Ninth Edition

Biology
Kenneth A. Mason
University of Iowa

Jonathan B. Losos
Apago PDF Enhancer
Harvard University

Susan R. Singer
Carleton College

based on the work of

Peter H. Raven
Director, Missouri Botanical Gardens; Engelmann
Professor of Botany, Washington University

George B. Johnson
Professor Emeritus of Biology, Washington University
Brief Contents

Preface v 29 Protists 567

30 Green Plants 588
Guided Tour x 31 Fungi 614
Contents xxi 32 Overview of Animal Diversity 633
33 Noncoelomate Invertebrates 649
Part I The Molecular Basis of Life 1 34
35
Coelomate Invertebrates 666
Vertebrates 693
1 The Science of Biology 1
2 The Nature of Molecules and the Properties of Water
3 The Chemical Building Blocks of Life 33
17 Part
VI Plant Form and Function 729

36 Plant Form 729

Part
II The Biology of the Cell 59 37
38
Vegetative Plant Development 753
Transport in Plants 769
4 Cell Structure 59 39 Plant Nutrition and Soils 786
5 Membranes 88 40 Plant Defense Responses 802
6 Energy and Metabolism 107 41 Sensory Systems in Plants 814
7 How Cells Harvest Energy 122 42 Plant Reproduction 839
8 Photosynthesis 147
9 Cell Communication 168
10 How Cells Divide 186 VII
Apago PDF Enhancer
Part Animal Form and Function 863

43 The Animal Body and Principles of Regulation 863

Part
III Genetic and Molecular Biology 207 44
45
The Nervous System 887
Sensory Systems 915
11 Sexual Reproduction and Meiosis 207 46 The Endocrine System 937
12 Patterns of Inheritance 221 47 The Musculoskeletal System 961
13 Chromosomes, Mapping, and the Meiosis–Inheritance 48 The Digestive System 981
Connection 239 49 The Respiratory System 1001
14 DNA: The Genetic Material 256 50 The Circulatory System 1018
15 Genes and How They Work 278 51 Osmotic Regulation and the Urinary System 1038
16 Control of Gene Expression 304 52 The Immune System 1055
17 Biotechnology 327 53 The Reproductive System 1084
18 Genomics 352 54 Animal Development 1105
19 Cellular Mechanisms & Development 372

Part
IV Evolution 396 Part VIII Ecology and Behavior 1132

20 Genes Within Populations 396 55 Behavioral Biology 1132

21 The Evidence for Evolution 417 56 Ecology of Individuals and Populations 1162
22 The Origin of Species 436 57 Community Ecology 1185
23 Systematics and the Phylogenetic Revolution 456 58 Dynamics of Ecosystems 1207
24 Genome Evolution 474 59 The Biosphere 1230
25 Evolution of Development 492 60 Conservation Biology 1256

Appendix A A-1
Part
V The Diversity of Life 507
Glossary G-1
26 The Tree of Life 507
Credits C-1
27 Viruses 528
28 Prokaryotes 545 Index I-1
iii
Contents

