INTRODUCTION

TO

BOTANY
Murray W. Nabors
University of Mississippi

J (

Universitats- und Landasbibliothek Darmstadt Bibliothek Biologie

PEARSON

Benjamin Cnmmings
San Francisco Boston Cape Town Hong Kong London Montreal Munich Paris Singapore New York Madrid Mexico City Sydney Tokyo Toronto

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BRIEF CONTENTS

Chapter 1 The World of Plants The Importance of Plants Plant Characteristics and Diversity Botany and the Scientific Method

Unit Two THE FUNCTIONS OF PLANTS

Chapter 7 Basic Plant Biochemistry The Molecular Components of Living Organisms Energy and Chemical Reactions Chemical Reactions and Enzymes Chapter 8 Photosynthesis An Overview of Photosynthesis Converting Light Energy to Chemical Energy: The Light Reactions Converting CO 2 to Sugars: The Calvin Cycle Chapter 9 Respiration An Overview of Nutrition Respiration Fermentation Chapter 10 Transport in Plants Molecular Movement Across Membranes Movement and Uptake of Water and Solutes in Plants Soil, Minerals, and Plant Nutrition Chapter 11 Plant Responses to Hormones and Environmental Stimuli Effects of Hormones Responses to Light Responses to Other Environmental Stimuli

Unit One THE STRUCTURE OF PLANTS

Chapter 2 Cell Structure and the Cell Cycle An Overview of Cells Major Plant Cell Organelles The Cytoskeleton: Controlling Cell Shape and Movement Membranes and Cell Walls The Cell Cycle and Cell Division Chapter 3 An Introduction to Plant Structure Basic Types of Plant Cells Tissues of Vascular Plants An Overview of Vascular Plant Organs An Overview of Plant Growth and Development Chapter 4 Roots, Stems, and Leaves: The Primary Plant Body Roots Stems . .. Leaves Chapter 5 Secondary Growth in Plants Secondary Growth: An Overview Growth Patterns in Wood and Bark Commercial Uses of Wood and Bark Chapter 6 Life Cycles and Reproductive Structures Plant Reproduction: An Overview Meiosis and Alternation of Generations Cone and Flower Structure Seed Structure Fruit Structure

Unit Three GENETICS AND GENE EXPRESSION

Chapter 12 Genetics Mendel's Experiments on Inheritance Beyond Mendel's Work

xvi

BRIEF CONTENTS Chapter 13 Gene Expression and Activation Gene Expression Differential Gene Expression Identifying Genes That Affect Development Chapter 14 Plant Biotechnology ; The Methods of Plant Biotechnology The Accomplishments and Opportunities of Plant Biotechnology Chapter 22 Gymnosperms An Overview of Gymnosperms Types of Living Gymnosperms Chapter 23 Angiosperms: Flowering Plants Sexual Reproduction in Flowering Plants The Evolution of Flowers and Fruits A Sampling of Angiosperm Diversity

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Unit Four EVOLUTION AND DIVERSITY

Chapter 15 Evolution History of Evolution on Earth Mechanisms of Evolution The Origin of Species Chapter 16 Classification Classification Before Darwin Classification and Evolution Major Groups of Organisms The Future of Classification Chapter 17 Viruses and Prokaryotes Viruses and the Botanical World Prokaryotes and the Botanical World Chapter 18 Algae Characteristics and Evolution of Algae Unicellular and Colonial Algae Multicellular Algae Chapter 19 Fungi Characteristics and Evolutionary History of Fungi ; The Diversity of Fungi Fungal Associations with Other Organisms Chapter 20 Bryophytes An Overview of Bryophytes Liverworts: Phylum Hepatophyta Hornworts: Phylum Anthocerophyta Mosses: Phylum Bryophyta Chapter 21 Seedless Vascular Plants The Evolution of Seedless Vascular Plants Types of Living Seedless Vascular Plants

Unit Five ECOLOGY

Chapter 24 Ecology and the Biosphere Abiotic Factors in Ecology .. . Ecosystems Chapter 25 Ecosystem Dynamics: How Ecosystems Work Populations . Interactions Between Organisms in Ecosystems Communities and Ecosystems .; . Chapter 26 Conservation Biology

Human Population Growth Human Impacts on Ecosystems The Future

Appendix A Basic Chemistry Appendix B Metric Conversions Appendix C Classification of Life Credits Glossary Index

Contents

1 The World of Plants

The Importance of Plants Photosynthesis sustains life on Earth Plants are our fundamental source of food Many medicines come from plants Plants provide fuel, shelter, and paper products Conservation biology is a critical area of research Biotechnology seeks to develop new plant products

