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Twelfth Edition

Biology
Kenneth A. Mason
University of Iowa

Jonathan B. Losos
William H. Danforth Distinguished University
Professor and Director, Living Earth Collaborative,
Washington University

Tod Duncan
University of Colorado Denver

Contributor:
Charles J. Welsh
Duquesne University
Based on the work of
Peter H. Raven
President Emeritus, Missouri Botanical Garden;
George Engelmann Professor of Botany Emeritus,
Washington University
George B. Johnson
Professor Emeritus of Biology, Washington
University
BIOLOGY, TWELFTH EDITION

Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2020 by McGraw-Hill Education. All rights
reserved. Printed in the United States of America. Previous editions © 2017, 2014, and 2011. No part of this publication may be reproduced
or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written consent of McGraw-Hill
Education, including, but not limited to, in any network or other electronic storage or transmission, or broadcast for distance learning.

Some ancillaries, including electronic and print components, may not be available to customers outside the United States.

This book is printed on acid-free paper.

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ISBN 978-1-260-16961-4 (bound edition)

MHID 1-260-16961-8 (bound edition)
ISBN 978-1-260-49470-9 (loose-leaf edition)
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All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.

Library of Congress Cataloging-in-Publication Data

Mason, Kenneth A., author. | Losos, Jonathan B., author. | Duncan, Tod, author.
Biology / Kenneth A. Mason, University of Iowa, Jonathan B. Losos,
 Washington University, Tod Duncan, University of Colorado, Denver;
contributors, Charles J. Welsh, Duquesne University.
Twelfth edition. | New York, NY : McGraw-Hill Education, [2020]
 | “Based on the work of Peter H. Raven, President Emeritus, Missouri
 Botanical Garden; George Engelmann, Professor of Botany Emeritus,
 Washington University, George B. Johnson, Professor Emeritus of Biology,
 Washington University.” | Includes index.
LCCN 2018036968| ISBN 9781260169614 (alk. paper) |
 ISBN 9781260565959
 LCSH: Biology—Textbooks.
 LCC QH308.2 .R38 2020 | DDC 570—dc23
 LC record available at https://lccn.loc.gov/2018036968

The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate an
endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education does not guarantee the accuracy of the information
presented at these sites.

mheducation.com/highered
Brief Contents

Committed to Excellence xi 27 Prokaryotes 557

Preparing Students for the Future xv 28 Protists 584
29 Seedless Plants 608
30 Seed Plants 623
31 Fungi 641
Part I The Molecular Basis of Life 1 32
33
Animal Diversity and the Evolution of Body Plans
Protostomes 687
664

1 The Science of Biology 1 34 Deuterostomes 720

2 The Nature of Molecules and the Properties of Water 18
3 The Chemical Building Blocks of Life 35
Part VI Plant Form and Function 762

Part II Biology of the Cell 63 35

36
Plant Form 762
Transport in Plants 788
4 Cell Structure 63 37 Plant Nutrition and Soils 807
5 Membranes 92 38 Plant Defense Responses 825
6 Energy and Metabolism 112 39 Sensory Systems in Plants 838
7 How Cells Harvest Energy 128 40 Plant Reproduction 866
8 Photosynthesis 154
9
10
Cell Communication 176
How Cells Divide 194
Part VII Animal Form and Function 900

41 The Animal Body and Principles of Regulation 900

Part III Genetic and Molecular Biology 217 42
43
The Nervous System 924
Sensory Systems 955
11 Sexual Reproduction and Meiosis 217 44 The Endocrine System 982
12 Patterns of Inheritance 231 45 The Musculoskeletal System 1006
13 Chromosomes, Mapping, and the Meiosis–Inheritance 46 The Digestive System 1026
Connection 250 47 The Respiratory System 1047
14 DNA: The Genetic Material 268 48 The Circulatory System 1066
15 Genes and How They Work 290 49 Osmotic Regulation and the Urinary System 1088
16 Control of Gene Expression 317 50 The Immune System 1106
17 Biotechnology 340 51 The Reproductive System 1135
18 Genomics 366 52 Animal Development 1157
19 Cellular Mechanisms of Development 389

Part IV Evolution 416

Part VIII Ecology and Behavior 1188

53 Behavioral Biology 1188

20 Genes Within Populations 416 54 Ecology of Individuals and Populations 1218
21 The Evidence for Evolution 443 55 Community Ecology 1242
22 The Origin of Species 463 56 Dynamics of Ecosystems 1265
23 Systematics, Phylogenies, and Comparative Biology 484 57 The Biosphere and Human Impacts 1289
24 Genome Evolution 504 58 Conservation Biology 1318

V
Appendix A
Part Diversity of Life on Earth 523
Glossary G-1
25 The Origin and Diversity of Life 523 Index I-1
26 Viruses 537

iii
About the Authors
Kenneth Mason maintains an association with the University of Iowa, Department of Biology after having served
as a faculty member for eight years. His academic positions, as a teacher and researcher, include the faculty of
the University of Kansas, where he designed and established the genetics lab, and taught and published on the
genetics of pigmentation in amphibians. At Purdue University, he successfully developed and grew large intro-
ductory biology courses and collaborated with other faculty in an innovative biology, chemistry, and physics
course supported by the National Science Foundation. At the University of Iowa, where his wife served as
©Kenneth Mason president of the university, he taught introductory biology and human genetics. His honor society memberships
include Phi Sigma, Alpha Lambda Delta, and, by vote of Purdue pharmacy students, Phi Eta Sigma Freshman
Honors Society.

Jonathan Losos is the William H. Danforth Distinguished University Professor in the Department of Biology
at Washington University and Director of the Living Earth Collaborative, a partnership between the university,
the Saint Louis Zoo and the Missouri Botanical Garden. Losos’s research has focused on studying patterns
of adaptive radiation and evolutionary diversification in lizards. He is a member of the National Academy
of Sciences, a fellow of the American Academy of Arts and Science, and the recipient of several awards,
including the Theodosius Dobzhanksy and David Starr Jordan Prizes, the Edward Osborne Wilson Naturalist
©Jonathan Losos Award, and the Daniel Giraud Elliot Medal, as well as receiving fellowships from the John Guggenheim and
David and Lucile Packard Foundations. Losos has published more than 200 scientific articles and has written
two books, Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles (University of California
Press, 2009) and Improbable Destinies: Fate, Chance, and the Future of Evolution (Penguin-Random
House, 2017).

Tod Duncan is a Clinical Assistant Professor at the University of Colorado Denver. He currently teaches first semester
general biology and coordinates first and second semester general biology laboratories. Previously, he taught general
microbiology, virology, the biology of cancer, medical microbiology, and cell biology. A bachelor’s degree in cell
biology with an emphasis on plant molecular and cellular biology from the University of East Anglia in England led to
doctoral studies in cell cycle control, and postdoctoral research on the molecular and biochemical mechanisms of DNA
alkylation damage in vitro and in Drosophila melanogaster. Currently, he is interested in factors affecting retention
©Lesley Howard and success of incoming first-year students in diverse demographics. He lives in Boulder, Colorado, with his two Great
Danes, Eddie and Henry.

iv
 Contents

Committed to Excellence xi 4.4 The Endomembrane System 73

Preparing Students for the Future
©Soames Summerhays/Natural Visions
xv 4.5 Mitochondria and Chloroplasts: Cellular
Generators 77
4.6 The Cytoskeleton 79
4.7 Extracellular Structures and Cell Movement 83
4.8 Cell-to-Cell Interactions 86

I The Molecular Basis

Part
of Life
1 The Science of Biology 1
1.1 The Science of Life 1
1.2 The Nature of Science 4
1.3 An Example of Scientific Inquiry: Darwin and
Evolution 8
1.4 Core Concepts in Biology 12
2 The Nature of Molecules and the
Properties of Water 18
2.1 The Nature of Atoms 19
2.2 Elements Found in Living Systems 23
2.3 The Nature of Chemical Bonds 24
2.4 Water: A Vital Compound 26
2.5 Properties of Water 29
2.6 Acids and Bases 30
3 The Chemical Building Blocks of Life 35
3.1 Carbon: The Framework of Biological Molecules 36
3.2 Carbohydrates: Energy Storage and Structural
Molecules 40
3.3 Nucleic Acids: Information Molecules 43
3.4 Proteins: Molecules with Diverse Structures and
Functions 46
3.5 Lipids: Hydrophobic Molecules 56
5 Membranes 92
5.1 The Structure of Membranes 92
5.2 Phospholipids: The Membrane’s Foundation 96
5.3 Proteins: Multifunctional Components 98
5.4 Passive Transport Across Membranes 100
5.5 Active Transport Across Membranes 103
5.6 Bulk Transport by Endocytosis and Exocytosis 106
6 Energy and Metabolism 112
6.1 The Flow of Energy in Living Systems 113
6.2 The Laws of Thermodynamics and
Free Energy 114
6.3 ATP: The Energy Currency of Cells 117
6.4 Enzymes: Biological Catalysts 118
6.5 Metabolism: The Chemical Description of Cell
Function 122
7 How Cells Harvest Energy 128
7.1 Overview of Respiration 129
7.2 Glycolysis: Splitting Glucose 133
7.3 The Oxidation of Pyruvate Produces
Acetyl-CoA 136
7.4 The Citric Acid Cycle 137
7.5 The Electron Transport Chain and
Chemiosmosis 140
7.6 Energy Yield of Aerobic Respiration 143
7.7 Regulation of Aerobic Respiration 144
7.8 Oxidation Without O2 145
7.9 Catabolism of Proteins and Fats 147
7.10 Evolution of Metabolism 149

©Dr. Gopal Murti/Science Source 8 Photosynthesis 154

8.1 Overview of Photosynthesis 154

Part
II Biology of the Cell 8.2 The Discovery of Photosynthetic
Processes 156
8.3 Pigments 158
4 Cell Structure 63 8.4 Photosystem Organization 161
4.1 Cell Theory 63 8.5 The Light-Dependent Reactions 163
4.2 Prokaryotic Cells 67 8.6 Carbon Fixation: The Calvin Cycle 167
4.3 Eukaryotic Cells 69 8.7 Photorespiration 170

v
 9 Cell Communication 176 14 DNA: The Genetic Material 268
9.1 Overview of Cell Communication 176 14.1 The Nature of the Genetic Material 268
9.2 Receptor Types 179 14.2 DNA Structure 271
9.3 Intracellular Receptors 181 14.3 Basic Characteristics of DNA Replication 275
9.4 Signal Transduction Through Receptor 14.4 Prokaryotic Replication 278
Kinases 182 14.5 Eukaryotic Replication 283
9.5 Signal Transduction Through G Protein–Coupled 14.6 DNA Repair 285
Receptors 186

10 How Cells Divide 194 15 Genes and How They Work 290
15.1 The Nature of Genes 290
10.1 Bacterial Cell Division 195
15.2 The Genetic Code 293
10.2 Eukaryotic Chromosomes 197
15.3 Prokaryotic Transcription 296
10.3 Overview of the Eukaryotic Cell Cycle 200
15.4 Eukaryotic Transcription 299
10.4 Interphase: Preparation for Mitosis 201
15.5 Eukaryotic pre-mRNA Splicing 301
10.5 M Phase: Chromosome Segregation and the Division
of Cytoplasmic Contents 203 15.6 The Structure of tRNA and Ribosomes 303
10.6 Control of the Cell Cycle 206 15.7 The Process of Translation 305
10.7 Genetics of Cancer 211 15.8 Summarizing Gene Expression 309
15.9 Mutation: Altered Genes 311
©Steven P. Lynch