5 Membranes 88
Part I The Molecular Basis of Life
1 The Science of Biology 1
5.1
5.2
5.3
The Structure of Membranes 88
Phospholipids: The Membrane’s Foundation 92
Proteins: Multifunctional Components 93
5.4 Passive Transport Across Membranes 96
1.1 The Science of Life 1
5.5 Active Transport Across Membranes 99
1.2 The Nature of Science 4
5.6 Bulk Transport by Endocytosis and Exocytosis 102
1.3 An Example of Scientific Inquiry: Darwin
and Evolution 8
1.4 Unifying Themes in Biology 12
6 Energy and Metabolism 107
6.1 The Flow of Energy in Living Systems 108
2 The Nature of Molecules 6.2 The Laws of Thermodynamics and Free Energy 109
6.3 ATP: The Energy Currency of Cells 112
and the Properties of Water 17 6.4 Enzymes: Biological Catalysts 113
2.1 The Nature of Atoms 18 6.5 Metabolism: The Chemical Description
2.2 Elements Found in Living Systems 22 of Cell Function 117
2.3 The Nature of Chemical Bonds 23
2.4 Water: A Vital Compound 25 7 How Cells Harvest Energy 122
2.5 Properties of Water 28 7.1 Overview of Respiration 123
2.6 Acids and Bases 29 7.2 Glycolysis: Splitting Glucose 127
Apago PDF Enhancer 7.3 The Oxidation of Pyruvate to Produce
3 The Chemical Building Blocks Acetyl-CoA 130
7.4 The Krebs Cycle 131
of Life 33
7.5 The Electron Transport Chain and Chemiosmosis 134
3.1 Carbon: The Framework
7.6 Energy Yield of Aerobic Respiration 137
of Biological Molecules 34
7.7 Regulation of Aerobic Respiration 138
3.2 Carbohydrates: Energy Storage
and Structural Molecules 38 7.8 Oxidation Without O2 139
3.3 Nucleic Acids: Information Molecules 41 7.9 Catabolism of Proteins and Fats 140
3.4 Proteins: Molecules with Diverse Structures 7.10 Evolution of Metabolism 142
and Functions 44
3.5 Lipids: Hydrophobic Molecules 53 8 Photosynthesis 147
8.1 Overview of Photosynthesis 147
8.2 The Discovery of Photosynthetic Processes 149

II Biology of the Cell

8.3 Pigments 151
Part 8.4 Photosystem Organization 154
8.5 The Light-Dependent Reactions 156
4 Cell Structure 59 8.6 Carbon Fixation: The Calvin Cycle 160
4.1 Cell Theory 59 8.7 Photorespiration 163
4.2 Prokaryotic Cells 63
4.3 Eukaryotic Cells 65 9 Cell Communication 168
4.4 The Endomembrane System 69 9.1 Overview of Cell Communication 168
4.5 Mitochondria and Chloroplasts: 9.2 Receptor Types 171
Cellular Generators 73
9.3 Intracellular Receptors 173
4.6 The Cytoskeleton 75
9.4 Signal Transduction Through Receptor Kinases 174
4.7 Extracellular Structures and Cell Movement 79
9.5 Signal Transduction Through G Protein-Coupled
4.8 Cell-to-Cell Interactions 82 Receptors 179

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 10 How Cells Divide 186 15.5 Eukaryotic pre-mRNA Splicing 289
15.6 The Structure of tRNA and Ribosomes 291
10.1 Bacterial Cell Division 187
15.7 The Process of Translation 293
10.2 Eukaryotic Chromosomes 189
15.8 Summarizing Gene Expression 297
10.3 Overview of the Eukaryotic Cell Cycle 192
15.9 Mutation: Altered Genes 299
10.4 Interphase: Preparation for Mitosis 193
10.5 M Phase: Chromosome Segregation and the Division
of Cytoplasmic Contents 194 16 Control of Gene Expression 304
10.6 Control of the Cell Cycle 198 16.1 Control of Gene Expression 304
16.2 Regulatory Proteins 305
16.3 Prokaryotic Regulation 308
16.4 Eukaryotic Regulation 312
Part
III Genetic and 16.5
16.6
16.7
Eukaryotic Chromatin Structure 316
Eukaryotic Posttranscriptional Regulation 317
Protein Degradation 322
Molecular Biology
11 Sexual Reproduction and Meiosis 207 17 Biotechnology 327
11.1 Sexual Reproduction Requires Meiosis 207 17.1 DNA Manipulation 327
11.2 Features of Meiosis 209 17.2 Molecular Cloning 330
11.3 The Process of Meiosis 210 17.3 DNA Analysis 335
11.4 Summing Up: Meiosis Versus Mitosis 215 17.4 Genetic Engineering 341
17.5 Medical Applications 343
12 Patterns of Inheritance 221 17.6 Agricultural Applications 346

