Campbell Biology, 3rd Canadian Edition PDF by Lisa A. Urry, Michael L Cain, Steven A Wasserman, Peter V Minorsky, Jane B Reece, Neil A Campbell, Fiona Rawle, Dion Durnford, Chris Moyes and Kevin Scott


Campbell Biology, Third Canadian Edition

By Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Jane B. Reece, Neil A. Campbell, Fiona Rawle, Dion Durnford, Chris Moyes and Kevin Scott

Campbell Biology, Third Canadian Edition

Detailed Contents:

1 Evolution, the Themes of Biology, and Scientific Inquiry 1 Inquiring About Life 1

CONCEPT 1.1 The study of life reveals common themes 3

Theme: New Properties Emerge at Successive Levels of Biological Organization 3

Theme: Life’s Processes Involve the Expression and Transmission of Genetic Information 6

Theme: Life Requires the Transfer and Transformation of Energy and Matter 8

Theme: From Molecules to Ecosystems, Interactions Are Important in Biological Systems 9

CONCEPT 1.2 The Core Theme: Evolution accounts for the unity and diversity of life 11

Classifying the Diversity of Life 11

The Tree of Life 15

CONCEPT 1.3 In studying nature, scientists make observations and form and test hypotheses 16

Exploration and Observation 16

Forming and Testing Hypotheses 17

The Flexibility of the Scientific Process 18

A Case Study in Scientific Inquiry: Investigating Coat Colouration

in Mouse Populations 18

Theories in Science 21

CONCEPT 1.4 Science benefits from a cooperative approach and diverse viewpoints 22

Building on the Work of Others 22

Science, Technology, and Society 23

The Value of Diverse Viewpoints in Science 24


Interview: Roberta Hamme 27

2 The Chemical Context of Life 30

A Chemical Connection to Biology 30

CONCEPT 2.1 Matter consists of chemical elements in pure form and in combinations called compounds 31

Elements and Compounds 31

The Elements of Life 31

Case Study: Evolution of Tolerance to Toxic Elements 32

CONCEPT 2.2 An element’s properties depend on the structure of its atoms 32

Subatomic Particles 32

Atomic Number and Atomic Mass 33

Isotopes 33

The Energy Levels of Electrons 34

Electron Distribution and Chemical Properties 36

Electron Orbitals 37

CONCEPT 2.3 The formation and function of molecules depend on chemical bonding between atoms 38

Covalent Bonds 38

Ionic Bonds 40

Weak Chemical Interactions 41

Molecular Shape and Function 42

CONCEPT 2.4 Chemical reactions make and break chemical bonds 43

3 Water and Life 47

The Molecule That Supports All of Life 47

CONCEPT 3.1 Polar covalent bonds in water molecules result in hydrogen bonding 48

CONCEPT 3.2 Four emergent properties of water contribute

to Earth’s suitability for life 48

Cohesion of Water Molecules 48

Moderation of Temperature by Water 49

Floating of Ice on Liquid Water 51

Water: The Solvent of Life 51

Possible Evolution of Life on Other Planets 53

CONCEPT 3.3 Acidic and basic conditions affect living organisms 54

Acids and Bases 54

The pH Scale 55

Buffers 56

Acidification: A Threat to Our Oceans 56

4 Carbon and the Molecular

Diversity of Life 60

Carbon: The Backbone of Life 60

CONCEPT 4.1 Organic chemistry is the study of carbon compounds 61

Organic Molecules and the Origin of Life on Earth 61

CONCEPT 4.2 Carbon atoms can form diverse molecules by bonding to four other atoms 62

The Formation of Bonds with Carbon 63

Molecular Diversity Arising from Variation in Carbon

Skeletons 64

CONCEPT 4.3 A few chemical groups are key to molecular function 67

The Chemical Groups Most Important in the Processes of Life 67

ATP: An Important Source of Energy for Cellular Processes 67

The Chemical Elements of Life: A Review 67

5 The Structure and Function of Large Biological Molecules 72

The Molecules of Life 72

CONCEPT 5.1 Macromolecules are polymers, built from monomers 73

The Synthesis and Breakdown of Polymers 73

The Diversity of Polymers 73

CONCEPT 5.2 Carbohydrates serve as fuel and building material 74

Sugars 74

Polysaccharides 76

CONCEPT 5.3 Lipids are a diverse group of hydrophobic molecules 78

Fats 78

Phospholipids 80

Steroids 81

CONCEPT 5.4 Proteins include a diversity of structures, resulting in a wide range of functions 81

Amino Acid Monomers 81

Polypeptides (Amino Acid Polymers) 84

Protein Structure and Function 84

CONCEPT 5.5 Nucleic acids store, transmit, and help express hereditary information 90

The Roles of Nucleic Acids 90

The Components of Nucleic Acids 90

Nucleotide Polymers 91

The Structures of DNA and RNA Molecules 92

CONCEPT 5.6 Genomics and proteomics have transformed biological inquiry and applications 93

DNA and Proteins as Tape Measures of Evolution 93


Interview: Jason Treberg 99

6 A Tour of the Cell 103

The Fundamental Units of Life 103

CONCEPT 6.1 Biologists use microscopes and the tools of biochemistry to study cells 104

Microscopy 104

Cell Fractionation 106

CONCEPT 6.2 Eukaryotic cells have internal membranes that compartmentalize their functions 107

Comparing Prokaryotic and Eukaryotic Cells 107

A Panoramic View of the Eukaryotic Cell 109

CONCEPT 6.3 The eukaryotic cell’s genetic instructions are housed in the nucleus and carried out by the

ribosomes 112

The Nucleus: Information Central 112

Ribosomes: Protein Factories 112

CONCEPT 6.4 The endomembrane system regulates protein traffic and performs metabolic functions 114

The Endoplasmic Reticulum: Biosynthetic Factory 114

The Golgi Apparatus: Shipping and Receiving Centre 116

Lysosomes: Digestive Compartments 117

Vacuoles: Diverse Maintenance Compartments 118

The Endomembrane System: A Review 118

CONCEPT 6.5 Mitochondria and chloroplasts change energy from one form to another 119

The Evolutionary Origins of Mitochondria and Chloroplasts 119

Mitochondria: Chemical Energy Conversion 120

Chloroplasts: Capture of Light Energy 120

Peroxisomes: Oxidation 122

CONCEPT 6.6 The cytoskeleton is a network of fibres that organizes structures and activities in the cell 122

Roles of the Cytoskeleton: Support and Motility 122

Components of the Cytoskeleton 123

CONCEPT 6.7 Extracellular components and connections between cells help coordinate cellular activities 128

Cell Walls of Plants 128

The Extracellular Matrix (ECM) of Animal Cells 128

Cell Junctions 129

CONCEPT 6.8 A cell is greater than the sum of its parts 131

7 Membrane Structure and

Function 137

Life at the Edge 137

CONCEPT 7.1 Cellular membranes are fluid mosaics of lipids and proteins 138

The Fluidity of Membranes 139

Evolution of Differences in Membrane Lipid Composition 140

Membrane Proteins and Their Functions 140

The Role of Membrane Carbohydrates in Cell-Cell Recognition 141

Synthesis and Sidedness of Membranes 142

CONCEPT 7.2 Membrane structure results in selective permeability 142

The Permeability of the Lipid Bilayer 143

Transport Proteins 143

CONCEPT 7.3 Passive transport is diffusion of a substance across a membrane with no energy investment 143

Effects of Osmosis on Water Balance 144

Facilitated Diffusion: Passive Transport Aided by Proteins 146

CONCEPT 7.4 Active transport uses energy to move solutes against their gradients 147

The Need for Energy in Active Transport 147

How Ion Pumps Maintain Membrane Potential 148

Cotransport: Coupled Transport by a Membrane Protein 149

CONCEPT 7.5 Bulk transport across the plasma membrane occurs by exocytosis and endocytosis 150