1
3 3 4 6 6 7 8

Unit One THE STRUCTURE OF PLANTS 23

Z Cell Structure and the Cell Cycle
An Overview of Cells Microscopes reveal the world of the cell The cell is the basis of an organism's structure and reproduction All cells are either prokaryotic or eukaryotic Cells p r o d u c e nucleic acids, proteins, carbohydrates, and lipids . Major Plant Cell Organelles The nucleus provides DNA "blueprints" for making proteins Ribosomes build proteins The endoplasmic reticulum is the site of most protein and lipid synthesis The Golgi apparatus completes and ships cell products Chloroplasts in green plant cells convert . solar energy into stored chemical energy , Mitochondria convert stored energy into , : energy to power the cell . . Microbodies aid in chemical reactions Vacuoles play a variety of roles in cell metabolism and cell shape The Cytoskeleton: Controlling Cell Shape and Movement . Micro tubules play an important role in cell movements Microfilaments help living cells change shape Motor proteins, or "walking molecules," cause movement Intermediate filaments help determine the permanent structure of cells

Plant Characteristics and Diversity 11 A set of characteristics distinguishes plants from other organisms . "11 Mosses are among the simplest types of plants 12 Ferns and their relatives are examples of seedless vascular plants . 14 Pine trees and other conifers are examples of nonflowering seed plants 15 Most plants are flowering plants with seeds protected in fruits . 1 5 Botany and the Scientific Method Botanists, like other scientists, test hypotheses Botany includes many fields of study Botanists also study algae, fungi, and disease-causing microorganisms Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
PLANTS

25
27 27 27 29 30 31 31 31 32 32 33 34 34 35 36 36 36 36 37

15 15 19 19 20 21 21 21 22 5 7 9 10

& PEOPLE: A Taste of Tea History

T H E INTRIGUING W O R L D OF PLANTS:

Black Pepper: Savior of Rotting Meat

CONSERVATION BIOLOGY: BIOTECHNOLOGY:

The Challenge of Forest Conservation Using Plants to Battle Bacteria

xvin

CONTENTS Membranes and Cell Walls Membranes are gatekeeping barriers around and within cells Cell walls protect plant cells and define cell shape Plasmodesmata are channels that connect plant cells The Cell Cycle and Cell Division The cell cycle describes the phases of a cell's life Mitosis and cell division7are involved ingrowth and reproduction Mitosis produces two daughter nuclei, each containing the same chromosome number as the original cell , Newcells typically become specialized Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
PLANTS

XIX

38 38 39 39 40 41 43

Roots anchor the plant and absorb water and minerals

,,

60 61 61 62 63 . 64 65 65 66 67

43 43 45 46 47 47 47

An Overview of Plant Growth and Development Embryos give rise to stems, leaves, and roots of adult seed plants Meristems enable plants to continue growing throughout their lives . Apical meristems initiate primary growth that makes roots and shoots longer Botanists are discovering how genes control the formation of apical meristems Apical meristems give rise to primary . meristems, which produce primary tissues Secondary growth from lateral meristems makes roots and stems thicker Some plants live for one growing season while others live for two seasons or longer Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More . 6

8 68 69 69

&

PEOPLE:

Pioneers of Microscopy
. .

28 42

THE I N T R I G U I N G W O R L D OF PLANTS:

Using Plant Cell Cultures

T H E I N T R I G U I N G W O R L D OF PLANTS:

3 An Introduction to Plant Structure

Basic Types of Plant Cells Parenchyma cells are the most common type of living differentiated cell Collenchyma cells provide flexible support Sclerenchyma cells provide rigid support Tissues of Vascular Plants The dermal tissue system forms the plant's outer protective covering The vascular tissue system conducts water, minerals, and food Ground tissue usually forms between . dermal and vascular tissues . An Overview of Vascular Plant Organs Stems position leaves for . maximum photosynthesis j Leaves function, in both photosynthesis and transpiration :

Flexible Fibers ' 5 3 PLANTS & P E O P L E : Cotton through the Centuries 55

48
50 50 50 51 52 53 54 58 59 .. 59 60

A Roots, Stems, and Leaves: The Primary Plant Body 70

Roots Taproot systems usually penetrate more deeply than fibrous root systems Root development occurs near the root tip The root cap protects the root apical meristem and helps the root penetrate the soil Absorption of water and minerals occurs mainly through the root hairs The primary structure of roots relates to obtaining water and dissolved minerals Some roots have specialized functions in addition to anchoring the plant and absorbing water and minerals Roots have cooperative relationships with other organisms . 72 72 73 73 74 74