12.1 The Mystery of Heredity 221

18 Genomics 352
12.2 Monohybrid Crosses: The Principle
of Segregation 224 18.1 Mapping Genomes 352
12.3 Dihybrid Crosses: The Principle of Independent 18.2 Whole-Genome Sequencing 356
Assortment 228 18.3 Characterizing Genomes 358
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12.4 Probability: Predicting the Results of Crosses 230 18.4 Genomics and Proteomics 362
12.5 The Testcross: Revealing Unknown Genotypes 231 18.5 Applications of Genomics 367
12.6 Extensions to Mendel 232
19 Cellular Mechanisms
13 Chromosomes, Mapping, and the of Development 372
Meiosis–Inheritance Connection 239 19.1 The Process of Development 372
13.1 Sex Linkage and the Chromosomal Theory 19.2 Cell Division 373
of Inheritance 240
19.3 Cell Differentiation 375
13.2 Sex Chromosomes and Sex Determination 241
19.4 Nuclear Reprogramming 380
13.3 Exceptions to the Chromosomal Theory
19.5 Pattern Formation 383
of Inheritance 244
19.6 Morphogenesis 390
13.4 Genetic Mapping 244
13.5 Selected Human Genetic Disorders 249

14 DNA: The Genetic Material 256

14.1
14.2
The Nature of the Genetic Material 256
DNA Structure 259
Part IV Evolution
20 Genes Within Populations 396
14.3 Basic Characteristics of DNA Replication 263
20.1 Genetic Variation and Evolution 396
14.4 Prokaryotic Replication 266
20.2 Changes in Allele Frequency 398
14.5 Eukaryotic Replication 271
20.3 Five Agents of Evolutionary Change 401
14.6 DNA Repair 273
20.4 Fitness and Its Measurement 405
15 Genes and How They Work 278 20.5 Interactions Among Evolutionary Forces 406
20.6 Maintenance of Variation 407
15.1 The Nature of Genes 278
20.7 Selection Acting on Traits Affected
15.2 The Genetic Code 282 by Multiple Genes 409
15.3 Prokaryotic Transcription 284 20.8 Experimental Studies of Natural Selection 411
15.4 Eukaryotic Transcription 287 20.9 The Limits of Selection 413
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21 The Evidence for Evolution 417
21.1 The Beaks of Darwin’s Finches:
Evidence of Natural Selection 418
21.2 Peppered Moths and Industrial Melanism:
More Evidence of Selection 420
Part V The Diversity of Life
26 The Tree of Life 507
21.3 Artificial Selection: Human-Initiated Change 422
26.1 Origins of Life 508
21.4 Fossil Evidence of Evolution 424
26.2 Classification of Organisms 512
21.5 Anatomical Evidence for Evolution 428
26.3 Grouping Organisms 514
21.6 Convergent Evolution and the Biogeographical
Record 430 26.4 Making Sense of the Protists 520
21.7 Darwin’s Critics 432 26.5 Origin of Plants 520
26.6 Sorting out the Animals 522
22 The Origin of Species 436
27 Viruses 528
22.1 The Nature of Species and the Biological
Species Concept 437 27.1 The Nature of Viruses 529
22.2 Natural Selection and Reproductive Isolation 441 27.2 Bacteriophages: Bacterial Viruses 533
22.3 The Role of Genetic Drift and Natural Selection 27.3 Human Immunodeficiency Virus (HIV) 535
in Speciation 443 27.4 Other Viral Diseases 539
22.4 The Geography of Speciation 444 27.5 Prions and Viroids: Subviral Particles 541
22.5 Adaptive Radiation and Biological Diversity 446
22.6 The Pace of Evolution 451 28 Prokaryotes 545
22.7 Speciation and Extinction Through Time 452
28.1 The First Cells 546
28.2 Prokaryotic Diversity 547
23 Systematics and the Phylogenetic 28.3 Prokaryotic Cell Structure 551
Revolution 456 28.4 Prokaryotic Genetics 554
23.1 Systematics 456 28.5 Prokaryotic Metabolism 559
23.2 Cladistics 458 28.6 Human Bacterial Disease 560
23.3 Apago PDF Enhancer
Systematics and Classification 461 28.7 Beneficial Prokaryotes 563
23.4 Phylogenetics and Comparative Biology 464
23.5 Phylogenetics and Disease Evolution 470 29 Protists 567
29.1 Eukaryotic Origins and Endosymbiosis 568
24 Genome Evolution 474 29.2 Defining Protists 571
29.3 Diplomonads and Parabasalids: Flagellated Protists
24.1 Comparative Genomics 474 Lacking Mitochondria 572
24.2 Whole-Genome Duplications 477 29.4 Euglenozoa: A Diverse Group in Which Some Members
24.3 Evolution Within Genomes 481 Have Chloroplasts 573
24.4 Gene Function and Expression Patterns 484 29.5 Alveolata: Protists with Submembrane Vesicles 576
24.5 Nonprotein-Coding DNA 29.6 Stramenopila: Protists with Fine Hairs 580
and Regulatory Function 485 29.7 Rhodophyta: Red Algae 582
24.6 Genome Size and Gene Number 486 29.8 Choanoflagellida: Possible Animal Ancestors 583
24.7 Genome Analysis and Disease Prevention 29.9 Protists Without a Clade 583
and Treatment 487
24.8 Crop Improvement Through Genome Analysis 489
30 Green Plants 588
30.1 Defining Plants 588
25 Evolution of Development 492
30.2 Chlorophytes and Charophytes: Green Algae 591
25.1 Overview of Evolutionary 30.3 Bryophytes: Dominant Gametophyte
Developmental Biology 492 Generation 593
25.2 One or Two Gene Mutations, New Form 495 30.4 Tracheophyte Plants: Roots, Stems, and Leaves 596
25.3 Same Gene, New Function 496 30.5 Lycophytes: Dominant Sporophyte Generation
25.4 Different Genes, Convergent Function 498 and Vascular Tissue 598
25.5 Gene Duplication and Divergence 499 30.6 Pterophytes: Ferns and Their Relatives 598
25.6 Functional Analysis of Genes 30.7 The Evolution of Seed Plants 602
Across Species 500 30.8 Gymnosperms: Plants with “Naked Seeds” 603
25.7 Diversity of Eyes in the Natural World: 30.9 Angiosperms: The Flowering Plants 606
A Case Study 501

