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Detailed Contents
1 Biology and the Tree of Life 1
1.1 What Does It Mean to Say That Something Is Alive? 1
1.2 The Cell Theory 2
Are All Organisms Made of Cells? 2 Where Do Cells Come From? 2
1.3 The Theory of Evolution by Natural Selection 4 What Is Evolution? 4
What Is Natural Selection? 4
1.4 The Tree of Life 5
Using Molecules to Understand the Tree of Life 6
How Should We Name Branches on the Tree of Life? 7
1.5 Doing Biology 8
The Nature of Science 8
Why Do Giraffes Have Long Necks? An Introduction to Hypothesis Testing 9
How Do Ants Navigate? An Introduction to Experimental Design 10
CHAPTER REVIEW 13
UNIT 1 THE MOLECULES OF LIFE 15
2 Water and Carbon: The Chemical Basis of Life 15
2.1 Atoms, Ions, and Molecules: The Building Blocks of Chemical Evolution 16
Basic Atomic Structure 16
How Does Covalent Bonding Hold Molecules Together? 17 Ionic Bonding, Ions, and the Electron-Sharing Continuum 18 Some Simple Molecules Formed from C, H, N, and O 19
The Geometry of Simple Molecules 20
Representing Molecules 20
Basic Concepts in Chemical Reactions 21
2.2 The Early Oceans and the Properties of Water 22 Why Is Water Such an Efficient Solvent? 22
How Does Water’s Structure Correlate with Its Properties? 22
Acid–Base Reactions Involve a Transfer of Protons 25
2.3 Chemical Reactions, Chemical Evolution, and Chemical Energy 27
How Do Chemical Reactions Happen? 27
What Is Energy? 27
Chemical Evolution: A Model System 29
How Did Chemical Energy Change during Chemical
Evolution? 33
2.4 The Importance of Carbon 33 Linking Carbon Atoms Together 34
Functional Groups 34
CHAPTER REVIEW 36
3 Protein Structure and Function 38
3.1 Early Origin-of-Life Experiments 39
3.2 Amino Acids and Polymerization 40
The Structure of Amino Acids 40
The Nature of Side Chains 40
How Do Amino Acids Link to Form Proteins? 42
3.3 Proteins Are the Most Versatile Large Molecules in Cells 45
3.4 What Do Proteins Look Like? 45
Primary Structure 46 Secondary Structure 46 Tertiary Structure 47
Quaternary Structure 48 Folding and Function 50
3.5 Enzymes: An Introduction to Catalysis 51
Enzymes Help Reactions Clear Two Hurdles 51
How Do Enzymes Work? 53
Was the First Living Entity a Protein Catalyst? 56
CHAPTER REVIEW 57
4 Nucleic Acids and the RNA World 59
4.1 What Is a Nucleic Acid? 59
Could Chemical Evolution Result in the Production of Nucleotides? 60
How Do Nucleotides Polymerize to Form Nucleic Acids? 61
4.2 DNA Structure and Function 62
What Is the Nature of DNA’s Secondary Structure? 62
DNA Functions as an Information-Containing Molecule 65
Is DNA a Catalytic Molecule? 65
4.3 RNA Structure and Function 66
Structurally, RNA Differs from DNA 66
RNA’s Structure Makes It an Extraordinarily Versatile Molecule 67
RNA Is an Information-Containing Molecule 67
RNA Can Function as a Catalytic Molecule 68
4.4 The First Life-Form 68
CHAPTER REVIEW 69
5 An Introduction to Carbohydrates 71
5.1 Sugars as Monomers 71
How Monosaccharides Differ 72
Monosaccharides and Chemical Evolution 73
5.2 The Structure of Polysaccharides 73
Starch: A Storage Polysaccharide in Plants 74
Glycogen: A Highly Branched Storage Polysaccharide in Animals 74
Cellulose: A Structural Polysaccharide in Plants 76
Chitin: A Structural Polysaccharide in Fungi and Animals 76
Peptidoglycan: A Structural Polysaccharide in Bacteria 76
Polysaccharides and Chemical Evolution 76
5.3 What Do Carbohydrates Do? 77
The Role of Carbohydrates as Structural Molecules 77
The Role of Carbohydrates in Cell Identity 77
The Role of Carbohydrates in Energy Storage 78
CHAPTER REVIEW 80
6 Lipids, Membranes, and the First Cells 82
6.1 Lipids 83
A Look at Three Types of Lipids Found in Cells 83
The Structures of Membrane Lipids 84
6.2 Phospholipid Bilayers 85
Artificial Membranes as an Experimental System 85
Selective Permeability of Lipid Bilayers 86
How Does Lipid Structure Affect Membrane Properties? 87
How Does Temperature Affect the Fluidity and Permeability of Membranes? 88
6.3 Why Molecules Move across Lipid Bilayers: Diffusion and Osmosis 89
Diffusion 89
Osmosis 90
6.4 Membrane Proteins 92
Evolution of the Fluid-Mosaic Model 92
Systems for Studying Membrane Proteins 94 Protein Transport I: Facilitated Diffusion via Channel Proteins 94
Protein Transport II: Facilitated Diffusion via Carrier Proteins 96
Protein Transport III: Active Transport by Pumps 97 Plasma Membranes and the Intracellular Environment 98
CHAPTER REVIEW 100
UNIT 2 CELL STRUCTURE AND FUNCTION 102
7 Inside the Cell 102
7.1 Bacterial and Archaeal Cell Structures and Their Functions 102
A Revolutionary New View 103
Prokaryotic Cell Structures: A Parts List 103
7.2 Eukaryotic Cell Structures and Their Functions 105
The Benefits of Organelles 107
Eukaryotic Cell Structures: A Parts List 107
7.3 Putting the Parts into a Whole 115
Structure and Function at the Whole-Cell Level 115
The Dynamic Cell 116
7.4 Cell Systems I: Nuclear Transport 116
Structure and Function of the Nuclear Envelope 116
How Are Molecules Imported into the Nucleus? 117
7.5 Cell Systems II: The Endomembrane System Manufactures and Ships Proteins 118 Studying the Pathway through the Endomembrane System 119
Entering the Endomembrane System: The Signal Hypothesis 120
Moving from the ER to the Golgi 122
What Happens inside the Golgi Apparatus? 122
How Do Proteins Reach Their Destinations? 122
7.6 Cell Systems III: The Dynamic Cytoskeleton 123
Actin Filaments 123
Intermediate Filaments 125
Microtubules 125
Flagella and Cilia: Moving the Entire Cell 127
CHAPTER REVIEW 129
8 Cell-Cell Interactions 131
8.1 The Cell Surface 132
The Structure and Function of an Extracellular Layer 132
The Cell Wall in Plants 132
The Extracellular Matrix in Animals 133
8.2 How Do Adjacent Cells Connect and Communicate? 134
Cell-Cell Attachments in Eukaryotes 135
Cells Communicate via Cell-Cell Gaps 138
8.3 How Do Distant Cells Communicate? 139
Cell-Cell Signaling in Multicellular Organisms 139
Signal Reception 140
Signal Processing 140
