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 FPO Co n t e n t s i n br i ef

CHAPTER 16 Bryophytes 366

CHAPTER 17 Seedless Vascular Plants 391
CHAPTER 18 Gymnosperms 430
CHAPTER 19 Introduction to the Angiosperms 457
CHAPTER 20 Evolution of the Angiosperms 477
CHAPTER 21 Plants and People 501
Preface xiii
The AnGIOSperm Plant Body:
INTRODUCTION 1 SECTION 5 Structure and Development 525
CHAPTER 1 Botany: An Introduction 2 CHAPTER 22 Early Development of the
Plant Body 526
SECTION 1 Biology of the Plant Cell 17 CHAPTER 23 Cells and Tissues of the
Plant Body 538
CHAPTER 2 The Molecular Composition
of Plant Cells 18 CHAPTER 24 The Root: Structure and
Development 558
CHAPTER 3 The Plant Cell and the Cell Cycle 38
CHAPTER 25 The Shoot: Primary Structure and
CHAPTER 4 The Movement of Substances Development 579
into and out of Cells 75
CHAPTER 26 Secondary Growth in Stems 614
SECTION 2 Energetics 93
CHAPTER 5 The Flow of Energy 94 SECTION 6
Physiology of Seed Plants 637
CHAPTER 6 Respiration 107 CHAPTER 27 Regulating Growth and Development:
The Plant Hormones 638
CHAPTER 7 Photosynthesis, Light, and Life 122
CHAPTER 28 External Factors and
Plant Growth 660
SECTION 33 Genetics and Evolution
SECTION 151
CHAPTER 29 Plant Nutrition and Soils 683
CHAPTER 8 Sexual Reproduction and Heredity 152
CHAPTER 30 The Movement of Water and
CHAPTER 9 The Chemistry of Heredity and Solutes in Plants 708
Gene Expression 174
CHAPTER 10 Recombinant DNA Technology, Plant SECTION 7 ECOLOGY

Biotechnology, and Genomics 192 On the Web: www.whfreeman.com/raven8e
CHAPTER 11 The Process of Evolution 209 CHAPTER 31 The Dynamics of Communities
and Ecosystems
SECTION 4 Diversity 233 CHAPTER 32 Global Ecology
CHAPTER 12 Systematics: The Science of
Biological Diversity 234 Appendix: Classification of Organisms A-1
CHAPTER 13 Prokaryotes and Viruses 256 Suggestions for Further Reading FR-1

CHAPTER 14 Fungi 278 Glossary G-1

Illustration Credits IC-1
CHAPTER 15 Protists: Algae and Heterotrophic
Protists 317 Index I-1
vi
 Co n t e n t s
Preface xiii Vacuoles 50
INTRODUCTION 1 Endoplasmic Reticulum 51
Golgi Apparatus 52
1 Botany: An Introduction 2
Cytoskeleton 54
Evolution of Plants 3
ESSAY: Cytoplasmic Streaming in
Evolution of Communities 9
Giant Algal Cells 56
Appearance of Human Beings 10
Flagella and Cilia 56
Cell Wall 56
The Cell Cycle 62
Interphase 64
Mitosis and Cytokinesis 65

4 The Movement of Substances

into and out of Cells 75
Principles of Water Movement 76
Cells and Diffusion 78
Osmosis and Living Organisms 79
ESSAY: Imbibition 80
Structure of Cellular Membranes 82
Transport of Solutes across Membranes 83
ESSAY: Patch-Clamp Recording in
SECTION 1 BIOLOGY OF THE PLANT CELL the Study of Ion Channels 84
17
Vesicle-Mediated Transport 86
2 The Molecular Composition of Plant Cells 18
Cell-to-Cell Communication 87
Organic Molecules 19
Carbohydrates 19
Lipids 22
Proteins 25
ESSAY: Vegetarians, Amino Acids,
and Nitrogen 25
Nucleic Acids 29
Secondary Metabolites 30

3 The Plant Cell and the Cell Cycle 38

Prokaryotes and Eukaryotes 39
ESSAY: Cell Theory versus Organismal
Theory 40
The Plant Cell: An Overview 42
Nucleus 42
Chloroplasts and Other Plastids 45 SECTION 2 ENERGETICS 93
Mitochondria 48 5 The Flow of Energy 94
Peroxisomes 49 The Laws of Thermodynamics 95
vii
viii   C ont e nts

Oxidation–Reduction 98 The Process of Meiosis 155

Enzymes 99 How Characteristics Are Inherited 159
Cofactors in Enzyme Action 101 Mendel’s Two Principles 162
Metabolic Pathways 102 Linkage 163
Regulation of Enzyme Activity 103 Mutations 164
The Energy Factor: ATP 104 Broadening the Concept of the Gene 166
Asexual Reproduction: An Alternative
6 Respiration 107 Strategy 169
An Overview of Glucose Oxidation 107 Advantages and Disadvantages of
Glycolysis 108 Asexual and Sexual Reprodution 170
The Aerobic Pathway 110 ESSAY: Vegetative Reproduction:
ESSAY: Bioluminescence 117 Some Ways and Means 171
Other Substrates for Respiration 117
Anaerobic Pathways 118 9 The Chemistry of Heredity and
ESSAY: The Botany of Beer 119 Gene Expression 174
The Strategy of Energy Metabolism 119 The Structure of DNA 174
DNA Replication 176
7 Photosynthesis, Light, and Life 122
From DNA to Protein: The Role of RNA 179
Photosynthesis: A Historical Perspective 122
The Genetic Code 180
The Nature of Light 125
Protein Synthesis 181
ESSAY: The Fitness of Light 126
Regulation of Gene Expression
The Role of Pigments 126
in Eukaryotes 186
The Reactions of Photosynthesis 129
The DNA of the Eukaryotic
The Carbon-Fixation Reactions 135
Chromosome 187
ESSAY: Global Warming: The Future
Transcription and Processing of
Is Now 140
mRNA in Eukaryotes 188
Noncoding RNAs and Gene Regulation 190

10 Recombinant DNA Technology, Plant

Biotechnology, and Genomics 192
Recombinant DNA Technology 192
Plant Biotechnology 198
ESSAY: Model Plants: Arabidopsis
thaliana and Oryza sativa 199
ESSAY: Totipotency 202
Genomics 205

11 The Process of Evolution 209

Darwin’s Theory 209
The Concept of the Gene Pool 211
The Behavior of Genes in Populations:
SECTION 3 GENETICS AND EVOLUTION 151 The Hardy-Weinberg Law 212
8 Sexual Reproduction and Heredity 152 The Agents of Change 212
Sexual Reproduction 153 Responses to Selection 214
The Eukaryotic Chromosome 154 ESSAY: Invasive Plants 217
 CONT ENTS    ix

The Result of Natural Selection: Adaptation 217 Reproduction and Gene Exchange 259
The Origin of Species 220 Endospores 261
How Does Speciation Occur? 221 Metabolic Diversity 261
ESSAY: Adaptive Radiation in Hawaiian Bacteria 263
Lobeliads 224 Archaea 269
The Origin of Major Groups of Organisms 229 Viruses 270
Viroids: Other Infectious Particles 275

14 Fungi 278
The Importance of Fungi 279
Characteristics of Fungi 281
ESSAY: Phototroprism in a Fungus 285
Microsporidians: Phylum Microsporidia 286
Chytrids: A Polyphyletic Group of Fungi
with Flagellated Cells 287
Zygomycetes: A Polyphyletic Group
of Filamentous Fungi 288
Glomeromycetes: Phylum
Glomeromycota 290
Ascomycetes: Phylum Ascomycota 291
Basidiomycetes: Phylum Basidiomycota 295
ESSAY: Predaceous Fungi 303
Symbiotic Relationships of Fungi 306
SECTION 4 DIVERSITY 233 ESSAY: From Pathogen to Symbiont:
Fungal Endophytes 307
12 Systematics: The Science of
Biological Diversity 234
15 Protists: Algae and Heterotrophic
Taxonomy: Nomenclature and
Protists 317
Classification 234
Ecology of the Algae 320
ESSAY: Convergent Evolution 239
ESSAY: Algae and Human Affairs 321
Cladistics 239
ESSAY: Red Tides/Toxic Blooms 323
Molecular Systematics 240
Euglenoids 324
ESSAY: Google Earth: A Tool for
Cryptomonads: Phylum Cryptophyta 324
Discovering and Protecting
Haptophytes: Phylum Haptophyta 326
Biodiversity 241
Dinoflagellates 327
The Major Groups of Organisms: Bacteria,
ESSAY: Coral Reefs and Global Warming 329
Archaea, and Eukarya 243
Photosynthetic Stramenopiles 330
Origin of the Eukaryotes 247
Red Algae: Phylum Rhodophyta 340
The Protists and Eukaryotic Kingdoms 248
Green Algae 345
Life Cycles and Diploidy 250
Heterotrophic Protists 358

13 Prokaryotes and Viruses 256 16 Bryophytes 366

Characteristics of the Prokaryotic Cell 257 The Relationships of Bryophytes to
Diversity of Form 259 Other Groups 367
x   C on t en ts

Comparative Structure and Reproduction Biochemical Coevolution 497

of Bryophytes 368
Liverworts: Phylum Marchantiophyta 373 21 Plants and People 501
Mosses: Phylum Bryophyta 378 The Rise of Agriculture 502
Hornworts: Phylum Anthocerotophyta 388 ESSAY: The Origin of Maize 510
The Growth of Human Populations 514
17 Seedless Vascular Plants 391 ESSAY: Biofuels: Part of the Solution,
Evolution of Vascular Plants 391 or Another Problem? 515
Organization of the Vascular Plant Agriculture in the Future 515
Body 392
Reproductive Systems 397
The Phyla of Seedless Vascular
Plants 398
ESSAY: Coal Age Plants 400
Phylum Rhyniophyta 402
Phylum Zosterophyllophyta 403
Phylum Trimerophytophyta 403
Phylum Lycopodiophyta 403
Phylum Monilophyta 409

18 Gymnosperms 430
Evolution of the Seed 430
Progymnosperms 432
Extinct Gymnosperms 433
Living Gymnosperms 435
Phylum Coniferophyta 437 SECTION 5 The Angiosperm Plant Body:
Other Living Gymnosperm Phyla: Structure and
Cycadophyta, Ginkgophyta, Development 525
and Gnetophyta 448 22 Early Development of the Plant Body 526
Formation of the Embryo 526
19 Introduction to the Angiosperms 457 The Mature Embryo 530
Diversity in the Phylum Anthophyta 457 Seed Maturation 532
The Flower 460 Requirements for Seed Germination 532
The Angiosperm Life Cycle 465 ESSAY: Wheat: Bread and Bran 533
ESSAY: Hay Fever 475 From Embryo to Adult Plant 534

20 Evolution of the Angiosperms 477 23 Cells and Tissues of the Plant Body 538
Angiosperm Ancestors 477 Apical Meristems and Their
Time of Origin and Diversification Derivatives 538
of the Angiosperms 478 Growth, Morphogenesis, and
Phylogenetic Relationships of the Differentiation 539
Angiosperms 478 Internal Organization of the Plant
Evolution of the Flower 482 Body 541
Evolution of Fruits 492 Ground Tissues 541
 CONT ENTS    xi

