BIOLOGICAL SCIENCE

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 Red-tailed Hawk, Buteo jamaicensis
The red-tailed hawk searches for a wide variety
of small prey, capturing them by swooping from
the air or diving from a stationary perch. Buteos
like the red-tailed hawk are adapted for predation,
with broad wings and tail for soaring flight, curved
talons for grasping prey, large forward-rotated eyes
for acute long-distance vision, and a sharp tearing
beak. The red-tailed hawk’s colouration, which is
darker above and paler underneath, camouflages
it from below. The red-tailed hawk’s opportunistic
hunting skills have helped it to be the most widely
distributed hawk in North America.

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 BIOLOGICAL SCIENCE
Second Canadian Edition

S C O T T F RE E M AN
University of Washington

M IK E H ARRIN G T O N
University of Alberta

J O AN S H ARP
Simon Fraser University

Toronto

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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

Vice‐President, Editorial Director: Gary Bennett

Senior Acquisitions Editor: Lisa Rahn
Senior Marketing Manager: Kim Ukrainec
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Cover Designer: Anthony Leung
Interior Designer: Anthony Leung
Cover Image: Getty Images/Deborah Harrison

Credits and acknowledgments for material borrowed from other sources and reproduced, with permission, in this
textbook appear on the appropriate page within the text or beginning on page C:1 of the backmatter.
Original edition published by Pearson Education, Inc., Upper Saddle River, New Jersey, USA. Copyright © 2011
Pearson Education, Inc. This edition is authorized for sale only in Canada.
If you purchased this book outside the United States or Canada, you should be aware that it has been imported without
the approval of the publisher or the author.
Copyright © 2014 Pearson Canada Inc. All rights reserved. Manufactured in the United States of America. This publication
is protected by copyright and permission should be obtained from the publisher prior to any prohibited reproduction,
storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording,
or likewise. To obtain permission(s) to use material from this work, please submit a written request to Pearson Canada Inc.,
Permissions Department, 26 Prince Andrew Place, Don Mills, Ontario, M3C 2T8, or fax your request to 416‐447‐3126, or
submit a request to Permissions Requests at www.pearsoncanada.ca.
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

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 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

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 Detailed Contents

About the Authors xx CANADIAN RESEARCH 2.1 The Carbon-Rich Tagish Lake
Preface to Instructors xxi Meteorite 37
CHAPTER REVIEW 38
Preface to Students: How to Use This Book xxxiv

1 Biology and the Tree of Life 1 3 Protein Structure and Function 40

1.1 What Does It Mean to Say That Something Is Alive? 1 3.1 Early Origin-of-Life Experiments 41
1.2 The Cell Theory 2 3.2 Amino Acids and Polymerization 42
Are All Organisms Made of Cells? 2 The Structure of Amino Acids 42
Where Do Cells Come From? 2 The Nature of Side Chains 42
1.3 The Theory of Evolution by Natural Selection 4 How Do Amino Acids Link to Form Proteins? 44
What Is Evolution? 4 3.3 Proteins Are the Most Versatile Large Molecules in
What Is Natural Selection? 4 Cells 46
CANADIAN RESEARCH 1.1 Artificial Selection on Bighorn Sheep CANADIAN RESEARCH 3.1 Designing New Proteins 47
in Alberta 6
3.4 What Do Proteins Look Like? 47
1.4 The Tree of Life 6 Primary Structure 48
Using Molecules to Understand the Tree of Life 7 Secondary Structure 48
How Should We Name Branches on the Tree of Life? 9 Tertiary Structure 49
1.5 Doing Biology 9 Quaternary Structure 50
Why Do Giraffes Have Long Necks? An Introduction to CANADIAN RESEARCH 3.2 Spider Silk Proteins 52
Hypothesis Testing 9 Folding and Function 52
How Do Ants Navigate? An Introduction to Experimental Design 11 3.5 Enzymes: An Introduction to Catalysis 54
CHAPTER REVIEW 13 Enzymes Help Reactions Clear Two Hurdles 55
How Do Enzymes Work? 56
Was the First Living Entity a Protein Catalyst? 59
UNIT 1 THE MOLECULES OF LIFE 16
CHAPTER REVIEW 60

2 Water and Carbon: The Chemical

Basis of Life 16 4 Nucleic Acids and the RNA World 62

4.1 What Is a Nucleic Acid? 62

2.1 Atoms, Ions, and Molecules: The Building Blocks of
Could Chemical Evolution Result in the Production of
Chemical Evolution 17 Nucleotides? 63
Basic Atomic Structure 17
How Do Nucleotides Polymerize to Form Nucleic Acids? 64
How Does Covalent Bonding Hold Molecules Together? 18
Ionic Bonding, Ions, and the Electron-Sharing Continuum 19 4.2 DNA Structure and Function 65
Some Simple Molecules Formed from C, H, N, and O 20 What Is the Nature of DNA’s Secondary Structure? 66
The Geometry of Simple Molecules 21 DNA Functions as an Information-Containing Molecule 67
Representing Molecules 21 Is DNA a Catalytic Molecule? 69
Basic Concepts in Chemical Reactions 22 4.3 RNA Structure and Function 69
2.2 The Early Oceans and the Properties of Water 23 Structurally, RNA Differs from DNA 69
Why Is Water Such an Efficient Solvent? 23 RNA’s Structure Makes It an Extraordinarily Versatile Molecule 70
How Does Water’s Structure Correlate with Its Properties? 23 RNA Is an Information-Containing Molecule 71
Acid–Base Reactions Involve a Transfer of Protons 26 RNA Can Function as a Catalytic Molecule 71
2.3 Chemical Reactions, Chemical Evolution, and 4.4 The First Life Form 71
Chemical Energy 28 CANADIAN RESEARCH 4.1 Designing New Deoxyribozymes 72
How Do Chemical Reactions Happen? 28 CHAPTER REVIEW 73
What Is Energy? 28
Chemical Evolution: A Model System 30
How Did Chemical Energy Change during Chemical Evolution? 34 5 An Introduction to Carbohydrates 75

2.4 The Importance of Carbon 34 5.1 Sugars as Monomers 75

Linking Carbon Atoms Together 35 How Monosaccharides Differ 76
Functional Groups 36 Monosaccharides and Chemical Evolution 76
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 5.2 The Structure of Polysaccharides 77 7.3 Putting the Parts into a Whole 126
Starch: A Storage Polysaccharide in Plants 78 Structure and Function at the Whole-Cell Level 126
Glycogen: A Highly Branched Storage Polysaccharide in Animals 78 The Dynamic Cell 126
Cellulose: A Structural Polysaccharide in Plants 78 7.4 Cell Systems I: Nuclear Transport 127
Chitin: A Structural Polysaccharide in Fungi and Animals 80 Structure and Function of the Nuclear Envelope 127
Peptidoglycan: A Structural Polysaccharide in Bacteria 80 How Are Molecules Imported into the Nucleus? 128
Polysaccharides and Chemical Evolution 80
7.5 Cell Systems II: The Endomembrane System
5.3 What Do Carbohydrates Do? 80 Manufactures and Ships Proteins 129
The Role of Carbohydrates as Structural Molecules 81
Studying the Pathway through the Endomembrane
The Role of Carbohydrates in Cell Identity 81
System 129
The Role of Carbohydrates in Energy Storage 81
Entering the Endomembrane System: The Signal
CANADIAN ISSUES 5.1 Raymond Lemieux and the Synthesis of
Hypothesis 131
Sugars 82
Moving from the ER to the Golgi 132
CANADIAN RESEARCH 5.1 Natural and Artificial Sweeteners 84
What Happens inside the Golgi Apparatus? 133
CHAPTER REVIEW 85 How Do Proteins Reach Their Destinations? 133
7.6 Cell Systems III: The Dynamic Cytoskeleton 134
Actin Filaments 134
6 Lipids, Membranes, and the First Cells 87
Intermediate Filaments 135
6.1 Lipids 88 Microtubules 136
A Look at Three Types of Lipids Found in Cells 88 CANADIAN RESEARCH 7.2 Pathogenic Bacteria Alter the
The Structures of Membrane Lipids 89 Cytoskeleton of Human Cells 137
Flagella and Cilia: Moving the Entire Cell 138
6.2 Phospholipid Bilayers 90
Artificial Membranes as an Experimental System 90 CHAPTER REVIEW 140
Selective Permeability of Lipid Bilayers 91
How Does Lipid Structure Affect Membrane Properties? 92
How Does Temperature Affect the Fluidity and Permeability of 8 Cell–Cell Interactions 143
Membranes? 93
8.1 The Cell Surface 144
CANADIAN ISSUES 6.1 Lipids in Our Diet: Cholesterol,
The Structure and Function of an Extracellular Layer 144
Unsaturated Oils, Saturated Fats, and Trans Fats 94
The Cell Wall in Plants 144
6.3 Why Molecules Move across Lipid Bilayers: Diffusion The Extracellular Matrix in Animals 145
and Osmosis 96 8.2 How Do Adjacent Cells Connect and
Diffusion 96
Osmosis 96
Communicate? 146
Cell–Cell Attachments in Eukaryotes 147
CANADIAN RESEARCH 6.1 Liposomal Nanomedicines 98
Cells Communicate via Cell–Cell Gaps 150
6.4 Membrane Proteins 99
8.3 How Do Distant Cells Communicate? 151
Evolution of the Fluid-Mosaic Model 99
Cell–Cell Signalling in Multicellular Organisms 151
Systems for Studying Membrane Proteins 101
Signal Reception 152
Protein Transport I: Facilitated Diffusion via Channel
Signal Processing 152
Proteins 102
CANADIAN RESEARCH 8.1 The Discovery of Insulin 156
Protein Transport II: Facilitated Diffusion via Carrier Proteins 104
Signal Response 158
Protein Transport III: Active Transport by Pumps 104
Signal Deactivation 158
Plasma Membranes and the Intracellular Environment 106
Cross-Talk: Synthesizing Input from Many Signals 158
CHAPTER REVIEW 107 Quorum Sensing in Bacteria 159
The Big Picture: Macromolecules 110 CANADIAN RESEARCH 8.2 How Do Intracellular Proteins Bind
to Receptor Tyrosine Kinases? 160
CHAPTER REVIEW 161
UNIT 2 CELL STRUCTURE AND FUNCTION 112

7 Inside the Cell 112

9 Cellular Respiration and Fermentation 163

9.1 The Nature of Chemical Energy and Redox

7.1 Bacterial and Archaeal Cell Structures and Their Reactions 164
Functions 112 The Structure and Function of ATP 164
A Revolutionary New View 113 What Is a Redox Reaction? 166
Prokaryotic Cell Structures: A Parts List 113
CANADIAN RESEARCH 7.1 Bacteria Cells Have Their Own
9.2 An Overview of Cellular Respiration 168
Cytoskeleton 115 9.3 Glycolysis: Processing Glucose to Pyruvate 169
7.2 Eukaryotic Cell Structures and Their Functions 116 Glycolysis Is a Sequence of 10 Reactions 169
The Benefits of Organelles 116 How Is Glycolysis Regulated? 170
Eukaryotic Cell Structures: A Parts List 117 9.4 Processing Pyruvate to Acetyl CoA 171
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 9.5 The Citric Acid Cycle: Oxidizing Acetyl 11.2 How Does Mitosis Take Place? 215
CoA to CO2 173 Proteins Needed for Mitosis 215
How Is the Citric Acid Cycle Regulated? 173 Cytokinesis Results in Two Daughter Cells 218
What Happens to the NADH and FADH2? 173 How Do Chromosomes Move during Mitosis? 218
9.6 Electron Transport and Chemiosmosis: Building 11.3 Control of the Cell Cycle 220
a Proton Gradient to Produce ATP 176 The Discovery of Cell-Cycle Regulatory Molecules 220
Components of the Electron Transport Chain 176 CANADIAN RESEARCH 11.1 Yoshio Masui and the
The Chemiosmosis Hypothesis 177 Discovery of MPF 222
How Is the Electron Transport Chain Organized? 178 Cell-Cycle Checkpoints Can Arrest the Cell Cycle 223
The Discovery of ATP Synthase 178 11.4 Cancer: Out-of-Control Cell Division 225
Organisms Use a Diversity of Electron Acceptors 179 Properties of Cancer Cells 225
CANADIAN RESEARCH 9.1 The ATP Synthase 180 Cancer Involves Loss of Cell-Cycle Control 226
9.7 Fermentation 181 CANADIAN RESEARCH 11.2 A Newly Discovered
CANADIAN ISSUES 9.1 Making Biofuels with Fermentation Property of Cancer Cells 227
and Anaerobic Respiration 183 CHAPTER REVIEW 229
9.8 How Does Cellular Respiration Interact with Other
Metabolic Pathways? 184 UNIT 3 GENE STRUCTURE AND EXPRESSION 232
Catabolic Pathways Break Down Molecules as Fuel 184
Anabolic Pathways Synthesize Key Molecules 185
CHAPTER REVIEW 185
12 Meiosis 232

