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 “I would like to dedicate this book to my grandparents, the best teachers one could hope for.”
—Mike Harrington

“For Yusef, who finds the world a fascinating place, and in memory of Yasmin, who found comfort in nature.”
—Joan Sharp

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10 9 8 7 6 5 4 3 2 1
Library and Archives Canada Cataloguing in Publication
Freeman, Scott, 1955-Biological science / Scott Freeman, Mike Harrington, Joan Sharp. — 2nd Canadian ed.
ISBN 978-0-321-78871-9
1. Biology—Textbooks. I. Harrington, Mike, 1968- II. Sharp, Joan Catherine, 1951- III. Title.
QH308.2.F73 2012 570 C2012-903875-X

ISBN 10: 0-321-78871-0

ISBN 13: 978‐0‐32‐178871‐9
Brief Contents

1 Biology and the Tree of Life 1

UNIT 1 THE MOLECULES OF LIFE 16 UNIT 6 THE DIVERSIFICATION OF LIFE 546

2 Water and Carbon: The Chemical Basis of Life 16 28 Bacteria and Archaea 546
3 Protein Structure and Function 40 29 Protists 571
4 Nucleic Acids and the RNA World 62 30 Green Algae and Land Plants 599
5 An Introduction to Carbohydrates 75 31 Fungi 635
6 Lipids, Membranes, and the First Cells 87 32 An Introduction to Animals 660
33 Protostome Animals 684
UNIT 2 CELL STRUCTURE AND FUNCTION 112
34 Deuterostome Animals 709
7 Inside the Cell 112 35 Viruses 742
8 Cell–Cell Interactions 143
UNIT 7 HOW PLANTS WORK 764
9 Cellular Respiration and Fermentation 163
10 Photosynthesis 188 36 Plant Form and Function 764
11 The Cell Cycle 212 37 Water and Sugar Transport in Plants 788
38 Plant Nutrition 809
UNIT 3 GENE STRUCTURE AND EXPRESSION 232
39 Plant Sensory Systems, Signals, and Responses 829
12 Meiosis 232 40 Plant Reproduction 860
13 Mendel and the Gene 252
14 DNA and the Gene: Synthesis and Repair 281 UNIT 8 HOW ANIMALS WORK 883
15 How Genes Work 300 41 Animal Form and Function 883
16 Transcription, RNA Processing, and Translation 314 42 Water and Electrolyte Balance in Animals 904
17 Control of Gene Expression in Bacteria 333 43 Animal Nutrition 924
18 Control of Gene Expression in Eukaryotes 347 44 Gas Exchange and Circulation 946
19 Analyzing and Engineering Genes 368 45 Electrical Signals in Animals 973
20 Genomics 392 46 Animal Sensory Systems and Movement 996

UNIT 4 DEVELOPMENTAL BIOLOGY 410

47 Chemical Signals in Animals 1019
48 Animal Reproduction 1041
21 Principles of Development 410 49 The Immune System in Animals 1065
22 An Introduction to Animal Development 426
23 An Introduction to Plant Development 440 UNIT 9 ECOLOGY 1088

UNIT 5 EVOLUTIONARY PROCESSES AND PATTERNS 455

50 An Introduction to Ecology 1088
51 Behavioural Ecology 1121
24 Evolution by Natural Selection 455 52 Population Ecology 1141
25 Evolutionary Processes 477 53 Community Ecology 1166
26 Speciation 503 54 Ecosystems 1193
27 Phylogenies and the History of Life 521 55 Biodiversity and Conservation Biology 1219