Vascular Tissues 544

Dermal Tissues 553

24 The Root: Structure and Development 558

Root Systems 559
Origin and Growth of Primary Tissues 560
Primary Structure 564
Effect of Secondary Growth on the
Primary Body of the Root 569
Origin of Lateral Roots 571
Aerial Roots and Air Roots 572
Adaptations for Food Storage:
Fleshy Roots 573
ESSAY: Getting to the Root of Organ
Development 574
SECTION 6 PHYSIOLOGY OF SEED PLANTS 637
25 The Shoot: Primary Structure
27 Regulating Growth and Development:
and Development 579
The Plant Hormones 638
Origin and Growth of the Primary
Auxins 639
Tissues of the Stem 580
Cytokinins 645
Primary Structure of the Stem 583
Ethylene 647
Relation between the Vascular Tissues
Abscisic Acid 649
of the Stem and Leaf 588
Gibberellins 650
Morphology and Structure of
Brassinosteroids 652
the Leaf 592 The Molecular Basis of Hormone Action 653
ESSAY: Leaf Dimorphism in Aquatic
Plants 596 28 External Factors and Plant Growth 660
Grass Leaves 598 The Tropisms 660
Development of the Leaf 599 Circadian Rhythms 665
ESSAY: Strong, Versatile, Sustainable Photoperiodism 668
Bamboo 603 The Floral Stimulus 674
Leaf Abscission 604 Vernalization: Cold and the Flowering
Response 674
Transition between the Vascular Systems
Dormancy 674
of the Root and Shoot 604
ESSAY: Doomsday Seed Vault:
Development of the Flower 604
Securing Crop Diversity 677
Stem and Leaf Modifications 607
Nastic Movements and Solar Tracking 678

26 Secondary Growth in Stems 614

29 Plant Nutrition and Soils 683
Annuals, Biennials, and Perennials 614 Essential Elements 684
The Vascular Cambium 615 Functions of Essential Elements 685
Effect of Secondary Growth on the The Soil 686
Primary Body of the Stem 617 Nutrient Cycles 691
Wood: Secondary Xylem 626 Nitrogen and the Nitrogen Cycle 692
ESSAY: The Truth about Knots 627 ESSAY: Carnivorous Plants 694
xii   C on t en ts

Phosphorus and the Phosphorus Cycle 700 Nutrient and Material Cycling
Human Impact on Nutrient Cycles Interactions between Organisms—
and Effects of Pollution 701 beyond Simple Trophic Relationships
Soils and Agriculture 702 Development and Change of Communities
ESSAY: Halophytes: A Future Resource? 703 and Ecosystems
ESSAY: Compost 704
Plant Nutrition Research 704 32 Global Ecology
ESSAY: The Water Cycle 705 Life on the Land
Rainforests
30 The Movement of Water and Solutes Deciduous Tropical Forests
in Plants 708 Savannas
Movement of Water and Inorganic
Deserts
Nutrients through the Plant Body 709
Grasslands
ESSAY: Green Roofs: A Cool Alternative 714
Temperate Deciduous Forests
Absorption of Water and Ions by Roots 717 Temperate Mixed and Coniferous Forests
Assimilate Transport: Movement of
Mediterranean Scrub
Substances through the Phloem 722
The Northernmost Forests—Taiga
and Boreal Forest
Arctic Tundra
A Final Word

Appendix: Classification of Organisms A–1

Suggestions for Further Reading FR–1

Glossary G–1

Illustration Credits IC–1

Index I–1

SECTION 7 ECOLOGY
On the Web: www.whfreeman.com/raven8e
31 The Dynamics of Communities
and Ecosystems
Ecosystem Energetics—Trophic Structure
 preface

A s we approached this revision of Biology of Plants, we

recognized that extensive work would be needed to address
the advances that have been made in all areas of plant biology.
From new molecular details about photosynthesis to the vast
differences in taxonomic relationships that have been revealed
by comparison of DNA and RNA sequences, to advances in
genomics and genetic engineering to an enhanced understanding
of the anatomy and physiology of plants, there have been excit-
ing developments in the field. This current edition of Biology of
Plants has undergone the most significant revision in its history,
with every topic scrutinized and, where necessary, revised and
updated.
While covering these advances, we have strengthened the
narrative by expanding and clarifying discussions; carefully
defining new terms; and adding new diagrams, photos, and
electron micrographs. Each chapter now begins with an at-
tractive photograph and informative caption that relates to the
chapter content but in a tangential way that often touches on
an environmental topic.
With each revision, we continue to pay special attention to
the book’s interlocking themes: (1) the functioning plant body
as the dynamic result of processes mediated by biochemical
interactions; (2) evolutionary relationships as valuable for un-
derstanding form and function in organisms; (3) ecology as an
integrated theme that pervades the book and emphasizes our
dependence on plants to sustain all life on Earth; and (4) mo-
lecular research as essential for revealing details about plant
genetics, cellular function, and taxonomic relationships.
impact of new molecular methods for studying plants,
resulting in the development of golden rice, as well as
plants that are resistant to herbicides, pesticides,
and diseases
• Chapter 11 (The Process of Evolution)—covers
recombination speciation (speciation not involving
polyploidy) and includes two new essays on
“Invasive Plants” and “Adaptive Radiation in Hawaiian
Lobeliads”
• Chapter 12 (Systematics: The Science of Biological
Diversity)—presents an expanded discussion of the
chloroplast as the main source of plant DNA sequence
data and introduces DNA barcoding and supergroups; new
essay on “Google Earth: A Tool for Discovering
and Protecting Biodiversity”
• Chapter 14 (Fungi)—reorganized and updated with the
latest classifications; includes nucleariids and the phyla
Microsporidia and Glomeromycota, as well as a new
phylogenetic tree of the fungi

Changes Reflecting Major Recent

Advances in Plant Science
Every chapter has been carefully revised and updated, most
notably:

• Chapter 7 (Photosynthesis, Light, and Life)—presents

an expanded discussion of light reactions, including an
updated diagram on the transfer of electrons and protons
during photosynthesis; new essay on “Global Warming:
The Future Is Now”

• Chapter 9 (The Chemistry of Heredity and Gene

Expression)—incorporates histone acetylation, DNA
methylation, epigenesis, and noncoding RNAs

• Chapter 10 (Recombinant DNA Technology, Plant The cedar-apple rust fungus alternates between two hosts, cedar
Biotechnology, and Genomics)—updates material on the and apple trees, causing damage to apple harvests (page 278).
xiv   PR EF A CE

• Chapter 15 (Protists: Algae and Heterotrophic Protists)—

incorporates the latest classifications, including a
phylogenetic tree showing the relationships of the algae;
new discussion of the cultivation of algae for biofuel
production and a new essay on “Coral Reefs and Global
Warming”

• Chapter 18 (Gymnosperms)—includes an expanded

discussion of double fertilization in the gnetophytes, as
well as a cladogram of the phylogenetic relationships
among the major groups of embryophytes and a new figure
depicting alternative hypotheses of relationships among
the five major lineages of seed plants

• Chapter 19 (Introduction to the Angiosperms)—follows

the classification recommended by the Angiosperm
Phylogeny Group and presents an expanded discussion
of embryo sac types

• Chapter 20 (Evolution of the Angiosperms)—presents an

expanded discussion of angiosperm ancestors and includes
new cladograms depicting the phylogenetic relationships
of the angiosperms Austrobaileya scandens is considered to have evolved separately
from the main lineage of angiosperms (page 480).

• Chapter 21 (Plants and People)—updated and revised to

include a new figure depicting independent centers of plant
domestication and discusses efforts to develop perennial
versions of important annual grains; new essay on
“Biofuels: Part of the Solution, or Another Problem?”

• Chapter 22 (Early Development of the Plant Body)—

the discussion of seed maturation and dormancy has been
revised, and considerable fine-tuning and updating have
occurred throughout this and the other anatomy chapters,
with an emphasis on structure/function relationships

• Chapter 23 (Cells and Tissues of the Plant Body)—the

presence of forisomes in sieve-tube elements of some
legumes has been added

• Chapter 24 (The Root: Structure and Development)—the

topic of border cells and their functions has been added

• Chapter 25 (The Shoot: Primary Structure and

Development)—includes a new discussion accompanied
by micrographs on leaf vein development and the ABCDE
model of flower development; new essay on “Strong,
Versatile, Sustainable Bamboo”

• Chapter 26 (Secondary Growth in Stems)—a new

diagram depicting the relationship of the vascular cambium
to secondary xylem and secondary phloem has been added

The Chinese brake fern removes arsenic from contaminated soils • Chapter 27 (Regulating Growth and Development: The
(page 391). Plant Hormones)—expanded discussions of the role of auxin