12.1 How Does Meiosis Occur? 233

Chromosomes Come in Distinct Types 233
10 Photosynthesis 188 The Concept of Ploidy 233
An Overview of Meiosis 234
10.1 Photosynthesis Harnesses Sunlight to Make The Phases of Meiosis I 237
Carbohydrate 188 The Phases of Meiosis II 238
Photosynthesis: Two Linked Sets of Reactions 189 A Closer Look at Prophase I 241
Photosynthesis Occurs in Chloroplasts 190 CANADIAN RESEARCH 12.1 The Proteins Required for
10.2 How Does Chlorophyll Capture Light Energy? 190 Prophase I of Meiosis 242
Photosynthetic Pigments Absorb Light 191 12.2 The Consequences of Meiosis 242
When Light Is Absorbed, Electrons Enter an Excited State 193 Chromosomes and Heredity 243
10.3 The Discovery of Photosystems I and II 195 Independent Assortment Produces Genetic Variation 243
How Does Photosystem II Work? 196 A Benefit of Crossing Over 244
How Does Photosystem I Work? 198 How Does Fertilization Affect Genetic Variation? 244
The Z Scheme: Photosystems II and I Work Together 198 12.3 Why Does Meiosis Exist? 245
10.4 How Is Carbon Dioxide Reduced to Produce The Paradox of Sex 245
Glucose? 200 The Purifying Selection Hypothesis 245
The Calvin Cycle Fixes Carbon 201 The Changing-Environment Hypothesis 246
The Discovery of Rubisco 202 12.4 Mistakes in Meiosis 247
Carbon Dioxide Enters Leaves through Stomata 203 How Do Mistakes Occur? 247
Mechanisms for Increasing CO2 Concentration Near Why Do Mistakes Occur? 248
Rubisco 204
CHAPTER REVIEW 249
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
13 Mendel and the Gene 252
Plants 206 13.1 Mendel’s Experimental System 252
CHAPTER REVIEW 207 What Questions Was Mendel Trying to Answer? 253
Garden Peas Served as the First Model Organism in Genetics 253
The Big Picture: Energy for Life 210
13.2 Mendel’s Experiments with a Single Trait 254
The Monohybrid Cross 254
11 The Cell Cycle 212 Particulate Inheritance 256
13.3 Mendel’s Experiments with Two Traits 258
11.1 Mitosis and the Cell Cycle 213
The Dihybrid Cross 258
What Is a Chromosome? 213
Using a Testcross to Confirm Predictions 260
Cells Alternate between M Phase and Interphase 214
The Discovery of S Phase 214 13.4 The Chromosome Theory of Inheritance 261
The Discovery of the Gap Phases 214 Meiosis Explains Mendel’s Principles 261
The Cell Cycle 214 Testing the Chromosome Theory 263

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 13.5 Extending Mendel’s Rules 265
Linkage: What Happens When Genes Are Located on the
16 Transcription, RNA Processing,
Same Chromosome? 265 and Translation 314
Do Heterozygotes Always Have a Dominant or Recessive
16.1 An Overview of Transcription 314
Phenotype? 267
Characteristics of RNA Polymerase 315
BOX 13.1 Quantitative Methods: Linkage 268
Initiation: How Does Transcription Begin? 316
How Many Alleles and Phenotypes Exist? 269
Elongation and Termination 317
Does Each Gene Affect Just One Trait? 269
Are Phenotypes Determined by Genes? 269 16.2 RNA Processing in Eukaryotes 318
What about Traits Like Human Height and Intelligence? 270 The Unexpected Discovery of Eukaryotic Genes in Pieces 318
RNA Splicing 319
13.6 Applying Mendel’s Rules to Humans 272 Adding Caps and Tails to Transcripts 320
Identifying Human Alleles as Recessive or Dominant 272
CANADIAN RESEARCH 13.1 The Genetics of Dog 16.3 An Introduction to Translation 321
Coat Colour 274 Ribosomes Are the Site of Protein Synthesis 321
Identifying Human Traits as Autosomal or Sex-Linked 275 Comparing Translation in Bacteria and Eukaryotes 321
How Does an mRNA Triplet Specify an Amino Acid? 321
CHAPTER REVIEW 276
16.4 The Structure and Function of Transfer RNA 323
What Do tRNAs Look Like? 324
14 DNA and the Gene: Synthesis and Repair 281 How Many tRNAs Are There? 325

14.1 What Are Genes Made Of? 282 16.5 The Structure and Function of Ribosomes 325
Initiating Translation 326
The Hershey–Chase Experiment 282
Elongation: Extending the Polypeptide 327
The Secondary Structure of DNA 283
Terminating Translation 327
14.2 Testing Early Hypotheses about DNA Synthesis: Posttranslational Modifications 329
The Meselson–Stahl Experiment 284 CANADIAN RESEARCH 16.1 RNA Synthesis in
14.3 A Comprehensive Model for DNA Synthesis 285 Mitochondria 329
How Does Replication Get Started? 287 CHAPTER REVIEW 330
How Is the Helix Opened and Stabilized? 287
How Is the Leading Strand Synthesized? 288
How Is the Lagging Strand Synthesized? 289 17 Control of Gene Expression in Bacteria 333
14.4 Replicating the Ends of Linear Chromosomes 292 17.1 Gene Regulation and Information Flow 333
CANADIAN RESEARCH 14.1 Telomeres, Telomerase, and Mechanisms of Regulation—An Overview 334
Cancer 294 Metabolizing Lactose—A Model System 335
14.5 Repairing Mistakes and Damage 294 17.2 Identifying Genes under Regulatory Control 336
Correcting Mistakes in DNA Synthesis 295 Replica Plating to Find Mutant Genes 336
Repairing Damaged DNA 296 Different Classes of Lactose Metabolism Mutants 337
Xeroderma Pigmentosum: A Case Study 296 Several Genes Are Involved in Lactose Metabolism 338
CHAPTER REVIEW 297 17.3 Mechanisms of Negative Control: Discovery of the
Repressor 338
The lac Operon 339
15 How Genes Work 300 Why Has the lac Operon Model Been So Important? 340
15.1 What Do Genes Do? 301 17.4 Mechanisms of Positive Control: Catabolite
The One-Gene, One-Enzyme Hypothesis 301 Repression 341
An Experimental Test of the Hypothesis 301 The CAP Protein and Binding Site 341
15.2 The Central Dogma of Molecular Biology 303 How Does Glucose Influence Formation of the CAP–cAMP
The Genetic Code Hypothesis 303 Complex? 341
RNA as the Intermediary between Genes and Proteins 303 CANADIAN RESEARCH 17.1 Bacterial Gene Expression and
Dissecting the Central Dogma 304 Probiotic Dairy Products 343
15.3 The Genetic Code 306 CHAPTER REVIEW 345
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 18 Control of Gene Expression in Eukaryotes 347
Point Mutation 309 18.1 Mechanisms of Gene Regulation in
Chromosome-Level Mutations 310 Eukaryotes—An Overview 348
CANADIAN RESEARCH 15.1 The Mutations Responsible for
Himalayan Fur Colour in Mink and Mice 311 18.2 Chromatin Remodelling 348
What Is Chromatin’s Basic Structure? 348
CHAPTER REVIEW 312 Evidence That Chromatin Structure Is Altered in Active Genes 349

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 How Is Chromatin Altered? 350 Using the Ti Plasmid to Produce Golden Rice 389
Chromatin Modifications Can Be Inherited 351 CHAPTER REVIEW 389
18.3 Initiating Transcription: Regulatory Sequences and
Regulatory Proteins 351
Some Regulatory Sequences Are Near the Promoter 351 20 Genomics 392
Some Regulatory Sequences Are Far from the Promoter 352
20.1 Whole-Genome Sequencing 392
The Role of Regulatory Proteins in Differential
How Are Complete Genomes Sequenced? 393
Gene Expression 354
Which Genomes Are Being Sequenced, and Why? 394
The Initiation Complex 354
Which Sequences Are Genes? 395
18.4 Posttranscriptional Control 356
20.2 Bacterial and Archaeal Genomes 396
Alternative Splicing of mRNAs 356
The Natural History of Prokaryotic Genomes 396
mRNA Stability and RNA Interference 357
Lateral Gene Transfer 397
How Is Translation Controlled? 358
Environmental Sequencing 397
Posttranslational Control 358
CANADIAN ISSUES 20.1 Genome Canada 398
18.5 How Does Gene Expression in Bacteria Compare with
20.3 Eukaryotic Genomes 398
That in Eukaryotes? 359 Parasitic and Repeated Sequences 399
18.6 Linking Cancer with Defects in Gene Regulation 360 Gene Families 401
Causes of Uncontrolled Cell Growth 360 Insights from the Human Genome Project 402
p53: A Case Study 361 CANADIAN RESEARCH 20.1 Human Genetic
CANADIAN RESEARCH 18.1 Chromatin Remodelling, Variation 404
Gene Transcription, and Cancer 362
20.4 Functional Genomics and Proteomics 406
CHAPTER REVIEW 363 What Is Functional Genomics? 406
The Big Picture: Genetic Information 366 What Is Proteomics? 406
Applied Genomics in Action: Understanding Cancer 407
CHAPTER REVIEW 408
19 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

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 24.3 The Process of Evolution: How Does Natural Selection
22 An Introduction to Animal Development 426
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 440 Weinberg Principle 478
The Gene Pool Concept 478
23.1 Gametogenesis, Pollination, and Fertilization 441 Deriving the Hardy–Weinberg Principle 478
How Are Sperm and Egg Produced? 441 The Hardy–Weinberg Model Makes Important Assumptions 479
Pollen–Stigma Interactions 441 How Does the Hardy–Weinberg Principle Serve as a Null
Double Fertilization 442 Hypothesis? 480
23.2 Embryogenesis 443 25.2 Types of Natural Selection 482
What Happens during Plant Embryogenesis? 443 Directional Selection 482
Which Genes and Proteins Set Up Body Axes? 445 Stabilizing Selection 483
23.3 Vegetative Development 446 Disruptive Selection 484
Meristems Provide Lifelong Growth and Development 446 Balancing Selection 485
Which Genes and Proteins Determine Leaf Shape? 447 25.3 Genetic Drift 485
CANADIAN RESEARCH 23.1 Apoptosis during the Formation of Simulation Studies of Genetic Drift 485
Plant Leaves 448 Experimental Studies of Genetic Drift 487
23.4 Reproductive Development 450 What Causes Genetic Drift in Natural Populations? 487
The Floral Meristem and the Flower 450 25.4 Gene Flow 489
The Genetic Control of Flower Structures 450 Gene Flow in Natural Populations 489
CHAPTER REVIEW 453 How Does Gene Flow Affect Fitness? 490
25.5 Mutation 490
UNIT 5 EVOLUTIONARY PROCESSES AND PATTERNS 455 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

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

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 Evaluating Molecular Phylogenies 640
30 Green Algae and Land Plants 599
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