v
Detailed Contents
About the Authors xx
Preface to Instructors xxi
Preface to Students: How to Use This Book xxxiv
1 Biology and the Tree of Life 1 3
1.1 What Does It Mean to Say That Something Is Alive? 1
1.2 The Cell Theory 2
Are All Organisms Made of Cells? 2
Where Do Cells Come From? 2
1.3 The Theory of Evolution by Natural Selection 4
What Is Evolution? 4
What Is Natural Selection? 4
CANADIAN RESEARCH 1.1 Artificial Selection on Bighorn Sheep
in Alberta 6
1.4 The Tree of Life 6
Using Molecules to Understand the Tree of Life 7
How Should We Name Branches on the Tree of Life? 9
1.5 Doing Biology 9
Why Do Giraffes Have Long Necks? An Introduction to
Hypothesis Testing 9
How Do Ants Navigate? An Introduction to Experimental Design 11
CHAPTER REVIEW 13
UNIT 1 THE MOLECULES OF LIFE 16
2 Water and Carbon: The Chemical
Basis of Life 16
2.1 Atoms, Ions, and Molecules: The Building Blocks of
Chemical Evolution 17
Basic Atomic Structure 17
How Does Covalent Bonding Hold Molecules Together? 18
Ionic Bonding, Ions, and the Electron-Sharing Continuum 19
Some Simple Molecules Formed from C, H, N, and O 20
The Geometry of Simple Molecules 21
Representing Molecules 21
Basic Concepts in Chemical Reactions 22
2.2 The Early Oceans and the Properties of Water 23
Why Is Water Such an Efficient Solvent? 23
How Does Water’s Structure Correlate with Its Properties? 23
Acid–Base Reactions Involve a Transfer of Protons 26
2.3 Chemical Reactions, Chemical Evolution, and
Chemical Energy 28
How Do Chemical Reactions Happen? 28
What Is Energy? 28
Chemical Evolution: A Model System 30
How Did Chemical Energy Change during Chemical Evolution? 34
2.4 The Importance of Carbon 34
Linking Carbon Atoms Together 35
Functional Groups 36
vi
CANADIAN RESEARCH 2.1 The Carbon-Rich Tagish Lake
Meteorite 37
CHAPTER REVIEW 38
Protein Structure and Function 40
3.1 Early Origin-of-Life Experiments 41
3.2 Amino Acids and Polymerization 42
The Structure of Amino Acids 42
The Nature of Side Chains 42
How Do Amino Acids Link to Form Proteins? 44
3.3 Proteins Are the Most Versatile Large Molecules in
Cells 46
CANADIAN RESEARCH 3.1 Designing New Proteins 47
3.4 What Do Proteins Look Like? 47
Primary Structure 48
Secondary Structure 48
Tertiary Structure 49
Quaternary Structure 50
CANADIAN RESEARCH 3.2 Spider Silk Proteins 52
Folding and Function 52
3.5 Enzymes: An Introduction to Catalysis 54
Enzymes Help Reactions Clear Two Hurdles 55
How Do Enzymes Work? 56
Was the First Living Entity a Protein Catalyst? 59
CHAPTER REVIEW 60
4 Nucleic Acids and the RNA World 62
4.1 What Is a Nucleic Acid? 62
Could Chemical Evolution Result in the Production of
Nucleotides? 63
How Do Nucleotides Polymerize to Form Nucleic Acids? 64
4.2 DNA Structure and Function 65
What Is the Nature of DNA’s Secondary Structure? 66
DNA Functions as an Information-Containing Molecule 67
Is DNA a Catalytic Molecule? 69
4.3 RNA Structure and Function 69
Structurally, RNA Differs from DNA 69
RNA’s Structure Makes It an Extraordinarily Versatile Molecule 70
RNA Is an Information-Containing Molecule 71
RNA Can Function as a Catalytic Molecule 71
4.4 The First Life Form 71
CANADIAN RESEARCH 4.1 Designing New Deoxyribozymes 72
CHAPTER REVIEW 73
5 An Introduction to Carbohydrates 75
5.1 Sugars as Monomers 75
How Monosaccharides Differ 76
Monosaccharides and Chemical Evolution 76
5.2 The Structure of Polysaccharides 77
Starch: A Storage Polysaccharide in Plants 78
Glycogen: A Highly Branched Storage Polysaccharide in Animals 78
Cellulose: A Structural Polysaccharide in Plants 78
Chitin: A Structural Polysaccharide in Fungi and Animals 80
Peptidoglycan: A Structural Polysaccharide in Bacteria 80
Polysaccharides and Chemical Evolution 80
5.3 What Do Carbohydrates Do? 80
The Role of Carbohydrates as Structural Molecules 81
The Role of Carbohydrates in Cell Identity 81
The Role of Carbohydrates in Energy Storage 81
CANADIAN ISSUES 5.1 Raymond Lemieux and the Synthesis of
Sugars 82
CANADIAN RESEARCH 5.1 Natural and Artificial Sweeteners 84
CHAPTER REVIEW 85
6 Lipids, Membranes, and the First Cells
6.1 Lipids 88
A Look at Three Types of Lipids Found in Cells 88
The Structures of Membrane Lipids 89
6.2 Phospholipid Bilayers 90
Artificial Membranes as an Experimental System 90
Selective Permeability of Lipid Bilayers 91
How Does Lipid Structure Affect Membrane Properties? 92
How Does Temperature Affect the Fluidity and Permeability of
Membranes? 93
CANADIAN ISSUES 6.1 Lipids in Our Diet: Cholesterol,
Unsaturated Oils, Saturated Fats, and Trans Fats 94
6.3 Why Molecules Move across Lipid Bilayers: Diffusion
and Osmosis 96
Diffusion 96
Osmosis 96
CANADIAN RESEARCH 6.1 Liposomal Nanomedicines 98
6.4 Membrane Proteins 99
Evolution of the Fluid-Mosaic Model 99
Systems for Studying Membrane Proteins 101
Protein Transport I: Facilitated Diffusion via Channel
Proteins 102
Protein Transport II: Facilitated Diffusion via Carrier Proteins 104
Protein Transport III: Active Transport by Pumps 104
Plasma Membranes and the Intracellular Environment 106
CHAPTER REVIEW 107
The Big Picture: Macromolecules 110
UNIT 2 CELL STRUCTURE AND FUNCTION 112
7 Inside the Cell 112
7.1 Bacterial and Archaeal Cell Structures and Their
Functions 112
A Revolutionary New View 113
Prokaryotic Cell Structures: A Parts List 113
CANADIAN RESEARCH 7.1 Bacteria Cells Have Their Own
Cytoskeleton 115
7.2 Eukaryotic Cell Structures and Their Functions 116
The Benefits of Organelles 116
Eukaryotic Cell Structures: A Parts List 117
7.3 Putting the Parts into a Whole 126
Structure and Function at the Whole-Cell Level 126
The Dynamic Cell 126
7.4 Cell Systems I: Nuclear Transport 127
Structure and Function of the Nuclear Envelope 127
How Are Molecules Imported into the Nucleus? 128
7.5 Cell Systems II: The Endomembrane System
Manufactures and Ships Proteins 129
Studying the Pathway through the Endomembrane
System 129
Entering the Endomembrane System: The Signal
Hypothesis 131
Moving from the ER to the Golgi 132
What Happens inside the Golgi Apparatus? 133
How Do Proteins Reach Their Destinations? 133
7.6 Cell Systems III: The Dynamic Cytoskeleton 134
87
Actin Filaments 134
Intermediate Filaments 135
Microtubules 136
CANADIAN RESEARCH 7.2 Pathogenic Bacteria Alter the
Cytoskeleton of Human Cells 137
Flagella and Cilia: Moving the Entire Cell 138
CHAPTER REVIEW 140
8 Cell–Cell Interactions 143
8.1 The Cell Surface 144
The Structure and Function of an Extracellular Layer 144
The Cell Wall in Plants 144
The Extracellular Matrix in Animals 145
8.2 How Do Adjacent Cells Connect and
Communicate? 146
Cell–Cell Attachments in Eukaryotes 147
Cells Communicate via Cell–Cell Gaps 150
8.3 How Do Distant Cells Communicate? 151
Cell–Cell Signalling in Multicellular Organisms 151
Signal Reception 152
Signal Processing 152
CANADIAN RESEARCH 8.1 The Discovery of Insulin 156
Signal Response 158
Signal Deactivation 158
Cross-Talk: Synthesizing Input from Many Signals 158
Quorum Sensing in Bacteria 159
CANADIAN RESEARCH 8.2 How Do Intracellular Proteins Bind
to Receptor Tyrosine Kinases? 160
CHAPTER REVIEW 161
9 Cellular Respiration and Fermentation 163
9.1 The Nature of Chemical Energy and Redox
Reactions 164
The Structure and Function of ATP 164
What Is a Redox Reaction? 166
9.2 An Overview of Cellular Respiration 168
9.3 Glycolysis: Processing Glucose to Pyruvate 169
Glycolysis Is a Sequence of 10 Reactions 169
How Is Glycolysis Regulated? 170
9.4 Processing Pyruvate to Acetyl CoA 171
DETAILED CONTENTS vii
 9.5 The Citric Acid Cycle: Oxidizing Acetyl
CoA to CO2 173
How Is the Citric Acid Cycle Regulated? 173
What Happens to the NADH and FADH2? 173
9.6 Electron Transport and Chemiosmosis: Building
a Proton Gradient to Produce ATP 176
Components of the Electron Transport Chain 176
The Chemiosmosis Hypothesis 177
How Is the Electron Transport Chain Organized? 178
The Discovery of ATP Synthase 178
Organisms Use a Diversity of Electron Acceptors 179
CANADIAN RESEARCH 9.1 The ATP Synthase 180
9.7 Fermentation 181
CANADIAN ISSUES 9.1 Making Biofuels with Fermentation
and Anaerobic Respiration 183
9.8 How Does Cellular Respiration Interact with Other
Metabolic Pathways? 184
Catabolic Pathways Break Down Molecules as Fuel 184
Anabolic Pathways Synthesize Key Molecules 185
CHAPTER REVIEW 185
10 Photosynthesis 188
10.1 Photosynthesis Harnesses Sunlight to Make
Carbohydrate 188
Photosynthesis: Two Linked Sets of Reactions 189
Photosynthesis Occurs in Chloroplasts 190
10.2 How Does Chlorophyll Capture Light Energy? 190
Photosynthetic Pigments Absorb Light 191
When Light Is Absorbed, Electrons Enter an Excited State 193
10.3 The Discovery of Photosystems I and II 195
How Does Photosystem II Work? 196
How Does Photosystem I Work? 198
The Z Scheme: Photosystems II and I Work Together 198
10.4 How Is Carbon Dioxide Reduced to Produce
Glucose? 200
The Calvin Cycle Fixes Carbon 201
The Discovery of Rubisco 202
Carbon Dioxide Enters Leaves through Stomata 203
Mechanisms for Increasing CO2 Concentration Near
Rubisco 204
How Is Photosynthesis Regulated? 205
What Happens to the Sugar That Is Produced by
Photosynthesis? 205
CANADIAN RESEARCH 10.1 Turning C3 Plants into C4
Plants 206
CHAPTER REVIEW 207
The Big Picture: Energy for Life 210
11 The Cell Cycle 212
11.1 Mitosis and the Cell Cycle 213
What Is a Chromosome? 213
Cells Alternate between M Phase and Interphase 214
The Discovery of S Phase 214
The Discovery of the Gap Phases 214
The Cell Cycle 214
viii DETAILED CONTENTS
11.2 How Does Mitosis Take Place? 215
Proteins Needed for Mitosis 215
Cytokinesis Results in Two Daughter Cells 218
How Do Chromosomes Move during Mitosis? 218
11.3 Control of the Cell Cycle 220
The Discovery of Cell-Cycle Regulatory Molecules 220
CANADIAN RESEARCH 11.1 Yoshio Masui and the
Discovery of MPF 222
Cell-Cycle Checkpoints Can Arrest the Cell Cycle 223
11.4 Cancer: Out-of-Control Cell Division 225
Properties of Cancer Cells 225
Cancer Involves Loss of Cell-Cycle Control 226
CANADIAN RESEARCH 11.2 A Newly Discovered
Property of Cancer Cells 227
CHAPTER REVIEW 229
UNIT 3 GENE STRUCTURE AND EXPRESSION 232
12 Meiosis 232
12.1 How Does Meiosis Occur? 233
Chromosomes Come in Distinct Types 233
The Concept of Ploidy 233
An Overview of Meiosis 234
The Phases of Meiosis I 237
The Phases of Meiosis II 238
A Closer Look at Prophase I 241
CANADIAN RESEARCH 12.1 The Proteins Required for
Prophase I of Meiosis 242
12.2 The Consequences of Meiosis 242
Chromosomes and Heredity 243
Independent Assortment Produces Genetic Variation 243
A Benefit of Crossing Over 244
How Does Fertilization Affect Genetic Variation? 244
12.3 Why Does Meiosis Exist? 245
The Paradox of Sex 245
The Purifying Selection Hypothesis 245
The Changing-Environment Hypothesis 246
12.4 Mistakes in Meiosis 247
How Do Mistakes Occur? 247
Why Do Mistakes Occur? 248
CHAPTER REVIEW 249
13 Mendel and the Gene 252
13.1 Mendel’s Experimental System 252
What Questions Was Mendel Trying to Answer? 253
Garden Peas Served as the First Model Organism in Genetics 253
13.2 Mendel’s Experiments with a Single Trait 254
The Monohybrid Cross 254
Particulate Inheritance 256
13.3 Mendel’s Experiments with Two Traits 258
The Dihybrid Cross 258
Using a Testcross to Confirm Predictions 260
13.4 The Chromosome Theory of Inheritance 261
Meiosis Explains Mendel’s Principles 261
Testing the Chromosome Theory 263
13.5 Extending Mendel’s Rules 265
Linkage: What Happens When Genes Are Located on the
Same Chromosome? 265
Do Heterozygotes Always Have a Dominant or Recessive
Phenotype? 267
BOX 13.1 Quantitative Methods: Linkage 268
How Many Alleles and Phenotypes Exist? 269
Does Each Gene Affect Just One Trait? 269
Are Phenotypes Determined by Genes? 269
What about Traits Like Human Height and Intelligence? 270
13.6 Applying Mendel’s Rules to Humans 272
Identifying Human Alleles as Recessive or Dominant 272
CANADIAN RESEARCH 13.1 The Genetics of Dog
Coat Colour 274
Identifying Human Traits as Autosomal or Sex-Linked 275
CHAPTER REVIEW 276
14 DNA and the Gene: Synthesis and Repair
14.1 What Are Genes Made Of? 282
The Hershey–Chase Experiment 282
The Secondary Structure of DNA 283
14.2 Testing Early Hypotheses about DNA Synthesis:
The Meselson–Stahl Experiment 284
14.3 A Comprehensive Model for DNA Synthesis 285
How Does Replication Get Started? 287
How Is the Helix Opened and Stabilized? 287
How Is the Leading Strand Synthesized? 288
How Is the Lagging Strand Synthesized? 289
14.4 Replicating the Ends of Linear Chromosomes 292
CANADIAN RESEARCH 14.1 Telomeres, Telomerase, and
Cancer 294
14.5 Repairing Mistakes and Damage 294
Correcting Mistakes in DNA Synthesis 295
Repairing Damaged DNA 296
Xeroderma Pigmentosum: A Case Study 296
CHAPTER REVIEW 297
15 How Genes Work 300
15.1 What Do Genes Do? 301
The One-Gene, One-Enzyme Hypothesis 301
An Experimental Test of the Hypothesis 301
15.2 The Central Dogma of Molecular Biology 303
The Genetic Code Hypothesis 303
RNA as the Intermediary between Genes and Proteins 303
Dissecting the Central Dogma 304
15.3 The Genetic Code 306
How Long Is a Word in the Genetic Code? 306
How Did Researchers Crack the Code? 307
15.4 What Is the Molecular Basis of Mutation? 309
Point Mutation 309
Chromosome-Level Mutations 310
CANADIAN RESEARCH 15.1 The Mutations Responsible for
Himalayan Fur Colour in Mink and Mice 311
CHAPTER REVIEW 312
16 Transcription, RNA Processing,
and Translation 314
16.1 An Overview of Transcription 314
Characteristics of RNA Polymerase 315
Initiation: How Does Transcription Begin? 316
Elongation and Termination 317
16.2 RNA Processing in Eukaryotes 318
The Unexpected Discovery of Eukaryotic Genes in Pieces 318
RNA Splicing 319
Adding Caps and Tails to Transcripts 320
16.3 An Introduction to Translation 321
Ribosomes Are the Site of Protein Synthesis 321
Comparing Translation in Bacteria and Eukaryotes 321
How Does an mRNA Triplet Specify an Amino Acid? 321
16.4 The Structure and Function of Transfer RNA 323
What Do tRNAs Look Like? 324
281 How Many tRNAs Are There? 325
16.5 The Structure and Function of Ribosomes 325
Initiating Translation 326
Elongation: Extending the Polypeptide 327
Terminating Translation 327
Posttranslational Modifications 329
CANADIAN RESEARCH 16.1 RNA Synthesis in
Mitochondria 329
CHAPTER REVIEW 330
17 Control of Gene Expression in Bacteria 333
17.1 Gene Regulation and Information Flow 333
Mechanisms of Regulation—An Overview 334
Metabolizing Lactose—A Model System 335
17.2 Identifying Genes under Regulatory Control 336
Replica Plating to Find Mutant Genes 336
Different Classes of Lactose Metabolism Mutants 337
Several Genes Are Involved in Lactose Metabolism 338
17.3 Mechanisms of Negative Control: Discovery of the
Repressor 338
The lac Operon 339
Why Has the lac Operon Model Been So Important? 340
17.4 Mechanisms of Positive Control: Catabolite
Repression 341
The CAP Protein and Binding Site 341
How Does Glucose Influence Formation of the CAP–cAMP
Complex? 341
CANADIAN RESEARCH 17.1 Bacterial Gene Expression and
Probiotic Dairy Products 343
CHAPTER REVIEW 345
18 Control of Gene Expression in Eukaryotes 347
18.1 Mechanisms of Gene Regulation in
Eukaryotes—An Overview 348
18.2 Chromatin Remodelling 348
What Is Chromatin’s Basic Structure? 348
Evidence That Chromatin Structure Is Altered in Active Genes 349
DETAILED CONTENTS ix
 How Is Chromatin Altered? 350
Chromatin Modifications Can Be Inherited 351
18.3 Initiating Transcription: Regulatory Sequences and
Regulatory Proteins 351
Some Regulatory Sequences Are Near the Promoter 351
Some Regulatory Sequences Are Far from the Promoter 352
The Role of Regulatory Proteins in Differential
Gene Expression 354
The Initiation Complex 354
18.4 Posttranscriptional Control 356
Alternative Splicing of mRNAs 356
mRNA Stability and RNA Interference 357
How Is Translation Controlled? 358
Posttranslational Control 358
18.5 How Does Gene Expression in Bacteria Compare with
That in Eukaryotes? 359
18.6 Linking Cancer with Defects in Gene Regulation 360
Causes of Uncontrolled Cell Growth 360
p53: A Case Study 361
CANADIAN RESEARCH 18.1 Chromatin Remodelling,
Gene Transcription, and Cancer 362
CHAPTER REVIEW 363
The Big Picture: Genetic Information 366
Using the Ti Plasmid to Produce Golden Rice 389
CHAPTER REVIEW 389
20 Genomics 392
20.1 Whole-Genome Sequencing 392
How Are Complete Genomes Sequenced? 393
Which Genomes Are Being Sequenced, and Why? 394
Which Sequences Are Genes? 395
20.2 Bacterial and Archaeal Genomes 396
The Natural History of Prokaryotic Genomes 396
Lateral Gene Transfer 397
Environmental Sequencing 397
CANADIAN ISSUES 20.1 Genome Canada 398
20.3 Eukaryotic Genomes 398
Parasitic and Repeated Sequences 399
Gene Families 401
Insights from the Human Genome Project 402
CANADIAN RESEARCH 20.1 Human Genetic
Variation 404
20.4 Functional Genomics and Proteomics 406
What Is Functional Genomics? 406
What Is Proteomics? 406
Applied Genomics in Action: Understanding Cancer 407