Exocytosis 150

Endocytosis 150

8 An Introduction to Metabolism 155

The Energy of Life 155

CONCEPT 8.1 An organism’s metabolism transforms matter and energy, subject to the laws of thermodynamics 156

Organization of the Chemistry of Life into Metabolic Pathways 156

Forms of Energy 156

The Laws of Energy Transformation 157

CONCEPT 8.2 The free-energy change of a reaction tells us whether or not the reaction occurs spontaneously 159

Free Energy Change, _G 159

Free Energy, Stability, and Equilibrium 159

Free Energy and Metabolism 160

CONCEPT 8.3 ATP powers cellular work by coupling exergonic reactions to endergonic reactions 162

The Structure and Hydrolysis of ATP 162

How the Hydrolysis of ATP Performs Work 163

The Regeneration of ATP 164

CONCEPT 8.4 Enzymes speed up metabolic reactions by lowering energy barriers 165

The Activation Energy Barrier 165

How Enzymes Speed Up Reactions 166

Substrate Specificity of Enzymes 167

Catalysis in the Enzyme’s Active Site 167

Effects of Local Conditions on Enzyme Activity 168

The Evolution of Enzymes 171

CONCEPT 8.5 Regulation of enzyme activity helps control metabolism 171

Allosteric Regulation of Enzymes 171

Localization of Enzymes within the Cell 173

9 Cellular Respiration and Fermentation 176

Life Is Work 176

CONCEPT 9.1 Catabolic pathways yield energy by oxidizing organic fuels 177

Catabolic Pathways and Production of ATP 177

Redox Reactions: Oxidation and Reduction 177

The Stages of Cellular Respiration: A Preview 180

CONCEPT 9.2 Glycolysis harvests chemical energy by oxidizing glucose to pyruvate 182

CONCEPT 9.3 After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic

molecules 183

Oxidation of Pyruvate to Acetyl CoA 183

The Citric Acid Cycle 184

CONCEPT 9.4 During oxidative phosphorylation, chemiosmosis couples electron transport to ATP

synthesis 186

The Pathway of Electron Transport 186

Chemiosmosis: The Energy-Coupling Mechanism 187

An Accounting of ATP Production by Cellular Respiration 189

CONCEPT 9.5 Fermentation and anaerobic respiration enable cells to produce ATP without the use of oxygen 192

Types of Fermentation 192

Comparing Fermentation with Anaerobic and Aerobic Respiration 193

The Evolutionary Significance of Glycolysis 193

CONCEPT 9.6 Glycolysis and the citric acid cycle connect to many other metabolic pathways 194

The Versatility of Catabolism 194

Biosynthesis (Anabolic Pathways) 195

Regulation of Cellular Respiration via Feedback Mechanisms 195

10 Photosynthesis 199

The Process That Feeds the Biosphere 199

CONCEPT 10.1 Photosynthesis converts light energy to the chemical energy of food 201

Chloroplasts: The Sites of Photosynthesis in Plants 201

Tracking Atoms through Photosynthesis: Scientific Inquiry 202

The Two Stages of Photosynthesis: A Preview 203

CONCEPT 10.2 The light reactions convert solar energy to the chemical energy of ATP and NADPH 204

The Nature of Sunlight 204

Photosynthetic Pigments: The Light Receptors 205

Excitation of Chlorophyll by Light 207

A Photosystem: A Reaction-Centre Complex Associated with

Light-Harvesting Complexes 207

Linear Electron Flow 209

Cyclic Electron Flow 210

A Comparison of Chemiosmosis in Chloroplasts and

Mitochondria 211

CONCEPT 10.3 The Calvin cycle uses the chemical energy of ATP and NADPH to reduce CO2 to sugar 213

CONCEPT 10.4 Alternative mechanisms of carbon fixation have evolved in hot, arid climates 215

Photorespiration: An Evolutionary Relic? 215

C4 Plants 215

CAM Plants 217

CONCEPT 10.5 Life depends on photosynthesis 218 The Importance of Photosynthesis: A Review 218

11 Cell Communication 222

Cellular Messaging 222

CONCEPT 11.1 External signals are converted to responses within the cell 223

Evolution of Cell Signalling 223

Local and Long-Distance Signalling 225

The Three Stages of Cell Signalling: A Preview 226

CONCEPT 11.2 Reception: A signalling molecule binds

to a receptor protein, causing it to change shape 227

Receptors in the Plasma Membrane 227

Intracellular Receptors 230

CONCEPT 11.3 Transduction: Cascades of molecular interactions relay signals from receptors to target

molecules in the cell 231

Signal Transduction Pathways 231

Protein Phosphorylation and Dephosphorylation 232

Small Molecules and Ions as Second Messengers 233

CONCEPT 11.4 Response: Cell signalling leads to regulation of transcription or cytoplasmic activities 236

Nuclear and Cytoplasmic Responses 236

Regulation of the Response 236

CONCEPT 11.5 Apoptosis integrates multiple cell-signalling

pathways 240

Apoptosis in the Soil Worm Caenorhabditis elegans 241

Apoptotic Pathways and the Signals That Trigger Them 241

12 The Cell Cycle 246

The Key Roles of Cell Division 246

CONCEPT 12.1 Most cell division results in genetically

identical daughter cells 247

Cellular Organization of the Genetic Material 247

Distribution of Chromosomes During Eukaryotic

Cell Division 248

CONCEPT 12.2 The mitotic phase alternates with interphase

in the cell cycle 249

Phases of the Cell Cycle 249

The Mitotic Spindle: A Closer Look 249

Cytokinesis: A Closer Look 253

Binary Fission in Bacteria 254

The Evolution of Mitosis 255

CONCEPT 12.3 The eukaryotic cell cycle is regulated by a molecular control system 256

The Cell Cycle Control System 256

Loss of Cell Cycle Controls in Cancer Cells 261


Interview: Julie Claycomb 267

13 Meiosis and Sexual Life Cycles 270

Variations on a Theme 270

CONCEPT 13.1 Offspring acquire genes from parents by inheriting chromosomes 271

Inheritance of Genes 271

Comparison of Asexual and Sexual Reproduction 271

CONCEPT 13.2 Fertilization and meiosis alternate in sexual life cycles 272

Sets of Chromosomes in Human Cells 272

Behaviour of Chromosome Sets in the Human Life Cycle 273

The Variety of Sexual Life Cycles 274

CONCEPT 13.3 Meiosis reduces the number of chromosome sets from diploid to haploid 275

The Stages of Meiosis 275

Crossing Over and Synapsis During Prophase I 278

A Comparison of Mitosis and Meiosis 278

CONCEPT 13.4 Genetic variation produced in sexual life cycles contributes to evolution 281

Origins of Genetic Variation Among Offspring 281

The Evolutionary Significance of Genetic Variation Within

Populations 282

14 Mendel and the Gene Idea 285

Drawing from the Deck of Genes 285

CONCEPT 14.1 Mendel used the scientific approach to identify two laws of inheritance 286

Mendel’s Experimental, Quantitative Approach 286

The Law of Segregation 287

The Law of Independent Assortment 290

CONCEPT 14.2 Probability laws govern Mendelian inheritance 292

The Multiplication and Addition Rules Applied to Monohybrid Crosses 293

Solving Complex Genetics Problems with the Rules

of Probability 293

CONCEPT 14.3 Inheritance patterns are often more complex than predicted by simple Mendelian genetics 294