76 77

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CONTENTS 80 81 82 83 83 85 86 87 88 88 89 90 90 91 93 96 97 98 98 98 The vascular cambium produces secondary xylem (wood) and secondary phloem 102 The cork cambium produces secondary dermal tissue 104 Bark consists of all the tissues external . to the vascular cambium ~ 105 Growth Patterns in Wood and Bark 107 The vascular cambium produces secondary xylem, secondary phloem, and ray parenchyma, as well as more vascular cambium 107 Sapwood conducts water and minerals, but heartwood does not .108 Growth rings in wood reflect the history of secondary growth in a tree ' 109 Dendrochronology is the science of tree ring dating and climate interpretation 109 Growth patterns in reaction wood counteract leaning . 110 The cork cambium is reformed as the stems and roots enlarge 110 Lenticels are pathways in the bark for gas exchange 112 Commercial Uses of Wood and Bark Wood is used mainly for fuel, paper products, and construction . Wood structure can be studied from three cutting planes Wood can vary in properties such as hardness and grain ' Latex, resin, and maple syrup are some products from wood fluids Commercial cork comes from the thick outer bark of some trees Trees are a renewable but limited natural resource . . . Summary Review Questions Questions for Thought and Discussion Evolution Connection 112 .112 113 114 115 . . 117 ,117 119 120 120 - 121 121 Bonsai Trees
.

Stems Botanists have developed zone and cell-layer models to describe stem growth In primary growth of most stems, the vascular tissue forms separate bundles " A transition region maintains vascular continuity between the root and stem Leaf primordia form in specific patterns on the sides of shoot apical meristems Stem variations reflect different evolutionary pathways Some stems have specialized functions in addition to support and conduction Leaves " A leaf primordium develops into a leaf through cell division, growth, and differentiation The leaf epidermis provides protection and regulates exchange of gas Mesophyll, the ground tissue in leaves, carries out photosynthesis - The vascular tissue in leaves is arranged in veins Leaf shapes and arrangements have environmental significance Abscission zones form in the petioles of deciduous leaves Some leaves have specialized functions in addition to photosynthesis and transpiration Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
T H E INTRIGUING WORLD OF PLANTS:

'.. Parasitic Roots _ ' PLANTS & PEOPLE: Stems and Roots as Foods EVOLUTION: Leaves That "Eat" Insects

79 87 , 95

5 Secondary Growth in Plants

Secondary Growth: An Overview Lateral meristems, cylinders of dividing cells, produce secondary vascular and secondary dermal tissue

To Learn More

99
101

PLANTS

&

PEOPLE:

102 111 ..116 118

T H E I N T R I G U I N G W O R L D OF P L A N T S :

101

Tree Clues to a Colonial Mystery PLANTS & PEOPLE: Different Roads to Producing Rubber CONSERVATION BIOLOGY: Sustainable Use of Wood Resources

CONTENTS

xxi

6 Life Cycles and Reproductive Structures 122

Plant Reproduction: An Overview 124 Asexual reproduction occurs through mitosis and results in offspring that are genetically / identical to each other and the parent 124 Sexual reproduction results in genetic variation 124 Meiosis and Alternation of Generations Daughter nuclei produced by meiosis have ' one copy of each chromosome Plant sexual life cycles feature both haploid and diploid multicellular forms Cone and Flower Structure In gymnosperms, some apical meristems produce cones In angiosperms, some apical meristems produce flowers A flower can consist of up to four types of modified leaves The number and symmetry of flower parts can vary Flowers can vary in the position of their ovaries Flower structures are examples of how natural selection modifies what is already present Seed Structure Seeds form from ovules on bracts of cones or in carpels of flowers Seeds nourish and protect developing embryos In seed germination, first the embryonic root grows through the seed coat, and then seedling formation begins Fruit Structure During seed development in a flowering plant, the ovary expands to become part or all of a fruit . Fruits can be categorized as simple, aggregate, or multiple A number of mechanisms disperse seeds and fruits to. new locations Summary Review Questions Questions for Thought and Discussion 126 .126 128 130 131 131 132 133 134 134 135 135 135

Evolution Connection To Learn More

BIOTECHNOLOGY:

145 145

Apomixis in Agriculture

137 138

T H E I N T R I G U I N G W O R L D OF PLANTS:

Tropical Fruits

Unit Two THE FUNCTIONS OF PLANTS 147

7 Basic Plant Biochemistry 149
151

135 137

The Molecular Components of Living Organisms Carbohydrates,Vwhich supply and store energy and serve as, structural building blocks, include sugars and polymers of sugars Proteins, which catalyze reactions and are structural building blocks, are polymers of amino acids The nucleic acids DNA and RNA, which code and express genetic information, are polymers of nucleotides Lipids are membrane components consisting mainly of carbon and hydrogen atoms derived from acetates and other molecules Secondary metabolites such as phenolics, alkaloids, and terpenoids often protect or strengthen plants Energy and Chemical Reactions Energy can be stored and can move or change matter Chemical reactions involve either a net input or a net output of free energy Redox reactions release energy as a result of movement of electrons between atoms or molecules ., The terminal phosphate bond in ATP releases energy when broken NADH, NADPH, and FADH2 are universal carriers of energy-rich electrons in living organisms