19
CHAPTER REVIEW 408
Analyzing and Engineering Genes 368

19.1 Case 1—The Effort to Cure Pituitary Dwarfism: Basic UNIT 4 DEVELOPMENTAL BIOLOGY 410
Recombinant DNA Technologies 368
Why Did Early Efforts to Treat the Disease Fail? 369
Steps in Engineering a Safe Supply of Growth Hormone 369
21 Principles of Development 410

19.2 Case 2—Amplification of Fossil DNA: The Polymerase 21.1 Shared Developmental Processes 411
Chain Reaction 374 Cell Proliferation 411
Requirements of PCR 374 Programmed Cell Death 412
PCR in Action 375 Cell Movement or Cell Growth 412
CANADIAN RESEARCH 19.1 Ancient DNA in Canada 376 Cell Differentiation 413
Cell–Cell Interactions 413
19.3 Case 3—Sanger’s Breakthrough Innovation: Dideoxy
DNA Sequencing 377 21.2 The Role of Differential Gene Expression
The Logic of Dideoxy Sequencing 378 in Development 413
“Next-Generation” Sequencing 379 Evidence That Differentiated Plant Cells Are
CANADIAN RESEARCH 19.2 Michael Smith and the Invention of Genetically Equivalent 413
Site-Directed Mutagenesis 379 Evidence That Differentiated Animal Cells Are
Genetically Equivalent 413
19.4 Case 4—The Huntington’s Disease Story: Finding How Does Differential Gene Expression Occur? 414
Genes by Mapping 381 CANADIAN RESEARCH 21.1 The First Cloned
How Was the Huntington’s Disease Gene Found? 381 Drosophila 415
What Are the Benefits of Finding a Disease Gene? 383
Ethical Concerns over Genetic Testing 383
21.3 Cell–Cell Signals Trigger Differential Gene
Expression 415
19.5 Case 5—Severe Immune Disorders: The Potential of Master Regulators Set Up the Major Body Axes 416
Gene Therapy 385 Regulatory Genes Provide Increasingly Specific Positional
How Can Novel Alleles Be Introduced into Human Cells? 385 Information 417
Using Gene Therapy to Treat X-Linked Immune Cell–Cell Signals and Regulatory Genes Are Evolutionarily
Deficiency 386 Conserved 419
Ethical Concerns over Gene Therapy 387 CANADIAN RESEARCH 21.2 Stem Cells and Stem
19.6 Case 6—The Development of Golden Cell Therapies 420
Rice: Biotechnology in Agriculture 387 21.4 Changes in Developmental Pathways Underlie
Rice as a Target Crop 388 Evolutionary Change 423
Synthesizing b-Carotene in Rice 388
CHAPTER REVIEW 424
The Agrobacterium Transformation System 388

x DETAILED CONTENTS
22 An Introduction to Animal Development 426
24.3 The Process of Evolution: How Does Natural Selection
Work? 464
22.1 Gamete Structure and Function 427 Darwin’s Four Postulates 464
Sperm Structure and Function 427 The Biological Definitions of Fitness and Adaptation 464
Egg Structure and Function 428 24.4 Evolution in Action: Recent Research on Natural
22.2 Fertilization 428 Selection 465
How Do Gametes from the Same Species Recognize Each Case Study 1: How Did Mycobacterium tuberculosis Become
Other? 429 Resistant to Antibiotics? 465
Why Does Only One Sperm Enter the Egg? 429 CANADIAN ISSUES 24.1 Evolution in Action: Do Hunting and
22.3 Cleavage 430 Fishing Select for Undesirable Traits? 467
Partitioning Cytoplasmic Determinants 431 Case Study 2: Why Are Beak Size, Beak Shape, and Body Size
Cleavage in Mammals 431 Changing in Galápagos Finches? 468
22.4 Gastrulation 432 24.5 Common Misconceptions about Natural Selection and
Formation of Germ Layers 432 Adaptation 471
Definition of Body Axes 433 Selection Acts on Individuals, but Evolutionary Change
Occurs in Populations 471
22.5 Organogenesis 434
Evolution Is Not Goal Directed 472
Organizing Mesoderm into Somites: Precursors of Muscle,
Organisms Do Not Act for the Good of the Species 472
Skeleton, and Skin 434
Limitations of Natural Selection 473
Differentiation of Muscle Cells 436
CANADIAN RESEARCH 22.1 Apoptosis during the CHAPTER REVIEW 474
Morphogenesis of Chick Embryos 436
CHAPTER REVIEW 438 25 Evolutionary Processes 477

25.1 Analyzing Change in Allele Frequencies: The Hardy–

23 An Introduction to Plant Development
23.1 Gametogenesis, Pollination, and Fertilization 441
How Are Sperm and Egg Produced? 441
Pollen–Stigma Interactions 441
Double Fertilization 442
23.2 Embryogenesis 443
What Happens during Plant Embryogenesis? 443
Which Genes and Proteins Set Up Body Axes? 445
23.3 Vegetative Development 446
Meristems Provide Lifelong Growth and Development 446
Which Genes and Proteins Determine Leaf Shape? 447
CANADIAN RESEARCH 23.1 Apoptosis during the Formation of
Plant Leaves 448
23.4 Reproductive Development 450
The Floral Meristem and the Flower 450
The Genetic Control of Flower Structures 450
CHAPTER REVIEW 453
UNIT 5 EVOLUTIONARY PROCESSES AND PATTERNS 455
440 Weinberg Principle 478
The Gene Pool Concept 478
Deriving the Hardy–Weinberg Principle 478
The Hardy–Weinberg Model Makes Important Assumptions 479
How Does the Hardy–Weinberg Principle Serve as a Null
Hypothesis? 480
25.2 Types of Natural Selection 482
Directional Selection 482
Stabilizing Selection 483
Disruptive Selection 484
Balancing Selection 485
25.3 Genetic Drift 485
Simulation Studies of Genetic Drift 485
Experimental Studies of Genetic Drift 487
What Causes Genetic Drift in Natural Populations? 487
25.4 Gene Flow 489
Gene Flow in Natural Populations 489
How Does Gene Flow Affect Fitness? 490
25.5 Mutation 490
Mutation as an Evolutionary Mechanism 490
Experimental Studies of Mutation 491

24 Evolution by Natural Selection 455 25.6 Nonrandom Mating 492

Inbreeding 493
24.1 The Evolution of Evolutionary Thought 456 Assortative Mating 494
Plato and Typological Thinking 456 Sexual Selection 495
Aristotle and the Great Chain of Being 456 CANADIAN RESEARCH 25.1 Evolution in Action: Kermode
Lamarck and the Idea of Evolution as Change through Time 456 Bears and Newfoundland Moose 496
Darwin and Wallace and Evolution by Natural Selection 456
CHAPTER REVIEW 500
24.2 The Pattern of Evolution: Have Species Changed
through Time? 457
Evidence for Change through Time 457 26 Speciation 503
Evidence of Descent from a Common Ancestor 459 26.1 How Are Species Defined and Identified? 503
Evolution’s “Internal Consistency”—The Importance of The Biological Species Concept 504
Independent Data Sets 463 The Morphospecies Concept 505

DETAILED CONTENTS xi
 The Ecological Species Concept 505
The Phylogenetic Species Concept 505
Species Definitions in Action: The Case of the Dusky
Seaside Sparrow 506
26.2 Isolation and Divergence in Allopatry 508
Dispersal and Colonization Isolate Populations 508
Vicariance Isolates Populations 509
26.3 Isolation and Divergence in Sympatry 509
Can Natural Selection Cause Speciation Even When Gene
Flow Is Possible? 509
How Can Polyploidy Lead to Speciation? 510
26.4 What Happens When Isolated Populations Come
into Contact? 513
Reinforcement 513
CANADIAN RESEARCH 26.1 Dolph Schluter Studies
New Species 514
Hybrid Zones 515
New Species through Hybridization 516
CHAPTER REVIEW 518
27 Phylogenies and the History of Life
Using Enrichment Cultures 552
Using Direct Sequencing 552
Evaluating Molecular Phylogenies 553
28.3 What Themes Occur in the Diversification of Bacteria
and Archaea? 555
Morphological Diversity 555
Metabolic Diversity 556
Ecological Diversity and Global Change 560
28.4 Key Lineages of Bacteria and Archaea 563
CANADIAN RESEARCH 28.1 Is There a Universal Tree of Life? 563
Bacteria 564
Archaea 564
■ Bacteria 7 Firmicutes 565
■ Bacteria 7 Spirochaetes (Spirochetes) 565
■ Bacteria 7 Actinobacteria 566
■ Bacteria 7 Chlamydiae 566
■ Bacteria 7 Cyanobacteria 567
■ Bacteria 7 Proteobacteria 567
■ Archaea 7 Crenarchaeota 568
■ Archaea 7 Euryarchaeota 568
CHAPTER REVIEW 569
521

27.1 Tools for Studying History: Phylogenetic Trees 521 29 Protists 571
How Do Researchers Estimate Phylogenies? 522 29.1 Why Do Biologists Study Protists? 572
How Can Biologists Distinguish Homology from Homoplasy? 522 Impacts on Human Health and Welfare 572
Whale Evolution: A Case History 524 Ecological Importance of Protists 574
27.2 Tools for Studying History: The Fossil Record 526 CANADIAN RESEARCH 29.1 How Will Phytoplankton Respond
How Do Fossils Form? 526 to Elevated CO2 Levels? 575
Limitations of the Fossil Record 527 29.2 How Do Biologists Study Protists? 577
Life’s Time Line 528 Microscopy: Studying Cell Structure 577
27.3 Adaptive Radiation 530 Evaluating Molecular Phylogenies 578
CANADIAN ISSUES 27.1 iBOL: The International Barcode of Life Discovering New Lineages via Direct Sequencing 578
Project 531 29.3 What Themes Occur in the Diversification of
Why Do Adaptive Radiations Occur? 532 Protists? 579
The Cambrian Explosion 534 What Morphological Innovations Evolved in Protists? 579
CANADIAN RESEARCH 27.1 The Burgess Shale: A Window into How Do Protists Obtain Food? 583
the Cambrian Explosion 536 How Do Protists Move? 585
27.4 Mass Extinction 538 How Do Protists Reproduce? 586
How Do Mass Extinctions Differ from Background Life Cycles—Haploid Dominated versus Diploid Dominated 587
Extinctions? 538 29.4 Key Lineages of Protists 588
The End-Permian Extinction 539 Amoebozoa 588
What Killed the Dinosaurs? 539 Excavata 588
CHAPTER REVIEW 542 Plantae 589
The Big Picture: Evolution 544 Rhizaria 590
Alveolata 590
Stramenopila (Heterokonta) 590
UNIT 6 THE DIVERSIFICATION OF LIFE 546 ■ Amoebozoa 7 Myxogastrida (Plasmodial Slime Moulds) 590
■ Excavata 7 Parabasalida 591
■ Excavata 7 Diplomonadida 591
28 Bacteria and Archaea 546 ■ Excavata 7 Euglenida 592
■ Plantae 7 Rhodophyta (Red Algae) 592
28.1 Why Do Biologists Study Bacteria and Archaea? 547 ■ Rhizaria 7 Foraminifera 593
Biological Impact 547
■ Alveolata 7 Ciliata 593
Medical Importance 548
■ Alveolata 7 Dinoflagellata 594
Role in Bioremediation 550
■ Alveolata 7 Apicomplexa 594
Extremophiles 551
■ Stramenopila 7 Oomycota (Water Moulds) 595
CANADIAN ISSUES 28.1 Bioremediation of Polluted Soils in
■ Stramenopila 7 Diatoms 595
Canada’s High Arctic 551
■ Stramenopila 7 Phaeophyta (Brown Algae) 596
28.2 How Do Biologists Study Bacteria and Archaea? 552 CHAPTER REVIEW 596