Signal Response 144
Signal Deactivation 144
Cross-Talk: Synthesizing Input from Many Signals 145
Quorum Sensing in Bacteria 145
CHAPTER REVIEW 146
9 Cellular Respiration and Fermentation 148
9.1 The Nature of Chemical Energy and Redox Reactions 149
The Structure and Function of ATP 149 What Is a Redox Reaction? 151
9.2 An Overview of Cellular Respiration 153
9.3 Glycolysis: Processing Glucose to Pyruvate 155
Glycolysis Is a Sequence of 10 Reactions 155
How Is Glycolysis Regulated? 156
9.4 Processing Pyruvate to Acetyl CoA 156
9.5 The Citric Acid Cycle: Oxidizing Acetyl CoA to CO2 158
How Is the Citric Acid Cycle Regulated? 158
What Happens to the NADH and FADH2? 160
9.6 Electron Transport and Chemiosmosis: Building a Proton Gradient to Produce ATP 161
Components of the Electron Transport Chain 161
The Chemiosmosis Hypothesis 162
How Is the Electron Transport Chain Organized? 163
The Discovery of ATP Synthase 164
Organisms Use a Diversity of Electron Acceptors 165
9.7 Fermentation 166
9.8 How Does Cellular Respiration Interact with Other Metabolic Pathways? 168
Catabolic Pathways Break Down Molecules as Fuel 168
Anabolic Pathways Synthesize Key Molecules 169
CHAPTER REVIEW 169
10 Photosynthesis 172
10.1 Photosynthesis Harnesses Sunlight to Make Carbohydrate 172
Photosynthesis: Two Linked Sets of Reactions 173
Photosynthesis Occurs in Chloroplasts 174
10.2 How Does Chlorophyll Capture Light Energy? 174
Photosynthetic Pigments Absorb Light 175
When Light Is Absorbed, Electrons Enter an Excited State 177
10.3 The Discovery of Photosystems I and II 179
How Does Photosystem II Work? 180
How Does Photosystem I Work? 182
The Z Scheme: Photosystems II and I Work Together 182
10.4 How Is Carbon Dioxide Reduced to Produce Glucose? 184
The Calvin Cycle Fixes Carbon 185
The Discovery of Rubisco 186
Carbon Dioxide Enters Leaves through Stomata 187
Mechanisms for Increasing CO2 Concentration 187 How Is Photosynthesis Regulated? 189
What Happens to the Sugar That Is Produced by Photosynthesis? 189
CHAPTER REVIEW 190
The Big Picture: Energy for Life 192
11 The Cell Cycle 194
11.1 Mitosis and the Cell Cycle 195
What Is a Chromosome? 195
Cells Alternate between M Phase and Interphase 196
The Discovery of S Phase 196
The Discovery of the Gap Phases 196
The Cell Cycle 196
11.2 How Does Mitosis Take Place? 197
Events in Mitosis 197
Cytokinesis Results in Two Daughter Cells 200
How Do Chromosomes Move during Mitosis? 201
11.3 Control of the Cell Cycle 202
The Discovery of Cell-Cycle Regulatory Molecules 203
Cell-Cycle Checkpoints Can Arrest the Cell Cycle 204
11.4 Cancer: Out-of-Control Cell Division 206
Properties of Cancer Cells 206
Cancer Involves Loss of Cell-Cycle Control 207
CHAPTER REVIEW 209
UNIT 3 GENE STRUCTURE AND EXPRESSION 211
12 Meiosis 211
12.1 How Does Meiosis Occur? 212 Chromosomes Come in Distinct Types 212
The Concept of Ploidy 212
An Overview of Meiosis 213 The Phases of Meiosis I 216
The Phases of Meiosis II 218 A Closer Look at Prophase I 219
12.2 The Consequences of Meiosis 220
Chromosomes and Heredity 221
Independent Assortment Produces Genetic Variation 221
The Role of Crossing Over 222
How Does Fertilization Affect Genetic Variation? 222
12.3 Why Does Meiosis Exist? 223
The Paradox of Sex 223
The Purifying Selection Hypothesis 224
The Changing-Environment Hypothesis 224
12.4 Mistakes in Meiosis 225
How Do Mistakes Occur? 225
Why Do Mistakes Occur? 226
CHAPTER REVIEW 227
13 Mendel and the Gene 230
13.1 Mendel’s Experimental System 230
What Questions Was Mendel Trying to Answer? 231
Garden Peas Served as the First Model Organism in Genetics 231
13.2 Mendel’s Experiments with a Single Trait 232
The Monohybrid Cross 232
Particulate Inheritance 234
13.3 Mendel’s Experiments with Two Traits 236
The Dihybrid Cross 236
Using a Testcross to Confirm Predictions 238
13.4 The Chromosome Theory of Inheritance 239
Meiosis Explains Mendel’s Principles 240
Testing the Chromosome Theory 241
13.5 Extending Mendel’s Rules 243
Linkage: What Happens When Genes Are Located on the Same Chromosome? 243
Do Heterozygotes Always Have a Dominant or Recessive Phenotype? 245
BOX 13.1 QUANTITATIVE METHODS: Linkage 245
How Many Alleles and Phenotypes Exist? 247
Does Each Gene Affect Just One Trait? 247
Are Phenotypes Determined by Genes? 247
What About Traits Like Human Height and
Intelligence? 248
13.6 Applying Mendel’s Rules to Humans 250
Identifying Human Alleles as Recessive or Dominant 250
Identifying Human Traits as Autosomal or Sex-Linked 251
CHAPTER REVIEW 253
14 DNA and the Gene: Synthesis and Repair 258
14.1 What Are Genes Made Of? 259
The Hershey-Chase Experiment 259
The Secondary Structure of DNA 260
14.2 Testing Early Hypotheses about DNA Synthesis: The Meselson-Stahl Experiment 261
14.3 A Comprehensive Model for DNA Synthesis 263
How Does Replication Get Started? 264
How Is the Helix Opened and Stabilized? 264
How Is the Leading Strand Synthesized? 265 How Is the Lagging Strand Synthesized? 266
14.4 Replicating the Ends of Linear Chromosomes 269
14.5 Repairing Mistakes and Damage 271
Correcting Mistakes in DNA Synthesis 271
Repairing Damaged DNA 272
Xeroderma Pigmentosum: A Case Study 272
CHAPTER REVIEW 274
15 How Genes Work 276
15.1 What Do Genes Do? 277
The One-Gene, One-Enzyme Hypothesis 277
An Experimental Test of the Hypothesis 277
15.2 The Central Dogma of Molecular Biology 279
The Genetic Code Hypothesis 279
RNA as the Intermediary between Genes and Proteins 279
Dissecting the Central Dogma 280
15.3 The Genetic Code 282
How Long Is a Word in the Genetic Code? 282
How Did Researchers Crack the Code? 283
15.4 What Is the Molecular Basis of Mutation? 285
Point Mutation 285
Chromosome-Level Mutations 286
CHAPTER REVIEW 287
16 Transcription, RNA Processing, and Translation 289
16.1 An Overview of Transcription 289
Characteristics of RNA Polymerase 290
Initiation: How Does Transcription Begin? 291 Elongation and Termination 292
16.2 RNA Processing in Eukaryotes 293
The Startling Discovery of Eukaryotic Genes in Pieces 293
