cambium cellfc
growth. Xylem rays are made up of ray tracheids, and ray
parenchyma and phloem rays are made of phloem
parenchyma. Ray cells may remain alive for several years,
perhaps 10 or more.
Anyone who has examined the stump of a cut tree has
observed annual rings (Figs. 4.21, 4.22, 4.23). One
annual ring represents the amount of secondary xylem
growth for one season. Thick rings mean maximum
growth has occurred during a season, and thin rings mean
minimal growth. The density (width) of these rings give an
companion cell (cc) indication of past climate conditions.
Microscopic examination of an individualgrowth ring
shows that during the early part of the growing season,
cells are large and have relatively thin walls. This part of
the growth ring is called the early wood (springwood).
sieve-tube member (p
Later in the season the cells become smaller in diameter
and have thicker walls; this is the late wood (summer
wood) (Figs. 4.23A 4.25/\). In certain trees, large vessel
members form only in the early wood and only small
vessel members occur in the late wood; this organization
is called ring porous (Fig. 4.23). In other trees the
occurrence of vessel members is uniform throughout the
growing season; this is diffuse porous wood (Fig. 4.24).
Many trees in tropical areas, where growing conditions are
uniform throughout the year, show no annual rings in their
wood.
branch is several years old, the inner part
After a tree or
of the stem usually becomes inactive and filled with resins.
This dense, central core is called the heartwood. Good
barrels and wooden tubs are made from heartwood,
because the resin makes this wood impermeable to water.
Outside the heartwood is a light-colored, less dense
region of active wood called sapwood (Fig. 4.22).

Two things contribute to the formation of heartwood.

One is the curious formation of structures called tyloses.
Tyloses are ingrowths of the primary wall of parenchyma
cells that grow through the pits of vessel members. The

walls of tyloses eventually become enlarged and may form

secondary walls that completely plug up the xylem.
Species of trees that have abundant parenchyma cells
adjacent to vessel members readily form tyloses; however
other species with more scattered parenchyma do not
(Fig. 4.26). The second contributing factor to forming
heartwood is the activity of rays. Rays apparently play an
important role in transporting resins from the active
sapwood into the central heartwood. Heartwood is a
repository for metabolic by-products, serving a function
parallel to the excretory system of animals.

Figure 4.19 Diagram as seen in radial section showing stages in Gymnosperm wood is anatomically simpler than
differentiation of vascular cambium cambium;
cells (c, cc, angiosperm wood. It is composed almost entirely of
companion cell; p
1
p
2 phloem; x 1
x2 x3 xylem).
, , , , ,

tracheids in the spring wood and of fiber-tracheids (cells

that are intermediate between fibers and tracheids) in the
summer wood. The gymnosperms are usually only
rays in

one cell layer thick and axial parenchyma is rare. Vessels

are absent, but there are many resin ducts. These
secretory ducts are long hollow tubes containing an
internal layer of cells that produce resin (Fig. 4.27).

chapter 4 I The Plant Body

60
 wood

'Qy ni rials

ml
fusiform

Figure 4.20 Vascular cambium. A, diagram showing relationship

of cambium and cambial initial to stem; note also the cell
orientations. The cells ofthe axial system are elongated vertically,
those of the ray system are elongated horizontally; B, tangential
section; C. cross section of quiescent cambium of locust
(Robinia), x300.

wood rays

annual ring tangential section

Figure 4.21 Portion of stem of oak (Ouercus), showing cross, Figure 4.22 Cross section of a branch of mulberry (Morus),
radial, and tangential sections and their gross characteristics, x'/2 .

x'/2 .

The Dicotyledonous Stem

61
Figure 4.23 Sections of wood of oak (Ouercus borealis). A,
cross section; B, tangential section; C, radial section, x40.

Figure 4.24 Cross section of diffuse-porous wood from popular

(Populus deltoides), x 1 50.

Figure 4.25 Sections of wood of redwood (Sequoia

sempervirens). A, cross section; B. tangential section; C, radial
section; D, scanning electron micrograph of radial section, x 100.

chapter 4 |
The Plant Body

62
xylem vessel
member fiber

Figure 4.26 Radial section of wood from white oak (Quercus

alba) that shows tyloses. Tyloses are ingrowths of parenchyma
cells that form secondary cell walls and may plug up xylem vessel
members, x35.

-wood ray fracheids resin duct wood parenchyma Formation of Cork (Periderm)

Cork Cambium. Figure 4.28 shows the origin of cork

cambium and the development of secondary tissues from
it.Cork cambium in most plants arises from outer cortical
cells. The outer derivative cells from the cork cambium

generally differentiate into cork cells, thus forming a layer

of cork beneath the epidermis. The inner cells are known
as phelloderm, and they are parenchymalike Cork
cells.

cambium may originate also from epidermal or phloem

cells.

Cork tissue (Figs. 4.28, 4.29), is composed of flattened,

thin-walled cells with no, or small, intercellular spaces. A
fattysubstance called suberin is deposited in the walls,
— secretory cells -
rendering the cells almost impermeable to water and
globule of resin
gases. Hence, this tissue provides protection for the stem
Figure 4.27 Resin duct in pine wood (Pinus) as seen in cross
against excessive loss of water and also against
section.
mechanical injury. The protoplasts of cork cells are short-
lived.

The Dicotyledonous Stem

63
 cuticle
epidermis
epidermis

first cork cell

cork cambium cork

phelloderm

cork cambium
cortex
collenchyma

parenchyma

cuticle

epidermis

cork

phloem
H cork cambium
phelloderm eel Figure 4.28 A, diagram showing origin of the first cork cambium
and the layer of cork. New cork cambia and new layers of
first
cork form in a similar manner each spring. B, light micrograph
cortex
showing cork cambium and newly formed cork in an elderberry
stem (Sambucus); x 200.

Bark. In a young stem, the bark is made up of the

following tissues, in this order, from the outside to the
inside: cork,cork cambium, phelloderm (if present),
cortex, and phloem (Figs. 4.28, 4.29). Microscopic
examination may reveal the presence of epidermal cells
still clinging to the cork. In old stems, the epidermis,
cortex, and primary phloem become separated from the
adjacent inner tissues by successively deeper layers of
cork formation. As this happens, tissues outside newly
formed cork die for lack of water and nutrients. They dry
up and eventually wither away.

Lenticels

An impervious layer of cork would effectively cut off the

oxygen supply of the living tissues beneath groups of it if

parenchyma cells called lenticels did not develop in

various places (Figs. 4.29, 4.2). They frequently originate
beneath the epidermal stomata of young stems. When a
cork cambium is formed, produces ordinary parenchyma
it

cells below these stomata in an outward direction. The

resulting loose aggregation of parenchyma tissue bursts
through the epidermis to form the lenticel. The air spaces
between the parenchyma cells permit gaseous
interchange

\ aafvf
xylem phloem The Monocotyledonous
Figure 4.29 Cross section of a portion of a young stem of Stem
elderberry (Sambucus). From the outside to the inside of the
stem, the following tissues may be seen: cork, cork cambium,
cortex, secondary phloem, and xylem. The vascular cambium is Primary Growth
probably the two layers of thin-walled cells on the phloem side of
the xylem. Note the lenticel in the cork, x 100. The angiosperms can be conveniently divided into two
major groups dicotyledonous and monocotyledonous
plants (Chapter 16). The primary difference between these
two groups is that seeds of monocotyledonous plants, for

chapter 4 The Plant Body

64
Figure 4.30 Arborescent monocotyledons. A, Calitornia tan palm
(Washingtonia filifera) in a native stand; B, palms, and presumably
Pandanus, have long-lived phloem; C, the Joshua tree (Yucca
brevifolia) has secondary growth but produces typical closed
vascular bundles trom meristematic tissue resembling
procambium strands.

example, grasses, contain only one cotyledon (seed leaf) Monocot stems are usually uniform in thickness from the
while dicotyledonous seeds contain two. In addition, they top to the bottom of the plant, asopposed to being
have certain other characteristics in their plant bodies that tapered like most dicot stems (Fig. 4.30).
are sometimes different. The apical meristem of a typical monocot looks like a

The Monocotyledonous Stem

65
 / primary •
thickening
r
meristem I

••••-f-
, procambium

shoot apex primary thickening

meristem
bases of leaves primary thickening meristem (Fig. 4.30/4). Further down
procambium the stem, parenchyma cells continue to divide and
enlarge, allowing for continued lateral stem enlargement.
This process is called diffuse secondary growth since it

does not involve an actual lateral meristem. In some

palms, the stem looks much thicker at the base than at
the top of the stem, but this thickening is caused partly by
the overlapping leaf bases of old leaves, and partly by the
presence, in some palms, of great masses of adventitious
aerial roots.

Other monocots like Agave, Cordyline, Sansevieria, and

Dracaena have true secondary growth that involves a
secondary meristem (Fig. 4.33). This secondary meristem
forms below the primary thickening meristem and extends
to the base of the plant. divides and forms only
It

parenchyma cells to the outside (secondary cortex). To

the inside, forms parenchyma cells and secondary
it
Figure 4.31 A, longitudinal section of Iris shoot apex to show
the primary thickening meristem, x88. 6, longitudinal diagram of vascular bundles (Fig. 4.33/4). The secondary vascular
the shoot apex of corn (Zea mays). bundles consist of a ring of xylem surrounding the
phloem, whereas the primary bundles consist of xylem
and phloem side by side.
small dome in a broad depression. Young leaves grow out
Secondary growth occurs in the roots of only one genus
of the depression and extend over the apical meristem.
of monocot Dracaena. A barklike layer is present in
Beneath it is the inverted saucer-shaped primary
some monocots. is derived from cortical parenchyma
It

thickening meristem (Figs. 4.31 A,B). This meristem may

cells that divide to form files of suberin-filled parenchyma
extend down the stem a short way, so that it forms new
cells.
cells upward to allow for an increase in length and
outward to allow for an increase in girth. Leaf traces
traverse the primary thickening meristem to connect
already existing bundles.
it to
Stem Modifications
The vascular bundles in monocots are usually scattered
in a random pattern throughout the ground tissue (Fig. Stems may become adapted for functions other than
4.32/A), but some monocot stems have a hollow core and support, conduction, and production of new growth. They
have bundles arranged in a ring similar to some dicot may, for instance, become attachment organs for vines;
stems (Fig. 4.32C). The terms pith and cortex are often they may carry on photosynthesis, store food or water,
not applicable in monocot stems. The tangled vascular and develop protecting devices.
bundle pattern in the nodes of monocot stems is complex
and is called a nodal plate (Fig 4.328).
Rhizomes
Secondary Growth A rhizome is a horizontal, underground stem. In most Iris

species, leaves and flowering stalks are produced at the

Most monocot stems lack secondary growth. Palm trees growing rhizome tip (Fig. 4.1 S). The leaves die a relatively
increase their thickness in the apex by the activity of the short distance back from the growing tip, so that many iris

chapter 4 |
The Plant Body

66
Figure 4.32 Internal anatomy of a monocotyledonous stem. A,
cross section of corn stem (Zea mays) showing the scattered
distribution of vascular bundles x 30. 6, longitudinal section
through a node in corn stem x40. C, cross section of wheat
stem (Triticum aestivum), note that in this plant the core of the
stem is hollow, x30.

'
-cambium

Figure 4.33 Cross section of a portion of monocotyledonous

stem, Dracaena sp., that has true secondary growth. Note that
the cambium forms discrete secondary vascular bundles and
parenchyma cells to the inside of the stem, and parenchyma cells
only toward the outside, x 13.

Stem Modifications

67
plants bear senescent leaves. Roots are also formed at
nodes, and they may remain for the life of the rhizome.
Other rhizomes, like Carina (Fig. 4.37A), produce upright
leafy stems with terminal flowers at every third node. The
intervening nodes are marked by only small sheath leaves

Corms
A shortened vertical, thickened underground stem is a
corm (Fig. 4.37). In Gladiolus, it consists of a short stem
with much stored food. Nodes are, as usual, indicated by
leaves; some bases are shown in Fig. 4.37B. Small buds
occur in axils of some of these leaves. In a median section
of a corm (Fig. 4.37Q one can distinguish between stored
food and the central portion containing a single bud that
will produce a single leafy, flowering shoot. Food stored in
a dormant corm is used in the production of the leafy
shoot. New corms will develop from axillary buds. In
addition, short underground stems may form, each giving
rise, at its tip, to a single small corm.

Bulbs
A corm in that food is stored in leafy
bulb differs from a
scales. The stem portion is small and has at least one
central terminal bud that will produce a single upright leafy
stem. In addition, there is at least one axillary bud that will
produce a bulb for the subsequent year. In the longitudinal
section of the sprouting daffodil bulb shown in Fig. 4.37D,
the stem is producing three leafy stalks, one of which is

forming a new bulb.

Food stored in the leafy scales of a bulb is used up by

the initial growth of a leafy shoot. Food to be stored for a

new bulb is supplied from a leafy shoot. The table onion is

a good example of a commercially valuable bulb.

Figure 4.34 Types of stem modifications, A and 6, stem tendrils;
C, F, and G, thorns; D and E, leaflike stems

Tubers
Tubers are enlarged terminal portions of slender rhizomes
(Figs. 4.36, 4.37E). The
(Solanum tuberosum), is a
potato,
good example. The potato plant possesses three types of are of two morphological sorts: leaf and stem. In the
stems: (a) ordinary aerial stems, (b) slender underground trumpet flower (Bignonia), for example, several uppermost
rhizomes, and (c) their enlarged tips, tubers. In the mature pairs of leaflets have no blades, but instead form very
potato the scar left where the tuber was broken from the slender leaf tendrils. Obviously, these are leaf tendrils. In
rhizome is clearly visible. On the potato tuber there are grapes (Vitis) and Virginia creepers (Parthenocissus
nodes and internodes, lateral buds, and a terminal bud. quinquefolia), tendrils are modified stems (Figs. 4.34A.B),
Buds develop into stems (Fig. 4.37E). The "eyes" of the as evidenced by their presence nodes in leaf axils.
at
tuber are groups of buds; each group along the sides In the Virginia creeper, each tendril ends in a knob that
represents a lateral branch with undeveloped internodes. flattens out when it comes in contact with a surface to
At the unattached "seed end" of the tuber, the "eye" is in which it adheres.
reality a terminal branch on which only one bud is strictly

terminal. In an elongated potato it is possible to make out

the spiral arrangement of the eyes, because there is only Cladodes
one eye at a node.
These are stems and
that are leaflike in form, are green,
perform the functions of leaves. They may bear flowers,
Stem Tendrils fruit, and temporary leaves. Examples of plants with

cladodes are Ruscus (Fig. 4.34D), Asparagus (Fig. 4.34E),

Tendrils are slender, coiling structures that are sensitive to Smilax, various species of cacti, and some orchids (e.g.,
contact stimuli and attach the plant to a support. Tendrils Epidendrum).

chapter 4 |
The Plant Body

68
Spines and Thorns Summary of Secondary Growth
Most spines and thorns ot plants are moditied stems or 1. The predominantly short life span of 3 to 5 years for

outgrowths of stems. Spines, however, occur in certain

cells, in general, limits the size and longevity of
plants,such as barberry (Berberis) and black locust individual plants having only primary growth. The
(Robinia pseudoacacia), and in a few cases even roots mechanism of secondary growth overcomes this

become limitation.
modified as spines. Good examples of stem
thorns are those of fire-thorn (Pyracantha) (Fig. 4.34F) 2. If all meristematic cells in a procambium strand
and honey locust They are borne
(Gleditsia) (Fig. 4.34G). become differentiated into primary vascular tissue, the

in the axils of leaves as ordinary branches are. Sometimes vascular strand is closed to further growth.
thorns bear leaves (Fig. 4.34F), which is further evidence 3. If there remains an active meristematic region between

that they are stems. Prickles, on the other hand, such as primary xylem and primary phloem, continued growth

those on rose stems, are merely epidermal outgrowths, is possible. These meristematic cells become the

somewhat like hairs. fascicular cambium.

4. An interfascicular cambium forms from ray
parenchyma between vascular strands.
cells

Stolons
_^_ 5. Union of fascicular and interfascicular cambium
produces a complete cylinder of vascular cambium.
Bermuda grass (Cynodon dactylon) has above-ground 6. Divisions in cambium are longitudinal, so that the stem
horizontal stems called stolons. These stems creep along now increases only in girth.

the ground, and at each node, shoots and roots arise. In 7. Cork cambium is short-lived; new cork cambiums may
strawberry (Fragaria) (Fig. 4.35) stolons, roots, and leaves arise each year, producing new layers of cork.
arise at every other node. 8. Cork cambium may originate in successively deeper
tissues from epidermis, cortex, and phloem.
9. The production of secondary vascular tissue thus
makes possible the attainment of great size and great
age by individual trees, even though the functioning
cells may be no older than those in a perennial
herbaceous plant such as an Iris.
10. Most monocot stems have only primary growth that
arises from a special primary thickening meristem.
This meristem causes increase in both height and
girth.

11. Palms supplement their lateral growth by the division

parenchyma cells throughout the stem.
of

12. Other monocots, such as Agave and Sansevieria, have

true secondary growth, with a type of cambium that
produces secondary vascular bundles and
parenchyma cells.
Figure 4.35 Runner of strawberry (Fragaria) x '/8 roots and
13. In woody and herbaceous stems, vascular tissues are
;

shoots are produced at every other node.

arranged in bundles, generally forming a circle. In

monocotyledonous plants, bundles are irregularly

distributed throughout the stem.
14. Rhizomes, bulbs, corms, stolons, and tubers are all

modified stems.

PART TWO
Leaves
Green plants and a few species of bacteria are the only
producers that use sunlight as an energy source; all other
inhabitants of the earth consume what the green plants
produce. In seed plants, leaves are the principal organs of
production. Chloroplasts within leaf cells are the sites that
trap light energy and use photosynthesis to convert it into
young tuber
chemical energy.
The water economy of plants calls for the absorption of
much more water than can be metabolized. This excess
potato seedling water is returned to the atmosphere by leaves. A second
Figure 4.36 A potato seedling showing development of young
role of leaves is transpiration. The structure of leaves is

tubers at the end of slender rhizomes. uniquely adapted to carry out these two primary roles.

Leaves

69
Figure 4.37 Plants illustrating continued development with only Leaves are of many shapes and sizes (Fig. 4.38).
primary growth. A, roots and shoots are produced at every third
Practically all leaves have veins for support and
node by the rhizome of Canna, x >/2 B, corm of Gladiolus, x 1
;

C, corm sectioned to show short stem with storage tissue of conduction, and a chlorenchyma tissue containing
current corm and disintegration of corm of preceding year, x 1 chloroplasts. Shape and size are such constant traits for
D, longitudinal section of young daffodil (Narcissus) plant,
showing two bulbs, one with two shoots, united by the short
many categories of plants that one may frequently identify
stem, x 1 E, potato tuber (Solanum), note spiral arrangement of
;
an unknown plant simply by its particular type of leaf. Leaf
the eyes and the short sprout, x 1 shape, however, is good diagnostic trait for
not always a
use in because may vary within a
plant identification, it

species, and because leaf shape may sometimes be

altered by its environment.
Leaf blades provide large surfaces for the absorption of
light energy and carbon dioxide, both of which are
required for photosynthesis. They are thin and hence no

chapter 4 The Plant Body

70
 stipules

stipules

Figure 4.38 Different kinds of leaves. A, poplar (Populus

deltoides); B, castor bean (Ricinus communis); C, oak (Ouercus
lobata); D, rose (Rosa odorata); E, Virginia creeper
(Parthenocissus quinquefolia); F, faba bean (Vicia faba), x '/2 .

cells lie from the surface. This form of the leaf

far The Blade
facilitates the absorption of light energy and the exchange
of carbon dioxide, oxygen, and water vapor between the Variable external features of the leaf blade are its overall
intercellular spaces of the leaf and the atmosphere. The shape: apex, margin, and base. This range of variation is

water that leaves utilizeand transpire comes from roots great, and many terms are employed by taxonomists to
and enters the leaf in the xylem tissue of veins. Food describe leaf shape accurately. Three of these terms seem
manufactured in leaves is transported out of the leaf in the important enough to us to mention here: leaf margin
phloem tissue of the same veins. In addition, veins may entire (smooth), dentate (toothed), or lobed (Figs.
add support to the leaf. 4.38F,AC).

Leaf Components The Petiole

Some leaves, such as those of peas (Pisum), beans

The leaves of dicotyledons are generally different from (Phaseolus), roses (Rosa), tulip trees (Liriodendron), and
those of monocotyledons. A typical foliage leaf of a plant many others, have two small, leaf like outgrowths at the
belonging to the dicotyledons is composed of two principal base of the petiole, known as stipules (Figs. 4.38D,F).
parts: (a)blade and (b) petiole (Fig. 4.38A). The blade is The petiole itself may be long or short, rounded or
thin and expanded, the petiole slender. The thin blade is occasionally flat. It is usually attached to the base of the
supported by a distinct network of veins that are leaf blade, but in some plants, such as Tropaeolum and
composed of vascular tissues and fibers. In addition to castor bean (Ricinus), it is attached at the middle of the
forming a supporting framework for softer tissues of the blade on the underside. This leaf is called a peltate leaf
blade, veins carry water, mineral salts, and food to and (Fig. The petiole is sometimes absent, as in Zinnia,
4.386).
from the leaf. the blade being mounted directly on the stem. Such a leaf

Leaf Components

71
 '^m ^^u
n
yi

— \ •

c>^LM
/
'

{ i
©^
r A S[ Ir

-i
r^V?
>A
j^wj^L/^
W*\-'
V^B \ L\— -

tt
\~~i -\^ ^M
'

*v i

Figure 4.39 Leaves, showing parallel and netted venation. A,

Canna about x '/5 B, portion of similar leaf, about x 10; C.
leaf, ;

maple (Acer) leaf, about x '/2 D, portion of similar leaf, about

;

x20.

is said to be sessile (Fig. 4.50D, juvenile Eucalyptus). union with the sheath, is carried around the stem in two
Monocotyledonous leaves such as grasses do not have earlike points, the auricles. Ligule and auricles may both
distinct petioles. Instead, the leaf is divided into two parts, be present, or one or the other may be absent.
sheath and blade. The blade is the typical thin, expanded
portion. The sheath is green, perhaps nearly as large as
the blade, and it completely sheaths the stem (Fig. 4.40/4).
Simple and Compound Leaves
In many species such as corn the sheath extends over at As to configuration of the blade, there are two kinds of
leastone complete internode. If the region of union leaves: (a) simple and compound. In a simple leaf
(b) the
between the blade and the sheath is examined carefully, a blade is all in one unit (Figs. 4.38C0 4.39AQ. In a
small flap of delicate tissue extending upward from the compound leaf the blade is composed of a number of
sheath may be seen, closely enveloping the stem. This is separate leaflike parts, the leaflets (Figs. 4.38D.F).
called the ligule (Fig. 4.400). It may, in some cases, serve Inasmuch as leaflets have the characteristics of a simple
to keep water and dirt from sifting down between stem and leaf, may sometimes be doubtful whether the structure
it is

sheath. In many species, of which barley (Hordeum) (Fig. a simple leaf or a leaflet, especially if the leaflets are large
4.400) is a good example, the base of the blade, at its or very numerous. A primary distinction is that buds occur

chapter 4 The Plant Body

72
 node

stem

blade

sheath auricles

Figure 4.40 Leaves of members of the grass family. A,

crabgrass (Digitaria sanguinatis), showing sheath and blade; S,
ligule and auricles of barley (Hordeum vulgare), about x y4 .

in the axils of leaves, but not in the axils of leaflets. from mechanical injury. The mesophyll is composed of
When the leaflets of a compound leaf arise from the parenchyma cells, most or
which contain all of
rachis (continuation of the petiole), as do the pinnae of a chlorophyll, and thus are able to carry on photosynthesis.
feather, the leaf is said to be pinnately compound, like the Veins possess xylem and phloem elements, which conduct
rose and bean (Fig. 4.38D.F); when the leaflets diverge water, inorganic salts, and foods. Fibers and collenchyma
from a common point at the tip of the petiole, the leaf is may be associated with conducting elements of midrib and
said to be palmately compound, like the horse chestnut larger lateral veins.
(Aesculus) and Virginia creeper (Fig. 4.39E). Compound The petiole has its own specialized structures that
leaves may be once, twice, or thrice compound. enable it to support the leaf blade, conduct food, water,
and inorganic and disconnect itself from the stem
salts, at

the close of the growing season without exposing living

Venation stem tissue to drying out or to infection.

The arrangement of veins of a leaf is called venation.

There are two principal types of venation: (a) parallel Epidermis
venation, usually characteristics of monocotyledonous
leaves (Figs. 4.39A6), and (b) netted venation, usually The epidermis covers the entire leaf surface and is

characteristic of dicotyledonous leaves (Figs. 4.39C.D). I

continuous with the surface of the stem to which the leaf

n etted venatio n there are one or more prominent veins is attached. In most leaves the epidermis is a single layer

from which smaller veins branch off to join with other of cells. It may consist of several kinds of cells: (a)

small veins, thus forming a conspicuous net. I n parallel^ ordinary epidermal cells, (b) guard cells, (c) hair cells
one or a few large veins (Fig. 4.44). Ordinary epidermal cells show a variety of
veined leaves, there usually is

with many veins branching from them. These veins run shapes, depending on the species. Epidermal cells are

parallel to each other and do not branch further. It is generally covered on their outer surfaces by a waxy
important to note that there are some dicotyledonous cuticle secreted by their protoplasts. The deposition of the
plants 'with parallel veined leaves and some cuticle frequently forms a pattern, quite specific for the

leaves on which it is found (Fig. 4.42).

monocotyledonous leaves with netted veins.
A guard cell is a special type of epidermal cell. Guard
cells occur in pairs, separated by an opening or pore (Fig.

Two guard cells plus a pore are called a stoma

Anatomy of the Foliage Leaf 4.43).
(plural = stomata). The size of stomata varies
considerably according to the species of plants and even
The anatomy of a leaf blade is best shown in section. In in any one plant. Here are some representative
Fig. 4.41 we observe three principal tissues: (a) measurements in microns (length x breadth): bean, 7 x
epidermis, (b) mesophyll (middle of leaf), and (c) veins 3; geranium (Pelargonium), 19 x 12; corn (Zea), 19 x

or vascular bundles. The epidermis usually consists of a 5. The number of stomata per unit area varies widely

single layer of cells that covers the entire leaf surface. It depending on the species of plant and the environmental
protects the tissues within the leaf from drying out and conditions under which it is growing. Usually more

Anatomy of the Foliage Leaf

73
 stoma with two guard cells upper epidermis

inter-
palisade parenchyma cellular
spaces

border parenchyma
guard cells

spongy parenchyma | ower gpiderm

Figure 4.41 Three-dimensional diagram of a section of a foliage

leaf, x20.

Epidermis

Figure 4.42 Cuticle on the epidermis of mala mujer

(Cnidoscolus) as seen with the scanning electron microscope. A,
upper surface, showing cuticular ridges, x 650. B, lower surface
at higher magnification, showing cuticular ridges between
stomata, X1100.

chapter 4 |
The Plant Body

74
ordinary epidermal ce

Figure 4.43 Views of stomata. A. diagram; 6, corn (Zea mays),

about x 1000; C, rice (Oryza sativa), x900.

thickened cell wall

stomata are found on the lower than on the upper surface. Table 4.1
Table 4.1 gives the average number of stomata per square Average Number of Stomata per Square Centimeter
centimeter on the upper and lower surfaces of some
common plants. Upper Lower
Stomata are widespread in the plant kingdom. With the Plant Epidermis Epidermis
exception of a few submerged aquatic plants, stomata are
present in all angiosperms and gymnosperms. Functional Alfalfa (Medicago) 16,900 13,800
stomata have been found in liverworts, mosses, horsetails, Apple (Malus) 29,400
club mosses, ferns, and cycads. In the angiosperms, they Bean (Phaseolus) 4,000 28,100
can occur on stems, petals, stamens, and pistils as well as Cabbage (Brassica) 14,100 22,600
on leaves. The major group of green plants that lack Corn (Zea) 5,200 6,800
stomata is the algae. Stomata are structurally similar in the English Oak (Quercus) 45,000
many different groups in which they are found. Nasturtium (Tropaeolum) 13,000
A cross-sectional view of the guard cells shows that the Oat (Avena) 2,500 2,300
walls are typically unevenly thickened, with the thicker, Potato (Solanum) 5,100 16,100
less elastic walls, adjacent to the pore. The stomata are Tomato (Lycopersicori) 1,200 13,000
the only openings in the leaf epidermis, and it is chiefly
through them that gases pass into or out of the leaf.

Although the cuticle is nearly impermeable to gases, small

amounts of gases pass directly through the outer wall and cells, including guard cells, has a cuticle (Fig. 4.43A) like
the cuticle of epidermal cells. The role of stomata, that of the epidermis of stems. It is effective in limiting
including the control of their movement in transpiration, is both the inward or outward movements of water vapor and
discussed Chapter 5.
in other gases. The cuticle is usually thicker on the
Guard cells occur in pairs; each is crescent-shaped or upperside of the leaf than on the underside.
semicircular in form, as seen in a surface view (Fig. Several different types of hairs grow out from the
4.43C). Chloroplasts occur in guard cells but are lacking epidermis of leaves and resemble hairs from the epidermis
in ordinary epidermal Both kinds of epidermal cells
cells. of stems. They may be unicellular or multicellular, simple
have long-lived protoplasts. The outer wall of epidermal or branched, scalelike or glandular. The unicellular hair,

Anatomy of the Foliage Leaf

75
Figure 4.44 Trichomes: A, Aleuntes leaf — branched; B, Croton
—
leaf simple, about x350; C, Phaseolus — gland x565.

the simplest kind, may be branched or unbranched. Intercellular spaces are much larger in spongy
Multicellular hairs may consist of a single row of cells, but parenchyma than in palisade parenchyma. Since the air
may also be branched 4.44AB). At the upper end
(Figs. spaces between mesophyll cells are interconnecting, many
glandular hairs bear a single large cell or a group of cells; cells are in contact withan intercellular space. Thus, most
some of these cells excrete ethereal oils, which often food-making have free access to carbon dioxide and
cells
impart a stickiness to leaves (Fig. 4.44C). oxygen. To facilitate gas exchange, large air spaces,
called substomatal chambers, are generally present
beneath each stoma (Fig. 4.43/4).
Mesophyll The leaves of some plants, especially those that grow in

very dry habitats, may have modified leaf structure. For

Mesophyll is the photosynthetic tissue between the upper example, their mesophyll cells are tightly packed together
and lower epidermis. It is parenchyma tissue, and is
and lack the differentiation between a palisade and spongy
traversed by veins. Chloroplasts are present in the layer. Such plants may also have a thick cuticle and
mesophyll cells, which, in some species, may be sunken stomata. All of these modifications are believed to
differentiated into two distinct layers: palisade reduce transpiration.
parenchyma and spongy parenchyma (Figs. 4.41, 4.45).
The palisade parenchyma is just below the upper
epidermis and usually consists of from one to several Vascular System
layers of narrow cells with their long axes at right angles
The spongy parenchyma extends from
to the leaf surface. The veins, or vascular bundles, form a network that
the palisadeparenchyma to the lower epidermis. Cells of extends throughout the leaf. The conducting elements are

the spongy parenchyma are irregular in shape and loosely xylem and phloem. Veins conduct water, mineral salts, and
arranged. foods, and also mechanically support the mesophyll

chapter 4 |
The Plant Body

76
 spongy parenchyma

lower epidermis

Figure 4.45 Photomicrograph of a cross section of a midrib of a

lilac(Syringa vulgaris) leaf, x 150.

tissue. In addition to the midrib and larger lateral veins that are characterized by the presence of an enlarged bundle
are visible to the naked eye, innumerable minute branch sheath (Fig. 4.466), and lack mesophyll differentiation into
veins can be seen with the aid of a microscope. The large palisade and spongy types.
veins contain vessels, tracheids, sieve tubes, companion
cells, and also some mechanical tissue (Fig. 4.45). Such
The Petiole
veins, in some leaves, may have both primary and
secondary vascular elements. In the petiole, phloem and xylem maintain their relative
In have both xylem and phloem
larger veins that positions; as in the stem the phloem is on the underside of
elements, the xylem is toward the upper surface of the the petiole (and leaf blade) and xylem ison the upperside.
leaf, and the phloem is toward the lower surface (Fig. One or more vascular bundles are embedded in

4.45).* Smaller veins have few vascular elements and few parenchyma. Fibers may be associated with vascular
or no mechanical elements. The very end of a vein is tissues of bundle^ and, not infrequently, groups of
usually a single tracheid with a spiral wall (Fig. 4.46/4). collenchyma cells occur beneath the epidermis.
Veinlets are usually surrounded by one or more layers of Collenchyma cells are also commonly present in leaves.
parenchyma cells, the bundle sheath, which may or may Usually they form part of the epidermis, but may be
not possess chloroplasts. Water and solutes must pass formed in the mesophyll near the leaf surface adjacent to
from conducting elements of the veinlet through the the vascular bundles.
bundle sheath in order to reach the mesophyll. The
smallest veinlet has an unbroken connection with vascular
Leaf Development
elements and stem to which the leaf
of the midrib, petiole,
is connected. Plants such as corn and other tropical

grasses use a photosynthetic pathway called C 4 Leaf development is intimately associated with the

photosynthesis (Chapter 6). Leaves of these plants usually differentiation of the young shoot apex.

Leaf Development

77
 The leaf is initiated on the flanks of the apex, as a slight
vein ending tracheid bulge resulting from the enlargement and division of
several cells in the outer layers of the apical meristem
(Fig. 4.47). This bulge continues to enlarge until it forms a
thin, elongated leaf primordium (Fig. 4.48/A). The flanks
of the primordium (the marginal meristem) will initiate the
formation of the leaf blade (Figs. 4.48S,Q. This region of
border parenchyma growth remains active until the leaf has fully formed. Most
cell divisions are completed early in leaf development; cell

enlargement accounts for most of the increase in leaf size.

spongy parenchyma
During the early stages of leaf formation, the cells
mmediately beneath each primordium also become
meristematic. These cells, the procambium, will form the
vascular tissue that connects the leaves to the stem's
vascular system (Fig. 4.47).
One problem that has continually intrigued plant
biologists is how the plant regulates the site of new leaf

initiation. The leaves of vascular plants are distributed on

the stem in four distinct patterns (phyllotaxy) — opposite,
alternate, whorled, and spiral (Fig. 4.49). The position of
a new leaf primordium is closely correlated with the
position of the next older primordium. This has been
proven by the experimental removal or isolation of the
youngest visible primordium on a shoot apex, causing the

epidermis

undle sheath

chloroplasts

xylem

Figure 4.46 Diagram of leaf vascular bundles. A, nofe the

parenchyma and the presence of only
tissue bordering the vein
annular wall thickenings in the tracheids at the vein tip. B, cross
section of a leaf with C 4 photosynthesis, note the prominent
bundle sheath cells.

chapter 4 |
The Plant Body

78
 n

leaf primordiom
Q

procambium

&
Figure 4.48 Growth of a young leaf. A, pencil stage, formation
of marginal meristems; B, activity of marginal
meristems initiates
expansion of a young leaf; C, continued lateral growth
lateral
through the activity of marginal meristems.

next formed primordium to develop closer to it. It is also

known, at least in ferns, that if the procambium beneath a
suspected leaf primordium is severed, the primordium will
not develop.

Leaf Shape
Figure 4.47 Diagram of a shoot apex showing the direct Internal Control
relationship between the initiation of a leaf and procambium
strand. The second leaf shows a definite primordium and the third
leaf is well-formed. The procambium strands associated with Leaf shape is under direct genetic control. Several single
these leaves are shown. gene mutations of tomato (Lycopersicon esculentum)
result in striking changes in leaf shape (Figs. 4.50A.B).

B opposite

Figure 4.49 Leaf arrangement on stem axis; A, alternate; 6,

opposite; C, whorled; D, spiral.

Leaf Shape

79
Leaves are also affected by the availability of internal different in shape from same
aerial leaves of the plants
growth regulators. When treated with the hormone (Fig. 4.50E). Similar changes may be induced in the aerial
gibberellic acid, thistle plants (Centaurea), for example, portion of the shoot by reducing the C0 2 content of the air

formed abnormal, entire leaves instead of normal, lobed and lowering the temperature.
leaves. Nitrogen starvation may also induce the plant to acquire
Leaf shape may also vary considerably with the leaf traits similar to those of sun leaves; water stress may
physiological age of plants. Thus, some plants have change the plant in a similar way. Thus, we see that
distinctive juvenile leaves for the first few years of growth. environmental stress is an important, but complex, factor.
Subsequent adult leaves of a different form are
characteristic of the older or adult plant (Figs. 4.50C,D).
Leaf Modifications

Environmental Factors Leaf shape may be considerably modified to perform

functions other than photosynthesis (Fig. 4.51).
Leaf shape is also influenced by environmental factors Bud scales are short, thick, sessile, often covered with
such as light, moisture, and temperature. The total dense hairs on the outer surface, and sometimes waxy or
intensity, wavelength, and daily duration of light have resinous. When present, they protect the delicate
separate but interrelated effects. Plants growing in intense meristematic tissue of the shoot tip and the rudimentary
sunlight (sun leaves) usually have thick leaves with a leaves from drying out.
thick palisade tissue and a dense, spongy parenchyma. The spines of various species of cacti and those of
Their intercellular spaces are small, and the epidermis is Fouquieria represent entire transformed leaves. In the
heavily cutinizedand generally glossy, with the stomata black locust (Fig. 4.51 A), the spines are stipules, rather
confined to the lower epidermis. These plants may also than leaves.
have woolly epidermal hairs. Leaves of the same species In some species of Lathyrus, the tendrils are
growing in shade (shade leaves) have contrasting traits. transformed leaflets; in other species, the whole leaf is

They are thin, possessing a single palisade layer and a transformed into a single tendril, and leaflike_stjpules

spongy parenchyma with many intercellular spaces. The perform the normal f unctions of th e leaves. In Bignonia
epidermis has a thin cuticle that is usually dull, and capreoiaia (trufnpeffiower), the thircfleafTet is transformed
stomata may be present on both upper and lower into a tendril (Fig. 4.516). Both leaf tendrils and stem
epidermal surfaces. tendrils serve to attach the plant to a support.
Light is required for the normal development of leaves, Leaves are sometimes modified as food or water storage
including the differentiation of chloroplasts to a state organs. The thick, fleshy bases of leaves that comprise
where they are capable of carrying on photosynthesis. In much of the daffodil (Narcissus) bulb (Fig. 4.37D)
darkness, the leaves of a typical dicot remain small and accumulate large quantities of food. Succulents found in

pale yellow while the stems grow long and slender. This deserts and saline soils have thick, fleshy leaves with
condition is called etiolation. Regular daily illumination special water-storage tissue.
that is intense enough to support photosynthesis is A striking adaptation of leaves to a special function
required for the continued healthy existence of leaves. occurs in insectivorous plants. In these plants, the leaves
Excessive shading usually results in the death of a leaf. have taken on forms and various structural features that
Submerged leaves of semi-aquatic plants may be vastly enable them to capture insects and obtain food from their

chapter 4 The Plant Body

80
 bodies. Well-known insectivorous plants are sundew
(Drosera, Fig. 4.51 E), pitcher plants (Darlingtonia, Fig.
4.51 C), and the Venus' fly trap (Dionaea muscipula, Fig.
4.51 D).
Leaves sometimes function effectively in vegetative
reproduction. In certain species of Bryophyllum (Fig.
4.51 F), patches of tissue located in notches along the leaf
margins remain meristematic. This tissue will eventually
develop small, new plants while the parent leaf is still
active. The little plants later drop from the leaf to the
ground, where under favorable conditions they may
develop into new individuals. Leaves of Begonia produce
plantlets by the differentiation of small clusters of
epidermal cells on the upper leaf surface.

Leaf Abscission

The separation of plant parts from the parent plant is a

normal, continually occurring phenomenon. Leaves fall,
fruits drop, flower parts wither and fall away, and even
branch tips or whole branches may be separated from the
parent plant as a normal part of its life. The fall of leaves
from woody dicotyledons in the autumn is the most
common example of this phenomenon. In practically all

cases, separation or abscission is the result of

differentiation in a specialized region known as the
Figure 4.50 Examples of induced leaf shape changes. A, B, abscission zone at, or close to, the base of the petiole
different leaf shapes induced in leaves of tomato (Lycopersicon
(Fig. 4.52). Parenchyma cells comprising the abscission
esculentum) by single gene mutations C, D, adult and juvenile
leaves of Eucalyptus sp., x '/4 and x '/2 E, examples of
.
layer may be smaller and lack lignin, compared to the
submerged and air forms of leaves of three aquatic plants. cells of adjacent tissues. Even vascular elements may be
shorter, and fibers may be absent from the bundle in the
abscission zone. These anatomical features definitely
make this zone an area of weakness.
Previous to the fall of leaves changes may normally
occur in this zone. Cell divisions, though apparently not
necessary, frequently take place, and produce a layer of
brick-shaped cells across the petiole. Actual separation of
the leaf may be brought about in several ways. In some

Leaf Abscission

81
 Figure 4.51 Examples of leaf modificafions. A, Robinia stipular
spines; 6, Bignonia tendrils; C, pitcher plant leaf (Darlingtonia
sp.); D, Venus fly trap (Dionaea sp.); E, sun dew (Drosera sp.); F,
foliar plantlets in Kalanchoe sp.

chapter 4 |
The Plant Body

82
 plantlets
A

Figure 4.52 The formation of the abscission zone

Leaf Abscission

83
species, the middle lamella is dissolved away and cells in both parenchyma tissues. Mesophyll is adapted for
separate. In other plants, walls and cells are dissolved. In photosynthesis. Some plants do not differentiate
a third small group ot plants, a layer ot cork torms across mesophyll into spongy and palisade regions.
the petiole so that the leat simply withers in place and is 5. Leaf shape under genetic and hormonal control, and
is

blown away. In all cases, a protective, corky layer ot cells is also influenced by light, moisture, and the
develops across the leat scar. It is continuous with the physiological age of the plant. Leaves may become
stem cork. Furthermore, the vessels are likely to become modified, serving as bud scales, spines, tendrils, the
plugged with tyloses or gums. These devices prevent the source of propagules, food or water storage organs,
tall ot a leaf from leaving an open wound or a point of and insect Such environmental factors as strong
traps.

entrance for organisms that might cause disease. and water stress can
light intensity, nutrient deficiency,

The abscission zone has two functions: (a) to bring affect leaf morphology and anatomy.

about the fall of the leaf or other plant part and (b) to 6. Leaf primordia develop in a definite spatial sequence on
protect the region of the stem from which the leaf has the flanks of the shoot apex.
fallen against insect damage or rot caused by bacteria or 7. The young leaf lamina is produced by marginal
fungi. meristems that are also responsible for the shape of the

Environmental cues are important in synchronizing leaf.

abscission with the seasons. These cues are most 8. Cell divisions are completed while the leaves are
commonly cold temperature or short days that in turn enclosed in the bud; leaf expansion results mainly from
induce hormonal changes that then affect the formation of cell enlargement.
the abscission zone (Chapter 8). 9. The formation of a definite abscission zone across a
petiole or fruit stem is responsible for leaf fall or fruit

drop. Environmental cues and hormonal changes affect

Summary — Leaves formation of the zone.

1. The chief functions of leaves are photosynthesis and

2.
transpiration.
Angiosperm leaves are generally supplied with flat, thin
PART THREE
blades that are attached to the stem by petioles or
sheaths. Veins strengthen the blades and transport food
Roots
and water. Leaf blades may be simple or compound;
leaf margins may be entire, dentate, or lobed.
3. A cross section through the blade of a typical leaf

shows the following tissues: upper epidermis, Functions of the Root

mesophyll, differentiated into palisade parenchyma and
spongy parenchyma, and lower epidermis. Generally, a
waxy cuticle coats the epidermis. Guard cells of the The functions of the root system are anchorage, storage,
epidermis form stomata that control gas exchange. conduction, and absorption. You need only walk through
4. Mesophyll tissue is comprised of palisade and spongy a stream bed and observe the exposed root systems of
parenchyma tissues and veins. Chloroplasts are present large trees, or attempt to pull weeds, to get a first-hand

Figure 4.53 Exposed roots of white elm.

chapter 4 |
The Plant B ody

84
pericycle

Figure 4.54 A. Cross-section of the root showing the primary

tissues. 6. Development of a root hair.

idea of the anchorage function of roots (Fig. 4.53). Plants

with unusual roots provide an anchorage function in
atypical ways. Adventitious roots on ivy, for example,
develop from the stems; these short roots have a flat end-
pad that anchors the vines to their growing surface. The
sticky substance that is secreted for this purpose tends to
erode building surfaces, offen requiring the ivy to be
pulled down before the building crumbles. Parasitic plants,
like dodder, anchor themselves by sinking haustorial roots
into their host's vascular tissue in order to tap its water
and nutrient supply.
The storage function of roots is seen in carrots, which
have a large tap root that undergoes secondary growth
(Fig. 4.55E). Its mass of secondary xylem, however, is

composed mostly of storage parenchyma cells stuffed with

water and carbohydrates. Root storage usually assists the
plant at some developmental stage, such as providing
nutrients during flower production. Sugar beets, for
instance, are biennial plants that begin to fill their storage
root with food at the end of the first growing season. This
stored material is partially used to assist the growth of
early shoots during the following season, but flowering
during summer uses most of the stored energy.
In a herbacious plant water is responsible for about 95%
of root Weight. Water is needed for all root processes and
for every metabolic reaction. Soil water contains dissolved
salts and minerals, such as potassium, sulfur, absorbed by the root hairs and epidermal cells and is then
phosphorous, calcium, magnesium, which are needed by passed on by diffusion and active selective processes to
the plant (Chapter 2). For these reasons, plants have the conducting elements (Fig. 4.54; also see Chapter 5).
developed an elaborate system for the absorption and
conduction of water (Chapter 5). Root hairs penetrate
between small soil particles and are in intimate contact Types of Root Systems
with them and the water film that surrounds them (Fig.
4.54). The cell walls of the root hairs and epidermal cells
are made of cellulose and pectin. The pectic coat on the When many grasses and small garden plants
pulled up,
outside of the cell wall makes the root hairs gummy, take withthem a massive clump of soil. This happens
facilitating their adherence to soil particles. Water is because the fibrous root system of these plants consists

Types of Root Systems

85
 *34

Figure 4.55 Different types of root systems. A, shallow,

spreading, fibrous root system; B, fibrous root system penetrating
the soil evenly from 1-1.5 m; C, tap root system, in which main
primary root penetrates soil 2.5 m or more; D, fibrous root system
developed from adventitious roots growing at lower nodes of
stem; E, tap root system in carrot (Daucus carota).

of severalmain roots that branch to form a dense mass of roots in the upper 15 cm (Fig. 4.56). However, these
intermeshed lateral roots (Fig. 4.55). plants may extend lateral roots well beyond the expansion
Vegetables with a large storage root, like the carrot, ofoverhead branches. In areas of closely packed trees, or
have a tap root system that consists of one main root on sandy soils, competition and low surface moisture may
from which lateral roots radiate (Fig. 4.55). Some desert reduce the amount of area covered and encourage deeper
plants have a rapidly growing tap root system that enables root penetration.
them to penetrate the soil quickly to reach deep sources
of water. The depth of root penetration of various plants is
quite remarkable. A Nebraska botanist, John
University of Development of Root Systems
Weaver, exhumed the entire root systems of 45 species of
native plants from the Midwest prairie. Only four species The seeds of higher plants contain a small undeveloped

maintained root systems primarily in topsoil; most of them plant, theembryo. When the seed germinates, the
had roots that extended to 4 m deep. Even in very hard embryonic root, or radicle, extends by the division and
soils, the depth of root penetration well exceeded the elongation of its cells to form the primary root (Fig. 4.57).
height of the above-ground plant. A typical annual plant, Tap root systems develop from only one enlarged primary
like corn or rye, will build an immense fibrous root system root, which then forms lateral or secondary roots.

in one growing season. In a study on rye, a single plant 50 Further branching of secondary roots give rise to
cm tall, with 80 tillers (shoot branches), had a root system succeeding orders of roots, for example, tertiary,
2 quarternary, etc. Fibrous root systems develop
amounting to 210 m of surface area compared to only 4 in a
2
to 5 m of the above-ground portion (Table 4.2). different way. The embryo of some grasses, like barley,

In trees and shrubs, most functioning roots are localized consists of not only one radicle, but also of several

in the upper 1 m of soil, with the majority of the feeder additional embryonic roots called seminal roots. The
seminal roots emerge soon after the radicle during
germination. They elongate rapidly, and soon it is not
possible to distinguish the primary root. Usually the
seminal roots do not persist, but in some grasses they

Table 4.2 may function throughout the plant's life.

Rye Root System Measurements Roots that develop from organs other than roots are
called adventitious roots. In most instances, adventitious
Root Number roots develop at the nodes There are several
of the stem.
Length (m)
common examples a young corn
of adventitious roots. In

Main roots 143 plant, soon after the emergence and development of a
65
Secondary roots 35,600 5,181
rudimentary root system, prop roots develop from the
Tertiary roots 2,300,000 174,947 shoot node nearest the soil level (Fig. 4.55). These prop
Quarternary roots 1 1,500,000 441,938 roots function as roots, but also assist in the support of

Total 14,000,000 609,570 the plant in the soil. Banyan (Ficus bengalensis) trees
grow in saline mud lagoons and tidal marshes.
in tropical

chapter 4 |
The Plant Body

86
 Pippin on M. II,
Figure 4 56 Root system of Cox's Orange
Note main
excavated when 16 years old from brick-earth soil.
vertical "sinkers.
scaffolding of roots about 25 cm deep and

Types of Ro ot Systems

87
Figure 4.57 Radish seedling (Raphanus sativus). A, photograph
showing root hair growth, x2; Sand C, diagrams showing
growth regions.

Branches of these trees form adventitious prop roots that divide; on the average, one daughter cell of each division
extend down into the mud where they enlarge and actually remains as part of the meristem and divides again, while
prop up the large branches. The banyan is considered a the other daughter cell differentiates into a mature,
sacred tree in some parts of India. Indian merchants have specialized cell.

in the past held open-air bazaars among the prop roots The central portion of the apical meristem is composed
and expansive branches of the banyan (Fig. 4.58). of a pellet-shaped region of cells known as the quiescent
In some instances, adventitious roots do not form at center (Fig. 4.59). These cells divide, but at an extremely
nodes. Pieces of stem, like a cane from a blackberry plant, slow rate. They act as a regulatory center by releasing
or a branch of willow, can be made to root from their cut dividing cells, initials, just fast enough to continuously
ends simply by placing them in moist soil. Leaves from maintain the root's shape and to keep pace with its

Begonia and several other species also can be rooted growth. The function of the quiescent center is unknown;
simply by soaking them in water. Many commercially one hypothesis, however, is that may be the location
it

important ornamentals, in fact, are reproduced by root where growth regulators are synthesized and released to
propagation from leaves or stems (Chapter 8). control development of the root as new cells are made.
Some initials divide and produce cells in front of the
apical meristem to form the root cap. The root cap is a
Internal Anatomy thimble-shaped region of cells that surrounds the apical
meristem and precedes as the root elongates and forces
it

its way through the soil. The root cap not only protects
Root Apex
the apical meristem and lubricates its passage through the
In our discussion of root structure let us first begin with soil, but also is the site that perceives gravity and controls

the embryonic root, the radicle. The root apical meristem the direction of root growth.
at the tip of the radicle is composed of small, regularly Initial cells behind the quiescent center divide to supply
shaped cells that are capable of dividing. The emergence the cells for the primary root tissues. These cells continue
of the radicle during germination is dependent on the to divide, so that the region of cell divisions may extend as
initiation of cell division in this region. After the radicle far as 1 .5 mm from the tip of the root. Beyond this region,
emerges, the cells of the root apical meristem continue to as far as 5 mm from the tip of the root, the cells are

chapter 4 |
The Plant Body

88
Figure 4.58 Banyan tree with ancient market under aerial prop
roots.

rapidly elongating in preparation to perform their mature their cell wall into the surrounding soil. Root hairs form in

functions. the zone of differentiation in the root. It is most likely that

The can be conveniently divided into regions of

root root hairs have as their principal function to reach and tap
cells (tissues) thatperform different and specific functions for water in the soil (Figs. 4.54, 4.59). New root hairs
in the mature root. These tissues originate from zones continually form as old ones die. In one rye (Secale) plant,
called primary meristems; they are the protoderm, 100 million new root hairs are estimated to be formed
ground meristem, and procambium (Fig. 4.59). These each day.
are similar to the primary meristems in the shoot. Ground meristem cells produce the cells of the cortex.
Derivative cells from these three primary meristems The cortex consists mostly of storage parenchyma and
elongate and differentiate into the three primary tissues: occasionally of sclerenchyma cells. The innermost cell

epidermis, cortex, and vascular tissue. layer of the cortex is the endodermis. Endodermal cell

walls are usually thin, except for a suberized band on the

radial and transverse walls, the Casparian strip (Fig.

Formation of Primary Tissues 4.60). Electron micrographs show that the plasmalemma
ofthe endodermal cell is fused to the Casparian strip. This
The protoderm consists of uniformly shaped meristematic arrangement creates a barrier in these walls, making them
cells. At some distance from the root apex, some cells of impermeable to the passage of water and forcing water to
the protoderm develop into root hairs by the extension of pass through the protoplasm of the endodermal cell layer.

Internal Anatomy

89
 procambium

cortex

primary phloem

^^ primary xylem

epidermis
« — pericycle

endodermis

region of
elongation

central cylinder

/''
. .
.

root cap

procambium

protoderm

ground meristem

apical meristem

quiescent center

root cap

Figure 4.59 Photographic (A) and diagrammatic (B)

representation ot a longitudinal-section through a root from the tip
to the initiation of the vascular tissue. C. Longitudinal-section
through root tip meristem.

chapter 4 |
The Plant Body

90
 Cosparian strip

pericycle

Figure 4.60 Three-dimensional view of two endodermal cells.

The suberized Casparian strip is shown in gray. Water from
solution outside the root wets the cellulose walls until it reaches
the Casparian strip. Since water cannot wet and cross the
Casparian strip, the cell walls inside the Casparian strip are wet
by water that has passed through the protoplast of the
endodermal cell.

This enables the root to regulate the movement of members and companion cells (in flowering plants); and
dissolved materials (Fig. 4.54). sieve cells (in conifers and ferns).
Endodermal mature regions of the root often
cells in The outer layer(s) of cells that comprise the stele is

develop partialsecondary cell walls. Passage cells called the pericycle. This cell layer(s) is meristematic and
adjacent to xylem points often do not form secondary plays several important roles in the life cycle of the root. In

walls, thus ensuring the continued passage of water. the shoot, recall that new leaves are formed at the surface
The vascular cylinder or stele develops from the and very near the apical meristem of the shoot. In roots,
procambium and forms the central core of the root. In lateral roots are instead initiated below the surface and at
dicotyledonous roots, the xylem occupies the middle of a distance from the meristem.
the stele, with the periphery consisting of alternating radii Initiation of lateral roots at particular locations is

of xylem and phloem monocotyledonous

(Fig. 4.61 A). In controlled by chemical growth regulators that cause
roots, the central core contains parenchyma cells (Fig. pericycle cells to begin to divide (Fig. 4.62). The lateral
4.61 B). The first matured xylem-conducting elements, the root primordia, which result, continue to form new cells
protoxylem, develop at the points of the radiating xylem. that in turn elongate. Endodermal cells subtending this

Dicot roots with two protoxylem points are called diarch region often also divide for a short time, contributing cells
roots, those with three points are and so on; most
triarch, to the tip of the new lateral root. As expands, the lateral
it

monocot roots have more than five protoxylem points and root pushes its way through and destroys the cortical cells
are therefore called polyarch. Metaxylem cells form within and the outer epidermis. As emerges, the new lateral
it

the mass of xylem to complete the central core. root becomes organized into an apical meristem with
The protoxylem is modified so that is capable of it primary meristems identical to its parent root. This system

transporting water while the root is elongating; that is, of lateral root formation may present a hazard in the
protoxylem cells must be able to withstand the forces that plant's development, because the lateral root forms a
move and still be flexible enough to elongate. This
water, wound in the cortex and epidermis of the primary root. If

is accomplished by the formation of a secondary cell wall the wound doesn't heal rapidly, is a good it point of entry
in the shape of annular rings or as a spiral (Fig. 4.1 4/4). for plant pathogens.
When the root has completely elongated, the metaxylem
cells mature. They no longer are required to elongate and,
consequently, form thick secondary cell walls with pitted Secondary Growth in Roots
areas through which lateral exchange may take place (Fig.
4.1 4/4). Protoxylem cells often become crushed while the Roots grow at such fast rates that the primary tissues are
metaxylem develops. Xylem of roots consists of vessel not capable of supplying the needs of a long-lived plant
members (in flowering plants), tracheids (in most ferns body. In these cases, secondary tissues are developed. As
and conifers), parenchyma, and fibers. in the stem, two secondary or lateral meristems are
Phloem cells form in the spaces between the protoxylem required to form these tissues, the vascular cambium and
arms. The protophloem is the first part of the vascular the cork cambium.
system to become functional. These cells form at the In roots, secondary growth is initiated by the active
periphery of the phloem and function primarily during root division of (residual) procambium cells between the arcs
elongation. Metaphloem cells develop toward the inside, of xylem and phloem. These cells form secondary xylem
and they function during the plant's adult life. Phloem of to the inside and phloem to the outside (Fig. 4.63). After a
roots consists of parenchyma and fibers; sieve-tube time, the crescent-shaped region of dividing cells joins

Internal Anatomy

91
 ,.'-

epidermis S :.
'

4 r

cortex

Y
'

3 ' '•' • 1
- •--• ,x;,i •• :
v.' •••' 8'

" ;' *****

."-•'• >
;" i, ' '"'

.-.:'' •. •'• •
. -.-. .

'-
.

vv::;-

epidermis

Figure 4.61 Cross section views of typical roots. A,

dicotyledonous root of Ranunculus sp., X83. B,
monocotyledonous root from corn (Zea mays), X79. endodermis cortical parenchyr

Figure 4.62 Branch roots, A, initiation of branch carrot (Daucus

carota) root through formation of meristematic cells in pericycle;
8 and C, enlargement of meristematic region; D, young root
pushing through cortex, X50.

chapter 4 |
The Plant Body

92
 It K
C)5 OX ^
4 An '
,cO ' QX^^c V. I
x

ndodermis

secondary phloe

pericycle

mary phloe

cambium

imary xylem

secondary xyle

Figure 4.63 Diagrammatic representation ot the development of xylem internally and secondary phloem externally (light gray). The
the secondary plant body in a root. A, at the completion of primary phloem is being pushed outward and a small amount of
primary growth a row of procambium cells remains (light green) pericycle (dark green) is still associated with it. D, a smooth circle
and a complete circle of pericycle is present (dark green). B, the of vascular cambium forms, producing secondary xylem and
procambium (light green) joins with the pericyclic cells outside phloem. The primary remains in the center of the stem, the
the xylem arms to form a continuous cyclinder of vascular primary p hloem has been crushed, and only a small amount of
cambium (light green). C, the vascular cambium forms secondary Jhe pericycie remains"

with the pericycle, which in turn also begins to divide. This

connection forms a complete ring of vascular cambium
around the entire vascular cylinder Secondary
(stele).

tissues are produced during the growing season to form

Special Roots
annual growth rings, just as in the stem. Continued growth
expands the root and finally causes the splitting, sloughing Roots of a great many plants do not have the general
off, and destruction of the cortex and epidermis. However, characteristics common to most roots. We have already
early stresses of expansion, plus the activity of growth discussed examples of adventitious roots that arise from
hormones, stimulates the pericycle between the xylem nonroot origins, the roots of parasitic plants, and the
arms to resume division and form the cork cambium. This uniquely shaped sucker roots of clinging ivy vines. Other
meristem forms cork cells to the outside, and parenchyma plants, especially those forming partnership with
to the inside, just as in the stem. microorganisms, have specialized root structures.

Special Roots

93
Mycorrhizae collapse of cortical cells is thought to cause contraction.
The vascular tissue in contracted roots forms a twisted,
Mycorrhizae are short roots common to many plants; they undulated mass (Fig. 4.64).
represent an association with a soil-borne fungus. Two
types of mycorrhizal roots may be found, depending on
Summary
whether the fungus penetrates into the root cells or not.
Ectotrophic types are found in roots of such trees as 1. The principal functions of roots are absorption of water
pines (Pinus), birches (Betula), willows (Salix), and oaks and nutrients,conduction of absorbed materials and
(Quercus). This type causes a drastic change in the root food, and anchorage of the plant in the soil.
shape (Fig. 4.64). These mycorrhizal roots are about 0.5 2. There are two general types of root systems: a fibrous
cm long, have no root cap, and exhibit a simple monarch root system and a tap root system.
stele. In addition, the fungus penetrates between the cell 3. Roots differ from all stems because they lack nodes
walls of the cortex and forms a covering sheath, or and internodes.
mantle, of fungal hyphae (Chapter 12) around the entire 4. The root tip is divided into four zones of specialization:
root. Mycorrhizal roots are short and forked and (a) root cap, which protects the (b) meristematic
sometimes are borne as tight clusters. Endotrophic region as it moves through the soil; (c) a region of

mycorrhizae do not form a mantle over the root, and the elongation; and (d) a region of differentiation
fungus actually enters the cortex cells. Mycorrhizal roots characterized externally by root hairs and internally by
are more efficient in mineral absorption, but they are the formation of primary vascular tissues.
apparently not absolutely essential for the growth of the 5. Soil water and nutrients move easily through epidermal
usual host plants. This known, because plants that are
is and cortical tissues.
artificially fed adequate nutrients can grow without 6. The endodermis forms the innermost cell layer of the
mycorrhizae. Mycorrhizae also may be beneficial to their cortex. A suberized band, the Casparian strip, in radial

host plants by secreting hormones or antibiotic agents that and transverse walls, completely encircles endodermal
reduce the potential of plant disease. Mycorrhizae are also cells.

discussed in Chapter 12. 7. Water cannot move across the Casparian strip.

Therefore, all water with dissolved nutrients inside the

endodermis has passed through the protoplasts of

Bacterial Nodules endodermal cells.

8. The pericycle, a row of cells internal to the
Some plants, like certain legumes, are capable of fixing endodermis, represents the outermost row of cells of
nitrogen, that is, changing N 2 in the soil into the NH 4 + the vascular cylinder; they have differentiated from
(ammonia) form of nitrogen that is usable by the plant. procambium. Cells of pericycle may eventually form
This is done by an unusual relationship between the part of vascular cambium or cork cambium or give rise
bacterium Rhizobium and the roots of legume plants. Root to branch roots.
cells are infected by the passage of a thin infection thread 9. In cross section, the primary xylem is star-shaped and
of the bacteria into the root hair cells of roots. The generally triarch to pentarch. Primary phloem arises
infection thread passes through the root hair by causing between the arms of primary xylem. There is no pith in
the cell membrane to enfold as penetrates the cell. The
it the roots of dicotyledons.
infection thread grows through the epidermal cell into the 10. Water and inorganic nutrients enter primary xylem
cortical cells. The bacteria then divide and also stimulate from parenchyma cells without passing through cells
the cortical cells to divide, thereby forming the root nodule of phloem.
(Fig. 4.64). The bacteria are the actual agents for fixing 11. Water and inorganic nutrients enter root-hair cells,
the nitrogen. pass through the cortex, are filtered by protoplasts of
endodermal cells, cross the pericycle, and move
directly into the vessels or tracheids of primary xylem.

Contractile Roots 12. In roots having secondary growth, a vascular cambium

isformed by pericyclic cells over xylem arms and
Roots of the dandelion (Taraxacum), ginseng (Panax procambial cells remaining between primary xylem and
quinquefolium), and some other plants are capable of phloem. This vascular cambium at first forms a wavy
contracting. The result is that above-ground parts are kept line in cross section. It eventually becomes circular,

near the soil surface. This contraction is caused by the and there is little difference in the appearance of wood
radial expansion of cells in the root cortex. Sometimes, the from root or stem.

chapter 4 |
The Plant Body

94
 ^^^^VV^^^-, - -

enlorged
root cortex
cells contain
bacteria

Figure 4.64 Views of root modifications. A, contractile root; 6,

branched habit of mycorrhizal root; C, longitudinal section
through one of the Y-branched mycorrhizal roots; D, cross
section through one root showing the fungal mantle; E, soybean
(Glycine max) root with bacterial nodules; F, cross section of root
and nodule.

Summary

95
 5 CHAPTER

5 the absorption and

transport systems

consumed by photosynthesis and

The living plant collects materials

environment and assembles them

from the
into plant cells
is

evaporation. In
lost

leaves, the water passes from the xylem

through

and organs. It therefore requires systems for vessels and tracheids to the mesophyll cells, from which it

absorbing materials, assembling new substances, and evaporates into the intercellular spaces and finally escapes
transporting materials within the plant. The assembly from the leaf as water vapor moving through the stomata.
systems, metabolism and photosynthesis, are discussed At most stages of its journey through the plant, water
Chapters 2 and 6. The present chapter deals with the molecules have the option of moving either through the
absorption and transport systems. protoplasts or through cell walls (Fig. 5.2). Thus, for
example, water first enters the walls of the root hair cells.
Consisting of a porous meshwork of cellulose microfibrils,
these walls readily accept water and allow
Water Absorption and movement. The epidermal walls touch cell walls in the
it rather free

Transport cortex, permitting the passage of water molecules from

cell to cell through the walls. In addition, there are water-
spaces between
filled cortical cells through which bulk
Green plants require only a few simple materials from the water may move.
environment: water, carbon dioxide, oxygen, and several Alternatively, water molecules may pass through the
minerals. Of these materials, water is the substance that plasma membranes of the cells and enter the protoplasts.
plants absorb and transport in the greatest quantities. The membranes offer little barrier to such movement of
Some absorbed water is consumed in metabolic
of the water in either direction. Once in the protoplast of a cell,

reactions such as photosynthesis and some of is it water may move to other cells by means of the
retained in the vacuoles as cells grow. But as much as protoplasmic bridges known as plasmodesmata.
98% of the water that enters the plant is lost through the The endodermis restricts the movement of water. The
shoot system by evaporation. This evaporative loss is endodermal cells are tightly pressed together and their
termed transpiration. Casparian (Chapter 4) present an impermeable layer
strips
Plants transpire a large amount of water. A single corn that water cannot cross. The endodermal barrier prevents
plant in Kansas, between May 5 and September 8, water from moving deeper into the root unless it travels
transpired 196 liters. It has been estimated that the through the protoplasts of the endodermal cells. This
amount of water lost by a grove of red maple trees in a limitation is considered an important means for giving the
growing season may be equivalent to a sheet of water 72 plant control, via the selective permeability of the plasma
cm deep over the entire grove. membrane, over the movement of solutes (not water)
An alfalfa plant may transpire 900 units of water for between the soil and the vascular system. This could also
each unit of dry matter produced, but a millet plant may be important in limiting the entry of toxic materials from
transpire 248 units of water for each unit of dry matter. the soil and the loss of important materials to the soil.
Although these values vary with the environmental In the xylem, the principal movement of water occurs as
conditions in which the plants are grown, it is evident that a bulk flow in the vessels and tracheids. Flow is probably
plants differ in their water economy. not appreciable within the secondarily thickened walls of
Plants draw their major water supplies from the soil these dead cells.
solution, the bulk water that lies between soil particles.

Water first enters the plant (Fig. 5.1) through the

epidermal cells of the root tip. The flow moves through the
cortex, the endodermis, and the pericycle, and finally into Water Potential and Water Movement
the xylem conduits: the vessels or tracheids. The liquid in
the xylem, a solution known as xylem sap, moves up the We have considered the path that water follows through
plant to any region where water is being removed: to the plant. Now we ask what is the force that drives the
growing shoots and fruits where water is accumulated in flow of water? The answer requires the concept of water
the vacuoles of enlarging cells, and to leaves where water potential.

96
 trocheid of leaf vein

pericycle cortex \_ epidermis

endodermis •

Figure 5.1 A, diagram showing water movement in a whole

bean plant. B, details of the xylem pathway.

Whenever a wet and a dry place are joined by a water If two ends of a water column are at different water
mass, water flows toward the dry region. This is the potentials, water will spontaneously move toward the end
principle behind the movement of water in plants. with the lower water potential. This is a formal way of
"Wetness" and "dryness" are relative terms. Pure water is saying that water tends to move from wet places to dry
perfectly wet, and a region that contains no water is places.
perfectly dry. However, between these extremes there are The concept of water potential is useful in explaining
many systems that contain water mixed with other osmosis and turgor pressure, two phenomena that are
materials. Physical chemists, who measure the degree of important in the mechanisms of water movement. Osmosis
wetness in a system, call it the water potential. Pure water is the diffusion of water across a differentially permeable
is assigned the highest potential; if anything is added to membrane (Fig. 5.3). It occurs when the solutions on the

the water, such as cellulose microfibrils or sugar two sides of the membrane differ in water potential. If the
molecules, the water potential decreases. In moist air, two solutions are at the same pressure and temperature,
where water is present as a gas, the water potential then the one with the higher solute concentration will have
increases more water is added to the air. The presence
if a lower water potential; it is less like pure water. If the
of other gases does not influence the water potential of membrane will not permit solutes to pass, water will then
humid air to any significant degree. move along the water potential gradient toward the

Water Absorption and Transport

97
 Casparian
strip

Figure 5.2 water and ion movement in outer root

Paths of
tissues. Two pathways are shown. Water and ions
alternative
move similarly but ions may be aided by transport mechanisms in
crossing plasma membranes. 1 = plasma membranes;
2 = protoplasm; 3 = plasmodesmata; 4 = cell walls; 5 = intercellular
space.

solution that contains the most solute particles. This water expansion is possible. Every incoming water molecule
flow across a membrane is osmosis; brings the two it adds to the hydrostatic pressure within the cell. This can
solutions closer to the same water potential. Unless an be appreciated by comparing the molecules to people
outside force intervenes, osmosis will continue until the trying to crowd into a full room. The added pressure is
two solutions achieve equal water potentials. Then termed turgor pressure. It is an outward pressure that is
equilibrium prevails. exerted on the cell wall by the crowded molecules in the
The living plant cell is in contact with a wet cell wall cell. Against this turgor pressure the wall exerts an

(Fig. 5.4). The water between the wall's carbohydrate opposite, inward-directed force called wall pressure. When
polymers is a dilute salt solution. On the other side of the the cell is swelling, turgor pressure exceeds wall pressure;
plasma membrane, the contents of the protoplast form a the wall yields and stretches. At equilibrium the turgor and
second more concentrated solution. Thus, water will tend wall pressures are equal and opposite.
to move by osmosis from the wall into the protoplast. This Turgor pressure together with the structure of the wall
transfer of water raises the volume of the protoplast. give the cell a rigid shape. Cells go limp when they lose
Against the swelling, the wall stretches a little but soon their turgor pressure through water loss. A plant with
comes to the limit of its elasticity and no more volume greatly reduced turgor pressure is said to be wilted.

membrane
membrane

• ?:•':•
:'P"
t
• • o.o :
. « ••••*• •
o
.

o
Y
:.o :- .• o\*
(

o , • .

• .
o ^•''" °
. • • * .'
cytoplasm
cell wall

Figure 5.3 System that would show osmosis. A membrane Figure 5.4 Conditions at the surface of the cell, affecting water
separates two solutions that contain water (o) and solute (•) movement. The solution in the cell wall contains fewer solutes
molecules. The pores in the membrane permit water but not than cytoplasm, but polymers such as cellulose are present.
solutes to pass. Water potential will be lower in the left-hand
solution; water will move into that solution.

chapter 5 |
The Absorption and Transport Systems

98
 If a cell or group of cells is immersed in a solution that Table 5.1
has a higher solute concentration than that of the cell sap, Factors That Affect Water Potential in a Solution
water diffuses outward, and the turgor pressure in the cell
is reduced. Although the volume of the cell decreases Water Potential Water Potential
somewhat, more striking is the withdrawal of the Is Increased by: Is Decreased by:
protoplast from the cell wall and the decrease in the size
of the vacuole. This phenomenon is called plasmolysis. Pressure Solutes
The space between the cytoplasm and the cell wall in a Heat Polymers (adhesion)
plasmolyzed cell is filled with the solution inwhich the
cells are immersed. If plasmolyzed cells are immersed in
water or in a solution whose concentration is less than
that of the cell sap, the cells regain their turgor; water participate in four hydrogen bonds with as many as four
molecules diffuse inward. The result is that the entire water
other water molecules.
A cell may die if pronounced and
plasmolysis is mass shows cohesion, a tendency to hang together. The
prolonged. For example, a heavy application of ordinary
if forces of cohesion not only join molecules of the bulk
salt is placed on the soil where weeds are growing, so solution into a loose unit, but also they join the water
that the root cells are surrounded by a solution of high mass to the layer of bound water adhering to the cell wall
concentration, water diffuses from the cells, and they polymers. The effect is as if the polymers exert an
become severely plasmolyzed. If this state is prolonged, anchoring influence on water. This effect weakens in

the roots die. The salt killed the roots, not because it was proportion to the distance water is from the wall polymers.
toxic to the root cells, but rather because severe The water in the saturated wall that is far from the
plasmolysis was brought about by the high concentration polymers acts like free water. But when this free water is
of the soil solution. In parts of the western United States withdrawn, the remaining water is closer to the polymers
where rainfall is low and the evaporation rate high, salts of and is more stronglybound by the forces of adhesion. In
the soil may accumulate on the surface and form what are brief, the adhesion forces exerted by wall polymers tend to
known as "alkali flats." In such soils, the salt make a dilute solution behave osmotically as if it were
concentration of the soil solution may be so high that more concentrated in solutes.
many crop plants cannot grow; only species that are we take capillary action and adhesion into account,
If it

especially adapted to tolerate high salt concentration are appears now that water movement between the wall and
able to survive. These plants are known as halophytes. the protoplast is controlled not only by the concentrations
For reasons that cannot be simply stated, turgor of solutes and by turgor pressure but also by the influence
pressure (any kind of pressure, for that matter) raises the of polymers in the cell wall. These factors are summarized

water potential of the solution. Water stops entering the for convenience in Table 5.1. Equilibrium prevails (no net

cell when the turgor pressure has raised the protoplast's water flow) when the sum of these effects on water
water potential to equal that of the water in the cell wall. potential is the same on both sides of the membrane. In
Then equilibrium prevails. Thus we see
an interplay that any other condition the direction of water movement (and
between pressure and solute concentrations determines its pace) varies with the balance of factors.

the direction of water flow. The effect of increased

pressure is to make a concentrated solution behave more
like a dilute solution The Mechanism of Water Movement in
In these matters, the carbohydrate polymers in the wall the Plant
do more than restrain the protoplast. They also influence
the potential of the water in the The effect is to
cell wall. Water moves within the plant toward any location where
decrease the water potential and the effect depends very the water potential is low; that is, toward any region where
strongly on the amount of water in the wall. In a the solute concentration is rising or where water is being
completely saturated wall, dripping wet, the effect of the removed. Photosynthesis both consumes water and
wall polymers is very slight. But when the wall begins to produces solutes, making the locations of photosynthesis
dry out, the water potential quickly becomes very sensitive low in water potential and the targets of water flow. The
to further drying. The drying wall develops an immensely water potential is also relatively low in the cells of growing
powerful tendency to keep its remaining water and to draw regions; here the cells have walls that continuously stretch
water from neighboring regions. This is the same principle under turgor pressure. This plasticity of the walls keeps
behind the absorptive powers of blotting paper. The water the turgor pressure from rising high enough to stop the
is said to move
response to capillary
in action. The osmotic movement of water into the cells. The incoming

driving force occurs because the wall polymers form water provides the bulk needed for growth.
bonds with the adjacent water molecules. Such binding In transpiration, water is moved by a third means. Here,
between water molecules and polymers is known as the primary event is the diffusion of water vapor from the
adhesion. The polymers may be viewed as having a humid air in the leaf to the dry air outside the leaf. This
tightly held blanket of bound water. movement leaves the air channels within the leaf with a
Equally important, water molecules tend to bond to one reduced water potential. Water evaporates from the moist
another. These intermolecular bonds, termed hydrogen walls of the leaf cells, obeying the rule of movement
bonds (Appendix), are weaker than the covalent bonds toward regions of low water potential. The walls are left

within a molecule. However, each water molecule can more dry and the remaining water drops in potential

Water Absorption and Transport

99
because of the capillary effect. Thus in transpiration, water work against gravity and resistance, a tall tree must be
movement is driven by withdrawing water from the wall able to survive with drier leaf cell walls than a short plant.
and exploiting the effect of capillary action or adhesion. Plant physiologists refer to this picture of water
Once a point of low water potential has been movement as the cohesion and transpiration-pull theory
established, the remaining question is how the motive (among other names). In this theory, note that water is
force is transmitted throughout the plant and into the soil being pulled, not pushed, and that the water column may
solution from which the water is obtained. Cohesion is a be under considerable tension. has often been asked
It

major part of the answer. The water mass within the whether water columns really can sustain the tensions
plant-soil system is a continuous whole that is bound needed for lifting water as high as the tops of tall trees,
together by the forces of cohesion. One may view the cell which may exceed 100 meters in height. Laboratory
walls that stand between the soil solution and the top of experiments have shown that thin columns of water do
the plant as a series of porous obstructions that exert a indeed have the required tensile strength and can
drag on the flow of water while leaving the forces of withstand pulling forces great enough to raise water up to
cohesion in operation. Thus, when the water potential is a height of 300 meters.
reduced by drying the walls of the leaf cells, the adhesion But the presence of a single, tiny gas bubble destroys
forces that attract nearby water molecules are effectively the tensile strength of a water column. With a bubble in

pulling on a chain of water molecules that extends all the the column, tension can raise water no higher than 9.75
way down to the soil solution and into each cell of the meters (at sea level). Beyond that point, further increases
plant. in tension merely expand the bubble with no further rise in

The forces that move water up the plant must work the water column.
against gravity as well as the drag (resistance) exerted by Under sufficient tension, a water column may
The
cell walls. taller the tree, the greater the weight of the spontaneously produce bubbles. This is called cavitation.
water columns that must be drawn up the xylem and the With sustained tension, bubbles will expand until they

greater the force that must be sustained to keep up a meet a such as a cell wall. Bubbles cannot cross
barrier
given flow rate. The driving force in transpiration is cell walls because the wall polymers exert too great an

generated by the drying leaf cell walls. Therefore, to do its attraction for water. In the plant, the conducting area of
the xylem is divided into a large number of microscopically
narrow vessels and tracheids. Cavitation may occur in
many of these during transpiration, but the bubble will be
confined to the vessel or tracheid of its origin and the rest
of the xylem can continue to function. When transpiration
stops (e.g., at night when stomata close), bubbles formed
by cavitation may be resorbed so that the blocked
conducting cell can function again.

Root Pressure and Guttation

It is an old observation that the rooted stump of a freshly

cut plant often "bleeds" xylem sap from the cut end. Such
"bleeding" represents the movement of water up the
xylem without transpiration as a driving force. The xylem
sap is pushed rather than pulled upward in this case. The
pushing force is termed root pressure. It is an osmotic
phenomenon: the xylem sap has a higher concentration of
ions than the soil solution because of the action of ion-
transport systems in the plasma membranes of root cells.
Thus water moves osmotically from the soil into the xylem.
This raises the volume of liquid in the xylem and forces it
up the stem. In this case the xylem contents are moved by
positive pressure rather than by tension.
Root pressure is unimportant as a source of water
movement when transpiration is occurring, but may be a
it

significant water-moving force in small plants when

transpiration is not taking place. For instance, when a
well-watered, vigorously growing tomato plant is placed
under a ceases as the atmosphere in
bell jar, transpiration

the jar becomes saturated. Continued absorption of water

by osmosis in the roots then results in a slow exudation of
water from the tips of the leaves (Fig. 5.5). This loss of
liquid from leaves is termed guttation. Many plants have
(Hordeum specialized openings called hydathodes at the tips of their
Figure 5.5 Guttation from tips of barley vulgare)
leaves. leaves, through which guttation liquid passes outward.

chapter 5 |
The Absorpti on and Transport Syst ems

100
How Roots "Locate" Water component of the air from which is absorbed (about it

0.03% of air is carbon dioxide). Correspondingly, there is

Without extensive irrigation or trequent rainfall, plants an advantage in being able to trap the largest possible
usually remove water from the soil faster than it can be fraction of the carbon dioxide that passes the plant in

replaced by movement from neighboring regions. Hence, currents of air.

for continued water absorption the roots must continually When air that contains carbon dioxide meets the plant
grow into new areas
have not yet been dried
that surlace, the only reason it enters the plant is that
Geotropism (Chapter 8) gives the growth of roots a photosynthesis keeps the carbon dioxide concentration in

general downward orientation, increasing the probability the tissues very low. This causes C0 2 to enter by diffusion
that the root will contact untapped reserves of water. In along the concentration gradient The steeper the gradient,
addition, pressures exerted by contact with soil particles the faster the flow of C0 2 into the plant and the larger a
often force roots to grow laterally and branch roots tend to fraction of the available C0 2 the plant will be able to trap
invade any moist soil they meet. from a moving air stream. The concentration gradient is

It is evident that roots do not "go in search of water." steepest when the photosynthetic cells are directly in

However, they grow only in moist soil. In practice, this contact with the air.

means that if the top 30 cm of soil are moist, and dry soil In line with these principles, the plant body tends to be
lies below it, the roots of plants will be confined to the structured in a way that brings as many photosynthetic
upper, moist soil. In irrigation practice, this relationship of cells as possible near to the outside air. The broad, flat

soil moisture and root growth is important. In irrigating a shape of the leaf greatly increases the area of plant tissue
garden or lawn with a hose, one is easily misled into that is close to the air. Within the leaf, air spaces form an
believing, because the soil surface is wet, that sufficient extensive system of channels, giving almost all the
water has been applied. Examination may reveal that at photosynthetic mesophyll cells direct contact with an air

depths of 12 cm or more the soil is very dry too dry for — space that leads to the outside of the leaf
the growth of roots. In growing plants, is nearly always it

desirable to stimulate maximum root growth and a root

system that attains its normal depth and spread. This may
be accomplished in part by keeping the soil moist to the
Control of Carbon Dioxide
proper depth. The loss of water from soils at levels below Absorption and Water Loss
the first few centimeters is essentially due to plants. Thus
one of the principal reasons for removing weeds from a
growing crop is that they deplete water from the soil. Cells that are directly exposed to dry air quickly lose too
If the soil solution increases in solute content, the rate much water by same air exposure
evaporation. Thus, the
of water absorption by the roots declines. When the that promotes C0 2 capture also raises dangerous
concentrations of the two solutions are the same, water problems of water loss. The balance between these two
intake ceases. And, if the soil solution becomes more factors has doubtless played a great part in the evolution
highly concentrated than the cell sap, owing to excessive of leaf structure.
applications of fertilizers or of saline water, water will be In land plants, fatal water loss is prevented by
withdrawn from the root cells. Moreover, root growth is impermeable cork on older stems and by an impermeable
inhibited at high salt concentrations and the roots are not waxy cuticle that covers the young shoots and leaves.
able to extend into new soil areas. These coatings limit gas exchange to special apertures,
the stomata, which can be opened and closed to get the
best possible balance between C0 2 capture and water
loss. A typical leaf may have as many as 6 million stomata
Carbon Dioxide Absorption (Chapter 4).

The stomata function in the following way (Fig. 5.6).

Each pore is lined by a pair of special epidermal cells
Carbon dioxide is often the factor that most limits plant known as guard cells. These cells are sausage-shaped;
growth: in most plants it is the sole source of carbon for they also differ from ordinary platelike epidermal cells in

building organic compounds, and it is only a minor having thicker walls and an ability to perform

K* in; water in

stimulus: light

> stimuli: dessication

high leaf C0 2

K~ out; water out , . .

low turgor stoma closed high turgor stoma open

Figure 5.6 Diagram summarizing the mechanism by which

+
guard cells open and close stomata The gain or loss of K
involves active transport; water follows by osmosis. G = guard cell;
S = stoma.

Control of Carbon Dioxide Absorption and Water Loss

101
photosynthesis. The walls of the guard cells are thickest a low water potential that they extract water from the
on the side that faces the pore. This feature allows the protoplasts as well as from the xylem. This removes the
stoma to be opened and closed by changing the turgor of turgor pressure needed to maintain leaf shape and the
the guard cells. When turgor pressure is low, the guard leaves wilt. Midday wilting is common in big-leafed herbs
cells press together and the stomatal pore is closed. An such as squash plants. is a temporary condition; normal
It

increase in turgor pressure stretches the walls of the turgor returns when the rate of transpiration decreases in

guard cell slightly, the thinner portions stretching the the evening and the walls within the leaf can regain
most. This causes the guard cells to become curved, moisture. Plants that are native to hot, dry areas often
opening the stomatal pore. have reinforcing sclerenchyma in their leaves, a feature
The turgor in the guard cells is controlled by adding and that makes the leaf less sensitive to drying.
removing solutes. If solutes are added, the water potential Certain plants called xerophytes (Chapter 9) are able to
decreases; water enters the guard cells from adjacent survive in extremely dry habitats, whereas certain other

cells, and the turgor pressure rises. The reverse happens plants succumb. Such surviving plants undoubtedly
if the guard cells lose solutes. maintain a balance between water outgo and water intake,
+
Guard up potassium (K ) ions from adjacent
cells take whereas in plants that fail to survive the dryness, the loss
+
cells as the stomata open. They release the K ions as of water, at least for a period, exceeds the absorption of
the stomata close. This ion transfer seems to depend on water. Plants adapted to dry habitats often have roots that
an active transport "pump" in the plasma membrane. The penetrate deeply into the soil; or, like many desert
+
movements of K probably play a major part in controlling cacti, they have an extensive surface root system that can
the stomata. rapidly absorb limited rainfall from spring or summer
Organic solutes may also be important. Thus in some showers. Many desert plants have special water storage
cases starch grains in the guard cells are partially tissue, as in the plants with fleshy stems and leaves that
converted to free sugar as the stomata are induced to are commonly known as succulents (Fig. 9.30).
open. The reverse occurs when the stomata are caused to Anatomical features that are advantageous from the
close. These facts show that the solute relations that point of view of preventing water loss are (a) the cuticle of
control the stomata are likely to be complex. leaves, young stems, and fruits, (b) sunken stomata, (c)
Control of stomatal opening by turgor pressure guards distribution of stomata, and (d) reduction of the transpiring
the plant against excessive water loss: if the leaves lose surface. In addition, stomatal behavior is an important
water too fast, they wilt — that is, they lose turgor. This factor in controlling water loss.
closes the stomata, limiting further water loss. (It has been Most of the water lost from a plant passes out through
discovered recently that the hormone abscisic acid is the stomata. Some water, however, is lost through the

rapidly produced by leaves when they experience water cuticle. Various modifications in leaf structure reduce
stress. This hormone may signal the stomata to close cuticular transpiration; for example, thickening of the outer
before the loss is severe.) wall of the epidermal cells and the presence of a waxlike
In two relevant environmental factors act as
addition, material, cutin, in this wall. Most plants of arid and semi-
control cues for the stomata. Carbon dioxide inside the arid climates have a thicker cuticle than do those of humid
leaf causes the stomata to close, and light causes stomata climates.
to open. The result is that the stomata open only when In some plants (Fig. 5.7), the stomata are below the
there is light for photosynthesis, and when the carbon general level of the leaf surface. When this condition

dioxide has been reduced to a point where more is occurs, the water vapor must diffuse through a relatively
needed. long and sometimes tortuous passageway, with the result
However, some plants have their light signals reversed, that the diffusion rate is lessened.
so that the stomata open and take in C0 2 at night while The leaves of many have stomata only on the
plants
closing in the daytime. As C0 2 enters these plants, it is undersurface on both surfaces, mostly on the lower.
or, if

stored by attachment to an organic carrier molecule. In Loss of water from leaves that have stomata only on the
daylight, with the stomata closed, the stored C0 2 is lower surface is usually less than loss from leaves that
gradually released by the carrier and used in normal have an equal number of them on both the upper and
photosynthesis. This reversed control system is especially lower sides.
common in succulent plants of dry regions. Its value is In most organs of transpiration are
plants, the principal
that less water is lost with the stomata open at night leaves. Therefore, any decrease in leaf surface will reduce
because the lower night temperatures yield slower transpiration and conserve the water absorbed by roots. In
evaporation. corn and some other monocots, the leaves roll up during
The rate of transpiration and its effect on the plant drought, thus exposing less surface to the air than do fully
depend on the humidity and temperature of the air. Low expanded leaves. Cacti and some euphorbias have no
humidity and high temperature promote evaporation, foliage leaves; in these plants the transpiring surface is

increasing the rate of transpiration and the extent of drying restricted to the stem surface.
in the walls of leaf cells. At moderate transpiration rates, The leaves of many plants are clothed with hairs. It's

the mesophyll cells of leaves have a solute concentration possible that hairs reflect light, reducing leaf temperature.
that is high enough to enable them to maintain turgor In some plants, such as common mullein, the hairs reduce
pressure even though the walls are partially dried. But transpiration. But experiments with some plants have
with increased rates of evaporation, as in the midday heat shown that water loss from the leaves is greater when the
of a dry summer day, the leaf cell walls may develop such hairs are present than when they are lacking.

chapter 5 |
The Absorption and Transport Systems

102
 spongy
parenchyma

fibers sunken stomata

epidermis
Figure 5.7 Sunken stomata in crypts of a yucca leaf. Note also
the thick epidermal cuticle and the large bundles of fibers.

micro-organisms the
Mineral Ion Absorption The rate of growth
in soil.

of plants is influenced greatly by

available nitrogen. Nitrogen is very mobile in the plant and
Principal Mineral Elements can be translocated from mature to immature regions of a
plant. An early symptom of nitrogen deficiency is a
A variety of mineral elements are needed by the plant. yellowing of leaves, particularly the older leaves; then
These and the typical quantities withdrawn from the soil follows a stunting in the growth of all parts of the plant

by wheat plants are shown in Table 5.2. The mineral (Fig. 5.8E).

elements, discussed below, are largely absorbed in the An excess of available nitrogen results in vigorous
form of ions dissolved in the soil water. vegetative growth and a suppression of food storage and

is an essential component of proteins, nucleic

Nitrogen of fruit and seed development.
acids,and many other molecules that play important Sulfur comprises a small but vital fraction of the atoms
metabolic roles. The atmosphere is rich in N 2 gas; it
in many protein molecules. As disulfide bridges (SS), the

comprises 78% of the air. Some 8000 kilograms of N 2 lie sulfur atoms aid in stabilizing the folded protein, while

above every square meter of land. However, ordinary sulfhydryl (SH) groups participate in the active sites of
green plants get their nitrogen mainly as nitrate ions some enzymes. Some enzymes require the aid of certain
(N0 3 ~) from the soil. To a lesser degree, ammonium ions small molecules that may contain sulfur. Finally, the
+ machinery of photosynthesis includes some sulfur-
(NH 4 ) also serves as a nitrogen source, and some plants
apparently can derive nitrogen from certain organic containing compounds (e.g., ferredoxin).
compounds in the soil. Ordinary green plants obtain sulfur in the form of sulfate
Nitrogenous fertilizers, both natural and commercial, are ions. When plants deficient in sulfur are given sulfur
usually the most important fertilizers applied to growing fertilizers, the most noticeable response is an increased
plants. The chief commercial nitrogenous fertilizers contain root development and deeper green color of the leaves.
nitrogen (a) in the nitrate form; (b) in the form of ammonia Sulfur deficiency symptoms are shown in Fig. 5.8F.
or its compounds; compounds, such as
(c) in organic Phosphorus is a component of nucleic acids, energy-
cottonseed meal, and (d) in the amide form, such as urea carrying molecules such as ATP, and the important
and calcium cyanamide. Organic nitrogen in complex phospholipids of cellular membranes. Some proteins also
molecules cannot be used directly by plants but must first contain phosphate groups. The highest concentrations of
be converted into available forms through the action of phosphorus occur in rapidly growing plant parts, such as

Table 5-2
Amounts (in kg. per hk.) of Macro- and Micronutrients Removed from the Soil
in One Growing Season by a Wheat Crop

Macronutrients Micronutrients

Element N K P Ca S Mg Fe Mn B Zn Cu
Amount 85 47 17 13 12 9 0.8 0.6 0.3 0.2 0.03

Mineral Ion Absorption

103
maturing fruits, seeds, and shoot tips. that some plants may require small amounts of Na and
Applications of phosphorus to deficient soils promote Ni.)
root growth and hasten maturation, particularly in cereals Iron ions are components of several electron-carrying
Phosphates (potash) are the principle source of compounds (cytochromes and ferredoxin) essential in

phosphorus for plants. A tomato plant grown in culture respiration and photosynthesis. In picking up and giving
solution without phosphorus is shown in Fig. 5.8C. off electrons, the iron ion is alternately reduced and
Potassium serves as an enzyme activator. Over 40 oxidized by other compounds.
enzymes have been found to require potassium for Although iron is not a component of chlorophyll, it is

maximum activity. In addition, potassium ions are essential for chlorophyll synthesis. Iron deficiency may be
important in controlling the stomata This is a mobile responsible for chlorosis (Fig. 5.8H). The quantity of iron
element, which will migrate from older tissues to required by plants is very small. For example, chlorosis of
meristematic regions. For example, during the maturing of pineapples in Hawaii, because of the unavailability of iron
a fruit crop potassium moves from leaves into fruits. in the soil, is cured by spraying the leaves with iron salt

Any water-soluble inorganic compound of potassium, solutions. Orchard trees suffering from iron chlorosis may
such as potassium sulfate, phosphate, or nitrate salts, can be cured by injecting iron compounds into the trunk or
be used by plants as a potassium source. Potassium applying iron salts to the soil.

deficiency symptoms in the tomato are shown in Fig. 5.8D. Boron deficiency causes varied and complex symptoms
Calcium is required by all ordinary green plants. is It such as decreased root and shoot elongation, inhibition of
one of the constituents of the middle lamella of the cell flowering, and darkening of tissues. The metabolic basis of
wall, where it occurs in the form of calcium pectate. boron action is not clear, though boron is known to play a
Calcium affects the permeability of cytoplasmic part in regulating carbohydrate breakdown. Deficiency
membranes and the hydration of colloids. Calcium may be diseases may be cured by applying very small quantities
found in combination with organic acids in the plant. (10-25 kg) of sodium tetraborate (borax) per acre.
Oxalic acid, for example, is a by-product of metabolism. It However, excessive amounts of boron in the soil or water
is a soluble substance and is toxic to the protoplasm if it may be toxic; in fact, borates are used as weed killers.
reaches a high concentration in the cell. When united with Zinc is an activator or a part of several enzymes. One of
calcium, however, the soluble oxalic acid is converted into the earliest effects of zinc deficiency is a drop in
the highly insoluble calcium oxalate, which does not injure production of the growth hormone auxin (Chapter 8),
the protoplasm. There is also evidence that calcium favors resulting in inhibition of growth. Deficiency diseases such
the translocation of carbohydrates and amino acids and as the well-known "little-leaf" of deciduous fruit trees (Fig.
encourages root development. Calcium deficiency is 5.9) can be cured by spraying the trees with zinc salts, by
frequently characterized by a death of the growing points injecting dilute solutions of zinc salts into the trunks, or by
(Fig. 5.8G) because it is not readily translocated in the driving zinc brads into the trunks.These correctives show
plant from mature to immature regions. General how small are the quantities of zinc needed by the plant.
disorganization of cells and tissues also results from Manganese activates several enzymes and also plays an
calcium deficiency. This effect is consistent with one of essential role in the oxygen-liberating steps of
the key roles of calcium — maintaining the normal structure photosynthesis. The most striking deficiency symptom is

of cell membranes. chlorosis (Fig. 5.10), but this chlorosis is somewhat

Magnesium is a constituent of chlorophyll; it occupies a different from that caused by iron deficiency. In iron

central position in the molecule (Fig. 6.4). Many enzyme

reactions, particularly those involving a transfer of
+2
phosphate (energy metabolism) are activated by Mg
ions. Magnesium is also vital to the function of ribosomes
in protein synthesis.
A deficiency in magnesium results in chlorosis — a lack
of chlorophyll and, therefore, foliage that is pale yellowish
rather than green (Fig. 5.8/). This disease is common in

cultivated plants and in many cases applications of

magnesium effect a cure. However, chlorosis can also be
caused by deficiencies in other elements such as iron.
The materials listed above provide elements that are
known as macronutrients (macro = large) because
plants require them in large quantities. The macronutrient
elements are C, H, O, N, P, S, K, Ca, and Mg.
In addition, plants require tiny amounts of several other

elements. These, known as micronutrients, include at

least those listed below. It is difficult to study the plant's
need for that element is needed only in trace
an element if

quantities,because the element may be hard to eliminate

from the nutrient medium or the air around the plant. Thus
Figure 5.9 Disease of peach (Prunus persica) known as "little
physiologists anticipate that in the future more essential caused by a deficiency of zinc. Branch at left untreated;
leaf,''
elements will be discovered. (There is already evidence branch at right cured by driving in zinc-coated nails.

chapter 5 |
The Absorption and Transport Systems G~
Cft)v£H rfl K
104
Confer f m»lyb&"< um
Figure 5.8 Tomato plants grown in the nutrient culture
solutions to show visual symptoms of mineral deficiencies
COLOR PLATE 2
A, complete solution; 6, deficiency symptoms in leaves; (continued)
C, minus phosphorus; D, minus potassium; E, minus
nitrogen; F, minus sulfur; G, minus calcium; H, minus iron
/, minus magnesium; J, minus all micronutrients.
All plants subjected to deficient solutions were first grown
in a complete nutrient solution for two weeks.
Figure 8.1 Above. Pea
seedlings grown in light and
COLOR PLATE 2
in darkness Note minimum
stem elongation and
maximum leat development in
light. The center two plants
have had only one day in
light, which caused
straightening of the stem
(hook-opening) and start of
leaf expansion and greening,
as compared to the fully
etiolated seedlings on the left.

Figure 8.2 /Above. Pea leaf

development as a function of
light. Fully developed green
leaves from plants in
continuous light; early stage
of development after one day
in light (center); etiolated leaf
from dark-grown plant
protected by plummular hook.
All eight days old.

Figure 8.3 Right: Corn

seedlings grown in light or in
darkness showing the long
mesocotyl growth that pushes
the coleoptile and its enclosed

first leaf and shoot apex to the

soilsurface. In the light-grown
seedlings the mesocotyls are
only a few millimeters long.

Figure 8.4 Right: Corn (Zea

mays) leaf development. A
seriesshowing the leaf
enclosed in the coleoptile,
emergence from the coleoptile
in darkness where unrolling is
prevented, an artificially
unrolled leaf, and a similar
leaf grown in light, fully
unrolled with greening
completed.
 some Australian soils that are low in molybdenum only
140 gm of Mo0 3 per hectare, applied once every 10
years, increased pasture yield by 6 to 7 times.
Chlorine participates in the oxygen-evolving steps of
photosynthesis. This element is present in the soil solution
as the very soluble CI" Because of the very small
ion.

quantities required by plants and its almost universal

occurrence in soils and air (salt spray travels long
distances), chlorine is never intentionally added as a
component of fertilizer. In fact rainfall may contain enough
chlorine to satisfy the requirements of plants.

The Absorption of Minerals

As mentioned above, the minerals required by plants are

normally absorbed by roots from the soil solution. There
are exceptions: the pitcher plant and the Venus' flytrap,
for example, capture insects that, on decomposition,
Figure 5.10 Chlorosis of tomato (Lycopersicon esculentum) leaf release minerals that are absorbed by the specialized
caused by a deficiency of manganese in the nutrient solution. leaves that serve as traps. Epiphytes (plants that use the
aerial parts of other plants as supports for growth and are
chlorosis the young leaves become white or yellow with not themselves rooted in the soil) may obtain minerals by
prominent green veins, while manganese chlorosis gives trapping airborne dust and debris. Highly specialized
the leaf a mottled appearance. Spraying or dusting crops parasitic plants such as dodder and mistletoe sink
with as little as 22 kg of manganese sulfate per hectare modified stems known as haustoria into the vascular
often cures deficiencies. tissues of host plants and obtain their minerals in that
Copper is a constituent of certain enzyme systems such fashion.
as ascorbic acid oxidase and cytochrome oxidase, and of Finally, many plants form cooperative associations with
the compound plastocyanin, which is part of the electron soil microorganisms. The microorganisms bring in minerals
transport chain in photosynthesis. Deficiency causes from the soil. In this respect two classes of symbiotic
abnormalities of growth (Fig. 5.11), especially in plants associations can be distinguished. Mycorrhizae (Fig.
growing in marsh and peat soils. 5.12) are associations in which a fungus invades the
Molybdenum is important in enzyme systems involved in tissues of the roots and, through its own absorptive
nitrogen fixation and nitrate reduction. Plants suffering capabilities, extracts minerals from the soil and brings
from molybdenum deficiency can absorb nitrate ions but them into the host plant. The fungus obtains vitamins or
cannot then metabolize this form of nitrogen. If nitrogen in other organic materials in exchange. In this association
the form of ammonium is supplied to these plants, the the fungus takes over the role of root hairs and the
effects of molybdenum deficiency are less severe. For formation of root hairs may be strongly suppressed.
Mycorrhizal associations are especially common in forest

trees on nutritionally poor soils.

Another kind of association is seen in nitrogen-fixing

nodules (Fig. 4.64E). Here, bacteria that can convert N2
to NH 4 + dwell within thickened regions (nodules) of roots.
The most common instance is that of legumes (e.g., clover
and alfalfa) where the bacteria are species of the genus
Fthizobium. Without the bacteria, the plants would be
unable to use the nitrogen that is present as N 2 in the soil
air. In return, the bacteria obtain photosynthetic products
that serve their own growth as well as providing the
energy needed for nitrogen fixation.
But aside from these special cases, it is absorption of
dissolved soil minerals by roots that provides the plant
with its mineral supply. Since the mineral ions are
passively carried along with the absorbed soil water, the
path of their movement through the plant is similar to that
of water. The dissolved soil minerals can pass through the
endodermal layer of the root only by going through the
plasma membrane and into the endodermal protoplasts.
This gives the plasma membranes an opportunity to
Figure 5.11 Tomatoes (Lycopersicon esculentum) growing in
solution with allessential chemical elements except copper.
screen the passage of minerals in and out of the vascular
Leaves at left were sprayed with solution containing copper. system, between the soil and the main plant body. The

Mineral Ion Absorption

105
 Figure 5.12 Mycorrhizal roots of Pinus
virginiana. A, diagram showing cross
section of root with attached fungal
hyphae; B, roots with fungal sheath.

extent to which this screening actually discriminates consumes ATP, which must be produced by respiration
between desirable and undesirable mineral elements is with a consequent demand for 2 and glucose.

undetermined, because it is clear from examination of

plants grown in soils with toxic minerals that poisonous
elements may enter the vascular system.
Oxygen Absorption
The plasma membranes of most cells, including those of
root cells, contain active transport systems that
discriminate between various ions. Some ions may Oxygen is required for respiration by all the cells in the
+
accumulate (e.g.,K ) to high concentrations inside the plant. Photosynthesis produces more than enough oxygen
protoplasts; others may be pushed back into the wall to meet the needs of nearby cells during the daylight
+
space (e.g., Na ). These active transport systems may hours, but much of this oxygen leaks away to the
raise the concentration of a given ion within the atmosphere and little of it is transported within the plant to
protoplasts as much as 1000 times above the away as the root tips. Consequently,
regions as far leaves
concentration in the soil solution. This may be a great aid may absorb oxygen from the air at night, and roots usually
to the plant in acquiring rare elements. However, the require a supply of oxygen from air trapped in the soil.

active transport systems have a cost: their operation The soil contains a reservoir of atmospheric gases in

Figure 5.13 Effect of aeration on root growth in tomato

(Lycopersicon esculentum). Left, plants growing in complete
nutrient solution through which air was bubbled; right, plants
growing in same solution without aeration.

chapter 5 l
The Absorption and Transport Systems

106
the spaces between solid particles. This trapped air may plants contain in the stem and roots large communicating
exchange with the open air above ground the soil has a it air spaces. Thus, air absorbed into the leaves and stems
suitably loose texture. Heavy or compacted soils are may reach the living cells of the root in sufficient quantity
deticient in aeration, and cultivation may improve the air Bald cypress and certain species in tropical mangrove
supply of roots. Roots may absorb oxygen directly from swamps develop special root branches that grow upward
the soil gases (the oxygen dissolves in the water of cell ends are above the water level.
until their These special
walls); alternatively, gaseous oxygen may dissolve in soil rootbranches have a central core of loose tissue through
water, from which is taken up along with the water and
it which air moves downward to the submerged organs.
minerals. There seems to be no special transport
mechanism for trapping soil oxygen; entry is by diffusion
along the concentration gradient that results from the
Transport of Organic Materials
consumption of oxygen in respiration.

The amount of air in the soil depends not only on the

pore space but also on the water content of the soil. If
Within a living plant, substances are constantly moving
water occupies the pore space, air is forced out. Water- from one place to another. Although they are interrelated,
soaked (saturated) soil contains practically no air except we can detect four types of movement based on the rates
the amount dissolved in water. If the soil about the roots is atwhich movement occurs: (a) slow diffusion of molecules
continuously water-soaked, plants die because of and ions; (b) moderate movement of materials carried by
insufficient oxygen and possibly as a result of the cytoplasmic streaming in living cells; (c) more rapid flow of
accumulation of carbon dioxide. Most species of land material in the sieve tubes; and (d) very rapid conduction
plants will grow normally with their roots in a water of water and mineral solutes in the xylem.
solution, if it is well-aerated by bubbling air through it (Fig.

5.13). Evidence indicates that inadequate soil aeration

results in a reduction in the rate of water and mineral Diffusion and Protoplasmic Streaming
absorption.
Most land plants, including agricultural plants, will not The movement of water or of solute molecules and ions
long survive with the root system submerged in unaerated into a cell across the cell wall is a relatively slow process.
water or surrounded by a soil that is water-soaked. But Diffusion occurs along activity gradients and may take
some plants flourish under such conditions. Among them place through plasmodesmatal connections between cells
are rice, various swamp and marsh plants, and the bald as well as directly through the permeable cellulose walls
cypress (Taxodium distichum, Fig. 5.14). Almost all such of most Once a solute molecule or ion diffuses into a
cells.

cell, its movement may be increased many times as

rate of
it is picked up by the protoplasmic stream. Cytoplasm may
stream at rates of a few to several hundred millimeters per
hour.The highest rate of protoplasmic streaming that has
been measured was observed in the protoplasm of a slime
mold in which the cytoplasm was streaming at a rate of
486 millimeters per hour.
In terms of the dimensions of a single cell, cytoplasmic
streaming is frequently very rapid, since a substance may
be transported across the length of a cell in a matter of
seconds. In terms of the whole plant, however, this is a
very slow process. It would take days, for instance, for a
molecule in the leaf to be carried upward to the growing
shoot tip or downward to the roots. Figure 5.15 shows the
way which one solute may be moving by diffusion and
in

protoplasmic streaming in one direction through a series

of cells while another solute may be moving in another
direction. Diffusion and protoplasmic streaming are the
chief methods by which solutes move through living plant
tissues, except in the sieve tubes.

The Mechanism of Phloem Conduction

The phloem sieve tubes move organic materials rapidly
throughout the plant. That the phloem is the major path of
food transport is shown by the results of ringing
experiments. When a ring of bark is removed from a stem
down to the cambium (thus removing the phloem), the

Figure 5.14 Aerial "stump roots" of bald cypress (Taxodium carbohydrate content and especially the sugar
distichum). concentration of the sap in the leaf, bark, and wood above

Transport of Organic Materials

107
 Solutes corried in cytoplosmic streom. Rote fost.

Diffusion from cell to cell

through wall or plasmodesmota.
Rate slow.

Figure 5.15 Diagram illustrating how two solutes may be moving

in opposite directions by diffusion and protoplasmic streaming in
the same tissue

the ring increases after a few hours. Below the ring, the along the sieve tube from regions of high pressure to
concentrations of these solutes in the bark and wood regions of low pressure.
decreases. These conditions would not prevail if foods When we consider this mechanism, it appears that the
were moving downward in the xylem tissues direction of movement in the phloem must always be from
Translocation of material in the phloem may reach a sites where solutes are being produced to sites where
rate of 100 centimeters per hour. When this is compared solutes are being consumed. Given this general rule, the
to the much lower rates of protoplasmic streaming, it. is direction of flow in the phloem may change from time to
evident that normal protoplasmic streaming is insufficient time depending on the activities of the plant. For example,
to cause this rapid transport. In fact, the mature functional during its first growing season the sugar beet transfers a
sieve-tube members have highly modified cellular contents great deal of sucrose from the leaves to the roots via the
(Chapter 3) in which protoplasmic streaming does not phloem. Next spring the process is reversed; sucrose
seem to occur. moves from the roots to the young, growing shoot system.
The contents of the sieve-tube members of one sieve Although sucrose is usually the principal organic
tube make up a continuous liquid system. The pores in the substance carried in the sieve tubes, other organic
sieve plates connect the sieve-tube members into a materials such as hormones and amino acids, and even
continuous tube. According to one widely accepted theory inorganic ions, may also be carried.
(the mass flow theory), water and solutes move together The phloem transport may be controlled by the
rate of
as one mass in the sieve tubes along their length. rates at which solutes are added and removed. But there
The mechanism of mass flow can be illustrated by the are at least two other mechanisms that may affect the
flow of photosynthetic products from the leaves to the rates of movement. These are discussed below.
roots or to other sites of storage and consumption (Fig. Callose, a special polysaccharide with unusual linkages
5.16). The process begins with the production of glucose between the glucose subunits, is deposited on the sieve
molecules in the mesophyll cells of the leaves. From this plates by the action of enzymes. Other enzymes remove
site of high concentration, glucose diffuses to the bundle callose from the same locations. The thickness of the
sheath cells thatline the leaf veins. These cells contain callose layer on the sieve plates is therefore dynamically
enzymes assemble glucose molecules into the double
that maintained. The thicker the callose, the smaller the pores
sugar, sucrose. An active transport system in the sheath in the sieve plates. To the extent that pore size influences
cells or the adjacent sieve tubes then transfers the the rate of flow in sieve tubes, regulation of the callose
sucrose molecules into the protoplast of the sieve tubes. system may affect the flow rates. Recent experiments have
As in all active transport, this process consumes ATP. The shown that the enzymes of the callosesystem are highly
accumulation of sucrose molecules in the sieve tubes sensitive to mechanical stimuli such as vibrations or
decreases the water potential of the phloem sap, causing crushing. Such stimuli cause an extensive buildup of
water to move into the sieve tubes from neighboring cells callose in a matter of seconds, a response that is slowly
such as the xylem elements. This increases the reversed when the stimuli cease.
hydrostatic pressure in the sieve tubes. This pressure Also a network of protein strands occurs in the
pushes the phloem sap en masse away from the leaf conducting space each sieve-tube element. Under
of
blade, down the petiole, and into the stem of the plant At normal conditions of smooth flow in the sieve tubes, the
any site where solutes (chiefly sucrose) are removed, the protein network seems to be firmly anchored to the ends
water potential increases and water diffuses out of the of the cells and does not affect the transport of materials.
sieve tubes and into adjacent cells and the xylem. This But abrupt changes in the flow rate, such as occur when
results in a lowered pressure and movement of solution a stem is cut. cause the protein network to rip loose and

chapter 5 |
The Absorption and Transport Systems

108
Figure 5.16 Mass flow hypothesis of phloem transport, in
mesophyll cells, photosynthesis (1) produces glucose, which
moves by diffusion (2) into bundle sheath cells. Water follows by
osmosis (3). Enzymes (4) in the bundle sheath convert glucose to
sucrose, which moves by active transport (5) into the sieve tube.
Water follows by osmosis (6), raising pressure in the sieve tube.
Elsewhere, sucrose diffuses (7) from the sieve tube; water follows
(8) by osmosis, reducing the local pressure. The sieve-tube
contents flow en masse (9) from high to low pressure regions.

pile up against the nearest sieve plate, partially blocking 9. Cohesive and adhesive forces between molecules
the sieve pores. Both the callose and the protein network prevent the xylem water columns from breaking under
may be significant in limiting the loss of phloem sap from tension.
stems and petioles that have been injured. 10. When there is no transpiration, water may enter the
xylem by osmosis, creating positive root pressure and
Summary pushing water up the plant.
1. Water is absorbed chiefly from the soil by roots and 11. Roots do not seek water, though growth is generally

moves throughout the plant in the xylem. directed downward. Roots grow only in moist soil.
12. Carbon dioxide is absorbed from the air, reaching leaf
2. Some water is consumed in metabolism or retained in

growth but most of it is lost by evaporation cells through stomata and intercellular spaces.
(transpiration). 13. Gas exchange is limited by cork and cuticles and by
3. The force that moves water is the tendency of water to stomata.
move from regions of high water potential to regions of 14. Stomata are opened and closed by raising and
low water potential. lowering the turgor pressure of guard cells. The
4. The water potential of a solution is raised by adding changes in pressure are achieved by pumping
pressure and is lowered by adding solutes or materials potassium ions into and out of the guard cells. Water
such as wall fibrils that absorb water. follows by osmosis. Light exposure and C0 2 depletion
5. The water potential of air increases as its humidity stimulate the "tomata to open.

increases. 15. Minerals enter the plant through the roots, moving
6. If two solutions are separated by a membrane that will passively with the water flow except when they meet
permit water but not solutes to pass, water will move membranes, which may present a barrier. Carriers
through the membrane from a high to a low water control ion movements through membranes.
potential. This water movement is called osmosis. 16. The impermeable Casparian strip, in the walls of the
7. Because plant cells are enclosed in walls, water endodermis, prevents water and solutes from using the
uptake creates turgor pressure, which gives the cell a cell walls as an avenue for moving between the

rigid shape. Wilting results from the loss of turgor. vascular system and outer root tissues. This permits
8. The transpiration-pull theory states that evaporation of membranes to control traffic.
water from the leaf pulls water upward through the 17. Leaves may absorb oxygen from the air at night. Roots
xylem. usually absorb 2 from air in the soil.

Summary

109
18. Organic solutes move through the phloem between osmotic entry of water. Opposite movements occur at

regions of production, storage, and use. the low pressure regions.