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31 Fungi 614 37 Vegetative Plant Development 753
31.1 Defining Fungi 614 37.1 Embryo Development 754
31.2 Microsporidia: Unicellular Parasites 618 37.2 Seeds 760
31.3 Chytridiomycota and Relatives: 37.3 Fruits 761
Fungi with Flagellated Zoospores 619 37.4 Germination 764
31.4 Zygomycota: Fungi that Produce Zygotes 620
31.5 Glomeromycota: Asexual Plant Symbionts 622 38 Transport in Plants 769
31.6 Basidiomycota: The Club (Basidium) Fungi 622
38.1 Transport Mechanisms 770
31.7 Ascomycota: The Sac (Ascus) Fungi 623
38.2 Water and Mineral Absorption 773
31.8 Ecology of Fungi 625
38.3 Xylem Transport 776
31.9 Fungal Parasites and Pathogens 629
38.4 The Rate of Transpiration 778
38.5 Water-Stress Responses 780
32 Overview of Animal Diversity 633 38.6 Phloem Transport 781
32.1 Some General Features of Animals 634
32.2 Evolution of the Animal Body Plan 636 39 Plant Nutrition and Soils 786
32.3 The Classification of Animals 640
39.1 Soils: The Substrates on Which Plants Depend 787
32.4 The Roots of the Animal Tree of Life 645
39.2 Plant Nutrients 790
39.3 Special Nutritional Strategies 792
33 Noncoelomate Invertebrates 649 39.4 Carbon–Nitrogen Balance and Global Change 795
33.1 Parazoa: Animals That Lack 39.5 Phytoremediation 797
Specialized Tissues 650
33.2 Eumetazoa: Animals with True Tissues 652
40 Plant Defense Responses 802
33.3 The Bilaterian Acoelomates 656
40.1 Physical Defenses 802
33.4 The Pseudocoelomates 661
40.2 Chemical Defenses 805
40.3 Animals that Protect Plants 809
34 Coelomate Invertebrates 666
40.4 Systemic Responses to Invaders 810
34.1 Phylum Mollusca: The Mollusks 666
34.2 Apago PDF 41
Phylum Nemertea: The Ribbon Worms 672 Enhancer
Sensory Systems in Plants 814
34.3 Phylum Annelida: The Annelids 673
41.1 Responses to Light 815
34.4 The Lophophorates: Bryozoa and Brachiopoda 676
41.2 Responses to Gravity 819
34.5 Phylum Arthropoda: The Arthropods 678
41.3 Responses to Mechanical Stimuli 821
34.6 Phylum Echinodermata: The Echinoderms 687
41.4 Responses to Water and Temperature 823
41.5 Hormones and Sensory Systems 825
35 Vertebrates 693
35.1 The Chordates 694 42 Plant Reproduction 839
35.2 The Nonvertebrate Chordates 695
42.1 Reproductive Development 840
35.3 The Vertebrate Chordates 696
42.2 Flower Production 842
35.4 Fishes 698
42.3 Structure and Evolution of Flowers 848
35.5 Amphibians 703
42.4 Pollination and Fertilization 851
35.6 Reptiles 706
42.5 Asexual Reproduction 857
35.7 Birds 712
42.6 Plant Life Spans 859
35.8 Mammals 716
35.9 Evolution of the Primates 721