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 Adaptations in Reproduction 690 ■ Chordata 7 Vertebrata 7 Mammalia 7 Eutheria (Placental
Metamorphosis 691 Mammals) 730
33.3 Key Lineages: Lophotrochozoans 691 ■ Chordata 7 Vertebrata 7 Reptilia 7 Lepidosauria (Lizards,
Snakes) 731
■ Lophotrochozoans 7 Rotifera (Rotifers) 692
■ Chordata 7 Vertebrata 7 Reptilia 7 Testudinia (Turtles) 731
■ Lophotrochozoans 7 Platyhelminthes (Flatworms) 692
■ Chordata 7 Vertebrata 7 Reptilia 7 Crocodilia (Crocodiles,
■ Lophotrochozoans 7 Annelida (Segmented Worms) 693
Alligators) 732
■ Lophotrochozoans 7 Mollusca 7 Bivalvia (Clams, Mussels,
■ Chordata 7 Vertebrata 7 Reptilia 7 Aves (Birds) 732
Scallops, Oysters) 695
■ Lophotrochozoans 7 Mollusca 7 Gastropoda (Snails, Slugs, 34.4 The Primates and Hominins 733
Nudibranchs) 696 The Primates 733
■ Lophotrochozoans 7 Mollusca 7 Polyplacophora CANADIAN ISSUES 34.1 Alberta during the Mesozoic Era 733
(Chitons) 697 Fossil Humans 736
■ Lophotrochozoans 7 Mollusca 7 Cephalopoda (Nautilus, The Out-of-Africa Hypothesis 739
Cuttlefish, Squid, Octopuses) 697 CHAPTER REVIEW 740
33.4 Key Lineages: Ecdysozoans 698
■ Ecdysozoans 7 Nematoda (Roundworms) 699
■ Ecdysozoans 7 Arthropoda 7 Myriapods (Millipedes, 35 Viruses 742
Centipedes) 700 35.1 Why Do Biologists Study Viruses? 743
■ Ecdysozoans 7 Arthropoda 7 Insecta (Insects) 700 Recent Viral Epidemics in Humans 743
■ Ecdysozoans 7 Arthropoda 7 Chelicerata (Spiders, Ticks, Current Viral Pandemics in Humans: HIV 744
Mites, Horseshoe Crabs, Daddy-Long-Legs, Scorpions) 703
35.2 How Do Biologists Study Viruses? 745
■ Ecdysozoans 7 Arthropoda 7 Crustaceans (Shrimp, Lobster,
Analyzing Morphological Traits 746
Crabs, Barnacles, Isopods, Copepods) 704
Analyzing Variation in Growth Cycles: Replicative and Latent
CANADIAN ISSUES 33.1 The First Census of Marine Life 705
Growth 746
CHAPTER REVIEW 707 Analyzing the Phases of the Replicative Cycle 748
35.3 What Themes Occur in the Diversification of
34 Deuterostome Animals 709 Viruses? 753
The Nature of the Viral Genetic Material 753
34.1 What Is an Echinoderm? 710 Where Did Viruses Come From? 754
The Echinoderm Body Plan 710 CANADIAN ISSUES 35.1 Viruses as Biological Control Agents 755
How Do Echinoderms Feed? 711 Emerging Viruses, Emerging Diseases 757
Key Lineages 712
35.4 Key Lineages of Viruses 759
■ Echinodermata 7 Asteroidea (Sea Stars) 712
■ Double-Stranded DNA (dsDNA) Viruses 759
■ Echinodermata 7 Echinoidea (Sea Urchins and
■ RNA Reverse-Transcribing Viruses (Retroviruses) 760
Sand Dollars) 713
■ Double-Stranded RNA (dsRNA) Viruses 760
34.2 What Is a Chordate? 713 ■ Negative-Sense Single-Stranded RNA ([-]ssRNA)
Three “Subphyla” 714 Viruses 761
Key Lineages: The Invertebrate Chordates 714 ■ Positive-Sense Single-Stranded RNA ([+]ssRNA) Viruses 761
■ Chordata 7 Cephalochordata (Lancelets) 715
CHAPTER REVIEW 762
■ Chordata 7 Urochordata (Tunicates) 715
34.3 What Is a Vertebrate? 716 UNIT 7 HOW PLANTS WORK 764
An Overview of Vertebrate Evolution 716
Key Innovations 718
Key Lineages 723 36 Plant Form and Function 764
■ Chordata 7 Vertebrata 7 Myxinoidea (Hagfish) and
Petromyzontoidea (Lampreys) 724 36.1 Plant Form: Themes with Many Variations 765
■ Chordata 7 Vertebrata 7 Chondrichthyes (Sharks, Rays, The Importance of Surface Area/Volume Relationships 765
Skates) 725 The Root System 766
CANADIAN RESEARCH 34.1 The Decline of Large, Predatory The Shoot System 768
Fishes in the World’s Oceans 726 The Leaf 770
CANADIAN RESEARCH 36.1 Does Phenotypic Plasticity of
■ Chordata 7 Vertebrata 7 Actinopterygii (Ray-Finned
Fishes) 727 Leaves Offer Protection against Herbivore Attack? 772
■ Chordata 7 Vertebrata 7 Actinistia (Coelacanths) and 36.2 Primary Growth Extends the Plant Body 774
Dipnoi (Lungfish) 728 How Do Apical Meristems Produce the Primary
■ Chordata 7 Vertebrata 7 Amphibia (Frogs, Salamanders, Plant Body? 774
Caecilians) 728 How Is the Primary Root System Organized? 775
■ Chordata 7 Vertebrata 7 Mammalia 7 Monotremata How Is the Primary Shoot System Organized? 776
(Platypuses, Echidnas) 729
36.3 Cells and Tissues of the Primary Plant Body 776
■ Chordata 7 Vertebrata 7 Mammalia 7 Marsupiala The Dermal Tissue System 778
(Marsupials) 730

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 The Ground Tissue System 778
The Vascular Tissue System 780 39 Plant Sensory Systems, Signals,
36.4 Secondary Growth Widens Shoots and Roots 782 and Responses 829
What Is a Cambium? 782 39.1 Information Processing in Plants 830
What Does Vascular Cambium Produce? 782 How Do Cells Receive and Transduce an External Signal? 830
What Does Cork Cambium Produce? 784 How Are Cell–Cell Signals Transmitted? 830
The Structure of a Tree Trunk 784 How Do Cells Respond to Cell–Cell Signals? 831
CHAPTER REVIEW 785 39.2 Blue Light: The Phototropic Response 832
Phototropins as Blue-Light Receptors 832
Auxin as the Phototropic Hormone 832
37 Water and Sugar Transport in Plants 788
39.3 Red and Far-Red Light: Germination and Stem
37.1 Water Potential and Water Movement 788 Elongation 836
What Is Water Potential? 789 The Red/Far-Red “Switch” 836
What Factors Affect Water Potential? 789 Phytochromes as Red/Far-Red Receptors 837
Calculating Water Potential 790 How Were Phytochromes Isolated? 837
Water Potentials in Soils, Plants, and the Atmosphere 791 CANADIAN RESEARCH 39.1 Plant Signalling Networks Help
37.2 How Does Water Move from Roots to Shoots? 792 Influence Proper Growth 838
Movement of Water and Solutes into the Root 793 39.4 Gravity: The Gravitropic Response 839
Water Movement via Root Pressure 794 The Statolith Hypothesis 840
Water Movement via Capillary Action 794 Auxin as the Gravitropic Signal 840
The Cohesion-Tension Theory 795 39.5 How Do Plants Respond to Wind and Touch? 841
37.3 Water Absorption and Water Loss 798 Changes in Growth Patterns 841
Limiting Water Loss 798 Movement Responses 841
Obtaining Carbon Dioxide under Water Stress 799 39.6 Youth, Maturity, and Aging: The Growth
CANADIAN RESEARCH 37.1 Ecological Pressures and the
Responses 842
Evolution of Drought Adaptation in Plants 799
Auxin and Apical Dominance 842
37.4 Translocation 800 Cytokinins and Cell Division 843
Tracing Connections between Sources and Sinks 801 Gibberellins and ABA: Growth and Dormancy 844
The Anatomy of Phloem 801 Brassinosteroids and Body Size 848
The Pressure-Flow Hypothesis 802 Ethylene and Senescence 848
Phloem Loading 803 An Overview of Plant Growth Regulators 849
Phloem Unloading 806
39.7 Pathogens and Herbivores: The Defence
CHAPTER REVIEW 807 Responses 851
How Do Plants Sense and Respond to Pathogens? 851
How Do Plants Sense and Respond to
38 Plant Nutrition 809 Herbivore Attack? 853
38.1 Nutritional Requirements of Plants 810 CHAPTER REVIEW 856
Which Nutrients Are Essential? 810 The Big Picture: How Vascular Plants Work 858
What Happens When Key Nutrients Are in Short Supply? 812
38.2 Soil: A Dynamic Mixture of Living and Nonliving
Components 813 40 Plant Reproduction 860
The Importance of Soil Conservation 814 40.1 An Introduction to Plant Reproduction 861
What Factors Affect Nutrient Availability? 814 Sexual Reproduction 861
38.3 Nutrient Uptake 816 The Land Plant Life Cycle 861
Mechanisms of Nutrient Uptake 816 Asexual Reproduction 863
Mechanisms of Ion Exclusion 818 40.2 Reproductive Structures 863
CANADIAN RESEARCH 38.1 Do Below-Ground Interactions When Does Flowering Occur? 864
between Plants and Fungi Influence Above-Ground The General Structure of the Flower 865
Interactions between Plants and Pollinators? 819 How Are Female Gametophytes Produced? 867
38.4 Nitrogen Fixation 822 How Are Male Gametophytes Produced? 867
The Role of Symbiotic Bacteria 822 40.3 Pollination and Fertilization 869
How Do Nitrogen-Fixing Bacteria Colonize Plant Roots? 823 Pollination 869
38.5 Nutritional Adaptations of Plants 824 CANADIAN RESEARCH 40.1 The Mating Strategies of
Epiphytic Plants 824 Flowering Plants 871
Parasitic Plants 825 Fertilization 873
Carnivorous Plants 825 40.4 The Seed 874
CHAPTER REVIEW 826 Embryogenesis 874

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 CANADIAN ISSUES 40.1 What Is the Effect of Agriculture on 42.4 Water and Electrolyte Balance in Terrestrial
Wild Bee Abundance and Crop Pollination? 875 Vertebrates 915
The Role of Drying in Seed Maturation 876 The Structure of the Kidney 915
Fruit Development and Seed Dispersal 876 The Function of the Kidney: An Overview 915
Seed Dormancy 878 Filtration: The Renal Corpuscle 916
Seed Germination 879 Reabsorption: The Proximal Tubule 917
CHAPTER REVIEW 880 Creating an Osmotic Gradient: The Loop of Henle 918
Regulating Water and Electrolyte Balance: The Distal Tubule
UNIT 8 HOW ANIMALS WORK 883 and Collecting Duct 920
CHAPTER REVIEW 922

41 Animal Form and Function 883

41.1 Form, Function, and Adaptation 884 43 Animal Nutrition 924

The Role of Fitness Trade-Offs 884 43.1 Nutritional Requirements 925
Adaptation and Acclimatization 884 Defining Human Nutritional Requirements 925
41.2 Tissues, Organs, and Systems: How Does Structure Meeting Human Nutritional Requirements 925
Correlate with Function? 886 CANADIAN ISSUES 43.1 Vitamin D Deficiency
Structure–Function Relationships at the Molecular and in Canada 926
Cellular Levels 886 43.2 Capturing Food: The Structure and Function of
Tissues Are Groups of Similar Cells That Function as a Unit 886 Mouthparts 929
Organs and Organ Systems 890 Mouthparts as Adaptations 929
41.3 How Does Body Size Affect Animal Physiology? 891 A Case Study: The Cichlid Jaw 929
Surface Area/Volume Relationships: Theory 891 43.3 How Are Nutrients Digested and Absorbed? 930
Surface Area/Volume Relationships: Data 892 An Introduction to the Digestive Tract 930
Adaptations That Increase Surface Area 894 An Overview of Digestive Processes 932
41.4 Homeostasis 894 The Mouth and Esophagus 933
Homeostasis: General Principles 894 The Stomach 934
The Role of Regulation and Feedback 895 The Small Intestine 936
The Cecum and Appendix 939
41.5 How Do Animals Regulate Body Temperature? 896 The Large Intestine 940
Mechanisms of Heat Exchange 896
Variation in Thermoregulation 896 43.4 Nutritional Homeostasis—Glucose as
Endothermy and Ectothermy: A Closer Look 897 a Case Study 940
Temperature Homeostasis in Endotherms 897 The Discovery of Insulin 940
Countercurrent Heat Exchangers 898 Insulin’s Role in Homeostasis 940
CANADIAN RESEARCH 41.1 Freeze-Tolerant Animals 900 Diabetes Can Take Several Forms 941
The Causes and Treatments of Diabetes 941
CHAPTER REVIEW 901
CANADIAN RESEARCH 43.1 Causes and Treatments of Diabetes
Mellitus Type 1 942
42 Water and Electrolyte Balance in CHAPTER REVIEW 944
Animals 904
42.1 Osmoregulation and Osmotic Stress 905 44 Gas Exchange and Circulation 946
What Is Osmotic Stress? 905 44.1 The Respiratory and Circulatory Systems 946
Osmotic Stress in Seawater 906
Osmotic Stress in Freshwater 906 44.2 Air and Water as Respiratory Media 947
Osmotic Stress on Land 906 How Do Oxygen and Carbon Dioxide Behave in Air? 947
How Do Cells Move Electrolytes and Water? 907 How Do Oxygen and Carbon Dioxide Behave in Water? 948

42.2 Water and Electrolyte Balance in Aquatic 44.3 Organs of Gas Exchange 949
Environments 908 Physical Parameters: The Law of Diffusion 949
How Do Sharks Excrete Salt? 908 How Do Fish Gills Work? 950
CANADIAN RESEARCH 42.1 The Bamfield Marine Sciences
How Do Insect Tracheae Work? 951
Centre and Research on Shark Osmoregulation 909 How Do Vertebrate Lungs Work? 952
How Do Freshwater Fish Osmoregulate? 910 Homeostatic Control of Ventilation 955

42.3 Water and Electrolyte Balance in Terrestrial 44.4 How Are Oxygen and Carbon Dioxide Transported
Insects 911 in Blood? 955
How Do Insects Minimize Water Loss from the Body Surface 911 Structure and Function of Hemoglobin 956
Types of Nitrogenous Wastes: Impact on Water Balance 912 CO2 Transport and the Buffering of Blood pH 958
CANADIAN RESEARCH 44.1 Dr. Peter Hochachka and
Maintaining Homeostasis: The Excretory System 913
Physiological Adaptation in Animals 959