Extending Mendelian Genetics for a Single Gene 294

Extending Mendelian Genetics for Two or More Genes 297

Nature and Nurture: The Environmental Impact on Phenotype 298

A Mendelian View of Heredity and Variation 299

CONCEPT 14.4 Many human traits follow Mendelian patterns of inheritance 300

Pedigree Analysis 300

Recessively Inherited Disorders 301

Dominantly Inherited Disorders 303

Multifactorial Disorders 304

Genetic Testing and Counselling 304

15 The Chromosomal Basis of Inheritance 312

Locating Genes Along Chromosomes 312

CONCEPT 15.1 Morgan showed that Mendelian inheritance

has its physical basis in the behaviour of chromosomes: Scientific Inquiry 314

CONCEPT 15.2 Sex-linked genes exhibit unique patterns of inheritance 315

The Chromosomal Basis of Sex 316

Inheritance of X-Linked Genes 317

X Inactivation in Female Mammals 318

CONCEPT 15.3 Linked genes tend to be inherited together because they are located near each other on the same

chromosome 319

How Linkage Affects Inheritance 319

Genetic Recombination and Linkage 320

Mapping the Distance Between Genes Using Recombination Data: Scientific Inquiry 323

CONCEPT 15.4 Alterations of chromosome number or

structure cause some genetic disorders 325

Abnormal Chromosome Number 325

Alterations of Chromosome Structure 326

Human Disorders Due to Chromosomal Alterations 327

CONCEPT 15.5 Some inheritance patterns are exceptions to standard Mendelian inheritance 329

Genomic Imprinting 329

Inheritance of Organelle Genes 330

16 The Molecular Basis of

Inheritance 334

Life’s Operating Instructions 334

CONCEPT 16.1 DNA is the genetic material 335

The Search for the Genetic Material: Scientific Inquiry 335

Building a Structural Model of DNA: Scientific Inquiry 337

CONCEPT 16.2 Many proteins work together in DNA replication and repair 340

The Basic Principle: Base Pairing to a Template Strand 340

DNA Replication: A Closer Look 342

Proofreading and Repairing DNA 346

Evolutionary Significance of Altered DNA Nucleotides 348

Replicating the Ends of DNA Molecules 348

CONCEPT 16.3 A chromosome consists of a DNA molecule packed together with proteins 350