in vascular differentiation and on hormone receptors and
signaling pathways for the plant hormones; new discussions
of brassinosteroids as a major class of plant hormones and
of strigolactones, which interact with auxin in regulating
apical dominance
• Chapter 28 (External Factors and Plant Growth)—
extensive revision of gravitropism, circadian rhythms, floral
stimulus, and thigmonastic movements; new discussion
of genes and vernalization, as well as hydrotropism,
phytochrome-interaction factors (PIFs), and the shade-
avoidance syndrome; new essay on “Doomsday Seed Vault:
Securing Crop Diversity”
• Chapter 29 (Plant Nutrition and Soils)—discusses
strategies involving nitrogen uptake by plants, along with
the new topics of beneficial elements, determinate and
indeterminate nodules, and strategies by plants for the
acquisition of phosphate; essay added on “The Water
Cycle”
Living organisms of the A soil horizon, or “topsoil” (page 690).
• Chapter 30 (The Movement of Water and Solutes in
Plants)—expanded discussion on hydraulic redistribution
and on the mechanisms of phloem loading, including the
polymer trapping mechanism; new essay on “Green Roofs:
A Cool Alternative”
• Chapter 31 (The Dynamics of Communities and
Ecosystems) and Chapter 32 (Global Ecology)—remain
PRE FACE    xv
online at www.whfreeman.com/raven8e. Fully illustrated,
these chapters have been thoroughly updated by Paul
Zedler of the University of Wisconsin, Madison
Media and Supplements
Companion Web Site
www.whfreeman.com/raven8e
For students, this free interactive Web site includes:
• Interactive study aids to help foster understanding
of important concepts from the text, including
multiple-choice quizzes, flashcards, plus interactive
figures and tables
• Animations and live-action videos illustrate several
topics from the text for a deeper understanding of the
more difficult concepts
• Two ecology chapters in easily downloadable and
printable PDF format, fully illustrated and thoroughly
updated with major contributions by Paul Zedler,
University of Wisconsin, Madison.
The dynamic eBook is customizable and fully integrates the
complete contents of the text and interactive media in a format
that features a variety of helpful study tools, including full-text
searching, note-taking, bookmarking, highlighting and more.
For instructors, the site includes all figures from the text
in both PowerPoint and JPEG formats, optimized for excellent
classroom projection. The site also includes the Test Bank by
Robert C. Evans, Rutgers University, Camden, which has been
thoroughly revised and organized by chapter in easy-to-edit
Word files. The Test Bank features approximately 90 questions
per chapter, structured as multiple-choice, true-false, and short-
answer questions that can be used to test student comprehen-
sion of all major topics in the textbook.
Instructor’s Resource DVD with Test Bank
Included here are all the resources from the site, with all text
images in JPEG and PowerPoint formats, plus the Test Bank in
easy-to-edit and -print Word format.
Overhead Transparency Set
The set includes full-color illustrations from the book, optimized
for classroom projection.
Laboratory Topics in Botany
Laboratory Topics in Botany offers several exercises within
each topic that can be selected for coverage that suits individual
course needs. Questions and problems follow each topic; refine-
ments and updating have been made throughout. Written by
Ray F. Evert and Susan E. Eichhorn, University of Wisconsin,
Madison; and Joy B. Perry, University of Wisconsin, Fox Valley.
xvi   PR EF A CE
Preparation Guide for Laboratory Topics in Botany
Preparation Guide for Laboratory Topics in Botany offers
helpful information on how to cover each topic, the length
of time needed to complete the material, the sources of sup-
plies and ordering schedules, how to set up the laboratory, and
ways to guide students in their laboratory work. Written by
Susan E. Eichhorn, University of Wisconsin, Madison; Joy B.
Perry, University of Wisconsin, Fox Valley; and Ray F. Evert,
University of Wisconsin, Madison.
Acknowledgments
We are grateful for the enthusiastic response we have received
from readers who have used previous editions of Biology of
Plants, either in English or in one of the six foreign languages in
which it has been published. As always, we have appreciated the
support and recommendations made by teachers who used the
last edition in their courses. We also wish to thank the following
people who provided valuable critiques of chapters or portions
of chapters for this edition:
Richard Amasino, University of Wisconsin, Madison
Paul Berry, University of Michigan
James Birchler, University of Missouri
Wayne Becker, University of Wisconsin, Madison
Clyde Calvin, Portland State University
Kenneth Cameron, University of Wisconsin, Madison
Nancy Dengler, University of Toronto
John Doebley, University of Wisconsin, Madison
Eve Emshwiller, University of Wisconsin, Madison
Thomas German, University of Wisconsin, Madison
Thomas Givnish, University of Wisconsin, Madison
Linda Graham, University of Wisconsin, Madison
Christopher Haufler, University of Kansas
David Hibbett, Clark University
Robin Kurtz, University of Wisconsin, Madison
Ben Pierce, Southwestern University
Scott Russell, University of Oklahoma, Norman
Dennis Stevenson, New York Botanical Garden
Joseph Williams, University of Tennessee, Knoxville
Paul Zedler, University of Wisconsin, Madison
The following people gave us helpful feedback as we planned
this edition:
Richard Carter, Valdosta State University
Sara Cohen Christopherson, University of Wisconsin, Madison
Les C. Cwynar, University of New Brunswick
Brian Eisenback, Bryan College
Karl H. Hasenstein, University of Louisiana at Lafayette
Bernard A. Hauser, University of Florida
Jodie S. Holt, University of California Riverside
George Johnson, Arkansas Tech University
Carolyn Howes Keiffer, Miami University
Jeffrey M. Klopatek, Arizona State University
Rebecca S. Lamb, Ohio State University
Monica Macklin, Northeastern State University
Carol C. Mapes, Kutztown University of Pennsylvania
Shawna Martinez, Sierra College
Austin R. Mast, Florida State University
Wilf Nicholls, Memorial University of Newfoundland
Karen Renzaglia, Southern Illinois University
Frances M. Wren Rundlett, Georgia State University
A. L. Samuels, University of British Columbia
S. E. Strelkov, University of Alberta
Alexandru M. F. Tomescu, Humboldt State University
M. Lucia Vazquez, University of Illinois at Springfield
Justin K. Williams, Sam Houston State University
Michael J. Zanis, Purdue University
We are very grateful to our artist, Rhonda Nass, for the exquisite
paintings that open each section and for her beautifully drawn
artwork. She has worked closely with us through many edi-
tions, and we value her ability to interpret our pencil sketches
and render them into drawings that are instructive and accu-
rate, as well as attractive. We are grateful to Rick Nass, who
contributed a number of expertly produced graphs. We also
thank Sarah Friedrich and Kandis Elliot, Media Specialists,
Department of Botany, University of Wisconsin, Madison, for
the preparation of digital images of photomicrographs and her-
barium specimens. Mark Allen Wetter, Collections Manager/
Senior Academic Curator, and Theodore S. Cochrane, Senior
Academic Curator, both of the Wisconsin State Herbarium,
Department of Botany, University of Wisconsin, Madison, were
very helpful in selecting and scanning herbarium specimens for
our use.
We would especially like to thank Sally Anderson, our
talented developmental editor, who has worked with us for five
editions now. We are grateful for her many contributions at
every stage of the process, from the early stages of planning
the new edition through the manuscript and proof stages to the
finished book. We have worked well together over the years,
and we thank her for her dedication to all aspects of making
this edition the most accurate and accessible of all.
We would also like to thank Richard Robinson, who has
written the engaging ecology-oriented essays that have been
added to this edition. The essays are highlighted in the text by
a green leaf, and they cover such topics as green roofs, invasive
plants, coral bleaching, the development of biofuels, and the use
of Google Earth to map and study biodiversity.
The preparation of the eighth edition has involved the
collaborative efforts of a large number of talented people at
W. H. Freeman and Company. Particular thanks go to Peter
Marshall, Publisher of Life Sciences, whose vision and support
have made this new edition possible; to Vivien Weiss, who has

skillfully managed the production process; to Elyse Rieder and
Bianca Moscatelli, who enthusiastically tracked down pho-
tographs; and to Blake Logan, who applied her design talents
to give this edition a new look. We especially want to thank
Sheridan Sellers for the remarkable work she has done in mak-
ing up the pages of this book—with her aesthetic and peda-
gogical talents, she has worked a miracle in placing the many
large and complicated illustrations into a coherent layout. And
we are grateful to Linda Strange, our long-time copyeditor
who, with good humor and a steady hand, manages to hold us
to a high standard in consistency and accuracy. We also thank
Marni Rolfes, associate editor, who competently handled the
day-to-day matters and kept us on track, and Bill Page, who
PREF ACE    xvii
coordinated the complicated illustration program. Our gratitude
also goes to Debbie Clare, Associate Director of Marketing,
who has been tireless in managing the sales and market-
ing efforts for this edition, and to Susan Wein, Production
Coordinator, for her many contributions during the complex
stages of production.
A great number of people, only some of whom are men-
tioned here, have contributed in many essential ways to this
revision, and we extend to them our deepest appreciation and
gratitude.
Ray F. Evert
Susan E. Eichhorn
INTRODUCTION

Mayapple (Podophyllum peltatum), which grows in open woods

and pastures in the contiguous United States and Canada, blooms
in early May. Utilizing the sun’s energy, the plant rapidly produces
stems, leaves, and flowers. Although called mayapple, the fruit,
which is yellowish when ripe, is actually a berry. The fruits are edible
and can be used in preserves and beverages, but the leaves and roots
are poisonous.

chapter outline
Evolution of Plants
Evolution of Communities
Appearance of Human Beings
“W hat drives life is . . . a little current, kept up by the
sunshine,” wrote Nobel laureate Albert Szent-Györgyi.
With this simple sentence, he summed up one of the greatest
marvels of evolution—photosynthesis. During the photosyn-
thetic process, radiant energy from the sun is captured and used
to form the sugars on which all life, including our own, de-
pends. Oxygen, also essential to our existence, is released as a
by-product. The “little current” begins when a particle of light
strikes a molecule of the green pigment chlorophyll, boosting
one of the electrons in the chlorophyll to a higher energy level.
The “excited” electron, in turn, initiates a flow of electrons that
ultimately converts the radiant energy from the sun to the chem-
ical energy of sugar molecules. Sunlight striking the leaves of
the water lily shown above, for example, is the first step in the
process leading to production of the molecules that make up the
flowers, leaves, and stems, as well as all the molecular compo-
nents that allow the plants to grow and develop.
Only a few types of organisms—plants, algae, and some
bacteria—possess chlorophyll, which is essential for a living
cell to carry out photosynthesis. Once light energy is trapped
in chemical form, it becomes available as an energy source to
all other organisms, including human beings. We are totally de-
pendent on photosynthesis, a process for which plants are ex-
quisitely adapted.
The word “botany” comes from the Greek botane-, meaning
“plant,” derived from the verb boskein, “to feed.” Plants, how-
ever, enter our lives in innumerable ways other than as sources
2
C hap t e r
1
Botany:
An Introduction
A change of habitat Although plants are primarily adapted for
life on land, some, such as the hardy water lily (Nymphaea fabiola),
have returned to an aquatic existence. Evidence of its ancestors’
sojourn on land includes a water-resistant waxy outer layer, or cuticle,
as well as stomata through which gases are exchanged, and a highly
developed internal transport system.
of food. They provide us with fiber for clothing; wood for fur-
niture, shelter, and fuel; paper for books (such as the page you
are reading at this moment); spices for flavor; drugs for medi-
cines; and the oxygen we breathe. We are utterly dependent on
plants. Plants also have enormous sensory appeal, and our lives
are enhanced by the gardens, parks, and wilderness areas avail-
able to us. The study of plants has provided us with great in-
sight into the nature of all life and will continue to do so in the
years ahead. And, with genetic engineering and other forms of
modern technology, we have entered the most exciting period
in the history of botany, where plants can be transformed, for
example, to resist disease, kill pests, produce vaccines, manu-
facture biodegradable plastic, tolerate high-salt soils, resist
freezing, and provide higher levels of vitamins and minerals in
food products, such as maize (corn) and rice.
CHEC K POINT S
After reading this chapter, you should be able to answer the following:
1. Why do biologists believe that all living things on Earth today
share a common ancestor?
2. What is the principal difference between a heterotroph and an
autotroph, and what role did each play on the early Earth?
3. Why is the evolution of photosynthesis thought to be such an
important event in the evolution of life in general?
4. What were some of the problems encountered by plants as they
made the transition from the sea to the land, and what structures
in terrestrial plants evolved to solve those problems?
5. What are biomes, and what are the principal roles of plants in an
ecosystem?
 Evolution of Plants 3

Evolution of Plants
Life Originated Early in Earth’s Geologic History
Like all other living organisms, plants have had a long history
during which they evolved, or changed, over time. The planet
Earth itself—an accretion of dust and gases swirling in orbit
around the star that is our sun—is some 4.6 billion years old
(Figure 1–1). It is believed that Earth sustained a lethal meteor
bombardment that ended about 3.8 to 3.9 billion years ago.
Vast chunks of rubble slammed into the planet, helping to keep
it hot. As the molten Earth began to cool, violent storms raged,
accompanied by lightning and the release of electrical energy,
and widespread volcanism spewed molten rock and boiling
water from beneath the Earth’s surface.
The earliest known fossils are found in rocks of Western 1–2 The earliest known fossils Obtained from ancient rocks
Australia about 3.5 billion years old (Figure 1–2). These micro- in northwestern Western Australia, these fossilized prokaryotes
fossils consist of several kinds of small, relatively simple fila- are dated at 3.5 billion years of age. They are about a billion years
mentous microorganisms resembling bacteria. About the same younger than the Earth itself, but there are few suitable older
age as these microfossils are ancient stromatolites—fossilized rocks in which to look for earlier evidence of life. More complex
microbial mats consisting of layers of filamentous and other organisms—those with eukaryotic cellular organization—did not
microorganisms and trapped sediment. Stromatolites continue evolve until about 2.1 billion years ago. For about 1.5 billion years,
to be formed today in a few places, such as in the warm, shal- therefore, prokaryotes were the only forms of life on Earth. These
low oceans off the shores of Australia and the Bahamas (see so-called microfossils have been magnified 1000 times.
Chapter 13). By comparing the ancient stromatolites with mod-
ern ones, which are formed by cyanobacteria (filamentous pho-
tosynthetic bacteria), scientists have concluded that the ancient on Mars, for example, whose early history paralleled that of
stromatolites were formed by similar filamentous bacteria. Earth. Strong evidence, first discovered by the Opportunity
Whether life originated on Earth or reached Earth through rover in 2004, indicated that water once flowed across the
space in the form of spores (resistant reproductive cells) or planet, raising the possibility that, at one time, Mars could
by some other means is problematic. Life may have formed have supported life (Figure 1–3). In 2008, the Phoenix Mars