151

154

157

158

160 163 163 163

137 139 141 143 144 144

164 165

166

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CONTENTS 167 167 Converting CO 2 to Sugars: The Calvin Cycle The Calvin cycle uses ATP and NADPH, from the light reactions to make sugar phosphates from CO 2 The Calvin cycle is relatively inefficient at converting CO 2 into sugars The enzyme rubisco also functions as an oxygenase, resulting in photorespiratiori The C 4 pathway limits the loss of carbon from photorespiration CAM plants store CO 2 in a C 4 acid at night for use in the Calvin cycle during the day Summary Review Questions : 185

Chemical Reactions and Enzymes Collision theory describes product formation by reactions in gases or liquids Enzymes position reactants, allowing reactions to occur with minimal activation energy or increase in temperature Cofactors such as coenzymes interact with enzymes to assist reactions Competitive and noncompetitive inhibition can slow or stop enzymatic reactions and pathways Enzymatic reactions are linked together into metabolic pathways Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
T H E I N T R I G U I N G W O R L D OF PLANTS:

185 187 188 189 191 192 193 194 194 194

168 170

170 172 172 173 173 174 174

Questions for Thought and Discussion Evolution Connection To Learn More

THE INTRIGUING WORLD OF PLANTS:

Forests Made of Carbohydrates ' 155 BIOTECHNOLOGY: Weapons Against Weeds 157 PLANTS & PEOPLE: Take Your Cofactors Every Day 171

Nonphotosynthetic Plants EVOLUTION: Evolution and O 2 Concentration

178 189

9 Respiration 8 Photosynthesis
An Overview of Photosynthesis Photosynthesis produces food, molecular building blocks, and O 2 , which support almost all life on Earth ' Photosynthesis'uses light energy to convert CO 2 and H 2 O into sugars The processes of photosynthesis and respiration are interdependent ....... Converting Light Energy to Chemical Energy: The Light Reactions Chlorophyll is the principal light-absorbing molecule of photosynthesis Light energy enters photosynthesis at locations called photosystems The light reactions produce O 2 , ATP, and NADPH . In the light reactions, ATP is synthesized using energy from chemiosmosis

195
197 197

175
177

177 178 179 180 180 182 183

An Overview of Nutrition All living organisms need sources of energy and carbon Plants use photosynthesis to store light energy in sugars and use respiration to transfer the energy from sugars to ATP . The breakdown of sugar to release energy can occur with or without oxygen Respiration Glycolysis splits six-carbon sugars into two molecules of pyruvate The Krebs cycle generates CO 2 , NADH, FADH2, andATP / The electron transport chain and oxidative phosphorylation transfer energy from the energy-rich electrons of NADH and FADH 2 toATP i The energy yield from respiration is high In some plants, the electron transport chain can generate excess heat . . . Plants, unlike animals, can make fatty acids into glucose

197 198 200 200 202

;202

205
206

184

207

CONTENTS Fermentation , In the absence of oxygen, pyruvate produced by glycolysis is converted to ethanol or lactate Some important industries rely on fermentation Fermentation has a low energy yield compared to that of respiration
Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More CONSERVATION BIOLOGY:
;

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208 208 208 209

210

Plants require 17 essential elements, . ; < most of which are obtained from soil Soil particles bind water and mineral ions Bacteria in the soil make nitrogen available to plants Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
THE INTRIGUING WORLD OF PLANTS:

227 229 230 232 233 233 234 234

211 211 212 . 2 1 2

The Power of Plants

200 204
:

Global Warming and the Greenhouse Effect

PLANTS & PEOPLE:

BIOTECHNOLOGY: Water-Efficient Crops PLANTS & PEOPLE: lustus von Liebig

219 221 228

Sucrose and Fructose: Sweeteners of Choice

THE INTRIGUING WORLD OF PLANTS:

A Father of Modern Agriculture

Skunk Cabbage

207

10 Transport in Plants
Molecular Movement Across Membranes Diffusion is the spontaneous movement of molecules down a concentration gradient Facilitated diffusion and active transport use proteins to assist in movement across membranes Exocytosis and endocytosis transport large molecules Osmosis is the movement of water across. . a selectively permeable membrane In plant cell growth, the osmotic potential inside the cell interacts with pressure generated by the cell wall - .'.. Movement and Uptake of Water and Solutes in Plants Water evaporation from leaves pulls water through the xylem from the roots Stomata control gas exchange and water loss for the plant . Sugars and other organic molecules move from leaves to roots in the phloem Soil, Minerals, and Plant Nutrition ; Soil is made of ground-up particles of rocks surrounded by negative charges that bind water and minerals