16 Control of Gene Expression 317

Part
III Genetic and Molecular 16.1
16.2
16.3
Control of Gene Expression 317
Regulatory Proteins 318
Prokaryotic Regulation 321
Biology 16.4 Eukaryotic Regulation 325
16.5 Chromatin Structure Affects Gene Expression 328
11 Sexual Reproduction and Meiosis 217 16.6 Eukaryotic Posttranscriptional Regulation 330
11.1 Sexual Reproduction Requires Meiosis 217 16.7 Protein Degradation 334
11.2 Features of Meiosis 219
11.3 The Process of Meiosis 220 17 Biotechnology 340
11.4 Summing Up: Meiosis Versus Mitosis 225 17.1 Recombinant DNA 340
17.2 Amplifying DNA Using the Polymerase Chain
12 Patterns of Inheritance 231 Reaction 345
17.3 Creating, Correcting, and Analyzing Genetic
12.1 The Mystery of Heredity 231
Variation 348
12.2 Monohybrid Crosses: The Principle of
17.4 Constructing and Using Transgenic Organisms 350
Segregation 234
17.5 Environmental Applications 354
12.3 Dihybrid Crosses: The Principle of Independent
Assortment 238 17.6 Medical Applications 356
12.4 Probability: Predicting the Results of Crosses 240 17.7 Agricultural Applications 360
12.5 The Testcross: Revealing Unknown Genotypes 241
12.6 Extensions to Mendel 242 18 Genomics 366
18.1 Mapping Genomes 366
13 Chromosomes, Mapping, and 18.2 Sequencing Genomes 370
the Meiosis–Inheritance 18.3 Genome Projects 373
18.4 Genome Annotation and Databases 374
Connection 250 18.5 Comparative and Functional Genomics 378
13.1 Sex Linkage and the Chromosomal Theory of 18.6 Applications of Genomics 383
Inheritance 251
13.2 Sex Chromosomes and Sex Determination 252
19 Cellular Mechanisms of
13.3 Exceptions to the Chromosomal Theory of
Inheritance 255 Development 389
13.4 Genetic Mapping 255 19.1 The Process of Development 389
13.5 Human Genetic Disorders 260 19.2 Cell Division 390

vi Contents
 19.3 Cell Differentiation 392 23.3 Systematics and Classification 489
19.4 Nuclear Reprogramming 397 23.4 Phylogenetics and Comparative Biology 493
19.5 Pattern Formation 400 23.5 Phylogenetics and Disease Evolution 499
19.6 Evolution of Pattern Formation 406
19.7 Morphogenesis 409 24 Genome Evolution 504
24.1 Comparative Genomics 504
©tamoncity/Shutterstock 24.2 Genome Size 508
24.3 Evolution Within Genomes 511

IV Evolution 24.4 Gene Function and Expression Patterns 515

Part 24.5 Applying Comparative Genomics 516

20 Genes Within Populations 416 ©Jeff Hunter/Getty Images

20.1 Genetic Variation and Evolution 416

20.2
20.3
20.4
Changes in Allele Frequency 418
Five Agents of Evolutionary Change 420
Quantifying Natural Selection 425
Part
V Diversity of Life
20.5 Reproductive Strategies 426 on Earth
20.6 Natural Selection’s Role in Maintaining
Variation 430 25 The Origin and Diversity
20.7 Selection Acting on Traits Affected by Multiple
Genes 432 of Life 523
20.8 Experimental Studies of Natural Selection 434 25.1 Deep Time 525
20.9  Interactions Among Evolutionary Forces 436 25.2 Origins of Life 525
20.10 The Limits of Selection 437 25.3 Evidence for Early Life 528
25.4 Earth’s Changing System 530
21 The Evidence for Evolution 443 25.5 Ever-Changing Life on Earth 531
21.1 The Beaks of Darwin’s Finches: Evidence of Natural
Selection 444 26 Viruses 537
21.2 Peppered Moths and Industrial Melanism: More Evidence 26.1 The Nature of Viruses 538
of Selection 446
26.2 Viral Diversity 542
21.3 Artificial Selection: Human-Initiated
Change 448 26.3 Bacteriophage: Bacterial Viruses 544
21.4 Fossil Evidence of Evolution 450 26.4 Viral Diseases of Humans 546
21.5 Anatomical Evidence for Evolution 454 26.5 Prions and Viroids: Infectious Subviral
Particles 552
21.6 Convergent Evolution and the Biogeographical
Record 456
21.7 Darwin’s Critics 458 27 Prokaryotes 557
27.1 Prokaryotic Diversity 558
22 The Origin of Species 463 27.2 Prokaryotic Cell Structure 562
22.1 The Nature of Species and the Biological Species 27.3 Prokaryotic Genetics 567
Concept 463 27.4 The Metabolic Diversity of Prokaryotes 571
22.2 Natural Selection and Reproductive Isolation 468 27.5 Microbial Ecology 573
22.3 The Role of Genetic Drift and Natural Selection in 27.6 Bacterial Diseases of Humans 575
Speciation 469
22.4 The Geography of Speciation 471 28 Protists 584
22.5 Adaptive Radiation and Biological Diversity 473
28.1 Eukaryotic Origins and Endosymbiosis 584
22.6 The Pace of Evolution 478
28.2 Overview of Protists 587
22.7 Speciation and Extinction Through Time 479
28.3 Characteristics of the Excavata 589
28.4 Characteristics of the Chromalveolata 592
23 Systematics, Phylogenies, and
28.5 Characteristics of the Rhizaria 598
Comparative Biology 484 28.6 Characteristics of the Archaeplastida 599
23.1 Systematics 484 28.7 Characteristics of the Amoebozoa 602
23.2 Cladistics 486 28.8 Characteristics of the Opisthokonta 603

Contents vii
29 Seedless Plants 608 34 Deuterostomes 720
29.1 Origin of Land Plants 608 34.1 Echinoderms 721
29.2 Bryophytes Have a Dominant Gametophyte 34.2 Chordates 723
Generation 611 34.3 Nonvertebrate Chordates 725
29.3 Tracheophytes Have a Dominant Sporophyte 34.4 Vertebrate Chordates 726
Generation 613
34.5 Fishes 728
29.4 Lycophytes Diverged from the Main Lineage
of Vascular Plants 616 34.6 Amphibians 733
29.5 Pterophytes Are the Ferns and Their 34.7 Reptiles 737
Relatives 617 34.8 Birds 742
34.9 Mammals 746
30 Seed Plants 623    34.10 Evolution of the Primates 751
30.1 The Evolution of Seed Plants 623
30.2 Gymnosperms: Plants with “Naked Seeds” 624 ©Susan Singer

30.3 Angiosperms: The Flowering Plants 628

30.4
30.5
Seeds 634
Fruits 635
Part
VI Plant Form and
31 Fungi 641 Function
31.1 Classification of Fungi 642
31.2 Fungal Forms, Nutrition, and Reproduction 643 35 Plant Form 762
31.3 Fungal Ecology 646 35.1 Organization of the Plant Body: An Overview 763
31.4 Fungal Parasites and Pathogens 650 35.2 Plant Tissues 766
31.5 Basidiomycota: The Club (Basidium) 35.3 Roots: Anchoring and Absorption Structures 772
Fungi 652 35.4 Stems: Support for Above-Ground Organs 776
31.6 Ascomycota: The Sac (Ascus) Fungi 654 35.5 Leaves: Photosynthetic Organs 781
31.7 Glomeromycota: Asexual Plant Symbionts 656
31.8 Zygomycota: Zygote-Producing Fungi 656 36 Transport in Plants 788
31.9 Chytridiomycota and Relatives: Fungi with
Zoospores 658 36.1 Transport Mechanisms 789
31.10 Microsporidia: Unicellular Parasites 659 36.2 Water and Mineral Absorption 792
36.3 Xylem Transport 795
32 Animal Diversity and the Evolution 36.4 Rate of Transpiration 797
36.5 Water-Stress Responses 799
of Body Plans 664
36.6 Phloem Transport 801
32.1 Some General Features of Animals 664
32.2 Evolution of the Animal Body Plan 666 37 Plant Nutrition and Soils 807
32.3 Animal Phylogeny 670
37.1 Soils: The Substrates on Which Plants Depend 807
32.4 Parazoa: Animals That Lack Specialized
Tissues 674 37.2 Plant Nutrients 811
32.5 Eumetazoa: Animals with True Tissues 677 37.3 Special Nutritional Strategies 813
32.6 The Bilateria 682 37.4 Carbon–Nitrogen Balance and Global Change 816
37.5 Phytoremediation 819
33 Protostomes 687
38 Plant Defense Responses 825
33.1 The Clades of Protostomes 688
38.1 Physical Defenses 825
33.2 Flatworms (Platyhelminthes) 689
38.2 Chemical Defenses 827
33.3 Rotifers (Rotifera) 692
38.3 Animals That Protect Plants 831
33.4 Mollusks (Mollusca) 693
38.4 Systemic Responses to Invaders 832
33.5 Ribbon Worms (Nemertea) 699
33.6 Annelids (Annelida) 700
33.7 Bryozoans (Bryozoa) and Brachiopods
39 Sensory Systems in Plants 838
(Brachiopoda) 703 39.1 Responses to Light 838
33.8 Roundworms (Nematoda) 705 39.2 Responses to Gravity 843
33.9 Arthropods (Arthropoda) 707 39.3 Responses to Mechanical Stimuli 845

viii Contents
 39.4 Responses to Water and Temperature 847 44 The Endocrine System 982
39.5 Hormones and Sensory Systems 849
44.1 Regulation of Body Processes by Chemical
Messengers 983
40 Plant Reproduction 866 44.2 Overview of Hormone Action 988
40.1 Reproductive Development 867 44.3 The Pituitary and Hypothalamus: The Body’s Control
40.2 Making Flowers 869 Centers 991
40.3 Structure and Evolution of Flowers 874 44.4 The Major Peripheral Endocrine Glands 996
40.4 Pollination and Fertilization 877 44.5 Other Hormones and Their Effects 1000
40.5 Embryo Development 882
40.6 Germination 888 45 The Musculoskeletal System 1006
40.7 Asexual Reproduction 891 45.1 Types of Skeletal Systems 1007
40.8 Plant Life Spans 893 45.2 A Closer Look at Bone 1009
45.3 Joints 1012
©Dr. Roger C. Wagner, Professor Emeritus of
Blologlcal Sciences, University of Delaware 45.4 Muscle Contraction 1013
45.5 Vertebrate Skeleton Evolution and Modes

VII Animal Form and

of Locomotion 1020
Part
46 The Digestive System 1026
Function 46.1 Types of Digestive Systems 1027
46.2 The Mouth and Teeth: Food Capture and Bulk
Processing 1029
41 The Animal Body and Principles 46.3 The Esophagus and the Stomach: The Early Stages
of Regulation 900 of Digestion 1030
46.4 The Intestines: Breakdown, Absorption, and
41.1 Organization of Animal Bodies 901 Elimination 1032
41.2 Epithelial Tissue 902 46.5 Accessory Organ Function 1035
41.3 Connective Tissue 905 46.6 Neural and Hormonal Regulation of the Digestive
  41.4 Muscle Tissue 908 Tract 1037
41.5 Nerve Tissue 909 46.7 Food Energy, Energy Expenditure, and Essential
41.6 Overview of Vertebrate Organ Systems 910 Nutrients 1038
41.7 Homeostasis 913 46.8 Variations in Vertebrate Digestive Systems 1042
41.8 Regulating Body Temperature 915
47 The Respiratory System 1047
42 The Nervous System 924 47.1 Gas Exchange Across Respiratory Surfaces 1048
42.1 Nervous System Organization 925 47.2 Gills, Cutaneous Respiration, and Tracheal
Systems 1049
42.2 The Mechanism of Nerve Impulse Transmission 928
47.3 Lungs 1052
42.3 Synapses: Where Neurons Communicate with Other
Cells 933 47.4 Structures, Mechanisms, and Control of Ventilation
in Mammals 1055
42.4 The Central Nervous System: Brain and
Spinal Cord 939 47.5 Transport of Gases in Body Fluids 1059
42.5 The Peripheral Nervous System: Spinal and Cranial
Nerves 946 48 The Circulatory System 1066
48.1 Invertebrate Circulatory Systems 1066
43 Sensory Systems 955 48.2 The Components of Vertebrate
Blood 1068
43.1 Overview of Sensory Receptors 956
48.3 Vertebrate Circulatory Systems 1071
43.2 Thermoreceptors, Nociceptors, and Electromagnetic
48.4 Cardiac Cycle, Electrical Conduction, ECG,
Receptors: Temperature, Pain, and Magnetic
and Cardiac Output 1074
Fields 958
48.5 Blood Pressure and Blood Vessels 1078
43.3 Mechanoreceptors I: Touch, Pressure, and Body
Position 959
43.4 Mechanoreceptors II: Hearing, Vibration, and
49 Osmotic Regulation and the Urinary
Balance 961 System 1088
43.5 Chemoreceptors: Taste, Smell, and pH 967 49.1 Osmolarity and Osmotic Balance 1088
43.6 Vision 969 49.2 Nitrogenous Wastes: Ammonia, Urea, and
43.7 Evolution and Development of Eyes 975 Uric Acid 1090