1–1 Life on Earth Of the nine planets in our solar system,
only one, as far as we know, has life on it. This planet, Earth, is
visibly different from the others. From a distance, it appears blue
and green, and it shines a little. The blue is water, the green is
chlorophyll, and the shine is sunlight reflected off the layer of
gases surrounding the planet’s surface. Life, at least as we know it,
depends on these visible features of Earth.
1–3 Life on Mars? This color-enhanced image shows a portion
of the Jezero Crater, a 25-mile-wide impact crater on northern
Mars that once held a lake. Claylike minerals (indicated in green)
were carried by ancient rivers into the lake, forming a delta.
Because clays are able to trap and preserve organic matter,
deltas and lakebeds are promising areas in which to search for
signs of ancient life on Mars.
4 C h a p te r 1    Botany: An Introduction

Lander found water ice in abundance near the surface. More-
over, its instruments monitored a diurnal water cycle: water
vapor, originating from the shallow subsurface water ice and
from water clinging to soil grains, is released into the Martian
atmosphere during the morning hours, and at night it condenses
and falls out by gravity. Most of the ice crystals evaporate as
they fall through the atmospheric boundary layer, but snowfall
on Mars has been observed.
No organic molecules or traces of previous or present bio-
logical activity were detected at the Phoenix landing site. How-
ever, one would expect organic molecules to be present in the
Martian soil, given the steady influx of certain types of mete-
orites that contain considerable quantities of organic material.
Meteorites that fall to Earth contain amino acids and organic
carbon molecules such as formaldehyde. We will continue to
assume, however, that life on Earth originated on Earth.
In 2011, NASA’s Mars Reconnaissance Orbiter satellite
found evidence of liquid water flowing down slopes and crater
walls during the warm month on Mars. The liquid is thought to
be highly salty and to occur just below the surface, where it is
protected from freezing in the frigid temperatures found on Mars
and from evaporating in the planet’s low air pressures. These
findings further raise the possibilities of finding life on Mars.
that all living things share a common ancestor: a DNA-based
microbe that lived more that 3.5 billion years ago. Near the end
of On the Origin of Species, Charles Darwin wrote: “Probably
all the organic beings which have ever lived on this earth have
descended from some one primordial form, into which life was
first breathed.”
Autotrophic Organisms Make Their Own Food, but
Heterotrophic Organisms Must Obtain Their Food from
External Sources
Cells that satisfy their energy requirements by consuming the
organic compounds produced by external sources are known
as heterotrophs (Gk. heteros, “other,” and trophos, “feeder”).
A heterotrophic organism is dependent on an outside source of
organic molecules for its energy. Animals, fungi (Figure 1–4),
and many of the one-celled organisms, such as certain bacteria
and protists, are heterotrophs.
As the primitive heterotrophs increased in number, they
began to use up the complex molecules on which their existence
depended—and which had taken millions of years to accumu-
late. Organic molecules in free solution (that is, not inside a cell)

Most Likely, the Forerunners of the First Cells Were Simple

Aggregations of Molecules
According to current theories, organic molecules, formed by the
action of lightning, rain, and solar energy on gases in the en-
vironment or spewed out of hydrothermal vents, accumulated
in the oceans. Some organic molecules have a tendency to ag-
gregate in groups, and these groups probably took the form of
droplets, similar to the droplets formed by oil in water. Such
assemblages of organic molecules appear to have been the fore-
runners of primitive cells, the first forms of life. Sidney W. Fox
and his coworkers at the University of Miami produced proteins
that aggregated into cell-like bodies in water. Called “proteinoid
microspheres,” these bodies grow slowly by the accumulation
of additional proteinoid material and eventually bud off smaller
microspheres. Although Fox likened this process to a type of re-
production, the microspheres are not living cells. Some research-
ers have suggested that clay particles, or even bubbles, may have
played a role in life’s origin on Earth by collecting chemicals
and concentrating them for synthesis into complex molecules.
According to current theories, these organic molecules may
also have served as the source of energy for the earliest forms
of life. The primitive cells or cell-like structures were able to
use these abundant compounds to satisfy their energy require-
ments. As they evolved and became more complex, these cells
were increasingly able to control their own destinies. With this
increasing complexity, they acquired the ability to grow, to re-
produce, and to pass on their characteristics to subsequent gen-
erations (heredity). Together with cellular organization, these
properties characterize all living things on Earth.
Today, just about all organisms use an identical genetic 1–4 A modern heterotroph This fungus, an orange-cap boletus
code to translate DNA into proteins (see Chapter 9), whether (Leccinum sp.) known as an Aspen mushroom, is growing on a
they are fungi, plants, or animals. It seems quite clear, there- forest floor in Colorado. Like other fungi, this boletus absorbs its
fore, that life as we know it emerged on Earth only once and food (often from other organisms).
 Evolution of Plants 5

1–5 Photosynthetic autotrophs Large-flowered

trilliums (Trillium grandiflorum), one of the first
plants to flower in spring in the deciduous woods
of eastern and midwestern North America, are seen
here growing at the base of birch trees. Like most
vascular plants, trilliums and birches are rooted in
the soil; photosynthesis occurs chiefly in the leaves.
Trillium produces flowers in well-lighted conditions,
before leaves appear on surrounding trees. The
underground portions (rhizomes) of the plant live
for many years and spread to produce new plants
vegetatively under the thick cover of decaying
material on the forest floor. Trilliums also reproduce
by producing seeds, which are dispersed by ants.

became more and more scarce, and competition began. Under the oceans and lakes reacted with dissolved iron and precipitated
pressure of this competition, cells that could make efficient use of as iron oxides (Figure 1–6). From about 2.7 to 2.2 billion years
the limited energy sources now available were more likely to sur- ago, oxygen began gradually to accumulate in the atmosphere.
vive than cells that could not. In the course of time, by the long, By about 700 million years ago, atmospheric levels of oxygen
slow process of elimination of the most poorly adapted, cells increased markedly, and began to approach modern levels dur-
evolved that were able to make their own energy-rich molecules ing the Cambrian period (570–510 million years ago).
out of simple inorganic materials. Such organisms are called This increase in oxygen level had two important conse-
autotrophs, “self-feeders.” Without the evolution of these early quences. First, some of the oxygen molecules in the outer layer
autotrophs, life on Earth would soon have come to an end. of the atmosphere were converted to ozone (O3) molecules.
The most successful of the autotrophs were those in which When there is a sufficient quantity of ozone in the atmosphere,
a system evolved for making direct use of the sun’s energy— it absorbs the ultraviolet rays—rays highly destructive to living
that is, the process of photosynthesis (Figure 1–5). The earli- organisms—from the sunlight that reaches the Earth. By about
est photosynthetic organisms, although simple in comparison 450 million years ago, the ozone layer apparently protected or-
with plants, were much more complex than the primitive het- ganisms sufficiently so they could survive in the surface layers
erotrophs. Use of the sun’s energy required a complex pigment of water and on the shores, and life emerged on land for the
system to capture the light energy and, linked to this system, a first time.
way to store the energy in an organic molecule.
Evidence of the activities of photosynthetic organisms
has been found in rocks 3.4 billion years old, about 100 mil-
lion years after the first fossil evidence of life on Earth. We can
be almost certain, however, that both life and photosynthetic
organisms evolved considerably earlier than the evidence sug-
gests. In addition, there seems to be no doubt that heterotrophs
evolved before autotrophs. With the arrival of autotrophs, the
flow of energy in the biosphere (that is, the living world and its
environment) came to assume its modern form: radiant energy
from the sun channeled through the photosynthetic autotrophs
to all other forms of life.

Photosynthesis Altered Earth’s Atmosphere, Which in Turn

Influenced the Evolution of Life
As photosynthetic organisms increased in number, they changed
the face of the planet. This biological revolution came about be-
cause photosynthesis typically involves splitting the water mol- 1–6 Banded iron formations These 2-billion-year-old red bands
ecule (H2O) and releasing its oxygen as free oxygen molecules of iron oxide (also known as rust), found at Jasper Knob in Michigan,
(O2). Prior to 2.2 billion years ago, the oxygen released into the are evidence of oxygen accumulation.
6 C h a p te r 1    Botany: An Introduction

Second, the increase in free oxygen opened the way to

a much more efficient utilization of the energy-rich carbon-
containing molecules formed by photosynthesis. It enabled or-
ganisms to break down those molecules by the oxygen-utilizing
process known as respiration. As discussed in Chapter 6, res-
piration yields far more energy than can be extracted by any
anaerobic, or oxygenless, process.
Before the atmosphere accumulated oxygen and became
aerobic, the only cells that existed were prokaryotic—sim-
ple cells that lacked a nuclear envelope and did not have their
genetic material organized into complex chromosomes. It is
likely that the first prokaryotes were heat-loving organisms
called “archaea” (meaning “ancient ones”), the descendants of
which are now known to be widespread, with many thriving
at extremely high temperatures and in acid environments hos-
tile to other forms of life. Bacteria are also prokaryotes. Some
archaea and bacteria are heterotrophic, and others, such as the
cyanobacteria, are autotrophic.
According to the fossil record, the increase of relatively 1–7 Evolution of multicellular organisms Early in the course of
abundant free oxygen was accompanied by the first appearance their evolution, multicellular photosynthetic organisms anchored
of eukaryotic cells—cells with nuclear envelopes, complex themselves to rocky shores. These kelp (Durvillaea potatorum), seen
chromosomes, and organelles, such as mitochondria (sites of res- at low tide on the rocks along the coast of Victoria and Tasmania,
piration) and chloroplasts (sites of photosynthesis), surrounded Australia, are brown algae (class Phaeophyceae), a group in which
by membranes. Eukaryotic organisms, in which the individual multicellularity evolved independently of other groups of organisms.
cells are usually much larger than those of the bacteria, appeared
about 2.1 billion years ago and were well established and diverse
developed relatively strong cell walls for support, as well as
by 1.2 billion years ago. Except for archaea and bacteria, all or-
specialized structures to anchor their bodies to the rocky sur-
ganisms—from amoebas to dandelions to oak trees to human be-
faces (Figure 1–7). As these organisms increased in size, they
ings—are composed of one or more eukaryotic cells.
were confronted with the problem of how to supply food to
the dimly lit, more deeply submerged portions of their bodies,
The Seashore Environment Was Important in the Evolution where photosynthesis was not taking place. Eventually, special-
of Photosynthetic Organisms ized food-conducting tissues evolved that extended the length
Early in evolutionary history, the principal photosynthetic or- of the bodies of these organisms and connected the upper,
ganisms were microscopic cells floating below the surface photosynthesizing parts with the lower, nonphotosynthesizing
of the sunlit waters. Energy abounded, as did carbon, hydro- structures.
gen, and oxygen, but as the cellular colonies multiplied, they
quickly depleted the mineral resources of the open ocean. (It is Colonization of the Land Was Associated with the
this shortage of essential minerals that is the limiting factor in
any modern plans to harvest the seas.) As a consequence, life
Evolution of Structures to Obtain Water and Minimize
began to develop more abundantly toward the shores, where the Water Loss
waters were rich in nitrates and minerals carried down from the The body of a plant can best be understood in terms of its long
mountains by rivers and streams and scraped from the coasts by history and, in particular, in terms of the evolutionary pressures
the ceaseless waves. involved in the transition to land. The requirements of a pho-
The rocky coast presented a much more complicated envi- tosynthetic organism are relatively simple: light, water, carbon
ronment than the open sea, and, in response to these evolution- dioxide for photosynthesis, oxygen for respiration, and a few
ary pressures, living organisms became increasingly complex minerals. On land, light is abundant, as are oxygen and car-
in structure and more diversified. Not less than 650 million bon dioxide, both of which circulate more freely in air than in
years ago, organisms evolved in which many cells were linked water. Also, the soil is generally rich in minerals. The critical
together to form an integrated, multicellular body. In these factor, then, for the transition to land—or as one investigator
primitive organisms we see the early stages in the evolution of prefers to say, “to the air”—is water.
plants, fungi, and animals. Fossils of multicellular organisms Land animals, generally speaking, are mobile and able to
are much easier to detect than those of simpler ones. The his- seek out water just as they seek out food. Fungi, though im-
tory of life on Earth, therefore, is much better documented from mobile, remain largely below the surface of the soil or within
the time of their first appearance. whatever damp organic material they feed on. Plants utilize
On the turbulent shore, multicellular photosynthetic or- an alternative evolutionary strategy. Roots anchor the plant in
ganisms were better able to maintain their position against the ground and collect the water required for maintenance of
the action of the waves, and, in meeting the challenge of the the plant body and for photosynthesis, while the stems provide
rocky coast, new forms developed. Typically, these new forms support for the principal photosynthetic organs, the leaves. A
 Evolution of Plants 7