213
215 215

1 1 Plant Responses to Hormones and Environmental Stimuli

235

216 216 216

217 219 219 223 224 . . 226

Effects of Hormones 237 Auxin plays a central role in cell enlargement and formation of new tissue 237 Cytokinins control cell division and differentiation and also delay aging 240 Gibberellins interact with auxins to regulate cell enlargement and stimulate seed germination 240 Abscisic acid causes seed dormancy and - regulates plant responses to drought . . . . 242 Ethylene allows the plant to respond to mechanical stress and controls fruit ripening and leaf abscission 243 Brassinosteroids are a newly discovered group of plant hormones that act like auxin 244 Additional compounds may play a role as plant hormones 244 Responses to Light 245 . Blue light absorption controls the growth of stems toward the light and the opening of stomata 245 Absorption of red and far-red light determines when seed germination, stem and root growth, and flowering occur 245

227

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CONTENTS Dihybrid crosses involve individuals that have different alleles for two specific genes ' Beyond Mendel's Work Mendel's laws also apply to crosses that involve more than two traits Some characters are not controlled by one dominant and one recessive allele The locations of genes affect inheritance patterns Genes interact with each other and with the environment Mendel's gene for height in peas controls the production of a growth-promoting hormone Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More & P E O P L E : A Brief Biography of Gregor Mendel PLANTS & P E O P L E : Genetics Before Mendel
PLANTS T H E I N T R I G U I N G WORLD OF PLANTS:

Photoperiodism regulates flowering and other seasonal responses Plants respond to repeating cycles of day and night .

246 249 250 250 251 251 253 254 255 256 256 257 257

266 268 269 269 272 273 274 275 276 276 276 276

Responses to Other Environmental Stimuli Roots and shoots respond to gravity Plants respond to mechanical stimuli, such as touch and wind ( Plants prepare for environmental conditions that prevent normal metabolism and growth Plants react to environmental stresses such as drought Plants deter herbivores and pathogens Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
BIOTECHNOLOGY:

Effects of Auxin and Cytokinins on Cultured Plant Cells Studying Photodormant Seeds The Arms Race Between Plants and Herbivores

241 247 253

T H E I N T R I G U I N G W O R L D OF PLANTS: EVOLUTION:

263 268 270

A Weed with Great Potential

Unit Three GENETICS AND GENE EXPRESSION 259

1 2 Genetics
Mendel's Experiments on Inheritance Making sense of Mendel's experiments requires a basic understanding of genes and chromosomes Monohybrid crosses involve individuals that have different alleles for a specific gene Segregation of alleles occurs during anaphase I of meiosis ' A testcross demonstrates the genotype of an individual with a dominant phenotype

1 3 Gene Expression and Activation

277
279 279 280 283 284 285 288 288 289 289 292

261
263

Gene Expression During replication, DNA is copied DNA codes for the structure of proteins During transcription, RNA is made from DNA During translation, a protein is made from messenger RNA Mutations can cause changes in gene expression Differential Gene Expression Gene expression is controlled at various levels Regulatory proteins control transcription Hormones and light can trigger the activation of transcription factors Identifying Genes That Affect Development Experiments on Arabidopsis illustrate the use of mutations to understand plant development ..

263 264 266 266

'

292

CONTENTS Transposons can be used to locate genes that affect development Homeotic genes control development in plants and animals
Summary Review Questions . -.'" i

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294 296 297 298 298 298 298

Genomics and proteomics will provide information needed for future efforts in genetic engineering Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
BIOTECHNOLOGY:

314 317 318 318 319 319

Questions for Thought and Discussion Evolution Connection To Learn More

THE I N T R I G U I N G W O R L D OF PLANTS:

Genetic Engineering of Salt-Tolerant Plants

310 312 315 316

The Closing of Stomata in Response to Drought Is a Typical STP . BIOTECHNOLOGY: DNA Microarrays

T H E I N T R I G U I N G W O R L D OF PLANTS:

291 293

Wide Crosses Between Plants BIOTECHNOLOGY: DNA Sequencing

BIOTECHNOLOGY: Analyzing DNA

Fragments and Solving Crimes

\4 Plant Biotechnology 299

The Methods of Plant Biotechnology Genes can be transferred between species through genetic engineering Plasmids often serve as vectors for gene transfer in plants Restriction enzymes and DNA ligase are used to make recombinant DNA Cloning produces multiple copies of recombinant DNA The polymerase chain reaction clones DNA without using cells Several methods can be used to insert cloned genes into plant cells In tissue culture, whole plants are grown from isolated cells or tissues r The Accomplishments and Opportunities of Plant Biotechnology Genetic engineering has made plants that are more resistant to pests and harsh soil conditions as well as more productive Transgenic plants contribute to human health and nutrition Genetically engineered crops require extensive field and market testing before they are released ' . Genetically engineered crops must be safe for the environment and for consumers The future holds many opportunities for plant biotechnology
. l-

301 301 302 302 303 303 304 306 307

Unit Four EVOLUTION AND DIVERSITY 321

15 Evolution
History of Evolution on Earth Fossils and molecular dating provide evidence of evolution Biogeography, anatomy, embryology, and physiology supply further evidence of evolution Chemosynthesis may have been the first event in the origin of life on Earth Prokaryotes were the predominant form of life for more than a billion years Plate tectonics and celestial cycles have shaped evolution on Earth - Extinction is a fact of life on Earth Mechanisms of Evolution Evolution is a change in the frequency of alleles in a population over time Most organisms have the potential to overproduce offspring Individuals in a population have many phenotypic differences

323
325 325

. : 327 327 328

307 309

329 331
332 332 334 335

311 .311 313

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CONTENTS 335 336 338 339 Major Groups of Organisms .360 Systematists have revised the number of . kingdoms . 360 Molecular data have led to identifying "super kingdoms" called domains 361 The domain Archaea and the domain Bacteria are two very different groups of prokaryotes 361 The domain Eukarya includes protists, animals, fungi, and plants 362 The Future of Classification New species remain to be discovered Systematists are studying speciation in action Molecular data will continue to provide insights into evolution ' The classification of organisms ' has practical benefits Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More 363 363 364 366 366 366 368 368 369 369

Some traits confer an adaptive advantage Natural selection favors individuals with the best-adapted phenotypes Evolution can occur rapidly In coevolution, two species evolve in response to each other ,

The Origin of Species 340 A biological species is a population of potentially interbreeding organisms 340 Both natural selection and geographical isolation drive speciation 342 Reproductive isolation can be prezygotic or postzygotic 342 Reproductive isolation in sympatric populations can occur because of polyploidy 343 Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
PLANTS

344 345 345 346 346

& PEOPLE: Germinating an Idea: . Evolution by Means of Natural Selection

326 336 339

THE INTRIGUING WORLD OF PLANTS: PLANTS

T H E INTRIGUING W O R L D OF P L A N T S :

Artificial Selection EVOLUTION: Plants of the Galapagos Islands

What's in a Plant Name? & PEOPLE: Linnaeus and the Lure of Plants

351 352

1 6 Classification
Classification Before Darwin ^ Classification of organisms dates back to ancient times Linnaeus laid the foundation for modern naming of species Classification and Evolution Systematists use a variety of characters to classify organisms Molecular data play a key role in phylogenetic classification Organisms are classified into a hierarchy Systematists form hypotheses about evolutionary relationships Cladograms are branching diagrams that show evolutionary relationships Systematists often disagree about how to classify organisms

347
349 349 350 351 353 354 354 356 358 359

1 7 Viruses and Prokaryotes

Viruses and the Botanical World Viruses are complexes of nucleic acid and protein that reproduce inside cells Viruses cause many important plant diseases Several approaches are used to prevent viral diseases in plants Viroids are infectious RNA molecules Prokaryotes and the Botanical World Prokaryotes are unicellular organisms with diverse characteristics ';'Some bacteria are photosynthetic, and some fix nitrogen Bacteria cause a variety of diseases in plants

370
372 372 374 375 376 376 376 377 379

CONTENTS Prokaryotes have many uses in industry, medicine, and biotechnology Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
PLANTS & PEOPLE: The Discovery of

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Summary 380 381 382 382 382 382

T H E I N T R I G U I N G W O R L D OF PLANTS: BIOTECHNOLOGY:

401 402 402 403 403

Review Questions Questions for Thought and Discussion Evolution Connection To Learn More

Watermelon Snow Algae as a Source of Fuel

397 399

Viruses in Tobacco
CONSERVATION BIOLOGY:

372 Cryptobiotic Crust 378

:

and Desert Varnish

THE I N T R I G U I N G W O R L D OF P L A N T S :