Contents ix
 49.3 Osmoregulatory Organs 1091 53.12 Altruism 1209
49.4 Evolution of the Vertebrate Kidney 1093 53.13  The Evolution of Group Living and Animal
49.5 The Mammalian Kidney 1095 Societies 1213
49.6 Hormonal Control of Osmoregulatory
Functions 1100 54 Ecology of Individuals and
Populations 1218
50 The Immune System 1106
54.1 The Environmental Challenges 1218
50.1 Innate Immunity 1106 54.2 Populations: Groups of a Single Species in One
50.2 Adaptive Immunity 1112 Place 1221
50.3 Cell-Mediated Immunity 1117 54.3 Population Demography and Dynamics 1224
50.4 Humoral Immunity and Antibody Production 1119 54.4 Life History and the Cost of Reproduction 1227
50.5 Autoimmunity and Hypersensitivity 1125 54.5 Environmental Limits to Population Growth 1230
50.6 Antibodies in Medical Treatment and 54.6 Factors That Regulate Populations 1232
Diagnosis 1127 54.7 Human Population Growth 1235
50.7 Pathogens That Evade the Immune System 1130
55 Community Ecology 1242
51 The Reproductive System 1135
55.1 Biological Communities: Species Living
51.1 Animal Reproductive Strategies 1135 Together 1243
51.2 Vertebrate Fertilization and Development 1138 55.2 The Ecological Niche Concept 1244
51.3 Structure and Function of the Human Male 55.3 Predator–Prey Relationships 1249
Reproductive System 1142
55.4 The Many Types of Species Interactions 1253
51.4 Structure and Function of the Human Female
55.5 Ecological Succession, Disturbance, and Species
Reproductive System 1146
Richness 1259
51.5 Contraception and Infertility Treatments 1150
56 Dynamics of Ecosystems 1265
52 Animal Development 1157
56.1 Biogeochemical Cycles 1266
52.1 Fertilization 1158
56.2 The Flow of Energy in Ecosystems 1272
52.2 Cleavage and the Blastula Stage 1162
56.3 Trophic-Level Interactions 1277
52.3 Gastrulation 1164
56.4 Biodiversity and Ecosystem Stability 1281
52.4 Organogenesis 1168
56.5 Island Biogeography 1284
52.5 Vertebrate Axis and Pattern Formation 1173
52.6 Human Development 1180 57 The Biosphere and Human
Impacts 1289
©K. Ammann/Bruce Coleman Inc./Photoshot 57.1 Ecosystem Effects of Sun, Wind, and Water 1289
57.2 Earth’s Biomes 1294

Part
VIII Ecology and 57.3
57.4
57.5
Freshwater Habitats 1297
Marine Habitats 1300
Human Impacts on the Biosphere: Pollution and
Behavior Resource Depletion 1304
57.6 Human Impacts on the Biosphere: Climate
Change 1310
53 Behavioral Biology 1188
53.1 The Natural History of Behavior 1189 58 Conservation Biology 1318
53.2  Nerve Cells, Neurotransmitters, Hormones, and 58.1 Overview of the Biodiversity Crisis 1318
Behavior 1190
58.2 The Value of Biodiversity 1323
53.3 Behavioral Genetics 1191
58.3 Factors Responsible for Extinction 1325
53.4 Learning 1193
58.4 An Evolutionary Perspective on the Biodiversity
53.5 The Development of Behavior 1194 Crisis 1336
53.6 Animal Cognition 1197 58.5 Approaches for Preserving Endangered Species and
53.7 Orientation and Migratory Behavior 1198 Ecosystems 1339
53.8 Animal Communication 1200
Appendix A
53.9 Behavior and Evolution 1203
53.10 Behavioral Ecology 1204 Glossary G-1
53.11 Reproductive Strategies 1207 Index I-1

x Content
Committed to Excellence
With the new 12th edition, Raven and Johnson’s Biology continues
the momentum built over the last four editions. We continue to pro-
vide an unmatched comprehensive text fully integrated with a con-
tinually evolving, state-of-the-art digital environment. We have
used this revision to recommit ourselves to our roots as the majors
biology text that best integrates evolution throughout. We have
added material emphasizing the relevance of evolution throughout
the ecology section, not only in all four ecology chapters, but also
in the chapters on behavior and conservation biology. In the animal
form and function section we have done extensive revision to mod-
ernize, and to emphasize evolution in the context of physiology.
Important contributions to this effort came from Dr. Charles Welsh
(Duquesne University), who provided his knowledge and experi-
ence to this important section. We have also moved the examples
and insights from the chapter devoted to the evolution of develop-
ment to place them into the appropriate context throughout the
book. This emphasizes the importance of evolution and develop-
ment by continually providing examples rather than gathering them
together in a single chapter.
We have also renewed our commitment to the ideas set forth
in the Vision and Change report from the AAAS, which provides a
framework for modern undergraduate biology education. This re-
port will have been with us for a decade coincident with our 12th
edition. One important idea articulated by Vision and Change was
an emphasis on core concepts. One of the key differences between
the way an expert organizes information in their brain compared to
a novice is that the expert has a conceptual framework in place to
incorporate new information. We have designed the new Connect-
ing the Concepts feature to address this disparity. We emphasize
core concepts in each chapter, then at the end of the chapter show
how these can be used to build a conceptual framework, and en-
courage the student to begin building their own. At the end of each
part of the book we expand this to show how core concepts are
interrelated and how a much larger conceptual framework is
constructed.
One unanticipated consequence of the Vision and Change
movement was how publishers chasing new approaches would
produce books so “feature-laden” as to be virtually unreadable by
the average student. We have not abandoned the idea that narra-
tive flow is important, even in a science textbook. While we
include a variety of features to improve student learning, they are
integrated into the text and not at the expense of the concise, ac-
cessible, and engaging writing style we are known for. We main-
tain the clear emphasis on evolution and scientific inquiry that
have made this a leading textbook of choice for majors biology
students.
Faculty want textbooks that emphasize student-centered ap-
proaches, and core concepts for the biological sciences. As a team,
we continually strive to improve the text by integrating the latest
cognitive and best practices research with methods that are known
to positively affect learning. We emphasize s­ cientific inquiry, in-
cluding an increased quantitative emphasis in the Scientific
Thinking figures. Our text continues to be a leader with an
organization that emphasizes important biological concepts, while
keeping the student engaged with learning outcomes that allow as-
sessment of progress in understanding these concepts. An inquiry-
based approach with robust, adaptive tools for discovery and
assessment in both text and digital resources provides the intellec-
tual challenge needed to promote student critical thinking and en-
sure academic success.
We continue to use our digital environment in the revision of
Biology. A major strength of both text and digital resources is assess-
ment across multiple levels of Bloom’s taxonomy that develops
critical-thinking and problem-solving skills in addition to com-
prehensive factual knowledge.
McGraw-Hill Education’s Connect® platform offers a
powerful suite of online tools that are linked to the text and in-
cludes new quantitative assessment tools. We now have avail-
able interactive exercises that use graphical data, controlled by
the student, to engage them in actively exploring quantitative
aspects of biology. Our adaptive learning system helps students
learn faster, study efficiently, and retain more knowledge of key
concepts.
The 12th edition continues to employ the aesthetically
stunning art program that the Raven and Johnson Biology text
is known for. Complex topics are represented clearly and suc-
cinctly, helping students to build the mental models needed to
understanding biology.
We continue to incorporate student usage data and input, de-
rived from thousands of our SmartBook® users. SmartBook “heat
maps” provided a quick visual snapshot of chapter usage data and
the relative difficulty students experienced in mastering the con-
tent. This “heat-mapping” technology is unique in the industry,
and allows direct editing of difficult areas, or problem areas for
students.
■■ If the data indicated that the subject was more difficult than
other parts of the chapter, as evidenced by a high proportion
of students responding incorrectly to the probes, we revised
or reorganized the content to be as clear and illustrative as
possible.
■■ In other cases, if one or more of the SmartBook probes
for a section was not as clear as it might be or did not
appropriately reflect the content, we edited the probe, rather
than the text.
We’re excited about the 12th edition of this quality textbook
providing a learning path for a new generation of students. All of
us have extensive experience teaching undergraduate biology, and
we’ve used this knowledge as a guide in producing a text that is up
to date, beautifully illustrated, and pedagogically sound for the stu-
dent. We are also excited about the continually evolving digital
environment that provides unique and engaging learning environ-
ment for modern students. We’ve worked hard to provide clear ex-
plicit learning outcomes, and more closely integrate the text with
xi
its media support materials to provide instructors with an excellent
complement to their teaching.
Ken Mason, Jonathan Losos, Tod Duncan
Cutting Edge Science
Changes to the 12th Edition
Part I: The Molecular Basis of Life
Chapter 1—New section added that elaborates on the core
concepts and prepares the student for the use of the Connecting
the Concepts feature.
Chapter 2—Edited for clarity, especially regarding atomic
structure and the periodic table.
Chapter 3—Edited for clarity especially regarding the structure
of nucleotides, the role of ATP in cells, and secondary structure
in proteins.
Part II: Biology of the Cell
Chapter 4—The section on the endomembrane system has been
completely rewritten. This includes new material on lipid
droplets. Material on adhesive junctions has been rewritten to
give a more evolutionary perspective.
Chapter 5—New material on proteins that can alter membrane
structure has been added. This provides information on how the
different cellular membranes can have different structures. Figure
on Na+/K+ pump was redone to address errors in mechanism.
Material on diffusion and facilitated diffusion was rewritten.
Chapter 6—The material on free energy and chemical reac-
tions was completely rewritten, including redoing the figures.
These changes significantly improve clarity and accuracy.
Material on the role of ATP in cells was rewritten for clarity.
Discussions of energy throughout the chapter were rewritten to
improve clarity and accuracy of chemical concepts.
Chapter 7—The nature and action of cofactors in redox
reactions and the role of ATP in cells were improved.
Chapter 8—The nature and structure of photosystems was
rewritten for clarity and accuracy.
Chapter 10—The section on chromosome structure was
completely rewritten to reflect new data and views of this
important topic. The material on cancer was expanded and
updated, producing a new section “Genetics of Cancer.” This
contains significant new information and pulls together
material on cancer from this chapter and others.
Part III: Genetic and Molecular Biology
The overall organization of this section remains the same. We
have retained the split of transmission genetics into two chapters
as it has proved successful for students.
xii Committed To Excellence
Chapter 11—Edited for clarity and readability for the student,
especially regarding the events of meiosis I.
Chapter 12—The material on extensions to Mendel was
rewritten for clarity and accuracy.
Chapter 13—The material on analyzing and mapping genetic
variation in humans was updated and rewritten. The section on
human genetic disorders was completely rewritten to reflect new
information, and to make more accessible for the student. A new
figure on imprinting in mouse was added to clarify this important
and difficult concept.
Chapter 14—The material on eukaryotic DNA replication was
rewritten and updated. Particular emphasis was placed on the
evolution of DNA replication. The section on DNA repair was
rewritten and updated and information on mismatch repair was
added.
Chapter 15—Content on process of transcription was rewritten
to reflect new data on elongation machinery. New data on
alternative splicing was included, along with information on the
integration of RNA modification during transcription. The
section on the nature of mutations was rewritten and includes
latest data on human mutation rates.
Chapter 16—Overview of control of eukaryotic transcription
was rewritten to reflect modern views. Continued updating of
the material on chromatin structure and the control of gene
expression. Material on control of gene expression at the level
of transcription was updated.
Chapter 18—New section added on the 1000 Genomes project
to illustrate how fast information on genetic diversity is accu-
mulating. The material on the wheat genome was updated,
which provides both new information and approaches to
complex genomes.
Chapter 19—Added a new section on the evolution of pattern
formation using new material and material from chapter 25.
This consolidates material on this subject, and provides a clear
vision for the student.
Part IV: Evolution
Chapter 20—The topic of sexual selection was moved into this
chapter from the Behavioral Biology chapter. Some material on
Lamarck was eliminated, natural selection was explicitly defined,
information on snp variation in humans and other animals was
added. New examples of pleiotropy were added, and new data on
how the speed of racehorses has not changed through time were
added along with a revised figure. A new section was added on the
role of sensory exploitation as a mechanism for traits to evolve
under sexual selection.
Chapter 21—A number of points were updated and an exam-
ple of vestigial traits involving the toenails of manatees was
added.
Chapter 23—The figure on the evolution of feathers in dino-
saurs was updated to incorporate new paleontological findings.
Discussion of HIV evolution and other points were also revised
in light of new science.
Chapter 24—Updated material on comparative genomics of
vertebrates. New data on Neanderthal and Denisovan genomes
have been added. Presentation of genes unique to humans has
been updated and edited for clarity.
Note: Evolution of Development (chapter 25 in the 11th edition)
was eliminated and material moved to other chapters, placing the
topic of evolution of development into the appropriate context.
This reflects the view that evolution and development are now so
clearly intertwined with all of biology that setting off the material
in a separate chapter no longer made sense.
Part V: Diversity of Life on Earth
Chapter 26—This chapter has been largely rewritten and now
includes material on viral diversity, classification, metagenomics,
and taxonomy. The latter part of the chapter now focuses on viruses
of medical importance to promote student engagement and interest.
Chapter 27—This chapter has been largely rewritten. In addition
to the traditional discussion of prokaryotic structure and function,
and taxonomy, there is new emphasis placed on microbial
ecology and medical microbiology with relevant examples.
Chapter 31—The chapter has been rewritten for clarity. The
chapter has also been reordered to bring material most relevant to
society to the front of the chapter. The reorganization includes
expanding and moving the fungal ecology up earlier in the chapter,
as well as expanding and moving the fungal parasites and patho-
gens up earlier in the chapter. The chapter now ends with the
coverage of fungal classification.
Chapter 32—Aspects of taxonomy and natural history were
updated in line with new findings.
Chapter 33—The presentation of taxonomic relationships was
revised as a result of new findings based primarily on molecular
phylogenetic studies, specifically with regards to Platyhelmin-
thes, lophotrochozoans (formerly Spiralia) and a few others.
New natural history information was included.
Chapter 34—The discussion of the evolutionary history of
vertebrates was substantially revised, especially the sections on
lobe-finned fishes/early tetrapods/early amniotes (emphasizing
now those terms, rather than referring to all of the early diverging
lineages as amphibians or reptiles). Also, the terminology about
human evolution was revised to acknowledge the new meaning of
“hominin” and “hominid.” A new paragraph on Homo naledi was
added to discuss recent discoveries.
Part VI: Plant Form and Function
There have been no major changes in the plant form and function
chapters. There has been overall editing for readability and
responding to recommendations by reviewers and users of the
11th edition.
Part VII: Animal Form and Function
Charles Welsh of Duquesne University, brought his expertise
in animal anatomy and physiology as a Contributor to the
Animal Form and Function Part in the 12th edition, placing
greater emphasis on evolutionary aspects of animal biology.
Chapter 41—The discussion of the evolution of tissues in
invertebrates and vertebrates was expanded, including the
addition of a phylogeny and an image of cnidarian tissues.
Chapter 42—The graph of an action potential was revised
and improved. Discussions and images of glial cells and
cranial nerves were added.
Chapter 43—The chapter was revised and reorganized
with regards to the general senses. The evolution of eyes
material found in chapter 25 in the 11th edition was moved
to this chapter with a revised phylogeny added. The
illustration depicting the evolution of the inner ear has been
revised to make it more clear, concise, and informative.
Chapter 44—Section 44.2 was formerly organized as action
of lipophilic vs. hydrophilic hormones. This has now been
reorganized to be a complete overview of how hormones
work. This organization should improve clarity for students.
Chapter 45—The chapter was extensively revised. This
included the addition of images for the human skeleton,
ossification, osteoporosis, invertebrate muscle, comparative
anatomy of flying vertebrates, and a new phylogeny that
reveals the evolution of various vertebrate skeletal characters.
Chapter 46—The structure of the latter half this chapter
was completely reorganized for better conceptual flow.
Chapter 47—The images for the bicarbonate buffering
system and the mechanics of breathing have been revised.
The discussion of lung volumes and capacities was expand-
ed with the addition of an accompanying figure.
Chapter 48—The chapter was reorganized and extensively
revised. Invertebrate circulatory systems is now the first
section in the chapter. The sections on Cardiac Cycle, ECG,
Electrical Conduction, and Cardiac Output have been reorga-
nized and revised. The discussions of blood vessels and blood
pressure are now in the same section. The phylogeny of the
evolution of vertebrate hearts has been revised.
Chapter 50—Material on innate immunity was updated
and rewritten for clarity. The coverage on effects of AIDS
was also updated to reflect new information.
Chapter 51—A discussion of some select invertebrate repro-
ductive strategies has been added, with accompanying images.
Committed To Excellence xiii
Chapter 52—A section detailing the classic experiments
regarding pattern formation in chick limb buds has been added.
This includes a discussion of AER, ZPA, FGF, Hox genes, and
Shh. The material on gene regulation from chapter 25 in the
11th edition has also been added.
Part VIII: Ecology and Behavior
Chapter 53—Stronger emphasis on phylogenetic and evolution-
ary perspectives was added throughout the chapter, including a
new section on evolution and behavior.
Chapter 54—Human population trends and other timely data were
updated to stay current. An evolutionary perspective on population
adaptation was added to the beginning of the chapter.
Chapter 55—An evolutionary perspective was added in several
places.
Chapter 56—New material on the impact of anthropogenic
changes on nutrient cycling was added. An evolutionary perspec-
tive to discussion of the species-area relationship was incorporated.
Chapter 57—Evolution was discussed more thoroughly in the
section on microclimate adaptation during adaptive radiation.
All of the data on biosphere impacts of humans were updated to
stay current.
Chapter 58—The chapter was substantially revised, including
much new discussion of the relevance of evolution to conserva-
tion biology, including the role of natural selection, the impor-
tance of phylogenetic perspectives, and how speciation can lead
to biodiversity hotspots.
A Note From the Authors
A revision of this scope relies on the talents and efforts of many
people working behind the scenes and we have benefited greatly
from their assistance.
Dr. Charles Welsh made significant contributions to the Animal
Form and Function section. He updated them to provide a more
modern perspective, and added new examples.
Beth Bulger was the copyeditor for this edition. She has la-
bored many hours and always improves the clarity and consis-
tency of the text. She has made significant contributions to the
quality of the final product.
We were fortunate to work again with MPS to update the art
program and improve the layout of the pages. Our close collabora-
tion resulted in a text that is pedagogically effective as well as
more beautiful than any other biology text on the market.
We have the continued support of an excellent team at
McGraw-Hill Education. Andrew Urban, preceded by Justin
Wyatt, the portfolio managers for Biology have been steady
leaders during a time of change. Senior Product Developer Liz
Sievers, provided support in so many ways it would be impossi-
ble to name them all. Kelly Hart, content project manager, and
David Hash, designer, ensured our text was on time and elegantly
designed. Kelly Brown, senior marketing manager, is always a
sounding board for more than just marketing, and many more
people behind the scenes have all contributed to the success of
our text. This includes the digital team, whom we owe a great
deal for their efforts to continue improving our Connect
assessment tools.
Throughout this edition we have had the support of spouses
and families, who have seen less of us than they might have
liked because of the pressures of getting this revision complet-
ed. They have adapted to the many hours this book draws us
away from them, and, even more than us, looked forward to its
completion.
In the end, the people we owe the most are the generations of
students who have used the many editions of this text. They have
taught us at least as much as we have taught them, and their ques-
tions and suggestions continue to improve the text and supple-
mentary materials.
Finally, we need to thank instructors from across the country
who are continually sharing their knowledge and experience with
us through market feedback and symposia. The feedback we re-
ceived shaped this edition. All of these people took time to share
their ideas and opinions to help us build a better edition of Biology
for the next generation of introductory biology students, and they
have our heartfelt thanks.