19. The mass flow hypothesis states that the solution in 20. The rate of movement in the phloem may also be
the sieve tubes moves en masse from regions of high affected by a protein network that can plug the sieve
turgor pressure to regions of low pressure. The high pores when sieve tubes are injured; and it is also
source pressures are thought to result from active affected by enzymes that control the size of the sieve
pumping of sucrose into sieve tubes, followed by pores by depositing and removing callose.

chapter 5 |
The Absorpt on and Transport _Systems
i

ncT
 6 CHAPTER

6 photosynthesis

The living body is a metabolic system that maintains process — that "fixed carbon dioxide, was required.
air,"

itself by consuming energy-rich fuel molecules. Thus, it in the light use

could be said that green plants
Many organisms (animals, fungi, and most bacteria) carbon dioxide and produce oxygen.
depend on supplies of fuel molecules that they find ready- But what was the fate of the carbon in the C0 2 ? Ingen-
made in the environment. The green plants and a few Housz answered this question in 1796 when he said that
forms of bacteria differ from them in this respect: they can the carbon went into the nutrition of the plant; that is, it
trap the energy of sunlight to produce their own fuel became part of the organic matter. Overall, these
molecules, using common
low-energy molecules such as discoveries can be summarized by writing the equation for
carbon dioxide and water as raw materials. The capture of photosynthesis as:
light to produce fuels is known as photosynthesis.
Ultimately, thewhole living world depends on carbon dioxide light
oxygen +
photosynthetic organisms for its energy. Without them the + water green plants organic matter
supply of chemical fuels would soon be exhausted.
Therefore plants have a primary importance in the The nature of the organic material formed in

economy of nature; this gives photosynthesis a unique photosynthesis could be guessed by observing that starch
value among metabolic pathways. seems appropriate, It
accumulates in leaves during photosynthesis and is used
therefore, to discuss photosynthesis in a separate chapter, up when the leaves are kept in darkness. Starch is built
and in more detail than was done for other phases of from glucose molecules. Assuming that glucose is the
metabolism. direct product of photosynthesis, we can write and
balance the overall equation for the process as:

The Discovery of 6CO + cu

6H 2
2
,
/~\
light energy

greenp)antcell
' C 6 H 12 6 + 60 2
Photosynthesis
This formulation tells us that water is required in the
same proportion as C0 2 . More advanced methods using
Prior to the early seventeenth century, the general belief radioactive tracers have confirmed this relationship, but
was that plants derived the bulk of their substance from have also shown that other sugars besides glucose are
soil humus. This idea was overthrown by a simple formed early in photosynthesis. Hence it is probably more
experiment performed by the Flemish physician and realistic to write photosynthesis as in the formula below.
chemist Jan van Helmont, who planted a willow branch in
light
a box and supplied rainwater to the plant as needed. In CO, H,0 (CH 2 0) + O;
five years it grew to a weight of 169 pounds (77 carbohydrate

kilograms). The soil had lost only 2 ounces (57 grams);

consequently, van Helmont reasoned that the plant If we look at the overall equation for photosynthesis, we
substance must have come from water. This was a logical might logically propose that the oxygen liberated is
deduction, though we now know it was not entirely released from the carbon dioxide molecule. However, van
correct. Neil, working at Stanford University, made a comparative
Our knowledge of photosynthesis was further improved study of the photosynthesis that occurred in several
by a religious reformer, philosopher, and spare-time organisms belonging to different groups of plants. The
naturalist, Joseph Priestley. In 1772, Priestley reported green and purple sulfur bacteria are able to use hydrogen
that a sprig of mint could restore confined air that had sulfide instead of water. Sulfur instead of oxygen is
been made impure by burning a candle. The plant liberated.
changed the air so that a mouse was able to live in it. The
light
experiment was not always successful, probably because CO, + 2H,S (CH 2 0) + H 2 + 2S
Priestley, who did not know the role of light, did not
always have adequate light. Seven years later, Jan Ingen-
Some other bacteria and some algae can use hydrogen
Housz noticed that air was revitalized only when green
instead of water to reduce carbon dioxide:
parts of plants were in the light.

It was three years 1782, that a Geneva pastor,

later, in light

(CH 2 0) + H 2
Jean Senebier, discovered another important part of the

111
 Comparing these cases, van Neil concluded that a .v>
general equation tor photosynthesis should be written as
follows:

light
CO,
'2 + 2FUA (CH 2 0) H2 + 2A
carbon hydrogen carbohydrate
dioxide donator

The hydrogen donator H 2 A can be H 2 0, H 2 S, H 2 , or any

other substance capable of donating hydrogen to C0 2 in

the process of photosynthesis.

To test van Neil's hypothesis, a group of biochemists
supplied plants with water that contained the unusual
18 16
oxygen isotope, instead of the common isotope 0.
18
The oxygen liberated in photosynthesis contained 0. If

plants were supplied instead with carbon dioxide that

18 16
contained 0, along with water that contained 0, then
18
the oxygen liberated did not contain 0. This
demonstrated that van Neil was correct: the oxygen
liberated in photosynthesis comes from water.

The Chloroplast — Site of

Photosynthesis

Although the overall equation of photosynthesis seems

simple, the discovery that 2 comes from H 2 rather than
from C0 2 makes it clear that there are hidden Figure 6.1 Electron micrograph of isolated chloroplasts of Vicia
complexities. A good understanding of photosynthesis faba, showing one plastid with stroma and one plastid that has
lost its outer envelope and stroma, x2500.
requires that the process be taken apart and studied piece
by piece. Plant biochemists are still working on this task.
One successful approach has been based on the
examination of chloroplasts, the organelles that perform
photosynthesis. Chloroplasts are abundant in the cells of
leaves and other green parts of plants; the green color is
organelles according to their density or mass. Again, trial

due to light-absorbing compounds or pigments within the and error are needed to determine the most effective
chloroplast. It has been mentioned (Chapter 3) that each procedure. The final result is a solution in which
chloroplast has three principal parts: (1) a surrounding chloroplasts are suspended almost free of other
envelope consisting of a pair of membranes; (2) a liquid organelles. The isolated chloroplasts can produce
called stroma enclosed by the envelope; and (3) a
that is carbohydrate and oxygen if they are given light, carbon

set of flattened membrane-lined sacs called thylakoids dioxide, and water.

(grana and frets) that are embedded in the stroma (Fig. The electron microscope has shown (Fig. 6.1) that
6.1). chloroplast suspensions contain both intact chloroplasts
Chloroplasts are very fragile, but if properly treated they that have all their membranes, and damaged chloroplasts
can be removed good working condition from the cell.
in that have lost their envelope and stroma and retain only
In the test tube, their operation can be explored without the thylakoids. This important fact makes it possible to
the confusion that goes with having other kinds of isolate thylakoids and to study their activity apart from the
organelles in action at the same time. stroma.
The procedure for isolating chloroplasts or other The thylakoids prove to be the part of the chloroplast
organelles (fractionation) begins with a method for that contains the green pigment. They cannot convert C0 2
breaking open Fragments of tissue are placed in a
cells. into carbohydrates. But if they are given light, water, and
and organic compounds. The
dilute solution of salts an organic compound that can accept electrons, the
precise composition of the medium is important, and must isolated thylakoids can produce 2
It appears therefore
.

be determined by trial and error. The cells are broken by that the thylakoids are the light-capturing and 2
-

grinding with sand in a mortar, or with a kitchen blender. producing part of the chloroplast (Fig. 6.2). Because of

This releases some of the chloroplasts intact, though theirdependence on light, the reactions of the thylakoids
others are broken. The ground material is filtered to are commonly called the light reactions. The reactions of
remove large solid fragments. Then the suspension of the stroma, which do not use light, are known as the dark
organelles is spun in a centrifuge to separate the reactions.

chapter 6 Photosynthesis

112
 C

(CH 0)
2

•
•
-yrU":..- •
' •

CO,

Figure 6.2 The raw materials and products ot the thylakoids and
the stroma.

C2 H 3 CH 3
The Thylakoid (Light) ,.C, c
I

Reactions
V^N N^.„<r Magnesium-
containing
The basic process in the thylakoids is a light-driven flow of / head portion

electrons (Fig. 6.3). Light energy displaces certain

electrons from their equilibrium locations, and their
H—
\
Mg >-
movements back toward
H3 C / **N'
equilibrium are channeled so as
to build useful, energy-rich molecules. The ability of
C C^ A
X' JC — CH 3

moving electrons to do work should not seem unusual, c

because we see in everyday life that electrical currents
can be used to do many kinds of construction work. H-C-H H-C- c=o
I

The two primary working components of the thylakoid H-C-H C— OCH,

are (1) a light-capturing system, and (2) a system of II

electron transport pathways or chains.

• -C?20 n
H 39
Lipid tail
portion

The Light-Capturing System

Figure 6.4 Chlorophyll a molecule.
The light trap consists of pigment molecules that are firmly
attached to the membranes of the thylakoids. The most chlorophyll molecules they will bind. In turn, the proteins
important pigment is chlorophyll (Fig. 6.4), a complex join into larger working groups in the membrane.
molecule that contains a light-absorbing "head" and a There are several different varieties of chlorophyll. Each
long inert tail. The head is a flat array of four connected differs according to the small groups that are attached to
rings, with a magnesium ion held in the center. The tail, a the light-absorbing ring. One of these, called chlorophyll
hydrocarbon chain, may help to hold the chlorophyll a (Fig. 6.4), is present in all chloroplasts and seems to be
molecule in position. Within the thylakoid membrane,
chlorophylls bind to proteins of several kinds (Fig. 6.5).
These proteins differ in the number and types of

light traps

H,0

electron
transport chains

Figure 6.5 Chlorophyll molecules (a,b) are bound to protein

H+ ,
2 NADPH molecules (circles) that in turn form groups (chlorophyll-protein
Figure 6.3 Light traps and electron transport chains are the two complexes) within the thylakoid membrane. This illustration is
major functional systems of the thylakoid. (diagrammatic). highly schematic; the detailed arrangements are unknown.

The Thylakoid (Light) Reactions

113
 »-heot excited-state
orbitals

^ > heat

energy
of
orbital

photon
absorption

ground-state
orbitals

400 500 600 700

Wovelength of light (nanometers)
blue green yellow orange red
Figure 6.8 Absorption of a photon by a pigment causes an
electron to move from
a low-energy, ground-state orbital in the
Figure 6 6 Absorption spectrum of chlorophyll. The graph molecule to a high-energy, excited-state orbital. The energy
shows the fraction of received light that is absorbed when the difference is the excitation energy, equal to the photon energy
pigment is exposed to various wavelengths of light. The relation The excited electron may give up energy to other molecules as
between wavelength and color of light is also shown. heat or light (fluorescence).

seems to be essential for photosynthesis. When light interacts with pigment molecules, the beam
In addition to chlorophyll a, the chloroplast contains of light acts aswere composed of light particles, called
if it

other pigments, such as carotenoids, phycocyanin, photons. Each photon carries a definite amount
phycoerythrin, and other chlorophylls. These are called (quantum) of energy, and there is a definite relationship
accessory pigments because, although they may between the quantum size and the wavelength of the light:
contribute to light capture in photosynthesis, they are not longer wavelengths mean less energy per photon. In
universal in occurrence and the light they absorb must be absorbing light, a pigment molecule must accept a whole
passed on to chlorophyll a before it can be used. photon or nothing at all. Whether absorption can occur
The color of a pigment is an indicator of its light- depends on the energy carried by the photon as well as
absorbing ability. For instance, chlorophyll absorbs red the internal structure of the molecule. The electrons in the
and blue light very effectively but is a poor absorber of molecule move about constantly in a set of pathways
green and yellow light. The absorption behavior is best known as orbitals. Each orbital is associated with a
shown by means of an absorption spectrum (Fig. 6.6). In definite amount of energy (Fig. 6.8); to be in a certain
this figure the subjective quality of color is supplemented orbital an electron must have exactly the right amount of
with a more exactly measurable property of light, the energy. At normal temperatures and in darkness the
wavelength. The accessory pigments differ from electrons within a molecule tend to occupy the orbitals
chlorophyll a in absorption behavior (Fig. 6.7) and may with the least energy, those closest to the atomic nuclei,
improve the photosynthetic efficiency by capturing some toward which the electrons are drawn by electrical forces.
of the light that the chlorophyll would have allowed to In this low-energy condition the molecule is said to be in

escape. the ground state. Absorption of a photon causes one of

The absorption properties of chlorophyll account for the the electrons to shift to an orbital of higher energy. The
green color of leaves. Sunlight is a mixture of all colors exact amount of additional orbital energy needed to make
(wavelengths) of light. From this mixture chlorophyll must be supplied by the photon: a photon with
this shift
subtracts the red and blue portions, leaving green light to too much or too little energy will not be absorbed.
be reflected or to pass through the leaf so that it may be Here, then, is the primary event of light capture and the
seen. primary conversion of light energy into chemical energy.
The pigment molecule after absorption is said to be in an
excited state; the pattern of electron movement within the
phycoerythrin molecule has been thrown out of equilibrium. We say that

phycocyanin
the energy has been converted to excitation energy.
light

Pigment molecules remain in the excited state only for a

small fraction of a second before giving up their extra
energy and returning to the ground state (Fig. 6.8). In
photosynthetic pigments, there are many closely spaced
orbitals, and the excited electron can dispose of its extra
energy by stepping through these orbitals, donating the
difference in energy to other molecules that collide with

400 500 600 700 the excited pigment. The effect is to increase the motion
Wavelength of light (nanometers) of the surrounding molecules. This extra motion appears
Figure 6 7 Absorption spectra of three accessory pigments. as heat, which is a waste to the system. Also, at any

chapter 6 J Photosynt hesis

114
moment the excited molecule may dispose of any called resonance transfer. Here the full quantum of
remaining excitation energy by giving oft a new photon. excitation energy (*) is passed from one pigment molecule
The emitted light is known as fluorescence and it, too, is to another without any loss.
a waste to the system. resonance
** *
pigment? + pigment^ , ranste
," pigment, + pigment!

In this event the pigment that originally had the excitation

Reaction Centers energy drops to ground state and the recipient pigment
its

Light absorption
becomes There is no transfer of electrons, no
excited.
is an organized
useful only if there is
emission of photons or chemical interaction between the
system available to exploit the excitation energy of
pigments. The exact mechanism of transfer is not known,
pigments before it can be converted to heat or fluorescent
but we do know that efficient transfer depends on the
light. This depends on the action of specially placed
pigments being bound close together.
chlorophyll a molecules that can give up an electron to an
acceptor compound when they receive excitation energy.
By contrast, ordinary pigments — including most Electron Transport
chlorophyll a molecules — do not ionize on excitation.
The ionizable chlorophyll a molecules occur at sites in
The events that follow the arrival of excitation energy at
the membrane called reaction centers. The detailed
the reaction centers are diagrammed in Fig. 6.10. Overall,
arrangement is not known, but each reaction center seems energy drives a flow
light of electrons along a system of
be bound to an antenna system, which is built of many
to
carriers from water to NADP + The . carriers are bound to
chlorophylls bound to proteins. The reaction center and its
membrane between
the the reaction centers. Though their
antenna form a photosystem. Two kinds of photosystems precise arrangement not known, the carriers seem to be
is
(I and II) are known. Each thylakoid has several hundreds +
organized so that the electron flow causes H to move
of them (Fig. 6.9). Photosystem I seems to contain six
from the stroma to the space within the thylakoid. This is
protein molecules with about seven chlorophyll a
analogous to charging a battery.
molecules per protein. One or two of the chlorophylls are +
The resulting difference in H concentration across the
at the reaction center. Other compounds may also be membrane represents a store of energy that is thought to
present in the photosystem, such as the unknown
drive the formation of ATP. According to current ideas, the
compound (Z, in Fig. 6.10) that accepts electrons from enzymes that form ATP are bound to the thylakoid
chlorophyll. Photosystem II is not as well known, but is
membrane and are arranged so that the formation of ATP
thought to be broadly similar to Photosystem + -
releases H to the stroma and OH to the space within
I.

As shown in Fig. 6.9, the thylakoid also contains units +

the thylakoid. The OFT combines with H to form H 2 0.
called light-harvesting complexes. These groups of +
These events would decrease the H difference across
pigment-bearing proteins do not have reaction centers, but the membrane, "discharging the battery" to form ATP.
collect light energy to feed the photosystems (mainly
Electrons move spontaneously along the electron
photosystem II). The light-harvesting complexes contain all transport chain because each carrier in the chain has a
the accessory pigments and some chlorophyll a. greater tendency to capture and hold electrons (a greater
electron affinity) than the carrier before it. This is shown in

The Transfer of Excitation Energy

Excitation energy finds its way from the antenna and light-

harvesting pigments to the reaction centers by a process

stroma space
photosystem
complex

photosystem II

complex

light-harvesting
complex space within thylakoid

Figure 6.9 Proposed model for the arrangement of

photosystems in the thylakoid membrane. Each complex is a
group of proteins to which chlorophyll molecules are bound. The
Photosystem and Photosystem complexes include reaction
I II

centers. Adapted from P. A. Armond, L. A. Staehelin, and C. J.

Arntzen, J. Cell Biol. 73:400-418 (1977).

The Thylakoid (Light) Reactions

115
 STROMA
+
H NADP + NADPH

SPACE WITHIN THYLAKOID

+
Figure 6.10 Proposed flow of electrons (green arrows) and H
in the thylakoid membrane. Overall, light is thought to drive a flow
+ across
of H the membrane in one direction and a flow of
electrons in the opposite direction. RC and RC are reaction
I II

centers; circles and PQ represent compounds that take part in

electron transport chains. Sizes of the parts are not drawn to
scale and locations are schematic.

Fig. 6.1 1 . Here the carriers with the greatest electron Chlorophyll supplies electrons for the transport chains.
affinity are lowest on the vertical scale. The oxidation- But ground-state chlorophyll, and the chlorophyll ion, have
reduction potential shown on this scale is an exactly a great tendency to hold their electrons. Only the excited,
measurable property of the carrier that is directly related un-ionized chlorophyll molecule can give up an electron to
to the electron affinity: the higher the potential, the lower the chain.
the electron affinity of the carrier. As Fig. 6.11 shows, the chain of carriers that starts with

-0.6

acceptor Z

ferredoxin ^~^
'

flavoprotein

NADI

photon
capture
s o.o

plastoquinone

photon
>
cytochrome f >^
capture plastocyanin
^N
chl

(photosystem I)

H 2 0.

1.0 chl

(photosystem II)

Figure 6.11 Oxidation-reduction potential diagram for the

electron carriers and Photosystems and II. The arrows show the
I

path of electrons in photosynthesis. Carriers high on the scale

tend to give up electrons to lower carriers; electrons lose energy
as they move to carriers that are lower on the potential scale.
Symbols: chl = chlorophyll a in the ground state;
chl* = chlorophyll a in the excited state.

chapter 6 |
Photosynthesis

116
Photosystem I ends by giving its electrons to the mobile This ATP formation in the thylakoids is called
compound NADP + which , picks up H
+
from the photophosphorylation. The ATP is released to the stroma,
surrounding solution at the same time: where it supplies part of the energy needed for sugar
production.
NADP + + H + + 2 electrons NADPH Since ATP carries more energy than the raw materials
from which it was made, clearly some process in the
The chlorophyll of Photosystem is left as a positive ion, I

thylakoids must have supplied the needed energy. At

which cannot accept photons or donate more electrons to
present the details are not known. We do know that under
the chain. These chlorophyll ions take electrons from the
normal conditions the energy is provided by the movement
chain fed by Photosystem II. This restores neutrality and
of electrons between Photosystems and (Fig. 6.12). I II

allows the chlorophyll of system to absorb another I

One view is that electron flow causes a gradient of H +

photon.
across the membranes, and ATP formation draws energy
Meanwhile, the chlorophyll of Photosystem II has been +
by discharging the H gradient (p. 1 15). In any case, we
left as an ion with a high electron affinity. The system II
say that ATP formation is somehow coupled to the flow of
chlorophyll ions get electrons from water molecules (Fig.
electrons. This process is known as noncyclic
6.1 1). Hydrogen ions and 2 are released from water photophosphorylation.
when the electrons are removed. The water molecule has If the supply of NADP + runs low, the electrons moved
a great electron affinity, but the chlorophyll ion has a still
by Photosystem spill into the chain from system
I and II

greater affinity. The precise mechanism that transfers

find their way back to the original chlorophyll of system I,

electrons from water to chlorophyll ions unknown. is

forming a closed cycle of electron movement that is driven
However, we do know that manganese ions and at least by excitation energy (Fig. 6.12). This cyclic electron flow
one enzyme are involved. may also cause coupled ATP formation. ATP formation in

this cycle is termed cyclic photophosphorylation.

Energy Storage by the Thylakoid

There is a close relation between the oxidation-reduction
Summary of the Thylakoid Reactions
potential of a carrier and the energy of its electrons: an
The events in the thylakoids are summarized in Fig. 6.12.
electron has more energy is attached to a carrier that
if it
The useful high-energy products are NADPH, 2 , and
is high on the potential scale. One can view the carriers in
ATP. Light acts by driving a flow of electrons from water
Fig. 6.1 1 as a series of stations that a moving electron
to NADP + The . removal of electrons splits water to release
must pass; the vertical position of the carrier represents +
2 and H . Finally, some of the energy flow during
the energy the electron has at each point in its travels.
electron transport is captured in the coupled formation of
With this view, we see that excitation gives the electron an
the ATP.
initial dose of energy; with each transfer along the chain

the electron loses some of its energy. The final product,

NADPH, carries electrons that
the original energy. NADPH is
still retain a large fraction of
a high-energy product of
C0 Reduction and the
2

the thylakoids that can carry high-energy electrons and Formation of Sugar
hydrogen to the stroma for use in building sugar
molecules. Having given up its load, the "empty" NADP
+ (Dark Reactions)
molecule can return to the thylakoid for more electrons.
2 is also a
high-energy product of the thylakoids. The
Two methods have been especially
sensitive analytical
oxygen atoms in 2 have a high affinity for electrons and
useful in showing the sequence of reactions by which
will take them from other molecules if the opportunity
sugar is built from C0 2 These new methods were the use
.

arises. The role of 2 in driving

respiration has already
of carbon dioxide in which the carbon was radioactive and
been discussed in Chapter 2.
the use of a special process, chromatography, with which
The thylakoids also condense ADP and inorganic
the investigator could easily and accurately separate
phosphate (Pi) to form the energy-rich molecules of ATP:
minute amounts of different organic compounds from one
v
ADP + Pi + energy ATP + H 2 another.

cyclic

\ e
xNADP + + H +
H,0
photosystem non-cyclic photosystem
1 ' 6
-
r a \ NADPH
1

chlorophyll e chlorophyll
hCO, ADP ATP
+ Pi + H O
Figure 6.12 Noncyclic photophosphorylation (ATP formation)
draws energy from the flow of electrons between Photosystems I

and II. In cyclic photophosphorylation the electrons start and end

in Photosystem I.

CO, Reduction and the Formation of Sugar (Dark Reactions)

117
 end of paper is in H;0

waih with butanol propionic acid

concentrated
and placed
J
on filter paper

partial separation of compounds

boiling alcohol

rotate paper

paper rotated

wash with phenol water

final separation of compounds

Place on x-ray film in dark. Develop film after two weeks.

Radioactive compounds on the paper are indicated by
darkened areas on the film.

final radiograph on the x-ray film

spots on film matched with paper

and compounds eluted and identified.

Figure 6 13 Method for labeling and isolating photosynthetic

14
products. Illuminated green cells are exposed to C0 2 then ,

dropped into boiling alcohol to extract the products that have

14
incorporated C. Chromatography follows as shown in the
diagram and described in the text. An autoradiograph completes
the process; identity of materials in the radioactive spots is
determined by comparison with the positions of spots made when
known compounds are subjected to similar chromatography.

chapter 6 |
Photosynthesis

118
 The problem confronting physiologists was to determine ADP, Pi
phosphoglyceraldehyde
the sequence of compounds through which carbon atoms NADPt,
taken into the leaf pass during the synthesis of sugar. NADPH, H +.
14
When the radioactive isotope of carbon ( C) became ATP
available, scientists treated leaves with carbon dioxide that phosphoglyceric acid
14
contained radioactive carbon ( C0 2 ) and determined the fructose 1,6-diphosphate

carbon compounds formed. This may easily be done by

the method outlined in Fig. 6.13. The mixture of qq ^S\ (carboxydismutase)

substances extracted from the leaf is first separated by

chromatography: A spot of the extract is placed on one ribulose 1 ,5-diphosphate
other

corner of a sheet of filter paper. One edge of the paper carbohydrates

is ADP-
then dipped in a carefully chosen solvent solution. The
solvent flows across the paper, carrying various materials ATP ribulose phosphate

from the leaf extract along at different and characteristic

rates. This converts the original spot of material into a row Figure 6.14 Diagram showing some steps in the carbon cycle of
of spots, each spot containing a different collection of photosynthesis, the Calvin, or C cycle. ,
3
molecules. The paper is now dried and rotated 90
combines with ribulose diphosphate, a five-carbon-atom
degrees; then another edge is dipped into a second
sugar phosphate continually being produced in the cell.
solvent solution. The second solvent differs from the first
Two molecules of phosphoglyceric acid (PGA), a three-
and causes solutes on the paper.
to migrate differently
carbon-atom compound, are produced.
Again as the solvent moves, it draws each spot out into a
2. Two molecules of PGA are reduced to form two
series of spots, each containing only one or a few
molecules of a three-carbon-atom sugar phosphate,
substances.
phosphoglyceraldehyde. The energy that drives this
By comparing the positions of spots on the finished
reaction comes from NADPH and ATP, and energy from
chromatograph with the positions reached by known
these molecules is stored in the newly formed triose
substances on similar runs, we can identify the substances
sugar. ADP and NADP are regenerated.
in each spot.
3. Two triose phosphates may be combined to form a six-
The method is completed by placing the chromatograph
carbon-atom sugar phosphate, fructose diphosphate.
in contact with a sheet of photographic film for several
By this process the plant has essentially added one
days. Spots that contain radioactive carbon expose the
C0 2 molecule to a five-carbon-atom sugar to produce
film. The developed film therefore shows which of the
14
one molecule of a six-carbon-atom sugar.
materials in the extract were produced from C0 2 in
4. As this process continues, some of the fructose
photosynthesis.
phosphate may be transformed through other reactions
A group of scientists at the University of California under
into other carbohydrates, including sucrose and starch.
the direction of Melvin Calvin were particularly successful
5. Some of the fructose phosphate molecules are used to
in using this technique for working out the early carbon
form new molecules of ribulose diphosphate, the
pathways of photosynthesis. Calvin eventually received the
compound that accepts C0 2 in step 1 Thus the whole .

Nobel Prize for this work.

process forms a cycle of reactions, with C0 2 from the
To introduce radioactive carbon into the photo-
air and hydrogen from water entering the cycle, and
synthesizing cells, these investigators placed an algal
various sugars being produced.
culture in a glass flask in the light, exposed the cells to
radioactive C0 2 , and killed the cells in boiling alcohol. By This carbon cycle of photosynthesis is often called the
14
varying the time after the plant was exposed to C0 2 ,
Calvin cycle or the C3 path. It is almost universally present
they could determine the first compounds synthesized in photosynthetic plants.
14
during exposure to C. There is a special added path of carbon fixation that is

If photosynthesis proceeds for an hour or so an in common in plants that have evolved in regions of high
atmosphere containing radioactive carbon dioxide, most of light intensity, high temperature, and drought. This is the
the labeled carbon will be found in carbohydrates (sugar C4 pathway, also called the Hatch-Slack pathway after the
or starch). If photosynthesis is stopped after only a few two Australian workers who studied it most extensively.
seconds, most of the labeled carbon is found in a three- In this pathway, C0 2 is first taken up by an enzyme
carbon acid containing phosphorus, phosphoglyceric acid called PEP carboxylase. This enzyme has a stronger
(PGA). affinity for C0 2 than does carboxydismutase. It adds C0 2
PGA is thus an intermediate between carbon dioxide to the three-carbon compound phosphoenolpyruvate
and sugar. By stopping photosynthesis at increasingly (PEP), to form oxaloacetic acid, a four-carbon-atom
longer intervals (from a few seconds to several minutes), compound. Later, oxaloacetic acid may be converted to
investigators have found that many carbon compounds two other C 4 acids, malic and aspartic. These acids may
become labeled with radioactive carbon. accumulate to high levels. They seem to serve chiefly as a
has been possible by means of such experiments to
It means of capturing C0 2 and storing it, because they later
draw up an outline representing the various reactions that release their C0 2 which is taken up by carboxydismutase
,

occur (Fig. 6.14). and used in the ordinary C 3 cycle to produce sugars. Thus
Here are the key points of this carbon cycle: the C 4 path is an addition, rather than an alternative, to
Catalyzed by the enzyme carboxydismutase, C0 2 the Calvin cycle.

CQ 2 Reduction and the Formation of Sugar (Dark Reactions)

119
 Under the hot and dry conditions ot the tropics, plants of high-producing grains. Unfortunately, these strains
that have the C 4 path are much more etticient at require high levels of fertilizer application — an expensive
extracting C0 2 from the air and building sugars than and pollution-creating practice.
plants that can use only the C 3 path. This efficiency is One aspect of photosynthesis that seems a likely point
reflected in the economic value of sugar cane, which has of attack for improving productivity is the process known
the C 4 path. Maize (corn) and sorghum are also C 4 plants, as photorespiration. In some plants, photorespiration may
among other examples from over 100 genera. These reoxidize and release as much as 30% of the C0 2 that has
include both dicots and monocots, which suggests that been incorporated in photosynthesis. Photorespiration
the C 4 pathway has arisen independently many times in differs from ordinary respiration in that it does not
evolution. generate ATP. At the present time it seems to be the result
of a flaw in the photosynthetic system. It involves the
one of the early products of the
oxidation of glycolic acid,
carbon reduction system. Investigations are currently
Efficiency of Photosynthesis underway to determine whether new strains of crop plants,
or new growing conditions, can be found that will have
lower rates of photorespiration and therefore better
Although the green leaf is the major organ used in the retention of the products of photosynthesis.

production of man's food, it is not particularly efficient in

utilizing the sun's energy. We speak of the efficiency of a

machine, such as a diesel engine, a gasoline motor, or an The Environment and

electric motor. We calculate the energy value of fuel used
by the machine and compare it with the machine's energy
Photosynthesis
output. Much of the fuel's potential energy is lost. The
ratio of energy outgo to energy intake represents The principal external conditions that affect the rate of
efficiency. Of the total radiant energy that falls upon green photosynthesis are (a) temperature; (b) light intensity,
leaves, about 80% is absorbed. Of the remaining 20%, a quality, and duration; (c) carbon dioxide content of the air;
part is reflected from the leaf surface and a part passes and (d) mineral elements in the soil.
through the leaf. Part of the radiant energy absorbed is
changed to heat and raises the temperature of the leaf; a
large part of the energy absorbed is used in transpiration; Temperature
the remainder of it is utilized in photosynthesis and stored
in carbohydrate molecules. Only about 0.5 to 3.5% of all Plants of cold climates carry on photosynthesis at much
the light energy that falls on a leaf is used in lower temperatures than do those of warm climates. The
photosynthesis. Although this is a very low percentage process is known to occur in certain evergreen species of

compared to machine efficiency, the supply of solar cold regions, even at temperatures below 0°C. Algae in

energy is continuous and abundant. the water of hot springs may carry on photosynthesis at a

However, there is reason to believe that the leaves of temperature as high as 75°C. Most ordinary temperate-
higher plants, under the most favorable conditions, should climate plants, however, function best between

be about 10 times more efficient in converting solar temperatures of 10 and 35°C. there is adequate lightIf

energy into chemical energy by photosynthesis. Research intensityand a normal supply of carbon dioxide, the rate
will be needed if we are to raise the photosynthetic of photosynthesis of most ordinary land plants increases

efficiency of plants growing out of doors to the efficiency with an increase in temperature up to about 25 °C; above

that is theoretically possible. The large research effort that

has been devoted to photosynthesis also aims for a basic
understanding of how nature converts light energy into
chemical energy. With this information we might someday
Light limiting
devise artificial alternatives to photosynthesis as a way to high light intensity

meet some of the world's food problems. And, too, some

of the lessons we learn about the plant's use of solar
energy could prove useful in our search for alternatives to
nuclear and fossil fuels. Light limiting

The productivity of plants is determined partly by the low light intensity

environment and partly by the hereditary characteristics of smperature limiting

low light intensity
the plant (e.g., its and metabolic capabilities).
structure
Thus one way to solve the problem of producing enough
food for the world population is to breed highly productive
approach already has been so
varieties of plants. This
successful in some areas (e.g., the development of highly Temperature increasing >
productive types of rice and wheat) that we speak of the Figure 6.15 Graph showing interaction of light and temperature
"green revolution." The importance of this approach was on the rate of photosynthesis. When light is limiting, the rate of
photosynthesis isindependent of temperature. When temperature
recognized by the recent award of the Nobel Prize to is limiting, the rate of photosynthesis is independent of light
Norman Borlaug, who succeeded in breeding new strains intensity.

chapter 6 |
Photosynthesis

120
this range there is a continuous fall in the photosynthetic 1. The living cells of all plants (both green and nongreen)
rate as the temperature is raised. At these higher and of all animals release carbon dioxide in the
temperatures, the length of exposure is of importance. At respiratory process (Chapter 2)
a given constant high temperature (for example, 40 °C), 2. The dead bodies of plants and animals, and the
the rate of photosynthesis decreases with time. excretions of animals, contain large quantities of carbon
Under conditions of low light intensity, an increase in and other elements in the form of organic compounds;
temperature beyond a certain minimum will not produce in the decay of these compounds, resulting from the
an increase in photosynthesis (Fig. 6.15). These activities of bacteria and fungi, large quantities of
conditions may occur in the winter in greenhouses. If the carbon dioxide are released to the atmosphere.
temperature is raised too high, the plants will suffer 3. Carbon dioxide is also added to the atmosphere when
because the has not been changed
rate of photosynthesis wood, coal, oil, gas, or any other carbon compound
but respiration has been increased by the higher burns.
temperature. 4. Carbon dioxide is released to the atmosphere from
mineral springs and volcanoes.
5. The oceans are important reservoirs of dissolved
Light carbon dioxide, and from them carbon dioxide probably
escapes whenever its concentration in the atmosphere
In discussing the effect of light upon the rate of
decreases.
photosynthesis, we must consider three elements:
(a) intensity, (b) quality (wavelengths), and (c) duration. If and temperature are favorable, the
light intensity
Given the proper temperature and sufficient carbon carbon dioxide atmosphere frequently limits the rate
of the
dioxide, carbohydrates produced by a given area of leaf of photosynthesis. This may be particularly true in
surface increase with increasing light intensity up to a greenhouses that are kept closed in the winter. Under
certain point (optimum light intensity), after which their these conditions, the carbon dioxide in the air may be
production decreases. reduced far below the 0.03% average.
Intense light appears to retard the rate of photo- It has been determined experimentally that at usual

synthesis. The usual light intensity in arid and semi-arid temperatures and light intensities, an artificial increase of
regions is well above the optimum for photosynthesis in carbon dioxide up to a concentration of 0.5% may result in
many plants, especially introduced crop plants. In these an increased rate of photosynthesis, but only for a limited
regions on days when the sky is overcast, the light period of time. It appears that this high level of carbon
intensity is probably closer to the optimum for photo- dioxide is harmful to plants; after 10 to 15 days' exposure,
synthesis than on clear, sunny days. Leaves on the the plants show injury.
surface of plants receive light of greater intensity than
those beneath that are shaded. Therefore, some of the
leaves receive light ofoptimum intensity, whereas others
Minerals
may receive light either above or below the optimum.

The chemical formula of chlorophyll a is C 55 H 72 5 N 4 Mg

Carbon Dioxide and of chlorophyll b is C 55 H 70 O 6 N 4 Mg. The synthesis of

chlorophyll, therefore, depends on a supply of nitrogen
The carbon dioxide utilized by land plants is absorbed by and magnesium, both derived from salts in the soil.
the leaves from the atmosphere. Moreover, chlorophyll is not formed unless iron is
Let us consider the problem in this way: If all the available, although this element is not a component of the

chlorophyll-bearing cells of the plants of the world are chlorophyll molecule. Leaves of plants deficient in

constantly taking carbon dioxide from the atmosphere nitrogen, magnesium, or iron are pale and yellow, a

during daylight, the quantities of this gas used must be condition termed chlorosis. This abnormal condition may

enormous, and there must necessarily be processes in also be caused by other factors, but whenever it does

nature that are continually replenishing this supply. It is occur, the rate of photosynthesis is lowered.
known that the amount
carbon dioxide in the air is low
of
(0.03%) and that it remains fairly constant. It has been Summary
estimated that a hectare of corn (25,000 plants) during a
growing season of 100 days will accumulate 6350 kg of 1. Photosynthesis is the primary energy-storing process of
carbon; all this carbon is derived from the carbon dioxide life. In it, light energy is stored as chemical energy in

of the atmosphere. It would require 23,300 kg of carbon organic compounds.

dioxide to furnish this quantity of carbon. These estimates 2. The raw materials of photosynthesis are carbon dioxide
serve to emphasize the fact that enormous quantities of and water; the products are sugar and oxygen (0 2 ).
carbon dioxide are used in the photosynthetic process of 3. Chloroplasts are organelles that perform photosynthesis
green plants. in green plants.
Obviously, the amount of carbon dioxide is limited. The 4. The energy of photons (light) removes electrons from
present atmospheric supply would be used up in about 22 chlorophyll. Some of the energetic electrons, plus
years were it not constantly being renewed. Through hydrogen ions from water, are taken up by NADP to
several natural processes carbon dioxide is continually form NADPH. The rest of the photoelectrons return to
released to the atmosphere: chlorophyll ions after passing along chains of electron

Summary

121
 carriers. During this transport, some of the electron compounds are used in a series of reactions that
energy is transferred to produce ATP. incorporate C0 2
and produce molecules of sugar. ATP
The electrons lost by chlorophyll in making NADPH are is used as an energy source. These steps are called the

replaced by electrons from water. This splits water dark reactions. Two major pathways of carbon fixation
+
molecules and 2
to release H . are the C 3 and C 4 pathways.
The reactions that require light to form ATP, NADPH, 8. The photosynthetic process captures only a small
and 2 are called the light reactions. They occur in fraction of the available light energy.
thylakoids of the chloroplasts. 9. Photosynthesis may be limited by the C0 2 supply, light,

In the stroma of the chloroplast, the H and electrons of temperature, minerals, and the hereditary efficiency of
NADPH are transferred to organic compounds. These the plant.

chapter 6 |
Photosynthesis

122
 CHAPTER

7 flowers, fruits
and seeds

PART ONE affected

angiosperm
in these ways and thus are important parts for
classification.
In addition, flowers have a high esthetic value, and the
The Flower resulting fruits and seeds are of major importance in food
production.
The essential steps of sexual reproduction, meiosis and
fertilization, take place in the flower. The complete sexual

The flower initiates the sexual reproductive cycle

Anthophyta (the flowering plants) and, in so
all
in cycle involves (a) the production of special reproductive
cells following meiosis, (b) pollination, (c) fertilization,
doing, it terminates the growth of the shoot bearing (d) fruit and seed development, (e) seed and fruit
the flower. With some plants —
annuals, biennials, and dissemination, and (f) seed germination. The seed

—
some perennials death follows flowering and seed set. completes the process of sexual reproduction in the
With most perennials the capacity for vegetative growth is angiosperms, and the embryo in the seed is the first stage
regained after sexual reproduction and the development of in the life cycle of new individuals.
the next generation. In woody perennials, provision is

always made for continued vegetative growth, through

mixed buds or associated shoot buds following flower Flower Structure
production. The function of the flower is to facilitate the
important events of gamete (haploid reproductive cell)

formation and fusion. A typical flower is composed of four whorls of modified

Of all the characteristics of flowering plants, the flower leaves: (a) sepals, (b) petals, (c)stamens, and (d) a
and fruit are the least affected by changes in the carpel or carpels, all attached to the modified stem end
environment. For instance, we have seen that leaf shape is that supports these structures, the receptacle (Fig. 7.1).
influenced by age, light, water, and nutrition. The basic The sepals enclose the other flower parts in the bud
morphology and anatomy of flowers and fruits are not and are generally green. All the sepals taken collectively

receptacle

Figure 7.1 A diagram of a longitudinal section of a flower of

Christmas rose (Helleborus). The perianth consists of two similar
whorls; there are numerous stamens arranged in a spiral on a
cone-shaped receptacle. Five separate carpels form the central
whorl of floral parts.

123
 carpels
gynoecium

anthers

Figure 7.2 The essential floral organs. A, Magnolia, showing

many separate stamens and separate carpels arranged in a spiral
on the receptacle; B, regal lily (Lilium regale), showing six
stamens with anthers and filaments, and three carpels in a single
pistil with stigma, style, and ovary; C, carpels suggestive of
foliage leaves. Sterculia plantanifolia: dehiscence of matured
ovary showing seed attached to the margins of the five leaflike
carpels, x%.
carpels

constitute the calyx. The petals are usually the

conspicuous, colored, attractive flower parts. Taken
together, the petals constitute the corolla.
The stamens form a whorl, lying inside the corolla. Each
stamen has a slender stalk or filament at the top of which
is an anther, the pollen-bearing organ. The whorl or

grouping of stamens is called the androecium (Fig. 7.2).

The carpel or carpels comprise the central whorl of
modified floral leaves (Fig. 7.2). Collectively, the carpels carpel
margins
are spoken of as the gynoecium. Each individual
structure in the gynoecium is referred to as a pistil. A pistil

may be composed of a single carpel, or of several united

carpels in the center of the flower.
There are generally three distinct parts to each pistil:
(a) an expanded basal portion, the ovary, in which are
borne the ovules; and (b) the style, a slender stalk
supporting (c) the stigma (Fig. 7.1).
The term perianth is applied to the calyx and corolla
collectively. It is frequently used to describe flowers, such
as the tulip (Tulipa), in which the two outer whorls are
similar in appearance. The individual parts of such a
perianth are called tepals. Carpels and Stamens as
Sometimes individual flowers or compact clusters of Modified Leaves
flowers will have a whorl of small leaves or bracts
subtending them. A collection of bracts subtending flowers The question arises as to why carpels and stamens are
is called an involucre. considered modified leaves. One reason is because the

chapter 7 I
Flowers, Fruits and Seeds

124
 pistil

points of
ovule attachment

Figure 7.3 Primitive pistil and stamen trom living plants in

southeast Asia. Apistil of Drimys piperita; B, cross section of
pistil (dotted line); C, pistil laid open; D, stamen of Degeneria

vitiensis; E, cross section of stamen.

early developmental stages of floral parts closely

resemble those of leaves. The second reason, also
Variations in Floral Structure
circumstantial and speculative, concerns the shape of

spore-bearing leaves in primitive angiosperms. If we were General Description

to examine the stamens of primitive flowers such as
Magnolia or Degeneria (Fig. 7.3) (see Chapter 16), we Complete and Incomplete Flowers
would see that the shape of the stamen is very leaflike. In The parts of a typical flower — sepals, petals, stamens, and
addition, the folded open carpels of primitive angiosperms carpels — are all attached to the receptacle. A flower with
are leaflike in appearance. a young pea pod, which is a
If
all four sets of floral leaves is complete
said to be a
single carpel, is opened carefully along its ventral suture flower. An incomplete flower lacks one or more of these
(toward the axis), the margins may be folded back, four sets. For example, there are (a) flowers that partially
showing the seeds, which have developed from ovules, or completely lack a perianth (Figs. 7.4A.B), (b) flowers
along the margin of a leaflike carpel (Fig. 7.1 ID). with carpels but no stamens, and (c) flowers with stamens
The relationship between carpels and stamens is even but no carpels (Fig. 7.5).
more striking in Sterculia platanifolia, because in this plant

there are five simple united only by their stigmas

Perfect and Imperfect Flowers
pistils

(Fig. 7.2C). When these stigmas mature, they open to Unisexual flowers are either staminate (stamen-bearing)
show five very leaflike carpels that bear seeds along their or pistillate (pistil-bearing). Unisexual flowers are said to
margins. Carpels may thus also be compared with leaves. be imperfect, whereas bisexual flowers are perfect. When
The evolutionary steps still remain speculative. staminate and pistillate flowers occur on the same

Variations in Floral Structure

125
 pistillate flowers

Figure 7.4 Flowers with perianth parts lacking. A, spathe of

calla (Zantedeschia aethiopica), sepals and calyx absent; B,
lily

flower of Clematis, petals absent and sepals prominent.

.4r»*

stigma

style

ovary

Figure 7.5 Imperfect flower. A, pistillate and B, staminate

flowers of squash (Cucurbita), x '/2-

chapter 7 l
Flowers, Fruits and Seeds

126
individual plant, as they do in corn (Zea), squash made up of petals of similar shape that radiate from the
(Cucurbita), walnut (Juglans) (Figs. 7.5AS, 7.6, 7.7), and center of the flower and are equidistant from each other.
many other species, the plant is said to be monoecious. Such flowers are said to be regular. In these cases, even
In corn (Fig. 7.5), for example, the tassel (borne at the top though there may be an uneven number of parts in the
of the stalk) consists of a group of staminate flowers, and perianth, any line drawn through the center of the flower
the young ear is a group of pistillate flowers. When will divide the flower into two similar halves. They may be

staminate and pistillate flowers are borne on separate exact duplicates or mirror images of each other.
individual plants, as in Asparagus, willow (Salix), and Irregular flowers, such as pea (Pisum) (Fig. 7.8C) and
many other species, the species (or the plant) is said to mints (Mentha) (Fig. 7.88), have whorls either (a) with
be dioecious. dissimilar flower parts, (b) with parts that do not radiate
from the center, or (c) are not equidistant from one
Floral Symmetry
another. In most of these flowers, only one line will divide

In many flowers such as those of columbine (Aquilegia) the flower in equal halves; the halves are usually mirror
(Fig. 7.8A) or cherry (Prunus) (Fig. 7.9S) the corolla is images of each other. Some flowers such as, bleeding

Figure 7.6 Flowers of corn (Zea mays). A, the tassel, x '/5 S, ;

exserted anthers of staminate flowers, x 1; C, pistillate flowers

forming a very young ear, x '/2 D, several pistillate flowers with
;

attached styles, x 1.

Variations in Floral Structure

127
 pistil late
flower

stigma

(male)
catkin

anthers
stamens

one stamen

Figure 7.7 English walnut (Juglans regia) flowers. A, staminate

flowers; B, staminate inflorescence (catkin); C, dehiscing anther;
D, pistillate flower.

heart (Dicentra), though irregular, may be bisected by any Elevation of Flower Parts
number of lines into similar mirror images.
The irregular bean or pea flower has a corolla com- In flowers of magnolia (Fig. 7.2A, 16.3) and tulip (Fig.
posed of the following (Fig. 7.8C): one broad con- 7.9A), the receptacle is convex or conical and the different

spicuous petal, the standard or banner; two narrower flower parts are arranged one on top of another. The
petals (wings), one on each side; and, opposite the gynoecium is thus situated on the receptacle above the
banner, two smaller petals that are united along their points of origin of the perianth parts and androecium. An
edges to form the keel. ovary in this position is said to be superior. In the daffodil
(Fig. 7.9C) the ovary appears to be below the apparent
points of attachment of the perianth parts and the
Union of Flower Parts stamens. This is an inferior ovary. With an inferior ovary,
the anatomy of the flower parts indicates that the lower
In the flower of columbine illustrated in Fig. 7.8A, all parts portions of the three outer whorls, calyx, corolla, and
of the flower are separate and each sepal,
distinct; that is, androecium have fused to form a tube or hypanthium.
petal, stamen, and carpel is attached at its base to the The ovary in the daffodil is completely adnate with the
receptacle. In many flowers, however, members of one or hypanthium.
more whorls are to some degree united with one another, In a flower with a superior ovary, sepals, petals, and

or are attached to members of other whorls. Union with stamens arise from the outer, lower portion of the concave
other members of a given whorl is termed connation. receptacle, below the point of origin of the carpels (Fig.
Union of flower parts from two different whorls is known 7.1). The perianth and stamens are hypogynous, or with
as adnation (Figs. 7.8A.C; 7.5A,B; 7.98.Q. A precise reference to the three outer whorls, a condition of
terminology has been developed to indicate these fusions. hypogyny exists. In a flower with an inferior ovary, the

For example, syncarpy refers to the situation where perianth and stamens appear to arise from the top of the
carpels of the gynoecium are connate. ovary, and they are epigynous.

chapter 7 I Flowers, Fruits and Seeds

128
Figure 7.8 Floral symmetry. A, regular flower of columbine
(Aquilegia); B, irregular flower of a mint (Salvia), x '/2 C, flowers
—
;

and essential organs flower or garden pea (Pisum sativum).

In some flowers such as the plum and apricot (Prunus apical meristem, and in the axil of each leaf a flower is

sp.), the hypanthium does not become adnate to the borne. The oldest flowers are at the base of the
ovary. The perianth parts and stamens arising from the rim inflorescence and the youngest are at the apex.
of cuplike hypanthium around the ovary are perigynous In a simple raceme, the flowers are on pedicels that are
(Fig. 7.98). about equal in length. In a spike (Fig. 7.10), the main axis
of the inflorescence is elongated, but the flowers, each in

the axil of a bract, are sessile (without a pedicel). The

Inflorescences catkin is a spike that usually bears only pistillate or

staminate flowers. The inflorescence as a whole is shed

In most flowering plants, flowers are borne in clusters or later. Examples are willow, cottonwood (Populus), and
groups. Morphologically, an inflorescence is a flower- walnut (Figs. 7.7, 16.7).
bearing branch or system of branches. In manuals for the In all the inflorescences just mentioned, the flowering
identification of flowering plants, there are many different axis is elongated. If it is short, flowers appear to be arising
terms descriptive of various kinds of inflorescence. Only umbrellalike from approximately the same level. An
the most common ones are discussed and illustrated here inflorescence of this kind, in which pedicels are of nearly

(Fig. 7.10). equal length, is umbel (Fig. 7.10). The onion

called an

A very simple type of inflorescence may be found in (Allium) is a good example. The head is an inflorescence

such plants as currant (Ribes) and radish (Raphanus). It is in which the flowers are sessile and crowded together on

called a raceme. In this type, the main axis has short a very short axis. Members of the family Asteraceae
branches, each of which terminates in a flower. Each (Compositae), including thistle (Cirsium) and sunflower,
flower is on a short branch stem called the pedicel. The have this type of inflorescence (Fig. 7.10).

main axis of a raceme continues to grow in length more or Inflorescences of the raceme type may be compound,
less indefinitely; the apical meristem persists. Primordia of that is, branched. A
branched raceme is called a panicle
leaves arise in the usual manner along the margin of this (Fig. 7.10). Compound spikes occur in wheat, rye, and

Variations in Floral Structure

129
 stigma

petals

anth

filament style

ovary Vhypanthium

ovary

sepals
calyx

Figure 7.9 Diagram of elevation of floral parts. A, hypogyny

(Tulipa); B, perigyny in cherry flower (Prunus avium); C, epigyny
in daffodil flower (Narcissus pseudonarcissus).

certain other grasses. (gynoecium) (Fig. 7.1). The perianth, composed of calyx
In contrast to the raceme types of inflorescences, just and corolla, a protective covering of the stamens and
is

described, is the cyme (Fig. 7.10). In the cyme, the apex pistils and when it is present it also serves to attract and
of the main axis produces a flower that involves the entire guide the movements of pollinators.
apical meristem; hence, that particular axis ceases to
elongate. Other flowers arise on lateral branches farther
down the axis of the inflorescence and, thus, usually the The Androecium
youngest of the flowers in any cluster occurs farthest from
the tip of the main stalk. The flower cluster of chickweed Each stamen consists of an anther supported by a filament
(Cerastium) is an example of the cyme. 1 A). The anther usually consists of four elongated
(Fig. 7.1

and connected lobes called pollen sacs. Early in the

development of the anther, the pollen sacs each contain a
Angiosperm Life Cycle mass of dividing cells called microsporocytes
(microspore mother cells) (Fig. 7.1 10). Each
microsporocyte divides by meiosis to form four haploid
The essential floral organs necessary for sexual microspores. The nucleus of each microspore then
(1 n)

reproduction are the stamens (androecium) and carpels divides by mitosis to form a two-celled pollen grain that

chapter 7 I Flowers, Fruits and Seeds

130
 O

',
s V
ss <:
7

*N <
7
°s <
7
% pedicel
7
bract
7
simple raceme spike panicle

bract

umbel

bract of involucre

receptacle

head
cyme
Figure 7.10 Diagram showing types of inflorescence.

contains a tube cell and a smaller generative cell (Fig. make intelligent guesses about relationships in their

The role of this two-celled, haploid, male

7.1 1E). evolution.
gametophyte (gamete-producing plant) is to produce After the pollen grains are mature, the anther wall splits
sperm nuclei necessary later for fertilization. open and the pollen are shed. In some manner (which will
The'pollen grain surrounded by an elaborate, ornate
is
be discussed later), the pollen are transported to the
cell wall. The pattern of the pollen wall can be intricate stigma of adjacent or distant flowers. This process is
and beautiful; at the same time it is useful both to the plant
called pollination. Once on the stigma, if the pollen and
and to plant taxonomists. The wall is composed of an
the stigma are genetically compatible, it will germinate to
extremely hard material called sporopollenin (Fig. 7.13).
form an elongate pollen tube The ornate
(Fig. 7.1 1F).
This material is so hard that pollen grains many thousands
pattern of the pollen grain wall serves as the means for
of years old still retain their same pollen wall texture and
receptive stigmas to recognize compatible pollen. The
pattern. (Fig. 7.12).
Remember that floral structures are relatively unaffected
troughs and ridges of the pollen wall (Fig. 7.13) apparently

by environmental changes. Consequently, the form of contain proteins in specific patterns and concentrations. If

such structures as pollen grains can be used to make the patterns correlate with those of the stigma, metabolic
genetic comparisons between plant species. Plant events are triggered within the pollen grain to stimulate the
taxonomists study pollen wall patterns and are able to pollen tube to grow into the pistil's tissue.

Angiosperm Life Cycle

131
 nther

'filament

filament

tetrad

tapetum
pollen sac

anther
6

mitosis

germinated pollen grain

generative ce

pollen tube
tube nucleus

sperm cells

microspores

tube nucleus D

Figure 7.11 Development of pollen from a microsporocyte to the

pollen grain. A, stamen; S, cross section of anther; C,
development of tetrad of cells from the pollen mother cell by
meiosis; D, four microspores; E, pollen grain; F, germination of
pollen grain into pollen tube.

Figure 7.12 Fossil (A) and modern (B) pollen of the same
genus.

chapter 7 I
Flowers, Fruits and Seeds

132
 *.»**_Z7.>- %.'

Figure 7.13 Pollen grains as seen with the scanning electron

Iris, X210; 8, day lily (Hemerocallis fulva), x275;
microscope. A,
C,cucumber (Cucumis sativus), x 870; D, ragweed (Ambrosia),
X1000.

Angiosperm Life Cycle

133
 -midrib

midrib

carpel

carpels

carpels

carpel (leaf)
margins placentae

Figure 7.14 Diagram comparing gynoecia consisting of a single

carpel and of three connate carpels. A, section of ovary of pea
consisting of a single carpel; B, section of ovary of Tulipa
showing three connate carpels.

The Gynoecium
or more fused carpels, may occur (Fig. 7.146).
The structure of the gynoecium depends on the number The ovary is a hollow structure having from one to
and arrangement of carpels comprising it (Fig. 7.8C). In several chambers, or locules (Figs. 7.14, 7.15). The
the pea flower for example, there is a single carpel number of carpels in a compound pistil is generally related
forming the gynoecium (Figs. 7.14, 7.18). It consists of to the number of stigmas (Fig. 7.9A), the number of
three parts: (a) the ovary, an expanded basal portion, locules and, sometimes, the number of faces of the ovary
(b) the style, a slender stalk, and (c) a hairy irregular tip (Fig. 7.1 4/\). The pea has a single stigma and the ovary

portion, the stigma. When the pistil is composed of one has one locule (Fig. 7.1 4/4). There are three stigmas in the
carpel, such as pea, is referred to as a simple pistil. In
it tulip gynoecium, the ovary is three-sided (Fig. 7.9/4),
many other instances a compound pistil, consisting of two contains three locules, and is composed of three carpels.

chapter 7 l
Flowers, Fruits and Seeds

134
Placentation egg cell associated with two synergid cells. Since it is

frequently difficult to differentiate the egg cell from the

The tissues within the ovary to which the ovules are other two cells, all three cells are sometimes referred to as
attached are called placentae (singular, placenta, Fig.
the egg apparatus. The two nuclei that migrated toward
7.16). The manner in which the placentae are distributed the center approach each other; these nuclei, known as
in the ovary termed placentation. When the placentae
is
polar nuclei (Figs. 7.1 5/, J), form a binucleate cell called
are on the ovary wall, as in the pea (Pisum) and bleeding
the endosperm mother cell. The three nuclei remaining
heart (Dicentra, Fig. 7.1 ID), the placentation is parietal. at the end of the embryo sac opposite the micropyle form
When they arise on the axis ot the ovary, which has the antipodal cells. The embryo sac, which may be
several locules as in lilies, Fuchsia, and tulip, the considered to be a seven-celled haploid plant, (female
placentation is axile (Fig. 7.1 7Q. Less frequently, the gametophyte), is now mature and ready for fertilization.
ovules are on the axis of a one-loculed ovary, in which
Not every species of flowering plant produces a seven-
event the placentation is central, as in the primrose family
embryo sac,
celled but there is always an egg cell and an
(Figs. 7.1 7A,B).
endosperm mother cell.

Style and Stigma

The style is a slender stalk that terminates in the stigma. It

is through stylar tissue that the pollen tube grows. In some

Fertilization
flowers, the style is very short or entirely lacking; in
Prior to fertilization, the pollen tube has grown down
others, it is long. In Zea mays, the corn silks are the styles
through the stigma and style and has entered the ovary
(Figs. 7.6C.D). In general, the style withers after
pollination, but in some plants (e.g., Clematis)
(Figs. 7.1 1F, 7.1 5J, 16.1). Many may
pollen grains
it persists
and becomes a structure that aids in the dispersal of the
germinate, and their pollen tubes may grow through the

The
pistil, but only one usually enters into the embryo sac for
fruit. stigmatic surface often has short cellular
fertilization. While the pollen tube is growing, the
outgrowths that aid in holding the pollen grains; and
generative cell within it divides by mitosis to form two
sometimes it secretes a sugary and sticky solution, the
stigmatic fluid. In many wind-pollinated plants, such as
sperm cells. In some plants, like sunflower, the sperm cells
form even before the pollen are shed from the anther.
the grasses, the stigma is much branched, or plumelike.
Reaching the ovary, the pollen tube grows toward one
The Ovule of the ovules, usually enters the micropyle, penetrates one
or more layers of nucellar cells, enters the embryo sac
The ovule, the structure that will eventually become the and approaches the egg cell and the
(Figs. 7. 15 J, 16.1),
seed, arises as adome-shaped mass of cells on the endosperm mother cell. These stages are not easy to
surface of the placenta.The outer cells of the dome study, but it appears that the tip of the tube ruptures, one
develop to form one or two protective layers, the sperm cell enters the egg, and the other sperm enters the
integuments (Figs. 7.1 5C, 16.1). The integuments do not endosperm mother cell. Within the egg, sperm and egg
fuse at the apex of the ovule, thus leaving a small nuclei fuse. Within the endosperm mother cell, the other
opening, the micropyle (Fig. 7.1 5Q. While this sperm nucleus and two polar nuclei fuse (Figs. 7.1 5/, J).
development is one of the internal dividing
taking place, This double fusion of egg with sperm and polar nuclei with
cells of the ovule, the megasporocyte, is enlarging in sperm is called double fertilization. The zygote and
preparation for meiosis. The tissue within which the primary endosperm cell result from these fusions, the
megasporocyte has differentiated is known as the antipodal cells and synergid cells will degenerate, and
nucellus. The ovule then is composed of one or two outer conditions are set for further development of the seed and
protecting integuments, along with the micropyle, nucellus, fruit.
and megasporocyte (Fig. 7.1 5E). One interesting question concerns how the pollen tube
is directed into the embryo sac. The answer has been
worked out partially for cotton (Gossypium). In the embryo
Embryo Sac Development sac of cotton, one or both of the synergid cells begin to
shrivel and die before the pollen tube enters. It has been
Meiotic division of the megasporocyte (megaspore suggested that this activity results in the flow of some
mother cell) is considered the first step in the development chemical substance from the synergid that possibly
of a seed. As a result of these divisions, a row of four cells influences the direction in which the pollen tube grows.
called megaspores are produced in the nucellus. As a Two observations favor this hypothesis. First, at the base

rule, three of the cells (the ones nearest the micropyle) of both synergids is found a highly convoluted cell wall,

disintegrate and disappear, whereas the one farthest from known as the filiform apparatus (Fig. 7.15/). Cell walls in

the micropyle enlarges greatly (Figs. 7.15D.E). This this configuration are commonly believed to be associated
megaspore now develops into the mature embryo sac. with active chemical secretion. The second bit of evidence
The usual stages are as follows: (a) a series of three is that the pollen tube actually penetrates one of the
mitotic divisions, forming an eight-nucleate embryo sac synergids and empties its contents into it. The two sperm
(Fig. 7.1 5/-/); (b) migration of nuclei, (Fig. 7.15/); (c) cell then pass through the incomplete upper cell wall of the
wall formation around nuclei (Fig. 7.15/). synergid; subsequently, one of them moves through the
At the micropylar end of the embryo sac there is one incomplete side wall of the egg to fertilize the egg cell, and

Angiosperm Life Cycle

135
 nucellus

micropyle
H micropyle

the other moves deeper into the embryo sac to fuse with the receptacle, sepals, or petals, to increased growth
the endosperm mother cell. and incorporation into the fruit.

The fusions that occur in the embryo sac initiate the

Thus, the importance of fertilization is to stimulate the
following changes in the entire ovary:
growth of certain floral parts and to bring about a
withering of others. The net result is the development of

1. Development of embryo plant.

the zygote to form the the ovary wall into the fruit and the ovule into the seed.
Development of the primary endosperm cell to form the
endosperm (the reserve food supply of the seed).
Development of integuments to form the seed coat. Pollination
Absorption or disintegration of nucellar tissue. In some
plants, however, a portion of the nucellus, rather than
the endosperm, may become the storage tissue of the Pollinationmay be brought about by either wind or
seed. Such tissue of nucellar origin is called perisperm. insects,and occasionally by water, birds, bats, and other
Development of ovary tissue forms the fruit. small mammals. Flower structure and pollen type are
Possible stimulation of accessory flower parts, such as generally adapted to one or the other of these pollinating

chapter 7 I Flowers, Fruits and Seeds

136
egg cell

egg cell

Figure 7.15 Diagram of embryo sac development. A, cross

section of a flower bud showing the four whorls, four-carpelate
ovary in center; B, enlarged view of ovary showing four carpels
and ovules; C, an ovule, megasporocyte in prophase of meiosis;
D, late telophase of second meiotic division; E, four megaspores,
three degenerating; F, two-nucleate embryo sac; G, four-nucleate
embryo sac; H, eight-celled embryo sac; /, mature embryo sac; J,
fertilization, synergids degenerating.

vectors.In insect pollination, adaptation may be very easily wind-borne. Their stigmas, in many cases, are
complex, indicating a long association between the insect feathery and expose a large surface to catch flying pollen.
vector and plants. Hayfever victims suffer from windborne pollen.

Living Vectors
Pollinating Vectors
Plants that are pollinated by living vectors usually possess
Wind
a colorful perianth that, together with other floral parts,

Pollen is carried chiefly by wind and insects. Wind may be arranged into a complex architecture (Figs. 7.8,
pollination is common in plants with inconspicuous They also produce both a sweet-tasting fluid, called
7.9).

flowers, such as in grasses (Fig. 7.6), walnuts (Fig. 7.7), nectar,and volatile compounds having distinctive odors.
oaks, and ragweeds (Ambrosia). Such plants usually Many investigators have shown that these characteristics
produce pollen in enormous quantities. The flowers usually are highly adapted for the attraction of pollinators.
lack odor and /or nectar and hence are unattractive to Bees, for example, are able to detect and distinguish
insects. Furthermore, their pollen is light and dry and between many odors and colors and degrees of

Pollination

137
 calyx
flower
stigma — 9

margin

funiculus

igma

Figure 7.16 Development of legume fruits. A, bean (Phaseolus

young pods, x 1 B, pistil from a pea
vulgaris) from flowers to ;

x 5; C, pea pod, unopened; D.opened

(Pisum sativum) flower,
pea pod showing developing seeds attached to carpel margins,
x1.

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Flowers, Fruits and Seeds

138
 another is called cross-pollination. Many species are
capable of both self- and cross-pollination; others are
obligate cross-pollinators.

Summary
1. The flower, the distinguishing structure of the
Anthophyta, is formed of four whorls of parts
specialized to carry out sexual reproduction, including
pollination, fertilization, and seed production.
2. A floral apex has lost its ability to elongate and its

potentiality for vegetative growth. The ability for

vegetative growth is not regained until the completion
of the two nuclear phenomena, meiosis and
fertilization.

3. The four whorls of floral leaves are (a) the calyx,

composed of sepals; (b) the corolla, composed of
petals; (c) the androecium, composed of stamens; and
(d) the gynoecium, composed of carpels.
4. There are two parts to a stamen, the pollen-producing
anther and the filament.
5. A single unit of the gynoecium is frequently called a
pistil. If it consists of one carpel, is a simple pistil;
it if

it consists of two or more fused carpels, is a it

compound pistil.

6. A pistil consists of three parts, the stigma, the style,

parietal and the ovary. The stigma is receptive to pollen, and
Figure 7.17 Three types of placentation. A and B, free central the ovary encloses the ovules. At the time of
(Primula); C, axile (Fuchsia); D, parietal (Dicentra). fertilization, the ovules consist of integuments,
nucellus, and embryo sac.
7. Meiosis takes place both in the anthers and in the
sweetness. Bees can distinguish at least three colors in ovule.
addition to ultraviolet, including the most common perianth 8. Pollen may be considered a two-celled haploid plant
shades (yellow and blue). Bees are directed to nectar- protected by a which is frequently elaborately
cell wall,

producing glands by splashes or lines of contrasting color. sculptured. Two sperms are produced by the
When the insect attempts to reach nectar or collect pollen, generative cell

floral architecture ensures that the insect transfers pollen 9. The embryo sac may be considered a seven-celled
to the stigma. In nasturtium, for example, nectar is held in haploid plant. Two of its cells, the egg cell and the
a long narrow tube or spur. A foraging bee directs its primary endosperm cell, are stimulated to develop by
proboscis down the tube and usually rubs it against the fusion with the sperm cells.

stigma along the way. In this way, pollen, which may have 10. Pollination is the transfer of pollen from anthers to
been picked up in earlier visits to other flowers, is stigmas. A pollen tube growing down the style to the
deposited on the stigma. embryo sac serves as a pathway by which the two
Orchid flowers have many very specialized mechanisms sperms reach the embryo sac.
to ensure pollination. The petal shape and color of species 11. Floral architecture facilitates pollination, which is most
of Ophrys closely resemble the appearance of a female frequently carried out by wind or insect vectors.
wasp or fly. Male wasps or flies, emerging from the pupal 12. Double fertilization, the union of one sperm with the
stage before the females, mistake the Ophrys flower for egg cell and of a second sperm with the primary
females. The male lands on the flower and attempts endosperm cell, occurs only in the Anthophyta.
copulation; repeated "pseudocopulation" results in 13. The embryo plant develops from the zygote. The
pollination. endosperm, developing from the fusion of sperm and
primary endosperm nucleus, supplies some
Types of Pollination
nourishment for embryo and sometimes for seed
There are essentially two different kinds of pollination, development.
determined by the genetic similarity of the plants involved. 14. Variation in floral architecture arises (a) from variation
If anthers and stigmas, essential organs in pollination, in number of parts in a given whorl; (b) from symmetry
have the same genetic constitution, the transfer of pollen in floral parts; (c) by connation of floral parts; (d) by
from the anther to the stigma in the same flower or plant adnation of floral parts; (e) by elevation of floral parts;

is called self-pollination. If two parent plants with different and (f) by the presence or absence of certain whorls.
genetic constitutions are involved, transfer of pollen from 15. Flowers are either solitary on flower stalks or grouped
the anther of the flower of one plant to the stigma of in various inflorescences such as heads, spikes,

Summary

139
 catkins, umbels, panicles, racemes, or cymes.
16. There are essentially two types of pollination: cross-
pollination, which results in much variation in the
\JkiSt
progeny, and self-pollination, which results in progeny ventral carpellary bundles
having great similarities.

PART TWO
The Fruit

A fruit is a ripened ovary of a flower; it may include other

floral parts. In many cases, fruit development depends on
pollination and the activity of indoleacetic acid, or other
growth substances.
Fruits are auxiliary structures in the sexual life cycle;
they occur only in the Anthophyta. Their development has
unquestionably been an important factor in the successful
evolution of land plants. Fruits protect seeds, aid in their

dissemination, and may be a factor in timing their

germination. Fruits are highly constant in structure and are
thus very important plant parts in the classification of the
Anthophyta. In everyday usage, the term "fruit" usually
refers to a juicyand edible structure, such as an apple,
plum, peach, cherry, orange (Citrus), or grape. Such
structures as string beans, eggplant (Solanum melongena),
okra (Hibiscus esculentus), squash, and cucumbers,
which are commonly called "vegetables," and "grams" of
corn, oats, wheat (Triticum), and other cereals are not
popularly thought of as fruits. However, all the above are
fruits in a botanical sense.

Development of the Fruit

Development of the fruit (pod) of pea and bean is

described as an example of the changes that may occur

during transformation of the ovary into a fruit. Pistils of
bean and pea flowers are each composed of one carpel Figure 7 18 Cross section of bean ovary after fertilization, x20.
(Fig. 7.17); that is, one ovule-bearing leaf. Recall that the

ovary of bean is formed by fusion of margins of the carpel

and that ovules are attached to these margins (Fig. ovules were not fertilized. If none within the bean ovary is
7.1 4A). A cross section of the bean ovary (Fig. 7.18) fertilized, the ovary does not enlarge. In most plants,
shows ovary wall, ovule, locule. The ovary wall may be normal fruit development takes place only if pollination is

divided into three distinct layers, (a) an outer epidermis, followed by fertilization and if fertilization is followed by
(b) an inner epidermis, and (c) a middle zone consisting of seed development. Some fruits, such as banana and navel
several layers of cells. There are three carpellary bundles, oranges, develop without fertilization. These fruits, called
one for each margin and one for the midrib opposite them. parthenocarpic fruits, are seedless and frequently
Fertilization initiates a series of changes in the embryo different in shape and taste from normal seeded fruits.
sac and other tissues of the ovule that lead to the
development of the seed. Tissues of the ovary wall
undergo marked changes into three layers. The fruit wall Kinds of Fruits
(developed from the ovary wall) is called the pericarp,
and the three more or less distinct parts, in order
beginning with the outermost, are exocarp, mesocarp, In classifying the different kinds of fruits, the following
and endocarp (Fig. 7.23S, shown in almond). When the criteria are taken into account: (a) the structure of the
pod is mature, floral structures such as pedicel, calyx, flower from which the fruit develops, (b) the number of

withered stamens, style and stigma, and even remnants of ovaries involved in fruit formation, (c) the number of
may also be present.
the corolla carpels in each ovary, (d) the nature of the mature
Mature pods usually show the presence of small, pericarp (dry or fleshy), (e) whether or not the pericarp
underdeveloped (abortive) ovules. It is probable that these splits (dehisces) at maturity, (f) if dehiscent, the manner

chapter 7 |
Flowers, Fruits and Seeds

140
 —

of splitting, and (g) the possible role that sepals or

receptacle may play in formation of the mature fruit.

Simple fruits are derived from a single ovary. They may

be dry or fleshy; the ovary may be composed of one
carpel or of two or more carpels; and the mature fruit may
be dehiscent or indehiscent. On the other hand,
aggregate fruits and multiple fruits are formed by
clusters of simple fruits. The difference between these
types of fruits depends on the number of flowers involved
in their formation. In strawberry (Fragaria) (Fig. 7.19/4) and
blackberry (Rubus) (Fig. 7.198), the aggregate fruit is

derived from the many ovaries of a single flower These

matured ovaries or fruits are all attached to a common
receptacle. Such groupings constitute aggregate fruits,
and the simple fruits comprising them may be classified
according to the scheme of classification of simple fruits

given below. Mulberry, (Morus), fig (Ficus), and pineapple

(Ananas comosus) are multiple fruits. Multiple fruits
achenes
consist of the enlarged ovaries of several flowers, more or
less grown together into one mass. In some plants
mulberry, for example — associated floral structures form a receptacle

part of the and in the fig the receptacle enlarges

fruit, to
become the sweet edible portion (Figs. 7. 20/4, C).

Simple Fruits

Pericarp Dry and Dehiscent drupes

Legume or Pod. This type of fruit, characteristic of

nearly all members pea family, Fabaceae
of the
(Leguminosae), arises from a single carpel, which at

maturity generally dehisces along both sutures (Fig. 7.17).

Pods may be spirally twisted or curved as in alfalfa.

*4 receptacle
Follicle. An example of follicle is the fruit of magnolia
(Figs. 7.21/4, 16.3). The follicle develops from a single
carpel, and opens along one suture, thus differing from
the pod, which opens along both sutures.

Capsule. Capsules are derived from compound ovaries,

thatis, an ovary composed of two or more united carpels.

Each carpel produces a few to many seeds. Capsules

dehisce in various ways (Figs. 7.21 B,C,D).

Silique. This is the type of fruit (Fig. 7.21 E)

characteristic of members of the mustard family,

Brassipaceae (Cruciferae). The silique is a dry fruit derived

from a superior ovary consisting of two carpels. At
maturity, the dry pericarp separates into three portions;
the seeds are attached to the central, persistent portion

Pericarp Dry and Indehiscent

Figure 7.19 Aggregate fruits. A, strawberry; B, blackberry.
Achene. Sunflower is an example of this type of fruit.

These fruits are commonly called "seeds," but as in

sunflower (Fig. 7.21 F), a carefully broken pericarp reveals

that the seed within is attached to the placenta by its stalk.
This pericarp may be separated easily from the seed coat,
that is, from the layer of cells just beneath it. Achenes are
indehiscent.

Kinds of Fruits

141
 fleshy receptacle

.pore
staminate flower

receptacle

drupes

Figure 7.20 The fruit and flowers of fig (Ficus carica). A, end of
fleshy receptacle, showing pore, which is lined with staminate
flowers; B, receptacle turned inside out, showing pistillate flowers
that have matured into small drupes, x 1 Diagram of a fig. C, D,
.

and E, inedible capri fig; C, section of receptacle showing location

of staminate and pistillate flowers, x 1 D, staminate flower, x3;
;

E, top: fertile long-styled pistillate flower of the edible Calimyrna

fig, x3; E, bottom: short-styled abortive pistillate flower, x3.

Figure 7.21 Simple fruits. A, Magnolia, follicle; B, Datura,

capsule; C, D, Tulipa, capsule; E, Lunaria, silique; F, Helianthus,
achene, G, Acer, samara.

Grain or Caryopsis. This is the fruit of the grass family, consisting of two carpels that split, when mature, along
Poaceae (Gramineae), which includes such important the midline into two one-seeded indehiscent halves.
plants as wheat, corn, and rice (Oryza). Like the achene,
the grain is a dry, one-seeded, indehiscent fruit (Fig. Nut. The term nut is popularly applied to a number of
7.26). It differs from the achene, however, in that pericarp hard-shelled fruits and seeds. A typical nut, botanically
and seed coat are firmly united all the way around. speaking, is a one-seeded, indehiscent dry fruit with a
hard or stony pericarp (the shell). Examples are the
Samara. This is a dry, indehiscent fruit, which may be chestnut (Castanea), walnut, and acorn (Fig. 7.23A). An
one-seeded, as in the elm (Ulmus), or two-seeded, as in acorn, the fruit of the oak, is partially enclosed by a
maple (Acer) (Fig. 7.21 G). These fruits are typified by an hardened involucral cup. The husk or shuck of the walnut,
outgrowth of the ovary wall,which forms a winglike which has been removed before the product reaches the
structure. market, is composed of involucral bracts, perianth, and the
outer layer of the pericarp. The hard shell is the remainder

Schizocarp. This is the fruit characteristic of the carrot of the pericarp.

family, Apiaceae (Umbelliferae), which includes such

Pericarp Fleshy
common plants as carrot (Daucus), celery (Apium), and
parsley (Petroselinum). The schizocarp is a dry fruit Drupe. Examples of this type of fleshy fruit are cherry,

chapter 7 I Flowers, Fruits and Seeds

142
 placenta

seeds

carpels

Kinds of Fruits

143
almond, peach, and apricot, all of which are members of a many living cells filled with juice.

genus in the Rosaceae (rose family). The olive (Olea) fruit Another berrylike fruit is that of members of the family
is also a drupe. The drupe is derived from a single carpel, Cucurbitaceae, which includes watermelon, squash,
and is usually one-seeded. However, one examines if pumpkin, and cucumber. It is called a pepo (Fig. 16.10)
young ovaries of flowers of almonds, cherries, plums, or The outer wall (rind) of the fruit consists of receptacle
other members of the group to which they belong, two tissue that surrounds and is fused with the exocarp. The
ovules will be found, one of which usually aborts (fails to flesh of the fruit is principally mesocarp and endocarp.
develop into a seed, Fig. 7.238). The drupe has a hard
endocarp consisting of thick-walled stone cells. The Pome. This type of fruit is characteristic of a subfamily
exocarp is thin, forming the skin, and the mesocarp forms of Rosaceae, which belong apple (Fig. 7.24) and pear.
to
the edible flesh. The pome is derived from an inferior ovary (Fig. 7.2AA).
The flesh is enlarged hypanthium, and the core has come
Berry. This fleshy type of fruit is derived from a from the ovary (Fig. 7.248).

compound ovary. Usually, many seeds are embedded in a

flesh, which is both endocarp and mesocarp, although the
line of demarcation may be difficult to see (Fig. 7.21 H).
The tomato is a common example of a berry.
The citrus fruit (lemon, orange, lime, and grapefruit) is a
Compound Fruits — Formed From
Several Ovaries
type of berry called a hesperidium. has a thick, leatheryIt

rind (peel), with numerous oil glands, and a thick juicy

Aggregate Fruits
portion composed of several wedge-shaped locules (Fig.
7.22). The peel of a citrus fruit is exocarp and mesocarp; An aggregate one formed from numerous carpels of
fruit is

the pulp segments are endocarp. The juice is in pulp sacs one These fruits, considered individually,
individual flower.
or vesicles; they are outgrowths from the endocarp walls may be classified as types of simple fruits. The strawberry
(Fig. 7.22C), and each mature vesicle is composed of flower has numerous separate carpels on a single

stigma

ovary

pericarp "peel"

oil glands

& seed

luice sacs

Figure 7.22 Flower and fruit (hesperidium) of orange (Citrus

sinensis). A, flower, dissected to show pistil and stamens; B,
lengthwise section of maturing ovary; C, cross section of mature
fruit.

chapter 7 |
Flowers, Fruits and Seeds

144
 mesocarp

endocarp

funiculus

abortive ovule
nucellus
integuments

ovary cavity

cotyledons

Figure 7.23 A, acorns of oak (Ouercus). The fruits are a type of

nut and the cups are fused involucrol Practs, x yA B, drupe,
.

almond (Prunus amygdalus).

calyx lobe

stamens and styles

endocarp

outer limit of
carpel (core line)
Figure 7.24 Flower and fruit (pome)
apple (Malus sylvestris).
of
A, median longitudinal section of flower showing inferior ovary
and fused (adnate) petals and sepals; B, median section of floral tube
mature fruit.

receptacle. The ovary of each carpel has one ovule, and Flowers of raspberry and blackberry, and other species
the ovary develops into a one-seeded dry fruit (achene). of Rubus have essentially the same structure as those of
The receptacle which these fruits are attached becomes
to strawberry. In these flowers, many separate carpels
fleshy; the whole structure, which we call a strawberry, is attached to one receptacle develop into small drupes
an aggregate of simple fruits, each an achene. The (Figs. 7.1 9AB; 16.9).
receptacle is stem tissue and consists of a fleshy pith and
Multiple Fruits
cortex with vascular bundles between them. The hull of

the strawberry fruit is composed of persistent calyx and A multiple one formed from individual ovaries of
fruit is

withered stamens. The achenes are usually spoken of as several flowers. These fruits, considered individually, may
seeds. be classified as types of simple fruits. The fig and

Kiods of Fruits

145
pineapple are examples of multiple fruits; the individual
fruits composing them are nutlets in fig and partheno-
Key to Fruits
carpic berries in pineapple. I. Fruit formed from a single ovary of one flower Simple
The fig fruit wean enlarged fleshy receptacle (Fig.
eat is fruits.

7.20). Its flowers are very small and are attached to the A. Pericarp fleshy.
inner wall of this receptacle. Both staminate and pistillate 1. The ovary wall fleshy and containing one or more
flowers occur and may be borne in the same or in carpels and seeds. Berry.
different receptacles. a.Ovary wall a hard rind. Pepo.
Each ovary develops into a nutlet that is embedded in b.Ovary wall with a leathery rind. Hesperidium.
the wall of the receptacle. Thus, the fig is derived from 2. Only a portion of the pericarp fleshy.
many flowers, all attached to the same receptacle. a. Exocarp thin; mesocarp fleshy; endocarp stony;
singleseed and carpel. Drupe.
Accessory Parts of Fruits b. Outer portion of pericarp fleshy, inner portion
papery, floral tube fleshy; several seeds and
In the pineapple, the edible portion is a thickened pulpy
central stemwhich berries are embedded. Tissues other
in
carpels. Pome.
B. Pericarp dry.
than the ovary wall which form part of a fruit, are referred
1. Dehiscent fruits.
to as accessory. Thus, much of the fleshy fruit of
pineapple, apple, and strawberry is accessory.
a. Composed of one carpel.
1. Splitting along two sutures. Legume.
2. Splitting along one suture. Follicle.
PART THREE b. Composed of two or more carpels.
1. Two or more carpels dehiscing in one of
four different ways. Capsule.
The Seed 2. Two carpels; separating at maturity, leaving
a persistent partition wall. Silique.
2. Indehiscent fruits.

Seed Development a. Pericarp bearing a winglike growth. Samara.

b. Pericarp not bearing a winglike growth.
1. Carpels, two to many, united when
The seed completes the process of reproduction initiated immature, splitting apart at maturity.

in the flower. The embryo developed from the zygote, and Schizocarp.
the seed coat from the integuments of the ovule. Recall 2. Carpels one, if more, not splitting apart at

that at the time of fertilization, egg cell, primary endosperm maturity; one-seeded fruits.

cell, embryo sac.

synergids, and antipodals constituted the a. Seed united to the pericarp all around.
The embryo sac is surrounded by nucellus, and all these Caryopsis or grain.
cells and tissues are enveloped by one or two integu- b. Seed not united to the pericarp all around.
ments. This complete structure is the ovule (Figs. 7.25A Fruit large, with thick, stoney wall. Nut.

7.15). Double fertilization involves (a) fusion of egg and Fruit small, with thin wall. Achene.
sperm nuclei to form a zygote nucleus and (b) fusion of II. Fruits formed from several ovaries.
polar nuclei with a second sperm nucleus to form a A. Fruits developing from one flower.
primary endosperm nucleus (Fig. 7.25S). Aggregate fruits (classify the individual fruits in key
After fertilization, the zygote nucleus remains quiescent for simple fruits).

for a time. The primary endosperm nucleus, however, B. Fruits developing from several flowers.

divides rapidly and soon builds up an endosperm tissue Multiple fruits (classify individual fruits in key for

which, at first, may lack cell walls (Fig. 7.25Q. The zygote simple fruits).

nucleus will divide only after the endosperm has

developed. The first cell divisions of the new generation
result in a filament of from four to eight cells (Fig. 7.25D).
Now the cell closest to the micropyle elongates, pushing
the other cells of the filament into the endosperm. At
about the same time, the cell furthest from the micropyle general, for there are many variations in details of embryo
divides at right angles to the axis of the filament (Fig. development.
7.25E). This is the first in a series of divisions that will While development of the embryo is taking place, the
finally result in the embryo of the seed. Further divisions nucellus, endosperm, and integuments are also
result in a globular stage, and finally in a heart-shaped undergoing changes that are characteristic of the group of
stage after the two cotyledons have developed (Fig. plants to which the seeds belong. In the great majority of
7.25G). The major distinction between embryos of plants, nucellus and endosperm are required only for the
dicotyledonous and monocotyledonous plants is the initial stages of development. This is particularly true of the
presence of cotyledons, two or one, respectively. The nucellus, which is generally used up as a nutritive source
embryo always consists, in addition, of an axis with a root in early stages. It persists as a food storage tissue, the
apex at one end and a shoot apex at the other. (Fig. perisperm, in seeds of sugar beet (Beta), and a few other
7.25H). The course of events just described is very species. The persistance of endosperm as a food reserve

chapter 7 I Flowers, Fruits and Seeds

146
 nuclear
ision
antipodal cells

Figure 7.25 Diagram of the development ot dicotyledonous

angiosperm seed. A, ovule after fertilization; B, after fusion of
gamete nuclei to form zygote and endosperm; C, free nuclear
divisions in the endosperm; D, filamentous stage of the
proempryo; E, elongation of suspensor cell and division of
proembryo cell; F, globular stage of the embryo; G, heart-shaped
stage of embryo development.

Seed Development

147
 seed coat
Endosperm

Coleoptile cotyledon

Aleurone shoot apex

layer
endosperm

radicle

Caryopsis
coat

Figure 7.27 A longitudinal section through an onion seed

Scutellum (Allium cepa) showing the embryo coiled within the endosperm.
(cotyledon)

Figure 7.26 Median section of caryopsis of yellow foxtail

(Setaria lutescens).

occurs in many monocotyledons (Figs. 7.26, 7.27). is It seed coat. In most dicotyledonous seeds food is stored in
highly developed in grasses, some of which like rice and cotyledons (Fig. 7.25) while in monocotyledonous seeds
corn are of major economic importance. The endosperm the endosperm is usually the principal food storage tissue.
persists as a food storage tissue in relatively few When food storage occurs within the embryo, the
dicotyledonous seeds; the castor bean (Ricinus normal vascular tissues of the embryo convey the
communis) is a good example. solubilized food to meristematic regions, where is it

Inmost seeds, integuments of the ovule become hard required for growth. Food stored in the endosperm,
and horny seed coats in the mature seed. The scanning outside the embryo, is in many cases absorbed through
electron microscope shows the seed coats to be variously normal protodermal or epidermal cells. In other cases, the
and sometimes beautifully sculptured (Fig. 7.28). cotyledons become specialized to act as absorbing organs
(Fig. 7.35). In most grasses, the entire cotyledon has
become a highly specialized absorbing organ known as
the scutellum (Fig. 7.26).
Kinds of Seeds

Seeds of angiosperms differ in two ways: (a) they have Common Bean
one or two cotyledons and (b) they store food either in the
embryo, in the endosperm, or more rarely, in nucellar The fruit of the bean plant is a pod; within it are seeds.
tissue, the perisperm, which lies between embryo and The external characters of the bean seed are more clearly

Figure 7.28 The coats of many seeds are characteristically

sculptured. A, morning glory (Convolvulus arvensis), x 23; B,
California poppy (Eschscholtzia californica), x 62.

chapter 7 I
Flowers, Fruits and Seeds

148
seen after soaking it in water. Points of interest are the
hilum, micropyle, and raphe (Fig. 7.29). The hilum is a
large oval scar, near the middle of one edge, left where
the seed broke from the stalk or funiculus when the
beans were harvested. The micropyle is a small opening in
the seed coat (integument) at one side of the hilum, which
was observed in the ovule as the opening through which
the pollen tube entered. The raphe is a ridge at the side of
the hilum opposite the micropyle. It represents the base of
the funiculus, which is fused with the integuments

Conducting tissue present in the funiculus will be

continued in the raphe. An elevation or bulge of the seed
coat adjacent to the micropyle marks the position of the
radicle (embryo root) within the seed.
When the seed coat of a soaked bean is removed, the

entire structure remaining is embryo; no endosperm is

present. The following parts of the embryo can be
observed: (a) a shoot consisting of two fleshy cotyledons,
a short axis (the hypocotyl) below the cotyledons, and a
short axis (the epicotyl) above the cotyledons, bearing
several minute foliage leaves and terminating in a shoot
tip; and (b) the root or radicle (Fig. 7.29).

Grasses
The so-called "seed" of corn, or of any other grass, is in

reality a fruit, the caryopsis. It is a one-seeded, dry,

indehiscent fruit with the pericarp (ovary wall) firmly
attached to the seed. Pericarp and seed coats are so
firmly attached to each other and to other tissues of the
grain that it is impossible to peel them away.
A longitudinal section of a caryopsis of yellow foxtail
(Setaria lutescens) is shown in Fig. 7.26.The endosperm
constitutes the bulk of the grain and iscomposed of (a) an
outermost layer (single row of cells) known as the
aleurone layer and (b) a starchy endosperm. Cells of the
aleurone layer contain proteins and fats but little or no
starch.
Like all embryos of other seed plants, the grass embryo
has an axis with a shoot apex and a root apex (Fig.
7.26). The shoot apex, together with several rudimentary
leaves, aresurrounded by a sheath, the coleoptile. The
rudimentary root (radicle)is also surrounded by a sheath,
radicle shoot
the coleorhiza. At the juncture of shoot and root is a very
short stemlike region. A relatively large part of the grass Figure 7 29 Bean (Phaseolus vulgaris) seed. A, external side
view; B, external face or edge view; C, embryo opened.
embryo is a single cotyledon, which has for a long time
been called the scutellum. It is a shield-shaped structure
that lies in contact with the endosperm. The outer cells of
the scutellum secreteenzymes that digest the stored foods
of the endosperm. These digested foods move from
endosperm cells through the scutellum to the growing
parts of the embryo. Unlike the bean, the cotyledon of
grasses remains within the seed during germination and
never develops into a green leaflike structure above Onion
ground.
It is noteworthy that the process of milling to make The onion seed consists of seed coats enclosing a small
polished white rice removes the outside caryopsis coat, amount of endosperm. The embryo is a simple axis, the
plus the entire protein-rich aleurone layer,and the outer radicle and single cotyledon being quite prominent. The
layers of the endosperm. This means that most of the shoot apex is located at about the midpoint of the axis and

nutritious protein is removed before the rice is eaten! The appears as a simple notch. The embryo is coiled within
removed material, composed of the outside layers and the the seed coats, the radicle usually pointing toward the
embryo axis, is sold as bran. micropyle in the seed coats (Fig. 7.27).

Kinds of Seeds

149
 What induces germination? Many seeds
Dissemination of Seeds and when
will germinate

provided with moisture, oxygen, and a favorable

Fruits temperature. The water content of seeds is low, between 5
and 10%. The cytoplasm with contained organelles is
Agents in Seed and Fruit Dispersal scarcely recognizable because is crowded between the
it

large amount of reserved food materials (Fig. 7.31). Water

The chief agents in seed and fruit dispersal are wind, is, at first, imbibed very rapidly, which results in the

water, and animals. swelling of the protoplasm with a reappearance of

organelles (Fig. 7.32). Increase in water content makes
Wind possible an increased metabolic activity. From this point in
time the seed must have an adequate water supply for
The structural modifications of seeds and fruits that aid in
survival.
dissemination by wind are of several types. The most
common of these are winged types (Fig. 7.30C) and
Many seeds not germinate even when supplied with
will

plumed types (Figs. 7.30A.B).

water, oxygen, and a favorable temperature. There are
various factors in different plants that are associated with
Water the breaking of dormancy. For instance, light is necessary
for the germination of some grass seed and for some
Fruits with a membranous envelope containing air like
strains of lettuce (Lactuca) and perhaps other plants.
those of sedges, or with a coarse, loose, fibrous outer
Water and gases must pass through the seed coats. If
coat, as in the coconut (Cocos), are well adapted for
seed coats restrict the movement of water and gases or
dispersal by water. A great variety of fruits and seeds float
are impermeable to one or the other or to both, the seed
in water, even though they lack special adaptations to
coats must decay, be broken, or be scratched, allowing
ensure buoyancy, and are readily transported long
water and oxygen to reach the embryo before germination
distances by moving water. In irrigated districts, irrigation
can start. In some seeds, the embryo itself fails to
water is a very important means by which weed seeds are
germinate when provided with favorable conditions. In
distributed.
orchids, the seeds are disseminated while embryos are
Animals rudimentary and the embryos must develop further before
germination.
Many seeds and fruits are carried by animals, both wild Many kinds of seeds are known that will not germinate
and domesticated. Seeds with beards, spines, hooks, or unless the seeds, while on a moist substrate, are
barbs adhere to hair of animals (Fig. 730E). An example is subjected for a time to temperatures close to freezing. At
bur clover (Medicago denticulata). the other extreme, some seeds will not germinate unless
Seeds of many plants pass through the digestive tract of they have been subjected to the rather high heat of a fire.
animals without having their viability impaired. Fleshy, Another type of dormancy is produced by the presence of
edible fruits may be eaten by birds (Fig. 7.30F) and natural chemical inhibitors. These occur in many fruits and
carried by them long distances, and then the seeds serve to keep the seeds dormant while they are enclosed
regurgitated or discharged with the excrement. Squirrels by the fruit. In other cases, inhibitors are present in the
carry nuts, such as those of walnut and hickory (Carya), seeds themselves or may be produced by decaying
and the seeds of pines. Seeds of some aquatic and marsh leaves, or forest litter. The influence of the plant growth
plants and of mistletoe (Phoradendron), which are covered substances on dormancy and seed germination is still
with a sticky material, are carried on the feet of birds. unclear.
The first indication that the processes of germination
have begun is generally the swelling of the radicle. In all

Seed Dormancy and cases, the radicle imbibes water rapidly and, bursting the
seed coats and other coverings that may be present,
Germination starts

to grow downward into the soil. This helps to assure that

the young seedling has an adequate supply of water and
Seeds can remain viable for remarkably long periods. In nutrients when the shoot breaks through the surface of
one study, begun in 1878, Dr. W. J. Beal of Michigan the soil. Although the succeeding steps of germination are
State University buried jars containing seeds from several essentially similar, there are variations. For instance, in

plant species. At 5- and 10-year intervals a jar was germination of beans, peas, and onions, a structure with a
opened and the seeds tested for germination. Most sharp bend or hook is first forced through the soil (Figs.

species remained viable for at least 10 years, and one 7.33, 7.34, 7.35), but the structure forming the hook is
species, the moth mullein (Verbascum blattaria), still different in each case. Once above the ground, the hook
germinated after more than 90 years. This, however, is no straightens. In the case of bean, a straightening of the
record for seed longevity. Seeds from the Oriental lotus hypocotyl raises cotyledons and shoot apex above ground
(Nelumbo nucifera) have been removed, still viable, from (Fig. 7.33); a lengthening and straightening of the epicotyl

archeological diggings known to be more than 1000 years will pull the shoot apex away from the cotyledons (Fig.

old! Seeds of a few other species have been recovered 7.33).

from cold and anaerobic deposits, dated at over 1000 Upon straightening of the pea epicotyl, however, the
years, and have also proven viable. is truly remarkable
It cotyledons remain in the ground, and only the apex and
that seeds can remain living over such long periods. first leaves will be raised upward (Fig. 7.34). Cotyledons

chapter 7 I
Flowers, Fruits and Seeds

150
Figure 7.30 Fruits and seeds showing various devices aiding in
dissemination. Wind: A, Clematis, x4; 6, dandelion (Taraxacum
vulgare), x 10; C, seed of Coulter's big-cone pine (Pinus
coulteri), x 2.
Attachment: D, cranesbill (Geranium), x 3; E, bur clover
(Medicago denticulata), x 5; F, fleshy edible fruit of Cottoneaster,
x4; Violent dehiscence of pericarp; G, vetch (Vicia sativa), x2.

Seed Dormancy and Germination

151
Figure 7.31 Yellow foxtail (Setaria lutescens) scutellum in dry Figure 7.32 Yellow foxtail (Setaria lutescens) radicle after 65
seed (caryopsis or grain), xOOOO. hours of germination, xOOOO.

seed coat

Figure 7.33 Stages in the germination of a bean (Phaseolus

vulgaris) seed.

chapter 7 l
Flowers, Fruits and Seeds

152
 plumule

seed coat

•^^ cotyledon

hypocotyl

radicle

Figure 7.34 Stages in the germination ot a pea (Pisum sativum)

seed.

cotyledon

vascular cylinder
cotyledon

seed coat

primary root

root cap

Figure 7.35 Germination ot an onion (Allium cepa) seed.

Seed Dormancy and Germination

153
that are raised above ground may carry on
photosynthesis.
In onion, the primary root first penetrates the soil.Then
a sharply bent cotyledon breaks the soil surface, and
slowly straightens out. The cotyledon of the onion is

tubular and base encloses the shoot apex (Fig. 7.35).

its

There is a small opening at the base of the cotyledon

through which the first leaf finally emerges. In grasses, the
situation is more complex. The shoot and root apices are
enveloped by tubular sheaths known as the coleoptile and
coleorhiza, respectively (Fig. 7.36). The primary root
rapidly pushes through the coleorhiza. The root continues
to grow, but is eventually replaced by adventitious roots
that arise from the lower nodes of the stem (Fig. 7.36).

The coleoptile elongates and emerges above ground,

becoming 2-4 cm long. At this time, the uppermost leaf
pushes its way through the coleoptile and, growing
rapidly, becomes part of the photosynthesizing shoot. adventitious
roots

Summary
1. Seeds may store food within or outside the embryo. In
most dicotyledons such as cucumber and bean, food is
stored in the two cotyledons. Cotyledons may also
serve as absorbing and, later, as photosynthesizing
organs. Food may be an endosperm rather
stored in

than in the cotyledons. monocotyledonous seeds,

In

food is stored in an endosperm.

2. In grasses, such as corn, a single cotyledon (scutellum) Figure 7.36 Germination of a grain of wheat (Tnticum).
has become a highly specialized absorption organ that
does not emerge from the seed. In seeds like those of
lilies and onion, the cotyledon emerges from the seed

coats and becomes green, but its tip continues to inhibitors, or the destruction of the seed coats, may be
absorb food from the endosperm. required in some instances.
3. Mature seeds are dormant and, depending on the 5. In germination, the cotyledons may be elevated above
species and the immediate environment of the seeds, the ground, sometimes to become photosynthetically
theymay remain viable and dormant from a few months active, or in the case of some monocotyledons, to
tomany years. continue absorption of the endosperm. In other cases,
4. Dormancy is usually broken by the provision of the cotyledons remain below the ground.
moisture, oxygen, and a favorable temperature. Other 6. In the grasses, the single cotyledon functions as a
factors, such as light, the removal of chemical specialized food-absorbing organ.

chapter 7 l
Flowers, Fruits and Seeds

154
 8 CHAPTER

8 the control of growth

and development

Which woodland?moreThe
a
are the alike: cattle in

cattle are decidedly

a herd or oaks
more
pattern adapted for movement through the soil (see Figs.
in 8.3, 8.4, Color Plate 2). The dark-grown seedling is said to
unitorm than the trees in terms of the number of be etiolated.
limbs and appendages they have. The systems that control In addition to leaf expansion, other changes are initiated
growth and development in the higher animal body are by light. The stem usually elongates very rapidly in
tightly programmed and conservative; they resist darkness (or in the soil) because light inhibits elongation.
influences from the environment far more than they adjust Supporting tissues of the vascular system are weakly
to them. developed in darkness when the soil would normally give
Plants are the opposite. They have stimulus-response all the support necessary. Light stimulates increased
systems that allow the environment to modify their path of development of xylem, by triggering the formation of
development. This is the principal way in which plants
enzymes that in turn catalyze lignin synthesis two or three
adjust to the environment; this is how they compensate for
hours after the start of illumination. This adds the strength
the lack of muscles. If in a poor location, they detect the
that is needed to support the stem above ground.
situation by means of environmental signals and they
Whereas light slows the elongation of much of the stem,
modify their morphology in a way that permits them to
cells on the lower surface of the hook increase their
persist. And whereas animals meet the threat of predators
elongation. This causes the stem to straighten so that the
by running away, the stationary plant suffers the attack
leaves are oriented upward (Fig. 8.1). These effects of
and later replaces the lost parts.
light are all initiated by phytochrome.

The first light received by the plant may be coming from

only one side. The usual response is for the stem or
Environmental Adaptation coleoptile to initiate a bending response that orients the tip
by the Young Plant toward the light.

The mesophyll cells of a leaf and the cortex of a stem

are programmed to delay the formation of the
The plant's use of environmental signals can be illustrated
photosynthetic apparatus until they are exposed to light
by tracing the behavior of a seedling as it starts the life of
(Figs. 8.1, 8.4). In darkness, the immature plastids contain
an independent plant.
a small amount of protochlorophyll, an incomplete form
Once the seed germinates, the root senses gravity and
of chlorophyll, which is attached to a special catalytic
directs growth downward, in the direction that usually
its
protein. On illumination, the protochlorophyll absorbs light
leads to water, minerals, and anchorage. The shoot grows
and is converted to chlorophyll. More protochlorophyll is
upward in response to gravity, toward the most likely
formed in its place. As chlorophyll accumulates,
location of light.
membranes in the plastids are organized into thylakoids
The newly emerging shoot is a delicate structure that
and the full battery of chloroplast enzymes is assembled.
must push its way through
and past obstacles. soil
The can be fully green within 36 to 48 hours (Figs.
leaf
Dicotyledonous plants do not expand their leaves until
8.2, 8.3). Phytochrome is involved in these events, too.
they reach the soil surface (see Color Plate 2, Figs. 8.1,

8.2). The cluster of leaf primordia and the apical meristem

The delay in developing mature chloroplasts prevents the
behind plant from needlessly spending stored food on plant parts
are usually inverted and pulled up through the soil

the recurved portion of the hypocotyl or epicotyl (the that may never receive light for photosynthesis.

hook).The delicate leaves and apex are thus protected. In As this introduction has shown, the systems that control
some monocot plants, the young leaves are rolled up and development in the plant can respond to signals from the

completely ensheathed in a protective tube, the environment. Physiologists have long felt that a study of

coleoptile. When the coleoptile reaches the surface, the these signals and the responses might lead us to valuable
leaves push out of the splitting coleoptile and unroll. Light, insights about the plant's developmental systems. Charles

received by a pigment called phytochrome, signals that Darwin, for example, held such a belief; and the book that

the shoot has reached the surface. Plants that are grown he and son Francis published in 1881 "The Power
his , of

in darkness do not receive this signal and grow in the Movement in Plants," helped to launch a revolution of

155
knowledge about developmental control. But to understand stimulate the reading of particular genes, as in the operon
we must tirst shitt our tocus to the level of
this revolution, mechanism (Chapter 2). According to another current idea
the cell. the hormones bind to membranes, affecting their
permeability to other molecules that in turn affect
regulatory enzymes and the reading of genes.
Hormones and the Control of Although we cannot define exactly how hormones work,
we can describe the kinds of developmental processes
Growth and Development that they help to control,and we can illustrate the
hormones in Development combines the results of
action.

The typical plant begins its life as a single cell, the zygote. three major processes: cell division, which produces new
Many cell divisions occur, and the organism becomes a cells; growth, which increases the size of cells and

colony of cells that cooperate to form and maintain an organs; and differentiation, the changes by which cells
integrated plant body. The cells in one region form a leaf; become specialized in structure and function. Each of
those in another region, a root or a flower. Some cells these processes can vary in speed and direction. And all
become green and perform photosynthesis; others lose of these variables must be controlled in each part of the

their nuclei,form sieve plates, and join the phloem. What plant body if development is to be normal. The following
causes the cells to develop so differently? What tells each passages outline what we know about the part that
cell the proper path for its development? And how do the hormones play.
cells achieve cooperation?
Acorns always produce oak trees; corn kernels always
produce corn plants. This shows that development
depends on hereditary information. At the outset, all the Auxins
information is contained in the zygote. Thus one might
suppose that cells develop differently because each cell Our knowledge of hormones began with the discovery of
gets only a fraction of the zygote's original store of the auxins, a class of hormones that were first detected
hereditary information. because they stimulate elongation growth in grass
But experiments have shown that mature parenchyma coleoptiles. The kind of experiment that led to their
cells can give whole plants, if the cells are removed
rise to discovery is shown in Fig. 8.6.
from their original location and are given suitable stimuli The only naturally occurring auxin to be identified so far
and raw materials. This demonstrates that mature cells is indoleacetic acid (IAA), which is shown in Fig. 8.5A

may contain all the hereditary information that the zygote But many man-made compounds have been developed
contained. Evidently, then, each cell uses only a part of its that affect plants in the same way as IAA, and these are

hereditary information. A common idea is that each cell also called auxins.
follows a particular reading program as it refers to the
hereditary information. This selective pattern of reading Control of Cell Enlargement by Auxin
guides the cell through a sequence of developmental
Cell enlargement is a process critical to the life of almost
changes. Cells come to differ because they follow different
every cell in a plant, and the ability of the plant to control
reading programs, but if so, how is the program
determined? this process precisely is central to growth and
The simplest suggestion is that cells follow a path of
morphogenesis. A plant cell may be thought
of as an

development response from the surrounding

inflatable bag (the protoplast) surrounded by a cover (the
in to signals
cell wall) that may be rigid or may be expanded under
environment. This agrees with the fact that seedlings can
modify their development according to cues such as light pressure, depending on its makeup. By using respiratory

and gravity. But, in addition, internal signals may pass energy and carrier systems, a variety of solutes are
between cells within the plant to coordinate their pumped into the vacuole, and water follows osmotically,
development. Such internal signals were postulated by creating turgor pressure. The turgor pressure in a typical

Charles and Francis Darwin, and much of modern plant cell might be maintained at five times the pressure of the

physiology has been concerned with identifying these surrounding air. For comparison, automobile tires are

signals and tracing their action. Today we know of five usually inflated tomore than three times atmospheric
classes of molecules that act as developmental signals in pressure. Whether or not growth occurs will depend on

plants. These compounds, called hormones, act at very both the pressure and the extensibility of the wall.
low concentrations and function in the plant only as As the cell grows, the wall is forced to stretch.
signals. Stretching hardens the wall and makes less extensible.
it

We do not yet know in detail how the hormones initiate The same thing happens with a stretched rubber band, as
their effects. But even small changes in their molecular one may easily discover. To continue growth, the wall
structures tend tochange their activity. This suggests that must therefore be continually softened. It is in this
hormones are precisely shaped to fit receptor sites, much softening process that auxin seems to exert its most
as modulators attach to regulatory enzymes (Chapter 2). immediate effect. The wall is built of carbohydrate
Proteins are likely candidates for receptors since other polymers, such as cellulose and pectin as well as protein.
types of molecules generally lack the structure-recognition These framework polymers are joined in a network by
capabilities that proteins have. One popular proposal is cross-links that are believed to limit the extent to which
that the hypothetical hormone receptors inhibit or the wall can be stretched. For continued growth,

chapter 8 I The Control of Growth and Development

156
 HC^^C- C— CH 2 — COOH
II

CH
^C N
H H

Indole acetic acid

A: Molecule of the auxin, indole acetic acid (IAA),

made up of two rings and a side chain
B: Molecule of gibberellic acid

CH 3
H 3 C^ CH 3 H
y OH H |

o^k ^^JL
CH 3
COOH

C: Zeatin D: The structure of abscisic acid (also called

dormin or abscisin II)

H \
^H
E: Ethylene

Figure 8.5 Plant hormones. A, indole acetic acid, IAA, the most
common natural auxin; B, gibberellic acid,
3
one of more than GA ,

40 very similar natural gibberellins; C, zeatin, one of several

natural cytokinins, D, abscisic acid, ABA; E, ethylene, a gaseous
growth regulator.

physiologists believe that the cross-links between wall addition to its wall-softening effect, auxin also appears to
polymers must continually be broken and re-formed in new play a role in stimulating and coordinating wall formation.
positions. By this theory the turnover of cross-links is the The mechanism for this effect of auxin is unknown.
basic softening process. Recent evidence suggests that in

some Controlling Auxin Concentration

hydrogen ions stimulate the breaking of
plant cells,
cross-links in the wall. The required hydrogen ions Elongation rates respond within minutes if auxin is added
originate in the protoplast and are pumped through the or removed. This emphasizes the importance of factors
plasma membrane and into the wall by a carrier system that control the concentration of auxin.
that is embedded in the membrane. In dark-grown plants Auxin is formed in regions of the plant separate from the
of several species, auxin has been found to stimulate the zones of rapid elongation. The tip of the coleoptile and the
action of this ion pump. Hence the higher the auxin stem tip with its cluster of young leaves make IAA from the
concentration, the faster hydrogen ions are supplied to the amino acid, tryptophan. The auxin is then moved down
wall, and the faster the cell grows. has long been known It the stem through the elongating region (Fig. 8.6). The
that auxin can act only respiration is also taking place.
if movement is called polar transport because it is a one-
This makes sense the ion pump in the membrane uses
if way, energy-requiring movement away from the tip.

ATP as an energy source. Typical rates of movement are 10 to 15 mm per hour,

There is evidence that in some cells, auxin may soften much slower than movement of materials in the xylem and
walls and promote growth by a mechanism other than phloem, which may exceed 1 meter per hour.
causing hydrogen ion secretion. This is true, for example, Furthermore, polar transport in the youngest stem regions
in light-grown pea stems. It is not yet known how auxin is in the phloem and
opposite to the direction of flow
promotes growth in such cases. xylem and has been shown to occur even in stem tissue
growth is to continue for a long period
If of time without from which all vascular elements were removed.
the wall thinning to the point where it breaks under turgor Thus, auxin moves down from the tip through the
pressure, new framework polymers must continually be growing zone, and the concentration in this stream
added. Materials are secreted through the membrane and establishes the growth rate of the tissue. Remove the tip
deposited in the wall. Respiratory activity is needed to of a coleoptile, the auxin source, and the remaining auxin

provide energy for both the synthesis and the secretion is rapidly drained out of the growing zone. Within a few

process. minutes the growth rate drops. Put back the tip or replace
The rate of wall secretion must be coordinated with the auxin supply with a block of gel containing IAA and
growth if the wall is to remain at a constant thickness. In growth is soon restored (Fig. 8.6). If, however, the auxin

Hormones and the Control of Growth and Development

157
 curvature no curvature curvature

tip
shielded base
shielded

photoreceptor
at tip
auxin moves
down shaded side
causing unequal
growth

auxin source
tip removed on one side

-=> tip

auxin replaceable
transported by auxin
growth down into
stops agar block

upright coleoptile
section

horizontal coleoptile section

14
S<&1
C-IAA-labeled
donor block

auxin transported
to lower side 65%

Auxin equal
on two sides No destruction of
IAA
L

Figure 8.6 Diagram of experiments showing how regulation of

auxin transport can result in phototropic curvature (A, B, C, L) or
in geotropic curvature (/, J. K), and the Avena curvature bioassay
used in estimating minute amounts of auxin (D, E, F, G, H).
Experiments in which IAA labeled with radioactive carbon was
used (/, J, K, L) show how geotropic and phototropic stimuli
cause auxin to be transported laterally. Unequal amounts of IAA
accumulate in the receiver blocks.

chapter 8 I The Control of Growth and Development

158
source placed on only one side of the cutoff stump, the
is pigment causes a lateral diversion of the auxin flow, as in
up below the auxin supply grow more than
cells that line geotropism. The result is a curvature toward the light.
those on the other side. This leads to a bending of the
coleoptile or stem. Apical Dominance

Tropic Curvatures In the typical pattern of growth, the growing tip of the

stem with its young leaves inhibits the sprouting of lateral

Plants can orient the direction of growth of organs in buds below the apex. This phenomenon is called apical
response to environmental cues. These growth responses dominance. In a related effect, called apical control, the
are called tropisms. When the shoot and root emerge main apex slows the growth of lateralbranches that do
from the seed, is of obvious advantage to direct shoot
it sprout. This dominance of apical growth over lateral
growth upward and root growth downward. The direction growth decreases with the distance from the tip of the
of gravitational force is a reliable cue. We do not yet know stem, and varies with the age of the plant, the genotype,
how the plant senses gravity, but many physiologists think nutrition, and other environmental factors. A plant with
the answer may be statoliths— particles in the cell that strong apical dominance (e.g., the sunflower) has little or
are heavy enough to sink to the bottom, comparable to the no branching. Weak apical dominance leads to a bushy
stones in the inner ear that give human beings a sense of appearance, as in tomato plants. In grasses, branching
the vertical. occurs at the very basal nodes of the stem. These
Colorless plastids, heavy with starch grains, may act as branches are called tillers and the process is tillering.
statoliths. When on its side, these
a plant is laid The suppressed lateral buds act as a reservoir of shoot
amyloplasts tumble to the lowermost side of the cell. tips that can rapidly take over growth to replace a
Once the shoot has sensed its orientation in space, we damaged main shoot tip. In garden pea seedlings, removal
know that the direction of auxin transport changes, so that of the growing tip is followed within four hours by
by the time the auxin has traveled from the tip of the stem increased metabolism in the first lateral bud below the old
to the main growing zone there may be twice as much tip. Another early event is the differentiation of xylem to

auxin moving through the lower half of the stem compared form a vascular connection between the bud and the main
with the upper side (Fig. 8.6). The greater growth on the stem. Completion of the vascular connection is followed by
lower side pushes the tip until is pointing up. The it growth of the new shoot.
amyloplasts settle back to their original position, and auxin The main shoot tip apparently gives off an inhibiting
transport becomes uniform on both sides again. The plant influence that passes down the stem. If the tip is replaced
requires a few minutes to sense its original change in by a supply of auxin in a lanolin paste or agar block, the
position, and by 15 minutes there is enough auxin lateral buds remain inhibited. This suggests that lateral
concentration difference to cause a curvature. In some buds are inhibited in the intact plant by auxin that the
rapidly growing seedlings, the stem may turn upright shoot tip produces. But paradoxically, after the branch has
within an hour from the time that it is placed horizontally. started to grow, auxin applications increase its growth and
This tropic curvature in response to gravity is called the active branch forms its own auxin supply. Thus auxin
geotropism. acts in opposite ways before and after the bud sprouts.
The geotropic curvature of roots downward has many Auxin inhibits the formation of vascular connections to the
aspects in common with the shoot response, except that lateral bud, a fact that may help to resolve the paradox.
there is is the hormone
considerable doubt that auxin This is an on differentiation, which is apart
effect of auxin
involved. It is cap produces some
clear that the root from its on growth. Here we see an example of a
effect
controlling substance that moves away from the tip to common observation: each plant hormone exerts a
control curvature. However, auxin is transported toward diversity of effects in different situations.
the root tip. The identity of the growth substance involved There are many other "growth correlations" where the
in root geotropism is being actively investigated. At present growth and development of one plant part is related to that

it seems likely that the controlling substance is a growth of another (Fig. 8.7). Thus apical control may cause lateral

inhibitor. organs such as branches, leaves, rhizomes, and stolons to

Phototropism is a change in the direction of growth grow horizontally or at some angle with respect to gravity
that results when exposed to light
the stem or coleoptile is that is different from the main shoot.
vertical habit of the

from one side (Fig. 8.6). The response depends on a Removal main shoot may be followed by a bending
of the

pigment system that absorbs blue and violet light. The of a nearby branch into an upright position as it takes over

nature of the pigment is unknown, but its absorption the dominant position in the plant. The direction of growth
suggests that it is not phytochrome or chlorophyll. In an of some rhizomes and stolons has been shown to depend

unevenly illuminated shoot, there is more pigment on hormones coming from the shoot tip.
activation on the lighted side than on the shaded side. One might suppose that continued production of auxin
Careful experiments have shown that IAA labeled with at the tip and its transport downward should lead to an

radioactive carbon and applied to the tips of coleoptiles accumulation somewhere. But no region of accumulation
moves symmetrically down the coleoptiles in the dark. has been found. Auxin is destroyed or inactivated along
Light treatments that cause curvature result in twice as the way. There are enzymes in plants that can destroy
much auxin moving down the shaded side as down the much more auxin than the plant can ever produce. And
lighted side, but the total amounts transported are the there are enzymes that inactivate auxin by tying to it

same as in dark controls. Thus somehow the light-sensing another molecule, forming an inactive compound:

Hormones and the Control of Growth and Development

159
 IAA and
gibberellic acid auxin and gibberellic acid synthesized
in young leaves and bud — move to

stem to control elongation

gibberellic acid controls cell

division in subapical region

auxin controls
differentiation
flowering stimulus moves from
leaves to buds to initiate flowers
abscisic acid made in

leaf in response to
water stress — closes
stomata, reduces water loss cytokinins move to leaves
from keeps root and
roots,

cytokinins made in young shoot growth in balance

fruit, necessary for growth

auxin and gibberellic acid promote

ethylene accumulates cambium in formation of
activity of
in mature fruit to secondary vascular tissues
induce ripening

ethylene and abscisic acid made

in senescing leaf promote
auxin moves abcission zone development
toward root IAA
tip

gibberellic acid and cytokinins

synthesized in roots move to
shoot and leaves

factor made in root tip

controls geotropism
of roots

Figure 8.7 A diagram illustrating some typical hormonal

interrelationships among various portions of the plant.

indoleacetic acid indoleacetylaspartic delicate operations on the shoot tip to study how xylem
+ aspartic acid acid (inactive) forms in the young leaf primordia and the nearby stem.
These studies show that a leaf primordium stimulates
Supplying the plant with high concentrations of auxin differentiation in the procambial strand leading to it (Fig.

frequently stimulates the synthesis of the enzymes 8.8/A). If the leaf primordium is sliced away, the same
responsible for inactivation. differentiation of vascular tissue in the stem can be
induced by applying auxin (Fig. 8.8C).
Cell Differentiation In Coleus stems, a wound that severs a vascular bundle
When xylem elements differentiate, the process is easily is followed by cell divisions and differentiation of xylem

seen because the changes in the cell are extreme. The elements from parenchyma cells around the wound. The
wall develops heavy thickenings with characteristic new xylem elements reconnect the injured bundle.
patterns, and eventually the protoplast dies. These visible Observed more closely, has been found that new
it

signs of change have made the xylem a favorite tissue for phloem cells are formed even earlier than the xylem cells
studying cell differentiation. in this regeneration process. These events require a
One experimental approach has been to perform supply of auxin that is normally transported out of the

chapter 8 |
The Control of Growth and Development

160
 leaf primordium

procambium procambium
strand

xylem does \^

differentiate

xylem does differentiate

leaf primordium removed

drop of IAA applied
here only
leaf primordium
not removed

notch

xylem does differentiate

xylem does not differentiate
C

Figure 8.8 Diagram of an experiment showing how a leaf

primordium provides a stimulus for xylem differentiation in the
procambium. The notch severing the procambium isolates the
above it. A, control with intact leaf primordium;
tissue of interest
6, leafprimordium removed; C, leaf primordium removed and a
drop of auxin substituted. Auxin is an effective replacement for
the stimulus from the primordium.

leaves above the wound and then down the stem. If the Whether the differentiating cells become xylem or
leaves are removed at the time of wounding, regeneration phloem elements in lilac callus depends on the
does not occur. The effect of the missing leaves can be concentration of sucrose that is applied. High
replaced by applying auxin to the petiole stumps. concentrations induce nothing but phloem; low
Further down the stem where elongation has ceased, concentrations induce only xylem; and intermediate levels
even an herbaceous plant like the tomato may have some stimulate both xylem and phloem to differentiate.
secondary growth taking place from a vascular cambium. Significantly, when both xylem and phloem are formed,
This cambial activity depends on a supply of auxin that is their arrangement is concentric; the xylem is toward the
received from the shoot tip. In woody plants that have center, as in the cross section of an intact stem.
been dormant over winter, the buds that sprout in the
spring give off auxin that stimulates the cambium. A wave
of cambial divisions can be traced down the stem,
following the flow of auxin. Gibberellins
Further advances concerning the differentiation process
have come with tissue cultures. Under suitable While European scientists of the 1 920s were occupied
parenchyma tissue from stems can
conditions, blocks of with the auxins, Japanese workers discovered another
be induced to continue cell divisions and growth without class of hormones. These hormones were first identified in

differentiation. This produces a uniform mass of studies of a disease of rice plants, the bakanae (foolish
unspecialized cells, a callus. Blocks of callus tissue made seedling) disease. Afflicted seedlings grow very tall and
from stems show a remarkable response to auxin and
lilac eventually fall over. The disease is caused by a fungus,
sucrose: if these two substances are applied to the top of Gibberella fujlkuroi. The abnormal growth could be
the block, a discontinuous ring of vascular tissue develops duplicated by treating normal seedlings with the liquid in

within the tissue block, at a distance below the surface. which the fungus has grown. The active agent in the liquid
The diameter of the ring and its distance from the droplet proved to be a group of substances that were given the
can be increased by raising the concentration of auxin. If name gibberellins. They form an extensive group of
auxin spreads by diffusion, its concentration must related compounds, variations on the basic structure
decrease with distance from the source. If we put these shown in Fig. 8.5S. Although these compounds were first
facts together, it seems that the signal for cells to become discovered in a disease, they have since been found as
vascular elements must involve their exposure to a natural hormones in a wide array of flowering plants. The

particular auxin concentration. bakanae disease was a result of oversupply.