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 44.5 The Circulatory System 961 How Do Muscles Contract? 1012
What Is an Open Circulatory System? 961 CHAPTER REVIEW 1016
What Is a Closed Circulatory System? 962
How Does the Heart Work? 964
CANADIAN ISSUES 44.1 The Risk Factors for Heart Attacks 967 47 Chemical Signals in Animals 1019
Patterns in Blood Pressure and Blood Flow 968
47.1 Cell–Cell Signalling: An Overview 1019
CHAPTER REVIEW 970 Major Categories of Chemical Signals 1020
Hormone Signalling Pathways 1021
What Makes Up the Endocrine System? 1022
45 Electrical Signals in Animals 973 Chemical Characteristics of Hormones 1023
45.1 Principles of Electrical Signalling 973 How Do Researchers Identify a Hormone? 1024
Types of Neurons in the Nervous System 974 47.2 What Do Hormones Do? 1025
The Anatomy of a Neuron 974 How Do Hormones Direct Developmental Processes? 1025
An Introduction to Membrane Potentials 975 How Do Hormones Coordinate Responses to Environmental
BOX 45.1 Quantitative Methods: Using the Nernst Equation to Change? 1027
Calculate Equilibrium Potentials 976 How Are Hormones Involved in Homeostasis? 1028
How Is the Resting Potential Maintained? 976
47.3 How Is the Production of Hormones Regulated? 1030
Using Microelectrodes to Measure Membrane Potentials 977
The Hypothalamus and Pituitary Gland 1031
What Is an Action Potential? 978
Control of Adrenaline by Sympathetic Nerves 1033
45.2 Dissecting the Action Potential 979 47.4 How Do Hormones Act on Target Cells? 1033
Distinct Ion Currents Are Responsible for Depolarization and
Steroid Hormones Bind to Intracellular Receptors 1034
Repolarization 979
Hormones That Bind to Cell-Surface Receptors 1035
How Do Voltage-Gated Channels Work? 979
CANADIAN ISSUES 47.1 Estrogens in the Environment 1036
How Is the Action Potential Propagated? 981
Why Do Different Target Cells Respond in Different Ways? 1038
45.3 The Synapse 983 CHAPTER REVIEW 1039
Synapse Structure and Neurotransmitter Release 984
What Do Neurotransmitters Do? 985
Postsynaptic Potentials 985 48 Animal Reproduction 1041
CANADIAN RESEARCH 45.1 David Suzuki and the Discovery
of the Genes Encoding Neuron Proteins 987 48.1 Asexual and Sexual Reproduction 1041
How Does Asexual Reproduction Occur? 1042
45.4 The Vertebrate Nervous System 989 Switching Reproductive Modes: A Case History 1042
What Does the Peripheral Nervous System Do? 989 Mechanisms of Sexual Reproduction: Gametogenesis 1043
Functional Anatomy of the CNS 989
How Does Memory Work? 991 48.2 Fertilization and Egg Development 1045
External Fertilization 1045
CHAPTER REVIEW 994 Internal Fertilization 1045
Unusual Aspects of Mating 1046
Why Do Some Females Lay Eggs While Others Give Birth? 1047
46 Animal Sensory Systems and Movement 996
48.3 Reproductive Structures and Their Functions 1048
46.1 How Do Sensory Organs Convey Information The Male Reproductive System 1048
to the Brain? 997 The Female Reproductive System 1050
Sensory Transduction 997
48.4 The Role of Sex Hormones in Mammalian
Transmitting Information to the Brain 998
Reproduction 1051
46.2 Hearing 998 Which Hormones Control Puberty in Mammals? 1052
How Do Sensory Cells Respond to Sound Waves and Which Hormones Control the Menstrual Cycle in
Other Forms of Pressure? 998 Mammals? 1053
The Mammalian Ear 999
48.5 Pregnancy and Birth in Mammals 1058
Sensory Worlds: What Do Other Animals Hear? 1001
Gestation and Early Development in Marsupials 1058
46.3 Vision 1002 Major Events during Human Pregnancy 1058
The Insect Eye 1002 How Does the Mother Nourish the Fetus? 1059
The Vertebrate Eye 1002 Birth 1061
CANADIAN RESEARCH 46.1 Why Do Wind Farms Kill Bats? 1004 CANADIAN ISSUES 48.1 Canada’s Assisted Human
Sensory Worlds: Do Other Animals See Colour? 1008 Reproduction Act 1062
46.4 Taste and Smell 1008 CHAPTER REVIEW 1063
Taste: Detecting Molecules in the Mouth 1008
Olfaction: Detecting Molecules in the Air 1009
46.5 Movement 1010
49 The Immune System in Animals 1065
Skeletons 1010 49.1 Innate Immunity 1066
Muscle Types 1012 Barriers to Entry 1066

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 The Innate Immune Response 1067 50.5 Biogeography: Why Are Organisms Found Where
49.2 The Adaptive Immune Response: Recognition 1069 They Are? 1110
An Introduction to Lymphocytes 1070 Abiotic Factors 1110
The Discovery of B Cells and T Cells 1071 CANADIAN ISSUES 50.1 Do Insect Outbreaks Contribute to
The Clonal-Selection Theory 1071 Climate Change? 1111
CANADIAN RESEARCH 49.1 Tak Wah Mak and the Discovery The Role of History 1113
of the T-Cell Receptor 1073 CANADIAN RESEARCH 50.2 Salmon Migration in a Warming
How Does the Immune System Distinguish Self from World 1114
Nonself? 1075 Biotic Factors 1115
Biotic and Abiotic Factors Interact 1116
49.3 The Adaptive Immune Response: Activation 1076
T-Cell Activation 1077 CHAPTER REVIEW 1118
B-Cell Activation and Antibody Secretion 1078
49.4 The Adaptive Immune Response: Culmination 1079
How Are Bacteria and Other Foreign Cells Killed? 1080
51 Behavioural Ecology 1121
How Are Viruses Destroyed? 1080 51.1 An Introduction to Behavioural Ecology 1121
Why Does the Immune System Reject Foreign Tissues Proximate and Ultimate Causation 1122
and Organs? 1080 Conditional Strategies and Decision Making 1122
Responding to Future Infections: Immunological Memory 1081 CANADIAN RESEARCH 51.1 Do Male Redback Spiders Benefit
49.5 What Happens When the Immune System Doesn’t from Being Eaten by Their Mates? 1123
Five Questions in Behavioural Ecology 1124
Work Correctly? 1083
Immunodeficiency Diseases 1083 51.2 What Should I Eat? 1124
Allergies 1083 Foraging Alleles in Drosophila melanogaster 1124
Optimal Foraging in White-Fronted Bee-Eaters 1125
CHAPTER REVIEW 1084
The Big Picture: How Humans Work 1086 51.3 Whom Should I Mate With? 1125
Sexual Activity in Anolis Lizards 1126
How Do Female Barn Swallows Choose Mates? 1127
UNIT 9 ECOLOGY 1088 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
50 An Introduction to Ecology 1088
51.5 How Should I Communicate? 1130
50.1 Levels of Ecological Study 1088 Honeybee Language 1131
Organismal Ecology 1089 Modes of Communication 1132
Population Ecology 1089 When Is Communication Honest or Deceitful? 1133
Community Ecology 1089 51.6 When Should I Cooperate? 1134
Ecosystem Ecology 1090 Kin Selection 1134
How Do Ecology and Conservation Efforts Interact? 1090 BOX 51.1 Quantitative Methods: Calculating the
50.2 Types of Aquatic Ecosystems 1090 Coefficient of Relatedness 1136
Nutrient Availability 1090 Reciprocal Altruism 1136
Water Flow 1091 An Extreme Case: Abuse of Non-Kin in Humans 1137
Water Depth 1091 CHAPTER REVIEW 1138
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
52 Population Ecology 1141

■ Freshwater Environments 7 Streams 1096 52.1 Demography 1141

■ Freshwater/Marine Environments 7 Estuaries 1097 Life Tables 1142
■ Marine Environments 7 The Ocean 1097 CANADIAN RESEARCH 52.1 Tyrannosaur Life Tables 1144
The Role of Life History 1145
50.3 Types of Terrestrial Ecosystems 1098
BOX 52.1 Quantitative Methods: Using Life Tables to Calculate
■ Terrestrial Biomes 7 Tropical Wet Forest 1100
Population Growth Rates 1146
■ Terrestrial Biomes 7 Subtropical Deserts 1101
■ Terrestrial Biomes 7 Temperate Grasslands 1102 52.2 Population Growth 1147
■ Terrestrial Biomes 7 Temperate Forests 1103 Quantifying the Growth Rate 1147
■ Terrestrial Biomes 7 Boreal Forests 1104 Exponential Growth 1148
■ Terrestrial Biomes 7 Arctic Tundra 1105 Logistic Growth 1148
BOX 52.2 Quantitative Methods: Developing and Applying
50.4 The Role of Climate and the Consequences of
Population Growth Equations 1149
Climate Change 1105
What Limits Growth Rates and Population Sizes? 1151
Global Patterns in Climate 1106
How Will Global Climate Change Affect Ecosystems? 1108 52.3 Population Dynamics 1152

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 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

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 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.

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 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 Canadian content throughout the book. Each chapter now
need to learn the language of biology and understand has at least one Canadian Research or Canadian Issues box.
fundamental concepts, but they also need to apply these con- We have chosen examples that both illustrate one of the main
cepts to new situations, analyze experimental design, synthesize concepts in the chapter and highlight the diversity of science
results, and evaluate hypotheses and data. being done in Canadian universities, colleges, and other or-
We wrote this book for instructors who embrace this chal- ganizations. These boxes now end with a “Think About It”
lenge—who want to help their students learn how to think like a question to allow students to test their understanding of the
biologist. The essence of higher education is to promote higher- material.
order thinking. Our job is to help students understand biological ⦁ The Big Picture These new two-page spreads are meant to
science at all six levels of Bloom’s taxonomy of learning. 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-
Analyze Evaluate Synthesize 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
Apply Big Picture: Macromolecules on pages 110–111). In addition,
the book’s MasteringBiology® website has 10 new concept
Explain map activities based on Big Picture content that will allow
you to explore the concepts and their connections with your
Remember
students during lectures.
⦁ BioSkills Students completing introductory biology need to
have acquired skills—the ability to read a graph, interpret an
Bloom’s Taxonomy. An annotated version of this graphic can be equation, understand the bands on a gel. The previous edition
found in “Preface to Students: How to Use This Book” at the front of
of Biological Science contained a series of appendices focused
this book.
on key skills for introductory biology students. Instructors and
students found them extraordinarily helpful. New in this edi-
The Evolution of a Textbook tion are BioSkills on using the metric system, common Latin
Evolution can be extremely fast in populations with short gen- and Greek roots, techniques for isolating and visualizing cell
eration times and high mutation rates. Biology textbooks are no components, cell and tissue culture methods, and model or-
exception. Generation times have to be short because the pace of ganisms. BioSkills are located in Appendix B.
research in biology and student learning is so fast. This book, in ⦁ Answer Key New to the Second Canadian Edition are sug-
particular, evolves quickly because it incorporates so many new gested answers to all questions and exercises in the textbook.
ideas with each edition. Some of these “alleles” are novel muta- Students asked us to make this important change between
tions, but most arrive via lateral transfer—from advisors, review- editions to make the book a more complete study tool. The
ers, friends, students, and the literature. answer key will allow them to self-check their understanding
while reading and when reviewing for exams. Answers are in
What’s New in This Edition Appendix A.
This revision was about making the book a better teaching and ⦁ Experiment Boxes This text’s hallmark has always been its
learning tool. To help students manage the mass of information emphasis on experimental evidence—on teaching how we
and ideas that is contemporary biology, we broke long para- know what we know. In the previous edition, key experi-
graphs into shorter paragraphs, made liberal use of numbered ments were converted to a boxed format so students could
lists and bulleted lists to “chunk” information and ideas, and easily navigate through the logic of the question, hypothesis,
broke out dozens of new sections and subsections. and test. In this edition, we added a new question to every
In addition, we came up with a long list of new or expanded experiment box to encourage students to analyze some aspect
features. of the experiment’s design.

