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Starr
Evers
Starr
Biology
Concepts & Applications
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10e
Biology
Concepts & Applications 10e Cecie Starr
Christine A. Evers
Lisa Starr
Copyright 2018 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203
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Biology
Concepts & Applications 10e
About the Cover Photo
Portuguese Man-of-War
Anyone unfamiliar with the biology of the
venomous Portuguese man-of-war would likely
mistake it for a jellyfish. Not only is it not a
jellyfish, it’s not even an “it,” but a “they.”
The Portuguese man-of-war is a siphonophore,
an animal made up of a colony of organisms
working together.
The man-of-war comprises four separate polyps.
It gets its name from the uppermost polyp, a
gas-filled bladder, or pneumatophore, which
sits above the water and somewhat resembles
an old warship at full sail. A man-of-war is also
known as a bluebottle for the purple-blue color
of its pneumatophores.
The tentacles are the man-of-war’s second
organism. These long, thin tendrils can extend
165 feet (50 meters) in length below the
surface, although 30 feet (10 meters) is more
the average. They are covered in venom-filled
nematocysts used to paralyze and kill fish and
other small creatures. For humans, a man-of-
war sting is excruciatingly painful, but rarely
deadly. But beware—even a dead man-of-war
washed up on shore can deliver a sting.
Muscles in the tentacles draw prey up to a
polyp containing the gastrozooids or digestive
organisms. A fourth polyp contains the
reproductive organisms.
A man-of-war is found, sometimes in groups
of 1,000 or more, floating in warm waters
throughout the world’s oceans. They have no
independent means of propulsion and either
drift on the currents or catch the wind with
their pneumatophores. To avoid threats on
the surface, they can deflate their air bags and
briefly submerge.
Australia • Brazil • Japan • Korea • Mexico • Singapore • Spain • United Kingdom • United States
Biology
Cecie Starr
Christine A. Evers
Lisa Starr
Biology: Concepts & Applications, © 2018, 2015, 2011 Cengage Learning
Tenth Edition
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Print Number: 01 Print Year: 2016
Contents in Brief
INTRODUCTION
1 The Science of Biology 2
UNIT 1 PRINCIPLES OF CELLULAR LIFE
2 Life’s Chemical Basis 22

3 Molecules of Life 36
4 Cell Structure 50
5 Ground Rules of Metabolism 76
6 Where It Starts—Photosynthesis 98
7 Releasing Chemical Energy 112
UNIT 2 GENETICS UNIT 6 HOW ANIMALS WORK
8 DNA Structure and Function 130 28 Animal Tissues and Organ Systems 480
9 From DNA to Protein 146 29 Neural Control 498
10 Control of Gene Expression 162 30 Sensory Perception 520
11 How Cells Reproduce 176 31 Endocrine Control 536
12 Meiosis and Sexual Reproduction 190 32 Structural Support and Movement 554
13 Patterns in Inherited Traits 204 33 Circulation 570
14 Human Inheritance 220 34 Immunity 590
15 Biotechnology 236 35 Respiration 614
36 Digestion and Human Nutrition 630
UNIT 3 P R I N C I P L E S O F E VO L U T I O N
37 Maintaining the Internal Environment 650
16 Evidence of Evolution 252 38 Reproduction and Development 666
17 Processes of Evolution 272
UNIT 7 PRINCIPLES OF ECOLOGY
18 Life’s Origin and Early Evolution 300
39 Animal Behavior 696
UNIT 4 E VO L U T I O N A N D B I O D I V E R S I T Y 40 Population Ecology 712
19 Viruses, Bacteria, and Archaea 314 41 Community Ecology 730
20 The Protists 330 42 Ecosystems 748
21 Plant Evolution 346 43 The Biosphere 764
22 Fungi 364 44 Human Effects on the Biosphere 784
23 Animals I: Major Invertebrate Groups 376 Appendix I 800
24 Animals II: The Chordates 400 Appendix II 801
Appendix III 802
UNIT 5 HOW PLANTS WORK Appendix IV 805
Appendix V 806
25 Plant Tissues 422 Appendix VI 808
26 Plant Nutrition and Transport 440 Appendix VII 808
27 Reproduction and Development Glossary 812
of Flowering Plants 454 Index 832
Detailed Contents
INTRODUCTION
1 THE SCIENCE OF BIOLOGY

1.1 Life Is More Than the Sum of Its Parts 4
Emergent Properties 4
Life’s Organization 4
1.2 Life’s Unity 6
Energy and Nutrients 6
Homeostasis 7
DNA Is Hereditary Material 7
1.3 Life’s Diversity 8
● Engage: Kristofer Helgen, National Geographic Explorer 9
1.4 Organizing Information About Species 10

A Rose by Any Other Name . . . 10
Distinguishing Species 10
1.5 The Science of Nature 12
Thinking About Thinking 12
The Scientific Method 12
1.6 Examples of Experiments in Biology 14
Research in the Real World 14
Potato Chips and Stomachaches 14
Butterflies and Birds 14
1.7 Analyzing Experimental Results 16
Sampling Error 16
Bias 17
Objectivity 17
1.8 The Nature of Science 18
What Science Is 18
What Science Is Not 18
APPLICATION: The Secret Life of Earth 19
UNIT 1 PRINCIPLES OF CELLULAR LIFE
2 LIFE’S CHEMICAL BASIS

2.1 Building Blocks of Matter 24
Atoms and Elements 24
Isotopes and Radioisotopes 24
● Engage: Kenneth Sims, National Geographic Explorer 25
2.2 Why Electrons Matter 26

Electrons Have Energy 26
The Importance of Vacancies 27
2.3 From Atoms to Molecules 28
Chemical Bonds 28
Ionic Bonds 28
Covalent Bonds 29
2.4 Hydrogen Bonding and Water 30
Hydrogen Bonds 30
Enric Sala/National Geographic Creative
Water’s Special Properties 30 General Features of Eukaryotic Cells 60
Water Is an Excellent Solvent 30 The Nucleus 61
Water Has Cohesion 31
4.6 The Endomembrane System 62
Water Stabilizes Temperature 31
A Variety of Vesicles 62
2.5 Acids and Bases 32
Endoplasmic Reticulum 62
Hydrogen Atoms and pH 32
Golgi Bodies 63
APPLICATION: Mercury Rising 33 4.7 Mitochondria 64
4.8 Chloroplasts and Other Plastids 65
3 MOLECULES OF LIFE 4.9 The Cytoskeleton 66
3.1 The Chemistry of Biology 38

Cytoskeletal Elements 66
Microtubules 66
Carbon, the Stuff of Life 38 Microfilaments 66
Metabolic Reactions 39 Intermediate Filaments 66
3.2 Carbohydrates 40 Cellular Movement 66
Carbohydrates in Living Systems 40 4.10 Cell Surface Specializations 68
Simple Sugars 40 Extracellular Matrices 68
Short-Chain Carbohydrates 40
Polysaccharides 40 Cell Junctions 69
4.11 The Nature of Life 70
3.3 Lipids 42
Lipids in Living Systems 42 Life Is Squishy 70
Fats 42 ● Engage: Dr. Kevin Peter Hand,
Phospholipids 43 National Geographic Explorer 71
Waxes 43
Steroids 43 APPLICATION: Food for Thought 71
3.4 Proteins 44
Building Proteins 44
From Structure to Function 44
Primary and Secondary Structure 44
Tertiary and Quaternary Structure 44
Finishing Touches 45
3.5 The Importance of Protein Structure 46
3.6 Nucleic Acids 47
APPLICATION: Fear of Frying 47
4 CELL STRUCTURE
4.1 What, Exactly, Is a Cell? 52
Cell Theory 52
Components of All Cells 52
Constraints on Cell Size 53
4.2 How We See Cells 54
● Engage: Dr. Aydogan Ozcan,
National Geographic Explorer 55
4.3 Cell Membrane Structure 56
The Fluid Mosaic Model 56
Proteins Add Function 56
4.4 Introducing the Prokaryotes 58
Structural Features 58
Biofilms 59
4.5 Introducing the Eukaryotic Cell 60
vii
Detailed Contents (continued)
5 GROUND RULES OF METABOLISM

5.1 Life Runs on Energy 78
Energy Disperses 78
Energy’s One-Way Flow 78
5.2 Energy in the Molecules of Life 80
Chemical Bond Energy 80
Why Earth Does Not Go Up in Flames 80
Energy In, Energy Out 81
5.3 How Enzymes Work 82
The Need for Speed 82
Influences on Enzyme Activity 83
5.4 Metabolic Pathways 84
Controls Over Metabolism 84 6.2 Sunlight as an Energy Source 102
Electron Transfers 85 Visible Light Drives Photosynthesis 102
To Catch a Rainbow 102
5.5 Cofactors 86
ATP—A Special Coenzyme 87 6.3 Light-Dependent Reactions 104
The Noncyclic Pathway 104
5.6 Diffusion and Membranes 88
The Cyclic Pathway 105
Factors That Affect Diffusion 88
Concentration 88 6.4 Light-Independent Reactions 106
Temperature 88 The Calvin–Benson Cycle 106
Size 88 Photorespiration 107
Charge 88
Pressure 88 6.5 Carbon-Fixing Adaptations of Plants 108
Osmosis 88 C4 Plants 108
Turgor 89 CAM Plants 108
5.7 Membrane Transport Mechanisms 90 APPLICATION: A Burning Concern 109
Transport Protein Specificity 90 ● Engage: Sanga Moses, National Geographic Explorer 109
Passive Transport 90
Active Transport 91
7 RELEASING CHEMICAL ENERGY
5.8 Membrane Trafficking 92
Vesicle Movement 92 7.1 Introduction to Carbohydrate Breakdown
Membrane Recycling 93 Pathways 114
Reaction Pathways 115
APPLICATION: A Toast to Alcohol Dehydrogenase 94
7.2 Glycolysis—Sugar Breakdown Begins 116
Glycolysis Reactions 116
Comparing Other Pathways 116
7.3 Acetyl–CoA Formation and the Citric Acid Cycle 118
7.4 Aerobic Respiration’s Big Energy Payoff 120
Electron Transfer Phosphorylation 120
Summing Up 121
7.5 Fermentation Pathways 122
Alcoholic Fermentation 122
Lactate Fermentation 123
7.6 Food as a Source of Energy 124
6 WHERE IT STARTS—PHOTOSYNTHESIS
Energy From Dietary Molecules 124
6.1 Overview of Photosynthesis 100 Fats 124
Autotrophs and Heterotrophs 100 Complex Carbohydrates 124
Proteins 124
Two Stages of Reactions 100
Where Photosynthesis Occurs 101 APPLICATION: Mitochondrial Malfunction 126
viii
10.3 Details of Body Form 168
UNIT 2 GENETICS
10.4 Gene Expression in Metabolic Control 170
8 DNA STRUCTURE AND FUNCTION The lac Operon 170
Lactose Metabolism in Mammals 170
8.1 The Discovery of DNA’s Function 132
Circadian Rhythms in Plants 171
Early Clues 132
10.5 Epigenetics 172
A Surprising Result 132
Final Pieces of Evidence 133 APPLICATION: Between You and Eternity 173
8.2 Discovery of DNA’s Structure 134
Building Blocks of DNA 134
DNA Sequence 135
8.3 Eukaryotic Chromosomes 136
Chromosome Structure 136
Chromosome Number 136
● Engage: Mariana Fuentes,
8.4 How Does a Cell Copy Its DNA? 138 1 1 HOW CELLS REPRODUCE
Semiconservative Replication 138 11.1 Multiplication by Division 178
Directional Synthesis 139 The Cell Cycle 178
8.5 Mutations and Their Causes 140 Mitosis Maintains the Chromosome Number 178
● Engage: Dr. Rosalind Franklin 141 Control Over the Cell Cycle 179
8.6 Cloning Adult Animals 142 Why Cells Divide by Mitosis 179
11.2 A Closer Look at Mitosis 181
APPLICATION: A Hero Dog’s Golden Clones 143
11.3 Cytoplasmic Division 182
11.4 Marking Time With Telomeres 183
9 FROM DNA TO PROTEIN
11.5 Pathological Mitosis 184
9.1 Introducing Gene Expression 148 The Role of Mutations 184
● Engage: Dr. John “Jack” Horner, Cancer 185
National Geographic Explorer 149 APPLICATION: Henrietta’s Immortal Cells 186
9.2 Transcription: DNA to RNA 150 ● Engage: Dr. Iain Couzin, National Geographic Explorer 187
Post-Transcriptional Modifications 151
9.3 RNA and the Genetic Code 152 12 MEIOSIS AND SEXUAL REPRODUCTION
The Messenger: mRNA 152
12.1 Why Sex? 192
The Translators: rRNA and tRNA 153
Introducing Alleles 192
9.4 Translation: RNA to Protein 154
On the Advantages of Sex 192
9.5 Consequences of Mutations 156
● Engage: Maurine Neiman,
APPLICATION: The Aptly Acronymed RIPs 158 National Geographic Explorer 193
12.2 Meiosis in Sexual Reproduction 194
10 CONTROL OF GENE EXPRESSION Meiosis Halves the Chromosome Number 194
How Meiosis Works 194
10.1 How Cells Control Gene Expression 164 Fertilization Restores the Chromosome Number 195
Switching Genes On and Off 164
12.3 A Visual Tour of Meiosis 196
DNA–Histone Interactions 164
Transcription 164 12.4 How Meiosis Introduces Variations in Traits 198
RNA Processing 165 Crossing Over 198
RNA Transport 165 Chromosome Segregation 199
Translation 165
Post-Translational Modification 165 12.5 Mitosis and Meiosis—An Ancestral Connection? 200
Protein Degradation 165 APPLICATION: How to Survive for 80 Million Years
10.2 Orchestrating Early Development 166 Without Sex 201
ix
1 3 PATTERNS IN INHERITED TRAITS 1 5 BIOTECHNOLOGY

13.1 Mendel, Pea Plants, and Inheritance Patterns 206 15.1 Cutting and Pasting DNA 238
Mendel’s Experiments 206 DNA Cloning 238
Inheritance in Modern Terms 207 Why Clone DNA? 239
13.2 Mendel’s Law of Segregation 208 15.2 Isolating Genes 240
13.3 Mendel’s Law of Independent Assortment 210 DNA Libraries 240
The Contribution of Crossovers 211 PCR 240
13.4 Beyond Simple Dominance 212 15.3 DNA Sequencing 242
Incomplete Dominance 212 The Human Genome Project 243
Codominance 212 15.4 Genomics 244
Epistasis 213
DNA Profiling 245
Pleiotropy 213
15.5 Genetic Engineering 246
13.5 Nature and Nurture 214
Genetically Modified Organisms 246
● Engage: Dr. Gay Bradshaw 215 Microorganisms 246
13.6 Complex Variation in Traits 216 Plants 246
Animals 247
APPLICATION: Menacing Mucus 217
15.6 Editing Genomes 248
Gene Therapy 248
1 4 HUMAN INHERITANCE CRISPR 248
14.1 Human Chromosomes 222 APPLICATION: Personal Genetic Testing 249
● Engage: Dr. Nancy Wexler 223
14.2 Examples of Autosomal Inheritance Patterns 224 UNIT 3 P R I N C I P L E S O F E VO L U T I O N
Autosomal Dominance 224 1 6 EVIDENCE OF EVOLUTION