xii DETAILED CONTENTS

30 Green Algae and Land Plants 599
Evaluating Molecular Phylogenies 640
Experimental Studies of Mutualism 641
30.1 Why Do Biologists Study the Green Algae and Land 31.3 What Themes Occur in the Diversification
Plants? 599 of Fungi? 643
Plants Provide Ecosystem Services 600 Fungi Participate in Several Types of Mutualisms 643
Plants Provide Humans with Food, Fuel, Fibre, Building CANADIAN ISSUES 31.1 Ectomycorrhizal Fungi Are Important
Materials, and Medicines 601 in Regeneration of Forest Stands Following
30.2 How Do Biologists Study Green Algae and Land Clear-Cutting 645
Plants? 602 CANADIAN RESEARCH 31.1 The Effect of Gap Size on
Analyzing Morphological Traits 602 Colonization of Conifer Seedling Roots by
Using the Fossil Record 603 Ectomycorrhizal Fungi 646
Evaluating Molecular Phylogenies 604 What Adaptations Make Fungi Such Effective
Decomposers? 647
30.3 What Themes Occur in the Diversification of Land Variation in Reproduction 648
Plants? 606 Four Major Types of Life Cycles 650
The Transition to Land, I: How Did Plants Adapt to Dry
Conditions? 606 31.4 Key Lineages of Fungi 652
Mapping Evolutionary Changes on the Phylogenetic Tree 608 ■ Fungi 7 Microsporidia 652
The Transition to Land, II: How Do Plants Reproduce in Dry ■ Fungi 7 Chytrids 653
Conditions? 609 ■ Fungi 7 Zygomycetes 654
CANADIAN RESEARCH 30.1 Flowering Plants and Their ■ Fungi 7 Glomeromycota 654
Pollinators 617 ■ Fungi 7 Basidiomycota (Club Fungi) 655
The Angiosperm Radiation 619 ■ Fungi 7 Ascomycota 7 Lichen-Formers 656
■ Fungi 7 Ascomycota 7 Non-lichen-Formers 657
30.4 Key Lineages of Green Algae and Land Plants 620
Green Algae 620 CHAPTER REVIEW 658
Nonvascular Plants (“Bryophytes”) 620
Seedless Vascular Plants 621
Seed Plants 621 32 An Introduction to Animals 660
■ Green Algae 7 Ulvophyceae (Ulvophytes) 622 32.1 Why Do Biologists Study Animals? 661
■ Green Algae 7 Coleochaetophyceae (Coleochaetes) 622 Biological Importance 661
■ Green Algae 7 Charophyceae (Stoneworts) 623 Role in Human Health and Welfare 661
■ Nonvascular Plants 7 Hepaticophyta (Liverworts) 623
■ Nonvascular Plants 7 Bryophyta (Mosses) 624
32.2 How Do Biologists Study Animals? 662
■ Nonvascular Plants 7 Anthocerophyta (Hornworts) 625
Analyzing Comparative Morphology 662
■ Seedless Vascular Plants 7 Lycophyta (Lycophytes, or Club
Evaluating Molecular Phylogenies 667
Mosses) 625 32.3 What Themes Occur in the Diversification of
■ Seedless Vascular Plants 7 Psilotophyta (Whisk Ferns) 626 Animals? 669
■ Seedless Vascular Plants 7 Equisetophyta (or Sphenophyta) Sensory Organs 669
(Horsetails) 626 Feeding 670
■ Seedless Vascular Plants 7 Pteridophyta (Ferns) 627 CANADIAN RESEARCH 32.1 The World’s Oldest Radula 672
■ Seed Plants 7 Gymnosperms 7 Cycadophyta (Cycads) 628 Movement 674
■ Seed Plants 7 Gymnosperms 7 Ginkgophyta (Ginkgoes) 628 Reproduction 676
■ Seed Plants 7 Gymnosperms 7 Redwood Group (Redwoods, Life Cycles 676
Junipers, Yews) 629
32.4 Key Lineages of Animals: Non-bilaterian Groups 678
■ Seed Plants 7 Gymnosperms 7 Pinophyta (Pines,
■ Porifera (Sponges) 679
Spruces, Firs) 629
■ Cnidaria (Jellyfish, Corals, Anemones, Hydroids) 680
■ Seed Plants 7 Gymnosperms 7 Gnetophyta
■ Ctenophora (Comb Jellies) 681
(Gnetophytes) 630
■ Acoelomorpha (Acoels) 681
■ Seed Plants 7 Anthophyta (Angiosperms) 630
CANADIAN ISSUES 30.1 Canada’s National Tree Seed CHAPTER REVIEW 682
Centre 631
CHAPTER REVIEW 632
33 Protostome Animals 684

33.1 An Overview of Protostome Evolution 685

31 Fungi 635 What Is a Lophotrochozoan? 685
What Is an Ecdysozoan? 686
31.1 Why Do Biologists Study Fungi? 636
Fungi Provide Nutrients for Land Plants 636 33.2 Themes in the Diversification of Protostomes 686
Fungi Speed the Carbon Cycle on Land 636 How Do Body Plans Vary among Phyla? 687
Fungi Have Important Economic Impacts 637 The Water-to-Land Transition 688
Adaptations for Feeding 689
31.2 How Do Biologists Study Fungi? 638
Adaptations for Moving 690
Analyzing Morphological Traits 638