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 ⦁ Art Program Recent research shows that students are more 3. Check Your Understanding boxes, at the end of key sec-
likely to interpret phylogenetic trees correctly if the trees tions, with a bulleted list of key points.
are designed with U-shaped branches instead of Y-shaped 4. Summary tables that pull information together in a compact
branches. We responded by redesigning every phylogenetic format that is easy to review and synthesize.
tree in the text. To make other subject areas more accessible
to visual learners, we enlarged figures, replaced hundreds of
photos with clearer images, and strove to streamline labels
Changes to Blue Thread Scaffolding
and graphics across the board. (More on improvements to the Each edition of this text has added tools to help students with
art program below.) metacognition—understanding what they do and don’t under-
stand. Novices like to receive information passively, and easily
⦁ MasteringBiology Quizzes MasteringBiology gives students persuade themselves that they know what’s going on. Experts are
round-the-clock access to quizzes. We developed 550 new as- skeptical—they want to solve some problems before they’re con-
signable questions based on the book’s “Blue Thread” ques- vinced that they know and understand an idea.
tions (more on the “Blue Thread” and its evolution below). In the previous edition, we formalized the metacognitive tools
We also developed a cumulative practice test to simulate what in Biological Science as a “Blue Thread” set of questions; in this
a real exam might be like. To help students keep up with their edition, we revised each question and put answers in the back of
reading, we created 55 new reading quizzes—one for each the book for easy student access.
chapter—that you can assign through MasteringBiology.
1. In-text “You should be able to’s” offer exercises on topics
⦁ MasteringBiology Experimental Inquiry Tutorials The call that professors and students have identified as the most dif-
to teach students about the process of science has never been ficult concepts in each chapter.
louder. In response, a team led by Tom Owens of Cornell 2. Caption Questions and Exercises challenge students to ex-
University developed 10 new interactive tutorials on clas- amine critically the information in a figure or table—not just
sic scientific experiments—ranging from Meselson–Stahl on absorb it.
DNA replication to the Grants’ work on Galápagos finches
3. Think About It questions test or expand on an important
and Connell’s work on competition. Students who use these
concept in each Canadian Research and Canadian Issues box.
interactive tutorials should be better prepared to think criti-
cally about experimental design and evaluate the wider im- 4. Check Your Understanding boxes present two to three
plications of the data—preparing them to do the work of real tasks that students should be able to complete in order to
scientists in the future. demonstrate a mastery of summarized key ideas.
5. Chapter Summaries include “You should be able to” prob-
⦁ MasteringBiology BioFlix Animations and Tutorials lems or exercises related to each of the key concepts declared
BioFlix™ are movie-quality, 3-D animations available on in the gold thread.
MasteringBiology. They focus on the most difficult core
6. End-of-Chapter Questions are organized around Bloom’s
topics and are accompanied by in-depth, online tutori-
taxonomy of learning, so students can test their understand-
als that provide hints and feedback to help guide student
ing at the knowledge, comprehension, and application levels.
learning. Thirteen BioFlix were available with the previous
edition of Biological Science. Five new BioFlix 3-D anima- The fundamental idea is that if students really understand
tions and tutorials have been developed for this edition— a piece of information or a concept, they should be able to do
on mechanisms of evolution, homeostasis, gas exchange, something with it. How do you get to Carnegie Hall? Practise.
population ecology, and the carbon cycle. As students mature as biologists-in-training and start taking
upper-division courses, most or all of this scaffolding can dis-
appear. By the time our students are in their fourth year, they
Changes to Gold Thread Scaffolding should have enough expertise to construct a high-level under-
The previous edition introduced a set of tools designed to help standing on their own. But if a well-designed scaffold isn’t there
with a chronic problem for novice learners: picking out impor- to get them started in their first and second years when they are
tant information. Novices highlight every line in the text and try novices, most will flounder. We have to help them learn how to
to memorize everything mentioned in lecture; experts instinc- become good students.
tively home in on the key unifying ideas.
For students to make the novice-to-expert transition, we have Supporting Visual Learners
to help them with features like:
Figures can help students, especially visual learners, at all levels
1. Key concepts that are declared at the start of each chapter, of Bloom’s Taxonomy—not only to understand and remember
highlighted with a key icon within the chapter, and reviewed the material, but also to exercise higher levels of critical thinking.
at the end of the chapter. The overall goal of the Second Canadian Edition art revision was
2. In-text highlighting, in gold, that directs their attention to to hone the figures for accessibility to help novice learners recog-
particularly important ideas. nize and engage with important visual information. In addition

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 to redesigning the previously mentioned phylogenetic trees, Kim Robert Holmberg, Athabasca University
Quillin led the effort to enhance virtually every other aspect of Andrea Kirkwood, University of Ontario Institute of Technology
the visual-teaching program. David LesBarreres, Laurentian University
Ivona Mladenovic, Simon Fraser University
⦁ Art and Photos Kim enlarged art and photographs in fig- Barbara Moon, University of the Fraser Valley
ures throughout the book to increase clarity by making details Blythe Nilson, University of British Columbia, Okanagan
physically easier to see. She also reduced the amount of detail Tanya Noel, York University
in labels and graphics to simplify, simplify, simplify. Robin Owen, Mount Royal University
Carol Pollock, University of British Columbia
⦁ Colour Use Kim continues to use colour strategically to
Melanie Rathburn, Mount Royal University
draw attention to important parts of the figures. In this revi-
Fiona Rawle, University of Toronto, Mississauga
sion, she boosted colour contrast in many figures to make the
Carla Starchuk, University of Alberta
art more vibrant and the details easier to see. Alexandra Venter, Athabasca University
⦁ Molecular Icons Kim redesigned many molecular icons to Usha Vivegananthan, Mohawk College
simplify their shapes. The overall contours are based on mo- Debbie Wheeler, University of the Fraser Valley
lecular coordinates, when available, to accurately represent Ken Wilson, University of Saskatchewan
size and geometry, but she smoothed the textures for a sim-
pler appearance—one that is more memorable and pleasing. Supplements Contributors
⦁ Molecular Models New molecular models have been intro- Instructors depend on an impressive array of support materials—
duced to help students visualize structure–function relation- in print and online—to design and deliver their courses. The stu-
ships. In Chapter 5, for example, redesigned 2-D line drawings dent experience would be much weaker without the study guide,
of sugars are now paired with 3-D ball-and-stick models. test bank, activities, animations, quizzes, and tutorials written by
⦁ “Pointers” The Second Canadian Edition figures still use the following individuals:
pointer annotations as a “whisper in the ear” to guide students
in interpreting figures, but Kim has replaced the hand with an Study on the Go—Nancy Flood, Thompson Rivers University
arrow to be more precise. PowerPoint and PRS Questions—Sharon Gillies, University of
the Fraser Valley
Testbank—Tamara Kelly and Nicole Nivillac, York University
Acknowledgments
Reviewers Book Team
The peer review system is the key to quality and clarity in sci- As coauthors on the Second Canadian edition of Biological Science,
ence publishing. In addition to providing a filter, the investment we would like to thank all the talented people who were involved
that respected individuals make in vetting the material—catching in the production of our textbook. This very professional team
errors or inconsistencies and making suggestions to improve the was headed by Gary Bennett, Vice President and Editorial Direc-
presentation—gives authors, editors, and readers confidence that tor. We are grateful for the guidance of both Michelle Sartor and
what they are publishing and reading meets rigorous professional Lisa Rahn, who replaced Michelle as Senior Acquisitions Editor.
standards. Ken Ko of Queen’s University shared the writing duties with
Peer review plays the same role in textbook publishing. The us on the topics of Canadian research on plant systems and on
time and care that this book’s reviewers have invested is a tribute gene regulation. We are impressed by the polished art produced
to their professional integrity, their scholarship, and their con- by Julia Hall from our hand-drawn scribbles.
cern for the quality of teaching. This edition has been revised Developmental Editor Joanne Sutherland patiently and expertly
and improved based on insights from the following individuals: provided guidance and encouragement throughout the process,
while the final version of the text was guided by Project Man-
Eric Alcorn, Acadia University ager Carrie Fox and Copyeditor Audra Gorgiev, directed by Lead
Greg Beaulieu, University of Victoria Project Manager Avinash Chandra, and effectively and efficiently
Todd Bishop, Dalhousie University managed by in-house Project Manager Rachel Thompson.
Peter Boag, Queen’s University It is always a genuine pleasure to work with Senior Market-
Dora Cavallo-Medved, University of Windsor
ing Manager Kim Ukrainec and Marketing Coordinator Kathie
Brett Couch, University of British Columbia
Kirchsteiger. These dedicated individuals supervise Pearson
Christine Dalton, University of the Fraser Valley
Nancy Flood, Thompson Rivers University
Canada’s talented sales reps, who listen to professors, advise the
Chris Garside, formerly of the University of Ontario Institute editorial staff, and get the book into student hands.
of Technology, now at University of Toronto, St. George Finally, we would like to offer our heartfelt thanks for the de-
Kim Gilbride, Ryerson University tailed suggestions from the reviewers who cast a careful eye over
Sharon Gillies, University of the Fraser Valley each draft chapter. Their thoughtful comments are very much
Anna Hicks, Memorial University appreciated.

PREFACE TO INSTRUCTORS xxiii

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 Serving a Community of Teachers and practice of evidence-based teaching into this textbook, and
welcome your comments, suggestions, and questions.
There is nothing that inspires us more than getting together with Thank you for considering this text, and for your work on be-
other biology instructors and “talking shop.” These meetings half of your students. We have the best jobs in the world.
may be during teaching workshops or less formal get-togethers.
SCOTT FREEMAN
While we all have our own personal teaching styles, these styles University of Washington
are a collection of ideas tested and refined with our colleagues—
MIKE HARRINGTON
or borrowed outright! University of Alberta
Research on biology education is gathering momentum, try- mjh@ualberta.ca
ing to catch up on the trail blazed by physics education research-
JOAN SHARP
ers, bringing the same level of rigour to our classrooms that we Simon Fraser University
bring to our lab benches and field sites. We try to bring the spirit jsharp@sfu.ca

xxiv PREFACE TO INSTRUCTORS

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 Content Highlights of the Second Canadian Edition

A
s discussed in the preface, a major focus of this revi- Unit 2 Cell Structure and Function
sion is to enhance the pedagogical utility of Biologi-
cal Science. New Canadian content has been added Chapter 7 Recent discoveries on bacterial cell structure is
to many of the chapters. Another major goal is to ensure that described in a new Canadian Research box. The relationships
the content reflects the current state of science and is accu- between chloroplasts and other plastids and between the lyso-
rate. In addition, every chapter has been rigorously evaluated somes and other endomembrane system components have
for discussions that, in the previous edition, may have been too been emphasized. Centrifugation is moved to BioSkills 11 in
complex or overly detailed. As a result of this scrutiny, certain Appendix B.
sections in every chapter have been simplified, content has Chapter 8 New sections on quorum sensing in bacteria and
been pruned judiciously, and the approach to certain topics has cross-talk among signal-transduction pathways have been
been re-envisioned to enhance student comprehension. In this added. Canadian research on both of these topics has also
section, some of the key content improvements to the textbook been included.
are highlighted.
Chapter 9 The discussions of mitochondrial structure, ATP
yield from glucose oxidation, and the role of GDP in the cit-
Unit 1 The Molecules of Life ric acid cycle have been updated. The introductory section on
cellular respiration has been simplified. The ATP synthase
Chapter 1 A new experiment on ant navigation and discus- enzyme is the subject of a new Canadian Research box.
sions of tree-based naming systems and artificial selection in
Chapter 10 A new section on regulation (inhibition) has
maize has been added. Coverage is expanded on the defin-
been added. The sections on C4 and CAM photosynthesis
ition of life.
now emphasize the role of these pathways in increasing CO2
Chapter 2 The descriptions of bond angles and the geometry concentrations versus water conservation. The Canadian re-
of simple molecules are simplified. Added is a discussion on search described in this chapter has been updated to describe
the hot-start hypothesis as well as a new Key Concept on the a project to improve photosynthesis in rice that is currently
nature of chemical energy. under way at the University of Toronto.
Chapter 3 This chapter has been streamlined by eliminating Chapter 11 The beginning of the chapter has been com-
discussion of optical isomers/chirality and reducing coverage pletely revised to include why and how each step of mitosis
of enzyme kinetics and reaction rates. New Canadian con- occurs. Mitosis is now presented in three ways: (1) the events
tent includes the impact of prions on the cattle industry and that define each stage, (2) the reason why the cell does what
new “designer proteins” being developed at the University of it does, and (3) how the chromosome behaviour is the result
Guelph. of microtubule, cohesin, condensin, and nuclear lamin pro-
Chapter 4 The discussion of RNA is expanded to include teins working in progression. The discussion on the role of
recently discovered roles for RNAs in cells. There are more activated MPF has been updated to include the triggering M
detailed explanations on how nucleotides are named and how phase of the cell cycle. Animal cell culture methods are moved
DNA molecules are measured. “Designer nucleotides” made to BioSkills 12 in Appendix B.
at McMaster University are the subject of a new Canadian Re-
search box. Also added is a new summary table (Table 4.1)
comparing DNA and RNA structure. Unit 3 Gene Structure and Expression
Chapter 5 A stronger emphasis on the link between elec- Chapter 12 The topic of crossovers has been expanded and
tronegativity of atoms and potential energy in, C ¬ C, C ¬ H, is the subject of a new Canadian Research box. The discus-
C ¬ O and bonds is developed. New ball-and-stick models sions of recombination rates and aneuploidy rates in humans
are added to clarify the differences in location and orientation are updated. New micrographs have been added to the phases
of functional groups. of meiosis figure (Figure 12.7).
Chapter 6 Coverage of secondary active transport has been Chapter 13 The linkage discussion and notation in fly crosses
expanded. Also included in this chapter is current research have been simplified. Sex-linkage is moved to the Mendelian
on the “first cell” and a discussion of nonrandom distribution section (Section 13.4 The Chromosome Theory of Inheritance),
of membrane proteins and phospholipids. and mapping is now covered in Box 13.1 Quantitative Methods:

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 Linkage. A new summary table (Table 13.3) presenting basic Chapter 23 A new section introducing basic concepts in angio-
vocabulary used in Mendelian genetics has been added. sperm gametogenesis has been added.
Chapter 14 A new space-filling model of DNA has been added
to Figure 14.4. The E. coli DNA polymerases I and III are now
described independently. Canadian research on the relationship
Unit 5 Evolutionary Processes and Patterns
between telomerase, telomeres, and cancer has been included. Chapter 24 A section on the internal consistency of diverse data
Chapter 15 Discussions on mutation in the melanocortin as evidence for evolution, including a new phylogeny and time
receptor (link to mouse-coat colour camouflage) and karyo- line of whale evolution, has been added. Figure 24.6, depicting
types of cancerous cells have been added. A new Canadian the evolution of the Galápagos mockingbird, and long-term data
Research box on fur colour in mink provides an opportunity on ground finches (Figure 24.18) are updated to reflect the most
for students to practise using the genetic codon table. current science. There is a new graph on the evolution of drug
resistance in pathogenic bacteria (Figure 24.14).
Chapter 16 The sections on transcription in bacteria and eu-
karyotes are now combined. The structure of the translation Chapter 25 The genetic drift example has changed from breed-
initiation complex in bacteria has been updated to reflect cur- ing in a small population on Pitcairn Island to coin flips simu-
rent science; snRNAs have been added to the discussion of lating mating in a single couple (using data from the author’s
RNA splicing. The subject of gene expression in organelles is classroom). The prairie lupine gene flow example is replaced
described within a new Canadian Research box. by recent work on an island population of the great tit, Parus
major. Notes on balancing selection, assortative mating, and
Chapter 17 The chapter was streamlined with the removal
interactions among evolutionary forces have been included.
of discussions of DNA fingerprinting and the structure of the
operator and DNA-binding proteins. Treatment of catabolite Chapter 26 The ecological species concept has been added to
repression/positive control has been trimmed. A practical ap- the species definitions included in the chapter. The speciation-
plication of bacterial gene expression done at the Université by-vicariance example has been changed from ratites to snap-
Laval has been added. ping shrimp, and the sympatric speciation example featuring
soapberry bugs has been changed to apple/hawthorn flies.
Chapter 18 Included in this chapter is a new summary table
(Table 18.1) comparing control of gene expression in bac- Chapter 27 The sections on adaptive radiation and mass
teria and eukaryotes. The chapter now describes the types of extinction have been completely reorganized. A new hypoth-
histone proteins in eukaryotes and new Canadian research esis for the cause of the Cambrian explosion is included, and
on the relationship between these proteins and cancer. Also detail on the “new genes, new bodies” hypothesis has been
added are discussions on ubiquitination and protein degrada- removed. Presentation of “Life’s Time Line” has been signifi-
tion, the importance of epigenetic inheritance (chromosome cantly overhauled (see Figures 27.8, 27.9, and 27.10). The
structure), and the histone code hypothesis. Burgess Shale fossil site is now introduced in this chapter.
Chapter 19 Figure 19.11 has been updated to show Sanger
sequencing done with fluorescently labelled nucleotides.
Southern/Northern/Western blots have moved to BioSkills 9 in
Unit 6 The Diversification of Life
Appendix B. The discussions on golden rice, the impact of GM The model organisms have been moved to BioSkills 14 in
crops, and SNP association studies for human diseases have Appendix B. Phylogenetic trees have been redrawn to reflect a
been updated with the most recent research. Notes on “next- horizontal orientation with U-shaped branches for easier com-
generation” sequencing technologies have been included. prehension.
Chapter 20 Human health applications now emphasize the Chapter 28 New information on mechanisms of pathogenicity
use of genomics and microarrays to study cancer. Several dat- is added. Extensive updates include new notes on archaeon–
asets are updated, including sequencing database totals. New eukaryote polymerases, the discovery of extensive biomass in
notes on miRNA genes, metagenomics, and the definition of the marine subfloor, an archaeon associated with a human
the gene have been added. disease, discovery of N-fixation and nitrification in archaea,
and bacteriorhodopsin’s role in phototrophy.
Chapter 29 A stronger emphasis on endosymbiosis as a
Unit 4 Developmental Biology theme in protist diversification has been threaded throughout
Chapter 21 The discussions of bicoid and regulatory gene this chapter.
cascades are simplified. New material on auxin as a master Chapter 30 New content on green algae as a grade and on
regulator in early development and the importance of apop- convergence in vascular tissue in mosses/vascular plants and
tosis have been added. gnetophytes/angiosperms has been added.
Chapter 22 The discussion about sea urchin fertilization and Chapter 31 The dynamic nature of mycelia, the importance
variation has been streamlined. of glomalin in soil, the role of mating types, and the discovery

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 of “multigenomic” asexual glomales all have new supporting on “talking trees” is included. The coverage of the receptors
material. for GA, auxin, ABA, and brassinosteroids and MeSA’s role
Chapter 32 The treatment of embryonic tissues, develop- in the SAR has been updated with the most current research.
mental patterns, the coelom, and body symmetry has been Plant tissue culture methods have been moved to BioSkills 12
updated to reflect the latest scientific thinking. A shift in em- in Appendix B.
phasis to the origin of the neuron and cephalization has been Chapter 40 Comments on day-length sensing and on pol-
implemented. A new Canadian Research box describes an early lination syndromes are new to this chapter.
Cambrian site in Jasper National Park, Alberta, that has yielded
microfossils identified as the oldest molluscan radulae.
Unit 8 How Animals Work
Chapter 33 New commentary on the independent transi-
tions to land as well as a clarified discussion on the nature of Chapter 41 New details on tissue types (especially connect-
the ecdysozoan–lophotrochozoan split are included. The dis- ive tissue) have been incorporated. The discussion of thermo-
cussion of annelids is updated to reflect recent results. A new regulation has been completely reorganized for a more logical
Canadian Issues box describes the First Census of Marine Life flow The research of Carleton University’s Ken Storey, who
and the Canadian Healthy Oceans Network (CHONe), which explores how some animals survive cold Canadian winters, is
worked with the census to establish a biodiversity database now included.
for Canada’s Pacific, Arctic, and Atlantic oceans. Chapter 42 The sections on the shark rectal gland and the
Chapter 34 The coverage of the echinoderm endoskeleton has mammalian loop of Henle have been revised to improve focus.
been expanded and a phylogeny of early tetrapods has been Chapter 43 A description of incomplete digestive systems is
added to the fin-to-limb transition figure (Figure 34.16). New now included, and coverage of comparative aspects of digest-
data have been incorporated in the evolution-of-fishes time ive tract structure and function has been expanded.
line (Figure 34.11). The treatment of the taxonomic status of
Chapter 44 Information on the types of circulatory systems
hagfishes and lampreys, evolution of the jaw (Figure 34.14),
and types of blood vessels has been consolidated. Details on
and H. sapiens migration (Figure 34.48) also include the most
surface tension and lung elasticity have been removed, while
recent data available. The emphasis on the adaptive signifi-
new content on countercurrent exchange in fish gills has been
cance of the amniotic egg has changed from watertightness
added.
to increased size and support. Emphasis in the discussion of
viviparity has changed to the adaptive advantage of embryo Chapter 45 The chapter and section introductions have been
portability and temperature control. The recent analysis of rewritten to introduce a comparative context and to make the
Ardipithecus ramidus as the first hominin, with data on esti- neuron-to-systems chapter organization more transparent.
mated body mass and braincase volume, has been included. New content on interspecific variation in nervous systems
has been added.
Chapter 35 The material on HIV phylogeny has been moved
to the section on emerging viruses. Chapter 46 The chapter has been shortened and its focus
sharpened by the removal of nonessential information. A new
Canadian Research box explores why large numbers of mi-
Unit 7 How Plants Work gratory bats are killed by turbines at wind farms in southern
Chapter 36 Surface-area-to-volume ratios have been added as Alberta.
a theme in root and shoot systems. New information on con- Chapter 47 New material on EPO abuse in athletes has been
tractile roots in Ficus and bulbs is incorporated into this chapter. included.
Chapter 37 New content on aquaporins and the transmem- Chapter 48 A new Canadian Issues box describes Canada’s
brane route to root xylem has been added, and coverage of why Assisted Human Reproduction Act. The section on sperm com-
air has such low water pressure potential has been expanded. petition includes new data from experiments on seed beetles.
New Canadian research is included that considers the adaptive
Chapter 49 The discussion of the V regions of BCRs and anti-
value of plastic responses of bluebunch wheatgrass under the
bodies and recombination in BCR/TCR genes has been sim-
increasingly dry conditions that climate change is bringing to
plified. New content on autoimmune disorders and diseases
many regions in the Canadian interior.
associated with immunosuppression, allergies, and immuno-
Chapter 38 The description of nitrogen fixation has been deficiency diseases has been added. The discussion of vaccina-
clarified. tion has been expanded.
Chapter 39 A new Canadian Research box explores how plant
signalling networks influence growth in plants. Figure 39.8 on
the acid-growth hypothesis has been redesigned, and the dis-
Unit 9 Ecology
cussion of polar auxin transport is simplified. New commen- Chapter 50 A new Canadian Research box explores whether
tary on the role of brassinosteroids in growth regulation and sockeye salmon stocks vary in their ability to cope with

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 increasing temperatures during migration. New information to explain the latitudinal gradient in species richness has been
on the importance of nutrient availability in aquatic ecosystems, expanded and clarified. Simon Fraser University paleontologist
with details on lake turnover and ocean upwelling, is included. Bruce Archibald and his colleagues have found a novel way to
A new section on the Wallace line has also been added. investigate the role of climatic factors in producing latitudinal
Chapter 51 The content in this chapter has been completely gradient in species richness, as explained in the new Canadian
reorganized to increase cohesiveness. It is presented as a ser- Research box.
ies of questions in behavioural ecology, with each question Chapter 54 The chapter was rewritten and reorganized to
addressed at the proximate and ultimate levels with separate sharpen its focus on human impacts. Sections on trophic cas-
case studies. Material on modes of learning, innate behaviour, cades and biomagnification have been added, as have recent
bat–moth interactions, sex change in wrasses, and acous- data on human appropriation of NPP, sources of nutrient
tic and visual signaling in red-winged blackbirds has been gain and loss, and the impact of ocean acidification on coral
trimmed. New content on animal eusociality and on child growth.
abuse in humans has been added. Chapter 55 New content on the impact of global climate
Chapter 52 Discussion of the hare–lynx cycle field experi- change and a new section on ways to preserve biodiversity are
ment has been reorganized for clarity, with new supporting now included. Two new boxes on quantitative methods have
“Results” data added to accompanying Figure 52.13. been added: one on estimating species numbers and species
Chapter 53 New content has been added on species richness losses and the other on population viability analysis. Discus-
and resistance of communities to invasion, the use of predators sion of Canada’s Species at Risk Act (SARA) has been moved
or parasites as biocontrol agents, and character displacement in to this chapter and is discussed in Canadian Issues 55.1. Re-
finches. The discussion of succession in Glacier Bay is reorgan- vised Canadian Issues 55.2 considers the fate of polar bears
ized and simplified. The discussion of alternative hypotheses in a warming Arctic.

Supplements

Instructor Resources ⦁ Canadian case studies picking up ideas raised in the Canadian
Research and Issues boxes are available to explore these stud-
All instructor resources are available on a flash drive (978-0- ies further and investigate how to apply them in the world.
321-72911-8) and can also be downloaded from the instructor Teaching notes include an Introduction, Learning Objectives,
resources area of MasteringBiology. Student Misconceptions, Classroom Management, Supple-
⦁ The entire textbook illustration program is available in JPEG mentary Questions, and References.
format with and without labels. Illustrations have been indi- ⦁ The Instructor Guide includes lecture outlines, active-learning
vidually enhanced for optimal in-class projection. lecture activities, answers to end-of-chapter questions, and in-
⦁ The entire illustration program is also available with editable novative material to help motivate and engage students.
labels and leaders in chapter-by-chapter Microsoft Power- ⦁ Test Bank and Computerized Test Bank questions are ranked
®
Point presentations. according to Bloom’s Taxonomy. Improved TestGen soft- ®
⦁ A second set of PowerPoint presentations offers lecture out- ware makes assembling tests much easier. The Test Bank is
lines for each chapter, augmented by key text illustrations and ®
also available in Microsoft Word format.
hyperlinks to animations.
⦁ A third set of PowerPoint presentations is layered to allow Student Resources
select key figures to be presented in a step-by-step manner. ⦁ The eText addresses the changing needs of students and in-
⦁ In-class active lecture questions correlated by chapter can be structors. Found within MasteringBiology, this electronic
used with any classroom response system and are available in version of the text links directly to animations, quizzes, and
PowerPoint format. videos.

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 ⦁ The Study Guide (978-0-321-82868-2) presents a breakdown of pre-quizzes and post-quizzes to test student’s understanding of
key biological concepts, and helps students focus on the fun- biology’s dynamic processes and concepts.
damentals of each chapter. It is designed in two parts to help
DISCOVERY VIDEOS Brief videos from the Discovery™ Channel
students study more effectively. Part I is intended as a “survival
on 29 different biology topics are available for student viewing
guide,” and Part II explores the material in the textbook, chap-
along with a corresponding video quiz.
ter by chapter.
VIDEOS Additional molecular and microscopy videos provide
MasteringBiology vivid images of processes of the cell.
Students who purchase a new copy of the text receive free access BIOSKILLS BioSkills (in Appendix B) provide background on
®
to MasteringBiology (www.masteringbiology.com), which con-
tains valuable videos, animations, and practice quizzes to help
key skills and techniques for introductory biology students. New
to the Second Canadian Edition are online questions that give
students learn and prepare for exams. students practice building their skill set.