Achondroplasia 224
Huntington’s Disease 224 16.1 Early Beliefs, Confusing Discoveries 254
Hutchinson–Gilford Progeria 225
The Autosomal Recessive Pattern 225 16.2 A Flurry of New Ideas 256
Tay–Sachs Disease 225 Evolution by Natural Selection 257
Albinism 225 16.3 Ancient Evidence 258
14.3 Examples of X-Linked Inheritance Patterns 226 Evidence in Rock Formations 258
The X-Linked Recessive Pattern 226 The Fossil Record 259
Red–Green Color Blindness 226
Duchenne Muscular Dystrophy 227 ● Engage: Dr. Paul Sereno,
Hemophilia 227 National Geographic Explorer 259
14.4 Changes in Chromosome Structure 228 16.4 Filling in Pieces of the Puzzle 260
Types of Change 228 Missing Links 261
Insertions 228
Duplications 228
Deletions 228
Inversions 228
Translocations 228
Chromosomes and Evolution 229
14.5 Changes in Chromosome Number 230
Down Syndrome 230
Sex Chromosome Changes 231
Turner Syndrome 231
XXX Syndrome 231
Klinefelter Syndrome 231
XYY Syndrome 231
14.6 Genetic Screening 232
APPLICATION: Shades of Skin 233
16.5 Drifting Continents, Changing Seas 262 17.11 Phylogeny 294
16.6 Putting Time Into Perspective 264 Cladistics 295
16.7 Evidence in Form 266 17.12 Applications of Phylogeny 296
Morphological Divergence 266 Conservation Biology 296
Morphological Convergence 267 Medical Research 296
16.8 Evidence in Function 268 APPLICATION: Directional Selection: Superbug
Patterns in Animal Development 268 Farms 297
APPLICATION: Reflections of a Distant Past 269
17 PROCESSES OF EVOLUTION
17.1 Alleles in Populations 274
Variation in Shared Traits 274
An Evolutionary View of Mutations 274
Allele Frequency 275
17.2 Genetic Equilibrium 276
Applying the Hardy–Weinberg Principle 276
Populations Evolve 277
17.3 Directional Selection 278
Selection for an Extreme Form 278
The Peppered Moth 278 18 LIFE’S ORIGIN AND EARLY EVOLUTION
Warfarin-Resistant Rats 279
18.1 Origin of Life’s Building Blocks 302
17.4 Stabilizing and Disruptive Selection 280
Selection for or Against an Intermediate Form 280 Conditions on Early Earth 302
Stabilizing Selection 280 Sources of Organic Subunits 302
Disruptive Selection 281 Lightning-Fueled Atmospheric Reactions 303
Reactions at Hydrothermal Vents 303
17.5 Natural Selection and Diversity 282 Delivery From Outer Space 303
Survival of the Sexiest 282
● Engage: Dr. Robert Ballard,
Maintaining Multiple Alleles 282 National Geographic Explorer 303
17.6 Genetic Drift and Gene Flow 284
18.2 Polymers to Protocells 304
Genetic Drift 284
Properties of Cells 304
Bottlenecks and the Founder Effect 284
Origin of Metabolism 304
Gene Flow 285
Origin of the Genome 304
17.7 Reproductive Isolation 286 Protocell Formation 305
17.8 Allopatric Speciation 288 18.3 Early Cellular Life 306
Speciation in Archipelagos 288 Traits of the First Cells 306
17.9 Other Speciation Models 290 Searching for Ancient Life 306
Sympatric Speciation 290 The Oldest Fossil Cells 306
Parapatric Speciation 291 Stromatolites and the Rise in Oxygen 306
● Engage: Dr. Julia J. Day, Early Eukaryotes 307
National Geographic Explorer 291 18.4 Origin of Eukaryotes 308
17.10 Macroevolution 292 Unique Eukaryotic Traits 308
Patterns of Macroevolution 292 A Mixed Heritage 308
Stasis 292 Evolution of Organelles 308
Exaptations 292 Nucleus and Endoplasmic Reticulum 308
Mass Extinctions 292 Mitochondria and Chloroplasts 309
Adaptive Radiation 292
18.5 Perspective on the Precambrian 310
Coevolution 292
Evolutionary Theory 293 APPLICATION: Looking for Life 311
xi
2 0 THE PROTISTS
20.1 Protist Classification and Characteristics 332
A Diverse Collection of Lineages 332
A Wide Array of Traits 332
Level of Organization 332
Metabolism 332
Habitats 333
Life Cycles 333
20.2 Protist Cell Anatomy 334
Standard Eukaryotic Features 334
Traits That Vary 334
Outer Coverings 334
Motility-Related Structures 334
Contractile Vacuoles 334
Chloroplasts 334
20.3 Excavates—Flagellated Cells 335
UNIT 4 E VO L U T I O N A N D B I O D I V E R S I T Y
Anaerobic Flagellates 335
1 9 VIRUSES, BACTERIA, AND ARCHAEA Trypanosomes and Euglenoids 335
20.4 The SAR Supergroup 336
19.1 The Viruses 316
Stramenopiles 336
Virus Structure 316
Alveolates 336
Viral Replication 316
Rhizarians 337
Bacteriophage Replication 316
HIV Replication 317 20.5 The Deadliest Protists 338
19.2 Viruses and Human Health 318 Life Cycle of Plasmodium 338
Common Viral Diseases 318 Health Effects of Malaria 339
Emerging Viral Diseases 318 Manipulating Its Hosts 339
New Flus—Viral Reassortment 319 ● Engage: Ken Banks, National Geographic Explorer 339
● Engage: Dr. Nathan Wolfe, 20.6 Archaeplastid Protists—Plant Relatives 340
National Geographic Explorer 319 Red Algae 340
19.3 Bacterial Structure and Function 320 Green Algae 341
Cell Size and Structure 320 20.7 Amoebozoans 342
Reproduction and Gene Exchange 320 Amoebas 342
Metabolic Diversity 321 Slime Molds 342
19.4 Bacterial Diversity and Ecology 322 20.8 Opisthokont Protists 343
Cyanobacteria 322
APPLICATION: Toxic Algal Blooms 343
Gram-Positive Bacteria 322
Proteobacteria 323
Spirochetes 323 2 1 PLANT EVOLUTION
19.5 Bacterial Effects on Human Health 324
21.1 Adapting to Life on Land 348
Normal Microbiota 324
Structural Adaptations 348
Toxins and Disease 324
Life-Cycle Changes 348
19.6 The Archaea 325
Pollen and Seeds 349
Comparisons With Bacteria 325
21.2 Nonvascular plants—Bryophytes 350
Archaeal Diversity 325
Mosses 350
19.7 Phylogeny of Prokaryotes 326
Liverworts and Hornworts 351
Identifying Species 326
21.3 Seedless Vascular Plants 352
Relationships Among Domains 326
Ferns and Close Relatives 352
APPLICATION: Sharing Viruses 327 Lycophytes 353
xii
21.4 History of the Vascular Plants 354 ● Engage: Dr. DeeAnn Reeder,
Tiny Branchers to Coal Forests 354 National Geographic Explorer 371
Rise of the Seed Plants 354 22.4 Human Uses of Fungi 372
● Engage: Jeff Benca, National Geographic Explorer 355
APPLICATION: Spread of Fungal Pathogens 373
21.5 What Are Gymnosperms? 356
Conifers 356 23 ANIMALS I: MAJOR INVERTEBRATE
Lesser-Known Lineages 357 GROUPS
21.6 Angiosperms—Flowering Plants 358
Flowers and Fruits 358 23.1 Animal Traits and Trends 378
Angiosperm Lineages 359 Shared Traits 378
Body Plan Variations 378
21.7 Keys to Angiosperm Success 360
Accelerated Life Cycle 360 23.2 Animal Origins and Diversification 380
Partnerships With Pollinators 360 Colonial Origins 380
Animal-Dispersed Fruits 360 Evidence of Early Animals 380
An Explosion of Diversity 380
APPLICATION: Saving Seeds 361
23.3 Sponges 381
23.4 Cnidarians 382
● Engage: Dr. David Gruber, National Geographic
Explorer 383
23.5 Flatworms 384
Free-Living Flatworms 384
Parasitic Flatworms 385
23.6 Annelids 386
Polychaetes 386
Leeches 386
Oligochaetes 387
Earthworm Body Plan 387
Earthworm Ecology 387
23.7 Mollusks 388
Gastropods 388
Bivalves 388
Cephalopods 389
23.8 Roundworms 390
23.9 Arthropod Characteristics 391
22 FUNGI
A Jointed Exoskeleton 391
22.1 Fungal Traits and Groups 366 Specialized Segments 391
Characteristics of Fungi 366 Metamorphosis 391
Organization and Structure 366 23.10 Arthropod Subgroups 392
Metabolism and Nutrition 366
Chelicerates 392
Spore Production 367
Millipedes and Centipedes 392
Major Groups 367
Crustaceans 393
22.2 Fungal Life Cycles 368
Insects 393
Zygote Fungi 368
23.11 Insect Diversity and Ecology 394
Sac Fungi 369
Characteristics of Insects 394
Club Fungi 369
Insect Ecology 394
22.3 Ecological Roles of Fungi 370
Health and Economic Effects 395
Nature’s Recyclers 370
23.12 Echinoderms 396
Beneficial Partners 370
Parasites and Pathogens 370 APPLICATION: Medicines From the Sea 397
xiii
UNIT 5 HOW PLANTS WORK
2 5 PLANT TISSUES
25.1 The Plant Body 424
● Engage: Dr. Mark Olson,
25.2 Plant Tissues 426
2 4 ANIMALS II: THE CHORDATES Simple Tissues 426

Parenchyma 426
24.1 Chordates—Vertebrates and Their Relatives 402 Collenchyma 426
Sclerenchyma 426
Chordate Characteristics 402
Complex Tissues 427
Invertebrate Chordates 402 Dermal Tissue 427
Overview of Chordate Evolution 403 Vascular Tissue 427
24.2 Fishes—Life in Water 404 25.3 Stems 428
Jawless Fishes 404 Internal Structure 428
Jawed Fishes 404 Variations on a Stem 429
Cartilaginous Fishes 404 Stolons 429
Bony Fishes 405 Rhizomes 429
24.3 Amphibians—The First Tetrapods 406 Bulbs 429
Corms 429
The Move to Land 406
Tubers 429
Modern Amphibians 406 Cladodes 429
Declining Diversity 407
25.4 Leaves 430
24.4 Amniotes 408 Internal Structure 430
24.5 Modern “Reptiles” 409 25.5 Roots 432
Lizards and Snakes 409 External Structure 432
Turtles 409 Internal Structure 432
Crocodilians 409
25.6 Primary Growth 434
24.6 Birds—Adapted to Flight 410 Primary Growth 434
24.7 Mammals 411 Secondary Growth 435
Furry or Hairy Milk Makers 411 25.7 Tree Rings and Old Secrets 436
Modern Subgroups 411
APPLICATION: Sequestering Carbon in Forests 437
24.8 Primates 412
Primate Characteristics 412
Modern Subgroups 413
2 6 PLANT NUTRITION AND TRANSPORT
24.9 A Human–Ape Comparison 414 26.1 Plant Nutrients and Availability in Soil 442
24.10 Early Hominins 415 Properties of Soil 442
24.11 The Genus Homo 416 How Soils Change 443
Early Homo Species 416 ● Engage: Dr. Jerry Glover,
Anatomically Modern Humans 416 National Geographic Explorer 443
Neanderthals and Denisovans 417 26.2 Root Adaptations for Nutrient Uptake 444
The Flores Hominins 417 The Function of Endodermis 444
● Engage: Drs. Louise Leakey and Meave Leakey, Mutualisms 445
National Geographic Explorers 417 26.3 Water Movement Inside Plants 446
24.12 Human Origin and Dispersal 418 Cohesion–Tension Theory 447
African Origins 418 26.4 Conserving Water 448
Across the Globe 418 26.5 Phloem Function 448
Mixing Genomes 418 Pressure Flow Theory 448
APPLICATION: Downsides to Walking Upright 419 APPLICATION: Leafy Cleanup 450
xiv
27 REPRODUCTION AND DEVELOPMENT ● Engage: Dr. Grace Gobbo,
OF FLOWERING PLANTS National Geographic Explorer 475
APPLICATION: Prescription: Chocolate 476
27.1 Flower Form and Function 456
● Engage: Dr. Dino Martins,
Floral Structure 456
Pollination 457
27.2 A New Generation Begins 458 UNIT 6 HOW ANIMALS WORK
Female Gamete Production 458
Male Gamete Production 458 28 ANIMAL TISSUES AND ORGAN SYSTEMS
Double Fertilization 458
28.1 Overview of Animal Body Plans 482
27.3 Seeds and Fruits 460 Levels of Organization 482
Seeds: Mature Ovules 460 Fluids of the Animal Body 482
Fruits: Mature Ovaries 460 Evolution of Animal Body Plans 483
27.4 Asexual Reproduction of Flowering Plants 462 28.2 Epithelial Tissues 484
Agricultural Applications 462 General Characteristics 484
27.5 Plant Hormones 463 Types of Epithelial Tissue 484
27.6 Auxin and Cytokinin 464 Epithelial Renewal and Cancer 485
27.7 Gibberellin 465 ● Engage: Dr. Brenda Larison,
27.8 Abscisic Acid and Ethylene 466
Abscisic Acid 466 28.3 Connective Tissues 486
Ethylene 467 General Characteristics 486
27.9 Early Development 468 Types of Connective Tissue 486
Breaking Dormancy 468 28.4 Muscle Tissues 488
After Germination 468 Skeletal Muscle 488
27.10 Movement 470 Cardiac Muscle 488
Environmental Triggers 470 Smooth Muscle 488
Gravity 470 28.5 Nervous Tissues 489
Light 470
28.6 Organ Systems 490
Contact 471
Organs and Organ Systems 490
27.11 Responses to Recurring Environmental Change 472
28.7 Integument 492
Daily Change 472
Components of Human Skin 492
Seasonal Change 472
Sun and the Skin 493
27.12 Responses to Stress 474
28.8 Negative Feedback in Homeostasis 494
Defending Against Disease 474
Plants Do Not Want to Be Eaten 474 APPLICATION: Growing Replacement Tissues 495
xv
29.7 Central Nervous System Tissues and Fluid 510
White Matter and Gray Matter 510
Cerebrospinal Fluid 510
29.8 The Spinal Cord 511
Structure and Function 511
Anesthesia and Injury 511
29.9 The Vertebrate Brain 512
● Engage: Dr. Diana Reiss,
29.10 The Cerebral Cortex 514
29.11 Emotion and Memory 515
The Limbic System 515
Making Memories 515
29.12 Studying Brain Function 516
Observing Electrical Activity 516
Monitoring Metabolism 516
Examining Brain Tissue 516
APPLICATION: Impacts of Concussions 517
3 0 SENSORY PERCEPTION
30.1 Detecting Sensory Stimuli 522
Diversity of Sensory Receptors 522
2 9 NEURAL CONTROL Assessing Sensory Input 522
Sensation Versus Perception 522
29.1 Organizations of Nervous Systems 500
30.2 General Senses 523
Invertebrate Nervous Systems 500
Somatic and Visceral Sensation 523
Nerve Nets 500
Bilateral Cephalized Systems 500 Pain and Pain Relief 523
The Vertebrate Nervous System 501 30.3 The Chemical Senses 524
29.2 Cells of the Nervous System 502 Sense of Smell 524

Three Types of Neurons 502 Sense of Taste 524
Neuroglia 502 30.4 Light and Vision 526
Diversity of Visual Systems 526
29.3 Membrane Potentials 503
Anatomy of the Human Eye 526
Resting Potential 503
Focusing Mechanisms 527
Action Potential 503
30.5 The Human Retina 528
29.4 A Closer Look at the Action Potential 504
Detecting Light 528
Reaching Threshold 504
Visual Processing 528
An All-or-Nothing Signal 504
30.6 Common Vision Disorders 529
Propagation Along an Axon 505
Color Blindness 529
29.5 The Synapse 506
Failure to Focus 529
Synapse Structure and Function 506
Loss of Vision 529
Signal and Receptor Variety 506
30.7 Hearing 530
Synaptic Integration 507
Properties of Sound 530
Drugs That Act at Synapses 507
Vertebrate Hearing 530
29.6 The Peripheral Nervous System 508
● Engage: Dr. Fernando Montealegre-Z,
Somatic Nervous System 508 National Geographic Explorer 531
Autonomic Nervous System 509 30.8 Balance and Equilibrium 532
Resting and Digesting 509
Fight or Flight 509 APPLICATION: Neuroprostheses 533
xvi
32.3 Bone Structure and Function 560
31 ENDOCRINE CONTROL
Bone Anatomy 560
31.1 Animal Hormones 538 Bone Formation and Turnover 560
Signals That Travel in Blood 538 Joints: Where Bones Meet 561
Types of Hormones 538 32.4 Skeletal Muscle Functions 562
Hormone Receptor Variation 539 32.5 Structure of Skeletal Muscle 563
31.2 The Human Endocrine System 540
32.6 The Sliding-Filament Model 564
31.3 The Hypothalamus and Pituitary Gland 542
32.7 Control of Muscle Contraction 565
Posterior Pituitary Function 542 The Motor Signal Pathway 565
Anterior Pituitary Function 543 Motor Units 565
Hormonal Growth Disorders 543
32.8 Fueling Muscle Contraction 566
31.4 Thyroid and Parathyroid Glands 544
Energy-Releasing Pathways 566
Metabolic and Developmental Types of Muscle Fibers 566
Effects of Thyroid Hormone 544
Hormonal Regulation of Blood Calcium Level 545 APPLICATION: Exercise and Inactivity 567
31.5 The Pineal Gland 545
31.6 The Adrenal Glands 546 3 3 CIRCULATION
Hormones, Stress, and Health 546
33.1 Circulatory Systems 572
31.7 The Gonads 547 Open and Closed Systems 572
31.8 The Pancreas 548 Vertebrate Circulatory Systems 572
Regulation of Blood Sugar 548 33.2 Human Circulatory System 574
Diabetes 549 The Pulmonary Circuit 574
31.9 Invertebrate Hormones 549 The Systemic Circuit 575
APPLICATION: Endocrine Disruptors 550 33.3 The Human Heart 576
The Cardiac Cycle 576
● Engage: Dr. Tyrone Hayes,
National Geographic Explorer 551 Setting the Pace 577
33.4 Components of Blood 578
Plasma 578
32 STRUCTURAL SUPPORT
Cellular Components 578
AND MOVEMENT Red Blood Cells 578
White Blood Cells 579
32.1 Animal Locomotion 556
Platelets 579
● Engage: Dr. Kakani Katija Young,
33.5 From the Heart to the Tissues 580
Rapid Transport in Arteries 580
32.2 Types of Skeletons 558 Adjusting Flow at Arterioles 580
Invertebrate Skeletons 558
33.6 Blood Pressure 581
The Vertebrate Endoskeleton 558
33.7 Capillary Exchange 582
33.8 Returning Blood to the Heart 583
Venules and Veins 583
Impaired Venous Return 583
33.9 Cardiovascular Disorders 584
Arrhythmias 584
Atherosclerosis 584
Heart Disease and Stroke 585
33.10 The Lymphatic System 586
Lymph Vascular System 586
Lymphoid Tissues 586
APPLICATION: When the Heart Stops 587
xvii
3 4 IMMUNITY 3 5 RESPIRATION
34.1 Integrated Responses to Threats 592 35.1 Respiratory Gas Exchange 616
Three Lines of Defense 592 Sites of Gas Exchange 616
The Defenders 593 Factors Affecting Gas Exchange 616
34.2 The First Line of Defense—Surface Barriers 594 35.2 Invertebrate Respiratory Organs 617
Biological Barriers 594 35.3 Vertebrate Respiratory Systems 618
Physiological and Anatomical Barriers 595 Fish Gills 618
34.3 Mechanisms of Innate Immunity 596 Vertebrate Lungs 618
Complement Activation 596 35.4 Human Respiratory System 620
Phagocytosis 596 The Airways 620
Inflammation 597 The Lungs 621
Fever 597 Muscles of Respiration 621
34.4 Antigen Receptors 598 35.5 How We Breathe 622
Antigen Receptor Diversity 599 The Respiratory Cycle 622
34.5 Overview of Adaptive Immunity 600 Control of Breathing 623
Two Arms of Adaptive Immunity 600 Choking—A Blocked Airway 623
Antigen Processing 601 35.6 Gas Exchange and Transport 624
34.6 Adaptive Immunity I: The Respiratory Membrane 624
An Antibody-Mediated Response 602 Oxygen Transport 624
Antibodies in ABO Blood Typing 603 Carbon Dioxide Transport 624
34.7 Adaptive Immunity II: ● Engage: Dr. Cynthia Beall,
The Cell-Mediated Response 604 National Geographic Explorer 625
Cytotoxic T Cells: Activation and Action 604
35.7 Respiratory Diseases and Disorders 626
The Role of NK Cells 605
Interrupted Breathing 626
● Engage: Dr. Mark Merchant, Lung Diseases and Disorders 626
National Geographic Explorer 605 Bronchitis, Asthma, and Emphysema 626
34.8 Immunity Gone Wrong 606
APPLICATION: Effects of Smoking 627
Overly Vigorous Responses 606
Allergies 606
Acute Illnesses 606 36 DIGESTION AND HUMAN NUTRITION
Autoimmunity 607
Immune Evasion 607 36.1 Evolution of Digestive Systems 632
34.9 AIDS 608 Sites of Digestion 632
Intracellular Digestion 632
HIV Revisited 608
Extracellular Digestion 632
Transmission 608
Testing 608 Sac or Tube? 632
Treatments 609 Specialized Regions 633
Testing 609 36.2 Human Digestive Tract 634
Prevention 609
36.3 Taking in Food 635
APPLICATION: Community Immunity 610 Mammalian Mouths 635
Bird Beaks 635
36.4 The Stomach 636
Structure and Function 636
Stomach Disorders 636
36.5 The Small Intestine 637
36.6 From Digestion to Absorption 638
Carbohydrate Digestion 638
Protein Digestion 638
Fat Digestion 638
Water Uptake 639
xviii
3 8 REPRODUCTION AND DEVELOPMENT
36.7 The Large Intestine 640
38.1 Mechanisms of Animal Reproduction 668
36.8 Metabolism of Absorbed Organic Compounds 641
Asexual or Sexual? 668
36.9 Vitamins and Minerals 642 Variation Among Sexual Reproducers 669
● Engage: Dr. Christopher Golden, 38.2 Stages of Animal Development 670
Gametes Form and Unite 670
36.10 Components of a Healthy Diet 644 Cleavage 670
Fruits, Vegetables, Whole Grains 644 Gastrulation 670
Heart-Healthy Oils 644 Tissues and Organs Form 671
Lean Meat and Low-Fat Dairy 645 38.3 Tissue and Organ Formation 672
Minimal Added Salt and Sugar 645 Cell Differentiation 672
36.11 Maintaining a Healthy Weight 646 Embryonic Induction 672
Apoptosis 673
APPLICATION: The Obesity Epidemic 647
Cell Migrations 673
38.4 Evolutionary Developmental Biology 674
37 MAINTAINING THE INTERNAL Constraints on Body Plans 674
ENVIRONMENT
Developmental Mutations 674
37.1 Water and Solute Balance 652 38.5 Overview of Human Development 675
Gains and Losses of Water and Solutes 652 38.6 Reproductive System of Human Females 676
Excretory Organs 653 Female Reproductive Anatomy 676
Nephridia 653 Egg Production and Release 677
Malpighian Tubules 653
38.7 Female Reproductive Cycles 678
Kidneys 653
Ovarian and Menstrual Cycles 678
37.2 Human Urinary System 654
From Puberty to Menopause 679
Components of the System 654
Estrous Cycles 679
Kidney Anatomy 654
38.8 Human Male Sexual Organs 680
The Nephrons 654
Tubular Components 655 Functions of the Testes 680
Blood Vessels 655 Ducts and Accessory Glands 680
37.3 Urine Formation and Concentration 656 ● Engage: Dr. Stewart C. Nicol,
Three Processes Form Urine 656 National Geographic Explorer 681
Glomerular Filtration 656 38.9 Bringing Gametes Together 682
Tubular Reabsorption 656 Sexual Intercourse 682
Tubular Secretion 656
The Sperm’s Journey 682
Concentrating the Urine 656
Fertilization 682
37.4 Kidney Disorders 658
38.10 Birth Control 684
37.5 Urine Tests 659
38.11 Sexually Transmitted Diseases 685
● Engage: Dr. Cheryl Knott,
38.12 Early Human Development 686
38.13 Emergence of Human Features 688
37.6 Maintaining Body Temperature 660
38.14 Prenatal Health and Nutrition 690
Heat Gains and Losses 660
Function of the Placenta 690
Modes of Thermoregulation 660
Effects of Maternal Health and Behavior 690
Vertebrate Responses to Cold 661 Maternal Nutrition 691
Vertebrate Responses to Heat 661 Teratogens 691
37.7 Human Thermoregulation 662 Investigating Birth Defects 691
Keeping Cool 662 38.15 Birth and Lactation 692
Staying Warm 662 Childbirth 692
Lactation 692
APPLICATION: Genetic Adaptation to
Differing Climates 663 APPLICATION: Reproductive Technology 693
xix
UNIT 7 PRINCIPLES OF ECOLOGY 40.5 Predation Effects on Life History 722