the vascular system, or conducting system, of the stem con-

ducts a variety of substances between the photosynthetic and
nonphotosynthetic parts of the plant body. The vascular system
has two major components: the xylem, through which water
passes upward through the plant body, and the phloem, through
which food manufactured in the leaves and other photosynthetic
parts of the plant is transported throughout the plant body. It is
this efficient conducting system that gives the main group of
plants—the vascular plants—their name (Figure 1–9).
Plants, unlike animals, continue to grow throughout their
lives. All plant growth originates in meristems, which are

20 μm
1–8 Stomata Open stomata on the surface of a Apical meristem
of shoot
tobacco (Nicotiana tabacum) leaf. Each stoma in the aerial
parts of the plant is regulated by two guard cells.

Leaflets

continuous stream of water moves upward through the roots

and stems, then out through the leaves. The outermost layer
of cells, the epidermis, of all the aboveground portions of the
plant that are ultimately involved in photosynthesis is covered
with a waxy cuticle, which retards water loss. However, the cu-
ticle also tends to prevent the exchange of gases between the Xylem
Node Phloem
plant and the surrounding air that is necessary for both photo-
Petiole
synthesis and respiration. The solution to this dilemma is found
in the stomata (singular: stoma), each consisting of a pair of Xylem Phloem
specialized epidermal cells (the guard cells), with a small open- Epidermis
Node
ing between them. The stomata open and close in response to
environmental and physiological signals, thus helping the plant Mesophyll
maintain a balance between its water losses and its oxygen and
Stoma Leaf
carbon dioxide requirements (Figure 1–8). Bud
Internode
In younger plants and in annuals—plants with a life span
of one year—the stem is also a photosynthetic organ. In longer-
Epidermis
lived plants—perennials—the stem may become thickened and Phloem
woody and covered with cork, which, like the cuticle-covered Xylem
Node
epidermis, retards water loss. In both annuals and perennials,
Cortex

1–9 A modern vascular plant Diagram of a young broad bean

Seed coat
(Vicia faba) plant, showing the principal organs and tissues of Stem
the modern vascular plant body. The organs—root, stem, and Cotyledon
Shoot system Xylem
leaf—are composed of tissues, which are groups of cells with Root system
distinct structures and functions. Collectively, the roots make up
the root system, and the stems and leaves together make up the Phloem
shoot system of the plant. Unlike roots, stems are divided into Epidermis
nodes and internodes. The node is the part of the stem at which Cortex
one or more leaves are attached, and the internode is the part of Root
the stem between two successive nodes. In the broad bean, the Lateral root
first few foliage leaves are divided into two leaflets each. Buds, or
embryonic shoots, commonly arise in the axils—the upper angle
between leaf and stem—of the leaves. Lateral, or branch, roots arise
from the inner tissues of the roots. The vascular tissues—xylem
and phloem—occur together and form a continuous vascular
system throughout the plant body. They lie just inside the cortex
in root and stem. The mesophyll tissue of leaves is specialized for
photosynthesis. In this diagram, a cotyledon, or seed leaf, can be Apical meristem
seen through a tear in the seed coat. of root
8 C h a p te r 1    Botany: An Introduction

(a) (b)

1–10 Examples of the enormous diversity of

biomes on Earth (a) The temperate deciduous forest,
which covers most of the eastern United States and
southeastern Canada, is dominated by trees that lose
their leaves in the cold winters. Here are paper birches
and a red maple photographed in early autumn in the
Adirondack Mountains of New York State. (b) Underlain
with permafrost, Arctic tundra is a treeless biome
(c) characterized by a short growing season. Shown here
are tundra plants in full autumn color, photographed
in Tombstone Valley, Yukon, Canada. (c) In Africa,
savannas are inhabited by huge herds of grazing
mammals, such as these zebras and wildebeests. The
tree in the foreground is an acacia. (d) Moist tropical
forests, shown here in Costa Rica, constitute the richest,
most diverse biome on Earth, with perhaps half of all
species of organisms on Earth found there. (e) Deserts
typically receive less than 25 centimeters of rain
per year. Here in the Sonoran desert in Arizona, the
dominant plant is the giant saguaro cactus. Adapted for
life in a dry climate, saguaro cacti have shallow, wide-
spreading roots, as well as thick stems for storing water.
(f) Mediterranean climates are rare on a world scale.
Cool, moist winters, during which the plants grow, are
followed by hot, dry summers, during which the plants
become dormant. Shown here is an evergreen oak
woodland on Mount Diablo in California.
(d)
 Evolution of Communities 9

(e) (f)

embryonic tissue regions capable of adding cells indefinitely to Evolution of Communities

the plant body. Meristems located at the tips of all roots and
The invasion of the land by plants changed the face of the
shoots—the apical meristems—are involved with the exten-
continents. Looking down from an airplane on one of Earth’s
sion of the plant body. Thus the roots are continuously reach-
great expanses of desert or on one of its mountain ranges, we
ing new sources of water and minerals, and the photosynthetic
can begin to imagine what the world looked like before the ap-
regions are continuously extending toward the light. The type
pearance of plants. Yet even in these regions, the traveler who
of growth that originates from apical meristems is known as
goes by land will find an astonishing variety of plants punctuat-
primary growth. On the other hand, the type of growth that re-
ing the expanses of rock and sand. In those parts of the world
sults in a thickening of stems and roots—secondary growth—
where the climate is more temperate and the rains are more fre-
originates from two lateral meristems, the vascular cambium
quent, communities of plants dominate the land and determine
and the cork cambium.
its character. In fact, to a large extent, they are the land. Rain-
During the transition “to the air,” plants also underwent
forest, savanna, woods, desert, tundra—each of these words
further adaptations that made it possible for them to reproduce
brings to mind a portrait of a landscape (Figure 1–10). The
on land. The first of these adaptations was the production of
main features of each landscape are its plants, enclosing us in
drought-resistant spores. This was followed by the evolution of
a dark green cathedral in our imaginary rainforest, carpeting the
complex, multicellular structures in which the gametes, or re-
ground beneath our feet with wildflowers in a meadow, moving
productive cells, were held and protected from drying out by a
in great golden waves as far as the eye can see across our imag-
layer of sterile cells. In the seed plants, which include almost
inary prairie. Only when we have sketched these biomes—nat-
all familiar plants except the ferns, mosses, and liverworts, the
ural communities of wide extent, characterized by distinctive,
young plant, or embryo, is enclosed within a specialized cov-
climatically controlled groups of plants and animals—in terms
ering (seed coat) provided by the parent. There the embryo is
of trees and shrubs and grasses can we fill in other features,
protected from both drought and predators and is provided with
such as deer, antelope, rabbits, or wolves.
a supply of stored food. The embryo, the supply of stored food,
How do vast plant communities, such as those seen on
and the seed coat are the components of the seed.
a continental scale, come into being? To some extent we can
Thus, in summary, the vascular plant (Figure 1–9) is char-
trace the evolution of the different kinds of plants and ani-
acterized by a root system that serves to anchor the plant in the
mals that populate these communities. Even with accumulat-
ground and to collect water and minerals from the soil; a stem
ing knowledge, however, we have only begun to glimpse the
that raises the photosynthetic parts of the plant body toward its
far more complex pattern of development, through time, of
energy source, the sun; and leaves, which are highly specialized
the whole system of organisms that make up these various
photosynthetic organs. Roots, stems, and leaves are intercon-
communities.
nected by a complicated and efficient vascular system for the
transport of food and water. The reproductive cells of plants are
enclosed within multicellular protective structures, and in seed
Ecosystems Are Relatively Stable, Integrated Units
plants the embryos are protected by resistant coverings. All of That Are Dependent on Photosynthetic Organisms
these characteristics are adaptations to a photosynthetic exis- Such communities, along with the nonliving environment of
tence on land. which they are a part, are known as ecological systems, or
10 C h a p te r 1    Botany: An Introduction