Reviewers for Biology, 12th edition

Carron Bryant East Mississippi Community
College
Mickael J. Cariveau University of Mount
Olive
Daniel Czerny Reading Area Community
College
Frank J. Dirrigl, Jr. University of Texas Rio
Grande Valley
Kathy McCann Evans Reading Area
Community College
Eric Ford East Mississippi Community
College-Golden Triangle
Mark Jonas Purchase College, SUNY
Kimberly Kushner Pueblo Community
College
Mark Levenstein University of Wisconsin,
Platteville
Cindy Malone California State University
Northridge
David McClellan University of Arkansas
Fort Smith
Shilpi Paul SUNY College at Old Westbury
Crima Pogge City College of San Francisco
Josephine Rodriguez The University of
Virginia’s College at Wise
Connie Rye East Mississippi Community
College
Devinder Sandhu USDA—Agricultural
Research Service
Ken Saville Albion College
Steven Shell The University of Virginia’s
College at Wise
Walter Smith The University of Virginia’s
College at Wise
Qiang Sun University of Wisconsin, Stevens
Point
Christopher Vitek University of Texas Rio
Grande Valley
D. Alexander Wait Missouri State University
Maureen Walter Florida International
University
Darla Wise Concord University

xiv Committed To Excellence

Preparing Students for the Future

Developing Critical Thinking with the Help of . . .

Scientific Thinking Figures
Key illustrations in every chapter highlight how the frontiers
of knowledge are pushed forward by a combination of hypoth-
esis and experimentation. These figures begin with a hypoth-
esis, then show how it makes explicit predictions, tests these
by experiment and finally demonstrates what conclusions can
be drawn, and where this leads. Scientific Thinking figures
provide a consistent framework to guide the student in the
logic of scientific inquiry. Each illustration concludes with
open-ended questions to promote scientific inquiry.
Data Analysis Questions
It’s not enough that students learn concepts and memorize
scientific facts, a biologist needs to analyze data and apply that
knowledge. Data Analysis questions inserted throughout the text
challenge students to analyze data and Interpret experimental
results, which shows a deeper level of understanding.
Inquiry Questions
Questions that challenge students to think about and engage in
what they are reading at a more sophisticated level.

SCIENTIFIC THINKING 32

Hypothesis: The plasma membrane is fluid, not rigid. 30

Temperature (°C)

Prediction: If the membrane is fluid, membrane proteins may

Body

diffuse laterally. 28
Test: Fuse mouse and human cells, then observe the distribution
of membrane proteins over time by labeling specific mouse and 26
human proteins. open habitat
24 shaded forest
Human
cell 24 26 28 30 32
Air Temperature (°C)

Mouse Figure 55.3 Behavioral adaptation. In open habitats, the

cell Intermixed Puerto Rican crested lizard, Anolis cristatellus, maintains a relatively
Fuse membrane proteins
cells constant temperature by seeking out and basking in patches of
sunlight; as a result, it can maintain a relatively high temperature even
when the air is cool. In contrast, in shaded forests, this behavior is not
possible, and the lizard’s body temperature conforms to that of its
surroundings.
Allow time for
mixing to occur (inset) ©Melissa Losos

Result: Over time, hybrid cells show increasingly intermixed proteins.

Inquiry question When given the opportunity, lizards
Conclusion: At least some membrane proteins can diffuse laterally in
the membrane.
? regulate their body temperature to maintain a temperature
optimal for physiological functioning. Would lizards in open
Further Experiments: Can you think of any other explanation for
habitats exhibit different escape behaviors from those of
these observations? What if newly synthesized proteins were inserted
lizards in shaded forest?
into the membrane during the experiment? How could you use this
basic experimental design to rule out this or other possible explanations?
Data analysis Can the slope of the line tell us something
about the behavior of the lizard?
Figure 5.5 Test of membrane fluidity.

xv
 contains a list of observations from the chapter that connects the
Connecting the Concepts secondary concept to the core concept.
There are two new but related features in Biology, 12th edition
At the chapter level:
that help students build a conceptual framework into which they
The Connecting the Concept shows the student a completed
can insert new knowledge. The Connecting the Concepts feature
concept (core concept, secondary concept, list of observations).
at the end of the chapters identifies core concepts that are
A second cog or gear is presented that lacks the list of observa-
related to material in the chapter. The conceptual framework
tions. The student is challenged to identify examples from the
begins with a core concept that is represented by a gear icon.
chapter that demonstrate how the secondary concept is related
Examples from the chapter that relate to the core concept are
to the core concept.
secondary concepts that are placed on the cogs. Each cog

CONNECTING THE CONCEPTS

This feature is intended to give you practice in organizing information using core concepts. We use a metaphor of gears and cogs to represent a conceptual
hierarchy with each core concept represented as a gear. Secondary concepts are the cogs, and tertiary concepts, which are particular examples from the chapter,
are presented as a list of bulleted points. Using the completed conceptual unit as a guide, build from material in the chapter a list of tertiary concepts that
support the open secondary concept.