An Experimental Study 722
3 9 ANIMAL BEHAVIOR Collapse of a Fishery 723
39.1 Factors Affecting Behavior 698 40.6 Human Population Growth 724
Foraging in Fruit Flies 698 Expansions and Innovations 724

Pair-Bonding in Voles 698 Fertility and Future Growth 725
Social Dominance in Cichlids 699 40.7 Economic Effects and Resource Consumption 726
39.2 Instinct and Learning 700 A Demographic Transition 726

Instinctive Behavior 700 Development and Consumption 727
Learned Behavior 700 APPLICATION: Managing Canada Geese 727
Time-Sensitive Learning 700
Conditioned Responses 700
Habituation 701 4 1 COMMUNITY ECOLOGY
Spatial Learning 701
Social Learning 701 41.1 Factors That Shape Communities 732
Imitative Learning 701 41.2 Mutualism 733
39.3 Animal Communication 702 41.3 Interspecific Competition 734
Types of Communication Signals 702 Types of Competition 734
Eavesdroppers and Fake Signals 703 Competitive Exclusion 734
● Engage: Dr. Isabelle Charrier, Resource Partitioning 735
National Geographic Explorer 703 ● Engage: Dr. Nayuta Yamashita,
39.4 Reproductive and Parenting Behavior 704 National Geographic Explorer 735
Mating Systems 704 41.4 Predation and Herbivory 736
Parental Care 705 Predator and Prey Abundance 736
39.5 Group Living and Social Behavior 706 Predator–Prey Arms Races 736
Benefits of Grouping 706 Plant–Herbivore Arms Race 737
Costs of Group Living 707 41.5 Parasites and Parasitoids 738
Evolution of Cooperation 707 Parasitism 738
39.6 Eusocial Animals 708 Parasitoids 739
Biological Pest Control 739
APPLICATION: Can You Hear Me Now? 709
Brood Parasitism 739
41.6 How Communities Change Over Time 740
4 0 POPULATION ECOLOGY
Ecological Succession 740
40.1 Characteristics of a Population 714 Effects of Disturbance 741
Size, Density, and Distribution 714 41.7 Single-Species Effects 742
Sampling a Population 715 Keystone Species 742
● Engage: Dr. Karen DeMatteo, Exotic and Endemic Species 742
National Geographic Explorer 715 41.8 Island Biogeography 744
40.2 Population Size and Exponential Growth 716 APPLICATION: Fighting Foreign Fire Ants 745
Immigration and Emigration 716
Zero to Exponential Growth 716
4 2 ECOSYSTEMS
Biotic Potential 717
40.3 Limits on Population Growth 718 42.1 The Nature of Ecosystems 750
Density-Dependent Factors 718 Producers and Consumers 750
Logistic Growth 718 Energy Flows, Nutrients Cycle 750
Density-Independent Factors 719 Trophic Structure 751
40.4 Life History Patterns 720 42.2 Depicting Trophic Structure 752
Timing of Births and Deaths 720 Food Webs 752
r-Selection and K-Selection 721 Ecological Pyramids 753
xx
42.3 Biogeochemical Cycles 754
44 HUMAN EFFECTS ON THE BIOSPHERE
42.4 The Water Cycle 754
● Engage: Jonathan Waterman, 44.1 Threatened and Endangered Species 786
National Geographic Explorer 755 Causes of Species Decline 786
Overharvesting 786
42.5 The Carbon Cycle 756
Habitat Degradation and Fragmentation 786
Carbon Reservoirs and Flow 756
The Unknown Losses 787
The Greenhouse Effect 757
● Engage: Dr. Paula Kahumbu,
42.6 The Nitrogen Cycle 758 National Geographic Explorer 787
Nitrogen Reservoirs and Flow 758
44.2 Desertification and Deforestation 788
Alterations to the Cycle 759
Desertification 788
42.7 The Phosphorus Cycle 760
Deforestation 788
APPLICATION: Toxic Transfers 761 44.3 Effects of Pollution 790
Talking Trash 790
43 THE BIOSPHERE Atmospheric Deposition 791
Acid Deposition 791
43.1 Global Air Circulation Patterns 766 Ammonium Deposition 791
Seasonal Effects 766 Mercury Deposition 791
Air Circulation and Rainfall 766 44.4 Ozone Depletion 792
Surface Wind Patterns 767 Depletion of the Ozone Layer 792
43.2 Oceans and Climate 768 Near-Ground Ozone Pollution 792
Ocean Currents 768 44.5 Global Climate Change 793
Regional Effects 768 44.6 Conservation Biology 794
43.3 Major Biomes 770 The Value of Biodiversity 794
Differences Among Biomes 770 Setting Priorities 794
Similarities Within a Biome 770 Preservation and Restoration 795
43.4 Tropical Rain Forests 771 44.7 Reducing Human Impacts 796
43.5 Cooler Climate Forests 772
APPLICATION: Citizen Scientists 797
Temperate Deciduous Forests 772
Coniferous Forests 772
43.6 Fire-Adapted Biomes 773
43.7 Deserts 774
Plant Adaptations to Drought 774
Desert Crust 774
43.8 Arctic Tundra 775
● Engage: Dr. Katey Walter Anthony,
43.9 Freshwater Ecosystems 776
Lakes 776
Nutrient Content and Succession 776
Seasonal Changes 776 APPENDICES
Streams and Rivers 777
The Role of Dissolved Oxygen 777 I Periodic Table of the Elements 800
43.10 Coastal and Coral Ecosystems 778 II The Amino Acids 801
Coastal Wetlands 778 III A Closer Look at Some Major Metabolic Pathways 802
Rocky and Sandy Seashores 778 IV A Plain English Map of the Human Chromosomes 805
Coral Reefs 779 V Restless Earth—Life’s Changing Geologic Stage 806
43.11 The Open Ocean 780 VI Units of Measure 808
APPLICATION: Effects of El Niño 781 VII Answers to Self-Assessments/Genetics Problems 808
xxi
Preface
A revolution in the way information is shared has fundamen- Concept Spreads
tally changed the nature of biological inquiry. Interdisciplin- The content of every chapter is organized as a series of con-
ary collaborations facilitated by instant, global access to data cepts, each explored in a section that is two pages or less.
and ideas have fostered entirely new areas of research, both Learning Objectives associated with each concept are phrased
theoretical and practical. New discoveries and new technolo- as activities that the student should be able to carry out after
gies emerging from these collaborations are altering the way reading the text.
biologists think about their work—and the field in general.
Realizing that a traditional life science education would Engage
not adequately prepare students for the changing field, the Our Engage feature associates chapter content with relevant
American Association for the Advancement of Science and research, while highlighting the diversity of the modern scien-
the National Science Foundation initiated a series of national tific community. Each illustrates one of six core competencies:
conversations among leading life scientists, policy makers, the ability to apply the process of science (Process of Science),
educators, and students. The result was a document, Vision use quantitative reasoning (Quantitative Reasoning), use
and Change in Undergraduate Biology Education, that calls for modeling and simulation (Complex Systems), tap into the
a fundamental change in the way life sciences are taught to all interdisciplinary nature of science (Interdisciplinary Science),
undergraduate students. A broad consensus recommends that communicate and collaborate with other disciplines (Collab-
science education become much more active, because personal orative Science), or understand the relationship between
experience with the process and limits of science better pre- science and society (Science & Society). Individuals whose
pares students to evaluate scientific content and differentiate work is spotlighted include well-established and newly
it from other information. A more concept-oriented approach minted scientists, as well as a few nonscientists; most are
that uses fundamental biological principles as a context for National Geographic Explorers or Grantees.
information (rather than the reverse) better prepares students
to understand rapid changes in the field. Our future citizens On-Page Glossary
and leaders will need this understanding to confront urgent An On-Page Glossary includes boldface key terms introduced
societal problems such as climate change, threats to biodiver- in each section. This running glossary, which can be used as
sity, and the global spread of disease. a convenient study aid, offers non-phonemic pronunciations
This book has been revised in alignment with “Vision and definitions in alternate wording. All glossary terms also
and Change” recommendations. As always, recent discover- appear in boldface in the Chapter Summary.
ies are integrated in an accessible and appealing introduction
to the study of life. In addition, this edition includes tools to Emphasis on Relevance
explore basic biological concepts from a variety of perspectives We continue to focus on real world applications, including
(molecular, cellular, organismal, ecological, and so on). social issues arising from new research and developments—
Our quest to educate and engage is shared by the National particularly the many ways in which human activities are
Geographic Society, with whom we have partnered for this continuing to alter the environment and threaten both human
book. You will see the fruits of the partnership in spectacular health and Earth’s biodiversity. Each chapter ends with an
photographs, informative illustrations, and text features that Application section that explains a current topic in light of the
highlight the wide variety of work supported by the society. chapter content, and also illustrates one of the core competen-
cies listed above.
Self-Assessment Tools
FEATURES OF THIS EDITION
Many figure captions include a Figure It Out question
Setting the Stage designed to engage students in an active learning process; an
Each chapter opens with a dramatic two-page photo spread. upside-down answer allows a quick check of understand-
A brief Links to Earlier Concepts paragraph reminds students ing. At the end of each chapter, Self-Assessment and Critical
of relevant information in previous chapters. A summary of Thinking questions provide additional self-assessment mate-
chapter content is organized and presented in terms of three rial. Another active-learning feature, the chapter-end Data
Core Concepts, each involving one of the following: evolution; Analysis class activity, sharpens analytical skills by asking stu-
information flow, exchange, and storage; structure and func- dents to interpret data presented in graphic or tabular form.
tion; pathways and transformation of energy and matter; or The data is related to the chapter material, and is taken from
systems. a published scientific study in most cases.
xxii
Chapter-Specific Changes consequences of HbS (E6V); and new art detailing the intron
This new edition contains 208 new photographs and 330 mutation that causes hairlessness in sphynx cats.
new or updated illustrations. In addition, the text of every
chapter has been updated and revised for clarity. A complete 10 Control of Gene Expression Now includes RNA inter-
section-by-section guide to new content and figures is avail- ference and microRNAs, and additional evidence for herita-
able upon request, but the highlights are summarized here. bility of epigenetic modifications. Application section has ex-
panded information about BRCA genes as tumor suppressors.
1 The Science of Biology Expanded material on the pro-
cess of science includes the concept that research is typically 11 How Cells Reproduce New ultra-high resolution confo-
nonlinear and unpredictable; text and table contrasting science cal live-cell images of mitosis by Dr. George von Dassow. Text
with pseudoscience; and text detailing the way theories can be and new art showing cytokinesis include ultrastructural de-
modified upon discovery of new data. tails/processes per current research and paradigms. Expanded
material on telomeres now includes telomere-associated trig-
2 Life’s Chemical Basis New table compares elemental gering and consequences of cell senescence.
composition of human body with Earth’s crust, seawater, and
the universe; effects of acid rain are now exemplified with dis- 12 Meiosis and Sexual Reproduction New 3D structured
solving shells of marine animals. illumination micrographs of meiosis in corn show synaptone-
mal complex detail in ultra-high resolution. Newly discovered
3 Molecules of Life New chapter opener illustrates forma- mechanism of gene acquisition by individual rotifers added to
tion of glycoaldehyde in interstellar gas; revised text and new Application essay.
art emphasizing protein structure–function relationship in-
cludes expanded discussion of amyloid fibrils and plaques. 13 Patterns in Inherited Traits Marfan syndrome dis-
cussion updated to reflect change in life expectancy due to
4 Cell Structure Added nonmotile cilia and their newly dis-
increased awareness, accompanied by new photo of Baylor’s
covered roles in cell signaling. New art shows ultrastructural
Isaiah Austin. New material details environmentally induced
details of cell junctions per recent discoveries. Expanded sec-
hemoglobin production in Daphnia; new photo shows
tion on the nature of life now includes theory of living systems.
green-to-red phenotype that accompanies the switch.
5 Ground Rules of Metabolism Coenzymes are now illus-
14 Human Inheritance Molecular pathogenesis of CF,
trated with ascorbic acid/scurvy example.
Huntington’s, and DMD updated to reflect current research.
6 Where It Starts—Photosynthesis Chapter has been
15 Biotechnology Heavily revised material includes mech-
reorganized for a better introductory sequence. Expanded
anism, application, and social implications of CRISPR-Cas9
discussion of the cyclic pathway emphasizes the interplay
between both versions of light reactions; expanded discussion gene editing system.
of photorespiration incorporates new research on its adaptive
16 Evidence of Evolution Cetacean evolutionary sequence
value. New Application discusses biofuels in context of anthro-
updated to reflect current accepted narrative.
pogenic CO2 and global warming.
17 Processes of Evolution New photo of velvet walk-
7 Releasing Chemical Energy New art and tables empha-
ing worm shooting streams of glue from its head illustrates
size the movement of energy and matter in aerobic respiration.
parapatric speciation. Expanded Application includes current
Expanded Application includes mechanisms of mitochondrial
statistics and research on generation and spread of antibiotic-
malfunction, narrative of affected child, and three-person IVF.
resistant bacteria associated with factory farms, and example
New Data Analysis concerns reprogramming of mitochondria
in brown fat by dietary fat overload. of cross-resistance to veterinary and human antibiotics.
18 Life’s Origin and Early Evolution Improved descrip-