THE BIG PICTURE New to the Second Canadian Edition, The Big GRAPHIT! Graphing tutorials show students how to plot, inter-
Pictures are interactive concept maps based on seven overarch- pret, and critically evaluate real data.
ing topics in biology that help students synthesize information Chapter 1
across broad concepts and not get lost in the details.
⦁ An Introduction to Graphing
Macromolecules (Chapters 2–6)
Chapter 50
⦁ How monomers are used to make macromolecules
⦁ Animal Food Production Efficiency and Food Policy
⦁ How macromolecules can be classified
⦁ Atmospheric CO2 and Temperature Changes
Energy (Chapters 9 and 10)
Chapter 52
⦁ How photosynthesis yields sugar
⦁ Age Pyramids and Population Growth
⦁ How cellular respiration yields ATP
⦁ How photosynthesis relates to cellular respiration Chapter 53

Genetic Information (Chapters 12–18) ⦁ Species Area Effect and Island Biogeography

⦁ How genes are expressed Chapter 55

⦁ How genetic information is copied and transmitted ⦁ Forestation Change

⦁ Global Fisheries and Overfishing
⦁ How genetic information changes
⦁ Municipal Solid Waste Trends in the United States
Evolution (Chapters 24–27)
⦁ Global Freshwater Resources
⦁ How species evolve
⦁ Prospects for Renewable Energy
⦁ How species form the tree of life
⦁ Global Soil Degradation
How Vascular Plants Work (Chapters 36–39)
⦁ How vascular plants capture light energy and take up CO2 WORD STUDY TOOLS New to the Second Canadian Edition are
⦁ How vascular plants obtain water and inorganic nutrients Latin and Greek root word flash cards to help students practise
the language of biology. In addition, an audio glossary provides
⦁ How vascular plants respond to hostile organisms
correct pronunciation to help students learn key terms intro-
How Humans Work (Chapters 41–47, 49) duced in the book.
⦁ How humans obtain nutrients and maintain homeostasis
CUMULATIVE TEST Every chapter offers 20 Practice Test ques-
⦁ How humans recognize and respond to hostile organisms tions that students can pool from different chapters into a Cumu-
Ecology (Chapters 50–55) lative Test to simulate a practice exam.
⦁ How organisms interact in their environment RSS FEEDS Real Simple Syndication directly links breaking news
⦁ How energy flows and nutrients cycle through ecosystems from four important sources: NPR (National Public Radio), Sci-
entific American, Science Daily News, and BioScience. Current
BIOFLIX™ BioFlix are 3-D movie-quality animations with care- articles reinforce the dynamic nature of science in our daily lives.
fully constructed student tutorials, labelled slide shows, study
sheets, and quizzes, that bring biology to life. eTEXT The eText of Biological Science, Second Canadian Edi-
tion, is available online 24/7 for students’ convenience. New an-
WEB ACTIVITIES Web Activities help students learn biologi- notation, highlighting, and bookmarking tools allow students to
cal concepts via simple, cartoon-style animations and contain personalize the material for efficient review.

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 STUDY ON THE GO At the end of every chapter, students will and link to a website containing Biological Science’s Study on the
find a QR code (a.k.a. quick response code) that links to Study Go content.
on the Go mobile content. Students can access text-specific re-
ScanLife
sources, including quizzes and flashcards, through their smart-
http://getscanlife.com
phones, allowing them to study whenever and wherever they
wish! NeoReader
Students can go to one of the sites below to see how to down- http://get.neoreader.com
load a free app to their smartphone that facilitates access to these
QuickMark
resources. Once the app is installed, the phone will scan the code
http://www.quickmark.com.tw

MASTERINGBIOLOGY MEDIA AT A GLANCE

BIOFLIX WEB ACTIVITIES VIDEOS BIOSKILLS
1 Biology and the Tree Artificial Selection; Introduction The Metric System;
of Life to Experimental Design Reading Graphs; Reading
a Phylogenetic Tree; Some
Common Latin and Greek
Roots Used in Biology

Unit 1 The Molecules of Life

2 Water and Carbon: The The Properties of Water Reading Chemical Structures;
Chemical Basis of Life Using Logarithms; Making
Concept Maps; Reading
Graphs

3 Protein Structure and Condensation and Hydrolysis An Idealized Alpha Helix (A);
Function Reactions; Activation Energy An Idealized Alpha Helix (B);
and Enzymes An Idealized Beta-Pleated
Sheet (A); An Idealized
Beta-Pleated Sheet (B)

4 Nucleic Acids and the Structure of RNA and DNA Stick Model of DNA; Surface Separating and Visualizing
RNA World Model of DNA Molecules; Biological
Imaging: Microscopy and
X-Ray Crystallography

5 An Introduction to Carbohydrate Structure and

Carbohydrates Function

6 Lipids, Membranes, Membrane Diffusion and Osmosis; Space-Filling Model of Biological Imaging:
and the First Cells Transport Membrane Transport Cholesterol; Stick Model of Microscopy and X-Ray
Proteins Cholesterol; Space-Filling Model Crystallography; Separating
of Phosphatidylcholine; Stick and Visualizing Molecules
Model of a Phosphatidylcholine

Unit 2 Cell Structure and Function

7 Inside the Cell Tour of an Animal Cell; Transport into the Nucleus; Confocal vs. Standard Separating Cell Components
Tour of a Plant Cell A Pulse-Chase Experiment Fluorescence Microscopy; by Centrifugation; Biological
Cytoplasmic Streaming; Imaging: Microscopy and
Crawling Amoeba X-Ray Crystallography;
Separating and Visualizing
Molecules

8 Cell–Cell Interactions Connexon Structure Separating and Visualizing

Molecules

9 Cellular Respiration Cellular Respiration Redox Reactions; Space-Filling Model of ATP

and Fermentation Glucose Metabolism (adenosine triphosphate);
Stick Model of ATP (adenosine
triphosphate)

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 MASTERINGBIOLOGY MEDIA AT A GLANCE (continued)
BIOFLIX WEB ACTIVITIES VIDEOS BIOSKILLS

10 Photosynthesis Photosynthesis Chemiosmosis; Photosynthesis; Space-Filling Model of

Strategies for Carbon Fixation Chlorophyll

11 The Cell Cycle Mitosis The Phases of Mitosis; Four Mitosis Separating and Visualizing
Phases of the Cell Cycle Molecules; Cell and Tissue
Culture Methods

Unit 3 Gene Structure and Expression

12 Meiosis Meiosis Meiosis; Mistakes in Meiosis Combining Probabilities; Using

Statistical Tests and Interpreting
Standard Error Bars

13 Mendel and Mendel’s Experiments; The Model Organisms; Combining

the Gene Principle of Independent Probabilities; Reading Graphs
Assortment

14 DNA and the Gene: DNA Replication DNA Synthesis Separating Cell Components
Synthesis and Repair by Centrifugation; Cell and
Tissue Culture Methods; Using
Logarithms; Reading Graphs

15 How Genes Work The One-Gene One-Enzyme

Hypothesis; The Triplet Nature
of the Genetic Code

16 Transcription, RNA Protein Synthesis RNA Synthesis; Synthesizing A Stick-and-Ribbon Rendering

Processing, and Proteins of a tRNA
Translation

17 Control of Gene The lac Operon Cartoon Model of the lac

Expression in Bacteria Repressor from E. coli

18 Control of Gene Transcription Initiation in Cartoon Model of the DNA- Biological Imaging:
Expression in Eukaryotes Binding Portion of TATA-Box Microscopy and X-Ray
Eukaryotes Binding Protein Interacting with Crystallography; Separating
DNA; Cartoon Model of the GAL4 and Visualizing Molecules
Transcription Factor from the
Yeast S. cerevisiae

19 Analyzing and Producing Human Growth Cartoon Model of the BamH1a Separating and Visualizing
Engineering Genes Hormone; The Polymerase Endonuclease Molecules
Chain Reaction

20 Genomics Human Genome Sequencing Model Organisms; Using

Strategies Logarithms

Unit 4 Developmental Biology

21 Principles of Early Pattern Formation in A Cartoon and Stick Model Model Organisms; Cell and
Development Drosophila of the Homeodomain of the Tissue Culture Methods
Engrailed Protein from
Drosophila Interacting
with DNA

22 An Introduction to Early Stages of Animal

Animal Development Development

23 An Introduction to Model Organisms

Plant Development

Unit 5 Evolutionary Processes and Patterns

24 Evolution by Natural Natural Selection for Antibiotic Reading a Phylogenetic Tree;

Selection Resistance Model Organisms; Reading
Graphs

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 MASTERINGBIOLOGY MEDIA AT A GLANCE (continued)
BIOFLIX WEB ACTIVITIES VIDEOS BIOSKILLS
25 Evolutionary Mechanisms of Evolution The Hardy–Weinberg Combining Probabilities;
Processes Principle; Three Modes of Using Statistical Tests and
Natural Selection Interpreting Standard Error
Bars; Reading Graphs

26 Speciation Allopatric Speciation; Speciation Reading a Phylogenetic Tree

by Changes in Ploidy

27 Phylogenies and the Adaptive Radiation Reading a Phylogenetic Tree

History of Life

Unit 6 The Diversification of Life

28 Bacteria and Archaea The Tree of Life Reading a Phylogenetic Tree;

Model Organisms

29 Protists Alternation of Generations A Crawling Amoeba Biological Imaging:

in a Protist Microscopy and X-Ray
Crystallography; Model
Organisms

30 Green Algae and Land Plant Evolution and the

Plants PhylogeneticTree

31 Fungi Life Cycle of a Mushroom

32 An Introduction to The Architecture of Animals

Animals

33 Protostome Animals Protostome Diversity Model Organisms

34 Deuterostome Deuterostome Diversity

Animals

35 Viruses The HIV Replicative Cycle Biological Imaging:

Microscopy and X-Ray
Crystallography; Separating
and Visualizing Molecules

Unit 7 How PlantsWork

36 Plant Form and Plant Growth

Function

37 Water and Sugar Water Transport in Plants Solute Transport in Plants Plasmolysis of Plant Cells
Transport in Plants

38 Plant Nutrition Soil Formation and Nutrient

Uptake

39 Plant Sensory Sensing Light; Plant Hormones; Cell and Tissue Culture
Systems, Signals, and Plant Defences Methods
Responses

40 Plant Reproduction Reproduction in Flowering

Plants; Fruit Structure and
Development

Unit 8 How Animals Work

41 Animal Form and Surface Area/Volume Using Logarithms

Function Relationships; Homeostasis

42 Water and Electrolyte The Mammalian

Balance in Animals Kidney

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 MASTERINGBIOLOGY MEDIA AT A GLANCE (continued)
BIOFLIX WEB ACTIVITIES VIDEOS BIOSKILLS

43 Animal Nutrition Homeostasis: Regulating The Digestion and Absorption Biological Imaging:
Blood Sugar of Food; Understanding Microscopy and X-Ray
Diabetes Mellitus Crystallography; Separating
and Visualizing Molecules

44 Gas Exchange and Gas Exchange Gas Exchange in the Lungs and
Circulation Tissues; The Human Heart

45 Electrical Signals in How Neurons Work; How Membrane Potentials; Action The Acetylcholine Receptor Using Logarithms
Animals Synapses Work Potentials

46 Animal Sensory Muscle Contraction The Vertebrate Eye; Structure The Acetylcholine Receptor
Systems and and Contraction of Muscle
Movement Fibres

47 Chemical Signals in Endocrine System Anatomy; Cartoon Model of the DNA Separating Cell Components
Animals Hormone Actions on Binding Motif of a Zinc Finger by Centrifugation
Target Cells Transcription Factor Binding to
DNA

48 Animal Reproduction Human Gametogenesis; Using Logarithms; Reading a

Human Reproduction Phylogenetic Tree

49 The Immune System The Inflammatory Response; Chemotaxis of a Neutrophil

in Animals The Adaptive Immune
Response

Unit 9 Ecology

50 An Introduction to Tropical Atmospheric

Ecology Circulation

51 Behavioural Ecology Homing Behaviour in Digger

Wasps

52 Population Ecology Population Ecology Modelling Population Growth;

Human Population Growth and
Regulation

53 Community Ecology Life Cycle of a Malaria Parasite;

Succession

54 Ecosystems The Carbon Cycle The Global Carbon Cycle

55 Biodiversity and Habitat Fragmentation Using Logarithms

Conservation Biology

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 Preface to Students: How to Use This Book

Focus on the Gold Thread

These red-tailed hawk chicks are

being fed by a parent. In three
years they will have grown and
had chicks of their own. Likewise
the pine tree they are nesting in
is also reproducing using seeds
within its pine cones. The birds,
the tree, and the organisms
present but too small to see
in this photograph all need to
produce offspring. Despite the
great diversity of life, all living
creatures share this and other
common properties.