17 Gene Expression: From Gene to Protein 355

The Flow of Genetic Information 355

CONCEPT 17.1 Genes specify proteins via transcription

and translation 356

Evidence from the Study of Metabolic Defects 356

Basic Principles of Transcription and Translation 357

The Genetic Code 359

CONCEPT 17.2 Transcription is the DNA-directed synthesis

of RNA: A closer look 362

Molecular Components of Transcription 362

Synthesis of an RNA Transcript 362

CONCEPT 17.3 Eukaryotic cells modify RNA after

transcription 365

Alteration of mRNA Ends 365

Split Genes and RNA Splicing 365

CONCEPT 17.4 Translation is the RNA-directed synthesis

of a polypeptide: A closer look 367

Molecular Components of Translation 368

Building a Polypeptide 370

Completing and Targeting the Functional Protein 372

Making Multiple Polypeptides in Bacteria and Eukaryotes 375

CONCEPT 17.5 Mutations of one or a few nucleotides can

affect protein structure and function 377

Types of Small-Scale Mutations 377

New Mutations and Mutagens 378

What Is a Gene? Revisiting the Question 380

18 Regulation of Gene Expression 385

Beauty in the Eye of the Beholder 385

CONCEPT 18.1 Bacteria often respond to environmental

change by regulating transcription 386

Operons: The Basic Concept 386

Repressible and Inducible Operons: Two Types of Negative Gene

Regulation 388

Positive Gene Regulation 389

CONCEPT 18.2 Eukaryotic gene expression is regulated at

many stages 390

Differential Gene Expression 390

Regulation of Chromatin Structure 391

Regulation of Transcription Initiation 392

Mechanisms of Post-Transcriptional Regulation 397

CONCEPT 18.3 Noncoding RNAs play multiple roles in

controlling gene expression 399

Effects on mRNAs by MicroRNAs and Small Interfering RNAs 399

Chromatin Remodelling and Effects on Transcription by ncRNAs 400

The Evolutionary Significance of Small ncRNAs 401

CONCEPT 18.4 A program of differential gene expression

leads to the different cell types in a multicellular organism 401

A Genetic Program for Embryonic Development 401

Cytoplasmic Determinants and Inductive Signals 402

Sequential Regulation of Gene Expression During Cellular

Differentiation 403

Pattern Formation: Setting Up the Body Plan 404

CONCEPT 18.5 Cancer results from genetic changes that

affect cell cycle control 408

Types of Genes Associated with Cancer 408

Interference with Normal Cell-Signalling Pathways 409

The Multistep Model of Cancer Development 411

Inherited Predisposition and Environmental Factors Contributing

to Cancer 414

The Role of Viruses in Cancer 415

19 Viruses 419

A Borrowed Life 419

CONCEPT 19.1 A virus consists of a nucleic acid surrounded

by a protein coat 420

The Discovery of Viruses: Scientific Inquiry 420

Structure of Viruses 420

CONCEPT 19.2 Viruses replicate only in host cells 422

General Features of Viral Replicative Cycles 422

Replicative Cycles of Phages 423

Replicative Cycles of Animal Viruses 426

Evolution of Viruses 428

CONCEPT 19.3 Viruses and prions are formidable pathogens

in animals and plants 430

Viral Diseases in Animals 430

Emerging Viruses 431

Viral Diseases in Plants 435

Prions: Proteins as Infectious Agents 435

20 DNA Tools and Biotechnology 438

The DNA Toolbox 438

CONCEPT 20.1 DNA sequencing and DNA cloning are

valuable tools for genetic engineering and biological inquiry 439

DNA Sequencing 439

Making Multiple Copies of a Gene or Other DNA Segment 441

Using Restriction Enzymes to Make a Recombinant DNA Plasmid 442

Amplifying DNA: The Polymerase Chain Reaction (PCR) and Its

Use in DNA Cloning 443

Expressing Cloned Eukaryotic Genes 445

CONCEPT 20.2 Biologists use DNA technology to study gene

expression and function 446

Analyzing Gene Expression 446

Determining Gene Function 450

CONCEPT 20.3 Cloned organisms and stem cells are useful

for basic research and other applications 452

Cloning Plants: Single-Cell Cultures 453

Cloning Animals: Nuclear Transplantation 453

Stem Cells of Animals 455

CONCEPT 20.4 The practical applications of DNA-based

biotechnology affect our lives in many ways 458

Medical Applications 458

Forensic Evidence and Genetic Profiles 461

Environmental Cleanup 462

Agricultural Applications 462

Safety and Ethical Questions Raised by DNA Technology 463

21 Genomes and Their Evolution 468

Combining Genomics with Traditional Ecological

Knowledge 468

CONCEPT 21.1 The Human Genome Project fostered

development of faster, less expensive sequencing

techniques 469

CONCEPT 21.2 Scientists use bioinformatics to analyze

genomes and their functions 470

Centralized Resources for Analyzing Genome Sequences 470

Identifying Protein-Coding Genes and Understanding Their

Functions 471

Understanding Genes and Gene Expression at the Systems Level 472

CONCEPT 21.3 Genomes vary in size, number of genes, and

gene density 474

Genome Size 474

Number of Genes 475

Gene Density and Noncoding DNA 475

CONCEPT 21.4 Multicellular eukaryotes have a lot of

noncoding DNA and many multigene families 478

Transposable Elements and Related Sequences 478

Other Repetitive DNA, Including Simple Sequence DNA 480

Genes and Multigene Families 480

CONCEPT 21.5 Duplication, rearrangement, and mutation of

DNA contribute to genome evolution 482

Duplication of Entire Chromosome Sets 482

Alterations of Chromosome Structure 482

Duplication and Divergence of Gene-Sized Regions of DNA 483

Rearrangements of Parts of Genes: Exon Duplication and Exon

Shuffling 484

How Transposable Elements Contribute to

Genome Evolution 487

CONCEPT 21.6 Comparing genome sequences provides clues

to evolution and development 487

Comparing Genomes 487


Interview: Maydianne Andrade 495

22 Descent with Modification:

A Darwinian View of Life 498

“Endless Forms Most Beautiful”—Charles Darwin 498

CONCEPT 22.1 The Darwinian revolution challenged traditional

views of a young Earth inhabited by unchanging species 499

Scala Naturae and Classification of Species 500

Ideas About Change over Time 500

Lamarck’s Hypothesis of Evolution 500

CONCEPT 22.2 Descent with modification by natural

selection explains the adaptations of organisms and the

unity and diversity of life 501

Darwin’s Research 501

CONCEPT 22.3 Evolution is supported by an overwhelming

amount of scientific evidence 506

Direct Observations of Evolutionary Change 507

Homology 509

The Fossil Record 511

Biogeography 512

What Is Theoretical about Darwin’s View of Life? 513

23 The Evolution of Populations 517

The Smallest Unit of Evolution 517

CONCEPT 23.1 Genetic variation makes evolution

possible 518

Genetic Variation 518

Sources of Genetic Variation 519

CONCEPT 23.2 The Hardy-Weinberg equation can be used

to test whether a population is evolving 520

Gene Pools and Allele Frequencies 521

The Hardy-Weinberg Equation 521

CONCEPT 23.3 Natural selection, genetic drift, and gene

flow can alter allele frequencies in a population 524

Natural Selection 525

Genetic Drift 525

Gene Flow 527

CONCEPT 23.4 Natural selection is the only mechanism that

consistently causes adaptive evolution 528

Natural Selection: A Closer Look 528

The Key Role of Natural Selection in Adaptive Evolution 530

Sexual Selection 530

Balancing Selection 531

Frequency-Dependent Selection 531

Heterozygote Advantage 532

Why Natural Selection Cannot Fashion Perfect Organisms 532

24 The Origin of Species 536

That “Mystery of Mysteries” 536

CONCEPT 24.1 The biological species concept emphasizes

reproductive isolation 537

The Biological Species Concept 537

Other Definitions of Species 540

CONCEPT 24.2 Speciation can take place with or without

geographic separation 541

Allopatric (“Other Country”) Speciation 541

Sympatric (“Same Country”) Speciation 543

Allopatric and Sympatric Speciation: A Review 546

CONCEPT 24.3 Hybrid zones reveal factors that cause

reproductive isolation 546

Patterns Within Hybrid Zones 546

Hybrid Zones and Environmental Change 547

Hybrid Zones over Time 547

CONCEPT 24.4 Speciation can occur rapidly or slowly and

can result from changes in few or many genes 550

The Time Course of Speciation 550

Studying the Genetics of Speciation 552

From Speciation to Macroevolution 553