Hormones and the Control of Growth and Development

161
Gibberellins and Stem Growth gibberellin are estimated by comparing their effect on
growth to that of known hormone concentrations.
The effect of gibberellin on a monocot (rice) was
Bioassays have been important in studying all the plant
described above. Some dicotyledonous plants have very
hormones, because plant materials are often sensitive to
few cell divisions in the subapical region of the stem tip
quantities of hormone that would be far too small to detect
during the early months of growth, but leaves are formed
chemically.
normally at the apical meristem. (Fig. 8.14). The result is a
shoot that has many leaves separated by very few short
Gibberellins and Enzyme Synthesis
internodes. A plant with this form is called a rosette,
whereas plants with elongate internodes are called As a grain such as barley or corn starts to germinate, the
caulescent. Rosette plants typically respond to gibberellin embryo begins to grow but has limited food reserved in
applications by rapid stem elongation, or bolting. The itself. The main reserves are in the starchy endosperm, a

bolting response can be caused naturally by an tissue with cells loaded with starch, reserve proteins, and
environmental factor such as winter cold or long days. some nucleic acids. A special layer of living cells, the
These factors trigger an increase in gibberellin, which aleurone layer, surrounds the main endosperm tissue and
promotes cell divisions as outlined above. is instrumental in digesting the stored materials to soluble
Auxins and gibberellins often act simultaneously to forms that can diffuse to the embryo. Early in germination,
control stem elongation. In some cases, gibberellin may the embryo synthesizes gibberellin (GA), which diffuses to
stimulate cell division, thus providing more cells on which the aleurone layer and triggers the synthesis of enzymes
auxin can act. In the coleoptile, gibberellins act at an early to digest stored materials in the endosperm. the embryo If

stage of development and the auxin-sensitive stage shown is removed from the seed prior to germination, very little

in Fig. 8.6 comes later in life. Woody stems need both digestion of the remaining endosperm takes place. Adding
auxin and gibberellin to maintain cambial activity. gibberellin in minute quantities to the embryoless
In Europe, commercial use is made of the growth endosperm induces the synthesis and secretion of
retardant CCC or Cycocel (2-chloroethyltrimethyl enzymes just as the embryo had provided the stimulus
if

ammonium chloride) to inhibit stem elongation in wheat (Fig. 8.10).

plants. Cycocel inhibits the production of gibberellin in the The barley aleurone layer can be isolated and studied
plant. Shorter, stronger stems result, and the plants are independently of the embryo and storage endosperm. It is

much more resistant to lodging, that is, to being knocked a collection of cells with no growth activities and no
down by wind and rain. Lodging makes harvesting difficult. division; it does have, however, a very active ability to

(Short stems in grains are also achieved genetically in synthesize a few proteins and to secrete them. As such it

breeding programs and are important factors Mexican in has been used to study the mechanism of action of
wheats and Philippine IR8 rice, plants of the "green gibberellin.
revolution," that have raised crop productivity in Asia and About 6 hours after adding gibberellin, the aleurone
Central America.) cells start to make enzymes that break down starch,
"Dwarf" forms in a variety of plants are often due to a proteins, and nucleic appearance of the
acids. Prior to the
diminished gibberellin synthesis or to an enhanced enzymes, new ribosomes and endoplasmic reticulum
production of compounds that oppose the action of membranes are formed, as well as enzymes involved in
gibberellin. Dwarf forms of corn and peas are used in membrane lipid synthesis. We have much to learn about
estimating the concentrations of gibberellins in extracts of the way in which gibberellin unleashes all these changes,
Such methods are called bioassays.
plants or plant parts. though it does appear that an activation of genes (new
Figure 8.9 shows a bioassay where quantities of enzyme synthesis) is involved.

0.0 0.0001 0.0005 0.001 0025 005 0.01 0.025 0.05

/ig GA 3 / PLANT

Figure 8.9 Dwarf peas (Pisum sativum) showing the promotion

of stem elongation by gibberellic acid applied seven days prior to
photograph. This response is used to bioassay the gibberellin
contents of plant extracts.

chapter 8 I The Control of Growth and Development

162
 Barley
Intact seed Embryo-less half-seed

embryo

embryo makes GA
which moves to
aleurone

amylase no amylase
synthesized

reserves
digested to no digestion
soluble forms of reserves

8: +GA C: -GA
amylase hydrolyzes
starch to glucose;
glucose taken up
by growing embryo

Figure 8.10 Diagram illustrating how gibberellin from the

embryo induces the synthesis of the starch-degrading enzyme,
a-amylase, in the aleurone layer.

In the manufacture of beer, the starch stored in the matter of trial and error. Such trials led to the discovery, in

grain endosperm must be hydrolyzed to a soluble form the mid-1950s, that cell division could be stimulated by
before its conversion into alcohol by the glycolytic adding adenine, one of the nucleic acid bases. Since then
enzymes of yeast. The natural production of amylase a variety of related compounds have been found to work
occurs in the malting barley. The addition of extra GA much better than adenine. These are the cytokinins, one
speeds up amylase synthesis by the aleurone enough to of which is shown in Fig. 8.5C. Some cytokinins are
be used commercially. synthetic, while others occur naturally, especially in young
developing fruits.

Besides affecting cell divisions, the cytokinins play a

part in controlling the initiation of plant organs. In tissue
Cytokinins
cultures, for example, cells of tobacco pith will enlarge if

Hormones of a third major class, the cytokinins, were supplied with nutrients and auxin, but will divide only if

discovered through work with tissue cultures. small amounts of a cytokinin are added. Moreover, by
Physiologists have found that keeping cells alive and varying the balance between auxin and cytokinin, it is

active after they have been removed from the plant possible selectively to initiate the development of roots and
depends on the exact constitution of the medium to wnich shoots (Fig. 8.11). High auxin-to-cytokinin ratios cause
the cells are transferred. Finding the best medium is a root initials to differentiate. A low ratio of auxin to cytokinin

Hormones and the Control of Growth and Development

163
 high cytokinin ratio

low cytokinin ratio

control

intermediate cytokinin ratio

intermediate cytokinin, low auxin

\^^Ly
continued growth as
callus

Figure 8.1 1 Diagram of the control of differentiation exerted by

interaction of auxin and cytokinin. Pieces of tobacco pith tissues
were aseptically grown on nutrient medium (tissue culture)
supplemented with various levels of the two hormones.

causes clumps of cells to become apical meristems that and root growth. The upward movement of cytokinins from
grow into shoots. Intermediate concentration ratios cause the root may help to maintain this balance. Increases in

callus to form. Shoots can be removed, rooted, and grown root growth cause more abundant cytokinin supplies,
to mature plants. which cause a corresponding increase in shoot growth.
Cytokinins also seem to contribute to apical dominance. Leaves also contribute to the shoot /root balance by
If a cytokinin is applied directly to a lateral bud that is producing carbohydrates and vitamins (especially vitamins
being suppressed by an active shoot tip, the bud may of the B group, needed as cofactors in respiration) that
grow out. The applied cytokinin promotes the formation of roots cannot build for themselves.
vascular connections between bud and stem, a process Cytokinins also prevent leaf deterioration, or
that is inhibited by auxin. Cytokinins appear to be made in senescence. a leaf is removed from a stem,
If

the tips of roots, and they travel upward in the xylem. senescence is initiated. The changes include the
They may accumulate at the cut end of a rooted stump, breakdown of storage carbohydrates, proteins, nucleic
where adventitious buds often form. The opposed acids, and chlorophyll; and a cessation of protein
movements of auxin and cytokinin may give each region of synthesis. the leaf is put under conditions where
If forms it

the plant a unique balance between quantities of these two adventitious roots, the senescence is halted. Applications
hormones, and their opposing effects on bud growth may of cytokinin have a similar effect in halting the
help to determine the pattern of branching as the plant deterioration of detached leaves. The maintenance of
grows. active RNA
and protein synthesis seems to be an
Plants normally maintain a close balance between shoot important part of cytokinin action in delaying senescence.

chapter 8 |
The Control of Growth and Development

164
Ethylene plumule emerges from the seed, the tip of the shoot has a
hooked curvature that protects tne apex. The young
Some it was the practice among lemon growers
years ago plumule synthesizes ethylene rapidly in darkness. The
to pick lemons green and to allow them to ripen in heated ethylene maintains the hook and prevents the leaves from
boxcars as they were shipped across the United States to enlarging. When the shoot breaks through the soil, light is

distant markets. This system worked nicely, until the leaky absorbed by phytochrome, which in turn triggers a
kerosene heaters in the treight cars were replaced by decrease in ethylene synthesis. The leaves are released
more efficient steam heaters. The shippers were dismayed from ethylene inhibition and expand. The hook "opens" to
to find that the lemons no longer ripened in time for present the leaves to sunlight, which further stimulates leaf
marketing. Investigators found that the kerosene stoves development. (Fig. 8.1).
had been leaking small quantities of a simple gas, While growing underground, the shoot encounters
ethylene, which acted as a ripening stimulus. This is one obstacles such as clods. The pressure of the obstacle
of many early observations that ethylene as a pollutant against the shoot causes a dramatic increase in ethylene
affects plant development. More recently, ethylene has production. The results can be simulated by gassing
been found to be a potent natural hormone that exposed seedlings with ethylene (Fig. 8.12). Ethylene
contributes to normal development in plants. Its structural induces the stem to swell and inhibits elongation. The
formula is shown in Fig. 8.5E. thickened stem can exert more upward force against the
Ethylene is built in the plant from the amino acid obstacle. If the stimulus is prolonged, the stem's geotropic
methionine, a constituent of all cells. Since ethylene has response is modified; grow almost horizontally.
it starts to
only limited solubility in the aqueous phase of a cell, it These responses increase the likelihood that the shoot will
diffuses into the atmosphere. Natural ethylene action in a penetrate or grow around the obstacle to reach the open
tissue depends on its continued manufacture. Its air.

concentration depends on the balance between synthesis Some of these effects involve changes in the direction
and diffusion. Enclosing plants especially fruits in a — — of cell growth, and they raise one of the most fundamental
tight container can cause the concentration of ethylene unsolved problems in plant development: the question of
made by the plant to increase to levels that have drastic what controls the direction of cell growth; what causes
effects on its growth and development. Effective levels are some cells to grow equally in all directions while others
often in the range of 0.1 to 1 part per million of air. Even become cylindrical or spindle-shaped. Studies of growing
the levels of ethylene present in urban air pollution are cells have suggested that the direction of growth is
sometimes enough to cause biochemical changes in determined when new cellulose microfibrils are added to
plants. the wall. If the microfibrils are laid down at random, the
The action of ethylene in controlling vegetative growth cell expands uniformly. If they are deposited around the

can be illustrated by its effect on the pea seedling. As the cell like the hoops around a barrel, the cell will grow

* 1

/
/
1
O on r\ n 05 1fl nr 20
\Ccre
0.040 0.080 0.160 0.320 0.640
1
ppm ETHYLENE

1- tial size (3 d< jys)

Figure 8 12 The response of dark-grown pea seedlings to

various levels of ethylene during four days (ppm = parts per
million ethylene in air). Internally generated ethylene under stress
can induce similar responses.

Hormones and the Control of Growth and Development

165
chiefly in length, because the hoop arrangement prevents to prevent ethylenefrom accumulating in the atmosphere
a large increase in cell diameter. in would trigger ripening. When a fruit goes
quantities that
What doesthis have to do with ethylene? In some ot the through the phase of rapid ripening, the production of
responses which ethylene induces a thickening, as in
in ethylene may increase many times beyond the rate that
the shoot that is trapped beneath an obstacle, the effect was initially necessary for it to reach the critical triggering

seems to depend on a change in the direction of cell level. Thus, "one bad apple in a barrel" producing
growth. Ethylene causes some of the cells to deposit their ethylene may trigger the ripening of all the apples in the
new cellulose microfibrils more randomly, resulting in the Or the ethylene produced by the mold on an
barrel.

consequences mentioned above. orange may be enough to trigger undesirable changes in

We have no idea at present how the protoplast, inside the rest of the oranges.
the plasma membrane, controls the orientation of wall
polymers that are laid down outside the plasma Abscisic Acid
membrane, or how an information store such as DNA can
specify such a directional process. But the effect of Many plants of the temperate zone become inactive or
ethylene seems to offer a step toward understanding these dormant during the cold season. In addition, seeds are
problems. usually dormant at the time of release. Even in active,
Another effect of ethylene is seen in the growth of photosynthesizing trees in midsummer, the terminal buds
young trees, where the relationship between height and on the young twigs may go dormant after producing a
diameter of the trunk is very much a function of the sufficient number of leaves to clothe the tree. These
motion it undergoes as a result of wind. If a young tree phenomena illustrate the importance of dormancy and its
stem is tied to a rigid stake, as is frequently the case in release in the plant's repertoire of control and capabilities.
container-grown trees, or is supported by other plants in a Although the mechanisms that occur during dormancy
crowded nursery, the usual response is a tall, slender, are not fully understood, one factor that is often involved is
weak stem. the stake is flexible enough to allow some
If thehormone abscisic acid. Abscisic acid (Fig. 8.5D) is so
movement in response to gentle winds, the plant has a named because is plentiful in leaves that have recently
it

sturdier form resembling one grown without support. The been dropped or abscised. Applications of abscisic acid

plant that grows without support from a young stage (often abbreviated ABA) to some plants will cause leaf and
typically has a shorter stem, larger in diameter, and with a fruit abscission.
tapered form coming from a wide base at the soil level. Some dormant buds and seeds are also rich in ABA,
Routine exposure to high winds can greatly increase this and their release from dormancy is correlated with a loss
response, as can be seen in trees on windy sea coasts or of ABA. In some seeds the ABA may be lost by leaching

mountain ridges. Controlled gassing of tree trunks with as seeds are exposed to rains. In other seeds and
ethylene stimulates enlargement, and it is thought that dormant buds, ABA may be gradually broken down by
stress-induced ethylene synthesis in the trunk contributes enzymes during the winter.

to this growth response. Just how ABA causes dormancy is not entirely clear. In

some instances seems that dormancy is actually due to

it

Ethylene and Fruit Ripening a lack of gibberellin. Where this is true, dormancy may be
overcome by adding gibberellin. ABA is removed (e.g.,
If

During the early development of a fruit, the seeds are by leaching), the level of gibberellin in the tissue increases
immature and delicate. Dispersal at this time would be as a prelude to the release from dormancy.
premature. Many fruits have a programmed pattern of When transpiration exceeds water absorption, the water
changes that convert the fruit from a seed-manufacturing content of leaves decreases, causing a loss of turgor and
to a seed-dispersal organ. These changes are called wilting. It has been shown that under water stress in less
ripening. than half an hour, the leaf may begin to make and
Often the fruit changes its color from green to yellow, accumulate abscisic acid. Within a few hours, several
orange, red, or blue. This increases visibility to potential species display a manifold increase in ABA concentration.
animal dispersal agents. To make this change chlorophylls Ithas also been shown that ABA fed to leaves with open
are broken down, and other pigments such as the red and stomata can start stomatal closure in just five minutes.
blue anthocyanins are synthesized. Another change is the Abscisic acid appears to act by interfering with the uptake
conversion of starch and organic acids to sugar so that or retention of potassium (or sodium) in guard cells. Since
the fruit becomes sweet. Some of the materials in the cell potassium ions are required to provide the high turgor that
walls are broken down; the cells now become more guard cells need to maintain their stomata open, a lack of
loosely bound to each other, resulting in a softer fruit. these ions will close the stomata. When water loss by
Volatile flavor components are synthesized, and these transpiration is thus retarded, turgor in the leaf as a whole
contribute to much of the flavor we value. One important is regained, and the leaf recovers from wilting. The high
link in this grand plan is the rise in ethylene to a critical level of ABA is metabolized away in one or two days and
concentration within the fruit. When this critical the stomata begin to open normally.
concentration is reached, ripening is initiated. Usually this
is signaled by, or coordinated with, the maturation of the Interactions Between Hormones
seeds.
Ethylene may cause ripening when it is not wanted. In For each class of hormone, there are examples where one
fruit held in cold storage, elaborate precautions are used hormone alone seems to control a particular process: for

chapter 8 |
The Control of Growth and Development

166
instance, the control of growth by auxin in the oat Flowering
coleoptile. But more usual for the various hormones to
it is

interact in complex ways to control development. This has Pineapple plants can be induced to flower and form fruits
already been seen in the control of root and shoot on a precise schedule by applying auxin, ethylene, or a
formation in tissue cultures by auxin and cytokinin. Also, compound that breaks down to release ethylene. The
the effects of ethylene and auxin are often interrelated. An auxin acts by inducing ethylene production in the plant

experimental treatment of tissues with high auxin levels This aids the pineapple industry in coordinating

often induces the synthesis of ethylene Ethylene may then

harvesting, processing, and shipping The same principle
inhibit the transport of auxin. These results suggest that
can be used at home; an ornamental bromehad house
plant can be induced to flower by putting in an airtight
auxin and ethylene levels may mutually control each other
it

in many tissues.
container with a ripe apple as an ethylene source.

The control of flower morphogenesis in cucurbits

Rooting of Cuttings
(cucumbers, squash, melons) is a complex case of
hormonal interaction. A typical cucurbit plant first forms In many commercially valuable plants the preferred
male (staminate) flowers; at later nodes it forms perfect method of propagation is by means of cuttings either —
flowers. Some produce plants with only female
strains because the seeds are hard to germinate, or because
(pistillate) flowers. It has been shown that applied auxin valuable hybrid properties are lost in seed formation, or

raises the internal ethylene level, and that the ethylene because cuttings produce a large plant faster than a
stimulates the initiation of female flower parts on flowers seedling. A "cutting" is a detached shoot. To form a new

that would normally lack them, or it tends to make pistillate plant, it must form roots, usually at the cut end (Fig. 8.13).

flowers from normally perfect flowers. Gibberellin, either in In some species cuttings spontaneously form roots only if

naturally high concentrations or artificially applied, they are kept moist. Other species are harder to root, and
promotes the initiation of anthers. There appears to be an for them various methods have been devised as rooting
ethylene-gibberellin balance that acts at the time of flower aids. Treatment of the cut end with auxins such as
initiation to determine which organs the flower will have. naphthaleneacetic acid or indolebutyric acid is often

Ethylene and auxin may interact in controlling the effective. The bases of the cuttings are immersed in the
abscission of leaves and fruits. Ethylene causes an growth regulator solution for 12 to 24 hours or dusted with
abscission layer to form and mature at the base of the a dry powdered preparation; they are then placed in a bed
petiole. Auxin both promotes ethylene formation and of planting compound or moist sand. The conditions
blocks ethylene action. If the auxin supply drops, as it
required for rooting (e.g., hormone concentration and
does an aging leaf, ethylene can act, the abscission
in

layer forms, and the leaf soon abscises.

The Practical Use of Hormones

Natural hormones and synthetic compounds that act like
them are used in growing and marketing plant products.
Several examples are mentioned below. Bear in mind that
these are only a sampling of the compounds and methods
used.

Fruit Ripening

Ripe bananas store poorly. Today it is common practice to

and store bananas while they are still green.
pick, ship,
Gassing with ethylene triggers ripening, and the bananas
will be ready to eat in five days.

Bud Sprouting
Stored potato tubers are prevented from sprouting by
treatment with auxins. This is a case of bud inhibition,

comparable to natural apical dominance.

Fruit Drop

Fruits such as apples, pears, and citrus may drop from the
trees before harvest time. The fruits can be induced to

cling to the trees for several days longer by spraying with Figure 8.13 The promotion of root initiation by the synthetic
synthetic auxinlike compounds, chiefly naphthyleneacetic auxin, indolebutyric acid, on American holly (Ilex opaca) cuttings

acid and 2,4-dichlorophenoxyacetic acid (2,4-D). These Row A, cuttings that stood in an aqueous solution of 0.01 %
indolebutyric acid for 17 hours before being placed in sandy
compounds apparently retard the formation of the rooting medium. Photographed after 21 days. Row B, untreated
abscission layer at the base of the petiole. controls

Hormones and the Control of Growth and Development

167
treatment time) vary with the species. Interested amateur Seed Germination
gardeners can buy these chemicals at retail nurseries.

Some seeds do not germinate in darkness. After a brief

Growth Regulators as Herbicides
soaking, a short exposure to red light will induce them to
Though auxin is essential in small quantities, an excess germinate. Red light acts by converting P r
to Pu which
, in

can cause serious physiological disturbances. Some turn relieves some unknown block in metabolism and
synthetic auxinlike compounds have proven to be very permits germination. If far-red light is given immediately
potent weed-killers (herbicides) for this reason. The so- after the red treatment, Pu is converted back to the
called "phenoxy" compounds are especially effective: for inactive P r
form and the seeds will not germinate. For this
instance, 2,4-D and 2,4,5-trichlorophenoxyacetic acid latter cancellation effect, the far-red treatment must occur
(2,4, 5-T). These compounds are selective, killing broad- before P,r has had time to act.
leafed plants and leaving the grasses relatively unharmed. Responses to light that are cancelled by a second dose
This makes them useful for destroying dicot weeds in are said to be photoreversible. Red/far-red
grain fields, dandelions in lawns, and brush in rangeland. photoreversibility is a unique phenomenon that is seen
only in events governed by phytochrome.

Lightand the Control of

Development Shoot Growth

The main signal to a seedling that ithas emerged from the

Plants have at least three pigment systems that initiate soil is the formation of P,r by light. The complex response
developmental changes in response to light. of the seedling has already been described. In these
Phototropism, the directional control of growth by light, de-etiolation responses the action of P may involve such
fr

involves an unknown pigment and has been described factors as the release of active gibberellin from an inactive
previously. The greening of plants when they break out of form and the activation of genes to form new enzymes.
the ground is initiated by a pigment called proto- When a plant finds itself almost continuously in the
chlorophyll, which was discussed at the beginning of the shade of other plants, the quality of light falling on the
chapter.
plantis changed in a way that can markedly affect the
The and by far the most versatile pigment system
third
phytochrome status of the tissue. As light passes through
that regulates development is the phytochrome system.
successive layers of leaves, the red wavelengths are
Phytochrome is a pigment that exists in two stable forms,
filtered out by chlorophyll, while the far-red wavelengths
active and inactive, each having a different color. The
pass through. In the shade created by other plants, light
inactive form absorbs red light most strongly and is called
rich in far-red tends to markedly lower the P,r
Pred or Pr and the active form is called P,ar red or Pu .
concentration. The result is more rapid stem elongation
,

because it most strongly absorbs the far-red light that lies

and an increased probability of growing out from under
just at the extreme of human visual sensitivity. When
the shade that is cast by other plants. Even in plants
either form absorbs light, it is converted to the other form.
grown in light, phytochrome is present, and experiments
have shown that exerts subtle controls on stem
it

elongation. In this example, a change in light quality can

slow destruction rapid destruction trigger a response that has obvious survival advantages.
enzymatic / rori Although the mechanism of action of phytochrome has
P-- — Pi, (active)
synthesis
not yetbeen established, some work suggests that it may
(dark reversion in some plants) be attached to membranes and may exert some control
over membrane function.

P continuously synthesized from precursors, and

r
is

slowly destroyed by enzymes. P,r may and exert its

exist
is
Photoperiodism and Flowering
effect for many hours, but is unstable and is more rapidly
destroyed by enzymes than is Pr In some plants, P,r is

spontaneously converted back to P, in darkness. Over most of earth's surface, there are pronounced
Continuous sunlight with its broad spectrum of seasonal differences in temperature, water supply, and

wavelengths constantly converts the available illumination. The developmental system of the flowering

phytochrome molecules back and forth between the Pr plant is equipped to prepare for these changes. Well
and Pu forms, so that a steady-state mixture results with before the weather is harsh, broad-leafed trees begin to
the P lPu ratio determined by the spectral composition of shut down the anabolic systems in the leaves. There is an
;

the light. orderly withdrawal; the leaves are stripped of some

Phytochrome has a wide variety of effects on plant nutrients, abscission layers are formed, and the leaves are
development, of which the following are important abandoned to the elements. If the plant can make
examples. underground storage organs, they are filled up before

chapt er 8 |
The Control o f Growth and D evelopment

168
there is a great danger of the loss of food. In addition the shown that the plants are really measuring the lengths of
metabolic system gradually shifts its composition to protect nights rather than days. But the names are too firmly
against frost damage before ice appears. The broad-leafed embedded in the language of physiologists to be changed.
tree buttons down for the winter and goes dormant. After Long-day plants usually flower in the spring when nights
the cold season, the plant still waits until it is well into the are growing shorter (Fig. 8.14). In these plants the signal
fair season before stirring to active life, disdaining the for flowering is sent from leaves to shoot tips when the
occasional day of false spring that occurs in late winter nights are less than a particular critical length. The critical
before the weather is reliable. night length is characteristic of the species and in general
Not all plants are so good at predicting seasons. But on is so that the plant has a suitable period of vegetative
set
the other hand, the preceding comments were scarcely a growth in the spring before flowering. Thus plants in the
sampling of the seasonal adjustments that plants can more northerly regions, where winter ends later, tend to
show. There are desert plants, tor example, that prepare have shorter critical night lengths; they therefore flower
themselves for a dangerously hot spell rather than a later in spring.
dangerously cold one. And more subtly, plants may time Short-day plants, such as many chrysanthemums,
their activities during the growing season in a way that typically time their flowering for fall or winter when the

minimizes competition for light, moisture, and pollinators. nights are progressively lengthening (Fig. 8.15). Here
This also may be based on a sensing of the seasons. again there is a critical night length, but in short-day
Of all the earth's seasonal variables, the most reliable is plants the signal for flowering occurs when nights become
the annual cycling of day and night lengths. Plants have longer than the critical length. These plants fail to flower in
been found to contain a system that measures the lengths the shorter days of early spring because at that time they
of the nights. The control of development by this timing are immature.
system is called photoperiodism. Presumably the advantage of flowering in the fall is that
All forms of photoperiodism are based on the system the plants have been able to accumulate photosynthetic
that monitors the day/night cycle. That system is located products all summer for better seed production. A well-set,
in the leaves. Given suitable light/dark cycles, accurate timing system should choose the optimum time
photoperiodically sensitive leaves send out stimuli through to initiate flowering so that the plant has the longest
the phloem to other parts of the plant: to the shoot apex, possible period of active growth while still allowing time for
where flowers or winter buds may be induced; or to seeds to mature before the cold sets in.

underground stems, which may be induced to form tubers. It is not entirely understood how the leaf measures
To explore photoperiodism, then, the principal focus of lengths of night. We do know that phytochrome signals
attention is on the leaves and their timing system. But to the end of the day. During the daylight hours there is
illustrate its operation in the passages below, let us take always some P,r present, but at night the Pfr gradually
the control of flowering as our example. disappears. It seems that the disappearance of Plr
In regard to flowering, some plants achieve their timing somehow "tells" the plant that night has begun. The next
without reference to light stimuli, relying instead on factors light exposure results in new P being formed; this acts as
lr

such as maturation. These plants are said to be day- a signal that the night is over. Since P formation requires
fr

neutral. Other plants, however, use photoperiodism to only a little light, even a brief flash of red light during the
time the flowering response. These plants fall into two night can seriously interfere with the timing of night
major groups that are traditionally designated as long-day length. As would be expected with a phytochrome-
and short-day plants. Actually, current research has

Figure 8.14 Effect of day length on behavior of the long-day

plant henbane (Hyoscyamus). Plants were grown with 8 hour
photoperiod until they were about a month old and were then
subjected to 24 photoperiods of 10, 11, 12, 13, 14, or 16 hours.
Initiation of flowers and accompanying stem elongation (bolting)
occurs on days longer than 1 2 hours.

Photoperiodism and Flowering

169
 run long enough to exceed the critical night length. The
answer determines whether or not the leaf will send a
flowering stimulus to the shoot apex.
In considering the flowering stimulus we have only
mentioned that it is made in the leaves and moved to the
apices. We do not yet know its chemical nature. Some
experiments strongly suggest that the same flowering
stimulus serves long-day, short-day, and day-neutral
plants. They apparently differ only in the conditions
required to bring about its synthesis.
Whatever the nature of the flowering stimulus, it must be
produced by a metabolic system that we might call a
stimulus generator. In short-day plants the stimulus
generator is activated if the clock passes its critical point
during one or more successive nights (Fig. 8.16). To
produce a short-day flowering habit, by contrast, the
stimulus generator must be set to produce its flowering
stimulus automatically if the clock does not pass the
critical point for one or more nights in succession. That is,

long-day and short-day plants differ according to whether

the clock activates or inhibits the stimulus generator when
the critical night length is exceeded.

Temperature and Development

Photoperiodism is not the only system that times flowering.

Some plants require a prolonged exposure to winter cold
as a prerequisite to flowering.
Figure 8.15 Effect of day length on behavior of
Chrysanthemum, a short-day plant. A, plant that received light of Plants having this type of control include both winter
natural short days of autumn and blossomed at the usual time. B, annuals and biennials. Winter annuals typically germinate
plant that received an hour of light near the middle of each night
during the winter so that the young plant
just prior to or
for several weeks beginning just before flower buds would
normally have been initiated; thus, each long dark period was passes through a period of weeks of exposure to low
divided into two short ones. This interruption was sufficient to temperatures prior to the warmer weather of spring and
delay flowering.
summer when they flower.
Biennials generally have a full season of vegetative

growth the first year, exposure to cold weather during the

mediated response, the effect of a red light flash at night next winter, and then they flower in the second spring and
can be cancelled when it is immediately followed by a summer. Without exposure to the low temperatures,
flash of far-red light. flowering be delayed or prevented (Fig. 8.17).
will

Though phytochrome signals the beginningand end of Gibberellins can sometimes substitute for the cold
night, it appears that in darkness some other component requirement. This promotion of flowering in response to
in many plants actually measures the time. This timing cold treatment is called vernalization. The cold treatment
system has been called the biological clock. Its nature is generally requires temperatures of 10°C or less for several
unknown, but it is highly accurate and insensitive to such weeks. Often, the cold requirement is followed by a
disturbing factors as differences in temperature and requirement for long days. This prevents the plant from
nutrition. Some plants can discriminate between night flowering too early in spring. The growing tip itself, rather
lengths that differ by as little as 10 minutes, a degree of than the leaves, may perceive the cold stimulus in

precision that allows flowering to occur within a week of vernalization.

the same date every year. "Winter" cereals, wheat and rye, are normally planted in

The biological clock is a metabolic system that the so that they germinate before the onset of winter
fall

completes a cycle approximately once each 24 hours. It and flower promptly in the following spring and summer
synchronizes many physiological functions on a daily aftermaking a minimum number of leaves. If they are
cycle and is present in organisms ranging from humans to planted in the spring and are not vernalized, flowering is

algae. delayed until many more leaves are formed. "Spring"

In regard to the timing of flowering, the disappearance cereal strains are normally planted in the spring and flower
of P fr
after nightfall somehow provides a time marker that promptly in response to long days. Winter cereal strains
the clock uses to begin to measure night length. At dawn can be artificially vernalized prior to a spring planting date
P lr
is re-formed and the timing is stopped. For the control by moistening the seeds and holding them at about 1 °C
of flowering, the important point is whether the clock has for a period of weeks.

chapter 8 I The Control of Growth and Development

170
 critical night
length

K
1
—>- inhibitory
signal
stimulus
generator
flowering
stimulus
1

1
to apex

promotory stimulus flowering

\ — signal generator stimulus
>~ to apex
1

Figure 8.16 Control of flowering in long-day (A) and short-day

(B) plants. Details are hypothetical. In both cases the timing of
night length is started by the disappearance of P after nightfall.
fr
In both, the same kind of flowering stimulus originates from a
metabolic "generator" in the leaves and moves to the shoot apex.
Whether a stimulus sent depends on whether the clock reaches
is
the critical night length measurement before being interrupted by
light. The plants act as if the clock sends a signal on reaching the
critical length; short-day and long-day plants differ as to whether
the signal promotes or inhibits the stimulus generator.

In the discussion of photoperiodism, we indicated that

many woody perennial plants use the photoperiod to
induce a dormant condition in the bud prior to the onset of
unfavorable weather. The bud typically goes through a
period of increasing dormancy until it can no longer be
rapidly reactivated by favorable environmental conditions
or a shock treatment such as defoliation. By fall may be it

in in which no treatment will activate

a state of rest other it

than the passage of time at low temperature, a process

called chilling. The chilling requirement generally requires
temperatures a little above freezing but below 10°C.
Normally, the requirement is met by midwinter and at that
time a branch brought into the greenhouse will have buds
bursting in a few weeks. The buds are no longer in a
condition of rest; they can respond to the warmer days of
spring with renewed growth. The photoperiod induced the
dormant condition, and the chilling process signaled that it

was safe to respond to springlike weather.

need 250 to 1000 hours of chilling
Fruit trees typically
to produce good growth and fruit production in spring and
summer. The lack of sufficient cold weather sets the Figure 8.17 Substitution of gibberellic acid for the cold
southern limit for peaches, apricots, cherries, plums, and requirement (winter) in the flowering of the biennial carrot
(Daucus carota). A, control plant under long days only; 6, long
apples, and for effective flowering of ornamentals such as days plus gibberellic acid, no cold treatment; C, long days plus
lilac (Syringa vulgaris). cold treatment, no gibberellic acid.

Temperature and Development

171
 Another temperature-related phenomenon is the Cytokinins control cell division in specific tissues, they
development ot frost hardiness. In the summertime, interact with auxin to control organ initiation and bud
temperatures that are a few degrees below freezing will kill growth, they help to prevent aging in leaves, and they
leaves and buds. During the fall, many plants gradually help to coordinate root and shoot growth.
develop resistance to temperatures as low as 50 °C below 10 Ethylene interacts with auxin to control stem
freezing. Short photopenods and exposure to temperatures elongation. It also helps to maintain the etiolated
just above freezing are the stimuli that induce frost growth habit in darkness and to regulate ripening in

resistance. many fruits.

The seeds of some species will not germinate until they 11 Abscisic acid helps to regulate the leaf's response to
have been exposed to several weeks of temperatures near water stress through stomatal closure, contributes to
freezing. Such a cold, wet treatment of seeds to promote seed dormancy, and promotes abscission in leaves
germination is called stratification. and fruits.
12. Light acts as a developmental signal when it is
Summary absorbed by specific photoreceptor pigments. The
1. Plants are able to respond to a wide variety of chief receptor pigments are phytochrome, the

environmental situations because they have systems phototropic pigment, and protochlorophyll.
for monitoring important environmental factors and 13 Phytochrome exists in active and inactive forms that

they have systems that modify the course of are interconvertible by red and far-red light.

development in response to these stimuli. Phytochrome senses when light is present or absent.
2. The control of seed germination and the control of When
stimulated by light, it starts seedlings on the

growth and development in the seedling are instances development of the above-ground mode of growth. It
in which developmental responses to light, gravity, may inhibit stem elongation, promote leaf expansion,
temperature, and other environmental cues have promote the germination of seeds, and participate in
survival value. many other developmental events.
3. The coordination of activities between various parts of 14 Photoperiodism, the control of development in
the plant, and the course of development within each response to the lengths of days and nights, is the
cell and organ, are controlled in part by the production basis for many seasonal adjustments in plants.

and distribution of hormones. 15. The photoperiodic timing system occurs in the leaves
4. Plant hormones are natural compounds that act in and seems to involve a "biological clock" that is
small quantities as signals to regulate development. governed by phytochrome.
5. Auxin, the gibberellins, the cytokinins, abscisic acid, 16 There are two main types of photoperiodically

and ethylene are the major plant growth hormones. controlled plants, as regards flowering: long-day and
6. Auxins promote cell enlargement by altering the cell short-day plants, which respond respectively to night
walls, making them more extensible. Growth is lengths shorter than critical or longer than critical.
controlled by factors that affect the synthesis, Many plants are day-neutral; in them, flowering is not

transport, and inactivation of auxin. Phototropic and photoperiodically controlled.

geotropic curvatures in the shoot are regulated by 17 Buds may be induced to go dormant by a
changing the transport of auxin to the growing cells. photoperiodic system and can be activated again by
7. Auxins also help to control other developmental favorable conditions only after passing through a
processes such as apical dominance, cell division, and prolonged chilling period, which is normally provided
differentiation of specific cell types in the vascular by the winter season.
system. 18 A prolonged period of low temperature may be needed
8. Gibberellins control stem elongation through effects on to vernalize certain plants, that is, to remove a block
cell elongation and division. They also affect seed that prevents the formation of flowers by the shoot
germination and cambial activity. In certain instances apex. Low temperature may also be a requirement for
they may induce the synthesis of enzymes. seed germination.

chapter 8 |
The Control of Growth and Development

172
 9 CHAPTER

9 plant ecology

Through evolution, existing plant species have composed of all the populations in a given habitat. An
achieved a balance with their environment. In each ecosystem consists of the living community plus the
type of habitat, certain species group together. nonliving factors of its environment.
Fossil records indicate that similar groups of species have
lived together for millions of years. The species of these
groups share incoming solar and
radiation, soil, water,
Components of the
nutrients toproduce a relatively constant amount of living
matter. They recycle nutrients from the soil to living tissue
Environment
and back to the soil again; and they divide the
environment's resources. Plant ecology attempts to explain
The environment of an organism includes all of the living
this balance between plants and their environment. For
and nonliving things around Some important
it.

example, what stresses does the environment put on

environmental components are moisture, temperature,
plants, and how do plants respond to them? Plant ecology light, soil, and living organisms, and all of them work
also attempts to determine how this natural balance may separately and jointly to influence plant distribution and
be artificially imitated or manipulated to improve human behavior. Species differ in their range of tolerance to these
life.
environmental factors. For example (Fig. 9.1), coast
Plant ecology deals with levels of biological organization redwood (Sequoia sempervirens) is restricted to a narrow
beyond the organism level: the population, the community, and adjacent southwestern Oregon,
strip of California
the ecosystem. A population is a group of closely related about 9500 square km, that experiences heavy summer
organisms that normally interbreed. A community is fog. Coast redwood is not tolerant of variations in moisture

Figure 9.1 Distribution of coast redwood (Sequoia sempervirens) and red maple (Acer rubrum) in the United States.

173
or temperature. Red maple (Acer rubrum), however, Moisture
occurs over about 4,000,000 square km of eastern North
America in wet, dry, cold, or warm environments. It is Water is a most important factor in determining both the
tolerant of wide variations in moisture and temperature. distribution of plants over the earth's surface and the
There are two types of environment. The macro- character of an individual plant. Probably no single factor
environment is influenced by the general climate, is so largely responsible for the abundance of plants in a
elevation, and latitude of the region. Weather Bureau data habitat as is the supply of water. So important are the
on rainfall, wind speed, and temperature are measures of water relationships of plants that various attempts have
the macroenvironment. Measurements are taken at a been made to classify plants on the basis of these
standard height of 1 .5 m above ground in a clear area relationships. One such classification divides plants into
away from buildings or trees. (a) xerophytes, which are able to live in very dry places,
The microenvironment is the environment that is close (b) hydrophytes, which live in water or in very wet soil,
enough to the surface of an organism or object to be and (c) mesophytes, which thrive best with a moderate
influenced by it. For example, bare soil tends to absorb water supply.
heat; consequently, the temperature just above or below Plants with xerophytic characteristics, which limit

the soil surface is much higher than air temperature (Fig. transpiration or in other ways closely control water
9.2). Light quality and quantity are much different for balance, occur in different climatic zones, especially
herbs beneath a forest canopy than for the leaves of the deserts. Xerophytic plants do not necessarily have a lower
canopy itself (Fig. 9.3). Air as far as 10 mm from the transpiration rate than do mesophytes when water is
surface of a leaf on a still day is less turbulent and higher ample, but they do possess one or more characteristics
in humidity than free air further away from the leaf. In that enable them to survive periods of drought. The most
marshy areas, where the water table is close to the effective of them are a thick cuticle, early or daytime
surface, minor dips or rises in the topography greatly stomatal closure, reduction of the transpiring surface, and
influence the root microenvironment. Plants growing in water storage tissue.
shallow depressions are often subject to more frequent The time of precipitation is as important to plant growth
frost than those growing on higher ground, because cold as the total annual amount. For example, tropical rain
air will settle in the depressions. Ecologists are convinced forests and savannahs may both receive the same
tropical
that the microenvironment is as important for plant growth total yearly rainfall —
let's say 200 cm —
but the forest
as the macroenvironment. receives an equal share of the total each month, while the
savannah has pronounced dry and wet seasons. The
120 • 120 result is that the two vegetation types are quite different:
100 — 100
:30 P M. tropical rain forests support tall, evergreen trees and vines
80 J3:30
A.M. : 1
- 80
Air
in profusion, but tropical savannahs support grasses,
S 60 60
•5 40 40 shrubs, or short trees, which are often deciduous.
1 20 — 20 Plants adapt to dry climates in several ways. Cactus
c
8 o plants (Fig. 9.30) develop water storage tissue in the stem
20 : 20
Soil
_ 40 and reduce transpiration by the loss of leaves; all their
40
60 1 1 1
1 1 1 1
1 1
60 photosynthetic activity is performed in stem tissue.
55 65 75 85 95 105 115 125 135 145 155
Mesquite shrubs (Prosopis) and salt-cedar trees (Tamarix)
°F
possess long roots that tap ground water, sometimes at
Figure 9.2 and soil temperatures at several positions just
Air
depths below 53 m. Epiphytes, growing on tree trunks
above and just below the soil surface. Temperatures were
measured during the warmest and coolest parts of the day. above the soil (Fig. 9.4), trap falling rain in leaf axils or in

Wavelength, microns
12.0X10'V
0.3 0.4 0.5 0.6 0.8 1.0 2.0 10.0

5 -
? 10.0 X10"
E
u

.E 8.0 X10" 5
E

Direct sunlight
Jf 6.0 X10" 5
o (Sea Level)
u
aT
5
c
o
4.0X10
o
~ I'":
2.0 X10" 5 Light transmitted

through vegetation^*"}

L
Figure 9.3 Distribution of solar radiation energy by wavelength.
Energy was measured in direct sunlight and beneath a forest
canopy.

chapter 9 |
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174
 Absorption of water by roots is slower in cold soil than in

warm soil. At high temperatures, evaporation may remove

much of the soil moisture before the new growth of
shallow roots can reach it, and transpiration will also be
stimulated, resulting in further rapid depletion of soil water.
Brief extremes in temperature may be more important in

determining plant distribution than are long-term moderate

temperatures. Palms and many cactus plants seem
excluded from areas that experience a specific frequency
of frost. California lilac (Ceanothus megacarpus) seeds
germinate poorly unless they have been exposed briefly to
high temperatures. Many temperate zone plants are not
frost-resistant until they have been exposed to moderately
low temperatures for a short time and thus "hardened."
Other plants require an alternation of day and night
temperatures (a thermoperiod) for best growth.
Topography and air temperature.
greatly influences soil
In mountains latitude and elevation combine to restrict

species to certain belts (as shown for mountain hemlock

in Fig. 9.5). Even at the same latitude and elevation, minor

differences in the direction of slope (aspect) create

different microenvironments. A gorge running east-west
through an Indiana forest was studied to illustrate the
effect of aspect on plant growth. The gorge was 65 m
wide at the top and 45 m deep; its sides supported
scattered trees, shrubs, and herbs. Meteorological
Figure 9.4 Tillandsia, a genus with many epiphytic species
common to tropical and subtropical regions of the western instruments were placed 1 5 cm above and below the soil
hemisphere. surface midway down each side. During spring, the south-
facing slope was found to exhibit a greater daily range of
topsoil temperature, a higher mean air temperature, a
dead on the surface of roots. These epiphytes are
cells
higher rate of soil water evaporation, and lower relative air
not parasites; they depend on trapped water and debris for
humidity than the north-facing slope. Of nine spring-
all growth requirements. Annual herbs adapt to drought by
flowering species present on both banks, the average
completing a brief life cycle only during periods of
flowering time was six days earlier on the south-facing
Boerhaavia repens of the Sahara
sufficient soil moisture.
bank. A similar difference in flowering time could be
Desert is a small annual that can go from seed to seed in
achieved on level ground only over a distance in latitude
10 to 14 days, but most annuals have a life span of 3 to 8
of 180 km.
months.
Temperature is only an estimate of the heat energy
There is some evidence to show that plants can absorb
available from solar radiation. Solar radiation at the limits
dew through their leaves, but the amount of water 2
of our atmosphere is equivalent to about 2 cal per cm per
obtained in this way is not great. Usually, dew, fog, or
min. Much of this radiation is absorbed, scattered, or
high humidity are of more value in reducing the rate of
reflected within the atmosphere, and only half may reach
transpiration, than by physically entering the plant.
the ground and heat it. In turn, the warm earth reradiates
(Exceptions include small epiphytes, certain annual plants,
mosses, and lichens.)
—
energy back to space terrestrial radiation. The difference
between solar (incoming) radiation and terrestrial
Plants may also utilize fog by condensing the mist into
(outgoing) radiation is termed net radiation. During a
large drops that trickle down the stem or drip from the
clear day at the equator, solar radiation is greater than
branches, increasing soil moisture considerably. The
terrestrial and, thus, net radiation is positive; during the
amount of water added to soil in this way by trees on the
night, the reverse is true and net radiation is negative. But,
San Francisco Peninsula was measured as 5 to 74 cm in
over a 24 hr period, a month, or a year, net radiation may
one summer, the exact amount within that range
be positive or negative, depending on the location. For
depending on the type of exposure. Normal precipitation
(rain) for the area is only 64 cm a year, with almost no
example (Fig. 9.6), at Aswan, Egypt a warm desert —
rain in summer.
climate — net radiation each month, with a peak
is positive
in June. Net radiation is positive only five months of the

year at Turukhansk, Siberia a cold subartic climate —

and —
the annual net radiation is very close to zero.
Temperature
Temperature influences moisture availability and the rate
at which chemical reactions occur. At freezing Light
temperatures, water changes to ice and is unavailable for
plant growth; consequently, in cold climates one of the Figure 9.3 shows how solar radiation is changed in quality

principal factors limiting growth may be drought. and quantity by its passage through green leaves. Light

Components of the Environment

175
 Altitude (ft)

11, 000

10,000

9,000

8,000

7,000

6,000

5,000

_ 4,000

'=_ 3,000

_ 2,000

1,000

50
Latitude [°N]

Figure 9.5 Distribution of mountain hemlock (Tsuga

mertensiana), showing the relationship between altitude and
latitude.

palisade and spongy mesophyll layers, and achieve

maximum rates of photosynthesis at much lower light

intensities than do sun leaves. Recent biochemical

Turukhansk photosynthetic
evidence has shown that the difference in

rates is due to a difference in the activity of the enzyme

responsible for C0 2 fixation.

When a large tree dies in a mature forest, the canopy is

opened and increased light intensity reaches the ground

beneath. This increased light radiation induces seedlings
there to begin rapid growth. Seedlings must be able to
survive many years of slow growth in shade before such
an opening "releases" them. Eastern hemlock (Tsuga
canadensis) is remarkably shade-tolerant. Mature trees of
this species live for 1000 years. They can retain the
capacity for rapid growth when released from shade to an
Figure 9.6 Monthly net radiation of Aswan, Egypt and age of 400 years. Saplings only 2 m tall and 2 to 3 cm in

Turukhansk, Siberia. diameter may have a ring count indicating an age of 60

years.
Trees that are not shade-tolerant may become
established and grow to maturity if some disturbance such
intensity on the forest floor may be only 1 to 5% that of
as fire or logging first removes the shade. Seedlings of
full sunlight. Some herbs deciduous forest complete
of the
shade-tolerant species grow beneath them, however, and
their life cycle in early spring, before the trees above them
replace them as they die, while their own seedlings do not
develop their complete foliage and reduce light intensity. pure
remain alive. For this reason, it is difficult to maintain
Leaves of plants that develop in shade exhibit different
stands of valuable shade-intolerant species such as teak,
morphology, anatomy, and physiology than those that
mahogany, Douglas fir, or the southeastern yellow pines.
develop in sunlight, even when both are attached to the
These species only invade disturbed sites and do not long
same plant. Shade leaves are larger, thinner, contain less
maintain themselves.
chlorophyll per gram of tissue, have less well-defined

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176
 Light is also reduced in intensity and quality by passage stages from a young soil in which the processes of soil
through water. Algae are not commonly found below a development are continuing to a mature soil that has
depth of 60 m except in very clear water. A record depth approached a steady state.
for algae may have been found in Lake Tahoe, California. Soil formation may proceed rapidly if the type of parent
A sample of water and bottom mud from a 1 40 m depth material and type of climate are favorable. Soils form
contained the alga Chara. Light intensity at that depth, rapidly where there is a warm, humid climate, forest
despite the water clarity, is only 0.1% of full sunlight. vegetation, flat topography, and a parent material that is
Brief exposure to light may be as important for some easily broken down. For example, Fort Kamenetz, built of
plants as long-term exposure. Recall from Chapter 8 that limestone slabs in the Ukrainian part of the Soviet Union,
seeds of some species can be stimulated to germinate in was abandoned 1699. Today, a mature soil 10 to 40 cm
in

the dark if exposed briefly to full sunlight; this

they are first thick has developed from the limestone. Soils form slowly
is done by simply shaking them for a few seconds in soil in regions with a cold, dry climate, grass vegetation,
held in a glass container. This treatment could be steeply sloping topography, and parent rock material that
duplicated nature by a plow turning the soil and seeds
in is not easily broken down. It has taken from 1000 to
in it, exposing the seeds to sunlight, then reburying
briefly 10,000 years for mature soils to develop in northern areas
them. Plowing aerates the soil, makes it more permeable scoured during the Wisconsin glaciation.
for root growth, and removes plants at the surface — all Plants influence soil development in several ways. They
conditions of advantage to an emerging seedling. move ions through the soil by absorbing them through
their roots. These ions accumulate in the leaves and are
returned to top soil after leaf abscission. Plant roots and
Soil shoots decay and add nitrogen and sometimes acids to
the soil. In addition, plant canopies shelter the soil surface,
Soil is the part of the earth's crust that has been changed reducing erosion and water loss.
by contact with the living and nonliving parts of the The texture of soil is determined by the mineral
environment. It is a weathered, superficial layer typically 1 fractions of a soil; we customarily speak of coarse sand,
to 3 m thick, made up of decomposed and partly finesand, silt, and clay. The size of soil particles
decomposed parent rock material with associated organic decreases in the order given (Table 9.1). Clay is not only
matter in various stages of decomposition. There are many the smallest mineral soil particle, but because of chemical
kinds of soils and many different soil conditions. The weathering it has also undergone the most change from
character of natural plant covering and the behavior of the parent rock.
crops depend on soil conditions as well as climatic Although most soils contain both mineral and organic
conditions. Environmental influences that operate through matter, the organic matter they contain constitutes only 2
soil are called edaphic factors; those that act on the plant to 5% of their weight. These predominantly mineral soils
through the atmosphere are called climatic factors. may be classified according to the relative amounts of the

Each environment creates a unique soil type. These soil sand, and clay they contain. Soils that contain roughly
silt,

types have their own history of development, morphology, equal amounts of sand, silt, and clay are called loams.

and chemical attributes. In the United States alone, there Depending on the relative amounts of each component in
are 10,000 soil types. the soil, we speak of clay, clay loam, silt loam, loam,
sandy loam, loamy sand, and sand textures (Table 9.2).
Soil Formation The suitability of a soil for plant growth is greatly
Most soils consist primarily of mineral particles that are influenced by its sand, silt, and clay content. Organic
soils are defined as soils having a layer, at least 30 cm
formed by a slow, continual process of weathering of the
thick, that is more than 30% organic matter.
parent rock. Mineral matter is derived from fragmented
rock. The kind of parent rock (e.g., granite, lava, Soil Water and Its Dissolved Substances
sandstone, limestone, and shale) and the degree of
An important difference among the various soils is their
weathering determine the nature of the mineral or clay and organic soils,
ability to hold water. It is greatest in
inorganic components of the soil.
and least in coarse sand.
Weathering may involve simply mechanical breaking of
the parent rock. For instance, the action of strong winds
or wave action may hurl sand against rock outcroppings Table 9.1
and wear them down, or water collecting in pores of the
Classification of Soil Mineral Matter According to Size
rocks may expand at freezing temperatures to create
of Particles (International System)
fissures that ultimately fragment the rock. Chemical
weathering results in more profound changes in the Range in Diameter
mineral matter itself. Atmospheric gases, such as C0 2 or
of Soil Particles,
S0 2 become dissolved in rainwater and produce acidic
,

Type of Particles mm
solutions that dissolve the parent rock material. Plant roots
secrete weak acids. Certain algae, bacteria, and lichens 2.0-0.2
Coarse sand
hasten the decomposition of the least resistant mineral 0.2-0.02
Fine sand
materials.
Silt 0.02-0.002
Weathering of the parent material, under the influence of 0.002 and smaller
Clay
climate and organisms, proceeds along a definite series of

Components of the Environment

177
Table 9.2
Composition of Three Soils According to the Size of
the Mineral Particles

Coarse Fine
Soil Type Sand Sand Silt Clay
(texture) % % % %
Sandy loam 67 18 6 9 15

Loam 27 32 21 20 '5

o
Clay 1 9 22 68 a
>

If water is the field, the spaces

applied to a soil in

between the (pore spaces) become filled

soil particles

only for a short time to the depth wetted. With drainage,

the water begins to move downward under the influence
of gravity. After a while, this movement downward stops;
the soil particles hold a certain amount of water against
4 5 6 7 8 9 10
the pull of gravity. The amount of water held by the soil

after drainage is called the field capacity of that soil. Figure 9.8 Relationship of relative nutrient availability to soil pH.

When a soil is at field capacity, or even at a moisture

content well below this amount, a film of water completely
surrounds each soil particle, and water also exists in the Acidity per se — that is, H+ or OH
-
ions — is not directly
form of wedges between soil particles. Clay particles and responsible for limiting plant growth. Rather, soil pH
organic matter in the soil are colloidal particles, and such affects the availability of plant nutrients in In an two ways.
particles hold water by imbibition. Water imbibed by soil acid hydrogen ions (FT) replace other cations (like
soil,
particles is much more difficult to remove from the soil K + Ca 2+ and Mg 2+ ) on the negatively charged clay
, ,

than water that exists as a film or as wedges. particles, allowing the nutrient ions to float free in the soil
Soil particles themselves, particularly finely divided clay water and to be easily leached from the soil as that water
and organic matter, act as giant negatively charged ions moves downward. Soil pH also affects the solubility of
that attract a cloud of positively charged ions such as plant nutrients. As shown in Fig. 9.8, some nutrients (e.g.,
Ca 2+ and K + Thus, colloidal soil particles, while not in
.
iron, manganese, and aluminum, increase in solubility as
solution themselves, serve as reservoirs to which many of pH decreases. Aluminum may even reach toxic levels in
the various ions essential for plant growth are loosely some acid soils. Other nutrients (e.g., calcium and
attached and from which these ions may be released into magnesium), increase in solubility as pH increases.
solution by a process known as cation exchange (Fig. Extremes of soil pH may also affect plant growth indirectly
9.7). by suppressing bacterial growth.
Acidity influences the physical properties of the soil, the
availability of certain minerals to plants, the biological
Soil Profile Development
activity in and consequently strongly influences
the soil, In wet temperate regions, such as those that support

plant growth. Certain plants such as azaleas, camellias, dense coniferous forests, soil development is principally
and cranberries grow best in acid soils; most plants do caused by percolating rainwater that continuously
best in soils near neutrality, while a few plants grow dissolves nutrient salts in the upper portion of the soil and
satisfactorily in slightly alkaline soils. carries them, along with particles of finely divided,
Acid such as those beneath northern coniferous
soils, chemically weathered mineral matter and bits or organic
trees that shed acidic foliage, range from pH 3.5 to 5.0; matter, downward into the lower levels of the soil. This
agricultural soils in the humid areas range from pH 5.0 to process is called leaching. In time, the soil consists of a
7.0; and soils in arid or saline regions may reach pH's as series of superimposed layers or horizons that differ in

high as 1 1 .0, but a range of 8.0 to 9.0 is more common. color, texture, and chemical attributes (Fig. 9.9).

2K +
7H + + Ca 2+
2+
M + NH +

Clay or organic matter Clay or organic matter

Hydrogen ions Various cations
with various ions with hydrogen as +
in soil solution in solution
as cations cations

Figure 9.7 Cation exchange on soil colloids.

chapter 9 |
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178
 Table 9 3
) litter The Derivation and General Significance of the Names
of the 10 Soil Orders

Order Derivation
Name of Name Characteristics

Entisol Artificially Soils of very limited

created development. Profile properties
largely inherited from the parent
material.

Inceptisol L. inceptum, Soils exhibiting moderate

beginning development. Identifying
horizons are of types that form
relatively quickly.

Aridisol L. aridus, Soils of arid regions. Limited

dry change in parent material
because of low climatic intensity.

Mollisol L. mollis, Soils with friable surface

soft horizons noticeably darkened by
organic matter. The degree of
base saturation is high.

Spodosol L. spodus, Soils with illuvial 62 horizons

wood ash having significant accumulations
of free Fe and Al oxides,
noncrystalline clays, and humus;
usually without structure and
Figure 9.9 Profile of a typical soil in a wet, temperate region.
often partially cemented.
The A horizon, except for a thin region just beneath the litter, is
leached of organic matter, clay, and salts. These leached Alfisol Artificially Soils containing a B horizon with
substances accumulate in the 6 horizon. The C horizon, removed created significant amounts of crystalline
from weathering processes, represents parent material from
which the A and 6 horizons have been formed. clays and generally a moderate
to high degree of base
saturation.

Ultisol L. ultimus, Soils containing a B horizon with

last significant amounts of crystalline
clays but generally with a low
The upper, or A, horizon of soil beneath a conifer forest degree of base saturation.
is rich in organic matter only in the few uppermost
centimeters, where freshly deposited organic litter is Oxisol Gr. oxid, Highly weathered soils
decomposing. The lower portion of the A horizon is sandy acid, sharp containing a B horizon
and light-colored, has a pH of 3 to 5, and may be deficient consisting primarily of
in nutrients, organic matter, and clay. Gradually, about 30 sesquioxides or 1:1 clays.

cm below the surface, the B horizon begins. The B

horizon is often reddish-brown from the oxidized iron that Vertisol L. verto, to Soils that form large cracks on
turn drying; self-plowing.
has accumulated in it; it is relatively rich in mineral
nutrients and clay and usually has a more neutral pH than
Histosol Gr. histos, Organic soils.
the A horizon. The B horizon, then, is a layer of
tissue
accumulation. Finally, 100 to 130 cmbelow the surface,
the C horizon, made up of slightly weathered parent
material, appears. The relative thickness of the A and B
horizons depends on the climate and amount of plant
cover. Soils like these are called spodosols (old name = Mollisols (old name = chernozem) develop beneath
podzols). There are nine other soil orders (Table 9.3), four grasslands that experience moderate rainfall and high
of which are discussed below. summer temperatures. Moisture is no longer sufficient to

Alfisols and ultisols lie beneath the deciduous forests of leach the soil severely; the pH is 7 to 8. Fibrous root
the eastern United States. They are related to spodosols in systems of grasses thoroughly permeate this soil, and
that they are leached and acidic, but not as extremely as their decay evenly enriches it with organic matter. An A

spodosals. horizon, containing about 10% organic matter by weight,

Components of the Environment

179
occupies the top 60 cm. may be almost black in the
It

upper part, but becomes dark brown below. Just beneath

the A horizon is a calcium carbonate accumulation,
leached trom above. The layer is sometimes called the A c
horizon. Below it is the parent material; there may be no B
horizon.
Oxisols (old name = laterites) torm in regions of high
rainfall with continuously warm temperatures, such as
those that support tropical rain forests. Mineral cycling is

rapid because of fast plant growth and fast litter

decomposition. Soil is acidic in pH. Removal of the forest

canopy quickly leads to soil deterioration because litter

decomposition no longer compensates for leaching.
Excessive amounts of fertilizer must be added the land is if

farmed. Silicates (sand) are leached, but clay and iron are
not; the iron in clay is oxidized to Fe 2 3 ,
giving the topsoil
a brilliant reddish color.

Fire
Figure 9.10 The pine savannah of the southeastern coastal
In the past 40 years, has been rediscovered as a
fire plain The pine shown is longleaf (Pinus palustris); the area is in
North Carolina.
major ecological factor. We say "rediscovered" because
primitive human beings were well aware of its effect and
learned to use it for their own purposes. It may well be
Longleaf pine is tolerant to fire, and dependent on it.

that their most important food crops, their domestic The seeds germinate in the fall soon after they drop to the
animals, their routes of migration, and some of their ground from the cones. During their first year of growth,
cultural attributes have been molded by natural and man- the seedlings are very sensitive to even the slightest fire.
made fires. Certainly, many animals are today attracted to However, during the next 2 to 4 years, longleaf pine
fire and exhibit behavioral patterns in relation to fire that seedlings are in the "grass" stage (Fig. 9.11). Most of the
imply thousands of years of evolution in response to it. growing is done by the roots, and the stem apex remains
Most natural fires are started by lightning, and in North close to the ground. The terminal bud is covered with a
America many vegetation types, including grassland, dense mat of hairs that protects it from surface fires that
chaparral, and at least some conifer forests, owe their may sweep the area during this time. Fire is even
distribution and community structure in large measure to beneficial at this stage, because it kills a particular fungal
lightning fires. On U.S. National Forest land, during the 22- disease that parasitizes the long needles.
year period 1945 to 1966, lightning fires accounted for By the time the seedling is 3 to 5 years old, a surface
64% of an average of 5000 fires each year. In
all fires,
fire isdesirable to remove grasses that may have covered
addition, early explorers and settlers invariably commented
on the frequency of fires in forests and grasslands. In the
words of E. V. Komerek, a recent investigator: "Lightning
fires are an integral part of our environment and though

they may vary in both time and space, they are

rhythmically in tune with global weather patterns. Our
environment can be called a fire environment."*
One of the first vegetation types shown to be dependent
on frequent fire for its maintenance in this country is the
pine savannah along the southeastern coastal plain (Fig.
9.10). Tall, scattered loblolly, slash, shortleaf, and longleaf
pines dominate the area, and a thick growth of grasses
(mainly Andropogon and Aristida), with some herbs, cover
the ground beneath. During the nineteenth century when
this vegetation was protected from fire, was noticed that it

the pines gave way to oak; in time the result was a thick
oak forest with little grass. As the pines were valuable for
lumber and turpentine, and the grass for forage,
landowners were concerned about this trend. But the
importance of fire was not conclusively demonstrated until

1930, with longleaf pine (Pinus palustris).

* Komarek, Sr. 1968, The nature of lightning fires, pp. 5-41.

E. V.
In Proceedings of the 7th Tall Timbers Fire Ecology Conference, Figure 9.11 Close-up of longleaf pine seedling in the "grass'
Tall Timbers Research Station, Tallahassee, Florida. stage.

chapter 9 |
Plant Ecology

180
the seedlings and shaded it trom the sun. The seedling Sierra Nevada (about 1 500 m elevation) today is no longer
will then make a spurt of stem growth so rapid, that by the stately and open. We now have evidence that in the
age of 8 to 9 years the young tree's canopy will be high absence of fire, these forests are transformed into
enough above the ground to be out of reach of surface crowded fir (Abies) and incense-cedar (Calocedrus)
fires. In addition, its thick bark can endure the heat of a forests, and that the longer fire is excluded the more

fire without permitting damage cambium.

to the fire is If
catastrophic is the eventual, inevitable fire.

kept from such an area for 15 years, grasses and young California experiences a high frequency of lightning

oak and pine saplings of other species become so dense strikes during dry, late summer, and it is impossible to

that reproduction of longleaf pine is suppressed. If fire

prevent fires from starting. A considerable amount of

continues to be kept out beyond that point, oak trees

litter — needles, twigs, and bark —
is shed each year. So

gradually replace the pines and form a closed forest.

long as fires move through an area frequently, say every
8 to 25 years, the fire does not become hot enough to
The standard practice today is to burn the grass and
reach the upper canopy and significantly damage the
pine debris about every four years. The grass is quickly
trees. The native trees are adapted to such frequent, light
able to regenerate itself, and pine reproduction is favored,
burns. The seedlings of many of them require contact with
while oak seedlings are killed. If longer periods go by
mineral soil in order to become established because litter
without fire, too much dry litter collects on the ground,
does not retain moisture. Fire, of course, clears the
increasing the hazard of setting a fire so hot that mature ground of litter. In controlled experiments, seedlings of
trees aredamaged. Of course, no matter how "cool" the ponderosa pine, Jeffery pine, and big tree are much more
fire, great care
is taken to control The cost for control it.
abundant on burned plots than on unburned plots. If the
burning amounts to considerably less than $1 per acre. seedlings are protected from fire for 10 to 15 years, they
On the west coast recognition of the importance of become tall enough and develop a bark that is thick
controlled burning lagged behind its perception on the enough to withstand a light surface However, should
fire.

east coast. In California, the absence of fire in some areas fire be excluded for much longer periods, so much brush

has produced potentially catastrophic conditions, and litter collects that even mature trees cannot withstand
especially in the mixed-conifer forest of the Sierra Nevada the flames that eventually come. Many magnificent stands
mountains. Before settlement by Europeans, natural fires of big tree — the most massive tree
(Sequoiadendron) in

swept these forests about once every eight years (as the world, and found nowhere else the world — are now, in

revealed when fire scars in tree trunks are dated by ring for this reason, in great danger of being destroyed by fire

(Fig. 9.13).
count). However, those lightning fires did not create
raging, rampaging, extremely hot fires, because there was
only a relatively small amount of litter on the ground. The
Biological Factors
appearance of those fire-adapted forests was stately and
open, according to early reports by travelers. As shown in It is typical for several plant and animal species to coexist

Fig. 9.12, however, much of the mixed-conifer belt of the in a given habitat. Very rarely does a single species, to the

Figure 9.12 Thick understory of shade-tolerant trees that has

developed in a Sierran mixed-conifer forest as a result of fire
exclusion.

Components of the Environment

181
 Amensalism, another form of biological interaction, may
be defined as the inhibition of one species by another.
Whereas competition results from the removal of a
resource, amensalism results from the addition of
something to the environment In the coastal hills of
southern California, sage shrubs (Salvia) cover the slopes,
and grasses carpet the valleys. Occasional pockets of
shrubs occur in the grassland. Figure 9.14 is an aerial
view of these pockets of shrubs. The ground beneath and
about the shrubs is bare of grass, and the grass is stunted
as far as 9 m from the shrubs. Since the zone of stunting
is well beyond the limits of shrub root growth, competition

between shrubs and grass for soil moisture is not the

cause of stunting. If clumps of grass are transplanted to
the bare zone, their growth is severely retarded. Analysis
of the soil beneath the shrubs, the bare zone, or the
grasses, shows no major differences in physical or
chemical attributes. The reason for this distribution is that
many volatile oils are emitted from sage leaves, and two of
the oils in particular —
cineole and camphor are very —
Figure 9.13 A
grove of giant sequoia, or big trees toxic to grass seedlings. When both fresh sage leaves and
(Sequoiadendron gigantea) in Yosemite National Park, California.
grass seeds are placed in a closed chamber, germination
is often reduced to zero, and radicle growth is

considerably reduced. These same toxins can be isolated

exclusion of all others, control a habitat. If all the species from the air around sage shrubs, from the natural soil, and
in a group are individually examined, they will be seen to from seed coats of grasses in the soil. The conclusion of
utilize different parts of the environment or alternate with some ecologists is that the bare areas are caused by
each other in time. Some plants (green plants) are toxins emitted by Salvia that inhibit the germination and

producers of carbohydrates, but others (parasitic and growth of grasses. More recent studies indicate that a
saprophytic fungi) are consumers or decomposers of it;
high intensity of foraging by birds and small mammals,
some animals feed on plants, others feed on insects, and which nest in the shrubs, also contribute to maintaining
others feed on small mammals; some plants are trees that the bare zones.
utilize full sunlight, others like ferns utilize weak light; Amensalism by chemical means may be very common
some animals are nocturnal, others are diurnal; some in nature. This happens because plants are leaky systems,

plants are evergreen, others deciduous; and so on. The passively contributing all sorts of substances to their

portion of the environment utilized by each species is environment. One investigator grew seedlings of 150
called its niche. The niche has often been called the species in a nutrient culture that included several
"occupational address" of a species. In a particular area radioactive elements as markers. The plants took up the
at a given time it is thought that one and only one species isotopes through their roots and transported them to all

can occupy or each niche. Two species that occupy

fill
parts of the plant, including the leaves. The plants were
similar niches compete strongly with each other. then exposed to a mist, and the water that condensed and
Competition is one form of biological interaction. It may
be defined as the decreased growth of two species or
Table 9.4
plants because of an insufficient supply of some necessary
Some Terms of
categories of Biological Interaction in
factor(s) (Table 9.4). Sometimes only a single factor, such
the Effect the Relationship "On"
Has on Each Partner.
as a mineral element, is lacking, but usually two or more
Indicates that the Two Are Not in Contact. No Effect =
factors are limiting and it is difficult to separate them and -
0, Stimulation = + Depression =
,

determine which one creates the greater competition

stress. Competition is very important in determining plant
Effect
distribution. Many species restricted to saline, dry, or
nutritionally poor soils would actually grow better on Off
On
"normal" soil if other plants were removed. These species
are restricted to their niches because they are poor Name of Interaction #1 =2 #3 =4
competitors comparison with the numerous plants that
in

populate more moderate niches. They will die because the Neutralism
other plants have faster root and shoot growth rates, and Competition - -
remove water and sunlight. Sometimes, introduced plants Mutualism + + - -
become widespread pests and actually replace native Protocooperation + +
species because they are better competitors than the Commensalism +
native species. This seems tobe the reason for the Amensalism -
enormous increase of cheatgrass (Bromus tectorum) in
Parasitism / herbivory + —
the intermountain region of the United States.

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182
Figure 9.14 Aerial photo of Salvia shrubs adjoining grassland.
The light bands between and around the shrubs are bare of most
plant cover.

ran off the leaves was collected for analysis. He found that and lichens and mycorrhizae are mutualistic associations
14 elements, 7 sugars, 23 amino acids, and 15 organic of fungiand higher plants. Epiphytes, mentioned earlier,
acids — all radioactive — had been leached from the plants. form a commensal relationship with the host tree
In nature, these substances would have been leached by Protocooperation is exhibited by adjacent members of
rain from the leaves and would have accumulated in the some species that form natural root grafts, thus sharing
soil. Other studies have shown that roots are similarly soil resources.
leaky.
Herbivory, another form of interaction, plays an obvious
part in plant distribution. A glance along pasture fences Ecology and Plant Populations
reveals that normally abundant but palatable plants are
absent from the pasture, yet are common outside weedy
it;

species, which are not palatable, are common in the Ecologists would like to use species as indicators of

pasture, yet are rare outside it. Grazing animals are particular environments. They would like to know, for

responsible for this difference. Plant defenses against example, that a certain species only grows in a certain
herbivores include: (1) a spatial distribution that makes it range of temperature or moisture. Then the temperature
difficult for herbivores to locate and damage host plants: and moisture patterns of an area could be accurately
(2) a life cycle timing, such as seasonal leaf drop, that predicted by just noting whether that plant species grows

makes the host plant unavailable to herbivores; and (3)

there. Unfortunately, most species are tolerant of a wide
the manufacture of substances that are distasteful or range of environments and they are not genetically
otherwise inhibit herbivores from feeding on host plants. uniform. Individuals of species X that grow in one
Parasitism is herbivory by a plant rather than by an environment are slightly different from others of species X
animal, but the result and type of interaction are the same. thatgrow in another environment. Their genetic variation
Animal pollinators form a mutualistic relationship with makes possible for them to compete successfully in
it

plants. The bee, moth, and beetle, which cross-pollinate several different environments.

flowers while feeding on nectar or pollen, are familiar In the early twentieth century, the Swedish botanist
examples. Lichens are mutualistic associations of fungi Gdte Turesson collected seeds of species that had a wide

Ecology and Plant Populations

183
 much higher light intensities during the growing season
than do arctic areas. Physiological ecotypes have resulted.
4
^^* For example, the optimum light intensity for photo-
synthesis in alpine plants is much higher than that for
3
- arctic plants (Fig. 9.15). The two ecotypes also differ
Arctic
morphologically in leaf color, leaf shape, and the
&
o
XoQ.
2
/ s' frequency of rhizome production.
/ // Applications of the ecotype concept have been
o +1
•
// /
/ /
especially useful in forestry. The U.S. Forest Service
o
conducts tests to determine the best ecotype of a species
// compensation point
.1 ° // to be used for reforestation in a given area. Seed is
Jo
/
DC / collected throughout a species' range and is germinated in
-1
/ a uniform garden; seedling growth is carefully watched,
and the ecotype best suited for the garden climate is
2 1 1 1 1 1
1 1

500 1000 1500 2000 3000 4000 5000 selected for nearby reforestation programs.
Light intensity, foot candles

Figure 9.15 Photosynthetic response to different light intensities

for alpine and arctic ecotypes of alpine sorrel (Oxyria digyna).
Ecology and Plant
Communities
—
range of habitats from lowland, southern, and central
Europe to northeastern Russia in the Ural Mountains. Some environments support only one species. Dense
When he germinated the seed and grew the plants in a clusters of cattail (Typha) grow in water-filled ditches, and
uniform garden in Akarp, Sweden, he found that members all other vascular species are excluded; widely spaced
of widespread species were not uniform. Plants that grew spinifex (Triodia) covershundreds of square km of arid
from seeds collected in warm, lowland sites often were Australia, and few other perennial species are present
taller and flowered later in the year than those that grew (Fig. 9.16). Most environments, however, support groups
from seeds collected in cold, northern sites. Yet these of species associated together in communities.
variants had very similar flower morphology and were Provided that distances are not too great, a given
interfertile, traits indicating that the variants were all part of community repeats wherever the environment is
itself
one species. Turesson called these variants within species suitable. For in the Smoky Mountains
example, high peaks
ecotypes. He hypothesized that they were ecologically of North Carolina all exhibit red spruce (Picea rubens),
and genetically adapted to particular environments. mountain ash (Sorbus americana), Fraser fir (Abies fraseri)
Evidence that has since accumulated indicates that and a ground carpeted with several moss and fern species
most wide-ranging species are composed of a continuum (Fig. 9.17). The spruce, fir, ash, mosses, and ferns are
of ecotypes, each differing slightly in morphology and /or important members (but not the only ones) of this high-
physiology. For example, alpine sorrel (Oxyria digyna) is a elevation forest. High elevations at different latitudes or on
small perennial herb that grows in rocky places above different continents support other species, so there are
timberline in mountains and north of timberline in the other types of high-elevation forest communities. There
arctic. Both the alpine and arctic environments experience are hundreds of nonforest communities as well. Along
long periods of freezing weather, but alpine areas receive slopes of the California Coast Range at moderate
elevation, a shrubby community of scrub oak (Quercus
dumosa), manzanita (Arctostaphylos), and Ceanothus
repeats itself. Black spruce (Picea mariana), Sphagnum
moss, and the shrub Labrador tea (Ledum palustre) are
repeatedly associated together in virtually every bog
across Canada.
Communities are named after their most common large
species — the plants that receive the full force of the
macroenvironment. The spruce and fir of the high-
mountain community in the Smoky Mountains occur in
greater numbers than does the mountain ash, and their
leaf canopies influence the microenvironment in which the

carpet of ferns and mosses grow. The spruce and fir are
said to dominate the community, and the community
name is spruce-fir. For similar reasons, we have black
spruce communities and scrub oak communities.
Communities exhibit a unique architecture or structure,
produced by the leaf canopies of their dominant and
subordinate species. In the tropical rain forest of British
Figure 9.16 A community composed of essentially a single
Guiana, one group of species forms a canopy at 30 m, a
species: spinifex, or porcupine grass (Triodia basedowii) in
central Australia. second group at 18 m, and a third at 9 m (Fig. 9.18). The

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184
 .

These fossil communities are so similar to modern

communities that we are able to reconstruct past climates
from them. A fossil community near the town of Goshen in
central Oregon, dated to the Eocene epoch (50 million
years ago), included the following genera: Drimys,
Magnolia, holly (Ilex), oak (Quercus), Ocotea, and fig

The closest living approximation

(Ficus). of this fossil
community today occurs 2400 km south of it in the
temperate uplands of Central America. Apparently, the
climate in North America is now much colder than it was
in the Eocene. The reconstruction of past vegetation types
and climates by the use of fossil evidence is called
paleoecology.
Another form of fossil evidence is pollen that has been
trapped in lake sediments. As pollen is shed near a lake,
some sinks to the bottom and is incorporated with silt and
organic matter that forms sediment. A chronological
sequence of pollen is thus preserved; the deeper the
pollen occurs in the sediment, the older the pollen. Pollen
of many species decay under such
is resistant to
anaerobic conditions, and may remain
intact for thousands
of years. Cores of sediments can be examined under the
microscope and the pollen identified as to genus, and
sometimes even to species. Within limits, paleoecologists
assume that the abundance of pollen in the core is
proportional to the abundance of that species in the past.

How May Communities Be Measured?

Figure 9.17 Subalpine forest in the mountains of North Carolina.
The trees are red spruce (Picea rubens) and fraser fir (Abies If communities are to be useful as tools with which to
fraseri). estimate the nature of the environment, they must be
described more completely than just by their name,
because minor variations in species composition indicate
ground floor is almost bare of shrubs or herbs. In
different environments. For example, in areas where
deciduous forest communities of the United States, there is
spruce and fir are both dominant and very abundant, the
one tree canopy layer at about 18 m, a layer of short trees
environment is probably different from where they are still
(like dogwood) at 3 m, a rich shrub layer at 1 m, and a
dominant but less abundant. How can the abundance of a
seasonally present layer of herbs on the ground. Some
species or the composition of a community be measured?
desert communities exhibit only a single (shrub) layer.
Fossil impressions millions of years old sometimes 1 One way community is to measure the
to characterize a
reveal communities very similar to those that exist today. amount ground covered by each species. The
of
amount of ground cover is important because it affects
the microenvironment of the community. To sample the
ground cover, one could stretch a tape measure across
the ground; the length of tape covered by each plant
canopy is proportional to the total ground cover of each
species. This method of determining ground cover is
called a line transect. Numerical information from line
transects often contradicts first impressions. In desert
shrub communities, for example, a person looking
across the landscape would think that the shrubs
covered much ground (Fig. 9.19), but such a
of the
estimate is inbecause of the low perspective. If
error
seen from above, when the canopy edges can be
projected perpendicularly to the ground, it is apparent
that actually only 10 to 20% of the ground is covered.
Forest communities, on the other hand, exhibit more
50 40 30 20 10
than 100% cover because of overlap in canopy layers
feet
at different levels.
Figure 9.18 Profile diagram of the architecture of a tropical rain 2. Another method of estimating cover, which is especially
forest in British Guiana. There are three tree strata, at
approximately 30, 18, and 9 m. Shrubs and ground herbs are
useful when by the use
plants are low to the ground, is

uncommon, but tree saplings are included in the 9 m stratum. of quadrats. Quadrats are variously shaped frames that

Ecology and Plant Communities

185
Figure 9.19 Desert scrub dominated by creosote bush (Larrea
tridentata), an abundant warm desert plant of North America.

can be placed on the ground directly over the plants.

The amount of ground covered by each species can be
estimated as a percentage of the area enclosed by the
quadrat frame.A community can be sampled by placing
quadrats random or at regularly spaced locations
at
throughout Usually, not more than 10% of the total
it.

community area is included in the quadrat samples. Figure 9.21 A stand of longleaf pine (Pinus palustris), about 50
3. Other methods of sampling, based on mathematical years old, showing a well-developed understory of broad-leaved
assumptions, can be used to determine whether species of oak and hickory. In time, the pines will die and be
replaced by maturing members of this understory
random, regularly (like
individuals are distributed at
trees in an orchard), or
clumps (Fig. 9.20).
in

Nonrandom distribution (regular or clumped) may result

invade following disturbance are annual and perennial
from propagation by asexual means, the method of
seed disperal, competition, or some pattern (like
herbs, which grow best in high light intensity, such as
terracing) in the microenvironment. Most temperate
horseweed (Conyza canadensis), Aster, and broomsedge
zone species are distributed in clumps. (Andropogon virginicus). These make up the pioneer
community. Pine seeds blow in from neighboring areas
during the first several years after the disturbance, and

Succession within five years there are many pine seedlings. As the
seedlings grow, they compete with the herbs for moisture,
As we have already mentioned, the microenvironment and their expanding canopies begin to change the
beneath a plant community is very different from that in microenvironment. Within 30 years of the disturbance, a
the open. Temperature, humidity, soil moisture, and light stand of pine results. Examination of the forest floor,
tall

are all affected by the canopy. Acommunity

stable however, shows many oak and hickory seedlings and very
consists of species whose seedlings can survive in its few pine seedlings. Pine seedlings grow poorly in shade
unique microenvironment; seedlings of other species do and under conditions of root competition, but those of oak
not survive. If the stable community is removed by some and hickory do well. Within 50 years of the disturbance,
catastrophe, leaving the soil exposed to full sunlight, the the hardwood seedlings form a well-defined understory
first species that colonize the site are not those of the old beneath the pines (Fig. 9.21). Whenever a pine tree dies,

community. They are seedlings of species adapted to it is replaced in the upper canopy by an oak or a hickory.

growth in full light intensity. Within 200 years of the disturbance, the forest again
In parts of the southeastern United States where consists only of mature oak and hickory trees, and the
oak-hickory communities are stable, the first species to forest floor is covered with many seedlings of the same

Figure 9.20 Diagrammatic representation of three types of plant

distribution: A, clumped; B, regular; C random.

chapt er 9 |
Pl ant Eco logy

186
trees.The shrubs and herbs associated with an oak-
hickory community are also present. Such a sequence of
change in plant communities is called a succession or
sere, and the stable community that is capable of
indefinitely maintaining itself is called a climax
community. In this case, oak-hickory is the climax
community. All the intermediate, temporary communities
are called serai stages.
Sometimes, climax communities are prevented from
becoming established because of a recurring disturbance.
There is some evidence that part of the prairie of the
central United States that greeted pioneers could have
been a forest were it not for recurring prairie fires. The
fires destroyed slow growing tree seedlings, but grasses
quickly regenerated or germinated. When fire is controlled,
prairie near the forest edge soon supports many tree
seedlings.
The first ecologists to elaborate successional
pathways — men like Europe and
Josef Paczoski in

Frederick Clements in had no

the United States —
experimental evidence for documentation. They could not
record changes as time passed, for most successions take
hundreds of years to complete. They determined the
existence of successional pathways by comparing a
climax community with adjacent small areas that had been
disturbed at known times in the past. They examined
freshly disturbed or denuded areas (as a result of, for
example, ice, landslides, fire, logging, and wind-throw),
which revealed the likely pioneer community, or the first

stage of succession. Sites that had been disturbed at

some time earlier (verified from records of fires, grazing,
Figure 9.22 Succession of communities about a pond near
or logging) exhibited later successional communities
Juneau, Alaska. Standing water in the pond supports leaves of
because the passage of time was greater. Complete cow lily (Nuphar); mats of Sphagnum moss and sedge (Carex)
sequences of succession were sometimes represented as encroach on the pond along its edge; beyond is a shrub-
dominated community of Empetrum, Cassiope, and Kalmia, with
bands of different communities about a gradually filling some mountain hemlock trees (Tsuga mertensiana); finally, sitka
bog or behind a retreating glacier (Fig. 9.22). The spruce (Picea sitchensis), a tall, dark-green tree, dominates the
rest of the area.
community closest to the lake or glacier was the most
recently established and represented the pioneer
community. The next closest community occupied soil that
toa different vegetation type than one dominated by
had been exposed for a longer time; at some earlier time it
shrubs or one dominated by conifers. On the other hand,
had supported the pioneer community, but enough time
many different communities dominated by different
had passed for the next successional stage to appear.
conifers belong to the same vegetation type. We will briefly
Beyond the second community was a third, representing
examine six major vegetation types: tundra, taiga,
the third successional stage, and so on until the climax
deciduous forest, tropical rain forest, savannah-prairie,
community was reached. Ring counts of trees closest to
and scrub.
the pond or glacier, coupled with estimates of the rate of
silting or of glacial movement, gave clues as to the time

necessary for each successional stage to be reached.

Tundra
The conclusions of these early workers have yet to be
experimentally or empirically verified, and only recently If one were to travel north or south toward the poles, it

have some of the causes of succession (such as would be evident that the trees gradually become stunted
competition, amensalism, and growth requirements of and less common, and finally disappear completely. At the
seedlings) been experimentally dealt with. same time, low shrubs, perennial herbs, and grasses
become dominant. The resulting meadowlike vegetation is
called tundra (Fig. 9.23). Annual plants are very rare, and
Vegetation Types of the World the reproduction of perennials is chiefly by vegetative
means such as rhizomes. Shrubs are dwarfed, gaining
normal height only in the protection of the lee of boulders
The term vegetation refers to the life form of plants or small hills. The winter wind, carrying ice, acts like a
dominating a community: trees, shrubs, or herbs; sand blast and prunes back stems wherever they project
deciduous or evergreen; coniferous or broadleaved. A beyond the boulder or hill.
community dominated by evergreen angiosperms belongs The warmest month has an average daily mean

Vegetation Types of the World

187
Figure 9.23 Alpine tundra, Beartooth Plateau, Wyoming, 3300 m
elevation.

temperature of 10°C or less and the growing season is The lower limit of these alpine areas depends on
short. Only 2 to 4 months of the year have average daily latitude. At 60° N in is at 900 m; at 45 °N in
Alaska, treeline
temperatures above freezing. Approximately the top 30 cm the Rocky Mountains, 3000 m; at 20°N in central
it is at
of soil thaws during the growing season and roots may Mexico, is at 4000 m; at 5°S in the Andes of South
it

freely penetrate it, but below this level soil water may be America, is back to 3600 m; and at 50°S in Chile,
it is it

permanently frozen; this soil is called permafrost. Annual back to 900 m. Because summer temperatures are cooler
precipitation is about 25 cm, very little falling as snow. The near the coast, timberline also depends on distance from
terrain is generally flat, but small depressions are common the ocean. Thus, timberline is lower in the Cascades than
and water tends them during the growing
to stand in in the Rockies.
season. Day length fluctuates from 24 hr in June (northern
Taiga
hemisphere) to none at all in December.
A similar tundra vegetation occurs in mountains at Just south of the tundra is the taiga, a broad belt of

elevations above The climate, however, is slightly

treeline. communities dominated by conifers. A similar belt occurs
different. Solar radiation is more intense, day length does in mountains, just below the alpine zone (Fig. 9.24). Trees

not fluctuate so extremely, and frosts are more common are slender, short (15-20 m), and relatively short-lived
during the growing season. Many plants are tinged with (less than 300 years), but the forests are dense. The taiga
red from abundant anthocyanin. This substance absorbs is made up of a patchwork of forests on upland sites that
some of the high light intensities that may damage the alternate with bogs formed in depressions. Soils are acidic
plant's photosynthetic apparatus. and relatively infertile spodosols; in northernmost or

Figure 9.24 Taiga, northern Canada.

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188
Figure 9.25 Coniferous forest, Olympic Peninsula, Washington.
Note the moss and lichen-covered trunks and branches and the
dense carpet of ground vegetation.

uppermost parts of the taiga the soil has permafrost below

a surface layer.
The growing season is 3 to 5 months long, and
temperatures above 30°C are occasionally reached.
Winter temperatures are very severe; the mean daily
temperature is below freezing for six months of the year,
and at Yakufsk, Siberia, the average January daily
temperature is -43°C. Annual precipitation is less than
Figure 9.26 Tropical rain forest, New Guinea. The dense, lower
50 cm, and the air is exceptionally dry. The taiga is vegetation is absent farther into the forest. Note the hanging
lianas.
monotonous and silent; animal life is not abundant.
At lower elevations in some mountains, and along the
coast (where temperature extremes are not so severe), summer rain. Annual precipitation ranges from 75 to 125
coniferous forests are much more luxurious. Trees are cm. Since the growing season may be as long as 200
taller and the undergrowth denser. Many favorite days, it is not surprising that much of the deciduous forest
recreation areas of the western United States are situated in Europe, Asia, and North America has been cut and
in these richer forests. The Olympic Peninsula of replaced with cropland.
Washington receives over 200 cm of annual precipitation Soils are alfisols in the north and west, ultisols in the
and supports the most spectacular coniferous forest in the south. (The latter have striking yellow or red colors in the
world (Fig. 9.25). Douglas fir, the most heavily cut lumber B horizon, which is often exposed if plowed.)
species of the United States, reaches its best growth in
this area.

Tropical Rain Forest

Deciduous Forest No other vegetation can quite equal the rain forest in

sheer diversity of species and complexity of community

In contrast to the taiga, the deciduous forest supports a structure (Fig. 9.26). The deciduous forest may exhibit 5
diversity of plant and is vibrant with animal activity.
life to 25 different tree species per acre, but the tropical rain
Deciduous angiosperm trees dominate communities of this forest supports 20 to 50. The tree canopies typically form
forest. When day length becomes shorter and nights three layers, and beneath this dense overstory there are
cooler in late summer, hormone concentrations in such very few shrubs and herbs. Light intensity on the ground
trees change, and the destruction of chlorophyll follows, is less than 1% of full sun. Most of the herbaceous plants

allowing other pigments to show through in brilliant fall grow as epiphytes on trunks and branches of trees. Vines
colors of red, yellow, and orange. In spring, before new cling to tree trunks and wind their way up to the canopies.
leaves have fullyexpanded from buds, beautiful and Seeds of a few species, such as the well-named strangler
distinctive herbaceous perennials come into flower. Shrubs fig, germinate in the canopy and grow down to the soil.

are common, but not dense. Many of the tree trunks flare out at the base in peculiar
These forests occur in areas with a cold (though not buttresses.
severe) winter, and a warm, humid summer. Snowfall may The species are evergreen. A given leaf may be shed or
be heavy, but most of the annual precipitation falls as a given bud may break at any time of the year. The leaves

Vegetation Types of the World

189
tend to be very large, with long, tapering tips that may
serve to drain the surface of moisture Rainfall is high,
often over 250 cm a year, but it is evenly distributed
throughout the year Temperature is also uniform
throughout the year, averaging 27°C. Warm temperatures
and high humidity couple to produce a climate oppressive
to humans. Afternoon showers, which reduce the
temperature, offer only short
f- arming peoples
relief.

of the rain forest

practiced agriculture on a cut-burn-cultivate-abandon plan.
have for centuries i
The vegetation is removed in a small area by cutting and
burning, cultivation follows for a few years, and the plot is

abandoned. Abandonment follows because crop growth

declines each year. The rain leaches nutrients from the
exposed topsoil and the nutrients are not replaced by litter Figure 9.28 The original steppe, or prairie vegetation of central

as they are beneath the normal forest canopy, as California, dominated by the perennial bunch grass Stipa pulchra

mentioned earlier in this chapter. Much of the rain forest

ofAsia and Africa has been disturbed in this way. More
vegetation or animal life remains today. Only from the
recent disturbance involves bulldozing and clear-cutting,
records of explorers and early settlers are ecologists able
especially in South America; the rain forest may be to reconstruct in their imagination what these thousands of
incapable of reinvading these areas. The vegetation that
square miles once looked like.
invades these abandoned plots is shrubby, dense, and
The eastern part, with relatively high rainfall, supported
often includes bamboo. It is this successional stage, which
some m
a dense "sea" of grass 1 to 2 tall. Here is a
eventually will give way to rain forest, that serves as the
description from a journal dated 1837:
"jungle" of television and films. Soils are typically in the
oxisol order.
The view from this mound beggars all description. An
. . .

ocean of prairie surrounds the spectator whose vision is

not limited to less than 30 or 40 miles. This great sea of
verdure is interspersed with delightfully varying

Savannah and Prairie undulations, like the vast swells of ocean, and every here
and there, sinking in the hollows or cresting the swells,
Forests giveway to grassland as rainfall decreases and appears spots of trees, as if planted by the hand of Art for
the purpose of ornamenting this naturally splendid scene.
becomes more seasonally distributed. At first, trees remain
close enough to retain a closed canopy, but gradually they This is now the corn belt and is probably one of the
become more widely spaced, and eventually they drop out most agriculturally fertile areas of the world. Productivity is

altogether except along river banks and canyons. so high that it permits us to export grain to many countries
Savannah is the term applied to grassland with widely around the world. Its mollisol soils are dark brown in color
spaced trees whose canopies cover less than 30% of the from the rich amount of organic matter left by decaying
ground (Fig. 9.27); steppe or prairie refers to grassland fibrous grass roots.
without trees (Fig. 9.28). The drier, western part supported shorter and sparser
Grasslands of the United States are heavily used for stands of grass, but there was still enough growth to allow
agriculture and grazing purposes, and little of the original graziers to cut and bale it as hay. Today, the western

Figure 9.27 African savannah in Kenya. The flat-topped trees

are Commiphora, which is the source of the biblical myrrh.

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190
prairiesupports a shorter, sparser cover ot grass. Reasons
for such a decline include too much grazing pressure by
herds of cattle and a slight drying trend in the climate of
the area over the past 75 years. Shrubs from the desert
and junipers from the foothills have invaded (possibly
because of fire suppression), and they have further
reduced grass cover.
Grasslands are the home of the largest terrestrial
herbivores and carnivores. Vast herds of bison that
roamed the North American prairie are gone, but animal
preserves in African savannahs still excite visitors by a
spectacular animal cast that includes eland, wildebeest,
giraffe, elephant, and lion.

Scrub
Scrub is vegetation dominated by shrubs, and it occurs in

areas with hot, dry summers and moderate winters. In

areas with high, but seasonal, precipitation, the shrubs are
tall and thorny, but if rainfall is only 40 to 50 cm a year,
scrub can be a nearly impenetrable thicket of shrubs with
evergreen, hard, spiny, small leaves (Fig. 9.29). Scrub of
this sort is found around the Mediterranean Sea, on
coastal hills of California and Chile, in southwestern
Australia, and at the tip of South Africa. Even though the
species of plants in these widely separate areas are Figure 9.30 Dese in southern Arizona with succulents
(cacti).
entirely different, the vegetation has come to look strikingly
similar. In the western U.S., this type of scrub is called
chaparral. germination and a colorful show of flowers a few weeks or
Desert scrub (Figs. 9.19 and 9.30) grows in areas of months later. Although desert scrub seems to be made up
rainfall below 25 cm a year. Shrubs are widely spaced and
mainly of perennial species, an actual count of all species
have far-reaching root systems (much of which runs just appearing throughout the year reveals nearly as many
beneath the soil surface). The shrubs may be associated annuals as perennials.
with succulents, plants that store water stems or
in

leaves, like the common cactus of the southwestern

United States. Some shrubs have deep roots that tap A Final Note
permanent supplies ground water, while others reduce
of
transpiration by dropping their leaves every dry period and We have described major vegetation types as if they have
developing new ones every rainy period. Seeds of desert sharp boundaries with, and differ greatly from, all other
annuals may remain viable for long periods in the soil until types. Actually, vegetation types typically grade into one
a favorable year, when rainfall is sufficient to trigger another or form complex mosaics that may extend for

Figure 9.29 Chaparral in the foothills of California

Vegetation Types of the World

191
thousands of hectares. Meeting zones of two or more
vegetation types are called ecotones.

Ecology and Ecosystem

Energy Flow
reflection heat
We have examined plant ecology at the population,
community, and vegetation levels. A final level of
organization is the ecosystem. The ecosystem of a
700,000 Kcal/m J
particular habitat includes the plant and animal
communities, the physical (nonliving) environment, and all

the interactions between them.

The organisms of an ecosystem can be divided into

three categories: producers, consumers, and

decomposers. Producers are green plants and certain
bacteria that perform photosynthesis. Consumers are
parasitic, herbivorous, or carnivorous plants and animals
transpiration jyj respiration
that feed on other plants or animals. Decomposers are net plont productivity

saprophytic bacteria, fungi, and certain animals that obtain

nutrition by breaking down dead organic matter. The three
groups form a complex web of interdependence:
consumers feed on producers, and decomposers feed on
the organic remains of both and convert them to lower-
energy, smaller molecules that are taken up again by
producers.
Caloric energy is also passed through the ecosystem,
and in this transfer organisms are linked to each other by
net steer growth
a food chain. Food chains are short in the arctic tundra:
meadow plants trap light energy and convert it to chemical
69Kcal/m !

energy (e.g., proteins or carbohydrates); caribou transfer

Figure 9.31 Loss of energy in western rangeland. Of 700,000
some of this energy to themselves, retaining some of it, kcal of radiant energy that reach each square meter of surface,
excreting some, and respiring some; and human beings only 0.1 % becomes available as plant food for the cattle grazing
transfer some of the caribou caloric energy by eating a the rangeland. Only 0.01% of the plant food consumed during an
eight month period is converted to meat available for human
fraction of the total caribou. Aquatic ecosystems exhibit consumption, that is, only 0.004% of the original incident solar
longer food chains: algae (plankton especially) produce energy.
the caloric energy; some is passed to the small animals
that feed on plankton, then to the small fish that eat the
respiration, waste matter, and inedible bones and hide,
plankton feeders, then to larger fish, and then to shore
leaving a net of only 69 kilocalories per square meter, of
birds or land animals like bear or man.
the original 700,000, that goes into meat. This represents
There is tremendous variation in net productivity from
about 139 kg of edible beef on the hoof per ha per year.
one ecosystem to another. Net productivity equals
calories produced in photosynthesis minus calories lost in
respiration. Desert ecosystems, for example, exhibit less
Pollution
than 0.5 gram of dry matter produced per square meter of
land per day; grasslands, 0.5 to 3.0; deciduous and Human beings add to the environment many substances
coniferous forest, 3 to 10; agricultural areas utilized all whose on plants and animals is poorly understood
effect
year (asin sugar cane production), 10 to 25. (Fig. 9.41). If organisms in food chains or ecosystems
Energy transfer through the ecosystem involves losses shared by humans are damaged by these substances,
each time energy passes from one type of organism to then humans may be indirectly, but nevertheless severely,
another. Figure 9.31 summarizes a very simple food chain damaged.
consisting of California rangeland on which steers are These substances are called pollutants. Some are
grazed. The range receives 700,000 kilocalories (kcal) of liquids or solids that have been introduced to waterways
sunlight energy per square meter per year. After deducting and may affect the metabolism of aquatic plants, such as
the energy lost by reflection and heating, the fraction algae (see Chapter 1 1). Others are gases released to the
spent by plants on their own respiration, and other energy atmosphere; they may affect the growth of wild or
channeled into roots and other unconsumable parts of the cultivated terrestrial plants. Among the most important of
plants, only 800 kilocalories are taken in by the grazing these air-borne pollutants are sulfur dioxide (S0 2 ), ozone
animals. The animals in turn spend nearly all their energy (0 3 ), and peroxyacetyl nitrate (C 2 H 3 5 N, also called PAN).
unproductively (from the point of view of human beings) in Sulfur is present in iron, copper, lead, and zinc ores;

chapter 9 I
Plant Ecology

192
during refining it is released as S0 2 . It is also released
trom the burning of coal. The S0 2 enters leaves through -v:o
:
---
, ,,c*::""o©"-.., ,

stomata and goes into solution on the wet mesophyll cell

CH,— C— O— O— N*
walls as sulfurous acid. Inside the cells, such vital
metabolic processes as protein synthesis are disrupted
because of the reducing nature of this acid. Ultimately,
-=> ' V
cells shrink and collapse, producing visible chlorotic •
*.w ....

lesions on the leaves. Chronic exposure of such sensitive

crops as alfalfa, barley, and lettuce to only 0.1 to 0.5 ppm
(parts per million) of S0 2 causes visible damage and
hydrocarbons + NO2+O2 + NO- —( PAN A

reduced yield. The effect of S0 2 on natural vegetation is

dramatically revealed in Copper Basin, Tennessee, where
early in this century a refinery freely released S0 2 before
its was understood. Over an area of perhaps
toxic nature
40 km 2 the original forest was destroyed (Fig. 9.32). The
barren land was so severely eroded that subsequent
+ O3
attempts to revegetate the area have been unsuccessful.
Ozone and PAN are important components of urban
smog. In large part they are a result of the incomplete
combustion of automobile fuel (Fig. 9.33). Our chemical
understanding of smog and its affect on plants started less
incomplete combustion
than 30 years ago. Ozone is naturally present in air, but gasoline >- hydrocarbons +NOj+H 0+02 2

smog contains 10 to 100 times the natural concentration,

and this concentration is sufficient to cause millions of
dollars of crop damage a year. Ozone enters through the
stomata, and its oxidizing nature causes many metabolic
malfunctions, including chloroplast breakdown. In the
Figure 9.33 The formation of ozone (CL) and peroxyacetal
nitrate (PAN) in smog. The formula for PAN is given at the top of
the diagram.

mountains near Los Angeles, air circulation takes sufficient

quantities ofozone to a height of 1700 m; this causes
needle drop, reduced growth, and the ultimate death of
trees in ponderosa pine forests (Fig. 9.34).
Other experiments in the Los Angeles Basin have
shown the effect of PAN on citrus. Established lemon and
orange trees were encased in miniature greenhouses.
Some greenhouses circulated normal, smoggy, outside air;
others circulated air that had first been filtered of PAN.
Over the period of six years, the average yearly fruit yield
of lemon trees was reduced 39% by PAN, and the orange
tree yield was reduced 64%. Evidently, PAN interferes with
the light reactions of photosynthesis and also with
respiration.
It is hoped that ecologists will eventually show us how

ecosystems can be rationally manipulated to

simultaneously satisfy all our needs: industry, urbanization,
and recreation. Knowledge must
agriculture, conservation,
be gained and compromises made between our needs and
what the environment can support.

Summary
1. Plant ecology is the study of how the environment
influences plant distribution and behavior. It is also the
study of the structure and function of nature. Ecology
deals with the population, community, and ecosystem
levels of organization.
2. The components of the environment include moisture,
Figure 9.32 Copper Basin, Tennessee. The normal eastern temperature, light, soil, fire, and living organisms. Each
deciduous forest was destroyed by sulfur fumes from a smelter in
the early twentieth century and revegetation attempts have been species utilizes a different part of the environment: its

unsuccessful. niche.

Summary

193
Figure 9.34 The effect of smog on ponderosa pine growth in a
mixed-conifer forest near the Los Angeles Basin. A, cross section
of a normal 30-year-old pine; B, the same age tree further
downslope, exposed to smog-polluted air.

3. The seasonal timing of rainfall may be as important as A plant community is composed of a group of
the annual total in determining the kind of vegetation associated species or populations that occupy a
present. particular type of environment. One or more of the
Slope aspect affects temperature in the species dominate the community,' but all of the species
microenvironment. It also affects the amount of net together combine to form a unique architecture of
radiation that reaches the plants. canopy layers. The structure and composition of
5. Light quantity and quality is changed as light passes communities can be objectively determined by the use
through a leaf canopy or through water. of sampling methods such as line transects or
6 Soil formation proceeds through mechanical and quadrats.
chemical weathering and through plant activity. 11. Climax communities perpetuate themselves, but
Colloidal soil particles are important as nutrient successional communities are transitory.
reservoirs, and soil pH affects their ability to retain Vegetation types of the world include tundra, taiga,
nutrients. Through time, soils in different regions deciduous forest, tropical rain forest, grassland, and
develop their own characteristic profiles. Soils can be scrub.
classified by profile characteristics into 10 orders and 13. An ecosystem consists of the plant and animal
thousands of soil types. communities of a habitat, their abiotic environment,
7. Fire is an important, natural environmental factor that and all the interactions between these components.
affects the distribution and architecture of many Organisms an ecosystem are tied together by food
of
vegetation types. chains. Damage done
to one component of a food
Forms of biological interaction include competition, chain (as through the action of pollution) may cause
amensalism, predation, and mutualism. indirect but serious damage to the entire ecosystem.
9. A taxonomic species includes genetically similar plants 14. Important atmospheric pollutants include sulfur
any wide-ranging taxonomic
that are interfertile. Within dioxide, ozone, and peroxyacetyl nitrate. The latter two
species, however, are many ecotypes that have are components of smog and cause millions of dollars
become adapted to different environments. of crop damage a year as well as ecological damage
to natural vegetation.

chapter 9 |
Plant Ecology

194
 10 CHAPTER

10 plant taxonomy
and evolution

taxonomy deals with the identification, naming, Linnaean (1753-present) period.

Plant
and classification of plants. The term Before the time of written history, human beings had
"identification" implies that many plant groups learned to identify many plants. These plants were
exist, and each group possesses unique characteristics so particularly useful as spices, medicines, foods, drugs, or
that it can be distinct from all other groups. These religious symbols. The written history of botany begins
characteristics, as we shall see, involve morphology, with the Greek Theophrastus (ca. 300 B.C.), a student of
anatomy, physiology, cytology, biochemistry, and Aristotle. He described and classified about 500 kinds of

geographic distribution. The term "classification" implies plants in the book De Historia Plantarum. He utilized
that plant groups can be ranked in some hierarchical morphological characters like habit (tree, shrub, herb),
relationship, based on similarities in their characteristics length of life (annual, biennial, perennial), corolla form
that reflect genetic relationships. Taxonomists are building (petals fused together or free), and ovary position
a library of the world's plant resources; they are making a (superior, inferior) to distinguish among his plants.
census of the unique characteristics of each of the half- Many of the taxonomists who followed the lead of
million species that make up the world's flora.* This Theophrastus through the dark ages and Renaissance are
census is far from complete. Many areas of the world have typified by Otto Brunfels (ca. 1500 A.D.) of Germany. He
not been botanically explored and their floras can only be too classified plants on the basis of gross morphology, but
estimated. he contributed to the record by expanding the list of
Taxonomists hope that, as the census becomes known and using drawings to accompany many of
plants
complete, patterns of similarity will become apparent the descriptions. Hans Weiditz was Brunfels' illustrator,
within this maze of plant diversity, and that these patterns and he did a magnificent job of carving accurate and
will enable them to reconstruct the evolutionary history of intricate woodcuts from which prints were made (Fig.

the present flora. They hope it be possible to

will 10.1). The descriptions and drawings of many species
determine which characteristics — outof the hundreds were compiled in books called herbals. Herbaceous plants
available — are most reliable in showing pathways of used for medicinal purposes were often emphasized in the
Are they characteristics of flower morphology,
evolution. herbals. It was common to assign medicinal properties to
wood anatomy, chemical composition, geographic a plant on the basis of its form. If the plant or parts of it

distribution, breeding behavior, or chromosome number? imitated the shape of an organ, then it was used for
Which traits in each category are primitive (imitating correcting ailments of that organ. For example, the simple
those of early, now-extinct plants)? Which traits are plant body of liverworts, which resembles the shape of a
advanced (derived in time from primitive traits)? When the liver, was used to treat liver ailments.
answers to such questions are known, it will be possible to Carolus Linnaeus (ca. 1750, Fig. 10.2) represents the
construct a phylogenetic or natural classification based culmination of pre-Darwinian thought. Born Carl Linne in
on genetic, evolutionary relationships. Such a southern Sweden, he attended the University of Uppsala
classification, in addition to summarizing the past, might and received an M.D. degree from the University of
prove useful in predicting future pathways of evolution. Harderwijk in the Netherlands. During his early days at
Uppsala, however, he had become interested in the
classification of plants and had developed his own system,
called the sexual system. was based almost entirely on
It

Historical Aspects the morphology and number of stamens and pistils, and
proved very efficient in classifying the many new plants
then being brought to Europe from Africa, America, and
Pre-Linnaean Period Asia and propagated in botanic gardens. After working on
the collections of many large botanic gardens, Linnaeus
The taxonomy can be very roughly divided into
history of published the book Species Plantarum in 1753. The work
a pre-Linnaean (300 B.C.-1753 A.D.) period and a post- included about 6000 species in 1000 genera.
Each species was described in Latin by a sentence
* Another meaning of "flora" is a book that lists all the plant
species found in a given region. Thus, floras have been published words that began with the genus name.
limited to twelve
for countries, states, counties, or even more local areas. Linnaeus considered this sentence (a polynomial) to be

195
 CXCVI Cowrafapc

Figure 10.3 Tradescantia virginiana (spider wort), x 2. Note the

jointed staminal hairs, from which the common name derives.

virginianum gramineum, common name Tradescantia

virginiana. Loosely translated, the polynomial means: The
fj 2>on t>em 2?ammen.
annual, herbaceous, upright Tradescantia from Virginia
Cojcfcnrc6na6c(/<Bo(c«cf.nat)/£ranc^ that has a grasslike habit, three petals, and [stamens with
Figure 10.1 Part of one page from Brunf el's sixteenth century hairs that are] spider-like, common name Tradescantia of
herbal, The plant illustrated is called stork's bill (probably Erodium Virginia. The genus name was in honor of John
cicutahum).
Tradescant, a gardener to King Charles I.

Nof surprisingly, the binomial became favored over the

polynomial by latertaxonomists, and was accepted as the
the official, scientific species name, but he also coined a
shortened form consisting of the genus name and one official, scientific name. Common names were left to
individual choice and not restricted to Latin. The usual
additional word from the polynomial. This shortened form
(a binomial) served as his common name for the plant. He
English common name for Tradescantia virginiana is

described spiderwort by this polynomial: Tradescantia

spiderwort (spider plant), which comes from the jointed

ephemerum phalangoides tripetalum non repens hairs on stamen filaments that resemble the jointed legs of
a spider (Fig. 10.3).

Post-Linnaean Period
After Darwin published Cn the Origin of Species in 1859,
emphasis in taxonomy shifted from a description of
species per se to: (a) a very selective description, based
on characters that reflected genetic (evolutionary)
relationships, and (b) the construction of a phylogenetic
classification scheme. Construction of phylogenetic
schemes demanded that decisions be made on which
traits are primitive and which are advanced. As there was

little or no fossil evidence to support any of several

hypotheses, early schemes were both numerous and often

in conflict. For example, the American C. E. Bessey

Table 10.1
Assumptions for the Besseyan Phylogenetic Scheme

Primitive Characters Advanced Characters

Plants woody Plants herbaceous

Flowers bisexual Flowers unisexual
Floral axis elongated Floral axis short
Floral parts spirally arranged Floral parts whorled
Floral parts numerous Floral parts few
Sepals or petals free Sepals or petals fused
Floral symmetry radial Floral symmetry bilateral
Figure 10.2 Linnaeus (1707-1778), the Swedish botanist who Fruit single Fruit aggregated
initiated the binomial system of nomenclature.

chapter 1Q l
Plant Taxonomy and Evolution

196
started with the assumptions listed in Table 10.1. Although
Bessey's assumptions are still widely accepted today,
some 60 years atter he proposed them, disagreements
continue to arise over specific interpretations. Adolph
Engler and Karl Prantl, living about the same time as
Bessey. constructed another classification scheme from
quite different assumptions.