Biology and the Tree of Life

1
I n essence, biological science is a search for ideas and observations that unify our
understanding of the diversity of life, from bacteria living in rocks a mile under-
ground to hedgehogs and humans. Chapter 1 is an introduction to this search.
The goals of this chapter are to introduce the nature of life and explore how biologists
go about studying it. The chapter also introduces themes that will resonate throughout
KEY CONCEPTS
Organisms obtain and use energy, are
made up of cells, process information,
replicate, and, as populations, evolve.
Key Concepts
Start with Key
this book: (1) analyzing how organisms work at the molecular level, (2) understanding The cell theory proposes that all Concepts on the
organisms in terms of their evolutionary history, and (3) helping you learn to think like organisms are made of cells and that all
a biologist. cells come from pre-existing cells. first page of every
Let’s begin with what may be the most fundamental question of all: What is life? The theory of evolution by natural
selection maintains that species change chapter. Read these
through time because individuals with
1.1 What Does It Mean to Say certain heritable traits produce more gold key points first
That Something Is Alive? offspring than other individuals do.
to familiarize yourself
A phylogenetic tree is a graphical
An organism is a life form—a living entity made up of one or more cells. Although
there is no simple definition of life that is endorsed by all biologists, most agree that
representation of the evolutionary
relationships between species. These
with the chapter’s
organisms share a suite of five fundamental characteristics: relationships can be estimated by
analyzing similarities and differences in
big ideas.
⦁ Energy To stay alive and reproduce, organisms have to acquire and use energy.
traits. Species that share distinctive traits
are closely related and are placed close to
⦁ Cells Organisms are made up of membrane-bound units called cells. A cell’s mem- each other on the tree of life.
brane regulates the passage of materials between exterior and interior spaces. Biologists ask questions, generate
⦁ Information Organisms process hereditary or genetic information, encoded in hypotheses to answer them, and design
units called genes, along with information they acquire from the environment. Right experiments or make observations that
now, cells throughout your body are using genetic information to make the mol- test the predictions made by competing
ecules that keep you alive; your eyes and brain are decoding information on this page hypotheses.
that will help you learn some biology.

When you see this checkmark, stop and test yourself. Answers are available in Appendix A. 1

MORE! Bulleted Lists Gold Highlighting Gold Key

Take note of bulleted lists Watch for important Material related
that “chunk” information information highlighted in to Key Concepts
and ideas. This will gold. Gold highlighting is will be signalled
help you manage the always a signal to slow down with a gold key.
information that you are learn- and pay special attention.
ing in the course.
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 SUMMARY TABLE 3.1 Protein Structure Summary Tables
Level Description Stabilized by Example: Hemoglobin
Primary The sequence of amino acids in a Peptide bonds Summary Tables pull information
Gly Ser Asp Cys
polypeptide
together in a compact format that is
Secondary Formation of α-helices and Hydrogen bonding between groups easy to review and synthesize.
β-pleated sheets in a polypeptide along the peptide-bonded backbone; One α-helix
thus, depends on primary structure

Tertiary Overall three-dimensional Bonds and other interactions

shape of a polypeptide (includes between R-groups, or between One of
hemoglobin’s
contribution from secondary R-groups and the peptide-bonded
subunits
structures) backbone; thus, depends on primary
structure

Quaternary Shape produced by combinations Bonds and other interactions Hemoglobin,

of polypeptides (thus, combinations between R-groups, and between which consists
of tertiary structures) peptide backbones of different of four
polypeptides; thus, depends on polypeptide
primary structure subunits

CHECK YOUR UNDERSTANDING Check Your Understanding

If you understand that . . .
The gold half of the Check Your Understanding boxes
⦁ Natural selection occurs when heritable variation in certain
summarizes important information from the section you
traits leads to improved success in reproduction. just read. Stop and ask yourself: Do I really understand every
⦁ Evolution is a change in the characteristics of a population bullet point?
over time.

You should be able to . . .

On the graph you just analyzed, describe the average kernel
protein content over time in a maize population where no
selection occurred.
Answers are available in Appendix A.

Summary of Key Concepts

The succinct Summary of Key
Concepts reviews important concepts
in short, manageable bullet points.

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 Practise with the Blue Thread.
Maximum speed of reaction
Rate of product formation

Substrate concentration
FIGURE 3.23 Kinetics of an Enzyme-Catalyzed Reaction. The
general shape of this curve is characteristic of enzyme-catalyzed Drawing Exercises
reactions.
Some caption questions and exercises
EXERCISE Label the parts of the graph that represent where
contain artwork from the textbook that
(1)
concentration and (2) most or all of the active sites present are you will be asked to draw on or modify.
occupied.

NEW! Suggested Answers

Caption Questions and Exercises Suggested answers for the Blue Thread
These challenge you to critically examine the Questions and Exercises are provided in
information in a figure or table—not just absorb it. Appendix A.

“You Should Be Able To”

Exercises
Text passages flagged with
blue type and the words “You Evolution occurs when heritable variation leads to differen-
should be able to” offer exercises tial success in reproduction. If you understand this concept,
on concepts that professors and you should be able to modify Figure 1.3 to show what hap-
students have identified as most pened when the same researchers selected individuals with the
difficult. These are the topics most lowest kernel protein content to be the parents of the next
students struggle with on exams. generation.

CHECK YOUR UNDERSTANDING

If you understand that . . .
Check Your Understanding
⦁ Natural selection occurs when heritable variation in certain
traits leads to improved success in reproduction. The blue half of the Check Your
⦁ Evolution is a change in the characteristics of a population Understanding boxes asks you to do
over time.
something with the information in the top
You should be able to . . . half. If you can’t complete these exercises,
On the graph you just analyzed, describe the average kernel
protein content over time in a maize population where no
go back and reread that section of the
selection occurred. chapter.
Answers are available in Appendix A.

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 Chapter Summaries
End-of-chapter “You should be able to” problems or exercises
help you review the key concepts declared in the gold thread.

Canadian Research 3.1 Designing New Proteins

Proteins are such useful macromolecules that scientists have used whereupon the enzyme portion will cut the hybrid protein in two,
them as tools in experiments for years. For example, the protein releasing the antimicrobial portion to fight the bacteria. Yada and
that makes jellyfish glow, green fluorescent protein, is used by his colleagues think that this hybrid protein may be used one day Think About It Questions
biologists to make different parts of cells visible with microscopes in either people or agriculturally important plants and animals.
(see BioSkills 10 in Appendix B). In fact, rather than rely on na-
ture to provide proteins with a desired activity, some scientists
Canadian Research and Canadian
SOURCE: Bryksa, B. C., Horimoto, Y., & Rada, R. Y. (2010). Rational redesign
have begun to engineer new proteins themselves.
Brian Bryksa, Yasumi Horimoto, and Rickey Yada from the
of porcine pepsinogen containing an antimicrobial peptide. Protein Engineering,
Design, & Selection, 23, 711–719.
Issues boxes each end with a
University of Guelph have made such a protein. It is a combina- question that will test or expand
tion of a cow protein that kills harmful bacteria and a pig enzyme Think About It: Why might this hybrid protein be better at
that works in the stomach and cuts up other proteins. The new treating infections than the antimicrobial protein by itself? on your understanding of an
protein is designed to travel to the location of a bacterial infection
important concept.

Steps to Understanding
Analyze Evaluate Synthesize End-of-chapter questions are scaled along Bloom’s
Taxonomy.

Apply TEST YOUR KNOWLEDGE

Explain Begin by testing your knowledge of new facts.

Remember
TEST YOUR UNDERSTANDING
Once you’re confident in your knowledge of the
Bloom’s Taxonomy material, demonstrate your understanding by
Bloom’s Taxonomy categorizes six levels answering the Test Your Understanding questions.
of learning competency. The Blue Thread
Questions and Exercises in the textbook APPLYING CONCEPTS TO NEW SITUATIONS
test on the higher levels of the scale—
Explain, Apply, Analyze, Evaluate, and Challenge yourself even further by applying your
Synthesize—to help you develop critical understanding of the concepts to new situations.
thinking skills and prepare you for exams.

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 Keep sight of the big picture.

Concept maps help you to keep sight of “big picture” relationships

among biological concepts.

NEW! Big Picture

Concept Maps
Seven remarkable Big
Picture concept maps
help you synthesize
information across the
chapters on energy,
genetics, evolution,
and ecology.

Check Your
Understanding
Check your under-
standing of these big
picture relationships
by answering the Blue
Thread Questions.

MasteringBiology®
Your professor may assign
interactive Big Picture
concept map exercises at
www.masteringbiology.com.

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 Learn to think like a scientist. Here’s how.

A unique emphasis on the process of scientific discovery and

experimental design teaches you how to think like a scientist
as you learn fundamental biology concepts.
Experiment Boxes EXPERIMENT

QUESTION: Why is the distribution of adult

Study Experiment Boxes Chthamalus restricted to the upper intertidal zone?

to help you understand how HYPOTHESIS: Adult Chthamalus are competitively excluded from the
lower intertidal zone.
experiments are designed NULL HYPOTHESIS: Adult Chthamalus do not thrive in the physical
conditions of the lower intertidal zone.
and give you practice
MasteringBiology ®
interpreting data.
EXPERIMENTAL SETUP:

Chthamalus in
www.masteringbiology.com upper intertidal zone
1. Transplant rocks
containing young
Chthamalus to

NEW! Experimental Inquiry Tutorials Mean tide level lower intertidal zone.

Semibalanus
in lower
Experimental Inquiry Tutorials based on some of biology’s most intertidal
zone 2. Let Semibalanus
seminal experiments can be found on www.masteringbiology.com. colonize the rocks.

Your instructor may assign these. They will give you practice analyzing
the experimental design and data, and help you understand reasoning
that led scientists from the data they collected to their conclusions. 3. Remove
Semibalanus from
half of each rock.
Some of the topics include: Monitor survival
of Chthamalus on
both sides.
• The Process of Science
Chthamalus
Chthamalus + Semibalanus
• Engelmann’s Photosynthesis and Wavelengths of Light
• Morgan’s Cross with White‐Eyed Males
PREDICTION: Chthamalus will survive better in the absence of
• Meselson‐Stahl’s Semiconservative Replication Semibalanus.
PREDICTION OF NULL HYPOTHESIS: Chthamalus survival will be low
• Steinhardt et al and Hafner et al’s Polyspermy and the same in the presence or absence of Semibalanus.

RESULTS:
• Grant’s Changes in Finch Beak Size 80
Percent age survival

• Went’s Phototropism and Auxin Distribution

of Chthamalus

60

• Coleman’s Obesity Gene 40

• Connell’s Competition in Barnacles 20

0
• Bormann, Likens et al’s Nutrient Cycling in Hubbard Brook Forest Competitor
absent
Competitor
present

CONCLUSION: Semibalanus is competitively excluding Chthamalus

from the lower intertidal zone.

FIGURE 53.6 Experimental Evidence for Competitive Exclusion.

SOURCE: Connell, J. H. (1961). The influence of interspecific competition and other
factors on the distribution of the barnacle Chthamalus stellatus. Ecology, 42, 710–723.
QUESTION Why was it important to carry out both treatments on
the same rock? Why not use separate rocks?

NEW! Source Citations

Each Experiment Box now cites the original
research paper, encouraging you to extend
your learning by exploring the primary
literature.

NEW! Experiment Box Questions

Each Experiment Box now includes a
question that asks students to analyze the
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 Bacteria NEW! Redesigned Phylogenetic Trees
Archaea
Practice “tree thinking” using these newly
AMOEBOZOA redesigned phylogenetic trees. Their
Lobose amoebae
U‐shaped, top‐to‐bottom format is consistent
Cellular slime moulds
with the way such trees are most commonly

UNIKONTA
Plasmodial slime
moulds depicted in the scientific literature.
OPISTHOKONTA
Fungi

Choanoflagellates

All eukaryotes Animals

are protists
except for the
fungi, animals,
EXCAVATA Expanded BioSkills Appendix
Parabasalids
EUKARYOTES and land plants
Diplomonads

Euglenids

Kinetoplastids

PLANTAE
Build skills that will be important to your success in future
Glaucophyte algae courses. At relevant points in the text, you’ll find references
Red algae to the expanded BioSkills Appendix that will help you learn
Green algae
Green and practice the following foundational skills:
plants
BIKONTA

Land plants

RHIZARIA • NEW! The Metric System

Foraminifera
• Reading Graphs
Chlorarachniophytes
• Reading a Phylogenetic Tree
ALVEOLATA
Ciliates • NEW! Some Common Latin and Greek Roots Used in
Biology
CHROMALVEOLATA

Dinoflagellates

Apicomplexa • Using Statistical Tests and Interpreting Standard Error Bars

STRAMENOPILA
Oomycetes
• Reading Chemical Structures
Diatoms • Using Logarithms
Brown algae • Making Concept Maps
• Separating and Visualizing Molecules
• Biological Imaging: Microscopy and X‐Ray Crystallography
• NEW! Separating Cell Components by Centrifugation
• NEW! Cell Culture Methods
• Combining Probabilities
• NEW! Model Organisms

PROCESS: PULMONARY CIRCULATION SYSTEMIC CIRCULATION

Informative Figures
1. Blood returns to 4. Blood returns to left
heart from body,
enters right atrium.
atrium from lungs.
Think through complex
5. Blood enters left
2. Blood enters
right ventricle.
Superior 6
Aorta ventricle. biological processes
vena cava
6. Blood is pumped
3. Blood is pumped
3 Pulmonary
veins from left ventricle with figures that clearly
from right ventricle to body.
to lungs. Pulmonary
artery 1. Blood returns to define concepts.
4. Blood returns to left heart from body,
Right Aortic 4 enters right atrium.
atrium from lungs. atrium Pulmonary valve
1 Left
valve
atrium
Right Left
atrioventricular atrioventricular
(AV) valve (AV) valve
Right Left
Inferior 2 ventricle 5 ventricle
vena cava

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