DETAILED CONTENTS xiii

 Adaptations in Reproduction 690
Metamorphosis 691
33.3 Key Lineages: Lophotrochozoans 691
■ Lophotrochozoans 7 Rotifera (Rotifers) 692
■ Lophotrochozoans 7 Platyhelminthes (Flatworms) 692
■ Lophotrochozoans 7 Annelida (Segmented Worms) 693
■ Lophotrochozoans 7 Mollusca 7 Bivalvia (Clams, Mussels,
Scallops, Oysters) 695
■ Lophotrochozoans 7 Mollusca 7 Gastropoda (Snails, Slugs,
Nudibranchs) 696
■ Lophotrochozoans 7 Mollusca 7 Polyplacophora
(Chitons) 697
■ Lophotrochozoans 7 Mollusca 7 Cephalopoda (Nautilus,
Cuttlefish, Squid, Octopuses) 697
33.4 Key Lineages: Ecdysozoans 698
■ Ecdysozoans 7 Nematoda (Roundworms) 699
■ Ecdysozoans 7 Arthropoda 7 Myriapods (Millipedes,
Centipedes) 700
■ Ecdysozoans 7 Arthropoda 7 Insecta (Insects) 700
■ Ecdysozoans 7 Arthropoda 7 Chelicerata (Spiders, Ticks,
Mites, Horseshoe Crabs, Daddy-Long-Legs, Scorpions) 703
■ Ecdysozoans 7 Arthropoda 7 Crustaceans (Shrimp, Lobster,
Crabs, Barnacles, Isopods, Copepods) 704
CANADIAN ISSUES 33.1 The First Census of Marine Life 705
CHAPTER REVIEW 707
34 Deuterostome Animals 709
34.1 What Is an Echinoderm? 710
The Echinoderm Body Plan 710
How Do Echinoderms Feed? 711
Key Lineages 712
■ Echinodermata 7 Asteroidea (Sea Stars) 712
■ Echinodermata 7 Echinoidea (Sea Urchins and
Sand Dollars) 713
34.2 What Is a Chordate? 713
Three “Subphyla” 714
Key Lineages: The Invertebrate Chordates 714
■ Chordata 7 Cephalochordata (Lancelets) 715
■ Chordata 7 Urochordata (Tunicates) 715
34.3 What Is a Vertebrate? 716
An Overview of Vertebrate Evolution 716
Key Innovations 718
Key Lineages 723
■ Chordata 7 Vertebrata 7 Myxinoidea (Hagfish) and
Petromyzontoidea (Lampreys) 724
■ Chordata 7 Vertebrata 7 Chondrichthyes (Sharks, Rays,
Skates) 725
CANADIAN RESEARCH 34.1 The Decline of Large, Predatory
Fishes in the World’s Oceans 726
■ Chordata 7 Vertebrata 7 Actinopterygii (Ray-Finned
Fishes) 727
■ Chordata 7 Vertebrata 7 Actinistia (Coelacanths) and
Dipnoi (Lungfish) 728
■ Chordata 7 Vertebrata 7 Amphibia (Frogs, Salamanders,
Caecilians) 728
■ Chordata 7 Vertebrata 7 Mammalia 7 Monotremata
(Platypuses, Echidnas) 729
■ Chordata 7 Vertebrata 7 Mammalia 7 Marsupiala
(Marsupials) 730
xiv DETAILED CONTENTS
■ Chordata 7 Vertebrata 7 Mammalia 7 Eutheria (Placental
Mammals) 730
■ Chordata 7 Vertebrata 7 Reptilia 7 Lepidosauria (Lizards,
Snakes) 731
■ Chordata 7 Vertebrata 7 Reptilia 7 Testudinia (Turtles) 731
■ Chordata 7 Vertebrata 7 Reptilia 7 Crocodilia (Crocodiles,
Alligators) 732
■ Chordata 7 Vertebrata 7 Reptilia 7 Aves (Birds) 732
34.4 The Primates and Hominins 733
The Primates 733
CANADIAN ISSUES 34.1 Alberta during the Mesozoic Era 733
Fossil Humans 736
The Out-of-Africa Hypothesis 739
CHAPTER REVIEW 740
35 Viruses 742
35.1 Why Do Biologists Study Viruses? 743
Recent Viral Epidemics in Humans 743
Current Viral Pandemics in Humans: HIV 744
35.2 How Do Biologists Study Viruses? 745
Analyzing Morphological Traits 746
Analyzing Variation in Growth Cycles: Replicative and Latent
Growth 746
Analyzing the Phases of the Replicative Cycle 748
35.3 What Themes Occur in the Diversification of
Viruses? 753
The Nature of the Viral Genetic Material 753
Where Did Viruses Come From? 754
CANADIAN ISSUES 35.1 Viruses as Biological Control Agents 755
Emerging Viruses, Emerging Diseases 757
35.4 Key Lineages of Viruses 759
■ Double-Stranded DNA (dsDNA) Viruses 759
■ RNA Reverse-Transcribing Viruses (Retroviruses) 760
■ Double-Stranded RNA (dsRNA) Viruses 760
■ Negative-Sense Single-Stranded RNA ([-]ssRNA)
Viruses 761
■ Positive-Sense Single-Stranded RNA ([+]ssRNA) Viruses 761
CHAPTER REVIEW 762
UNIT 7 HOW PLANTS WORK 764
36 Plant Form and Function 764
36.1 Plant Form: Themes with Many Variations 765
The Importance of Surface Area/Volume Relationships 765
The Root System 766
The Shoot System 768
The Leaf 770
CANADIAN RESEARCH 36.1 Does Phenotypic Plasticity of
Leaves Offer Protection against Herbivore Attack? 772
36.2 Primary Growth Extends the Plant Body 774
How Do Apical Meristems Produce the Primary
Plant Body? 774
How Is the Primary Root System Organized? 775
How Is the Primary Shoot System Organized? 776
36.3 Cells and Tissues of the Primary Plant Body 776
The Dermal Tissue System 778
 The Ground Tissue System 778
The Vascular Tissue System 780
36.4 Secondary Growth Widens Shoots and Roots 782
What Is a Cambium? 782
What Does Vascular Cambium Produce? 782
What Does Cork Cambium Produce? 784
The Structure of a Tree Trunk 784
CHAPTER REVIEW 785
37
Water and Sugar Transport in Plants 788
37.1 Water Potential and Water Movement 788
What Is Water Potential? 789
What Factors Affect Water Potential? 789
Calculating Water Potential 790
Water Potentials in Soils, Plants, and the Atmosphere 791
37.2 How Does Water Move from Roots to Shoots? 792
Movement of Water and Solutes into the Root 793
Water Movement via Root Pressure 794
Water Movement via Capillary Action 794
The Cohesion-Tension Theory 795
37.3 Water Absorption and Water Loss 798
Limiting Water Loss 798
Obtaining Carbon Dioxide under Water Stress 799
CANADIAN RESEARCH 37.1 Ecological Pressures and the
Evolution of Drought Adaptation in Plants 799
37.4 Translocation 800
Tracing Connections between Sources and Sinks 801
The Anatomy of Phloem 801
The Pressure-Flow Hypothesis 802
Phloem Loading 803
Phloem Unloading 806
CHAPTER REVIEW 807
38
Plant Nutrition 809
38.1 Nutritional Requirements of Plants 810
Which Nutrients Are Essential? 810
What Happens When Key Nutrients Are in Short Supply? 812
38.2 Soil: A Dynamic Mixture of Living and Nonliving
Components 813
The Importance of Soil Conservation 814
What Factors Affect Nutrient Availability? 814
38.3 Nutrient Uptake 816
Mechanisms of Nutrient Uptake 816
Mechanisms of Ion Exclusion 818
CANADIAN RESEARCH 38.1 Do Below-Ground Interactions
between Plants and Fungi Influence Above-Ground
Interactions between Plants and Pollinators? 819
38.4 Nitrogen Fixation 822
The Role of Symbiotic Bacteria 822
How Do Nitrogen-Fixing Bacteria Colonize Plant Roots? 823
38.5 Nutritional Adaptations of Plants 824
Epiphytic Plants 824
Parasitic Plants 825
Carnivorous Plants 825
CHAPTER REVIEW 826
39 Plant Sensory Systems, Signals,
and Responses 829
39.1 Information Processing in Plants 830
How Do Cells Receive and Transduce an External Signal? 830
How Are Cell–Cell Signals Transmitted? 830
How Do Cells Respond to Cell–Cell Signals? 831
39.2 Blue Light: The Phototropic Response 832
Phototropins as Blue-Light Receptors 832
Auxin as the Phototropic Hormone 832
39.3 Red and Far-Red Light: Germination and Stem
Elongation 836
The Red/Far-Red “Switch” 836
Phytochromes as Red/Far-Red Receptors 837
How Were Phytochromes Isolated? 837
CANADIAN RESEARCH 39.1 Plant Signalling Networks Help
Influence Proper Growth 838
39.4 Gravity: The Gravitropic Response 839
The Statolith Hypothesis 840
Auxin as the Gravitropic Signal 840
39.5 How Do Plants Respond to Wind and Touch? 841
Changes in Growth Patterns 841
Movement Responses 841
39.6 Youth, Maturity, and Aging: The Growth
Responses 842
Auxin and Apical Dominance 842
Cytokinins and Cell Division 843
Gibberellins and ABA: Growth and Dormancy 844
Brassinosteroids and Body Size 848
Ethylene and Senescence 848
An Overview of Plant Growth Regulators 849
39.7 Pathogens and Herbivores: The Defence
Responses 851
How Do Plants Sense and Respond to Pathogens? 851
How Do Plants Sense and Respond to
Herbivore Attack? 853
CHAPTER REVIEW 856
The Big Picture: How Vascular Plants Work 858
40 Plant Reproduction 860
40.1 An Introduction to Plant Reproduction 861
Sexual Reproduction 861
The Land Plant Life Cycle 861
Asexual Reproduction 863
40.2 Reproductive Structures 863
When Does Flowering Occur? 864
The General Structure of the Flower 865
How Are Female Gametophytes Produced? 867
How Are Male Gametophytes Produced? 867
40.3 Pollination and Fertilization 869
Pollination 869
CANADIAN RESEARCH 40.1 The Mating Strategies of
Flowering Plants 871
Fertilization 873
40.4 The Seed 874
Embryogenesis 874
DETAILED CONTENTS xv
 CANADIAN ISSUES 40.1 What Is the Effect of Agriculture on
Wild Bee Abundance and Crop Pollination? 875
The Role of Drying in Seed Maturation 876
Fruit Development and Seed Dispersal 876
Seed Dormancy 878
Seed Germination 879
CHAPTER REVIEW 880
UNIT 8 HOW ANIMALS WORK 883
41 Animal Form and Function 883
41.1 Form, Function, and Adaptation 884
The Role of Fitness Trade-Offs 884
Adaptation and Acclimatization 884
41.2 Tissues, Organs, and Systems: How Does Structure
Correlate with Function? 886
Structure–Function Relationships at the Molecular and
Cellular Levels 886
Tissues Are Groups of Similar Cells That Function as a Unit 886
Organs and Organ Systems 890
41.3 How Does Body Size Affect Animal Physiology? 891
Surface Area/Volume Relationships: Theory 891
Surface Area/Volume Relationships: Data 892
Adaptations That Increase Surface Area 894
41.4 Homeostasis 894
Homeostasis: General Principles 894
The Role of Regulation and Feedback 895
41.5 How Do Animals Regulate Body Temperature? 896
Mechanisms of Heat Exchange 896
Variation in Thermoregulation 896
Endothermy and Ectothermy: A Closer Look 897
Temperature Homeostasis in Endotherms 897
Countercurrent Heat Exchangers 898
CANADIAN RESEARCH 41.1 Freeze-Tolerant Animals 900
CHAPTER REVIEW 901
42 Water and Electrolyte Balance in
Animals 904
42.1 Osmoregulation and Osmotic Stress 905
What Is Osmotic Stress? 905
Osmotic Stress in Seawater 906
Osmotic Stress in Freshwater 906
Osmotic Stress on Land 906
How Do Cells Move Electrolytes and Water? 907
42.2 Water and Electrolyte Balance in Aquatic
Environments 908
How Do Sharks Excrete Salt? 908
CANADIAN RESEARCH 42.1 The Bamfield Marine Sciences
Centre and Research on Shark Osmoregulation 909
How Do Freshwater Fish Osmoregulate? 910
42.3 Water and Electrolyte Balance in Terrestrial
Insects 911
How Do Insects Minimize Water Loss from the Body Surface 911
Types of Nitrogenous Wastes: Impact on Water Balance 912
Maintaining Homeostasis: The Excretory System 913
xvi DETAILED CONTENTS
42.4 Water and Electrolyte Balance in Terrestrial
Vertebrates 915
The Structure of the Kidney 915
The Function of the Kidney: An Overview 915
Filtration: The Renal Corpuscle 916
Reabsorption: The Proximal Tubule 917
Creating an Osmotic Gradient: The Loop of Henle 918
Regulating Water and Electrolyte Balance: The Distal Tubule
and Collecting Duct 920
CHAPTER REVIEW 922
43 Animal Nutrition 924
43.1 Nutritional Requirements 925
Defining Human Nutritional Requirements 925
Meeting Human Nutritional Requirements 925
CANADIAN ISSUES 43.1 Vitamin D Deficiency
in Canada 926
43.2 Capturing Food: The Structure and Function of
Mouthparts 929
Mouthparts as Adaptations 929
A Case Study: The Cichlid Jaw 929
43.3 How Are Nutrients Digested and Absorbed? 930
An Introduction to the Digestive Tract 930
An Overview of Digestive Processes 932
The Mouth and Esophagus 933
The Stomach 934
The Small Intestine 936
The Cecum and Appendix 939
The Large Intestine 940
43.4 Nutritional Homeostasis—Glucose as
a Case Study 940
The Discovery of Insulin 940
Insulin’s Role in Homeostasis 940
Diabetes Can Take Several Forms 941
The Causes and Treatments of Diabetes 941
CANADIAN RESEARCH 43.1 Causes and Treatments of Diabetes
Mellitus Type 1 942
CHAPTER REVIEW 944
44 Gas Exchange and Circulation 946
44.1 The Respiratory and Circulatory Systems 946
44.2 Air and Water as Respiratory Media 947
How Do Oxygen and Carbon Dioxide Behave in Air? 947
How Do Oxygen and Carbon Dioxide Behave in Water? 948
44.3 Organs of Gas Exchange 949
Physical Parameters: The Law of Diffusion 949
How Do Fish Gills Work? 950
How Do Insect Tracheae Work? 951
How Do Vertebrate Lungs Work? 952
Homeostatic Control of Ventilation 955
44.4 How Are Oxygen and Carbon Dioxide Transported
in Blood? 955
Structure and Function of Hemoglobin 956
CO2 Transport and the Buffering of Blood pH 958
CANADIAN RESEARCH 44.1 Dr. Peter Hochachka and
Physiological Adaptation in Animals 959
44.5 The Circulatory System 961
What Is an Open Circulatory System? 961
What Is a Closed Circulatory System? 962
How Does the Heart Work? 964
CANADIAN ISSUES 44.1 The Risk Factors for Heart Attacks 967
Patterns in Blood Pressure and Blood Flow 968
CHAPTER REVIEW 970
45 Electrical Signals in Animals 973
45.1 Principles of Electrical Signalling 973
Types of Neurons in the Nervous System 974
The Anatomy of a Neuron 974
An Introduction to Membrane Potentials 975
BOX 45.1 Quantitative Methods: Using the Nernst Equation to
Calculate Equilibrium Potentials 976
How Is the Resting Potential Maintained? 976
Using Microelectrodes to Measure Membrane Potentials 977
What Is an Action Potential? 978
45.2 Dissecting the Action Potential 979
Distinct Ion Currents Are Responsible for Depolarization and
Repolarization 979
How Do Voltage-Gated Channels Work? 979
How Is the Action Potential Propagated? 981
45.3 The Synapse 983
Synapse Structure and Neurotransmitter Release 984
What Do Neurotransmitters Do? 985
Postsynaptic Potentials 985
CANADIAN RESEARCH 45.1 David Suzuki and the Discovery
of the Genes Encoding Neuron Proteins 987
45.4 The Vertebrate Nervous System 989
What Does the Peripheral Nervous System Do? 989
Functional Anatomy of the CNS 989
How Does Memory Work? 991
CHAPTER REVIEW 994
46 Animal Sensory Systems and Movement 996
46.1 How Do Sensory Organs Convey Information
to the Brain? 997
Sensory Transduction 997
Transmitting Information to the Brain 998
46.2 Hearing 998
How Do Sensory Cells Respond to Sound Waves and
Other Forms of Pressure? 998
The Mammalian Ear 999
Sensory Worlds: What Do Other Animals Hear? 1001
46.3 Vision 1002
The Insect Eye 1002
The Vertebrate Eye 1002
CANADIAN RESEARCH 46.1 Why Do Wind Farms Kill Bats? 1004
Sensory Worlds: Do Other Animals See Colour? 1008
46.4 Taste and Smell 1008
Taste: Detecting Molecules in the Mouth 1008
Olfaction: Detecting Molecules in the Air 1009
46.5 Movement 1010
Skeletons 1010
Muscle Types 1012
How Do Muscles Contract? 1012
CHAPTER REVIEW 1016
47 Chemical Signals in Animals 1019
47.1 Cell–Cell Signalling: An Overview 1019
Major Categories of Chemical Signals 1020
Hormone Signalling Pathways 1021
What Makes Up the Endocrine System? 1022
Chemical Characteristics of Hormones 1023
How Do Researchers Identify a Hormone? 1024
47.2 What Do Hormones Do? 1025
How Do Hormones Direct Developmental Processes? 1025
How Do Hormones Coordinate Responses to Environmental
Change? 1027
How Are Hormones Involved in Homeostasis? 1028
47.3 How Is the Production of Hormones Regulated? 1030
The Hypothalamus and Pituitary Gland 1031
Control of Adrenaline by Sympathetic Nerves 1033
47.4 How Do Hormones Act on Target Cells? 1033
Steroid Hormones Bind to Intracellular Receptors 1034
Hormones That Bind to Cell-Surface Receptors 1035
CANADIAN ISSUES 47.1 Estrogens in the Environment 1036
Why Do Different Target Cells Respond in Different Ways? 1038
CHAPTER REVIEW 1039
48 Animal Reproduction 1041
48.1 Asexual and Sexual Reproduction 1041
How Does Asexual Reproduction Occur? 1042
Switching Reproductive Modes: A Case History 1042
Mechanisms of Sexual Reproduction: Gametogenesis 1043
48.2 Fertilization and Egg Development 1045
External Fertilization 1045
Internal Fertilization 1045
Unusual Aspects of Mating 1046
Why Do Some Females Lay Eggs While Others Give Birth? 1047
48.3 Reproductive Structures and Their Functions 1048
The Male Reproductive System 1048
The Female Reproductive System 1050
48.4 The Role of Sex Hormones in Mammalian
Reproduction 1051
Which Hormones Control Puberty in Mammals? 1052
Which Hormones Control the Menstrual Cycle in
Mammals? 1053
48.5 Pregnancy and Birth in Mammals 1058
Gestation and Early Development in Marsupials 1058
Major Events during Human Pregnancy 1058
How Does the Mother Nourish the Fetus? 1059
Birth 1061
CANADIAN ISSUES 48.1 Canada’s Assisted Human
Reproduction Act 1062
CHAPTER REVIEW 1063
49
The Immune System in Animals 1065
49.1 Innate Immunity 1066
Barriers to Entry 1066
DETAILED CONTENTS xvii
 The Innate Immune Response 1067
49.2 The Adaptive Immune Response: Recognition 1069
An Introduction to Lymphocytes 1070
The Discovery of B Cells and T Cells 1071
The Clonal-Selection Theory 1071
CANADIAN RESEARCH 49.1 Tak Wah Mak and the Discovery
of the T-Cell Receptor 1073
How Does the Immune System Distinguish Self from
Nonself? 1075
49.3 The Adaptive Immune Response: Activation 1076
T-Cell Activation 1077
B-Cell Activation and Antibody Secretion 1078
49.4 The Adaptive Immune Response: Culmination 1079
How Are Bacteria and Other Foreign Cells Killed? 1080
How Are Viruses Destroyed? 1080
Why Does the Immune System Reject Foreign Tissues
and Organs? 1080
Responding to Future Infections: Immunological Memory 1081
49.5 What Happens When the Immune System Doesn’t
Work Correctly? 1083
Immunodeficiency Diseases 1083
Allergies 1083
CHAPTER REVIEW 1084
The Big Picture: How Humans Work 1086
UNIT 9 ECOLOGY 1088
50
An Introduction to Ecology 1088
50.1 Levels of Ecological Study 1088
Organismal Ecology 1089
Population Ecology 1089
Community Ecology 1089
Ecosystem Ecology 1090
How Do Ecology and Conservation Efforts Interact? 1090
50.2 Types of Aquatic Ecosystems 1090
Nutrient Availability 1090
Water Flow 1091
Water Depth 1091
CANADIAN RESEARCH 50.1 The Future of Canada’s Lakes and
Wetlands 1092
■ Freshwater Environments 7 Lakes and Ponds 1094
■ Freshwater Environments 7 Wetlands 1095
■ Freshwater Environments 7 Streams 1096
■ Freshwater/Marine Environments 7 Estuaries 1097
■ Marine Environments 7 The Ocean 1097
50.3 Types of Terrestrial Ecosystems 1098
■ Terrestrial Biomes 7 Tropical Wet Forest 1100
■ Terrestrial Biomes 7 Subtropical Deserts 1101
■ Terrestrial Biomes 7 Temperate Grasslands 1102
■ Terrestrial Biomes 7 Temperate Forests 1103
■ Terrestrial Biomes 7 Boreal Forests 1104
■ Terrestrial Biomes 7 Arctic Tundra 1105
50.4 The Role of Climate and the Consequences of
Climate Change 1105
Global Patterns in Climate 1106
How Will Global Climate Change Affect Ecosystems? 1108
xviii DETAILED CONTENTS
50.5 Biogeography: Why Are Organisms Found Where
They Are? 1110
Abiotic Factors 1110
CANADIAN ISSUES 50.1 Do Insect Outbreaks Contribute to
Climate Change? 1111
The Role of History 1113
CANADIAN RESEARCH 50.2 Salmon Migration in a Warming
World 1114
Biotic Factors 1115
Biotic and Abiotic Factors Interact 1116
CHAPTER REVIEW 1118
51 Behavioural Ecology 1121
51.1 An Introduction to Behavioural Ecology 1121
Proximate and Ultimate Causation 1122
Conditional Strategies and Decision Making 1122
CANADIAN RESEARCH 51.1 Do Male Redback Spiders Benefit
from Being Eaten by Their Mates? 1123
Five Questions in Behavioural Ecology 1124
51.2 What Should I Eat? 1124
Foraging Alleles in Drosophila melanogaster 1124
Optimal Foraging in White-Fronted Bee-Eaters 1125
51.3 Whom Should I Mate With? 1125
Sexual Activity in Anolis Lizards 1126
How Do Female Barn Swallows Choose Mates? 1127
51.4 Where Should I Live? 1129
How Do Animals Find Their Way on Migration? 1129
Why Do Animals Move with a Change of Seasons? 1130
51.5 How Should I Communicate? 1130
Honeybee Language 1131
Modes of Communication 1132
When Is Communication Honest or Deceitful? 1133
51.6 When Should I Cooperate? 1134
Kin Selection 1134
BOX 51.1 Quantitative Methods: Calculating the
Coefficient of Relatedness 1136
Reciprocal Altruism 1136
An Extreme Case: Abuse of Non-Kin in Humans 1137
CHAPTER REVIEW 1138
52 Population Ecology 1141
52.1 Demography 1141
Life Tables 1142
CANADIAN RESEARCH 52.1 Tyrannosaur Life Tables 1144
The Role of Life History 1145
BOX 52.1 Quantitative Methods: Using Life Tables to Calculate
Population Growth Rates 1146
52.2 Population Growth 1147
Quantifying the Growth Rate 1147
Exponential Growth 1148
Logistic Growth 1148
BOX 52.2 Quantitative Methods: Developing and Applying
Population Growth Equations 1149
What Limits Growth Rates and Population Sizes? 1151
52.3 Population Dynamics 1152
 How Do Metapopulations Change through Time? 1152 Positive and Negative Feedback 1213
Why Do Some Populations Cycle? 1153 Impact on Organisms 1213
BOX 52.3 Quantitative Methods: Mark–Recapture Productivity Changes 1214
Studies 1154 CHAPTER REVIEW 1216
How Does Age Structure Affect Population Growth? 1156
CANADIAN RESEARCH 52.2 The Snowshoe Hare–Lynx Cycle
What Questions Remain? 1156
Analyzing Change in the Growth Rate of Human Populations 1159
55 Biodiversity and Conservation Biology 1219