Do species that now resemble each other represent

progeny from a common ancestor, or is possible that it

they started from quite different sources and simply

evolved to look alike 9 These two possibilities are
diagrammed in Fig. 10.4. It is seen that species A and B,
which are very similar today, actually arose from quite
different stocks. Similar environments, however, have
shaped them along convergent paths. This pattern is
termed convergent evolution. A striking example of
convergent evolution is the present similarity between
cactus plants of the southwestern United States and
certain Euphorbia species of Africa. Both groups grow in
hot, arid climates, and, as seen in Fig. 10.5, both have a
similar morphology. They have no leaves, their stems are
ribbed and swollen and contain water-storage tissue, and
they are armed with spines. Yet, other traits, like flower
morphology and geographic distribution, are so different
that is clear they must have arisen from different stocks.
it

On the other hand, B and C, which are today less

similar than A and B, actually did evolve from a common
ancestor. This pattern is called divergent evolution. As
time passes, 8 and C will become more and more
different. The same pattern will be repeated by Cand D,
but at this point in time they are still quite similar.
If a certain group of plants arose from a single ancestral
type, the group is referred to as monophyletic. If a group
arose from several different ancestral types, the group is Figure 10.5 Convergent evolution of members of the Cactaceae
of the south-western United States and members of the
(left)
referred to as polyphyletic. (Of course, the terms are
Euphorbiaceae (right) of Africa, x yl0 .

relative, because ultimately all life can be traced back to

some one, first cell that evolved in the primaeval seas.)
Bessey considered each separate group of seed plants
(e.g., the Anthophyta, Coniferophyta, Ginkgophyta) to be monophyletic
largest group of plants that is definitely is
monophyletic. Some taxonomists considered all the seed the family. (See the next section.)
plants together to be monophyletic. Still others did not At present, we still lack a classification scheme that all

think that even one group, the Anthophyta, was botanists accept. For convenience and consistency, this
monophyletic. Unless fossil evidence is at hand, it is very book follows a polyphyletic scheme by Scagel et al.

difficult to separate the effects of convergent and divergent (1967) and Bessey's views (Table 10.3).
evolution. At present, most taxonomists agree that the

How Are Plants Classified?

A 6 D C
Over 550,000 species of plants clothe the earth and
inhabit most of the waters to a depth of approximately 75
Present
Time m. What is a species 9 A species consists of groups of
morphologically and ecologically similar plants; these
groups may be called populations. Since the populations
within a species are reproductively isolated from the
populations of other species, genetic information is not
exchanged between them. Within a typical species,
Degree of Similarity
however, the plants are very similar in many aspects of

Figure 10.4 Hypothetical pathways of species evolution. Xand theirmorphology and interbreed freely. Their genetic
Y represent ancestral forms from which A, B, C, and D have constitutions are similar, even identical; they are very
since evolved. The paths leading to A and 6 represent convergent
evolution; the paths leading to B and C represent divergent closely related. Such populations of closely related,
evolution. interbreeding individuals constitute a species.

How are Plants Classified?

197
 Thus, the word "similar," in a natural classification,
means organisms with genetic constitutions indicating
-anthers
relationship. This system
is exemplified in the complete

standard classification of thesweet pea given in Table 10.2. Note

the characteristic ending for each category.
-•-androecium

Plant Groups
Broad groups of plants that include several divisions may
be arranged. Although these categories are meaningful,
they do not have taxonomic importance. For instance, all
plants can be divided in two groups: those with nuclei (the
eukaryotes, Fig. 10. IB) and those without nuclei (the
prokaryotes, Fig. 10. 7 A). A large group of primitive plants,
mainly aquatic and filamentous, may be set apart from
other plants by a very distinctive characteristic: the sexual
stigma
cells (eggs and sperms; that is, the gametes) are
Figure 10.6 Diagram showing traits that are important in the produced in structures not protected by a jacket of
classification of the sweet pea (Lathyrus odoratus L). A, flower,
exploded view; B, 10 stamens, 9 attached together and 1 free; C,
vegetative or sterile cells (Fig. 10. 7C). These plants may
pod with seeds. be referred to collectively as thallophytes. The
thallophytes constitute eight algal, one fungal, and a single

bacterial division (Fig. 10.8). Fungi and most bacteria lack

The most basic requirement of a biological classification chlorophyll. The orriy unifying characteristics of all the
is that show genetic relationships among the organisms
it remaining plants is that (1) the reproductive cells are
species may have some
classified. Several different protected by a jacket of sterile cells and (2) the embryo
characteristics common, and an occasional mating may
in derives its first nourishment from the gametophyte. A small
take place between members of two different but similar group of these plants typically lacks vascular conducting
species. Such genetically related species are grouped tissue.They are the liverworts and mosses; the
together in genera (s. genus). A comparison of some bryophytes (Fig. 10.9). All other plants have developed a
genera shows that they have traits in common. Genera specialized conducting tissue and are known as vascular
with similar traits are genetically related, but less closely plants.
than species within a genus; they are grouped in families. The less advanced vascular plants, called the lower
Plants in same family have common characteristics
the vascular plants, do not produce seeds. Examples include
that indicate they may all have evolved from the same the wisk fern (Psilotum, Fig. 10.10), horsetails (Equisetum,
ancestor. For instance, members of the pea family have Fig. 10.11), club mosses (Lycopodium, Fig. 10.12), and
flower parts pods like pea or bean pods, and
in fives, true ferns (Fig. 10.13). These comprise four divisions.
dissected leaves like pea or locust leaves. Many members The more advanced vascular plants bear seeds and are
of the family have flowers resembling the sweet pea, with seed plants. These are
frequently referred to as the
10 stamens, nine of them grown together (Fig. 10.6). separated into gymnosperms (Fig. 10.14), whose seeds
Families are grouped together in orders, and orders are are borne naked on a cone scale, and angiosperms (Fig.
grouped in classes. At the top level it is customary to 10.15), whose seeds are protected within an ovary.
divide all plants into divisions. Differences in The most conspicuous division of gymnosperms are the
photosynthetic pigments, manner of leaf development, conifers (Coniferophyta). Together with the angiosperms
structure of the conducting (vascular) tissues, and (division Anthophyta), theydominate the present flora. We
methods of reproduction are the most important elements may angiosperms into two subclasses. The
divide the
in separating plants into divisions. seeds of one subclass have two seed leaves, or
cotyledons, as seen in the peanut or bean. These are the
Table 10.2 Dicotyledonae. The seeds of the other subclass (onions,

Sweet Pea lilies, grasses, orchids) have a single seed-leaf. These are
the Monocotyledonae.

Classification Level Name Ending The relationship between the divisions and classes is

shown by the key in Table 10.3.

Specific epithet Odoratus —
Genus Lathyrus —
Family Fabaceae aceae Herbaria and Botanic Gardens
Order Rosales ales
Subclass Dicotyldeonae ae Several thousand new species of plants are discovered

Class Angiospermae ae and described every year. The plant specimens may have
Division (phylum) Anthophyta phyta been collected in parts of the world previously unexplored
Kingdom Plantae not consistent by botanists, although new species are being discovered
every year in areas that botanists have traversed many

chapter 10 |
Plant Taxonomy and Evolution

198
 -nucleoplasm

sterile jacket

fertile cells

dicryosome

Figure 10.7 Diagrams to show differences between groups of

plants. A, a prokaryotic cell (a blue-green alga); 6, a eukaryotic
cell (the green alga Chlorella), C, a group of reproductive cells
lacking a protective jacket (the brown alga Ectocarpus); D, a
group of reproductive cells provided with a protective jacket (the
liverwort Riccia).

times. These specimens ultimately fall into the hands of a

taxonomist who specializes in the group (family or genus)
to which the plant in question belongs. This botanist sets Methods of Taxonomic
out to identify the species by comparing with allied
forms, herbarium specimens, or published descriptions.
it

He
Research
may find that the plant has characteristics similar to those
of a species previously described, or he may find that the Traditional (Morphological)
planthas characters so different from those of any known
species that he concludes that it is a new species (species About 20 years ago, the geneticist C. H. Waddington said
novum, sp. nov.) that has never been described. it is "an empirical fact that living organisms do not vary

Accordingly, he describes this new species and gives it a continuously over the whole range, but they fall into more
name. The description is published one of the
in Latin in or less well-defined groups, which are commonly called
many recognized botanical journals, and the specimen or species." This philosophy is quite appropriate for
specimens he used in making the description are properly traditional taxonomists. They realize that the love of
labeled and placed in one or more of the herbaria of the classifying nature often putsboundary lines between
world. This specimen is known as the type specimen. things where boundaries simply do not exist; but when it
Herbaria (s. herbarium) are storehouses of pressed, comes to species, they are convinced from extensive field
dead, preserved plants. In contrast, botanic gardens, or observations that discrete species do exist and that
arboretums, "store" living plants. Some of the great classification at this level is not arbitrary or artificial.
botanic gardens of today, such as the Kew Gardens in Traditional taxonomists primarily examine plant
England, have a long history of distributing valuable crop morphology, searching for a few traits that consistently
and ornamental species to areas far from their native enable them to separate plants into "well-defined groups."
habitats. Sometimes the species actually grow better in In pre-Darwin days, a variety of traits were utilized, but
their new habitats because insect or fungal parasites are now the choice is limited to those which presumably are
absent. (a) genetically controlled (rather than environmentally

How are Plants Classified?

199
 n

Figure 10.8 Pigments, hence color, are of importance in

distinguishing divisions ot thallophytes. A, Amanita muscana is in
the Mycota; it lacks chlorophyll but does have a bright red
pigment in the cap. B, Ulva (sea lettuce) is in the Chlorophyta,
has chlorophylls a and b, and is green C, Postelisa (sea palm) is
in the Phaeophyta, and has chlorophylls a and c but they are
masked by a brown pigment. D, Iridophycus, and E, Porphyra,
are in the Rhodophyta, and they have chrlorophyll a and a red
pigment.

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200
Figure 10.9 The hair cap moss, Polytrichum, is a representative
of the Bryophyta. Figure 10.11 Horsetail (Equisetum telmateia L), the only genus
inthe division Sphenophyta.

Figure 10.10 Psilotum nudum, a living species closely

resembling fossils of the earliest vascular plants, found in
sedimentary rocks laid down about 400 million years ago.

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201
 a
1
Hi
^^3

^ afii'fMwC

Ik:
Figure 10.12 Lycopodium obscurum L (running pine), a
representative of one of the five genera in the division Lycophyta.
wm- i*9^.. k^^l *-* ^ '

',

**Sr^ ^* **•>* j

3k '••-
P'*
-

A
J'-y^i^C

Figure 10.14 The Coniferophyta. A, Pinus jeffreyi (Jeffrey pine),

closely related to P. ponderosa (ponderosa pine), the most
abundant western states pine. 6, an open cone of P. monophylla
(single-leaf pinon pine), showing two seeds on a scale.

Figure 10.13 The fern Polypodium vulgare, a representative of

the division Pterophyta.

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 —

characters to serve in determining the number of groups.

The characters they select show discontinuities and are
most helpful in separating related groups.

Anatomical
Anatomical studies of particular tissue and cell types
within the vascular system reveal evolutionary patterns.
Vessel elements, for example, show the changes
summarized in Fig. 10.16. Evidence for this set of
hypotheses comes from examination of fossils, of lower
plants that have vessels (such as Selaginella), and of
correlations between changes of vessel characters with
changes in other characters such as flower morphology
(i.e., plants with primitive flower morphology also show

primitive vessel characters). Similar hypotheses of

evolution have been applied to other cell types, such as
fibers.

Biochemical
We have already seen (Table 10.3) that algal divisions are
separated by biochemical characteristics, such as the type
of chlorophyll or accessory pigments, the major
Figure 10.15 The angiosperms, or Anthophyta. A, the flower
and Citrus sinensis (orange). 6, the fruit of Capsicum
fruit of
component form
of the cell wall, or the principal of food
frutescens var. grossum (bell pepper), showing seeds within the stored. Only within the 20 years, however, has
last
matured ovary.
biochemistry been regularly applied as a taxonomic tool
for higher groups of plants. Members of closely related
taxa are chemically treated to extract particular

controlled) and (b) conservative in the evolutionary compounds, and the amounts (or presence versus
sense. Flower and fruit morphology, for example, meet absence) of the compounds extracted are compared. The
these criteria, but leaf size does not. Leaf size, and other more chemically similar that taxa are, the closer their

traits that can be modified by the environment, are said to genetic relationship is presumed to be.

be plastic or variable. Almost every category of compound has

biochemical
It should not be supposed that traditional taxonomists
been used at one time or another in these studies
are limited at the start of a study to the examination of sugars, amino acids, fats, oils, alkaloids, alcohols,

only a few They examine many characters, measure

traits.
terpenes, and phenols —
and they have been isolated by
many plants, and keep records of all of them. They paper chromatography, electrophoresis, fractionation or,
eventually, however, subjectively select only a few more recently, gas chromatography. Many of these

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203
Table 10.3
Synopsis of Plant Divisions Covered in This Book

Common names
Divisions or examples

A. Plants without roots, stems, or leaves, and no protective jacket of thallophytes

vegetative cells around the developing reproductive cells
8. Plants lacking chlorophyll
C. Prokaryotes Schizomycophyta bacteria
C. Eukaryotes
D. Vegetative stage of naked cytoplasm Myxomycota slime molds
D. Vegetative stage with a cell wall, mostly filamentous Eumycota fungi
B. Plants with chlorophyll algae
C. Chlorophyll a alone
D. Prokaryotes Cyanophyta blue-green algae
D. Eukaryotes
E. With water-soluble blue and red pigments Rhodophyta red algae
E. With plastid pigment xanthophyll Xanthophyta
C. Chlorophylls a and c
D. Large seaweeds with a cell wall of cellulose Phaeophyta brown algae
D. Small, mostly unicells, cell wall of cellulose, two anterior Chrysophyta golden algae
unequal flagella

D. Small, mostly unicells, cell wall of silica Bacillariophyta diatoms

D. Small unicells lacking a cell wall or with cellulose plates, Pyrrophyta
two lateral unequal flagella

C. Chlorophylls a and b
D. Unicells without a cell wall Euglenophyta Euglena
D. Greatly diversified forms with cell walls Chlorophyta green algae
A. Protective jacket of vegetative cells present; roots, stems, and
leaves may be present
B. Plants lacking vascular tissue Bryophyta liverworts, mosses
B. Plants with vascular tissue vascular plants
C. Plants without seeds lower vascular plants
D. Roots absent Psilophyta Psilotum
D. Roots present
E. Small leaves, no gap at union with stem, leaves do not
unroll as they open
Stems not jointed
F. Lycophyta club mosses
Stems jointed
F. Sphenophyta horsetails
E. Leaves well-developed, gap at union with stem, leaves Pterophyta ferns
unroll as they open

C. Plants with seeds seed plants

D. Seeds not covered gymnosperms
E. Seeds in cones
F. Trees palmlike Cycadophyta cycads
F. Trees conical Coniferophyta conifers
E. Seeds not in cones like those above

F. Leaves fan-shaped and deciduous; trees Ginkgophyta Ginkgo

F. Leaves not fan-shaped; shrubs, vines, or prostrate Gnetophyta Ephedra,
perennials Gnetum,
Welwitschia
D. Seeds covered Anthophyta angiosperms
b
E. Two-seed leaves Dicotyledonae dicotyledons
b
E. One-seed leaf Monocotyledonae monocotyledons

a
A few bacterial species do have chlorophyll.
"Classes.

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204
 turnedoff, and the gel is "developed." The gel is

immersed in a solution that contains a substrate upon

which the enzymes under study act. The solution also
contains a stain that couples with the product of the
enzyme-substrate reaction. The result is that bands of
color on the gel mark the locations of very specific
enzymes. The distance that the enzymes have moved from
their origin depend on their charge and shape; the number
of bands reveals the number of enzymes capable of
reacting with the substrate (Fig. 10.17).

Biological

Biological taxonomists are interested in determining what

the natural biotic units are. By natural biotic units they
mean populations of plants that maintain their
distinctiveness because of biological barriers that
genetically isolate them from other populations. These
isolating barriers may be due to breeding behavior (time of
flowering, type of pollinator), habitat or geographic
isolation, or to the inability to form fertile hybrids with
closely related groups (sometimes because of differences
in chromosome number or chromosome morphology).
Biological taxonomists often deal with populations at,

and below, the rank of species because they are vitally

concerned with detecting divergent evolution at an early
stage. They are interested in discovering the ways in
which species become distinct and in which they are able
to maintain their distinctiveness even though living in close
proximity or in similar habitats. Biological taxonomists are
referred to as biosystematists, and their field as
biosystematics.
Many species that are widespread and occupy a variety
of habitatsmay be composed of several genetically distinct
subspecies. The subspecies are all interfertile, but hybrids
Figure 10.16 Vessel elements from members of the Anthophyta.
A, primitive type from Liriodendron tulipifera (tulip tree); B, are not common because hybrids are less suited to the
advanced type from Quercus (oak). habitats than are parental types. Drs. Clausen, Keck, and
Hiesey of the Carnegie have shown the
Institution

existence of subspecies in plants such as five-finger

compounds are high in molecular weight and are
(Potentilla glandulosa), which grows in California from the
structurally complex. It is reasoned by biochemical
coast near Stanford to high elevations near timberline in
taxonomists that these complicated molecules probably
the Sierra Nevada mountains.
evolved at only one time and
in only one original group of
Clausen, Keck, and Hiesey dug up plants of five-finger
plants; therefore, species or genera or families that
that were growing all along this range and transplanted
possess the same complicated molecules are undoubtedly
related to this original group of plants.
One of the most widely used methods for comparing
taxonomic similarity of organisms at the molecular level is
gel electrophoresis. This method detects differences
between enzymes on the basis of
(or other proteins)
differences in their electrical charge and shape. Such
differences are the result of changes in the amino acid
sequence of the molecule. Since amino acids are coded
for by small segments of the DNA molecule, amino acid
differences are thought to be equivalent to differences
between single genes.
A small amount of liquid extract of crushed leaves (or of
any other plant organ desired) is placed in a notch cut
into a gelatinous material. This gel, often made from potato
starch, either fills a tube or coats a glass plate. The
Figure 10.17 A "developed" gel of 11 Clarkia genotypes. The
extract is placed at one end of the gel, and an electric enzyme bands have migrated downward from the "top" of the
current is sent across it. After several hours, the current is gel.

Methods of Taxonomic Research

205
them together in a uniform garden at Stanford. In the Dandelions and herbaceous annuals, on the other hand,
uniform garden, any differences among the plants from invest a high proportion of theirenergy in seed production
would be due to genetic control They
different locations and have a short life cycle. Each seed is and often
small
found morphological differences: timberline plants were wind-disseminated, which means that the young seedling
much shorter than lowland plants. They also found has very little food reserve to use while it establishes itself

physiological differences: the timberline plants flowered as a plant. The result most seedlings perish. But,
is that

earlier in the summer than lowland plants. When five- because the reproductive capacity of the few survivors is
finger plants from different locations were grown together so high, the population density is maintained. This is

in j garden near timberline, these differences proved called an r strategy. The letters r and K are components in

critical to survival: lowland plants that did not become an equation for population growth rate. Obviously, both K
dormant were killed during the winter, and lowland plants and r strategies can lead to success in an evolutionary
that flowered late in summer were nipped by frost before sense. The question is, under what set of environmental
seeds could be produced. Clausen, Keck, and Hiesey conditions will each strategy be more successful? The
named the lowland group P. glandulosa ssp. concept of r and K strategies was first proposed in 1962
( = subspecies) typica and the timberline group P.
by the very innovative ecologist Robert MacArthur.
glandulosa ssp. nevadensis (Fig. 10.18).

Biosystematists are also interested in comparing taxa

Numerical
(taxon, singular; plants of any taxonomic rank) on the
basis of their survival strategies. All the approaches to taxonomy just discussed share one
One example can be seen by
of different strategies feature in common: each
subjectively selects one form of
comparing an oak with a dandelion. Oaks and other large similarity on which
base conclusions, and each weights
to
perennial plants devote a lot of energy to the accumulation its own choice of evidence as more important toward

of great bulk that can withstand many environmental understanding relationships than any other form of
stresses. The price oak "pays" for its size is a long evidence. Traditionalists weight morphological evidence,
juvenile period of development when reproduction is nil, biosystematists weight breeding behavior, and so on.
and even after that period the amount of energy put into Another approach to taxonomy is to consider all forms
flower and seed production is low, compared to the of evidence with equal emphasis; all evidence has equal
energy invested in the total mass of the plant. Seeds weight. This is a statistical approach and is called
(fruits) are large and are disseminated by gravity, water, or numerical taxonomy. A basic tenet of numerical
animals. Seed production and seedling establishment can taxonomists is that a great deal of evidence is required to
be very low for many years in succession because of poor objectively separate taxa from utilize 50 each other. They
environmental factors, or internal factors, but the to 300 characteristics for each study.
population density remains constant because of the long The numerical taxonomist does not admit that discrete
life span of individuals. This is called a K strategy by species exist in nature. He/she insists that when many
ecologists. traits instead of a few are considered, a continuum of

Figure 10.18 Growth of Potentilla glandulosa ssp. typica after

one growing season at each transplant garden. This coastal
subspecies, or ecotype, grew best in its normal environment at
Stanford (left); grew very poorly at timberline (right) and was
it

by the following winter temperatures. Middle plant was

killed
grown at an intermediate elevation.

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206
variation willemerge. Boundaries between species can be little changed from its original form; it may be the
erected by taxonomists, and these boundaries may be microscopic cell wall of a spore or pollen grain, which is

objectively selected; but the boundaries are arbitrary and also very resistant. But most plant parts are not hard, and
artificial in the final analysis, according to numerical plant fossils seldom consist of unchanged structures. They
taxonomists. The viewpoints of traditional and numerical often are simply the impressions of leaf or stem fragments
taxonomists, then, are quite different. Interestingly, thai —
were trapped in mud mud that later was
classification schemes arrived at by numerical means often compressed into sedimentary rock (Fig. 10.19). To yield a
show good agreement with those arrived at by traditional good fossil, plant parts must settle in quiet waters, then be
means. Can we conclude that experienced, traditional buried under silt or volcanic ash to lie trapped in
taxonomists are just as effective as the computers used in surroundings unfavorable to decay. They are seldom
numerical studies? entombed where shed but, more typically, are first carried
by water or wind for some distance. the plant part isIf

large enough, for example, a tree trunk, the interior of the

Evolution cell may be replaced by silica quartz crystals. The tissue

pattern and anatomical details are faithfully preserved (Fig.

10.20). In addition, degradation products from tissue
In a broad sense the term evolution refers to a process
decay may remain trapped as "chemical fossils." The
involving gradual changes. It is well known that neither
porphyrins from chlorophyll degradation, for example, are
animals, nor plants, nor cities, nor states remain the same
very inert and long-lasting. Their presence in sediments
through time. We may speak of the evolution of the means
indicates that photosynthetic plants were present at the
of transportation, the evolution of human clothing, the
time the sediments were laid down.
evolution of mountains and valleys, and the evolution of
It is important to note that plant fossils consist almost
many other nonorganisms.
remains that grew in or near water or
entirely of plant
Organic evolution pertains to the gradual changes that
were carried by streams into lakes, bays, or other sites of
have taken place in living organisms. Coal, for instance, is
deposition. The remains of plants growing in arid or
formed from the remains of plants that were different in
mountainous regions are rarely deposited where
appearance from those growing today. The student of
conditions are favorable for their preservation as fossils.
organic evolution may be interested in, among other
Either it is too dry or the site is undergoing erosion and
things, accounting for the disappearance from the earth's
flora of the Coal Age plants and the appearance of the
the deposits are of very short duration. This means that
the geological record of plants must probably remain
modern flowering plants.
incomplete and that the records we do have mainly
represent the floras of lowlands and moist habitats.

The Geologic History of Plants Considerable evidence indicates that the great
environmental extremes prevalent in arid or semiarid
Our understanding of plant history comes from the mountainous areas are especially favorably to rapid
examination of fossils. A fossil may be a shell or bone, evolution. This factor, we shall see, may be very important

Figure 10.19 An imprint of a leaf of Pecopteris, a common plant

of the Pennsylvania geologic period, about 300 million years ago.

Evolution

207
Figure 10.20 A thin section ot petrified wood, as seen through a
light microscope. Tracheid lumens are filled with silica stone.

in connection with the problem of determining the origin of conditions were perfect for preservation of the organisms.
the angiosperms and may serve to explain peculiarities They were preserved in the sediment in a siliceous
that surround the first records of the angiosperms. solution that later crystallized into rock called chert, much
as a modern biological specimen is preserved by being

When Did the World Begin? embedded in plastic. The soft bodies of these early
organisms were not distorted because the silica matrix of
When no
did the world begin? Earth scientists have chert is incompressible. Today, these beds of chert, up to
sure answer, but the calculations of astronomers and 130 m thick, are exposed. When thin sections are placed
physicists, coupled with known age of meteorite fragments
on a microscope slide, the perfectly preserved organisms
and of rocks from the moon, lead us to estimate that the
(microfossils) can be seen. The chert itself has not been
surface of the earth cooled to a crust about 4.5 to 5.0
dated, but rock layers above and below have. By it

billion years ago. This primordial crust probably contained

inference, the age of the chert is estimated to be 3.2
uplands, ocean basins, plateaus, and mountainous
billion years.
volcanoes. Volcanic eruptions filled the atmosphere with
Many unusually shaped structures, which may or may
carbon dioxide, methane, carbon monoxide, nitrogen,
not be organisms, have been seen in this chert (called the
hydrogen, and strong-smelling ammonia and hydrogen
Fig Tree Formation), but only two are seen frequently
sulfide. Free oxygen was absent. The first rains washed,
enough and in great enough detail to leave no doubt that
dissolved, and eroded the uplands, carrying many
they once lived. One is a rod-shaped bacteriumlike cell,
nutrients to the young oceans.
called Eobacterium isolatum (Fig. 10.21), the other is
Life, or some sort of half-way form of life, probably
possibly a blue-green alga, Archaeosphaeroides
originated very early. It may have consisted of aquatic,
barbertonensis. Organic residues were also found in the
smali, self-replicating, heterotrophic cells that lived on the
rich,anaerobic oceanic soup around them. However, our chert. Analysis of them showed the presence of certain
present methods of detecting remains of ancient life hydrocarbons that can most reasonably be regarded as
require that the fossils be preserved in rocks. So far, the breakdown products of chlorophyll. Analysis also revealed
oldest rocks found on earth are 3.4 billion years old. This that the ratio of carbon-13 to carbon-12, two natural

means that we have no record of the first 1 to 1 .5 billion isotopes of carbon, was lower than the ratio in today's
years of earth's history (Table 10.4). atmosphere. When plants photosynthesize, they "prefer"

Aquatic life did exist 3 billion years ago, for we have to use carbon-12; this means that their tissue and fossil

fossil evidence of it. Some of the oldest rocks known lie residues have a low carbon-13 to carbon-12 ratio,

exposed near the gold-mining town of Barberton, on the corresponding to the ratio in the chert. This combination
border between the Republic of South Africa and of direct and indirect evidence strongly suggests that plant
Swaziland. About 3.2 billion years ago, this area was a life existed 3.2 billion years ago.
shallow, warm sea or embayment. Living things existed in A similar, but richer, collection of microfossils has been
thin sheets at the bottom of a silica-rich sea. Apparently, preserved in another chert deposit, the Gunflint Formation,

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208
Table 10.4
6
Geologic Time and the Dominance of Different Plant Groups through Time

Began (millions of
Era Period Epoch or part years ago) Dominants

Cenozoic Quaternary Recent Last 1 2,000 years

Pleistocene 2.5 Flowering Plants

Teritiary Pliocene 7
Miocene 26
Oligocene 38
Eocene 54
Paleocene 65
Mesozoic Cretaceous Upper 90

Lower 136 Gymnosperms

Jurassic Upper 166

Lower 190

Triassic Upper 200

Lower 225

Paleozoic Permian Upper 260

Lower 280 Lower vascular plants

Carboniferous Pennsylvanian 325

Mississippian 345

Devonian Upper 360

Middle 370

Lower 395 Algae

Silurian 430

Ordovician 500

Cambrian 570

Proterozoic Precambrian 4,500-5,000

aShaded sections were times of great evolutionary changes in the plant kingdom. The time scale is not drawn to scale. All named epochs
do not appear on this chart.

along the Ontario shore of Lake Superior. By dating layers and Nostoc; (b) bacteria; (c) eukaryotic fungi; and
above and below it, geologists estimate this deposit to be (d) eukaryotic green algae. Cytological details are
2 billion years old. Filaments are most abundant, and preserved so well that stages of cell division can even be

some resemble modern blue-green algae (Fig. 10.22). detected (Fig. 10.23), but you should be aware that the
Small spheres that might be colonial blue-green algae or interpretation of cell details is not uniform, and some
spores are also common. Analysis of the organic residue biologists claim that the "nuclei" are artifacts and that the
and determination of the carbon isotope ratio show that cells are those of prokaryotes.
photosynthetic organisms were present. The origin of If fossil eukaryotes, then perhaps they could
these are
photosynthesis was important; most scientists believe that reproduce sexually. Sexual reproduction brings with it
the 2 of the atmosphere resulted from it. and a more rapid rate of evolution.
variation in offspring It

The first eukaryotic plants may have appeared 1 billion enabled a greater diversity of forms to develop and
years ago. Fossils in the Bitter Springs Formation, about proliferate. A spurt of evolution resulted. By the end of the
65 km northeast of Alice Springs, in the heart of Australia, Proterozoic era (Table 10.4), the seas had become
are rich and varied, and indicate that at least four groups crowded with life, including many forms of animals and
of organisms existed at that time: (a) prokaryotic plants. The rate of oxygen formation had increased; by the
filamentous blue-green algae, akin to modern Oscillatoria end of the Proterozoic era the level of oxygen in the air

Evolution

209
 effects were minor. It was a time when a climate much like
that in today's tropics dominated the entire world. It was a
time of enormous evolutionary change in the plant
kingdom. However, the rate of evolution did not proceed at
a constant pace during the entire Paleozoic era. Most of
the major changes in plant form appeared in only 25 to 50

million years, during the upper-Silurian to mid-Devonian

periods. It was then that plants came to dominate the land
and that the oxygen level of the air may have reached

10% of the present level.

Through the Cambrian, Ordovician, and most of the
Silurian periods, algal forms continued to dominate the
plant kingdom. Blue-green and green algae were most
abundant, but lime-encrusted red algae and some brown
algae resembling Laminaria and Fucus were also present.
The lime-encrusted forms secreted calcium carbonate just
Figure 10.21 Oldest known bacterium, Eobacterium isolatum, as modern forms do.
from the Fig Tree Formation, about 3.2 billion years old.

Invasion of the Land: Silurian and Devonian

may have been 1% of its present level. Oxygen not only Periods
supports life; it also screens out ultraviolet radiation from
the sun.The radiation can be damaging to cell activities. What are some of the adaptations required for plant life on
This amount of oxygen —
1% of present level produced — a dry land? In water, plants need no complex structures for
sufficient filter of radiation to permit life in all but the top 2 support nor for the uptake of nutrients. The surrounding
to3 cm of water. Life on land, except in sheltered places, water buoys them up and bathes them with soluble
would still have been impossible. nutrients. In contrast, land plants must not only develop
A "sudden" evolutionary spurt 600 million years ago roots for the absorption of nutrients and the elaboration of
marked the beginning of the long Paleozoic era, a time stiff tissue for support, they must provide pathways for

when much of the land was low-lying, and inland seas water and nutrient transport —
xylem and phloem.
moderated the climate so that seasonal and latitudinal Reproductive cells on land must be carried by agents

Figure 10.22 The filamentous blue-green alga Animikiea septata

from the Gunflint Formation, 2 billion years old.

Figure 10.23 Sequence of fossil impression from the Bitter

Springs Formation arranged to show cell division.

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210
other than water, and they must be thick-walled and Leaves had also evolved, changing from very small
cutinized to avoid desiccation. All external surfaces of land epidermal outgrowths of the primitive vascular plants, to
plants, in fact, must be adapted to reduce water loss; larger true leaves having a branching vascular system, as
these adaptations can take the form of stomata and found in Aneurophyton and Archeopteris. The "true"
cuticle. leaves are called megaphylls; the epidermal outgrowths
Despite tempting fragments of evidence, such as are called microphylls.
cutinized spores and bits of xylem dating back to the
Cambrian period, the first undisputed fossils of terrestrial, Coal Age Forests and Seed Plants
vascular plants do not appear until the upper-Silurian
period. But what appeared at that time was not just one Plant development during the Devonian period not only
Many types suddenly occur the modified the vegetative size of plants, it affected their
type of vascular plant. in

representatives of the Psilophyta, Lycophyta,

reproduction as well. The seed plants developed in
first
fossil record:
the upper Devonian period, but it was not until the Coal
Sphenophyta, and pre-ferns. Thus we can surely believe
that the first vascular plant evolved many millions of years Age (Mississippian and Pennsylvanian epochs, also known
before these many types. Did they all evolve from a as the Carboniferous period) that seed plants became
common ancestor or did they evolve from several separate abundant.

lines? At this time there is not enough evidence to answer Extremely lush, swamp forests dominated the Coal Age
the question. landscape (Fig. 10.26A). Because the land was low, minor
The early vascular plants consisted of slender forking changes in sea level successively inundated, buried, then

stems, with or without leaflike appendages, and with supported one forest after another. The buried organic
sporangia either at the tips of the branches or along their remains have become compressed and changed through
sides. Perhaps the most primitive example is Cooksonia, time. Today, they form the coal, gas, and some of the oil

discovered in 395 million-year-old fossil beds in reserves of the world. These sources of energy are the

Czechoslovakia and Wales, from the end of the Silurian chemically changed, fossil remains of Coal Age forests
period (Fig. 10.24). It was a plant probably less than 10 and, for that reason, they are referred to as fossil fuels.

cm tall, made up of dichotomizing branches less than 4 Burial of plants and transformation into fossil fuel have
mm in diameter that terminated with sporangia. The continuously taken place throughout time, but not at the
spores had a waxy cuticle, indicating that they were rate it did during the Coal Age (Fig. 10.268).
adapted for dissemination on land. The xylem in the stem These forests were dominated by Lycophyta. In the
was a solid, central core; there was no pith. We know Devonian period, this division evolved rapidly into both
nothing of the root system. Modern, living wisk fern herbaceous and woody types; in the Coal Age, the woody
(Psilotum, see Fig. 10.10) still retains many of these types reached tree stature. One example is Lepidodendron
primitive features. (Fig. 10.27), which is up to 35 m tall and with a trunk 1 m
From this primitive, herbaceous start, evolution across. Straplike leaves and sporangia occurred near the
progressed rapidly to the first trees only 25 million years ends of the branches. A cross section of the trunk reveals
later. Aneurophyton, for example, was up to 12 m tall. pith, primary and secondary xylem, cambium, and an
Archeopteris was more than 30 and up to 1 .5 m in m tall enormous amount of cortex and cork. Very little of the

diameterat the base (Fig. 10.25). Both had evolved pith, stem area served for conduction or strengthening. The
cambium, and the capacity to produce considerable roots were dichotomously branched.
secondary xylem. Since the petrified wood samples lack Second in abundance were giant horsetails, such as
annual rings it is not possible to determine their age. Catamites (Fig. 10.28). Catamites was smaller than
Lepidodendron, but was still 10 m tall and had a trunk
about 30 cm in diameter. Whorls of branches developed,
sporangia from which formed smaller branches; and from these
branches leaves developed at the nodes. The upright
stems arose from a horizontal rhizome system.
Third in abundance —
not tall, but dominating the forest
floor —
were ferns and seed ferns. Seed ferns were
fernlike, but in addition bore seeds on their large leaves

instead of spores. Exactly from what plant group the ferns

evolved is not clear, for they appear rather suddenly in the
fossil record in the Coal Age.
Gymnosperms (other than seed ferns) were fourth in
abundance, but these were forms quite unlike modern
gymnosperms and were not abundant. Gymnosperms did
not become abundant until 100 million years later.
Herbaceous forms of Lycophyta and Sphenophyta were
common, but members of the Psilophyta — direct
descendants of the first, weak, vascular plants to climb

Figure 10.24 A reconstruction of Cooksonia, one of the first

onto land only 70 million years earlier — were rare,

vascular plants. The above-ground portion, shown here, was less although we have only a few fossils from that time. The
than 10 cm tall. lower vascular plants dominated world vegetation during

Evolution

211
Figure 10.25 Reconstructions of large Devonian trees. A,
Aneurophyton, 7 to 13 m tall; B, Archeopteris, about 30 m tall.

the Coal Age just as thoroughly as the algae dominated the Mesozoic era was the age of the gymnosperms.
the world in previous ages. The dominance of the lower However, the group that dominates the world today, the
vascular plants was short, however, and great extinctions flowering plants, must have been evolving all through the
lay ahead. Mesozoic era. Pieces of wood, leaf impressions, and
pollen scattered through the geologic record as far back
Permian Extinctions and on to the Mesozoic Era seem
as the Triassic period to be angiosperm in nature.

The 375 million-year-long Paleozoic era was marked by But the first uncontestable appearance of fossil

two concentrated periods of evolution: a 25 million year angiosperms is in the Cretaceous period.
period of innovation in the lower-Devonian period during The Cretaceous period was a third interval of rapid

which weak vascular herbs led to forest trees; and a 50 evolution, undoubtedly because of major climatic and
million year period of major extinctions in the Permian geologic changes that took place. Two major biological
period. These changes were associated with increasing changes at the close of the Cretaceous period were (a) a
oxygen in the air; the extinction may have been caused by spread of the flowering plants, and (b) extinction of the
a cooling and drying climate and an uplifting of the land. dinosaurs. A number of other changes in the plant world
In many ways, the sudden rise and fall of the Coal Age accompanied all this (e.g., the extinction or near-extinction

forests are just as striking and mysterious as the rise and of several gymnosperm groups), but the rise of the
fall of the dinosaurs many years later. The Permian angiosperms was the greatest change. Where had they
extinctions marked the start of the Mesozoic era. come from, and why did they evolve so quickly? We may
The plants that replaced the lower vascular plants in never know the answers. At present, one leading theory is
dominance were gymnosperms: the true conifers that flowering plants evolved over a long period of time
(Coniferophyta), Cycadophyta, and Ginkgophyta. Most of This evolution took place in tropical uplands where fossil

chapt er 10 |
Plant Taxonomy and Evolution

212
 preservation in sediments is rare. These early angiosperms
may have occupied warm, seasonally wet, rocky slopes
with great variations in microhabitats resulting trom
ditterences in exposure, elevation, drainage, and soil type.

This environmental variability could have been a stimulus

to evolution. As the land lifted and Cretaceous seas
withdrew, new lowlands could have been rapidly invaded
by diverse forms of flowering plants that had evolved on
mountain slopes. By the time the Cenozoic era began, 65
million years ago, the conifers dominated only the cold
temperate and polar regions, and to the angiosperms
belonged everything else.

Climatic Change in the Cenozoic Era

The gradient of temperature with latitude that we have

today apparently did not exist at the beginning of the
Cenozoic era. Looking at fossil deposits of marine
plankton, we can estimate the ocean surface temperatures
from the equator to the north pole at past times. In the
mid-Cretaceous period, the difference from equator to pole
was only 1 1 °C, but by the end of the Cretaceous period, it

Figure 10.26 Coal Age forests, and the fossil fuel results. A,
reconstruction of such a forest, courtesy of the Field Museum of
Natural History. Note the seeds attached to fernlike leaf in left
center. B, cartoon relating the age of dinosaurs wrongly with the
coal age. Dinosaurs flourished 100-200 million years after the
coal age.

Evolution

213
 pith

primary xylem

secondary xylem

cambium

Figure 10.27 Lepidodendron, a dominant of the Coal Age forest.

A, reconstruction of the entire plant, about 30 m B, cross
tall;

section of the trunk.

had grown to 15°C. The difference continued to grow of vascular plants appear to be expanding in terms of
during the Cenozoic era. We
can guess that similar diversity and abundance: ferns and angiosperms. Both
temperature changes occurred on land. In the past, the groups have survived great physical changes of the earth,
climate over the globe was broadly zoned and without yet their evolution seems to have been stimulated by these
temperature extremes, but today many climatic zones exist stresses, while many other forms have become extinct or
between the pole and the equator. Figure 10.29 shows survive as tenuous remnants.
how these zones may have formed and shifted during the By the end of the Ice Age the human species began to
last 60 million years. During Pleistocene glaciation, the play a role in the evolution and distribution of plants.
temperature gradient was at a peak. Although the evidence for seed
earliest archeological
The Pleistocene Ice Age, which ended only a moment such as grain, beans,
agriculture (cultivation of annuals
—
ago in geologic time about 12,000 years produced — and squash) takes us back to 9000 B.C., people may have
another period of extinctions. Glaciated areas were been cultivating root crops (perennials propagated by
scraped clean of plants, and they are still today being cuttings and harvested for starch in the "roots", such as
slowly revegetated. sweet potato and taro) in southern Asia as long ago as
In the Recent period, conifers have become more and 13,000 B.C. Some of these root crops have been
more restricted in the land area they dominate. The asexually propagated for so many years that they have
flowering plants, however, continue to evolve, especially in lost or nearly lost the capacity for sexual reproduction,
harshly cold or dry environments. Today, only two groups and it is doubtful that they could survive in nature.

chapter 10 |
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214
 People have also accidentally domesticated and favored
the evolution of certain other plants, the weeds. These
plants grow well in disturbed or trampled soil, in waste
areas rich in nitrogen, or in cropland. Some of them have
evolved seeds that imitate the size of crop seeds, so they
may not be separated during the threshing or sieving of
crop seeds. When the next season's crop is sown, the
weeds are sown inadvertently along with it.

Charles Darwin and Evolution

Many biologists since the time of the Greek philosophers

have attempted to explain the mechanism of evolution.
Great progress has been made during the nineteenth and
twentieth centuries. Our better understanding was made
possible chiefly because the works of Charles Darwin and
Gregor Mendel provided a basis that enabled biologists to
approach the problem of evolution in an entirely different
way.
Darwin, as a result of many years of careful study and
observation of a large number of plants and animals,
emphasized the following points:

1. The numbers of plants or animals may increase in a

geometric ratio. For example, a given plant may
produce 1000 seeds. If each seed grows into a new
plant and each new plant produces 1 000 seeds, 1
million new plants could result. A third generation from
the one original plant would result in 1 billion plants.

This example shows that plants (and animals also) have

the potential to increase their numbers at a tremendous
rate.
2. Actually, the number of individuals of a given plant or

animal remains fairly is no such

constant. There
tremendous increase in the number of individuals that
seed production seemingly makes possible.
3. No two individual plants or animals are identical; there
is variation.

Reasoning from these observations, Darwin arrived at

these conclusions:

1. Any given population is usually able to reproduce many

more young individuals than can adequately be raised
in the region it occupies. Therefore, a struggle for
existence occurs among the individuals.
2. In the struggle for existence, only those individuals
survive that, because of their particular variation, are
best adapted to their immediate environment. Thus, a
natural selection takes place; the unfit do not survive or
do not reproduce.
3. These selected variations may be inherited, that is,
Figure 10.28 Catamites, another Coal Age forest tree. A,
reconstruction of an entire plant, including rhizome and roots, of passed from one generation to another and, thus, may
C. carinatus. B, detail of whorled leaves of the form species gradually give rise to new species.
Annularia radiata, which are very similar to the leaves of C.
carinatus. The leaves are about 1 to 2 cm long.
The Sources of Variation

Since variation plays such an important part in plant

Annual grains have been selected for productivity, rather evolution, let us examine the sources of variation a little

than natural survival, and the result is they have become more closely.
so changed from their wild relatives that it is difficult to One source of variation is mutation, the spontaneous
determine where, and from what stock, they were first transformation of a gene (Fig. 10.30). Mutation does not
domesticated. create new genetic material (i.e., new DNA); it simply

Evolution

215
 \\ WARM
V^TEMPERATE
subtropical''

Figure 10.29 Climatic and vegetation zones in North America

during the past 40 million years. A, Oligocene; 8, Miocene; C,
Pliocene; D, at the peak ot the Pleistocene Ice Age; E, present.

chapter 10 |
Plant Taxonomy and Evolution

216
 Figure 10.30 Examples of mutations. A, common wild sunflower;
B, mutant sunflower, known as sun gold; C, variation in the seed
coats of beans.

changes the arrangement ot the existing material so that translocations. A final powerful source of variation is

the enzymes and other proteins encoded by the DNA are recombination, the reshuffling of chromosomes during
no longer quite the same. A mutated gene can back- sexual reproduction (see chapter 3).
mutate to the original condition also. Although several
environmental tactors, such as radiation, can cause a
mutation, most mutations seem to result trom cellular
"mistakes" in copying the DNA molecule during cell Table 10.5
division. Mutation Rates of Different Genes in Corn (Zea)
Mutations are a universal tact ot They are known to
lite.

occur in every plant and animal that has been studied. Number of Mutations
This does not mean that a given gene is relatively unstable Gene per 1,000,000 Gametes
and is likely to mutate very otten. Rates ot mutation for an
individual gene, or locus, probably average one for every Seed color, not purple 492
100,000 cells. The mutation rate does vary from gene to Seed color inhibitor 106
gene, however, as shown for corn (Table 10.5), where Purple seed color 11
one gene may have a mutation rate 500 times that of Sugary seed 2.4
another. Shrunken seed 1.2
Other sources of variation include chromosome Waxy seed Less than 0.1
aberrations such as deletions, duplications, inversions,

Evolution

217
 The Role of Natural Selection collected and sown back at sea level, normal tall plants
would There had been no genetic change only a
result. —
Mutation and recombination produce new patterns of temporary, plastic response to a harsh environment. Such
heredity, which result in variation among temporary changes are called phenotypic changes. In
individuals. However, for an entire population of individuals contrast, genotypic changes (changes in the genes) are
in a species to progressively change into something new, passed on to offspring.
more than this raw material is needed. Some force or Darwin's concept of evolution, expressed over a
pressure must be exerted on the population so that the hundred years ago, is the theory accepted today, not
abundance of certain mutations becomes higher and Lamarck's. Its differences with Lamarck's are illustrated in
higher, until all or most members of the species possess Fig. 10.31. Basically, Darwin's concept is this: Variation

it. How does this happen? The presently accepted answer exists in the initial population; an environmental stress

had its own evolution, over the course of 150 years of places certain individuals at an advantage; because those
debate and experimentation. The answer is based on the individuals survive or reproduce more successfully, they
idea of natural selection. leave more offspring that carry the same genetic traits; the
At the beginning of the nineteenth century, the French abundance of the advantageous traits in this way
naturalist Jean Baptiste Lamarck (1744-1829) was a increases in every generation, but variation still persists.
liberal — for his —
day in his thoughts on evolution. This process of directed change is called natural
Lamarck did not believe that all species were of the same selection.
age, that all were created together at one time. Neither did
he believe that the same species always existed. He
believed that new ones were forming all the time as a How Species Remain Distinct
result of changing environments. This stand was quite
Evolution proceeds most rapidly in a population of
heretical, considering that Linnaeus 50 years earlier had
organisms if it is "isolated" from other populations. But we
practically founded "modern" taxonomy on the principles
mean a very special kind of isolation: reproductive
that there was only one time of creation, and that all the
isolation. If plants in one isolated population cannot breed
species in the world from that time on were fixed and
with similar plants in other populations, then natural
constant. Lamarck thought that new traits and new
selection will produce a distinctive species in the shortest
species could evolve from old ones if a species were
amount of time. This species will from allremain distinct
placed under stress. For example, if a tall plant at sea level
others, once has formed (Fig. 10.32). Without isolation,
it
were transplanted to a severe, timberline habitat, the
crossbreeding would dilute the abundance of new genes
climate would stunt it. This stunted plant would shed
that are of most value in one particular environment; all
seeds and the new seedlings would also be stunted, even
if grown back at sea level. Characters acquired during the
members of a species would continue to be more or less
alike and not as well adapted to extremes within their
lifetime of one plant would be passed on to succeeding

generations.
range, as they could be if isolated. Mediocrity —
Lamarck himself did no experimentation to bolster his
population of generalists — would result. One large
population of generalists is not always as successful a
theory. Other botanists did, however. Bonnier in France
strategy for survival as several small populations of
made and sea-level
reciprocal transplants of alpine
specialists.
species. He grew a number of plants of each species to a
convenient size at Paris, then cloned them (split them into
How can reproductive isolation be established? Table

pieces) and planted them in plots in the Alps, the

10.6 lists the most important isolating mechanisms. The
Pyrenees, and Paris. They were not planted in gardens,
prezygotic mechanisms have to do with preventing pollen
but were put in among natural vegetation. The plots were
of one population from reaching the stigmas of another.
This can be accomplished by separating the populations in
sometimes fenced, sometimes not; watering and
fertilization were not practiced. The plots were periodically
space (isolated valleys or separating continents), time

visited for a number of years (1884-1920) and any

(different seasons of flowering), or biology (different

changes in the plants noted. He concluded that a number pollinating insects, which do not visit both flowers). On a

of lowland species were transformed into related alpine or

world scale, continental drift (the gradual spreading apart

subalpine species during the years of the study. Also

of continents over the past 200 million years) may have
around 1 920, the American ecologist Frederic Clements been the most important isolating mechanism. Postzygotic
established a similar series of "gardens" from Pike's Peak mechanisms prevent normal offspring from developing,

He too concluded that some

to the California shore.
even though pollination occurs. The hybrids may be weak
lowland species were transformed to related high-elevation
or sterile, for example, or the developing embryos can
abort.
species, and vice versa.
However, the most carefully documented and controlled Hybrids are often weak in nature, or at least they are
transplant experiments were last conducted, between 1920 less well-adapted to a given habitat than either parent. So
and 1940, by Clausen, Keck, and Hiesey in California, as the most common type of hybrid in nature is not a "pure"
mentioned earlier in this chapter. Their detailed study of hybrid, halfway between each parent. Instead, the
some 60 taxonomically diverse species showed not one successful hybrids are often the result of backcrossing
case of a lowland species becoming an alpine species, or between the original pure hybrid and either or both
the reverse. A lowland species at timberline might become parental types (Fig. 10.33). The few pure hybrids in this
dwarf or prostrate or stunted, but if its seeds were way produce an entire series of partial hybrids between

chapter 1Q |
Plant Taxonomy and Evolution

218
 .

original
short-necked
ancestor

longer neck

DARWIN'S GIRAFFE

natural selection
favors longer
necks: character
passed to next
generations

original group
variation in neck
length

necks longer but

still variable

Figure 10.31 Comparison of Lamarck's and Darwin's concepts

of evolution

themselves and original parental types. This is called 3. A species is a convenient unit of information. Species
introgressive hybridization, and it creates great diversity that show a close genetic relationship are grouped
(hybrid swarms) that may spell success for the group as a into a genus. Related genera are grouped into a
whole. family, families into orders, orders into classes, and
classes into divisions. For the sake of worldwide
uniformity, each species is described in Latin and its
Summary name is written in Greek or Latin as a binomial: the
genus (capitalized) followed by the specific epithet
1 As a floral survey of the world becomes more
(not usually capitalized).
complete, the goal of taxonomy becomes the 4. The book can be
21 plant divisions covered in this
construction of a phylogenetic classification scheme grouped which include
into several categories,
thatsummarizes the evolutionary history of our flora.
prokaryotes, eukaryotes, thallophytes, liverworts and
2. The history of taxonomy can be divided into a pre-
mosses, lower vascular plants, higher vascular plants,
Darwinian period and a post-Darwinian period.
seed plants, gymnosperms, and angiosperms. Two
Linneaus culminated the pre-Darwinian period (ca.
classes of angiosperms are the Monocotyledonae and
1753) and organized plant nomenclature. During the the Dicotyledonae.
post-Darwinian period there were several attempts to 5. Dried, pressed, and labeled specimens of all named
classify the plantkingdom on a natural (phylogenetic) species of plants are kept in herbaria. Living examples
basis; schemes by Bessey and Engler and Prantl are are grown in botanic gardens (arboretums).
examples. At present, we still lack a classification 6. Taxonomic research may take any of several
scheme accepted by all botanists because the directions, utilizing patterns of morphology, anatomy,
evolutionary history of plants is not completely known. biochemistry, and breeding behavior, any of which
 Table 10.6
Summary of Some Important Reproductive Isolating
Mechanisms

A. Prezygotic mechanisms: prevention of pollination or

fertilization

1. Separation in space: two populations

live far from

each other so cannot be transferred, or

that pollen
they live close but are separated by barriers
(mountains, bodies of water), or they occupy
different habitats (lowlands versus uplands), or
drifting continents move them apart.

2. Separation in time: two populations flower at different

times of the year, or if they flower at the same time,
the pollen is shed before the stigmas are receptive.

3. Biological because the pollinating insect

separation:
or animal is each population,
different for is it

unlikely that cross-pollination will occur, even the if

plants are close to each other; or the flowers may

have the same pollinator but open at different times
of the day (morning versus evening).

4. Physiological differences: the pollen of one is unable

togrow through the style of the other, or it grows
more slowly than pollen of the other.
Overlapping ranges,
but distinct species mechanisms: prevention
B. Postzygotic of normal offspring

Figure 10.32 Diagram showing the sequence of events which

development
leads to the formation of new races, subspecies, and species.
One species (green, at top) extends into new habitats, and 1. Incompatibility of zygotic or embryonic tissue with
populations at the extremes become modified into races or that of the mother plant produces seed abortion.
subspecies. If the races or subspecies become isolated, they may
evolve further and become incompatible with the original species; 2. Hybrids (F,) are completely inviable, or considerably
at this point they are recognized as distinct species. The weakened.
reproductive barrier will keep the species distinct, even if later
migration brings them back into contact (bottom). 3. Hybrids are vigorous but sterile.

4. F, is vigorous, but successive generations (F2 , etc.)

Parent Type A are weak or sterile.

may be used to show genetic relationships between

taxa.The numerical taxonomist utilizes information
from all fields and does not give extra weight to any
particular type of information. This is in contrast to the
work of other taxonomists who choose subjectively to
weight particular types of information. Biosystematists,
for example, emphasize the importance of breeding
behavior; they often deal with populations at the
species and subspecies levels.
First "Pure Plants can also be characterized or described by their
50% A
life cycle pattern. For example, herbaceous annuals
may exhibit an "r" type life cycle "strategy," while
long-lived woody perennials may exhibit a "K" type
"strategy."
Organic evolution is the gradual change that has taken
place in living organisms.
The fossil record reveals that prokaryotic life began
Hybrid H
more than three billion years ago, in the Proterozoic
Parent Type B
era. Eukaryotic life began more than one billion years
Figure 10.33 Introgressive hybridization. The degree of shading ago. The fossil record became richer, more varied,
(horizontal lines in the circles) shows the degree of similarity of
and more complex in the early part of the Paleozoic
hybrids to parental types A and B. Introgressive hybrids D, E, F,
G, and H would be more numerous in nature than "pure" hybrid era, as algal forms and the first terrestrial plants
C. evolved. The Cambrian period saw the evolution of

chapter 1Q |
Plant Taxonomy and Evo lution

220
 tree forms and seed plants. Later in the Paleozoic era, containing them.
Coal Age forests were dominated by lower vascular Mutations causing considerable phenotypic change
plants. Climatic and geological changes next brought are likely to kill or to greatly weaken the plant
dominance of the world's vegetation to the because they upset the delicate equilibrium existing
gymnosperms and then to the angiosperms in the between the plant and its environment.
Mesozoic era. The angiosperms have dominated most Hybrids differ from their parents because of the
terrestrial habitats, in all kinds of climatic extremes, resulting new combinations of genes. Hybrid
during the Cenozoic era (the last 65 million years). swarms, resulting from introgressive hybridization,
10. A modern theory of the mechanism of evolution can are common in some taxa.
be outlined as follows: If the variants produced by mutations or
a. Genes, contained in the chromosomes, are largely hybridization are better adapted to the environment
responsible for the development, structure, and than the parent plants, the parent plants may be
metabolism of plantsand animals. replaced by the new forms. Many complicated
b. The complement of genes does not remain factors are involved in this replacement.
absolutely constant. Mutations (changes in New species are formed by processes that divide a
chromosomes and genes) occur, which modify the population, ultimately leading to reproductive
structure and metabolism of the individuals isolation.

Summary

221
 1 1 CHAPTER

11 algae

are photosynthetic, nonvascular plants. They Grouping these species under the general
Algae
include some the most primitive plants on earth.
of
all

classification of algae is highly artificial because there

They may exist as single cells, as loosely does form and metabolism
exist a far greater diversity of
organized clumps ot cells (colonies), as flat, leaflike among them than among the bryophytes and the vascular
sheets, or as intricately branched and intertwined plants. It is difficult to find definitive traits that are common
filaments (Fig. 11.1). Only rarely, in the kelps, do algal to all algae yet absent from all other divisions. Possibly the
bodies become large and complex enough to differentiate only structure possessed by all higher plants that rarely
their tissues into organs that resemble roots, stems, and occurs in the algae is a protective jacket of sterile cells
leaves (Fig. 1 1 .2). Approximately 25,000 species of algae surrounding the developing gametes. In addition, most
have been identified, but many more, especially those in algalbodies are relatively undifferentiated. Their aquatic
the oceans, remain to be described for the first time. It is and semi-aquatic environments lack survival pressure for
estimated that there may be 10 times this number of the evolution of vascular and supporting tissue. All the
species living today. cells of most algae can carry on photosynthesis and

eye spot >

— nucleus

Chlamydomonas

chloroplast

•nucleus

cytoplasm

Volvox

Spirogyra Volvolcine Line

222
 Cou

Siphonous Line

Fucus furcata

Porphyro sp.

Figure Algal diversity. Unicellular forms include (A)

11.1
Chlamydomonas in the Chlorophyta. Colonial forms include (B)
Gonium and (C) Volvox, both in the Chlorophyta. Filamentous
forms can be separate and relatively simple in differentiation, as
(D) Spirogyra, or (E) Hydrodictyon, or nonseptate and
differentiated into prostrate, anchoring filaments and upright,
photosynthetic filaments as in (F) Caulerpa, all in the
Chlorophyta. Sheetlike forms include (G) Porphyra in the
Rhodophyta. More differentiated, larger forms with leaflike fronds
include seaweeds and kelps such as (H) Fucus and (I) Ptilota
Ptilota filiciana
filiciana.
 growing symbiotically with other plants or animals. Algae
occur in the most severe habitats on earth. Some species
grow on snow in perpetually freezing temperatures; others
thrive in hot springs at temperatures of 70°C or more; a
few live in exiremely saline bodies of water, such as the
Great Salt Lake in Utah; yet others survive the pressure
+
and low light intensity conditions of 100 m below the
surface of lakes or seas. They have even been detected 1

km from ground zero, 6 months after a 20 kiloton atomic

bomb tower test in Nevada, and they were considered to
be survivors of the explosion.
Probably the most commonly noticed natural urban
habitats for algae are the sides of glass fish tanks. They
blades may also generally be found around leaking faucets and in

garden or park pools that are not kept "pure" with

chemicals. The "bloom" occurring during the summer on
many lakes or the scum found on ponds is actually algae.
Microscopic forms occur in most natural waters, including
the top 75 m of the ocean. Here they constitute a primary
food source for marine animals of all types and account
for about 50% of the oxygen released into the atmosphere
through photosynthesis.

Economic Importance of Algae

Algae are important in two basic, but quite different ways.

They are important to the entire biosphere because of the
ecologically vital functions they perform. These functions
include the production of carbohydrates, which places the
algae at the base of food chains, the fixation of nitrogen,
and reef building. They are also economically important to
people because they serve as food, fodder, and fertilizer,

and also have many industrial and pharmaceutical uses.

Ecological Functions

Algae as Producers

In aquatic ecosystems, algae are the major producers.

Shallow bodies of water may depend on productivity by
higher plants such as rushes, water lilies, duckweed, or
the like, or they may be fueled by a constant supply of
decomposing plant material (detritus) washed down from
the surrounding land. About 70% of the globe is

Laminaria andersonii covered by deep water.

nonterrestrial and most of that is

Figure 11.2 Example of a kelp, Laminaria andersonii, in the Here algae are the only producers. They have been called
Phaeophyta. the grasses of the oceans, converting solar energy into
chemical energy that is passed up through the rich marine
food chain, and releasing oxygen as an important by-
obtain water and nutrients directly from their surroundings. product to both water and atmosphere.
Algae lack true roots, stems, and leaves (with vascular Algae (and other life) occupy several distinct habitats in
tissue); such a body is such a
called a thallus. Plants with aquatic ecosystems. One habitat, along rocky oceanic
body are called thallophytes, as described in Chapter 10. coasts, is the intertidal zone (Fig. 1 1 .3). This zone is

Algae can be found nearly everywhere. They float in air especially severe for the growth of algae, because the
or water. They are attached to tree trunks or branches, to plants are alternately exposed by low tide and inundated
the bottom of streams, to soil particles, or to rocky by high tide. When exposed, the plants are beset by
intertidal cliffs battered by surf. They are also found desiccation, high temperatures, high light intensity, and

chapter 11 |
Algae

224
increased salinity. When the tide comes in, all aquatic algae have adjusted their metabolism to the light

environmental factors change abruptly. In winter, the at this depth. Figure 1 1 .7 shows the action spectrum of
plants may be locked in ice. The neritic zone is below the photosynthesis for the green alga sea lettuce (Ulva

intertidal, but is still relatively shallow and near shore. taeniata), which grows high up in the intertidal zone;
Species of attached algae in the intertidal and neritic superimposed on the same graph is the action spectrum
zones have their own particular distribution: some occur at for the red alga Mynogramme spectabilis, which grows at

the upper end, others occur in the middle, and others in much greater depths. You can see that the peak has
the lower region (Fig. 1 1 .4). Their distribution probably shifted and condensed to center around the blue-green

reflects different degrees of adaptation to the stresses of region, 440 to 680 m/x. Special accessory pigments

exposure, for those species at the upper part of the (phycobilins) present in the red algae, trap the light of

zone are exposed more frequently and for longer

intertidal blue-green wavelengths, eventually transferring this energy
periods of time than species farther down. Experiments to chlorophyll.

tend to confirm this hypothesis: algae that grow in the

upper part of the intertidal zone are actually more resistant
Productivity and Pollution

to drying, and to fluctuations in temperature, light intensity,

The density of phytoplankton is not very high in the open
and salinity.
ocean, perhaps a few thousand cells per liter, but the
Gradually, with increasing depth, the quality and
oceanic expanse is enormous and this results in a high
quantity of penetrating lightbecomes less and less
annual rate of gross productivity: 32.6 x 10 16 kcal (3
favorable for photosynthesis. The compensation depth or
quintillion, 260 quadrillion). This is about three times the
compensation intensity is the point where positive annual productivity of all the world's grassland and
growth is no longer possible (Fig. 1 1 .5). In deep bodies of
pasture, and four times that of all cropland.
between the surfaces and the
water, the habitat
As great as this productivity is, however, it is limited by
compensation depth is dominated by floating algae, the many environmental factors such as light, temperature,
phytoplankton. and nutrients. these limiting factors are ameliorated,
If

Phytoplankters are usually microscopic, but they do productivity will increase manyfold. A natural process of
include the large, floating Sargassum seaweed, namesake increasing productivity and a changing environment takes
of the Sargasso Sea off Bermuda. The microscopic forms water because
place very slowly in most bodies of silt

are typically unicellular and maintain buoyancy by storing accumulates, making the depth shallower and the water
oil droplets or by developing fine projections that extend uniformly warmer, and vegetation encroaches along the
out from the cell wall (Fig. 1 1 .6). The average life span of edges and contributes increasing amounts of detritus. This
a given cell is probably measured in hours or days; if it process is called eutrophication, and it ordinarily takes
does not reproduce the protoplast dies and the cell wall centuries, millenia, or eons. Human activities, however,
remains will sink to the bottom. Most phytoplankters are mainly through the disposal of industrial, agricultural, and
members of the Bacillariophyta (diatoms) and Pyrrhophyta residential wastes, can telescope eutrophication into a
(dinoflagellates) divisions, but the Chlorophyta (greens), matter of decades. This is called cultural eutrophication.
Cyanophyta (blue-greens), and Euglenophyta are also As nutrients increase, algal and bacterial activity
represented. increase, resulting in more turbidity and lower light.

The quality of light changes with depth. Since red light Oxygen becomes limiting near the bottom; this limitation
is completely absorbed in the upper layers, a blue-green plus siltation prove fatal to fish eggs of some species.
twilight prevails further down. Experiments show that Algal species replace one another in an aquatic

Figure 1 1 .3 The intertidal zone, exposed at low tide, along the

central coast of California.

Economic Importance of Algae

225
 125

125

^
Laminaria
spp
J^>
Costaria
cosfala
jili
Zosfera
marina
Agardhiella
f
coulter/
^
Diafom
crust
*
Rhodymenia
perfuse
1
Nereocysf/s
&
Fauchea
spp.
turf forming
species

Figure 1 1 .4 Representative transects of benthic (bottom

attached) algae in the near-shore region of Puget Sound,
Washington. Dark substrate is rocky.

succession: generally greens are replaced by blue-greens

Full that may impart disagreeable odors or tastes to the water.
-12
10"" 10"'1 u"
in- in-8
10" 8 in - *
10"° in" 4
10"" 10"'
1
i i I 1 1 1 i i 1 I T 1
~sun As the base of the food chain changes, so does the entire
1

chain. After 100 years of accelerating industrial and

residentialdevelopment along its shores, Lake Erie had
400
compensation for '/ changed from a relatively clean lake with important fishing
i
phytoplankton • (
and recreational values to an aesthetic and economic loss.
800
Plankton and other algae increased six- or sevenfold in
some places, and the composition shifted toward the
noxious biue-greens. Oxygen content of bottom waters
1200
declined 67%. Commercial catches of trout, whitefish,
herring, sauger, walleye, and blue pike declined by 99.9%,
while such fish as carp, shad, alewives, and smelt
1600 - 1 limit for '/color vision in man
increased. Recently, the rate of pollution has declined and
water quality has improved.
2000
Red Tides and Blooms
2400 -
One of the results of cultural eutrophication is the episodic
_ \S limit for vision
(
for man appearance of red tides or blooms: the rapid growth of

hi iii phytoplankton populations until theybecome dense

2800 i i i i i i i i i

enough to color the water. This phenomenon can occur in

Figure 1 1 .5 The penetration
of sunlight into the clearest ocean
bodies of water that range from small ponds to lakes to
water. The compensation intensity for most phytoplankton is
1 /100th to 1 /1000th the intensity of full sunlight at the ocean large coastal regions. Marine occurrences are often
surface; compensation depth is about 550 ft (170 m). reported along the southwest coast of India, southwest

chapter 11 j
Algae

226
 1

Africa,southern California, Texas, Florida, Peru, and grown to such an extent in Japan that it is cultivated in
Japan. Factors associated with red tides or blooms are shallow, intertidal bays (Fig. The Polynesians in
1 1 .9).

long hours of sunlight, shallow, warm, offshore water, and Hawaii were known to have utilized and named at least 75
high levels of nitrogen and phosphorus. species of limu (algae) as food sources. Some rare
Red most often caused by dinoflagellate
tides are species were cultivated for the nobility in marine fish

species of Gymnodinium or Gonyaulax. Both species ponds. Dulce (the red seaweed Rhodymenia palmata) has
produce a water-soluble toxin of high potency that affects been known as a food for 12 centuries in the British Isles.

animal nervous systems. It is related to curare poison, Itwas the Irish who discovered that small quantities of
extracted from certain tropical flowering plants, and it is 10 Irish moss (the red alga Chondrus crispus), when boiled

times as effective as cyanide. Massive fish kills and the with milk, would produce a jelly dessert that the French
poisoning of shellfish result from red tides. A 1947 episode later called blanc mange. Brown algae off the coast of
off Florida killed an estimated 500 million fish. California (mainly Macrocystis pyrifera, Fig. 11.12) have
Other blooms are caused by the chrysophyte been harvested for their content of iodine, which is added
Prymnesium parvum and blue-green species of in trace amounts to the diet to prevent goiter, an

Microcystis, Anabaena, Nostoc, Aphanizomenon, enlargement of the thyroid gland in the neck.
Gloeotrichia, and Oscillatoha. The blooms are not It is not as food or medicine, however, that algae are

necessarily blue-green in color, however; a species of most important today. With some exception, they do not
Oscillatoria causes red blooms, which give the Red Sea its have much nutritive value in fact, their major—
name. Some of the plants above also produce toxins. A constituents are largely indigestible. Algae are used more
mere 72 g of Anabaena toxin would cause the death, as condiments, garnishes, or desserts than as staple
within two minutes, of an adult human. foods, much as we use lettuce, watercress, celery, or
whipped cream. Iodine now is regularly obtained from
Nitrogen Fixation other sources and added to table salt. Claims as to the

Some bacteria and about a fourth of all blue-green algal health-giving value of seaweed do not have much

species are able to assimilate or "fix" elementary nitrogen, foundation in fact.

N 2 The
. details of this metabolic pathway are still not
Fodder and Fertilizer
clear, but this overall reaction can be written:

Seaweeds not only contain such important trace elements

as iodine, but they contain large amounts of potash
N 2 + 3H 2 2NH,
(potassium), nitrogen, phosphorus, and other
characteristics of good fertilizer or cattle feed
The NH 3 can then be taken up by organisms, whereas supplements. In historic times, the North American Indians
the N2 cannot. All forms of biological nitrogen fixation, and the Scotch-Irish used Irish moss as a fertilizer to build
bacterial as well as algal, contribute on the average 1 up poor soils for such diverse crops as corn and potatoes.
kilograms of nitrogen per hectare (ha) per year to Seaweed has more recently been shown to compare
terrestrial surfaces around the world. In exceptionally favorably in nutrition to barnyard manure: it enhances
favorable areas, such as tropical grasslands, the amount germination, increases the uptake of nutrients in plants,

added may be as high as 227 kg/ha. Blue-green algae and seems to impart a degree of frost, pathogen, or insect
are cultivated in rice paddies and in some other kinds of resistance. Macrocystis (Fig. 10.12) continues to be
agriculture as a form of free fertilizer. Living cells secrete harvested off the California coast for use as a cattle feed
freeamino acids and peptides into the soil or water supplement.
medium, and dried algal crusts can be plowed into the
land. Specialized, thick-walled cells called heterocysts
Cell Walls and Cell Wall Extracts
conduct most of the fixation; the resulting organic nitrogen
Peculiar characteristics of the cell walls of diatoms,
then diffuses to other cells in the filament (Fig. 1 1 .8) or
browns, and reds have led to many recent industrial,
out to the environment.
pharmaceutical, and dietary advances. It is in these areas
that the algae have their greatest economic value. The
characteristics result in products such as diatomite, agar,
Economic Uses carrageenan, and algin, each of which we shall consider
individually.
Algae as Food or Medicine

Seaweed — marine algae of moderate size, which usually Diatomite. Diatomite, also called diatomaceous earth, is

grow in the intertidal or neritic zones — forms an important a chalky, sedimentary rock composed of the cell wall
part of the human diet and medicine chest in several parts remains of unicellular algae called diatoms. Diatoms, you
of the world. In the orient, seaweed harvesting has been recall, are important members of the phytoplankton (Fig.
known for 5000 years. Shen Nong, the legendary Chinese 1 1 .6). One of the richest deposits of diatomite is located
"father of medicine," prescribed seaweed for certain near Lompoc, California (Fig. 11.10). About 15 million
ailments in 3600 B.C. Some 3000 years later, Confucius years ago, that area was submerged beneath a warm,
praised its curative value. For centuries, the Japanese shallow sea. As diatoms flourished and died, their remains
have used algae as a healthful, tasty supplement to their accumulated in bottom sediments. These particular
rice diet. The demand for nori (the red alga Porphyra) has remains persisted intact because the wall is not composed

Economic Importance of Algae

227
 upper valve

Asteromphalus elegans

Navicula digiioradiaia

Figure 1 1 Examples of phytoplankters. A to D, diatoms

.6
Asteromphalus elegans, Navicula digitoradiata, Asterionella
formosa, and Biddulphia biddulphia. E and F, dinoflagellates
Ceratocorys aultii and Ceratium sp.

chapte r 11 |
A lgae
228
 spines ' »

girdle

girdle view Ceratocorys aultii

Aslerionella formosa

girdle section

plates

i&a valve
flagellae

Ceratium sp.

Biddulphia biddulphia

Economic Importance of Algae

229
 100

red algae growing

al some depth
\
\

//\
>
60
\

x / \\
/
/ \ \
\ /
\

\ / \
• /
/

/
s^ /
^^--^ \ \

jr ^ \
'green , 1

surface-growing
algae \\
\\
N
1
( I 1
i i i -\j

400 440 480 520 560 600 640 680 720 760
wavelength in m/i

Figure 1 1 .7 Action spectrum for photosynthesis in a green,

surface-growing alga (solid line) and in a red alga growing at
some depth (dashed line).

v * .
Q fessaF"
•
1

-
•

wP Figure 1 1 .8 Ultrastructure of the heterocyst of the blue-green

^ alga Anabaena. A, filaments of A. azollae, x880, showing

enlarged heterocysts in different stages of development. B,
electron micrograph of a heterocyst of A. cylindrica, x 22,000,
showing constrictions at heterocyst poles, three layers outside the
cell wall, and contorted thylakoids (photosynthetic membranes) in

| the cytoplasm. C, an enlargement of B, showing

microplasmodesmata connecting the cytoplasm of the heterocyst
with that of the adjoining vegetative cell.

chapteMi
J Algae

230
Figure 1 1 .9 Nori (Porphyra tenera, Rhodophyta) culture, Sendai
Prefecture, Honshu Island, Japan. A, distance view, showing the
extent of the hibi nets in January, about the time of harvest. 6,
closer view, showing the hibi nets about 30 cm above mean low
water in September, at the beginning of nori cultivation.

of carbohydrates such as cellulose; rather it is 95 percent improve flow, and increase stability. In those ways it is

silica. used in cement, stucco, plaster, grouting, dental

The walls of a diatom fit together like two halves of a impressions, paper, asphalt, paint, and pesticides.
Petri dish and the case is perforated with
glasslike Diatomite is also used as an abrasive.
hundreds of microscopic pores (Fig. 11.11). Each half is

called a frustule and the region of overlap is the girdle. Agar. Less than 100 years ago, a physician's wife, Frau
Under the electron microscope, it is possible to see that Fanny Eilshemius Hesse, discovered the use of agar as a
even the pores have pores. The perforations form bacterial culture medium. She passed the information on
exquisite, symmetrical designs, each design peculiar to a to her husband, and he to Robert Koch, and Koch to the
given species. Diatoms extract the opaline silica from the world via his important scientific writings in the late
surrounding water by a process that is still not nineteenth century.
understood. more recent geologic time, the Lompoc
In Agar (or agar-agar) is a polysaccharide, analogous to
area was above sea level and the diatom deposit
uplifted starch or cellulose but chemically quite different. It is

was revealed by erosion. Today, such companies as found in the walls of certain red algae. Mainly species of
General Refractories and Johns-Manville mine the Gelidium and Gracilaria seaweeds are commercially
diatomite for industrial and pharmaceutical use. harvested, divers descending 3 to 12 m in the warm near-
Diatomite makes a superior filter or clarifying material shore waters off Japan, China, California, Mexico, South
because the microscopic pores create a large surface America, Australia, and the southeastern United States.
area (0.5 lbs — —
230 g contain the area of a football field), In addition to its bacteriological use, agar is important in

and the rigid walls are incompressible. Diatomite is inert the bakery trade. When added to icing, it retards drying in

and can be added to many materials to provide bulk, the open air or prevents running in cellophane packages.

Economic Importance of Algae

231
 10 Aerial view of diatomite deposits near Lompoc,
California.

Because it is virtually indigestible, agar is also used wall by weight. Water is strongly adsorbed to algin,
medically as a bulk-type laxative. resulting in a thick solution. For example, one tablespoon
of algin added to a quart (one liter) of pure water will
Carrageenan. This is another polysaccharide from the increase the viscosity to approximately that of honey. In
walls of red algae. It is mainly harvested from Irish moss nature, algin may be valuable to intertidal browns during
and the substance takes its name from the town of exposure by enabling water to be retained in and on the
Carragheen, County Cork, along the south shore of plant. Commercially, however, it has many uses (Table
Ireland, where its properties were first discovered. 11.1).
Carrageenan reacts with the proteins in milk to make a Hundreds of species possess algin, but only a few are
stable,creamy, thick solution or gel. Consequently, it is commercially harvested: Macrocystis pyrifera along the
used in ice cream, whipped cream, fruit syrups, chocolate California coast, and species of Laminaria, Fucus, and
milk, custard, evaporated and even macaroni.
milk, bread, Ascophyllum off England. Macrocystis is the largest kelp
It is added to dietetic, low-calorie foods to bring back the (a general term for large, marine, brown algae) known,
appropriate mouth "feel" of nondietetic foods. and it any multicellular plant
has the fastest growth rate of
Carrageenan is also used in toothpaste, pharmaceutical in the world. From a single-celled zygote, grows into a it

jellies, lotions of many sorts and as a whiskey chaser for a young plant rooted on a rocky bottom 6 to 30 m below the
cold "cure." Irish moss is commercially harvested in this surface, and then into a 60 m long mature giant much of —
country in Maine, principally by two companies: Kraft its shoot floating on the surface —
in the course of a single
Foods and Marine Colloids. growing season (Fig. 11.12). At this stage is it

differentiated into a rootlike region (the holdfast or

Aigin. Since this long-chain polymer is made up of hapteron) that anchors the plant to bottom substrate, a
repeating organic acid units, it is also called alginic acid. It
stemlike region (the stipe), and numerous leaflike fronds
is the principal wall component of brown algae, that arise all along the stipe and possess gas-filled
constituting up to 40% of the middle lamella and primary bladders at their base that increase buoyancy. Each stipe,

c hapter 1 1 |
A lgae
232
 lower valve (frustule) — open chamber
'

Figure 11.11 Cell wall details of the triangular diatom

Triceratium favus.

E cono mic Importance of Algae

233
Table 11.1 Differences in cell wall construction, number and
Some of the Products in Which Algln Is Used placement of flagellae, and type of chlorophyll are quite

fundamental and significant We shall briefly survey seven

Food Products Industrial of the divisions.

Bakery icings and Water base paints

meringues Wall joint cements
Salad dressings Welding rod coatings Cyanophyta: the Blue-Green Algae
French dressings Textile print paste
Dietetic dressings Textile sizing
Unique among all the algal divisions, the blue-greens are
prokaryotic and are thus most closely related to bacteria.
Pickle relish Latex creaming and
Meat sauces and pepper thickening In fact,some taxonomists place them in the bacteria,
sauces Adhesives calling them the Cyanobacteria. They lack an organized
Paper coatings nucleus, membrane-bound organelles, and endoplasmic
Orange concentrates
reticulum. They do, however, contain gas vacuoles,
Fruit drinks Corrugated paper
photosynthetic membranes, and storage granules of
Dietetic beverages and Paper sizing
material that has been likened to lipid, glycogen, and
drinks Paper cartons for foods,
protein (Fig. 1 1.13). The wall resembles that of certain
Beer soaps, and detergents
bacteria, being composed of muramic acid, glucosamine,
Ice cream and related Food wrappers
Waxed cardboard cartons glutamic acid, glucose, xylose, mannose, and several
frozen desserts
Boiler compounds
other mureins and sugars. Cellulose is not present.
Egg nog
Creamed cottage cheese Can sealing compounds Virtually all free-living blue-green algae are surrounded by
Cream cheese Battery plate separators a mucilaginous sheath (Fig. 1 1 .14).

Morphological diversity is limited in the Cyanophyta to

Pasteurized cheese Mold release coatings
spreads Wax cleaner polishes
and simple filamentous forms (Fig.
unicellular, colonial,

Canned buttered vegetables Ceramic glaze 11.14). There are no flagellated cells, and sexual

Canned chow mein reproduction has never been documented. Instead,

Sugar beet clarification
Canned meat stews Finger paints asexual reproduction by fragmentation of filaments, the

Cake mixes formation of resting cells, or simple fission predominates.

Pharmaceutical Movement is passive except in some filamentous forms,
Puddings, pie and cake
Antibiotic tablets and which and glide by an unexplained mechanism.
rotate
fillings
suspensions Some forms, such as Nostoc or Anabaena, possess
Fountain syrups
Dental impression scattered specialized cells called heterocysts, which are
Buttered pancake syrups
compounds the site of nitrogen fixation, as mentioned earlier (Fig.
Berry syrups
Toothpaste 11.7).
Chip dips and mixes
Surgical jellies
Instant dessert mixes
Mineral oil emulsions
Chocolate drink
Delicatessen salads
Medicated rubbing Rhodophyta: the Red Algae
ointments
Candy
Tranquihzing tablets Cyanophyta and Rhodophyta have a few characteristics in
Dessert and salad gels
Hand lotion common. Both possess only chlorophyll a and identical
Breading batters
Facial beauty masque accessory pigments called phycobilins, neither possesses
flagellated cells, and both form a nonstarch storage
product related to amylopectin. The differences, however,
are much more striking. The Rhodophyta are eukaryotic,

growing as much as 30 cm a day, may live only a year or possess cellulose walls, and exhibit some of the most
less, even though it has the potential for continuous tip complex life cycles, incorporating both sexual and asexual
growth. Other stipes continue to arise from the holdfast, reproduction, of any group of plants. Their range of
however, and an average plant might live 4 to 10 years. morphological diversity is also greater than that of the
Macrocystis has been called the sequoia of the sea; blue-greens, including sheetlike forms and complex, finely
indeed, grows in such towering, dense stands or
it
branched filamentous forms with holdfast and stipe (Fig.

1 1 .1). In some species, pits connect the cytoplasm of

"beds," that a scuba diver may feel he is swimming
through a forest. It is the fast growth of Macrocystis, adjacent cells.

combined with its dense beds, that make

occurrence in it
The reds are almost exclusively marine, and are most

commercially harvestable. Barges visit the beds every six abundant in warm water, often extending to some depth.
weeks, cutting the tops one meter below the surface, then Again like the Cyanophyta, some forms are lime-encrusted
taking them to shore for processing. and participate in tropical reef building.

Algal Classification Euglenophyta

This division is a small one, consisting mainly of
As shown in Table 1 1 .2, there are many algal divisions unicellular, motile species (Fig. 11.1). Members of this
that differ in biochemistry, cytology, and habitat. division have several protozoan features, such as the lack

chapter 1 1 |
Algae

234
 Macrocysfis pyrifera

Figure 11.12 The kelp Macrocystis pyrifera, including details of

the blades (C and D) and holdfast (B).

Algal Classification

235
Table 1 1 .2

Major Characteristics of 10 Algal Divisions 9

Chlorophylls Number
and Storage of
Division Accessory Cell Wall Product Flagelia Habitats Species Notes
Pigments

Cyanophyta a + Mureins and Cyanophyte none All,but ofen 1500 Asexual reproduction
(blue-greens) phycobilins sugars + starch (
= polluted only; prokaryotic;
(phycocyanin sheath amylopectin) water; 75% prominent in blooms;
and marine some fix N
phycoerythrin)
Rhodophyta a (possibly + Cellulose + Floridean starch none 98% 4000 Includes some
(reds) d) + sometimes (= amylopectin) marine; seaweeds; complex
phycobilins agar or oftenwarm, life cycles
carrageenan deep water
Chrysophyta a + c + Cellulose Fats, oils, 2, unequal, Mainly 300
(golden algae, fucoxanthin chrysolaminarin anterior fresh water
class
Chrysophyceae
only)

Xanthophyta a ( + c in Cellulose or Oil or 2, various Mainly 400 Sometimes placed in

(yellow-greens) some) none; chrysolaminarin fresh water Chrysophyta
sometimes
with Si and
diatomlike
Euglenophyta a + b None Paramylon 1-3, equal, Mainly 450 Many animal-like
(paramylum) anterior polluted properties
fresh water

Chlorophyta a + b Cellulose Starch 2, equal, Widest 7000 Possible precursor

(greens) anterior distribution of higher plants
of any
division

Charophyta a + b Cellulose Starch 2, equal, Often 250 Differentiated into

(stoneworts) anterior bottom nodes and
dwellers in internodes;
fresh water gametangia with
sterile jacket,
sometimes put in
Chlorophyta
Bacillariophyta a + c + Silica + Fats, oils, generally Mainly 8000 + Prominent in
(diatoms) fucoxanthin pectin chrysolaminarin none aquatic phytoplankton;
sometimes put in
Chrysophyta
Pyrrhophyta a + c + None or Starch(?) 2, unequal, 93% marine 1000 Prominent in
(dinoflagellates, peridinin plates of lateral phytoplankton
class cellulose
Dinophyceae
only)

Phaeophyta a + c + Cellulose + Laminarin, 2, unequal, 99.7% 1500 No unicellular forms;

(browns) fucoxanthin algin mannitol lateral marine; includes the kelps
often, (another name for
shallow brown seaweeds)
water

a A Few Small Groups Have Not Been Included.

chapter 1 1 |
Algae

236
 degree of differentiation. This type of habit is called
heterotrichy. Other forms as shown in Fig. 11.1, are
unicellular, filamentous (with or without cross-walls),
colonial, and sheetlike.

Bacillariophyta: the Diatoms

We have already discussed these organisms earlier in the

chapter as one of the major components of phytoplankton.

They exist as single cells (Figs. 1 1 .6, 11.11 )
— sometimes
stalked and sedentary, sometimes floating free — or as
filaments. As seen in a face view, their silica walls are
two main orders,
either circular or elongate, giving rise to
Centrales and Pennales. Some diatoms are capable of a
gliding motion even though cilia and flagella are absent.
Diatoms contain chlorophyll a and c, store carbohydrate
as oil, and their unusual silicate wall is embedded in a
pectin matrix. They are mainly aquatic, but are also found
in soil.

Pyrrhophyta: the Dinoflagellates

Together with the diatoms, this group dominates the

phytoplankton. Most of its species are unicellular,
flagellated, and marine (Fig. 11.6), but a few are colonial
or filamentous. These organisms in great density cause
"red tides." The cells are either naked, that is, with only a
Figure 11.13 Ultrastructure of the blue-green alga Anabaena
flos-aquae, x 73,500, showing microfibrils in the outersheath, a thickened membrane around them as in Euglenophyta, or
four-layered cell wall, gas vesicles in longitudinal section (rods) they are enclosed by a unique cellulose wall that
and cross section (circles), photosynthetic membranes, dark-
staining lipid bodies, and many granular ribosomes.
resembles patches or armor plating stuck together. Usually
two flagella are present, both emerging from the same
pore, but otherwise different. One flagellum is flat and
ribbonlike and encircles the cell in a transverse groove; it
of a cell wall (although the outer cytoplasm may be
is responsible for rotation and some forward movement. A
modified for rigidity), rapid movement, and the ability to
second flagellum trails behind and provides forward
produce a nonstarch food reserve called paramylon. Some
movement while at the same time acting as a rudder.
Euglenophyta, in addition, lack chloroplasts and either
and c and a brown
Dinoflagellates contain chlorophyll a
engulf or absorb food. Exposure to ultraviolet radiation can
pigment, peridinin, which gives the groupits green-brown
halt chloroplast division but not cell division; the result is
or orange-brown color. True starch appears to be stored.
that within a few generations some progeny cells will lack
Some forms are luminescent and contribute to the glow of
chloroplasts. These cells will continue to live as
water when it is disturbed at night, as in the wake of a
heterotrophs and produce progeny just like themselves,
ship.
revealing what a minor genetic difference there sometimes
is between "plants" and "animals."

Phaeophyta: the Brown Algae

Chlorophyta: the Green Algae
The browns include filamentous forms, sheetlike forms,
The green algae are predominantly fresh-water forms, but and large kelps <th more complex differentiation in
•

they also exist in salt water, on snow, in hot springs, on anatomy and morphology than any other algae (Figs. 11.1,
on branches, and on the leaves of terrestrial plants.
soil, 1 1 .2, 11.12, 11 .15). Unicellular forms are unknown.

They form the second-largest division within the algae. Chlorophyll a and c are present, plus the accessory
(Diatoms are the largest.) pigment fucoxanthin, which gives these plants their
group shares several important traits
Cytologically, this characteristic brownish color (though color can range
with higher plants: chlorophyll a and b, similar accessory from olive-brown to golden-brown to practically black).
pigments, starch as a storage product, and cellulose cell Carbohydrate is stored as mannitol or laminarin, not as
walls. For this reason, some scientists hypothesize that starch. The cell walls are composed of cellulose and often
ancient Chlorophyta served as the ancestors of all higher a great deal of algin as well.
plants. The most complex greens have well-developed Kelps such as Macrocystis (Fig. 11.12) are complex
prostrate portions and upright portions, indicating a high anatomically as well as morphologically. If the stipe is

Algal Classification

237
19

** ®* -*
ts
•
A

Figure 11.14
.%
Diversity of form in the Cyanophyta. A and B,
loosely organized colonies of Gloeocystis sp. and Fisherella
musicela, both x800; note the pronounced gelatinous sheath
around Gloeocystis. C, a colony of the filamentous Nostoc sp., \
x 880, with enlarged heterocysts visible, all embedded in a
gelatinous matrix. D, filaments of Oscillatoria sp. (large cells) and
Anabaena sp. (small cells, with terminal and intercalary
heterocysts), x880. E, the spiral filamentous Arthrospira sp.; two
filaments are intertwined along one portion of the figure, x 880.

*« j?

"
x
'I it/
% < r

chapter 1 1 |
Algae

238
 sectioned and examined under the microscope, several
regions are apparent (Fig. 11.16). Cells in the outermost
layer not only are protective, but they remain meristematic
and contain chloroplasts as well. To distinguish this
diverse tissue trom epidermis, it is given the name
meristoderm.
A broad cortical region is composed ot parenchyma-like
cells. Mucilage-secreting cells torm definite canals through
the cortex. Loosely packed filaments of cells fill the central
region, the medulla. Some inner cortex cells next to the
medulla appear to function as sieve-tube members: they
have sieve plates, form callose, adjoin one another to form
continuous tubes, and are known to translocate mannitol
by a mechanism resembling that found in vascular plants
(Fig. 1 1.17). However, these cells contain no tissue that
resembles xylem. The value of a photosynthate-conducting
system in these large plants is easy to perceive. The mass
of floating fronds considerably reduces the penetration of
light into the water below; consequently the lower part of

the stipe and the holdfast may be below the compensation

depth. They must be nourished by food translocated from
above.
'
Unlike many of the other algae, the dominant generation
of the kelps is diploid. The dominant generation is also
larger and more complex than any other alga.
>

\ Reproduction

Knowledge of reproductive processes in the algae is

incomplete and largely confined to the Rhodophyta,

Phaeophyta, and Chlorophyta. In some other divisions,
detailed knowledge is known only for a very few species,
genera, or families. There is no typical life cycle for the
algae as a group.

Asexual Reproduction: Vegetative

Vegetative reproduction is widespread in many forms and
is probably used in all divisions. Indeed, in those algal
groups where sexual reproduction is unknown or rare,
asexual reproduction may be the only form of
reproduction.
The longitudinal splitting of a biflagellate motile Euglena
into two, still motile, but uniflagellate daughter cells is
simple vegetative reproduction. The diatoms may divide
vegetatively for up to five years. Recall that the cells have
two sections that fit into each other like the top and
bottom of a Petri dish. After division, a new wall forms
within the old wall (Fig. 11.18). This means that the new
cell receiving the lower and smaller wall will be smaller
than the parent cell. Eventually, a minimum size is reached
and division stops. Full size is regained at the time of
sexual reproduction (Fig. 11.18).
Vegetative reproduction in filamentous forms occurs by
a simple fragmentation of the filament. In some species,
specialized cells seem
be associated with the breaking
to
of the thallus. In species of the Cyanophyta, notably
Oscillatoria (Fig. 1 1.15), the point at which fragmentation
occurs is marked by a dead cell. Separation at these dead

Reproduction

239
Figure 11.15 Aan example of a kelp, or brown seaweed:
Postelsia palmaeformis (sea palm), showing organs that resemble
roots, stems, and leaves. B, young plant.

chapter 11 |
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240
 transition filamentous
zone medula mucilage layer

Figure 11.16 Anatomical details of the stipe of Postelsia. A,

diagrammatic cross section, showing the major regions and the
location of detail drawings B and C. B, details of the meristoderm
region, cortex with mucilage canals, and inner cortex with sieve-
tubelike cells in a transition region near the central medulla. C,
end wall of a sieve-tubelike cell, showing pores
detail of the
through^ which mannitol and other substances can flow.

Reproduction

241
 Most zoospores are pear-shaped or spherical.
Depending on the species, they have two, four, or many
flagella. After liberation from the sporangium, zoospores

are actively motile for as short a time as a few minutes or

for as long as three days. Their movement and periods of

activity are frequently affected by light. After their period of

activity they settle to the bottom of a pond or culture tank,
lose their flagella, and secrete a cell wall, and by cell

division begin to develop a new thallus.

Sexual Reproduction
Sexual reproduction is responsible for variation within a
population. Since sexual reproduction is frequently
associated with resistant cells that carry the plants over
seasons unfavorable to growth, all of the new plants in a
given population formed by sexual reproduction will have
different genotypes when growth is resumed; those plants
best adapted to the new growing conditions will become
established.
The cells that fuse in sexual reproduction are known as
gametes. Gametes have the haploid or n number of
chromosomes. The single cell resulting from the fusion is
a zygote, and has a diploid or 2n number of
chromosomes. By cell division, the originally one-celled
zygote can produce a diploid plant (the sporophyte
generation). The number of chromosomes is returned to
the haploid, or n, number by meiosis. Reproductive cells
Figure 11.17 Photograph of sieve-tubelike cells, with swollen
junctures, from the kelp Laminaria groenlandica, about x 900. resulting from meiosis, with the n number of
chromosomes, may be called meiospores. Meiosis occurs
at different places in the sexual cycle, and the cells in

which occurs have been called meiocytes. Meiospores

it

can be motile or nonmotile. If they lodge in an appropriate

cells results in short filaments that have a gliding motion.
environment, they germinate and by cell division produce
This enables them to change their position in the
a haploid plant (the gametophyte generation). The
mucilaginous mass which the filaments grow, and
in
gametophyte generation produces the sexual cells called
gradually to enlarge the mass. Other species may modify
gametes.
somatic cells by adding a thick wall, and these cells may
Gametes are produced in cells called gametangia. A
function as resting spores, living through a hostile period
gametangium, in the most unspecialized case, may be
of time in dormancy when all other somatic cells die.
indistinguishable from an ordinary vegetative cell, except
that several additional mitoses occur within it, giving rise
Asexual Reproduction: Mitospores to 16 to 32 cells. In the unicellular, motile green alga
Chlamydomonas, these cells strongly resemble the
Asexual spores are always preceded by mitosis and may vegetative cells, but they may be smaller (Figs. 11.19,
be called mitospores to distinguish them from spores 1 1 .21). In the simple filamentous species of the green alga

preceded by meiosis, called meiospores. The latter Ulothrix (Figs. 1 1 .20, 1 1 .21), an ordinary vegetative cell

constitute a stage in sexual reproduction and will be gives rise to 8 to 64 flagellated cells. When only eight
considered later. Since only mitotic divisions are involved,' motile cells are formed, they germinate directly, acting
the genotypes of all asexual spores arising from one and simply as zoospores (mitospores), and the cell in which
the same parent are identical with each other and with the they are formed is called a sporangium. However, when a
parent plant. They will produce a population of genetically greaternumber of cells is formed, the cells are smaller
identical individuals. When a population has the genotype and they behave as gametes that fuse in pairs. When
best fitted to a given growing condition, or locality, it will gametes resemble each other, they are called
multiply by asexual reproduction and populate that locality. isogametes, and the species that produces them is
Mitospores may be either motile or nonmotile. Motile isogamous (Fig. 11.21). Thus, Chlamydomonas and
spores are called zoospores and nonmotile spores are Ulothrix are isogamous.
aplanospores (or, in Polysiphonia,
frequently called In some species, the gametes differ in size, one being
carpospores). Mitospores are produced in specialized slightly smaller than the other. Gametes with only a slight
cells called sporangia. In many algae, the sporangia show differencein size are said to be anisogametes

little, if any, difference appearance from ordinary

in (heterogametes), and species expressing this state are
vegetative cells. The number of spores produced depends anisogamous species or exhibit anisogamy
on the species, but it is typically 16 to 64. (heterogamy) (Fig. 11.21).

chapter 1 1 |
Algae

242
 __
1—3
o
X« m

O iii.i.i.i.
'
•" "v

i . tV
r >.i.i.i.i-i.i.i.i-i-i.i.cy

r
\

n
A
mitosis

zygote

isogametes

Figure 11.18 A gametic life cycle as represented by a diatom.

A, the progeny cells continually become smaller if, after cell
division, the new cell forms within the silica shell of the old cell.
B, meiosis occurs; three of the nuclei with the n number of
chromosomes degenerate and the single remaining haploid cell
becomes a gamete. C, fusion of the two gametes to form a
zygote. -

In other species, the gametes not only differ in size but The sperms are liberated from the antheridium and swim
in degree of motility. One type of gametangium produces toward the oogonium. One sperm enters the oogonium,
many gametes called sperm. A sperm-
small, motile either through the funnel end or by a breaking down of
producing gametangium is known as an antheridium. the oogonial cell wall. Union of egg and sperm now
Another vegetative cell may become quite enlarged, ensues; this process is known as fertilization. When
sometimes becoming flask-shaped. The protoplast retracts gametes differ in size and activity as they do in Laminaria,
slightly from the cell wall. has become an egg cell, and
It Fucus, and Polysiphonia, the plants exhibit oogamy (Fig.
the cell in which it is produced is known as an oogonium. 11.21).

Reproduction

243
 sporangium

conjugation

Figure 11.19 A zygotic life cycle is represented by the green

alga Chlamydomonas. Meiosis occurs during the germination of
the zygote; thus the zygote is the only diploid (sporophytic) cell.

How does a sperm cell find an egg cell? It could happen In many of the more primitive algae, meiosis

by chance, much as wind-blown pollen lands on stigmas accompanies the germination of the zygote. In this case,
of flowers of its own species. However, there is the mature plants are haploid and the diploid stage is
considerable evidence that the female gametangia or limited to the zygote. This is a zygotic life cycle. It occurs
gametes in plants produce chemical agents that attract the in many Chlorophyta and in a few primitive Rhodophyta.
sperms to the close proximity of the egg cells. For In most plants, meiosis and fertilization take place in

instance, a solution prepared from the mature female distinct generations. One generation is haploid and
filaments of the alga Oedogonium may be drawn up into a produces gametes, the other is diploid and produces
capillary tube. When the tube is placed in a suspension of meiospores. This is a sporic life cycle. The plants of the
sperms, the sperms will collect at the tip of the capillary sporophyte and gametophyte generations may be identical
tube and some of them will eventually enter the tube. This in appearance; if so, there is an alternation of

attraction seems to be species-specific for Oedogonium. isomorphic generations, as occurs in some Chlorophyta,
Once the gametes fuse, a diploid zygote cell is formed. primitive Phaeophyta, and all higher Rhodophyta. In
Further cell division will produce the sporophyte alternation of heteromorphic generations, the haploid
generation. plants of the gametophyte generation have different forms
from the diploid plants of the sporophyte generation. This
Alternation of Generations
occurs in a few Chlorophyta and in all of the more
Fertilization constitutes one of the two critical steps of a advanced Phaeophyta, and in the rest of the plant

complete sexual life cycle; meiosis is the second critical

kingdom beyond the algae.
step. We may distinguish three types of life cycles,
depending on the relation of meiosis to fertilization. Gametic Life Cycle. The diatoms are unicellular forms
In most animals, meiosis results directly in gamete that may divide by cell division for periods as long as five
formation. In a sense, the entire gametophyte is composed years. With each division, one of the cells is smaller than
of gametes. In this case, the mature individuals are diploid the parent cell (Fig. 11.18). At the end of a period of
and the haploid stage is limited to the gametes. This is a occurs and four gametes are
mitotic divisions, meiosis
gametic life cycle. It occurs in the diatoms and in the produced, not which may function (Fig. 11.18).
all of
order Codiales of the Chlorophyta. Depending on the species, these most generally are

chapter 1 1 |
Algae

244
 escaping isogamete

ye spot

zygote

resting zygospore zoospores

mitospores)

germinating zygospore
meiosis)

meiospores

germinating zoospore

Figure 1 1 .20 A zygotic life cycle as represented by the

filamentous green alga Ulothrix. As with Chlamydomonas, only
the zygote is diploid.

/ anisogamy

heterogametes j^^\

Figure 1 1 .21 Differences in gametes. A, in isogamy the gametes

are identical; S, in anisogamy the gametes are slightly different in
size or activity; C, in oogamy, the gametes are very different in
size and activity and are called sperms and eggs.

Reproduction

245
isogametes, but they may be anisogametes or even eggs stimulated to grow. It enlarges and gives rise to a
and sperm. After fusion, the zygote increases greatly in surrounding protective case formed of haploid filaments
size and, again depending on the species, may begin a (cystocarp, Fig. 1 1 .235). Thus, the first sporophyte
rest period. The germination of the zygote is by mitotic cell generation (carposporophyte) in Polysiphonia and many
division. Note that in diatoms, the products of meiosis are other Rhodophyta is composed of short diploid filaments

gametes, not meiospores. The gametes themselves are the that produce diploid spores. This sporophyte is protected
only haploid cells in the life cycle. by an envelope of gametophyte cells. When discharged,
the diploid carpospores give rise to a second sporophyte
(tetrasporophyte), identical in appearance to the two
Zygotic Life Cycle. Chlamydomonas and Ulothrix are
gametophyte plants. Meiosis occurs in sporangia in the
examples of a zygotic life cycle (Figs. 11.19, 11 .20). In
sporophyte plant in cells formed between rows of axial
each of them, two haploid gametes fuse to produce a
and pericentral cells (Fig. 1 1 23G). Each sporangium
diploid zygote. Meiosis occurs in the germination of the
produces four meiospores (tetraspores) that on
zygote, which is the only diploid cell in the life cycle.
germination give rise to male or female gametophyte
plants similar in appearance to the sporophyte plants.
Sporic Life Cycle: Alternation of Isomorphic
Generations. Ectocarpus (Phaeophyta) and
Polysiphonia (Rhodophyta) will serve to illustrate Sporic Life Cycle: Alternation of Heteromorphic
alternation of isomorphic generations. In Ectocarpus (Fig. Generations
11.22), specialized reproductive cells are formed. The
Kelp. Alternation of heteromorphic generations occurs in
gametangia are grouped together, forming elongated,
both the Chlorophyta and Phaeophyta but is most highly
slightly curved structures. Each cell in this structure is a
developed in the Phaeophyta, particularly in the kelps of
gametangium that has resulted from mitotic divisions and
the order Laminariales. The sporophyte generation of the
will produce one motile gamete with the haploid
kelps is large and well known; its vegetative characteristics
chromosome number. They conjugate to form the usual
diploid zygote. The zygote, upon germination, will grow
have been discussed in some detail earlier in the chapter.

into a diploid sporophyte (Fig. 1 1 .22C) whose thallus is

The sporangia arise from meristematic cells, or the
meristoderm, that form the outermost layer of cells on the
identical in appearance to that of the gametophyte thallus.
fronds (Fig. 1 1 .246). The sporangia usually arise in
Two different kinds of sporangia are formed on this
groups (sori) and are accompanied by an elongation of
sporophyte plant. One of them resembles exactly the
adjacent cells that serve to protect the developing
group of gametangia found on the gametophyte plant. Like
sporangia (Fig. 11.24). Depending on the species, each
the gametangia, all sporangium cells have been formed
sporangium produces 8 to 64 motile meiospores.
through mitotic divisions, and each cell will produce a
motile diploid zoospore that can germinate directly into a
The meiospores germinate into male or female
gametophyte plants, both of which consist of a small
new diploid plant (Fig. 1 1 .226). The other type of
sporangium found on the sporophyte plant is spherical; branched filament (Figs. 1 1 24C,D). The antheridia are
after meiosis, will produce numerous motile zoospores or
it
produced in large numbers, either singly or in groups, at
the ends of short branches. There appear to be no
meiospores that have the haploid chromosome number.
specialized oogonia, as eggs may be produced in any of
These meiospores grow into new gametophyte plants (Fig.
the cells of the female gametophyte (Fig. 1 1 .246). The
1 22A). Haploid gametes, haploid meiospores, and diploid
zoospores are all identical in appearance. eggs are usually discharged from the oogonium before
fertilization, but remain attached to it for some time. The
egg is thus fertilized outside of the oogonium in open sea
Polysiphonia. In the Rhodophyta, an alternation of water.
isomorphic generations is complicated by a second,
sporophyte phase. The delicate feathery thalli
distinctive of
male and female gametophytes and one sporophyte of Fucus. Alternation of heteromorphic generations has
Polysiphonia are identical in appearance and growth (Fig. reached an extreme in Fucus (Fig. 1 1 .25). The diploid
1 1 .23). No motile cells are produced. The nonmotile generation is the conspicuous one (Fig. 11.1), and the
sperm (spermatia) are produced in great profusion near haploid phase has been reduced to a few haploid cells.
the tips of male gametophyte filaments (Fig. 1 1 23C). The These do not develop into free-living vegetative
oogonia (carpogonia) form near the tips of the female gametophytes but become transformed into sperm and
gametophyte. The oogonia are flask-shaped cells with eggs and fuse to produce a new diploid generation (Fig.
long slender necks (trichogynes) that protrude from a 11.25).
protecting envelope formed from the cells at the bases of The ends of the diploid strap-shaped thallus of Fucus
the oogonia (Fig. 1 1 .23D). Spermatia are carried by water are swollen and notched. They bear small raised areas
currents to make contact with the necks of the oogonia. (Fig. 1 1 .25/4). A section through these areas shows them
Nuclei enter the necks and pass downward to unite with to be small (conceptacles) that open by small
cavities
the female nuclei. conical pores to the sea water (Fig. 1 1 .256). Unilocular
Fertilization stimulates the development of an array of sporangia arise in these cavities; and they are of two
short diploid filaments (the carposporophyte), which will types. Microsporangia are formed profusely at the ends of

eventually produce diploid mitospores (carpospores). The short branching filaments, and megasporangia are large
original protective envelope of haploid filaments is also spherical cells separated from the wall of the cavity by a

chapter 11 |
Algae

246
 a —

S^tJ diploid filament

gametophytes sporophyte

Figure 1 1 .22 Sporic life cycle, alternation of isomorphic

generations differing only in having diploid and haploid
chromosome numbers, as represented by the filamentous brown
alga Ectocarpus.

single cell (Fig. 1 1 .25C). Sterile hairs are numerous, cells will differentiate into a sperm and be discharged into

surround the sporangia and may protrude outward the cavity.

through the pore. In the Pacific Coast species of Fucus, The megasporangium also contains a single large cell

both types of sporangia occur in the same cavity, but in

a megasporocyte. Meiosis produces four meiospores, and

the Atlantic Coast species they occur on separate plants.

each meiospore undergoes a mitosis without cell division;
the result is four binucleate cells, each one of which is a
The microsporangium contains a single large cell — megagametophyte. Ultimately, cell walls do form and an
microsporocyte (Fig. 1 1 .25F). Meiosis takes place, and
additional mitosis occurs, resulting in eight cells that
four haploid cells are produced. Each can be eggs (Fig. 1 1 .25E.O). The
further differentiate into eight
simultaneously considered a meiospore and a unfertilizedeggs are discharged from the old
microgametophyte. Each microgametophyte will rapidly megasporangium walls and they pass, with the sperms,
divide by four mitoses, giving rise to 16 cells (a total of 64 outside the cavity into the open sea where fertilization
cells within the old sporangium walls). Each of the 16 (64) takes place.

Reproduction

247
 gametophyte

Figure 1 1 .23 Sporic life cycle, alternation of isomorphic

generations, as represented by the red alga Polysiphonia. A, male
gametophyte plant. S, female gametophyte plant. C, production of
spermatia at the tip of male gametophyte. D, the carpogonium
(oogonium) is protected by filaments and spermatia become
attached to the protruding trichogyne. E, the zygote produces a
body of 2n carpospores, and the gametophytic cystocarp
enlarges and surrounds the carposporophyte. F, a sporophyte,
identical in appearance to the gametophyte, arises from the
carpospores. G, some pericentral cells give rise to meiocytes
which, through meiosis, produce four meiospores (tetraspores).
H, the vegetative structure of both gametophytes and the
sporophyte are identical.

chapter 11 |
Algae

248
 meiospores

Figure 1 1 .24 A sporic life cycle, with alternation of

heteromorphic generations, as shown by the brown alga
Laminaha.

Reproduction

249
 conceptacles

Figure 1 1 .25 A sporic life cycle, as shown by Fucus. A,

dichotomously branching blade with swollen tips. B, cross section
of tip showing cavities (conceptacles) C, the conceptacles
contain sterile hairs (paraphyses), shorter filaments bearing many
small antheridial meiocytes, and large oogonial meiocytes on
short stalks. D, detail of filaments bearing sperm. E, detail of
oogonial meiocyte after meiosis. F and G, meiosis in male
meiocyte. H, mitotic division resulting in a compound
microgametophyte. /, each cell functions as a sperm, shown
leaving the microsporangium, J, K, and L, meiosis in the oogonial
meiocyte. M, a single mitosis following meiosis. N, an eight-celled
megagametophyte forms. O, the eight cells leave the
megasporangium and function as eggs. P, fertilization takes place
in the open water.

chapter 1 1 |
Algae

250
Reproduction

251
Summary plants because they share the same chlorophylls and
storage products as higher plants. Their species exhibit
1. Algae are photosynthetic, nonvascular, relatively a great range of morphological diversity: unicellular
undifferentiated organisms. Their gametangia are motile, unicellular nonmotile, colonial, filamentous
unicellular — that is, they lack a jacket of sterile without cross-walls, filamentous with cross-walls,
protecting cells around the developing gametes heterotrichous forms, and sheetlike forms. The
2. The algae consist of more than 25,000 species Pyrrhophyta include the organisms which, in great
belonging to 10 divisions. Although most species are cause certain kinds of red tides. The
density,
aquatic, the algae as a group are found in a great Phaeophyta include the largest, most anatomically
diversity of habitats including some extreme complex algae, called kelps. Photosynthate is
environments. translocated in some kelps through cells that closely
3. Ecological importance of the algae centers around their resemble sieve cells. Tissue resembling xylem,
role as phytoplankton at the base of aquatic food however, is absent.
chains. Water pollution can lead to cultural Reproduction in the algae includes asexual and sexual

eutrophication, red tides,and blooms. Some blue-green types.There is no typical life cycle for the algae as a
algae are also important because they fix atmospheric group. Asexual reproduction may involve cell division in
nitrogen into forms that can be utilized by other unicellular forms, fragmentation of filaments, or the
organisms. production of mitospores. The Cyanophyta only
4. The algae are economically important mainly because reproduce asexually. Sexual reproduction may involve
of their cell wall materials (agar, algin, carrageenan), isogamy, anisogamy, or oogamy, and the alternation of
but they have some medical, food, fodder, and fertilizer generations that results may be of identical-looking
uses. generations (isomorphic generations) or of different-
5. Algal classification is based on biochemical and looking generations (heteromorphic generations). In any
cytological traits, as well as gross morphology. The case, the gametophyte generation is haploid and the
Cyanophyta are the only prokaryotic algae. The sporophyte generation is diploid.
Rhodophyta share some biochemical traits with the In a gametic life cycle, mature plants are diploid and the
Cyanophyta but, in contrast, exhibit complex life cycles haploid stage is limited to the gametes. In a zygotic life

and a greater range of morphological diversity. The cycle, mature plants are haploid and the diploid stage is
Euglenophyta are mainly unicellular and have some limited to the zygote. In many algae, however, there are
protozoan traits, such as the absence of a cell wall. The two distinct, independent generations that alternate in a
Chlorophyta are thought to be precursors of higher sporic life cycle.

chapt er 11 |
A lgae
252
 12 CHAPTER

12 the mycota (fungi)

We commonly meet the fungi as mushrooms,

mildews, molds, and the organism that causes
athlete's foot. A few fungi provide food for
they produce for sexual production. Since there
simple way
is no

summarize the differences between the

to
various groups without discussing their life cycles, we
humans, but many more compete with us for food sources shall survey each group individually.
and several of them use us as their own living food
supply. The baking and brewing industries have an ancient
partnership with the fungus known as yeast Subdivision Myxomycotina
(Saccharomyces), and the medical profession has the
makings of another long partnership with fungi such as
Penicillium notatum for their production of penicillin and The slime molds, which there are some 300 known
of
other antibiotics. It is doubtful whether any other group of species, have a vegetative body that consists of a slimy
organisms is associated with human beings in as widely mass of naked protoplasm without an external wall. The
varied an array of relationships. body has no definite shape; it creeps slowly by amoeboid
movement on rotting tree trunks or dead leaves, engulfing
bacteria and the spores of other fungi, or absorbing
nutrients from dead organic matter. The absence of cell
Classification of the Fungi
walls in most stages, the amoeboid movement, and the
ability to take in solid food are characteristics that are
usually associated with animals. A good case may be
The fungi are eukaryotic organisms that reproduce by
made molds are really animals
for the idea that the slime
means of spores and that cannot perform photosynthesis.
that belong to the same group as the amoeba. It is in
There are more than 200,000 known species of fungi.
regard to reproduction that they are most like plants: they
They are classified as a single division, the Mycota, which
produce spores with cellulose walls.
is subdivided in different ways by different taxonomists.
There are two major groups of myxomycete fungi. The
The scheme shown below will be used in this text.
acellular slime molds have a vegetative body called a
Plasmodium (Fig. 12.1). It is a large, easily visible mass of
protoplasm with no internal division into cells
Division Mycota (the fungi) (ie, it is coenocytic). The Plasmodium originates as a

I. Subdivision Myxomycotina (slime molds) single cell, an amoeba or a flagellated cell. Feeding,

II. Subdivision Eumycotina (true fungi) growth, and nuclear division proceed without cytokinesis

A. Class Oomycetes (egg fungi) toproduce the large Plasmodium. Sometimes individual
B. Class Zygomycetes (zygote fungi)
Plasmodia fuse and sometimes two amoebae fuse to form
C. Class Ascomycetes (sac fungi) a diploid zygote that develops into a Plasmodium.

D. Class Basidiomycetes (club fungi)

Eventually the Plasmodium moves to a drier location and
E. Class Fungi Imperfecti (imperfect fungi) puts forth stalked masses of protoplasm (sporangia) in

which the contents become divided by cellulose walls to

All the fungi are heterotrophs: they must obtain food form spores. The spores are discharged from the
from the environment. Most are saprophytes. That is, their sporangia and spread by the wind. In the presence of
food is taken from nonliving sources such as plant and water, they germinate; the wall is ruptured, and the
animal wastes and dead bodies. The rest are parasites, contents escape in the form of a flagellated or amoeboid
organisms that take organic materials directly from other cell to repeat the cycle.
living organisms. There are even a few fungi that are The cellular slime molds differ from the acellular types
parasites on other fungi! in that they are not coenocytic. A pseudoplasmodium
The Myxomycotina are separated from the rest of the (pseudo = false) forms from many separate amoebae.
fungi because their active cells lack cell walls. The These amoebae originally live separately, feeding and
Eumycotina have cell walls, giving the organisms a more dividing, until eventually one or more of them begin to emit
or less rigid body shape and plantlike habits. a chemical signal (cyclic AMP) that diffuses into the
Mycologists (those who specialize in the study of fungi) surroundings and induces neighboring amoebae to
subdivide the Eumycotina on the basis of the structures migrate toward them. When the amoebae meet, they stick

253
 together; however, they retain their plasmalemmas and do
spore not fuse. This mass of separate cells, the
pseudoplasmodium, now moves about as a unit. As in the
germination acellular forms, the cellular slime molds reproduce by
means of spores in stalked sporangia.