RNA Splicing 294
Adding Caps and Tails to Transcripts 295 16.3 An Introduction to Translation 295
Ribosomes Are the Site of Protein Synthesis 295
Comparing Translation in Bacteria and Eukaryotes 296 How Does an mRNA Triplet Specify an Amino Acid? 297
16.4 The Structure and Function of Transfer RNA 297
What Do tRNAs Look Like? 299
How Many tRNAs Are There? 299
16.5 The Structure and Function of Ribosomes 300
Initiating Translation 301
Elongation: Extending the Polypeptide 301
Terminating Translation 302
Post-Translational Modifications 304
CHAPTER REVIEW 304
17 Control of Gene Expression in Bacteria 307
17.1 Gene Regulation and Information Flow 307
Mechanisms of Regulation—An Overview 308
Metabolizing Lactose—A Model System 309
17.2 Identifying Genes under Regulatory Control 310
Replica Plating to Find Mutant Genes 310
Different Classes of Lactose Metabolism Mutants 311
Several Genes Are Involved in Lactose Metabolism 312
17.3 Mechanisms of Negative Control: Discovery of the Repressor 312
The lac Operon 313
Why Has the lac Operon Model Been So Important? 314
17.4 Mechanisms of Positive Control: Catabolite Repression 314
The CAP Protein and Binding Site 315
How Does Glucose Influence Formation of the CAP–cAMP Complex? 316
CHAPTER REVIEW 317
18 Control of Gene Expression in Eukaryotes 319
18.1 Mechanisms of Gene Regulation in Eukaryotes— An Overview 320
18.2 Chromatin Remodeling 320
What Is Chromatin’s Basic Structure? 320
Evidence That Chromatin Structure Is Altered in Active Genes 321
How Is Chromatin Altered? 322
Chromatin Modifications Can Be Inherited 323
18.3 Initiating Transcription: Regulatory Sequences and Regulatory Proteins 323
Some Regulatory Sequences Are Near the Promoter 323
Some Regulatory Sequences Are Far from the Promoter 324
The Role of Regulatory Proteins in Differential Gene
Expression 326
The Initiation Complex 326
18.4 Post-Transcriptional Control 328
Alternative Splicing of mRNAs 328
mRNA Stability and RNA Interference 329
How Is Translation Controlled? 330
Post-Translational Control 330
18.5 How Does Gene Expression in Bacteria Compare with That in Eukaryotes? 331
18.6 Linking Cancer with Defects in Gene Regulation 332
Causes of Uncontrolled Cell Growth 332
p53: A Case Study 332
CHAPTER REVIEW 333
The Big Picture: Genetic Information 336
19 Analyzing and Engineering Genes 338
19.1 Case 1—The Effort to Cure Pituitary Dwarfism: Basic Recombinant DNA Technologies 338
Why Did Early Efforts to Treat the Disease Fail? 339
Steps in Engineering a Safe Supply of Growth Hormone 339
Ethical Concerns over Recombinant Growth Hormone 343
19.2 Case 2—Amplification of Fossil DNA: The Polymerase Chain Reaction 344
Requirements of PCR 344
PCR in Action 345
19.3 Case 3—Sanger’s Breakthrough Innovation: Dideoxy DNA Sequencing 346
The Logic of Dideoxy Sequencing 346
“Next Generation” Sequencing 347
19.4 Case 4—The Huntington’s Disease Story: Finding Genes by Mapping 348
How Was the Huntington’s Disease Gene Found? 348
What Are the Benefits of Finding a Disease Gene? 350
Ethical Concerns over Genetic Testing 350
19.5 Case 5—Severe Immune Disorders: The Potential of Gene Therapy 351
How Can Novel Alleles Be Introduced into Human Cells? 351
Using Gene Therapy to Treat X-Linked Immune Deficiency 352
Ethical Concerns over Gene Therapy 354
19.6 Case 6—The Development of Golden Rice: Biotechnology in Agriculture 354
Rice as a Target Crop 355
Synthesizing (3-Carotene in Rice 355
The Agrobacterium Transformation System 355
Using the Ti Plasmid to Produce Golden Rice 356
CHAPTER REVIEW 356
20 Genomics 359
20.1 Whole-Genome Sequencing 359
How Are Complete Genomes Sequenced? 360
Which Genomes Are Being Sequenced, and Why? 361
Which Sequences Are Genes? 362
20.2 Bacterial and Archaeal Genomes 363
The Natural History of Prokaryotic Genomes 363
Lateral Gene Transfer 364
Environmental Sequencing 364
20.3 Eukaryotic Genomes 365
Parasitic and Repeated Sequences 365
Gene Families 367
Insights from the Human Genome Project 368
20.4 Functional Genomics and Proteomics 370
What Is Functional Genomics? 370
What Is Proteomics? 371
Applied Genomics in Action: Understanding Cancer 371
CHAPTER REVIEW 372
UNIT 4 DEVELOPMENTAL BIOLOGY 374
21 Principles of Development 374
21.1 Shared Developmental Processes 375 Cell Proliferation 375
Programmed Cell Death 376
Cell Movement or Cell Growth 376 Cell Differentiation 377
Cell-Cell Interactions 377
21.2 The Role of Differential Gene Expression in Development 377
Evidence That Differentiated Plant Cells Are Genetically
Equivalent 377
Evidence That Differentiated Animal Cells Are Genetically
Equivalent 377
How Does Differential Gene Expression Occur? 378
21.3 Cell-Cell Signals Trigger Differential Gene Expression 379
Master Regulators Set Up the Major Body Axes 379
Regulatory Genes Provide Increasingly Specific Positional Information 381
Cell-Cell Signals and Regulatory Genes Are Evolutionarily Conserved 383
Common Signaling Pathways Are Active in Many Contexts 383
21.4 Changes in Developmental Pathways Underlie Evolutionary Change 384
CHAPTER REVIEW 385
22 An Introduction to Animal Development 388
22.1 Gamete Structure and Function 389
Sperm Structure and Function 389
Egg Structure and Function 390
22.2 Fertilization 390
How Do Gametes from the Same Species Recognize Each Other? 391
Why Does Only One Sperm Enter the Egg? 391
22.3 Cleavage 392
Partitioning Cytoplasmic Determinants 393
Cleavage in Mammals 393
22.4 Gastrulation 394
Formation of Germ Layers 394
Definition of Body Axes 395
22.5 Organogenesis 396
Organizing Mesoderm into Somites: Precursors of Muscle, Skeleton, and Skin 396
Differentiation of Muscle Cells 398
CHAPTER REVIEW 399
23 An Introduction to Plant Development 401
23.1 Gametogenesis, Pollination, and Fertilization 402
How Are Sperm and Egg Produced? 402
Pollen–Stigma Interactions 402
Double Fertilization 403
23.2 Embryogenesis 404
What Happens during Plant Embryogenesis? 404
Which Genes and Proteins Set Up Body Axes? 406
23.3 Vegetative Development 407