• Positively charged soil can Plant

nutrients must be actively de s
transported into roots due con certa toxify
to their sequestration by env tami in
iro nat
ility
ilab t

anionic soil particles.

ava lan
nt ine p s

nm ed
e
d pro oil

ent
nut eterm perti

• Porous soils leach water s

S

rapidly and can contribute

to water stress.
rie

• The chemical properties of Life is subject Living systems

clay make it adsorb water
and minerals tightly. to chemical and transform
• The water potential of the physical laws energy & matter
soil affects the transport of
minerals into the root.
• Low soil pH can cause toxic
aluminum to leach from
rocks.
• Salt accumulation in soil
can affect soil water
potential and cause loss of
plant cell turgor.

At the Part level:
As valuable as that exercise is, the full understanding of a
conceptual framework and how that helps students see the
connections to core concepts is when the chapter-ending
Connecting the Concepts are pulled together. This happens at
the Part level, which themselves present a higher level to the
conceptual framework. When these are built, students see how
topics that appear unrelated fit into the conceptual framework
of the core concepts. Once students begin to see these connec-
tions, the topics and information in biology make
more sense.

xvi Preparing Students for the Future

 Connecting the Concepts Part VI Plant Form and Function
Vascular plants are comprised of roots and shoots, which in turn are made of three principal tissue types. Each of these tissues has distinct
cell types that express the genes needed to produce the proteins necessary for their specialized functions. Plants move fluids using differ-
ences in solute concentration and pressure. Plant form is often an evolutionary compromise between competing needs such as maximizing
the surface area of leaves for photosynthesis while minimizing water loss when exchanges gases. The reproductive structures of plants are
organized into flowers that have evolved to facilitate the dissemination of genetic information.
Each Connecting the
Concept unit (a Core
concept, secondary concept,
and bulleted list) is picked• The
up from the end-of-chaptertranspiration
features. This reinforces the
• The rate of osmosis limits water
overarching hierarchy of the
• The combined effects of solute potential
Core concepts, tying
together seemingly unrelated of plant cells.
• Water transport from roots to shoots is
material.
Connecting th
Vascular plants are comprised o
cell types that express the gen
ences in solute concentration a
the surface area of leaves for p
organized into flowers that hav
• Leaves are arranged on stems to maximize light capture.
• Stems may have secondary growth to provide support to the plant body.
• Axillary buds produced by the shoot apical meristem allow leaves or flowers to be produced
on the stem.
• Horizontal stems allow a plant to spread laterally above ground.
• Tubers can be packed with starch for storage purposes.
• Gametes are produced in the gametophytes of
flowers. • Flattened stems of some cacti capture light energy for photosynthesis.
• The calyx protects the budding flower.
• The petals collectively form the corolla and their • Gibberellins, a family of growth • Gametes are produced in
colors attract animal pollinators. hormones, can be produced by flowers.
• Wind-pollinated plants don’t have elaborate corollas bacteria infecting certain plants’ • The calyx protects the bud
because they don’t need to attract pollinators. roots and influence plant growth.
rs mo Stem • The petals collectively form
• The long stamens make pollen more accessible to we l
Flo e wel for
di s a
var car fied s nd
• Allelopathy is a form of signaling
where one plant releases
colors attract animal pollin
animal pollinators or wind. ar ted tion iety ry o tem • Wind-pollinated plants don
• The carpel houses the female reproductive structures
with the elongated style being more accessible to rep
p c
ada rodu
of f ut a s
unc
tion
compounds that inhibit seed
germination or the growth of because they don’t need t
s neighboring plants.
pollinators or pollen carried by the wind.
• Toxins produced by plants
communicate to potential predators
• The long stamens make p
animal pollinators or wind
cohesion and adhesion of water
molecules allows forces generated by Structure
that the plant is not safe to eat.
• Chemical signals can modulate the
• The carpel houses the fem
with the elongated style b
pollinators or pollen carrie
to move water great
distances in plants.
determines behaviors of insects that protect
plants from predation.
function
movement into roots, but is accelerated
• Chemicals released by plants as a
wound response can attract insects
• The cohesion and adhesio
molecules allows forces g
to defend the plant against
by facilitated diffusion through aquapo- transpiration to move wat
rins. herbivores.
distances in plants.
• The plant hormone jasmonic acid
and pressure potential determine the
direction of water movement into and out
transduces long distance wound
response signals in plant bodies.
• The rate of osmosis limits
movement into roots, but
by facilitated diffusion thro
rins.
Signaling
driven by a gradient of water potential sa
nd
mediates • The combined effects of s
with lowest values in the leaves.
• Chemical and physical properties of
sic try
Phy emis e r
ch ictat wate lant
plant health Students will see how the and pressure potential de
direction of water movem
d t of he p
membranes and cell walls restrict the
movement of solutes through the plant. v e m en nd t
o u same Core concepts are of plant cells.
• Water transport from roots
mo nd ar
• Positively charged soil
into
a
found throughout the book, driven by a gradient of wa
with lowest values in the l
determine plant
nutrients must be Life is subject Living systems establishing the conceptual • Chemical and physical pro
availability
properties
nutrient
actively transported membranes and cell walls
to chemical and
Soil
depend on
into roots due to their
sequestration by physical laws information framework into which they movement of solutes throu
anionic soil particles.
• Porous soils leach
water rapidly and can
transactions can insert new knowledge. • Positively charged soil
nutrients must be
contribute to water
stress. actively transported
Information into roots due to their
• The chemical can be sequestration by
properties of clay communicated anionic soil particles.
make it adsorb water in non-
and minerals tightly. chemical • Porous soils leach
• The water potential of ways water rapidly and can
the soil affects the contribute to water
transport of minerals stress.
into the root. • Light can be perceived by plant cell receptors such as Pfr.
• The chemical
• Low soil pH can cause • Signal transduction pathways communicate information received in light signals to plant response mechanisms. properties of clay
toxic aluminum to • Plants can respond to perceived light with changes in gene expression. make it adsorb water
leach from rocks. • Differences in received light wavelength can cause specific plant growth responses. and minerals tightly.
• Salt accumulation in • The environment can signal seeds to germinate using light of specific wavelengths. • The water potential of
soil can affect soil the soil affects the
water potential and • Light containing blue wavelengths can signal phototropic responses.
cause loss of plant transport of minerals
• Some plants can change behavior based on the day/night cycle.
cell turgor. into the root.
• Gravitational fields can trigger directional growth responses.
• Low soil pH can cause
• Some plants can respond to touch. toxic aluminum to
leach from rocks.
• Salt accumulation in
soil can affect soil
water potential and
Preparing Students for the Future xvii cause loss of plant
cell turgor.
Strengthen Problem-Solving Skills with Connect®
Detailed Feedback in Connect®
Learning is a process of iterative development, of making
mistakes, reflecting, and adjusting over time. The question and
test banks in Connect® for Biology, 12th edition, are more than
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ences that systematically build student learning over time.
For many students, choosing the right answer is not
necessarily based on applying content correctly; it is more a
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questions in a decision-tree-style framework to scaffold
xviii Preparing Students for the Future
learning, where each step models and reinforces the learning
process.
The feedback for each higher level Blooms question
(Apply, Analyze, Evaluate) follows a similar process: Clarify
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Unpacking the Concepts
We’ve taken problem solving a step further. In each chapter,
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and test banks are broken out by the steps of the detailed
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steps, such as “What is the key concept addressed by the
question?” before proceeding to answer the question. A
professor can choose which version of the question to include
in the assignment based on the problem-solving skills of the
students.

Graphing Interactives
To help students develop analytical skills, Connect® for Biology,
12th edition, is enhanced with interactive graphing questions.
Students are presented with a scientific problem and the
opportunity to manipulate variables, producing different results
on a graph. A series of questions follows the graphing activity
to assess if the student understands and is able to interpret the
data and results.

Quantitative Question Bank

Many chapters also contain a Quantitative Question Bank.
These are more challenging algorithmic questions, intended to
help your students practice their quantitative reasoning skills.
Hints and guided solution options step students through a
problem.

Preparing Students for the Future xix

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Part I The Molecular Basis of Life

CHAPTER
1
The Science of Biology

Chapter Contents
1.1 The Science of Life
1.2 The Nature of Science
1.3 An Example of Scientific Inquiry:
Darwin and Evolution
1.4 Core Concepts in Biology

Y
©Soames Summerhays/Natural Visions

Introduction
You are about to embark on a journey—a journey of discovery about the nature of life. More than 180 years ago, a young English
naturalist named Charles Darwin set sail on a similar journey on board H.M.S. Beagle; a replica of this ship is pictured here. What
Darwin learned on his five-year voyage led directly to his development of the theory of evolution by natural selection, a theory that has
become the core of the science of biology. Darwin’s voyage seems a fitting place to begin our exploration of biology—the scientific
study of living organisms and how they have evolved. Before we begin, however, let’s take a moment to think about what biology is
and why it’s important.