8 DNA Structure and Function New micrographs and re-
vised art reinforce DNA structure and clarify mutations. tions of the Hadean, Archean, and Phanerozoic eons. Added
information about Miller–Urey’s experiments that simulated
9 From DNA to Protein Expanded section on mutations conditions around volcanoes. Updated discussion of protocells.
includes material on a beneficial hemoglobin mutation (E6K, Deleted coverage of Jeon’s study of endosymbionts in amoebas.
HbC) that offers resistance to malaria without the health New subsection summarizes events of the Precambrian.
xxiii
Preface (continued)
Chapter-Specific Changes (continued) involving resveratrol. New Application involves benefits of

plant secondary metabolites using as an example the discovery
19 Viruses, Bacteria, and Archaea Improved art compar- of epicatechin in cocoa via the low incidence of hypertension
ing viral structures. Updated information about Ebola, AIDS, among the Kuna tribe, with emphasis on similar signaling
West Nile virus. Added information about the Zika virus. pathways in plants and humans.
New figure shows viral reassortment. Added figure illustrating
the three common bacterial shapes. New table lists common 28 Animal Tissues and Organ Systems New summary
bacterial diseases. New subsection discusses metagenomics, table describing tissue types. Improved graphic illustrates rela-
relationships among prokaryotes, and the relationship between tive volumes of the fluid components of a human body. Added
prokaryotes and eukaryotes. information about brown fat versus white fat and white matter
and gray matter. New graphic illustrates properties of stem
20 The Protists New table lists the various protist groups cells. Updated information about research on and clinical use
and their traits. Chapter now organized around the major eu- of induced pluripotent stem cells (IPSCs).
karyotic supergroups. New overview of protist cell structure.
29 Neural Control New overview of intracellular signal-
New information about African trypanosomiasis and expand-
ing. Revised/reorganized coverage of the peripheral nervous
ed coverage of malaria.
system. New photo of a paralyzed veteran using a robotic exo-
21 Plant Evolution Revised life cycle graphics throughout skeleton. New subsection covers tissues and fluid of the CNS.
the chapter; improved figure illustrating generalized process of Updated Application with the latest findings about brain dam-
seed production. age among professional football players.
22 Fungi Added text and table comparing traits of fungi, 30 Sensory Perception Simplified figure for human olfac-
plants, and animals. New photos of athlete’s foot and ring- tion. Added frontal view to illustration of visual accommo-
worm. Updated information about white nose syndrome. New dation. New graphic depicts the anatomy of the retina and
information about medical use of psilocybin. includes light-channeling neuroglia. Application now covers
both cochlear and retinal implants.
23 Animals I: Major Invertebrate Groups More on the
31 Endocrine Control Added information about sites
similarities between choanoflagellates and animals. Coverage
of human steroid hormone production (gonads/adrenals).
of placozoans deleted. New figure illustrates bivalve (clam)
Added information about hormone type (amino acid or ste-
anatomy, and sea star anatomy figure has been updated.
roid) to Table 31.1. Updated figure depicting feedback control
Expanded coverage of roundworms as a model organism. New
of thyroid hormone.
information about overharvesting of krill and about copepods
as reservoirs for cholera-causing bacteria. 32 Structural Support and Movement Improved figures
depicting locomotion of fly and earthworm. Revised figure
24 Animals II: The Chordates Updated lancelet art. New
showing the structure of skeletal muscle.
subsection and figure devoted to declining amphibian diversity.
New figure of avian skeleton. Updated discussion of fos- 33 Circulation Updated photo depicting measurement of
sil hominins and evidence of interbreeding among humans, blood pressure. Expanded coverage of venule function. New
Neandertals, and Denisovans. art depicts atherosclerosis. Added discussion of heart attack
symptoms and of causes and symptoms of stroke. New Data
25 Plant Tissues Many new photos. Expanded Application Analysis exercise on hypertension and risk of stroke.
material includes carbon release by decomposition, relative sta-
bility of dead plant matter, and CO2 and climate change. 34 Immunity Text updates reflect current research on role
of keratinocytes as immune cells. New Application essay that
26 Plant Nutrition and Transport Many new photos and details vaccination and benefits of herd immunity features a
updated art pieces. Mechanism of regulation of water flow narrative by a mother whose unvaccinated child has perma-
through bordered pits updated per current research. nent health consequences of contracting measles.
27 Reproduction and Development of Flowering 35 Respiration Improved photo of insect tracheal system.
Plants Revisions include addition/illustration of nastic move- Increased emphasis on evolutionary trends in discussion of
ments, and explanation of Phylloxera resistance in American vertebrate lungs. Updated information about risks for SIDS.
grapevines based on enhanced hypersensitive response Added information about risks of vaping.
xxiv
36 Digestion and Human Nutrition Expanded discussion
of sponge digestion with new graphic. New graphic shows
diet-related variation in bird beaks. Improved photo of interior
of small intestine. New graphics depict peristalsis and seg-
mentation. Updated coverage of human nutrition and factors
affecting weight.
37 Maintaining the Internal Environment Improved

description of variations in types of nitrogenous wastes. New
photo of donor/recipient in living donor kidney transplant.
New information about how diseases affect composition
of urine. Added information about human body hair as a
temperature-related adaptation.
38 Reproduction and Development Improved description

of apoptosis. Reorganized discussion of human reproduction;
female now precedes male. New information about genetics
of bird beak and human facial development, post-fertilization
epigenetic reset, mechanism of Ru-486, developmental effects Acknowledgments
of Zika virus, and banking of sperm and eggs. New Data Writing, revising, and illustrating a biology textbook is a major under-
Analysis exercise about regional variations in male infertility. taking for two full-time authors, but our efforts constitute only a small
part of what is required to produce and distribute this one. We are truly
39 Animal Behavior New information about oxytocin fortunate to be part of a huge team of very talented people who are just
and human autism and about epigenetic effects in a variety as committed to creating and disseminating this exceptional science
of contexts. Improved honeybee dance language figure. New education product.
photo/information about tent caterpillars, a pre-social spe- Biology is not dogma; the fantastic amount of research in the field
cies. Application now covers effects of shipping noise on whale makes paradigm shifts common. Only with the ongoing input of our
communication. many academic reviewers and advisors (see following page) can we
continue to tailor this book to the needs of instructors and students
40 Population Ecology Introduction now explains applica- while integrating new information and models. We continue to learn
tions of population ecology. Updated human population sta- from and be inspired by these dedicated educators.
tistics. Improved graphic of intermediate disturbance model. This book benefits from a collaborative association with the
National Geographic Society in Washington D.C. The Society has
41 Community Ecology New figure shows sundew plants
graciously allowed us to enhance our presentation with its extensive
and spiders as competitors for insect prey. New discussion resources, including beautiful maps and images, Society explorer and
of study that showed character displacement in Galápagos grantee materials, and online videos.
finches. At Cengage, Hal Humphrey managed all of the inhouse produc-
tion operations. April Cognato’s efforts, vision, and leadership brought
42 Ecosystems Updated information about current level of
this book forward with Vision and Change; Jake Warde provided
atmospheric carbon dioxide; expanded discussion of sources of
insightful suggestions, kind support, and encouragement. Kellie Petruz-
fossil fuels.
zelli coordinated nearly all of the development of study, testing, and
43 The Biosphere Improved description of how Earth’s online active learning resources. Grace Davidson orchestrated a con-
shape influences wind direction. More details about plant tinuous flow of files, photos, and illustrations while managing sched-
adaptations to coastal life. New coverage of ocean acidification ules, budgets, and whatever else happened to be on fire. (Grace, thank
and other threats to reefs. you as always for your patience and dedication.) Cheryl DuBois at
Lumina Datamatics, and Christine M. Myaskovsky and Betsy Hatha-
44 Human Effects on the Biosphere Added information way at Cengage Learning, were responsible for photo research. Valuable
about habitat fragmentation, the Great Pacific Garbage Patch, suggestions from copyeditor Anita Wagner Hueftle and proofreader
ill effects of ammonium and mercury deposition, nitrous ox- Diane Miller again kept our text clear and concise.
ide’s effect on the ozone layer, and the leakage of methane from
pipelines. —Lisa Starr, Chris Evers, and Cecie Starr 2016
Tom McHugh/Science Source.

xxv
Reviewers for This Edition
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xxvi
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ENERGY EFFICIENCY OF BIOFUEL PRODUCTION 25
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xxvii
Biology
1
The Science
of Biology
Links to Earlier Concepts
Whether or not you have studied biology, you already have
an intuitive understanding of life on Earth because you are
part of it. Every one of your experiences with the natural
world—from the warmth of the sun on your skin to the love
of your pet—contributes to that understanding.
Core Concepts
Interactions among the components of a biological system
give rise to complex properties.
We can understand life by studying it at

increasingly inclusive levels, starting with
atoms that compose matter, and extending to
the biosphere. Each level is a biological system
composed of interacting parts. Interactions
among the components of a system give rise
to complex properties not found in any of the components.
Interactions among organisms and their environment result in
the movement of matter and energy.
Evolution underlies the unity and diversity of life.
Shared core processes and features that are

widely distributed among organisms provide
evidence that all living things are linked by
lines of descent from common ancestors. All
biological systems are sustained by the
exchange of matter and energy; all store,
retrieve, transmit, and respond to information essential for life.
The field of biology relies upon experimentation and the

collection and analysis of scientific evidence.
Science addresses only testable ideas about

observable events and processes. Observation,
experimentation, quantitative analysis, and
critical thinking are key aspects of research in
biology. Carefully designed experiments that
yield objective data help researchers unravel
cause-and-effect relationships in complex biological systems.
Near a tent serving as a makeshift laboratory, herpetologist

Paul Oliver records the call of a frog on an expedition to
New Guinea’s Foja Mountains cloud forest. 3
Photograph by Tim Laman/National Geographic Creative.

1.1 Life Is More Than the Sum of Its Parts
LEARNI NG OBJ ECTI VES
tasks 5. For example, a flower is an organ of reproduction
Describe the successive levels of life’s organization.
in plants; a heart, an organ that pumps blood in animals. An
Explain the idea of emergent properties and give an example.
organ system is a set of interacting organs and tissues that
fulfill one or more body functions 6. Examples of organ
systems include the aboveground parts of a plant (the shoot
system), and the heart and blood vessels of an animal (the
EMERGENT PROPERTIES circulatory system).
Biologists study life. What, exactly, is “life”? We may never Unique types of organisms—California poppies, for
actually come up with a concise definition, because living example—are called species. A population is a group of
things are too diverse, and they consist of the same basic com- interbreeding individuals of the same species living in a given
ponents as nonliving things. When we try to define life, we area. For example, all California poppies growing in Califor-
end up with a long list of properties that differentiate living nia’s Antelope Valley Poppy Reserve form a population 8.
from nonliving things. These properties often emerge from A community consists of all populations of all species in a
the interactions of basic components. To understand how given area. The Antelope Valley Reserve community includes
that works, take a look at the groups of squares in Figure 1.1. California poppies and all other plants, as well as animals,
The property of “roundness” emerges when the component microorganisms, and so on 9. Communities may be large or
squares are organized one way, but not other ways. Another small, depending on the area defined. The next level of orga-
example is a complex behavior called swarming. When hon- nization is the ecosystem, which is a community interacting
eybees swarm, they fly en masse to establish a hive in a new with its physical and chemical environment 0. Earth’s largest
location. Each bee is autonomous, but the new hive’s location ecosystem is the biosphere, and it encompasses all regions of
is decided collectively based on an integration of signals from the planet’s crust, waters, and atmosphere in which organisms
hivemates. The swarm’s collective intelligence is the emergent live a.
property in this example.
A characteristic of a system (a colony of bees swarming,
Figure 1.1 The same
for example) that does not appear in any of the system’s com- materials, assembled
ponents (individual bees) is called an emergent property. in different ways,
The idea that structures or systems with emergent properties form objects with dif-
ferent properties.
can be assembled from the same components is a recurring
The property of
theme in our world—and also in biology. “roundness” emerges
when the squares are
assembled one way, but
LIFE’S ORGANIZATION not the others.
Biologists view life as having nested levels of organization;

interactions among the components of each level give rise to atom Smallest unit of a substance; consists of subatomic particles.
biosphere (BY-oh-sfeer) All regions of Earth where organisms live.
emergent properties (Figure 1.2). This organization begins
cell Smallest unit of life.
with atoms. Atoms are the smallest units of a substance; they
community All populations of all species in a defined area.
and the fundamental particles that compose them are the
ecosystem A community interacting with its environment.
building blocks of all matter 1. Atoms bond together to form
emergent property (ee-MERGE-ent) A characteristic of a system
molecules 2. There are no atoms unique to living things, but that does not appear in any of the system’s component parts.
there are unique molecules. A cell 3, which is the smallest molecule (MAUL-ick-yule) Two or more atoms bonded together.
unit of life, consists of many of these “molecules of life.” organ In multicelled organisms, a structure that consists of tissues
Some cells live and reproduce independently. Others engaged in a collective task.
do so as part of a multicelled organism. An organism is organism (ORG-uh-niz-um) An individual that consists of one or
an individual that consists of one or more cells 7. In most more cells.
multicelled organisms, cells are organized as tissues 4. A organ system In multicelled organisms, set of interacting organs
that carry out a particular body function.
tissue consists of specific types of cells in an arrangement
population Group of interbreeding individuals of the same species
that allows the cells to collectively perform some function— living in a defined area.
protection from injury (dermal tissue) or movement (muscle species (SPEE-sheez) Unique type of organism.
tissue), for example. An organ is a structure composed of tissue In multicelled organisms, specialized cells organized in a
tissues that collectively carry out a particular task or set of pattern that allows them to perform a collective function.
4 INTRODUCTION
– 6 organ system
A set of interacting
1 atom organs. The shoot
+ Atoms and the particles
that compose them
system of this poppy
plant includes its above-
make up all matter. ground parts: leaves,
– flowers, and stems.
2 molecule 7 multicelled organism

Atoms join other atoms Individual that consists of
in molecules. This is more than one cell. Cells
a model of a water of this California poppy
molecule. The molecules plant make up its shoot
special to life are much system and root system.
larger and more complex
than water.
3 cell 8 population
The cell is the smallest Group of single-celled
unit of life. Some, like or multicelled individuals
these plant cells, live and of a species in a given
reproduce as part of a area. This population of
multicelled organism; California poppy plants is
others do so on their own. in California’s Antelope
Valley Poppy Reserve.
4 tissue 9 community
Organized array of cells All populations of all
that interact in a collec- species in a specified
tive task. This is dermal area. These plants are
tissue on the outer sur- part of the community
face of a flower petal. in the Antelope Valley
Poppy Reserve.
5 organ 0 ecosystem
Structural unit of inter- A community interacting
acting tissues. Flowers with its physical environ-
are the reproductive ment through the transfer
organs of some plants. of energy and materials.
Sunlight and water sus-
tain the community in the
Antelope Valley.
Figure 1.2 Levels of life’s organization.

New emergent properties appear at each a biosphere
successive level. The sum of all ecosys-
?
tems: every region of
FIGURE IT OUT Earth’s waters, crust,
At which level does the emergent property of “life” and atmosphere in
appear?
Answer: The cell which organisms live.
C R E D I T S : (2) 3, 4: © Umberto Salvagnin, www.flickr.com/photos/kaibara.; 5: California Poppy, © 2009, Christine M. C H A P T E R 1 THE SCIENCE OF BIOLOGY 5
Welter; 6: Lady Bird Johnson Wildflower Center; 7: Michael Szoenyi/Science Source; 8: James Randklev/Exactostock-1672/
SuperStock; 9: © Sergei Krupnov, www.flickr.com/photos/7969319@N03; 10: © Mark Koberg Photography; 11: NASA.
1.2 Life’s Unity
Distinguish producers from consumers. 1 producer 2 consumer acquiring energy
acquiring energy and nutrients by eating a producer
Define homeostasis and explain why it is important for
and nutrients from
sustaining life.
the environment
List some functions that are guided by an organism’s DNA.
Explain how DNA is the basis of similarities and differences among
organisms.
All living things share a particular set of key features. You

already know one of these features: Because the cell is the
smallest unit of life, all organisms consist of at least one cell.
For now, we introduce three more: All living things require
ongoing inputs of energy and raw materials; all sense and
respond to change; and all use DNA as the carrier of genetic
information (Table 1.1).
ENERGY AND NUTRIENTS

Not all living things eat, but all require nutrients on an ongo-
ing basis. A nutrient is a substance that an organism acquires
from the environment in order to support growth and sur-
vival. Both nutrients and energy are essential to maintain the
organization of life, so organisms spend a lot of time acquiring
ENERGY IN
them. However, the source of energy and the type of nutrients SUNLIGHT
required differ among organisms. These differences allow
4 Producers harvest energy from
us to classify all living things into two categories: produc- the environment. Some of that energy
ers and consumers (Figure 1.3). A producer makes its own flows from producers to consumers.
food using energy and simple raw materials it obtains from
nonbiological sources 1. Plants are producers. By a process PRODUCERS
called photosynthesis, plants can use the energy of sunlight plants and other self-feeding organisms
to make sugars from carbon dioxide (a gas in air) and water.
Consumers, by contrast, cannot make their own food. A
3 Nutrients that get
consumer obtains energy and nutrients by feeding on other incorporated into the cells
organisms 2. Animals are consumers. So are decompos- of producers and consumers
ers, which feed on the wastes or remains of other organisms. are eventually released back into
the environment (by decomposi-
Leftovers from consumers’ meals end up in the environment,
tion, for example). Producers
where they serve as nutrients for producers. Said another way, then take up some of the
nutrients cycle between producers and consumers 3. released nutrients.
TABLE 1.1
CONSUMERS
Some Key Features of Life animals, most fungi, many protists, bacteria
Cellular basis All living things consist of one or more cells.