25 The History of Life on Earth 555

Dinosaurs of a Feather 555

CONCEPT 25.1 Conditions on early Earth made the origin

of life possible 556

Synthesis of Organic Compounds on Early Earth 556

Abiotic Synthesis of Macromolecules 557

Protocells 557

Self-Replicating RNA 558

CONCEPT 25.2 The fossil record documents the history of life 558

The Fossil Record 558

How Rocks and Fossils Are Dated 560

The Origin of New Groups of Organisms 560

CONCEPT 25.3 Key events in life’s history include the

origins of unicellular and multicelled organisms and the

colonization of land 562

The First Single-Celled Organisms 564

The Origin of Multicellularity 565

The Colonization of Land 566

CONCEPT 25.4 The rise and fall of groups of organisms

reflect differences in speciation and extinction rates 567

Plate Tectonics 568

Mass Extinctions 570

Adaptive Radiations 572

CONCEPT 25.5 Major changes in body form can result from

changes in the sequences and regulation of developmental genes 574

Effects of Developmental Genes 575

The Evolution of Development 576

CONCEPT 25.6 Evolution is not goal oriented 578

Evolutionary Novelties 578

Evolutionary Trends 579



Interview: Laura Hug 583

26 Phylogeny and the Tree of Life 586

Investigating the Tree of Life 586

CONCEPT 26.1 Phylogenies show evolutionary

relationships 587

Binomial Nomenclature 588

Hierarchical Classification 588

Linking Classification and Phylogeny 589

Visualizing Phylogenetic Relationships 589

CONCEPT 26.2 Phylogenies are inferred from morphological

and molecular data 591

Morphological and Molecular Homologies 591

Sorting Homology from Analogy 591

Evaluating Molecular Homologies 591

CONCEPT 26.3 Shared characters are used to construct

phylogenetic trees 592

Cladistics 592

Maximum Parsimony and Maximum Likelihood 594

Interpreting Phylogenetic Trees 595

Applying Phylogenies 597

CONCEPT 26.4 An organism’s evolutionary history is

documented in its genome 599

Gene Duplications and Gene Families 599

Genome Evolution 600

CONCEPT 26.5 Molecular clocks help track evolutionary time 600

Molecular Clocks 600

Applying a Molecular Clock: The Origin of HIV 601

CONCEPT 26.6 Our understanding of the tree of life

continues to change based on new data 602

From Two Kingdoms to Three Domains 602

The Important Role of Horizontal Gene Transfer 603

27 Bacteria and Archaea 607

Masters of Adaptation 607

CONCEPT 27.1 Structural and functional adaptations

contribute to prokaryotic success 608

Cell-Surface Structures 608

Endospores 609

Motility 610

Internal Organization and DNA 611

Reproduction 612

CONCEPT 27.2 Rapid reproduction, mutation, and

genetic recombination promote genetic diversity in

prokaryotes 612

Rapid Reproduction and Mutation 612

Genetic Recombination 612

CONCEPT 27.3 Diverse nutritional and metabolic

adaptations have evolved in prokaryotes 615

The Role of Oxygen in Metabolism 616

Nitrogen Metabolism 616

Metabolic Cooperation 616

CONCEPT 27.4 Prokaryotes have radiated into a diverse set

of lineages 617

An Overview of Prokaryotic Diversity 617

Bacteria 620

Archaea 620

CONCEPT 27.5 Prokaryotes play crucial roles in the

biosphere 621

Chemical Recycling 621

Ecological Interactions 622

CONCEPT 27.6 Prokaryotes have both beneficial

and harmful impacts on humans 623

Mutualistic Bacteria 623

Pathogenic Bacteria 623

Prokaryotes in Research and Technology 624

28 Protists 628

You Are What You Eat 628

CONCEPT 28.1 Most eukaryotes are single-celled

organisms 629

Structural and Functional Diversity in Protists 629

Four Supergroups of Eukaryotes 629

Endosymbiosis in Eukaryotic Evolution 632

Endosymbiosis and the Spread of Photosynthesis 632

How Does an Endosymbiont Evolve into an Organelle? 633

CONCEPT 28.2 Excavates include protists with modified

mitochondria and protists with unique flagella 635

Diplomonads and Parabasalids 635

Euglenozoans 635

CONCEPT 28.3 SAR is a highly diverse group of protists

defined by DNA similarities 637

Stramenopiles 637

Oomycetes (Water Moulds and Their Relatives) 639

Alveolates 640

Rhizarians 641

CONCEPT 28.4 Red algae and green algae are the closest

relatives of land plants 645

Red Algae 645

Green Algae 645

CONCEPT 28.5 Unikonts include protists that are closely

related to fungi and animals 647

Amoebozoans 648

Opisthokonts 650

CONCEPT 28.6 The relationships of some protists to other

eukaryotes is uncertain 650

Haptophytes 650

Cryptomonads 651

CONCEPT 28.7 Protists play key roles in ecological

communities 652

Symbiotic Protists 652

Photosynthetic Protists 652

29 Plant Diversity I: How Plants

Colonized Land 657

The Greening of Earth 657

CONCEPT 29.1 Plants evolved from green algae 658

Morphological and Molecular Evidence 658

Adaptations Enabling the Move to Land 658

Derived Traits of Plants 659

The Origin and Diversification of Plants 659

CONCEPT 29.2 Mosses and other nonvascular plants have

life cycles dominated by gametophytes 663

Bryophyte Gametophytes 664

Bryophyte Sporophytes 666

The Ecological and Economic Importance of Mosses 666

CONCEPT 29.3 Ferns and other seedless vascular plants

were the first plants to grow tall 668

Origins and Traits of Vascular Plants 668

Classification of Seedless Vascular Plants 672

The Significance of Seedless Vascular Plants 672

30 Plant Diversity II: The Evolution

of Seed Plants 678

Transforming the World 678

CONCEPT 30.1 Seeds and pollen grains are key adaptations

for life on land 679

Advantages of Reduced Gametophytes 679

Heterospory: The Rule Among Seed Plants 680

Ovules and Production of Eggs 680

Pollen and Production of Sperm 680

The Evolutionary Advantage of Seeds 681

Evolution of the Seed 682

CONCEPT 30.2 Gymnosperms bear “naked” seeds, typically

on cones 682

The Life Cycle of a Pine 682

Evolution of Gymnosperms 686

Gymnosperm Diversity 686

CONCEPT 30.3 The reproductive adaptations of angiosperms

include flowers and fruits 686

Characteristics of Angiosperms 686

Angiosperm Evolution 690

Evolutionary Links Between Angiosperms and Animals 692

Angiosperm Diversity 692

CONCEPT 30.4 Human welfare depends greatly on seed plants 695

Products from Seed Plants 695

Threats to Plant Diversity 695

31 Fungi 698

Brewer’s Yeast and Climate Change 698

CONCEPT 31.1 Fungi are heterotrophs that feed

by absorption 699

Nutrition and Ecology 699

Body Structure 699

Specialized Hyphae in Mycorrhizal Fungi 700

CONCEPT 31.2 Fungi produce spores through sexual or

asexual life cycles 702

Sexual Reproduction 702

Asexual Reproduction 702

CONCEPT 31.3 The ancestor of fungi was an aquatic, singlecelled,

flagellated protist 703

The Origin of Fungi 703

Basal Fungal Groups 703

The Move to Land 704

CONCEPT 31.4 Fungi have radiated into a diverse set of lineages 704

Chytrids 704

Zygomycetes 706

Glomeromycetes 707

Ascomycetes 707

Basidiomycetes 709

CONCEPT 31.5 Fungi play key roles in nutrient cycling,

ecological interactions, and human welfare 711

Fungi as Decomposers (Saprotrophs) 711

Fungi as Mutualists 711

Fungi as Parasites 712

Practical Uses of Fungi 714

32 An Overview of Animal Diversity 717

Welcome to Your Kingdom 717

CONCEPT 32.1 Animals are multicellular, heterotrophic

eukaryotes with tissues that develop from embryonic layers 718

Nutritional Mode 718

Cell Structure and Specialization 718

Reproduction and Development 718

CONCEPT 32.2 The history of animals spans more than half

a billion years 719

Steps in the Origin of Multicellular Animals 719

Neoproterozoic Era (1 Billion–542 Million Years Ago) 720

Paleozoic Era (542–251 Million Years Ago) 720

Mesozoic Era (251–65.5 Million Years Ago) 722

Cenozoic Era (65.5 Million Years Ago to the Present) 722

CONCEPT 32.3 Animals can be characterized by “body plans” 723

Symmetry 724

Tissues 724

Body Cavities 724

Protostome and Deuterostome Development 725

CONCEPT 32.4 Views of animal phylogeny continue to be

shaped by new molecular and morphological data 726

The Diversification of Animals 726

Future Directions in Animal Systematics 728

33 An Introduction to Invertebrates 731

Life Without a Backbone 731

CONCEPT 33.1 Sponges are basal animals that lack true tissues 735