This is the most exciting time to be studying biology in the history

1.1 The Science of Life of the field. The amount of information available about the natural
world has exploded in the last 42 years, since the construction of
the first recombinant DNA molecule. We are now in a position to
ask and answer questions that previously were only dreamed of.
Learning Outcomes The 21st century began with the completion of the sequence
1. Compare biology to other natural sciences. of the human genome. The largest single project in the history of
2. Describe the characteristics of living systems. biology took about 20 years. Yet less than 15 years later, we can
3. Characterize the hierarchical organization of sequence an entire genome in a matter of days. This flood of se-
living systems. quence data and genomic analysis are altering the landscape of
biology. These and other discoveries are also moving into the
clinic as never before, with new tools for diagnostics and treat-
ment. With robotics, next-generation DNA sequencing technolo-
gies, advanced imaging, and analytical techniques, we have tools
available that were formerly the stuff of science fiction.
In this text, we attempt to draw a contemporary picture of the
science of biology, as well as provide some history and experimen-
tal perspective on this exciting time in the discipline. In this intro-
ductory chapter, we examine the nature of biology and the
foundations of science in general to put into context the informa-
tion presented in the rest of the text.
Biology unifies much of natural science
The study of biology is a point of convergence for the information
and tools from all of the natural sciences. Biological systems are
the most complex chemical systems on Earth, and their many func-
tions are both determined and constrained by the principles of
chemistry and physics. Put another way, no new laws of nature can
be gleaned from the study of biology—but that study does illumi-
nate and illustrate the workings of those natural laws.
The intricate chemical workings of cells can be understood
using the tools and principles of chemistry. And every level of bio-
logical organization is governed by the nature of energy transac-
tions first studied by thermodynamics. Biological systems do not
represent any new forms of matter, and yet they are the most com-
plex organization of matter known. The complexity of living sys-
tems is made possible by a constant source of energy—the Sun.
The conversion of this radiant energy into organic molecules by
photosynthesis is one of the most beautiful and complex reactions
known in chemistry and physics.
CELLULAR LEVEL
Atoms Molecule Macromolecule Organelle
H
N
H
N
C
O
0.2 μm
2 part I The Molecular Basis of Life
The way we do science is changing to grapple with increas-
ingly difficult modern problems. Science is becoming more interdis-
ciplinary, combining the expertise from a variety of traditional
disciplines and emerging fields such as nanotechnology. Biology is at
the heart of this multidisciplinary approach because biological prob-
lems often require many different approaches to arrive at solutions.
Life defies simple definition
In its broadest sense, biology is the study of living things—the
­science of life. Living things come in an astounding variety of
shapes and forms, and biologists study life in many different ways.
They live with gorillas, collect fossils, and listen to whales. They
read the messages encoded in the long molecules of heredity and
count how many times a hummingbird’s wings beat each second.
What makes something “alive”? Anyone could deduce that a
galloping horse is alive and a car is not, but why? We cannot say,
“If it moves, it’s alive,” because a car can move, and gelatin can
wiggle in a bowl. They certainly are not alive. Although we cannot
define life with a single simple sentence, we can come up with a
series of seven characteristics shared by living systems:
■■ Cellular organization. All organisms consist of one or
more cells. Often too tiny to see, cells carry out the basic
activities of living. Each cell is bounded by a membrane that
separates it from its surroundings.
■■ Ordered complexity. All living things are both complex and
highly ordered. Your body is composed of many different
kinds of cells, each containing many complex molecular
structures. Many nonliving things may also be complex, but
they do not exhibit this degree of ordered complexity.
Cell Tissue Organ
100 μm
 ■■ Sensitivity. All organisms respond to stimuli. Plants grow
toward a source of light, and the pupils of your eyes dilate
when you walk into a dark room.
■■ Growth, development, and reproduction. All organisms
are capable of growing and reproducing, and they all possess
hereditary molecules that are passed to their offspring,
ensuring that the offspring are of the same species.
■■ Energy utilization. All organisms take in energy and use it
to perform many kinds of work. Every muscle in your body
is powered with energy you obtain from your diet.
■■ Homeostasis. All organisms maintain relatively constant internal
conditions that are different from their environment, a process
called homeostasis. For example, your body temperature
remains stable despite changes in outside temperatures.
■■ Evolutionary adaptation. All organisms interact with other
organisms and the nonliving environment in ways that
influence their survival, and as a consequence, organisms
evolve adaptations to their environments.
Living systems show hierarchical organization
The organization of the biological world is hierarchical—that is,
each level builds on the level below it:
1. The cellular level. At the cellular level (figure 1.1),
atoms, the fundamental elements of matter, are joined
together into clusters called molecules. Complex
biological molecules are assembled into tiny structures
called organelles within membrane-bounded units
we call cells. The cell is the basic unit of life. Many
­independent organisms are composed only of single
ORGANISMAL LEVEL
Organ system Organism Population
cells. Bacteria are single cells, for example. All
animals and plants, as well as most fungi and algae,
are multicellular—composed of more than one cell.
2. The organismal level. Cells in complex multicellular
organisms exhibit three levels of organization. The most
basic level is that of tissues, which are groups of similar
cells that act as a functional unit. Tissues, in turn, are
grouped into organs—body structures composed of
several different tissues that act as a structural and
­functional unit. Your brain is an organ composed of
nerve cells and a variety of associated tissues that form
protective coverings and contribute blood. At the third
level of organization, organs are grouped into organ
systems. The nervous system, for example, consists of
sensory organs, the brain and spinal cord, and neurons
that convey signals.
Figure 1.1 Hierarchical organization of living systems.
Life forms a hierarchy of organization from atoms to complex
multicellular organisms. Atoms are joined together to form molecules,
which are assembled into more complex structures such as organelles.
These in turn form subsystems that provide different functions. Cells
can be organized into tissues, then into organs and organ systems such
as the goose’s nervous system pictured. This organization then extends
beyond individual organisms to populations, communities, ecosystems,
and finally the biosphere. (Organelle): ©Keith R. Porter/Science Source; (Cell):
©STEVE GSCHMEISSNER/Getty Images; (Tissue): ©Ed Reschke; (Organism):
©Russell Illig/Getty Images; (Population): ©George Ostertag/age fotostock;
(Species): ©iStockphoto/Getty Images; ©Pictureguy/Shutterstock; (Community):
©Ryan McGinnis/Alamy; (Ecosystem): ©Robert and Jean Pollock; (Biosphere):
Source: NASA
POPULATIONAL LEVEL
Species Community Ecosystem Biosphere
chapter 1 The Science of Biology 3
 3. The populational level. Individual organisms can
be categorized into several hierarchical levels within
the living world. The most basic of these is the
population—a group of organisms of the same species
living in the same place. All populations of a particular
kind of organism together form a species, its members
similar in appearance and able to interbreed. At a higher
level of biological organization, a biological community
consists of all the populations of different species living
together in one place.
4. The ecosystem level. At the highest tier of biological
organization, populations of organisms interact with
each other and their physical environment. Together
populations and their environment constitute an
ecological system, or ecosystem. For example, the
biological community of a mountain meadow interacts
with the soil, water, and atmosphere of a mountain
ecosystem in many important ways.
5. The biosphere. The entire planet can be thought of as an
ecosystem that we call the biosphere.
As you move up this hierarchy, the many interactions occurring at
lower levels can produce novel properties. These so-called
emergent properties may not be predictable. Examining individ­
ual cells, for example, gives little hint about the whole animal.
Many weather phenomena, such as hurricanes, are actually emer-
gent properties of many interacting meteorological variables. It is
because the living world exhibits many emergent properties that it
is difficult to define “life.”
This description of the common features and organization of
living systems provides an introduction for our exploration of biol-
ogy. Before we continue, we will consider the broader question of
the nature of science itself.
Learning Outcomes Review 1.1
Biology as a science brings together other natural sciences, such
as chemistry and physics, to study living systems. Life does not
have a simple definition, but living systems share a number of
properties that together describe life. Living systems can be
organized hierarchically, from the cellular level to the entire
biosphere, with emergent properties that may exceed the sum of
the parts.
■■ Can you study biology without studying other sciences?
1.2 The Nature of Science
Learning Outcomes
1. Compare the different types of reasoning used by biologists.
2. Demonstrate how to formulate and test a hypothesis.
Much like life itself, the nature of science defies simple descrip-
tion. For many years scientists have written about the “scientific
4 part I The Molecular Basis of Life
method” as though there is a single way of doing science.
This oversimplification has contributed to confusion on the part of
nonscientists about the nature of science.
At its core, science is concerned with developing an increas-
ingly accurate understanding of the world around us using observa-
tion and reasoning. To begin with, we assume that natural forces
acting now have always acted, that the fundamental nature of the uni-
verse has not changed since its in­ception, and that it is not changing
now. A number of complementary approaches allow understanding
of natural phenomena—there is no one “scientific method.”
Scientists also attempt to be as objective as possible in the
interpretation of the data and observations they have collected.
Because scientists themselves are human, this is not completely
possible, but because science is a collective endeavor subject to
scrutiny, it is self-correcting. One person’s results are verified by
others, and if the results cannot be repeated, they are rejected.
Much of science is descriptive
The classic vision of the scientific method is that observations lead
to hypotheses that in turn make experimentally testable predic-
tions. In this way, we dispassionately evaluate new ideas to arrive
at an increasingly accurate view of nature. We discuss this way of
doing science later in this section but it is important to understand
that much of science is purely descriptive: In order to understand
anything, the first step is to describe it completely. Much of biol-
ogy is concerned with arriving at an increasingly accurate descrip-
tion of nature.
The study of biodiversity is an example of descriptive sci-
ence that has implications for other aspects of biology in addition
to societal implications. Efforts are currently under way to classify
all life on Earth. This ambitious project is purely descriptive, but it
will lead to a much greater understanding of biodiversity as well as
the effect our species has on biodiversity.
One of the most important accomplishments of molecular
biology at the dawn of the 21st century was the completion of
the sequence of the human genome. Many new hypotheses
about human biology will be generated by this knowledge,
and many experiments will be needed to test these hypotheses,
but the determination of the sequence itself was descriptive
science.
Science uses both deductive
and inductive reasoning
The study of logic recognizes two opposite ways of arriving at
logical conclusions: deductive and inductive reasoning. Science
makes use of both of these methods, although induction is the
­primary way of reasoning in hypothesis-driven science.
Deductive reasoning
Deductive reasoning applies general principles to predict spe-
cific results. More than 2200 years ago, the Greek scientist
­Eratosthenes used Euclidean geometry and deductive reasoning
to accurately estimate the circumference of the Earth (­figure 1.2).
Deductive reasoning is the reasoning of mathematics and ­phi-
losophy, and it is used to test the validity of general ideas in all
 Figure 1.2 Deductive reasoning: How Eratosthenes estimated
the circumference of the Earth using deductive reasoning. 1. On a
Sunlight Height of Light rays day when sunlight shone straight down a deep well at Syene in Egypt,
at midday obelisk parallel
Eratosthenes measured the length of the shadow cast by a tall obelisk in the city
of Alexandria, about 800 kilometers (km) away. 2. The shadow’s length and the
obelisk’s height formed two sides of a triangle. Using the recently developed
Well a principles of Euclidean geometry, Eratosthenes calculated the angle, a, to be 7°
Distance between and 12´, exactly 1⁄50 of a circle (360°). 3. If angle a is 1⁄50 of a circle, then the
cities = 800 km distance between the obelisk (in Alexandria) and the well (in Syene) must be
Length of equal to 1⁄50 the circumference of the Earth. 4. Eratosthenes had heard that
shadow
it was a 50-day camel trip from Alexandria to Syene. Assuming a camel
travels about 18.5 km per day, he estimated the distance between obelisk
and well as 925 km (using different units of measure, of course).
5. Eratosthenes thus deduced the circumference of the Earth to be
50 × 925 = 46,250 km. Modern measurements put the distance from
the well to the obelisk at just over 800 km. Using this distance
a Eratosthenes’s value would have been 50 × 800 = 40,000 km. The
actual circumference is 40,075 km.

Observation

Question
branches of knowledge. For example, if all mammals by defini-
tion have hair, and you find an animal that does not have hair,
then you may conclude that this animal is not a mammal. A bi- Hypothesis 1
Hypothesis 2 Reject
ologist uses deductive reasoning to infer the species of a speci- Potential
Hypothesis 3 Experiment hypotheses
hypotheses
men from its characteristics. Hypothesis 4 1 and 4
Hypothesis 5

Inductive reasoning
Remaining Hypothesis 2 Reject
In inductive reasoning, the logic flows in the opposite direction, possible Hypothesis 3 Experiment hypotheses
from the specific to the general. Inductive reasoning uses specific hypotheses Hypothesis 5 2 and 3
observations to construct general scientific principles. For example,
if poodles have hair, and terriers have hair, and every dog that you Last remaining
observe has hair, then you may conclude that all dogs have hair. In- possible Hypothesis 5
hypothesis
ductive reasoning leads to generalizations that can then be tested. Modify hypothesis
Inductive reasoning first became important to science in the 1600s
in Europe, when Francis Bacon, Isaac Newton, and others began to Predictions
use the results of particular experiments to infer general principles
about how the world operates.
An example from modern biology is the role of homeobox Experiment 1 Experiment 2 Experiment 3 Experiment 4
genes in development. Studies in the fruit fly, Drosophila melano-
gaster, identified genes that could cause dramatic changes in de-
velopmental fate, such as a leg appearing in the place of an antenna. Predictions
These genes have since been found in essentially all multicellular confirmed
animals analyzed. This led to the general idea that homeobox
genes control developmental fate in animals. Figure 1.3 How science is done. This diagram illustrates
how scientific investigations proceed. First, scientists make
Hypothesis-driven science observations that raise a particular question. They develop a number
of potential explanations (hypotheses) to answer the question. Next,
makes and tests predictions they carry out experiments in an attempt to eliminate one or more of
Scientists establish which general principles are true from among these hypotheses. Then, predictions are made based on the remaining
the many that might be true through the process of systematically hypotheses, and further experiments are carried out to test these
testing alternative proposals. If these proposals prove inconsistent predictions. The process can also be iterative. As experimental results
with experimental observations, they are rejected as untrue. are performed, the information can be used to modify the original
Figure 1.3 illustrates the process. hypothesis to fit each new observation.