Requirement Life is sustained by ongoing inputs of 5 All of the energy that enters the world
for energy and energy and nutrients. of life eventually flows out of it, mainly as
nutrients heat released back to the environment.
Homeostasis Living things sense and respond
appropriately to change.
DNA is hereditary Genetic information in the form of DNA Figure 1.3 The one-way flow of energy and cycling of
material is passed to offspring. materials through the world of life.
6 INTRODUCTION (3) top, © Victoria Pinder, http://www.flickr.com/photos/vixstarplus; bottom producer, Zhemchuzhina/

C R E D I T:
Shutterstock.com; bottom consumer, ImageZebra/Shutterstock.com.
Unlike nutrients, energy is not cycled. It flows in one tions in their internal environment within ranges that favor
direction: from the environment 4, through organisms, and cell survival. Homeostasis is the name for this process, and it
back to the environment 5. This flow maintains the organi- is one of the defining features of life.
zation of every living cell and body, and it also influences how
individuals interact with one another and their environment.
The energy flow is one-way, because with each transfer, some
DNA IS HEREDITARY MATERIAL
energy escapes as heat, and cells cannot use heat as an energy With little variation, the same types of molecules perform the
source. Thus, energy that enters the world of life eventually same basic functions in every organism. For example, infor-
leaves it (we return to this topic in Chapter 5). mation in an organism’s DNA (deoxyribonucleic acid) guides
ongoing cellular activities that sustain the individual through
its lifetime. Such functions include development: the pro-
HOMEOSTASIS cess by which the first cell of a new individual gives rise to a
An organism cannot survive for very long unless it can multicelled adult; growth: increases in cell number, size, and
respond appropriately to specific stimuli inside and outside of volume; and reproduction: processes by which organisms
itself. For example, humans and some other animals normally produce offspring. Inheritance, the transmission of DNA to
perspire (sweat) when the body’s internal temperature rises offspring, occurs during reproduction. All organisms inherit
above a certain set point (Figure 1.4). The moisture cools the their DNA from one or more parents.
skin, which in turn helps cool the body. Individuals of every natural population are alike in most
All of the internal fluids that bathe the cells in your body aspects of body form and behavior because their DNA is
are collectively called your internal environment. Temperature very similar: Humans look and act like humans and not like
and many other conditions in that environment must be kept poppy plants because they inherited human DNA, which
within certain ranges, or your cells will die (and so will you). differs from poppy plant DNA in the information it carries.
By sensing and adjusting to change, all organisms keep condi- Individuals of almost every natural population also vary—just
a bit—from one another: One human has blue eyes, the next
has brown eyes, and so on. Such variation arises from small
differences in the details of DNA molecules, and herein lies
the source of life’s diversity. As you will see in later chapters,
differences among individuals of a species are the raw material
of evolutionary processes.
consumer (kun-SUE-murr) Organism that gets energy and

nutrients by feeding on tissues, wastes, or remains of other
organisms.
development (dih-VELL-up-ment) Process by which the first cell
of a multicelled organism gives rise to a multicelled adult.
DNA Deoxyribonucleic (dee-ox-ee-ribe-oh-nuke-LAY-ick) acid;
molecule that carries hereditary information; guides development
and other activities.
growth In multicelled species, an increase in the number, size, and
volume of cells.
homeostasis (home-ee-oh-STAY-sis) Process in which organisms
keep their internal conditions within tolerable ranges by sensing and
responding appropriately to change.
inheritance (in-HAIR-ih-tunce) Transmission of DNA to offspring.
nutrient (NEW-tree-unt) Substance that an organism acquires
from the environment to support growth and survival.
photosynthesis (foe-toe-SIN-thuh-sis) Process by which producers
use light energy to make sugars from carbon dioxide and water.
Figure 1.4 Living things sense and respond to their environ- producer Organism that makes its own food using energy and
ment. Sweating is a response to an internal body temperature that nonbiological raw materials from the environment.
exceeds the normal set point. The response cools the skin, which in reproduction (ree-pruh-DUCK-shun) Processes by which
turn helps return the internal temperature to the set point. organisms produce offspring.
C R E D I T: (4) iStockphoto.com/gvillani. C H A P T E R 1 THE SCIENCE OF BIOLOGY 7
1.3 Life’s Diversity
collection of eukaryote groups that are not plants, animals, or
List two characteristics of prokaryotes.
fungi. Collectively, they vary dramatically, from single-celled
Name the four main groups of eukaryotes.
consumers to giant, multicelled producers.
Fungi (singular, fungus) are eukaryotic consumers that
secrete substances to break down food externally, then absorb
Differences in the details of DNA molecules are the basis nutrients released by this process. Many fungi are decompos-
of a tremendous range of differences among types of organ- ers. Most fungi, including those that form mushrooms, are
isms. Various classification schemes help us organize what multicellular. Fungi that live as single cells are called yeasts.
we understand about the scope of this variation, which is Plants are multicelled eukaryotes, and the vast major-
an important aspect of Earth’s biodiversity. For example, ity of them are photosynthetic producers that live on land.
organisms can be grouped on the basis of whether they Besides feeding themselves, plants also serve as food for most
have a nucleus, which is a saclike structure containing a other land-based organisms.
cell’s DNA. Bacteria (singular, bacterium) and archaea Animals are multicelled consumers that ingest other
(singular, archaeon) are organisms whose DNA is not con- organisms or components of them. Unlike fungi, animals
tained within a nucleus. All bacteria and archaea are single- break down food inside their body. They also develop
celled, which means each organism consists of one cell through a series of stages that lead to the adult form. All ani-
(Figure 1.5A,B). Collectively, these organisms are the most mals actively move about during at least part of their lives.
diverse representatives of life. Different kinds are producers
or consumers in nearly all regions of Earth. Some inhabit
such extreme environments as frozen desert rocks, boiling
sulfurous lakes, and nuclear reactor waste. The first cells on
Earth may have faced similarly hostile conditions.
Traditionally, organisms without a nucleus have been
classified as prokaryotes, but the designation is now used
only informally. This is because bacteria and archaea are
less related to one another than we once thought, despite
their similar appearance. Archaea turned out to be more
closely related to eukaryotes, which are organisms whose
DNA is contained within a nucleus. Some eukaryotes live
as individual cells; others are multicelled (Figure 1.5C).
Eukaryotic cells are typically larger and more complex than
bacteria or archaea.
There are four main groups of eukaryotes: protists,
fungi, plants, and animals. Protist is the common term for a
A Bacteria are the most numerous organisms on Earth. Clock-
animal Multicelled consumer that breaks down food inside its wise from upper left, a bacterium with a row of iron crystals that
body, develops through a series of stages, and moves about during serves as a tiny compass; a common resident of cat and dog
part or all of its life. stomachs; photosynthetic bacteria; types found in dental plaque.
archaea (are-KEY-uh) Group of single-celled organisms that lack a
nucleus but are more closely related to eukaryotes than to bacteria.
bacteria The most diverse and well-known group of single-celled
organisms that lack a nucleus.
eukaryote (you-CARE-ee-oat) Organism whose cells
characteristically have a nucleus.
fungus Single-celled or multicelled eukaryotic consumer that
breaks down material outside itself, then absorbs nutrients released
from the breakdown.
plant A multicelled, typically photosynthetic producer. B Archaea resemble bacteria, but are more closely related to
prokaryote (pro-CARE-ee-oat) Single-celled organism without a eukaryotes. Left, an archaeon that grows in sulfur hot springs.
nucleus. Right, two types of archaea from a seafloor hydrothermal vent.
protist Common term for a eukaryote that is not a plant, animal,
or fungus. Figure 1.5 A few representatives of life’s diversity.
8 INTRODUCTION C R E D I T S : (5A) top left, Dr. Richard Frankel; top right, Science Source; bottom left, www.zahnarzt-stuttgart.com;
bottom right, © Susan Barnes; (5B) left, Eye of Science/Science Source; right, © Dr. Harald Huber, Dr. Michael Hohn,
Prof. Dr. K.O. Stetter, University of Regensburg, Germany.
ENGAGE: PROCESS OF SCIENCE
Protists are a group of extremely diverse eukaryotes that

range from microscopic free-living cells (left) to giant multicelled
seaweeds (right).
Figure 1.6 On a survey in New Guinea, Kristofer Helgen finds a critically

endangered long-beaked echidna. The only known living populations of
this animal occur in New Guinea, and all are in rapid decline.
Fungi are eukaryotic consumers that secrete substances to

break down food outside their body. Some are single-celled (left),
Kristofer Helgen
but most are multicelled (right).
National Geographic Explorer Kristofer Helgen discovers new
animals. Deep in a New Guinea rain forest (Figure 1.6). High
on an Andean mountainside. Resting in a museum’s specimen
drawer. “Conventional wisdom would have it that we know
all the mammals of the world,” he notes. “In fact, we know so
little. Unique species, profoundly different from anything ever
discovered, are out there waiting to be found.” His own efforts
prove this. Helgen himself has discovered approximately 100 new
Plants are multicelled eukaryotes, most of which are species of mammals previously unknown to science. “Since I was
photosynthetic. Nearly all have roots, stems, and leaves. three years old, I’ve been transfixed by animals,” he recalls. “Even
then, my excitement revolved around figuring out how many
different kinds there were.”
Helgen’s search plunges him into the wild on almost every
continent. Yet about three times as many new finds are made
within the walls of museums. “An expert can go into any large
natural history museum and identify kinds of animals no one knew
existed,” he explains. When only a few specimens of a species
exist, and reside in museums scattered across the globe, sheer
Animals are multicelled eukaryotes that ingest other organisms logistics often prevent researchers from pinpointing a new find.
or their parts, and they actively move about during part or all of Helgen recently discovered a specimen of a long-beaked echidna
their life cycle.
in a London museum. It had been collected from Australia in 1901,
C Eukaryotes are single-celled or multicelled organisms whose misidentified, and left forgotten in the bottom of a drawer. Long-
DNA is contained within a nucleus. Eukaryotes include protists, beaked echidnas were thought to be extinct in Australia for at least
plants, fungi, and animals.
5,000 years. Helgen’s discovery that one was alive in 1901 means a
population might still exist there, waiting to be discovered.
C R E D I T S : (5C) Protists: top left, Courtesy of Allen W.H. Bé and David A. Caron; bottom left, MI Walker/Science Source; right, C H A P T E R 1 THE SCIENCE OF BIOLOGY 9
© worldswildlifewonders/Shutterstock.com; Plants: left, © Jag_cz/Shutterstock.com; right, Martin Ruegner/Radius Images/Getty
Images; Fungi: left, London Scientific Films / Exactostock-1598/SuperStock; right, Kichigin/Shutterstock.com; Animals: left, AAAS;
middle, Thomas Eisner, Cornell University; right, iStockphoto.com/keeweeboy; (6) Tim Laman/National Geographic Creative.
1.4 Organizing Information About Species
An eighteenth-century naturalist, Carolus Linnaeus,
Explain how organisms are named in the Linnaean system.
standardized a naming system that we still use. By the Lin-
Describe the way species are classified in taxa.
naean system, each species is given a unique two-part scien-
List the taxa from species to domain. tific name. The first part of a scientific name is the genus
Describe the “biological species concept” and explain its drawbacks. (plural, genera), which is defined as a group of species that
share a unique set of features. Combined with the second part
of the name (the “specific epithet”), it designates one species.
Thus, the dog rose now has one official name, Rosa canina,
A ROSE BY ANY OTHER NAME . . . that is recognized worldwide.
Each time we discover a new species, we name and classify The genus name is always capitalized, and the specific
it, a practice called taxonomy. Taxonomy began thousands epithet is not. Both are always italicized. Consider Panthera,
of years ago, but naming species in a consistent way did not a genus of big cats. Lions belong to the species Panthera leo.
become a priority until the eighteenth century. At the time, Tigers belong to a different species in the same genus (Pan-
European explorers who were just discovering the scope of thera tigris), and so do leopards (P. pardus). Note how the
life’s diversity started having more and more trouble commu- genus name may be abbreviated after it has been spelled out.
nicating with one another because species often had multiple
names. For example, the dog rose (a plant native to Europe,
Africa, and Asia) was alternately known as briar rose, witch’s
DISTINGUISHING SPECIES
briar, herb patience, sweet briar, wild briar, dog briar, dog The individuals of a species share a unique set of traits.
berry, briar hip, eglantine gall, hep tree, hip fruit, hip rose, hip For example, giraffes normally have very long necks, brown
tree, hop fruit, and hogseed—and those are only the English spots on white coats, and so on. These are morphologi-
names! Species often had multiple scientific names too, in cal (structural) traits. Individuals of a species also share
Latin that was descriptive but often cumbersome. The sci- biochemical traits (they make and use the same molecules)
entific name of the dog rose was Rosa sylvestris inodora seu and behavioral traits (they respond the same way to certain
canina (odorless woodland dog rose), and also Rosa sylvestris stimuli, as when hungry giraffes feed on tree leaves). We can
alba cum rubore, folio glabro (pinkish white woodland rose rank a species into ever more inclusive categories based on
with smooth leaves). some subset of traits it shares with other species. Each rank,
wild carrot marijuana apple prickly rose dog rose

domain Eukarya Eukarya Eukarya Eukarya Eukarya
kingdom Plantae Plantae Plantae Plantae Plantae
phylum Magnoliophyta Magnoliophyta Magnoliophyta Magnoliophyta Magnoliophyta
class Magnoliopsida Magnoliopsida Magnoliopsida Magnoliopsida Magnoliopsida
order Apiales Rosales Rosales Rosales Rosales
family Apiaceae Cannabaceae Rosaceae Rosaceae Rosaceae
genus Daucus Cannabis Malus Rosa Rosa
species carota sativa domestica acicularis canina
10
Figure 1.7 Taxonomy of five species that are related at different
levels. Each species has been assigned to ever more inclusive groups,
or taxa: in this case, from genus to domain.
INTRODUCTION
? FIGURE IT OUT
Which of the plants shown here are in the same
family?
Answer: Apple, prickly rose, and dog rose
C R E D I T S : (7) from left, © xania.g, www.flickr.com/photos/52287712@N00; underworld/Shutterstock.com; © Sylvie

Bouchard/Shutterstock.com; Courtesy of Melissa S. Green, www.flickr.com/photos/henkimaa; © Grodana Sarkotic.
or taxon (plural, taxa), is a group of organisms that share a Archaea Eukarya Protists Plants Fungi Animals Bacteria
unique set of inherited traits. Each category above species—
genus, family, order, class, phylum (plural, phyla), kingdom,
and domain—consists of a group of the next lower taxon
(Figure 1.7). Using this system, we can sort all life into a few
categories (Figure 1.8 and Table 1.2). Later chapters return to Figure 1.8 The big picture of life. This diagram summarizes one
details of these and other classification systems. hypothesis about how all life is connected by shared ancestry. Lines
indicate proposed evolutionary connections between the domains.
It is easy to tell that humans and dog roses are different
?
species because they appear very different. Distinguishing FIGURE IT OUT
other species that are more closely related may be much more Protists, plants, fungi, and animals are at which
challenging (Figure 1.9). In addition, traits shared by mem- taxonomic level(s)?
Answer: All are kingdoms
bers of a species often vary a bit among individuals.

TABLE 1.2
How do biologists decide whether similar-looking organ-
isms belong to different species? The short answer to that All of Life in Three Domains
question is that they rely on whatever information is available. Single cells, no nucleus. Evolutionarily closer to
Archaea
Early naturalists studied anatomy and distribution—essen- eukaryotes than to bacteria.
tially the only methods available at the time—so species were Eukaryotic cells (with a nucleus). Includes single-
Eukarya
named and classified according to what they looked like and celled and multicelled species of protists, plants,
where they lived. Today’s biologists are able to compare traits fungi, and animals.
that the early naturalists did not even know about, including Bacteria Single cells, no nucleus.
biochemical ones such as DNA molecules.
Consider that the information in a molecule of DNA
changes a bit each time it passes from parents to offspring, and
it has done so since life began. Over long periods of time, these
tiny changes have added up to big differences between species
such as humans and dog roses. Thus, differences in DNA are
one way to measure relative relatedness: The fewer differences
between species, the closer the relationship. For example, we
know that the DNA of humans is more similar to chimpanzee
DNA than it is to cat DNA, so we can assume that chimpan-
zees and humans are closer relatives than cats and humans.
Every living species known has DNA in common with every
Figure 1.9 Four butterflies, two species: Which are which?
other species, so every living species is related to one extent or
another. Unraveling these relationships has become a major The top row shows two forms of the species Heliconius melpomene;
the bottom row, two forms of H. erato.
focus of biology (later chapters return to this topic).
H. melpomene and H. erato never cross-breed; DNA comparisons
The discovery of new information sometimes changes
confirmed that they are truly different species. The alternate but
the way we distinguish a particular species or how we group similar patterns of coloration evolved as a shared warning signal to
it with others. For example, Linnaeus grouped plants by the predatory birds that these butterflies taste terrible.
number and arrangement of reproductive parts, a scheme that
resulted in odd pairings such as castor-oil plants with pine breed, produce fertile offspring, and do not interbreed with
trees. Having more information today, we place these plants other groups. This “biological species concept” is useful but
in separate phyla. not universally applicable. For example, we may never know
Evolutionary biologist Ernst Mayr defined a species as whether populations of some species could interbreed because
one or more groups of individuals that potentially can inter- impassable geographical barriers keep them separated. As
another example, populations often continue to interbreed
genus (JEE-nuss) A group of species that share a unique set of even as they diverge, so the exact moment at which two popu-
traits.
lations become two species is often impossible to pinpoint. We
taxon (TAX-on) A rank of organisms that share a unique set of
traits. return to speciation and how it occurs in Chapter 17, but for
taxonomy (tax-ON-oh-me) Naming and classifying species in a now it is important to remember that a “species” is a conve-
systematic way. nient but artificial construct of the human mind.
C R E D I T S : (9) From Meyer A., Repeating Patterns of Mimicry. PLoS Biology Vol. 4, No. 10, e341 doi:10.1371/journal. C H A P T E R 1 THE SCIENCE OF BIOLOGY 11
pbio.0040341.
1.5 The Science of Nature
a bit of extra awareness. There are many ways to do it. For
Describe critical thinking and give some examples of how to do it.
example, you might ask yourself some of the following ques-
Distinguish between inductive and deductive reasoning.
tions while learning something new:
Use suitable examples to explain dependent and independent variables.
Explain how a control group is used in an experiment. ▶ What message am I being asked to accept?
List the tasks that are part of the scientific method. ▶ Is the message based on facts or opinion?
▶ Is there a different way to interpret the facts?
▶ What biases might the presenter have?
▶ How do my own biases affect what I’m learning?
THINKING ABOUT THINKING
Most of us assume that we do our own thinking, but do we, Questions like these are a way of being conscious about learn-
really? You might be surprised to find out how often we let ing. They can help you decide whether to allow new informa-
others think for us. Consider how a school’s job (which is to tion to guide your beliefs and actions.
impart as much information to students as quickly as possi-
ble) meshes perfectly with a student’s job (which is to acquire
as much knowledge as quickly as possible). In this rapid-fire
THE SCIENTIFIC METHOD
exchange of information, it can be very easy to forget about Critical thinking is a crucial part of science, the systematic
the merit of what is being exchanged. Any time you accept study of the physical universe and how it works. Biology, the
information without evaluating it, you let someone else think branch of science concerning past and present life, comprises
for you. hundreds of specializations (Table 1.3 and Figure 1.10). A
Critical thinking is the deliberate process of judging the line of inquiry in biology typically begins with a researcher’s
quality of information before accepting it. “Critical” comes curiosity about something observable in nature, such as, say,
from the Greek kriticos (discerning judgment). When you use an unusual decrease in the number of birds occupying a
critical thinking, you move beyond the content of new infor- particular area. The researcher reads about what others have
mation to consider supporting evidence, bias, and alternative discovered before making a hypothesis, a testable explanation
interpretations. How does the busy student manage this? for a natural phenomenon. An example of a hypothesis would
Critical thinking does not necessarily require extra time, just be: The number of birds is decreasing because the number
TABLE 1.3
A Few Research Specializations in Biology