Meristems Provide Lifelong Growth and Development 407
Which Genes and Proteins Determine Leaf Shape? 408
23.4 Reproductive Development 409
The Floral Meristem and the Flower 409
The Genetic Control of Flower Structures 409
CHAPTER REVIEW 412
UNIT 5 EVOLUTIONARY PROCESSES AND PATTERNS 414
24 Evolution by Natural Selection 414
24.1 The Evolution of Evolutionary Thought 415
Plato and Typological Thinking 415
Aristotle and the Great Chain of Being 415
Lamarck and the Idea of Evolution as Change
through Time 415
Darwin and Wallace and Evolution by Natural Selection 415
24.2 The Pattern of Evolution: Have Species Changed through Time? 416
Evidence for Change through Time 416
Evidence of Descent from a Common Ancestor 418
Evolution’s “Internal Consistency”—the Importance of Independent Datasets 422
24.3 The Process of Evolution: How Does Natural Selection Work? 422
Darwin’s Four Postulates 423
The Biological Definitions of Fitness and Adaptation 424
24.4 Evolution in Action: Recent Research on Natural Selection 424
Case Study 1: How Did Mycobacterium tuberculosis Become Resistant to Antibiotics? 424
Case Study 2: Why Are Beak Size, Beak Shape, and Body Size Changing in Galápagos Finches? 426
24.5 Common Misconceptions about Natural Selection and Adaptation 429
Selection Acts on Individuals, but Evolutionary Change Occurs in Populations 429
Evolution Is Not Goal Directed 430
Organisms Do Not Act for the Good of the Species 430
Limitations of Natural Selection 431
CHAPTER REVIEW 432
25 Evolutionary Processes 435
25.1 Analyzing Change in Allele Frequencies: The Hardy-Weinberg Principle 436
The Gene Pool Concept 436
Deriving the Hardy-Weinberg Principle 436
The Hardy-Weinberg Model Makes Important
Assumptions 437
How Does the Hardy-Weinberg Principle Serve as a Null Hypothesis? 438
25.2 Types of Natural Selection 440
Directional Selection 440
Stabilizing Selection 441
Disruptive Selection 442
Balancing Selection 442
25.3 Genetic Drift 443
Simulation Studies of Genetic Drift 443
Experimental Studies of Genetic Drift 445
What Causes Genetic Drift in Natural Populations? 445
25.4 Gene Flow 447
Gene Flow in Natural Populations 447
How Does Gene Flow Affect Fitness? 448
25.5 Mutation 448
Mutation as an Evolutionary Mechanism 448
Experimental Studies of Mutation 449
25.6 Nonrandom Mating 450 Inbreeding 450
Sexual Selection 452
CHAPTER REVIEW 456
26 Speciation 458
26.1 How Are Species Defined and Identified? 458
The Biological Species Concept 459
The Morphospecies Concept 460
The Phylogenetic Species Concept 460
Species Definitions in Action: The Case of the Dusky Seaside Sparrow 461
26.2 Isolation and Divergence in Allopatry 462
Dispersal and Colonization Isolate Populations 463
Vicariance Isolates Populations 464
26.3 Isolation and Divergence in Sympatry 464
Can Natural Selection Cause Speciation Even When Gene Flow Is Possible? 465
How Can Polyploidy Lead to Speciation? 465
26.4 What Happens When Isolated Populations Come into Contact? 468
Reinforcement 468
Hybrid Zones 468
New Species through Hybridization 470
CHAPTER REVIEW 472
27 Phylogenies and the History of Life 474
27.1 Tools for Studying History: Phylogenetic Trees 474
How Do Researchers Estimate Phylogenies? 475
How Can Biologists Distinguish Homology from
Homoplasy? 475
Whale Evolution: A Case History 477
27.2 Tools for Studying History: The Fossil Record 479
How Do Fossils Form? 479
Limitations of the Fossil Record 480
Life’s Time Line 481
27.3 Adaptive Radiation 484
Why Do Adaptive Radiations Occur? 484
The Cambrian Explosion 486
27.4 Mass Extinction 488
How Do Mass Extinctions Differ From Background Extinctions? 489
The End-Permian Extinction 489
What Killed the Dinosaurs? 490
CHAPTER REVIEW 492
The Big Picture: Evolution 494
UNIT 6 THE DIVERSIFICATION OF LIFE 496
28 Bacteria and Archaea 496
28.1 Why Do Biologists Study Bacteria and Archaea? 497
Biological Impact 497
Medical Importance 498
Role in Bioremediation 500
Extremophiles 501
28.2 How Do Biologists Study Bacteria and Archaea? 501
Using Enrichment Cultures 501
Using Direct Sequencing 502
Evaluating Molecular Phylogenies 503
28.3 What Themes Occur in the Diversification of Bacteria and Archaea? 504
Morphological Diversity 504
Metabolic Diversity 506
Ecological Diversity and Global Change 509
28.4 Key Lineages of Bacteria and Archaea 512
Bacteria 512
Archaea 512
? Bacteria > Firmicutes 513
? Bacteria > Spirochaetes (Spirochetes) 513
? Bacteria > Actinobacteria 514
? Bacteria > Chlamydiae 514
? Bacteria > Cyanobacteria 515
? Bacteria > Proteobacteria 515
? Archaea > Crenarchaeota 516
? Archaea > Euryarchaeota 516
CHAPTER REVIEW 517
29 Protists 519
29.1 Why Do Biologists Study Protists? 520
Impacts on Human Health and Welfare 520
Ecological Importance of Protists 522
29.2 How Do Biologists Study Protists? 524
Microscopy: Studying Cell Structure 524
Evaluating Molecular Phylogenies 525
Discovering New Lineages via Direct Sequencing 525
29.3 What Themes Occur in the Diversification of Protists? 526
What Morphological Innovations Evolved in Protists? 526
How Do Protists Obtain Food? 529
How Do Protists Move? 532
How Do Protists Reproduce? 533
Life Cycles—Haploid- versus Diploid-Dominated 533
29.4 Key Lineages of Protists 536
Amoebozoa 536
Excavata 536
Plantae 536
Rhizaria 536
Alveolata 537
Stramenopila (Heterokonta) 537
? Amoebozoa > Myxogastrida (Plasmodial Slime
Molds) 537
? Excavata > Parabasalida 538
? Excavata > Diplomonadida 538
? Excavata > Euglenida 539
? Plantae > Rhodophyta (Red Algae) 539
? Rhizaria > Foraminifera 540
? Alveolata > Ciliata 540
? Alveolata > Dinoflagellata 541
? Alveolata > Apicomplexa 541
? Stramenopila > Oomycota (Water Molds) 542
? Stramenopila > Diatoms 542
? Stramenopila > Phaeophyta (Brown Algae) 543
CHAPTER REVIEW 543
30 Green Algae and Land Plants 546
30.1 Why Do Biologists Study the Green Algae and Land Plants? 546
Plants Provide Ecosystem Services 547
Plants Provide Humans with Food, Fuel, Fiber, Building Materials, and Medicines 548