Part VII Animal Form

Part VI Plant Form and Function
36 Plant Form 729
and Function
36.1 Organization of the Plant Body: An Overview 730
43 The Animal Body and Principles
36.2 Plant Tissues 733 of Regulation 863
36.3 Roots: Anchoring and Absorption Structures 739 43.1 Organization of the Vertebrate Body 864
36.4 Stems: Support for Above-Ground Organs 743 43.2 Epithelial Tissue 865
36.5 Leaves: Photosynthetic Organs 747 43.3 Connective Tissue 868
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 43.4 Muscle Tissue 870 49 The Respiratory System 1001
43.5 Nerve Tissue 872
49.1 Gas Exchange Across Respiratory Surfaces 1002
43.6 Overview of Vertebrate Organ Systems 872
49.2 Gills, Cutaneous Respiration,
43.7 Homeostasis 876 and Tracheal Systems 1004
43.8 Regulating Body Temperature 878 49.3 Lungs 1006
49.4 Structures and Mechanisms of Ventilation
44 The Nervous System 887 in Mammals 1009
44.1 Nervous System Organization 888 49.5 Transport of Gases in Body Fluids 1012
44.2 The Mechanism of Nerve Impulse Transmission 890
44.3 Synapses: Where Neurons Communicate 50 The Circulatory System 1018
with Other Cells 896 50.1 The Components of Blood 1018
44.4 The Central Nervous System: 50.2 Invertebrate Circulatory Systems 1022
Brain and Spinal Cord 901
50.3 Vertebrate Circulatory Systems 1023
44.5 The Peripheral Nervous System:
Sensory and Motor Neurons 909 50.4 The Four-Chambered Heart
and the Blood Vessels 1026
50.5 Characteristics of Blood Vessels 1030
45 Sensory Systems 915 50.6 Regulation of Blood Flow and Blood Pressure 1034
45.1 Overview of Sensory Receptors 916
45.2 Mechanoreceptors: Touch and Pressure 917 51 Osmotic Regulation
45.3 Hearing, Vibration, and Detection
of Body Position 920
and the Urinary System 1038
45.4 Chemoreceptors: Taste, Smell, and pH 925 51.1 Osmolarity and Osmotic Balance 1038
45.5 Vision 928 51.2 Osmoregulatory Organs 1040
45.6 The Diversity of Sensory Experiences 933 51.3 Evolution of the Vertebrate Kidney 1042
51.4 Nitrogenous Wastes: Ammonia, Urea,
and Uric Acid 1044
46 The Endocrine System 937 51.5 The Mammalian Kidney 1045
46.1 Regulation of Body Processes 51.6 Hormonal Control of Osmoregulatory Functions 1050
by Chemical Messengers 938
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46.2 Actions of Lipophilic Versus Hydrophilic
Hormones 943 52 The Immune System 1055
46.3 The Pituitary and Hypothalamus: 52.1 Innate Immunity 1055
The Body’s Control Centers 946 52.2 Adaptive Immunity 1061
46.4 The Major Peripheral Endocrine Glands 951 52.3 Cell-Mediated Immunity 1066
46.5 Other Hormones and Their Effects 955 52.4 Humoral Immunity and Antibody Production 1068
52.5 Autoimmunity and Hypersensitivity 1075
47 The Musculoskeletal System 961 52.6 Antibodies in Medical Treatment and Diagnosis 1077
47.1 Types of Skeletal Systems 962 52.7 Pathogens That Evade the Immune System 1079
47.2 A Closer Look at Bone 963
47.3 Joints and Skeletal Movement 967 53 The Reproductive System 1084
47.4 Muscle Contraction 969 53.1 Animal Reproductive Strategies 1084
47.5 Modes of Animal Locomotion 975 53.2 Vertebrate Fertilization and Development 1087
53.3 Structure and Function of the Human Male
48 The Digestive System 981 Reproductive System 1091
53.4 Structure and Function of the Human Female
48.1 Types of Digestive Systems 982 Reproductive System 1094
48.2 The Mouth and Teeth: Food Capture 53.5 Contraception and Infertility Treatments 1098
and Bulk Processing 984
48.3 The Esophagus and the Stomach:
The Early Stages of Digestion 985 54 Animal Development 1105
48.4 The Intestines: Breakdown, Absorption, 54.1 Fertilization 1106
and Elimination 987 54.2 Cleavage and the Blastula Stage 1110
48.5 Variations in Vertebrate Digestive Systems 990 54.3 Gastrulation 1112
48.6 Neural and Hormonal Regulation 54.4 Organogenesis 1116
of the Digestive Tract 993
54.5 Vertebrate Axis Formation 1122
48.7 Accessory Organ Function 994
54.6 Human Development 1125
48.8 Food Energy, Energy Expenditure,
and Essential Nutrients 995
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 57.3 Predator–Prey Relationships 1192
57.4 The Many Types of Species Interactions 1196