52.4 How Can Population Ecology Help Endangered 55.1 What Is Biodiversity? 1220
Biodiversity Can Be Measured and Analyzed at Several
Species? 1161
Levels 1220
Using Life Table Data 1161
How Many Species Are Living Today? 1221
Preserving Metapopulations 1163
BOX 55.1 Quantitative Methods: Extrapolation
CHAPTER REVIEW 1163 Techniques 1222
55.2 Where Is Biodiversity Highest? 1223
53 Community Ecology 1166 Hotspots of Biodiversity and Conservation 1223

53.1 Species Interactions 1166 55.3 Threats to Biodiversity 1224

Three Themes 1167 Changes in the Nature of the Problem 1224
Competition 1167 CANADIAN ISSUES 55.1 SARA—Canada’s Species at Risk
Consumption 1171 Act 1224
Mutualism 1176 How Can Biologists Predict Future Extinction Rates? 1229
CANADIAN ISSUES 55.2 Polar Bears in a Warming Arctic 1230
53.2 Community Structure 1178 BOX 55.2 Quantitative Methods: Population Viability
How Predictable Are Communities? 1178 Analysis 1232
How Do Keystone Species Structure Communities? 1180
55.4 Why Is Biodiversity Important? 1233
53.3 Community Dynamics 1181 Economic Benefits of Biodiversity 1233
Disturbance and Change in Ecological Communities 1181 Biological Benefits of Biodiversity 1234
Succession: The Development of Communities after An Ethical Dimension? 1236
Disturbance 1182
55.5 Preserving Biodiversity 1236
53.4 Species Richness in Ecological Communities 1185 Designing Effective Protected Areas 1237
Predicting Species Richness: The Theory of Island Beyond Protected Areas: A Comprehensive Approach 1237
Biogeography 1185 CANADIAN RESEARCH 55.1 A Solution to the Problem of
Global Patterns in Species Richness 1186 Habitat Fragmentation 1238
BOX 53.1 Quantitative Methods: Measuring Species
Diversity 1187 CHAPTER REVIEW 1242
CANADIAN RESEARCH 53.1 Why Is Biodiversity Higher in The Big Picture: Ecology 1244
the Tropics? 1189
CHAPTER REVIEW 1190 APPENDIX A: Answers A:1
APPENDIX B: BioSkills B:1

54 Ecosystems 1193
1 The Metric System B:1
2 Reading Graphs B:2
54.1 How Does Energy Flow through Ecosystems? 1194
3 Reading a Phylogenetic Tree B:4
Why Is NPP So Important? 1194
Solar Power: Transforming Incoming Energy to 4 Some Common Latin and Greek Roots Used in Biology B:6
Biomass 1194 5 Using Statistical Tests and Interpreting Standard Error Bars B:6
Trophic Structure 1195 6 Reading Chemical Structures B:7
CANADIAN ISSUES 54.1 The Ecological Lessons of the
7 Using Logarithms B:9
Balsam Fir Food Web 1196
Energy Transfer between Trophic Levels 1197 8 Making Concept Maps B:9
Trophic Cascades and Top-Down Control 1198 9 Separating and Visualizing Molecules B:10
Biomagnification 1199 10 Biological Imaging: Microscopy and X-Ray Crystallography B:13
Global Patterns in Productivity 1201
What Limits Productivity? 1202 11 Separating Cell Components by Centrifugation B:16
12 Cell and Tissue Culture Methods B:17
54.2 How Do Nutrients Cycle through Ecosystems? 1204
Nutrient Cycling within Ecosystems 1204 13 Combining Probabilities B:18
CANADIAN RESEARCH 54.1 Can Predators Increase Nutrient 14 Model Organisms B:19
Cycling? 1205 Glossary G:1
Global Biogeochemical Cycles 1208
Credits C:1
54.3 Global Warming 1211
Understanding the Problem 1211
Index I:1

DETAILED CONTENTS xix

About the Authors

SCOTT FREEMAN received his Ph.D. in Zoology from the University of Washington and was sub-
sequently awarded an Alfred P. Sloan Postdoctoral Fellowship in Molecular Evolution at Princeton
University. His current research focuses on the scholarship of teaching and learning—specifically
(1) how active learning and peer teaching techniques increase student learning and improve perfor-
mance in introductory biology and (2) how the levels of exam questions vary among introductory
biology courses, standardized postgraduate entrance exams, and professional school courses. He has
also done research in evolutionary biology on topics ranging from nest parasitism to the molecular
systematics of the blackbird family. Scott teaches introductory biology for majors at the University of
Washington and is coauthor, with Jon Herron, of the standard-setting undergraduate text Evolution-
ary Analysis.

MIKE HARRINGTON completed his B.Sc. and Ph.D. in the Zoology Department of the University
of British Columbia. His graduate work on Drosophila chromatin structure combined classical and
molecular genetics. He is presently a Faculty Lecturer in the Biological Sciences Department at the
University of Alberta. He teaches cell biology at the first- and second-year levels and genetics at the
second-, third-, and fourth-year levels. His teaching goals are (1) to find ways to incorporate current
scientific research into introductory courses, (2) to develop new ways to expand a course’s bound-
aries with online material, and (3) to use clicker classroom response systems to teach content with
questions.