30.2 How Do Biologists Study Green Algae and Land Plants? 549
Analyzing Morphological Traits 549
Using the Fossil Record 550
Evaluating Molecular Phylogenies 551
30.3 What Themes Occur in the Diversification of
Land Plants? 553
The Transition to Land, I: How Did Plants Adapt to Dry
Conditions? 553
Mapping Evolutionary Changes on the Phylogenetic Tree 555
The Transition to Land, II: How Do Plants Reproduce in Dry
Conditions? 556
The Angiosperm Radiation 564
30.4 Key Lineages of Green Algae and Land Plants 566
Green Algae 566
Non-Vascular Plants (“Bryophytes”) 567
Seedless Vascular Plants 567
Seed Plants 567
? Green Algae > Ulvophyceae (Ulvophytes) 568
? Green Algae > Coleochaetophyceae (Coleochaetes) 568
? Green Algae > Charophyceae (Stoneworts) 569
? Non-Vascular Plants > Hepaticophyta (Liverworts) 569
? Non-Vascular Plants > Bryophyta (Mosses) 570
? Non-Vascular Plants > Anthocerophyta (Hornworts) 571
? Seedless Vascular Plants > Lycophyta (Lycophytes, or Club
Mosses) 571
? Seedless Vascular Plants > Psilotophyta (Whisk Ferns) 572
? Seedless Vascular Plants > Equisetophyta (or Sphenophyta) (Horsetails) 572
? Seedless Vascular Plants > Pteridophyta (Ferns) 573
? Seed Plants > Gymnosperms > Cycadophyta (Cycads) 574
? Seed Plants > Gymnosperms > Ginkgophyta
(Ginkgos) 574
? Seed Plants > Gymnosperms > Redwood group (Redwoods, Junipers, Yews) 575
? Seed Plants > Gymnosperms > Pinophyta (Pines, Spruces, Firs) 575
? Seed Plants > Gymnosperms > Gnetophyta
(Gnetophytes) 576
? Seed Plants > Anthophyta (Angiosperms) 576
CHAPTER REVIEW 577
31 Fungi 579
31.1 Why Do Biologists Study Fungi? 580
Fungi Provide Nutrients for Land Plants 580
Fungi Speed the Carbon Cycle on Land 580
Fungi Have Important Economic Impacts 581
31.2 How Do Biologists Study Fungi? 582
Analyzing Morphological Traits 582
Evaluating Molecular Phylogenies 584
Experimental Studies of Mutualism 586
31.3 What Themes Occur in the Diversification of Fungi? 586
Fungi Participate in Several Types of Mutualisms 586
What Adaptations Make Fungi Such Effective
Decomposers? 589
Variation in Reproduction 590
Four Major Types of Life Cycles 591
31.4 Key Lineages of Fungi 594
? Fungi > Microsporidia 594
? Fungi > Chytrids 595
? Fungi > Zygomycetes 595
? Fungi > Glomeromycota 596
? Fungi > Basidiomycota (Club Fungi) 596
? Fungi > Ascomycota > Lichen-Formers 597
? Fungi > Ascomycota > Non-Lichen-Formers 598
CHAPTER REVIEW 599
32 An Introduction to Animals 601
32.1 Why Do Biologists Study Animals? 602
Biological Importance 602
Role in Human Health and Welfare 602
32.2 How Do Biologists Study Animals? 603
Analyzing Comparative Morphology 603
Evaluating Molecular Phylogenies 607
32.3 What Themes Occur in the Diversification
of Animals? 610 Sensory Organs 610 Feeding 610
Movement 613 Reproduction 615 Life Cycles 615
32.4 Key Lineages of Animals: Non-Bilaterian Groups 617
? Porifera (Sponges) 618
? Cnidaria (Jellyfish, Corals, Anemones, Hydroids) 619
? Ctenophora (Comb Jellies) 620
? Acoelomorpha (Acoels) 620
CHAPTER REVIEW 621
33 Protostome Animals 623
33.1 An Overview of Protostome Evolution 624
What Is a Lophotrochozoan? 624
What Is an Ecdysozoan? 625
33.2 Themes in the Diversification of Protostomes 625
How Do Body Plans Vary among Phyla? 626
The Water-to-Land Transition 627
Adaptations for Feeding 628
Adaptations for Moving 628
Adaptations in Reproduction 630
33.3 Key Lineages: Lophotrochozoans 630
? Lophotrochozoans > Rotifera (Rotifers) 631
? Lophotrochozoans > Platyhelminthes (Flatworms) 631
? Lophotrochozoans > Annelida (Segmented Worms) 633
? Lophotrochozoans > Mollusca > Bivalvia (Clams, Mussels,
Scallops, Oysters) 634
? Lophotrochozoans > Mollusca > Gastropoda (Snails, Slugs, Nudibranchs) 635
? Lophotrochozoans > Mollusca > Polyplacophora (Chitons) 636
? Lophotrochozoans > Mollusca > Cephalopoda (Nautilus, Cuttlefish, Squid, Octopuses) 636
33.4 Key Lineages: Ecdysozoans 637
? Ecdysozoans > Nematoda (Roundworms) 638
? Ecdysozoans > Arthropoda > Myriapods (Millipedes, Centipedes) 639
? Ecdysozoans > Arthropoda > Insecta (Insects) 639
? Ecdysozoans > Arthropoda > Chelicerata (Spiders, Ticks,
Mites, Horseshoe Crabs, Daddy Longlegs, Scorpions) 642
? Ecdysozoans > Arthropoda > Crustaceans (Shrimp, Lobster,
Crabs, Barnacles, Isopods, Copepods) 643
CHAPTER REVIEW 644
34 Deuterostome Animals 646
34.1 What Is an Echinoderm? 647
The Echinoderm Body Plan 647
How Do Echinoderms Feed? 648
Key Lineages 649
? Echinodermata > Asteroidea (Sea Stars) 649
? Echinodermata > Echinoidea (Sea Urchins and Sand
Dollars) 650
34.2 What Is a Chordate? 650
Three “Subphyla” 651
Key Lineages: The Invertebrate Chordates 651
? Chordata > Cephalochordata (Lancelets) 652
? Chordata > Urochordata (Tunicates) 652
34.3 What Is a Vertebrate? 653
An Overview of Vertebrate Evolution 653
Key Innovations 655 Key Lineages 660
? Chordata > Vertebrata > Myxinoidea (Hagfish) and
Petromyzontoidea (Lampreys) 661
? Chordata > Vertebrata > Chondrichthyes (Sharks, Rays, Skates) 662
? Chordata > Vertebrata > Actinopterygii (Ray-Finned
Fishes) 662
? Chordata > Vertebrata > Actinistia (Coelacanths) and Dipnoi (Lungfish) 663
? Chordata > Vertebrata > Amphibia (Frogs, Salamanders, Caecilians) 664
? Chordata > Vertebrata > Mammalia > Monotremata (Platypuses, Echidnas) 665
? Chordata > Vertebrata > Mammalia > Marsupiala (Marsupials) 665
? Chordata > Vertebrata > Mammalia > Eutheria (Placental
Mammals) 666
? Chordata > Vertebrata > Reptilia > Lepidosauria (Lizards,
Snakes) 666
? Chordata > Vertebrata > Reptilia > Testudinia (Turtles) 667
? Chordata > Vertebrata > Reptilia > Crocodilia (Crocodiles,
Alligators) 667
? Chordata > Vertebrata > Reptilia > Aves (Birds) 668
34.4 The Primates and Hominins 668 The Primates 668
Fossil Humans 670
The Out-of-Africa Hypothesis 672
CHAPTER REVIEW 673 35 Viruses 675
35.1 Why Do Biologists Study Viruses? 676
Recent Viral Epidemics in Humans 676
Current Viral Epidemics in Humans: HIV 677
35.2 How Do Biologists Study Viruses? 678
Analyzing Morphological Traits 679