Part VIII Ecology and Behavior

55 Behavioral Biology 1132
57.5 Ecological Succession, Disturbance, and
Species Richness 1202

58 Dynamics of Ecosystems 1207

55.1 The Natural History of Behavior 1133 58.1 Biogeochemical Cycles 1208
55.2 Nerve Cells, Neurotransmitters, Hormones, 58.2 The Flow of Energy in Ecosystems 1214
and Behavior 1134
58.3 Trophic-Level Interactions 1219
55.3 Behavioral Genetics 1135
58.4 Biodiversity and Ecosystem Stability 1223
55.4 Learning 1137
58.5 Island Biogeography 1226
55.5 The Development of Behavior 1139
55.6 Animal Cognition 1141
59 The Biosphere 1230
55.7 Orientation and Migratory Behavior 1142
55.8 Animal Communication 1144 59.1 Ecosystem Effects of Sun, Wind, and Water 1230
55.9 Behavioral Ecology 1147 59.2 Earth’s Biomes 1235
55.10 Reproductive Strategies and Sexual 59.3 Freshwater Habitats 1238
Selection 1150 59.4 Marine Habitats 1241
55.11 Altruism 1154 59.5 Human Impacts on the Biosphere:
55.12 The Evolution of Group Living Pollution and Resource Depletion 1245
and Animal Societies 1157 59.6 Human Impacts on the Biosphere:
Climate Change 1250
56 Ecology of Individuals
60 Conservation Biology 1256
and Populations 1162
60.1 Overview of the Biodiversity Crisis 1257
56.1 The Environmental Challenges 1162
60.2 The Value of Biodiversity 1261
56.2 Populations: Groups of a Single Species
in One Place 1165 60.3 Factors Responsible for Extinction 1264
56.3 Population Demography and Dynamics 1168 60.4 Approaches for Preserving Endangered Species and
Ecosystems 1275
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56.4 Life History and the Cost of Reproduction 1171
56.5 Environmental Limits to Population Appendix A A-1
Growth 1173
56.6 Factors That Regulate Populations 1175 Glossary G-1
56.7 Human Population Growth 1178 Credits C-1
Index I-1
57 Community Ecology 1185
57.1 Biological Communities: Species Living Together 1186
57.2 The Ecological Niche Concept 1188

xxvi table of contents