CONCEPT 33.2 Cnidarians are an ancient phylum of eumetazoans 736

Medusozoans 737

Anthozoans 737

CONCEPT 33.3 Lophotrochozoans, a clade identified by

molecular data, have the widest range of animal body forms 739

Flatworms 739

Rotifers 742

Lophophorates: Ectoprocts and Brachiopods 743

Molluscs 744

Annelids 747

CONCEPT 33.4 Ecdysozoans are the most species-rich animal

group 750

Nematodes 750

Arthropods 751

CONCEPT 33.5 Echinoderms and chordates are

deuterostomes 759

Echinoderms 759

Chordates 761

34 The Origin and Evolution of Vertebrates 765

Half a Billion Years of Backbones 765

CONCEPT 34.1 Chordates have a notochord and a dorsal,

hollow nerve cord 766

Derived Characters of Chordates 766

Lancelets 767

Tunicates 768

Early Chordate Evolution 769

CONCEPT 34.2 Vertebrates are chordates that have a

backbone 769

Derived Characters of Vertebrates 769

Hagfishes and Lampreys 770

Hagfishes 770

Lampreys 770

Early Vertebrate Evolution 771

CONCEPT 34.3 Gnathostomes are vertebrates that have jaws 772

Derived Characters of Gnathostomes 772

Fossil Gnathostomes 773

Chondrichthyans (Sharks, Rays, and Their Relatives) 773

Ray-Finned Fishes and Lobe-Fins 775

CONCEPT 34.4 Tetrapods are gnathostomes that have

limbs 777

Derived Characters of Tetrapods 777

The Origin of Tetrapods 777

Amphibians 779

Salamanders 779

Frogs 779

Caecilians 779

Lifestyle and Ecology of Amphibians 779

CONCEPT 34.5 Amniotes are tetrapods that have

a terrestrially adapted egg 782

Derived Characters of Amniotes 782

Early Amniotes 783

Reptiles 783

CONCEPT 34.6 Mammals are amniotes that have hair and

produce milk 788

Derived Characters of Mammals 789

Early Evolution of Mammals 789

Monotremes 790

Marsupials 790

Eutherians (Placental Mammals) 791

CONCEPT 34.7 Humans are mammals that have a large brain

and bipedal locomotion 796

Derived Characters of Humans 796

The Earliest Hominins 797

Australopiths 797

Bipedalism 798

Tool Use 798

Early Homo 799

Neanderthals 800


Interview: Jacqueline Monaghan 805

35 Plant Structure, Growth, and

Development 808

Why Did the Palm Tree Cross the Road? 808

CONCEPT 35.1 Plants have a hierarchical organization

consisting of organs, tissues, and cells 809

Basic Vascular Plant Organs: Roots, Stems, and Leaves 809

Dermal, Vascular, and Ground Plant Tissues 812

Common Types of Plant Cells 813

CONCEPT 35.2 Different meristems generate cells for

primary and secondary growth 816

CONCEPT 35.3 Primary growth lengthens roots and shoots 818

Primary Growth of Roots 818

Primary Growth of Shoots 818

CONCEPT 35.4 Secondary growth increases the diameter of

stems and roots in woody plants 822

The Vascular Cambium and Secondary Vascular Tissue 823

The Cork Cambium and the Production of Periderm 824

Evolution of Secondary Growth 824

CONCEPT 35.5 Growth, morphogenesis, and cell

differentiation produce the plant body 825

Model Organisms: Revolutionizing the Study of Plants 826

Growth: Cell Division and Cell Expansion 826

Morphogenesis and Pattern Formation 828

Gene Expression and Control of Cell Differentiation 828

Shifts in Development: Phase Changes 829

Genetic Control of Flowering 830

36 Resource Acquisition and Transport

in Vascular Plants 834

Overview 834

CONCEPT 36.1 Adaptations for acquiring resources were key

steps in the evolution of vascular plants 835

Shoot Architecture and Light Capture 835

Root Architecture and Acquisition of Water and Minerals 837

CONCEPT 36.2 Different mechanisms transport substances

over short or long distances 837

The Apoplast and Symplast: Transport Continuums 837

Short-Distance Transport of Solutes across Plasma

Membranes 838

Short-Distance Transport of Water Across Plasma

Membranes 838

Long-Distance Transport: The Role of Bulk Flow 841

CONCEPT 36.3 Transpiration drives the transport

of water and minerals from roots to shoots via the

xylem 842

Absorption of Water and Minerals by Root Epidermal

Cells 842

Transport of Water and Minerals into the Xylem 842

Bulk Flow Transport via the Xylem 842

Xylem Sap Ascent by Bulk Flow: A Review 846

CONCEPT 36.4 The rate of transpiration is regulated by stomata 846

Stomata: Major Pathways for Water Loss 847

Mechanisms of Stomatal Opening and Closing 847

Stimuli for Stomatal Opening and Closing 848

Effects of Transpiration on Wilting and Leaf Temperature 848

Adaptations That Reduce Evaporative Water Loss 848

CONCEPT 36.5 Sugars are transported from sources to sinks via the phloem 849

Movement from Sugar Sources to Sugar Sinks 849

Bulk Flow by Positive Pressure: The Mechanism of Translocation

in Angiosperms 850

CONCEPT 36.6 The symplast is highly dynamic 852

Changes in Plasmodesmatal Number and Pore Size 852

Phloem: An Information Superhighway 852

Electrical Signalling in the Phloem 852

37 Soil and Plant Nutrition 855

The Corkscrew Carnivore 855

CONCEPT 37.1 Soil contains a living, complex ecosystem 856

Soil Texture 856

Topsoil Composition 856

Soil Conservation and Sustainable Agriculture 857

CONCEPT 37.2 Plant roots absorb essential elements from the soil 860

Essential Elements 860

Symptoms of Mineral Deficiency 861

Improving Plant Nutrition by Genetic Modification 862

CONCEPT 37.3 Plant nutrition often involves relationships

with other organisms 863

Bacteria and Plant Nutrition 863

Fungi and Plant Nutrition 867

Vertebrates and Plant Nutrition 868

Epiphytes, Parasitic Plants, and Carnivorous Plants 869

38 Angiosperm Reproduction and

Biotechnology 873

Canola (Canadian Oil Low Acid): A Canadian Invention 873

CONCEPT 38.1 Flowers, double fertilization, and fruits are

unique features of the angiosperm life cycle 874

Flower Structure and Function 874

Methods of Pollination 875

The Angiosperm Life Cycle: An Overview 875

Seed Development and Structure: A Closer Look 878

Sporophyte Development from Seed to Mature Plant 881

Fruit Structure and Function 882

CONCEPT 38.2 Flowering plants reproduce sexually,

asexually, or both 883

Mechanisms of Asexual Reproduction 883

Advantages and Disadvantages of Asexual Versus Sexual

Reproduction 885

Mechanisms That Prevent Self-Fertilization 886

Totipotency, Vegetative Reproduction, and Tissue Culture 887

CONCEPT 38.3 Humans modify crops by breeding and

genetic engineering 888

Plant Breeding 889

Plant Biotechnology and Genetic Engineering 889

The Debate over Plant Biotechnology 891

39 Plant Responses to Internal

and External Signals 895

Stimuli and a Stationary Life 895

CONCEPT 39.1 Signal transduction pathways link signal

reception to response 896

Reception 897

Transduction 897

Response 898

CONCEPT 39.2 Plant hormones help coordinate growth,

development, and responses to stimuli 899

A Survey of Plant Hormones 900

CONCEPT 39.3 Responses to light are critical for plant

success 908

Blue-Light Photoreceptors 908

Phytochromes as Photoreceptors 909

Biological Clocks and Circadian Rhythms 910

The Effect of Light on the Biological Clock 911

Photoperiodism and Responses to Seasons 912

CONCEPT 39.4 Plants respond to a wide variety of stimuli

other than light 914

Gravity 914

Mechanical Stimuli 914

Environmental Stresses 915

CONCEPT 39.5 Plants respond to attacks by herbivores

and pathogens 919

Defences against Pathogens 919

Defences Against Herbivores 921


Interview: Matt Vijayan 925

40 Basic Principles of Animal Form

and Function 928

Diverse Forms, Common Challenges 928

CONCEPT 40.1 Animal form and function are correlated at

all levels of organization 929

Evolution of Animal Size and Shape 929

Exchange with the Environment 929

Hierarchical Organization of Body Plans 931

Coordination and Control 935

CONCEPT 40.2 Feedback control maintains the internal

environment in many animals 936

Regulating and Conforming 936

Homeostasis 936

CONCEPT 40.3 Homeostatic processes for thermoregulation

involve form, function, and behaviour 938

Endothermy and Ectothermy 939

Variation in Body Temperature 939

Balancing Heat Loss and Gain 940

Acclimation and Acclimatization 943

Physiological Thermostats and Fever 944

CONCEPT 40.4 Energy requirements are related to animal

size, activity, and environment 945

Energy Allocation and Use 945

Quantifying Energy Use 945