Analyzing Variation in Growth Cycles: Replicative and Latent Growth 679
Analyzing the Phases of the Replicative Cycle 681
35.3 What Themes Occur in the Diversification of Viruses? 686
The Nature of the Viral Genetic Material 686
Where Did Viruses Come From? 686
Emerging Viruses, Emerging Diseases 688
35.4 Key Lineages of Viruses 689
? Double-Stranded DNA (dsDNA) Viruses 690
? RNA Reverse-Transcribing Viruses (Retroviruses) 691
? Double-Stranded RNA (dsRNA) Viruses 691
? Negative-Sense Single-Stranded RNA ([—]ssRNA)
Viruses 692
? Positive-Sense Single-Stranded RNA ([+]ssRNA)
Viruses 692
CHAPTER REVIEW 693
UNIT 7 HOW PLANTS WORK 695
36 Plant Form and Function 695
36.1 Plant Form: Themes with Many Variations 696
The Importance of Surface Area/Volume Relationships 696
The Root System 697
The Shoot System 699
The Leaf 701
36.2 Primary Growth Extends the Plant Body 704
How Do Apical Meristems Produce the Primary Plant Body? 704
How Is the Primary Root System Organized? 705
How Is the Primary Shoot System Organized? 706
36.3 Cells and Tissues of the Primary Plant Body 706
The Dermal Tissue System 707 The Ground Tissue System 708
The Vascular Tissue System 710
36.4 Secondary Growth Widens Shoots and Roots 712
What Is a Cambium? 712
What Does Vascular Cambium Produce? 713
What Does Cork Cambium Produce? 713
The Structure of a Tree Trunk 714
CHAPTER REVIEW 715
37 Water and Sugar Transport in Plants 717
37.1 Water Potential and Water Movement 717
What Is Water Potential? 718
What Factors Affect Water Potential? 718
Calculating Water Potential 719
Water Potentials in Soils, Plants, and the Atmosphere 720
37.2 How Does Water Move from Roots to Shoots? 721
Movement of Water and Solutes into the Root 722 Water Movement via Root Pressure 723
Water Movement via Capillary Action 723
The Cohesion-Tension Theory 724
37.3 Water Absorption and Water Loss 727
Limiting Water Loss 727
Obtaining Carbon Dioxide under Water Stress 728
37.4 Translocation 728
Tracing Connections between Sources and Sinks 728
The Anatomy of Phloem 729
The Pressure-Flow Hypothesis 730
Phloem Loading 731 Phloem Unloading 733
CHAPTER REVIEW 735
38 Plant Nutrition 737
38.1 Nutritional Requirements of Plants 738
Which Nutrients Are Essential? 738
What Happens When Key Nutrients Are in Short Supply? 740
38.2 Soil: A Dynamic Mixture of Living and Nonliving Components 741
The Importance of Soil Conservation 742
What Factors Affect Nutrient Availability? 742
38.3 Nutrient Uptake 744
Mechanisms of Nutrient Uptake 744
Mechanisms of Ion Exclusion 746
38.4 Nitrogen Fixation 748
The Role of Symbiotic Bacteria 749
How Do Nitrogen-Fixing Bacteria Colonize Plant Roots? 749
38.5 Nutritional Adaptations of Plants 750 Epiphytic Plants 750
Parasitic Plants 751
Carnivorous Plants 751
CHAPTER REVIEW 752
39 Plant Sensory Systems, Signals, and Responses 755
39.1 Information Processing in Plants 756
How Do Cells Receive and Transduce an External Signal? 756
How Are Cell-Cell Signals Transmitted? 756
How Do Cells Respond to Cell-Cell Signals? 757
39.2 Blue Light: The Phototropic Response 758
Phototropins as Blue-Light Receptors 758
Auxin as the Phototropic Hormone 759
39.3 Red and Far-Red Light: Germination and Stem Elongation 762
The Red/Far-Red “Switch” 763
Phytochromes as Red/Far-Red Receptors 763
How Were Phytochromes Isolated? 763
39.4 Gravity: The Gravitropic Response 764
The Statolith Hypothesis 764
Auxin as the Gravitropic Signal 765
39.5 How Do Plants Respond to Wind and Touch? 766
Changes in Growth Patterns 766
Movement Responses 766
39.6 Youth, Maturity, and Aging: The Growth Responses 767
Auxin and Apical Dominance 767
Cytokinins and Cell Division 768
Gibberellins and ABA: Growth and Dormancy 769
Brassinosteroids and Body Size 773
Ethylene and Senescence 773
An Overview of Plant Growth Regulators 774
39.7 Pathogens and Herbivores: The Defense
Responses 776
How Do Plants Sense and Respond to Pathogens? 776
How Do Plants Sense and Respond to Herbivore Attack? 778
CHAPTER REVIEW 781
40 Plant Reproduction 783
40.1 An Introduction to Plant Reproduction 784
Sexual Reproduction 784
The Land Plant Life Cycle 784
Asexual Reproduction 786
40.2 Reproductive Structures 786
When Does Flowering Occur? 787
The General Structure of the Flower 788
How Are Female Gametophytes Produced? 790
How Are Male Gametophytes Produced? 790
40.3 Pollination and Fertilization 792 Pollination 792
Fertilization 794
40.4 The Seed 795
Embryogenesis 796
The Role of Drying in Seed Maturation 797
Fruit Development and Seed Dispersal 797 Seed Dormancy 798
Seed Germination 799
CHAPTER REVIEW 800
UNIT 8 HOW ANIMALS WORK 803
41 Animal Form and Function 803
41.1 Form, Function, and Adaptation 804
The Role of Fitness Trade-Offs 804
Adaptation and Acclimatization 804
41.2 Tissues, Organs, and Systems: How Does Structure Correlate with Function? 806
Structure-Function Relationships at the Molecular and Cellular Levels 806
Tissues Are Groups of Similar Cells That Function as a Unit 806
Organs and Organ Systems 810
41.3 How Does Body Size Affect Animal Physiology? 811
Surface Area/Volume Relationships: Theory 811
Surface Area/Volume Relationships: Data 812
Adaptations That Increase Surface Area 814
41.4 Homeostasis 814
Homeostasis: General Principles 814
The Role of Regulation and Feedback 815
41.5 How Do Animals Regulate Body Temperature? 816
Mechanisms of Heat Exchange 816
Variation in Thermoregulation 816
Endothermy and Ectothermy: A Closer Look 817
Temperature Homeostasis in Endotherms 817
Countercurrent Heat Exchangers 818
CHAPTER REVIEW 820
42 Water and Electrolyte Balance in Animals 822
42.1 Osmoregulation and Osmotic Stress 823
What Is Osmotic Stress? 823
Osmotic Stress in Seawater 824
Osmotic Stress in Freshwater 824
Osmotic Stress on Land 825
How Do Cells Move Electrolytes and Water? 825
42.2 Water and Electrolyte Balance in Aquatic Environments 826
How Do Sharks Excrete Salt? 826
How Do Freshwater Fish Osmoregulate? 827
42.3 Water and Electrolyte Balance in Terrestrial Insects 828