Subdivision Eumycotina

All but the simplest of the true fungi (Eumycotina) have a

vegetative body composed of branching tubes (Fig. 12.2)
called hyphae (hypha, singular). True cell walls, produced
by the protoplast, surround the hyphae. The fungi are
eukaryotes whose cell contents are similar to those of
plants except that plastids and usually dictyosomes are
missing. Most of the space within the hypha is occupied
by large vacuoles.
Hyphae grow only at their tips; there the walls are soft
and extensible under the turgor pressure from the
protoplast. As a region of wall is left behind the growing
tip, it becomes thicker and thus highly resistant to further

stretching. Branches originate as bulges in the sides of

the growing hyphal tips, where the walls are still soft. The
branches behave exactly as their parent hyphae. Hence
the system of hyphae progressively extends and ramifies
karyogamyl as it absorbs nutrients and water from the substratum. The
(zygote) hyphae possess a "chemical sense" and can orient their
Figure 12.1 Life cycle of a myxomycete. growth and branching toward sources of raw materials
such as sugars, amino acids, water, and minerals. The
orientation of growth by chemical signals is called
chemotropism. Its effect here is to cause the fungus to

septate hyphae nonseptate (coenocytic) hyphae

mycelium

haustoria
mycelium /-

Figure 12.2 Various aspects of the fungal vegetative body.

chapter 12 |
The Mycota

254
progressively invade food sources such as moist slices of
bread and decaying stored fruits.

The system of connected hyphae is the fungus

organism. Mycologists call it a mycelium (Fig. 12.2). It is

comparable Plasmodium of the slime mold. There is

to the
some cooperation between neighboring hyphae, but the
degree of cooperation rapidly declines with distance.
Hence as the mycelium grows and spreads, its advancing
hyphae tend to form local groups that function
independently of one another, as they belonged to if

entirely separate mycelia. Older parts of the mycelium may

be abandoned, their usable contents withdrawn, and their
empty walls sealed off by cross-walls that resemble
abandoned tunnels. Such an event, or the accidental
shearing of mycelia into parts by outside forces, readily
converts a single mycelium into many mycelia. Thus,
fragmentation is a real and useful mode of vegetative Figure 12.3 Water mold growing on a hemp seed, x3.

reproduction that is available to nearly all fungi.

Mycelia often show sophisticated responses in which
development keyed to environmental limitations and
is
cell. Septa, or crosswalls (septum, singular), may form
opportunities. These responses use light, gravity and
occasionally in old hyphae of some species, and normally
chemical signals to direct growth and to select the best
the reproductive cells are cut off from the vegetative
times for reproduction.
hyphae by crosswalls.
True fungi are solution feeders: they can take in only
The Oomycetes are divided into several orders.
small molecules. In many cases their food requirements
Specialists do not agree completely and the number of
are very simple, consisting of no more than an organic
recognized orders depends on the authority one is
carbon compound such as a sugar, plus water, minerals,
consulting. We shall consider the orders Saprolegniales
and perhaps a few vitamins (enzyme cofactors, required
and Peronosporales. Note that the names of orders end in
only in small amounts, which the fungus cannot build for
-ales.
itself.) The vitamin most often needed by fungi is the same
Vitamin B, that is essential in human nutrition. Despite the Saprolegniales
simplicity of their needs, mycelia can exploit a wide range
of organic materials as food sources if they are available. Fungi of this order usually live in fresh or salt water;

Food sources are often hard materials such as wood, but hence the common name water molds. Most are
also include insoluble storage polymers such as seed saprophytic. Even the species that attack the gills of fish
proteins and starches. These materials must be rendered grow on tissues that have been weakened or suffered
soluble before they can diffuse through the walls of the injury. They may be easily cultivated by placing small

hyphae and reach the protoplast. Hyphae build and pieces of meat, egg albumin, radish seeds, or dead flies in
secrete enzymes that hydrolyze proteins to amino acids a dish of pond water (Fig. 12.3). After the mycelium has
and polysaccharides to free sugars. Working outside the ramified through the substratum, hyphae grow outward

hyphae, these enzymes require water as a reactant and as into the water. Reproductive cells are produced by these

a medium for support and transport. Hence mycelia tend hyphae.

to be active only in moist places.
Asexual Reproduction

With ample food supply, there is little tendency to produce

Class Oomycetes reproductive cells. If a well-developed mycelium is

transferred to distilled water which a food supply is

in

The Eumycotina include some fungi that have swimming lacking, asexual sporangia will usually appear. Sporangia
cells and some that do not. This distinction seems are formed by a cross-wall cutting off the tip of a hypha
fundamental, and mycologists group all fungi with motile from the rest of the mycelium (Fig. 12.4A). The
cells into a class that in this book we
Oomycetes call sporangium is multinucleate. The protoplasm of the
(other names are available; e.g., Mastigomycotina). sporangium divides into a large number of spores, each
Many Oomycetes cause severe plant diseases; others with one nucleus (Figs. 12.46, C). Upon maturity, the
infect fish, insects, or even humans. Many members are spores are discharged from the sporangium. In most
saprophytes or weak parasites. Some are minute forms of species these spores have flagella that permit swimming
one to several cells, and a few of these smaller sorts are movements, and are therefore known as zoospores. In
reported to attack small aquatic animals. Most Saprolegnia, each zoospore has two flagella attached to
representatives of the Oomycetes develop a more or less its anterior end that enable it to swim actively (Fig. 12.4D).

extensive mycelium of indefinite form (Fig. 12.3). After a time, the zoospores settle down, lose their flagella,
The hyphae of actively growing Oomycetes are develop a cellulose wall, and pass through a resting
generally coenocytic (Fig. 12.2). The whole mycelium period. When they have resumed activity, they escape
consists of a single, very highly branched multinucleate from the wall; the two newly developed flagella are now

Subdivision Eumycotina

255
 flagella

immature sporangium

Figure 12.4 Zoosporangia of Saprolegnia. A, immature

sporangium;
Dorangium; B, mature sporangium; C, discharge of zoospores; D,
a zoospore

attached laterally, and zoospores swim about for a period. antheridium touches an oogonium, it produces a short
If they come to rest on a suitable substance, each sends hypha called a fertilization tube that punctures the
out a hypha that penetrates this substance. oogonial wall, and comes in contact with one or more
In a few species of water molds, one or even both eggs. If the oogonium contains several eggs, the
zoospore stages are suppressed. When zoospores emerge fertilization tube usually branches, sending a branch to
from the sporangia, they may germinate directly into a each egg. Nuclei (male gametes) from the antheridium
new mycelium. In certain other species, the spores never migrate into each egg, and fertilization ensues. The zygote
leave the sporangia but germinate while still enclosed develops a heavy wall, becoming an oospore, and usually
within it, and the germ tubes pierce the old sporangial will not germinate for several months, even under

still other species, spores are not even formed; the

wall. In favorable conditions. Upon germination, the oospore sends
sporangia germinate directly into a coenocytic mycelium. out new hyphae, which rapidly grow into a typical
mycelium.
Sexual Reproduction Some oomycete species are bisexual, both kinds of
gametangia being borne on the same mycelium. But
When stimulated to reproduce sexually, the mycelium
others have separate sexes. In these species, the
produces specialized hyphae called oogonia and antheridia and oogonia do not form unless a suitable
antheridia in which gametes are formed. The nuclei in the partner is available. The system that signals the presence
vegetative mycelium are diploid; meiosis occurs in the
of a partner has attracted considerable scientific interest
specialized sexual hyphae to produce the haploid nuclei
because it is a clear-cut case where environmental signals
needed for gametes. cause major developmental changes. By placing male and
Egg cells are formed inside the oogonia, which are female mycelia in the same tank of water but separated by
spherical cells at the tips of short side hyphae (Fig. 12.5).
a membrane, it has been found that the female constantly
When mature, each oogonium is three to four times the emits very small quantities of a compound that functions
diameter of an ordinary hypha and may contain from one as a sex hormone. The male sensitively detects this
to twenty eggs. Each spherical egg contains one nucleus.
hormone and responds by producing antheridia. The
The antheridia are also formed at the tips of hyphae
in turn give off a second hormone that diffuses
antheridia
(Fig. 12.5).Each is separated from the rest of the hypha to the female mycelium and induces it to produce oogonia.
by a cross-wall. The antheridia are usually similar in Additional hormones induce the antheridium to grow
diameter to ordinary hyphae. it, and produce the
toward the oogonium, cling to
The antheridium grows toward an oogonium; it is
fertilization tubes. It has not yet been discovered how the
directed in its growth by a chemical signal (a sex hormones produce these responses, but special sets of
hormone) released by the oogonium. When the genes are probably called into play.

chapter 12 |
The Mycota

256
 the day had no idea of the cause and were skeptical when
botanists discovered that a fungus was the culprit. Since
then it has been found that Phytophthora infestans can be
controlled (it still exists in potato fields) by spraying
infected fields with fungicide, as well as by carefully
oogonia
disposing of infected tubers.
The mycelium of Phytophthora grows between the cells

septum within a leaf or stem (Fig. 12.6). Parasitic hyphae

(haustoria) penetrate the cells. Aerial sporangia arise on
long aerial hyphae called sporangiophores that extend
out through the stomata of the infected plant. The
theridia sporangia break loose and are disseminated by air

currents.
Some of the sporangia eventually come to rest on the
hyphae leaves of susceptible plants. When moisture on the leaf

surface is sufficient, zoospores escape from them. They

soon send out small hyphae called germ tubes that
penetrate the host tissues and bring about new infections.
Downy mildews are a group of fungi in the
Peronosporales that cause infections on many cultivated
and wild plants (Fig. 12.7). They are easily identified by
the sporangiophores that may, in severe cases, nearly
cover the leaf. In the laboratory, the sporangia can usually
be induced to germinate by floating them on cool water.
A small group of Peronosporales, the "white rusts,"
develop sporangia beneath the epidermis of such crop
antheridia
plants as mustards and spinach. We mention them
because their sporangiophores do not grow out of the
stomata as in the downy mildews. Instead, they collect in
pustules under the epidermis of the stem or leaf.

Sporangia are cut off in chains from the tips of the

sporangiophores. They accumulate in large numbers and
wall of
finally rupture the epidermis, forming creamy-white
oogonium pustules (Fig. 12.8).
Sexual reproduction in the Peronosporales is similar to
that observed in the Saprolegniales.

Figure 12.5 Gametangia in Saprolegnia. A, oogonia and Class Zygomycetes

antheridia with associated hyphae; B, antheridia, oogonium with
many eggs, tertilization tubes in place.
The Zygomycetes are the simplest group of fungi that lack
motile cells. They owe their name to the trait of converting
the zygote into a resting spore called a zygospore (Fig.
12.10). These are chiefly terrestrial fungi that form an
Peronosporales extensive coenocytic mycelium. In contrast to the
Nearly all the Peronosporales are obligate parasites. In Oomycetes and the green plants, the walls of the
general, heavy-walled oospores carry them over zygomycete mycelium have chitin as their fibrous
unfavorable periods, and various sorts of asexual spores framework. Chitin is a polysaccharide in which the sugar
bring about rapid multiplication under suitable conditions. subunits have been modified by the addition of a
The most famous member of the Peronosporales is nitrogenous group. The same substance is responsible for
Phytophthora infestans, (Fig. 12.6), a fungus that indirectly the strength of insect exoskeletons.
caused the deaths by starvation of at least 250,000 people The hyphae of the vegetative mycelium invade the food
in Ireland between 1843 and 1847, and induced another substrate and also grow on the surface, forming a cottony
million Irish people to move to the United States. This mass (Fig. 12.9). When the fungus is well established, it

fungus causes a disease of the potato called late blight. commences sexual reproduction with the formation of
In those years Ireland had virtually a one-crop economy, specialized sexual hyphae (Fig. 12.10). These hyphae
featuring the potato. Phytophthora was present in the grow toward one another, touch, and fuse at the tips.
potato fields for many years prior to the starvation years, Special enzymes dissolve the wall at the contact point so
gradually spreading, but conditions were never right for that the protoplasm of the two hyphae flows together in a
maximum growth. Then, starting with 1843, there followed single mass. At the same time, a crosswall forms behind
a period of unusually warm, humid summers during which the contact point on each side. This walls off a single
the fungus nearly annihilated the potato crops. The large cell with many nuclei from each parent. The nuclei
disease kills foliage and rots the potato tubers. Farmers of fuse in pairs to form diploid zygote nuclei. The

Subdivision Eumycotina

257
Figure 12.6 Asexual reproduction of Phytophthora infestans.
The vegetative mycelium invades the spongy mesophyll of a
potato leaf. Sporangiophores (A, B) grow through stomata.
Multicellular sporangia (B, C) form. Each sporangium breaks off
as a unit (C) and is carried by wind to another leaf surface (D).
Zoospores emerge from the sporangium (D) and swim in the film
of moisture on the leaf surface before encysting and becoming
dormant (E). The dormant spore later germinates (F) and
produces a germ tube that may penetrate the epidermis through
cell walls or stoma, to produce a new mycelium within the leaf.
By repeating these events the spores can quickly infect a whole
potato field.

surrounding cell wall thickens and the cell becomes a

dormant spore (a zygospore). The diploid nuclei undergo

meiosis to produce a new generation of haploid nuclei
before the zygospore germinates. In all the life history of
the zygomycete fungus, only the zygospore is diploid.
In some species of Zygomycetes, a single mycelium can

produce zygospores without a partner. But many species

require a mating between sexual hyphae of two different
mycelia of opposite "sexes." The partners look alike and
contribute equally to mating, so mycologists designate
them as " + " and "-" mating types rather than "male"
and "female." The mycelia advertise their mating type to
one another by emitting sex hormones, as in the
oomycetes.
The Zygomycetes reproduce asexually by means of
sporangia, which form at the tips of aerial
sporangiophores (Fig. 12.11). The sporangiophore swells
at the and a crosswall forms below the swelling. The
tip

multinucleate protoplasm in the swollen sporangium then

divides into many spores. Members of the class differ
greatly in the details of asexual reproduction: for instance, Figure 12.7 Downy mildew (Bremia) on lettuce leaf.

chapter 12 |
The Mycota

258
Figure 12.8 Albugo on shepherd's purse.

young zygospore

zygospore

-"'/> .'" .3" Figure 12.10 Sexual reproduction in Rhizopus stolonifer. (From
Cryptogamic Botany, vol /4/gae and Fungi, by G. M. Smith.
I,

Copyright 1 955, McGraw-Hill Book Company. Used with

Figure 12.9 Rhizopus on peach, x4. permission of McGraw-Hill Book Company.)

the common bread mold, Rhizopus nigricans, forms sporangiophores, or sporangiophores that branch and
hyphae (stolons) that arch through the air
special, thick bear several sporangia.
and are anchored to the substrate at intervals by finer Many of the Zygomycetes have sophisticated
branch hyphae or rhizoids (Fig. 12.1 1). Clumps of mechanisms that use light and gravity to direct
sporangiophores arise at each anchor point. Other sporangiophore growth so that the spores will be brought
members of the Zygomycetes produce solitary Other mechanisms may
to locations suitable for dispersal.

Subdivision Eumycotina

259
 developing sporangia

Figure 12.11 Asexual reproduction in Rhizopus stolonifer.

time the formation of sporangiophores on the basis of to the ( + ) and (-) nuclei pair but do
ascogonium. The
temperature, illumination, and nutritional conditions. not fuse. The ascogonium produces hyphae into
fertilized

Spores are dispersed by various means. In some which the pairs of nuclei migrate. Hyphae with the paired
species the sporangial wall breaks down at maturity and nuclear arrangement are not truly diploid (2n), nor are
the light, dry spores are spread by wind. In others the they haploid (1n). Rather, their nuclear condition is

spores form a sticky mass that is carried by animals. symbolized as (n+ n). An (n + n) hypha is known as a
Members genus Pilobolus build up pressure in the
of the dikaryon (di = two; karyon = nucleus). Since these n+n
sporangiophore until the sporangium as a whole is hyphae produce asci, they are called ascogenous
discharged explosively and travels several meters through hyphae.
the air (Pilobolus literally means "hat thrower.") The ascogenous hyphae may grow and branch before
Most Zygomycetes are saprophytes, though a few
of the they develop into asci. The formation of the ascus is
are parasitic on other members of the same class. If left outlined in Fig. 12.14.
uncontrolled some, such as Rhizopus, cause major losses Most ascomycetes concentrate their asci within a
of stored foods. Few of them cause human diseases. complex reproductive structure called an ascocarp (Fig.

12.15).
The ascocarp, composed of both vegetative and ascus-
Class Ascomycetes bearing hyphae, is characteristic of the species. It may be
microscopic or as much as 15 cm in diameter. There are
The Ascomycetes are named for their trait of producing three general types of ascocarps:
sexual spores inside a sac (ascus) like marbles in a
1. Cleistothecium: hollow, completely closed sphere
transparent bag (Fig. 12.14). Ascus is derived from the
(Figs.12.15AS).
Greek word for sac (plural asci).
2. Perithecium: hollow, flask-shaped body with narrow
The an ascomycete typically begins with the
life of
opening (Figs. 12.15C.D).
germination of a haploid spore to produce a vegetative
3. Apothecium: open, cup-shaped body (Figs. 12.15E.F).
mycelium. As in the Zygomycetes, the walls contain chitin
instead of cellulose. But in the Ascomycetes the mycelium The asci line the inner surface of the ascocarp. This
is divided into cells by cross-walls. The cross-walls have a surface layer is the hymenium or fertile layer. Sterile

hole in the center (Fig. 12.12) that allows the passage of haploid cells, called paraphyses, also arise in the
organelles from one cell to the next; however until sexual hymenium (Fig. 12.16).
reproduction occurs, each cell usually maintains just one The hymenial layer is surrounded and protected by
haploid nucleus. haploid hyphae that form a layer called the peridium.
Sexual reproduction begins with the fusion of two cells The fertilization of the ascogonium is usually the signal
(Fig. 12.13). Many ascomycete fungi are self-incompatible; that starts the formation of an ascocarp. Therefore in the
fusion occur only if the cells come from mycelia of
will ascomycetes, n+n
hyphae are usually confined to the
opposite mating types. In some species, fusion can occur reproductive body and do not contribute to the vegetative
between any two hyphae, while in others, fusion involves mycelium. This contrasts with the Basidiomycetes, to be
specialized multinucleate hyphae called ascogonia and discussed later.

antheridia, and still others produce small free cells called The ascocarp is composed of both haploid and n+n
spermatia instead of antheridia. hyphae. The result is a multicellular body that is
When contact is made, nuclei pass from the antheridium considerably more complex in structure than anything

chapter 12 |
The Mycota

260
Figure 12.12 A small portion of a hypha of Rhizoctonia solani
showing a septum, with pore, through which a mitochondrion
appears to be passing, x 24,000.

( + ) spore

ascogenous
hypha

(— ) spore

Figure12.13 Diagram of conjugation and formation of n-

ascogenous hyphae.

found in the Oomycetes and Zygomycetes. For this reason light enough for long-distance travel on the wind. Spores

the Ascomycetes are often called higher fungi and the produced by such a pinching process are called conidia.
former groups lower fungi. The hyphae that produce them are conidiophores
Although sexual reproduction is important for producing (-phore = bearer). Various species differ greatly in the
new genetic combinations in the Ascomycetes, asexual branching of the conidiophores. Conidia are usually
reproduction more important as a means of
is produced in immense numbers. Cold generally kills them.
multiplication and dispersal. Most species reproduce But within a single summer, the growth of mycelia is so
asexually by pinching off the end cells of aerial hyphae dozens of generations of conidia and mycelia can
fast that
(Figs. 12.17). These cells develop resistant walls and are be formed and epidemics of fungal growth can occur.

Subdivision Eumycotina

261
 ascogenous
hypha

Figure 12.14 Ascus formation. A, paired nuclei divide by

mitosis; hypha develops J shape. B, crosswalls form; penultimate
cell isn+n. C, nuclear fusion (karyogamy) in penultimate cell. D,
meiosis divides 2n zygote into four haploid nuclei. E, each
nucleus undergoes mitosis. F, walls form around haploid nuclei,
yielding eight ascospores inside ascus (old hyphal wall.)

Figure 12.15 Diagram showing three types of ascocarps. A and

B, cleistothecia, X500; C and D, perithecia, x150; E and F,
apothecia, E x '/4l F x 10. (From Comparative Morphology of
Fungi, by E. A. Gauman; copyright 1928, McGraw-Hill Book
Company. Used with permission of McGraw-Hill Book Company.
D, redrawn from E. A. Gauman, The Fungi. A Description of Their
Morphological and Evolutionary Development. Hafner Publishing
Company, New York, 1952.)

chapter 12 |
The Mycota

262
 ( + ) spore spore

Figure 12.16 Diagram of a cross-section of an apothecium.

(Redrawn from Fundamentals of Cytology, by L. W. Sharp.
Copyright 1943 by McGraw-Hill Book Company, New York. Used
with permission of McGraw-Hill Book Company.)

palisade parenchyma

haustoria
Figure 12.17 Powdery mildew on leaf surface.

Subdivision Eumycotina

263
Examples of Ascomycetes mitosis and budding (Fig. 12.18). Later, meiosis finally
intervenes to restore the haploid condition. Yeasts differ
There are about 15,000 species of Ascomycetes, and so
greatly in the relative time spent in the diploid and haploid
many variations that even an introductory survey of them
states. Some strains are diploid most of the time, a highly
would require many pages. Thus in the present book we
unusual trait among the sac fungi.
will consider only a few illustrative examples.

Powdery mildews. These are a group of ascomycete

Yeasts. The simplest ascomycetes are the yeasts. genera that owe their common name to the development
Saccharomyces cerevisiae is the yeast of baking and of a powdery appearance on the leaves of infected plants.
brewing. This fungus is important because converts it
All the powdery mildews are obligate parasites; they can
sugar (glucose) into alcohol and carbon dioxide. This trait
only grow on living plants. The genus Erysiphe is
is energy from glucose when molecular
useful in gaining representative (Fig. 12.17).
oxygen is not available (Chapter 2). In the presence of The mycelium is generally confined to the surface of the
oxygen, yeast (like other organisms) can completely leaves, flowers, or fruits. Haustoria penetrate epidermal
oxidize glucose to carbon dioxide and water; although it
and parenchyma cells, from which they secure
gains much extra energy this way, does not produce the
it
nourishment. At first the mycelium appears like a delicate
economically valuable alcohol. cobweb. Eventually, it assumes a white powdery
Yeasts are normally single-celled. After cell division the appearance because of the development of numerous
new cells separate; thus ordinary mitotic division is a form conidiospores that serve to spread the fungus from one
of asexual reproduction (Fig. 12.18). In older cultures the plant to another during the growing season.
cells may hang together and form short chains. Some Sulfur dusted on the host plants at this stage prevents
yeasts double and pinch in half. More often,
in size infections from these spores.
though, division is a process of budding: new cells grow For sexual reproduction, small black cleistothecia are
out from the mother cell much as a small bubble would formed, each enclosing one or several asci (Figs.
form if a piece of thin rubber were made to expand The ascospores are
12.15/4,6). usually discharged by
through a small opening in some heavier material. The force from the cleistothecium.
small "bubbles" are called buds. They enlarge and finally
Although the powdery mildews may not kill their host,
separate from the parent cell.
theyweaken it and greatly reduce the crop yield. Powdery
Ascus formation in yeasts is very simple. Two mildews cause diseases of apples, grasses, grains,
compatible yeast cells fuse, the nuclei fuse, and meiosis grapes, cherries, and many other plants.
follows. The original wall forms the ascus. There is no
ascocarp. Monilinia. Brown rot of stone fruits, a very severe
Most Ascomycetes do nothing more with the diploid disease of peaches, cherries, plums, apricots, and
zygote nucleus than to divide it by meiosis, producing nectarines, is caused by a disk fungus (Monilinia, Fig.
haploid nuclei. Some yeasts follow the same pattern. But 12.19, Color Plate 3), which infects mainly blossoms and
in others, the diploid cell may reproduce asexually by fruits. The ascocarps of the order to which Monilinia

n + n
conjugation

Figure 12.18 Life cycle of Saccharomyces cerevisiae

chapter 12 |
The Mycota

264
Figure 12.19 A, apothecia of
the brown rot fungus COLOR PLATE 3
(Monilinia fructicola), x '/2
;

6, brown rot on cherry

compared with a healthy fruit,

x '/2 .

Figure 12.33 Various types

of lichens. A, the conspicu-
ous green foliose lichen is
Peltigera aphthosa and it is
surrounded by the white
branches of the fruticose
lichen Cladonia sp.; B, crus-
tose lichen Acarospora flava
colors a rock on the skyline
of a Sierra Nevada ridge; C,
Ramalina reticulata, a fruti-
cose lichen, clothes the
branches of an oak in the
foothills of the Coast Ranges
along the Pacific Coast.

:\\^ ~
Figure 13.11 Habit of
Marchantia A, x '/6 8, x 1 ;
COLOR PLATE 3
Note gemmae cups. (continued)
C, anthendial heads, x 2.
D, archegonial heads, x 1.5.
E, under surtace of an
archegonial head; the
sporophytes are enveloped by
a protective perianth
Immature sporophytes are
green; ripening sporophytes
show yellow, x 3.
belongs are cup- or disk-shaped (apothecia) (Fig. even cause extensive decay mines and
of timbers in

12.15E.F; 12.19/4— Color Plate They may be as much

3). buildings. Specialized tissues and organs may be formed,
as 15 cm in diameter, depending on the species, and are some of the n + n hyphae becoming specialized to
sometimes brilliantly colored. Members ot a related genus, transport food and water, and others becoming tough and
Morchella (the morels), are edible. woody.
The vegetative body of such a fungus is a mycelium that
Claviceps. The genus Claviceps
is parasitic on grasses, starts from a haploid spore and invades its substratum as

including grains. A
dormant mycelium that replaces the in previous groups. The walls of the mycelium contain

mature grain is known as ergot. It possesses several chitin rather than cellulose, and the hyphae are almost

alkaloids that have medicinal properties. Ergot constricts completely septate. There is a pore in the center of each
the blood vessels, particularly those that pass into the cross-wall but an elaborate caplike structure covers the

hands and feet, thus depriving the extremities of a normal pore so that organelles probably cannot pass between
blood supply. During humid summers in Central Europe, cells.

Claviceps may infect rye heavily. In centuries past, before These fungi are not noted for their asexual reproduction.

the nature of the fungus was understood, ergot would be Instead, sexual reproduction is highly developed.

milled along with the grains of rye. The contaminated Sexual reproduction begins with the conjugation or
flour, which might contain as much as 10% of powdered fusion of hyphae to produce a dikaryon with the paired,
mycelium, would be baked in bread. A continued diet of n+n nuclear condition. Some species are homothallic,
bread from this flour resulted in much misery. Because of conjugation taking place between any two cells of any two
the contraction of the blood vessels and the limited supply hyphae or even between two cells of the same hypha.
of blood reaching feet or hands, gangrene sets in. Hands, Other species are heterothallic and conjugation will only
arms, and legs die and finally drop off. The disease was occur between hyphae from mycelia of different mating
known as "Holy Fire" because of a burning sensation in types.

the extremities. Today, ergot is a valued drug used to Once formed, the dikaryon may grow and branch to

control hemorrhage, particularly during childbirth, and to form a long-lived n+n vegetative mycelium. The dikaryon
treat migraine. may persist for many years, perhaps even for centuries. If

With the knowledge of the poisonous nature of the conditions are not right, the production of sexual spores
ergot, diseased grain no longer milled and Claviceps is
is and basidiocarps may be postponed indefinitely.
not of concern indiet. However, cattle may
human In the n+n mycelium, division occurs in such a way

occasionally feed on grasses infected by ergots and thus that all the cells maintain the n+n condition. This requires
be poisoned. a special set of events (Fig. 12.20). Growth of the tip cell

of the accompanied by division of both nuclei.

hypha is

The tip cell develops a J-shape and one of the nuclei

Class Basidiomycetes migrates into the end of the J. Cross-walls are laid down
as shown in Fig. 12.20. The lateral cell bends around,
Based on the complexity and size of their reproductive fuses with the trailing cell, and the n+n condition is
structures, the most advanced of the higher fungi fall in restored in that cell. This happens each time a new cell is
the class Basidiomycetes. These include all the formed at the tip of the hypha. The curved and fused
mushrooms, puffballs, and the bracket fungi that grow on lateral cell is termed a clamp connection. Of all the fungi,
the trunks of trees. These elaborate structures are called only the Basidiomycetes form a dikaryotic vegetative
basidiocarps. mycelium with clamp connections.
But not all the Basidiomycetes have such elaborate Although both haploid and n+n mycelia develop
reproductive bodies. The real feature that determines extensively, the n+n phase is of special interest because
whether a fungus belongs to this class is the precise way it gives rise to the spore-bearing body, or basidiocarp, in

in which sexual meiospores are formed. Rather than being which the basidia are developed. The hyphae form a
enclosed in a common sac, here the spores formed after tangled mass in the substrate or host and emerge to form
meiosis are carried on separate stalks called sterigmata
(sterigma singular) (Fig. 12.22). The cell that undergoes
meiosis and produces the spores is called a basidium,
because in some cases it is club-shaped (basidium is
Greek for "club"). The meiospores are called
basidiospores.
The Basidiomycetes consist of two main groups or
subclasses, which will be considered in turn.

Subclass Homobasidiomycetidae
The Homobasidiomycetidae comprise all the species that
form elaborate fruiting bodies or basidiocarps. Some of
these are parasitic but most of them are saprophytic.
Some form mycorrhizae, symbiotic associations with the
roots of higher plants, previously discussed in Chapter 5.
Figure 12.20 Formation of a clamp connection, which maintains
Others obtain their carbohydrates from wood and may the dikaryotic condition in the basidiomycete hypha.

Subdivision Eumycotina

265
rlQUr© 1 c..c.\ f* :us brunnescens, the common edible
mushroom, x 1,

a basidiocarp —
the mushroom, puffball, or bracket fungus. along with cytoplasm. The formation of a resistant wall
The basidiocarp of a typical mushroom such as Agaricus completes each spore.
brunnescens (Fig. 12.21) consists of a short upright stalk If the gill is very young, developing basidia on which
or stipe, attached at its base to a mass of mycelium and basidiospores have not yet been formed may be observed;
expanding on top into a broad cap or pileus. The Figure 12.23 shows a section through three gills. Note two
underside of the pileus of Agaricus is formed by thin gills stages of development of basidia and the loose tangle of
radiating outward from the stipe. These gills are lined with hyphae that form the center of the gill. When a
a hymenium or spore-bearing layer. basidiospore is discharged, it is shot horizontally from its

Basidia develop from the tipmost cells of hyphae in the position on the basidium straight into the space between
hymenium Here the paired nuclei finally fuse
(Fig. 12.22). two gills. A basidiospore resembles a toy rubber balloon,
which immediately
to give a diploid (2n) nucleus, having a large volume for its weight. It is shot from the
undergoes meiosis. Sterigmata form at the tip of the basidium with a force sufficient to carry it about midway
basidium. The haploid nuclei that result from meiosis are between the two gills. It then falls straight downward. It

squeezed through the narrow necks of the sterigmata has been estimated that some basidiocarps may discharge

sterigma basidiospore n

fusion nucleus '

basidii

Figure 12.22 Basidiospore formation as it occurs in a

mushroom (Homobasidiomycetidae). (From Cryptogamic Botany:
Vol. I Algae and Fungi, by G. M. Smith; copyright 1955, McGraw-
Hill Book Company; after H. Kniep, Sexualitat der Niederen
Pflanzen, copyright G. Fischer Verlag, Jena 1928; and after
Researches on Fungi, Vol. VII The Sexual Process in the
Uredinales, by A. H. R. Buller; University of Toronto Press 1922.
Used with permission from McGraw-Hill Book Company, Gustav
Fischer Verlag, the Royal Society of Canada, and the University of
Toronto Press.)

chapter 12 |
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266
 Not Agaricales have gills. In some, the lower surface
all

of the cap or bracket has many small pores that extend

upward to form small tubules (Fig. 12.24). The hymenium
lines the tubules. Many bracket fungi found on the trunks
of forest trees are such pore fungi.
Some bracket fungi cause serious damage to forest
trees and lumber in mines and wooden buildings. During
World War much green lumber had to be used; because
II,

fungi were able to feed on the improperly cured lumber,

much loss resulted. One of the best-known fungi in this
group is Fomes applanatus (Fig. 12.25). causes a
It

disease of forest trees known as white-mottled rot, to

which many hardwoods and conifers are susceptible.
The basidiocarp of Fomes applanatus is a very elegant
shelf-shaped bracket or conk. The conks are perennial
and grow to a large size. They are gray on top, and the
lower surface has millions of pores lined with a creamy-
white hymenium.
Most of the bracket fungi of the forest normally live in
Figure 12.23 Section showing three gills of a mushroom.
the dead heartwood, to which they gain entrance by
wounds. They are, however, able to invade the living cells
of sapwood, which they may destroy.
as many as a million spores a minute for several days.
Other types of basidiocarps also occur, such as those in
The Agaricales include not only edible fungi but also
puffballs, and bird's-nest fungi.
some of the most deadly of poisonous mushrooms. Those
of the genus Amanita (Color Plate 1 ) are especially
Subclass Heterobasidiomycetidae
dangerous; it is said that the Eskimos used to pay high
prices for dried specimens that, when taken in extremely The fungi of this subclass are much less familiar to the
small quantities, induce hallucinations. Experimentation is average person than those of the group just described,
ill-advised, however, because small dosages can be toxic. because they do not produce elaborate basidiocarps. But
Although poisonous mushrooms are much less common economically they may be much more important. This
than benign ones, mycologists usually advise enthusiasts subclass includes some of the most virulent crop
to consultan expert before sampling mushrooms they destroyers: the rusts and smuts, which attack cereal
have gathered from nature. grains such as wheat and corn and cost millions of dollars

Figure 12.24 The underside of the pore fungus Polyporus

brumalis, x '/3 .

Subdivision Eumycotina

267
 Puccinia graminis has two hosts: a grass, and a shrub
known as barberry (Berberis). Various strains of the
fungus attack different grasses. Variety avenae attacks
oats and variety tritici attacks wheat. These host choices
are hereditary with the fungus.
The two hosts have different roles in the life of the
fungus (Fig. 12.26). Roughly speaking, the grass serves to
multiply and spread the existing genetic types of the
fungus population. This is achieved by asexual
reproduction using food reserves that are drained from the
leaves and stems of the grass plant. The grass is usually
not killed, but its vitality is impaired and its seed
production reduced.
The barberry host is the sexual ground for the rust
fungus. Here, matings between different mycelia produce
new genetic combinations. These in turn move out to
invade the grain fields. The barberry plant is not killed by
the infection.
To survey the life cycle of wheat rust (Fig. 12.27), let us
begin with the basidiospores. These spores are carried by
air currents in the spring, and they can only attack
barberry shrubs. They germinate on barberry leaf
Figure 12.25 Basidiocarp of Fomes applanatus, x'/j.
surfaces, and form a germ tube that penetrates the
epidermis.
A mycelium develops in the tissues of the host.

in lost harvests and preventive costs. Most members of

Pustules, called spermagonia, appear on the upper
surface of leaves. A spermagonium is pear-shaped and
this subclass are parasites on plants.
Some of the Heterobasidiomycetidae may have two has a small opening through the upper epidermis to the
hosts; that is, separate phases of their life history are Hyphae in the
exterior of the leaf (Fig. 12.27/\).

passed on different plants. Such forms are heteroecious. spermagonium cut off large numbers of spermlike cells,
If only one host is required to complete their life history, the spermatia. These are forced out of the spermagonia

they are said to be autoecious. through the pores.

The rust fungi owe their name to the rusty color of some Puccinia graminis is heterothallic; both plus and minus
of their spores, which form in abundance on the leaves of mating types appear on barberry leaves. These develop
infected grasses. An example of this group is Puccinia from plus or minus basidiospores. Spermatia are
graminis. It causes common wheat rust, and is a persistent transferred by wind or insects to adjacent spermagonia,
and costly pest. where they come in contact with receptive hyphae.

bosidium- -+- basidiospores --mycelium

/ (2n) In) in)

karyogamy

f
teliospores (n+n)

receptive spermatia (n)

hyphae (n)

conjugation

mycelium (n+n)
— aeciospores-4-
(n + n)

wheat host barberry host

Figure 12.26 Summary of Puccinia graminis life cycle.

chapter 12 |
The Mycota

268
spermatid fertilize compatible receptive hyphae

Figure 12.27 Stages in the life cycle of wheat rust, Puccinia

graminis. A, formation of receptive hyphae and their conjugation
on barberry; 6, pustules (aecia) on barberry leaf; C, section
through aecia and spermagonia on barberry; D, pustule (sorus)
on wheat stem, producing red uredospores; E, the same
mycelium on wheat later produces black teliospores as seen in
this section view.

Subdivision Eumycotina

269
 basidiospores

karyogamy
+

meiosis »•

Figure 12.28 Germination of a teliospore, Puccinia graminis. A,

teliospore with binucleate cells; B, 2n cells; C, meiosis and
germination of a cell produces a hypha with four haploid nuclei;
D, crosswalls form and basidiospores are extruded.

A plus ( + ) spermatium fuses with a cell of a minus (-) germinate and produce basidiospores as in Puccinia.
receptive hypha, and vice versa. An n+n mycelium Haploid mycelia from these spores develop on the corn
results. In its formation, the process of genetic plants and conjugate to restore the n+n condition that
recombination has been completed. leads to teliospores. Resistant strains of corn effectively
How do mycelia get from barberry to wheat? The newly limit this pest.

formed n+n mycelium on the barberry leaf invades the Another group smut fungi can invade only the pistils
of
spongy mesophyll. Close to the lower epidermis the of flowers in grain plants. These are the blossom-infecting
mycelium forms pustules called aecia (Figs. 12.278,Q. smuts. Ustilago tritici and U. avenae attack wheat and
Here n+n spores (aeciospores) are formed asexually. oats, respectively. The germinating spore builds a small
They are released when the pustule breaks through the mycelium that gently invades the grain seed. When the
leaf epidermis. Aeciospores, borne by the wind, attack infected seed is planted, the contained mycelium grows in
only wheat plants. pace with the plant, even accelerating its growth. But
Hyphae from the germinating aeciospores enter the when flowers are formed masses of smut spores form in
wheat plant through its stomata. A delicate mycelium place of the seeds and are spread by wind to the slower-
forms with haustoria that penetrate the cells of the leaf growing normal plants in the neighborhood.
and extract nutrients. About 10 days after infection, red Still other smuts infect only young seedlings. This is true

n+n uredospores form. The pustules or uredia of Tilletia tritici, which invades wheat plants. These also
containing these spores open up (Fig. 12.27D). Carried on replace the mature grain with masses of spores.
wind currents, the uredospores attack other wheat plants
at a rate that can attain epidemic proportions. Wheat rust
gets its name because of these red lesions (uredia) in the Class Fungi Imperfecti
stems.
As the wheat begins to mature, the production of About 24,000 species of fungi, in some 1200 genera, are
uredospores gives way to the production of black known only by their asexual stages. The class name,
same mycelium (Fig. 12.275). Whereas
teliospores by the Fungi Imperfecti, arises from the custom of calling the
uredospores and aeciospores are killed by winter cold, sexual stages of fungi perfect stages and the asexual
teliospores are not. They are overwintering spores. Each stages imperfect stages. Since only the imperfect stages
cell of the teliospore begins with the dikaryotic (n+n) of this large group of fungi are known, they are called the
nuclear condition, but during winter, fusion occurs to form Fungi Imperfecti.
a zygote. Meiosis occurs in the warm, moist weather of The structure of the hyphae, which are septate, and of

spring. Then each cell of the teliospore puts out a single spores, suggest thatmany imperfect fungi may be
short hypha, the basidium (Fig. 12.28), and four Ascomycetes; others may be Basidiomycetes. The Fungi
basidiospores are extruded. No host is involved at this Imperfecti may be considered to be Ascomycetes or
stage; germination occurs at any moist surface where the Basidiomycetes whose sexual stages have not been
spore happens to land. With this event the fungus has observed or no longer exist. Occasionally, sexual stages
completed its cycle. are discovered in species that have previously been
The smuts are another group of basidiomycetes that classified as imperfects. When this happens, the fungus is
attack grain crops. Their hallmark the replacement of
is renamed, but usually the old name for the imperfect
grains by black masses of spores that have a sooty look; stages is kept too. For instance, the imperfect fungus
hence the name. Penicillium vermiculatum is now known to form asci similar
Smuts vary in their method of attacking the host. to those of the ascomycete genus Talaromyces; hence
Ustilago maydis spores cause local infection of corn plants this fungus is also called Talaromyces vermiculatum.
(Fig. 12.29). The mycelium stays near the point of Since the classification of imperfect fungi is very
infection and produces tumors where spores are formed. doubtful, the groupings are designated as form genera or
The spores are n+ n teliospores that survive the winter to form families to show that the members do not

chapter 12 |
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270
Figure 12.29 Smut disease of corn caused by Ustilago maydis.

necessarily have a natural or family relationship. For purified, these enzymes are important industrial
instance, some species have been named simply on the preparations. Aspergillus oryzae is used in the preparation

basis of the host upon which they were found. of ricewine and soybean sauces. Several species of
Only a single form group will be mentioned, the form Aspergillus are important in cheese manufacture. A

order Moniliales; it is the largest in the class and has over disease resembling tuberculosis is caused by Aspergillus

10,000 form species. fumigatus, and several other Aspergillus species cause
diseases in plants. Strains of Aspergillus flavus form
Penicillium and Aspergillus are probably the most
compounds called aflatoxins that are toxic to animals.
widespread fungi. They are the common blue, green, and
Penicillium notatum has won deserved fame because
black molds that occur on citrus fruits, jellies, and
penicillin, a drug derived from it, the growth of
will inhibit
preserves. Their conidia are in the air and soil
bacteria without injuring human Substances such
tissue.
everywhere.
as penicillin, formed by one organism and inhibiting the
In Penicillium, the spores are formed on profusely
growth of other organisms, are called antibiotics. Our
branched conidiophores (Fig. 12.30A). In Aspergillus, the knowledge of them dates back to 1870, but their
tip of the conidiophore swells and conidia form in long significance was not fully appreciated until the discovery
chains radiating from this swollen tip (Figs. 12.30B.C). of penicillin.
Enzymes secreted by these fungi are particularly active In 1928, Alexander Fleming investigated extracts of
in digesting starch and other carbohydrates. When Penicillium; after demonstrating their antibiotic properties,

conidia

conidia

conidiophores —

Figure 12.30 Diagrams of different forms of conidiophores. A,

Penicillium; B, C, Aspergillus; B, three-dimensional appearances;
C, optical section through 8.

Subdivision Eumycotina

271
 The Lichens
The lichens are composite plants composed of algae and
fungi. The algal components are generally single-celled
forms belonging to either the Chlorophyta or the
Cyanophyta. When free from the fungus, the algae may
exist normally.
The fungal component is generally an Ascomycete, but
algal associations with Basidiomycetes are also known.
The fungal component of the lichen cannot live separated
from the alga unless supplied with a special nutritive
medium.
The association of a fungus and an alga in the lichen is

generally believed to be symbiotic; that is, both the

fungus and the alga derive benefit from the association.
The alga furnishes food for the fungus, which supplies
moisture, shelter, and minerals for the alga.
A section through a lichen thallus reveals four distinct
layers. The top and bottom layers consist of a compact
mass of intertwining fungal filaments. The algal cells form
a green layer beneath the top mass of fungal filaments,
and a loose layer of hyphae lies directly below the algal
Figure 12.31 Nematode trapping loops of Dactylaria. (Redrawn
from E. A. Gauman, Fungi, A Description of Their Morphological cells (Fig. 12.32).
Features and Morphological Development, Hafner Publishing Lichens are widespread. They form luxuriant growths on
Company, New York, 1950.)
the frozen, northern tundras, where they supply feed
("reindeer moss") for animals. In the United States they
frequently cover rocks, trees and boards exposed to sun
and wind (Figs. 12.33AS.C, Color Plate 3). They are slow
he gave the antibiotic the name penicillin. It was not until but efficient soil formers; rocks are disintegrated slowly by
after 1940 that a concentrated effort was made to produce their action. Lichens are pioneer plants, appearing before
penicillin on a commercial scale, for use in World War II. any other plants on recent lava flows. Lichens and mosses
Many thousands of pounds of mycelium of Peniclllium accumulate soil and organic matter sufficient to allow
notatum have been grown, and the species has been herbs and later shrubs and trees to become established.
subjected to intensive study, yet no sexual stages have For convenience in classifying, lichens may be divided
ever been observed. The absence of sexual reproduction into three groups simply on the basis of their vegetative
presents the geneticists who are studying Penicillum body. This is a highly artificial grouping. Probably the
notatum with a difficult breeding problem. Nevertheless, most conspicuous lichens are members of the fruticose
they have succeeded in growing strains that produce type. They are formed by small tubules or branches.
many times more penicillin per pound than do the original Fruticose lichens may make extensive and attractive
strains. growths on oaks or the dead branches of certain pines
Several forms in the Moniliales are adapted to capture (Fig. 12.33C). Some lichens have a leaflike plant body.
and destroy microscopic animals. A few trap nematode
worms by forming small rings that quickly constrict when
stimulated by the contact of a nematode crawling through
them. A nematode-trapping form is shown in Fig. 12.31.
Since the sexual process has not been observed to
occur in the Fungi Imperfecti and genetic variation is not
thought to be a part of asexual reproduction, the question
may be asked: Is genetic variation possible in the Fungi
Imperfecti? The answer is, apparently, yes, at least in the
laboratory. Careful genetic analysis has produced
evidence that mycelia of imperfect fungi may sometimes
fuse and bring together nuclei of different origin. By
unknown mechanisms, nuclei may be formed genes
of
from each of the original kinds of nuclei. Meiosis appears
not to be involved; hence this way of forming recombinant
nuclei is called the parasexual cycle. It is well-established
in laboratory cultures where it may account for the great NOSTOC (an alga)
variability of some Fungi Imperfecti. It has not been
established that the process is significant in wild
populations. Figure 12.32 Cross section through a lichen thallus.

chapter 12 |
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272
They constitute the foliose type and are common in male reproductive cells and may have flagellated
moister climates, where their growths on tree trunks or on spores. The mycelium may be diploid. Many species
fallen logsare extensive (Fig. 12.33/4). However, the are aquatic. Some are important plant pathogens.
crustose lichens are the most common type, as they grow 5. The remaining classes of true fungi lack motile cells
extensively on rocks, bark, poles, boards, and other hard and are classified by their mode of sexual
surfaces. Indeed, any bright coloring on the dark rocks reproduction.
(Fig. 12.33S) and gray granites of our northern mountains, 6. The zygote fungi (Class Zygomycetes) mate by fusing
on unpainted wood, or on bark of trees is quite likely to be special hyphae. The zygote becomes a spore.
caused by a growth of crustose lichens. Sporangia are produced for asexual reproduction. The
mycelium is haploid, coenocytic, and is usually
terrestrial.Most are saprophytes.
Asexual Reproduction 7. The sac fungi (Class Ascomycetes) form meiospores
inside a sac (the ascus). Vegetative body may be
Lichens may multiply simply by the distribution of small unicellular (yeasts) or a multicellular mycelium that is
pieces of the vegetative thallus. Many lichens produce divided into cells by perforated cross-walls. The
special powdery bodies, composed of both a fungus and vegetative mycelium is haploid. Fusion of sexual
an alga, that serve as a means of vegetative reproduction.
hyphae produces dikaryotic (n+ri) hyphae that
These bodies are called soredia (singular, soredium). produce asci. Most species form their asci within an
Both the algal and the fungal components may produce ascocarp composed of many haploid and dikaryotic
spores.The algae may grow independently, but the if
hyphae. Asexual reproduction is usually by conidia.
young hyphae do not find the required algal cells when Most species are terrestrial saprophytes but some are
fungal spores germinate, they soon die.
dangerous plant pathogens.
8. The club fungi (Class Basidiomycetes) produce
meiospores on the surface of special cells, often club-
Sexual Reproduction shaped, called basidia. Mushrooms and brackets are
the fruiting bodies (basidiocarps) of fungi in the
Sexual reproduction is characteristic of the type of fungus
subclass Homobasidiomycetidae. The basidiocarps
present in the lichen. Since the fungal component is an
consist entirely of dikaryotic hyphae. Dikaryons form
Ascomycete in the great majority of lichens, ascocarps
by fusion of haploid hyphae. The dikaryotic condition
with asci and ascospores are formed. Upon the
is maintained by clamp connections. The vegetative
germination of the ascospore, the germ tube must find its
proper algal associate
mycelium may be haploid or dikaryotic. Most fungi in
in order to survive.
this subclass are terrestrial saprophytes. The rusts and
smuts (Subclass Heterobasidiomycetidae) lack
Summary basidiocarps. They are plant parasites and may be
1. The fungi (Division Mycota) are heterotrophic serious pathogens. Some of them use two plant
eukaryotes that lack photosynthesis; typically species as alternate hosts at different stages of the life

reproduce by spores; and have cell walls during at cycle.

least part of thelife cycle. About 200,000 species are 9. The imperfect fungi (Class Fungi Imperfecti) have no
known. known sexual reproduction. Reproduction is asexual,
2. Slime molds (Subdivision Myxomycotina) lack cell usually by conidia. Most form a septate terrestrial
walls except in spores. The vegetative body has mycelium. This class includes the most common
amoeboid motion and engulfs solid food particles. household molds and the fungi that produce penicillin

3. True fungi (Subdivision Eumycotina) have cell walls and fine cheeses.
throughout the life cycle and feed only by absorbing 10. The lichens are symbiotic associations in which a
dissolved solutes from solution. Some are unicellular fungus and an alga produce a joint body. Widely
but most form a vegetative body (the mycelium) varied forms occur. Reproductive structures include
consisting of branched filaments (hyphae). Five separate fungal and algal spores as well as bodies
Classes of true fungi are discussed. (soredia) that include both alga and fungus. Some
4. The egg fungi (Class Oomycetes) have flagellated lichens are important pioneers on bare rock.

Summary

273
 13 CHAPTER

13
parenchyma tissue occur in some
The division Bryophyta includes hornworts, mosses,
and liverworts. Perhaps the two greatest differences exception of one stage in the life
forms. With the
history of mosses, the
between algae and bryophytes are the general bryophyte plant body is never filamentous. It is composed
structure of the plant bodies and the occurrence of of blocks or sheets of cells that form a parenchymatous
multicellular gametangia. It will be recalled that the plant Another important difference is that the sporophyte
tissue.

body of thallophytes is generally composed either of single generationin the bryophytes is attached to the

cells (or filaments of cells), or of intertwining filaments, gametophyte and it remains there throughout its life time.
resulting in a more or less simple body. Layers of

Table 13.1 General Characteristics

Tabulation of Differences between Thallophytes and
Bryophytes
The comparative characteristics of the thallophytes and
Thallophytes Bryophytes bryophytes are shown in Table 13.1 The multicellular
female gametangia characteristic of all members of the
1. Mostly aquatic 1. Mostly terrestrial, but Bryophyta produce but a single gamete, the egg, which is
moist habitat surrounded by a layer of protecting cells. Male gametangia
of the bryophytes produce numerous sperms, but they are
2. A few of the kelps have 2. Some mosses have cells
elements that resemble suggesting sieve
always surrounded by a protective layer of parenchyma
cells;

sieve cells otherwise food conducted

cells, called a sterile jacket.

in relatively
Water is required by bryophytes, as in the algae, to

undifferentiated cells
effect fertilization. A definite alternation of generations
occurs in all members of the Bryophyta, with the
3. None have water- 3. Simple water-conducting
sporophyte (diploid) generation always more or less
conducting elements cells (not vessels or
dependent on the gametophyte (haploid) generation. In all
tracheids)
Bryophyta, an embryo sporophyte, consisting of a
4. In general, no specialized 4. Rizoids anchor and spherical or elongated mass of tissue, is formed directly
water- and nutrient- absorb water and mineral after the germination of the zygote.
absorbing tissue; rhizoids nutrients Asexual spores are not formed in Bryophyta. Some
and haustoria may occur members of this division reproduce asexually either by
in some fungi fragmentation or by multicellular units called gemmae. A
5. Mostly filamentous or a Only one stage of mosses haploid plant, producing gametes, alternates with a diploid
lacework of intertwining is filamentous; all others plant; meiosis occurs in the diploid plant, and haploid
filaments; parenchyma in are formed of spores containing n sets of chromosomes in each
1

a few parenchyma cells nucleus are formed. Thus, a haploid gamete-producing

6. A definite alternation of Allhave an alternation of plant —the gametophyte —
regularly alternates with a

generations in many heteromorphic diploid spore-producing plant —

the sporophyte, as shown
forms generations below.

7. Both sporophytes and Gametophyte

gametophytes nutritionally independent; sporophyte
independent nutritionally dependent
8. Gametangia are either Gametangia are always
single cells or groups of composed of gamete-
single cells not producing cells covered
accompanied by a jacket by a jacket of sterile
of sterile vegetable cells vegetative cells

9. Spores often water- Spores wind-dispersed

dispersed

274
Economic Importance Class Musci
Bryophytes occur on soil, trunks of trees, and rocks; many
species are truly aquatic. They are not extremely important General Characteristics
economically, but they do have an interesting if short,
ethnohistory. Native American Indians in Utah, for Although the mosses are small plants, they are
instance, ground up mosses, such as Mnium and Bryum, nevertheless conspicuous. They frequently cover rather
into a paste and applied it as a poultice to treat burns and large areas of stream banks.They grow on rocks and
bruises. In Europe, the growth of mosses such as and are sometimes submerged in streams. Although
trees,
Dicranoweisia was encouraged on shingle roofs to make some mosses are able to resist considerable drought,
them watertight. (Figs. 13.1, 13.2A 13.5) all require moisture for active
Sphagnum is by far the most important moss growth and reproduction.
economically. The surface layers of stems and "leaves" of Mosses as a class show great structural uniformity.
Sphagnum are composed of alternating files of living green Funaria and Mnium and Sphagnum are examples (Figs.
cells and large empty hyaline cells (Fig. 13.2). The hyaline 13.1, 13.2A, 13.5). Gametophytes of nearly all species
cells are capable of absorbing water or other fluids in have two growth stages: (a) a creeping, filamentous stage
great quantities as high as 20 times their own weight. In (the protonema) (Fig. 13.1), from which is developed
the 1880s Germany was discovered, quite by
in it
(b) the moss plant with an upright or horizontal stem
Sphagnum moss pads make excellent
accident, that bearing small, spirally arranged green leaves (Fig. 13.5).
absorbent bandages. A workman in Kiel lacerated his arm
while working in a peat moor. Since he had no other

bandage material, he packed his wound with dry peat

(Sphagnum). Ten days later, when he was able to obtain Gametophyte
treatment, the wound had healed. After that, disinfected
bags filled with dry Sphagnum became a common wound Germinating spores of mosses do not develop directly into
dressing in Germany. The use of Sphagnum as wound a leafy gametophyte but first become a filamentous
dressings was not popular outside Germany until World structure. This early stage ofgametophyte is called the
War At that time they were used on a large scale by the
I.
protonema (Fig. 13.1). The protonema is not a permanent
Allied Armies as well. The British, for example, used structure, although it may branch considerably under
1,000,000 Sphagnum dressings each month. Sphagnum favorable conditions and cover a rather large area of soil,
dressings were used very little in the United States, but the sometimes forming a green coating resembling algal
American Red Cross did publish inquiries and instructions growth. Cells composing the protonema contain numerous
on finding useful Sphagnum. More recently, Sphagnum is chloroplasts (Fig. 13.1). At various locations, cell divisions
still used as packing for vegetables, in potting mix to along the protonema produce swollen regions called
increase the water holding capacity of the soil; and old buds. From them develop the upright gametophyte stage.
compressed Sphagnum is used as a fuel in the form of a
The mature stem of the leafy gametophyte shows a
low grade coal known as peat.
wide range of differentiation, depending on the species,
age, and environment. Elongate cells called rhizoids
anchor the gametophyte to the substrate (Figs. 13.1, 13.5,
13.7). These structures are not roots, and apparently not
Classification
involved in absorption. The simpler mosses, like Tetraphis

The Bryophyta are divided into three classes: Hepaticae (Fig. 13.2E) have an outer layer of thick-walled epidermal
cells (stereids), surrounding an undifferentiated cortex
(liverworts), Anthocerotae (hornworts), and Musci
(mosses). The Hepaticae are the simplest and perhaps the made up of parenchymalike cells. Epidermal cells of the

most primitive bryophytes. They consist of a flat, peat moss Sphagnum (Figs. 13.28 to 13. 2D) are large and
ribbonlike, green thallus that produces gametes and a empty, with pores that open to the outside.
meiospore-producing capsule or sporangium usually Some mosses, such as Mnium (Figs. 13.3A, 13.36,

embedded within thegametophyte thallus. The name 13.4) have a central conducting strand in their stems. It is

liverwort is very old, having been used in the ninth made up of elongated, thin-walled hydroids, which are
century. It was probably applied to these plants because dead, empty cells that conduct water (Figs. 13.36, 13.4).
of their fancied resemblance to the liver and the belief (the Their end walls are oblique and sometimes very thin,
"Doctrine of Signatures") that plants resembling human perforated with pores, or partly dissolved. Experiments
organs would cure diseases of the organs they resembled. with dyes show that translocation of water can occur in

At any rate, a prescription for a liver complaint in the these cells. Some moss "leaf" midribs also contain
1500s called for "liverworts soaked in wine." hydroids, but only in one genus are these known to

Mosses are the best-known class of Bryophyta. They connect with the hydroids of the stem. Hydroids show
have simple "stems" and "leaves," and sporophytes are some resemblances to tracheids, but lack specialized
borne on green gametophytes. They have either an pitting and lignified walls. No lignin has been detected in

upright or prostrate leafy gametophyte, and may form bryophytes.

extensive mats on moist, shaded soil. The sporophyte is A few of the most specialized mosses also contain cells
raised above the leafy gametophyte. resembling the sieve cells of vascular plants. Between the

Class Musci

275
Figure 13.1 Funaria. A, germination of spores; 6, protonema; C,
protonema with bud.

epidermis and central strand, elongated cells with oblique contain chloroplasts that become orange-red when the
end walls (leptoids) may occur (Fig. 13.4). These cells are antheridium ripens. The sperms consists mainly of an
alive, but their nuclei are degenerate and inactive. They elongated nucleus and each has two flagella. The
have many plasmodesmata in their end walls, and callose antheridia are surrounded by club-shaped, multicellular,
may be present. Studies show that sugars may be sterile hairs (paraphyses) with conspicuous chloroplasts.
translocated through these cells. The archegonia have a long neck, a thickened middle
It is important to emphasize that: (1) most mosses do portion called the venter, which contains the egg, and a
not contain leptoids; (2) there are important differences long stalk (Fig. 13.6Q. When sufficient moisture is
between hydroids and tracheids, and between leptoids and present, sperms are extruded from the antheridium. The
is as yet no firm evidence to show
sieve cells; (3) there neck cells of the archegonium separate, opening a
these cells are phylogenetically related to conducting cells passage to the egg. Sperms swim down, possibly
of vascular plants. responding to a chemical signal, and one fuses with the
egg. Mosses are dependent on free water for fertilization.
Sexual Reproduction

The gametophytes of many mosses are monoecious Sporophyte

(homothallic); that is, both antheridia (sperm-producing)
and archegonia (egg-producing) are produced by the Soon after fertilization, the zygote begins to develop into a
same gametophyte (Fig. 13.5). Some mosses are spindle-shaped embryo that differentiates into a
dioecious (heterothallic); in other words, antheridia and sporophyte consisting of a foot, seta, and sporangium or
archegonia are produced on separate gametophytes. In capsule. The foot penetrates the base of the venter and
Mnium, shoots bearing antheridia are easily recognized, grows into the apex of the leafy shoot. It absorbs water
by the surrounding leaves that spread around them and nourishment from the gametophyte for its growth and
somewhat like petals of a flower. The group of antheridia development. The seta elongates rapidly, raising the
appears as an orange spot in the center of the terminal sporangium 1 cm or more above the top of the leafy
cluster of leaves (Figs. 13.5, 13.6). gametophyte (Fig. 13.7/4). The old archegonium increases
Antheridia of most true mosses (Fig. 13.66) are in size as the sporophyte enlarges. When the seta
enclosed by cells forming a sterile jacket. These cells elongates, the top of the expanded archegonium is torn

chapter 13 |
Bryophytes

276
Figure 13.2 A, habit of sporophyte plant of Sphagnum
papillosum, x2. 6, external view of stem showing large hyaline
cells, x450. C, cross section of branch stem showing external
hyaline cells and thick-walled cells of central axis, x590. D, leaf
cells; note the narrow chlorophyll containing cells and the larger
outside -«- -*~ center
hyaline cells with peculiar bandlike thickenings, x330. E, cross
section of Tetraphis pellucida stem to show its simple structure, Figure 13.4 Partial stem portion showing "phloemlike" leptoids
x70. and "xylemlike" hydroids.

central
region

Figure 13.3 Longitudinal sections through gametophyte stems

of mosses. A, stem of Mnium showing central strand of
sclerenchymalike cells, x 100; B, cross section of Mnium seta (1)
thick-walled stereids; (2) thin-walled cortex; (3) central region of
partially collapsed hydroids, x 72.

Class Musci

277
 paraphyses

antheridial head, archegonial head

lage leaves

Figure 13.5 Mnium, showing location of gametangia, x1

Figure 13.6 Microscopic views of moss gametangia. A,

antheridial heads of Polytnchum, x 1 S, antheridial head of
,

Mnium, x 35; C, archegonium surrounded by paraphyses

(Mnium), x50.

chapter 13 |
Bryophytes

278
 capsule

spores

leafy gametophyte

occur in the epidermis covering the lower half of the

sporangium. The sterile tissue forming the inner portion of
the sporangium may conveniently be divided into three
regions, each of which may be recognized by the type of
cells comprising it. The fourth region, composed of
sporogenous cells, forms a layer around the columella
(Fig. 13.7S). The cells of the sporogenous tissue
(sporocytes) may increase in number by mitosis; each cell

contains the diploid number of chromosomes. Meiosis

takes place next, and haploid spores containing ^n
chromosomes result. They are the first cells of the
gametophyte generation.
The columella projects upward, forming a small dome
above the main mass of the sporangium. The four or five
outer layers of cells of this dome differentiate into a dry,
Figure* 13.7 Mnium, showing gametophytes with attached
A, brittlecap called the operculum (Figs. 13.76, 13.8). The
sporophyles. B, median section through a mature but unopened
cells immediately beneath the operculum form a double
capsule of Mnium, x 25.
row of triangular peristome teeth (Figs. 13.88.C). The
broad bases of the teeth are attached to the thick-walled
deciduous cells that form the annulus around the upper
from its point of attachment to the gametophyte and end of the sporangium. When the sporangium matures
elevated with the sporangium. The upper end of the old and becomes dry, thin-walled cells of the annulus break
archegonium, now known as the calyptra, remains for a down, allowing the operculum to fall away and expose the
time as a covering for the sporangium (Fig. 13.7A). peristome teeth. By this time, most of the thin-walled cells
The sporangium of mosses (Figs. 13.7, 13.8) may within the columella have collapsed and the cavity thus
measure 1 to 3 mm in diameter and 2 to 6 mm long. It is formed is filled with a loose mass of spores. Peristome
surrounded by an epidermal layer composed of cells teeth are rough and are very sensitive to the amount of
similar to those in the epidermis of higher plants. Stomata moisture in the 13.86,Q. When they are wet or
air (Figs.

Class Musci

279
 annulus spores
annulus

operculum

capsule

nsrome

Figure 13.8 Scanning micrograph of a capsule of Funaria. A,

capsule with operculum in place; B, operculum has been shed,
peristome teeth partially open, spores attached to them; C,
peristome teeth fully open, capsule empty, x 300.

the atmospheric humidity is very high, they bend into the

cavity of the sporangium; when
dry, they straighten and lift
out some of the spores, which are then disseminated by
airmovements. If a spore comes to rest on moist soil and
if and temperature are favorable,
illumination it germinates 10. The embryo develops into a sporophyte consisting of a
and grows into a protonema. and sporangium. The sporangium contains
foot, seta,

sporogenous tissue and several types of sterile tissues,

Summary of Moss Life History and among which are the operculum, annulus, and
Characteristics peristome.
11. Sporocytes, each containing 2n chromosomes, form
1. The gametophyte consists of (a) a filamentous,
from sporogenous cells.
branched, algal-like structure called a protonema, and 12. Sporocytes undergo meiosis. Four spores result from
(b) leafy shoots that develop from buds on the
each sporocyte
protonema. The shoots consist of a stalk bearing
13. When spores germinate, they form the protonema.
rhizoids at its lower end and leaves throughout its
14. In the central strand of some mosses, eg, Sphagnum
length. The gametophyte
is green and able to
and Mnium, there are specialized cells called hydroids
synthesize food. gametophyte nuclei are haploid.
All
thatmay be involved in water conduction.
2. Antheridia and archegonia develop at the apex of the
15. Cells specialized for sugar transport, called leptoids,
leafy gametophyte.
are also found in some mosses.
3. Each antheridium produces hundreds of sperms.
4. A single egg is formed in the venter of each
archegonium.
5. Gametes (eggs and sperms) each contain 1 n Class Hepaticae
chromosomes.
6. When sufficient moisture is present, mature sperms General Characteristics and Distribution
are extruded from the antheridium. The neck of the
mature archegonium is open and sperms swim down it Hepaticae are commonly known as liverworts. There are
to theegg in the venter of the archegonium. some 8500 species of liverworts. The great majority of
7. The egg is fertilized by one sperm. them grow in moist, shady localities. The gametophyte is
8. The zygote, as a result of the union of gametic nuclei, the prominent plant, and the sporophyte is partially or

is diploid. wholly dependent on the gametophyte. The gametophyte

9. An embryo sporophyte, consisting of a spindle-shaped is green and may grow either as a flat ribbon or as a leafy
mass of undifferentiated cells, partially dependent on shoot. In either event, the plant body is frequently called a
the gametophyte, develops from the zygote. thallus, even though its internal structure does not

chapter 13 |
Bryophytes

280
 Antheridia. Antheridia of different representatives of the
Bryophyta are similar, though they may vary somewhat in

shape. In Riccia, they are pear-shaped and composed of

two types of cells, (a) fertile and (b) sterile (Fig. 13.108).
Fertile cells give rise to sperm, which are relatively
numerous, small, and dense with protoplasm. Sterile cells
form a protective jacket, one cell in thickness, around the
fertile cells. The antheridia are connected to the

gametophyte by a short stalk (Fig. 13.10A).

Mature sperm consist mainly of an elongated nucleus
with two long flagella. They may be shot with considerable
force from the mature antheridium or they may be
extruded slowly in a single mucilagenous mass. In any
event, they do not leave the antheridium until enough
moisture is present to allow them to swim about.

Archegonia. The archegonium of Riccia is a flask-

shaped structure consisting of two parts, (a) an expanded
basal portion, the venter, and (b) an elongated neck
(Figs. 13.10CD). Four cover cells are located at the top
of the neck. Each archegonium contains a single egg cell,
Figure 13.9 Habit of Riccia, x1.
which is located in the venter. A short stalk attaches the
archegonium to the gametophyte. Archegonia of a similar
structure are found in other Bryophyta.
correspond to that of the thallophytes. Of the four orders Shortly before the egg cell is mature, the cover cells
in the Hepaticae, we shall consider briefly the separate. At the same time, the cells in the center of the

characteristics of two genera, Riccia and Marchantia, both neck dissolve, so that an open canal connects the venter
belonging to the order Marchantiales. with moisture outside the archegonium. Free-swimming

Gametophytes of the Marchantiales are small, green, sperms move toward certain chemical substances formed
ribbon-shaped plants (Figs. 13.9, 13.11). They branch by the archegonium. Several sperms may enter the
regularly by a simple forking at the growing tip, resulting in archegonium, but only one sperm fertilizes the egg in the
a number of Y-shaped branches. In some species, a venter.

rosette may be formed. The upper surface of the thallus is

composed of cells adapted for photosynthesis and Sporophyte
arranged to form air chambers that open to the surface by
As a result of fusion of sperm and egg nuclei, the zygote
a pore (Figs. 13.10A 13.12C,D). Several types of storage
contains the diploid, or 2n, set of chromosomes. Mitosis
cells generally make up the lower surface. Rhizoids,
now proceeds in a more or less orderly manner until a
specialized elongated cells, extend downward from the
spherical mass of some 30 or more similar cells is formed
lowermost layer of cells and anchor the gametophyte to
(Fig. 13.10D). These cells are partially dependent on the
the substrate. The thallus is thus differentiated into distinct
gametophyte. This mass of undifferentiated cells
upper and lower portions. Scales, frequently brown or red,
comprising the young sporophyte is an embryo. is It
are also formed on the lower surface (Figs. 13.107*).
located within the venter of the archegonium.
Further development of the embryo involves
differentiation of an outer layer of cells to form a protective
Riccia jacket of sterile tissue surrounding a mass of cells that are
capable of forming spores. This spore-forming tissue is
Riccia is a widely distributed genus, and although it
called sporogenous tissue. The sporogenous cells
requires water for active growth, most species can tolerate
continue to divide by mitosis until many have formed.
considerable drought. Several species are aquatic,
These cells, now known as sporocytes, will divide by
growing either on mud or on the surface of small ponds.
meiosis to produce four spores with the haploid

Gametophyte chromosome number. Remember that meiosis involves two

cell divisions and results in a reduction in the
The gametophyte is a small green thallus, frequently chromosomes by one half (see Chapter 3).
forming a rosette (Fig. 13.9). Tissue on the lower side is The mature sporophyte of Riccia (Fig. 13.10E) is called
composed of colorless cells that may contain starch. a capsule. is composed of a jacket of sterile cells
It

Tissue on the upper side consists of vertical rows of enclosing a mass of spores. The sporophyte wall breaks
chlorophyll-bearing cells, between which are air chambers down when the spores are mature. These spores remain
(Fig. 13.10/4). The gametangia are embedded in deep, embedded in the gametophyte thallus and are not released
lengthwise depressions or furrows, on the upper surface until after the death and decay of the gametophyte. As

and archegonia are usually found

of the thallus. Antheridia each spore germinates, it grows into a typical
on the same gametophyte (homothallic). gametophyte plant.

Class Hepaticae

281
 neck canal

Figure 13.10 Riccia. A, cross section through a thallus:

gametangia are on the upper surlace of the thallus between the
photosynthetic filaments or in a notch in the thallus, x8. 8, an
antheridium; note jacket of sterile cells surrounding the
developing sperm cells. C, archegonium on thallus, neck canal
open, egg cell in venter. D, young sporophyte developing in
venter of old archegonium. E, sporophyte with spore mother cells
still enclosed in venter of old archegonium, F, meiosis occurs

within the sporophyte. G, the venter now contains fragments of

the old spore case (sterile cells of the sporophyte) and spores. H,
a spore. B through G, about x50; H, x100.

chapter 13 |
Bryophyles

282
<z==w

Class Hepaticae

283
 spore

Figure 13.12 Life cycle diagram and habit of Marchantia

polymorpha. A, cross section view of thallus. Note the pores and
air chambers containing columns of chlorophyllous cells. B, habit
of male (left) and female (right) thalli with antheridiophores and
archegoniophores respectively. C, section through head of
antheridiophore to show antheridia. D, section through head of
archegoniophore to show archegonia. E, enlarged view of
antheridium. F, enlarged view of an archegonium. G, immature
sporophyte plant. H, mature sporophyte composed of foot, seta,
and sporangium. sporangium releasing spores and elaters. J,
/,

spore germination to form new gametophyte thallus. K, asexual

reproduction by gemmae.

chapter 13 |
Bryophyles

284
 1

Significant Steps in the Life History of chloroplasts. As in Riccia, the lower surface of the thallus

Riccia is composed of colorless cells, some of which are

modified for storage. Rhizoids, which anchor the thallus,
1. The gametophyte, a plant, absorbs
small, green, flat grow from the cells covering the lower surface. Several
water and mineral salts from the and carbon soil rows of scales are also attached to this lower surface (Fig.
dioxide from the air. It contains chlorophyll and can 13.12L).
synthesize food. All gametophytic nuclei are haploid
(contain "In set of chromosomes). Asexual Reproduction. Marchantia reproduces
2. Gametangia (antheridia and archegonia) develop in a asexually in two ways: (a) Older parts of the thallus die
furrow on the upper surface of the gametophyte. and younger portions, no longer attached, develop into
3. Each antheridium produces thousands of sperms. new individual plants; and (b) small cups, known as
4. A single egg is formed in the venter of each gemmae cups, form on the upper surface, and small
archegonium. disks of green tissue, called gemmae, grow from the
5. Gametes (eggs and sperms) contain 1 n set of bottom of these cups (Figs. 13.1 1 S 13.12/vf). Gemmae,
chromosomes. when mature, break off from the thallus and are distributed
6. When sufficient moisture is present, mature sperms into nearby areas. Raindrops are often the agents that
are extruded from antheridia. The neck of the mature break off gemmae and scatter them away from the thallus.
archegonium is opened and sperms swim down to the New gametophyte plants grow from gemmae.
egg in the venter.
7. Each egg is fertilized by a single sperm. Sexual Reproduction. Marchantia is heterothallic; that
8. The zygote, as a result of the union of two haploid is, gametophytes have either antheridia or archegonia.

gametic nuclei, is diploid.

The gametangia, however, are very similar in structure to
9. An embryo sporophyte, consisting of a spherical mass those of Riccia. Antheridia are pear-shaped bodies
dependent on the
of undifferentiated cells, partially composed of a jacket of sterile tissue surrounding sperms
gametophyte, develops from the zygote. Each nucleus (Fig. 13.128,C,D). They are borne on disks raised above
of the embryo sporophyte is diploid (2n). the thallus on slender stalks, called antheridiophores
10. The cells of the embryo differentiate into a sporangium (Figs. 13.11C, 13.12A). Antheridiophores are modified
that consists of a jacket of sterile cells enclosing a portions of the thallus having furrows, rhizoids, and air
mass of fertile cells. chambers. Antheridia develop in cavities on the upper
11. Sporocytes, each with 2n nuclei, form in sporangia. surface of the disk, the youngest antheridia being close to
12. Sporocytes undergo meiosis. A quartet of spores forms the outer margin of the disk (Fig. 13.12B). Mature sperms
from each sporocyte. All sporocytes are 2n and spores are extruded in a mucilagenous mass.
are 1 n.
Archegonia are flask-shaped and have the same
13. Upon death and decay of old gametophytes, spores structures as those of Riccia —
venter, neck, and cover
are liberated from the capsules and new gametophytes cells (Figs. 13.1 1 D, 1 3.1 2G,H). They are borne on
from them. specialized branches called archegoniophores. The disk
at the top of the archegoniophore typically has eight or
Marchantia more lobes. In development, the lobes bend downward
Marchantia may grow in large mats on moist rocks and and then grow inward toward the stalk. Thus, the
youngest portion of an older disk bearing archegonia is on
soil inshady locations. It is widely distributed and fairly
the underside and close to the stalk (Figs. 13.1 E,
common on banks of cool streams. It is somewhat better
13.12E.F). Archegonia develop in the lower surface of the
adapted to growing on land than is Riccia, but
considerable moisture is still required for active growth
disk but in tissue similar to that found on the upper
surface of the thallus. Fingerlike processes grow out from
and for fertilization. For all references below to Figure
13.11, see Color Plate 3.
between the lobes. The archegonia mature, and
fertilization takes place when the disk of the
Gametophyte archegoniophore is but slightly elevated above the thallus.

The gametophyte of Marchantia differs from that of Riccia

Sporophyte
in gametangia on special disks are raised some
that
distance above the vegetative thallus (Figs. 13.1 1 C.D, Zygotes develop, as in Riccia, into diploid embryos, which
13.12AE). The thallus of Marchantia is similar to that of are spherical masses of undifferentiated, dependent tissue
Riccia, but much coarser. It is strap-shaped, with a (Fig. 13.12H,/). Subsequent growth, however, is more
prominent midrib and shows dichotomous branching (Figs. complicated than in Riccia. Some cells of the embryo
13.1 1 AS)- The tips of the branches are notched. An divide to form a mushroomlike growth that becomes
individual thallus may be 1 to 1.5 cm broad. Their length embedded in gametophyte tissue. This growth is called the
(1 to 1 .5 cm) and the degree of branching depend on foot. Cells in the central portion of the embryo form a
growth conditions. It is usual for new growth to overlap seta. The remaining lower cells develop into a
the older decaying ends of adjacent thalli. On its upper sporangium (Fig. 13.12/-/,/). Lengthening of the seta
surface are polygonal areas, with a small but conspicuous suspends the sporangium below the disk. While this
pore in the center (Figs. 13.12L). These areas, with their development is taking place, the stalk of the
air pores, mark the outlines of air chambers, each of archegoniophore elongates, lifting the disk with its
which is filled with short filaments of cells containing archegonia well above the thallus (Figs. 13.11D.E, 13.12/).

Class Hepaticae

285
 Fertilization stimulates enlargement ot the old
Class Anthocerotae
archegonium, which keeps pace with the enlargement of
the developing sporophyte. As a result, the sporophyte is
continually enclosed within the archegonium, which, The Anthocerotae have the simplest gametophytes of the
because of its increase in size and change in function and Bryophyta. They are small, green thallus plants with little
shape, is now known as the calyptra. In addition, the internal differentiation of vegetative tissues. They are
surrounding gametophyte tissue produces two other slightly lobed, with numerous rhizoids growing from the

envelopes that protect the sporophyte. lower surface (Fig. 13.13). The antheridia are similar in
The mature sporophyte (Fig. 13.12/) is composed of structure to those encountered among the Hepaticae (Fig.
three parts, foot, seta, and sporangium. The foot is an 13.14A). They are located in roofed chambers in the
absorbing organ. The seta serves to lower the sporangium upper portion The archegonia are
of the thallus.

away from the archegonial disk and thus to facilitate embedded within the thallus and are in direct contact with
distribution of spores. Before meiosis, the sporangium or the vegetative cells surrounding them (compare Figs.
capsule composed of a jacket of sterile cells
is 13.10C, 13.148).
surrounding a mass of sporocytes, among which are a The sporophyte (Figs. 13.13, 13.15) of the Anthocerotae
number of sterile elongated cells (Figs. 13.1 2/, J). Four is in striking contrast to those of the Hepaticae. The
spores, each containing 1 n chromosomes, develop from subepidermal cells contain chloroplasts, and typical

each sporocyte. The elongated sterile cells are stomata are found in the epidermis. A
embedded in
foot

transformed into spiral elements, called elaters, that the thallus serves as an absorbing organ. The sporangium
change shape under the influence of varying moisture is an upright elongated structure. Sporogenous tissue

conditions. Their twisting motion as they imbibe and lose forms a cylinder parallel with the elongated axis of the
water aids in dispersal of spores (Figs.13.12/,J,K). sporangium. Spores mature in progression from the top
Dissemination of spores is aided by two structural down. A meristematic region (i.e., region of continuous

features not found in Riccia: (a) the sporangium of the cell division) lies just above the foot and continually adds
sporophyte hangs from the lower side of the raised new cells to the base of the sporangium. Its presence
archegonial disk (Figs. 3.1 1E), and (b) elaters help to means that spores may be produced over long periods.
empty the spore case. Spores develop immediately into Under exceptionally favorable growing conditions, the
new gametophytes they land in an appropriate habitat.
if
sporophyte may lengthen greatly. Some sporogenous
tissue at the base of the sporangium may be replaced by a
conspicuous conducting strand. The foot enlarges and,
Significant Steps in the Life History of through decay of the gametophyte, comes into more or
Marchantia less direct contact with the soil. Such sporophytes are
1. The green gametophyte thallus absorbs water and capable of surviving independently for some time.

mineral salts from soil and carbon dioxide from air.

2. Gametangia (antheridia and archegonia) are borne on
upright branches called antheridiophores and
archegoniophores.
3. The zygote is diploid (2n).
4. The embryo sporophyte, consisting of a spherical
mass of undifferentiated cells, is formed from the
zygote.
5. Cells of the embryo sporophyte differentiate into foot,
seta,and sporangium.
6. The foot is an absorbing organ. is embedded in the It

gametophyte, from which receives nourishment. it

7. The sporangium consists of a jacket of sterile cells

sporophyte
surrounding a mass of sporogenous tissue
interspersed with elongated sterile cells.
8. The seta is a stalk that, in lengthening, lowers the
sporangium below the surface of the archegonial disk.
9. Repeated mitotic cell divisions in sporogenous tissue
result in sporocytes each one of which contains 2n
chromosomes.
10. Sporocytes divide by meiosis into tetrads of spores;
each spore contains 1 n chromosomes.
11. The elongated sterile cells are transformed into spiral
elaters.
gametophyte

12. The presence of elaters and the hanging position of

the sporophyte aid in disseminating spores.
13. Spores germinate immediately into new gametophyte Figure 13.13 Anthoceros gametophyte with upright, dependent
plants. sporophyte, x3.

chapter 13 |
Bryophyles

286
 antheridial chamber

antheridium

sperm cells

gametophyte

ventral canal
ce

egg ce

Figure 13.14 Section through an antheridium (A) and an

archegonium (B) ot Anthoceros, x 50.

elaters

spore
tetrads

sporophyte

Figure 13.15 Longitudinal medium section through sporophyte

of Anthoceros.

Class Anthocerotae

287
 14 CHAPTER

14 lower
vascular plants

contrast to algae, tungi, liverworts, and mosses, the roots. The vascular tissue is simple (Fig. 14.2), consisting
Insporophytes possess a well-
ot vascular plants of poorly developed phloem and xylem. An endodermis,
developed vascular system that serves to conduct with a conspicuous Casparian strip, separates the central
water, mineral salts, and food. There are 10 divisions of vascular tissue from the outer cortex. The xylem contains
vascular plants; six are seed-bearing and four do not bear spiral and scalariform tracheids.
seeds. The latter four divisions, which together make up Sporangia are borne in axils of some of the scalelike
the lower vascular plants, are the Psilophyta, Lycophyta, leaves (Fig. 14.1). Meiosis occurs in the sporangia,
Sphenophyta,* and Pterophyta. forming tetrads of meiospores. When the meiospores are
The number of families and genera in these divisions is ultimately released from the sporangia, they may land in a
small, and their members do not form a very conspicuous suitable habitat, germinate, and divide by mitosis
part of the current land flora. The Pterophyta is currently repeatedly to form a gametophyte.
the largest of the four divisions; it comprises the ferns, Psilophyta gametophytes are little more than a
which are prominent in certain cool, moist habitats. nonphotosynthetic axis of parenchymatous tissue several
However, plant fossils indicate that there was a period in millimeters long (Fig. 14.3). Xylem and phloem are rarely
the earth's history when the members of the Lycophyta, present. Gametophytes may grow on the surface of soil,
Sphenophyta, and Pterophyta formed a large and rock, or bark, or may grow within the soil. Nutrients are
dominant flora (Chapter 10). provided via a symbiotic or parasitic relationship with a
Although they are land plants, they still require free fungus, which ramifies throughout the tissue.
water for fertilization. The sporophyte is the dominant Gametangia are scattered over the surface of the
generation and — except youngest stages in the
for the gametophyte: flask-shaped archegonia with eggs, and
formation of an embryo —
it is independent of the spherical antheridia with spirally coiled, multiflagellate
gametophyte. Gametophytes are small, and some may sperm (Fig. 14.4).
lack chlorophyll. In its simplest form, the plant body of the After a sperm swims through free water and reaches an
sporophyte is an axis. Meristematic tissue terminates the egg, fertilization occurs and a diploid zygote cell begins
shoots and roots. Most lower vascular plants are dividing. The young sporophyte (embryo) consists of a
herbaceous perennials, but some ferns reach tree stature shoot-root axis attached to the gametophyte by a foot.
and have some woody tissue. While the shoot-root portion is assuming form, the foot
enlarges by repeated cell divisions, sending haustorial
outgrowths into the gametophytic tissue. The foot, by
virtue of its and organization, is well suited for the
position
Division Psilophyta functions of anchorage and absorption until the shoot-root
axis becomes physiologically independent and a mature
sporophyte results.

This division is represented in the existing flora by a single

family, the Psilotaceae, with two genera, Psilotum and
Tmesipteris. They are rare plants found mainly in the Division Lycophyta
tropics, one form of Psilotum growing as far north as
Florida. Figure 14.1 shows the simple plant body of
Psilotum nudum. Roots are not present and leaves The Lycophyta are well-represented in the northern
(microphylls) are small and scalelike. These are not "true" hemisphere by three genera: Lycopodium, Selaginella, and
leaves (megaphylls, see Chapter 10); they are instead Isoetes. We shall consider only the first two genera.
more like epidermal outgrowths. The branching, upright There are many fossil Lycophyta belonging to several
stem is slightly flattened and contains chlorophyll. A orders, Lepidodendrales being one of the best known.
fungus is always associated with the branched rhizome, Now-extinct representatives of this order were forest trees,
which is clothed with many rhizoids. There are no true some of which bore seeds and possessed megaphylls
(Chapter 10).
* Wechoose to use the old designation, Sphenophyta, for the The leaves of the living members of this division are
division to which the horsetails belong, because the new
generally small and usually arranged in a spiral. Leaf gaps
proposed designation, Arthrophyta, might be confused with
Anthophyta or Arthropoda. are never formed at the junction of stem and leaf in the

288
 -

epidermis
parenchyma
erenchyma

parenchyma y
endodermis

sporangium

parenchyma -

epidermis

Figure 14.2 Cross section of Psilotum stem. A, diagram to show

the arrangement of tissues. B, enlarged sector showing cellular
detail.

subtropical and tropical forests. They cannot grow in arid

habitats. Several species grow the eastern and
in

northwestern United States, but none occur in the more

arid states of the southwest. Some eastern species are in

Figure 14.1 Psilotum nudum. A, plant; note sporangia on danger of extinction because they are popular as
branches at left. B, end of a branch, showing scale leaves and Christmas decorations.
sporangia at nodelike swellings.
Mature Sporophyte

Lycophyta, so that a living stem of a member of this

The main stem branches freely and is prostrate. Upright
division has a core of xylem. The leaves are microphylls; stems, approximately 20 cm in height, grow from the

the central vein does not branch. horizontal stem. Both types of stems are sheathed with
small green leaves. Small but well-developed adventitious
roots arise irregularly from the underside of the horizontal

Lycopodium stem. As in many higher plants, the primary root, which

grows from the embryo, is not long-lived.
Approximately 400 species are in this genus. Most are The xylem in a Lycopodium stem occurs as a complex
trailing plants, many forming short, upright branches that system of anastomosing strands, so that its distribution in
loosely resemble pine seedlings (Figs. 14.5, 14.6). They the stem varies greatly in different cross sections of the
are frequently called "ground pine" or "club moss." stem. The phloem is present between the strands of xylem
Although widely distributed, they are most abundant in and thus, too, forms a complex system of anastomosing

Division Lycophyta

289
Figure 14.3 Gametophyte of Psilotum.

The xylem is composed of tracheids, whereas the

strands. species, they differ from sterile leaves in size, shape,
phloem contains sieve cells and some parenchyma cells. position, and color. Such modified sporophylls are
An endodermis encircles the vascular cylinder. grouped together closely at the ends of stems, forming a
Lycopodium stems also lack pith, as do roots of most cone or strobilus (Figs. 14.5, 14.6, 14.8).
higher vascular plants.
Leaves of Lycopodium show a well developed epidermis Gametophyte. The meiospores germinate and develop
with stomata, a mesophyll with many air spaces, and a into gametophytes. Gametophytes in some species grow
midvein (Fig. 14.7). above ground and are green; in other species,
gametophytes are subterranean and lack chlorophyll. The
Reproduction
gametophytes are always associated with a fungus. Both
Lycopodium may reproduce asexual ly or through an male and female gametangia are found on the same
alternation of generations involving distinctive gametophyte. Gametangia are borne on the upper portion
gametophytic and sporophytic generations. Both of the gametophyte (Fig. 14.8). Fertilization occurs when
generations are independent and the sporophyte sufficient free water is present to allow sperms to swim to
generation is the dominant generation. Meiospores are mature archegonia.
borne in sporangia in or near the leaf axils (Figs. 14.5,
14.6), sheathing the stem. Spores (Fig. 14.8) and Sporophyte Embryo. The embryo sporophyte possesses
sporangia of a given species are all alike. Leaves bearing (a) a well-developed foot, (b) rudiments of a short primary
sporangia are called sporebearing leaves, or sporophylls. and (d) a short shoot apex
root, (c) leaf primordia, (Fig.

In some species, they closely resemble ordinary nonspore- 14.8). The embryo grows directly into the mature
bearing leaves in their structure and appearance. In other sporophyte plant.

chapter 14 I Lower Vascular Plants

290
 *

#%L,

Figure 14.4 Closer view of Psilotum gametophyte, showing

globular antheridia (male gametangia) and cross-shaped neck
cells of archegonia (female gametangia).

Selaginella crevices in the higher Sierra Nevada and is able to

withstand periods of drought. The more delicate
Selaginella species resemble those of Lycopodium in their Selaginella emmeliana grows best in a humid environment;
general appearance but are smaller. They number more it is grown as an ornamental plant.
frequently
than 700. Although they are widely distributed, most of The vascular system in stems of Selaginella consists of
them are tropical; a few grow in temperate zones. Some one to several branching strands, each having a central
species are adapted to withstand periods of drought and core of xylem surrounded by phloem. The cross sections
hence may grow in relatively dry localities. in Fig. 14.9 show a single strand in a stem of Selaginella.

Itoccurs in a large air space and is supported by strands

Mature Sporophyte of radially arranged endodermal cells. Vessels are present

As in Lycopodium, the sporophyte of Selaginella generally in the xylem of several species of Selaginella. Parenchyma

consists of a branched, prostrate stem with short, upright and sclerenchyma tissue form a cortex, which is bounded
branches, usually only 10 cm high. In some species, the externally by an epidermis.

stem is and slender and taller. Both horizontal and

upright
Reproduction
upright stems are sheathed with small leaves in four
longitudinal rows or ranks. As in Lycopodium, spores are borne in sporangia, which
Two species of Selaginella are shown in Fig. 14.5. One grow in or near axils of sporophylls. Although sporophylls
of them, Selaginella watsonii, grows in exposed rock do not differ greatly in appearance from sterile leaves,

Division Lycophyta

291
 lE^^TI

/
- ^^ *

H » ^»

Figure 1 4.5 Two species of Selaginella. A, S. watsonii, a

montane species growing in crevices in granite; B, S. emmeliana,
native to tropical America and requiring a moist, warm climate.

they are always grouped to form cones, or strobili, at the The megaspore (containing the female gametophyte)
ends of upright branches. may be shed fromthe cone or strobilus at almost any
Two types of sporangia are formed: megasporangia stage of the development of the gametophyte. In some
(Greek, mega, large) and microsporangia (Greek, micro, species, it may be retained in the strobilus until well after
small). As the names indicate, megasporangia produce any event, fertilization occurs only when
fertilization. In

larger spores. A single strobilus usually contains both from rain or dew, is present,
sufficient water, either
types of sporangia. allowing sperms to swim to archegonia.
Within a developing megasporangium, all but one of the
megasporocytes (potential spore-producing cells) Male Gametophyte. Upon germination, microspores
degenerate. This remaining megasporocyte, nourished in divide into two cells. One of them, the prothallial cell, is
part by fluid resulting from the degenerating spores and in small and does not divide further; it represents the
part by cells surrounding the spore cavity, increases vegetative portion of the male gametophyte. The other cell,
greatly in size. During meiosis, four large spores, called by repeated divisions, develops into an antheridium
megaspores, are formed from the megasporocyte. Each composed of a jacket of sterile cells enclosing 128 or 256
megaspore may germinate and give rise to a female bilflagellated sperms. This development takes place within
gametophyte (megagametophyte; Fig. 14.10). the microspore wall. Microspores are shed from the
Only a few microsporocytes within the developing microsporangium midway in the development of the male
microsporangium degenerate. The approximately 250 gametophyte and grow to maturity without direct
microsporocytes that remain undergo meiosis, each connection with parent sporophyte or soil. Usually,
forming four small spores, or microspores (Fig. 14.10). microspores sift down to the bases of the
The production by a given species of two distinct types megasporophylls below the microsporophylls (Fig. 14.10).

of —
meiospores megaspores and microspores is called — In this position, they are close to the developing female
heterospory. Thus, Selaginella is heterosporous. gametophytes. The sperms escape when the microspore
Lycopodium, which produces but one spore type, is said wall ruptures.
to be homosporous. Sperms swim which grow on that
to the archegonia,
portion of the female gametophyte protruding from the
Female Gametophyte. Repeated cell divisions within the megaspore (Fig. 14.10). Fertilization ensues, and the
megaspore result in the female gametophyte, which is resulting zygote initiates the diploid or sporophyte
contained within the megaspore wall until it nears maturity. generation.
Its increase in size eventually ruptures the megaspore wall,
and a small cushion of colorless gametophytic tissue Sporophyte Embryo. Of the two cells formed by the first
protrudes from the megaspore along the lines of rupture division of the zygote, only one develops into an embryo.
(Fig. 14.10). Archegonia develop on the protruding The othercell grows into an elongated structure, the

cushion. suspensor, which pushes the developing embryo into

chapter 14 | Lower Vascular Plants

292
Figure 14.6 Sporophylls in Lycopodium. A and B, sporophylls
similar to vegetative leaves in L. selago. C, sporophylls different
from vegetative leaves and grouped in terminal strobili in L.

obscurum.

Figure 14.7 A, cross-section of Lycopodium leaf. B, stomate

detail. C,vein detail.

Division Lycophyta

293
 egg eel

Figure 14.8 Stages in the lite cycle of Lycopodium. A, L.

annotinum; B, section of sporophyll showing location of
sporangia; C, spore; D, gametophyte; E, section of gametophyte
showing location of gametangia; F, antheridium; G, sperm; H,
archegonium; section of embryo; J, underground gametophyte
/,

with young sporophyte.

chapter 14 I Lower Vascular Plants

294
 Si
mature sporophyte \l\«

Division Lycophyta

295
 epidermis

cortex

f xylem .
stele- ^T
phoem-U—
^
endodermis

Figure 14.9 Cross section of a stem of Selaginella. A, diagram

showing stem tissues; a single, flattened cylinder of
distribution of
vascular tissue is supported in an air space by filaments of
endodermal cells and the cortex is composed of parenchyma and
sclerenchyma. B, detail showing cellular structure.

epidermis

sclerenchyma <

> cortex

> cortex

pericycle
stele

chapter 14 l

Lower Vascular Plants

296
gametophytic tissue, where there is a food supply (Fig. each vascular bundle, or encircle a ring of vascular
14.10). bundles. In some species, there is also an inner
The embryo is a structure with (a) a foot, (b) a root, endodermis.
(c) two embryonic leaves, and (d) a shoot (Fig. 14.10). In
embryo is held by the
certain species of Selaginella, the
megaspore and retained within the strobilus. does not It
Reproduction
pass into a dormant state, as do embryos of seeds, but
continues to grow. Young sporophytes may be found In all species of Equisetum, the sporangium-bearing
extending from the strobilus of parent sporophytes. If the organs, sporangiophores, are specialized structures very
developing embryo were to pass into a period of different from ordinary leaves. They are grouped together
dormancy while being held by the parent sporophyte, a in strobili, or cones (Fig. 14.1 1) at the summit of main
condition would arise that approaches the seed habit. upright branches and occasionally on lateral branches. In

most species, strobili are borne on ordinary vegetative

shoots; in a few species, they are formed only on special
Division Sphenophyta fertile shoots.
The sporangiophores are stalked, shield-shaped
structures borne at right angles to the main axis of the
Members of this division once grew very abundantly, as cone. The cone may be compared to a pole to which open
their good representation in the fossil record shows. umbrellas have been fastened, the handles of the
Today, the division is represented by a single genus umbrellas being at right angles to the pole. Sporangia are
(Equisetum) of about 25 species (Fig. 14.1 1). Many attached to the underside of the shield (umbrella), close to
species inhabit cool, moist places, but Equisetum arvense its edge. They extend horizontally inward, toward the axis
also grows in dry habitats. Most species are characterized cone. Four meiospores with the haploid number
of the of
by the presence of silica in the epidermis of stems. chromosomes are formed from each sporocyte. All
Because of this trait, they were used in colonial days to meiospores are morphologically alike. Equisetum,
scour pots and pans and hence were called scouring therefore,is homosporous. Upon maturity the meiospores

rushes. The genus is also commonly known as the are discharged from the sporangia. The spores are fragile
horsetails. and normally live but a few days.

Gametophytes
Mature Sporophyte Gametophytes are small green bodies about the size of a
pinhead and consist of a cushionlike base with many
The sporophyte of Equisetum is the dominant phase of its
erect, delicate lobes (Fig. 14.13). They are easily cultured
life history. In one tropical species, the sporophyte is
in the laboratory.
vinelike and may reach 7 m in length. Usually, 1 m
Gametangia are similar to those of Lycopodium.
represents the maximum upright growth. All species are
Fertilization takes place only when there is free water.
perennials and have a branched rhizome from which
Sperms are spiral shaped and multiciliate.
upright stems arise. Stems, depending on the species,
The zygote develops directly into an embryo. No
may branch either profusely or sparingly. In either case
suspensor is formed. The embryo is similar to those
they are straight and marked by vertical ridges and distinct
already described, except that the foot is small or lacking.
nodes (Fig. 14.1 1). Bases
nodes are sheathed by
of
whorls of small simple leaves. Many branches may also
form at each node. The microphyllous leaves are much
reduced in size, nongreen and, in many species, short- Division Pterophyta
lived. The stems are green and are, therefore, the organs

of food manufacture. Roots occur at the nodes of the

rhizomes, but they may also develop from stem nodes if The division Pterophyta comprises the ferns, some 10,000
the stem becomes procumbent on a moist surface. species, most of which are shade-loving plants of small
Except at nodes, stems of Equisetum are hollow and the size, their upright leaves, or fronds, generally being their
ribs make prominent markings on their outer most prominent feature. Most species native to temperate
circumferences. The ridges are formed of sclerenchyma zones have an underground rhizome with leaves and roots
tissue and not only project outward, but extend inward arising at nodes (Fig. 14.14); a few are small, floating
almost to the small vascular bundles (Fig. 14.12). There aquatics without rhizomes. Some ferns of tropical regions
are air spaces, or canals, between vascular bundles. do grow into fairly large trees. All Pterophyta have a
Photosynthetic tissue lies between these air canals and definite alternation of generations, with both sporophyte
the thin, outer layer of sclerenchyma tissue and epidermis. and gametophyte being autotrophic plants.
Vascular strands are also marked by a canal (Fig. 14.12). The Pterophyta comprise four orders, the largest of
Arms of xylem extend outward from this canal, and which, the Filicales, includes the true ferns. The Filicales
phloem tissue lying outward from the canal is present are divided into 1 1 or more families, of which the largest
between radial arms of xylem. Some species contain and best-known in the United States is the Polypodiaceae.
vessels in the xylem. An endodermis is present, but its Most of the discussion below deals with representatives of
location varies from species to species; it may surround this family.

Division Pterophyta

297
 microsporophylls

N
Figure 14.10 Stages in the life cycle of Selaginella. A, shoot of
S. wafson// showing two strobili; B, microsporophyll with
microsporangium; C, megasporophyll with megasporangium; D,
strobili showing sporophylls with sporangia; E, microsporangium
discharging microspores; F, megasporangium discharging
megaspores; G, microspores sift downward and lodge in axils of
megasporophylls close to megaspores; H, a microspore; /, section
of a microspore; J, male gametophyte with microspore; K,
antheridium with developing sperm cells; L, mature sperm cells;
M, female gametophyte within megaspore; N, section of female
gametophyte showing location of archegonia and a young embryo
pushed by its suspensor into the vegetative tissue of the female
gametophyte; O, the young embryo sporophyte generally remains
embedded in vegetative gametophyte tissue.

chapter 14 I
Lower Vascular Plants

298
microspore

Divisi on Pteroph yta

299
Figure 14.11 Stems of Equisetum telmateia. A, fertile stems with
note much-branched, adjacent sterile stems. B, strobilus
strobili;
with sporangiophores. C, enlarged view of the stem, showing
three nodes, vertical ridges, and reduced leaves.

chapter 14 l
Lower Vascular Plants

300
 epidermis
cortex

endodermis

vailecular
canal
central canal

carinal canal

sclerenc
cortex
parenchy

canal

carinal

Figure 14.12 Cross section of Equisetum stem. A, diagram

showing arrangement of tissues; 8, enlarged view showing details
of cellular structure.

Division Pterophyta

301
 Mature Sporophyte
The sporophyte, which is the dominant generation of all

ferns,possesses an underground stem or rhizome from

which leaves and adventitious roots arise (Fig. 14.14).
Leaves are the most prominent part of the fern plant and
they vary greatly and form. Young fern leaves are
in size

and consist chiefly of meristematic

rolled into tight spirals
tissue (Fig. 14.15). As the leaf matures, it unwinds from
the base upwards. The upright, expanded portion is
mature, but the leaf continues to grow by cell division at
its coiled tip. In certain species, the apical meristem may
remain active for years, resulting in leaves that are nearly
3 m long. The uncoiled, fully expanded fern leaf lacks any
residual meristematic tissue.
Most fern leaves are compound, although simple types
exist inall groups. The most common type of fern leaf has

Figure 14.13 Gametophyte of Equisetum. a stout or rigid petiole, which is prolonged to form a rachis
from which leaflets (pinnae and pinnules) arise (Fig.
14.15).
The vascular tissue of ferns is organized into one or
more vascular strands, each having a core of xylem,
surrounded by phloem and separated from the ground
parenchyma by an endodermis. These vascular bundles
are interconnecting, so that, as in Lycopodium, the precise

arrangement will vary from section to section. However,

there are arrangements characteristic of various genera.
pinnule
For instance, in Polypodium there is a ring of small
bundles placed well out from the center of the rhizome
(Fig. 14.16). Each strand is formed of a central core of

xylem, surrounded by phloem, and the whole has a

bounding endodermis. The xylem consists largely of
tracheids, vessels being known in only two genera. Sieve
cells occur in the phloem.
A typical fern leaf has a well-developed epidermis with
chloroplasts in the epidermal cells and stomata on the
lower surface. The mesophyll may be relatively
undifferentiated or divided into palisade and spongy
parenchyma layers (Fig. 14.16).

Reproduction

Vegetative Reproduction

Vegetative reproduction may occur in one of two ways:

(a) by death and decay of the older portions of the

rhizome and the subsequent separation of the younger

growing ends; and (b) by the formation of deciduous leaf-
borne "buds," which become detached and grow into
new plants. Such buds occur in only a few genera. Some
clumps of ferns are thought to be hundreds of years old.

Sexual Reproduction
In the sexual life cycle, independent sporophyte and

gametophyte generations alternate with each other. The

vegetative structure of the sporophyte has been described
roots above. Spores are borne in sporangia, which ordinarily
develop on the lower surface or margins of fronds (Fig.
14.17). Not all leaves are fertile (i.e., spore-producing),
and the fertile leaves in some species are dissimilar in
Figure 14.14 A mature fern sporophyte, showing
typical,
rhizome below ground and fronds above ground. Lady fern, structure to the sterile (non-spore producing) leaves. The
Athyrium felix-femina. distribution of sporangia on the leaf surface varies

chapter 14 l
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302
 Figure 14.15 Diversity of form in fern fronds. A, lightly rolled tip
of a young frond of the tree fern Cyathea; B, adder's tongue
(Ophioglossum vulgatum); C, moonwort (Botrychium lunaria); D,
grape fern (Botrychium multifidum); E, golden back fern
(Pityrogramma triangularis); F, spleenwort (Asplenium
pinnatlfidum); G, leather leaf (Polypodium scouleri); H, holly fern
(Polystichum lonchitis; five-finger fern (Adiantum pedatum); J,
I,

lip fern (Cheilanthes covillei); and K, cliff brake (Pellaea

compacta).

Ophioglossum vulgatum Botrychium lunaria Botrychium multifidum Pityrogramma Asplenium pinnatifidum

(adder's tongue) (moonwort) (grape fern) triangularis (spleenwort)
(golden back fern)

Polypodium scouleri Polystichum lonchitis Adiantum pedatum Cheilanthes covillei Pellaea compacta
(leather leaf fern) (holly fern) (five-finger fern) (lip fern) cliff brake)

Division Pterophyta

303
 epidermis

phloem
-^— xylem

cortex

xylem

phloem
pericycle

endodermis

Figure 14.16 Fern anatomy. A, Polypodium stem cross section,

showing relationship of tissues; B, a higher magnification of A,
showing cellular detail; C, leaf cross section of Polypodium.

phloem

considerably in different genera and species (Fig. 14.17). sporangia and aids in dispersal of ripe spores (Fig. 14.18).
Sporangia may (a) cover much of the lower surface, The young sporangium is filled with sporocytes. Meiosis
(b) be grouped in sori (singular, sorus) and grow in a results, as always, in spores with a reduced number of
definite relationship with veins, or (c) grow only along chromosomes. Ferns, with the exception of two families,
margins of the leaf. the Marsileaceae and the Salviniaceae, are homosporous.
When sporangia are grouped together in sori, a These two families are widely distributed, small, aquatic,
structure called the indusium (Fig. 14.18), which is and unfernlike in appearance.
sometimes umbrella-like, may be present, thereby
protecting young and developing sporangia. Frequently, Gametophyte. Gametophytes, or prothallia, of the
marginal sporangia are protected by the curled edge of Polypodiaceae are small, flat, green heart-shaped

the leaf which forms a false indusium.

itself, structures (Fig 14.19) with rhizoids on their lower surface
The sporangium is a delicate, watch-shaped case, (Fig. 14.18). In most species, they apparently mature

consisting of a single layer of epidermal cells, only one rapidly and are not long-lived. Antheridia and archegonia

row of which possesses heavy walls. This row, which are borne on the same prothallus. Antheridia are formed
nearly encircles the sporangium in the Polypodiaceae, is when the prothallus is very young and are scattered over
the annulus; it functions in opening dried mature its lower surface. Normally, 32 sperms develop within

chapter 14 | Lower Vascular Plants

304
 each antheridium.
Archegonia torm later than antheridia and are usually
clustered close to the notch, also on the undersurface of
the gametophyte (Fig. 14.18).
Fertilization occurs when free water is present and
sperms are thus able to swim to the neck of the
archegonium. The resulting zygote is diploid and rapidly
develops into an embryo sporophyte comprising a foot,
root, stem, and leaf (Fig. 14.18). The embryo develops
directly into a young sporophyte, without a dormancy
period.

Summary
1. Lower vascular plants is a general name for vascular
plants that do not produce seeds. This group consists
of four divisions, the Psilophyta, Lycophyta,
Sphenophyta, and Pterophyta. The latter division, the
most species and is the most
ferns, contains the
widespread.
2. The Psilophyta is represented by only two genera,
Figure 14.17 Diversity in sori. A, Pellaea, showing sporangia in
a band close to leaflet margin; B, Woodwardia, showing sori Psilotum and Tmesiptris. Sporophytes of Psilotum
parallel to midrib and covered with indusia; C, Polypodium, consist of a branched, underground rhizome with
showing round sori; D, Polystichum, showing round sori with
centrally attached indusia; E, Asplenium, showing sporangia in
rhizoids (no true roots) and an upright, green, branched
long rows on veins. stem with scalelike microphylls. It is homosporous and

Sum mary
305
 archegonia

Figure 14.18 Stages in the life cycle of a fern. A, fern frond; B.

section through a sorus, showing indusium, sporangia, and
spores; C, spore; D, gametophyte; E, antheridium; F, section of
antheridium showing sperm; G, open antheridium discharging
sperm; H, archegonium; archegonium receptive to sperm; J,
/,

section of gametophyte showing a zygote with the venter of the

archegonium; K, section of gametophyte and embryo; L,
gametophyte with a young sporophyte still attached.

chapter 14 I Lower Vascular Plants

306
sperm
cover cells

Summary

307
 the meiospores germinate to produce
nonphotosynthetic, small gametophytes. Free water is

required for fertilization. The embryo sporophyte is at

first attached to the gametophyte by an absorbing

organ (the foot), and it depends for nourishment on the
gametophyte.
3. The Lycophyta is represented by only two genera in
this book: Lycopodium and Selaginella. Isoetes (the
quillworts) was not discussed, nor were two other
genera, Stylites and Phylloglossum.
4. Sporophytes of Lycopodium (club mosses) are less
than 20 cm tall and exhibit true roots, stems, and
leaves. The stems are sheathed with small leaves that
contain an epidermis with stomata, a mesophyll with
intercellular air spaces, and a central vein. Sporangia
are borne in the axils of leaves: they may be clustered
near the branches in a strobilus (cone). The
tips of

genus is homosporous, and the meiospores produce

small gametophytes that may or may not have
chlorophyll. Free water is required for fertilization, and
the embryo sporophyte is at first attached to the
gametophyte by a foot, as in Psilotum.
5. Sporophytes of Selaginella are similar in appearance to
those of Lycopodium. Vessels are present in the xylem
Figure 14.19 Photographs of fern gametophytes. A a view from of some species. Sporangia are borne in the axils of
beneath, showing rhizoids, antheridia (dots), and archegonia leaves and they are clustered into strobili. Sporangia
(near notch); B, a gametophyte (G) with attached young
sporophyte (S) emerging and overtopping the gametophyte. and spores are of two kinds. Megasporangia produce
megaspores that germinate and grow (within the
ruptured spore wall) into female gametophytes.
Microsporangia produce microspores that germinate
and grow (within the spore wall) into male
gametophytes. Microspores may sift down the strobilus

chapter 14 l
Lower Vascular Plants

308
 to lie near megaspores, then they rupture to release 7. Sporophytes of Pterophyta (ferns) are typically below
sperm; if free water is present fertilization can be 1 m in height, but some reach tree stature. In most
accomplished in the strobilus. The embryo is at first cases, the stem is completely underground as a
pushed deep within the female gametophyte by a rhizome, and only the large leaves and petioles are
suspensor, and nutrition is gained from the above ground. A few ferns are small, floating aquatics.
gametophyte through a foot. Selaginella is Vessels are presentin some species. Leaves may
heterosporous. and spongy parenchyma. Sporangia are
exhibit palisade
Sporophytes of Sphenophyta (horsetails) are about 1m produced on specialized leaves in some species, or on
tall and consist of an underground rhizome, with true
the undersides of all leaves in The
other species.
roots, and an upright, jointed, silica-rich stem.
sporangia may be clustered into compact groups called
Microphylls and branches arise in whorls at the nodes.
soriand covered with an indusium. Most ferns are
Vessels are present in some species. Sporangia are
homosporous and, if so, the meiospores germinate and
grouped into complex strobili. The division is
grow into small, photosynthetic, heart-shaped
homosporous. Meiospores germinate and grow into
gametophytes (prothallia). Free water is necessary for
small, photosynthetic gametophytes. Free water is
fertilization, and the embryo sporophyte is much like
required for fertilization. The embryo stage is similar to
that of Psilotum. that of Psilotum.

Table 14.1
Comparison of Psllophyta, Lycophyta, Spenophyta, and Pterophyta

Generation Psilotum Lycopodlum Selaginella Equlsetum Ferns

Sporophyte A branching Prostrate Prostrate branching Rhizome, upright, Rhizome, roots, and
stem, scale branching stem stem with upright jointed stem, small leaves
leaves, no roots with upright branches, roots, and leaves
branches, roots, leaves, also scales
and leaves
Herbs or Herbs Herbs Herbs Herbs or trees
epiphytes
Simple vascular Simple vascular Simple vascular Simple vascular Stem: several strands of
system system system, but vessels system but vessels vascular tissue each
are present may be present with xylem surrounded
by phloem; Vessels:
only in two genera;
Sieve cells: no
companion cells in
phloem
One type of One type of Megasporangia and One type of One type of sporangium
sporangium sporangium microsporangia sporangium (mostly)
Homospory Homospory Heterospory Homospory Homospory (mostly)
No sporophylls Sporophylls Sporophylls present Sporangiophores Leaves bear spores in

present sporangia
No cone A cone in some Cone Cone composed of No cone
species sporangiophores
Embryo develops Embryo develops Embryo develops Embryo develops Embryo develops
attached to attached to within female attached to attached to
gametophyte gametophyte gametophyte on gametophyte gametophyte
sporophyll

Gametophyte Irregular Irregular to Male gametophyte: A single green Heart-shaped, small,

subterranean tapered one prothallial cell, thallus resembling green, completely
structure, subterranean and an antheridium Anthoceros independent thallus
associated with structure, within microspore
fungus associated with
fungus
Gametangia Gametangia Female Gametangia Antheridium of
embedded in embedded in gametophyte: small embedded in approximately 6 to 10
thallus thallus amount of tissue neck of
thallus, cells, archegonia
withinmegaspore, archegonium partially embedded in
cushion protrudes in protruding gametophyte
which gametangia
are embedded
Motile sperms Motile sperms Motile sperms Motile sperms Motile sperms

Summary

309
 15 CHAPTER

15 gymnosperms

The higher vascular plants possess both vascular

and seeds. Gymnosperms (conifers and their
tissue
Coniferophyta. There are nine families and 550 species.
They comprise pines, hemlocks, firs, spruces, junipers,
relatives) and angiosperms (flowering plants) yews, redwoods, and many others. Not all of them bear
together make up the higher vascular plant group. These cones, yet the cone such a conspicuous feature of
is

plants dominated the world's vegetation during the many of them that the division has been named the
Cenozoic era (Chapter 10). Coniferophyta or cone bearers. Most conifers form true
Gymnosperms are a major source of lumber and paper cones. Juniper berries are in reality cones with fleshy
pulp in North America. This group is represented by such adhering scales. In yews, seeds are surrounded at the
common trees as pines (Pinus), Spruces (Picea), firs base by a more or less pulpy, berrylike body and are not
(Abies), and true cedars (Cedrus), all of which possess borne in cones. In either case, the seeds are naked, not
well-developed seed-bearing cones (Fig. 15.1). Other being surrounded by an ovary wall.
gymnosperms, such as Ginkgo, yews (Taxus), and The following brief descriptions of several common
Mormon tea or joint-fir (Ephedra), do not bear cones. genera enable us to introduce the more important conifers
A trait common to all gymnospermous plants is the in some of the seven families.

absence of a protecting case, that is, an ovary wall,

Family Pinaceae
around the seeds. In pines and other cone-bearers, seeds
are borne on the surface of scales that comprise the cone Pinus (Pines). Pines are usually large trees. Some
and, though well-protected by the scales, they are not members of bristlecone pine (Pinus longaeva) are the
surrounded by floral parts. In yews, seeds are partially oldest living things, over 5000 years old. The leaves are
surrounded by a red berrylike structure (Fig. 15.2). In needlelike,two or more growing together (except in Pinus
Ephedra, ovules are borne in axils of short bracts (Fig. monophylla) in a fascicle or group, which is sheathed at
15.2) and in Ginkgo, naked ovules are attached to ends of the base (Fig. 15.1). The cones vary greatly in size and
short branches (Fig. 15.2). By contrast, the seed of an shape and are very characteristic of the species to which
angiosperm is surrounded by a matured ovary wall. they belong.
Seeds lacking protection of an ovary wall are said to be
—
"naked" thus the name gymnosperm, derived from two Abies (Firs). Firs are stately trees of a symmetrical,
Greek words gymnos (naked) and sperm (seed). Fossil cylindrical or pyramidal shape. The leaves are flat and
Lepidodendrales, belonging to the Lycophyta, bore seeds linear; in cross section they are relatively broad, without
more than 200 million years ago, thus indicating that the marked angles. The cones are erect on the branches and
seed habit is neither of recent geological origin, nor shatter at maturity (Fig. 1 5.3).
indicative of a single evolutionary line.
There are four divisions of living gymnosperms, Picea (Spruces). These trees closely resemble the firs,
encompassing more than 600 species. Of these divisions, but they can be distinguished by the position of the leaves
we shall only briefly consider: (a) the Cycadophyta, or on the branchlets, by the angular appearance of the
cycads; (b) the Ginkgophyta, composed of only one leaves in cross section, and by the cones that are hanging
species, the maiden-hair tree; and (c) the Gnetophyta, an and do not shatter at maturity (Fig. 15.4).
artificial grouping of three unusual genera, Gnetum,

Ephedra, and Welwitschia. We will discuss a fourth Tsuga (Hemlocks). The trees of this genus are
division, the Coniferophyta or conifers, in much more pyramidal with slender horizontal branches. The leaves
detail, and we begin with that group. are usually two-ranked, linear, flat, and with a short
petiole.They resemble the leaves of firs but are much
shorter. The cones are small.
Division Coniferophyta
Pseudotsuga (Douglas Fir). Only two species occur in

the United States. One of them, the Douglas fir

Classification (Pseudotsuga menziesii), is the most important timber tree

of the United States. It may grow to a height of 60 m, with
Without exception, the well-known and economically a trunk diameter of 3 m. Its leaves are flat, like those of
important gymnosperms of temperate zones belong to the the true firs, but have white lines on either margin and a

chapter 15 |
Gymnosperms

310
Figure 15.1 Pines. A, Pinus jefireyi (Jeffrey pine), showing the
trunk of one tree and the crown of a second tree. B, mature
ovulate cones of P. ponderosa (ponderosa pine), a closely related
species. C, fascicle of ponderosa pine needles.