JOAN SHARP received her B.A. and B.Sc. from McGill University and her M.Sc. from the University of
British Columbia. She is a Senior Lecturer at Simon Fraser University, where she teaches Introduction to
Biology, General Biology, Ecology, and Vertebrate and Invertebrate Biology. Her teaching and research
interests include a number of areas: (1) Prior or newly acquired misconceptions interfere with student
success in building meaningful biological understanding. It is important to understand common miscon-
ceptions and to develop activities that allow students to address and correct their misconceptions. Concept
inventories can be used to measure students’ learning gains to assess the success of teaching strategies tar-
geting student misconceptions. (2) Students’ written work can serve as a starting point to address areas of
misunderstanding and to help students refine and express biological ideas. (3) Case studies engage students
with key concepts by using meaningful real-world scenarios. The use of clickers allows the implementation
of case studies in large lecture courses, facilitating small group discussion and increasing student learning.

Illustrator
KIM QUILLIN combines expertise in biology and information design to create lucid visual repre-
sentations of biological principles. She received her B.A. in Biology at Oberlin College and her Ph.D.
in Integrative Biology from the University of California, Berkeley (as a National Science Foundation
Graduate Fellow), and taught undergraduate biology at both schools. Students and instructors alike
have praised Kim’s illustration programs for Biological Science, as well as Biology: A Guide to the Natu-
ral World by David Krogh and Biology: Science for Life by Colleen Belk and Virginia Borden, for their
success in the visual communication of biology. Kim is a Lecturer in the Department of Biological Sci-
ences at Salisbury University.

xx
Preface to Instructors
T ⦁ Canadian Content We have updated and expanded the
his book is for instructors who want to help their stu-
dents learn how to think like a biologist. Our students
need to learn the language of biology and understand
fundamental concepts, but they also need to apply these con-
cepts to new situations, analyze experimental design, synthesize
results, and evaluate hypotheses and data.
We wrote this book for instructors who embrace this chal-
lenge—who want to help their students learn how to think like a
biologist. The essence of higher education is to promote higher-
order thinking. Our job is to help students understand biological
science at all six levels of Bloom’s taxonomy of learning.
Analyze Evaluate Synthesize
Apply
Explain
Remember
Bloom’s Taxonomy. An annotated version of this graphic can be
found in “Preface to Students: How to Use This Book” at the front of
this book.
The Evolution of a Textbook
Evolution can be extremely fast in populations with short gen-
eration times and high mutation rates. Biology textbooks are no
exception. Generation times have to be short because the pace of
research in biology and student learning is so fast. This book, in
particular, evolves quickly because it incorporates so many new
ideas with each edition. Some of these “alleles” are novel muta-
tions, but most arrive via lateral transfer—from advisors, review-
ers, friends, students, and the literature.
What’s New in This Edition
This revision was about making the book a better teaching and
learning tool. To help students manage the mass of information
and ideas that is contemporary biology, we broke long para-
graphs into shorter paragraphs, made liberal use of numbered
lists and bulleted lists to “chunk” information and ideas, and
broke out dozens of new sections and subsections.
In addition, we came up with a long list of new or expanded
features.
Canadian content throughout the book. Each chapter now
has at least one Canadian Research or Canadian Issues box.
We have chosen examples that both illustrate one of the main
concepts in the chapter and highlight the diversity of science
being done in Canadian universities, colleges, and other or-
ganizations. These boxes now end with a “Think About It”
question to allow students to test their understanding of the
material.
⦁ The Big Picture These new two-page spreads are meant to
help students see the forest for the trees. They are concept
maps that focus on particularly critical areas—Energy, Ge-
netic Information, Evolution, Macromolecules, Ecology, and
How Multicellular Organisms Work. Each synthesizes con-
tent and concepts from an array of chapters and includes ex-
ercises for students to complete. You’ll recognize these pages
readily—their edges are coloured black (for example, see The
Big Picture: Macromolecules on pages 110–111). In addition,
the book’s MasteringBiology® website has 10 new concept
map activities based on Big Picture content that will allow
you to explore the concepts and their connections with your
students during lectures.
⦁ BioSkills Students completing introductory biology need to
have acquired skills—the ability to read a graph, interpret an
equation, understand the bands on a gel. The previous edition
of Biological Science contained a series of appendices focused
on key skills for introductory biology students. Instructors and
students found them extraordinarily helpful. New in this edi-
tion are BioSkills on using the metric system, common Latin
and Greek roots, techniques for isolating and visualizing cell
components, cell and tissue culture methods, and model or-
ganisms. BioSkills are located in Appendix B.
⦁ Answer Key New to the Second Canadian Edition are sug-
gested answers to all questions and exercises in the textbook.
Students asked us to make this important change between
editions to make the book a more complete study tool. The
answer key will allow them to self-check their understanding
while reading and when reviewing for exams. Answers are in
Appendix A.
⦁ Experiment Boxes This text’s hallmark has always been its
emphasis on experimental evidence—on teaching how we
know what we know. In the previous edition, key experi-
ments were converted to a boxed format so students could
easily navigate through the logic of the question, hypothesis,
and test. In this edition, we added a new question to every
experiment box to encourage students to analyze some aspect
of the experiment’s design.
xxi
Another random document with
no related content on Scribd:
 The plant called “life everlasting” is one which grows in dry, open,
sunny places. It clothes its leaves with silky hairs, and so prevents
them from throwing off too quickly the small amount of water its roots
are able to provide. Without this silky coat, the sun would suck its
leaves quite dry of water.
Sometimes a leaf has only a few of the little leaf mouths through
which most of the water passes. As these mouths are wide open
only in the sunlight, and as often the rest of the leaf is covered with a
thick skin which prevents the water from slipping away (as a little of it
nearly always does) through the cell walls, such a leaf will hold its
water supply and keep fresh for a long time. Such leaves as these
we find on what we call “evergreen” plants. The pines and hemlocks
which light up the woods all winter have these thick-skinned, few-
mouthed leaves, which throw off so little water that even when the
ground is frozen hard, and gives no drinking water to the roots, they
are able to keep fresh by the careful way in which each one hoards
its own little supply.
 WOOLLY AND “DUSTY” LEAVES

C URIOUSLY enough, some plants put on a hairy coat for just the
opposite reason from the one which makes life everlasting
clothe itself in that fashion. Life everlasting fears lest its leaves throw
off their water, or perspire too quickly.
Down by the stream that runs through the meadow grow great
clusters of the pink-flowered steeple bush. If you look at the lower
sides of the leaves of the steeple bush, you see that they are very
woolly. As this wool is not between the sun and the leaf blade, it
cannot be meant to protect the leaves from the heat of the sun; and
indeed in this wet meadow, close to the river, never mind how quickly
the leaves throw off their water, the roots can have no difficulty in
finding close by more than enough to make good the loss. No, the
fact is that these leaves need to throw off water very freely indeed to
make room for the ever-fresh supply that is pushing up the stem, and
their woolly covering is intended to help them do this very thing. Its
object is to aid perspiration. In swampy places the moisture rises
every night from the wet ground, and settles on the plants about. The
little mouths on the under surfaces of the leaves of the steeple bush
would soon be clogged with the moisture rising from below, if they
were not protected in some way; and if they became so clogged,
they could not throw off the water with which the whole plant is
charged. Thus, by having this thick coat of hair, the water that
otherwise would cling to the outer surface of the leaf blade is kept at
a distance from the little mouths, and these are not interrupted in the
performance of a duty so necessary to the health of the plant.
This same habit of coating its lower leaf surfaces with hair, you
notice in the speckled or swamp alder, a shrub which grows also in
wet places, and therefore runs the same risk of having its leaf
mouths clogged with water.
So when you see only the upper surface of a leaf covered with
hair, you can guess that the object of the plant is to prevent too much
perspiration; but when you see only its lower side clothed in this
same way, you can guess that the plant fears too little perspiration.
Sometimes you find a plant with leaves which have a coating of
what looks almost like dust on one or both of their surfaces. This
dust we call “bloom.” We see it in apples and grapes, as well as on
leaves. It is made up of a waxy material which is put forth by the
plant just as it puts forth hair. This bloom the plant uses also as a
help to free perspiration. By thus clothing its leaves it shields the little
mouths from water clogging; and so you can be sure that the little
mouths have not been filled with water, and thus prevented from
doing their work.
The cabbage leaf has mouths on both of its surfaces, and so both
sides are covered with this protecting bloom. If you dip a cabbage
leaf in water and then shake it, the drops roll off and leave it quite
dry.
 PRICKLES AND POISON

L EAVES need to protect themselves from other enemies than too

great heat and too much water.
We found that the prickly armor of the thistle enabled it to live in
pastures where the cattle had killed most of the other plants.
Many animals like to eat green leaves, so we are not surprised to
find that plants invent different ways of protecting themselves.
One look at the leaf of the thistle is enough to persuade us that it
would not be very good eating.
The red-berried holly, with which we decorate our churches and
houses at Christmas time, is another plant with prickly leaves.
 Some plants cover their leaves with bristles, which the cattle dread
almost as much as the stout prickles.
As we read before, the mullein defends its leaves by a fuzzy coat
of hair. Such an armor as this is less warlike than that of the thistle,
but quite as effective.
Other plants fill their leaves with juices which are either poisonous
or unpleasant.
It seems as if animals guessed the presence of these unfriendly
juices by the plant’s smell, for they will munch the different growing
things all about such a plant as this, and leave the harmful leaves
severely alone.
The nettles cover their leaves with stinging hairs. These stiff hairs
break off when handled, burying themselves in the flesh, and
sending out a burning acid that punishes severely the meddler, man
or beast, as it may happen to be.
By this time I think you realize that leaves are well worth noticing.
And when you have looked at a leaf so fully as to be able to carry in
your mind its outline, I hope you will then discover whether it wears a
coat of hair, or a dusty bloom, or a prickly armor, or a thick,
evergreen skin, and that you will decide what enemies it is trying to
escape.
 SOME CRUEL TRAPS

H AVE you ever seen a leaf like the one in this picture (Fig. 152)?
It is shaped something like a pitcher; and the plant on which it
grows has been named the “pitcher plant.”

Fig. 152

The pitcher plant lives in low, wet place, such as the shaded
swamp, or the marsh down by the lake.
On account of its curious leaves it is brought to the cities, and is
sold on the street corners or at the florists’.
In June comes the great flower of the pitcher plant. Sometimes
this is a dull red; again it is a delicate pink or perhaps a light green;
and it has faint, pleasant fragrance.
Next June I hope that some of you children will find these beautiful
flowers and these curious leaves.
Why should a leaf be shaped like a pitcher, do you suppose?
These leaves are not only pitcher-like in shape, but also in their
way of holding water; for if you succeed in discovering a settlement
of pitcher plants, you will find that nearly every pitcher is partly filled
with rain water. Usually this water is far from clear. It appears to hold
the remains of drowned insects; and sometimes the odor arising
from a collection of these pitcher plants is not exactly pleasant.
 Perhaps you wonder how it happens that dead insects are found
in every one of these pitchers; and possibly you will be surprised to
learn that apparently these curious leaves are built for the express
purpose of capturing insects.
It is easy to understand that these odd leaves are not so well fitted
as more simple ones to cook the plant’s food in the sun, or to take
carbon from the air; but if they are unfitted to provide and prepare
ordinary food, possibly they are designed to secure food that is
extraordinary.
It seems likely that the pitcher plant is not content to live, like other
plants, upon the simple food that is taken in from the earth and from
the air. We are led to believe that it wishes something more
substantial; that it needs a meat diet; and that to secure this, it
teaches its leaves to capture flies and insects in order that it may
suck in their juices.
These leaves are veined in a curious and striking fashion. The
bright-colored veins may convince the insects of the presence of the
sweet nectar in which they delight. At all events, in some way they
are tempted to enter the hollow leaf; and, once they have crawled or
tumbled down its slippery inner surface, they find it impossible to
crawl back again, owing to the stiff hairs, pointing downward, which
line the upper part of the pitcher.
Even if they have wings, it is difficult for them to fly upward in so
straight a line as would be necessary to effect their escape.
When tired out in their efforts to get out of this cruel trap, they fall
into the water at the bottom of the pitcher, and are drowned. Their
bodies decay and dissolve; and it is thought that this solution is
taken in by the leaf, and turned over to the plant as food.
It is just the old, sad story of the spider and the fly, you see, only
now it is the pitcher and the fly.
But be sure to examine one of these pitchers if you possibly can,
and then you will understand better how the whole thing is managed.
The leaf in this picture (Fig. 153), for it is a leaf, you cannot find in
our North American swamps. It grows on a plant called Nepenthes, a
plant which lives in hot countries far from the United States.