ecosystems. An ecosystem is a kind of corporate entity made

up of transient individuals. Some of these individuals, the
larger trees, live as long as several thousand years; others, the
microorganisms, live only a few hours or even minutes. Yet the
ecosystem as a whole tends to be remarkably stable (although
not static). Once in balance, it does not change for centuries.
Our grandchildren may someday walk along a woodland path
once followed by our great-grandparents, and where they saw
a pine tree, a mulberry bush, a meadow mouse, wild blueber-
ries, or a robin, these children, if this woodland still exists, will
see roughly the same kinds of plants and animals in the same
numbers.
An ecosystem functions as an integrated unit, although
many of the organisms in the system compete for resources.
Virtually every living thing, even the smallest bacterial cell
or fungal spore, provides a food source for some other living
organism. In this way, the energy captured by green plants is
transferred in a highly regulated way through a number of dif-
ferent types of organisms before it is dissipated. Moreover, in-
teractions among the organisms themselves, and between the
organisms and the nonliving environment, produce an orderly
1–11 The clock face of biological time Life appears relatively
early in the Earth’s history, sometime before 6:00 a.m. on a 24-hour
cycling of elements such as nitrogen and phosphorus. Energy
must be added to the ecosystem constantly, but the elements scale. The first multicellular organisms do not appear until the
are cycled through the organisms, returned to the soil, decom- twilight of that 24-hour day, and the genus Homo is a very late
posed by soil bacteria and fungi, and recycled. These transfers arrival—less than a minute before midnight.
of energy and the cycling of elements involve complicated
sequences of events, and in these sequences each group of or-
ganisms has a highly specific role. As a consequence, it is im-
possible to change a single component of an ecosystem without the plants that invaded the land—have changed the surface of
the risk of destroying the balance on which the stability of the the planet, shaping the biosphere according to their own needs,
ecosystem depends. ambitions, or follies.
At the base of productivity in almost all ecosystems are With the cultivation of crops, starting about 10,500 years
the plants, algae, and photosynthetic bacteria. These organisms ago, it became possible to maintain growing populations of
alone have the ability to capture energy from the sun and to people that eventually built towns and cities. This develop-
manufacture organic molecules that they and all other kinds of ment (reviewed in detail in Chapter 21) allowed specialization
organisms may require for life. There are roughly half a mil- and the diversification of human culture. One characteristic of
lion kinds of organisms capable of photosynthesis, and at least this culture is that it examines itself and the nature of other liv-
20 times that many heterotrophic organisms, which are com- ing things, including plants. Eventually, the science of biology
pletely dependent on the photosynthesizers. For animals, in- developed within the human communities that had been made
cluding human beings, many kinds of molecules—including possible through the domestication of plants. The part of biol-
essential amino acids, vitamins, and minerals—can be obtained ogy that deals with plants and, by tradition, with prokaryotes,
only through plants or other photosynthetic organisms. Further- fungi, and algae is called botany, or plant biology.
more, the oxygen that is released into the atmosphere by pho-
tosynthetic organisms makes it possible for life to exist on the Plant Biology Includes Many Different Areas of Study
land and in the surface layers of the ocean. Oxygen is necessary The study of plants has been pursued for thousands of years, but
for the energy-producing metabolic activities of the great ma- like all branches of science, it became diverse and specialized
jority of organisms, including photosynthetic organisms. only during the twentieth century. Until the late 1800s, botany
was a branch of medicine, pursued chiefly by physicians who
used plants for medicinal purposes and who were interested in
Appearance of Human Beings determining the similarities and differences between plants and
Human beings are relative newcomers to the world of living or- animals for that purpose. Today, however, plant biology is an
ganisms (Figure 1–11). If the entire history of the Earth were important scientific discipline that has many subdivisions: plant
measured on a 24-hour time scale starting at midnight, cells physiology, which is the study of how plants function, that is,
would appear in the warm seas before dawn. The first multicel- how they capture and transform energy and how they grow and
lular organisms would not be present until well after sundown, develop; plant morphology, the study of the form of plants;
and the earliest appearance of humans (about 2 million years plant anatomy, the study of their internal structure; plant tax-
ago) would be about half a minute before the day’s end. Yet onomy and systematics, involving the naming and classifying of
humans more than any other animal—and almost as much as plants and the study of the relationships among them; cytology,
 Appearance of Human Beings 11

the study of cell structure, function, and life histories; genet-
ics, the study of heredity and variation; genomics, the study of
the content, organization, and function of genetic information
in whole genomes; molecular biology, the study of the struc-
ture and function of biological molecules; economic botany, the
study of past, present, and future uses of plants by people; eth-
nobotany, the study of the uses of plants for medicinal and other
purposes by indigenous peoples; ecology, the study of the rela-
tionships between organisms and their environment; and paleo-
botany, the study of the biology and evolution of fossil plants.
Included in this book are all organisms that have tradi-
tionally been studied by botanists: not only plants but also
prokaryotes, viruses, fungi, and autotrophic protists (algae).
Nonphotosynthetic eukaryotes and protists have tradition-
ally been the province of zoologists. Although we do not re-
gard algae, fungi, prokaryotes, or viruses as plants, and shall
not refer to them as plants in this book, they are included here
because of tradition and because they are normally considered
part of the botanical portion of the curriculum, just as botany
itself used to be considered a part of medicine. Moreover, both
prokaryotes (e.g., nitrogen-fixing bacteria) and fungi (e.g.,
mycorrhizal fungi) form important, mutually beneficial sym-
biotic relationships with their plant hosts. Virology, bacteriol-
ogy, phycology (the study of algae), and mycology (the study
of fungi) are well-established fields in their own right, but they
still fall loosely under the umbrella of botany.
A Knowledge of Botany Is Important for Dealing
with Today’s—and Tomorrow’s—Problems
In this chapter, we have ranged from the beginnings of life on
this planet to the evolution of plants and ecosystems to the de-
velopment of agriculture and civilization. These broad topics
are of interest to many people other than botanists, or plant bi-
ologists. The urgent efforts of botanists and agricultural scien-
tists will be needed to feed the world’s rapidly growing human
population (Figure 1–12), as discussed in Chapter 21. Modern
plants, algae, and bacteria offer the best hope of providing a re-
newable source of energy for human activities, just as extinct
plants, algae, and bacteria have been responsible for the mas-
sive accumulations of gas, oil, and coal on which our modern
industrial civilization depends. In an even more fundamental

1–12 Growth of the human population Over the last 10,000 years, the human population has grown from
several million to approximately 6.5 billion. A significant increase in the rate of population growth occurred as
a result of the cultivation of plants as crops, and an even more dramatic increase began with the advent of the
Industrial Revolution, which started in the middle of the eighteenth century and continues to the present.
   The consequences of the rapid growth of the human population are many and varied. In the United States
and other parts of the developed world, they include not only the sheer numbers of people but also heavy
consumption of nonrenewable fossil fuels and the resulting pollution—both as the fuels are burned and as a
result of accidents such as oil spills at drilling sites and during transport. In less developed parts of the world,
the consequences include malnutrition and, all too often, starvation, coupled with a continuing vulnerability to
infectious diseases. The consequences for other organisms include not only the direct effects of pollution but
also—and most important—the loss of habitat.
12 C h a p te r 1    Botany: An Introduction
sense, the role of plants, along with that of algae and photo-
synthetic bacteria, commands our attention. As the producers
of energy-containing compounds in the global ecosystem, these
photosynthetic organisms are the route by which all other liv-
ing things, including ourselves, obtain energy, oxygen, and the
many other materials necessary for their continued existence.
As a student of botany, you will be in a better position to assess
the important ecological and environmental issues of the day
and, by understanding, help to build a healthier world.
In this second decade of the twenty-first century, it is clear
that human beings, with a population of 6.5 billion in 2010 and
a projected population of 9 billion by 2050, are managing the
Earth with an intensity that would have been unimaginable a
few decades ago. Every hour, manufactured chemicals fall on
every square centimeter of the planet’s surface. The protec-
tive stratospheric ozone layer formed 450 million years ago
(a)
(c)
has been seriously depleted by the use of chlorofluorocarbons
(CFCs), and damaging ultraviolet rays penetrating the depleted
layer have increased the incidence of skin cancer in people
all over the world. Moreover, it has been estimated that, by
the middle of this century, the average temperature will have
increased between 1.5° and 4.5°C due to the greenhouse ef-
fect. This global-warming phenomenon—the trapping of heat
radiating from the Earth’s surface out into space—is intensi-
fied through the increased amounts of carbon dioxide, nitrogen
oxides, CFCs, and methane in the atmosphere resulting from
human activities. And most seriously, a large portion of the
total number of species of plants, animals, fungi, and micro-
organisms is disappearing during our lifetime—the victims of
human exploitation of the Earth—resulting in a loss of biodi-
versity. All of these trends are alarming, and they demand our
utmost attention.
(b)
1–13 Phytoremediation (a) Sunflowers growing on a lake
contaminated with radioactive cesium and strontium following the
1986 Chernobyl nuclear disaster in Ukraine, then part of the USSR.
Suspended from styrofoam rafts, the sunflowers’ roots are able to
remove up to 90 percent of the contaminants in 10 days. (b) Poplar
(Populus spp.) and willow (Salix spp.) trees growing on a fuel-
contaminated site in Elizabeth, North Carolina. The deep-rooted
trees draw the contaminants up through their stems and leaves,
reducing the need for mechanical pumping and treatment of
contaminated groundwater. (c) Naturally occurring selenium
accumulates in ditches containing run-off from irrigated croplands,
creating bodies of standing water that are poisonous to wildlife,
especially migrating birds. Furthermore, plants grown in the
high-selenium soil that results from evaporation of the water are
toxic. Pickleweed (Salicornia bigelovii), a salt marsh plant, is highly
efficient at removing selenium, which is absorbed by the plant and
then released into the atmosphere to be dispersed by prevailing
winds. For the endangered salt marsh harvest mouse, seen here,
pickleweed is a staple food.
 Appearance of Human Beings 13

Marvelous new possibilities have been developed during the

past few years for the better utilization of plants by people, and
we discuss these developments throughout this book. It is now
possible, for example, to clean up polluted environments through
phytoremediation (Figure 1–13), to stimulate the growth of
plants, to deter their pests, to control weeds in crops, and to form
hybrids between plants with more precision than ever before.
The potential for exciting progress in plant biology grows
with every passing year, as additional discoveries are made and
new applications are developed. The methods of genetic engi-
neering, discussed in Chapter 10, make it possible to accom-
plish the astonishing feat of transferring genes from a virus, a
bacterium, an animal, or a particular plant into an entirely dif-
ferent species of plant in order to produce specific desirable
characteristics in the recipient plant. These so-called transgenic
plants, which contain genes from entirely different species, can
be made to exhibit new and extraordinary properties. By in-
serting maize and bacterial genes into the rice nucleus, for ex-
ample, a more nutritious rice, with higher levels of β-carotene,
can be produced (see Figure 10–1). Another area of investiga-
tion is working toward increasing the iron content of rice. Both
of these developments show promise for improving the health
of the vast numbers of poorly nourished people with rice-
dependent diets. In addition, pest-resistant varieties of maize 1–14 Transgenic plants Citrus seedlings were transformed by
and cotton have been developed by transferring genes from a insertion of flower-initiation genes from Arabidopsis, the small
soil bacterium that attacks the caterpillars that cause major crop flowering plant of the mustard family that is widely used for genetic
losses. The transformed maize and cotton, with their ability to research. The six-month-old transgenic seedling on the right has
express the bacterial genes, kill the caterpillars that might at- developed flowers, whereas the control seedling on the left has not
tack them, allowing growers to reduce their use of pesticide. and will take years to flower and set fruit.
The Hawaiian papaya industry was saved by development
of transgenic papaya trees able to resist the papaya ringspot
virus (see Figure 10–13). Other improvements involve trans- track in New York City, originally slated for demolition, has
genic soybeans that can tolerate Roundup, an herbicide that been saved and now serves as a popular urban destination
kills both broad-leaved weeds and untransformed soybeans. known as the High Line (Figure 1–15a). Running for roughly
In addition, citrus plants have been transformed to flower in a mile, the park is planted primarily with the types of wildflow-
6 months rather than the usual 6 to 20 years, thereby reduc- ers, grasses, shrubs, and trees that grew along the track during
ing the time needed for a citrus tree to set fruit (Figure 1–14). the decades when it was no longer in use. A meandering path-
Attempts are being made to increase the efficiency of photo- way follows the route of the original track, and the setting, with
synthesis and thereby increase crop yields, and to enhance the its views of the Hudson River, as well as of the city life below,
“glossiness” of crops by selectively breeding for plants with draws millions of visitors a year. Another example of the rec-
waxier leaves. This increased waxiness would benefit the plant lamation of an industrial site is a decommissioned military air-
by reducing water loss and, by increasing the reflectivity of the base at Magnuson Park in Seattle, Washington (Figure 1–15b).
surfaces of the crop, could result in a slight cooling of summer Cleared of asphalt and converted to thriving wetlands, the area
temperatures in central North America and Eurasia. with its newly created ponds and thickets of native plants has
Hopes for the future include, among numerous possibili- attracted a variety of wildlife. Winding trails invite visitors to
ties, biodegradable plastics, trees with higher fiber content for experience the peaceful setting as they learn about the essential
paper manufacture, plants with increased levels of healthy oils role played by wetlands habitats.
and anti-cancer proteins, and vaccines that can be produced in As we turn to Chapters 2 and 3, in which our attention nar-
plants, holding the promise of someday being able to deliver rows to a cell so small it cannot be seen by the unaided eye, it
hepatitis B vaccine, for example, in bananas. These methods, is important to keep these broader concerns in mind. A basic
first applied in 1973, have already been the basis of billions of knowledge of plant biology is useful in its own right and is es-
dollars in investments and increased hope for the future. Dis- sential in many fields of endeavor. It is also increasingly rel-
coveries still to be made will undoubtedly exceed our wildest evant to some of society’s most crucial problems and to the
dreams and go far beyond the facts that are available to us now. difficult decisions that will face us in choosing among the pro-
In addition, we have come to appreciate, even more, the posals for diminishing them. To quote from an editorial in the
importance of green spaces to our increasingly complex lives. 19 November 2010 issue of Science: “Plants are essential to
In cities, former industrial sites are being skillfully developed the survival of our planet—to its ecology, biodiversity, and cli-
into parks of various types. An abandoned elevated railroad mate.” Our own future, the future of the world, and the future
14 C h a p te r 1    Botany: An Introduction