How Do Insects Minimize Water Loss from the Body Surface? 828
Types of Nitrogenous Wastes: Impact on Water Balance 829
Maintaining Homeostasis: The Excretory System 830
42.4 Water and Electrolyte Balance in Terrestrial Vertebrates 832
The Structure of the Kidney 832
The Function of the Kidney: An Overview 832
Filtration: The Renal Corpuscle 832
Reabsorption: The Proximal Tubule 834
Creating an Osmotic Gradient: The Loop of Henle 835
Regulating Water and Electrolyte Balance: The Distal Tubule and Collecting Duct 837
CHAPTER REVIEW 839 43 Animal Nutrition 841
43.1 Nutritional Requirements 842
Meeting Basic Needs in Humans 842
Studying Nutrient Requirements 842
43.2 Capturing Food: The Structure and Function of Mouthparts 843
Mouthparts as Adaptations 843
A Case Study: The Cichlid Jaw 844
43.3 How Are Nutrients Digested and Absorbed? 845
An Introduction to the Digestive Tract 845
An Overview of Digestive Processes 846
The Mouth and Esophagus 847
The Stomach 848
The Small Intestine 851
The Cecum and Appendix 854
The Large Intestine 854
43.4 Nutritional Homeostasis—Glucose as a Case Study 856
The Discovery of Insulin 856
Insulin’s Role in Homeostasis 856
Diabetes Can Take Several Forms 856
The Type 2 Diabetes Mellitus Epidemic 857
CHAPTER REVIEW 858
44 Gas Exchange and Circulation 861
44.1 The Respiratory and Circulatory Systems 861
44.2 Air and Water as Respiratory Media 862
How Do Oxygen and Carbon Dioxide Behave in Air? 862
How Do Oxygen and Carbon Dioxide Behave in Water? 863
44.3 Organs of Gas Exchange 864
Physical Parameters: The Law of Diffusion 864
How Do Fish Gills Work? 865
How Do Insect Tracheae Work? 866
How Do Vertebrate Lungs Work? 867
Homeostatic Control of Ventilation 870
44.4 How Are Oxygen and Carbon Dioxide Transported in Blood? 870
Structure and Function of Hemoglobin 871
CO2 Transport and the Buffering of Blood pH 873
44.5 The Circulatory System 874
What Is an Open Circulatory System? 875
What Is a Closed Circulatory System? 875
How Does the Heart Work? 877
Patterns in Blood Pressure and Blood Flow 882
CHAPTER REVIEW 883
45 Electrical Signals in Animals 885
45.1 Principles of Electrical Signaling 885
Types of Neurons in the Nervous System 886
The Anatomy of a Neuron 886
An Introduction to Membrane Potentials 887
BOX 45.1 QUANTITATIVE METHODS: Using the Nernst
Equation to Calculate Equilibrium Potentials 888
How Is the Resting Potential Maintained? 888
Using Microelectrodes to Measure Membrane Potentials 890
What Is an Action Potential? 890
45.2 Dissecting the Action Potential 891
Distinct Ion Currents Are Responsible for Depolarization and Repolarization 891
How Do Voltage-Gated Channels Work? 891
How Is the Action Potential Propagated? 893
45.3 The Synapse 895
Synapse Structure and Neurotransmitter Release 895
What Do Neurotransmitters Do? 896
Postsynaptic Potentials 897
45.4 The Vertebrate Nervous System 899
What Does the Peripheral Nervous System Do? 899
Functional Anatomy of the CNS 900
How Does Memory Work? 902
CHAPTER REVIEW 904
46 Animal Sensory Systems
and Movement 907
46.1 How Do Sensory Organs Convey Information to the Brain? 908
Sensory Transduction 908
Transmitting Information to the Brain 909
46.2 Hearing 909
How Do Sensory Cells Respond to Sound Waves and Other Forms of Pressure? 909
The Mammalian Ear 910
Sensory Worlds: What Do Other Animals Hear? 912
46.3 Vision 913
The Insect Eye 913
The Vertebrate Eye 914
Sensory Worlds: Do Other Animals See Color? 917
46.4 Taste and Smell 918
Taste: Detecting Molecules in the Mouth 918
Olfaction: Detecting Molecules in the Air 919
46.5 Movement 920
Skeletons 920
Muscle Types 921
How Do Muscles Contract? 922
CHAPTER REVIEW 926
47 Chemical Signals in Animals 929
47.1 Cell-to-Cell Signaling: An Overview 929
Major Categories of Chemical Signals 930
Hormone Signaling Pathways 931
What Makes Up the Endocrine System? 932
Chemical Characteristics of Hormones 933
How Do Researchers Identify a Hormone? 934
47.2 What Do Hormones Do? 935
How Do Hormones Direct Developmental Processes? 935
How Do Hormones Coordinate Responses to
Environmental Change? 937
How Are Hormones Involved in Homeostasis? 938
47.3 How Is the Production of Hormones
Regulated? 940
The Hypothalamus and Pituitary Gland 940
Control of Epinephrine by Sympathetic Nerves 943
47.4 How Do Hormones Act on Target Cells? 943
Steroid Hormones Bind to Intracellular Receptors 943
Hormones That Bind to Cell-Surface Receptors 945
Why Do Different Target Cells Respond in Different Ways? 947
CHAPTER REVIEW 948
48 Animal Reproduction 950
48.1 Asexual and Sexual Reproduction 950
How Does Asexual Reproduction Occur? 951
Switching Reproductive Modes: A Case History 951
Mechanisms of Sexual Reproduction: Gametogenesis 952
48.2 Fertilization and Egg Development 954
External Fertilization 954
Internal Fertilization 954
Unusual Aspects of Mating 955
Why Do Some Females Lay Eggs while Others Give Birth? 956
48.3 Reproductive Structures and Their Functions 957
The Male Reproductive System 957
The Female Reproductive System 959
48.4 The Role of Sex Hormones in Mammalian Reproduction 960
Which Hormones Control Puberty in Mammals? 961
Which Hormones Control the Menstrual Cycle in Mammals? 962
48.5 Pregnancy and Birth in Mammals 967
Gestation and Early Development in Marsupials 967
Major Events during Human Pregnancy 967
How Does the Mother Nourish the Fetus? 968
Birth 970
CHAPTER REVIEW 971
49 The Immune System in Animals 973
49.1 Innate Immunity 974
Barriers to Entry 974
The Innate Immune Response 975
49.2 The Adaptive Immune Response: Recognition 977
An Introduction to Lymphocytes 978
The Discovery of B Cells and T Cells 979
The Clonal-Selection Theory 979
How Does the Immune System Distinguish Self from Nonself? 983
49.3 The Adaptive Immune Response: Activation 984
T-Cell Activation 984
B-Cell Activation and Antibody Secretion 986
49.4 The Adaptive Immune Response: Culmination 987
How Are Bacteria and Other Foreign Cells Killed? 987
How Are Viruses Destroyed? 987
Why Does the Immune System Reject Foreign Tissues and Organs? 988