Minimum Metabolic Rate and Thermoregulation 946

Influences on Metabolic Rate 946

Torpor and Energy Conservation 948

41 Animal Nutrition 951

The Need to Feed 951

CONCEPT 41.1 An animal’s diet must supply chemical energy

and building blocks 952

Essential Nutrients 952

Dietary Deficiencies 954

Assessing Nutritional Needs 956

CONCEPT 41.2 Food processing involves ingestion,

digestion, absorption, and elimination 956

Digestive Compartments 958

CONCEPT 41.3 Organs specialized for sequential stages

of food processing form the mammalian digestive

system 959

The Oral Cavity, Pharynx, and Esophagus 960

Digestion in the Stomach 960

Digestion in the Small Intestine 962

Absorption in the Small Intestine 963

Absorption in the Large Intestine 964

CONCEPT 41.4 Evolutionary adaptations of vertebrate

digestive systems correlate with diet 965

Dental Adaptations 965

Stomach and Intestinal Adaptations 965

Mutualistic Adaptations 966

Mutualistic Adaptations in Herbivores 968

CONCEPT 41.5 Feedback circuits regulate digestion, energy

storage, and appetite 969

Regulation of Digestion 969

Regulation of Energy Storage 969

Regulation of Appetite and Consumption 971

Obesity and Evolution 972

42 Circulation and Gas Exchange 975

Trading Places 975

CONCEPT 42.1 Circulatory systems link exchange surfaces

with cells throughout the body 976

General Properties of Circulatory Systems 976

Open and Closed Circulatory Systems 977

Evolution of Vertebrate Circulatory Systems 978

CONCEPT 42.2 Coordinated cycles of heart contraction drive

double circulation in mammals 981

Mammalian Circulation 981

The Mammalian Heart: A Closer Look 981

Maintaining the Heart’s Rhythmic Beat 982

CONCEPT 42.3 Patterns of blood pressure and flow reflect

the structure and arrangement of blood vessels 983

Blood Vessel Structure and Function 984

Blood Flow Velocity 984

Blood Pressure 985

Capillary Function 987

Fluid Return by the Lymphatic System 988

CONCEPT 42.4 Blood components function in exchange,

transport, and defence 989

Blood Composition and Function 989

Cardiovascular Disease 992

CONCEPT 42.5 Gas exchange occurs across specialized

respiratory surfaces 994

Partial Pressure Gradients in Gas Exchange 994

Respiratory Media 994

Respiratory Surfaces 994

Gills in Aquatic Animals 995

Tracheal Systems in Insects 996

Lungs 997

CONCEPT 42.6 Breathing ventilates the lungs 999

How an Amphibian Breathes 999

How a Bird Breathes 999

How a Mammal Breathes 999

Control of Breathing in Humans 1000

CONCEPT 42.7 Adaptations for gas exchange include

pigments that bind and transport gases 1001

Coordination of Circulation and Gas Exchange 1001

Respiratory Pigments 1002

Respiratory Adaptations of Diving Mammals 1003

43 The Immune System 1008

Recognition and Response 1008

CONCEPT 43.1 In innate immunity, recognition and response

rely on traits common to groups of pathogens 1009

Innate Immunity of Invertebrates 1009

Innate Immunity of Vertebrates 1011

Evasion of Innate Immunity by Pathogens 1014

CONCEPT 43.2 In adaptive immunity, receptors provide

pathogen-specific recognition 1014

Antigen Recognition by B Cells and Antibodies 1014

Antigen Recognition by T Cells 1015

B Cell and T Cell Development 1016

CONCEPT 43.3 Adaptive immunity defends against infection

of body fluids and body cells 1019

Helper T Cells: A Response to Nearly All Antigens 1020

B Cells and Antibodies: A Response to Extracellular Pathogens 1020

Cytotoxic T Cells: A Response to Infected Cells 1023

Summary of the Humoral and Cell-Mediated Immune

Responses 1023

Immunization 1024

Active and Passive Immunity 1025

Antibodies as Tools 1025

Immune Rejection 1026

CONCEPT 43.4 Disruptions in immune system function can

elicit or exacerbate disease 1027

Exaggerated, Self-Directed, and Diminished Immune

Responses 1027

Evolutionary Adaptations of Pathogens That Underlie Immune

System Avoidance 1029

Cancer and Immunity 1032

44 Osmoregulation and Excretion 1035

A Balancing Act 1035

CONCEPT 44.1 Osmoregulation balances the uptake and loss

of water and solutes 1036

Osmosis and Osmolarity 1036

Osmotic Challenges 1036

Energetics of Osmoregulation 1039

Transport Epithelia in Osmoregulation 1039

CONCEPT 44.2 An animal’s nitrogenous wastes reflect its

phylogeny and habitat 1040

Forms of Nitrogenous Waste 1040

The Influence of Evolution and Environment on Nitrogenous

Wastes 1041

CONCEPT 44.3 Diverse excretory systems are variations

on a tubular theme 1042

Excretory Processes 1042

Survey of Excretory Systems 1042

CONCEPT 44.4 The nephron is organized for stepwise

processing of blood filtrate 1046

From Blood Filtrate to Urine: A Closer Look 1046

Solute Gradients and Water Conservation 1047

Adaptations of the Vertebrate Kidney to Diverse Environments 1049

CONCEPT 44.5 Hormonal circuits link kidney function, water

balance, and blood pressure 1052

Antidiuretic Hormone 1052

The Renin-Angiotensin-Aldosterone System 1053

Coordinated Regulation of Salt and Water Balance 1054

45 Hormones and the Endocrine

System 1057

The Body’s Long-Distance Regulators 1057

CONCEPT 45.1 Hormones and other signalling molecules

bind to target receptors, triggering specific response

pathways 1058

Intercellular Communication 1058

Chemical Classes of Intercellular Signalling Factors 1059

Cellular Response Pathways 1060

Multiple Effects of Hormones 1062

CONCEPT 45.2 Feedback regulation and coordination with

the nervous system are common in endocrine systems 1064

Simple Endocrine Pathways 1064

Simple Neuroendocrine Pathways 1064

Feedback Regulation 1064

Coordination of the Endocrine and Nervous Systems 1065

Thyroid Regulation: A Hormone Cascade Pathway 1068

Hormonal Regulation of Growth 1069

CONCEPT 45.3 Endocrine glands respond to diverse stimuli

in regulating homeostasis, development, and behaviour 1070

Parathyroid Hormone and Vitamin D: Control of Blood Calcium 1070

Adrenal Hormones: Response to Stress 1070

Sex Hormones 1072

Hormones and Biological Rhythms 1074

46 Animal Reproduction 1078

Pairing Up for Sexual Reproduction 1078

CONCEPT 46.1 Both asexual and sexual reproduction occur

in the animal kingdom 1079

Mechanisms of Asexual Reproduction 1079

Variation in Patterns of Sexual Reproduction 1079

Reproductive Cycles 1080

Sexual Reproduction: An Evolutionary Enigma 1081

CONCEPT 46.2 Fertilization depends on mechanisms that

bring together sperm and eggs of the same species 1082

Ensuring the Survival of Offspring 1082

Gamete Production and Delivery 1083

CONCEPT 46.3 Reproductive organs produce and transport

gametes 1085

Human Male Reproductive Anatomy 1085

Human Female Reproductive Anatomy 1086

Gametogenesis 1087

CONCEPT 46.4 The interplay of tropic and sex hormones

regulates mammalian reproduction 1090

Hormonal Control of the Male Reproductive System 1091

Hormonal Control of Female Reproductive Cycles 1091

Human Sexual Response 1094

CONCEPT 46.5 In placental mammals, an embryo develops

fully within the mother’s uterus 1094

Conception, Embryonic Development, and Birth 1094

Maternal Immune Tolerance of the Embryo and Fetus 1098

Contraception and Abortion 1098

Modern Reproductive Technologies 1099

47 Animal Development 1103

A Body-Building Plan 1103

CONCEPT 47.1 Fertilization and cleavage initiate embryonic

development 1104

Fertilization 1104

Cleavage 1107

CONCEPT 47.2 Morphogenesis in animals involves specific

changes in cell shape, position, and survival 1109

Gastrulation 1109

Developmental Adaptations of Amniotes 1113

Organogenesis 1114

Cellular Mechanisms in Morphogenesis 1115

CONCEPT 47.3 Cytoplasmic determinants and inductive

signals contribute to cell fate specification 1117

Fate Mapping 1117

Axis Formation 1119

Cell Fate Determination and Pattern Formation by Inductive

Signals 1121

Climate Change and Development 1124

48 Neurons, Synapses,

and Signalling 1129

Lines of Communication 1129

CONCEPT 48.1 Neuron organization and structure reflect

function in information transfer 1130

Neuron Structure and Function 1130

Introduction to Information Processing 1130

CONCEPT 48.2 Ion pumps and ion channels establish the resting potential of a neuron 1132