19 Fungi

404

The Huge Importance of Tiny Photosynthesizers 379

18Algae

384
386 386 387 388 388 389 391

Characteristics and Evolutionary History of Fungi 406 A combination of morphological and developmental characteristics distinguish fungi from other organisms 406 Fungi probably evolved from flagellated protists 407 The Diversity of Fungi Chytridiomycetes (phylum Chytridiomycota) produce flagellated reproductive cells Zygomycetes (phylum Zygomycota) form resistant zygosporangia prior to meiosis Ascomycetes (phylum Ascomycota) produce sexual spores in sacs called asci . Basidiomycetes (phylum Basidiomycota). produce sexual spores on club-shaped cells called basidia Fungal Associations with Other Organisms Lichens are associations of fungi and photosynthetic algae or bacteria Some fungi form mutualistic associations with insects Summary Review Questions 408 408 408 411

Characteristics and Evolution of Algae Algae are distinguished by their photosynthetic pigments and other characteristics Endosymbiosis played a key role in the evolution of algae Unicellular and Colonial Algae Euglenoids (phylum Euglenophyta) have a pellicle beneath the plasma membrane Many dinoflagellates (phylum Dinophyta) have hard cellulose plates Diatoms (phylum Bacillariophyta) form cell walls of silica Yellow-green algae (phylum Xanthophyta) are important members of freshwater phytoplankton Golden-brown algae (phylum Chrysophyta) form unique, dormant spores Cryptomonads (phylum Cryptophyta) use ejectisomes for sudden escape Haptophytes (phylum Prymnesiophyta) ; have a distinctive, moveable haptonema MulticelUular Algae In many brown algae (phylum Phaeophyta), alternate'generations are heteromorphic Red algae (phylum Rhodophyta) have complex life cycles with three multicellular phases Green algae (phylum Chlorophyta) share a common ancestor with plants

415 419 420 421 422 423 423 423 423

392 392 393 393 394 394 395 397

Questions for Thought and Discussion Evolution Connection To Learn More

T H E I N T R I G U I N G W O R L D OF PLANTS:

Fungi That Live on Dung

CONSERVATION BIOLOGY:

Dutch Elm Disease

410 414
417

PLANTS & PEOPLE:.Growing Mushrooms

XXVU1

CONTENTS

2 0 Bryophytes
An Overview of Bryophytes Bryophytes were among the first land plants Bryophytes have many similarities to green algae in the class Charophyceae and to vascular plants In bryophytes, alternation of generations involves a dominant gametophyte and attached sporophyte .. . Bryophytes play important ecological roles " Many bryophyte species tolerate drought conditions Liverworts: Phylum Hepatophyta Liverwort gametophytes can be either -thalloid or leafy A liverwort life cycle demonstrates dominance of the gametophyte Hornworts: Phylum Anthocerophyta The hornwort life cycle features a hornlike sporophyte The evolutionary history of hornworts, as with other bryophytes, is being debated Mosses: Phylum Bryophyta . There are three main classes of mosses The life cycle of Polytrichum demonstrates characteristic features of mosses Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
EVOLUTION:

424
426 426

427

Land plants arose from green algae in the class Charophyceae Three phyla of extinct vascular plants appear in the fossil record beginning 430 million years ago _ In living seedless vascular plants, alternation of generations involves independent gametophytes and sporophytes . Types of Living Seedless Vascular Plants Whisk ferns comprise most of the living members of phylum Psilotophyta Living members of phylum Lycophyta include club mosses, spike mosses, and quillworts Horsetails are the living members of phylum Sphenophyta Phylum Pterophyta consists of ferns, the largest group of seedless vascular plants Summary Review Questions Questions for Thought and Discussion

443

444

446 448 448 449 453 454 459 460 460 460 460

428 429 430 430 430 431 433 .433 433 434 435 .436 438 439 . 439 439 439 427 436

Evolution Connection To Learn More

T H E I N T R I G U I N G W O R L D OF P L A N T S :

Alternative Life Cycles EVOLUTION: Telomes and Origins of Sporangia

448 456

2 2 Gymnosperms

461
.463 463 465

Bogs

T H E I N T R I G U I N G W O R L D OF P L A N T S :

Unusual Mosses

An Overview of Gymnosperms Seed plants have significant selective advantages Living gymnosperms are related to extinct plants from the Paleozoic and Mesozoic erasIn gymnosperms and other seed plants, dependent gametophytes develop within the parent sporophyte The pine life cycle illustrates basic features of gymnosperm reproduction .