(a) (b)
1–15 Greening of abandoned industrial sites (a) The High Line in New York City is built on an
abandoned railroad track elevated above a newly resurrected neighborhood of restaurants, galleries,
and shops. Remnants of the original track can be seen among the shrubs, perennials, grasses, and trees
planted along the popular promenade. (b) A recently created wetlands situated on a former airbase
at Magnuson Park in Seattle, Washington, offers a rich habitat of native plants and various species of
wildlife, including dragonflies, frogs, ducks, owls, hawks, shorebirds, and warblers.

of all kinds of plants—as individual species and as components
of the life-support systems into which we all have evolved—
depend on our knowledge and ability to critically assess the in-
formation we are given. Thus, this book is dedicated not only
to the botanists of the future, whether teachers or researchers,
but also to the informed citizens, scientists, and laypeople alike,
in whose hands such decisions lie.
Summary
Photosynthesis Is the Process by Which the Sun’s Energy
Is Captured to Form Organic Molecules
Only a few kinds of organisms—plants, algae, and some bacte-
ria—have the capacity to capture energy from the sun and use
it to form organic molecules by the process of photosynthesis.
Almost all life on Earth depends, directly or indirectly, on the
products of this process.
The Chemical Building Blocks of Life Accumulated in the
Early Oceans
The planet Earth is some 4.6 billion years old. The oldest
known fossils date back 3.5 billion years and resemble today’s
filamentous bacteria. Although the process by which living or-
ganisms arose is a matter of speculation, there is general agree-
ment that life as we know it probably emerged on Earth only
once—that is, all living things share a common ancestor.
Heterotrophic Organisms Evolved before Autotrophic
Organisms, Prokaryotes before Eukaryotes, and
Unicellular Organisms before Multicellular Organisms
Heterotrophs, organisms that feed on organic molecules or on
other organisms, were the first life forms to appear on Earth.
Autotrophic organisms, those that could produce their own food
by photosynthesis, evolved no less than 3.4 billion years ago.
Until about 2.1 billion years ago, the prokaryotes—archaea and
bacteria—were the only organisms that existed. Eukaryotes, with
larger, much more complex cells, evolved at that time. Multicel-
lular eukaryotes began to evolve at least 650 million years ago,
and they began to invade the land about 450 million years ago.
With the advent of oxygen-producing photosynthesis, in
which water molecules are split and oxygen is released, oxygen
began to accumulate in the atmosphere. The presence of this
free oxygen enabled organisms to break down the energy-rich
products of photosynthesis by aerobic respiration.
Colonization of the Land Was Associated with the
Evolution of Structures to Obtain Water and Minimize
Water Loss
Plants, which are basically a terrestrial group, have achieved
a number of specialized characteristics that suit them for life
on land. These characteristics are best developed among the
members of the dominant group known as the vascular plants.
Among these features are a waxy cuticle, penetrated by special-
ized openings known as stomata through which gas exchange
 Questions 15

takes place, and an efficient conducting system. This system

consists of xylem, in which water and minerals pass from the
roots to the stems and leaves, and phloem, which transports
the products of photosynthesis to all parts of the plant. Plants
increase in length by primary growth and expand in girth by
secondary growth through the activity of meristems, which are
embryonic tissue regions capable of adding cells indefinitely to
the plant body.
Ecosystems Are Relatively Stable, Integrated Units That
Are Dependent on Photosynthetic Organisms
As plants have evolved, they have come to constitute biomes,
great terrestrial assemblages of plants and animals. The interact-
ing systems made up of biomes and their nonliving environments
are called ecosystems. Human beings, which appeared about
2 million years ago, developed agriculture about 10,500 years
ago and thus provided a basis for the huge increase in their pop-
ulation levels. Subsequently, they have become the dominant
ecological force on Earth. Humans have used their knowledge of
plants to foster their own development and will continue to do so
with increasingly greater importance in the future.
QUESTIONS
1. What was the likely source of the raw material incorporated into
the first life forms?
2. What criteria would you use to determine whether an entity is a
form of life?
3. What role did oxygen play in the evolution of life on Earth?
4. What advantages do terrestrial plants have over their aquatic
ancestors? Can you think of any disadvantages to being a
terrestrial plant?
5. Plants enter our lives in innumerable ways other than as sources
of food. How many ways can you list? Have you thanked a green
plant today?
6. A knowledge of botany—of plants, fungi, algae, and bacteria—is
key to our understanding of how the world works. How is that
knowledge important for dealing with today’s and tomorrow’s
problems?

Genetic Engineering Allows Scientists to Transfer Genes

between Entirely Different Species
With the advent of genetic engineering, it became possible for
biologists to transfer genes from one species into an entirely
different species. Genetic engineering has already resulted in
the development of transgenic plants with desirable traits such
as increased nutritive value and resistance to certain diseases
and pests.
Another random document with
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 THE FALCON.
Scene.—An Italian Cottage. Castle and Mountains seen through
Window.

Elisabetta discovered seated on stool in window darning. The

Count with Falcon on his hand comes down through the door at
back. A withered wreath on the wall.

Elisabetta.

So, my lord, the Lady Giovanna, who hath been away so

long, came back last night with her son to the castle.

Count.

Hear that, my bird! Art thou not jealous of her?

My princess of the cloud, my plumed purveyor,
My far-eyed queen of the winds—thou that canst soar
Beyond the morning lark, and howsoe’er
Thy quarry wind and wheel, swoop down upon him
Eagle-like, lightning-like—strike, make his feathers
Glance in mid heaven.

[Crosses to chair.

I would thou hadst a mate!

Thy breed will die with thee, and mine with me:
I am as lone and loveless as thyself.

[Sits in chair.

Giovanna here! Ay, ruffle thyself—be jealous!

Thou should’st be jealous of her. Tho’ I bred thee
The full-train’d marvel of all falconry,
And love thee and thou me, yet if Giovanna
Be here again—No, no! Buss me, my bird!
The stately widow has no heart for me.
Thou art the last friend left me upon earth—
No, no again to that.

[Rises and turns.

My good old nurse,

I had forgotten thou wast sitting there.

Elisabetta.

Ay, and forgotten thy foster-brother too.

Count.

Bird-babble for my falcon! Let it pass.

What art thou doing there?

Elisabetta.

Darning, your lordship.

We cannot flaunt it in new feathers now:
Nay, if we will buy diamond necklaces
To please our lady, we must darn, my lord.
This old thing here (points to necklace round her neck),
they are but blue beads—my Piero,
God rest his honest soul, he bought ’em for me,
Ay, but he knew I meant to marry him.
How couldst thou do it, my son? How couldst thou do it?

Count.

She saw it at a dance, upon a neck

Less lovely than her own, and long’d for it.

Elisabetta.

She told thee as much?

 Count.

No, no—a friend of hers.

Elisabetta.

Shame on her that she took it at thy hands,

She rich enough to have bought it for herself!

Count.

She would have robb’d me then of a great pleasure.

Elisabetta.

But hath she yet return’d thy love?

Count.

Not yet!

Elisabetta.

She should return thy necklace then.

Count.

Ay, if
She knew the giver; but I bound the seller
To silence, and I left it privily
At Florence, in her palace.

Elisabetta.

And sold thine own

To buy it for her. She not know? She knows
There’s none such other——

Count.
 Madman anywhere.
Speak freely, tho’ to call a madman mad
Will hardly help to make him sane again.

Enter Filippo.

Filippo.

Ah, the women, the women! Ah, Monna Giovanna, you here
again! you that have the face of an angel and the heart of a—
that’s too positive! You that have a score of lovers and have not
a heart for any of them—that’s positive-negative: you that have
not the head of a toad, and not a heart like the jewel in it—that’s
too negative; you that have a cheek like a peach and a heart
like the stone in it—that’s positive again—that’s better!

Elisabetta.

Sh—sh—Filippo!

Filippo (turns half round).

Here has our master been a-glorifying and a-velveting and a-

silking himself, and a-peacocking and a-spreading to catch her
eye for a dozen year, till he hasn’t an eye left in his own tail to
flourish among the peahens, and all along o’ you, Monna
Giovanna, all along o’ you!

Elisabetta.

Sh—sh—Filippo! Can’t you hear that you are saying behind

his back what you see you are saying afore his face?

Count.

Let him—he never spares me to my face!

Filippo.
 No, my lord, I never spare your lordship to your lordship’s
face, nor behind your lordship’s back, nor to right, nor to left, nor
to round about and back to your lordship’s face again, for I’m
honest, your lordship.

Count.

Come, come, Filippo, what is there in the larder?

[Elisabetta crosses to fireplace and

puts on wood.

Filippo.

Shelves and hooks, shelves and hooks, and when I see the
shelves I am like to hang myself on the hooks.

Count.

No bread?

Filippo.

Half a breakfast for a rat!

Count.

Milk?

Filippo.

Three laps for a cat!

Count.

Cheese?

Filippo.
 A supper for twelve mites.

Count.

Eggs?

Filippo.

One, but addled.

Count.

No bird?

Filippo.

Half a tit and a hern’s bill.

Count.

Let be thy jokes and thy jerks, man! Anything or nothing?

Filippo.

Well, my lord, if all-but-nothing be anything, and one plate of

dried prunes be all-but-nothing, then there is anything in your
lordship’s larder at your lordship’s service, if your lordship care
to call for it.

Count.

Good mother, happy was the prodigal son,

For he return’d to the rich father; I
But add my poverty to thine. And all
Thro’ following of my fancy. Pray thee make
Thy slender meal out of those scraps and shreds
Filippo spoke of. As for him and me,
There sprouts a salad in the garden still.
(To the Falcon.) Why didst thou miss thy quarry yester-even?
To-day, my beauty, thou must dash us down
Our dinner from the skies. Away, Filippo!