Fig. 153

The leaf in the picture is full grown, and all ready for its work of
trapping animals. Before it was old enough to do this, the lid which is
now lifted was laid nicely across the opening to the pocket, and so
prevented any unseasonable visits.
Sometimes these pockets are so large as to be able to hold and to
hide from sight a pigeon. They are gayly colored, and the rim around
their border is covered with a sugary, tempting juice. So you can
guess that the animals in search of nectar are not slow in accepting
the invitation offered by color and sweets, and that some of these
are imprudent enough to venture across the sticky edge. In this
event they are pretty sure to lose their footing on the slippery inner
surface of the pocket, and to fall into the watery liquid with which it is
filled. Even if they do not slip immediately, their efforts to crawl back
over the rim are defeated by a row of teeth such as you see in the
picture.
The liquid at the bottom of the leaf is not rain water, as in the
pitcher plant. It is given out by the leaf itself; and it contains an acid
which dissolves the animals’ bodies, so that their more nourishing
parts can easily be taken in by certain little cells which line the lower
part of the pocket, and which have been brought up to this work.
 Fig. 154

The next picture (Fig. 154) shows you a water plant. It is called the
“bladderwort,” because of the little bags or bladders which you see
growing from the branches under water. The little bladders are traps
set for water animals, which swim into them in their wish, perhaps, to
escape some enemy. But they are quite unable to swim out again;
for the door into the bladder is transparent, and looks like an open
entrance with a nice hiding place beyond. It opens easily from the
outside, but is so arranged that it will not open from within. So when
the poor little animal hurriedly swims into what seems to it a cozy
resting spot, and draws a long breath of relief at getting safe inside, it
is hopelessly caught, and must slowly starve to death, for there is no
chance of escape. It may live for nearly a week in this prison; but at
last it dies. Its body decays, and is taken in as food by the cells set
apart for that purpose.
 Strangely enough, though we ourselves do not hesitate to kill
animals for food, and sometimes, I am sorry to say, for nothing but
amusement, we give a little shiver of disgust when we find these
plants doing the same thing. Some lines that came out in one of the
magazines a few years ago express this feeling:—

“What’s this I hear

About the new Carnivora?
Can little plants
Eat bugs and ants
And gnats and flies?
A sort of retrograding!
Surely the fare
Of flowers is air,
Or sunshine sweet.
They shouldn’t eat
Or do aught so degrading.”
 MORE CRUEL TRAPS

T HE plants about which we read in the last chapter do not take

any active part in capturing insects. They set their traps, and
then keep quiet. But there are plants which lay hold of their poor
victims, and crush the life out of them in a way that seems almost
uncanny.
This leaf (Fig. 155) belongs to a plant which lives in North
Carolina. It is called Venus’s flytrap.

Fig. 155

You see that the upper, rounded part of the leaf is divided by a rib
into two halves. From the edges of these rounded halves run out a
number of long, sharp teeth; and three stout bristles stand out from
the central part of each half. When an insect alights upon this
horrible leaf, the two halves come suddenly together, and the teeth
which fringe their edges are locked into one another like the fingers
of clasped hands.
The poor body that is caught in this cruel trap is crushed to pieces.
Certain cells in the leaf then send out an acid in which it is dissolved,
and other cells swallow the solution.
After this performance the leaf remains closed for from one to
three weeks. When finally it reopens, the insect’s body has
disappeared, and the trap is set and ready for another victim.
 The next picture (Fig. 156) shows you a little plant which is very
common in our swamps,—so common that some of you ought to find
it without difficulty next summer, and try upon it some experiments of
your own.

Fig. 156

It is called the “sundew.” This name has been given to it because

in the sunshine its leaves look as though wet with dew. But the pretty
drops which sparkle like dew do not seem so innocent when you
know their object. You feel that they are no more pleasing than is the
bit of cheese in the mouse trap.
When you see this plant growing in the swamp among the
cranberry vines and the pink orchids, you admire its little white
flowers, and its round red-haired leaves, and think it a pretty,
harmless thing. But bend down and pluck it up, root and all, out of
the wet, black earth. Carry it home with you, and, if you have a
magnifying glass, examine one of its leaves.
The picture (Fig. 157) shows you a leaf much larger than it is in
life. The red hairs look like pins stuck in a cushion, and the head of
each pin glistens with the drop that looks like dew.
 Fig. 157

But the ants and flies do not take these drops for dew. They
believe them to be the sweet nectar for which they long, and they
climb or light upon the leaves in this belief.
And then what happens?
The next two pictures will show you (Figs. 158, 159).

Fig. 158

Fig. 159
 The red hairs close slowly but surely over the insect whose legs
are already caught and held fast by the sticky drops it mistook for
nectar, and they hold it imprisoned till it dies and its juices are
sucked in by the leaf.
I should like you to satisfy yourselves that these leaves act in the
way I have described. But a bit of fresh meat will excite the red hairs
to do their work quite as well as an insect, and I hope in your
experiments you will be merciful as well as inquiring.
So you see that the little sundew is quite as cruel in its way as the
other insect-eating plants. But its gentle looks seem to have
deceived the poet Swinburne, who wonders how and what these
little plants feel, whether like ourselves they love life and air and
sunshine.

“A little marsh-plant, yellow-green

And tipped at lip with tender red,
Tread close, and either way you tread,
Some faint, black water jets between
Lest you should bruise its curious head.

“You call it sundew; how it grows,

If with its color it have breath,
If life taste sweet to it, if death
Pain its soft petal, no man knows,
Man has no sight or sense that saith.”
 THE FALL OF THE LEAF

Y OU know that in autumn nearly all the leaves fall from the trees.
To be sure, a few trees (such as the pines and hemlocks) and
some plants (such as the laurel and wintergreen and partridge vine)
do hold fast their leaves all winter; but these are so few as compared
with the many plants which lose their leaves, that they hardly count.
Perhaps you never stopped to wonder why most plants get rid of
their leaves before winter comes on; but you feel pretty sure now
that there is some good reason for a habit that is adopted by nearly
all the plants that live in this part of the country.
When we were talking about the way in which leaves defend
themselves from different dangers, we found that evergreen leaves,
the leaves which hold fast to the tree and keep fresh all winter,
manage to keep their water safe inside their cells by wearing a very
thick skin, and by not having too many little leaf mouths. For when a
leaf has a thin skin and a great many mouths, its water leaks away
very quickly. And if many such leaves should remain upon a plant
into the winter, might it not happen that they would let off all its water
at a time when its roots could not find any more in the frozen
ground? And thus might not the leaves kill the plant by draining it
quite dry?
So you can see why it is well for most plants to shed their leaves
before winter comes on and the root’s drinking water is turned into
ice.
But when a plant is about to shed its leaves, it takes care not to
waste the precious food which they hold. This food it draws back into
its stem and roots, laying it away in safe places beneath the buds
which are to burst another year.
It is this action on the part of the plant which changes the color of
the leaves every fall. That material which makes them green is
broken up, and part of it is taken away. That which is left is usually
yellow or brown or reddish, and gives the leaves the beautiful colors
we see in our October woods.
So whenever you see the woods changing color, losing their fresh
green and turning red and yellow, you can be sure that the trees
have begun to prepare for winter. You know that they are stowing
away their food in warmer, safer places than can be supplied by the
delicate leaves. And when all the food has been drawn out of the
leaves, and packed away in the right spots, then the plant finishes a
piece of work it began some time before. This piece of work is the
building-up of a row of little cells just where the leafstalk joins the
stem or branch. When this row is complete, it acts almost like a
knife, loosening the stalk from the stem.
Then the leaf’s life work is over; and with the first breeze, the
empty shell, which is all that is left, breaks away from the parent
plant, and drifts earthward.
 Part VI—Flowers

THE BUILDING PLAN OF THE CHERRY

BLOSSOM

O NE day your teacher brought to school a branch broken from

the cherry tree. This she placed in water, standing the tumbler
on the sunny window sill; and now its buds have burst into a glory of
white blossoms.
To-day I want you to study the flower of the cherry; for if you know
all about this flower, which is put together in a rather simple way, one
that is easy to study, it will not be so difficult for you to understand
other, less simple flowers.
You may wonder why I do not wait till the cherry tree outside is in
blossom; but if we waited till May, other flowers, which are not built
on quite so simple a plan, would have come and gone, and you
would not have been able to understand them so well as if you had
first studied the simple make-up of the cherry blossom.
Last fall we learned a little about this flower, but we had only its
picture to help us in our work: so I think it well to begin all over again.
In looking at the cherry blossom (Fig. 160), we should first notice
the green cup which holds the rest of the flower.
This cup is divided into five green leaves.
During the babyhood of the flower, when it was quite too young to
face the cold, windy world, these green leaves were folded together
so as to shut away from all harm its more delicate parts.
 Above this green cup we see a circle made up of five white leaves.
These pretty leaves are spread outward as if they were quite proud
of themselves, and eager to attract attention.
And that is just what they are trying to do; for the cherry blossom is
not wise enough to know that here in the schoolroom there are no
bee visitors to bring it yellow dust, and to help it grow into a cherry.
These leaves are the little handkerchiefs which the cherry tree, just
like the apple tree we read about long ago, uses in signaling the
bees.

Fig. 160

Within the circle of white leaves you see a quantity of what we

named “pins with dust boxes.” You remember that these dust boxes
hold the powdery material which is as wonderful as Cinderella’s fairy
godmother in its power to do strange and surprising things.
And in the very center of the flower you find a single “pin,” as we
called it, with a flat top which is not a dust box.
But you remember that at the foot of this pin is another sort of box,
a seedbox (Fig. 161).
And you have not forgotten that it is on the flat top of this pin that
the bee brushes the yellow dust which gives new life to the seed
below, and turns the little case of the seedbox into the juicy cherry.
So now what do we find in the cherry blossom? We find
1. A green cup cut above into separate leaves.
 2. A circle of white leaves.
3. Some pins with dust boxes.
4. One pin with a seedbox.

Fig. 161

Here you have the plan on which the cherry blossom is built (for
flowers, like houses, are built on different plans), and the building
plan of the cherry blossom is one of the simplest of all. So it is well,
before studying more difficult flowers, to feel quite at home with this
one. And you must try to remember first what work each part of the
flower is expected to perform; for you see that the leaves of the
green cup, the pretty white leaves, the pins with dust boxes, and the
pin with a seedbox, have each and all their special task,—a task
which they alone are able to accomplish.
Now, in talking about a flower it is troublesome to use a great
many words where one would answer every purpose, so I will tell
you what these different parts of the flower have been named; and
by taking a little trouble to remember these names, we can save a
good deal of time.
The green cup is called the “calyx.”
“Calyx” is a Greek word meaning “cup.”
The circle of leaves which grow above the green cup or calyx is
called the “corolla.”
“Corolla” comes from a word which means “crown.”
The pins with dust boxes are called “stamens.”
“Stamen” comes from a word meaning “to stand.”
The pin with a seedbox is called the “pistil.”
 “Pistil” is another form of the word “pestle.” A pestle is an
instrument used in the drug shops for pounding and mixing
medicines. You might ask to look at one the next time you are sent to
the drug shop, and then you can see for yourselves if it really looks
like its namesake, the pin with a seedbox.
Perhaps at first you may find it a little difficult to bear in mind these
four words with their meanings; but soon they will become quite
easy, and will save you much trouble.
Green cup,—calyx.
Circle of flower leaves,—corolla.
Pins with dust boxes,—stamens.
Pin with seedbox,—pistil.
If you remember the names of these four parts of the flower, how
the different parts look, and what they do, you will have made a good
start in the study of flowers.