Division Coniferophyta

311
 groove along the upper surface. The cones are 5 to 1 1 cm
long, pendulous, and easily recognized by bracts
extending outward below each scale.

Larix (Larches). Trees belonging to this genus grow in

the cooler portions of the Northern Hemisphere. They

differ from most other members of the Pinaceae in that
they are deciduous. The American larch, or tamarack, is a
tall tree frequently found in bogs. The needles are short,
linear, and grouped in crowded clusters on short spurs.
Figure 15.2 Gymnosperms without typical cones. A, branches
of Taxus with seeds; note berrylike aril surrounding seed. B, On leading shoots, however, the needles are arranged
branch and seeds of Ginkgo biloba. C, a cluster of female strobili spirally. The cones are small and persistent (Fig. 15.5).
of Ephedra; note "pollination drops" at the trip of two cones.

Family Cupressaceae

This family contains the junipers, cypresses, and false

cedars (true cedars, Cedrus species, are in the
Pinaceae). The leaves are borne singly, but are usually

chapter 15 |
Gymnosperms

312
 Figure 15.3 Abies. Ovulate cone and branch. Note arrangement
of needles.

p
m^
i*
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-r/^
w«S*- 3%^^
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K.£*2 BBjfca^fc
"a"^
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E^SSl
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s^k t^ Jill.
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._- -v—BC? ^K
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iSEBE*

Figure 1 5.4 Picea. /A, branch of P. alba with ovulate cones; 6,

leaves of Picea.

small and scalelikeand are crowded closely around the common and the record height being 112 m. Redwood
stems (Fig. 15.6). The cones may be woody, as in has outstanding lumber qualities, not the least of which is

Cupressus, or fleshy, as in Juniperus. The cone of juniper its resistance to decay. Consequently, logging has
is often called a "berry," but it is a modified cone, removed 90% of all virgin redwood forest in the past 150
composed of fleshy scales that completely enclose the years. Only half of the remaining acreage is protected
seeds (Fig. 15.6). from future logging. Sierra bigtree (Sequoiadendron
giganteum) is the most massive of any plant or animal
Family Taxodiaceae
species in the world; its trunk at breast height may reach
This family contains the bald cypress, redwood, and Sierra 9.8 m in diameter and its height above 82 m. The trunk
bigtree. Redwood (Sequoia sempervirens) may be the alone of the General Sherman tree weighs 625,000 kg
tallest tree in the world, individuals over 60 m being (over 680 tons).

Division Coniferophyta

313
 ^2p* »|*M # •

PRLS
L* ±Sri*^wlLJ

v ^H^^B^MpI
Jt8l^>fejSv^j
B*Sk hHr

Figure 15.5 Larix lyallii A, branch with needles; B, mature

ovulate cones.

MUBR V . • r^tf paw

1 ^^PtJ? Ij^HK ^^^^^*l

&
aw

n
* *0 -

^
^^«
V/«VY^
**• .

- *

•/i

Figure 15.6 Juniperus occidentalis growing at 2200 m in the

Sierra Nevada Mountains of California. /A, tree; S, branch with
staminate cones; C, branch with ovulate cones.

chapter 15 |
Gymnosperms

314
Family Taxaceae Sporophyte
All species of pines are trees. They range in size and age
With two exceptions, ovulate cones are not borne by
yew from scrubby, 70-year-old fire-adapted species such as
members of the family. The seed of this family so
strongly resembles a drupe or a nut that it is usually called
knobcone pine (Pinus attenuata) to large, straight trunked
The embryo protected by an outer flesh called an
ponderosa pine (Pinus ponderosa, Fig. 15.1) 500 years
a fruit. is
old, to twisted, slow-growing bristlecone pine (Fig. 15.7)
aril, and an inner hard pit. Both these tissues may be
derived from the integuments of the ovule; hence, the
5000 years old.

structure is morphologically a naked seed. As shown in

Conifer wood has no vessels. A three-dimensional
reconstruction from a small piece of Sequoia is
Fig. 15.2, the outer red ground pine (Taxus
aril of
canadensis) almost encloses the seed. It drops away when
diagrammed in Fig. 15.8. It is extremely regular. Tracheids
are elongated cells that appear in cross section as square,
the seed is mature.
Native yews are common only in a few places in the
hollow cells. Those formed in the spring are largest in

diameter; as the season progresses, newly formed

United States. Possibly the best-known yew is the English
tracheids are smaller in diameter with thicker walls. The
yew, which, because of the excellent bows that were
heavy-walled cell is called a fiber-tracheid. Note xylem
made from its wood, is closely linked with English history
and folklore. Yews are now widely cultivated.
rays are composed of brick-shaped parenchyma cells,

sometimes with dark-staining contents. Other parenchyma

cells may run in vertical columns; these are called axial

Life History of a Conifer parenchyma.

Many gymnosperms produce resin. Typically, resin
The following outline of a gymnosperm life cycle is based occurs in long resin ducts that are surrounded by
largely on the genus Pinus. It differs from life cycles of parenchyma cells (Fig. 15.9). Resin may play a role by
other genera mainly in requiring three summers for inhibiting the activity of wood-boring insects. Sequoia does
completion. Significant stages of pollination, fertilization, not produce resin.
and maturation of the seed occur within a decimeter of In the phloem, companion cells are missing, but
each other on a given branch, thus making it an ideal otherwise, sieve cells, fibers, and parenchyma cells are
subject for class study. present.

Figure 15.7 Pinus longaeva (bristlecone pine) in the White

Mountains of California, growing near timberline.

Division Coniferophyta

315
 bordered pits- ray
Figure 15.8 Block diagram of secondary xylem of redwood
(Sequoia sempervirens).

wood ray tracheids resin duct wood parenchyma The needle leaves of Pinus show considerable
adaptation to drought (Fig. 15.10): sunken stomata, thick
cuticle, fibrous epidermis, close-packed mesophyll, and
veins only in the center of a thick leaf. Such modifications
are important in winter, when soil moisture is frozen and

the evergreen leaves are subjected to drying winds.

All conifers produce two kinds of spores microspores
—
and megaspores borne in cones that are
morphologically distinct. The two types of cones are
known, respectively, as staminate (or pollen) and ovulate
(or seed) cones.

Staminate (Pollen) Cone

Staminate cones average 10 mm or less in length, by 5

globule of resin- 1 "— secretory cells - mm in diameter. They are borne in groups, usually on

Figure 15.9
lower branches of trees (Fig. 15.1 1). Each cone is
Resin duct in pine wood, as seen in cross section.
composed of a large number of small scales
(microsporophylls) arranged spirally on the axis of the
cone. Two microsporangia develop on the under surface
of each scale (Fig. 15.11).

Stages of microspore development are quite similar to

those of Selaginella and to spores of mosses and ferns.
Microsporocytes, which are surrounded by a nutritive cell

chapter 15 |
Gymnosperms

316
 1

parenchyma
^*f^^/- mesophyll
schlerenchyma

schlerenchyma

mesophyll

epidermis hlerenchyma
Pinus

Figure 15.10 Cross section of a two-needled pine leaf. The inset

(above) shows the entire cross section, with major tissues
outlined; the large drawing shows cellular detail of a wedge from
the epidermis in to one of the vascular bundles at the center.

layer, the tapetum, undergo meiosis, and four Female Gametophyte. The megasporocyte soon divides
microspores result. As usual, each microspore contains by meiosis. Four megaspores, usually arranged in a single
the haploid number of chromosomes. The nucleus within row of four cells, result from each megasporocyte. The
the newly formed microspore divides several times by nucleus within each megaspore has the haploid number of
mitosis, forming a pollen grain that contains two viable, chromosomes. Generally, only one of the four megaspores
haploid nuclei and vestiges of several vegetative cells develops into a female gametophyte; the other three
(Figs. 15.11, 15.12). Enormous numbers of pollen grains degenerate. Germination of the remaining megaspore and
are finally shed from the microsporangium. They are light growth of the female gametophyte progresses very slowly.
in weight and bear two wings that may facilitate dispersal Several months are required in most conifers, and 13
by wind. months are required in pine.
The development of the female gametophyte takes place
Ovulate (Seed) Cone entirely within the ovule. There are approximately 1

The ovulate cone, when mature, is the well-known cone of mitotic divisions of the megaspore before cell walls begin

pines and other conifers. Each cone is composed of an to appear between the newly formed nuclei. Walls do

axis upon which are borne several woody scales in a gradually form, however, resulting in a small mass of

spiral fashion. Ovulate cones develop at tips of young gametophytic tissue, completely enclosed by the diploid
branches in early spring (Fig. 15.13). Two ovules, each cells of the ovule. The adjacent sporophytic tissue of the
enclosing a single megasporangium, develop on the upper ovule, the nucellus, is in part digested by the developing
surface of ovuliferous scales (Figs. 15.12, 15.13). female gametophyte.
The ovules first appear as small protuberances on the While cell walls are being laid down in the developing
upper surface of this scale, close to the axis of the cone. female gametophyte, two or more archegonia differentiate
A protective layer of cells, the integument, develops early at its micropylar end. Figs. 15.12 and 15.14 show that the
on the outer surface of the ovule. In the end of each ovule ovule at this stage consists of integuments, nucellus, and
near the axis of the cone, there is a small opening, the female gametophyte containing several archegonia, each
micropyle, through which pollen grains may enter. with its enclosed egg. Directly beneath the micropyle is a
In pine, one megasporocyte lies in the center of each space, the micropylar chamber. Nucellar tissue lies
ovule (Fig. 15.12); several are contained in ovules of between the micropylar chamber and the archegonia.
redwoods and cypresses. Like microsporocytes, the
megasporocyte is surrounded by a nutritive tissue called Pollination
the nucellus (Fig. 15.12). The nucellus is in reality a
megasporangium because surrounds the area where
it It will be recalled that pollination in flowers is the transfer
megaspores arise. of pollen from anther to stigma; in conifers, it is the

Division Coniferophyta

317
 Figure 15.1 1 Staminate cones of P/nus. A
end of a branch with
a cluster of staminate cones holding ripe pollen. B, longitudinal
section of one staminate cone, showing microsporangia attached
to the underside of each scale with pollen grains inside. C,
scanning electron micrograph of pollen grains (probably shrunken
from treatment in the SEM).

transfer of pollen from staminate cone to ovulate cone.

Conifer pollen is windblown. In many species, ovulate
cones are borne on higher branches of the tree and
staminate cones are concentrated on lower branches;
since pollen does not blow directly upward, cross
pollination is usual.
Pollination occurs in most conifers in early spring soon

after the ovulate cone emerges from the dormant bud

(Table 15.1) — about the time of meiosis. At this age,
scales of young cones turn slightly away from the axis so
that pollen grains can sift down to the axis of the cone.
Here they come in contact with a sticky substance
secreted by the ovule. As this material dries, it draws
some pollen grains through the micropyle into the
micropylar chamber where they lodge close to the
developing female gametophyte (Fig. 15.13). The pollen
grain germinates and develops slowly into a tubular male
gametophyte as grows through the nucellus. Thus, male
it

and female gametophytes of conifers develop to maturity

in close proximity within the ovule. They are both

mm
dependent on nucellar tissue for nourishment and

WMT %
protection.
Several nuclear divisions occur
walls are formed. Two
in the tube, but no
of the last-formed nuclei are
nuclei. This branched pollen tube, containing two
sperm
sperm
cell

nucleiand several vegetative nuclei, is the male

gametophyte (Fig. 15.12).
-«*A
Fertilization and Embryo Formation
Development male and female gametophytes is so
of
coordinated that the egg is formed and ready for

fertilization when the pollen tube, containing two sperm

nuclei, has reached the archegonium. In pine, this

development requires about one year. At the time of
• " i 7 fertilization, pine cones are generally green and the scales
'"V tightly closed, showing a spiral pattern of arrangement

(Fig. 15.13). The sperm nuclei, together with other

protoplasmic contents of the pollen tube, are discharged

chapter 15 |
Gymnosperms

318
 ,

directly into theegg cell. Sperm nuclei do not possess becomes packed with food to be used not only for the
ciliaand hence are not actively motile. One sperm nucleus growthof the embryo but also as a reserve in the seed.
comes in contact with the egg nucleus and unites with it. It will be recalled gametophyte usually
that the female
The nonfunctioning sperm nucleus and the other contains two or more archegonia. Since the egg in each

protoplasmic material discharged into the egg cells soon archegonium may be fertilized and since each of the four
undergo disorganization. embryo-forming cells may give rise to an embryo, eight or
The formation of the embryo is preceded by the more embryos may develop in every seed. Normally,
development of a relatively elaborate proembryo. This however, only one embryo survives, but seeds with two
structure consists of four tiers of four cells each (Fig. well-formed embryos are not rare.
15.12). The four cells farthest from the micropylar end of The mature embryo consists of several cotyledons or
the proembryo may each develop into an embryo. The seed leaves, epicotyl, hypocotyl, and radicle or
intermediate cells are the suspensor cells; they elongate rudimentary root (Fig. 15.12). The embryo is embedded,
greatly and push the embryo cells deep into the female as previously mentioned, in the enlarged female
gametophyte (Fig. 15.12). While this development is taking gametophyte. Both embryo and female gametophyte are
place, the female gametophyte continues to grow, surrounded by a papery shell and a hard protective seed
digesting most of the rest of the nucellus. It enlarges and coat, both formed from the integument of the ovule. The

Table 15.1
A Generalized Life History of a Pine

Pine tree (sporophyte) Time of occurrence

Staminate cone Ovulate cone Summer, year

Microsporocytes Megasporocytes

Meiosis Spring, year 1

Microspores Megaspores

Pollen
(young male
gametophyte)

Pollination Early summer,

year 1

Pollen tube
(mature male) Female
gametophyte) gametophyte

Sperm nuclei Egg nucleus

Fertilization Late spring, year 2

I
Zygote

I
Proembryo Summer, year 2
\
Embryo
(
Seed Fall, year 2

Division Coniferophyta

319
 ovule v

integument

Figure 15.12 States in the life cycle of a pine, A, young ovulate

cone; 6, scale from ovulate cone showing two ovules on upper
surface; C, section of ovulate cone showing megasporocyte cell
and pollen in pollen chamber; D, staminate cone; E, scale from
staminate cone; F, cross section of staminate scale; G, winged
pollen grain;H, development of male gametophyte in pollen grain;
/, pollen tube penetrating nucellus; J, formation of linear tetrad of
megaspores; K, female gametophyte with egg cell and pollen
tube; L, suspensor and proembryo sporophyte within female
gametophyte; M, proembryo; N, section of seed showing seed
coats, female gametophyte, and embryo sporophyte; O, winged
seed; P, seedlings.

chapter 15 |
Gymnosperms

320
staminate
strobilus

Division Coniferophyta

321
 m^%

¥/
E

megasporeocyte
open
pollen chamber

Figure 15.13 Ovulate cones of Pinus. A, longitudinal section of

entire cone at the time of pollination, less than 1 cm long; note
the pollen chambers with open micropyles. B, close-up view of
the one ovule from A. C, an ovule with a pollen grain in the pollen
chamber. D, cone at the time of embryo development, scales
closed.
V
bract pollen grains

whole structure is the seed (Fig. 15.12). In many pines, years, and some may remain embedded in the old mature
the seeds are winged. cones for six years or more. Heat causes the resin coating
Normally, in pines, the seed matures some 12 months the cones of some species to melt, allowing the cones to
after fertilization; two years intervene between initiation of open and release the seeds. This behavior has
ovules and formation of seeds. Young pine seedlings (Fig. considerable survival value, since large numbers of seeds
15.12) have several cotyledons. A generalized life history are released after fires, and injured trees are thus replaced
shown in Table 15.1
of a pine is by the young ones. In many species, however, seeds are
Gymnosperm seeds may remain dormant for many shed soon after they are mature.

chapter 1 5 |
Gymnosperms
322
 pollen tube

ntegument

L- micropylar chamber

egg eel

archegonium

female gametophyte

Figure 15.14 Section through tip of ovule after pollination but

before fertilization.

The Conifer Life Cycle in Perspective retained in the tissue of the gametophyte parent and its

early growth takes place within this tissue. There is also

The pine life cycle, representative of the Coniferophyta, is additional protection given to developing gametes: the

part of a general life cycle trend from Bryophyta to gametangia have a sterile jacket of cells around the
Anthophyta. Quite simply, that trend is: an increasing gametes.
importance and complexity of the sporophyte generation In the lower vascular plants, the sporophyte becomes

and at the same time a decreasing size and degree of the dominant generation —
dominant in terms of life span,
size, and anatomical complexity. It also becomes an
independence of the gametophyte generation.
As a generalization, thallophytes are part of this trend. independent phase. During the embryo stage, the
In the algae and fungi, the gametophyte generation tends
sporophyte is still attached to and nourished by the

to be dominant; that is, a given species tends to spend

gametophyte, but ultimately it becomes established in the
soil and is from then on independent of the gametophyte.
most of its active life span in the haploid state.
Multicellular gametangia are retained, as is the
Furthermore when sperm and egg (or + and - gametes)
requirement for free water for fertilization.
join to produce a zygote, that cell is not retained or
In the conifers, the size, complexity, and longevity of the
protected within parental tissue. Recall the life cycle of
sporophyte generation become even more extreme, and
Fucus, where the eggs and sperm are released from
the gametophyte generation becomes dependent on the
gametangia and meet in the open water; the zygote then
sporophyte and protection. Both the male
for nutrition
begins dividing into a young plant while still in the open
gametophyte (pollen tube) and the female gametophyte
water.
are parasitic on the nucellus (2n tissue) and incapable of
In the Bryophyta, the gametophyte generation is still
manufacturing their own food or of existing in the open.
dominant and it tends to be more anatomically and
They exhibit very little differentiation, and antheridia are
morphologically complex than the sporophyte.
not recognizable. The gametes are protected to a greater
The sporophyte is still not an independent entity, because
degree than in lower vascular plants in that the egg and
it on the gametophyte for nutrition during its
partly relies
the pollen tube that carry sperm to egg always lie within
entire life. In gametophyte is photosynthetic
contrast, the
surrounding tissue.
and completely independent and can reproduce asexually
Pollination is traditionally regarded as an adaptation to a
to continue growth indefinitely (indeed, in some mosses,
dry, terrestrial habitat. Furthermore, free
water is not
the sporophyte generation is rarely seen). Cells that
required for Another innovation is additional
fertilization.
resemble tracheids and sieve cells are present in some protection for the embryo by its enclosure in a dispersal
species. There is more protection given to the young package (the seed) complete with stored food and a hard
sporophyte (embryo) than in thallophytes: the zygote is
seed coat.

Division Coniferophyta

323
 high, high
(Fig. 15.2) that are divided into two lobes. The sperms are
_..( >.
flagellated. In the United States the Ginkgo is commonly
j
planted as a city sidewalk tree.

3
31

Division Gnetophyta
|

i ' \'
I (.1

3
/ "q This division consists of only three genera, and in some
3 respects their traits place them as intermediate between
1

3
the gymnosperms and the angiosperms. For example, they
>

!
contain vessels in the xylem, the ovules appear to be

low_
"'•••()
surrounded by two integuments, and the pollen-producing
T T T
structures superficially resemble stamens. All these traits

Thollophytes Bryophytes Lower Conifers are shared by angiosperms and absent in other
vascular
plonts
gymnosperms, yet true flowers and fruit are absent
seeds are still borne naked.
Figure 15.15 Life cycle trend from thallophytes to conifers,
highly diagrammatic. Sporophyte, solid line; gametophyte, dotted Only one genus, Ephedra (joint-fir, Mormon tea), is
line. found in North America. Its 40 species are distributed in

warm temperate parts of the Mediterranean region, India,

China, the southwestern deserts of the United States, and
Figure 15.15 summarizes the shift in life cycle patterns mountainous parts of South America. It is a shrubby
that we have seen from thallophytes to conifers. xerophyte, with whorls of small deciduous leaves at
prominent joints; most photosynthesis is accomplished in

the green stems (Fig. 15.17).

Division Cycadophyta The other genera look quite different from Ephedra and
exhibit important life cycle differences. Their inclusion into
one division is a matter of convenience for classficiation
Some 200 million years ago members of the Cycadophyta
and the resulting division is quite artificial. Welwitschia is
formed an extensive portion of the earth's flora, probably
another xerophyte, distributed along the southwest coast
constituting the food supply for at least some of the
of Africa. It has two long, straplike leaves that trail along
herbivorous dinosaurs. Today, there remain 9 to 10
the soil surface (Fig. 15.18) Gnetum is a tropical genus of
genera of cycads with about 100 species growing in
30 species, mainly climbing lianas. The leaves look very
widely separated areas of the earth's surface but largely
much like typical dicot leaves.
confined to the tropics. Only one genus, Zamia, occurs
naturally in the continental United States; it is found in

southern Florida. Various genera are, however, cultivated Summary

outdoors in warmer regions, and in greenhouses
elsewhere. 1. The gymnosperms are higher vascular plants and
Two cycads, Cycas revoluta and Dioon spinulosum, are possess both vascular tissue and seeds. Seeds aid
shown in Fig. were grown in the conservatory
15.16. Both plant dispersal, protection of the embryo, and

at Golden Gate Park, San Francisco. These trees are

in establishment of the young sporophyte. A second
palmlike in appearance; in fact, Cycas revoluta is known advancement is pollination, which permits sperms to
as Sago palm. In Zamia, the genus native to Florida, the reach eggs while surrounded by protecting tissue and
trunk is largely subterranean (Fig. 15.16), but in Cycas it in the absence of free water.
forms a single straight bole. The trees grow slowly, a 2 m 2. Of the four divisions of gymnosperms, Cycadophyta,
high specimen being perhaps as much as 1000 years old. Ginkgophyta, Gnetophyta, and Coniferophyta, the latter
That these trees are really gymnosperms and not palms contains the most species, is the most widespread, and
is readily evident from their large cones (Fig. 15.16). A contributes the most to modern vegetation types. The
primitive feature of the division is the presence of Coniferophyta contains seven families, several of which
flagellated sperm. were discussed. The Pinaceae contains pines, firs,
spruces, hemlocks, Douglas fir, larch and true cedars

(Cedrus spp.). The Cupressaceae contains junipers,

Division Ginkgophyta cypresses, and false cedars. The Taxodiaceae contains
bald cypress, redwood, and Sierra bigtree. The
Taxaceae contains the yews. Other families are better
Only one living representative, the maiden-hair tree represented in the southern hemisphere.
(Ginkgo biloba), remains of this very ancient division of 3. Pinus (pine) life cycle was examined as representative
plants. It has been reported as growing wild today in of the Coniferophyta. The sporophyte is large and
forests of remote western China. Ginkgo has, however, anatomically complex. Vessels are absent, but
been grown for centuries on Chinese and Japanese considerable secondary xylem is produced. Leaves are
temple grounds and is now cultivated in many countries. It modified for arid conditions. Pine is heterosporous, the
is a large tree with characteristic small, fan-shaped leaves microspores being produced in small (staminate) cones

chapter 15 |
Gymnosperms

324
 subterranean
stem

Figure 15.16 Cycads. A, Cycas revoluta showing a

microsporangiate cone. 6, Dioon spinulosum showing ovulate
cone. C, Zamia showing ovulate cone and subterranean stem.

on lower branches. Megaspores are produced in other nucellar cell functions as a megasporocyte, and, of the
(ovulate) cones on upper branches; these cones are at four resulting megaspores, only one survives. It begins
tirst small but over a two year period mature into large, to divide to form the female gametophyte at the same
woody cones. time that pollination (the transfer of pollen from male to

The microspores begin within the

female cone) occurs. At maturity, the female
dividing while still
gametophyte is a relatively undifferentiated,
spore wall. Atter two divisions, and the elaboration of
multicellular, nonphotosynthetic thallus embedded in the
air-filled wings on the outside of the thickened spore
nucellus. At one end are several archegonia.
wall, growth ceases and the immature male
6. If the pollen grain is drawn through the micropyle (an
gametophytes are liberated These are
to the wind.
opening in the integument), it comes to lodge against
pollen grains. Some, by chance, sift down into young
the nucellus. It germinates and a pollen tube begins to
ovulate cones and are sealed in when the cone scales parastically grow toward the female gametophyte
close. through the nucellus. Finally, a third cell division
Two ovules are present on each scale in the ovulate produces two sperm and the formation of the male
cone. An ovule consists of an outer integument, gametophyte is complete. No antheridium is
nucellar tissue, and the female gametophyte. One recognizable.

Summary

325
Figure 15.17 Ephedra vindis, a member of the Gnetophyta. A, a
portion of the woody shoot; B, a close-up of A, showing two
scalelike leaves attached to a young, green stem.

7. Fertilization is accomplished when the pollen tube dominance and complexity of the gametophyte have
ruptures near an archegonium and one sperm unites declined.
with an egg. The and
resulting zygote divides The Cycadophyta contain about 100 species of tropical,
an embryo. A mature embryo lies
differentiates into slow-growing, palmlike plants. The Ginkgophyta
surrounded by female gametophyte tissue, some consists of a single species that may now be extinct in
remnants of the nucellus, and a seed coat (from the the wild but is widely cultivated. The Gnetophyta exhibit
integument). The embryo is dormant until the seed is some traits that link them more closely to the flowering
shed and comes to lie in an appropriate plants than to other gymnosperm groups. Only one of
microenvironment. its three genera, Ephedra, is found in North America.
8. In general, life cycles from thallophytes to conifers have This latter division is quite artificial in terms of
shown a definite trend or pattern: thedominance and classification.
complexity of the sporophyte have increased and the

chapter 15 l
Gymnosperms

326
 .

^s#ft&*

r-
B
i^'^." .'_-_

mego porongiate
=

cones

bracts

microsporangiate
cones

We/wi'fschio mirabilis

Figure 15.18 Welwitschia, growing in the Namib Desert of

southwestern Africa. A, one of the larger plants, about 1 .5 m tall
and carbon dated at 1 500 years old; S, closer view of smaller
plant, showing the two leaves emerging from the nearly buried
stem; C, male (left) and female (right) cones borne near the
growing base of the long, leathery leaves. The plants are
unisexual.

Sum mary
327
 1 6 CHAPTER

16 angiosperms

The angiosperms, or flowering plants, are the

dominant plants ot the world today. They include
(Fig. 16.1D).

cytoplasm,
Each sperm nucleus, with its associated
a sperm cell.
is

nearly all the crop plants of orchard, garden, and Free water is not needed for fertilization in angiosperms;
field.Hardwood forests, shrublands, grasslands, and motile, ciliated sperms are not produced by any members

deserts are composed chiefly of flowering plants. They of this division. The pollen tube with sperm cells and tube

show great variation in form, from simple stemless, free- nucleus, present, constitute the mature male
if

floating duckweed (Lemna) through a whole series of gametophyte.

herbaceous types to shrubs and trees such as oaks and
beeches (Fagus). We have already studied the structure
and physiology of the angiosperm plant body in Female Gametophyte
considerable detail. Angiosperms constitute the
Anthophyta, which is divided into two classes, An enlarged cell, the megasporocyte, within nucellar
Monocotyledonae and Dicotyledonae. young ovule undergoes meiosis and forms
tissue of a four

The outstanding and unique structure of the megaspores arranged in a row. While this process is
angiosperms is the flower (Fig. 16.1). The flower is a occurring, integuments form about the nucellar tissue,

shoot bearing floral leaves. In a complete flower (Fig. resulting in the formation of an ovule. The three

16.1), the floral leaves are sepals, petals, stamens, and megaspores closest to the micropyle generally
carpels. Some flowers have only the essential reproductive disintegrate. The remaining megaspore undergoes three
successive mitotic divisions, resulting in seven cells of the
structures, stamens and carpels. Other flowers are
unisexual (Fig. 16.7): they have either stamens or carpels. embryo sac or female gametophyte: one egg cell, two
All flowering plants produce seeds except some that
synergid cells, one endosperm mother cell with two polar
human beings have modified so that they are now nuclei, and three antipodal cells (Fig. 16.1 A).

seedless (e.g., seedless grapes and bananas). In

angiosperms, seeds are borne within a closed structure,
the ovary, which eventually becomes the fruit. Fertilization and Seed Development
With penetration of the pollen tube into the mature embryo
sac, the stage set for fertilization. In angiosperms,
Review of Life Cycle fertilization
is

involves not only the union of the egg cell with

the sperm cell but, in addition, the union of a second
The life cycle of the angiosperms was thoroughly sperm cell with the endosperm mother cell to form the
discussed in Chapter 7. At this point let us simply review primary endosperm cell. Since there are two nuclei in the
its most important aspects. endosperm mother cell, the nucleus of the primary
endosperm cell will have three sets of chromosomes. The
fusions of two sperm cells, one with the egg, the second
Male Gametophyte with the endosperm mother cell, is double fertilization.
The resulting zygote generally develops directly into a
The anther is the part of the stamen responsible for small proembryo, from one end of which a typical embryo
production of pollen. Microsporocytes form within the develops. The fate of the primary endosperm cell depends
pollen sac (Fig. 16.1). They divide by meiosis to form on the species. This cell gives rise to an endosperm that
haploid (1n) microspores, which in turn divide mitotically plays some role in nourishing the developing embryo. In a
to form a generative and a tube nucleus (Fig. 16.1 Q. A few genera, notably the grasses, the endosperm enlarges
heavy, sculptured wall forms about the microspore and a and persists in the seed to form the source of nourishment
mature pollen grain results. The pollen is now shed and for the young seedling. A seed always includes an embryo
conveyed in one fashion or another (Chapter 7) to a and a food supply to enable the young seedling to
stigma where germinates. A pollen tube penetrates the
it establish itself, except in rare cases like orchids where the
stigma (Figs. 16.1 AD) and grows through the style, down embryo remains quite rudimentary. Food is generally
to the ovule within the ovary. The generative nucleus stored either in cotyledons of the embryo or in endosperm.

usually divides within the pollen tube into two sperm nuclei With the germination of a seed and the development of

328
 perianth
I corolla

Figure 16.1 A, diagram of a flower; B, cross section of an

anther; Cmature pollen grain; D, germinating pollen grain.

Review of Life Cycle

329
Table 16.1 thallophytes, bryophytes, lower vascular plants, and seed
Generalized Life History of an Anglosperm plants appears in Fig. 15.15.

Plant (sporophyte) In all Pterophyta the sporophyte is the dominant

generation, and both gametophyte and sporophyte
Stamen Carpel generations are independent. In gymnosperms and
(microsporophyll) (megasporophyll) angiosperms, there occurs a further reduction in size and
I \ complexity of the gametophyte to a condition of complete
Pollen sacs of anther Nucellus of ovule dependency on the parent sporophyte. This is
(microsporangia) (megasporangium) accompanied by an overall increase in the general
t \
complexity of the sporophyte. can be stated that, as a
It

Microsporocytes Megasporocytes guiding principle, the more advanced forms in these

divisions are more highly adapted to grow and flourish in a
(Meiosis) dry land habitat than are the more primitive forms. The
most obvious developments designed to accomplish this
Microspores Megaspores are: (a) adaptations thatremove any dependence of the
plants on free moisture for fertilization, and (b)
I

Pollen adaptations, both vegetative and reproductive, that enable

(young male the plant to grow and reproduce on dry land. Examples
gametophyte) are given below.

(Pollination) Protection of the Female Gametophyte

Pollen tube Embryo sac The gynoecium of the angiosperm flower is essentially a

(mature male (female modified leaf, or leaves, enclosing an ovule, or ovules, in

gametophyte) gametophyte) which the female gametophyte is be found. The ovary,

to

Sperm
/
cell
\Sperm cell Egg
/
cell
\
Endosperm
as we know, eventually develops into a fruit. This
enclosure of ovules and, subsequently, seeds by an ovary
mother cell wall is a new development that sets angiosperms apart

\ from all other groups of plants. The female gametophyte

(embryo sac) has gained, in the ovary wall, an added

(Double fertilization)

Zygote
/ N Primary
protective barrier. However, this has necessitated further
adaptations to enable sperms to pass through this added
protection: (a) development of a receptive stigma and (b)
endosperm cell
growth of a pollen tube through the style.

Proembryo
I
Size of Gametophytes
Embryo Endosperm
Reduction in size of gametophytes has proceeded to a

point in which the male gametophyte, the pollen tube,

Seed
consists of one vegetative nucleus and two sperm cells
t
(Fig. 16.1D). The female gametophyte is a seven-celled
Plant (sporophyte)
structure, one of whose cells is an egg cell. A second cell,
the endosperm mother cell, is an innovation in that it, too,
is receptive to a sperm cell. The remaining five cells are

vegetative cells (Fig. 16.1 A). It should be pointed out that

the seedling into a flowering plant, the life cycle of an embryo sacs of some angiosperms have more cells, while
angiosperm is completed. The essential steps are shown a few of them have less, but all have an egg cell and an
in Table 16.1. endosperm mother cell.

Comparison Angiosperm of Nourishment for the Developing Embryo

Life Cycle with more Double fertilization is still another difference between the

Primitive Plants life cycle of the angiosperms and more primitive forms. In

the latter, nourishment for the young developing

sporophyte is generally provided by the female
Certain comparative details between structures and life gametophyte (Figs. 14.10M,/V) or, in the case of many
cycles of members of the gymnosperms and angiosperms ferns, by the simple prothallus, or gametophyte (Fig.
are given in Table 16.2. Details concerning the Psilophyta, 14.18J). In angiosperms, food for the developing seedling
Lycophyta, Sphenophyta, and Pterophyta are given in is stored either in the endosperm or in the cotyledons of

Table 14.1. A general comparison of life cycles in the embryo itself.

chapter 16 |
Angiosperms

330
Table 16.2
Comparison of the Conifers and Flowering Plants

Generation Gymnosperms Angiosperms

Roots, stems, and leaves present Roots, stems, and leaves present
Trees and shrubs, no herbs Trees, shrubs, and herbs
Stem: one core of xylem surrounded by phloem, Stem: one core of xylem surrounded by phloem, or
vessels inone small division only several vascular strands with phloem exterior
Sieve cells without companion cells Companion cells and sieve-tube members
Pith present in stems, not in roots Pith in stems, not in roots, of dicots; in some
monocot roots
Sporophyte Cambium develops in all species Cambium in perennial forms; absent generally in

annuals
Staminate and ovulate cones, generally, with Flowers, stamens, and carpels
staminate and ovulate scales
Heterospory Heterospory
Microspores in microsporangia Microspores in anther sacs (megasporangia)
Megaspores or nucellus Megaspores in nucellus
Ovules present, exposed on scales Ovules present, enclosed within carpel
Embryo develops within a seed Embryo develops within a seed
Fertilization Double fertilization absent; no endosperm Double fertilization, resulting in zygote and primary
endosperm cell

gametophyte
Pollen tube: male Pollen tube: male gametophyte
Female gametophyte within ovule Female gametophyte (embryo sac) within ovule
Antheridium absent Antheridium absent
Sperm cells formed in pollen tube Sperm cells formed in pollen tube

Gametophyte Motilesperms in several primitive genera, but free

water not needed for fertilization
Nonmotile sperms in all other forms Nonmotile sperms
A very greatly reduced archegonium completely Archegonium only suggested by synergid cells of
embedded in the female gametophyte the embryo sac
Embryo within female gametophyte and seed coat Embryo within seed coats, remains of nucellus,
and endosperm

Fruit themore advanced and specialized forms have

presumably arisen (Chapter 10).
Fertilization stimulates tissues around the zygote to begin
further development. In angiosperms, the ovary wall is

stimulated to produce a and the fruit is a

fruit, The Besseyan System
development originating with, and characteristic of,
angiosperms. The protection of the seed from its
Many systems of classifying plants have been proposed. A
system that has found much favor with American botanists
surroundings by the ovary wall is accompanied by various
devices for bringing it under conditions favorable for
was developed by Charles E. Bessey (1900s). His system
regards the order Ranales as being the most primitive. In
germination.
this order, the Magnoliaceae is one of the most primitive
families, with the Ranunculaceae somewhat more

Evolution in the Angiosperms advanced. The Christmas rose (Helleborus), magnolias

(Fig. 16.3), and buttercups (Ranunculus) are

representatives of these families.

Ifangiosperms represent the culmination of an extensive
If we consider the arrangement of flower parts in
evolutionary development, it is likely that within the
magnolias and buttercups to be most primitive, then it is
division anthophyta itself, evolution has been and still is
possible to compare them to other families to determine
active. In other words, families of angiosperms must be
their apparent relationships. The principal tendencies in
related and should be possible to arrange the families in
it
evolution of the flower, according to the Besseyan system.
a sequence that would give some indication of
are as follows:
relationships. Some families will be found to be more
primitive than others; and from the more primitive types 1. From an elongated to a shortened floral axis.

Evolution in the Angiosperms

331
2. From a spiral to a whorled condition of tloral parts. complete our discussion with the line of
3. From numerous and separate stamens and carpels to monocotyledonous families. should be further noted that
It

few and connate stamens and carpels. lines of development are not straight but branched and
4. From numerous and separate sepals and petals to few treelike. The arrangement of the families discussed here

and connate sepals and petals. as they occur in the Besseyan system is shown in Fig.
5. From complete and perfect flowers to incomplete and 16.2.
imperfect flowers.
6. From regular to irregular flowers.
7. From hypogyny to epigyny.
Magnoliaceae to Lamiaceae
According to the Besseyan view, there were at least In this line of ascent (Fig. 16.2) all flowers, in all families
three main lines of advance from the primitive Ranalian are hypogynous. Syncarpy (the fusion of carpels)
type of flower. These lines culminated in (a) mints occurred very early, as did a reduction in number of floral

(Lamiaceae), (b) asters (Asteraceae), and (c) orchids parts. Other types of connation and the change to
(Orchidaceae). irregular flowers appeared somewhat later but still early in
the line of ascent. Based on their floral characteristics,
primarily, is possible to show an evolutionary connection
it

Selected Families of between the following families: Magnoliaceae,

Ranunculaceae, Malvaceae, Brassicaceae, Solanaceae,
Angiosperms Salicaceae, and Lamiaceae (Fig. 16.2).
The Malvaceae is a clearly marked family with relatively
primitive characters as shown by its regular, perfect, and
Let us now examine selected families of angiosperms with
hypogynous flowers. There are numerous stamens with
two objectives in mind: (a) to learn the characteristics of
connate anthers, and the five or more carpels are
some important angiosperm families and (b) to discover
connate. In Solanaceae, the flowers, while perfect and
how these families fit into the Besseyan system of
regular, show connation of sepals, petals, and carpels,
angiosperm classification. In our discussion of these
with stamens adnate to the corolla tube. The Salicaceae is
matters so far, floral characteristics have been
a family in which reduction in both number and size of
emphasized. The emphasis is placed on floral parts
floral parts has resulted in its advanced condition. The
because their form and structure are little influenced by
flowers are imperfect and grouped in catkins, and both
the external environment. They also are structurally more
calyx and corolla are lacking. The pistillate flowers are
constant from generation to generation than are vegetative
parts.Because of this constancy in structure, floral parts hypogynous and, in staminate flowers, the number of
stamens has been reduced to one or two. The Lamiaceae,
have a much greater value in both plant identification and
or mints, represent the most advanced family in this line of
in tracing evolutionary relationships than do vegetative
ascent. The flowers are hypogynous, with an irregular
characters. However, the latter cannot be neglected.
symmetry; there is a reduction in number of parts to four
Since the magnolia and buttercups are thought to be
or two; and sepals, petals, and carpels are connate. The
primitive forms, we shall begin our discussion with these
stamens are not only adnate to the corolla tube but are
families, then proceed to follow through a line of ascent
also highly specialized for insect pollination. Let us
ending in the mint family. We shall consider the line
consider some of the general characteristics of these
culminating in the Asteraceae (Compositae)* and
families.
* According to rules of plant nomenclature family names should

end in -ceae. Traditionally, the names of certain families such as Magnoliaceae

Cruciferae and the Compositae have not ended this way. In this
chapter the currently more acceptable synonym names ending in This group shares with the Ranunculaceae the distinction
-ceae will be used. of being very ancient. The tulip tree, Liriodendron, as

asteraceae
orchidaceae (compositae)

apiaceae
iridaceae solanaceae
(umbelliferae)

poaceae
cucurbitaceae salicaceae
(graminae)

fabaceae
(leguminosae

alvaceae

magnoliaceae

Figure 16.2 Diagram showing the evolutionary tendencies in the

Angiospermae according to the Besseyan system.

chapter 16 |
Angiosperms

332
 seeds

Figure 16.3 Magnolia grandiflora flower and fruit with hanging

seeds.

shown by remains, once had a very wide

fossil Ranunculaceae
distribution. Most species of the Magnoliaceae are trees or
Most of these are herbs but there are a few small shrubs
fairly large shrubs. Magnolias themselves are magnificent
and woody climbers. Leaves are frequently compound, a
trees, Magnolia grandiflora grows to 30 m or more with
feature that has led to the common name of crowfoot
stiff, large, evergreen leaves and large white blossoms 18
family. There are about 50 genera and approximately 1900
to 20 cm across. The magnolias are largely warm-climate
species growing largely in the North temperate and arctic
species, but tulip trees will tolerate northern winters (Fig.
regions. Many of our common garden flowers belong to
16.3). There are some 12 genera and 230 species* in the Among these are Delphinium, Aquilegia,
this family.
family.
Paeonia, Ranunculus, Anemone, and Clematis. The marsh
marigold (Caltha) and Hepatica, common spring flowers,
* The estimates of genus-species numbers were taken from are members of this family. One species, Aconitum
Willis, J.C., A
Dictionary of the Flowering Plants and Ferns, 8th napellus, yields the drug aconite, a cardiac and respiratory
edition, revisedby H.K. Airy Shaw, 1973, Cambridge University
Press. Note that species numbers from other authors may differ sedative, and some species of Delphinium are poisonous
depending on the system used to calculate their estimates. to livestock.

Selected Families of Angiosperms

333
Brassica haber

Figure 16.4 Brassica haber branch and flower.

Brassicaceae (Cruciferae) viewpoint of politics, agriculture, and industry (Fig. 16.5).

The cotton of commerce occurs as long hair or fuzz on

This is a large, distinct family of many cultivated forms, as
seeds, which are borne in large capsules. Herbs, shrubs
well as some that are noxious weeds. Cabbage,
and and such ornamentals as Hibiscus, okra,
trees,
cauliflower, broccoli, brussels sprouts, kohlrabi, and kale
hollyhocks, and numerous weeds are members of this
are all horticultural varieties of a single species, Brassica
family.
oleracea (Fig. 16.4). Wild mustard, another Brassica
species, is a sometimes noxious weed in grain fields, Solanaceae
though sometimes used as cover crop in orchards. A large family with many tropical forms, it is also well-
Radishes, turnips, and stocks are other members of this represented in temperate regions. There are about 90
family, as are many garden herbs. The family has a genera and over 2000 species, some 1700 of the latter
worldwide distribution in temperate and subarctic zones; being a single genus, Solarium. To this family belong
in

all are herbs. There are about 375 genera and 3200 tomatoes, potatoes, tobacco, eggplant, peppers, Petunias,
species. and Salpiglossis. Although the family is of worldwide
distribution, most forms were brought
of the cultivated
Malvaceae
under domestication in the Western Hemisphere. There
This is a large family of 95 genera and 1000 species. are many poisonous and drug plants in the family, such as
Cotton, taken from seed coats of various members of the belladonna and atropine. Even such a common plant as
genus Gossypium, makes this family important from the the tomato was long supposed to be poisonous, as its

chapter 16 |
Anglosperms

334
 it,, mv /-f %

Figure 16.5 A, Gossypium sp. (cotton) shoot with flowers. S,

splitflower showing carpels. C, dehisced capsule (bole) showing
cotton "fiber," which is actually made of seed coat hairs. D,
cotton seed with attached "fibers."

scientific name Lycopersicon, which means "wolf peach," members. Peppermint, spearmint, thyme, sage, and
seems to indicate (Fig. 16.6).There are many erect and lavender are examples. There are about 180 genera with
climbing herbs in the family, also some shrubs and small 3500 species well-distributed over the surface of the earth.
trees. There are herbs and shrubs in the family, generally with
characteristic square stems and opposite leaves.
Salicaceae
The willow family is comprised of three genera (Salix,
Magnoliaceae to Asteraceae
willows; Populus, poplars; Chosenia, an asiatic shrub),
with about 530 species. They are mostly trees, but some The second line of ascent involves the Fabaceae,
shrubby forms are known. They are very abundant in the Rosaceae, Apiaceae, and Asteraceae, with an offshoot
Northern Hemisphere, mostly in temperate zones. Baskets family, the Cucurbitaceae. This large group of families is
are woven from branches of the basket willow, and paper characterized by an early change from hypogyny to
pulp is obtained from trunks of one species. Willows are epigyny. Following this there is a division into two
common along water courses (Fig. 16.7), frequently sublines, one being marked by a lack of connation and a
overhanging or even choking mountain streams, to the ill retention of regular flowers. In the other subline, connate
comfort of fishermen. Pussy willow is a familiar example of and irregular flowers both occur in advanced forms.
this family. The Rosaceae (rose family) is one of the more primitive
families in this line of evolutionary development. There
Lamiaceae (Labiatae)
exists a considerable variety of flower structure within the
Mints (Fig. 16.8) represent the supposed highest advance family. The more primitive members are regular, perfect,
of one angiosperm evolution. It is a
of the three lines of with numerous parts and with such a slight degree of
large family of considerable economic importance, largely perigyny as to be recognizable only with careful
because of volatile oils produced by certain of its observation. This is exemplified by the boysenberry (Fig.

Selected Families of Angiosperms

335
 ocule

seed

Figure 16.6 Solanaceae. A, branch of Lycopersicon showing

leaf, flower, and buds; B, lontitudinal section of flower of
Lycopersicon; C, cross section of tomato fruit.

16.9). Changes within the family involve a reduction in the characters as loss of floral parts, imperfect flowers,
number of floral parts, a shift from slight perigyny to true epigyny, and connation.
perigyny and to epigyny. Connation also occurs. Members
of the family generally have regular flowers.
Rosaceae
Fabaceae (pea family) are considered to be a more Whereas grasses and legumes supply bread and
advanced family than Rosaceae, even though their flowers vegetables of basic importance, the rose family supplies
are hypogynous. There is a reduction in the number of fruit for dessert and roses for decoration. There are more
parts, the gynoecium having only one carpel; connation of than 2000 species in this family and over
00 genera, not
1

stamens and of some petals occurs, and the corolla parts counting the almost numberless cultivated forms of roses,
are irregular. Apiaceae represent a further advance over peaches, apples, cherries, boysenberries (Fig. 16.9),
Rosaceae in that the flowers are epigynous and exhibit almonds, and so on. The family is of worldwide distribution
syncarpy (Fig. 16.9). This line of ascent finds its climax in and somewhat heterogeneous. Many of the principles of
the Asteraceae, whose flowers all possess advanced angiosperm evolution can be demonstrated with its
characters, such as reduction in number of parts, members. There are trees, shrubs, and herbs in the family.
and some
modification of parts (pappus), irregular flowers, Leaves are usually alternate and bear stipules.
imperfect flowers, connation each whorl, and adnation.
in
Cucurbitaceae
The Cucurbitaceae culminate a branch line from the
rose order. Flowers of Cucurbitaceae have such advanced The gourd or melon family is of worldwide distribution in at

chapter 16 |
Angiosperms

336
 gland

Figure 16.7 Willow (Salix) inflorescence and individual flowers;

A, male catkin, x 1; B, single male flowers, X15; C, single female
flowers, X15.

of Western Hemisphere origin. Cucumbers and melons

have probably come from Africa and central Asia. Other
species appear to have been first cultivated in the tropics
of Asia, Polynesia, and India. They frequently use tendrils
to climb and are annual or perennial vines. Most are rapid-
growing and frost-tender. There are about 1 10 genera with
640 species. Stems are usually soft and hairy or prickly.
The generally simple leaves are large and sometimes
deeply cut. Flowers are usually unisexual.

Fabaceae (Leguminosae)
The legume family has such distinctive characteristics that
its members can frequently be recognized with only little

experience one of the three largest

(Fig. 16.1 1). It is

families, with 600 genera and 12,000 species. All major

growth forms are represented: herbs, both annuals and
perennials, shrubs, vines, and trees. It is of worldwide
Figure 16.8 Lamiaceae. Irregular hypogynous flower of mint, distribution. While less heterogeneous than the rose
showing union of sepals and petals. family, considerable variation is found among its various
members. It is a family of considerable importance in

supplying food for humans and their animals. Many

least warmer regions of the world, and various peoples legumes are used for ornaments, from shade trees to cut
have selected different forms for domestication (Fig. flowers. Some lumber is obtained from the black locust,

16.10). It is also probably the only group in which fruits and some of the tropical species furnish wood for fine
are highly prized for ornamental purposes and for use as cabinet work. Association of the nitrogen-fixing bacteria
containers of various types. Pumpkins and squashes are with roots of legumes places this family in a unique

Selected Families of Angiosperms

337
 Rubus

Figure 16.9 Rubus sp. (boysenberry) flowering branch with one fancy vinegar and nectar for desert honey. Leaves are of
flower split open to show the fleshy receptacle.
various shapes.

position relative to maintenance of soil fertility. Peas, Monocotyledonous Line from

beans, peanuts, clovers, and lupines are common Magnoliaceae to Orchidaceae
herbaceous legumes; wisteria is a vine, brooms and
redbuds are shrubs or low trees, and locusts and acacias There are many obvious differences between the
are trees. The Mimosa genus alone has some 450 dicotyledonous plants such as the Magnolias and the
species, including the sensitive plant of the florist shop; monocotyledonous line of evolution. These have been
there are others of varying habit from tall trees to low discussed and are briefly reviewed in Table 16.3.
herbs. Leaves of legumes are prevailingly pinnately
compound and quite generally with stipules. Sometimes a
leaflet will be modified to a tendril.
Table 16.3
Tabulation of Differences between Monocotyledons
Asteraceae (Compositae)
and Dicotyledons
The Asteraceae is the largest family of angiosperms (Fig.
16.12). Only a few of these plants are woody. In this Monocotyledons
herbaceous plants there are around 900 genera
family of
and about 13,000 known species. The family is not only of 1. One cotyledon or seed leaf.
worldwide distribution but in most places is relatively 2. Generally marked parallel leaf venation.
abundant. The family is not noted for its food plants; 3. Flower parts typically in groups of three or multiples of
endive (Cichorium), artichokes (Cynara), chicory, lettuce, three.
and sunflower form most of its contribution in this respect. 4. Vascular bundles of stems scattered throughout a
Neither is it famous as a producer of drugs or other cylindrical mass of ground tissue.
commercial products. One species, safflower (Carthamus), 5. Vascular cambium lacking in most forms.
is now being grown for oil and members of the genera

Taraxacum and Parthenium are being considered as

Dicotyledons
possible sources of latex for rubber. Paging through a
seed catalog or a visit to a florist shop will show members Two cotyledons or seed leaves.
1.
of this family in their full grandeur. Dahlia,
2. Generally marked netted venation of leaves.
Chrysanthemum, Aster, Zinnia, Tagetes, Gaillardia,
3. Flower parts typically in groups of four or five.
Ageratum and many others are representatives of this
4. Vascular bundles of stems usually arranged in the form
family. Dandelions color lawns, and various wild species
of a cylinder.
add to fall infest of meadows and fields. There are 400
5. Vascular cambium present in forms having secondary
species of the genus Artemisia, that cover arid portions of
growth.
the world and supply, among other things, tarragon for

chapter 16 I Angiosperms

338
 filaments

Figure 16.10 Cucurbitaceae. A, Cucurbita pepo, flowering and

stem; B, C. pepo, fruit; D, C. maxima, staminal column; D,
fruiting
C. maxima stigma, longitudinal section through pistil, corolla tube
removed; E, C. maxima, cross section through ovary; F, C.
maxima, style and stigma.

However, some primitive monocotyledons, particularly characters of grasses include hypogyny, regular flowers,
certain water weeds such as arrowheads and water and only connation of carpels. The loss of perianth parts
plantains, have much in common with buttercups and and the reduction in number of stamens and carpels mark
marsh ^marigolds. The families selected to represent the them as a more advanced family than the Liliaceae.
monocots are Liliaceae, Poaceae, Iridaceae, and
Liliaceae
Orchidaceae. Lilies differ from magnolias in all distinctive

monocot characteristics. In addition, they show a Unlike grasses, lilies are grown largely for ornamental

reduction in the number of floral parts and a connation of

purposes, although two genera, Allium (e.g., onions,

show a single advance over lilies in that

carpels. Irises garlic, and leeks) and Asparagus, are grown extensively
for food. Some species yield drugs, and others have
they are epigynous. Many irises are regular, perfect, and
with separate perianth parts. Stamens show no connation. poisonous properties that may cause trouble in pastures or
Syncarpy occurs as does in lilies. Orchids show
it
on ranges of western states (Fig. 16.13). There are about
evolutionary advance in that they have irregular flowers
4100 species in some 280 genera. Most lilies grow from a
very highly specialized for insect pollination. Stamens have bulb or a bulblike organ and flower in a single growing

been reduced season, after which the shoot dies down. A few, such as
number to one or two.
in

The order which grasses belong may have arisen

to
Joshua trees, are woody perennials. Tulips, hyacinths, day
lilies, and aloes are other examples of the lily genera.
from the order to which lilies belong. The primitive

Selected Families of Angiosperms

339
 anthers

Figure 16.11 Fabaceae. A, inflorescence and leaf of Lathyrus;

B, exploded view of Lathyrus corolla; C, essential organs of
Lathyrus flower; D, cross section of ovary; E, pod of Lathyrus.

Iridaceae

The iris family contains about 60 genera and 800 species.

They are all herbaceous, largely perennial forms, usually
with rhizomes, bulbs, or corms, as in lilies. Leaves and
flowering stalks last for only one season. Some of the
choicest florists' plants occur in this family Iris (Fig.
receptacle
16.14), Gladiolus, Freesia, Crocus, Watsonia, and so forth.
bracts of involucre
Orchidaceae
It is agreed by all that orchids represent the most

advanced family in the Monocotyledoneae (Fig. 16.15).

They are all herbaceous, occurring throughout the world
largely in the tropics, but with a few genera extending into
colder temperate regions. There are some 1 7,000
recognized species in several hundred genera, making this
one of the three largest families of angiosperms. All forms
are perennial, with tuberous, bulbous, or otherwise
thickened roots. They may be erect, prostrate, or climbing.
A few are saprophytic and lack chlorophyll; others are
bracts of involucre epiphytes, growing on trees without benefit of soil.

Poaceae (Gramineae)
Humans and other mammals have been associated with
Figure 16.12 Composite flower and fruit of sow thistle
(Sonchus). A, flowering head; B, single ray flower; C, mature the grass family for far more years than those of recorded
achenes. history. It is probably not an overstatement to say that

chapter 16 |
Angiosperms

340
 ovule

Calachotius luieus

Figure 16.13 Calachortus luteus. A, flowering branch; 6, flower;

C, cross section through ovary.

Selected Families of Angiosperms

341
 petals

Iris douglasiana

Figure 16.14 Iris douglasiana. A, entire flowering plant; 6, flower

showing interior ovary; C, section through ovary.

chapter 16 |
Anglosperms

342
sepals

Figure 16.15 Cattleya sp. (Orchid).

Selected Families of Angiosperms

343
without the grass family human civilization, as we know it, vegetative characteristics of grasses have been described
could not have developed. Different grasses were in some detail in Chapter 7 and shown in Fig. 7.6.

domesticated by the three centers of civilization; wheat,

rye, barley, and oats by peoples of the Mediterranean Summary of Angiosperm Evolution
region and Southwest Asia; corn by natives of the Western Angiosperm
1. families, according to the Besseyan system
Hemisphere; and and millet (Eleusine) by peoples in
rice
of classification, are all derived from the primitive order
the Orient. Grasses are grown for human and animal Ranales, to which buttercups and magnolias belong.
consumption, for ornament, and for uses in arts and 2. Important changes in floral evolution, according to the
industry. There are over 620 genera and about 10,000 Besseyan system, are as follows: (a) from a spiral to a
species. Most grasses are herbaceous, either annuals or whorled arrangement of floral parts; (b) from many
perennials; a few bamboos are climbers and some are parts to few or even to a loss of parts; (c) from separate
woody and up to 20 m tall. Grasses of one kind or another floral parts to connate parts; (d) from a regular flower to
grow in all kinds of soil and situations. Grasses may be an irregular flower; (e) from hypogyny to epigyny.
conveniently divided into six groups according to their use. 3. According to the Besseyan system of classification
(a) Bamboos, mostly evergreen, stout perennials, are used there are three branched lines of ascent from the
for construction in many parts of the Orient, and some Ranales.
have been introduced into warmer parts of the United 4. The first line of ascent following the Ranales includes:

States as ornamentals, (b) Cereals and some other annual Malvaceae (mallow family), Brassicaceae (mustard
grasses supply grain and forage for both human beings family), Solanaceae (potato family), Salicaceae (willow
and animals, (c) Sugar-producing species, such as sugar family), and Lamiaceae (mint family).
cane, are strong, upright perennials growing only in the 5. The second line comprises: Rosaceae (rose family),
tropics, (d) Sod-forming grasses, perennials that cover Fabaceae (pea family), Apiaceae (carrot family), and
many square miles of the earth's surface, are used in Asteraceae, with Cucurbitaceae (melon family) as an
lawns and meadows and for forage, (e) Bunch grasses, offshoot from Rosaceae.
common in semi-arid regions, are perennials that do not 6. The third line comprises the Monocotyledoneae in the
form rhizomes and do not produce a turf as sod grasses following order: Liliaceae (lily family), Iridiaceae (iris
do. There is a large group of grasses grown for
(f) family),and Orchadaceae (orchid family). The Poaceae
ornamental purposes, such as pampas grass. Flowers, and (grass family) is an offshoot from the Liliaceae.

chapter 16 |
Angiosperms

344
 APPENDIX

basic ideas of chemistry

The living body

called molecules.
is built of very small units of matter
There are thousands of kinds of
Some of the
below, along with the
elements most prominent
letter symbols
in life

that chemists
are listed

use to
molecules, which differ in size, shape, behavior, indicate their presence in a molecule:
and role in the body. The largest molecules carry
hereditary information and are slender threads that may Name of element Symbol Number of Protons
reach 1 mm in length. But closer to the average size are
molecules of table sugar (sucrose): about three million * Hydrogen H 1

molecules of sucrose would have to be placed end to end, * Carbon C 6

to span the printed word "cube." The living body builds * Nitrogen N 7
nearly all of its own molecules, by rearranging the parts of *Oxygen 8
simpler molecules taken from the environment. The Sodium Na 11
systems in the body that build new molecules are Magnesium Mg 12
themselves collections of molecules. * Phosphorus P 15
Sulfur S 16
Chlorine CI 17
What Are Molecules Made Of? Potassium K 19
Calcium Ca 20
Iron Fe 26
A molecule is built of still smaller units of matter called
atoms. An atom in turn contains three kinds of material
particles, electrons, protons, and neutrons. The The elements that are starred provide almost all the nuclei
electrons in an atom move separately around a group of of biological molecules. These nuclei combine in many
protons and neutrons that are joined together to form a different combinations, along with electrons, to form
single unit called the atomic nucleus. There are many thousands of compounds.
kinds of nuclei, which differ in the numbers of protons and Chemists indicate the composition of a molecule by
neutrons they contain. With rare exceptions nuclei are using the elemental symbols to show the kinds of nuclei
extremely stable. They are never modified by the events present. Subscript numbers indicate how many nuclei
that occur in the living body. there are of each kind. Thus the symbol CH 4 means a
A molecule that contains nuclei of differing kinds is molecule that has four hydrogen atoms and one carbon
called a chemical compound. In contrast a unit of matter atom (the subscript "1 " is assumed no other number is
if

is called an element all of its nuclei contain the same

if
supplied).
number of protons. Several elemental substances were
obtained and given common names before the discovery
of atomic structure: silver, gold, iron, sulfur, oxygen, and
others. The nuclei of an element may differ in the number What Holds the Parts of the
of neutrons they contain, but this difference does not
influence the chemical behavior of the nucleus. The Molecule Together?
neutrons seem to be important chiefly in holding the
nucleus together as a unit. (Nuclei with the same number
of protons but differing numbers of neutrons are called Protons, neutrons, and electrons are far smaller than the
isotopes. An number of neutrons far from
isotope with a atom as a whole. Protons and neutrons are about equal in
the average may be unstable and may spontaneously weight, and both are about 1840 times heavier than an
decompose, a process known as radioactive decay. Such electron. Thus almost all the weight or mass of an atom is
a nucleus splits into parts, releasing much energy. Decay in the nucleus. Nevertheless, the nucleus is extremely tiny.

events are rare in the molecules of life and will not be If an atom were magnified to the size of a house, the
considered further.) nucleus would be about as large as a pinhead. Since the

A1
electrons are also very small, this means that the atom (or Figure A.1

the molecule) is mostly empty space.

The forces that hold the nuclei and electrons together in nuclei

a molecule are electrical. Each electron carries a unit of

negative electrical charge, and each proton has an equal-
no
sized unit of positive electrical charge. Neutrons carry
charge. Two one another they carry
particles attract if
-bonding orbital
opposite charges and repel or push one another away if
the/ carry charges of the same sign. These forces grow
stronger as the particles approach closer. (But the
attraction between an electron and a nucleus turns into a
repulsion they approach too close. This keeps the
if

electrons and nuclei from colliding.)

Because of these electrical forces, each electron and
nucleus in the molecule experiences a combination of
pulls and pushes from all the other particles at all times.
The molecule holds together because the attraction nucleus

between nuclei and electrons is enough to overcome the

forces of repulsion. non-bonding orbital

A nucleus can hold in its vicinity about as many

electrons as it has protons. Therefore most molecules Figure A.2

have an equal number of protons and electrons. Such a
molecule is said to be electrically neutral, because any
oriented. Some spend all their time near a
of the electrons
force that its electrons exert on a distant object is just
single nucleus (Figure A.1). These electrons are said to be
countered by an opposite force exerted by the protons.
in a nonbonding orbital. Other electrons travel between
Though most molecules are electrically neutral, there
two or more nuclei. These are bonding electrons. The
are many kinds of molecules that have more protons than
region of space they used is called a bonding orbital
electrons or vice-versa. These molecules are called ions.
(Figure A.2). The two nuclei that share the bonding
In symbolizing an ion, chemists indicate the number and
electrons are said to be joined by a covalent bond. The
sign of the excess charge with a superscript: for example,
+ sharing of electrons results in a powerful force holding the
K indicates the potassium ion, with one more proton than two nuclei
-2 at a stable distance from one another.
the number of electrons; or S0 4 for the sulfate ion, with
Covalent bonds within a molecule are shown by
two more electrons than protons. Ions with a positive
structural formulas, where the nuclei are represented by
charge are cations; those with a net negative charge,
the elemental symbols and a covalent bonding orbital is
anions. Ions that have the same charge repel one another
shown by a line between the nuclei; for example, C-N.
at a distance; those with opposite signs tend to approach
Electrons that are in nonbonding orbitals are usually not
and stay close together. When two opposite ions are in
shown in the structural formula.
contact, chemists often say they are joined by an ionic
It is important to know how orbitals form around the
bond. and
nuclei of hydrogen (H), carbon (C), nitrogen (N),
oxygen (O), since these make up the bulk of the
What Determines the Shape molecules of life. H is the simplest nucleus, consisting of
just one proton. The charge on the proton is so weak that
of a Molecule? only one orbital forms around the H nucleus. C, N, and O
nuclei have six, seven, and eight protons, respectively,
Nuclei are heavy and take up definite positions in the and they exert a much stronger attraction for electrons.
molecule. Their locations can be shown accurately in Each of these nuclei is the center for five orbitals. Since
drawings or models. But the electrons continually move at these three elements are almost alike with respect to the
high speed around and between the nuclei. Chemists kinds of orbitals that form around them, they are
describe this situation by saying that the nuclei are considered as a group below.
embedded in a cloud of electrons. Thus molecules do not One of the five orbitals is a spherical region that lies
have sharp boundaries. But two molecules repel one very close to the nucleus (Figure A.3). The electrons in

another if they get too close, because the two electron this orbital are nonbonding electrons and are usually
clouds repel one another. The size of a molecule is the
distance at which this repulsion becomes great enough to
prevent an oncoming molecule from moving any closer.
Though electrons do not settle in definite positions, they
have regions where they spend most of their time. The
nucleus (C, N or O)
region where a given electron can usually be found is
called an orbital. The orbitals do not have sharp
inner non-bonding
boundaries but they can be shown in diagrams by means orbital (2 electrons)

of shaded areas.
The nuclei act as centers about which the orbitals are Figure A3

appendix A |
Basic Chemistry

A2
 6

nucleus

w
Figure A.4
outer orbital (has 2 lobes)

9
omitted trom diagrams of molecules. But in studying this
nonbonding orbital we encounter a general rule that
Figure A.
applies to all orbitals: each orbital contains exactly two
electrons when the nucleus is part of a stable molecule.
An orbital cannot hold more than two electrons and, for if enough full-time (unshared) and half-time (shared)
some reason an orbital loses one of its electrons, there is electrons to equal its own charge; and (b) this is achieved

a powerful tendency for the molecule to rearrange itself or in way that places two electrons in-each orbital. The
a
to react with other molecules so that each orbital again same rule applies to all other cases where C occurs in
has two electrons. biological molecules, and it applies equally to H, N, and O.
Most commonly, the four remaining orbitals all have the Notice also that the four bonds made by a C atom tend
same shape: each is approximately dumbbell-shaped, with to be rigidly spaced; the angle between any two of the
the nucleus at the narrow point and with one lobe much bonds is about 109°. This consequence of orbital
larger than the other (Figure A.4). The smaller lobe can be repulsion doesmuch to fix the internal geometry of the
ignored. These be oriented in
four equal orbitals tend to molecule. No drawing of a molecule on a flat page can
space so that their axes are as far from one another as adequately show these angles, so chemists frequently
possible (Figure A.5). This happens because the electrons resort to physical models by using parts such as balls to
of each orbital push those of neighboring orbitals away. represent nuclei and springs or sticks to show covalent
(But actually the outer orbitals overlap more than the bonds.
diagrams indicate, so that electrons shield the nucleus The structural rules seen above can explain why
from external contact equally in all directions.) nitrogen (N) nuclei commonly form three covalent bonds,
In the case of carbon, all four of the outer orbitals are as in ammonia (NH 3 ) (Figure A. 8). The N nucleus has four
bonding orbitals. This seen most simply with CH 4 or
is ,
electrons to itself: two in the inner nonbonding orbital and
methane, which is shown in Figure A. 6 with the structural two in an outer orbital that are not shared with other
formula and as it would appear in a ball-and-stick model. nuclei (this is also a nonbonding orbital) (Figure A.9).
The C nucleus with its inner electrons forms the center of Also, the N nucleus shares six electrons in the three

the molecule (Figure A.7). Each of the outer, oblong bonding orbitals. Counting these half-time electrons, there
orbitals has an H nucleus embedded in one end, with the are a total of seven negative charges near the N nucleus
C nucleus at the other end. Therefore C forms four at any moment, matching the seven protons of the
covalent bonds. Every H nucleus shares two electrons. nucleus. The N nucleus forms only three covalent bonds
These shared electrons might be considered as half-time rather than four because the extra proton in the N nucleus
electrons from the point of view of the hydrogen nucleus. is two electrons in one of the outer
sufficient to hold
Together, they add up to placing one full unit of negative orbitals without sharing with another atom. As in methane,
charge near the H nucleus, to balance the proton's unit of there are 10 protons and 10 electrons in the ammonia
positive charge. The C nucleus has the two electrons of molecule. Notice that the usual repulsion between the
the inner orbital to itself, and also shares four pairs of half- orbitals occurs, so that the three bonds between N and H
time electrons in the bonding orbitals. Therefore six form a pyramidal arrangement.
electrons are likely to be near the C nucleus at any Similar rules apply to the behavior of oxygen (O), as
moment, balancing the six positive charges of the nucleus. by water (H 2 0) (Figure A. 10). Oxygen forms
illustrated
Overall, the molecule is neutral; it has 10 electrons and 10 only two bonds rather than four because the O nucleus
protons. has enough protons to hold pairs of electrons in two of
In summary, methane illustrates two important rules of the outer orbitals without having to share with other nuclei
molecular structure: (a) each nucleus tends to control (Figure A.1 1). The eight protons of the O nucleus are

2 electrons in non-bonding orbital

each orbital

v>
H (1 +) nucleus

v^
Figure A.5 Figure A.7

Basic Chemistry

A3
Figure A.
u^
Figure A. 13

two non-bonding
orbitals (each
has 2 electrons) Since an orbital can contain only two electrons, the
double bond consists of two orbitals (Figure A. 13). One of

v the orbitals has three parts: an oval region between the

two C nuclei plus a small lobe on either end. The other
orbital consists of two sausage-shaped regions. The two
electrons in this orbital spend part of their time in one
Figure A.
sausage-shaped region; part of the time in the other.
Remember that the diagram of an orbital represents only
the places where the electrons spend most of their time;
they can pass between two lobes but they spend little time
in the intervening space.

C yJ
When two
other bonds
nuclei are joined by a double bond,
lie in a single plane. This is

arrangement that is maintained because the orbitals in the

a rigid
all their

double bond resist being twisted. By contrast, a single

Figure A. 10
bond forms an axis about which the joined parts of a
molecule may freely rotate. Figure A. 14 illustrates this
arrangement with the substance ethane, C 2 H 6 Rotation .

does not change the angle between two neighboring

3 non-bonding orbitals bonds, but rotation lends a good deal of variability to the
(two electrons each)
shape of a molecule if there are several nuclei joined into
chains by single bonds.

How Molecules Behave:

Polarity, Hydrogen Bonds,
Figure A. 11
and Solubility
balanced by six full-time electrons in the three nonbonding
orbitals, plus four half-time electrons that are shared in the Molecules may attract or repel one another because of
bonds between O and H. Again, there are 10 protons and their electrical charges. This is important in establishing
10 electrons in the molecule. how molecules will behave as a group in the body mass.

A less common but still important orbital arrangement is Even neutral molecules may have local regions of
seen in molecules like ethylene, C2 H 4 (Figure A. 12). Each positive and negative charge. This happens because
C nucleus has the usual pair of inner, nonbonding electrons that are shared between two unlike nuclei
electrons and shares two pairs of electrons in bonds with usually spend more of their time closer to one nucleus
H shared between the
nuclei. But also, four electrons are than the other. A measure called electronegativity
two C nuclei. The sharing of four electrons is known as a expresses the tendency of a nucleus to dominate in
double bond. Chemists represent a double bond by two means a stronger
sharing electrons: a higher value
lines between the nuclei. Double bonds such as C=N, tendency hoard the electrons. Electronegativities
to for

C=C, and C=0

are common in the molecules of life. some common elements are:

rT ^H

Figure A. 12

appendix A I Basic Chemistry

A4
 The H of one water molecule is attracted to the O of a
Element Electronegativity
neighboring water molecule, because of their opposite
charges (Figure A. 15). This attraction represents a weak
3.5
bond called a hydrogen bond. It is only about 1 /16 as
N 3.0
strong as the covalent bond between the H and O within a
C 2.5
water molecule. A hydrogen bond is indicated in diagrams
S 2.5
by a dotted line.
H 2.1
Each two outer nonbonding orbitals in a water
of the
P 2.1
molecule can become involved in a hydrogen bond, and
each H of the molecule can form another hydrogen bond.
Because of these differences in electronegativity, a Therefore each water molecule can, and does, form as
molecule that contains a mixture of nuclei tends to be many as four hydrogen bonds with neighboring water
negative near the N and O nuclei, and positive near the C, molecules or other polar molecules that may be present.
H, P, and S nuclei. A molecule that has such internal The force exerted by these bonds allows water molecules
charge separations is said to be polar. There is a degree to move about only by breaking and re-forming hydrogen
of polarity in every bond between two unlike nuclei, but bonds one at a time.
between C and H the polarity is insignificant whereas in Other molecules besides water can form hydrogen
bonds such as O-H, C-O, C-N, and N-H the polarity is bonds. This happens when a hydrogen nucleus that is
high enough to affect the behavior of the molecule toward sharing electrons with or N comes close to another O
other molecules. Water is a strongly polar molecule. The or N, with the three nuclei approximately in a straight line.
local charge concentrations are less than the value of a Since most biological molecules contain O-H or N-H, they
whole electron, and are shown in diagrams by the can make hydrogen bonds with one another or with water.
symbols S(-) and 8( + ): A mass of pure substance consists of many molecules
of the same kind packed regularly (a crystal) or irregularly
8(-)
(an amorphous solid or liquid). On contact with water, the
O
molecules may mass to become surrounded
leave such a
/ \
H H by water molecules. The mass is said to dissolve; the
8( + ) 8( + )
molecules that become embedded in the water are called
solute molecules; the water is said to act as a solvent;
and the resulting mixture of molecules is a solution. The
number of solute molecules in a given volume of solvent is

the solute concentration. Concentrations are usually

expressed in terms of molarity. For example, a solution
*.
\j may be 0.01 molar, abbreviated as 0.01 M. A M solution
o contains, by definition, 6.023
(one mole) one liter of
in solution.
x 10 23 molecules
A 0.01
1

M solution
of solute

contains 1 /1 00 as much solute as a 1 M solution, and so

h
Q on.
For most solutes, there
solute molecules that a given solvent
is a limit to the concentration of
will accept. Excess
solute molecules spontaneously aggregate to form masses
such as crystals. A solution with the limiting solute
Figure A.14 The substance ethane, C H as it would appear in concentration is said to be saturated. The solubility of a
2 6 ,

a ball-and-stick model. One of the H atoms has been shaded to

show how rotation around the axis of the central (C - C) bond substance is the concentration at the point of saturation.
can affect the internal geometry of the molecule. Solubility is high when the solute molecules are attracted

\/
\U-
•

Yv / S^

H
/ -H
|
^ ^J

Figure A. 15 Several water molecules joined by hydrogen bonds.

(Dotted lines represent hydrogen bonds.) On the left, structural
formulas; on the right, the arrangement as it would look using
ball-and-stick models.

Basic Chemistry

A5
to solvent molecules more strongly than they are attracted What governs the occurrence of chemical reactions 9 Most
to one another and more strongly than solvent molecules biological molecules are stable and resist change. This is
attract one another because the nuclei and electrons fall into an arrangement
A general rule governing solubility is that water will that is the best compromise between the forces of
accept polar but not nonpolar molecules. Polarity is a attraction and repulsion. Any small change from this

factorbecause the water molecules form a network, joined condition will create an imbalance of forces that tends to
by hydrogen bonds between one another. A solute can restore the initial condition. As illustrated in Figure A. 17,

move into this network if it can form hydrogen bonds with there may be several different stable arrangements that a
water; that is, if it is polar. If not, the network of water given collection of nuclei and electrons may fall into, given
molecules resists being parted to make room for the solute the chance. Some of these arrangements are more stable
molecules. Ions often are particularly soluble: solids than others, butall have a degree of stability for the

composed of positive and negative such as NaCI,

ions, reasons given above. To go from one arrangement to
often readily break up in contact with water because each another, the molecule would have to pass through an
ion acquires a blanket of oriented water molecules. intermediate arrangement that can be reached only by
Masses of nonpolar molecules dissolve freely in solvents working against electrical forces. The intermediate
such as oil or gasoline, which are themselves made of condition is a highly unstable arrangement that will

nonpolar molecules. There are attractions between the spontaneously give way to any other arrangement where
polar solute molecules in a mass that prevent them from the forces are again balanced. A molecule that is in the
leaving the mass
be surrounded by nonpolar solvent
to process of changing is called an activated intermediate.
molecules. These facts may be summarized by the dictum,
"like dissolves like." But molecules differ widely in their
degree of polarity and correspondingly they differ widely in O
the degree of solubility in various solvents. II II II

C-H C-H C-O-H

I
I
I

How Molecules Behave: c=o C-O-H C-H

I
II

Chemical Reactions H-C-H C C

I
/ \ / \
H H H H H
A chemical reaction is an event in which new kinds of
molecules are built by rearranging the parts of old Figure A. 17 Three possible structures that could occur with the
molecules Chemists symbolize a reaction by means of an formula C H 4 2 .

arrow, with the initial molecules (reactants) at the tail and

the final molecules (products) at the head. A collision between two molecules may set the stage for
Some common types of reactions are illustrated in an exchange of parts, as illustrated in Figure A.18. But the
Figure A. 16. The simplest are rearrangements or exchange first requires the two molecules to form an
isomerizations, which a single molecule undergoes an
in unstable activated complex.
internal reorganization. There are also addition reactions,
Activated complexes form only when outside forces
where two molecules join to form a larger molecule. The push the molecules out of their stable arrangements. The
reverse is a decomposition reaction. The most common required force is most commonly provided by the force of
kind of reaction is the transfer, in which two molecules collision between two molecules. Momentum carries the
meet and exchange or transfer parts to form two new
molecules.

H o- -H
\ /
II
O C
I isomenzation
H-i IX C
II

/ \
H H

H H H
\ / I

c H H-C-H
II + I |
addition
c H H-C-H
/ \ I

H H H

o o
II II

R-C-O-H + R'-O — H R-C-O-R' + H2 transfer

Figure A. 16 Examples showing three general types of reactions.

appendix A |
Basic Chemistry

A6
 O H+ O O
II
H+
II
OH
II II

0— P-0 H-0-P-O-H
I I

H— Q-P-O-P-0- H-0- P-0 P-O-H +

x
o o ,0
I

o o +H 0.
/ \ \ / .-•
\ \
H H H H H H H H H

activated complex

Figure A. 18 The reaction between pyrophosphoric acid and

water, illustrating an activated complex. The bonds shown with
dotted lines in the activated complex are those that will be
affected by the reaction. As written, the reaction is a hydrolysis,
one of several kinds of transfer reactions.

molecules together until their shapes become distorted, The difference in energy between the activated
just as a basketball is flattened on one side at the moment intermediate and the separate molecules is termed the
of impact with the floor. Sufficient distortion will bring the activation energy. This is the amount of energy that must
molecules into the activated state enabling a chemical be supplied in the collision if there is to be a reaction
reaction to follow. A lesser degree of distortion allows the event. The higher the hill, the less likely is the collision to
colliding molecules to rebound without reacting. produce a reaction.
Chemists routinely use energy concepts to discuss Whether the collision causes a reaction will depend on
chemical reactions. A typical graph of the energy relations the nature of the molecules, their orientation when they
in a reaction shown in Figure A. 19. The molecules
is hit, and how hard they hit. Most collisions at the
contain a certain amount of stored or potential energy in temperatures of life fail to bring about chemical reactions
their stable condition. The activated complex contains because of defects in one of these factors.
more potential energy. This reflects the fact that force In principle, all chemical reactions are reversible. Using
must be applied (work must be done) to bring the stable Figure A.19 as an example, if A and B can collide to form

molecule into the activated condition. Moving molecules an activated complex, so can C and D collide and form the
have kinetic energy, or energy of motion, in addition to same kind of activated complex. The latter can break apart
their stored potential energy. A collision between two to form either A and B, or C and D. The back reaction
molecules converts this kinetic energy into potential would look the same as if a film of the forward reaction
energy: the colliding molecules are momentarily at rest in were being run backward. As shown in Figure A. 18,
a distorted state. When the molecules rebound from the reversibility is shown in reaction diagrams by putting two
collision, some of the potential energy is converted back opposed arrows at each step.
into kinetic energy. In this process the total amount of Which of the two opposed reactions will run the faster?
energy remains constant. Energy cannot be created or The rate of a reaction depends on the frequency of
destroyed by the reactions in life, though the form of the collisions, hence on the concentrations of the reacting
energy may be changed. molecules. If we refer again to Figure A.19, the reaction
The products of a reaction may contain, between them, from left to right depletes the supply of A and B so that
either more or less potential energy than the original the reaction rate declines with time. The products C and D
reactants; the difference is made up by a compensating accumulate and one another increasingly
collide with
change in the amount of kinetic energy carried by the often, raising the rate of theback reaction. Thus the
molecules. In Figure A.19 the products have less potential overall rate, the difference between forward and back
energy and therefore more kinetic energy than the reactions, depends on the prevailing balance of
reactants. The kinetic energy of the molecules is known as concentrations. Equilibrium occurs when the
heat. Thus heat is produced as the reaction in Figure A.19 concentrations of all the molecules are just sufficient for
progresses. the forward and back reactions to run at the same rate.

At equilibrium the overall rate is zero; the opposed

reactions cancel.
activated complex
What determines the balance of concentrations at

equilibrium? most reactions (Figure A.19) there is a

In

difference in potential energy between the reactant and

Ea = activation energy product molecules. This gives the back reaction a different
activation energy requirement than the forward reaction: in
the reaction of Figure A.6, the back reaction has a higher
energy hill to climb. A higher activation energy
requirement means that fewer collisions will have enough
energy to form the activated complex. Therefore more
collisions must occur per unit time if the back reaction rate
Figure A.19 Potential energy changes that occur during a is to equal the forward reaction rate. The rates will only be
reaction event in which molecules A and B collide, form an
activated complex, and react to form C and D. Graph shows the equal when there are more of the low-energy molecules
potential energy of (A + B), the activated complex, and (C + D). than the high-energy molecules. In Figure A.19, C and D

Basic Chemistry

A7
willbe more abundant than A and B at equilibrium. HA + H 9 H 3O
f
+ A
These ideas suggest that reactions should have the
Acids vary widely The very strong acid HCI
in strength.
overall eftect of replacing molecules that are rich in +
(hydrochloric) completely breaks up into CI" and H 3
potential energy with molecules that are lower in potential
when contacts water. A weak acid remains mostly un-
it
energy. Nevertheless, living bodies do form many
ionized, as HA. Water and acetic acid (which causes the
molecules that are higher in potential energy than the raw
sourness in vinegar) are both weak acids.
materials from which they are made. This is possible
A base is a molecule that reduces the concentration of
because the energy relations discussed above refer only
hydronium in OH" is a base. There
water. For example,
to the sum of potential energy, taking both product
are several ways
which a substance can decrease the
in
molecules together. As long as this sum is greater for A
hydronium concentration. Thus B~ can be an ion that
and B than for C and D, the latter will tend to be the more
reacts as:
abundant at equilbrium. C is very low in potential
If

energy, D may be richer in potential energy than A or B. B + H 3 0+ BH + H P

In biological systems, reactions rarely come to
The OH~ ion acts in this way. As another example, BOH
equilibrium because various materials are continually
may be a compound that ionizes in water to release OH".
being withdrawn or supplied by other reactions. Also, back
The OH" then reacts as shown above.
reactions are often insignificant because their activation
Instead of talking about hydronium, chemists often
energy requirements may be too high to meet in collisions
abbreviate by limiting their attention to the H+ ion itself,
at body temperature.
ignoring the part played by H 2 in carrying the proton (as

H 3 + ). From this viewpoint, pH refers to the H +

+
Special Reactions: concentration. But free H never occurs in the solution.

Acids and Bases

Functional Groups
Acidity results from a special type of transfer reaction,
involving water and a hydrogen nucleus (proton). When The molecules of life are often large and complex. But
two water molecules are joined by a hydrogen bond, . they are easier to understand when we use the concept of
occasionally the hydrogen nucleus jumps from one functional groups. A functional group is a part of the
molecule to the other: molecule that is especially likely to participate in chemical
reactions.
q
The functional groups carboxyl
H ||

/ ( -C-O-H
H-0 H-0 + + H
H-0 H O
O
hydronium hydroxyl
II

and phosphoryl O-P—O-H

I

+
The products are H 3 , known as hydronium, and OH", O-H
called the hydroxyl ion. A meeting between hydronium
and a hydroxyl ion usually results in a return to the two
+
water molecules: the ionization of water strongly favors the act as acids. These can donate H to water:

two un-ionized molecules.

The concentration of hydronium ions determines the II II

acidity of the solution. More hydronium means a more

R-C-O-H + H,0 R-C-0 + H 3 0"

acidic solution. Chemists express the hydronium

concentration by a measure known as the pH. The scale
runs from to 14. A change of 1 unit on the pH scale is a O
ten-fold change in hydronium concentration. The lower the II II

R-0 P— O-H + 2H,0 * R-O-P- -0~ + 2H 3 0"

pH, the higher is the hydronium concentration. Thus a
I I

solution with pH 2 has 10 times as much hydronium as a O-H 0"

solution with pH 3. Pure water has a pH of 7. Most living
The carbon of carboxyl and the phosphorus of
material have a pH value near 7. Stomach juice in human
phosphoryl tend to have a partial positive charge; the
beings is very acidic, with a pH around 1 Even at this
attached oxygens have partially drawn away the shared
.

strongly acidic pH, though, there are many more neutral

electrons, leaving the C and P nuclei relatively exposed to
water molecules than hydronium ions in the solution: pH 1
attack by other molecules. Therefore these nuclei tend to
represents about 1 hydronium for every 180 neutral water
chemical reactions. By contrast, C
be involved often in
molecules.
nuclei that are only bonded to other C or H nuclei are less
An acid is a compound that raises the hydronium subject to chemical attack.
concentration inwater. Any compound that can donate a
proton to a water molecule is an acid. If we use the H
symbol HA for a generalized acid, the ionization is: The amino group ( — acts as a base. The
\

appendix A |
Basic Chemistry

A8
nonbonding orbital of the N atom can accept a proton Since O nuclei have a high affinity for electrons and will
from water or hydronium: hoard the electrons that they share, the change in
bonding that occurs in this reaction causes the uppermost
H H C to lose a fraction of a negative charge (it is oxidized)
/ + while the lower C makes a matching gain in negative
R-N + H3 + - R-N-H + H2
\ charge (it is reduced). The molecule as a whole is neither
I

H oxidized nor reduced since it has not lost or gained

electrons.
As the example illustrates, the electron-attracting
last

capacity of O
causes groups to be oxidized when O
becomes attached to them, whereas a replacement of O
by H is a reduction because H tends to donate electrons.

Hydroxyl ( — O— H) and carbonyl 1^ — ° ) are also

Oxygen nuclei rarely are poorer in electrons than when
twoof them are competing for electrons in an
functional groups. They do not form ions appreciably 2

unless they are joined together as carboxyl, but the

molecule. With its high electronegativity, O will readily give

oxygen atoms in these groups pull electrons from

up its position in 2 to form bonds with C and H, which do
not compete as well for electrons. The possibilities for
adjoining C nuclei, creating polarity and exposing the C
nuclei to attack. A common reaction is that of a carboxyl
drawing electrons from have been maximallyC and H
with a hydroxyl group, to give a product called an ester
exploited by O in and H 2 0. Therefore
the molecules C0 2
these are very stable molecules. In them, C and H are
plus a water molecule (See Figure A. 16.) Such reactions,
highly oxidized and O is highly reduced. Compounds such
which give off a water molecule, are called
as sugars,
condensations. The opposite, cleavage of a molecule by
adding water, is a hydrolysis reaction. An example of a
hydrolysis is shown in Figure A.18. Both condensations H H H OH H
and hydrolysis reactions are transfer reactions. I I I I I

H-C-C-C-C-C-C=0
I I I I I I

Oxidation and Reduction OH OH OH H OH H

An oxidation-reduction is a reaction in which electrons which have C— C and C — H C and H bonds, contain
are transferred from one group or molecule to another. and therefore are more
nuclei that are richer in electrons
The entity that loses the electrons is said to be oxidized; reduced than the C and H in C0 2 and H 2 0. A compound
the one that gains the electrons is reduced. Often such a such as sugar is highly reduced. The more bonds C and H
reaction also involves a transfer of atomic nuclei. have with one another rather than with O, the more
A simple example is the ionization of chlorophyll, an reduced the compound is. Reduced compounds
important event in photosynthesis. After absorbing a unit spontaneously react with 2 (with a suitable catalyst) to
molecule passes an
of light energy, the chlorophyll give products which the C and H are more fully
in

electron to a compound that acts as an electron acceptor: combined with O and therefore are more highly oxidized.
This occurs because it transfers electrons from C and H to
chlorophyll + acceptor O. The reverse, the removal of O, is unfavorable unless
+
-* chlorophyll + acceptor" energy is supplied by the environment. Therefore in the
presence of 2 reduced compounds are rich in energy as
,

Here chlorophyll is oxidized and the acceptor is reduced. compared to oxidized products. The stored energy is often
Somewhat more complex is the formation of the called redox energy and may be defined as energy
compound NADPH from NADP + another event in ,
associated with an unfavorable distribution of electrons
photosynthesis: between the nuclei. This redox energy is released when a

+ +
reaction allows the electrons to assume a more favorable
2Z~ + NADP + H,0 2Z + NADPH + H 2 arrangement.
In this reaction, which probably proceeds by several steps,
NADP + receives two electrons, with Z as the electron
-
donor. NADP + also gains a proton. The donor Z is
+
oxidized; NADP becomes reduced.
Oxidation and reduction can also occur during an
internal reorganization of a molecule. For example:

OH O
II

C-H
I

H-C-H
I
I

c=o H-C-OH
I
I

R R

Basic Chemistry

A9
 B APPENDIX
genetics supplement

In many flowering plants, segments of a stem can be goal of genetics has been to deduce the laws that govern
rooted to give a set of plants with identical heredity, a inheritance in matings. This is useful in plant breeding,
clone. Clones occur widely in nature because many plants where one must plan breeding programs to give
can reproduce vegetatively (e.g., by runners or rhizomes). predictable improvements in the plants; it is also useful in

But plants of almost all species also reproduce sexually, exploring the fundamental mechanisms of heredity.
by means of matings. In such a mating (Figure B.1), A good breeding program requires plants whose
meiosis in one diploid parent produces haploid egg cells, matings can be controlled. The possibility of accidental
while meiosis in the other parent produces haploid nuclei pollination by wind or insects, or must
of self-fertilization,
that are incorporated into pollen grains. The egg cells and be considered. In self-fertilization, pollen and eggs come
the generative cells of pollen are gametes. Egg and pollen from the same parent.
nuclei meet and fuse in the act of fertilization, to yield a Success in a breeding program also depends on the
diploid zygote.* The zygote is the first cell of the offspring choice of traits to be studied. Traits are observable
or progeny plant. It contains all the hereditary information characteristics such as color of petals and shape of
that will guide the young plant's development. Half of that leaves. Some traits are easier to study than others. For
information came from the pollen, and half from the egg. example, the weight of seeds produced by a plant is partly
determined by heredity, but also it is strongly influenced
by the environment. Even in a constant environment,
diploid diploid
though, seed weights show many
intergrades between the
(2n) (2n) PARENTS
adult adult heaviest and the lightest.Seed weight, oil or starch
content, growth rates, and so on are quantitative traits.
Such traits commonly show continuous variation, in
which the differences between plants are distributed
haploid widely and evenly about a mean (Fig. B.1 3). Traits with
haploid
(In)
(In)
GAMETES continuous variation are hard to study because the
pollen
egg cell
nucleus
progeny of matings are hard to classify and patterns of
heredity are complex.
By contrast, some traits show discontinuous variation.
fertilization
Thus a population of garden pea plants might contain
diploid
some individuals that form red flowers and some with
(2n) purple flowers, but no intergrades. Traits that show
zygote
discontinuous variation are easy to study because the
progeny can be assigned to definite classes and the
mitotic various types occur in simple ratios. Historically, studies of
divisions; PROGENY
development
discontinuous traits gave us the key to understanding
heredity. Today, it is still easiest to learn genetics by
starting with discontinuous traits.

Traits That Show

Figure B.1
Discontinuous Variation
With a clone, one knows exactly what to expect from
one generation to another; each generation will be
The first systematic breeding program was done in the
preceding one, except for variations that
identical to the
mid-nineteenth century by Gregor Mendel. He chose the
the environment causes. But matings make predictions
garden pea, partly because many varieties of peas were
more difficult. The progeny plant can be expected to differ
available and partly because their matings could be
from both of its parents in many ways, unless the two
controlled. The stigmas mature before the anthers and
parents carried identical hereditary information. A major
before the flowers open. At this stage the flowers can be
* The details of meiosis are presented on pried open, the immature anthers can be removed, and
pp. 38-43, while
gamete formation and fertilization are described on pp. 130-140. pollen from another plant can be applied to the stigma.

B1
Fertilization occurs in the closed flower, with the stigma
shielded from stray pollen.
Each variety of peas that Mendel studied had its own P, cross:
P:
9 Round X
a Wrinkled

set of unique hereditary features. In a field sown with

seeds of one variety, all the progeny for generation after I
F II Rou nd
generation would have the features typical of that variety. ,

The
traits
plants
under study;
in such a field are said to "breed true" for the
collectively, the plants form a true-
F
, 9 Round X
a Round

breeding strain.
F cross

75% Round
The Monohybrid Cross 25% Wrinkled

Figure B.3
The problem in genetics is to explore what will happen
when plants of two true-breeding strains are mated. This is

easiest to approach if the plants differ in only one trait.

Matings of this kind are called monohybrid crosses. shows that the F, plants did contain the information for

(Actually, the parental plants may differ in other

wrinkled seeds, but the information was suppressed.
characteristics as well, but so long as attention
To explain the patterns of inheritance that he saw,
is confined
Mendel proposed the scheme shown in Fig. B.4. He
one trait, such as flower
to just color, the mating is a
monohybrid cross.) suggested that the hereditary information for seed shape

Figure B.2 shows the kind of results that Mendel

is carried by particles. There were two versions of the

obtained monohybrid crosses with peas. The original

in
particle. One kind (call it R) carries information making
for
round seeds. The other (r) carries information for making
parents are called the P Generation. One parental plant in
Fig. B.2 came from a strain that always produces round
wrinkled seeds. Mendel proposed that each gamete
seeds; the other from a strain that always forms wrinkled
carries one of these particles, so that the zygote contains

seeds. The progeny are the First Filial or F Generation.

two of them. To account for the suppression of the
1

In this cross, Mendel found that all of the F, plants formed

"wrinkled" he proposed that the factor R is
trait,

round seeds. This seems surprising at first, because dominant over r when the two are both present in the
having received hereditary information for making both same organism. That is, the presence of R prevents r from
round and wrinkled seeds, one might expect some of the affecting the shape of seeds.

progeny to form wrinkled seeds. Either the hereditary

information for wrinkled seeds has been suppressed, or it

never entered the zygote during fertilization. P: Round Wrinkled

RR rr

P generation meiosis I meiosis I

true-breeding true-breeding
Round-seeded Wrinkled-seeded gametes
plants plants

( pollen ) egg cells

fertilization

development
Figure B.4

"

F generation

Round-seeded plants Today, Mendel's particles are called genes. They are
known be made of the substance DNA. DNA can be
to
Figure B.2 modified many ways, though new modifications
in

(mutations) in any one gene occur only rarely (see

An additional cross (Fig. B.3) proves that the Chapter 2 and pp. 215-217). The different versions of the
information for making wrinkled seeds was indeed present gene proposed by Mendel are slightly different segments
in the F, plants. were mated with one
Here the F, plants ofDNA, carrying information for the same trait. Different
another. In such a cross, the F, plants are said to be self- same gene are now called alleles.
versions of the
crossed or selfed, and their progeny are the F 2 Because there are many ways to mutate a gene, many
Generation. Although all the F, plants formed round different alleles are possible. But a diploid cell carries only

seeds, some of the F 2 plants made wrinkled seeds. This two copies of each gene, so that only two alleles for that

Traits That Show Discontinuous Variation

B2
gene can be present. (There are exceptions to this rule, partly by the laws of chance, which govern the random
but they can be ignored in elementary studies.) combinations of the various gamete types.
In modern language, we would rephrase Mendel's As anyone knows who has played games with dice or
proposal by saying that allele R (round seeds) is dominant cards, the laws of chance leave room for variation. With
over allele r (wrinkled seeds); allele r is said to be many tosses of a coin, "heads" will come up very nearly
recessive to R. This pair ot alleles is said to show 50% of the time. But with just a few tosses the percentage
classical dominance. We
have not explained how R may be quite far from the ideal 50% figure. Likewise in
suppresses r; we have merely developed a picture that counting the progeny of a mating, we do not expect the
heips to predict the progeny in matings. Classical numbers to be exactly as predicted; deviation is likely to
dominance has been tound between many pairs ot alleles, be greater, the smaller the number of progeny. Geneticists
affecting many traits in many plant and animal species. have adopted statistical methods that help to estimate how
The idea of dominance between alleles explains why far the real results of a cross are likely to deviate from
some of the F 2 plants produced wrinkled seeds (Fig. B.3). ideal predictions.
The analysis is shown in Fig. B.5. Each nucleus in the A note on terminology: the combination of alleles
parental plants contains the allele combination Rr. Meiosis present in a plant is known as the genotype, while the
separates the alleles, so that half of the gametes contain observable trait itself — seed shape
in this case is the —
allele R and half contain allele r. Gametes with these phenotype. True-breeding plants have two copies of the
alleles meet at random in matings, so that zygotes are same allele, RR or rr. They are said to be homozygous
formed with all the possible combinations. These are for this gene. Plants that carry two different alleles, such
shown in the four boxes in Fig. B.5, along with the seed as Rr, are heterozygous.
shapes that the progeny will form. Three of the four
combinations result in round-seeded plants; the fourth The Testcross
combination, rr, yields plants with wrinkled seeds.
The dominance mechanism makes its easy to recognize
plants that are homozygous for the recessive allele,
because they are the only plants that show the recessive
X
Rr Round Rr Round phenotype. But plants with genotypes RR and Rr are
identical in appearance; both produce round seeds. How
can we deduce the true genotype of a round-seeded plant,
so as to make accurate predictions in matings that use the
plant?
The easiest approach is to use a testcross. In a
R testcross, one parent has only recessive alleles for the
trait being studied. The other parent has the dominant

phenotype, but its genotype is unknown. The testcross for

RR Rr
seed shape in peas is diagrammed in Fig. B.6. There it

Round Round has been assumed that the unknown parent has the
egg cells
genotype Rr. The recessive parent forms only gametes
with the allele r. The other parent produces gametes with
Rr rr
allele R and an equal number with allele r. Only two
Round Wrinkled
progeny genotypes are possible; they are shown in the
boxes. the unknown parent is heterozygous, half the
If
Figure B.5
progeny form round seeds and half form wrinkled seeds. If
the unknown parent were homozygous RR, all the
To be useful, genetic analyses should predict not only progeny in the testcross would have formed round seeds.
the kinds of progeny that will occur, but also their relative The testcross is useful because the alleles contributed by
abundance or frequency. Looking back to Fig. B.3, one the recessive parent do not interfere with the expression
can see that 25% of the progeny formed wrinkled seeds. of the other parent's alleles. The progeny phenotypes
Why 25%? To understand this, one must be aware that directly reveal the unknown parent's gamete types. From
chance plays a major part in bringing the gametes this information we can easily deduce the unknown
together. The contained alleles do not affect a gamete's parent's genotype.
chances of meeting and fusing with another gamete. Since
meiosis forms equal numbers of R eggs and r eggs, the Incomplete Dominance
laws of chance assure that the two types of eggs will be
involved in matings equally often. Thus half the progeny Though many traits show classical dominance when pairs

should contain an r egg nucleus. By chance, half of these of alleles are brought together in matings, many others do
will combine with pollen that carry allele r, since meiosis not. One exception is shown in Fig. B.7. When plants from

also assures that r pollen are just as abundant as R a red-flowered strain of snapdragon (Antirrhinum) are
is 25%, hence 25% of the F
pollen. Half of half 2 plants
are crossed with plants from a white-flowered strain, the F,
the rr which form wrinkled seeds. The frequency of
type, progeny all have pink flowers. Here Mendel's idea of
a given progeny type is set partly by the mechanics of dominance does not apply. But the F plants still do 1

meiosis, which fixes the frequency of gamete types; and contain the information for making red and white flowers.

Traits That Show Discontinuous Variation

B3
 Rr Round rr Wrinkled

gametes gametes

/ \

R r

RR Rr
gametes <

Red Pink

o
gametes

Rr

Figure B 6 Pink White

Figure B.8
When the F, snapdragons are selfed (Fig. B.7, bottom),
half the progeny torm pink tlowers, but the rest produce
red or white flowers.
a strain of corn with high resistance to fungus attack but
with poor grain yield may be crossed with another strain
White
that has low fungus resistance but good yield. Such a
Red
cross, involving two traits, is a dihybrid cross. The goal
P, cross
would be produce plants that yield well and are
to
disease-resistant. But the progeny might also include
All pink
plants that combine poor yield with poor resistance. How
many of the progeny can be expected to have the
desirable combination of features? The answer will
Pink Pink
determine the effectiveness of the breeding program.
In a monohybrid cross, we were dealing with only one
F, cross gene had two alleles. In dihybrid crosses, two genes
that

25% Red are involved, each with two alleles. It is important to

50% Pink
distinguish clearly between "genes" and "alleles"
25% White
because confusion between these terms will make it
Figure B.7 impossible to understand inheritance in dihybrid crosses.
The genes are carried on chromosomes, with many
Although dominance does not apply, these results can genes per chromosome, governing many traits. Each kind
be explained easily with the allele theory of inheritance. In of chromosome in the haploid set carries a particular

Fig. B.8, the red-flowered plants are assigned the group of genes, which are arranged in a definite sequence
genotype RR; white-flowered plants have the genotype rr. along the chromosome. Thus two genes on the same
The Ft progeny all have the genotype Rr, which is found chromosome are physically coupled together. They are
by observation to give a pink-flowered phenotype. In the said to be linked. Two genes that are carried on different
next cross, the pink-flowered parents (with genotype Rr) chromosomes are unlinked. Linkage affects the formation
form equal numbers of R and r gametes. These unite at of gametes, which in turn limits the occurrence of certain
random so that four combinations result. One combination progeny types. Therefore one must know whether two
(RR) yields red-flowered plants; two combinations (Rr) genes are linked or not, good predictions are to be made
if

give pink, and the fourth combination (rr) produces white- about their behavior in matings.
floweVed plants. One would expect 25% red, 50% pink, In garden peas, the shape and color of the seeds are

and 25% white. This matches the observed progeny in the governed by a pair of unlinked genes. As to shape, round
F 2 generation (Fig. B.7). (R) is dominant over wrinkled (r). In seed color, yellow (Y)

Pairs of alleles such as this one, where the is dominant over green (y). Figure B.9 shows how a plant

heterozygous condition gives a different phenotype than that is heterozygous for both genes produces gametes.
two homozygous phenotypes, are said
either of the to Remember is preceded by a duplication of the
that meiosis
show codominance or incomplete dominance. chromosomes, so each chromosome entering meiosis
that
has two identical chromatids. * Then in prophase of the
first meiotic division, homologous chromosomes join
Dihybrid Crosses and Linkage together. The resulting tetrad is a group of four

Many breeding programs are concerned with bringing two * It may be worthwhile to review Fig. 3.32 and the associated text
ormore qualities together in the same plant. For example, on pp 42-43, where meiosis is shown in more detail).

Traits That Show Continuous Variation

B4
 parental parent
r sporocyte sporocyte
|2n) (2n)

gametes gametes gametes gametes

(
R
> c
y
)
WU R
)
y P L

Y W r
(
P
I
1
)

Figure B.9 Figure B.10

chromatids. In Fig. B.9 the long chromosomes form one coupling between p and I. When meiosis is complete, half
tetrad; the short chromosomes form another. The tetrads the gametes have the genotype PL and half have the
then line up at the equator of the spindle. But tetrads genotype pi.

move independently of one another. Therefore the two Some cells take the right branch of Fig. B.10. Here, an
can assume two alternative positions relative to
tetrads event called a crossover occurs in the tetrad: two of the
one another. one arrangement (the left branch in Fig.
In homologous chromatids break at the same point. Then the
B.9), meiosis gives two gametes with genotype RY and broken ends rejoin in such a way that the chromatids
two with genotype ry. The other arrangement (right have traded parts. Tracing through the rest of meiosis,
branch, Fig. B.9) results in two gametes with genotype Ry one can see that the crossover has introduced two new
and two with genotype rY. Only one of the possible classes of gametes. The chromatids that took part in the
arrangements can occur in any one cell that undergoes crossover will result in gametes with the genotypes PI and
meiosis. But meiosis occurs in many cells. Chance pL. The chromatids that were not involved in the
governs the positions that the tetrads take, so that the two crossover will form gametes with genotypes PL and pi.
arrangements will be equally common. Therefore, if the Meiosis with a crossover has formed four kinds of
two genes are unlinked, the four gamete types will be gametes, whereas meiosis without a crossover gave only
equally numerous. two kinds of gametes. When a large number of cells
Next, let us consider the formation of gametes when the undergo meiosis, only a minority will have crossovers
two genes are linked. This can be illustrated with another between the two genes. Therefore, the effect of linkage is
pair of genes in the garden pea: flower color, with purple to reduce the frequency of two classes of gametes out of
(P) dominant over red (p); and pollen shape, with long (L) the four.
dominant over round (I). Figure B.10 shows the formation As Figs. B.9 and B.10 show, linkage affects the
of gametes in a plant that is heterozygous for both genes. frequency of gamete types. However, we cannot directly
Here we assume that the parent has allele P on the same see the genotypes of gametes. How, then, can we
chromosome as allele L, and allele p is on the same determine whether the genes in a pair are linked or
chromosome as allele I. unlinked? The easiest method is to subject the doubly
Most cells that undergo meiosis will follow the sequence heterozygous plant to a testcross.
of events in the left branch of Fig. B.10. Pairing brings the Figure B.1 1 shows what happens in the testcross when
homologous chromosomes together in a tetrad. The the genes are unlinked. As always in a testcross, one
coupling between P and L is maintained, as is the parent has only recessive alleles. This allows the other

Traits That Show Discontinuous Variation

B5
 parent's alleles to be fully expressed in the progeny. The
genes are unlinked, so the heterozygous parent has
produced four equally common kinds of gametes.
Therefore the progeny also include four equally common
phenotypes.
Contrast this with the behavior of linked genes in a
testcross (Fig. B.12). Again there are four classes of
gametes
progeny, but two classes are more common than the other

(=D two. The

by crossovers
One can see
rare types are
in
due
meiosis.
to gametes

that the testcross of the double

that were formed

heterozygote can be useful indetermining whether two

25%
D CD genes are linked or not. But whatever the outcome, the
testcrossshows the relative frequencies of the various
round, yellow
gamete types that a plant produces. This is useful
information for breeding programs, because the gamete
Y 25% frequencies can be used to predict progeny genotype and
DCD phenotype frequencies. The mathematical methods for
round, green making these predictions cannot be described in the
gametes <
space we have here, but they are simple and can easily
be learned from any elementary genetics text.
25% Though Fig. B.10 shows only one crossover between
DCD
two chromatids, it should be mentioned that more than
wrinkled, yellow one crossover can occur in a given tetrad during a given
meiosis. Also, even though a single crossover involves
only two chromatids, additional crossovers may involve
25%
DCD other chromatids. Chance plays a large part in setting the
wrinkled, green
locations where crossovers will occur along the
chromosome. Thus crossovers occur less often between
Figure B.11 two closely spaced genes than between two distant genes.
Two or more crossovers may occur between genes that
occur at opposite ends of the chromosome, so that in
testcrosses the genes may appear to be unlinked.
Geneticists have developed mathematical methods for
taking these factors into account in planning crosses and
predicting results. Such methods have also been used to
map the chromosomes, showing the locations where
genes occur.

gametes

Traits That Show

Continuous Variation
P L 43%

The preceding discussion has concentrated on traits that

purple, long
show discontinuous variation. But continuous variation is
the rule with many traits. For example, the seeds of corn

P 1 7% (Zea mays) vary in protein content. In a study of this trait,

( I 1
produced seeds high in protein were mated
plants that

purple, round
with plants that produce low-protein seeds. The F,
gametes <
contained progeny with a wide range of seed protein
content (Fig. B.13). This pattern suggests that protein
P L 7% content might be controlled by environmental factors
( I )

rather than heredity. Undoubtedly the environment does

red, long
play a part. But another kind of experiment shows that
heredity is involved too. A field of corn was planted and

P 1 43% seeds from the crop were used to replant the field again
the next year —but only seeds from plants that gave the

red, round
best protein yields were used. Every year, a similar
selection and replanting was done. Figure B.14 shows that
Figure B.12 the selection process gradually raised the average protein

Traits That Show Continuous Variation

B6
 discrete classes? Aside from environmental etfects, the

high-protein corn low-protein corn central explanation seems to be that several genes
contribute to the control of the trait. Two or more alleles
might be present in the population for each of these
genes. In matings, then, gametes and zygotes with many
combinations of alleles Each combination
could result.

might give a somewhat different phenotype with respect to

Percent the trait that is being studied. many genes are involved,
If

of
*Y the effect of changing any one gene's alleles might be too
progeny
small to detect easily against the background of
environmental effects. For plant improvement in such
cases, selection programs like the one in Fig. B.14 may be
25
Protein content of important, together with specific crosses between strains
seed (%of seed weight)
that have been selected for favorable features.
Figure B.13

Maternal Inheritance

Allthe patterns of heredity mentioned so far have

concerned genes that are carried in the nucleus, with a
pollen grain and an egg cell making equal contributions.
These include the great majority of cases. But anomalous
patterns of heredity sometimes occur, in which the
maternal (egg-producing) parent determines a trait in the
progeny and the pollen seems to make no contribution.
This can result from at least two causes. First, there are
cases in which fertilization is bypassed and the flower
forms seeds that start from a purely maternal cell (usually
diploid) instead of a zygote. This process, called
apomixis, occurs in the lawn dandelion. It is really a form
20 40 of asexual reproduction and does not belong in a
Number of generations discussion of matings, but we mention here because it

Figure B.14. The on seed protein content in

effect of selection such cases can greatly confuse an attempt at controlled
Zea mays. Data from Woodworth, C. M., E. R. Leng, and R. W. breeding. Second, plastids contain some genes that
Jugenheimer. 1952. Fifty generations of selection for protein and
oil in corn. Agron. J. 44:60-65. contribute to the development of the chloroplasts. These
genes are passed from generation to generation by
duplication and division of the plastids; their inheritance is
distinct from that of nuclear genes. Since egg cells
content of seeds produced in this field. In a parallel contribute plastids to the zygote while pollen cells usually
experiment, another field was replanted year after year, do not, only the maternal parent has a part to play in

using only the seeds from plants that had the lowest plastid inheritance. Effects of the chloroplast genes are
protein yield in the preceding year's crop. In this especially noted in the inheritance of variegated color
experiment, the continued selection gradually decreased patterns in the leaves of some plants (e.g., variegated
the average level of seed protein. These experiments show Mirabilis).Here the egg cell contains both normal plastids
that protein content in the seeds depends on heredity: and plastids that cannot develop into green chloroplasts
otherwise the protein content should not have been so because of defective genes. Some cells in the embryo
affected by selection of the seeds. come have only the defective plastids. These cells
to
If quantitative traits such as seed protein content are divide further to produce the distinctive white or yellow
under hereditary control, why don't the progeny fall into patches on the leaves and stem of the mature plant.

Maternal Inheritance

B7
 APPENDIX

table of metric equivalents

Exponents IV. Units of Volume

Very large or very small numbers are often written in liter (I) = 10 3 ml = 1.06 U.S. liquid qt = 0.91 U.S. dry qt
exponential form to save space. Thus, in the number = 0.88 British qt
3 3 =
10 ,
the exponent is the superscript 3. 10 is short for milliliter, cubic centimeter (ml or cm 3 ) = 0.03 fluid oz
1,000; or 1 followed by 3 zeroes. The number 10~ 2 0.06 in.
3

represents 0.01 ; the digit 1 is placed 2 places to the right

of the decimal.
V. Miscellany; Units of Pressure, Light
Intensity, Energy, Work, Force
Common Prefixes
2
9 bar = 0.99 atm of pressure = 14.54 lb/in.
nano = 10~ (one billionth)
10~ lux (L) - 0.09 ft-c of light intensity
micro = 6
(one millionth)
= -3 joule (j)
= 0.735 ft-lb of work = 9.5 x 10~ 4 British
milli 10 (one thousandth)
3 Thermal Units (BTU)
kilo = 10 (one thousand times) -7
erg = 10 joule
mega = 10 6 (one million times) 3
watt (w) = 1.3 x 10" hp
calorie, gram-calorie (cal) = amount of heat needed to
I. Units of Length raise the temperature of 1 g of water 1 °C from 14.5 to
15.5°C
kilometer (km) = 10
3
m = 0.62 mile kilocalorie, kilogram-calorie (kcal) = 1000 c = 3.97 BTU
meter (m) = 1 .09 yd = 3.28 ft
(1 BTU is the amount of heat needed to raise 1 lb of
decimeter (dm) = 10~ m = 3.90 in. 1

water 1°F)
-2
centimeter (cm) = 10 m = 0.39 in. solar constant (radiation at Earth's outer atmosphere
3
millimeter (mm) = 10~ m = 0.04 in. = 2 cal/cm 2 /min = 13,400 = 4.06 x 10 7
-6 edge) ft-c
micron, micrometer (n) = 10 m BTU/m 2 /yr
-9
millimicron, nanometer (m/x or nm) = 10 m dyne = force required to accelerate 1 g at the rate of 1

angstrom (A) = 10 10
m cm /sec /sec = g-cm/sec
2
1
5 2
newton = 10 dyne = kg-m/sec 1

II. Units of Weight

metric ton = kg
6
10 g = 1.10 short
1 tons = 0.98 VI. Units of Temperature
= 2200 lb
long ton
3 one degree (1 °C) = °K = .8°F
Celsius, Centigrade
kilogram (kg) = 10 g =2.20 lb 1 1

gram (g) = 0.035 oz absolute zero = -273°C = 0° Kelvin = -460°F

_3 boiling point of water = 100°C = 373°K = 212°F
milligram (mg) = 10 g = 0.01 5 grain
freezing point of water = 0°C = 273 °K - 32 °F
to convert from °F to °C: (°F - 32) x 0.56 = °C
III. Units of Area

are (a) = 100 m 2 = 1075.84 ft 2

hectare (ha) = 100 are = 2.47 acres

C1
 D APPENDIX

glossary

Action spectrum (F. acte, a thing done). A graph relating

Abbreviation Meaning
the degree of physiological response (e.g., phototropism,
A.S. Anglo-Saxon photosynthesis) caused by different wavelengths of light.