chapter 1 The Science of Biology 5

 After making careful observations, scientists construct a
hypothesis, which is a suggested explanation that accounts for
those observations. A hypothesis is a proposition that might be
true. Those hypotheses that have not yet been disproved are re-
tained. They are useful because they fit the known facts, but they
are always subject to future rejection if, in the light of new infor-
mation, they are found to be incorrect.
This is usually an ongoing process with a hypothesis chang-
ing and being refined with new data. For instance, geneticists
George Beadle and Edward Tatum studied the nature of genetic
information to arrive at their “one-gene/one-enzyme” hypothesis
(see chapter 15). This hypothesis states that a gene represents the
genetic information necessary to make a single enzyme. As inves-
tigators learned more about the molecular nature of genetic infor-
mation, the hypothesis was refined to “one gene/one polypeptide”
because enzymes can be made up of more than one polypeptide.
With still more information about the nature of genetic informa-
tion, other investigators found that a single gene can specify more
than one polypeptide, and the hypothesis was refined again.
Testing hypotheses
We call the test of a hypothesis an experiment. Suppose you enter
a dark room. To understand why it is dark, you propose several
hypotheses. The first might be, “There is no light in the room be-
cause the light switch is turned off.” An alternative hypothesis
might be, “There is no light in the room because the lightbulb is
burned out.” And yet another hypothesis might be, “I am going
blind.” To evaluate these hypotheses, you would conduct an ex-
periment designed to eliminate one or more of the hypotheses.
For example, you might test your hypotheses by flipping the
light switch. If you do so and the room is still dark, you have dis-
proved the first hypothesis: Something other than the setting of the
light switch must be the reason for the darkness. Note that a test
such as this does not prove that any of the other hypotheses are
true; it merely demonstrates that the one being tested is not. A suc-
cessful experiment is one in which one or more of the alternative
hypotheses is demonstrated to be inconsistent with the results and
is thus rejected.
As you proceed through this text, you will encounter many
hypotheses that have withstood the test of experiment. Many will
continue to do so; others will be revised as new observations are
made by biologists. Biology, like all science, is in a constant state of
change, with new ideas appearing and replacing or refining old ones.
Establishing controls
Often scientists are interested in learning about processes that are
influenced by many factors, or variables. To evaluate alternative hy-
potheses about one variable, all other variables must be kept constant.
This is done by carrying out two experiments in parallel: an experi-
mental treatment or group and a control treatment or group. In
the experimental treatment, one variable is altered in a known way to
test a particular hypothesis. In the control treatment, that variable is
left unaltered. In all other respects the two experiments are identical,
so any difference in the outcomes of the two experiments must result
from the influence of the variable that was changed.
Much of the challenge of experimental science lies in de-
signing control experiments that isolate a particular variable from
other factors that might influence a process.
6 part I The Molecular Basis of Life
Using predictions
A successful scientific hypothesis needs to be not only valid but
also useful—it needs to tell us something we want to know. A hy-
pothesis is most useful when it makes predictions because those
predictions provide a way to test the validity of the hypothesis. If
an experiment produces results inconsistent with the predictions,
the hypothesis must be rejected or modified. In contrast, if the pre-
dictions are supported by experimental testing, the hypothesis is
supported. The more experimentally supported predictions a hy-
pothesis makes, the more valid the hypothesis is.
As an example, in the early history of microbiology it was
known that nutrient broth left sitting exposed to air becomes con-
taminated. Two hypotheses were proposed to explain this observa-
tion: spontaneous generation and the germ hypothesis. Spontaneous
generation held that there was an inherent property in organic mol-
ecules that could lead to the spontaneous generation of life. The
germ hypothesis proposed that preexisting microorganisms that
were present in the air could contaminate the nutrient broth.
These competing hypotheses were tested by a number of ex-
periments that involved filtering air and boiling the broth to kill
any contaminating germs. The definitive experiment was per-
formed by Louis Pasteur, who constructed flasks with curved
necks that could be exposed to air, but that would trap any con-
taminating germs. When such flasks were boiled to sterilize them,
they remained sterile, but if the curved neck was broken off, they
became contaminated (figure 1.4).
SCIENTIFIC THINKING
Question: What is the source of contamination that occurs in a flask
of nutrient broth left exposed to the air?
Germ Hypothesis: Preexisting microorganisms present in the air
contaminate nutrient broth.
Prediction: Sterilized broth will remain sterile if microorganisms are
prevented from entering flask.
Spontaneous Generation Hypothesis: Living organisms will
spontaneously generate from nonliving organic molecules in broth.
Prediction: Organisms will spontaneously generate from organic
molecules in broth after sterilization.
Test: Use swan-necked flasks to prevent entry of microorganisms. To
ensure that broth can still support life, break swan-neck after sterilization.
Broken
neck
of flask
Flask is sterilized Unbroken flask Broken flask becomes
by boiling the broth. remains sterile. contaminated after
exposure to germ-laden air.
Result: No growth occurs in sterile swan-necked flasks. When the
neck is broken off, and the broth is exposed to air, growth occurs.
Conclusion: Growth in broth is of preexisting microorganisms.
Figure 1.4 Experiment to test spontaneous generation
versus germ hypothesis.
Another random document with
no related content on Scribd:
The Project Gutenberg eBook of Sunfire!
This ebook is for the use of anyone anywhere in the United States
and most other parts of the world at no cost and with almost no
restrictions whatsoever. You may copy it, give it away or re-use it
under the terms of the Project Gutenberg License included with this
ebook or online at www.gutenberg.org. If you are not located in the
United States, you will have to check the laws of the country where
you are located before using this eBook.

Title: Sunfire!

Author: Edmond Hamilton

Illustrator: Virgil Finlay

Release date: November 20, 2023 [eBook #72181]

Language: English

Original publication: New York, NY: Ziff-Davis Publishing Company,

1962

Credits: Greg Weeks, Mary Meehan and the Online Distributed

Proofreading Team at http://www.pgdp.net

*** START OF THE PROJECT GUTENBERG EBOOK SUNFIRE! ***

 SUNFIRE!

By EDMOND HAMILTON

Illustrated by FINLAY

He was walking in the pine grove, with

the resinous smell of the trees in his
nostrils. Once he had met a smell
vaguely like it, far away from Earth.
Forget about that, a voice said in his
mind, but he would never forget.

[Transcriber's Note: This etext was produced from

Amazing Stories September 1962.
Extensive research did not uncover any evidence that
the U.S. copyright on this publication was renewed.]
Everything in the old house seemed just the same as it had been
before he went to space.
It was incredible, thought Hugh Kellard, standing in the front hall and
looking around the silent, sunlit rooms, how little it had changed. The
life was gone out of it now, all the people and voices and the comings
and goings when his grandfather still lived and he had visited here.
But that had been long ago, and he was amazed that so much
remained still untouched.
"Like travelling into the past," thought Kellard, "to come back to this
part of Earth."
He was tired, in body and mind and nerve, and he stood for a while,
just staring. The agent who cared for the old place had let him in and
gone away, and there was not a sound in the house. He walked into
the living-room where his grandfather's desk still stood beneath a
window, and looked out. The window faced northward, along the
California coastal cliffs that run north along Morro Bay to Big Sur. The
Pacific foamed and surged against the huge broken stones beneath
the cliffs, and the hills, somber now with a tinge of autumn,
shouldered massively up toward the east from the cliff road. It all
looked as lonely as ever, no other houses in sight but this gray,
weatherbeaten house that had faced the sea-wind and the sea-fog for
over a hundred years.
Kellard walked back along the hall. On its walls still hung the ornately
framed family photographs which his grandfather had stubbornly kept
in place. His great-grandfather, and his great-aunt something, and all
the rest of them, on back into the shadows. They were all there, they
had not been touched, nothing in the house had been touched, just
as his grandfather's will had enjoined. Keep the old house, he had
said. Some of the family will be back some day.
The old man had been right, he thought. One of the family had come
back at last, one who had roamed farther than almost anybody on
Earth.
"But that's all done with," he told himself. "Here I am, and here I stay.
I'm through with space."

He started through the rooms, opening windows, letting in light and

air. The furniture was faded and old-fashioned, but the place was not
dusty, the agent had seen that it was kept in shape. Kellard picked
one of the big upstairs bedrooms for himself, and brought in the
blankets and cartons and luggage from the car. He went into the
utility room and turned on the power-unit, remembering as he did so
how his grandfather had disliked and distrusted the unit, how he had
refused to have one until the electric wires were all gone and there
was no other way to get power. He checked the stove and freezer,
shoved his cartons of food into the latter, and then looked around and
wondered what to do next.
Standing in the silent house, he wondered suddenly if he had been
foolish to quit everything and come back to Earth and this old place?
No, he thought heavily. Mercury ended it for me. I made my decision
and that is that. Forget it.
He strode abruptly out of the house and started walking. And after a
little while the dark weight in his mind, the somber knowledge, faded
and receded in the new-found, old-remembered interest of the things
about him.
His way took him across the road, past the shabby barns and up
sloping pastures where once his grandfather had kept the fine horses
he bred. Then he was in among the pines, climbing more steeply, with
the resinous smell of the trees strong in his nostrils. That smell he
had never forgotten, and once he had met a smell vaguely like it, far
away from Earth—
Forget all that, Kellard.
The trees took him in and he walked through a dapple of sun and
shadows. A deer slipped away through the pines ahead of him, and
quail burst up from almost under his feet. He remembered a grove of
bigger pines farther up the slope, and an old man and a boy walking
up to them. How long ago was that? He had been fifteen—and he
was thirty-two now. Seventeen years. Still, he thought he could find
the place.
He found it. The big pines were still there, for people did not use
wood much any more. The rough dark giants stood at dignified
intervals from each other, and he sat down with his back against the
massive trunk of the biggest.
Funny, he thought. When I was a boy sitting here dreaming about the
future and what I was going to do, I never once imagined that some
things would stay much the same. The whole world would somehow
be miraculously transformed—but it wasn't. This tree was here when
men first reached the Moon, and Mars, and Venus and the rest, but it
didn't know about that, it didn't change because of it.

Kellard sat for a long time, still wrapped in a gray weariness, his
emotions in a numb trance. He sat listening to the distant, uneasy
murmur of the sea, until the sunset light shafting through the trees
dazzled his eyes, and then he got up and went back down to the
house. He heated food, ate it, and then went out to the porch in front
of the house and sat watching the sun sink toward the vast golden
sheet of the Pacific. He thought of the little dot close to the sun that
he could not see, the little world and the strange, terrible place upon it
where Morse and Binetti had died.
The telephone rang.
Kellard did not stir, and it rang and rang again.
Go ahead and ring your head off, he thought. You're not getting me
back. I told you. I've had it.
The ringing stopped. The sun sank and darkness came with the hosts
of wheeling stars, and there was no sound but the vast voices rolling
in from sea, as Kellard sat staring and drinking.
He finally got up, as the fog started coming in. He moved with gravity,
feeling much better. He went in and turned on the lights, and then
looked at the faces that stared from the long row of framed
photographs.
He raised the bottle to them in a gesture of salutation.
"You see, Kellards, that your prodigal son—or great-grandson—has
come home again from space."
He gravely drank, and continued to stand looking along the faded
faces.
"You were lucky—you know that? Back in your time, there were
hopes, and dreams, and man's road would go on forever, from
triumph to triumph everlasting. But that road was a blind alley, all the
time, even if I'm the only one who knows it."
The faces looked back at him, unchanging, but he read reproach in
their steady gaze, their lined features.
"I'm sorry," said Kellard. "You had your own troubles, I know. I
apologize, Kellards. I am very tired and a little drunk, and I am going
to bed."

The next morning he was making coffee when there came a banging
of the old-fashioned knocker on the front door. A certain tightness
came into Kellard's face. He had expected them to send some one.
He had not expected the man who stood at the door. He was not in
Survey uniform, although he was the highest brass there was. He
was a big, slow-moving man with a heavy face and blue eyes that
seemed mild if you didn't know him.
"Well," said Kellard. And after a moment, "Come on in."
Halfrich came in. He sat down and looked interestedly around at the
old room and furniture.
"Nice," he murmured. Then he looked at Kellard and said, "All right,
let's have it. Why did you quit?"
Kellard shrugged. "It was all in my letter of resignation. I'm getting a
bit old and tired for Survey, I—"
"Bull," said Halfrich. "It was something about that crack-up on
Sunside, wasn't it?"
Kellard said slowly, "Yes. The deaths of Binetti and Morse, and the
after-effects of that shock, made me feel I didn't have it any more."
Halfrich looked at him. "You've had crack-ups before. You've seen
men die. You've had almost as many years in Survey as I have, and
you've taken as many jolts. You're lying, Kellard."
Kellard got up, and walked a few steps and swung around again.
"So I'm lying. I want out, and what difference does it make why?"
"It makes a difference," Halfrich said grimly. "I remember from away
back at Academy, even though you were two years after me. You
were the space-craziest cadet there was. You spouted the glories of
the conquest of space until we were all sick of it. You haven't
changed in all the years in Survey—until now. I want to know what
can change a man like that."
Kellard said nothing. He went to the window and looked out at the
long rollers coming endlessly in and crashing against the rocks.
"What did you see on Sunside, Kellard?"
He turned around sharply at that.
"What do you mean? What would there be to see there, but hot rocks
and volcanoes and a cross-section of hell generally? It's all in my
report."
Halfrich sat like a judge, and spoke like one pronouncing sentence.
"You saw something, you met something there. You covered by
tearing out the film of the automatic sweep-camera. Whatever it had
recorded, you didn't want us to see, did you?"
Kellard came toward him and spoke angrily and rapidly. "Do you
realize that we flamed out and crashed there? A crash like that can
do damage. It killed Binetti and mortally injured Morse, and smashed
the sweep-camera."
Halfrich nodded. "That's what we thought, at first. But the radar-
sweep had an automatic recorder too. It was something new. Binetti
knew about it, as communications officer, but I guess he hadn't told
you, or you'd have smashed it too. Its record shows something."