Field Focus
Astrobiology Potential life elsewhere in the universe
Biogeography Distribution of life on Earth
Bioinformatics Development of tools to analyze data
Botany Plant structure and processes
Cell biology Cell structure and processes
Ecology Interactions among organisms, and among
organisms and their environment
Ethology Animal behavior
Genetics Inheritance
Marine biology Life in saltwater habitats
Medicine Human health
Paleontology Life in the ancient past
Structural biology Architecture-dependent function of large
biological molecules
Figure 1.10 Example of scientific research in biology. Left, National Geographic Explorer Tierney Thys travels the world’s oceans to study the giant sun-
fish (mola). “When it comes to fishes, the mola really pushes the boundary of fish form,” she says. “It seems a somewhat counterintuitive design for plying
the waters of the open seas—a rather goofy design—and yet the more I learn about it, the more respect and admiration I have for it.”
12 INTRODUCTION C R E D I T: (10) © Mike Johnson.
of cats is increasing. Making a hypothesis is an example of Test results—data—offer a quantifiable way of evalu-
inductive reasoning, which means arriving at a conclusion ating the hypothesis. Data that validate the prediction are
based on one’s observations. Inductive reasoning is the way evidence in support of the hypothesis. Data that show the
we come up with new ideas about groups of objects or events. prediction is invalid are evidence that the hypothesis is flawed
Note that a scientific hypothesis must be testable in the real and should be revised. A necessary part of science is report-
world; consider how, for example, there would be no way we ing one’s data and conclusions in a standard way, such as in a
could test a hypothesis that the number of birds is decreasing peer-reviewed journal article. The communication gives other
because undetectable aliens are taking them to another planet scientists an opportunity to evaluate the tested hypotheses for
(Section 1.8 returns to the topic of testability). themselves, both by checking the conclusions drawn and by
A prediction, or statement of some condition that repeating the experiments.
should occur if the hypothesis is correct, comes next. Making Forming a hypothesis based on observation, and then
predictions is called the if–then process, systematically testing and evaluating the hypothesis, are col-
in which the “if ” part is the hypothesis, lectively called the scientific method (Table 1.4).
and the “then” part is the prediction: If the
number of birds is decreasing because the TABLE 1.4
number of cats is increasing, then removing The Scientific Method
cats from the area should stop the decline. 1. Observe some aspect of nature.
Using a hypothesis to make a prediction is
2. Think of an explanation for your observation
a form of deductive reasoning, the logical (in other words, form a hypothesis).
process of using a general premise to draw 3. Evaluate the hypothesis.
a conclusion about a specific case. a. Make a prediction based on the hypothesis (If . . . then).
Next, a researcher tests the “then” part of the prediction. b. Test the prediction using experiments or surveys.
Some predictions are tested by systematic observation; others c. Analyze the results (data).
require experimentation. Experiments are tests designed to 4. Decide whether the results support your hypothesis or not
determine whether a prediction is valid. In an investigation (form a conclusion).
of our hypothetical bird–cat relationship, the researcher may 5. Report your experiment, data, and conclusion to the
scientific community.
remove all cats from the area. If working with an object or
event directly is not possible, experiments may be performed biology The scientific study of life.
on a model, or analogous system (for example, animal dis- control group Group of individuals identical to an experimental
eases are often used as models of similar human diseases). group except for the independent variable under investigation.
A typical experiment explores a cause-and-effect relation- critical thinking Evaluating information before accepting it.
ship using variables. A variable is something that varies: a data (DAY-tuh) Experimental results.
characteristic that differs among individuals, say, or an event deductive reasoning Using a general idea to make a conclusion
that differs over time. An independent variable is defined about a specific case.
or controlled by the person doing the experiment (in the dependent variable In an experiment, a variable that is
presumably affected by an independent variable being tested.
bird–cat experiment, the independent variable is the presence
experiment A test designed to evaluate a prediction.
or absence of cats). A dependent variable is presumed to be
experimental group In an experiment, a group of individuals
influenced by the independent variable (the dependent variable who have a certain characteristic or receive a certain treatment.
in the bird–cat experiment would be the number of birds). hypothesis (hi-POTH-uh-sis) Testable explanation of a natural
Biological systems are complex because they involve phenomenon.
many interdependent variables. It can be difficult to study independent variable Variable that is controlled by an
one variable separately from the rest. Thus, biology research- experimenter in order to explore its relationship to a dependent
variable.
ers often test two groups of individuals simultaneously. An
inductive reasoning Drawing a conclusion based on observation.
experimental group is a set of individuals that have a certain
model Analogous system used for testing hypotheses.
characteristic or receive a certain treatment. This group is
prediction Statement, based on a hypothesis, about a condition
tested side by side with a control group, which is identical to that should occur if the hypothesis is correct.
the experimental group except for one independent variable: science Systematic study of the physical universe.
the characteristic or the treatment being tested. Any differ- scientific method Making, testing, and evaluating hypotheses.
ences in experimental results between the two groups is likely variable (VAIR-ee-uh-bull) In an experiment, a characteristic or
to be an effect of changing the variable. event that differs among individuals or over time.
C R E D I T: (in text) panbazil/Shutterstock.com. C H A P T E R 1 THE SCIENCE OF BIOLOGY 13
1.6 Examples of Experiments in Biology
LEARNI NG OBJ ECTI VE POTATO CHIPS AND STOMACHACHES
Give an example, real or hypothetical, of an experiment in which a
dependent variable is affected by an independent variable. In 1996 the U.S. Food and Drug Administration (the FDA)
approved Olestra®, a fat replacement manufactured from
sugar and vegetable oil, as a food additive. Potato chips were
the first Olestra-containing food product to be sold in the
RESEARCH IN THE REAL WORLD United States. Controversy about the chip additive soon
Particularly in biology, scientific research rarely proceeds in raged. Many people complained of intestinal problems after
a direct, start-to-finish fashion as Table 1.4 might suggest. eating the chips, and thought that the Olestra was at fault.
Scientists often describe their work as a nonlinear process of Two years later, researchers at the Johns Hopkins University
exploration, asking questions, testing hypotheses, and chang- School of Medicine designed an experiment to test whether
ing directions. Hypotheses are often wrong and experimental Olestra causes cramps.
results are often unpredictable. Research usually raises more The researchers made the following prediction: if Olestra
question than it answers, so there never really is an end- causes cramps, then people who eat Olestra should be more
point. The unpredictability might be frustrating at times, but likely to get cramps than people who do not eat it. To evalu-
researchers typically say they enjoy their work because of the ate their prediction, they used a Chicago theater as a “labora-
surprising twists and turns it takes. tory.” They asked 1,100 people between the ages of thirteen
To give you a sense of how biology experiments work, and thirty-eight to watch a movie and eat their fill of potato
we summarize two published studies here. chips. Each person received an unmarked bag that contained
13 ounces of chips.
In this experiment, the individuals who received Olestra-
containing potato chips constituted the experimental group,
and individuals who received regular chips were the control
A Hypothesis group. The independent variable was the presence or absence
Olestra ® causes intestinal cramps. of Olestra in the chips.
A few days later, the researchers contacted everyone and
B Prediction collected reports of any post-movie gastrointestinal prob-
People who eat potato chips made with Olestra will be more
lems (the dependent variable). Of 563 people making up
likely to get intestinal cramps than those who eat potato chips
made without Olestra. the experimental group, 89 (15.8 percent) complained about
cramps. However, so did 93 of the 529 people (17.6 percent)
C Experiment Control Group Experimental Group making up the control group—who had eaten the regular
Eats regular Eats Olestra chips. People were about as likely to get cramps whether or
potato chips potato chips not they ate chips made with Olestra. These results showed
that the prediction was invalid, so the researchers concluded
D Results 93 of 529 people 89 of 563 people that eating Olestra does not cause cramps (Figure 1.11).
get cramps later get cramps later
(17.6%) (15.8%)
BUTTERFLIES AND BIRDS
E Conclusion The peacock butterfly is a winged insect named for the large,
Percentages are about equal. People who eat potato chips
colorful spots on its wings. In 2005, researchers reported the
made with Olestra are just as likely to get intestinal cramps
as those who eat potato chips made without Olestra. results of experiments investigating whether certain behaviors
These results do not support the hypothesis. help these butterflies defend themselves against insect-eating
birds. The study began with the observation that a rest-
ing peacock butterfly sits motionless with its wings folded
(Figure 1.12A). The dark underside of the wings provides
Figure 1.11 The steps in a scientific experiment to determine
if Olestra causes cramps. A report of this study was published in the appropriate camouflage. However, when a peacock butterfly
Journal of the American Medical Association in January 1998. sees a predator approaching, it exposes its brilliant spots by
?
repeatedly flicking its wings open and closed (Figure 1.12B).
FIGURE IT OUT At the same time, it moves the hindwings in a way that pro-
What was the dependent variable in this experiment? duces a hissing sound and a series of clicks. A colorful, moving,
Answer:Whether or not a person got cramps
noisy insect is usually very attractive to insect-eating birds, so
14 INTRODUCTION CREDITS: (11) photo, © Superstock.
the researchers were curious about why the peacock butterfly
moves and makes noises only in the presence of predators.
After they reviewed earlier studies, the scientists made two
A With wings folded,
hypotheses that might explain the wing-flicking behavior: a peacock butterfly
resembles a dead leaf,
Hypothesis 1: Peacock butterflies flick their wings in the so it is appropriately
presence of a predator because it exposes brilliant wing camouflaged from
predatory birds.
spots, thereby reducing predation. Peacock butterfly
wing spots resemble owl eyes, and anything that looks
like owl eyes is known to startle insect-eating birds.
Hypothesis 2: Peacock butterflies hiss and click when
they flick their wings because these sounds reduce
predation by birds. The sounds may be an additional B When a predatory
defense that startles insect-eating birds. bird approaches, a
butterfly flicks its
The researchers then used their hypotheses to make the fol- wings open and
closed, revealing
lowing predictions:
brilliant spots and
producing hissing
Prediction 1: If exposing brilliant wing spots by wing- and clicking sounds.
flicking reduces predation by insect-eating birds, then
removing the wingspots from peacock butterflies should
make them more likely to get eaten.
Prediction 2: If the hissing and clicking sounds pro- C Researchers tested
duced during wing-flicking reduce predation by insect- whether the wing
flicking and sound-
eating birds, then silencing peacock butterflies should making behaviors of
make them more likely to get eaten. peacock butterflies
affected predation
Next came the experiments. The researchers used a black by blue tits (a type
marker to cover up the wing spots of some butterflies. Other of songbird).
butterflies had the sound-making part of the wings removed
with scissors. A third group had both treatments: Their spots
were covered and wings were cut. Each butterfly was then put
into a large cage with a hungry blue tit (Figure 1.12C), and Experimental Treatment Number of Butterflies Eaten
the pair was watched for thirty minutes. Spots painted out 5 of 10 (50%)
Figure 1.12D lists the results. All of the butterflies with Wings cut 0 of 8 (0%)
unmodified wing spots survived, regardless of whether they Spots painted, wings cut 8 of 10 (80%)
made sounds. These results were consistent with the first None 0 of 9 (0%)
hypothesis: By exposing brilliant spots, peacock butterfly * Proceedings of the Royal Society of London, Series B (2005) 272:
wing-flicking behavior decreases predation by blue tits. 1203–1207.
In contrast, a large proportion of butterflies without D The researchers painted out the spots of some butterflies, cut
spots got eaten, whether or not they could make sounds. the sound-making part of the wings on others, and did both to
Experimental results were not consistent with the second a third group; then exposed each butterfly to a hungry blue tit
hypothesis that peacock butterfly sounds reduce predation by for 30 minutes. Results support only the hypothesis that peacock
butterfly spots deter predatory birds.
birds. Other questions raised by these results offer an example
of how research leads to more research: Do other predatory Figure 1.12 Testing the defensive value of two peacock but-
birds respond differently to the sounds than blue tits? If not, terfly behaviors.
?
do the sounds reduce predation by other organisms (such as
mice) that eat peacock butterflies? If sound-making is unre- FIGURE IT OUT
lated to predation, what is its function? Several additional What was the dependent variable in this series of
experiments?
Answer: Getting eaten
experiments would be necessary to answer these questions.
CREDITS: (12A) © Matt Rowlings, www.eurobutterflies.com; (12B) © Adrian Vallin; (12C) © Antje Schulte. C H A P T E R 1 THE SCIENCE OF BIOLOGY 15
Another random document with
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—Généreux comme un voleur! dit le comte.
—Voyons, voyons, ce n’est pas répondre, repartit Max en se
levant à son tour. Veux-tu de la femme ou n’en veux-tu pas?
—Je n’en veux pas.
—Alors je la prends.
—Un instant! s’écria Henri en étendant la main; il me semble que
je suis bien quelqu’un ou quelque chose ici, et que j’ai des droits
comme un autre. Qui est-ce qui a tué le mari?
—Au fait, c’est un antécédent, dit en riant le comte.
Un gémissement se fit entendre à ce mot. Je tournai les yeux du
côté où il venait: une femme était étendue sur un lit à colonnes, les
bras et les jambes liés aux quatre supports du baldaquin. Mon
attention avait été tellement absorbée sur un seul point, que je ne
l’avais pas aperçue d’abord.
—Oui, continua Max; mais qui les a attendus au Havre? qui est
accouru ici à franc étrier pour vous avertir?
—Diable! fit le comte, voilà qui devient embarrassant, et il
faudrait être le roi Salomon en personne pour décider qui a le plus
de droits, de l’espion ou de l’assassin.
—Il faut pourtant que cela se décide, dit Max. Tu m’y as fait
penser, à cette femme, et voilà que j’en suis amoureux maintenant.
—Et moi de même, dit Henri. Ainsi, puisque tu ne t’en soucies
pas, toi, donne-la à celui de nous deux que tu voudras.
—Pour que l’autre m’aille dénoncer à la suite de quelque orgie
où, comme aujourd’hui, il ne saura plus ce qu’il fait, n’est-ce pas?
Oh! que non, messieurs. Vous êtes beaux, vous êtes jeunes, vous
êtes riches, vous avez dix minutes pour lui faire la cour. Allez, mes
don Juan.
—A la cour près, ce que tu viens de dire est une idée, répondit
Henri. Qu’elle choisisse elle-même celui qui lui conviendra le mieux.
—Allons, soit, répondit Max; mais qu’elle se dépêche. Explique-
lui cela, toi qui parles toutes les langues.
—Volontiers, dit Horace. Puis, se tournant vers la malheureuse
femme:—Milady, lui dit-il dans l’anglais le plus pur, voici deux
brigands de mes amis, tous deux de bonne famille, au reste, ce dont
on peut vous donner la preuve sur parchemin, si vous le désirez, qui,
élevés dans les principes de la secte platonique, c’est-à-dire du
partage des biens, ont commencé par manger les leurs; puis,
trouvant alors que tout était mal arrangé dans la société, ont eu la
vertueuse idée de s’embusquer sur les grandes routes où elle
passe, pour corriger ses injustices, rectifier ses erreurs et équilibrer
ses inégalités. Depuis cinq ans, à la plus grande gloire de la
philosophie et de la police, ils s’occupent religieusement de cette
mission, qui leur donne de quoi figurer de la manière la plus
honorable dans les salons de Paris, et qui les conduira, comme cela
est arrivé pour moi, à quelque bon mariage qui les dispensera de
continuer de faire les Karl Moor et les Jean Sbogar. En attendant,
comme il n’y a dans ce château que ma femme, et que je ne veux
pas la leur donner, ils vous supplient bien humblement de choisir,
entre eux deux, celui qui vous conviendra le plus; faute de quoi, ils
vous prendront tous les deux. Ai-je parlé en bon anglais, madame, et
m’avez-vous compris?...
—Oh! si vous avez quelque pitié dans le cœur, s’écria la pauvre
femme, tuez-moi! tuez-moi!
—Que répond-elle? murmura Max.
—Elle répond que c’est infâme, voilà tout, dit Horace; et j’avoue
que je suis un peu de son avis.
—Alors..... dirent ensemble Max et Henri en se levant.
—Alors, faites comme vous voudrez, répondit Horace; et il se
rassit, se versa un verre de vin de Champagne et but.
—Oh! tuez-moi donc! tuez-moi donc! s’écria de nouveau la
femme en voyant les deux jeunes gens prêts à s’avancer vers elle.
En ce moment, ce qu’il était facile de prévoir arriva: Max et Henri,
échauffés par le vin, se trouvèrent face à face, et, poussés par le
même désir, se regardèrent avec colère.
—Tu ne veux donc pas me la céder? dit Max.
—Non! répondit Henri.
—Eh bien! alors, je la prendrai.
—C’est ce qu’il faudra voir.
—Henri! Henri! dit Max en grinçant des dents, je te jure sur mon
honneur que cette femme m’appartiendra!
—Et moi, je te promets sur ma vie qu’elle sera à moi; et je tiens
plus à ma vie, je crois, que tu ne tiens à ton honneur.
Alors ils firent chacun un pas en arrière, tirèrent leurs couteaux
de chasse et revinrent l’un contre l’autre.
—Mais, par grâce, par pitié, au nom du ciel, tuez-moi donc! cria
pour la troisième fois la femme couchée.
—Qu’est-ce que vous venez de dire? s’écria Horace toujours
assis, s’adressant aux deux jeunes gens d’un ton de maître.
—J’ai dit, répondit Max en portant un coup à Henri, que ce serait
moi qui aurais cette femme.
—Et moi, reprit Henri, pressant à son tour son adversaire, j’ai dit
que ce serait non pas lui, mais moi; et je maintiens ce que j’ai dit.
—Eh bien! murmura Horace, vous en avez menti tous les deux;
vous ne l’aurez ni l’un ni l’autre.
A ces mots, il prit sur la table un pistolet, le leva lentement dans
la direction du lit et fit feu: la balle passa entre les combattans et alla
frapper la femme au cœur.
A cette vue, je jetai un cri affreux et je tombai évanouie, et aussi
morte en apparence que celle qui venait d’être frappée.
XIII.
Lorsque je revins à moi j’étais dans le caveau: le comte, guidé