D. Dutch
Active solute absorption. The intake of dissolved
dim. diminutive
materials by cells or organelles against an electrochemical
F. French
potential gradient and requiring a supply of metabolic
Gr. Greek energy.
It. Italian
Adaptation (L. ad, to + aptare, to fit). Adjustment of an
L. Latin
organism to the environment.
Lapp. Lappland
M.E. Medieval English Adenine. A purine base present in nucleic acids and
M.L. Medieval Latin nucleotides.
N.L. New Latin Adhesion (L. adhaerere, to stick to). A sticking together
O.F. Old French of unlike things or materials.
O.E. Old English
Adnation (L. adnasci, to grow to). In flowers, the growing
R. Russian
together of two or more whorls to a greater or less extent;
Sp. Spanish
compare connation.
ADP. Adenosine diphosphate
A. Abbreviation tor an angstrom, 0.0001 ot a micron;
there are 10,000,000 angstroms in a millimeter, 10 in a Adsorption (L. ad, to + sorbere, to suck in). The
millimicron, and 10 in a nanometer. concentration of molecules or ions of a substance at a
surface or an interface (boundary) between two
Abscisic acid. A hormone variously inducing abscission,
substances.
dormancy, stomatal closure, growth inhibition, and other
responses in plants. Abbreviated ABA. Advanced (M.E. advaunce, to forward). Said of a
taxonomic trait thought to have evolved late in time from
Abscission zone (L. abscissus, cut otf). Zone ot delicate
some more primitive trait.
thin-walled cells extending across the base ot a petiole,
the breakdown ot which disjoins the leat or truit trom the Adventitious (L. adventicius, not properly belonging to).

stem. Referring to a structure arising from an unusual place:

buds at other places than leaf axils, roots growing from
Absorb (L. ab, away + sorbere, to suck in). To suck up,
stems or leaves.
to drink up, or to take in; in plant cells materials are taken
in (absorbed) in solution. Aeciospore (Gr. aikia, injury + spore). One of the

dikaryotic asexual spores of rust fungi.

Absorption spectrum. A graph relating the ability ot a
substance to absorb light of various wavelengths. Aecium (plural, aecia) (Gr. aikia, injury). In rust, a sorus
that produces aeciospores.
Accessory (M.L. accessorius, an additional appendage).
Something aiding or contributing in a secondary way, such Aerate. To supply or impregnate with common air, such
as buds in addition to the main axillary bud. as by bubbling air through a culture solution.

Achene. Simple, dry, one-seeded indehiscent fruit, with Aerobe (Gr. aer, air + bios, life). An organism that uses
seed attached to ovary wall at one point only. molecular oxygen in its respiratory process.

Acid. A substance that can donate a hydrogen ion. Aerobic respiration. Respiration involving molecular
Actinomorphic (Gr. aktis, ray + morphe, form). Said of
oxygen.
flowers of a regular or star pattern, capable of bisection in Agar (Malay agaragar). A gelatinous substance obtained
two or more planes into similar halves. mainly from certain species of red algae.

D1
Aggregate fruit (L. ad, to + gregare, to collect; to bring borne in a vessel; thus a group of plants whose seeds are
together). A fruit developing from the several separate borne within a matured ovary.
carpels of a single flower; e.g., a strawberry; compare with
Anisogamy (Gr. an, prefix meaning not + isos, equal +
multiple fruit.
gamete, spouse). The condition in which the gametes,

Alcohol (M.L. from Arabic al-kuhl, a powder for painting though similar in appearance, are not identical. Compare
eyelids; later applied, in Europe, to distilled spirits which with heterogamy.
were unknown in Arabia). A product of the distillation of Annual (L. annualis, within a year). A plant that completes
wine or malt; any one of a class of compounds analogous its life cycle within one year and then dies.
to common alcohol; the ending ol designates a member of
Annual ring. Ring of xylem in wood, which indicates an
this class of compounds.
annual increment of growth in trees growing in temperate
Aleurone layer (Gr. aleuron, flour). The outermost cell regions.
layer of the endosperm of wheat and other grains.
Annular vessels (L. annularis, a ring). Vessels with
Alfisol. Modified podsol soil, typical of the northern part of rings of secondary wall material.
the deciduous forest.
Annulus (L. anulus or annulus, a ring). In ferns, a row of
Alga (plural, algae) (L. alga, seaweed). A member of the specialized cells in a sporangium, of importance in
large group of thallus plants containing chlorophyll and opening of the sporangium; in mosses, thick-walled cells
thus able to synthesize carbohydrates. along the rim of the sporangium to which the peristome
Algin. A long-chain polymer of mannuronic acid found in
teeth are attached.

the cell walls of the brown algae. Anther (M.L. anthera, from the Gr. anthros, meaning
flower). Pollen-bearing portion of stamen.
Alkali (Arabic alqili, the ashes of the plant saltwort). A
substance with marked basic properties. Antheridiophore (Gr. anthros, flowers + Gr. phoros, to
bear). In some liverworts, an elongate structure, bearing
Allele (Gr. allelon, of one another, mutually each other).
antheridia.
Variant forms of a gene.
Antherldium (anther + -idion, a Gr. dim. ending, thus a
Alpine (L. Alpes, the Alps Mountains). Meadowlike
little anther). Male gametangium or sperm-bearing organ
vegetation at high elevation, above tree line.
with a sterile jacket of cells around the spermatocytes.
Alternate. Referring to bud or leaf arrangement in which
Anthocyanin (Gr. anthros, a flower + kyanos, dark blue).
there is one bud or one leaf at a node.
A blue, purple, or red vacuolar pigment.
Alternation of generations. The alternation of haploid
Anticlinal cell division. Cell division where the newly
(gametophytic) and diploid (sporophytic) phases in the life
formed cell wall is perpendicular to the axis of the organ
cycle ofmany organisms; the phases (generations) may
surface.
be morphologically quite similar or very distinct, depending
on the organism. Antipodal (Gr. anti, opposite + pous, foot). Referring to
cells at the end of the embryo sac opposite that of the egg
Amensalism (L. a, not + mensa, form of
table). A
apparatus.
biological interaction in which one organism is inhibited by
another, by the addition of something to the environment. Apex (L. apex, a tip, point, or extremity). The tip, point, or
angular summit of anything: the tip of a leaf; that portion
Amino acid One of the building blocks of a protein.
of a root or shoot containing apical and primary
Ammonlflcatlon (Ammon, Egyptian sun god, near whose meristems.
temple ammonium salts were first prepared from camel
Apical dominance. The inhibition of lateral buds or
dung + L. facere, to make). Decomposition of amino meristems by the apical meristem.
acids, resulting in the production of ammonia.
Apical meristem. A mass of meristematic cells at the very
Amyloplast (L. amylum, starch + plastos, formed). tip of a shoot or root.
Cytoplasmic organelle specialized to store starch.
Abundant
Aplanospore (Gr. a, not + planetes, wanderer + sporos,
in roots in storage organs such as tubers.
seed, spore). A nonmotile spore, one that is carried
Anabolism (Gr. ana, up + metabolism). The constructive passively by wind, water, or other organisms.
phase of metabolism, in which more complex molecules
Apomixis (Gr. apo, away from + mixis, a mingling). The
are built from simpler substances.
production of offspring in the usual sexual structures
Anaerobe (Gr. a, without + aer, air + bios, life). An without the mingling and segregation of chromosomes.
organism able to live in the absence of free oxygen, or in
Apotheclum (Gr. apotheke, a storehouse). A cup-shaped
greatly reduced concentrations of free oxygen.
or saucer-shaped open ascocarp.
Anaphase (Gr. ana, up + phais, appearance). That stage
Arboretum (L. arbor, tree, also arboretum, a place grown
in mitosis in which half chromosomes or sister chromatids with trees). A place, often outdoors, set aside for the
move to opposite poles of the cell.
and shrubs as
display of living plants, including herbs well
Androecium (Gr. andros, man + oikos, house). The as trees, contrasts with an herbarium, which displays
aggregate of stamens in the flower of a seed plant. dead, preserved remains of plants.

Angiosperm (Gr. angion, a vessel + sperma from Archegoniophore (Gr. archgonos, founder of a race +
speirein, to sow, hence a seed or germ). Literally a seed Gr. phoros, to bear). Elongate structure found on some

appendix D |
Glossary

D2
liverworts that bears archegonia. Type of meiospore borne by basidia in the
Basidiomycetes.
Archegonlum (L. dim. of Gr. archegonos, literally a little

founder of a race). Female gametangium or egg-bearing Basidium (plural, basidia) (M.L. basidium, a little

organ, in which the egg is protected by a jacket of sterile pedestal). A specialized reproductive cell of the

cells. Basidiomycetes in which nuclei fuse and meiosis occurs. It

may be a special club-shaped cell, a short filamentous

Aril(ML. arillus, a wrapper for a seed). An accessory
cell, or a short four-celled filament.
seed covering formed by an outgrowth at the base of the
ovjle in Taxus.
Benthon (Gr. benthos, the depths of the sea). Attached
aquatic plants and animals, collectively.
Ascus (plural, ascl) (Gr. askos, a bag). A specialized cell,
Berry. A simple fleshy fruit, the ovary wall fleshy and
characteristic of theAscomycetes, in which two haploid
including one or more carpels and seeds.
nuclei fuse, immediately after which three (generally)
divisions occur, two of which constitute meiosis, resulting Biennial (L. biennium, a period of two years). A plant that

in eight ascospores still contained within the ascus.

requires two years to complete its life cycle. Flowering is

normally delayed until the second year.

-ase. A chemical suffix indicating an enzyme.
Bifacial leaf (L. bis, twice + fades, face). A leaf having
Asexual (Gr. a, without + L. sexualis, sexual). Any type upper and lower surfaces and anatomy distinctly different.
of production not involving the union of gametes or
Binomial (L. binominis, two names). Two-named; in
meiosis.
biology each species is generally indicated by two names,
Aspect (L. aspectus, appearance). The direction of slope the genus to which belongs and its own species name.
it

of a surface, as a hillside with a south-facing aspect. Bloassay (Gr. bios, life + L. exagere, to weigh or test).
Assimilation (L. assimilare, to make like). The To test for the presence or quantity of a substance by
transformation of food into protoplasm. using an organism's response as an indicator.

Atom (Gr. atomos, indivisible). A unit of matter, consisting

Biological barrier. A barrier to crossing (hybridization) of

of a dense, central nucleus surrounded by a number of

plants caused by differences in pollination vector or timing
in flower opening, in contrast to physiological barriers
negatively charged electrons that are in constant motion.
(incompatability of pollen with stigma or style) or
The nucleus consists of several positively charged protons
ecological barriers (habitats too far apart).
and uncharged neutrons.
Biology (Gr. bios, life + logos, word, speech, discourse).
ATP. Adenosine triphosphate, a high-energy organic
The science that deals with living things.
phosphate of great importance in energy transfer in

cellular reactions.
Biosystematics (Gr. bios, life + synistanai, to place
together). A field of taxonomy that emphasizes breeding
Auricles (L. auricula, dim. of auris, ear). In grasses, small behavior and chromosome characteristics.
projections that grow out from the opposite side of the leaf
Blotic (Gr. biotikos, relating to life). Relating to life.
sheath at its upper end where it joins the blade.
Biotln. A vitamin of the B complex.
Autotrophic (Gr. auto, self + trophein, to nourish with
Bladder (O.E. blaedre, a blister). A gas-filled sac whose
food). Pertaining to an organism that is able to
buoyancy keeps some aquatic plants upright.
manufacture its own food.
Bloom (Gr. blume, flower). A flower; also the coloring of a
Axial system (L. axis, axle). System of cells in secondary
body of fresh water by a high density of microscopic algal
tissues that are oriented parallel to the long axis of the
organisms.
stem, forms from fusiform initials of the vascular cambium.
Bordered pit. A pit in a tracheid or vessel member having
Axil (L. axilla, armpit). The upper angle between a petiole a distinct rim of the cell wall overarching the pit
of a leaf and the stem from which it grows. membrane.
Axlle placentatlon. Condition where ovules arise on the Botanic garden. See Arboretum.
axis of an ovary with several locules. Botany (Gr. botane, plant, herb). The science dealing with

Axillary bud. A bud formed in the axil of a leaf. plant life.

Bract (L. bractea, a thin plate of precious metal). A

Auxin (Gr. auxein, to increase). A hormone that
type of
modified leaf, from the axil of which arises a flower or an
regulates many aspects of plant growth and development.
inflorescence.
Banner bandum, a standard). Large, broad, and
(M.L.
Bud (M.E. budde, bud). An undeveloped shoot, largely
conspicuous petal of legume type of flower.
meristematic tissue, generally protected by modified scale-
Bark (Swedish bark, rind). The external group of tissues, leaves. Also a swelling on a yeast cell that will become a
from the cambium outward, of a woody stem or root. new yeast cell when released.
+ A
Base. A substance that can accept a proton (H ). Also, Bud scale. modified protective leaf of a bud.
the purine and pyrimidine groups in nucleic acids and Bud scar. A scar left on a twig when the bud or bud
nucleotides are collectively called bases; they act as scales fall away.
bases.
Bulb (L. bulbus, a modified bud, usually underground). A
Basidlospore (M.L. basidium, a little pedestal + spore). short, flattened, or disk-shaped underground stem, with

Glossary

D3
many fleshy scale-leaves filled with stored food. spores produced in a carpogonium.
Bundle scar. Scar where conducting strands passing
left Carposporophyte (Gr. karpos, fruit + sporos, seed,
out of the stem into the leaf stalk were broken off when spore + phuton, plant). One of two sporophyte
the leaf fell. generations in certain red algae; grows attached to the

Bundle sheath. Sheath of parenchyma cells that surround gametophyte generation, in contrast to the
the vascular bundles of leaves, sometimes called border tetrasporophyte.
parenchyma. Caruncle (L. caruncula, dim. of caro, flesh, wart). A
C3 cycle. The Calvin-Benson cycle of photosynthesis, in spongy outgrowth of the seed coat, especially prominent
which the first stable products after C0 2 fixation are three- in the castor bean seed.

carbon molecules. The overall process builds sugar from Caryopsls (Gr. karyon, a nut + opsis, appearance). A
C0 2 ATP, and NADPH.
, simple, dry, one-seeded, indehiscent fruit, with pericarp

C4 cycle. The Hatch-Slack cycle of photosynthesis, in firmly united all around to the seed coat.
which the first stable products after C0 2 fixation are four- Casparian strip. Suberized strip that impregnates the
carbon molecules. radial and transverse walls of endodermal cells.

Callose (L. callum, thick skin + ose, a suffix indicating a Catabolism (Gr. kata, down + metabolism). The phase of
carbohydrate). An amorphous polysaccharide deposited metabolism in which complex substances are broken
around pores in sieve tube members. down into simpler molecules, the chief role being to
Callus (L. callum, thick skin). Mass of thin-walled cells, provide chemically reactive materials for use in anabolism
usually developed as the result of wounding or in tissue and to provide ATP for cellular work.
cultures. Catalyst (Gr. katelyein, to dissolve). A substance that
Calorie (L. calor, heat). The amount of heat needed to accelerates a chemical reaction but that is not used up in

raise the temperature of 1 g of water 1 °C (usually from the reaction.

14.5 to 15.5°C), also called gram-calorie; 1000 calories = Cation exchange (Gr. kata, downward). The replacement
1 kilocalorie. of one positive ion (cation) by another, as on a negatively

Calyptra (Gr. kalyptra, a veil, covering). In bryophytes, an charged clay particle.

envelope covering the developing sporophyte, formed by Catkin (literally a kitten, apparently first used in 1578 to
growth of the venter of the archegonium. describe the inflorescence of the pussy willow). A type of
Calyx (Gr. kalyx, a husk, cup). Sepals collectively; inflorescence, really a spike, generally bearing only

outermost flower whorl. pistillate flowers or only staminate flowers, which

eventually fall from the plant entire.
Cambium cambium, one of the alimentary body fluids
(L.

supposed to nourish the body organs). A layer, usually Caulescent (Gr. kaulos, a plant stem). A plant whose
regarded as one or two cells thick, of persistently stem bears leaves separated by visibly elongated
meristematic tissue, giving rise to secondary tissues, internodes, as opposed to a rosette plant.

resulting in growth in diameter. Cell (L. cella, small room). The smallest unit of material in

Canopy (Gr. kanopeion, a cover over a bed to keep off the organism that is capable of self-reproduction. It is

gnats). The leafy portion of a tree or shrub. surrounded by a plasma membrane and contains a store
of DNA together with a metabolic system.
Capillaries (L. capillus, hair). Very small spaces, or very
fine bores in a tube. Cell wall. A layer of material, chiefly elongated polymers,
that is laid down outside the plasma membrane of most
Capsule (L. capsula, dim. of capsa, a case). A simple,
plant cells, and that serves to protect the protoplast and to
dry, dehiscent fruit, with two or more carpels.
limit its expansion.
Carbohydrate (chemical combining forms, carbo, carbon
Cellulose (cell + ose, a suffix indicating a carbohydrate).
+ hydrate, containing water). Compounds with the
A polysaccharide occurring in the cell walls of the majority
general formula Cn (H 2 0)„ or C n H 2nO„
of plants; it is composed of hundreds of simple sugar
Carbon fixation. The enzymatic reaction in which C0 2 is
molecules, glucose, linked together in a characteristic
attached to a receiver compound such as ribulose manner.
diphosphate, thereby adding to the supply of organic
Cenozoic (Gr. kainos, recent + zoe, life). The geologic
carbon. Occurs chiefly in photosynthesis.
era extending f m 65 million years ago to the present.
Carotene (L. carota, carrot). A reddish-orange plastid
Centromere (L. centrum, center + Gr. meros, part).
pigment.
Specialized part of chromosomes where spindle fibers
Carotenoids. A class of fat-soluble compounds (lipids)
attach andwhere two chromatids connect. Two
that includes carotenes as well as xanthophylls; most of kinetochores, one on each chromatid, compose one
them absorb light and appear yellow, orange or red. centromere.
Carpel (Gr. karpos, fruit). A floral leaf bearing ovules Chalaza (Gr. chalaza, small tubercle). The region on a
along the margins. seed at the upper end of the raphe where the funiculus
Carpogonium (Gr. karpos, fruit + gonos, producing). spreads out and unites with the base of the ovule.
Female gametangium (in red algae). Chaparral (Sp. chaparro, an evergreen oak). A
Carpospore (Gr. karpos, fruit + spore). One of the vegetation type characterized by small leaved, evergreen

appendix p |
Glossary

D4
shrubs growing together into a nearly impenetrable scrub; Citric acid cycle. A system of reactions that contributes
shrubby oaks are found in chaparral of California and the to the catabolicbreakdown of fuels in respiration and that
Mediterranean region, but other genera are typical of provides building materials for a number of important
chaparral in Chile, South Africa, and Australia. anabolic pathways. Also called the Krebs cycle and the

Chemotroplsm (chemo + Gr. tropos, a turning). tricarboxylic acid (TCA) cycle.

Influence of a chemical substance on the direction of Cladode (Gr. kladodes, having many shoots). A branch
growth. resembling a foliage leaf.
Chernozem (R. cherny, black + zem, earth). A soil Class (L. classis, one of the six divisions of Roman
characteristic of some grassland vegetation in warm areas people). A taxonomic group below the division level but
with moderate rainfall; dark in color because of a high
above the order level.
content of organic matter; a molllsol.
Clay. Soil particles less than 2 microns in diameter,
Chlasma (Gr. chiasma, two lines placed crosswise). The
composed mainly oxygen
of aluminum (Al), (O), and
cross formed by breaking, during prophase I of meiosis, of
silicon (S).
two nonsister chromatids of homologous chromosomes
and the rejoining of the broken ends of different Clelstothecium (plura, clelstothecia) (Gr. kleistos, closed
chromatids. + thekion, a small receptacle). The closed, spherical
ascocarp of the powdery mildews.
Chltln (Gr. chiton, a coat of mail). A polymer in which the
monomer unit is the modified sugar /V-acetyl glucosamine; Climax community. The last stage of a natural
it is the principal stiffening material in the cell walls of succession; a community capable of maintaining itself as
most fungi and in the exoskeletons of insects and long as the climate does not change.
crustaceans.
Clone The aggregate of
(Gr. klon, a twig or slip).
Chlamydospore (Gr. chlamys, a horseman's or young individual organisms produced asexually from one sexually
man's coat + spore). A heavy-walled resting asexual produced individual.
spore.
Closed bundle. A vascular bundle lacking residual
Chlorenchyma (Gr. chloros, green + procambium.
-enchyma, a suffix meaning tissue). Parenchyma tissue
Coal Age. The Carboniferous period, beginning 345
possessing chloroplasts.
million years ago and ending 280 million years ago.
Chlorophyll (Gr. chloros, green + phyllon, leaf). The
green pigment found in the chloroplast, important in the
Coalescence (L. coalescere, to grow together). A
condition in which there is union of separate parts of any
absorption of light energy in photosynthesis.
one whorl of flower parts; synonyms are connation and
Chloroplast (Gr. chloros, green + plastos, formed).
cohesion.
Specialized cytoplasmic body, containing chlorophyll, in

which occur important reactions of starch or sugar

Coenocyte (Gr. koinos, shared in common + kytos, a

synthesis.
vessel). A plant or filament whose protoplasm is

continuous and multinucleate and without any division by

Chlorosis (Gr. chloros, green + osis, diseased state).
walls into separate protoplasts.
Failure of chlorophyll development, because of a
nutritional disturbance or because of an infection of virus, Coenzyme. A substance, usually nonprotein and of low
bacteria, or fungus. molecular weight, necessary for the action of some
enzymes.
Chromatid (chromosome + -id, L. suffix meaning
daughters of). The half-chromosome during prophase and Cohesion (L. cohaerere, to stick together). Union or
metaphase of mitosis, and between prophase and I
holding together of parts of the same materials; the union

anaphase of meiosis.
II of floral parts of the same whorl, as petals to petals.

Chromatin (Gr. chroma, color). Substance in the nucleus Coleoptile (Gr. koleos, sheath + ptilon, down, feather).
that readily takes artificial staining; also, that portion that The first leaf in germination of grasses, which sheaths the
bears the determiners of hereditary characters; made up succeeding leaves.
of DNA and protein.
Coleorhiza (Gr. koleos, sheath + rhiza, root). Sheath that
Chromoplast (Gr. chroma, color + plastos, formed). surrounds the radicle of the grass embryo and through
Specialized plastid containing yellow or orange pigments. which the young root bursts.
Chromosome (Gr. chroma, color + soma, body). A Collenchyma (Gr. kolla, glue + -enchyma, a suffix,
condensed mass of chromatin, visible during cell division. derived from parenchyma and denoting a type of cell or

Cilia (singular, clllum) (Fr. cil, an eyelash). Protoplasmic tissue). A stem tissue composed of cells that fit rather
hairs which, by a whiplike motion, propel certain types of closely together and with walls thickened at the angles of
unicellular organisms, gametes, and zoospores through the cells.
water.
Colloid (Gr. kolla, glue + eidos, form). Referring to
Clsterna (plural, clsternae) (L. cisterna, a reservoir). A matter composed of particles, ranging in size from 0.0001
flattened sac, composed of a continuous surrounding to 0.000001 millimeter, dispersed in some medium, as clay
membrane, together with the enclosed space. particles in soil.

Glossary

D5
Colony (L. colonia, a settlement). A growth form originally of quite distinct parents, coming to appear more
characterized by a group of closely associated, but poorly and more alike through time because of selection
differentiated, cells; sometimes filaments can be pressure.
associated together in a colony (as in Nostoc), but more Cork. An external, secondary tissue impermeable water
to
typically unicells are associated in a colony.
and gases.
Community (L. communitas, a fellowship). All the
Cork cambium. The cambium from which cork develops.
populations within a given habitat; usually the populations
are thought of as being interdependent. Corm (Gr. kormos, a trunk). A short, solid, vertical,
enlarged underground stem in which food is stored.
Companion cells. Cell associated with sieve-tube
members. Corolla (L. corolla, dim. of corona, a wreath, crown).
Petals, collectively; usually the conspicuous colored flower
Compensation depth compensare, (L. to
whorl.
counterbalance). That depth, in a body of water, at which
light intensity is too low for photosynthesis of floating or Cortex (L. cortex, bark). Primary tissue of a stem or root
submerged plants to exceed respiration. bounded externally by the epidermis and internally in the
stem by the phloem and in the root by the pericycle.
Competition (L. strive together). A form
competere, to of
biological interaction in which both organisms (at least Cotyledon (Gr. kotyledon, a cup-shaped hollow). Seed
initially) decline in growth or success because of the leaf; two, generally storing food in dicotyledons; one,
insufficient supply of some necessary factor(s). generally a digestive organ in the monocotyledons.

Complete flower. A flower having four whorls of floral Covalent bond. A binding force that holds two atoms
leaves: sepals, petals, stamens, and carpels. together, due to the sharing of electrons.

Compound leaf. A leaf whose blade is divided into Crlstae (L. crista, a crest). Crests or ridges, used here to
several distinct leaflets. designate the infoldings of the inner mitochondrial

Conceptacle (L. conceptaculum, a receptacle). A cavity membrane.

or chamber of a frond (of Fucus, for example) in which Cross-pollination. The transfer of pollen from a stamen to
gametangia are borne. the stigma of a flower on another plant, except in clones.

Cone (Gr. konos, a pine cone). A fruiting structure Crossing-over. The exchange of corresponding segments
composed of modified leaves or branches, which bear between chromatids of homologous chromosomes.
sporangia (microsporangia, megasporangia, pollen sacs,
Cultural eutrophicatlon (Gr. eu, good, well + trephein,
or ovules), and are frequently arranged in a spiral or four-
Organic pollution of bodies of water resulting
to nourish).
ranked order; for example, a pine cone.
from mankind's activities; results in oxygen depletion and
Cone scale. The flat, woody parts of pine cones that a change in the biota; occurs over a shorter period of time
spiral out from the central axis and bear the ovules (and than natural eutrophication.
later seeds) on their upper surface; each is subtended by
Cuticle (L. cuticula, dim. of cutis, the skin). Waxy layer on
a sterile bract; strictly speaking, a cone scale is not
outer wall of epidermal cells.
equivalent to a megasporophyll but, instead, is thought to
represent a modified branch system. Cutin (L. cutis, the skin). Waxy substance that is but
slightly permeable to water, water vapor, and gases; a
Conduction (L. conducere, to bring together). Act of
major part of the cuticle.
moving or conveying water through the xylem in plant
organs. Cutinlzatlon. Impregnation of cell wall with a substance
called cutin.
Conidla (singular, conidlum) (Gr. konis, dust). Asexual
reproductive cells of fungi, arising by the cutting off of Cyme (Gr. kyma, a wave, a swelling). A type of
terminal or lateral cells of special hyphae, or by being inflorescence in which the apex of the main stalk or the

pushed out from a flask-shaped cell. axis of the inflorescence ceases to grow quite early,
relative to the laterals.
Conidiophore (conidia + Gr. phoros, bearing).
Conidium-bearing branch of hypha. Cystocarp (Gr. kystos, bladder + karpos, fruit). A
peculiar diploid spore-bearing structure formed after
Conifer (cone + L. ferre, to carry). A cone-bearing tree;
fertilization in certain red algae.
in the'coniferophyta.

Conjugation (L. conjagatus, united). Process of sexual Cytochrome (Gr. kytos, a receptacle or cell + chroma,
reproduction involving the fusion of isogametes or of color). A class of several electron-transport proteins

specialized cell extensions. serving as carriers in mitochondrial oxidations and in

photosynthetic electron transport.

Connation (L. connatus, to be born together). Condition
in flower where there is a union of similar parts of any one Cytokinesis (Gr. kytos, a hollow vessel + kinesis,

whorl of appendages; synonym of coalescence. motion). Division of cytoplasmic constituents at cell

division.
Conservative (L. conservare, to keep). Said of a
taxonomic trait whose expression is not modified to any Cytoklnin. A class of hormones that participate in

great extent by the external environment; a trait that is controlling many developmental processes in plants.

constant unless its genetic base is changed. Cytology (Gr. kytos, a hollow vessel + logos, word,
Convergent evolution. Process of successive progeny, speech, discourse). The science dealing with the cell.

appendix D |
Glossary

D6
Cytoplasm (Gr. kytos, a hollow vessel + plasma, form). Diffusion (L. diffusus, spread out). The movement of
All the protoplasm of a protoplast outside the nucleus. molecules from a region of higher concentration to a
DNA region of lower concentration.
Cytoslne. A pyrimidine base found in and RNA.
Deciduous (L. deciduus, falling). Referring to trees and Digestion (L digestio, dividing, or tearing to pieces, an
shrubs that lose their leaves in the fall.
The processes of rendering food
orderly distribution).
available for metabolism by breaking it down into simpler
Decomposer (L. de, from + componere, to put toqether).
compounds, chiefly through actions of enzymes.
An organism that obtains food by breaking down dead
organic matter into simpler molecules. Dlkaryon (Gr. di, two + karyon, nut). A hypha or
mycelium in which each cell contains two haploid nuclei,
Decomposition (L. de, to denote an act undone +
the two usually derived from different parent organisms.
componere, to put together). A separation or dissolving
The dikaryotic condition is often abbreviated as the n +n
into simpler compounds; rotting or decaying.
condition.
Dehiscent (L. dehiscere, to split open). Opening
Dinoflagellate (Gr. dinein, to whirl + L. flagellum, a
spontaneously when ripe, splitting into definite parts.
whip). The common name for members of the algal
Deletion (L. deletus, to destroy, to wipe out). Used here to division Pyrrhophyta; the organisms are typically
designate an area, or region, lacking from a chromosome. unicellular and motile, with cell walls made up of

Dendrogram (Gr. dendron, tree + gramme, what is overlapping plates.

written or drawn). A graph showing relationship between Dioecious (Gr. dis, twice + oikos, house). Unisexual;
things at different levels of similarity; the graph resembles having the male and female elements in different
the limbs of a tree.
individuals.
Denitriflcation (L. de, to denote an act undone + nitrum,
Diploid (Gr. diploos, double + oides, like). Having a
nitro, a combining form indicating the presence of nitrogen
double set of chromosomes, or referring to an individual
+ facere, to make). Conversion of nitrates into nitrites, or
containing a double set of chromosomes per cell; usually
into gaseous oxides of nitrogen, or even into free nitrogen.
a sporophyte generation.
Deoxyrlbose nucleic acid (DNA). Hereditary material; the
Divergent evolution. Process of successive progeny,
DNA molecule carries hereditary information.
originally of quite similar parents, coming appear more
to
Desert scrub (M.E., schrubbe, shrub). A vegetation type and more different through time because of isolation and
characterized by evergreen or drought-deciduous shrubs selection pressure.
growing together rather openly generally in an area with
Division. A major portion of the plant kingdom; equivalent
annual precipitation below 25 cm.
to phylum.
Detritus (L. detritus, worn away). Particulate organic
DNA. Deoxyribonucleic acid.
matter released in the process of decomposition of dead
organisms or parts of organisms (such as plant litter). Dominant (L. dominah, to rule). Referring, in ecology, to
species of a community that receive the full force of the
Development (F. developper, to unfold). Changes in the
macroenvironment; usually the most abundant of such
plant body that result from controlled processes of cell
species.
division, cell growth, and cell differentiation.

Diatom Member Dormant (L. dormire, to sleep). Being in a state of

(Gr. diatomos, cut in two). of a group of
reduced physiological activity such as occurs in seeds,
golden brown algae with silicious cell walls fitting together
buds, etc.
much as do the halves of a pill box.
Dorsiventral (L. dorsum, the back + venter, the belly).
Diatomite. Diatomaceous earth; that is, sedimentary
Having upper and lower surfaces distinctly different, as
deposits made up of the silica wall remains of diatoms.
many leaves do.
Dicotyledon (Gr. dis, twice + kotyledon, a cup-shaped
hollow). A plant whose embryo has two cotyledons. Double bond. A covalent bond that involves four
electrons.
Dlctyosome (Gr. diktyon, a net + soma, body). One of
thecomponent parts of the Golgi apparatus; in plant cells Drupe (L. drupa, an overripe olive). A simple, fleshy fruit,

a complex of flattened double lamellae. derived from a single carpel, usually one-seeded, in which
the exocarp is thin, the mesocarp fleshy, and the
Differentially permeable. See Selectively permeable.
endocarp stony.
Differentiation (L. differre, to carry different ways).
Early wood. That portion of an annual ring formed during
Developmental change of a cell leading to the presence of
spring, characterized by large cells and thin walls, also
features that equip the cell for performing specialized
called spring wood.
functions.
Ecology (Gr. o/7<os, home + logos, discourse). The study
Diffuse porous. Wood with an equal and random
of life in relation to environment.
distribution of large xylem vessel members throughout the
growth season. Ecosystem (Gr. oikos, house + synistanai, to place
An inclusive term for a
together). living community and all
Diffuse secondary growth. Secondary growth, such as in
the factors of its nonliving environment.
palm trees, that is caused by a proliferation of
parenchyma cells and not by vascular cambium. Ecotype (Gr. oikos, house + typos, the mark of a blow).

Glossary

D7
Genetic variant within a species that is adapted to a Endosperm mother cell. One of the seven cells of the
particular environment yet remains intertertile with all other mature embryo sac, containing the two polar nuclei and,
members of the species. after reception of a sperm cell, giving rise to the primary

edaphos, Pertaining to endosperm cell from which the endosperm develops.

Edaphlc (Gr. soil). soil conditions
that influence plant growth. Environment (O.F., environ, around). The living and
A nonliving factors that surround a given organism or
Egg (A.S. aeg, egg). female gamete.
community of organisms.
Elater (Gr. elater, driver). An elongated, spindle-shaped,
sterile, hygroscopic cell in the sporangium of a some Enzyme (Gr. en, in + zyme, yeast). A protein that acts as

Bryophyta sporophytes. a catalyst to speed chemical reactions.

Electron (Gr. elektron, gleaming in the sun, from L. Epicotyl (Gr. epi, upon + kotyledon, a cup-shaped

electrum, amber, from which electricity was first produced hollow). The upper portion of the axis of embryo or

by friction). An elementary particle of matter bearing a unit seedling, above the cotyledons.
of negative electrical Low in mass and in constant
charge. Epidermis upon + derma, skin). A superficial
(Gr. epi,
rapid motion, electrons repel one another and are layer of cells occurringon all parts of the primary plant
attracted to the positively charged atomic nucleus. The body: stems, leaves, roots, flowers, fruits, and seeds; it is
motion of the electrons defines the size and chemical absent from the root cap and not differentiated on the
properties of the atom or molecule. apical meristems.

Electron transport chain. A membrane-bound system Eplgyny (Gr. epi, upon + gyne, woman). The
that controls the flow of electrons from reduced to arrangement of floral parts in which the ovary is
oxidized compounds, so that some of the energy carried embedded in the receptacle so that the other parts appear
by the electrons is used to form ATP. The chain consists to arise from the top of the ovary.
of several compounds (carriers) that alternately accept
Epiphyte (L. epi, upon + phyton, a plant). A plant that
and donate electrons. Found in mitochondria and
grows upon another plant, but is not parasitic.
chloroplasts.

A microscope uses a beam of

ER. Endoplasmic reticulum.
Electron microscope. that
electrons rather than light to produce a magnified image. Ethylene. C 2 H 4 a hormone that participates
,
in the control

Element (L. elementa, the first principles). In modern of many developmental processes in plants.
chemistry, a substance that cannot be divided or reduced Etiolation (F. etioler, to blanch). A condition involving
by any known chemical means to a simpler substance; 92 increased stem elongation, poor leaf development, and
natural elements are known, of which gold, carbon, lack of chlorophyll, found in plants growing in the
oxygen, and iron are examples; several, including absence, or in a greatly reduced amount, of light.

Plutonium, have been formed in atomic piles.

Eukaryote (L. eu, true + karyon. A nut, referring in
Embryo (Gr. en, in + bryein, to swell). A young modern biology to the nucleus); any organism
sporophytic plant, while still retained in the gametophyte characterized by having cellular organelles, including a
or in the seed. nucleus, bounded by membranes.
Embryo sac. The female gametophyte of the Eutrophicatlon (Gr. eu, good, well + trephein, to
angiosperms; generally a seven-celled structure; the seven nourish). Pollution of bodies of water resulting from slow,
cells are two synergids, one egg cell, three antipodal cells
natural, geological, or biological processes such as
(each with a single haploid nucleus), and one endosperm siltation or encroachment of vegetation or accumulation of
mother cell with two haploid nuclei. detritus, also called natural eutrophication.

Endocarp (Gr. endon, within + karpos, fruit). Inner layer

Evapotranspiratlon (L. evaporare, e, out of + vapor,
of fruit wall (pericarp).
vapor + F. transpirer, to perspire). The process of water
Endodermis (Gr. endon, within + derma, skin). The layer loss in vapor form from a unit surface of land both directly
of living cells, with various characteristically thickened and from leaf surfaces.

walls and no intercellular spaces, which surrounds the

an The development
Evolution (L. evolutio, unrolling). of a
vascular tissue in nearly all roots and certain stems and group
race, genus or other larger of plants or animals.
leaves.
Exine (L. exterus, outside). Outer coat of pollen.
Endogenous (Gr. endon, within + genes, born).
Produced within the cell or organism. Exocarp (Gr. exo, without, outside + karpos, fruit).

Outermost layer of fruit wall (pericarp).

Endoplasmic reticulum (Gr. endon, within + plasma,
anything formed or molded; L. reticulum, a small net). Exogenous (Gr. exe, out, beyond + genos, race, kind).
A system of membrane-bound cysternae found in the Produced outside of, originating from, or due to external
cytoplasm. causes.

Endosperm endon, within + sperma, seed). The

(Gr. Facultative (L. facultas, capability). Referring to an
nutritive tissue formed within the embryo sac of seed organism having the power to live under a number of
plants; it is often consumed as the seed matures, but certain specific conditions, e.g., a facultative parasite may
remains in the seeds of corn and other cereals. be either parasitic or saprophytic.

appendix D |
Glossary

D8
Family (L. familia, family). In plant taxonomy, a group of Frond (L. frons, branch, leaf). A synonym for a large
genera; families are grouped in orders. divided leaf, especially for a fern leaf.

Fascicle (L. fasciculus, a small bundle). A bundle of Fruit (L. frucfus, that which is enjoyed, hence product of
leaves arising from one point on the stem. the soil, trees, cattle, etc.). A matured ovary; in some seed
Fascicular cambium. Cambium within vascular bundles. plants other parts of the flower may be included; also
applied, as fruiting body, to reproductive structures of
Fermentation (L. fermentum, a drink made from
some fungi.
fermented barley, beer). Catabolic breakdown of fuels by a
process that does not involve molecular oxygen. Frustule (L. frustulum, little piece). A diatom cell,

composed of two overlapping halves (valves).

Fern (O.E., fearn, fern). Common name for members of
the division Pterophyta, part of the lower vascular plants. Fucoxanthin (Gr. phykos, seaweed + xanfhos, yellowish
brown). A brown pigment found in brown algae.
Ferredoxln. An electron-transferring protein containing
iron, involved in photosynthesis and in nitrogen fixation. Fungus (plural fungi) (L. fungus, a mushroom). A
eukaryotic organism that lacks plastids and that
Fertilization (L. fertilis, capable of producing fruit). The
reproduces by means of spores.
union of egg and sperm.
An elongated, tapering,
Funiculus (L. funiculus, dim. of funis, rope or small cord).
Fiber (L. libra, a fiber or filament).
A stalk of the ovule, containing vascular tissue.
thick-walled strengthening cell occurring in various parts
of plant bodies. Fusiform initials (L. fusus, spindle + form). Meristematic

Flber-tracheid. Xylem elements found in pine that are

cells in the vascular cambium that develop into xylem and
phloem cells comprising an axial system.
structurally intermediate between tracheids and fibers.

Field capacity. The amount of water retained in a soil

Gametanglum (Gr. gametes, a husband, gamete, a wife

(generally expressed as percent by weight) after large

+ angeion, a vessel). Organ bearing gametes.
capillary spaces have been drained by gravity. Gamete (Gr. gametes, a husband, gamete, a wife). A
protoplast that fuses with another protoplast to form the
Filament (L. filum, a thread). Stalk of stamen bearing the
anther at its tip; also, a slender row of cells (certain
zygote in the process of sexual reproduction.

algae). Gametic life cycle. A life cycle in which the haploid

Flagellum (plural, flagella) (L. flagellum. a whip). A long,
phase is limited to gametes, as in some diatoms and many
slender whip of protoplasm. animals.

Flora (L. floris, a flower). An enumeration of all the Gametophyte (gamete + Gr. phyton, a plant). The
species that grow in a region; also, the collective term for gamete-producing plant.
all the species that grow in a region. Gel (L. gelare, to freeze). Jellylike colloidal mass.
Floret (F. fleurette, a dim. of fleur, flower). One of the Gemma (plural, gemmae) (L. gemma, a bud). A small
small flowers that make up the composite inflorescence mass of reproductive tissue in some fungi and bryophytes.
(head) or the spike of the grasses.
Gene (Gr. genos, race, offspring). A group of base pairs
Flower (F. fleur, L. flos, a flower). Floral leaves grouped in the DNA molecule that determines one or more
together on a stem and adapted for sexual reproduction in hereditary characters.
the angiosperms.
Gene recombination. The appearance of gene
Follicle (L. folliculus, dim. of follis, bag). A simple, dry, combinations in the progeny different from the
dehiscent fruit, with one carpel, splitting along one suture. combinations present in the parents.
Food (A.S. foda). Any organic substance that furnishes Generation (L. genus, birth, race, kind). Any phase of a
energy and building materials directly for vital processes. life cycle characterized by a particular chromosome
Food chain. The path along which caloric energy is number, as the gametophyte generation and the
transferred within a community (from producers to sporophyte generation.
consumers to decomposers). Genetics (Gr. genesis, origin). The science of heredity.

Foot (O.E., fot, foot). That portion of the sporophyte of Genotype (gene + type). The assemblage of genes in an
bryophytes and lower vascular plants which is sunk in
organism.
gametophyte tissue and absorbs food parasitically from
Genus (plural, genera) (Gr. genos, race, stock). A group
the gametophyte.
of structurally and phylogenetically related species.
Fossil (L. fossio, a digging). Any impression, impregnated
Geotropism (Gr. ge, earth + tropos, turning). A growth
remains, or other trace of an animal or plant of past
curvature induced by gravity.
geological ages that has been preserved in the earth's
crust. Germination (L. germinare, to sprout). The beginning of
growth of a seed, spore, or other once-dormant structure.
Fossil fuel. Hydrocarbon deposits, currently mined and
refined for use as fuel (coal, gas, oil), which were Gibberelllns. A class of hormones that participate in
originally deposits of detritus of now-extinct plants. controlling many developmental processes in plants.

Fret (O. F. frette, latticework). Flattened membrane sacs Girdle (O.E. gyrdel, enclosure, girdle). That region of a

that connect grana in chloroplasts. frustule where the two valves overlap.

Glossary

D9
Glucose (Gr. glykys, sweet + ose, a suffix indicating a Helix (Gr. helix, anything twisted). Anything having a
carbohydrate). A common hexose sugar. spiral form.

Glume (L. gluma, husk). An outer and lowermost bract of Hemlcellulose. A class of polysaccharides of the cell
a grass spikelet. wall, built of several different kinds of simple sugars linked
Glycogen (Gr. glykys, sweet + genes, born). A in various combinations.
polysaccharide built of glucose, akin to starch, serving as Herb (L. herba, grass, green blades). A seed plant that
a food reserve in animals and fungi and some prokaryotes. does not develop woody tissues.
Glycolysis (Gr. glykys, sweet + lysis, a loosening). Herbaceous (L. herbaceus, grassy). Referring to plants
Decomposition of sugar compounds without involving free having the characteristics of herbs.
oxygen; early steps of respiration.
Herbal (L. herba, grass). A book that contains the names
Golgl body (Italian cytologist Camillo Golgi 1844-1926, and descriptions of plants, especially those which are
who first described the organelle). In animal cells, a thought to have medicinal uses.
complex perinuclear region thought to be associated with
Herbarium (L. herba, grass). A collection of dried and
secretion; in plant cells a series of flattened plates, more pressed plant specimens.
properly called dlctyosomes.
Herbicide (L. herba, grass or herb + cidere, to kill). A
Grana (singular, granum) (L. granum a seed). Structures
chemical used to kill plants, frequently chemically related
within chloroplasts, seen as a series of parallel lamellae
to a hormone.
with the electron microscope.
Heredity (L. hereditas, being a heir). The transmission of
Ground cover. The area of ground covered by a plant morphological and physiological characters of parents to
when its canopy edge is projected perpendicularly down. their offspring.
Ground meristem (Gr. meristos, divisible). A primary
Heterobasidiomycetidae (Gr. heteros, other +
meristem that gives rise to cortex and pith.
Basidiomycete). A subclass of Basidiomycetes with
Ground tissues. Category of primary tissues variable basidia, never club-shaped cells.
(parenchyma, collenchyma, and sclerenchyma) that + cytis, a
Heterocyst (Gr. heteros, different bag). An
provide such basic functions as storage, support, and enlarged colorless cell that may occur in the filaments of
secretion.
certain blue-green algae; associated with nitrogen fixation.
Growth (A.S. growan, probably from Old Teutonic gro,
Heteroeclous (Gr. heteros, different + oikos, house).
from which grass is also derived). An irreversible increase Referring to fungi that cannot carry through their complete
in size.
life cycle unless two different host species are present.
Growth retardant. A chemical (such as cycocel, CCC) Heterogametes (Gr. heteros, different + gamete).
that selectively interferes with normal hormonal promotion Gametes dissimilar from each other in size and behavior,
of growth — but without appreciable toxic effects.
like egg and sperm.

Guanine. A purine base found in DNA and RNA. Heterogamy (Gr. heteros, different + gamos, union or
Guttatlon (L. gutta, drop, exudation of drops). Exudation reproduction). Reproduction involving two types of
of water from plants, in liquid form. gametes.
Gynoeclum (Gr. gyne, woman + oikos, house). The Heterospory (Gr. heteros, different + spore). The
aggregate of carpels in the flower of a seed plant. condition of producing microspores and megaspores.

Gymnosperm (Gr. gumnos, naked + sperma, seed, Heterothallic (Gr. heteros, different + thallus). Referring
common name for a group of divisions,
sperm). The which male gametangia and female
to species in
including the Coniferophyta, which bear exposed seeds, in gametangia are produced by different individual plant
contrast to the flowering plants that bear the seeds bodies.
enclosed in a fruit (mature ovary). Heterotrichy (Gr. heteros, different + trichos, a hair). In
Haplold (Gr. haploos, single + oides, like). Having a the algae, the occurrence of two types of filaments, erect
single complete set of chromosomes, or referring to an and prostrate.
individual or generation containing such a single set of
Heterotrophic (Gr. heteros, different + trophein, to
chromosomes per cell; usually a gametophyte generation. nourish with food). Referring to a plant obtaining
Hapteron (Gr. haptein, to fasten). An individual branch of nourishment from outside sources.
the hold fast organ of a kelp; also called haptere (plural,
Heterozygous (Gr. heteros, different + zygon, yoke).
hapteres). Having two different alleles for a given gene in the diploid
Haustorium (plural, haustorla) (ML. haustrum, a pump). cell.

A projection that acts as a penetrating and absorbing Hexose (Gr. hexa, six + -ose, suffix indicating
organ. A carbohydrate with
carbohydrate). six carbon atoms
Head. An inflorescence, typical of the composite family, in (e.g., C 6 H 12 6 ).
which flowers are sessile and grouped closely on a Higher vascular plant. Common name for extant seed-
receptacle. producing plants; hence now includes only gymnosperm
Heartwood. Wood in the center of old secondary stems and angiosperm divisions although in the past other
that is plugged with resins and tyloses and is not active. groups did produce seeds.

appendix D |
Glossary

D10
Hllum (L. hilum, a trifle). Scar on seed, which marks the food). A condition of overgrowth or excessive development
place where the seed broke from the stalk. of an organ or part.

Homobasldiomycetidae (Gr. homo, the same + Hypha (plural, hyphae) (Gr. hyphe, a web). A slender,
Basidiomycete). A subclass of Basidiomycetes with a elongated, threadlike cell or filament of cells of a fungus.
typical club-shaped cell as a basidium. Hypocotyl (Gr. hypo, under + kotyledon, a cup-shaped
Homologous chromosomes (Gr. homologos, the same). hollow). That portion of an embryo or seedling between
Members of a chromosome pair; they may be the cotyledons and the radicle or young root.
heterozygous or homozygous.
Hypogyny (Gr. hypo, under + gyne, female). A condition
Homospory (Gr. homos, one and the same + spore). in which the receptacle is convex or conical, and the
The condition of producing one sort of spore only. flower parts are situated one above another in the

Homothalllc (Gr. homos, one and the same + thallus). following order, beginning with the lowest: sepals, petals,

Referring to species in which male gametangia and female stamens, carpels.

gametangia are produced. By the same individual plant Hypothesis (Gr. hypothesis, foundation). A tentative
body. theory or supposition provisionally adopted to explain
Homozygous homos, one and the same + zygon,
(Gr. certain facts and to guide in the investigation of other
yoke). Having two identical alleles for a given gene in the facts.

diploid cell. IAA. Indoleacetic acid.

Hormogonla (singular, hormogonlum) (Gr hormos, Imbibition (L. imbibere, to drink). The absorption of
necklace + gonos, offspring). Short filaments, the result liquids or vapors into the ultramicroscopic spaces or pores
of a breaking apart of filaments of certain blue-green algae found in such materials as cellulose or a block of gelatine;
at the heterocysts. an adsorption phenomenon.
Hormone (Gr. hormaein, to excite). A compound that is Imperfect flower. A flower lacking either stamens or
normally produced by a plant and whose sole function is pistils.
to act as a signal in controlling development.
Imperfect fungi. Fungi reproducing only by asexual
Humidity, relative (L. humidus, moist). The ratio of the means.
weight of water vapor in a given quantity of air, to the total
Incomplete flower. A flower lacking one or more of the
weight of water vapor that quantity of air is capable of
four kinds of flower parts.
holding at the temperature in question, expressed as
percent. Indehlscent (L. in, not + dehiscere, to divide). Not
opening by valves or along regular lines.
Humus (L. humus, the ground). Decomposing organic
matter in the soil. Indicator species. A species that has a narrow range of
tolerance for one or more environmental factors so that,
Hybrid (L. hybrida, offspring of a tame sow and a wild
from its occurrence at a site, one can predict these factors
boar, a mongrel). The offspring of two plants or animals
at that site (e.g., nutrient availability or summer
differing in at least one Mendelian character; or the
temperatures).
offspring formed by mating two plants or animals that
differ genetically. Induslum. Membranous growth of the epidermis of a fern
leaf that covers a sorus.
Hydathode (Gr. hydro, water + O.E. thoden, stem or
thyddan, to thrust). A structure, usually on leaves, which Infect (L. infectus, to put into, to taint with morbid matter).
releases liquid water during guttation. Specifically to produce disease by such agents as bacteria
or viruses.
Hydrogen acceptor. A substance capable of accepting
hydrogen atoms or electrons in the oxidation-reduction Inferior ovary. An ovary more or less, (sometimes
reactions of metabolism. completely) attached to the calyx and corolla.

Hydrogen bond. A weak bond that occurs between a Inflorescence (L. inflorescere, to begin to bloom). A
hydrogen atom and an oxygen or nitrogen atom when the flower cluster.
hydrogen is already covalently bonded to another oxygen Inheritance (O.F. enheritance, inheritance). The reception
or nitrogen atom. The hydrogen bond is not covalent. or acquisition of characters or qualities by transmission of
Hydrolysis (Gr. hydro, water + lysis, loosening). Reaction parent to offspring.
of a compound with water, attended by decomposition into Inorganic. Referring to compounds that do not contain
less complex compounds. both carbon and hydrogen.
Hydrophyte (Gr. hudor, water + phuton, plant). A plant
Integuments (L. integumentum, covering). Cell layers
that normally grows in a wet habitat. around ovule that develop into the seed coat.
Hymenlum (Gr. hymen, a membrane). Spore-bearing Interphase between + phase). Period between
(L. inter,
tissue in various fungi.
mitotic divisions, consists of G1 pre-DNA synthesis phase;
,

Hypanthlum (L. hypo, under + Gr. anthos, flower). S, DNA synthesis; and G2, post-DNA synthesis phase.
Fusion of calyx and corolla part way up their length to
Intercalary (L. intercalare, to insert). Descriptive or
form a cup, as in many members of the rose family. meristematic tissue or growth not restricted to the apex of
Hypertrophy (Gr. hyper, over + trophein, to nourish with an organ, i.e., growth at nodes.

Glossary

D11
Intercellular (L. inter, between + cells). Lying between Cellular membranes, frequently those seen in

cells. choloroplasts.

Interfascicular cambium (L. inter, between + fasciculus, Lamina (L. lamina, a thin plate). Blade or expanded part
small bundle). Cambium that develops between vascular of a leaf.

bundles. Late wood. That portion of an annual ring which formed

Internode (L. inter, between + nodus, a knot). The region during summer and fall, characterized by small diameter
of a stem between two successive nodes. cells with thick walls, also called summer wood.
Intlne (L. intus, within). The innermost coat of a pollen Lateral bud. A bud that grows out of the side of a stem.

grain. Laterite (L. later, a brick). A soil characteristic of rain

Intracellular (L. intra, within + cell). Lying within cells. forest vegetation; color is red from oxidized iron in the A
horizon; in the oxisol soil order.
Introgresslve hybridization (L. intro, to the inside +
gress, walk + hybrida, halfbreed). Back-crossing between Latex (L. latex, juice). A milky secretion.

complete or partial hybrids and the original parental stock. Leach (O.E. leccan, to moisten). To extract a soluble or

Involucre (L. involucrum, a wrapper). A whorl or rosette moveable substance (as ions or clay particles or bits of

of bracts surrounding an inflorescence.

organic matter) by causing water to filter down through a
material (as a soil horizon).
Ion. An atom or molecule that has a net negative or
Leaf. Lateral outgrowth of stem axis, which is the usual
positive electrical charge because the number of electrons
primary photosynthetic organ, and in the axil of which
does not equal the number of protons.
may be bud.
Ionic compound. A substance whose molecules readily
Leaf axil. Angle formed by the leaf stalk and the stem.
break up into ions when placed in contact with water.
Leaf scar. Characteristic scar on stem axis made after
Irregular flower. A flower in which one or more members
leaf abscission.
of at least one whorl are of different form from other
members of the same whorl; zygomorphic flower. Leaflet. Separate part of the blade of a compound leaf.

Isobilateral leaf (Gr. isos, equal + L. bis, twice, twofold

Leaf primordium (L. primordium, a beginning). A lateral
+ lateralis, pertaining to the side). A leaf having the upper outgrowth from the apical meristem, which will become a
leaf.
and lower surfaces and anatomy essentially similar.

Isodiametric (Gr. isos, equal + diameter). Having

Legume (L. legumen, any leguminous plant, particularly
bean). A simple, dry dehiscent fruit with one carpel,
diameters equal in all directions, as a ball.
splitting along two sutures.
Isogametes (Gr. isos, equal + gametes). Gametes similar
in size and behavior.
Lemma (Gr. lemma, a husk). Lower bract that subtends a
grass flower, but above the glumes.
Isogamy (Gr. isos, equal + gamete, spouse). The
Lenticel (M.L. lenticella, a small lens). A structure of the
condition in which the gametes are identical.
bark that permits the passage of gas inward and outward.
Isolating barriers. Any barrier to the crossing
Leucoplast (Gr. leukos, white + plastos, formed). A
(hybridization) of plants; includes biological, physiological,
colorless plastid.
and ecological categories.
Liana (F. liane from Her, to bind). A plant that climbs upon
Isomers (Gr. isos, equal + meros, part). Two or more
other plants, depending upon them for mechanical
compounds having the same molecular formula but
support, a plant with climbing shoots.
different internal structure; for example, glucose and
fructose, both having the formula C 6 H 12 .
Lichen (Gr. leichen, thallus plants growing on rocks and
6
trees).A composite plant consisting of a fungus living
K selection. Natural selection that favors long-lived, late-
symbiotically with an alga.
maturing individuals that devote a small fraction of their
Lignification (L. lignum, wood + facere, to make).
resources into reproduction; tree species are K strategists.
Impregnation of a cell wall with lignin.
Karyogamy (Gr. karyon, nut + gamos, marriage). The
Lignin (L. lignum, wood). An organic substance or group
fusion of two nuclei.
of substances impregnating the cellulose framework of
Keel (A.S. ceol, ship). A structure of the legume type of certain plant cell walls.
flower, made up of two petals loosely united along their
Ligule (L. ligula, dim. of lingua, tongue). In grass leaves,
edges.
an outgrowth from the upper and inner side of the leaf
Kelp. (M.E. culp, seaweed). A collective name for any of blade where it joins the sheath.
the large brown marine algae.
Line transect. A method of sampling vegetation by
Kinetochore. (Gr. kinein, to move + chorein, to move stretching a tape along a straight line and measuring the
apart). Specialized portion of chromosome, marks point of canopy cover of plants beneath that line or which cut
spindle fiber attachment, related to mechanism of through a vertical plane described by that line.
chromosome movement.
Linkage. The grouping of genes on the same
Krebs cycle. See Citric acid cycle. chromosome.
Lamella (plural, lamellae) (Gr. lamin, a thin blade). Linked characters. Characters of a plant or animal

appendix p |
Glossary

D12
controlled by genes grouped together on the same resulting in four megaspores; synonymous with megaspore
chromosome. mother cell of other texts.

Lipase (Gr. lipos, fat + -ase, suffix indicating an enzyme). Megasporophyll (Gr. megas, large + spore + Gr.
Any enzyme that breaks fats into glycerin and fatty acids. phyllon, leaf). A leaf bearing megasporangia.

Lipids (Gr. lipos, fat + L. ides, suffix meaning son of; now Melocyte (meiosis + Gr. kyios, currently meaning a cell).

used sense of having the quality of). An artificial

in Any cell in which meiosis occurs.

grouping of compounds consisting of substances that are make

Meiosis (Gr. meioun, to smaller). Two special cell
insoluble in water and soluble in fat solvents.
divisions occurring in the life cycle of every sexually
Liverwort (liver + M.E. wort, a plant, literally, a liver plant, reproducing plant and animal, halving the chromosome
so named in medieval times because of its fancied number and effecting a segregation of genetic
resemblance to the lobes of the liver). Common name for determiners.
the Class Hepaticae of the Bryophyta
Meiospore (meiosis + spore). Any spore resulting from
Loam (O.E. lam or Old Teutonic lai, to be sticky, clayey). the meiotic divisions.
A particular soil texture class, referring to a soil having
Membrane (L. membrana, skin covering the separate
30-50% sand, 30-40% silt, and 10-25% clay.
members of the body). A limiting protoplasmic surface,
Lobed leaf (Gr. lobos, lower part of the ear). A leaf consisting of protein and lipid, which surrounds cellular
divided by clefts or sinuses. organelles.

Locule (L. loculus, dim. of locus, a place). A cavity of the Meristem (Gr. meristos, divisible). Undifferentiated tissue,
ovary in which ovules occur. the cells of which are capable of active cell division and
Lodlcules (L. lodicula, a small coverlet). Two scalelike differentiation into specialized tissues.

structures that lie at the base of the ovary of a grass Meristoderm (meristem + epidermis). The outer
flower. meristematic cell layer (epidermis) of some Phaeophyta.
Lower vascular plant. Common name for extant vascular Mesocarp (Gr. mesos, middle + karpos, fruit). Middle
plants thatdo not produce seeds (even though in the past layer of fruit wall (pericarp).
some of their members did produce seeds); the
Mesophyll (Gr. mesos, middle + phyllon, leaf).
Psilophyta, Lycophyta, Sphenophyta, and Pterophyta
Parenchyma tissue of leaf between epidermal layers.
divisions.
Mesophyte (Gr. mesos, middle + phuton, plant). A plant
Lumen (L. lumen, light, an opening for light). The cavity of
that normally grows in moist habitats.
the cell within the cell walls.

Lysis (Gr. lysis, a loosening). A process of disintegration

Mesozoic (Gr. mesos, middle + zoe, life). A geologic era
beginning 225 million years ago and ending 65 million
or cell destruction.
years ago.
Macroenvironment (Gr. makros, large + O.F. environ,
about). The environment due to the general, regional
Metabolism (M.L. from Gr. metabolos, to change). The
overall set of chemical reactions occurring in an organism
climate; traditionally measured some 1 .5 m above the
or cell.
ground and away from large obstructions.
Macronutrient (Gr. makros, large + L. nutrire, to
Metabolite (Gr. metabolos, changeable + ites, one of a

nourish). An essential element required by plants in

group). A chemical that is a normal cell constituent

relatively large quantities.

capable of entering into the biochemical transformations
within living cells
Mating types. A term applied to organisms that show no
male-female differentiation. Two individuals that can
visible Metamorphic rock (Gr. meta, change + morphe, shape
mate belong to different mating types. Mating types are or form). One of three major categories of rock; rocks

often designated as ( + ) and (-). whose original structure or mineral composition has been
changed by pressures or temperatures in the earth's crust.
Medulla (L. medulla, marrow). The filamentous center of
certain lichens and kelp blades and stipes. Metaphase (Gr. meta, after + phsis, appearance). Stage
of mitosis during which the chromosomes, or at least the
Megaphyll (Gr. megas, great + phyllon, leaf). A leaf
kinetochores, lie in the central plane of the spindle.
whose trace is marked with a gap in the stem's vascular
system. Metaxylem. Last formed primary xylem.

Megasporangiate cone. In gymnosperms, a cone that Microbody (Gr. mikros, small + body). A cellular

produces megaspores and, ultimately, seeds; synonyms organelle, always bound by a single membrane, frequently
include female cone, ovulate cone, seed cone. spherical, from 20 to 60 nanometers in diameter,
containing a variety of enzymes.
Megasporangium megas, large + sporangium).
(Gr.
Sporangium that bears megaspores. Microcapillary space. Exceedingly small spaces, such as
those found between microfibrils of cellulose.
Megaspore (Gr. megas, large + spore). The meiospore
of vascular plants, which gives rise to a female Microenvironment (Gr. mikros, small + O.F. environ,
gametophyte. The environment close enough to the surface of a
about).
living or nonliving object to be influenced by it.
Megasporocyte megas, large + spora, seed + L.
(Gr.
cyta, vessel). A diploid cell in which meiosis will occur, Microfibrils (Gr. mikros, small + fibrils, dim. of fiber).

Glossary

D13
Very small fibers of the cell wall. Molecule (F. mdle, + clue, a dim.; literally, a little mass).
Microfossll (Gr. mikros, small + L. fossilis, dug up). A unit of matter, the smallest portion of an element or a
Fossils of microscopic organisms, only visible when thin compound that retains chemical identity with the

sections of rock are examined. substance in mass; the molecule usually consists of a
union of two or more atoms, some organic molecules
Micrometer (Gr. mikros, small + metron, measure). One
-6
of a meter, or 0.001 millimeter; also called
containing a very large number of atoms.
millionth (10 )

a micron, and abbreviated jum. Molllsol (L. mollis, soft + solum, soil, solid). One of the

Micronutrlent (Gr. mikros, small + L. nutrire, to nourish).

ten world soil orders, characterized by containing more
An essential element required by plants in relatively small
than 1% organic matter in the top 17.5 cm and associated

quantities.
with grassland vegetation; synonymous with chernozem.

Mlcrophyll (Gr. mikros, small + phyllon, leaf). A leaf Monocotyledon (Gr. monos, solitary + kotyledon, a cup-

whose trace is not marked with a gap in the stem's shaped hollow). A plant whose embryo has one cotyledon.

vascular system; microphylls are thought to represent Monoecious monos, solitary + oikos, house).
(Gr.
epidermal outgrowths. Having the reproductive organs in separate structures, but
Micropylar chamber. The space between the micropyle borne on the same individual.
and the nucellus; sealed off from the outside when the Monophyletlc (Gr. mono, single + phyle, tribe). Said of
micropyle closes after pollination. organisms having a common (but sometimes quite
Micropyle (Gr. mikros, small + pulon, orifice, gate). A ancient) ancestor.
pore leading from the outer surface of the ovule between Morphogenesis (Gr. morphe, form + L. genitus, to
the edges of the integuments down to the surface of the produce). The changes in body form that occur during
nucellus.
development of an organism.
Microsporanglate cone. In gymnosperms, a cone that
Morphology (Gr. morphe, form + logos, discourse). The
produces microspores and, ultimately, pollen; synonyms
study of form and its development.
include male cone, pollen cone, and staminate cone.
Moss (L. muscus, moss). A bryophytic plant.
Mlcrosporanglum (plural, mlcrosporangia) (Gr mikros,
iittle + sporangium). A sporangium that bears Multiciliate (L. multus, many + F. cil, an eyelash). Having
microspores. many cilia present on a sperm or spore or other type of
ciliated cell.
Microspore (Gr. mikros, small + spore). A spore which,
in vascular plants, gives rise to a male gametophyte. Multiple fruit. A cluster of matured ovaries produced by
separate flowers; e.g., a pineapple.
Microsporocyte (Gr. mikros, small + spora, seed + L.

cyta, vessel). A diploid cell in which meiosis will occur, Mutation (L. mutare, to change). A sudden, heritable
resulting in four microspores; synonymous with change appearing in an individual as the result of a

microspore mother cell. change in genes or chromosomes.

Microsporophyll (Gr. mikros, little + spore + Gr. Mutualism (L. mutuus, reciprocal). A form of biological
A leaf bearing microsporangia
phyllon, leaf). interaction in which both organisms must associate
Microtubule (Gr. mikros, small + tubule, dim. of tube). A together for continued success of both.
tubule 25 nm in diameter and of indefinite length, Mycelium (Gr. mykes, mushroom). The mass of hyphae
occurring in the cytoplasm of many types of cells. forming the body of the fungus.
Middle lamella (L. lamella, a thin plate or scale). Thin Mycology (Gr. mykes, mushroom + logos, discourse).
layer separating two adjacent protoplasts and remaining as The branch of botany dealing with fungi.
a distinct cementing layer between adjacent cell walls.
Mycorrhiza (Gr. mykos, fungus + riza, root). A symbiotic
Millimeter. The 0.001 part of a meter, equal to 0.0394
association between a fungus and usually the root of a
inch.
higher plant.
Mitochondrion (plural, mitochondria) (Gr. mitos, thread
NAD. Nicotinamide adenine dinucleotide, a coenzyme
+ chondrion, a grain). A membrane-bounded organelle
capable of being reduced.
associated with intracellular respiration.
Mitosis (plural, mitoses) (Gr. mitos, a thread). Nuclear
NADH. Reduced NAD.
division, involving coiling and equal distribution of NADP. Nicotinamide adenine dinucleotide phosphate, a
chromosomes into derivative nuclei; the
duplicate coenzyme capable of being reduced.

chromosome number remains constant. Compare with NADPH. Reduced NADP.

meiosis.
Naked bud. A bud not protected by bud scales.
Mitospore (mitosis + spore). A spore forming after
Nanometer (Gr. nanos, small). One millionth (10~ 6 ) of a
mitosis.
millimeter, equals 10 angstroms; abbreviated nm.
Mixed bud. A bud containing both rudimentary leaves and
flowers. Natural biotic unit. A species defined by isolating barriers
rather than by morphological features; the species of
Molecular biology. A field of biology that emphasizes the
biosystematists.
interaction of biochemistry and genetics in the life of an
organism. Natural classification. A classification scheme that is

appendix D |
Glossary

D14
based on the phylogenetic nature of the organisms bounded organelle that contains most of the DNA in

an artificial classification, which

classified; contrasts with eukaryotic cells.

separates organisms on the basis of convenient traits, but Numerical taxonomy. A field of taxonomy that does not
fails to show the evolutionary relationships among the
place subjective weight on any particular type of evidence
organisms. shows
that relationships between taxa.
Natural selection. The effect of the environment in
Nut (L. nux, nut). A dry, indehiscent, hard, one-seeded
channeling the genetic variation of organisms down
fruit, generally produced from a compound ovary.
particular pathways.
Obligate anaerobe. An organism that cannot live in the
Nectar (Gr. nektar, drink of the gods). A fluid rich in
presence of oxygen.
sugars secreted by nectaries, which are often located near
flowers.
Obligate parasite. An organism that can only live as a
or in
parasite.
Nectar guide. A mark of contrasting color or texture that

may serve to guide pollinators to nectaries within the Obligate saprophyte. An organism that can only live as a
flower. saprophyte.

Nectary (Gr. nektar, the drink of the gods). A nectar- Oogamy (Gr. oion, egg + gamete, spouse). The condition

secreting gland. inwhich the gametes are different in form and activity, i.e.,
sperms and eggs.
Net productivity. The arithmetic difference between
caloriesproduced in photosynthesis and calories lost in Oogonium (L. dim. of Gr. oogonos, literally, a little egg

respiration. layer). Female gametangium of egg-bearing organ not

protected by a jacket of sterile cells, characteristic of the
Net radiation. The arithmetic difference between incoming
thallophytes.
solar radiation and outgoing terrestrial radiation.

Net venation. Veins of leaf blade visible to the unaided

Oospore (Gr. oion, an egg + spore). A resistant spore
developing from a zygote resulting from the fusion of
eye, branching frequently and joining again, forming a
heterogametes.
network.

Neutron (L. neuter, neither). An uncharged particle found Open bundle. A vascular bundle with residual

in the atomic nucleus of all elements except hydrogen. procambium.

Niche (It. nicchia, a recess in a wall). The functional Operculum (L. operculum, a lid). In mosses, cap of

relationship of an organism to its ecosystem. sporangium.

Nitrification (L. nitrum, nitro, a combining form indicating Opposite. Referring to leaf arrangement in which there
the presence of nitrogen + facere, to make). Change of are two leaves opposite each other at a node.
ammonium salts into nitrates through the activites of Order (L. ordo, a row of threads in a loom). A taxonomic
certain bacteria. category below class and above family.
Nitrogen fixation. The process of reducing N 2 gas into Organ (L. organum, an instrument or engine of any kind).
ammonia and incorporating it into the protoplast; A part or member of a plant body adapted for a particular
accomplished only by certain prokaryotes. function.

nm. See nanometer. Organelle. A membrane-bound specialized region within a

n + n. See Dlkaryon. cell such as the mitochondrion or dictyosome.
Node (L. nodus, a knot). Slightly enlarged portion of the Organic. Referring to compounds that contain both
stem where leaves and buds arise, and where branches carbon and hydrogen; also generally referring to the
originate. material products of living organisms.

Nonseptate. Descriptive of hyphae or algal filaments Organic evolution. See Evolution.

lacking crosswalls.
Organism. An individual living body.
Nucellus (L. nucella, a small nut). Tissue composing the
-ose. A chemical suffix indicating a carbohydrate.
young ovule,
chief part of the in which the embryo sac
develops; megasporangium. Osmosis (Gr. osmos, a pushing). Diffusion of a solvent
through a differentially permeable membrane.
Nucleic acid. A polymeric molecule consisting of subunits
called nucleotides, linked together in a chain. Ovary (L. ovum, an egg). Enlarged basal portion of the

pistil, which becomes the fruit.

Nucleolus (L. nucleolus, a small nucleus). Dense
protoplasmic body in the nucleus. Ovulate. Referring to a cone, scale, or other structure
bearing ovules.
Nucleosides. Components of nucleic acids consisting of a
base and a sugar; in DNA, the sugar is deoxyribose, and Ovulate cone. See Megasporangiate cone.
in RNA, ribose; the bases adenine, guanine, and cytosine Ovule (F. ovule, from L. ovulum, dim. of ovum, egg). A
occur in both DNA and RNA; the base thymine occurs in rudimentary seed, containing, before fertilization, the
DNA; the base uracil occurs in RNA. female gametophyte, with egg cell, all being surrounded
Nucleotide. A nucleoside to which a phosphate unit is by the nucellus and one or two integuments.
attached. Ovullferous (ovule + L. ferre, to bear). Referring to a
Nucleus (L. nucleus, kernel of a nut). A membrane- scale or sporophyll bearing ovules.

Glos sary

D15
Oxidation. The removal of electrons or hydrogen, or the Parthenocarpy (Gr. parthenos, virgin + karpos, fruit).

addition of oxygen to a compound. The development of fruit without fertilization.

Oxisol (Gr. oxus, sharp, sour + L. solum, soil, solid). One Parthenogenesis (Gr. parthenos, virgin + genesis,
of the ten world soil orders, characterized by a soil The development of a gamete into a new individual
origin).
horizon at least 30 cm thick that is highly weathered, rich without fertilization.
in clay, but low in nutrients, and red in color, and
associated with tropical vegetation; synonymous with Passive solute absorption. Absorption due only to forces
of simple diffusion.
laterite.

P and P r abbreviations
fr
. for the far-red (FR) or red (R) Pathogen (Gr. pathos, suffering + genesis, beginning).
absorbing forms of phytochrome (P). An organism that causes a disease.
P-protein. Proteinaceous contents of phloem sieve tube Pathology (Gr. pathos, suffering + logos, account). The
members, sometimes called slime. study of diseases, their effects on plants or animals, and
Palea (L. palea, chaff). Upper bract that subtends a grass their treatment.

flower. Pathway. A sequence of chemical reactions, each

Paleoecology (Gr. palaios, ancient). A field of ecology governed by an enzyme, that gradually transform a
that reconstructs past vegetation and climate from fossil starting molecule into some final product.
evidence.
Peat (M.E. pete, of Celtic origin, a piece of turf used as
Paleozoic (Gr. palaios, ancient + zoe, life). A geologic Any mass of semicarbonized vegetable tissue, such
fuel).
era beginning 570 million years ago and ending 225 as Sphagnum, formed by partial decomposition in water.
million years ago.
Pectin (Gr. pektos, congealed). A class of polymers of the
Palisade parenchyma. Elongated cells, containing many cell wall that are built chiefly of partially oxidized sugars.
chloroplasts, found just beneath the upper epidermis of
leaves. Pedicel (L. pediculus, a little foot). Stalk or stem of the
individual flowers of an inflorescence.
Palmately veined (L. palma, palm of the hand).
Descriptive of a leaf blade with several principal veins Peduncle (L. pedunculus, a late form of pediculus, a little
spreading out from the upper end of the petiole. foot). Stalk or stem of a flower that is borne singly; or the

Panicle (L. panicula, a tuft). An inflorescence, the main

main stem of an inflorescence.

axis of which is branched, and whose branches bear Penicillin. An antibiotic derived from the mold Penicillium.
loose racemose flower clusters.
Pentose (Gr. pente, five + OSE). A five-carbon sugar,
Pappus (L pappus, woolly, hairy seed or fruit of certain
C5 H 10 O 5 .

plants). Scales or bristles representing a reduced calyx in

composite flowers. PEP. Phosphoenolpyruvic acid, or phosphoenolpyruvate.

Parallel venation. Type of venation in which veins of a Perennial (L. perennis, lasting the whole year through). A
leaf blade that are clearly visible to the unaided eye are plant that lives from year to year.

parallel to each other.

Perfect flower. A flower having both stamens and pistils.

Paraphysls (plural, paraphyses) (Gr. para, beside +

Perianth (Gr. peri, around, about + anthos, flower). The
physis, growth). A sterile, slender, multicellular hair
petals and sepals taken together.
growing beside fertile cells in certain thallophytes and
bryophytes. Pericarp (Gr. peri + karpos, fruit). Fruit wall, developed
from ovary wall.
Parasite (Gr. parasitos, one who eats at the table of
another). An organism deriving its food from the living Pericllnal cell division. Cell division where the newly
body of another plant or an animal. formed cell wall is parallel to the axis of the organ.

Parenchyma (Gr. parenchein, an ancient Greek medical Pericycle (Gr. peri + kyklos, circle). Tissue, generally of
term meaning to pour beside and expressing the ancient root, bound externally by the endodermis and internally by
concept that the liver and other internal organs were the phloem.
formed by blood diffusing through the blood vessels and
coagulating, thus designating ground tissue). A tissue
Periderm (Gr. peri + Gr. derma, skin). Protective tissue
that replaces the epidermis after secondary growth is
composed of cells that usually have thin walls; site of most
initiated. Consists of cork, cork cambium, and phelloderm.
essential processes such as photosynthesis, secretion,
and storage. Perldium (plural, peridia) (Gr. peridion, a little pouch).
Parent material. The original rock or depositional matter External covering of the hymenium of certain fungi; in

from which the soil of a region has been formed. Myxomycetes, the hardened envelope that covers the
sporangium.
Parietal (F. parietal, attached to the wall, from L. paries,

wall). Belonging connected with, or attached to the

to, Perlgyny (Gr. peri + gyne, a female). A condition in
wall of a hollow organ or structure, especially of the ovary which the receptacle is more or less concave, at the
or cell. margin of which the sepals, petals, and stamens have
Parietal placentation. A type of placentation in which their origin, so that these parts seem to be attached

placentae are on the ovary wall. around the ovary; also called half-inferior.

appendix D J Glossary

D16
Perlsperm. Nutritive tissue in some seeds that torms from Photosynthesis (Gr. photos, light + syn, together +
the nucellus. tithenai, to place). A process inwhich light energy is used
Peristome (Gr. peri + stoma, a mouth). mosses, a
In
to drive the formation of organic compounds.
fringe of teeth about the opening of the sporangium. Phototroplsm (Gr. photos, light + Gr. tropos, a turning).

+ A Influence of light on the direction of plant growth.

Perltheclum (Gr. peri theke, a box). spherical or
flask-shaped ascocarp having a small opening. Phycobilins (Gr. phucos, seaweed). Red and blue
accessory pigments characteristic of blue-green and red
Permafrost (L. permanere, to remain + A.S. freosan, to
algae.
freeze). Soil thatpermanently frozen; usually found
is

some distance below a surface layer that thaws during Phycoblliprotelns. Pigments found in the red and blue-
warm weather. green algae; a phycobilin associated with a protein.

Permeable (L. permeabilis, that which can be penetrated). Phycocyanin (Gr. phykos, seaweed + kyanos, blue). A
Said of a membrane, cell, or cell system through which blue phycobilin pigment occurring in blue-green algae.
substances may diffuse. Phycoerythrln (Gr. phykos, seaweed + erythros, red). A
red phycobilin pigment occurring in red algae.
Peroxysome. An organelle of the microbody class that
contains enzymes capable of destroying peroxides. Phylogenetic classification. See Natural classification.
Petal (Gr. petalon, a flower leaf). One of the flower parts, Phylogeny (Gr. phylon, race or tribe + genesis,
usually conspicuously colored. beginning). The evolution of a group of related individuals.

Petiole (L. petiolus, a little foot or leg). Stalk of leaf. Phylum (Gr. phylon, race or tribe). A primary division of
the animal or plant kingdom.
PGA. 3-Phosphoglyceric acid, a three-carbon compound
formed by the interaction of carbon dioxide and a five- Physiology (Gr. physis, nature + logos, discourse). The
carbon compound, ribulose diphosphate; the reaction science of the functions and activities of living organisms.
yields two molecules of PGA for each molecule of ribulose Phytobenthon (Gr. phyton, a plant + benthos, depths of
diphosphate; PGA is the first stable product of carbon the sea). Attached aquatic plants, collectively.
fixation in the C3 cycle of photosynthesis.
Phytochrome. A pigment found in green plants; it is

Phelloderm (Gr. phellos, cork + derma, skin). A layer of associated with the control of development in response to
cells formed in the stems of some plants from the inner light stimuli.
cells of the cork cambium.
Phytoplankton (Gr. phyton, a plant + planktos,
Phellogen (Gr. phellos, cork + genesis, birth). Cork wandering). Free-floating plants, collectively.
cambium, a cambium giving rise externally to cork and in
Pigment. A substance that absorbs visible light, hence,
some plants internally to phelloderm.
appears colored.
Phenotype (Gr. phaneros, showing + type). The bodily Pileus (L. pileus, a cap). Umbrella-shaped cap of fleshy
characteristics of an organism. fungi.

Phloem (Gr. phloos, bark). Food-conducting tissue, Pinna (plural, pinnae) (L. pinna, a feather). Leaflet or
consisting of sieve tubes members or sieve cells, division of a compound leaf (frond).
companion cells, parenchyma, and fibers.
Pinnately veined (L. pinna, a feather + vena, a vein).
Phosphoenolpyruvate (PEP) carboxylase. The enzyme Descriptive of a leaf blade with single midrib from which
responsible for the fixation of inorganic C0 2 into smaller veins branch off, somewhat like the divisions of a
oxaloacetic acid in a dark reaction of the C4 feather.
photosynthesis cycle.
Pioneer community. The first stage of a succession.
Phosphorylation. A reaction in which phosphate is added
Pistil (L. pistillum, a pestle). Central organ of the flower,
to a compound, e.g., the formation of ATP from ADP and
typically consisting of ovary, style, and stigma.
inorganic phosphate.
Pistillate flower. A flower having pistils but no stamens.
Photon. A quantum of light; the energy of a photon is

proportional to its frequency: E =

hi> where E is energy;
Pit. A thin area of a secondary cell wall.

27
h, Planck's constant, 6.62 x 10~ erg-second; and v is Pith. The parenchymatous tissue occupying the central
the frequency. portion of a stem.

Photoperiod (Gr. photos, light + period). The optimum Placenta (plural, placentae) (L. placenta, a cake). The
length of day or period of daily illumination required for tissue within the ovary to which the ovules are attached.
the normal growth and maturity of a plant. Placentation (L. placenta, a cake + -tion, state of).

Photophosphorylation. A reaction in which light energy Manner in which the placentae are distributed in the
is converted into chemical energy in the form of ATP ovary.

produced from ADP and inorganic phosphate. Plankton (Gr. planktos, wandering). Free-floating aquatic
Photoreceptor (Gr. photos, light + L. receptor, a
plantsand animals, collectively; generally microscopic.
receiver). A light-absorbing molecule involved in Plasma membrane. The membrane that separates the
converting light into some metabolic (chemical energy) living protoplast from the external environment; found in all

form, e.g., chlorophyll and phytochrome. cells.

Glossary

D17
Plasma lemma (Gr. plasma, anything formed + lemma, a Polymer. A molecule that is made by coupling together
husk of a fruit). A synonym for plasma membrane. many small molecules (monomers) that are similar to one
another.
Plasmodesma (plural, plasmodesmata) (Gr. plasma,
something formed + desmos, a bond, a band). Fine Polymerization. The chemical union of monomers to
protoplasmic thread passing through the wall that produce a polymer.
separates two protoplasts. Polynomial (Gr. polys, many + L. nomen, name).
Plasmodium (Gr. plasma, something former + mod. L. Scientific name for an organism composed of more than
odium, something of the nature of). In Myxomycetes, a two words; compare to binomial.
coenocytic mass of protoplasm, with no surrounding wall. Polynucleotides (Gr. polys, much, many). Long-chain
molecules composed of units (monomers) called
Plasmogamy (Gr. plasma, anything molded or formed +
gamos, marriage). The fusion of protoplasts, not nucleotides; nucleic acid is a polynucleotide.
accompanied by nuclear fusion. Polyphyletic (Gr. polys, many + phyle, tribe). Referring
to organisms that did not have an ancestor in common.
Plasmolysis (Gr. plasma, something formed + lysis, a
loosening). The separation of the cytoplasm from the cell Polyploid (Gr. polys, many + ploos, fold). Referring to a

wall due to removal of water from the protoplast. plant or tissue with more than two complete sets of
chromosomes per cell.
Plastid (Gr. plastis, a builder). A class of organelles,
including the chloroplast and several related kinds of Polyribosome (Gr. polys, many + ribosomes). An
bodies; the latter kinds are associated with storage of food aggregation of ribosomes; frequently simply polysome.

materials and some of them (chromoplasts) are highly Polysaccharides (Gr. polys, much, many + sakcharon,
pigmented. sugar). Polymeric molecules composed of units
(monomers) of a sugar; starch and cellulose are
Plastoqulnone. A quinone, one of a group of compounds
polysaccharides.
involved in the transport of electrons during
photosynthesis in chloroplasts. Pome (Gr. pomme, apple). A simple fleshy fruit, the outer
portion of which is formed by the floral parts that surround
Plumule (L. plumula, a small feather). The first bud of an
the ovary.
embryo or that portion of the young shoot above the
cotyledons. Population (L. populus, people). A group of closely
related, interbreeding organisms.
Podzol (R. pod, under + zola, ashes). A soil
Pore spaces. Spaces between soil particles that may be
characteristic of taiga vegetation; color of the A horizon is
with air or water and into which root hairs may
filled
gray because of excessive leaching; in the spodosol soil
penetrate; the larger the soil particles, the larger the pore
order.
space.
Polar transport. The directed movement within plants of
Prairie (L. pratum, meadow). Grassland vegetation, with
compounds hormones) predominantly in one
(usually
trees essentially absent; often considered to have more
direction; polar transport overcomes the tendency for
rainfall than does the steppe.
diffusion in all directions.
Predation (L. predatio, plundering). A form of biological
Polarity (Gr. pol, an axis). The observed differentiation of
interaction in which one organism is destroyed (by
an organism, tissue, or cell into parts having opposed or
ingestion); parasitism is a form of predation.
contrasted properties or form.
Primary (L. primus, first). First in order of time or
Pollen (L. pollen, fine flour). The germinated microspores development.
or partially developed male gametophytes of seed plants.
Primary pltfield. Thin areas of primary cell walls.
Pollen mother cell. See Microsporocyte.
Primary endosperm cell. A cell of the embryo sac after
Pollen profile. A
diagrammatic summary of the sequence fertilization, generally containing a nucleus resulting from
and abundance of pollen types that have been fusion of the two polar nuclei with a sperm nucleus; the
chronologically trapped in sediments. endosperm develops from this cell.

Pollen tube. The parastic, complete male gametophyte of Primary meristems. Meristems of the shoot or root tip

seed-plants, which grows through the nucellus of a giving rise to the tissues of the primary plant body.
gymnosperm ovule or through the pistil of an angiosperm. Primary tissues. Those tissues, epidermis, xylem, phloem,
Pollination.The transfer of pollen from a stamen or and ground tissues, which form from primary meristems.
staminate cone to a stigma or ovulate cone. Primitive (L. primus, first). Referring to a taxonomic trait

thought to have evolved early in time.

Pollinium (L. pollentis, powerful or pollinis, fine flour +
ium, group). A mass of pollen that sticks together and is Primordlum (L. primus, first + ordiri, to begin to weave;
transported by pollinators as a mass; present in orchids literally beginning to weave, or to put things in order). The
and milkweeds. beginning or origin of any part of an organ.

Pollutant (L. polluere, to dirty + lutum, mud). An Procamblum (L. pro, before + cambium). A primary
unnatural, human-related substance that is introduced to meristem that gives rise to primary vascular tissues and, in

the environment; may be gas, liquid, or solid, easily most woody plants, to the vascular cambium.
broken down or long-lasting. Producer (L. producere, to draw forward). An organism

appendix p |
Glossary

D18
thatproduces organic matter for itself and other organisms A denser body occurring within the chloroplasts of certain
(consumers and decomposers) by photosynthesis. algae and liverworts and apparently associated with starch
deposition.
Proembryo (L. pro, before + embryon, embryo). A group
of cells arising from the division of the egg
fertilized cell Pyrlmldlnes. A group of compounds in which carbon and
before the cells that are to become the embryo are nitrogen atoms form a ring structure; includes the
recognizable. compounds cytosine, thymine, and uracil.

Prokaryotes (L pro, before + Gr. karyon, a nut, referring Quadrat (L. quadrus, a square). A frame of any shape
in modern biology to the nucleus). Primitive organisms, that, when placed over vegetation, defines a unit sample
bacteria, and blue-green algae, which do not have the area within which the plants may be counted or measured.
DNA separated from the cytoplasm by an envelope. Quantum (L. quantum, how much). An elemental unit of

Prophase (Gr. pro, before + phasis, appearance). An energy.

early stage in nuclear division, characterized by coiling of Quiescent center (L. quiescere, to rest). Disk-shaped
the chromosomes and formation of the mitotic spindle. region of root apex containing slowly dividing cells.

Proplastld (Gr. pro, before + plastid). A type of plastid, r selection. Natural selection that favors short-lived, early-
occurring generally in meristematic cells, which will maturing individuals which devote a large fraction of their
develop into a chloroplast. resources into reproduction; annual herbs are r strategists.
Protease (protein + -ase, a suffix indicating an enzyme). Raceme racemus, a bunch of grapes). An
(L.
An enzyme breaking down a protein. inflorescence in which the main axis is elongated and the

flowers are born on pedicels that are about equal in

Protein (Gr. proteios, holding first place). A polymeric
length.
molecule made of subunits called amino acids which are
linked together in a chain by a type of bond called the Rachllla (Gr. rhachis, a backbone + L. dim, ending -ilia).

peptide bond. Shortened axis of spikelet.

Proterozoic (Gr. protero, before in time + zoe life). The Rachls (Gr. rhachis, a backbone). Main axis of spike; axis
earliest geologic era, beginning about 4.5-5 billion years of fern leaf (frond) from which pinnae arise; in compound
ago and ending 570 million years ago; also called the leaves, the extension of the petiole corresponding to the
Precambrian era. midrib of an entire leaf.

Protochlorophyll (Gr. protos, first + chloros, green + Radicle (L. radix, root). Portion of the plant embryo that
phyllos, leaf). One of the precursors of chlorophyll. develops into the primary or seed root.

Prothallus (L. pro, before + Gr. thallos, young shoot, Random plant distribution. A distribution of a plant

sprout). A synonym for the gametophyte generation of species within an area such that the probability of finding
ferns; also called prothallium; the prothallial cells of some an individual at one point is the same for all points.

male gametophytes are sterile and thought to represent a Raphe (Gr. rhaphe, seam). Ridge on seeds, formed by the
much reduced vegetative body. stalk of the ovule, in those seeds inwhich the funiculus is
Protoxylem. First formed primary xylem. sharply bent at the base of the ovule.

Protoderm (Gr. protos, first + derma, skin). A primary Raphides (Gr. rhaphis, a needle). Fine, sharp, needlelike

meristem that gives crystals.

rise to epidermis.

Proton (Gr. proton, first). The nucleus of a hydrogen atom Ray initials. Meristematic cells in the vascular cambium
is a single positively charged particle, the proton; the
that develop into xylem and phloem cells comprising the
ray system.
nucleus of other elements consists of protons and
all
-24
neutrons; the mass of a proton is 1.67 x 10 gram. Ray system (L. radius, a beam or ray). System of cells in
secondary tissues that are oriented perpendicular to the
Protonema (plural, protonemata) (Gr. protos, first +
long axis of the stem, form from ray initials of the vascular
nema, a thread). An algal-like filamentous growth; an early
cambium.
stage in development of the gametophyte of mosses.
Receptacle (L. receptaculum, a reservoir). Enlarged end
Protoplasm (Gr. protos, first + plasma, something
of the pedicel or peduncle to which other flower parts are
formed). Living substance.
attached.
Protoplast (Gr. protoplastos, formed first). The organized
Recombination (L. re, repeatedly + combinatus, joined).
living unit of a single cell.
The mixing of genotypes that results from sexual
Pseudopodlum (Gr. pseudes, false + podion, a foot). In reproduction.
Myxomycetes, an armlike projection from the body by
Red tide. The coloring of nearshore, marine water by a
which the plant creeps over the surface.
high density of mircroscopic algal organisms that may
Purines. A
group of compounds in which carbon and additionally release toxic byproducts into the water.
nitrogen atoms form a double ring structure, one ring with
Reduction (F. reduction, L. reductio, a bringing back).
six atoms, the other with five atoms; includes the
Any chemical reaction involving the removal of oxygen
compounds adenine and guanine.
from or the addition of hydrogen or an electron to a
Pyrenold (Gr. pyren, the stone of a fruit + L. o'ides, like). substance.

Glossary

D19
Regular flower. A flower in which the corolla is made up Samara (L. samara, the fruit of the elm). Simple, dry, one-
shaped pefals equally spaced and
of similarly radiating or two-seeded indehiscent fruit with pericarp bearing a
from the center of the flower. winglike outgrowth.

Reproduction (L. re, repeatedly

producere, to give + Sand. Soil particles between 50 and 2000 microns in

birth to). The process by which plants and animals give diameter.
rise to offspring.
Saprophyte (Gr. sapros, rotten + phyton, a plant). Ar.
Reproductive isolation. The separation of populations in organism deriving its food from the dead body or the
time or space so that genetic flow between them is cut off. nonliving products of another plant or animal.

Residual meristem. Meristematic region near the tip of Sapwood. Peripheral wood that actively transports.
the shoot apex that remains after differentiation of the pith Savannah (Sp. sabana, a large plain). Vegetation of
and cortex. scattered trees in a grassland matrix.
Resin duct. Resin canal; in conifers, continuous tubes Shizocarp (Gr. schizein, to split + karpos, fruit). Dry fruit
lined with secretory cells that run through the sapwood; with two or more united carpels that split apart at maturity.
they function as repositories for metabolic byproducts, but
Sclereids (Gr. skleros, hard). Sclerenchyma cells having
may have an ecological use as deterrents to wood-boring
variably shaped, heavily lignified cell walls.
insects and they have economic value as a source of
turpentine and other naval stores. Sclerenchyma (Gr. skleros, hard + -echyma, a suffix
denoting tissue). A strengthening tissue composed of cells
Respiration (L. re, repeatedly, + spirare, to breathe). In
with heavily lignified cell walls.
the cell, the catabolic process by which sugars and other
and broken down, with some
fuels are oxidized of the
Scrub (A.S. scrob, a shrub). Vegetation dominated by
energy captured in the formation of ATP. shrubs; described as thorn forest in areas with moderate
rainfall, or as chaparral or desert in areas with low rainfall.
Rhizoid (Gr. rhiza, root + L. o'i'des, like). One of the
Scutellum (L. scutella, a dim. of scutum, shield). Single
cellular filaments that perform the functions of roots.
cotyledon of grass embryo.
Rhizome (Gr. rhiza, root). An elongated, underground,
Seaweed. Any large, marine alga; usually a red or brown
horizontal stem.
alga; includes the kelps.
Rhizophores (Gr. rhiza, root + phoros, bearing). Leafless
Secondary tissues. Those tissues, xylem, phloem, and
branches that grow downward from the leafy stems of
periderm, that form from secondary meristems.
certain Lycophyta and give rise to roots when they come
into contact with the soil. Secretory structures. Any of a number of specialized
such as nectaries, glands, and
plant structures
Ribose. A pentose sugar.
hydathodes that secrete secondary plant substances.
Ribosomes (ribo, from RNA + Gr. somatos, body). Small
Rock formed from
Sedimentary rock (L. sedere, to sit).
particles 10-20 nm in diameter, containing RNA and
material deposited as sediment, then physically or
protein; active in protein synthesis.
chemically changed by compaction and hardening while
Ring porous. Wood with large xylem vessel members buried in the earth crust.
mostly in early wood; compare with diffuse porous. which may be sown). Popularly
Seed (A.S. sed, anything
Ripening (A.S.perhaps related to reap). Changes in a
rifi, as originally used, anything that may be sown; i.e., "seed"
fruit that follow seed maturation and that prepare the fruit potatoes, "seeds" of corn, sunflower, etc.; botanically, a
for its function of seed dispersal. seed is the matured ovule without accessory parts.

RNA. Ribonucleic acid. Seed coat. A hardened, outer layer of the seed, derived
from the integument(s) of the ovule, and which functions
Root (A.S. rot). The descending axis of a plant, normally
to prevent mechanical disturbance to and water loss from
below ground, serving to anchor the plant and absorb and
conduct water and mineral nutrients.
the embryo; it may also regulate germination in several
ways.
Root cap. A thimblelike mass of living cells covering and
Seed plant. Common name for members of the
protecting the apical meristems of a root; site of
gymnosperm and angiosperm groups, and for extinct
perception of gravity in geotropism.
members of other groups that produced seeds.
Root hairs. Epidermal projections of root cells in region of
Selectively permeable. Referring to a membrane that
maturation, provide means to increase the absorptive
permits some kinds of molecules to pass through while not
surface of root.
allowing other kinds of molecules to pass through.
Root pressure. Pressure in the xylem arising as a result
Self-pollination. Transfer of pollen from the stamens to
of osmosis in the root.
the stigma of either the same flower or flowers on the
Rootstock. An elongated, underground, horizontal stem. same plant.

Rosette. A shoot with a very short stem, composed of Seminal root. The root or roots forming from primordia
several unelongated internodes with fully expanded leaves. present in the seed.

Runner. A stem that grows horizontally along the ground Sepals (ML. sepalum, a covering). Whorl of sterile, leaf-
surface. like structures that usually enclose the other flower parts.

appendix p |
Glossary

D20
Septate (L. septum, fence). Divided by crosswalls into Soil (L. solum, The uppermost stratum of the
soil, solid).

cells or compartments. earth's crust, which has been modified by weathering and
Septum (L. septum, fence). Any dividing wall or partition; organic activity into (typically) three horizons: an upper A
frequently a crosswall in a fungal or algal filament. horizon that is leached, a middle B horizon in which the
leached material accumulates, and a lower C horizon that
Serpentine (L. serpens, a serpent). Referring to soil
is unweathered parent material.
derived from metamorphic parent material characterized
(among other things) by low calcium (Ca), high Soil texture. Refers to the amounts of sand, silt, and clay
magnesium (Mg), and a greenish-gray color. in a soil, as a sandy loam, loam, or clay texture.

Sessile (L. sessilis, low, dwarf, from sedere, to sit). Sitting, Solute (L. solutus, from solvere, to loosen). A dissolved
referring to a leaf lacking a petiole or a flower or fruit substance.
lacking a pedicel. Solution. A homogeneous mixture, the molecules of the
Seta (plural, setae) (L. seta, a bristle). In bryophytes, a dissolved substance (e.g., sugar), the solute, being
short stalk of the sporophyte, which connects the foot and dispersed between the molecules of the solvent (e.g.,
the capsule. water).

Sexual reproduction. Reproduction that requires meiosis Solvent. A substance, usually a liquid, having the
and fertilization for a complete life cycle. properties of dissolving other substances.
Shade tolerance. The ability to grow slowly in the shade Soredium (plural, soredia) (Gr. soros, a heap). A sexual
of an overstory canopy; an essential characteristic for a reproductive body of lichens, consisting of a few algal
climax species. cells surrounded by fungal hyphae.
Sheath. Part of leaf that wraps around the stem, as in
Sorus (plural, sori) (Gr. soros, a heap). A cluster of
grasses. sporangia in ferns.

Shoot (derivation uncertain, but early referring to new Species (L. species, appearance, form, kind). A group of
plant growth; 1450, "Take a feyr schoyt of blake thorne"). individuals usually interbreeding freely and having many
A young branch that grows out from the main stock of a characteristics in common.
tree, or the young main portion of a plant growing above

ground.
Sperm (Gr. sperma, the generative substance or seed of a
male animal). A male gamete.
Shoot tip. Terminal portion of the shoot containing apical
and primary meristems and cells in early stages of Spermagonium (plural, spermagonla) (Gr. sperma,
differentiation.
sperm + gonos, offspring). Flask-shaped structure
characteristic of the sexual phase of the rust fungi;
Sibling species. Species morphologically nearly identical
bearing receptive hyphae and spermatia.
but incapable of producing fertile hybrids.

Side-chain. A Spermatophyte (Gr. sperma, seed + phyton, plant). A

part of a polymer that extends laterally from
seed plant.
the main chain.
Spike (L. spica, an ear of grain). An inflorescence in
Sieve cell. A long and slender sieve element without a
which the main axis is elongated and the flowers are
companion cell, with relatively unspecialized sieve areas,
sessile.
and with tapering end walls that lack sieve plates; found in
gymnosperms and ferns. Splkelet (L. spica, an ear of grain + dim. ending -let).

Sieve plate. Perforated wall area in a sieve-tube member The unit of inflorescence in grasses; a small group of
through which pass strands connecting sieve-tube grass flowers.

protoplasts. Spindle (A.S. spinel, and instrument employed in spinning

Sieve tube. A series of sieve-tube members forming a thread by hand). Referring in mitosis and meiosis to the
long cellular tube specialized for the conduction of food spindle-shaped intracellular aggregate of microtubules
materials. involved in chromosome movement.

Sieve tube members. An enucleate phloem cell primarily Spodosol (Gr. spodos, wood ashes; R. pod, under +
responsible for photosynthate transport; separated from zola, ashes). One of the ten world soil orders,
other sieve tube members by sieve plates. characterized by an ashy, sandy, bleached, acidic A2
horizon and associated mainly with coniferous forest
Sillque (L. siliqua, pod). The fruit characteristic of
Brassicaceae (mustards); two-celled, the valves splitting vegetation; synonomous with podzol.

from the bottom and leaving the placentae with a partition Sporangiophore (sporangium + Gr. -phore, a root of
stretched between. phorein, to bear). A branch bearing one or more
Silt. Soil particle between 2 and 50 jum in diameter. sporangia.

Simple pit. Pit not surrounded by an overarching border; Sporangium (spore + Gr. angeion, a vessel). Spore
in contrast to bordered pit.
case.

Single bond. A covalent bond that involves the sharing of Spore (Gr. spora, seed). A reproductive cell that develops
two electrons. into a plant without union with other cells.

Siphonous line. A line of evolutionary development in the Sporlc life cycle. A life cycle that includes alternation of
algae in which mitosis is not followed by cytokinesis; this generations, the sporophyte and gametophyte being more
results in an elongated multinucleate, coenocytic filament. than zygote and gametes, respectively.

Glossary

D21
Sporocyte (spore + L. cyta, vessel). A diploid or haploid Substrate. A molecule that engages in a reaction
undergo mitosis or meiosis
cell that will to produce spores. catalyzed by ar) enzyme.
Sporophore (spore + Gr. phorein, to bear). The truiting Succession (L. successio, a coming into the place of
body of fleshy and woody fungi, which produces spores. another). A sequence of changes in time of the species
Sporophyll (spore + Gr. phyllon, leaf). A spore-bearing that inhabit an area, from an initial pioneer community to a

leaf.
final climax community.

Sporophyte (spore 4- Gr. phyton, a plant). In alternation Succulent. A plant with fleshy, water-storing parts.

which meiosis occurs and

of generations, the plant in Sucrose. Table sugar, a disaccharide made of a molecule
which thus produces meiospores. ofglucose linked to a molecule of fructose.

Spring wood. See Early wood. A simple carbohydrate such

Sugar. as glucose.

Stamen (L. stamen, the standing-up things or a tuft of Summer wood. See Late wood.
thready things). Flower structure made up of an anther Superior ovary. An ovary completely separate and free
(pollen-bearing portion) and a filament. from the calyx.
Staminate cone. See Microsporangiate cone. Suspensor (L. suspendere, to hang). A cell or chain of

Stamlnate flower. A flower having stamens but no pistils. cells developed from a zygote whose function is to place
the embryo cells in an advantageous position to receive
Starch (M.E. sterchen, to stiffen). A polysaccharide
food.
composed of glucose; the chief food storage material of
many plants. Suture (L. sutura, a sewing together; originally the sewing
together of flesh or bone wounds). The junction, or line of
Statolith (Gr. statos, standing + lithos, stone) An
junction, of contiguous parts.
organelle that moves to its position in a cell as a result of
gravity, thus providing an initial sensing of, or orientation Symbiosis (Gr. syn, with + bios, life). An association of
to, gravity by a cell. two different kinds of living organisms.

Stele (Gr. stele, a post). The central cylinder, inside the Sympetaly (Gr. syn, with + petalon, leaf). A condition in

cortex, of roots and stems of vascular plants. which petals are united.

Stem (O.E. stemn). The main body of the portion above Synandry (Gr. syn, with + andros, a man). A condition in

ground of tree, shrub, herb, or other plant; the ascending which stamens are united.

axis, whether above or below ground, of a plant, in Syncarpy (Gr. syn, with + karpos, fruit). A condition in

contradistinction to the descending axis or root. which carpels are united.

Steppe (R. step, a lowland). An arid grassland vegetation. Synergids (Gr. synergos, toiling together). The two nuclei

Sterlgma (plural, sterigmata) (Gr. sterigma, a prop). A at one end of the embryo sac, which, with the third (the

slender, pointed protuberance at the end of a basidium, egg), constitute the egg appartus.

which bears a basidiospore. Synsepaly (Gr. syn, with + sepals). A condition in which
Stigma (L. stigma, a prick, a spot, a mark). Receptive
sepals are united.

portion of the style to which pollen adheres. Taiga (Teleut taiga, rocky mountainous terrain). A broad
The stem portion of a
northern belt of vegetation dominated by conifers; also, a
Stipe (L. stipes, post, tree trunk).

kelp, to which are attached bladders and blades. similar belt in mountains just below alpine vegetation.

Stipule (L. stipula,dim of stipes, a stock or trunk). A Tannin. A substance that has an astringent, bitter taste.

leaflike structure from either side of the leaf base. Tapetum (I. tapete, carpet). The tissue that lines
developing pollen sacs (microsporangia) of seed plants; it
Stolon (L. stolo, a shoot). A stem that grows horizontally
degenerates and provides nutrition to the tissue within.
along the ground surface.

Stoma Taxon (plural, taxa) (Gr. taxis, order). A general term for
stomata) (Gr. stoma, mouth). Epidermal
(plural,
any taxonomic rank, from subspecific to divisional.
structure on stems and leaves composed of two guard
cells plus the small pore between them, through which Taxonomy (Gr. taxis, arrangement + nomos, law).

gases pass. Systematic botany; the science dealing with the

describing, naming, and classifying of plants
Strobilus (Gr. strobilos, a cone). A number of modified,
spore-producing scales or leaves (sporophylls) grouped TCA cycle. See Citric acid cycle.
together on an axis. Teliospore (Gr. telos, completion + spore). Resistant
Stroma (Gr. stroma, a bed or covering). A mass of spore characteristic of the Heterobasidiomycetidae, in
protecting vegetative filaments; the background substance which karyogamy and meiosis occur and from which a
of chloroplasts, probably the location of the carbon cycle basidium develops.
of photosynthesis. Tellum (plural, telia) (Gr. telos, completion). A sorus of

Style (Gr. stylos, a column). Slender column of tissue that teliospores.

arises from the top of the ovary and through which the Telophase (Gr. telos, completion + phase). The last

pollen tube grows. stage of mitosis, in which daughter nuclei are reorganized.
Suberln (L. suber, the cork oak). A waxy material found in Tendril (L. tendere, to stretch out, to extend). A slender
cell walls of cork tissue and endodermis. coiling organ that aids in the support of stems.

appendix p |
Glossary

D22
Terminal bud. A bud at the end of a stem. Transpiration (F. transpirer, to perspire). The giving off of
water vapor from the surface of leaves
Testa (L. testa, brick, shell). The outer coat of the seed.
Trichogyne (Gr. trichos, a hair + gyne, female)
Tetrad (Gr. tetradeion, a set of four). A group of four,
Receptive hairlike extension of the female gametangium in
usually referring to the meiospores immediately after
the Rhodophyta and Ascomycetes.
meiosis.

Tetraplold (Gr. tetra, four + ploos, fold). Having four sets

Trlchome (Gr. trichoma, a growth of hair). A short
filament of cells.
of chromosomes per nucleus.
Trlose (Gr. treis, three + -ose, suffix indicating a
Tetraspores (Gr. tetra, four + spores). Four spores
carbohydrate). Any three-carbon sugar.
formed by division of the sporocyte.
+ sporos, seed, spore
Tritium.A hydrogen atom, the nucleus of which contains
Tetrasporophyte (Gr. tetra, four 3
one proton and two neutrons; is written as H; the moreit
+ phuton, plant). One
two sporophyte generations in
of
common hydrogen nucleus consists only of a proton.
certain red algae; produces meiospores in tetrad clusters.
Tropical rain forest. Vegetation with several tree strata,
Tetrasporlne line (tetraspore + L. suffix -ine, like). A line
characteristic of tropical lowland regions.
of evolutionary development in the algae in which mitosis
is directly followed by cytokinesis, resulting in a filament, Tropism (Gr. trope, a turning). An orientation of the

thallus, or complex plant body of varied form. direction of growth in an organ, guided by an external
stimulus such as light or gravity.
Thallophytes (Gr. thallos, a sprout + phyton, plant). A
division of plants whose body is a thallus, i.e., lacking Tuber (L. tuber, a bump, swelling). A much-enlarged,
roots, stems, and leaves. short, fleshy underground stem tip.

Thallus (Gr. thallos, a sprout). Plant body without true Tundra (Lapp, tundar, hill). Meadowlike vegetation at low
roots, stems, or leaves. elevation in cold regions that do not experience a single
month with average daily maximum temperatures above
Thermoperlod (Gr. therme, heat + periods, a cycle). A 50 °F.
difference in temperature between day and night.
Turgid (L. trugidus, swollen, inflated). Swollen, distended;
Thymidine. A nucleoside incorporated in DNA, but not in
referring to a cell that is firm due to water uptake.
RNA.
3
Turgor pressure (L. turgor, a swelling). The pressure
Thymidine H. Tritiated or radioactive thymidine.
within the cell resulting from the absorption of water into
Thymine. A pyrimidine occurring in DNA, but not in RNA. the vacuole and the imbibition of water by the protoplasm.

Tiller (O.E. telga, a branch). A grass stem arising from a Tylosis (plural, tyloses) (Gr. tylos, a lump or knot). A
lateral bud at a basal node; tillering is the process of tiller growth of one cell into the cavity of another.
formation.
Type specimen. The herbarium specimen selected by a
Tissue. A group of cells that perform a collective function. taxonomist to serve as a basis for the naming and
descriptions of a new species.
Tonoplast (Gr. tonos, stretching tension + plastos,
molded, formed). The cytoplasmic membrane bordering Ultisol. A modified podsol soil, with red or yellow B
the vacuole; so-called by de Vries, as he thought it horizon, representative of the southern deciduous forest.
regulated the pressure exerted by the cell sap. Umbel (L. umbella, a sunshade). An inflorescence, the
Toxin (L. toxicum, poison). A poisonous secretion of a individual pedicels of which all arise from the apex of the

plant or animal. peduncle.

Tracheid (Gr. tracheia, windpipe). An elongated, tapering Unavailable water. Water held by the soil so strongly that
xylem cell, with lignified pitted walls, adapted for root hairs cannot readily absorb it.

conduction and support. Unicell (L. unus, one + cell). An organism consisting of a

Tracheophytes (Gr. tracheia, windpipe + phyton, plant). single cell; generally used in describing algae.
Vascular plants. Uniseriate (L. unus, one + M.L. seriatus, to arrange in a

Trait. A distinctive definable characteristic; a mark of series). Said of a filament having a single row of cells.

individuality. Uracil. A pyrimidine found in RNA but not in DNA.

Transfer cells. Specialized cells modified by their cell wall Uredium (plural, uredla) (L. uredo, a blight). A sorus of
projections for efficient short distance transport. uredospores.

Transfusion tissue (L. trans, across + fundere, to pour, Uredospore (L. uredo, a blight + spore). A red, one-
to melt). In pine needles, the tissue surrounding the celled summer spore in the life cycle of the rust fungi.
which may serve to transfer water, nutrients,
central veins,
A
Vacuole (L. dim. of vacuus, empty). watery solution of
and food between the vascular tissue and the mesophyll.
various substances forming a portion of the protoplast
Translocation (L. trans, across + locare, to place). The distinct from the protoplasm.
transfer of food materials or products of metabolism.
Vascular (L. vasculum, a small vessel). Referring to any
Transmit. To pass or convey something from one person, plant tissue or region consisting of or giving rise to
organism, or place to another person, organism, or place. conducting tissue, e.g., bundle, cambium, ray.

Glossary

D23
Vascular bundle. A strand of tissue containing primary water molecules in any other system; the activity of these
xylem and primary phloem (and procambium present) it water molecules may be greater (positive) or less
and sometimes enclosed by a bundle sheath of (negative) than the activity of the water molecules under
parenchyma or fibers. standard conditions.
Vascular cambium. Cambium giving rise to secondary
Weathering. Physical and chemical change in parent
phloem and secondary xylem. material that leads to soil formation.
Vascular plant (L. vasculum, small vessel). The common
name for any plant that has xylem and phloem; includes Weed (A.S. weod, used at least since 888 in its present
meaning). Generally a herbaceous plant or shrub not
the higher and lower vascular plants but not the kelps
(which possess sieve tubelike cells) or the bryophytes
valued for use or beauty, growing where unwanted, and

(some of which contain cells resembling tracheids and regarded as using ground or hindering the growth of more
desirable plants.
sieve tubes).

Vegetation (L. vegetare, to quicken). The plant cover that Whorl. A circle of three or more flower parts, or of leaves.
clothes a region; it is formed of the species that make up
Whorled. Referring to bud or leaf arrangement in which
the flora, but is characterized by the abundance and life
there are three or more buds or three or more leaves at a
form (tree, shrub, herb, evergreen, deciduous plant, etc.)
node.
of certain of them.
Venation (L. vena, a vein). Arrangement of veins in leaf Wings. Lateral petals of legume type of flower.

blade.
Wood (M.E., wode, wude, a tree). Secondary xylem.
Venter (L. venter, the belly). Enlarged basal portion of an
Xanthophyll (Gr. xanthos, yellowish brown + phyllon,
archegonium in which the egg cell is borne.
leaf). A yellow chloroplast pigment.
Ventral canal cell. The cell just above the egg cell in the
archegonium. Xerophyte (Gr. xeros, dry + phuton, plant). A plant that
normally grows in dry habitats.
Ventral suture (L. ventralis, pertaining to the belly). The
line of union of the two edges of a carpel. Xylem (Gr. xylon, wood). A plant tissue consisting of

Vernalization (L. vernalis, belonging to spring + izare, to tracheids, vessel elements, parenchyma cells, and fibers;

make). The promotion of flowering by naturally or wood.

artificially applied periods of extended low temperature;
Zoosporangium (Gr. zoon, an animal + sporangium). A
seeds, bulbs, or entire plants may be so treated.
sporangium bearing zoospores.
Vessel (L. vasculum, a small vessel). A series of xylem
elements whose function it is to conduct water and Zoospore (Gr. zoon, an animal + spore). A motile spore.

mineral nutrients. zygon a yoke + morphe, form).

Zygomorphic (Gr.
Vessel element. A xylem cell derived from the vascular Referring to bilateral symmetry; said of organisms, or a
cambium or procambium; a portion of a vessel. flower, capable of being divided into two symmetrical
Vitamins (L. vita, life + amine). Naturally occurring halves only by a single longitudinal plane passing through
organic substances, akin to enzymes, necessary in small the axis.

amounts for the normal metabolism of plants and animals. Zygospore (Gr. zygon, a yoke + spore). A thick-walled
Volvocine line (Volvox + L. suffix -ine, like). A line of resistant spore developing from a zygote resulting from the
evolutionary development in the algae in which the cells fusion of isogametes.
remain separate, as in Volvox, and are never connected to
form a filament or flattened thallus.
Zygote (Gr. zygon, a yoke). A protoplast resulting from
the fusion of gametes.
Water potential. Refers to the difference between the
water molecules in pure distilled water at
activity of Zygotic life cycle. A life cycle in which the diploid phase
standard temperature and pressure, and the activity of is represented only by the zygote.

appendix D |
Glossary

D24
 E APPENDIX

illustration credits

Noncredited figures are either original to this book

or are used with permission from T. E. Weier et ai,
Botany: An Introduction to Plant Science, 5th Ed.,
Wiley and Sons, New York. Figure numbers are
given in boldface type.

Chapter 1: Introduction 4.11 Courtesy of A. S. Crafts.