A cold feeling came over Kellard. He had thought that he had covered
everything, but he had calculated from insufficient data.
He kept his nerve. A radar record was not like a photograph, they
couldn't prove much from that, they certainly couldn't guess the truth
from it. They must not guess the truth.
He laughed mirthlessly. "A radar record made on Sunside isn't worth
the paper it's on. The storms of radiation there make radar practically
unreliable."
Halfrich was watching him keenly. "But not entirely. And over and
above the static and the fake bogies, the record shows quite clearly
that you went outside the ship after the crash, that you walked about
a thousand yards, and that you were approached by some things that
register vaguely but unmistakably."
He paused and then he asked, "Who—or what—did you meet there,
Kellard?"
Kellard was cold inside, but all the same he made a disgusted sound
that he hoped was convincing.
"Who would I meet on Sunside? Beautiful lightly-clad maidens? After
all, you know, it's only four hundred degrees Centigrade there, and
practically no atmosphere, and nothing much else but solar radiation
and hot rock and volcanoes. I tell you, the radar record is worthless."
Halfrich was studying him with that mild estimating look that Kellard
knew well, and didn't like at all. It was the look that came into
Halfrich's face when friendship didn't matter and the good of the
Survey did.
"You're still lying," he said. "You met or saw something there. And it
did something to you—something that made you resign. Something
that's taken all the life and eagerness out of you."
"Oh, hell, be reasonable!" said Kellard angrily. "You know no kind of
life can exist on Sunside. My mission was the second time even
Survey has landed there. Pavlik's mission, the first, didn't see
anything. Neither did I. Quit dreaming it up. Go back to Mojave and
your job, and leave me be."

Halfrich rose. "All right," he said. "I'll go back to the base. And you're
going with me."
"Oh, no," said Kellard. "I'm through, quit, resigned."
"Your resignation has not been accepted," Halfrich told him. "You're
still liable to Survey discipline. You'll obey orders just as you always
did, or you'll go up before a court-martial."
"So that's it," said Kellard.
Halfrich nodded. "That is it. I don't like to do this. You're an old friend.
But—"
"But the Survey comes first," Kellard said, between his teeth.
"The Survey," said Halfrich, "comes first. It has to. It's why we've got
stations on Venus and Mars and Ganymede, not to say the Moon. It's
why we'll someday be able to hit for deep space and the starworlds.
And when one of my best officers suddenly goes off the deep end
and won't say why, I'll damn well wring it out of him. Whatever you
found on Mercury doesn't belong to you, it belongs to us, and we'll
have it."
Kellard looked at him and started to say something and didn't, and
then turned his back on Halfrich and looked out the window at the
sea. In a low voice he said,
"Let it be, John. I'm telling you now, you'll be sorry if you don't."
There was no answer to that at all, and the silence was his answer.
He turned back around.
"All right, you have a rope around my neck. I'll go back to base with
you. I'll tell you not one thing more than here."
"In which case," Halfrich said, "we'll go on out to Sunside, and you'll
go right along with us."
A rage born of desperation came to Kellard. He had tried to spare
people this—Halfrich, the Survey, the whole human race. But they
would not let it be so. Damn them, he thought, if they must do this,
they have it coming to them.
"All right," he said flatly. "I'll get my jacket. I take it that you have a flier
waiting."

The fast flier, less than an hour later, whizzed down over the gaunt
mountains and across the desert, and the glitter and splendor of
Mojave Base sprang up to meet them. The tall ships shone like silver,
and something about them, something about the feel of the place,
made you think that this bit of desert did not belong to Earth at all but
was part of space, a way-station, the first way-station of all, to the
stars.
That, thought Kellard, was what he had thought when he had first
come here, years ago. And it had not been just a youngster's passing
enthusiasm, it had deepened and strengthened through all the years
of work and danger—until Sunside. And oh God, he thought, why did
I have to go there, at that place, at that moment. I could have lived
my whole life and done my work, all of us could have, without ever
dreaming the truth.
He knew now that he had no choice. He must go back to Sunside
with them. For even if he told them the truth, they would not believe,
they would insist on going to see for themselves. He would keep
silent, and that was all he could do now.
Four days later a Y-90 experimental cruiser, outfitted for space
research and with full anti-heater equipment, took off from Mojave.
Kellard had kept silent. And still silent he sat in his recoil-harness and
took the jolts, and heard Halfrich grunting beside him, and viciously
hoped that that he was not liking it.
Halfrich had brought along a consulting biophysicist, a keen-faced
man of middle age named Morgenson, who did not look as though he
was enjoying the mission either. But the three-man crew of the little Y-
90 were young men in their twenties. They spoke to Halfrich and to
Kellard as though they were heroes out of legend, for in the Survey
twelve to fifteen years of space-missions was an age.
It was only after they had gone a long way and a long time through
the sunwashed spaces that one of the three, Shay, the navigator,
ventured to put a question to Kellard.
"You were with the first mission to Ganymede, sir, weren't you?"
Kellard nodded. "Yes, I was."
"Wouldn't that have been something!" said Shay. "I mean, to be the
first."
"It was something," said Kellard.
"Maybe someday I——" Shay began, and broke off and then went on,
"I mean, if the star-drive is perfected as soon as some people say it
will be, I could maybe be one of the first ones out there? Sir?"
"You could be," said Kellard. "Someone's going to be first. The stars
are waiting. All we have to do is go out there and keep going, and the
stars will be ours, just like the planets here are, all ours, forever and
amen."
Shay looked at him puzzledly, and shuffled, and then went away.
Halfrich had been listening, and watching. He said, "Did you have to
slap the kid's face?"
Kellard shrugged. "What did I say? I was merely repeating what
everyone feels, these days. The glory of the conquest of space."
"I'd give a lot," Halfrich said, "to know what's riding you. We'll soon
reach Sunside and we'll find out, but I wish you'd tell me now."
"All right," said Kellard. "I'll tell you. I've been disinherited. That's
what's wrong with me."

He would say nothing more, nor did Halfrich ask him another
question, until the Y-90 was far in past the orbit of Venus and going
into its pattern of approach.
"I assume," said Halfrich, "that you bear none of us any personal ill-
will. If there is anything dangerous awaiting us, now would be the
time to tell us."
Kellard considered. "You're going to land, I suppose, at the same spot
where we crashed."
"Of course."
"Then land," said Kellard. "As far as I know, there is not a thing there
to harm you."
In the scanner, he watched Mercury swing slowly toward them, a tiny
crescent of white that was hard to see against the Sun. For here the
Sun was a monster thing, fringed with writhing flames, paling the
stars, drenching this whole area with radiation that already would
have killed them but for the ship's anti-heaters.
Kellard remembered that when he had come this way before, Binetti
had quoted something, a line from William Blake's poems, he had
said. "The desire of the moth for the star." And that was what we
were, he thought. Three little moths, going right into the furnace, and I
was the only one to get out of it, but now I'm going back.
The Y-90 went into its landing pattern. It skimmed over the dark side
of Mercury, the black cliffs and peaks and chasms that never saw the
Sun, and then light seemed to burst ragingly up from all the horizon
ahead of them, and they were over Sunside.
In old days this little world had been called "the moon of the Sun,"
and it looked like it, the same stark, lifeless rock plains and ridges
and cracks, the fang-like look of pinnacles in a place where no
atmosphere eroded anything. But the Moon was cold and still,
whereas Sunside seemed to throb with sullen hidden fires. Volcanoes
spewed ash and lava, and the infernal storm of radiation from
overhead made everything quiver in a shimmering haze. The
indicator board told them that the temperature of the outside hull was
climbing to four hundred as the Y-90 went down.
And the wide valley that haunted his dreams opened up ahead.
Across it the squat volcanic cones still dribbled ash and dust and it
was all just as it had been when he had last looked back from the
relief cruiser that had come from Venus Station to take him off. And
there gleamed bright on its floor the crumpled wreck in which Binetti
and then Morse had died.
Kellard's gaze flew to the place north of the wreck, the tumbled, odd-
shaped rocks. He felt his palms sweating. Maybe there would be
nothing. After all, could it all happen again?
They set down, and after the crashing rocket uproar, the steady throb
of the anti-heaters was an anti-climactic sound.
"You've got the armor ready?" Halfrich asked of Morgenson.
The biophysicist nodded nervously. "Three suits, with their anti-heater
equipment tested on and off all the way out."
"One suit stays here, for emergencies," Halfrich said. "Kellard and I
will go out, when there's something to go out for. First, we'll make
observations."

The recording telescope-cameras and the radar, Halfrich ordered

focused on the place of the odd-shaped rocks. And then, sitting there
on Sunside, they watched. They waited.
Nothing.
Kellard's hopes began to rise. He was right, he told himself, it couldn't
happen again.
"How long," he asked, "are we going to sit waiting for nothing
because a radar made a screwy record? If those anti-heaters quit for
five minutes, we're fried."
Halfrich looked at him bleakly. "I'll tell you how long. Till you tell the
truth, and we see the truth for ourselves. That's how long."
Kellard shrugged. "If that's the way you want it. I would tell you to go
to hell except that we're already there."
They watched and waited some more.
Morgenson said, on a rising note of excitement, "There's something
——"
Halfrich got to the 'scope fast. Kellard, looking through the scanner,
saw the geyser of flame that was beginning to pour up from the rocks.
It grew slowly, but steadily, in height.
"What is it?" Halfrich asked him.
"Can't you see for yourself?" said Kellard. "There's a blowhole out
there and it throws off burning gases from the interior. It did it twice
while I was waiting in the wreck."
Halfrich said, "It's in the same location where radar recorded you
before, with those other blips. There's something about this—We'll go
have a look."
"If you must," said Kellard. "You'll find it's just what I've said."
They got into the heat-armor. It was a clumsy outfit, for it had to have
room for an efficient anti-heater, and the long tube of the heat-
discharge was a nuisance. Kellard had spent days in one of these
suits, waiting for the relief ship after the crack-up, and he did not like
the feel of it at all.
Halfrich tested the radio and then said, "All right, Shay, lock us out
and stand by. Morgenson, you keep watching."
They stepped upon Sunside.
There beat down upon them such a storm of radiation, such cataracts
of heat and light, that instinctively they bowed their heads as before a
deluge. It took an effort of will to step forward through that tempest,
but Halfrich made it. They walked, slowly and heavily, and at first they
saw only the blackened rocks beneath their feet, and the little puddles
and rivulets of molten lead, and their own massive armored feet
plodding.
Then, as they went forward, they straightened against the impact.
Through the face-plate of his armor, dimmed by the many-layered
filters, Kellard saw the column of flame ahead. It was a hundred feet
high now, and growing higher, and though there was no air-borne
sound on this almost airless world, the sound of it came through the
rocks and the soles of their feet, a throbbing and roaring that quivered
through all their bodies.

They reached the tumbled rocks, and stopped. And now the fire-
fountain was so lofty that they had to lean back their heads to look at
its topmost crest. Some unthinkable diastole and systole of the fiery
planet was at work, and this periodic geyser of flame was its result.
The rocks shook and roared, and the fires raged higher, and Kellard
thought again, what devil is in the blood of our race that drives us to
places like this where we should not be?