par le cri que j’avais poussé et par le bruit de ma chute, m’avait sans
doute trouvée dans le laboratoire, et, profitant de mon
évanouissement, qui avait duré plusieurs heures, m’avait transportée
dans cette tombe: il y avait près de moi, sur une pierre, une lampe,
un verre, une lettre: le verre contenait du poison; quant à la lettre, je
vais vous la dire:
—Hésitez-vous à me la montrer, m’écriai-je, et n’êtes-vous
confiante qu’à demi?
—Je l’ai brûlée, me répondit Pauline; mais, soyez tranquille: je
n’en ai pas oublié une parole.
«Vous avez voulu que la carrière du crime fût complète

pour moi, Pauline: vous avez tout vu, tout entendu: je n’ai
donc plus rien à vous apprendre: vous savez qui je suis, ou
plutôt ce que je suis.
»Si le secret que vous avez surpris était à moi seul, si
nulle autre vie que la mienne n’était en jeu, je la risquerais
plutôt que de faire tomber un seul cheveu de votre tête. Je
vous le jure, Pauline.
»Mais une indiscrétion involontaire, un signe d’effroi
arraché à votre souvenir, un mot échappé dans vos rêves,
peut conduire à l’échafaud non-seulement moi, mais encore
deux autres hommes. Votre mort assure trois existences: il
faut donc que vous mouriez.
»J’ai eu un instant l’idée de vous tuer pendant que vous
étiez évanouie, mais je n’en ai pas eu le courage, car vous
êtes la seule femme que j’aie aimée, Pauline: si vous aviez
suivi mon conseil, ou plutôt obéi à mes ordres, vous seriez à
cette heure près de votre mère. Vous êtes venue près de moi:
ne vous en prenez donc qu’à vous de votre destinée.
»Vous vous réveillerez dans un caveau où nul n’est
descendu depuis vingt ans, et dans lequel, d’ici à vingt ans
peut-être, nul ne descendra encore. N’ayez donc aucun
espoir de secours, car il serait inutile. Vous trouverez du
poison près de cette lettre: tout ce que je puis faire pour vous
est de vous offrir une mort prompte et douce au lieu d’une
agonie lente et douloureuse. Dans l’un ou l’autre cas, et
quelque parti que vous preniez, à compter de cette heure,
vous êtes morte.
»Personne ne vous a vue, personne ne vous connaît;
cette femme que j’ai tuée pour mettre Max et Henri d’accord
sera ensevelie à votre place, ramenée à Paris dans les
caveaux de votre famille, et votre mère pleurera sur elle,
croyant pleurer sur son enfant.
»Adieu, Pauline. Je ne vous demande ni oubli ni
miséricorde: il y a longtemps que je suis maudit, et votre
pardon ne me sauverait pas.»
—C’est atroce! m’écriai-je; ô mon Dieu, mon Dieu! que vous avez
dû souffrir!
—Oui. Aussi tout ce qui me resterait à vous raconter ne serait
que mon agonie: ainsi donc...
—N’importe, m’écriai-je en l’interrompant, n’importe, dites-la.
—Je lus cette lettre deux ou trois fois: je ne pouvais pas me
convaincre moi-même de sa réalité. Il y a des choses contre
lesquelles la raison se révolte: on les a devant soi, sous la main,
sous les yeux; on les regarde, on les touche, et l’on n’y croit pas.
J’allai en silence à la grille; elle était fermée; je fis deux ou trois fois
en silence le tour de mon caveau, frappant ses murs humides de
mon poing incrédule; puis je revins m’asseoir en silence dans un
angle de mon tombeau. J’étais bien enfermée; à la lueur de la lampe
je voyais bien la lettre et le poison; cependant je doutais encore; je
disais, comme on se le dit quelquefois en songe: Je dors, je vais
m’éveiller.
Je restai ainsi assise et immobile jusqu’au moment où ma lampe
se mit à pétiller. Alors une idée affreuse, qui ne m’était pas venue
jusque là, me vint tout-à-coup; c’est qu’elle allait s’éteindre. Je jetai
un cri de terreur et m’élançai vers elle: l’huile était presque épuisée.
J’allais faire dans l’obscurité mon apprentissage de la mort.
Oh! que n’aurais-je pas donné pour avoir de l’huile à verser dans
cette lampe. Si j’avais pu l’alimenter de mon sang, je me serais
ouvert les veines avec mes dents. Elle pétillait toujours; à chaque
pétillement, sa lumière était moins vive, et le cercle de ténèbres,
qu’elle avait éloigné lorsqu’elle brillait dans toute sa force, se
rapprochait graduellement de moi. J’étais près d’elle, à genoux, les
mains jointes; je ne pensais pas à prier Dieu, je la priais, elle...
Enfin elle commença de lutter contre l’obscurité, comme j’allais
bientôt moi-même commencer de lutter contre la mort. Peut-être
l’animais-je de mes propres sentimens; mais il me semblait qu’elle
se cramponnait à la vie, et qu’elle tremblait de laisser éteindre ce feu
qui était son âme. Bientôt l’agonie arriva pour elle avec toutes ses
phases: elle eut des lueurs brillantes, comme un moribond a des
retours de force; elle jeta des clartés plus lointaines qu’elle n’avait
jamais fait, comme au milieu de son délire l’esprit fiévreux voit
quelquefois au-delà des limites assignées à la vue humaine; puis la
langueur de l’épuisement leur succéda; la flamme vacilla pareille à
ce dernier souffle qui tremble aux lèvres d’un mourant; enfin elle
s’éteignit, emportant avec elle la clarté, qui est la moitié de la vie.
Je retombai dans l’angle de mon cachot. A compter de ce
moment, je ne doutai plus: car, chose étrange, c’était depuis que
j’avais cessé de voir la lettre et le poison que j’étais bien certaine
qu’ils étaient là.
Tant que j’avais vu clair, je n’avais point fait attention au silence:
dès que la lumière fut éteinte, il pesa sur mon cœur de tout le poids
de l’obscurité. Au reste, il y avait quelque chose de si funèbre et de
si profond, qu’eussé-je eu la chance d’être entendue, j’eusse hésité
peut-être à crier. Oh! c’était bien un de ces silences mortuaires qui
viennent s’asseoir pendant l’éternité sur la pierre des tombes.
Une chose bizarre, c’est que l’approche de la mort m’avait
presque fait oublier celui qui la causait: je pensais à ma situation,
j’étais absorbée dans ma terreur; mais je puis le dire, et Dieu le sait,
si je ne pensai pas à lui pardonner, je ne songeai pas non plus à le
maudire. Bientôt je commençai à souffrir de la faim.
Un temps que je ne pus calculer s’écoula, pendant lequel
probablement le jour s’était éteint et la nuit était venue: car, lorsque
le soleil reparut, un rayon, qui pénétrait par quelque gerçure du sol,
vint éclairer la base d’un pilier. Je jetai un cri de joie, comme si ce
rayon m’apportait un espoir.
Mes yeux se fixèrent sur ce rayon avec tant de persévérance,
que je finis par distinguer parfaitement tous les objets répandus sur
la surface qu’il éclairait: il y avait quelques pierres, un éclat de bois
et une touffe de mousse: en revenant toujours à la même place, il
avait fini par tirer de terre cette pauvre et débile végétation. Oh! que
n’aurais-je pas donné pour être à la place de cette pierre, de cet
éclat de bois et de cette mousse, afin de revoir le ciel encore une
fois à travers cette ride de la terre.
Je commençai à éprouver une soif ardente et à sentir mes idées
se confondre: de temps en temps des nuages sanglans passaient
devant mes yeux, et mes dents se serraient comme dans une crise
nerveuse; cependant j’avais toujours les yeux fixés sur la lumière.
Sans doute elle entrait par une ouverture bien étroite, car lorsque le
soleil cessa de l’éclairer en face, le rayon se ternit et devint à peine
visible. Cette disparition m’enleva ce qui me restait de courage: je
me tordis de rage et je sanglotai convulsivement.
Ma faim s’était changée en une douleur aiguë à l’estomac. La
bouche me brûlait; j’éprouvais le désir de mordre; je mis une tresse
de mes cheveux entre mes dents, et je la broyai. Bientôt je me sentis
prise d’une fièvre sourde, quoique mon pouls battît à peine. Je
commençai à penser au poison: alors je me mis à genoux et je
joignis les mains pour prier; mais j’avais oublié mes prières:
impossible de me rappeler autre chose que quelques phrases
entrecoupées et sans suite. Les idées les plus opposées se
heurtaient à la fois dans mon cerveau; un motif de musique de la
Gazza bourdonnait incessamment à mes oreilles; je sentais moi-
même que j’étais en proie à un commencement de délire. Je me
laissai tomber tout de mon long et la face contre terre.
Un engourdissement, produit par les émotions et la fatigue que
j’avais éprouvées, s’empara de moi: je m’assoupis, sans que le
sentiment de ma position cessât de veiller en moi. Alors commença
une série de rêves plus incohérens les uns que les autres. Ce
sommeil douloureux, loin de me rendre quelque repos, me brisa. Je
me réveillai avec une faim et une soif dévorantes: alors je pensai
une seconde fois au poison qui était là près de moi, et qui pouvait
me donner une fin douce et rapide. Malgré ma faiblesse, malgré mes
hallucinations, malgré cette fièvre sourde qui frémissait dans mes
artères, je sentais que la mort était encore loin, qu’il me faudrait
l’attendre bien des heures, et que de ces heures les plus cruelles
n’étaient point passées: alors je pris la résolution de revoir une fois
encore ce rayon de jour qui, la veille, était venu me visiter, comme
un consolateur qui se glisse dans le cachot du prisonnier. Je restai
les yeux fixés vers l’endroit où il devait paraître: cette attente et cette
préoccupation calmèrent un peu les souffrances atroces que
j’éprouvais.
Le rayon désiré parut enfin. Je le vis descendre pâle et blafard:
ce jour-là le soleil était voilé sans doute. Alors tout ce qu’il éclairait
sur la terre se représentait à moi: ces arbres ces prairies, cette eau
si belle; Paris, que je ne reverrais plus; ma mère, que j’avais quittée
pour toujours, ma mère, qui déjà peut-être avait reçu la nouvelle de
ma mort et qui pleurait sa fille vivante. A tous ces aspects et à tous
ces souvenirs, mon cœur se gonfla, j’éclatai en sanglots et je fondis
en pleurs: c’était la première fois depuis que j’étais dans ce caveau.
Peu à peu le paroxysme se calma, mes sanglots s’éteignirent, mes
larmes coulèrent silencieuses. Ma résolution était toujours prise de
m’empoisonner; cependant je souffrais moins.
Je restai, comme la veille, les yeux sur ce rayon tant qu’il brilla;
puis, comme la veille, je le vis pâlir et disparaître. Je le saluai de la
main... et je lui dis adieu de la voix, car j’étais décidée à ne pas le
revoir.
Alors je me repliai sur moi-même et me concentrai en quelque
sorte dans mes dernières et suprêmes pensées. Je n’avais pas fait
dans toute ma vie, comme jeune fille ou comme femme, une action
mauvaise: je mourais sans aucun sentiment de haine ni sans aucun
désir de vengeance: Dieu devait donc m’accueillir comme sa fille, la
terre ne pouvait me manquer que pour le ciel. C’était la seule idée
consolante qui me restât; je m’y attachai.
Bientôt il me sembla que cette idée se répandait non-seulement
en moi, mais autour de moi; je commençai d’éprouver cet
enthousiasme saint qui fait le courage des martyrs. Je me levai tout
debout et la tête vers le ciel, et il me sembla que mes yeux perçaient
la voûte, perçaient la terre et arrivaient jusqu’au trône de Dieu. En ce
moment, mes douleurs mêmes étaient comprimées par l’exaltation
religieuse; je marchai vers la pierre où était posé le poison, comme
si je voyais au milieu des ténèbres; je pris le verre, j’écoutai si je
n’entendais aucun bruit, je regardai si je ne voyais aucune lumière;
je relus en souvenir cette lettre qui me disait que depuis vingt ans
personne n’était descendu dans ce souterrain, et qu’avant vingt ans
peut-être personne n’y descendrait encore; je me convainquis bien
dans mon âme de l’impossibilité où j’étais d’échapper aux
souffrances qui me restaient à endurer, je pris le verre de poison, je
le portai à mes lèvres et je le bus, en mêlant ensemble, dans un
dernier murmure de regret et d’espérance, le nom de ma mère, que
j’allais quitter, et celui de Dieu que j’allais voir.
Puis je retombai dans l’angle de mon caveau; ma vision céleste
s’était éteinte, le voile de la mort s’étendait entre elle et moi. Les
douleurs de la faim et de la soif avaient reparu; à ces douleurs
allaient se joindre celles du poison. J’attendais avec anxiété cette
sueur de glace qui devait m’annoncer ma dernière agonie... Tout-à-
coup j’entendis mon nom; je rouvris les yeux et je vis de la lumière:
vous étiez là, debout à la grille de ma tombe!... vous, c’est-à-dire le
jour, la vie, la liberté... Je jetai un cri et je m’élançai vers vous. Vous
savez le reste.
Et maintenant, continua Pauline, je vous rappelle sur votre
honneur le serment que vous m’avez fait de ne rien révéler de ce
terrible drame tant que vivra encore un des trois principaux acteurs
qui y ont joué un rôle.
Je le lui renouvelai.
X I V.
La confidence que m’avait faite Pauline me rendait sa position

plus sacrée encore. Je sentis dès lors toute l’étendue que devait
acquérir ce dévoûment dont mon amour pour elle me faisait un
bonheur; mais en même temps je compris quelle indélicatesse il y
aurait de ma part à lui parler de cet amour autrement que par des
soins plus empressés et des attentions plus respectueuses. Le plan
convenu entre nous fut adopté; elle passa pour ma sœur et
m’appela son frère: cependant j’obtins d’elle, en lui faisant
comprendre la possibilité d’être reconnue par quelque personne qui
l’aurait rencontrée dans les salons de Paris, qu’elle renonçât à l’idée
de donner des leçons de langue et de musique. Quant à moi,
j’écrivis à ma mère et à ma sœur que je comptais rester pendant un
an ou deux en Angleterre. Pauline éleva encore quelques difficultés
lorsque je lui fis part de cette décision; mais elle vit qu’il y avait pour
moi un tel bonheur à l’accomplir, qu’elle n’eut plus le courage de
m’en parler, et que cette résolution prit entre nous force de chose
convenue.
Pauline avait hésité longtemps pour décider si elle révélerait ou
ne révélerait pas son secret à sa mère, et si, morte pour tout le
monde, elle serait vivante pour celle à qui elle devait la vie: moi-
même je l’avais pressée de prendre ce parti, faiblement il est vrai:
car il m’enlevait, à moi, cette position de protecteur qui me rendait si
heureux à défaut d’un autre titre; mais Pauline, après y avoir réfléchi,
avait repoussé, à mon grand étonnement, cette consolation, et
quelques instances que je lui eusse faites pour connaître le motif de
son refus, elle avait refusé de me le révéler, prétendant qu’il
m’affligerait.
Cependant nos journées passaient ainsi, pour elle dans une
mélancolie qui semblait parfois n’être point sans charmes, pour moi
dans l’espérance, sinon dans le bonheur; car je la voyais de jour en
jour se rapprocher de moi par tous les petits contacts du cœur, et,
sans s’en apercevoir elle-même, elle me donnait des preuves lentes
mais visibles du changement qui s’opérait en elle: si nous travaillions
l’un et l’autre, elle à quelque ouvrage de broderie, moi à un dessin
ou à une aquarelle, il m’arrivait souvent, en levant les yeux vers elle,
de trouver les siens fixés sur moi: si nous sortions ensemble, l’appui
qu’elle me demandait d’abord était celui d’une étrangère à un
étranger; puis, au bout de quelque temps, soit faiblesse, soit
abandon, je la sentais peser mollement à mon bras; si je sortais
seul, presque toujours, en tournant le coin de la rue Saint-James, je
l’apercevais de loin à la fenêtre, regardant du côté où elle savait que
je devais venir: tous ces signes, qui pouvaient simplement être ceux
d’une familiarité plus grande, et d’une reconnaissance plus
continuelle, m’apparaissaient à moi comme des révélations d’une
félicité à venir; je lui savais gré de chacun d’eux, et je l’en remerciais
intérieurement, car je craignais, si je le faisais tout haut, de lui faire
apercevoir à elle-même que son cœur prenait peu à peu l’habitude
d’une amitié plus que fraternelle.
J’avais fait usage de mes lettres de recommandation, et, tout
isolés que nous vivions, nous recevions parfois quelque visite: car
nous devions fuir à la fois et le tumulte du monde et l’affectation de
la solitude. Parmi nos connaissances les plus habituelles était un
jeune médecin qui avait acquis, depuis trois ou quatre ans, à
Londres, une grande réputation pour ses études profondes de
certaines maladies organiques: chaque fois qu’il venait nous voir, il
regardait Pauline avec une attention sérieuse, qui, après son départ,
me laissait toujours quelques inquiétudes; en effet, ces belles et
fraîches couleurs de la jeunesse dont j’avais vu son teint autrefois si
riche, et dont j’avais d’abord attribué l’absence à la douleur et à la
fatigue, n’avaient point reparu depuis la nuit où je l’avais trouvée
mourante dans ce caveau, ou, si quelque teinte revenait colorer
momentanément ses joues, c’était pour leur donner, tant qu’elle y
demeurait, un aspect fébrile plus inquiétant que la pâleur elle-même.
Il arrivait aussi parfois que tout-à-coup, sans cause comme sans
régularité, elle éprouvait des spasmes qui la conduisaient à des
évanouissemens, et que, pendant les jours qui suivaient ces
accidens, une mélancolie plus profonde s’emparait d’elle. Enfin ils se
renouvelèrent avec une telle fréquence et une gravité si visiblement
croissante, qu’un jour que le docteur Sercey était venu nous faire
une de ses visites habituelles, je l’arrachai aux préoccupations
qu’éveillait toujours en lui la vue de Pauline, et lui prenant le bras, je
descendis avec lui dans le jardin.
Nous fîmes plusieurs fois sans parler le tour de la petite pelouse;
puis enfin nous vînmes nous asseoir sur le banc où Pauline m’avait
raconté cette terrible histoire. Là, nous restâmes un moment pensifs;
enfin j’allais rompre le silence, lorsque le docteur me prévint:
—Vous êtes inquiet sur la santé de votre sœur? me dit-il.
—Je l’avoue, répondis-je, et vous-même m’avez laissé
apercevoir des craintes qui augmentent les miennes.
—Et vous avez raison, continua le docteur, elle est menacée
d’une maladie chronique de l’estomac. A-t-elle éprouvé quelque
accident qui ait pu altérer cet organe?
—Elle a été empoisonnée...
Le docteur réfléchit un instant.
—Oui, c’est bien cela, me dit-il, je ne m’étais point trompé: je
vous prescrirai un régime qu’elle suivra avec une grande exactitude.
Quant au côté moral du traitement, il dépend de vous; procurez à
votre sœur le plus de distraction possible. Peut-être est-elle prise de
la maladie du pays, et un voyage en France lui ferait-il du bien.
—Elle ne veut pas y retourner.
—Eh bien! une course en Écosse, en Irlande, en Italie, partout où
elle voudra; mais je crois la chose nécessaire.
Je serrai la main du docteur, et nous rentrâmes. Quant à
l’ordonnance, il devait me l’envoyer à moi-même. Je comptais, pour
ne pas inquiéter Pauline, substituer sans rien dire le régime qui lui
serait prescrit à notre manière de vivre ordinaire; mais cette
précaution fut inutile; à peine le docteur nous eût-il quittés, que
Pauline me prit la main.
—Il vous a tout avoué, n’est-ce pas? me dit-elle. Je fis semblant
de ne pas comprendre, elle sourit tristement.—Eh bien! continua-t-
elle, voilà pourquoi je n’ai pas voulu écrire à ma mère: à quoi bon lui
rendre son enfant pour qu’un an ou deux après, la mort vienne la lui
reprendre? C’est bien assez de pleurer une fois ceux qu’on aime.
—Mais, lui dis-je, vous vous abusez étrangement sur votre état:
c’est une indisposition, et voilà tout.
—Oh! c’est plus sérieux que cela, répondit Pauline avec son
même sourire doux et triste, et je sens que le poison a laissé des
traces de son passage et que je suis atteinte gravement; mais
écoutez-moi, je ne me refuse pas à espérer. Je ne demande pas
mieux que de vivre: sauvez-moi une seconde fois, Alfred. Que
voulez-vous que je fasse?
—Que vous suiviez les prescriptions du docteur: elles sont
faciles; un régime simple mais continu, de la distraction, des
voyages.
—Où voulez-vous que nous allions? je suis prête à partir.
—Choisissez vous-même le pays qui vous est le plus
sympathique.
—L’Ecosse, si vous voulez, puisque la moitié de la route est faite.
—L’Ecosse, soit.
Je fis aussitôt mes préparatifs de départ, et trois jours après nous
quittâmes Londres. Nous nous arrêtâmes un instant sur les bords de
la Twed pour la saluer de cette belle imprécation que Schiller met
dans la bouche de Marie Stuart:
«La nature jeta les Anglais et les Ecossais sur une