1.1 Photo courtesy of R. M. Liebaert. 4.12 A, courtesy of J. Cronshaw and K. Esau, J. Cell Biol. 38,
1.2 Photo courtesy of Dr. N. J. Lang. 25; B, as above, p. 292.
1.3 Photo courtesy of Dr. N. J. Lang. 4.13 A through D redrawn from Carl C. Forsaith, The
1.4 Photo courtesy of D. Dreyfus. Technology ofNew York State Timbers, New York State
1.5 Photo courtesy of R. M. Liebaert. College of Forestry Publication 18.
1.6 Photo courtesy of R. M. Liebaert. 4.14 6, courtesy of L. M. Srivastava.
1.7 Photo courtesy of D. Brown. 4.16 After McArthur.
4.23 Slides courtesy of Triarch Products.
Chapter 2: Metabolism 4.25 Slides courtesy of D. Graham; stereoscan courtesy of D.
2.6
'

From Scientific American, April 1972. Copyright 1972 by Hess.

Richard E. Dickerson and Irving Geis.
4.30 A, courtesy of Richard Parker, National Audubon Society;
2.8 From V. A. Greulach and J. E. Adams, Plants: An B, courtesy of Diamon T. Smithers, National Audubon
Introduction to Modern Botany, 3rd Ed., John Wiley & Society.
Sons, Inc., 1976. Used with permission. New
4.31 From K. Esau, Plant Anatomy, 2nd Ed., Wiley, York.
4.42 Courtesy of G. Breckon.
Chapter 3: The Plant Cell
4.43 A, from R. M. Holman and W. W. Robbins, A Textbook of
3.1 From Hooke, Micrographia (1664). The Council of the
General Botany, Wiley, New York; B, courtesy of C. H. Lin;
Royal Society of London for Improving Natural Knowledge.
C, courtesy of W. W. Thomson.
3.4 Courtesy of Prof. Harry T. Horner, Jr.
4.44 A and B, courtesy of S. Lynch.
3.5 Courtesy of Prof. James P. Braselton.
4.48 Redrawn after G. S. Avery, Amer. J. Bot. 20, 565.
3.6 Courtesy of Prof. Harry T. Horner, Jr.
4.50 A and B, courtesy of Tomato Genetics Cooperative; E,
3.7 Courtesy of Prof. Harry T. Horner, Jr
drawing by Peggy-Ann Kessler Duke in V. A. Greulach,
3.9 Courtesy of Prof. Chin Ho Lin.
Plant Function and Structure, Macmillan, New York.
3.10 Courtesy of J. B. Pantastico.
4.51 D, courtesy of W. Russell; E, courtesy of P. Jones.
3.11 Courtesy of Prof. Harry T. Horner, Jr.
4.53 From H. J. Fuller and O. Tippo, College Botany, Holt,
3.12 Courtesy of Prof. Harry T. Horner, Jr.
Rinehart, and Winston, New York.
3.15 Courtesy of E. G. Cutter.
4.55 E, from L. Kutschera, Wurzelatlas, Geschaftsfuhrer des
3.17 Courtesy of Prof. Harry T. Horner, Jr.

3.18 A and B from Newcomb, Protoplasma 84, 3,

DLG-Verlages, Frankfurt.
E. H. 1 1

Copyright Springer-Verlag. 4.56 From Rogers and Head in W. J. Whittington (Ed.), Root

3.19 Courtesy of D. Branton. Growth, Plenum Press, New York. Also courtesy of East

3.25 Chromosoma 25, 249. Copyright Mailing Research Station.

Courtesy of A. Bajer,
Springer-Verlag. 4.58 Drawn by Sue MacLeod.
3.27 Courtesy of A. Bajer. 4.62 A through D, after K. Esau.
3.28 Courtesy of P. K. Hepler and W. T. Jackson, J. Cell Biol.
38, 437.
3.30 Courtesy of M. C. Ledbetter. Chapter 5: The Absorption and Transport Systems
5.9 Photo courtesy of E. L. Proebsting.
3.31 B, courtesy of Gankin.
5.10 Photo courtesy of D. R. Hoagland.

Chapter 4: The Plant Body 5.11 Photo courtesy of D. R. Hoagland.

4.1 C, courtesy of E. B. Risley. 5.12 Redrawn from McDougall, Plant Ecology, 4th Ed., Lea and
4.9 A and B redrawn from Carl C. Forsaith. The Technology of Febiger, 1949, in V. A. Greulach and J. E. Adams, Plants:
New York State Timbers, New York State College of An Introduction to Modern Botany, 3rd Ed., John Wiley and
Forestry Publications 18; F, after A. S. Foster. Sons, Inc., 1976. Used with permission.
4.10 From J. S. Pate and B. E. S. Gunning, Protoplasma 68, 5.13 Photo courtesy of D. R. Hoagland
140. 5.14 Photo courtesy of U. S. Forest Service.

E1
Chapter 7: Flowers, Fruits and Seeds 10.22 Courtesy of J. W. Schopf.
7.3 Redrawn from A. S. Foster and E. M. Gifford, Jr., 10.23 From J. W. Schopf, Journal of Paleontology 42:651 -688,
Comparative Morphology of Vascular Plants, W. H. and courtesy of the Society of Economic Paleontologists
Freeman, San Francisco; also from W. Bailey and A. C. I. and Mineralogists.
Smith, J. Arnold Arboretum 23, 256-265, 1942; also from 10.24 Redrawn from H. P. Banks, Evolution and Plants of the
J. E. Canright, Amer. J. Botany 39, 484-497, 1952. Past, Wadsworth, Belmont, California. Reprinted by
7.13 Courtesy of D. Hess. permission of the publisher.
7.16 A and B, redrawn from Priestly and Scott, An Introduction 10.25 A, redrawn from Goldring, W Scientific Monthly 24,
to Botany, 4th Ed., Longmans Group Ltd. 515-527; B, redrawn from C. B. Beck, American Journal
7.24 B, from W. W. Robbins, The Botany of Crop Plants, of Botany 49, 376, 1962.
McGraw-Hill, New York. Used with permission of McGraw- 10.26 A, courtesy of Field museum of Natural History; 6,
HillBook Company. cartoon courtesy of Sidney Harris.
7.27 Redrawn after Wilhelm Troll. 10.27 Redrawn from M. Hirmer, Handbuch der Palaobotanik 1,

7.28 Courtesy of D. Hess and H. Drever. 182-232.

7.35 A and B, redrawn after Wilhelm Troll. 10.28 Redrawn from M. Hirmer, Handbuch der Palaobotanik 1,
409-452, 1927.
Chapter 8: The Control of Growth and Development 10.29 Redrawn from E. Dorf, American Scientist 48, 341-364,
8.9 Photo courtesy of L. Rappaport.
1960. Reprinted by permission, American Scientist,
8.12 Photo courtesy of J. Goeschl and H. Pratt. Journal of Sigma Xi, the Scientific Research Society of
8.13 Photo courtesy of U.S. Department of Agriculture. North America.
8.14 Photo courtesy of Plant Industry Station, Crops Research 10.30 A and courtesy of C. Heiser; C, courtesy of
6, F. Smith.
Division, Agricultural Research Service, U.S. Department of 10.31 Redrawn from J. M. Savage, Evolution, Holt, Rinehart and
Agriculture.
Winston, New York.
8.15 Photo courtesy Crops Research
of Plant Industry Station,
10.32 Adapted from G. L. Stebbins, Processes of Organic
Division, Agricultural Research Service, U.S. Department of
Evolution, 2nd Ed., Prentice-Hall, Englewood Cliffs, New
Agriculture.
Jersey. Copyright 1971, p. 89, by permission of Prentice-
8.17 Photo courtesy of A. Lang. Hall, Inc.

Chapter 9: Plant Ecology

9.1 From Silvics of Forest Trees of the United States, U.S. Chapter 11: Algae
Department of Agriculture Handbook No. 271, Washington, 11.4 Redrawn from M. Neushul, Ecology 48, 90. Copyright
1 967 by the Ecological Society of America.
D.C.
9.2 Redrawn from W. D. Billings, Plants and the Ecosystem, 11.5 Redrawn from M. N. Hill, BioScience 78(10), 965,
published by the American Institute of Biological
Wadsworth Publishing Company, Belmont, California.
Sciences.
9.3 Redrawn from D. M. Gates, Ecology 46:1-13. Copyright,
1965, by the Ecological Society of America.
11.7 Redrawn from F. T. Haxo and L. R. Blinks, Journal of
General Physiology 33, 389-442, The Rockefeller
9.4 Courtesy of the New York Botanical Garden.
University Press.
9.6 Redrawn from D. M. Gates, Energy Exchange in the
11.8 All courtesy of N. J. Lang.
Biosphere, Harper and Row, New York.
11.9 Courtesy of I. Abbott.
9.12 Courtesy of H. Biswell.
11.10 Courtesy of Johns-Manville Corporation.
9.14 Courtesy of C. H. Muller.
11.13 Courtesy of N. J. Lang.
9.15 Redrawn from H. A. Mooney and W. D. Billings, Ecological
Monographs 37:1-29. Copyright, 1961, by the Ecological
11.14 A and 6, courtesy of D. Brandon; C, D, and E, courtesy of
N. J. Lang.
Society of America.
9.17 Courtesy of U. S. Forest Service.
11.16 6,redrawn from R. F. Scagel etal., An Evolutionary
Survey of the Plant Kingdom, 1 965, Wadsworth
9.18 Redrawn from P. W. Richards, Tropical Rain Forest,
Publishing Co., Inc., Belmont, California, 94002. Reprinted
Cambridge University Press.
by permission of the publisher.
9.21 Courtesy of U. S. Forest Service.
11.17 Courtesy of K. Schmitz.
9.22 Courtesy of J. Major.
9.23 Courtesy of J. Major.
11.18 A and C, redrawn after G. M. Smith, Freshwater Algae of

9.24 Courtesy of W. D. Billings.

the United States, 1933, McGraw-Hill, New York. Used
9.26
with permission of McGraw-Hill Book Co.
Courtesy of G. Webster.
9.27 Courtesy of O. A. Leonard.
11.19 Redrawn after F. Moewns.
9.28 Courtesy of J. Major.
11.23 Redrawn from R. F. Scagel, et al., An Evolutionary Survey

9.30
of the Plant Kingdom, 1965, Wadsworth Publishing Co.,
Courtesy of Ansel Adams.
Inc., Belmont, California, 94002. Reprinted by permission
9.31 '
From R. M. Love, "The rangelands of the western United
of the publisher.
States," Scientific American 222, 94. Copyright, 1970, by
Scientific American, Inc. All rights reserved.
11.25 Fthrough P, redrawn from R. F. Scagel, etal., An
Evolutionary Survey of the Plant Kingdom, 1 965,
9.34 From J. R. McBride, etal., California Agriculture
Wadsworth Publishing Co., Inc., Belmont, California,
29(12):8-9.
94002. Reprinted by permission of the publisher.

Chapter 10: Plant Taxonomy and Evolution

10.2 Courtesy of the Library of the New York Botanical Chapter 12: The Mycota
Garden. 12.1 Redrawn from C. J. Alexopoulous, Introductory Mycology,
10.8 With the assistance of D. Brown. John Wiley & Sons, Inc., New York, 1962.
10.17 Courtesy of L. D. Gottlieb. 12.4 From The Saprolegniaceae: With Notes On Other Water
10.18 Courtesy of W. M. Hiesey. Molds by William C. Coker, The University of North
10.19 Courtesy of the Field Museum of Natural History. Carolina Press, 1923. By permission of the publisher.
10.21 Courtesy of E. S. Barghoorn. 12.5 From The Saprolegniaceae: With Notes On Other Water

appendix e I Illustration Credits

E2
 Molds by William C. Coker, The University of North Botany, Macmillan, New York. Reprinted with permission
Carolina Press, 1923. By permission of the publisher. of Macmillan Publishing Co.,
Inc. Copyright 1949 by L H.
12.6 Based on L. R. Jones, N. J. Giddins and B. F. Lutman, Bailey,renewed 1977 by E. Z. Bailey.
"Investigations of the Potato Fungus," Vermont Exp. Sta. 13.15 Redrawn from R. M. Holman and W. W. Robbins, A
Bull. 168, 9, 1912 Textbook of General Botany, John Wiley & Sons, New
12.7 Photo courtesy of Plant Pathology Department, University York.
of California, Davis.
12.8 From R. M. Holman and W. W. Robbins, A Textbook of Chapter 14: Lower Vascular Plants
General Botany, John Wiley & Sons, New York. 14.3 Courtesy of D. Brandon.
12.9 Photo courtesy of R. N. Campbell. 14.4 Courtesy of D. Brandon.

12.12 Photo courtesy of C. E. Bracker and E. E. Butler. 14.8 B, redrawn from M. G. Sykes, Annals of Botany (London)
12.15 A through C; E, F: from Comparative Morphology of 22, 63; C, redrawn from E. Pritzel in A. Engel and K.
Fungi, by E. A. Gauman, Copyright 1928, McGraw-Hill Prantl, E, G, and H,
Die naturlichen pflanzenfamilien;
Book Co. Used with permission of McGraw-Hill Book Co. redrawn from Bruchmann, Flora 101, 220, with
H.
D, redrawn from E. A. Gauman, The Fungi: A Description permission of Gustav Fischer Verlag, New York; J,
of Their Morphological and Evolutionary Development, redrawn from material supplied by A. J. Eames.
Hafner Publishing Co., New York, 1952. 14.10 E, H, J, K, and L, redrawn from R. A. Slagg, American
I,

12.17 Redrawn from Selecta Fungorum Carpologia of the Journal of Botany 19, 106-7; J, and O redrawn from H.
/,

Brothers Tulasne, L. R. and C, Vol. 1861-65. I, Bruchmann, Flora 104, 180, with permission of Gustav
Translated into English by W. B. Grove; A. H. R. Buller Fischer Verlag, New York.
and C. L. Shear, Eds., Oxford (Clarendon Press) 1931. 14.13 Redrawn from E. R. Walker, The Botanical Gazette 92, 7,

Copyright Commonwealth Mycological Institute, Ferry copyright by the University of Chicago Press.
Lane, Kew, Richmond, Surrey, TW9 3AF, U. K. Used with 14.18 E, F,and G, redrawn from M. E. Hartmann, The Botanical
permission. Gazette 91 259, copyright by the University of Chicago
,

12.19 Photo courtesy of J. Ogawa. Press; Hand redrawn from D. H. Campbell, 1905,
/,

12.23 Redrawn from A. H. R. Buller, Researches on Fungi, Vol. Mosses and Ferns, Macmillan, New York; J and K,
Ill, The Production and Liberation of Spores in redrawn from R. M. Holman and W. W. Robbins, A
Hymenomycetes and Uredineae. Copyright Longman Textbook of General Botany, Wiley, New York.
Group Ltd., Harlow, Essex (U. K.). Used with permission. 14.19 A, courtesy of D. Brandon.
12.24 Photo courtesy of Roche.
12.25 Photo courtesy of Brownell. Chapter 15: Gymnosperms
12.27 A, from A. H. R. Buller, Researches on Fungi, Vol. Vll, 15.4 A, courtesy of the American Museum of Natural History;
The Sexual Process in the Uredinales. Royal Society of B, redrawn from L.H. Bailey, The Cultivated Conifers in
Canada, University of Toronto Press, 1 922. Used with North America, Macmillan, New York.
permission of the Royal Society of Canada and University 15.12 G, H, and redrawn from Coulter and Chamberlain,
/,

of Toronto Press. D, photo courtesy of E. E. Butler. Morphology of the Gymnosperms, University of Chicago
12.30 A, after K. B. Raper and D. F. Alexander, J. Elisha Press, with permission from the University of Chicago
Mitchell Scientific Society, Vol. 61, August 1945. Used Press.
with permission. 15.14 Slide courtesy of Triarch Products.
12.31 Redrawn from E. A. Gauman, Fungi: A Description of 15.16 Courtesy of the Field Museum of Natural History.
Their Morphological Features and Morphological 15.18 Courtesy of A. Addicott.
Development, Hafner Publishing Co., New York, 1950.
12.32 Redrawn from Darbishire. Chapter 16: The Angiosperms
16.1 Cand D, redrawn from Bonnier and Sablon, Cours de
Chapter 13: The Bryophyta Botanique, Librairie Generale de I'Enseignement.
13.1 Redrawn from W. P. Schimper, Recherches sur les 16.2 Courtesy of McMinn.
mousses, Strasbourg, 1948. 16.6 Redrawn from L. H. Bailey, Manual of Cultivated Plants,
13.2 Redrawn from R. F. Scagel et ai, An Evolutionary Survey Macmillan, New York. Copyright 1949 by L. H. Bailey,

of the Plant Kingdom, 1965, Wadsworth Publishing Co., renewed 1977 by E. Z. Bailey.
Inc., Belmont, California, 94002. Reprinted by permission 16.10 Redrawn from L. H. Bailey, Manual of Cultivated Plants,
of the publisher. Macmillan, New York. Copyright 1949 by L. H. Bailey,
13.3 8, courtesy of C. Laning. renewed 1977 by E. Z. Bailey.
13.4 C. Hebant, 1967, Naturalia Monspeliensia Ser. Bot. 18, 16.11 Redrawn from L. H. Bailey, Manual of Cultivated Plants,
301. Macmillan, New York. Copyright 1949 by L. H. Bailey,
13.8 Courtesyof W. Russell and D. Hess. renewed 1977 by E. Z. Bailey.
13.14 Redrawn from G. M. Smith, F. M. Gilbert, G. S. Bryan, 16.12 Redrawn from E. Korsmo, Unkrauter Im Ackerbau Der
R. I. Evans, and J. F. Stauffer. A Textbook of General Neuzeit, 1930, Springer.

Illustration Credits

E3
INDEX TO GENERA

This index includes all the genera Anabaena (Cyanophyta), 227, 230, Beet, see Beta
mentioned in the text. Species are not 234, 237, 238 Begonia (Begoniacea), 81, 88
shown, thus a given set of entries for a Ananas (Bromeliaceae), 104, 141, 146, Berberis (Berberidaceae) 69, 268-70
genus may refer to more than one 167 Bermuda grass, see Cynadon
species. In parentheses following the Andropogon (Poaceae), 180 Beta (Chenopodiaceae), leaf 30, 85,
genus is the family (for genera in the Anemone (Ranunculaceae), 333 108, 146
Anthophyta) or the class (for genera in Aneurophyton (f), 211-14 Betula (Betulaceae), 94
the Mycota) or the division (for all other Animikiea (f), 210 Biddulphia (Bacillariophyta), 228
genera). Common names are listed, An nu Ian a (f), 215 Bignonia (Bignoniaceae), 68, 80, 82
but they give only cross-references to Anthoceros (Bryophyta), 286, 287, 309 Bigtree, see Sequoiadendron
the scientific name, and all page en- Aphanizomenon (Cyanophyta), 227 Bindweed, see Convolvulus
tries appear after the scientific name. Apium (Apiaceae), 142 Birch, see Betula
Common names selected are those Apple, see Malus Blackberry, see Rubus
preferred by Bailey (Manual of Culti- Apricot, see Prunus Black crowberry, see Empetrum
vated Plants) or Munz and Keck (A Aquilegia (Ranunculaceae), 127, 129, Black locust, see Robinia
California Flora). 333 Black spruce, see Picea
Arabidopsis (Brassicaceae), 38 Blood lily, see Haemanthus
Abies (Coniferophyta), 181, 184, 310, Arachis (Fabaceae), 338 Blossom infecting smut, see Ustilago
313, 324 Archeopteris (f), 211-214 Blue mold, see Penicillium
Acacia (Fabaceae), 338 Archaeospheroides (f), 208 Boerhaavia (Nyctaginaceae), 175
Acer (Aceraceae), 96 Aristida (Poaceae), 180 Botrychium (Pterophyta), 303
distribution, 173, 174 Armeria (Plumbaginaceae), 53 Box elder, see Acer
fruit, 142 Artemisia (Asteraceae), 338 Boysenberry, see Rubus
leaf, 46, 49 Arthrospira (Cyanophyta), 238 Brassica (Brassicaceae), 75, 92, 257,
Aconitum (Ranunculaceae), 333 Ash, see Fraxinus 334
Adder's Tongue, see Ophioglossum Asparagus (Liliaceae), 68, 127, 339 Bread mold, see Rhizopus
Adiantum (Pterophyta), 303 Aspen, see Populus Bremia (Oomycete), 258
Aesculus (Hippocastanaceae), 49, 73 Aspergillus (Fungi Imperfecti), 271 Bristlecone pine, see Pinus
Agardiella (Rhodophyta), 226 Asplenium (Pterophyta), 303, 305 Broad bean, see Vicia
Agaricus (Basidiomycetes), 266-267 Aster (Asteraceae), 338 Broccoli,see Brassica
Agave (Agavaceae), 66, 69 Asterionella (Bacillariophyta), 228 Brome grass, see Bromus
Ageratum (Asteraceae), 338 Asteromphalus (Bacillariophyta), 228 Bromus (Poaceae), 182
Ailanthus (Simarubaceae), 49 Avena (Poaceae), 270, 334 Brown rot fungus, see Monilinia
Albugo (Oomycetes), 259 fruit, 140 Brussel sprouts, see Brassica
Aleurites (Euphorbiaceae), 76 growth, 158, 167 Bryophyllum (Crassulaceae), 81
Alfalfa, see Medicago leaf, 75 Bryum (Bryophyta), 275
Alisma (Alismaceae), 339 Avicennia (Verbenaceae), 107 Buckeye, see Aesculus
Allium (Liliaceae), bulb, 68 Butchers broom, see Ruscus
flower, 129, 339 Baldcypress, see Taxodium Buttercup, see Ranunculus
root, 29 Bamboo, see Bambusa
seed, 148, 149 Bambusa (Poaceae), 190, 344 Cabbage, see Brassica
seedling, 151, 153, 154 Banana, see Musa Catamites (f), 211, 215

Aloe (Liliaceae), 339 Banyan, see Ficus California lilac, see Ceanothus
Alpine sorrel, see Oxyria Barberry, see Berberis California poppy, see Eschscholzia
Alternanthera (Amaranthaceae), 57 Barley, see Hordeum Calla lilly, see Zantesdeschia
Amanita (Basidiomycete), 199, 267 Basswood, see Tilia Callixylon (f), form genus of
Ambrosia (Chenopodiaceae), 133, 137 Bean, see Phaseolus Archeopteris
American larch, see Larix Beard grass, see Andropogon Calochortus (Liliaceae), 341
American laurel, see Kalmia Beech, see Fagus Calocedrus (Coniferophyta), 181
Caltha (Ranunculaceae), 333, 339 Cordyline (Aguaceae), 66 Eleusine (Poaceae), 344
Cannabis (Moraceae), 68, 70, 255 Corn, see Zea Empetrum (Empetraceae), 187
Capsella (Brassicaceae), 259 Cortaderia (Poaceae), 344 Eobacterium (f), 208, 210
Capsicum (Solanaceae), 334 Costaria (Phaeophyta), 226 Ephedra (Gnetophyta), 310, 312, 326
Carex (Cyperaceae), 187 Cotton, see Gossypium Epidendrum (Orchidaceae), 69
Carrot, see Daucus Cottonwood, see Populus Equisetum (Sphenophyta), 198, 201,
Carthamus (Asteraceae), 338 Couch whip, see Fouquieria 297, 300-01, 309
Carya (Jugiandaceae), 150, 186 Cow lily, see Nuphar Ergot, see Claviceps
Cassiope (Ericaceae), 187 Crabgrass, see Digitaria Erodium (Geraniaceae), 196
Castanea (Fagaceae), 142 Creosote bush, see Larrea Erysiphe (Ascomycete), 263, 264
Castor bean, see Ricinus Crocus (Iridaceae), 340 Eschscholzia (Papaveraceae), 148
Catalpa (Bignoniaceae), 46, 48, 49 Croton (Euphorbiaceae), 76 Eucalyptus (Myrtaceae), 72, 81
Cattleya (Orchidaceae), 343 Cucumber, see Cucumis Euglena (Chlorophyta), 204, 239
Caulerpa (Chlorophyta), 223 Cucumis (Cucurbitaceae), 133, 154, Euphorbia (Euphorbiaceae), 102, 197
Cauliflower, see Brassica 167. 337
Ceanothus (Rhamnaceae), 175, 184 Cucurbita (Cucuribitaceae), 102, 167, Fagus (Fagaceae), 328
Cedar, see Cedrus 214, 337, 339 Fauchea (Rhodophyta), 226
Cedrus (Coniferophyta), 310, 312, 324 flower, 127 Ficus (Moraceae), fossil community,
Celery, see Apium fruit, 140 185
Centaurea (Asteraceae), 80 ultrastructure, 55 germination, 189
Cerastium (Caryophyllaceae), 130 Cupressus (Coniferophyta), 313, 317, fruit, 141, 142, 145-146
Ceratium (Pyrrhophyta), 228 324 root, 86, 89
Ceratocorys (Pyrrhophyta), 228 Cuscuta (Cuscutaceae), 85, 105 Field mushroom, see Agaricus
Cercis (Fabaceae), 338 Cyathea (Pterophyta), 303 Fig. see Ficus
Chain fern, see Woodwardia Cycas (Cycadophyta), 324, 325 Filaree, see Erodium
Chara (Chlorophyta), 23, 117 Cynodon (Poaceae), 69 Fir, see Abies

Cheatgrass, see Bromus Cypress, see Cupressus Firethorn, see Pyracantha

Cheilanthes (Pterophyta), 303 Fischerella (Cyanophyta), 238
Cherry, see Prunus Dactylaria (Mycota), 272 Five-finger, see Potentilla
Chestnut, see Castanea Daffodil, see Narcissus Five-finger fern, see Adiantum
Chickweed, see Cerastium Dahlia (Asteraceae), 338 Flyagaric, see Amanita
Chlamydomonas (Chlorophyta), 223, Dandelion, see Taraxacum Fomes (Basidiomycete), 267, 268
242, 244-^16 Darlingtonia (Sarraceniaceae), 81, 82, Fouquieria (Fouquiereaceae). 80
Chlorella (Chlorophyta), 199 105 Foxtail, see Setaria

Chondrus (Rhodophyta), 227, 232 Datura (Solanaceae), 142 Fragaria (Rosaceae), fruit, 141, 144,
Chosenia (Salicaceae), 335 Daucus (Apiaceae), 86, 142 146
Christmas rose, see Helleborus Day-lily, see Hemerocallis stem, 69
Chrysanthemum (Asteraceae), 338 Death angel, see Amanita Fraser fir, see Abies
photoperiod, 169, 170 Degeneria (Degeneriaceae), 125 Fraxinus (Oleaceae), 46
Cichorium (Asteraceae), 338 Delphinium (Ranunculaceae), 333 Freesia (Iridaceae), 340
Cinquefoil, see Potentilla Dicentra (Fumariaceae), 128, 129, 139 Fuchsia (Onagraceae), 135, 139
Cirsium (Asteraceae), 129 Dicranoweisia (Bryophyta), 275 Fucus (Phaeophyta), 210, 223, 232,
Citrus (Rutaceae), 165, 166. 167, 203, Digitaria (Poaceae), 73 243, 323
271 Dionaea (Droseraceae), 81, 82, 105 life history, 246-247, 250
fruit, 140, 144 Dioon (Dycadophyta), 324, 325 Funaria (Bryophyta), 275, 276, 280
smog, 193 Dodder, see Cuscuta
Cladonia (Lichen), 272 Douglas fir, see Pseudotsuga Gaillardia (Asteraceae), 338
Clarkia (Onagraceae), 205 Downy mildew, see Bremia Garlic, see Allium
Claviceps (Ascomycetes), 265 Dracena (Agavaceae), 66, 67 Gelidium (Rhodophyta). 231
Clematis (Ranunculaceae), 333 Dragon's-blood tree, see Dracena Giant kelp, see Macrocystis
flower, 126. 135 Drimys (Magnoliaceae), 125, 185 Gibberella fujikuroi (Mycota), 161
seed dispersal. 150 Drosera (Droseraceae), 81, 82 Gigartina (Rhodophyta), 226
Cliff brake, see Pellaea Duckweed, see Lemna Ginkgo (Ginkophyta), 204, 310, 312,
Clover, see Trifolium Dulce, see Rhodymenia 324
Club moss, see Lycopodium Dusty miller, see Centaurea Ginseng, see Panax
Cnidoscolus (Euphorbiaceae), 74 Dutchman's breeches, see Dicentra Gladiolus (Iridaceae), 70, 340
Coconut, see Cocos Gleditsea (Leguminoseae), 69, 236
Cocos (Palmaceae), 150 Echeveria (Crassulaceae), 46 Gloeotrichia (Cyanophyta), 227
Coleus (Labiatae), 160 Ectocarpus (Phaeophyta), 199, 246, Glycine (Fabaceae), 95, 271
Colocasia (Araceae), 214 248 Gnetum (Gnetophyta), 204, 310, 324
Columbine, see Aquilegia Eel grass, see Zostera Golden back see Pityrogramma
fern,

Convolvulus (Convolulaceae), 148 Eggplant, see Solanum Gomum (Chlorophyta), 223

Cooksonia (f), 211 Elderberry, see Sambucus Gonyaulax (Pyrrhophyta), 227

Index to Genera
12
Gooseberry, see Ribes Juncus (Cyperaceae), 224 Mala mujer, see Cnidoscolus
Gossypium (Malvaceae), 135, 334, 335 Juniper, see Juniperus Malus (Rosaceae), 75, 87, 140, 144,
Gracilaria (Rhodophyta), 231 Juniperus (Coniferophyta), 310, 313, 145, 146, 166, 167, 171, 264, 336
Grape, see Vitis 314, 324 Mangrove, see Avicennia
Green mold, see Aspergillus Maple, see Acer
Ground-pine, see Lycopodium, Taxus Kalanchoe (Crassulaceae), 82 Marchantia (Bryophyta), 280, 284-286
Guayule, see Parthenium Kale, see Brassica Marigold, see Tagetes
Gum tree, see Eucalyptus Kalmia (Ericaceae), 187 Mariposa lily, see Calochortis
Gymnodinium (Pyrrhophyta), 227 Kelp, see Laminaria Marsh marigold, see Caltha
Kohlrabi, see Brassica Medicago (Leguminoseae), 21, 24, 96,
Haemanthus (Amaryllidaceae), 35-37 105
Haircap moss, see Polytrichum Labrador tea, see Ledum smog, 193
Hedera (Araliaceae), 85 Lactuca (Asteraceae), 258, 338 stem, 58, 59
Helianthus (Asteraceae), 32, 217, 338 leaf, 190 stomata, 75
flower, 129, 135 seed, 150 Melon, see Cucurbita
fruit, 141, 142 Laminaria (Phaeophyta), 210, 224, 226, Mentha (Lamiaceae), 127, 335
stem, 20, 57, 159 232, 242, 243, 249 Mesquite, see Prosopis
Helleborus (Ranunculaceae), 123, 331 Larch, see Larix Mexican tea, see Ephedra
Hemerocallis (Liliaceae), 133, 339 Larix (Coniferophyta), 312, 314, 324 Microcystis (Cyanophyta), 227
Hemlock, see Tsuga Larkspur, see Delphinium Millet, see E leu cine; Pennicetum

Hemp, see Cannabis Larrea (Zygophyllaceae), 186 Mimosa (Fabaceae), 338

Hepatica (Ranunculaceae), 333 Lathyrus (Fabaceae), 340 Mint, see Mentha
Hibiscus (Malvaceae), 140, 334 classification, 198 Mistletoe, see Phoradendron
Hickory, see Carya leaf, 80 Mnium (Bryophyta), 275-280
Holly, see Ilex Lavender, see Lavendula Monilinia (Ascomycetes), 264-65
Holly fern, see Polystichum Lavendula (Lamiaceae), 335 Monk's hood, see Aconitum
Honey locust, see Gleditsia Leather leaf, see Polypodium Monotropa (Pyrolaceae), 1
Honey mesquite, see Prosopis Ledum (Ericaceae), 184 Moonwort, see Botrychium; Lunaria
Hordeum (Poaceae), 344 Leek, see Allium Morchella (Ascomycetes), 265
leaf, 72, 73 Lemna (Lemnaceae), 223, 328 Morel, see Morchella
growth, 162, 163 Lepidodendron (f), 211, 214 Mormon tea, see Ephedra
root, 86 Lettuce, see Lactuca Morus (Moraceae), 61, 141
smog, 193 Lilac, see Syringa Mother-in-law's tongue, see
Hornwort, see Anthoceros Li Hum (Liliaceae), 39, 124, 198, 339 Sanseviena
Horse chestnut, see Aesculus Lily,see Lilium Mountain ash, see Sorbus
Horsetail, see Equisetum Linden, see 7'ilia Mountain hemlock, see Tsuga
Hyacinth, see Hyacinthus Lip fern, see Cheilanthes Mountain sorrel, see Oxyria
Hyacinthus, (Liliaceae), 339 Liriodendron (Magnoliaceae), leaf, 71 Mulberry, see Morus
Hydrocharis (Hydrocharitaceae), 27 taxonomy, 205, 332 Mullein, see Verbascum
Hydrodictyon (Chlorophyta), 223 Liverwort,see Marchantia; Riccia Musa (Musaceae), 140, 167
Loblolly Pine,see Pinus Mushroom, see Agaricus
Ilex (Aquifoliaceae), 185 Longleaf Pine, see Pinus Mustard, see Brassica
Incense-cedar, see Calocedrus Longpod beans, see Vicia Myriogramme (Rhodophyta), 225
Indian pipe, see Monotropa Lotus, see Nelumbo
Ipomoea (Convolvulaceae), 214 Lunaria (Crucifereae), 142 Narcissus (Amaryllidaceae), 128, 130
Iridophycus (Rhodophyta), 200 Lupinus (Fabaceae), 338 leaf, 80
Iris (Iridaceae), 339, 340, 342 Lycopersicon (Solanaceae), 334, 335, stem, 68, 70
flower, 133 336 Nasturtium, see Tropaeolum
stem, 46, 66, 69 leaf, 75, 79, 81 Nectarine, see Prunus
Irish moss, see Chondrus nutrition, 100, 104, 105, 106 Needlegrass, see Stipa
Isoejes (Lycophyta), 288, 308 stem, 58, 159, 161 Nelumbo (Nymphaeaceae), 150
Ivy, see Hedera Lycopodium (Lycophyta), 198, 202. Nematode trapping fungus, see
288-291, 293-294, 308-309 Dactylaria
Jeffrey pine, see Pinus Nereocystis (Phaeophyta), 226
Jimson weed, see Datura Macrocystis (Phaeophyta), 227, 232, Nicotiana (Solanaceae), 54, 163, 164,
Joint fir, see Ephedra 234, 235, 237, 239 334
Joshua tree, see Yucca Magnolia (Magnoliaceae), 331, 332, Nori, see Porphyra
Juglans (Juglandaceae), bud 46, 49 333, 339 Nostoc (Cyanophyta), 209, 227-, 234
flower, 127, 129, 137 flower, 124, 125,128 Nuphar (Nymphaeaceae), 187
fruit, 142 fossil community, 185

seed, 150 fruit, 142 Oak, see Quercus

stem, 48 stem, 56 Oat, see Avena
twig, 47 Maidenhair tree, see Ginkgo Ocotillo, see Fouquieria

Index to Genera
13
Oedogonium (Chlorophyta), 244 stem, 45, 46, 56, 63 Radish, seeRaphanus
Onion, see Allium succession, 186 Ragweed, see Ambrosia
Ophioglossum (Pterophyta), 303 Pisum (Leguminoseae), 32 Ranunculus (Ranunculaceae), 92, 331
Ophrys (Orchidaceae), 139 flower, 125, 127, 129, 134, 135, 138 332, 333, 339
Orange, see Citrus fruit, 138, 140 Raphanus (Crucifereae), 255
Oryza (Poaceae), 107, 120, 161, 162, growth, 157, 159, 162, 165 flower, 129
271, 227, 334 leaf, 71 root, 88

fruit, 142 seedling, 150, 153 Raspberry, see Rubus

leaf, 75 Pitcher plant, see Darlingtonia Red bud, see Cercis
seed, 148, 149 Pityrogramma (Pterophyta), 303 Red foxtail grass, see Setaria
Oscillatoria (Cyanophyta), 209, 227, Plane tree, see Platanus Red spruce, see Picea
238, 239, 240 Platanus (Platanaceae), 49 Redwood, see Sequoia
Oxyria (Polygonaceae), 184 Plum, see Prunus Reindeer moss, see Cladonia
Poker plant, see Tritoma Rhizobium (Schizomycophyta), 94, 105
Paeonia (Ranunculaceae), 333 Poly podium (Pterophyta), 202, 303, Rhizoctonia (Fungi Imperfecti), 261
Painted tongue, see Salpiglossis 304, 305 Rhizopus (Zygomycetes), 259
Palm, see Washingtonia Polypody, see Polypodium Rhodymenia (Rhodophyta), 226, 227
Pampas grass, see Cortaderia Polyporus (Basidiomycetes), 267 Ribes (Saxifragaceae), 129
Panax (Araliaceae), 94 Polysiphonia (Rhodophyta), 242, 243, Riccia (Bryophyta), 199, 281, 282, 285
Parthenium (Asteraceae), 338 246, 248 Rice, see Oryza
Parthenocissus (Vitaceae), 68, 71, Polystichum (Pterophyta), 303, 305 Ricinus (Euphorbiaceae), 71, 148
73 Polytrichum (Bryophyta), 201, 278 Robinia (Fabaceae), leaf, 80, 82
Pea, see Lathy rus; Pisum Ponderosa Pine, see Pinus stem, 60, 69
Peach, see Prunus Poplar, see Populus taxonomy, 338
Peanuts, see Arachis Poppy, see Eschscholzia Rock kelp, see Fucus
Pear, see Pyrus Populus (Salicaceae), 335 Rosa (Rosaceae), 71, 336
Peat moss, see Sphagnum flower, 129 Rose, see Rosa
Pellaea (Pterophyta), 303, 305 stem, 62, 70, 71 Rose-mallow, see Hibiscus
Pennicetum (Poaceae), 96 Porcupine grass, see Stipa Rubus (Rosaceae), 88, 141, 145, 335,
Penicillium (Fungi Imperfecti), 253, Porphyra (Rhodophyta), 200, 223, 227, 336, 338
271-72 231 Ruscus (Liliaceae), 68
Peonia, see Paeonia Postelsia (Phaeophyta), 200, 240, 241 Rush, see Juncus
Peony, see Paeonia Potato, see Solanum Rye, see Secale
Pepper (the vegetable), see Capsicum Potentilla(Rosaceae), 205-206
Peppermint, see Mentha Primrose, see Primula Saccharum (Poaceae), 120, 344
Petroselinum (Apiaceae), 142 Primula (Primulaceae), 139 Saccharomyces (Ascomycetes), 11,
Petunia (Solanaceae) 334 Prosopis (Leguminoseae), 174 273
163, 253, 265,
Phaseolus (Leguminoseae), 25, 214 Prunus (Rosaceae), 46, 171, 259, 264, Safflower, see Carthamus
fruit, 138, 140 336 Sage, see Salvia
leaf, 76 flower, 127, 129, 130 Sagebrush, see Artemisia
seed, 149 fruit, 140, 142, 145 Sago-palm, see Cycas
seedling, 150, 151 nutrition, 104 Salix (Salicaceae), 111, 127, 129, 335,
shoot apex, 50 Prymnesium (Chrysophyta), 227 337
stomate, 75 Pseudotsuga (Coniferophyta), 176, root, 88, 194
Phoradendron (Loranthaceae), 105, 189, 310, 324 Salpiglossis (Solanaceae), 334
150 Psilotum (Psilophyta), 198, 201, 204, Salt cedar, see Tamarix
Phylloglossum (Lycophyta), 308 211, 288-291, 305, 309 Salvia (Labiateae),335
Phytophthora (Oomycetes), 257, 258 Ptilota (Rhodophyta), 223 amensalism, 182, 183
Picea (Coniferophyta), 310, 313, 324 Puccinia graminis (Basidiomycetes), flower, 129
subalpine forest, 184, 185 268-270 Sambucus (Caprifoliaceae), 64, 89
succession, 187 Pyracantha (Rosaceae), 69 Sansevieria (Agauaceae), 66
Pigweed, see Chenopodium Pyrus (Rosaceae), 144, 167 Saprolegnia (Oomycetes), 255, 256,
Pine, see Pinus 257
Pineapple, see Ahanas Quercus (Fagaceae), 156, 272 Sargassum (Phaeophyta), 225
Pink, see Armeria chaparral, 184 Scouring-rush, see Equisetum
Pinus (Coniferophyta), classification, flower, 137 Sea lettuce, see Ulva
202, 310, 311 fossil community, 185 Sea-palm, see Postelsia
fire response, 180, 181 fruit, 142, 145 Secale (Poaceae), 89, 170, 265, 344
life cycle, 315-326 leaf, 71, 75 Sedge, see Carex
root, 94,106 root, 94 Selaginella (Lycophyta), 288, 291 292, ,

seed, 150 stem, 61, 62, 63 296-298, 308, 309

seed dispersal, 151 succession, 180, 181, 186 cells, 24

shade tolerance, 176 taxonomy, 205, 206 taxonomy, 203

smog, 193, 194 see Isoetes
Quillwort, Sensitive plant, see Mimosa

Index to Genera
14
Sequoia (Coniferophyta), 313, 315, Tagetes (Asteraceae), 338 Ustilago (Basidiomycete), 270, 271
316, 317 Talaromyces (Ascomycetes), 270
distribution, 173 Tamarack, see Larix Venus Fly Trap, see Dioneae
stem, 45, 62 Tamarisk, see Tamarix Verbascum (Scrophulariaceae), 102,
Sequoiadendron (Coniferophyta), 181, Tamarix (Tamaricaceae), 174 150
313 Taraxacum (Compositeae), 94, 151, Vicia (Fabaceae), 32, 71, 112
Setaria (Poaceae), 148, 149, 151 168, 338 Virginia creeper, see Parthenocissus

Sheep see Rumex

sorrel, Taro, see Colocasia Virgin's-Bower, see Clematis
Shepherd's purse, see Capsella Taxodium (Coniferophyta), 107, 313 Vitis (Vitaceae), 68, 140, 264
Shortleaf Pine, see Pinus Taxus (Coniferophyta), 310, 312, 315, Volvox (Chlorophyta), 223
Slash Pine, see Pinus 324
Walnut, see Juglans
Smilax (Liliaceae), 68 Tetraphis (Bryophyta), 275, 277
Washingtonia (Palmaceae), 64, 146
Smuts, see Tilletia; Ustilago Thorn-apple, see Datura
Water mold, see Saprolegnia
Solarium (Solanaceae), 140. 257, 258, Thistle, see Centaurea; Cirsium Water net, see Hydrodictyon
334 Thyme, see Thymus
Water plantain, see Alisma
leaf, 75 Thymus (Lamiaceae), 335
Watsonia (Irid^ceae), 340
stem, 68, 69, 70 Tilia (Tiliaceae), 56
Welwitschia (Gnetophyta), 204, 310,
Sorbus (Rosaceae), 184 Tillandsia (Bromeliaceae), 175
324, 327
Sorghum (Poaceae), 120 77//ef/a (Basidiomycetes), 270
Wheat, see Triticum
Soybean, see Glycine Tmesipteris (Psilophyta), 305
Wheat rust, see Puccinia
Spanish bayonet, see Yucca Tobacco, see Nicotiana
Whiskfern, see Psilotum
Spearmint, see Mentha Tomato, see Lycopersicon
White mottled rot, see Fomes
Sphagnum (Bryophyta), 187, 275, 277, Tradescantia (Commelinaceae), 44,
White rust, see Albugo
280 196
Willow, see Salix
Spiderwort, see Tradescantia Tree fern, see Cyatheae
Windflower, see Anemone
Spinacea (Chenopodiaceae), 257 Tree-of-Heaven, see Ailanthus
Wiregrass, see Aristida
Spinach, see Spinacea Triceratium (Bacillariophyta), 233
Wisteria (Fabaceae), 338
Spirogyra (Chlorophyta), 223 Trifolium (Fabaceae), 51, 105, 338
Woodwardia (Pterophyta), 305
Spleenwort, see Asplenum Triticum (Poaceae), 120, 170, 267, 270,
Spruce, see Picea 344
Yeast, See Saccharomyces
Spurge, see Euphorbia cell, 162
23,
Yucca (Liliaceae), 65, 103, 339
Squash, see Cucurbita embryo, 154
Yew, see Taxus
Sterculia (Sterculiaceae), 124, 125 fruit, 140, 142

Stipa (Poaceae), 190 stem, 67 Zamia (Cycadophyta), 324, 325

Storksbill,see Erodium Tropaeolum (Tropaeolaceae), 71, 75, Zantedeschia (Araceae), 126
Strangler fig, see Ficus 139 Zea (Poaceae), 21, 22, 96, 267, 270,
Strawberry, see Fragaria Trumpet flower, see Bignonia 271, 217, 344
Stylites (Lycophyta), 308 Tsuga (Coniferophyta), 310, 324 embryo, 148, 154
Sugar beet, see Beta distribution, 175, 176 flower, 127, 135
Sugar cane, see Saccharum shade tolerance, 176 fruit, 140, 142

Sundew, see Drosera succession, 187 growth, 162

Sunflower, see Helianthus Tulip, see Tulipa leaf, 75, 77, 102

Sweet pea, see Lathyrus Tulipa (Liliaceae), 128, 130, 134, 144, photosynthesis, 120, 121
Sweet potato, see Ipomoea 339 root, 92
Sword fern, see Polystichum see Liriodendron
Tulip tree, seed, 148
Sycamore, see Platanus Twayblade, see Ophrys seedling, 154
Syringa (Oleaceae), 46, 171 stem, 66, 67
leaf, 48, 77 Ulothrix (Chlorophyta), 242,245-246 Zinnia (Asteraceae), 71, 338
stem, 161 Ulva (Chlorophyta), 200, 225, 226 Zostera (Zosteraceae), 226

Index to Genera
15
SUBJECT INDEX

This index includes terms, topics, Aldehyde. 3 Annual rings. 60, 181. 187, 194, 211
place names, persons, and plant Aleurone layer. 149. 162, 162 Annulus, of fern sporangium, 304
names above the genus level. The Alfisol. 179, 189 of moss sporangium, 279, 280, 281
generic index should be consulted for Algae, 1, 75, 120, 170, 177, 192. 209, Anther, 124, 128, 130, 139, 167, 317.
other plant names. Not all persons 222-252, 234. 236. 272. 273, 274. 328-330
mentioned in the text are listed here. 288. 323 Antheridiophores, 284, 285, 286
Pages on which terms have been blue-green, see Cyanophyta Antheridium, 243, 256, 257. 260, 276,
defined are shown in bold face. Glos- brown, see Phaeophyta 280, 285, 286, 288-309. 323. 325
sary entries have not been indexed. classification of, 198 Anthocerotae. 274, 275, 286-287
distribution, 177 Anthocyanin. 30. 166, 188
ABA, see Abscisic acid economic importance, 227, 231, Anthophyta, 197, 198, 203, 204, 205.
Abscisic acid, 102, 157. 166, 172 232, 234 209, 211-215. 221, 288, 310, 317,
Abscission. 24 fossil. 220. 227. 231 323, 324, 328-344
leaf. 81, 83-84, 166, 168, 172, 191 green, see Chlorophyta Antibiotics. 253. 271
Absorption, 96-110 red. see Rhodophyta Anticlinal, 36
oxygen, 106-107 reproduction, 239, 242-252 Anticodon. 14, 15
solutes by roots, 103-106 Algin, alginic acid, 232, 236, 237, 252 Antipodal cells, 135, 146, 328
water, 96-101, 166 Alkaloids, 265 Apex, root, 88
Accessory fruit, 146 Allele. 42 shoot, 77-79, 170, 171, 172.
Acetic acid, 376 Allosteric site, 12 Apiaceae. 142, 336, 344
Acetyl group. 11 Alternation of generations. 243-252, Apical dominance. 45. 159-160, 164,
Achene, 141, 142, 145, 146 274, 290, 297 167, 172
Acidity, 376 heteromorphic generations. 244 Apical meristems, 47, 50, 88-89, 149,
Activated complex, 374, 375 isomorphic generations, 244 154, 155. 162, 164
Active site, 8, 14, 103 see also plant group in question Aplanospores, 242
Active transport, 101, 102. 106. 108. Amensalism, 182-183, 194 Apothecium, 260, 262, 263. 265
109 Amino acid, 5, 6, 8, 14-16, 108. 157. Aquatic plants. 80. 81. 192, 198, 208,
Adenine. 12, 163 165, 183. 203, 205, 227. 254, 255 222-252, 273. 297. 304, 309
Adenosine diphosphate, see ADP Amino group, 5, 376 Arboretum, see Botanic gardens
Adenosine monophosphate, see AMP Ammonia, 227. 371 Archegonia, 276, 280, 281, 285, 286,
Adenosine triphosphate, see ATP Ammonium. 103, 105. 208 288-309, 317, 325, 326, 331
Adhesion. 99, 100, 109 AMP. 8. 253 Archegoniophores, 284. 285, 286
Adnation, 128, 332 Amylase. 163 Aridisol. 179
ADP. 9-16 Amylopectin, 234, 236 Aril. 310, 312, 315
Aecia, 269, 270 Amyloplast, 20. 21. 25, 26, 32. 159 Arthrophyta, 288. See also
Aeciospore, 270 Anabolism. 8-12, 16. 168 Sphenophyta
Aflatoxin, 271 Anatomy, 45-95. 195. 203. 207. 219. Ascocarp, 260, 264, 273
Agar, 231-232, 236, 252 221. 237. 289-290, 293. 296. 297. Ascogenous hyphae, 260, 261
Agaricales, 267 301. 304, 315-317, 323. 324 Ascomycetes, 260-265, 270, 272, 273
Age, cell, 54, 60. 69 Anaphase, in meiosis, 39, 40, 42 classification, 253
earth, 208 in mitosis, 34, 35, 43 Ascospores, 262, 273
plant, 176, 211, 304, 310, 315. 324. Androecium, 124, 128. 130-33. 139 Ascus, 260, 262, 264, 270, 273
327 Angiosperm, classification, 198 Asexual reproduction, see
seed, 150, 191 evolution 237, 331-344 Reproduction; or plant in question
Agriculture, 190. 192, 193, 194. 214, life cycle, 130-140, 328-331 Aspartic acid, 119
224 see also Anthophyta Aspect, slope, 175, 194
Air chambers, 281 Anion, 370 Asteraceae. 129. 332, 335, 336, 344
Alcohol, 4, 11, 16, 163. 203 Anisogamy, 242, 245. 246. 252 Atmospheric pressure, 156
Alcoholic fermentation, 11, 271 Annual plants. 170, 175, 191, 195, 337 Atoms. 369-377
ATP, 4, 8-16, 102 103, 106, 108, 117, gametophyte, 274-276, 281, 285 Carbon monoxide, 208
119-120, 122,. 157 sporophyte, 274-286 Carbonyl, 377
Atropine, 334 Bud, 56, 161, 164, 167, 171, 172, 276, Carboxyl group, 5, 6, 376, 377
Auricles, 72, 73 318 Carnauba wax, 4
Autoecious, 268 accessory, 46 Carotenoids, 1 14
Auxin, 104, 156-161, 162, 163, 164, adventitious, 46, 164 Carpel, 123, 124, 125, 130, 134-135,
167. 172 apical,166 137, 138, 139, 140, 141, 146, 330,
Axil, leaf, 45 arrangements, 45-47 332
axillary, 68, 72-73, 159 Carpogonium, 246
Bacillariophyta, 204, 225-229, dormant, 46 Carpospore, 242, 246
231-233, 236, 239, 243, 244, 246 lateral, 68 Carposporophyte, 246, 248
Bacteria. 1, 1 1, 15, 16, 17, 79, 94, 95. moss, 275, 280 Carrageanan, 252, 232, 236
121, 177, 178, 192, 227, 253 scales, 45, 56, 80, 84 Carrier electron transport, 109,
fossil, 208, 209 scars, 45 115-117, 122
nitrogen-fixing, 105, 337 terminal, 180 Carrier hypothesis, 157
photosynthetic, 1 1 trace, 57 Caryopsis, 142, 146, 148, 149, 151,
Banner, 128 Budding, 264 264, 265, 270
Bark, 64, 107, 181, 273 Bulb, 68, 69, 70, 340 Casparian strip, 85, 89, 91 , 94, 96, 109,
Base, 12-16, 376 Bundle, 288
pairs, 13, 14. 43, sheath, 77, 108, 109 Catabolism, 8-12, 16
sequence, 15, 16, 43 vascular, 53, 57-59, 66-67, 73, Catalysis, 7, 16, 377
Basidiocarp, 265, 267, 268, 273 76-78, 140, 160, 317 Catalysts, 8, 11, 16
Basidiomycetes, 253, 260, 265, 273 Catkin, 128, 129, 140, 332, 337
Basidiospore, 266, 270 C3 , 119, 120, 122 Cation, 370
Basidium, 265, 266, 270, 273 C4 , 77, 78, 119, 120, 122 Caulescent, 162
13
Benthic, 226 C, 208, 209 Cavitation, 100
Berry, 144,146 Cacti, 102, 174, 191, 196 CCC, 162
Besseyan system of classification, 219, Calcium, 104, 369 Cell, 1, 98-102, 104, 109, 112, 156,
331-332. 344 Calcium carbonate, 210 160, 162
Betacyanin, 30 Calcium oxalate, 104 cycle, 32-34
Biennal plant, 195 Callose, 54-55, 108, 109, 110, 239, 276 division, 25, 135, 156, 160, 161, 162,
Binomial, 219 Callus, 36, 161, 164 163, 172, 209, 217, 244-252, 253,
Bioassay, 158, 162 Calorie, 192 265, 292, 318
Biological barriers, 205 Calvin, M. 119 eukaryotic, 2, 17, 32, 198, 204, 209,
Biological clock, 170, 171 Calvin Cycle, see C3 219, 220, 234, 273
Biological interactions, 181-183, 193, Calyptra, 279, 286 growth, 156-157
194 Calyx, 123-124, 126, 130, 139, 140, plate, 36, 37
Biosystematics, 205-206, 220 332 polarity of division, 37
Bird's nest fungus, 267 Cambium, 181, 211, 331 prokaryotic, 2, 17, 198, 204, 209,
Bitter Springs Formation, 209 cork, stem, 48, 69, 91, 93, 94 219, 220, 234
Bladders, 232 fascicular, 58, 59, 69 prothallial. 292, 309
Blade, algal, 232, 235 interfascicular, 58, 59, 69 sap, 30
Blooms, algal, 224, 226-227, 252 vascular, root, 91, 93. 94 structure and function, 17^42, 159,
Bog, 184, 188, 312 stem, 48, 58-61, 67, 69, 161, 162, 230
Bolting, 162 172, 338 tube, 131
Bond, Cambrian, 209, 210, 211, 220-221 wall, 1, 25, 30-32, 36, 38, 96,
chemical, 8, 99, 369-370, 372, 377 Camphor, 182 98-100, 104, 106, 107, 109, 131,
covalent, 370, 371, 373 Cap, see Pileus 135, 139, 156, 157, 160, 165, 172,
double, 372 Capillary 99, 193, 203, 204, 207, 231, 232, 233,
hydrogen, 372, 373, 374, 376 action 106 234, 236, 237, 292, 304, 308, 317
ionic, 370 Capsule, 141, 142, 146, 275, 276, 281 wall expansion, 98, 156
Borlaug, G., 120 Carbohydrates, 3, 8, 16, 99, 104, 120, Cellulose, 1, 4, 18, 31, 32, 56, 91, 98,
Boron, 104 156, 164, 192, 265, 271, 224 107, 156, 165, 166, 204, 231, 234,
Botanic gardens, 198-199 conduction, 104, 107-109 236, 253, 255, 260
Bracket fungi, 265, 273 manufacture of, 112, 119, 121 fibrils, 36, 38, 96, 97
Bracts, floral, 124 Carbon, 3, 11, 101-102, 111, 119, 121, Cenozoic, 209, 213-216, 221, 310
Brassicaceae, 141, 332, 334, 344 369-372, 376, 377 Centrales, 237
Bromeliaceae, 167 Carbon dioxide, 3, 8-11, 15, 96, Centromere, 34
Bryophyta, 1, 75, 200-201, 219, 222, 101-102, 107, 109, 121, 122, 208, Charophyta 236
274-287, 323 377 Chaparral, 180, 191
characteristics, 274 in photosynthesis, 111-112, 117-120 Cheese, 271, 273
classification, 198, 204 Carbon fixation, 119, 122 Chemical energy, 113, 114, 121, 156,
economic importance, 275 Carboniferous, 209, 211 157

Subject Index
18
Chemical reactions. 5. 7-16. 374-377 Collenchyma. 52, 53, 57. 77 227. 230, 236. 239. 252. 271
Chemotropism. 254, 276. 281 Colloids. 104. 178, 194 Cycadophyta. 75. 204. 212. 225. 310,

Chernozem. 179 Colonial bodies. 234. 237. 252 324-326

Chiasma. 39 Colony, 222 Cycads, see Cycadophyta
Chilling. 171, 172 Columella. 279 Cycocel, 162
Chitin. 1. 257. 265 Commensalism. 182 Cyme. 130, 140
Chlorenchyma, 51, 70 Community. 173, 184-187. 193, 194 Cystocarp, 246, 248
Chlorine. 105. 369 climax, 187, 194 Cytochrome, 7, 104
Chlorophyll. 104. 113-117. 121. 159. dominants of. 184, 185. 194 Cytokinesis, 33-36. 43
168. 207. 208. 225, 252. 290. 308. fossil. 185 Cytokinin. 157. 163-164. 167. 172
377 measurement of. 185-187 Cytology. 32, 195. 234. 252
break down, 164, 166. 189 pioneer, 186, 187 Cytoplasm. 18. 98. 99. 150. 204, 266,

formation. 155 successional, 185-187, 194 230. 234

kinds. 113-114. 200, 204. 234. 236. Companion cells. 55. 56. 60, 77. 91. Cytosine. 12
237 309. 315. 331.
Chlorophyta. 199. 200, 204. 209, 210. Compensation depth, 225, 226. 239 2,4-D, 167, 168
223. 225. 226. 230. 236. 237. 242. Compensation intensity. 225, 226 Dark reactions. 112, 117-120. 122
244. 252. 272 Competition, 169. 182, 183, 194 Darwin C. 155. 156, 196, 199, 215,
Chloroplasts. 2. 18. 21. 24. 26. 32. 51. Compositae, see Asteraceae 218, 219
69. 75. 84. 112. 121. 237. 239. Compound, chemical. 369 Deciduous, 312
275. 286 Concentration, solute. 373 Decomposers, 182. 192
break down of. 193 Conceptacles. 246 Defoliation, 171

Chlorosis, 104, 105. 121, 193 Cone. 204. 290, 292. 297. 308. 309 Dehiscence, 140
Chromatids, 34, 35, 40, 42, 44 Conidia, 261, 271, 273 Dehydration. 6
Chromatography, 203 Conidiophores. 261, 271 Denaturation. 6
Chromoplasts, 26. 69 Coniferophyta, 198. 202. 204. 212. 214. Deoxyribose nucleic acid, see DNA
Chromosomes. 34, 217. 221. 297. 317 267. 310-326 Desert, 102. 169. 185, 186, 191, 192

deletion, 217 197

classification. 225
Detritus, 224,

duplication, 217 Conifers, see Coniferophyta Development. 103, 123. 155-172. 221
homologous, 38. 39, 42, 43 Conjugation. 261, 265. 269. 270 Devonian, 209. 210-212
inversion, 217 Conk. 267 Diatom, see Bacillariophyta
Chrysolamarin, 236 Connation. 128, 332. 335. 339 Diatomite. 227, 231. 232
Chrysophyceae, 236 Consumers. 182, 192 Dicotyledonae. 155, 198. 204. 219,
Chrysophyta, 204. 227, 236 Continental drift. 218 324. 328
Cineole, 182 Copper. 92. 105 Dictyosome, 17, 21, 22. 27. 54. 254
Cisternae, 27. 32 Cork. 17. 31. 63-64. 84. 101. 109. 211 Differentiation. 47, 57-58. 159.

Citric acid cycle. 10, 11. 16 Cork cambium. 48, 69. 91. 93. 94 160-161. 163. 164, 165, 172
Cladode, 68 Corm. 68, 69, 70, 340 Diffuse porous, 60, 62

Clamp connection. 265, 273 Corolla, 124, 130. 140. 332 Diffuse secondary growth. 66
Class, 198, 219 Cortex, root. 85, 89. 91. 92. 94. 96 Diffusion. 99. 101. 102. 107. 108. 109.

Classification, 1, 140. 195-207, stem, 51, 56, 59. 64. 66. 69. 145. 161
218-219 155, 211.239. 241. 275. 288. 291. Dikaryon, 260, 265, 270. 273
natural, 195, 219 296 see Pyrrhophyta
Dinoflagellates,

phylogenetic, 195, 219 Cotyledon. 146. 148, 149, 150. 154. Dinophyceae. 236
Clay, 177-178 204, 319, 322, 328 Dinosaur. 212
Cleistothecium, 260, 262, 264 Coupled reaction, 1 17 Dioecious. 127, 276
Climatic change. 191. 208, 209, 211, Cover cells, 281. 285 Diploid. 33, 42. 257-273. 244-252.
212, 214. 216, 221 Cretaceous. 209. 211-213 239. 274. 286. 305. 317
Climatic factors, 176. See also Cristae. 23. 32 Disaccharide, 4
Environmental factors Crossing over. 43 Disease, fungal. 253. 255. 257 258.
Climax. 187 Cruciferae, see Brassicaceae 259. 267-270, 271, 273
Club see Basidiomycetes
fungi, Crystals. 25. 31. 32. 373 Dissemination, 135, 150. 166, 206
Coal Age, 206, 211-215. 221 Cucurbitaceae, 144. 336. 339 344 Disulfide bridge. 6, 103
Coalescence, see Connation Cultivation. 107 Division, cell, see Cell, division
Codon, 14-16 Cupressaceae. 312-314 plant. 198, 200-203, 204, 211-213,

Coenocytic, 253, 255. 25^. 273 Curare. 227 234-237. 253, 274. 288. 297. 309.
Coenzyme A. 11 Cuticle. 51, 102, 103. 109. 211 310. 323-324. 330-331
Cofactor. 255 of leaf, 73, 74. 75. 101. 316 DNA, 12-16, 24, 32. 44. 166. 205. 215.

Cohesion, 99, 100, 109 Cutin, 4,30, 31, 51, 102 217
and tranpiration pull theory, 100, 109 Cuttings, 167 bases, 12
Coleoptile. 149, 154, 155, 156, 157, Cyanide. 227 helix. 44
159, 162, 167 Cyanobacteria. see Cyanophyta mutations. 217. 218. 221
Coleorhiza, 149, 154 Cyanophyta, 1, 199, 204. 208. 210. protein synthesis. 217

Subject Index
19
 replication, 42-44 conversion in photosynthesis, 115, leaf, 74, 84, 101, 102, 316, 317
Doctrine of signatures, 275 121 Musci, 257, 258, 264, 268, 270
Dominant, 326 exitation, 114, 115, 117 Phaeophyta, 239
Dormancy, 3, 46, 150, 154, 161, 166, flow in ecosystems, 192 root, 89, 91
169. 171, 172, 206, 258, 265, 305, kinetic, 375 stem, 64, 69
318, 322, 326 light, 113, 114, 122 Epigyny, 128, 130, 332, 335, 339
Double helix, 12, 13 NADPH, 117, 119, 121, 122 Epiphytes, 105, 174-75, 183, 189, 309,
Drought tolerance, 101-103 potential, 375, 376 340
Drupe, 142, 145, 146 Redox, 377 Equilibrium, 375, 376
respiration, 156, 157 Ergot, 265
Earth, age of, 208 Engler, A., 219 Ester, 377
Ecology, 173-194, 234 Entisol, 179 Ethylene, 157, 165-166, 167, 172, 372
paleo-, 185 Environmental factors, 150, 154, Etiolation, 80, 155, 168, 172
Ecosystem, 173, 192-193, 194 173-183, 188, 236, 256 Etioplast, 25, 26
Ecotone, 192-193 affecting growth and development, Euglenophyta, 204, 234, 235-236, 252
Ecotype, 184, 194, 206 80, 155, 172 Eukaryote, 2, 17, 32, 198, 204, 209,
Edaphic factors, 176, 193, 194 dew, 175 219, 220, 234, 273
Egg, 135, 146, 198, 243, 244, 245, 246, fire, 180-181, 187, 193, 194, 322 Eumycotina, 204, 253-273
248, 250, 256, 257, 274, 280, 281 fog, 173, 175 Euphorbiaceae, 197
285, 317, 319, 320, 323, 324, 326, frost, 188, 206 Eutrophication, 225-226, 252
328, 330 gravity, 100, 255, 259 Evaporation, 96, 102, 109
Elater, 286 humidity, 175, 186, 190 Evolution, 2, 173, 195, 207-220
Electron, 11, 104, 114, 115, 116, 117, ice, 187 algae, 252
121, 369, 370, 371 , 372, 374, 376, light, 109, 121, 122, 150, 159, 168, Anthophyta, 209, 212-215, 221,
377 169, 175-177, 184, 186, 189, 193, 331-344
acceptor, 115, 377 194, 224, 225, 226, 242, 255, 259, convergent, 197
affinity, 11, 115-117 260 divergent, 197
115-117, 122, 377
carrier, macro-, 174 mechanism, 215, 217-221
microscope, 18 micro-, 174, 326 monophyletic, 197
transport system, 11, 113, 115, 116, moisture, 99, 102, 109, 150, 168, origin of higher plants, 236, 237
117 169, 186/189, 190, 191, 194,225, polyphyletic, 197
Electronegativity, 372, 373, 377 253-255, 257, 272, 316 vessels, 275
Electrophoresis, 203, 205 nutrition, 260 Excited state, 114, 116
Element, 369 past, 216 Exocarp, 140, 144, 146
Elements essential for plant growth, pollutants, 192-194 Extinction, periods of, 212
103-105. See also Nutrition radiation, 210, 237
Embryo, 136, 139, 220, 281, 286, salinity, 224, 225 Fabaceae, 141, 198, 336, 337, 340,
288-309, 330, 331 soil, 120, 122, 186, 272, 292, 316, 344
angiosperm, 146-150, 162, 163 323 Family, 198, 219
bryophyte, 323 solar radiation, 174, 175-177, Fascicle, 310, 311
gymnosperm, 315, 318-322, 324, 188-194, 210, 237 Fascicular cambium, 58, 59, 69
326 temperature, 6, 97, 102, 119, Fat, 4, 16, 149, 203, 236
lower vascular plants, 288, 290, 120-121, 122, 162, 168, 169, Fatty acid, 4
292-293, 294-295, 297, 298-299, 170-172, 174, 175, 186, 187-188, Feedback, 12, 16
305, 306-307, 309 189, 190, 193, 194, 213-216, 224, Fermentation, 16
sac, 135-137, 139, 146, 328, 330 225, 226, 227, 260, 261, 270 Fern, 184, 198, 202, 211, 214, 316
Endocarp, 140, 144, 146 water, 290, 292, 305, 308, 309, 323, pre-, 211
Endodermis, 85, 89, 91, 94, 96, 105, 324 seed, 211
109, 296, 297 wind, 162, 187, 253, 268, 272, 316 see also Pterophyta
Endoplasmic reticulum, 17, 23, 27, 32, Enzymes, 5-1 6, 43, 1 03, 1 05, 1 08, 1 09, Ferredoxin, 103, 104
162 110, 117, 119, 149, 155, 159, Fertilization, 123, 131, 135, 136, 137,

Endosperm, 36, 136, 139, 146, 147, 162-163, 166, 168, 172, 176, 205, 139, 140, 146, 147, 220, 243,
148, 149, 162, 163, 328, 330 217, 271 244-252, 256-273, 276, 286,
mother cell, 135, 328, 330 activators, 104 288-309, 318-323, 326
primary endosperm cell, 136, 139, amino acid activating, 14 double, 135, 139, 146, 328-331
146, 328, 330, 331 regulatory, 12, 15, 16, 156 tube, 256-258
primary endosperm nucleus, 139, Eocene, 185, 209 Fertilizers, 101, 103, 105, 120, 180,

146 Epicotyl, 149, 150, 155, 319 224, 227

Energy, 16 Epidermal cells, 275 Fiber, 53, 77, 91, 203, 316
activation, 375, 376 Epidermal hairs, 51 , 53, 56, 75, 76, 1 02 phloem, 103, 315
ATP, see ATP Epidermis, 47, 48, 57, 63, 73-76, 96, stem, 52, 56, 57
carrier, 156 102, 148, 279, 290, 291, 297, 304, tracheid, 60, 315
chemical, 9, 10, 111, 113, 114, 119, 308 xylem, 55, 56
120, 121, 153 fruit, 140 Fibrils, cellulose, 36, 38

Subject Index
no
Fig tree formation, 208, 210 Lycophyta, 211, 288 218, 221, 256
Filacales, 297, 302-309 micro, 208-229 mutations, 79, 81, 217, 221
Filament, 222, 223 pollen, 212 recombination, 217, 270
of anther, 124, 130, 139 Psilophyta, 201, 211 Generative cell, 131, 139, 328
Filamentous, body, 198, 204, 209, 234, record, 208-215, 220-221 Genetic information, 12-16
237, 243, 246, 248, 252, 274, 280 seeds, 211, 214 Genetics, 195, 199, 205
Fire, see Environmental factors seed plants, 221 Genotype, 159, 183, 218, 242
Flagella, 1 , 25, 204, 234, 236, 237, 242, Sphanophyta, 297 Genus, 198, 219
246, 252, 253, 257, 273, 276, 288, wood, 212 Geotropism, 101, 155, 158, 159, 165,
297, 324 Fragmentation, 234, 239, 252, 255 172
Flavor,166 Frets, 24, 32 Germination, 150-153, 154, 162, 191
Flemming, A., 271-272 Frond, 232, 235, 297, 303, 306 bud, 161, 164, 167, 171, 177, 189
Flora, 195 Frost hardening, 172 pollen grain, 131, 132, 318, 325
Flower, 75, 123-140, 156, 264, 324, Fructose, 4 spores, 275, 276, 280, 288, 290, 308,
328, 329 Fruit, 123, 136, 140-146, 203, 324, 328, 309, 317 ,

bracts, 124 330, 331 seed, 28, 123, 155, 159, 165, 166,
buds, 137, 169, 170 abscission, 167, 255, 264 167. 168, 170, 172, 180, 227
color, 30 aggregate, 141, 144-145, 146, 196 spores, 253, 258, 260, 268, 270, 273
complete, 125 dehiscence, 140 zygote, 244, 246, 274
development, 124-125, 167 development, 138, 140, 163 Germ tube, 257, 268, 273
essential organs, 123-124 dissemination, 206 Gibberellin, 80, 157, 161-163, 166,
evolution, 124-125, 344 drop, 167 167, 168, 170, 171, 172
imperfect, 125, 126, 128, 142, 146, indehiscent, 149 Gills, of mushroom, 266, 267
196, 332 multiple, 141, 145-146 Ginkgophyta, 197, 204, 310, 324, 326
incomplete, 125, 126, 332 parthenocarpy, 140, 146 Girdle, 231
induction, 206 ripening, 165, 166, 167, 172 Glaciation, 214, 216
inflorescences, 128, 129-131 simple, 141-144 Glands, 144
irregular, 127, 128, 129, 332, 335 types, 140-146 Glucose, 3, 4, 9, 10, 1 1 30, 106, 108,
morphology, 123-129, 203 Frustale, 213 109, 111, 119, 234
parts, number of, 332, 338 Fucoxanthin, 236, 237 Glutamic acid, 234
perfect, 125, 167, 196, 332, 339 Functional group, 376 Glycerol, 4
pistilate, 125, 126, 128, 146, 167, Fungi club, see Basidiomycetes Glycogen, 234
332 egg, 253 Glycolic acid, 120
regular, 127, 129, 332, 339 Imperfecti, 253, 270-272, 273 Glycolysis, 9-11, 16, 163
staminate, 125, 126, 128, 142, 146, rust, 267, 273 Glyoxysome, 25, 28
167, 332 sac, see Ascomycetes Gnetophyta, 204, 310, 324, 326
symmetry, 127-129, 332 smut, 267, 270, 273 Golgi apparatus, see Dictyosome
Flowering, 104, 168-170, 172 true, 253 Grain, see Caryopsis
induction, 206, 220 white rust, 257 Gramineae, see Poaceae
Fluorescence, 114, 115 zygote, 253 Grana, 24, 32, 112-117. See also
Follicle, 141, 142, 146 see also Mycota Thylakoid
Food chain, 192, 194, 226 Fungicide, 257 Grass, 137, 264, 265
Foot, Bryophyta, 276, 280, 285, 286 Funiculus, 149 fruit, see Caryopsis
lower vascular plants, 288, 290, 297, Fusiform initials, 58 Grassland, 179, 180, 187, 190-191,
305, 308, 309 Fusion, 261, 264, 265, 272, 273 192, 194, 227
Forest, conifer, 178, 180, 181, Gravity, 156, 159, 172
188-189, 192, 194 G1, 32-33, 36, 43, 44 Greening, 168
deciduous, 179, 185, 186, 192, 193, G2, 32-33, 36, 43, 44 Green revolution, 120, 162
194 GA, see Gibberellin Ground meristem, 47-48, 56
fossil, 211-215 Gametangia, 198, 222, 236, 242, 243, Ground pine, see Lycophyta
subalpine, 184-185, 188-189 252, 257, 274, 278, 281 , 285, 286, Ground state, 114, 115, 116
tropical rain, 174, 180, 184-185, 288-309, 323 Growth, 101, 103, 104, 105, 109,
189-190, 194, 289 Gamete, 123, 147, 198, 243, 244-252, 155-172
Form taxa, 270 255-273, 274, 323 cell, 156-157, 159, 172
Formula, structural, 370, 371 Gametic life cycle, 243, 244-246, 252 control of, 155-172
Fossils, 2, 132, 185, 203, 207-208, 297, Gametophyte, 242, 244-252, 274-276, secondary, 161, 166
333 281 285, 280, 285, 286, 288-309,
,
regulators, see Hormones
algae, 209, 210, 220, 227-231 323, 324, 326 Guanine, 12, 14
angiosperms, 209 female, 135,317, 318, 319, 325, Guard cells,' 51, 56, 73, 75, 84, 101,
Calamitales, 211-215 328-331 102, 109, 166
ferns, 211, 214 male, 131, 320, 328-331 Gums, 30
fuel, 120, 211 Gemmae, 274 Gunflint formation, 208-209, 210
gymnosperms, 209, 211, 212 cup, 285 Guttation, 100
Lepidodendrales, 211, 310 Gene, 12, 13, 14, 42, 156, 162, 168, Gymnosperms, 209, 211, 212, 219,

Subject Index
111
 221, 310-326. 331 Horsetail, see Sphenophyta K, selection, 206, 220
anatomy, 60 Humus, 1 1 Karyogamy, 262
classification, 198 Hyaline cell, 275, 277 Keel, 128
conifers, see Coniferophyta Hybrid, 167, 184, 210-219, 220-221 Kelp, 222, 223, 224, 226, 232, 235.
Cycads, 212 Hybridization, 197, 210-219, 220-221 236, 237-242, 246, 274
fossil, 209, 211, 212 Hydathodes, 100 Ketone, 3
Ginkgophyta, 212 Hydrocarbon, 208 Kinetic energy, 8
Gynoecium, 124, 128, 130, 134-135, Hydrogen, 3, 9, 10, 11, 16, 119, 369, Kinetochore, 34, 36, 40, 42
139, 336 370. 371, 372, 376, 377 Krebs cycle, 10
bonds, 6, 12, 13, 99, 372-374, 373,
3 376
H, see Tritium Labiatae, see Lamiaceae
Hairs, epidermal, 102 carrier, 11, 112 Lamarck, J. 218, 219
B.,

leaf, 73, 75, 76 ions, 117, 122, 157 Lamella, middle, 232
root. 85. 89, 96, 105 sulfide, 111,208 Lamiaceae. 332, 335. 337. 344
stem, 51, 53. 56 Hydroids, 275, 277, 280 Laminariales, 246
Halophytes. 99 Hydrolysis, 4, 9. 162, 163, 375. 377 Laminann, 236
Haploid, 33. 42, 242, 244-252, Hydronium, 376, 377 Laterite, 180
256-273. 280. 281. 286. 317-328 Hydrophytes. 174 Laurie acid. 4
Hapteron, see Holdfast Hydroxyl, 376, 377 Leaching, 166, 178, 190
Hardening, 169 Hymenium, 260, 267 Lead, 192
Hardwoods, 267 Hypanthium. 128, 129, 144 Leaf. 69-84, 96, 156, 204, 211, 222.
Hatch-Slack pathway, see C4 Hyphae, 106, 254-273 224, 240, 257, 258, 263, 264, 268,
Haustoria, 105, 264, 270. 274 ascogeneous. 260, 261 291. 293, 298, 308, 310. 312-313,
Head (type of inflorescence). 129 receptive, 268, 269 316. 324, 326, 327, 331
Heartwood, 60, 267 Hypocotyl, 149, 155, 319 abscission, 81. 83-84. 167, 172, 191
Heat, 375 hook, 150, 155, 165 airspaces, 76, 101
Helix, DNA. 44 Hypogyny, 128, 130, 332, 339 anatomy, 73-79, 84, 101. 317
Protein, 6 arrangement, 46, 78, 79, 275
Hepaticae. 75. 198. 199, 275, 282-286. axil, 45, 129, 290, 308
Ice Age. 214. .216
288 blade, 70-71, 84
Imbibition, 178
Herbals, 195 buds, 72-73, 84
Indole acetic acid (IAA), see Auxin
Herbaria, 198-199 color, 189
Indusium, 304, 305. 306. 309
Herbicides. 168 compound. 72-73. 84. 198
Inflorescences, 128, 129-131, 139,337
Herbivory, 182-183, 191 destruction, 193
Ingen-housz, J., 111
Heredity, 2, 7, 79, 120, 131, 156. 166, development, 57-58, 77-79, 84. 155,
Inheritance, 166
217-218, 369 159, 162. 165-166, 172, 198
Inhibitors, 150, 154, 162, 167, 182
Hesperidium, 144, 146 fern, 75, 304, 305, 306, 309
Inorganic compounds, 3
Heterobasidiomycetidae, 267-270. function, 69, 77, 84, 101, 102. 106,
Insectivorous plants. 80
273 108, 120, 164, 168, 169, 170, 172
Inseptisol. 179
Heterocyst, 227, 230, 234, 238 gaps, 204, 288-289
Integument, 135, 136, 139, 146, 148,
Heteroecious. 268 gymnosperm, 75
149, 317, 319, 325, 328
Heterogamy, 242, 252 initiation, 77-79
Intercellular space, 51, 63, 64, 76, 96,
Heterospory, 292, 324, 331 juvenile. 80. 81
98, 107, 109
Heterothallic, 265, 268, 276, 285 kelp. 224. 232. 235. 239, 240
Interfascicular cambium, 58, 69
Heterotrichy, 237, 252 lower vascular plants, 288, 289, 290,
Internode, 45, 68. 162, 236
Heterotrophic, 253, 273 291, 293, 297. See also Leaf, fern
Interphase. 33, 40
Hexose, 3 modification, 80-81, 84, 102
Intertidal zone. 224-225. 232
Higher vascular plants, 219. 324 monocotyledonous, 72. 73
Involucre, 124, 142
Hilum, 149 morphogenesis, 77-79, 84
Iodine. 227
Histosol, 179 morphology, 69-73, 79-81 84 .

Ion, 3, 98. 103. 107, 108, 117. 157, 166,

Holdfast, 232, 235, 239 moss. 277
370, 374, 376, 377
Homeostasis, 19 peltate. 71
Ionization,376
Homobasidiomycetidae, 266, 273 petiole. 71, 73. 77. 84
Iridaceae, 340
Homospory, 292, 297, 304. 305, 308, photosynthesis. 69, 77, 111-122
Iron, 11, 104, 121. 180, 192, 369
309 pnmordia, 57, 78-79, 84, 129, 155,
Isogamy, 242, 245. 246, 252
Homothallic, 265, 276. 281 160-161
Isomer, 4
Hormones, 80, 84, 91 . 93, 94, 1 02, 1 04, role in stem differentiation, 57-58
Isotopes. 369
108, 140, 156-158, 164, 172, 189, scars, 45, 49
256 sessile. 71-72

interactions between, 166-167 Jacket of sterile cells, see Sterile jacket shade. 176
mechanism of action, 156 Jungle, 190 sheath, 72, 73, 84
practical use of. 167-168 Jurassic. 209 simple, 72-73. 84
Homworts, see Anthocerotae Juvenility. 80, 81 sun, 176

Subject Index
112
 trace, 57 Psilophyta, 309, 316, 317, 319, 324, 330, 331
venation, 72, 73, 338 325 lycophyta, 308
Leaflet, 72, 305 Pterophyta, 298 pine, 316,319, 320, 324, 325
Legume, 2. See also Pod Selaginella, 292, 297 Pterophyta, 298
Leguminosae, see Fabaceae Megasporocyte, 135, 137, 247, 292, Selaginella, 292
64
Lenticel, 45, 317, 319, 320, 325, 328 Microsporocyte, 130, 132, 248, 316,
Lepidodendrales, 288, 310 Megasporophyll, 298 317, 319, 328
Leptoids, 276, 277, 280 Meiocyte, 242, 248, 250 Microsporophyll, pine, 316
Leucoplasts, 25, 32 Meiosis, 33, 38-^4, 123, 132, 137, 139, Microtubule, 25, 28, 32, 34, 36, 37, 38,
Liana, 189 243, 244, 244-252, 256-273, 274, 43
Lichens, 175, 177, 189, 272-273 279, 288-309, 317, 318, 319 Middle lamella, 36, 43, 84, 104, 232
Lifecycle, see organism in question Meiospores, 40, 265, 273, 242, Midrib, 77
Light, 102, 111-117, 120, 121, 122, 244-252, 275, 288-309 Mildews, 253
175-177 Membrane, 5, 19, 22, 24, 27, 31, downy, 257, 258
absorption, 115, 377 97-100, 103, 104, 109, 113, 115, powdery, 263, 264, 265
by chlorophyll, 1 14 155, 156, 157, 162, 168, 234 Mimicry, 139
reactions, 112, 113-117, 122, 193 Meristems, 47, 56, 104, 286 Minerals, 3, 15, 96, 109, 120, 122, 155,
sensitivity to, 155, 156, 159, 165, apical, 47, 50, 88-89, 129, 130, 139, 254, 255, 272
168-169, 171-172 149, 154-155, 162, 164 absorption, 103-107
trap, 112, 113-115 ground, 47, 56, 89 principle elements, 103-105
see also Environmental factors marginal, 78, 84 Miocene, 209, 216
Lignification, 56 meristoderm, 239, 241, 246 Mississippian, 209, 211
Lignin, 30, 31, 32, 53, 155, 275 primary, 47, 50, 89 Mitochondria, 17, 21, 22, 23, 32, 54,
Ligule, 72, 73 primary thickening, 66 261
Liliaceae, 344 procambium, 47, 48, 56, 57, 58, 59, Mitosis, 33-38, 135, 244-252, 279,
Line transect, 185, 194 69, 79, 89, 93, 94 288, 317. See also Cell, division
Linnaeus, C, 195-96 protoderm, 47 Mitospore, 242, 244-252
Lipid, 4, 8, 11, 16, 113, 162, 234, 237 residual, 57 Modulator, 12, 156
Liverwort, see Hepaticae subapical region, 162 Molarity, 373
Loam, 177 Meristoderm, 239, 241, 246 Molds, 253, 273
Locule, 134, 140 Mesocarp, 140, 144, 146 slime, 1, 107, 204, 253, 254, 273

Lodging, 162 Mesophyll, 73, 76, 84, 96, 101, 102, water, see Saprolegniales
Lower vascular plants, 209, 219, 108, 109, 155, 193, 258, 270, 290, Molecule, 3, 369-377
288-309, 323 308, 316 structure, 370-372
Lumen, 53 Mesophytes, 174 Mollisol, 179-180, 190

Lycophyta, 75, 202, 288, 305, 308, 330 Mesozoic, 209, 212-213, 221 Molybdenum, 105
Metabolism, 3-16, 109, 111, 131, 150, Moniliales, 271-272
Macroenvironment, 174 159, 168-170, 195, 203, 252, 221, Monocots and dicots, compared, 338
Macronutrient elements, 103, 104 222, 225, 234 Monocotyledonae, 198, 204, 219, 328,
Magnesium, 3, 369
104, 113, 121, control of, 11-16 338, 340, 344
Magnoliaceae, 331, 332-333, 335 phases, 8-16 families, 332
338 Metaphase, in meiosis, 39, 40, 42 leaf, 72, 73, 155
Malic acid, 1 19 in mitosis, 34, 35, 43 stems, 64-67, 69
Malvaceae, 332, 334, 344 Metaphloem, 91 Monoecious, 127, 276
Manganese, 104, 105, 117 Metaxylem, 91 Monophyletic, 197
Mannitol, 237, 239 Methane, 371 Morphogenesis, 57-58, 77-79,
Mannose, 234 Methionine, 165 156-157
Marchantiales, 280 Microbody, 17, 25, 28, 32 Morphology, 45-47, 184, 195, 206,
Marsileaceae, 304 Microenvironment, 174 234, 237, 303, 323
Mass flow, 108, 110 Microfibril, 96, 97, 165-166, 237 Mosses, 169, 175, 184, 187, 189, 198,
Mating type, 258, 265 Microgametophyte, 247 272, 316, 323
Meadow, 187 Micronutrient elements, 104, 227 Mucilage canal, 241
Medulla, 239, 241 Microphyll, 211, 276, 288, 289, 305, Muramic acid, 234
Megaphyll, 211, 288 309 Murein, 235-236
Megasametophyte, 292 Micropyle, 135, 146, 149, 317, 322, 328 Musci, 275-280, 288, 316, 323
Megasporangium, 246 Microscope, electron, 18 Mushroom, 265-267, 273
Lycophyta, 308 history, 17 Mutation, 79, 81, 215, 217, 221
pine, 317 Microsporangium, 246, 248, 250, 331 Mutualism, 182-183, 194
Pterophyta, 298 Lycophyta, 308 Mycelium, 255-273
Selaginella, 292, 309 pine, 316, 317, 318 Mycology, 253
Megaspore, 137 Pterophyta, 298 Mycorrhizae, 94, 95, 105, 106, 183, 265
Anthophyta, 328, 330, 331 Selaginella, 292, 309 Mycota, 1. 75, 105, 121, 161, 182, 183,
Lycophyta, 308, 309 Microspore, Anthophyta, 132, 328, 192, 198, 199, 200, 208, 253-273,

Subject Index
113
 265, 274, 288, 290, 309 transport, 107-109 Pericycle, 85, 91, 93, 94, 96
Myxomycotina, 204, 252-254, 273 Osmosis, 97, 99, 100, 101, 102, 109, Periderm, 63-64
110, 156 Pendinin, 237
NAA, 167 Ovary, 124, 128, 134, 135, 136, 137, Pendium, 260
NAD(DPN), 3, 8, 9, 10, 11, 16, 377 139, 140, 141, 142, 144, 145. 146, Perigyny, 129, 130, 335
NADP(TPN), 117, 119, 121, 122 195, 203, 310, 328. 330. 331 Perisperm, 136, 146, 148
Natural selection. 215, 217 inferior,128 Peristome, 279, 280
Needles, 310, 314 superior, 128 Perithecium, 260, 261
Neck, 276, 280, 281, 285 Ovule, angiosperm, 124, 135, 137, Permafrost, 188, 189
Nectar, 137, 139 139-140, 144-148, 328, 330 Permeability, 12, 96, 97, 104, 156
Net radiation, 175, 176. See also, gymnosperm, 317, 318, 320, 322, Permian, 209, 212
Environmental factors; Solar 325 Peronosporales, 255
radiation Oxalic acid, 104 Peroxisome, 25, 28
Neritic, 225 Oxaloacetic acid, 11, 119 Petals, 123, 127, 136, 139. 145, 196
Neutralism. 182 Oxidation, 16, 377 color. 30
Neutron, 369, 370 Oxidation-reduction potential, 116, 117 Petiole. 53. 71, 73. 77, 84, 108. 161,
Niche, 182, 193 Oxisol. 179-180, 190 167, 309
Nicotinamide adenine dinucleotide, Oxygen. 3, 10, 11, 16, 96, 111. 112, pH. 178, 179, 180. 194. 376
see NADP or NAD 117, 121, 122, 150. 208, 209, 211, Phaeophyta, 199, 200, 204, 210, 226,
Nitrate, 103 224, 369-372, 376. 377 227, 232, 236. 237-242. 244,
reduction, 105 absorption, 106-107, 109, 150 246-252
Nitrogen, 3, 103, 121, 215, 370-372. soil. 106-107 Phenotype, 218, 221
377 Ozone, 192, 194 Phloem, 56, 57, 60, 64, 66, 77, 90, 91,
deficiency, 103 94, 160, 161, 210, 288, 293, 309,
fixation, 94, 105, 224. 227, 234, 252, Paleocene, 209 315, 331
337 Paleoecology, 185 conduction, 107-109, 110
storage, 5 Paleozoic, 209-212, 220-221 differentiation, 156
Node, 45, 68, 69, 86, 94, 236, 300, 309 PAN, 192, 194 primary, 51 , 53-55, 69
Nodules, bacterial, 2, 94, 95, 105 Panicle, 129, 140 sap, 108
Nucellus, 135, 136, 139, 146, 317, 318. Paramylon. 236, 237. 250 secondary, 59, 63, 93
319, 325, 326, 330, 331 Paraphyses. 250, 260, 276, 278 translocation, 157, 169
Nuclear pore, 25 Parasexual cycle. 272 Phosphate. 8, 10, 11, 12. 103. 104.
Nucleic acid, 16, 103. 162-164 Parasitism, 85, 93, 105, 182-183, 199, 117 376
Nucleolus, 25 253, 255, 257, 260, 264, 265, 273. Phosphoenolpyruvic acid (PEP), 199
Nucleoplasm, 25, 32. 34, 35, 42 288, 323, 325 carboxylase, 119
Nucleotide, 8, 12, 13 Parenchyma, 51-53, 55. 56. 57. 58, 59, Phosphoglyceric acid (PGA), 119
Nucleus, 17,19,21, 22, 25, 32, 54, 55, 66, 77, 78, 85, 91, 156, 160, 161, Phospholipid, 5, 103
255-273
156, 198, 209, 239. 264. 274, 288, 291. 296, 309 Phosphorus. 3, 103, 104, 119. 227.
atomic. 369-374, 376, 377 axial. 315 369, 372, 376
Nut, 142, 145. 146, 315 border, 77. 108, 109 Phosphorylation, cyclic. 117
Nutrition, 103-105, 159, 178, 190, 210, palisade, 76, 84, 309 noncyclic, 1 17
225, 252. 260. 309, 323 ray. 315 in photosynthesis, 1 17
spongy, 76, 84, 309 in respiration, 10
Oil. 182. 203. 236. 237 xylem, 56 Phosphoryl group, 5
Oligocene, 209, 216 Parthenocarpy, 140, 146 Photon, 114, 115, 117. 121
Oogamy, 243, 245 Pathogenic, 227 Photoperiodism, 168, 169, 170, 171,
Oogonium, 243, 248, 256, 257 Peat, 105, 275 172, 188, 189
Oomycetes. 253, 255-258, 261, 273 Pectates. see Pectin Photophosphorylation, 117
Oospore. 256, 257 Pectin, 32. 36, 53. 85. 104. 156. 236. Photorespiration, 120
Operculum of moss sporangium, 279, 237 Photosynthesis, 1, 8, 16, 69. 96. 99,
280 Pedicel, 129, 140 101, 102, 103, 104, 105, 106, 109,
Operon mechanism, 15. 16, 156 Peltate. 71 111-122, 154, 156, 166. 169, 184.
Orbitals, 114, 115, 370-373. 377 Penicillin. 253, 271, 273 188, 193, 207. 208, 224, 225, 230,
excited state, 114, 116 Pennales, 287 252, 273, 297. 308. 323. 324, 377
ground state, 114, 115, 116 Pennsylvanian, 209, 211 C3 cycle, 119, 120, 122
Orchidaceae, 198, 332. 340 Pentose, 4 C4 cycle, 119, 120, 122
Order, 198, 219 Pepo, 144, 146 chlorophyll, 104, 113-117, 121,
Ordovician, 210 Peptide, 227 377
Organ. 45. 47 bond, 5, 15 conditions affecting, 120-122
Organelles, 12, 16, 17, 23-29, 31, 54, Perianth, 123, 124, 125, 126, 128, 129, development, 154, 155
55, 112, 121, 150 130, 137, 142. 339 discovery, 111-112
Organic acid, 30, 183, 232 Pericarp, 140, 142, 146, 149 efficiency, 114, 120, 122
Organic substances, 3-6, 110 Periclinal, 36 enzymes, 1 19

Subj ect Index

114
 1 ,

food production, 120-22 Plantae, 198 Predation/parasitism, 194

general equation, 111, 112 Plasmalemma, 19, 27, 30, 31, 53. 89. Pressure, atmospheric, 99, 156
history, 111-112 96, 98, 100, 102, 105, 106, 166, root, 100, 109
photosynthetic bacteria, 1 1 237, 254 turgor, 97-99, 102, 108, 109, 110,
pigments, 112, 113, 114, 115, 198 Plasma membrane, see Plasmalemma 156, 157, 254
plastids,see Chloroplasts Plasmodesmata, 23, 31 32, 54, 55, 96,
, wall, 98
products, 111-112, 117-120 98, 107, 230, 276 Prickles, 69
sugars, 117, 119, 120, 121, 122 Plasmodium, 253, 255 Priestley, J., 111
Photosystem, I, II, 115, 116, 117 Plasmolysis, 99 Primary plant body, 47
Phototropism, 158, 159, 172 Plastid, 17, 24-25, 26, 54, 112, 155, Primary thickening meristem, 66
Phycobilin, 114, 225,236 159, 254 Primordia, bud, 50
Phycocyanin, 114, 236 Plastocyanin, 105 leaf, 78, 79, 84, 129, 155, 161
Phycoerythrin, 114, 236 Pleistocene. 209, 214, 216 Procambium root, 89, 93, 94
Phylogeny, 195, 219, 236, 237, 276 Pliocene, 209, 216 stem, 47, 48, 56, 57, 58, 59, 69, 79,
Phylum, see Division Plumule, 165 160, 161
Phytochrome, 155, 165. 168, 169, 170, Poaceae, 142, 198, 340, 344 Producers, 182, 192, 224-226
172 Pod, 105, 125, 138, 141, 146, 148-149, Productivity, 190, 192, 225-226
Phytoplankton, 225-229. 231, 236, 237 198 Proembryo, flowering plant, 147, 328.
Picograms, 32 Podzol, 179, 194 330
Pigments, 112, 113, 114, 115, 155, Polarity, cell, 36, 37 pine, 319, 320
159, 166. 168, 172, 198, 200. 204 chemical, 4, 5, 6, 372, 373, 374, 377 Prokaryotes, 2, 17, 198, 204, 209, 219,
accessory, 114, 115, 236 Polar nuclei, 135, 146, 328 220, 234
Pileus, 266, 267 Polar transport, 157, 158, 159, 167 Prophase, meiosis, 39, 40, 42
Pinaceae 310-311, 313, 315-326 Pollen, 130, 132, 133, 135, 136-140, mitosis, 33-34, 35
Pioneer species, 272, 273 207, 244, 317, 328, 329, 330 Proplastid, 25, 26, 32
Pistil, 124, 125, 130, 131, 134-135, fossil, 132, 185, 212 Protease, 12
139, 140, 195, 270 germination, 131, 132, 135, 318, 325 Protein, 2, 5, 1 1, 103, 131, 149, 155,
Pit, 55, 60 pine, 317-318, 319, 322 156, 162, 164, 192,205,217, 232.
bordered, 55, 56 tube, 131, 135, 139, 318, 323, 325, 234, 255
simple, 53, 55 326, 331 P-, 54
Pith, 51 , 56, 59, 94, 1 45, 1 63, 1 64, 21 1 Pollen mother cells, see structure, 5, 6, 7
290, 331 Microsporocyte synthesis, 12-16, 32, 104. 162, 164,
ray, 53, 56. 59 Pollen sac, 130 193
Placenta, 135, 141 Pollination, 123, 131, 136-137, Proterozoic, 209, 220
Placentation, 135, 139 139-140, 169, 310, 317-318, 319, Prothallia, of ferns, 304-309
axile, 135 323, 324, 325, 330 of Psilotum, 292
central, 135 animal, 137, 138-139, 332, 339 Protochlorophyll, 155, 168, 172
parietal, 135 cross, 139, 140, 183, 220 Protocooperation, 148, 182-183
Plankton, 192, 213. See also self-,139, 140 Protoderm, root, 89
Phytoplankton vectors, 130 stem, 47. 56
Plant, annual, 58, 170, 175, 191, 195, wind, 135, 137 Proton, 369-372, 376, 377
337 Pollution, 120, 165, 192-194, 225-226, Protonema, 275, 276, 280
aquatic, 297, 304, 309 236, 252 Protoplasm, 98, 104, 150, 253, 255,
biennial, 170, 195 Polymer, 5, 12, 13, 98, 99, 157, 166 257, 258
classification, 195 Polymerization, 4 streaming, 107, 108
community, 184-187 Polynomial, 195 Protoplast, 19,31,96, 98, 99, 105, 106,
day-neutral, 169, 172 Polypeptide, 5, 6 156, 160, 166, 225, 243
diseases, 253, 255, 257-259, Polyphyletic, 197 Protoxylem, 9
267-271, 273 Polypodiaceae, 297, 301-308 Pseudoplasmodium, 253, 254
distribution, 195 Polyribosome, 23, 32 Psilophyta, 204, 210, 288, 305, 330
herbaceous, 338 Polysaccharide, 4 Psilotaceae, 288
insectivorous, 105 Polysome, 23, 32 Pteridospermophyta, 204
long-day, 169, 171, 172 Pome, 144 Pterophyta, 75, 202, 204, 288, 297,
lower vascular, 209, 219, 288-309, Population, 173, 183-184, 193, 197, 302-309, 316, 330
323 205-207, 217, 221 Puffballs, 265, 267
perennial, 45, 58, 171, 337 Pore fungus, 267 Purine, 13
pitcher, 105 Porphyrin, 207 Pyrimidine, 13
populations. 183-184 Potassium, 3, 102, 104, 109, 166, 369, Pyrrophyta, 204, 225, 227-228, 236,
seeds, 219, 277 370 252
short-day, 169, 170, 171, 172 Prairie, 187, 190-191. See also Pyruvate, pyruvic acid, 10, 11, 16
succulent, 80, 102, 191 Grassland
vascular, 198, 201, 209, 211, 219, 219
Prantl, K., Quadrat, 185-186, 194
222, 239 Precambnan, 209 Quantum, light, 114, 115

Subject Index
115
Quaternary, 209 240, 265, 288, 297, 305, 308, 309 coat, 136, 141, 146, 148, 149, 154,
Quiescent center, 88 adventitious, 86, 88, 154, 164 182, 217, 319, 320, 323, 326, 331
anatomy, 85, 88-95 Coniferophyta, 315, 320, 322, 323
r, selection, 206, 220 apical meristem, 88-89 development, 123, 144, 146-154,
Race, 220 cap, 88, 90, 159 169, 172, 217, 268, 328, 330, 331
Raceme, 129, 140 competition, 186 dissemination, 150, 166, 186
Radicle, 86, 88, 149, 150, 319 contractile, 95 dormancy, 150
Radioactive decay, 369 cork, 91, 93, 94 endosperm, 146-148
Ranales, 331, 344 cortex, 85, 89, 92, 94, 96 food storage, 255
Ranunculaceae, 331, 332, 333 crop, 214 fossil, 214, 221
Raphe, 149 cuttings, 167-168 germination, 150-153, 154, 155, 159,
Ray, initials, 60 differentiation, 86, 88 162, 165, 166, 167, 170, 172, 180,
pith, 53, 56, 59 external morphology, 85-88 191, 227
vascular, 58, 60-62, 315-316 functions, 84-95, 96, 99, 100, 101, plants, 198, 211, 219, 221. See also
Reaction centers, 115, 116 106, 107 Angiosperms; Gymnosperms
Recent, 209 growth, 155, 159, 163, 167, 177 Seedling, 155, 156, 159, 161, 165, 168,
Receptacle, 123, 124, 128, 136, 141, hairs, 85, 88, 89, 94, 96, 105 172, 176, 182, 186, 320, 322
142, 145, 146 lateral, 86, 91, 92, 94 by smuts, 270
infection
Recombination, 43, 270, 218, 272 meristem, 89 Senescence, 164, 172
Red tides, 226-227, 237, 252 nodules, see Nodules Sepals, 123, 126, 136, 139, 140, 145,
Reduction, 377 origin, 86, 88 196
Reef, 224, 234 pressure, 100 Septum, 255
Reproduction, asexual (vegetative), 66, primary, 86 Sere, 187
68, 69, 81, 88, 167, 184, 239, 242, prop, 86, 88 Sessile, 71-72, 129
252, 255, 258, 261 , 264, 265, 268, quiescent center, 88 Seta, 277, 280, 285, 286
270, 290 secondary, 86 Shade tolerance, 176
fragmentation, 234 239, 274, 285 seminal, 86 Sheath, 77, 78
sexual, 123-136, 146-148, 214, systems, 85-86, 94, 174, 179, 191 bundle, 77, 78
242-252, 256, 259, 264, 265, 268, vascular cambium, 91, 93, 94 leaf, 72, 73, 84
270, 272, 273, 285, 290 Rooting,164/167-168 Shoot, 155, 158, 160, 161, 163, 164,
Reproductive isolation, 218, 220, 221 Rosaceae, 144, 336, 344 165, 168, 169, 170, 171, 172, 232,
Resin, 60, 315 Rosales, 198 288, 290. See also Stem.
ducts, 60, 63, 315 Rosette plant, 45, 46, 152 Sieve, cell, 55, 91 , 252, 274, 309, 315,
Resonance transfer, 119 Runners, 69 323, 331
Respiration, 9-11, 16, 32, 104, 106, plate, 54, 108, 109, 156, 239, 241
107, 121, 156, 157, 164, 192, 193 Salicaceae, 332, 335, 344 tube, 53-55, 56, 57, 77, 107, 108,
Rhizoids, Bryophyta, 274, 275, 280, Salinity, 224, 225 109, 110
285, 286 Salviniaceae, 304 tube member, 53-55, 57, 91, 239,
lower vascular plants, 259, 288, 304, Samara, 142, 146 241, 242, 331
308 Sand, 177-178 Silica, 204, 207, 208, 231, 237, 243,
Rhizomes, 66, 68, 69, 70, 159, 187, Saprolegniales, 255-257 297
211, 340 Saprophyte, 182, 192, 253, 260, 265, Silique, 141, 142, 146
Equisetum, 297 273, 340 Silt, 177-178
Psilotum, 289, 291 305 , Sapwood, 60, 267 Silurian, 209, 210-211
Rhodophyta, 200, 204, 225, 226, 227, Savannah, 174, 180, 190-191 Smog, see Environmental factors,
230, 231 , 234, 236, 244, 246, 249, Scale, 281, 310, 317, 318, 322, 325 pollutants
252 Scar, bud scale, 45 Sodium, 166, 369
Ribonucleic acid, see RNA bundle, 48 Soil, 3, 101, 103, 104, 155, 165, 168,
Ribose, 4, 13 leaf, 45 177-180, 193, 224, 227, 237, 292,
Ribosome, 13, 14, 15, 16, 17, 20, 22, Schizocarp, 142 316, 323
23, 24, 32, 104, 162, 237 Sclereids, 52, 53, 56, 57 air, 106-107
Ribulose disphosphate, 119 Sclerenchyma, 53, 57, 102, 296, 297 cation exchange, 178
Ring porous, 60 Scrub, 194 colloids,178
Ripening, 166, 167, 172 chaparral, 191 formation, 177-178
RNA, 13, 14, 16 desert, 185-186, 191, 192 horizons, 178, 179
-ase, 20 Scutellum, 143, 149, 154 leaching, 178
m-, 13, 14, 15, 16 Seaweed, 223, 226, 227, 231, 236, loam, 177
polymerase, 13, 15, 16 244, 246. See also Kelp mineral, 177
ribosomal, 13 Secondary plant body, 58-66 moisture, 186
synthesis, 13, 15, 16, 164 Secretion, 157, 162, 271 orders, 179, 194
t-, 14, 15, 16 Secretory cells, 53 organic matter, 177
Root, 84-95, 96, 98, 100, 101, 105, Seed, 135, 138, 139, 146, 198, 203, parent material, 177
109, 156, 159, 204, 211, 222, 224, 204, 206, 215, 305, 324, 328 peat, 105

Subject Index
116
 1

pH, 178, 179, 180, 194 Starch, 4, 54-55, 119, 149, 158, 166, Substomatal chamber, 51, 76
podzol, 194 205, 231, 236, 237, 255, 271 Substrate, 8
pore spaces, 178 floridean, 236 Succession, 186-187, 194, 225-226
profile, 178-180, 194 grains, 24 Succulence, 80, 102, 191
salinity, 224 hydrolysis, 162, 163 Sucrose, 4, 30, 108, 109, 110, 119,
solution, 96, 99, 100, 101, 105, 107 photosynthesis, 1 1 161, 369. See also Sugar
temperature, 174 Statolith, 159 Sugar, 3, 8, 9, 10, 11, 16, 97, 107, 117,
texture, 177-178 Stele, 91. See also Tissue, vascular 119, 120, 121, 122, 166, 183, 203,
types, 177, 194 Stem, 204, 222, 224, 240, 257, 269, 217, 234, 236, 254, 255, 257, 276,
water, 109 270, 275, 324, 325, 326, 331 369, 377
see also Environmental factors anatomy, 47-58 formation, 117-120
Solanaceae, 334, 336, 344 apex, 77-79, 157 Sulfate, 103, 370
Solar radiation, 174, 175-177, 188 Bryophyta, 275 Sulfur, 3, 103, 111, 192, 194, 264,369,
Solubility, 5, 372-374 Coniferophyta, 315-316 373
Solutes, 96, 98-1 00, 1 03-1 06, 1 07, 1 08, development, 160, 161, 164, 172 Suspensor, 147
109, 110, 156, 157, 273, 372-374 functions, 45,169 of Lycophyta, 309
Solution, 372-374 growth, 157, 159, 166 of pine, 319
Solvent, 373-374 kelp, 224, 232, 233, 235, 237, 239, of pterophyta, 298
Soredia, 273 240, 241, 242 of Selaginella, 292
Sorus, 269, 246 Lycophyta, 290, 291, 296, 308 of Sphenophyta, 297
of fern, 304, 305, 309 modification, 66, 68-70 Symbiosis, 2, 105, 272, 273, 288
Spathe, 128 monocotyledon, 64-67 Symmetry of flower, 127-129
Species, 1, 194, 197-198, 199, morphology, 45-47 Syncarpy, 332. 339
205-207, 217-218, 219, 220, 221 primary growth, 47-57 Synergid cells, 135, 137, 146, 328
number of, 196, 222, 236, 253, 270, Psilophyta, 288, 289
252, 310, 324, 326 Pterophyta, 304 2,4,5-T, 168
pioneer, 272 secondary growth, 58-66 Taiga, 188-189, 194
subspecies, 220 Sphenophyta, 297, 300, 301 Tannins, 30
Sperm, 131, 135, 139, 146, 198, 243, storage in, 45 Tapetum, 316-317
244, 245, 248, 250, 274, 276, 280, Steppe, 190-191. See also Grassland Taxaceae, 324
285, 288-309, 318, 319, 324, 325, Stereid, 275, 277 Taxodiaceae, 206, 313, 324
326, 328, 330, 331 Sterigma, 265, 266 Taxonomy, 131, 195-202, 218-219
Spermagonia, 269 Sterile jacket, 274, 276, 281, 285 anatomical, 203
Spermatia, 260, 268, 270, 246, 248 Stigma, 124, 125, 131, 134, 135, 139, biochemical, 203, 204
Sphenophyta, 75, 201, 204, 211, 288, 140, 220, 317, 328, 330 biological, 205-206
297, 305, 309, 330 fluid, 135 morphological, 140
Spike, 129, 139 Stipe, 232, 234, 239, 240, 266 numerical, 206-207, 220
Spindle, 25, 34, 35, 43 Stipule, 71, 80 Teliospores, 269, 270
Spines, 69, 80, 84, 197 Stolons, 69, 159, 259 Telophase, in meiosis, 39, 40, 42
Spodosol, 179, 188 of Rhizopus, 259 in mitosis, 35, 36, 43

Sporangiophore, 258, 259, 260, 297, Stomata, 51, 75, 96, 100, 101, 102, Temperature, see Environmental
300, 309 104, 109, 211, 258, 270, 279, 286, factors
Sporangium, 253-273, 211, 242, 275, 290, 308, 316 Tendrils, 68, 80, 84, 337
276, 280, 285, 286, 288-309 closure, 174 Tepals, 124
Spore, 207, 209, 253-273, 244-252, distribution, 73, 75 Tertiary, 209
274, 279, 285, 288-309 leaf, 73, 84 Tetraspores, 246
germination, 260, 270, 273, 275, 276, mosses, 75, 279 Tetrasporophyte, 246
280, 288, 290, 317 movement, 101-102, 166, 172 Thallophytes, 198, 219, 224, 274, 281,
nesting, 242 stem, 64 323, 324, 326
see also Reproduction; or plant in sunken, 76, 102, 103 Thallus, 275, 280, 285, 286
question transpiration, 101 Thermoperiod, 175
Sporic life cycle, 246-252 Stone cells, 52, 144 Thorns, 68-69, 191
Sporocyte, 279, 280, 281, 285 Stoneworts, see Charophyta Thylakoid, 24, 1 12, 1 13, 1 15, 122, 155,
Sporophyll, 290, 291-292, 293, 294, Stratification, 171 230
309 Strobilus, Coniferophyta, 320, 322 energy storage by, 117
Sporophyte, 239, 244-252, 274-286, Cycadophyta, 324 reactions of, 112
288-309, 319, 320, 323, 324, 326, Lycophyta, 290, 297 Thymidine, 44
330 Pterophyta, 298 Thymine, 12, 13
Sporopollenin, 131 Sphenophyta. 297, 300 Type specimen, 199
Spur, 310 see also Cone Tiller, 159
Stamen, 123, 124, 125, 128, 129, Stroma, 24, 32, 112, 1 13, 1 17, 122 Timberline, 184. 187-188, 206, 218
130-133, 139, 140, 145, 195, 198, Style, 124, 134, 135, 139, 140, 328, 330 Tissue, 18, 47, 222
328, 330, 332 Suberin, 4, 30, 31, 32, 63 cambium, 48, 338

Subject Index
117
 collenchyma, 77 Tyloses, 60, 63, 84 role in plant, 96, 168
cork, 63 storage, 102
Ultisol, 179, 189
cortical, 85, 89 stress, 166, 172
Ultraviolet radiation, 210, 237 transport, 96-101
culture, 161, 163, 164, 167
Umbel, 129, 140
endodermal, 89 see also, Environmental factors
Umbelliferae, see Apiaceae
epidermal, 47, 88 Waxes, 4, 217
Unicell, 222-223, 234, 237, 242, 244,
parenchyma, 78, 288 Weathering, 177, 194
252 Weed,
phloem, 51, 288 101, 182, 183, 215
Unisexual, 327, 328. See also Flower,
Whorled leaves, 46
primary, 48-57, 89-91
and pistillate staminate Wilting, 98, 102, 166
sclerenchyma, 53
Units of measurement, 378 Wind, 166
secondary, 48
Uracil, 13, 14
sporogenous, 279, 281, 286 dispersal agent, 137, 162
Uredinia, 270 see also Environmental factors
vascular. 53-57, 89, 96, 155, 161,
Uredospores, 269, 270 Wood, 255, 265
198, 204, 224, 288, 296, 324, 338
Uridine, 13
Tonoplast, 19, 31, 32 anatomy, 62
Torus, 55 diffuse porous, 60, 62
Vacuole, 18, 19, 21, 30-32, 99, 156,
Toxin, 182, 271, 227 early, 60
234, 254
TPN, see NADP Vascular bundle, see Bundle
heart, 60

Traces, 57 late, 60
Vascular cambium, see Cambium
Tracheids, 55-57, 77, 91 96, , 1 00, 208, petrified, 208, 211-212
Vascular plants, see Plant
274, 288, 317, 323 ring porous, 60
Vascular rays, 58
Transcription, 13, 14, 15 sap, 60
Vascular system, 76-77, 155. See also
Transfer cells, 53 spring, 60
Tissue
Transfer reaction, 374, 375 summer, 60
Vegetation, world types, 187-192, 194,
Translation, 13, 16 Wound healing, 160, 161
216
Translocation, 107-109, 110, 157, 239,
Veins, 73, 76, 78, 84, 108. 305, 316
252, 276 Xanthophylls, 204
Venation, 72, 73, 338
Transpiration, 69, 96, 99, 100, 102,
285 Xanthophyta, 236
Venter, 276, 280, 281,
109, 120, 175, 191
Vernalization, 170, 172
Xerophytes, 102, 174
and absorption, 102 Vertisol, 179 Xylem, 100, 102, 107, 108, 109, 155,
conditions affecting rate, 101-102 157, 159, 164,210,239,252,276,
Vessels, 56, 57, 77, 96, 100, 309
cuticular, 102 288, 308, 309, 315, 317, 323, 324,
elements, 55-57, 91, 205
osmotic factors, 96-102 331
evolution, 203
pull, 99-100 differentiation, 160, 161
lower vascular plants and
rate, 96, 1 74 leaf, 77
gymnosperms, 291, 315, 324
regulation by guard cells, 101-102, lower vascular plants, 288, 289-290,
Vines, 189, 297
166 291, 293, 297
Vitamin, 164, 255
stomatal, 101-102 primary, 51, 55-56, 69
Vitamin Bi, 255
Transport, 96-110 ray, 315
active, 101, 102, 106, 157 Wall, cell, pressure, 98 root, 85, 93. 94, 96
polar, 157, 158, 159, 167 primary, 56 sap, 96, 100
Triassic, 209, 212 secondary patterns, 56, 57, 96, 160 secondary, 59, 66, 316, 324
Trichogyne, 246, 248 see also Cell, wall Xylose, 234
Trichomes, 76 Water, 3, 9, 10, 1 1, 15, 96-102,
Triose, 4 121-122, 155, 371, 373 Yeast, 11, 163, 253, 265, 273
phosphate, 119 absorption, 96-101, 109, 166
Tropical rain forest, see Forest adhesion, 99, 100 Zeatin, 157
Tropism, 159 bound, 99 Zinc, 104, 152
Tryptophan, 157 cohesion, 99-100, 373 Zoosporangium, 256
Tube cell, 131,328 loss, 102 Zoospores, 255, 256, 258, 246, 242
Tuber, 68-70, 169, 257 movement, mechanism, 99-100, Zygomycetes, 253, 260, 261 273 ,

Tumor, 270 101-102, 109 Zygospore, 257, 258

Tundra, 187, 194 photosynthesis, 111, 112, 176 Zygote. 135, 136, 139, 147, 156, 220,
alpine, 184, 188 potential, 96-101, 102, 108, 109 232, 243, 253, 244-252, 255-273,
arctic, 184, 187-188, 192, 272 quantity transpired, 96 274, 276, 280, 285, 288-309, 319,
Turgor pressure, 97-99, 102, 109, 156, reactions, 374, 375, 376, 377 323, 326, 328, 330. 331
157, 166, 254 respiration, 9-1 Zygotic life cycle, 244, 252

Subject Index
118
ISBN 0-471-02114-8