Responding to Future Infections: Immunological Memory 989
49.5 What Happens When the Immune System Doesn’tWork Correctly? 990
Immunodeficiency Diseases 990
Allergies 990
CHAPTER REVIEW 991
50 An Introduction to Ecology 993
50.1 Areas of Ecological Study 993
Organismal Ecology 994
Population Ecology 994
Community Ecology 994
Ecosystem Ecology 995
How Do Ecology and Conservation Efforts Interact? 995
50.2 Types of Aquatic Ecosystems 995
Nutrient Availability 995
Water Flow 996
Water Depth 996
? Freshwater Environments > Lakes and Ponds 997
? Freshwater Environments > Wetlands 998
? Freshwater Environments > Streams 999
? Freshwater/Marine Environments > Estuaries 1000
? Marine Environments > The Ocean 1000
50.3 Types of Terrestrial Ecosystems 1001
? Terrestrial Biomes > Tropical Wet Forest 1003
? Terrestrial Biomes > Subtropical Deserts 1004
? Terrestrial Biomes > Temperate Grasslands 1005
? Terrestrial Biomes > Temperate Forests 1006
? Terrestrial Biomes > Boreal Forests 1007
? Terrestrial Biomes > Arctic Tundra 1008
50.4 The Role of Climate and the Consequences of Climate Change 1008
Global Patterns in Climate 1009
How Will Global Climate Change Affect Ecosystems? 1011
50.5 Biogeography: Why Are Organisms Found Where They Are? 1013
Abiotic Factors 1013
The Role of History 1013
Biotic Factors 1015
Biotic and Abiotic Factors Interact 1015
CHAPTER REVIEW 1017
51 Behavioral Ecology 1019
51.1 An Introduction to Behavioral Biology 1019
Proximate and Ultimate Causation 1020
Conditional Strategies and Decision Making 1020
Five Questions in Behavioral Ecology 1021
51.2 What Should I Eat? 1021
Foraging Alleles in Drosophila melanogaster 1021
Optimal Foraging in White-Fronted Bee-Eaters 1022
51.3 Who Should I Mate With? 1022
Sexual Activity in Anolis Lizards 1023
How Do Female Barn Swallows Choose Mates? 1024
51.4 Where Should I Live? 1026
How Do Animals Find Their Way on Migration? 1026
Why Do Animals Move with a Change of Seasons? 1027
51.5 How Should I Communicate? 1027
Honeybee Language 1028
Modes of Communication 1029
When Is Communication Honest or Deceitful? 1029 51.6
When Should I Cooperate? 1031
Kin Selection 1031
BOX 51.1 QUANTITATIVE METHODS: Calculating the Coefficient of Relatedness 1032
Reciprocal Altruism 1033
An Extreme Case: Abuse of Non-Kin in Humans 1034
CHAPTER REVIEW 1035
52 Population Ecology 1037
52.1 Demography 1037
Life Tables 1038
The Role of Life History 1039
BOX 52.1 QUANTITATIVE METHODS: Using Life Tables to Calculate Population Growth Rates 1040
52.2 Population Growth 1041
Quantifying the Growth Rate 1041
Exponential Growth 1042
Logistic Growth 1042
BOX 52.2 QUANTITATIVE METHODS: Developing and Applying Population Growth Equations 1043
What Limits Growth Rates and Population Sizes? 1044
52.3 Population Dynamics 1046
How Do Metapopulations Change through Time? 1046
Why Do Some Populations Cycle? 1047
BOX 52.3 QUANTITATIVE METHODS: Mark-Recapture Studies 1048
How Does Age Structure Affect Population Growth? 1050
Analyzing Change in the Growth Rate of Human
Populations 1052
52.4 How Can Population Ecology Help Endangered Species? 1053
Using Life-Table Data 1054
Preserving Metapopulations 1055
CHAPTER REVIEW 1056
53 Community Ecology 1058
53.1 Species Interactions 1058
Three Themes 1059
Competition 1059
Consumption 1063
Mutualism 1068
53.2 Community Structure 1070
How Predictable Are Communities? 1070
How Do Keystone Species Structure Communities? 1072
53.3 Community Dynamics 1073
Disturbance and Change in Ecological Communities 1073
Succession: The Development of Communities after Disturbance 1074
53.4 Species Richness in Ecological Communities 1077
Predicting Species Richness: The Theory of Island Biogeography 1077
Global Patterns in Species Richness 1078
BOX 53.1 QUANTITATIVE METHODS: Measuring Species Diversity 1079
CHAPTER REVIEW 1080
UNIT 9 ECOLOGY 993
54 Ecosystems 1083
54.1 How Does Energy Flow through Ecosystems? 1083
Why Is NPP So Important? 1084
Solar Power: Transforming Incoming Energy to Biomass 1084
Trophic Structure 1085
Energy Transfer between Trophic Levels 1086
Trophic Cascades and Top-Down Control 1087
Biomagnification 1088
Global Patterns in Productivity 1089
What Limits Productivity? 1090
54.2 How Do Nutrients Cycle through Ecosystems? 1092
Nutrient Cycling within Ecosystems 1092
Global Biogeochemical Cycles 1094
54.3 Global Warming 1098
Understanding the Problem 1098 Positive and Negative Feedback 1099
Impact on Organisms 1099
Productivity Changes 1101
CHAPTER REVIEW 1103
55 Biodiversity and Conservation Biology 1105
55.1 What Is Biodiversity? 1106
Biodiversity Can Be Measured and Analyzed at Several Levels 1106
How Many Species Are Living Today? 1107
BOX55.1 QUANTITATIVE METHODS: Extrapolation Techniques 1108
55.2 Where Is Biodiversity Highest? 1109
Hotspots of Biodiversity and Endemism 1109 Conservation Hotspots 1110
55.3 Threats to Biodiversity 1110
Changes in the Nature of the Problem 1110
How Can Biologists Predict Future Extinction Rates? 1114
BOX 55.2 QUANTITATIVE METHODS: Population Viability Analysis 1115
55.4 Why Is Biodiversity Important? 1117
Economic Benefits of Biodiversity 1117
Biological Benefits of Biodiversity 1117
An Ethical Dimension? 1120
55.5 Preserving Biodiversity 1120
Designing Effective Protected Areas 1120
Beyond Protected Areas: A Comprehensive Approach 1121
CHAPTER REVIEW 1123
The Big Picture: Ecology 1126
APPENDIXA: BioSkills B:1
1 The Metric System B:1
2 Reading Graphs B:2
3 Reading a Phylogenetic Tree B:4
4 Some Common Latin and Greek Roots Used in Biology B:6
5 Using Statistical Tests and Interpreting Standard Error Bars B:7
6 Reading Chemical Structures B:8
7 Using Logarithms B:9
8 Making Concept Maps B:10
9 Separating and Visualizing Molecules B:11
10 Biological Imaging: Microscopy and X-Ray Crystallography B:13
11 Separating Cell Components by Centrifugation B:16
12 Cell and Tissue Culture Methods B:17
13 Combining Probabilities B:19
14 Model Organisms B:19
APPENDIX B: Answers A:1
Glossary G:1
Credits C:1
Index I:1