planche étendue au milieu de l’Océan: elle la sépara en deux
parties inégales, et voua ses habitans au combat éternel de
sa possession. Le lit étroit de la Twed sépare seul les esprits
irrités, et bien souvent le sang des deux peuples se mêla à
ses eaux: la main sur la garde de leur épée, depuis mille ans
ils se regardent et se menacent debout sur chaque rive:
jamais ennemi n’opprima l’Angleterre, que l’Ecossais n’ait
marché avec lui; jamais guerre civile n’embrasa les villes de
l’Ecosse, sans qu’un Anglais n’ait approché une torche de ses
murailles, et cela durera ainsi, et la haine sera implacable et
éternelle jusqu’au jour où un même parlement unira les deux
ennemies comme deux sœurs, et où un seul sceptre
s’étendra sur l’île tout entière.»
Nous entrâmes en Ecosse.

Nous visitâmes, Walter Scott à la main, toute cette terre poétique
que, pareil à un magicien qui évoque des fantômes, il a repeuplée
de ses antiques habitans, auxquels il a mêlé les originales et
gracieuses créations de sa fantaisie: nous retrouvâmes les sentiers
escarpés que suivait, sur son bon cheval Gustave, le prudent
Dalgetty. Nous côtoyâmes le lac sur lequel glissait, la nuit, comme
une vapeur, la Dame blanche d’Avenel. Nous allâmes nous asseoir
sur les ruines du château de Lochleven, à l’heure même où la reine
d’Ecosse s’en était échappée, et nous cherchâmes sur les bords de
la Tay le champ clos où Torquil du Chêne vit tomber ses sept fils
sous l’épée de l’armurier Smith, sans proférer d’autre plainte que ces
mots, qu’il répéta sept fois: Encore un pour Eachar!...
Cette excursion sera éternellement pour moi un rêve de bonheur
dont jamais n’approcheront les réalités de l’avenir: Pauline avait une
de ces organisations impressionnables comme il en faut aux artistes,
et sans laquelle un voyage n’est qu’un simple changement de
localités, une accélération dans le mouvement habituel de la vie, un
moyen de distraire son esprit par la vue même des objets qui
devraient l’occuper: pas un souvenir historique ne lui échappait; pas
une poésie de la nature, soit qu’elle se manifestât à nous dans la
vapeur du matin ou le crépuscule du soir, n’était perdue pour elle.
Quant à moi, j’étais sous l’empire d’un charme; jamais un seul mot
des événemens accomplis n’avait été prononcé entre nous depuis
l’heure où elle me les avait racontés; pour moi, le passé disparaissait
parfois comme s’il n’avait jamais existé. Le présent seul qui nous
réunissait était tout à mes yeux: jeté sur une terre étrangère, où je
n’avais que Pauline, où Pauline n’avait que moi, les liens qui nous
unissaient se resserraient chaque jour davantage par l’isolement;
chaque jour je sentais que je faisais un pas dans son cœur, chaque
jour un serrement de main, chaque jour un sourire, son bras appuyé
sur mon bras, sa tête posée sur mon épaule, était un nouveau droit
qu’elle me donnait sans s’en douter pour le lendemain; et plus elle
s’abandonnait ainsi, plus, tout en aspirant chaque émanation naïve
de son âme, plus je me gardais de lui parler d’amour, de peur qu’elle
ne s’aperçût que depuis longtemps nous avions dépassé les limites
de l’amitié.
Quant à la santé de Pauline, les prévisions du docteur s’étaient
réalisées en partie; cette activité que le changement des lieux et les
souvenirs qu’ils rappelaient entretenaient dans son esprit, détournait
sa pensée des souvenirs tristes qui l’oppressaient aussitôt qu’aucun
objet important ne venait l’en distraire. Elle-même commençait
presque à oublier, et à mesure que les abîmes du passé se
perdaient dans l’ombre, les sommets de l’avenir se coloraient d’un
jour nouveau. Sa vie, qu’elle avait crue bornée aux limites d’un
tombeau, commençait à reculer ses horizons moins sombres, et un
air de plus en plus respirable venait se mêler à l’atmosphère
étouffante au milieu de laquelle elle s’était sentie précipitée.
Nous passâmes l’été tout entier en Écosse; puis nous revînmes à
Londres: nous y retrouvâmes notre petite maison de Piccadilly, et ce
charme que l’esprit le plus enclin aux voyages éprouve dans les
premiers momens d’un retour. Je ne sais ce qui se passait dans le
cœur de Pauline, mais je sais que, quant à moi, je n’avais jamais été
si heureux.
Quant au sentiment qui nous unissait, il était pur comme la
fraternité: je n’avais pas, depuis un an, redit à Pauline que je
l’aimais, depuis un an Pauline ne m’avait point fait le moindre aveu,
et cependant nous lisions dans le cœur l’un de l’autre comme dans
un livre ouvert, et nous n’avions plus rien à nous apprendre.
Désirais-je plus que je n’avais obtenu?... Je ne sais; il y avait tant de
charme dans ma position, que j’aurais peut-être craint qu’un bonheur
plus grand ne la précipitât vers quelque dénoûment fatal et inconnu.
Si je n’étais pas amant, j’étais plus qu’un ami, plus qu’un frère; j’étais
l’arbre auquel, pauvre lierre, elle s’abritait, j’étais le fleuve qui
emportait sa barque à mon courant, j’étais le soleil d’où lui venait la
lumière; tout ce qui existait d’elle existait par moi, et probablement le
jour n’était pas loin où ce qui existait par moi existerait aussi pour
moi.
Nous en étions là de notre vie nouvelle, lorsqu’un jour je reçus
une lettre de ma mère. Elle m’annonçait qu’il se présentait pour ma
sœur un parti non-seulement convenable, mais avantageux: le
comte Horace de Beuzeval, qui joignait à sa propre fortune vingt-
cinq mille livres de rente qu’il avait héritées de sa première femme,
mademoiselle Pauline de Meulien, demandait Gabrielle en
mariage!...
Heureusement j’étais seul lorsque j’ouvris cette lettre, car ma
stupéfaction m’eût trahi: cette nouvelle que je recevais n’était-elle
pas bien étrange en effet, et quelque nouveau mystère de la
Providence ne se cachait-il pas dans cette bizarre prédestination qui
conduisait le comte Horace en face du seul homme dont il fût
connu? Quelque empire que je fusse parvenu à prendre sur moi-
même, Pauline ne s’en aperçut pas moins, en rentrant, qu’il m’était
arrivé, pendant son absence, quelque chose d’extraordinaire; au
reste, je n’eus pas de peine à lui donner le change, et, dès que je lui
eus dit que des affaires de famille me forçaient de faire un voyage
en France, elle attribua tout naturellement au chagrin de nous
séparer l’abattement dans lequel elle me retrouvait. Elle-même pâlit
et fut forcée de s’asseoir: c’était la première fois que nous nous
éloignions l’un de l’autre depuis près d’un an que je l’avais sauvée;
puis il y a, entre cœurs qui s’aiment, au moment d’une séparation,
quoique en apparence courte et sans danger, de ces pressentimens
intimes qui nous la font inquiétante et douloureuse, quelque chose
que la raison dise pour nous rassurer.
Je n’avais pas une minute à perdre; j’avais donc décidé que je
partirais le lendemain. Je montai chez moi pour faire quelques
préparatifs indispensables. Pauline descendit au jardin, où j’allai la
rejoindre aussitôt que ces apprêts furent terminés.
Je la vis assise sur le banc où elle m’avait raconté sa vie. Depuis
ce temps, je l’ai dit, comme si elle eût été réellement endormie dans
les bras de la mort, ainsi qu’on le croyait, aucun écho de la France
n’était venu la réveiller, mais peut être approchait-elle du terme de
cette tranquillité, et l’avenir pour elle allait-il douloureusement se
rattacher à ce passé que tous mes efforts avaient eu pour but de lui
faire oublier. Je la trouvai triste et rêveuse; je vins m’asseoir à son
côté; ses premiers mots m’apprirent la cause de sa préoccupation.
—Ainsi vous partez? me dit-elle.
—Il le faut, Pauline, répondis-je d’une voix que je cherchais à
rendre calme, vous savez mieux que personne qu’il y a des
événemens qui disposent de nous, et qui nous enlèvent aux lieux
que nous voudrions ne pas quitter d’une heure, comme le vent fait
d’une feuille. Le bonheur de ma mère, de ma sœur, le mien même,
dont je ne vous parlerais pas s’il était le seul compromis, dépendent
de ma promptitude à faire ce voyage.
—Allez donc, reprit Pauline tristement; allez, puisqu’il le faut;
mais n’oubliez pas que vous avez en Angleterre aussi une sœur qui
n’a pas de mère, dont le seul bonheur dépend désormais de vous, et
qui voudrait pouvoir quelque chose pour le vôtre!...
—Oh! Pauline! m’écriai-je en la pressant dans mes bras, dites-
moi, doutez-vous un instant de mon amour? croyez-vous que je ne
m’éloigne pas le cœur brisé? croyez-vous que le moment le plus
heureux de ma vie ne sera pas celui où je rentrerai dans cette petite
maison qui nous dérobe au monde tout entier?... Vivre avec vous de
cette vie de frère et de sœur, avec l’espoir seulement de jours plus
heureux encore, croyez-vous que ce n’était pas pour moi un bonheur
plus grand que je n’avais jamais osé l’espérer?..... oh! dites-moi, le
croyez-vous?...
—Oui, je le crois, me répondit Pauline; car il y aurait de
l’ingratitude à en douter. Votre amour a été pour moi si délicat et si
élevé, que je puis en parler sans rougir, comme je parlerais d’une de
vos vertus... Quant à ce bonheur plus grand que vous espérez,
Alfred, je ne le comprends pas!... Notre bonheur, j’en suis certaine,
tient à la pureté même de nos relations; et plus ma position est
étrange et sans pareille peut-être, plus je suis déliée de mes devoirs
envers la société, plus, pour moi-même, je dois être sévère à les
accomplir...
—Oh! oui... oui, lui dis-je, je vous comprends, et Dieu me punisse
si j’essayais jamais de détacher une fleur de votre couronne de
martyre pour y mettre en place un remords! mais enfin il peut arriver
tels événemens qui vous fassent libre... La vie même adoptée par le
comte, pardon si je reviens sur ce sujet, l’expose plus que tout
autre...
—Oh! oui... oui, je le sais... Aussi, croyez-le bien, je n’ouvre
jamais un journal sans frémir... L’idée que je puis voir le nom que j’ai
porté figurer dans quelque procès sanglant, l’homme que j’ai appelé
mon mari menacé d’une mort infâme... Eh bien!... que parlez-vous
de bonheur dans ce cas-là, en supposant que je lui survécusse?...
—Oh! d’abord... et avant tout, Pauline, vous n’en seriez pas
moins la plus pure comme la plus adorée des femmes..... N’a-t-il pas
pris soin de vous mettre à l’abri de lui-même, si bien qu’aucune
tache de sa boue ni de son sang ne peut vous atteindre?... Mais je
ne voulais point parler de cela, Pauline! Dans une attaque nocturne,
dans un duel même, le comte peut trouver la mort... Oh! c’est
affreux, je le sais, de n’avoir d’autre espérance de bonheur que celle
qui doit couler de la blessure ou sortir de la bouche d’un homme
avec son sang et son dernier soupir!... Mais enfin, pour vous-
même... une telle fin ne serait-elle pas un bienfait du hasard... un
oubli de la Providence?
—Eh bien?... dit en m’interrogeant Pauline.
—Eh bien! alors, Pauline, l’homme, qui, sans conditions, s’est fait
votre ami, votre protecteur, votre frère, n’aurait-il pas droit à un autre
titre?

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1 The Science of Biology 2

UNIT 1 PRINCIPLES OF CELLULAR LIFE

2 Life’s Chemical Basis 22

UNIT 2 GENETICS UNIT 6 HOW ANIMALS WORK

1 THE SCIENCE OF BIOLOGY

1.4 Organizing Information About Species 10

UNIT 1 PRINCIPLES OF CELLULAR LIFE

2 LIFE’S CHEMICAL BASIS

2.2 Why Electrons Matter 26

3.1 The Chemistry of Biology 38

APPLICATION: Fear of Frying 47

5 GROUND RULES OF METABOLISM

1 3 PATTERNS IN INHERITED TRAITS 1 5 BIOTECHNOLOGY

14.2 Examples of Autosomal Inheritance Patterns 224 UNIT 3 P R I N C I P L E S O F E VO L U T I O N

Autosomal Dominance 224 1 6 EVIDENCE OF EVOLUTION

APPLICATION: Shades of Skin 233

APPLICATION: Reflections of a Distant Past 269

UNIT 5 HOW PLANTS WORK

2 4 ANIMALS II: THE CHORDATES Simple Tissues 426

29.2 Cells of the Nervous System 502 Sense of Smell 524

UNIT 7 PRINCIPLES OF ECOLOGY 40.5 Predation Effects on Life History 722

Foraging in Fruit Flies 698 Expansions and Innovations 724

39.2 Instinct and Learning 700 A Demographic Transition 726

18 Life’s Origin and Early Evolution Improved descrip-

Chapter-Specific Changes (continued) involving resveratrol. New Application involves benefits of

37 Maintaining the Internal Environment Improved

38 Reproduction and Development Improved description

Tom McHugh/Science Source.

Stanton Belford Mark Hens Sean Rice

Solomon Gebru Jonas Okeagu Gordon Woolam

Joel Gluck Jennifer O’Neil David Wessner

Marla Gomez Peter J. Park Ovid Wong

Nazanin Z. Hebel Marc Perkins Xiaoning Zhang

MindTap for Biology: Concepts & Applications 10e

ASSIGN AND GRADE CONTENT THAT MATTERS

The researchers grew a mixture of native perennial grasses 15

abandoned. They measured the usable energy in biofuels made 10

“The technical support provided by

Instructor Companion Site Everything you need for

We can understand life by studying it at

Evolution underlies the unity and diversity of life.

Shared core processes and features that are

The field of biology relies upon experimentation and the

Science addresses only testable ideas about

Near a tent serving as a makeshift laboratory, herpetologist

Photograph by Tim Laman/National Geographic Creative.

Biologists view life as having nested levels of organization;

2 molecule 7 multicelled organism

Figure 1.2 Levels of life’s organization.

All living things share a particular set of key features. You

ENERGY AND NUTRIENTS

Cellular basis All living things consist of one or more cells.

6 INTRODUCTION (3) top, © Victoria Pinder, http://www.flickr.com/photos/vixstarplus; bottom producer, Zhemchuzhina/

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C R E D I T: (4) iStockphoto.com/gvillani. C H A P T E R 1 THE SCIENCE OF BIOLOGY 7

Protists are a group of extremely diverse eukaryotes that