466 466

2 1 Seedless Vascular Plants

440

The Evolution of Seedless Vascular Plants 442 Seedless vascular plants dominated the landscape around 350 million years ago 442

Types of Living Gymnosperms 469 Phylum Coniferophyta contains conifers, which are the dominant forest trees in cooler climates 470 Phylum Cycadophyta contains cycads, which resemble tree ferns or palms 474 Phylum Ginkgophyta contains one living species 475 Phylum Gnetophyta contains three diverse. genera found in tropical forests or in deserts 476

CONTENTS Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
BIOTECHNOLOGY:

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r-

477 478 478 479 479

Unit Five ECOLOGY

2 4 Ecology and the Biosphere

503
505
507 507 508 511 513 513 513 515 518 521 522 . 523 523 523

Improving and Protecting Trees 472 474

THE INTRIGUING W O R L D OF PLANTS:

The Wollemi Pine: A Living Fossil -

2 3 Angiosperms: Flowering Plants

Sexual Reproduction in Flowering Plants 482 Angiosperms, like gymnosperms, have a dominant sporophyte and dependent gametophyte 482 Self-pollination and cross-pollination are both common in angiosperms 484 The Evolution of Flowers and Fruits The selective advantages of flowering plants account for their success Flowers have evolved as collections of highly modified leaves The evolution of angiosperms began during the Mesozoic era , During the Cretaceous period, angiosperms. spread rapidly throughout.the world A Sampling of Angiosperm Diversity Phylum Anthophyta contains more than 450 families, classified mainly by flower structure Several families illustrate the diversity of floral and fruit structure Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More Superweeds EVOLUTION: The Origins of Domesticated Corn, Wheat, and Rice
BIOTECHNOLOGY: THE INTRIGUING W O R L D OF PLANTS:

Abiotic Factors in Ecology Abiotic factors are physical variables in an organisms's environment The tilt of Earth's axis causes seasons and affects temperatures The atmosphere circulates in six global cells The rotation and topography of Earth affect global patterns of wind and precipitation Ecosystems The biosphere can be divided into biogeographic realms and biomes Terrestrial biomes are characterized by rainfall, temperature, and vegetation Light penetration, temperature, and nutrients are important abiotic factors in aquatic biomes Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More

486 487 488 488 494 494

495 497 500 501 501 502 502 485 496 498

T H E I N T R I G U I N G W O R L D OF PLANTS: CONSERVATION BIOLOGY:

Weeds El Nino and La Nina

509 512

2 5 Ecosystem Dynamics: How Ecosystems Work 524

Populations The reproductive characteristics of plants create challenges in studies of plant populations The distribution of plants in a population may be random, uniform, or clumped 526 526 526

A Recently Discovered Orchid

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CONTENTS

Age distributions and survivorship curves describe the age structure of populations The growth of populations over time is limited by environmental resources The growth of plant populations depends on reproductive patterns Interactions Between Organisms in Ecosystems Commensalism and mutualism are interactions in which at least one species benefits Predation, herbivory, and parasitism are interactions in which at least one species is harmed Plants compete for resources with members of their own and other species ' ' Communities and Ecosystems Communities can be characterized by species composition and by vertical and horizontal species distribution Apparently uniform environments are often . composed of different microenvironments A moderate level of disturbance can increase the number of species in an ecosystem Ecological succession describes, variation in communities over time The energy stored in photosynthetic organisms passes inefficiently to other organisms in the same ecosystem Biological magnification increases the concentration of some toxic substances at higher trophic levels Water and nutrients cycle between biotic and abiotic components of ecosystems Human activity has fragmented stable ecosystems into distinctive patches Summary Review Questions Questions for Thought and Discussion Evolution Connection To Learn More
T H E I N T R I G U I N G W O R L D OF P L A N T S : CONSERVATION BIOLOGY:

527 528 529 531 531 531 532

2 6 Conservation Biology 548

Human Population Growth 550 Human population is increasing exponentially 550 Increased food production will involve genetically altered plants, improved growing practices, and more efficient food distribution systems 550 Human Impacts on Ecosystems The presence and activities of large human populations disturb ecosystems The geographic information system provides a new tool to record changes in ecosystems The Future The future of human interactions with ecosystems can be modeled on worst-case or best-case scenarios Achieving a best-case scenario for the biosphere would involve a marked reversal of current trends A number of problems would have to be overcome to reverse current trends of ecosystem destruction and modification It is important to establish models of success in promoting ecosystem restoration Summary Review Questions 552 552 560 562

534

534 536 537 537

562

563

564 '< 564 567 568 568 568 569

540

541 542 543 545 546 546 547 547

Questions for Thought and Discussion Evolution Connection To Learn More

CONSERVATION BIOLOGY:

Land Races 553 559 565 570 577 578 580 583 602

and Seed Banks

T H E I N T R I G U I N G W O R L D OF PLANTS:

Kudzu BIOTECHNOLOGY: Recreating Lost Worlds Through Genomics and Proteomics

Appendix A Basic Chemistry Appendix B Metric Conversions Appendix C Classification of Life Credits Glossary Index

Ant Plants 532 Figs in the Forest 535 EVOLUTION: Primary Succession after a Volcanic Eruption 538