[Exit, followed by Filippo.

Elisabetta.

I knew it would come to this. She has beggared him. I always

knew it would come to this! (Goes up to table as if to resume
darning, and looks out of window.) Why, as I live, there is Monna
Giovanna coming down the hill from the castle. Stops and
stares at our cottage. Ay, ay! stare at it: it’s all you have left us.
Shame upon you! She beautiful! sleek as a miller’s mouse! Meal
enough, meat enough, well fed; but beautiful—bah! Nay, see,
why she turns down the path through our little vineyard, and I
sneezed three times this morning. Coming to visit my lord, for
the first time in her life too! Why, bless the saints! I’ll be bound to
confess her love to him at last. I forgive her, I forgive her! I knew
it would come to this—I always knew it must come to this!
(Going up to door during latter part of speech and opens it.)
Come in, Madonna, come in. (Retires to front of table and
curtseys as the Lady Giovanna enters, then moves chair
towards the hearth.) Nay, let me place this chair for your
ladyship.

[Lady Giovanna moves slowly down

stage, then crosses to chair,
looking about her, bows as she
sees the Madonna over fireplace,
then sits in chair.

Lady Giovanna.

Can I speak with the Count?

Elisabetta.
 Ay, my lady, but won’t you speak with the old woman first, and
tell her all about it and make her happy? for I’ve been on my
knees every day for these half-dozen years in hope that the
saints would send us this blessed morning; and he always took
you so kindly, he always took the world so kindly. When he was
a little one, and I put the bitters on my breast to wean him, he
made a wry mouth at it, but he took it so kindly, and your
ladyship has given him bitters enough in this world, and he
never made a wry mouth at you, he always took you so kindly—
which is more than I did, my lady, more than I did—and he so
handsome—and bless your sweet face, you look as beautiful
this morning as the very Madonna her own self—and better late
than never—but come when they will—then or now—it’s all for
the best, come when they will—they are made by the blessed
saints—these marriages.

[Raises her hands.

Lady Giovanna.

Marriages? I shall never marry again!

Elisabetta (rises and turns).

Shame on her then!

Lady Giovanna.

Where is the Count?

Elisabetta.

Just gone
To fly his falcon.

Lady Giovanna.

Call him back and say

I come to breakfast with him.
 Elisabetta.

Holy mother!
To breakfast! Oh sweet saints! one plate of prunes!
Well, Madam, I will give your message to him.

[Exit.

Lady Giovanna.

His falcon, and I come to ask for his falcon,

The pleasure of his eyes—boast of his hand—
Pride of his heart—the solace of his hours—
His one companion here—nay, I have heard
That, thro’ his late magnificence of living
And this last costly gift to mine own self,

[Shows diamond necklace.

He hath become so beggar’d, that his falcon

Ev’n wins his dinner for him in the field.
That must be talk, not truth, but truth or talk,
How can I ask for his falcon?

[Rises and moves as she speaks.

O my sick boy!
My daily fading Florio, it is thou
Hath set me this hard task, for when I say
What can I do—what can I get for thee?
He answers, “Get the Count to give me his falcon,
And that will make me well.” Yet if I ask,
He loves me, and he knows I know he loves me!
Will he not pray me to return his love—
To marry him?—(pause)—I can never marry him.
His grandsire struck my grandsire in a brawl
At Florence, and my grandsire stabb’d him there.
The feud between our houses is the bar
I cannot cross; I dare not brave my brother,
Break with my kin. My brother hates him, scorns
The noblest-natured man alive, and I—
Who have that reverence for him that I scarce
Dare beg him to receive his diamonds back—
How can I, dare I, ask him for his falcon?

[Puts diamonds in her casket.

Re-enter Count and Filippo. Count turns to Filippo.

Count.

Do what I said; I cannot do it myself.

Filippo.

Why then, my lord, we are pauper’d out and out.

Count.

Do what I said!

[Advances and bows low.

Welcome to this poor cottage, my dear lady.

Lady Giovanna.

And welcome turns a cottage to a palace.

Count.

’Tis long since we have met!

Lady Giovanna.

To make amends
I come this day to break my fast with you.
 Count.

I am much honour’d—yes—

[Turns to Filippo.

Do what I told thee. Must I do it myself?

Filippo.

I will, I will. (Sighs.) Poor fellow!

[Exit.

Count.

Lady, you bring your light into my cottage

Who never deign’d to shine into my palace.
My palace wanting you was but a cottage;
My cottage, while you grace it, is a palace.

Lady Giovanna.

In cottage or in palace, being still

Beyond your fortunes, you are still the king
Of courtesy and liberality.

Count.

I trust I still maintain my courtesy;

My liberality perforce is dead
Thro’ lack of means of giving.

Lady Giovanna.

Yet I come
To ask a gift.

[Moves toward him a little.

 Count.

It will be hard, I fear,

To find one shock upon the field when all
The harvest has been carried.

Lady Giovanna.

But my boy—
(Aside.) No, no! not yet—I cannot!

Count.

Ay, how is he,

That bright inheritor of your eyes—your boy?

Lady Giovanna.

Alas, my Lord Federigo, he hath fallen

Into a sickness, and it troubles me.

Count.

Sick! is it so? why, when he came last year

To see me hawking, he was well enough:
And then I taught him all our hawking-phrases.

Lady Giovanna.

Oh yes, and once you let him fly your falcon.

Count.

How charm’d he was! what wonder?—A gallant boy,

A noble bird, each perfect of the breed.

Lady Giovanna (sinks in chair).

What do you rate her at?

 Count.

My bird? a hundred
Gold pieces once were offer’d by the Duke.
I had no heart to part with her for money.

Lady Giovanna.

No, not for money.

[Count turns away and sighs.

Wherefore do you sigh?

Count.

I have lost a friend of late.

Lady Giovanna.

I could sigh with you

For fear of losing more than friend, a son;
And if he leave me—all the rest of life—
That wither’d wreath were of more worth to me.

[Looking at wreath on wall.

Count.

That wither’d wreath is of more worth to me

Than all the blossom, all the leaf of this
New-wakening year.

[Goes and takes down wreath.

Lady Giovanna.

And yet I never saw

The land so rich in blossom as this year.
 Count (holding wreath toward her).
Was not the year when this was gather’d richer?

Lady Giovanna.

How long ago was that?

Count.

Alas, ten summers!

A lady that was beautiful as day
Sat by me at a rustic festival
With other beauties on a mountain meadow,
And she was the most beautiful of all;
Then but fifteen, and still as beautiful.
The mountain flowers grew thickly round about.
I made a wreath with some of these; I ask’d
A ribbon from her hair to bind it with;
I whisper’d, Let me crown you Queen of Beauty,
And softly placed the chaplet on her head.
A colour, which has colour’d all my life,
Flush’d in her face; then I was call’d away;
And presently all rose, and so departed.
Ah! she had thrown my chaplet on the grass,
And there I found it.

[Lets his hands fall, holding wreath

despondingly.

Lady Giovanna (after pause).

How long since do you say?

Count.

That was the very year before you married.

Lady Giovanna.
When I was married you were at the wars.

Count.

Had she not thrown my chaplet on the grass,

It may be I had never seen the wars.

[Replaces wreath whence he had

taken it.

Lady Giovanna.

Ah, but, my lord, there ran a rumour then

That you were kill’d in battle. I can tell you
True tears that year were shed for you in Florence.

Count.

It might have been as well for me. Unhappily

I was but wounded by the enemy there
And then imprison’d.

Lady Giovanna.

Happily, however,
I see you quite recover’d of your wound.

Count.

No, no, not quite, Madonna, not yet, not yet.

Re-enter Filippo.

Filippo.

My lord, a word with you.

Count.
 Pray, pardon me!

[Lady Giovanna crosses, and passes

behind chair and takes down
wreath; then goes to chair by
table.

Count (to Filippo).

What is it, Filippo?

Filippo.

Spoons, your lordship.

Count.

Spoons!

Filippo.

Yes, my lord, for wasn’t my lady born with a golden spoon in

her ladyship’s mouth, and we haven’t never so much as a silver
one for the golden lips of her ladyship.

Count.

Have we not half a score of silver spoons?

Filippo.

Half o’ one, my lord!

Count.

How half of one?

Filippo.
 I trod upon him even now, my lord, in my hurry, and broke
him.

Count.

And the other nine?

Filippo.

Sold! but shall I not mount with your lordship’s leave to her
ladyship’s castle, in your lordship’s and her ladyship’s name,
and confer with her ladyship’s seneschal, and so descend again
with some of her ladyship’s own appurtenances?

Count.

Why—no, man. Only see your cloth be clean.

[Exit Filippo.

Lady Giovanna.

Ay, ay, this faded ribbon was the mode

In Florence ten years back. What’s here? a scroll
Pinn’d to the wreath.
My lord, you have said so much
Of this poor wreath that I was bold enough
To take it down, if but to guess what flowers
Had made it; and I find a written scroll
That seems to run in rhymings. Might I read?

Count.

Ay, if you will.

Lady Giovanna.

It should be if you can.

(Reads.) “Dead mountain.” Nay, for who could trace a hand
So wild and staggering?

Count.

This was penn’d, Madonna,

Close to the grating on a winter morn
In the perpetual twilight of a prison,
When he that made it, having his right hand
Lamed in the battle, wrote it with his left.

Lady Giovanna.

Oh heavens! the very letters seem to shake

With cold, with pain perhaps, poor prisoner! Well,
Tell me the words—or better—for I see
There goes a musical score along with them,
Repeat them to their music.

Count.

You can touch

No chord in me that would not answer you
In music.

Lady Giovanna.

That is musically said.

[Count takes guitar. Lady Giovanna

sits listening with wreath in her
hand, and quietly removes scroll
and places it on table at the end
of the song.

Count (sings, playing guitar).

“Dead mountain flowers, dead mountain-meadow flowers,

Dearer than when you made your mountain gay,
Sweeter than any violet of to-day,
Richer than all the wide world-wealth of May,
To me, tho’ all your bloom has died away,
You bloom again, dead mountain-meadow flowers.”

Enter Elisabetta with cloth.

Elisabetta.

A word with you, my lord!

Count (singing).

“O mountain flowers!”

Elisabetta.

A word, my lord! (Louder).

Count (sings).

“Dead flowers!”

Elisabetta.

A word, my lord! (Louder).

Count.

I pray you pardon me again!

[Lady Giovanna, looking at wreath.

Count (to Elisabetta.)

What is it?

Elisabetta.
 My lord, we have but one piece of earthenware to serve the
salad in to my lady, and that cracked!

Count.

Why then, that flower’d bowl my ancestor

Fetch’d from the farthest east—we never use it
For fear of breakage—but this day has brought
A great occasion. You can take it, nurse!

Elisabetta.

I did take it, my lord, but what with my lady’s coming that had
so flurried me, and what with the fear of breaking it, I did break
it, my lord: it is broken!

Count.

My one thing left of value in the world!

No matter! see your cloth be white as snow!

Elisabetta (pointing thro’ window).

White? I warrant thee, my son, as the snow yonder on the

very tip-top o’ the mountain.

Count.

And yet to speak white truth, my good old mother, I have seen
it like the snow on the moraine.

Elisabetta.

How can your lordship say so? There, my lord!

[Lays cloth.

O my dear son, be not unkind to me.

And one word more.