The Resting Membrane Potential 1132

Modelling the Resting Membrane Potential 1133

CONCEPT 48.3 Action potentials are the signals conducted by axons 1134

Hyperpolarization and Depolarization 1135

Graded Potentials and Action Potentials 1135

Generation of Action Potentials: A Closer Look 1136

Conduction of Action Potentials 1137

CONCEPT 48.4 Neurons communicate with other cells at synapses 1139

Generation of Postsynaptic Potentials 1139

Summation of Postsynaptic Potentials 1141

Termination of Neurotransmitter Signalling 1142

Modulated Signalling at Synapses 1142

Neurotransmitters 1142

49 Nervous Systems 1147

Command and Control Centre 1147

CONCEPT 49.1 Nervous systems consist of circuits

of neurons and supporting cells 1148

Glia 1149

Organization of the Vertebrate Nervous System 1150

The Peripheral Nervous System 1151

CONCEPT 49.2 The vertebrate brain is regionally

specialized 1153

Arousal and Sleep 1153

Biological Clock Regulation 1156

Emotions 1156

Functional Imaging of the Brain 1158

CONCEPT 49.3 The cerebral cortex controls voluntary

movement and cognitive functions 1158

Information Processing 1158

Language and Speech 1160

Lateralization of Cortical Function 1160

Frontal Lobe Function 1160

Evolution of Cognition in Vertebrates 1160

CONCEPT 49.4 Changes in synaptic connections underlie

memory and learning 1161

Neural Plasticity 1162

Memory and Learning 1162

Long-Term Potentiation 1163

CONCEPT 49.5 Many nervous system disorders can be

explained in molecular terms 1164

Schizophrenia 1165

Depression 1165

The Brain’s Reward System and Drug Addiction 1166

Alzheimer’s Disease 1166

Parkinson’s Disease 1167

Future Directions 1167

50 Sensory and Motor

Mechanisms 1170

Sense and Sensibility 1170

CONCEPT 50.1 Sensory receptors transduce stimulus

energy and transmit signals to the central nervous

system 1171

Sensory Reception and Transduction 1171

Transmission 1172

Perception 1172

Amplification and Adaptation 1172

Types of Sensory Receptors 1173

CONCEPT 50.2 The mechanoreceptors responsible for

hearing and equilibrium detect moving fluid or settling

particles 1175

Sensing of Gravity and Sound in Invertebrates 1175

Hearing and Equilibrium in Mammals 1175

Hearing and Equilibrium in Other Vertebrates 1179

CONCEPT 50.3 Visual receptors in animals depend on

light-absorbing pigments 1180

Evolution of Visual Perception 1180

The Vertebrate Visual System 1181

CONCEPT 50.4 The senses of taste and smell rely on similar

sets of sensory receptors 1186

Taste in Mammals 1187

Smell in Humans 1188

CONCEPT 50.5 The physical interaction of protein filaments

is required for muscle function 1189

Vertebrate Skeletal Muscle 1189

Other Types of Muscle 1195

CONCEPT 50.6 Skeletal systems transform muscle

contraction into locomotion 1196

Types of Skeletal Systems 1196

Types of Locomotion 1197

51 Animal Behaviour 1203

The How and Why of Animal Activity 1203

CONCEPT 51.1 Discrete sensory inputs can stimulate both

simple and complex behaviours 1204

Fixed Action Patterns 1204

Migration 1204

Behavioural Rhythms 1205

Animal Signals and Communication 1205

CONCEPT 51.2 Learning establishes specific links between

experience and behaviour 1207

Experience and Behaviour 1208

Learning 1208

CONCEPT 51.3 Selection for individual survival and

reproductive success can explain most behaviours 1213

Foraging Behaviour 1213

Mating Behaviour and Mate Choice 1214

CONCEPT 51.4 Genetic analyses and the concept of inclusive

fitness provide a basis for studying the evolution of

behaviour 1219

Genetic Basis of Behaviour 1219

Genetic Variation and the Evolution of Behaviour 1220

Altruism 1221

Inclusive Fitness 1222

Evolution and Human Culture 1223


Interview: Erin Bertrand 1227

52 An Introduction to Ecology

and the Biosphere 1230

Life on and in the Ice 1230

CONCEPT 52.1 Earth’s climate varies by latitude and season

and is changing rapidly 1231

Global Climate Patterns 1231

Regional and Local Effects on Climate 1231

Microclimate 1235

Global Climate Change 1236

CONCEPT 52.2 The structure and distribution of terrestrial

biomes are controlled by climate and disturbance 1237

Climate and Terrestrial Biomes 1238

General Features of Terrestrial Biomes 1239

Disturbance and Terrestrial Biomes 1239

CONCEPT 52.3 Aquatic biomes are diverse and dynamic

systems that cover most of Earth 1244

Zonation in Aquatic Biomes 1244

CONCEPT 52.4 Interactions between organisms and the

environment limit the distribution of species 1245

Dispersal and Distribution 1250

Abiotic Factors 1251

Biotic Factors 1252

CONCEPT 52.5 Ecological change and evolution affect one

another over time 1254

53 Population Ecology 1258

Sable Island, a Natural Laboratory 1258

CONCEPT 53.1 Dynamic biological processes influence population density, dispersion, and demographics 1259

Density and Dispersion 1259

Demographics 1261

CONCEPT 53.2 The exponential model describes population growth in an idealized, unlimited environment 1264

Change in Population Size 1264

Exponential Growth 1264

CONCEPT 53.3 The logistic model describes how a population grows more slowly as it nears its carrying capacity 1265

The Logistic Growth Model 1266

The Logistic Model and Real Populations 1267

CONCEPT 53.4 Life history traits are products of natural selection 1268

Evolution and Life History Diversity 1268

“Trade-offs” and Life Histories 1269

CONCEPT 53.5 Density-dependent factors regulate population growth 1271

Population Dynamics 1272

CONCEPT 53.6 The human population is no longer growing exponentially but is still increasing rapidly 1275

The Global Human Population 1275

Global Carrying Capacity 1277

54 Community Ecology 1282

Dynamic Communities 1282

CONCEPT 54.1 Community interactions are classified by whether they help, harm, or have no effect on the species

involved 1283

Competition 1283

Exploitation 1285

Positive Interactions 1288

Facilitation 1290

CONCEPT 54.2 Diversity and trophic structure characterize biological communities 1290

Species Diversity 1290

Diversity and Community Stability 1291

Trophic Structure 1291

Species with a Large Impact 1293

Bottom-Up and Top-Down Controls 1294

CONCEPT 54.3 Disturbance influences species diversity and composition 1296

Characterizing Disturbance 1297

Ecological Succession 1298

Human Disturbance 1300

CONCEPT 54.4 Biogeographic factors affect community diversity 1300

Latitudinal Gradients 1300

Area Effects 1301

Island Equilibrium Model 1301

CONCEPT 54.5 Pathogens alter community structure locally and globally 1302

Pathogens and Communities 1303

Community Ecology and Zoonotic Diseases 1303

55 Ecosystems and Restoration

Ecology 1308

Engineering Salmon 1308

CONCEPT 55.1 Physical laws govern energy flow and chemical cycling in ecosystems 1309

Conservation of Energy 1309

Conservation of Mass 1310

Energy, Mass, and Trophic Levels 1310

CONCEPT 55.2 Energy and other limiting factors control primary production in ecosystems 1311

Ecosystem Energy Budgets 1311

Primary Production in Aquatic Ecosystems 1313

Primary Production in Terrestrial Ecosystems 1315

CONCEPT 55.3 Energy transfer between trophic levels is typically only 10% efficient 1317

Production Efficiency 1317

Trophic Efficiency and Ecological Pyramids 1317

CONCEPT 55.4 Biological and geochemical processes cycle nutrients and water in ecosystems 1319

Biogeochemical Cycles 1319

Decomposition and Nutrient Cycling Rates 1322

CONCEPT 55.5 Restoration ecologists help return degraded ecosystems to a more natural state 1323

Bioremediation 1324

Biological Augmentation 1325

Restoration Projects Worldwide 1325

56 Conservation Biology and Global Change 1330

What Is Going Wrong with the Right Whales? 1330

CONCEPT 56.1 Human activities threaten Earth’s biodiversity 1331

Three Levels of Biodiversity 1331

Biodiversity and Human Welfare 1333

Threats to Biodiversity 1334

CONCEPT 56.2 Population conservation focuses on

population size, genetic diversity, and critical habitat 1336

Small-Population Approach 1336

Field Study: The Greater Prairie Chicken and the Extinction Vortex 1337

Field Study: Analysis of Grizzly Bear Populations 1338

Declining-Population Approach 1338

Field Study: Decline of the Rufa Red Knot 1339

Weighing Conflicting Demands 1340

CONCEPT 56.3 Landscape and regional conservation help sustain biodiversity 1340

Landscape Structure and Biodiversity 1340

Establishing Protected Areas 1341

Urban Ecology 1344

CONCEPT 56.4 Earth is changing rapidly as a result of human actions 1344

Acid Precipitation 1344

Nutrient Enrichment 1345

Toxins in the Environment 1346

Greenhouse Gases and Climate Change 1347

Depletion of Atmospheric Ozone 1347

CONCEPT 56.5 Sustainable development can improve human lives while conserving biodiversity 1351

Sustainable Development 1351

Field Study: Sustainable Development in Costa Rica 1352

The Future of the Biosphere 1353

APPENDIX A Answers A-1

APPENDIX B Periodic Table of the Elements B-1

APPENDIX C A Comparison of the Light Microscope

and the Electron Microscope C-1

APPENDIX D Classification of Life D-1

APPENDIX E Scientific Skills Review E – 1



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