Biology EIGHTH EDITIONEldra P. Solomon Linda R. Berg Diana W. MartinUniversity of South Florida St. Petersburg College Rutgers University Australia • Canada • Mexico • Singapore • Spain • United Kingdom • United States
DEDICATIONTo our families, friends, and colleagues who gave freely of theirlove, support, knowledge, and time as we prepared this eighthedition of BiologyEspecially toRabbi Theodore and Freda BrodAlan and JenniferChuck and Margaret
ABOUT THE AUTHORSELDRA P. SOLOMON has written LINDA R. BERG is an award-winning DIANA W. MARTIN is the Directorseveral leading college-level textbooks teacher and textbook author. She re- of General Biology, Division of Lifein biology and in human anatomy and ceived a B.S. in science education, an Sciences, at Rutgers University, Newphysiology. Her books have been trans- M.S. in botany, and a Ph.D. in plant Brunswick Campus. She received anlated into more than 10 languages. physiology from the University of Mary- M.S. at Florida State University, whereDr. Solomon earned an M.S. from the land. Her research focused on the evo- she studied the chromosomes of relatedUniversity of Florida and an M.A. and lutionary implications of steroid biosyn- plant species to understand their evolu-Ph.D. from the University of South thetic pathways in various organisms. tionary relationships. She earned a Ph.D.Florida. Dr. Solomon taught biology and Dr. Berg taught at the University of at the University of Texas at Austin,nursing students for more than 20 years. Maryland at College Park for 17 years where she studied the genetics of theShe is adjunct professor and member of and at St. Petersburg College in Florida fruit fly, Drosophila melanogaster, andthe Graduate Faculty of the University for 8 years. During her career, she taught then conducted postdoctoral researchof South Florida. introductory courses in biology, botany, at Princeton University. She has taught In addition to being a biologist and and environmental science to thousands general biology and other courses atscience author, Dr. Solomon is a bio- of students. At the University of Mary- Rutgers for more than 20 years and haspsychologist with a special interest in the land, she received numerous teaching been involved in writing textbooks sinceneurophysiology of traumatic experience. and service awards. Dr. Berg is also the 1988. She is immensely grateful that herHer research has focused on the relation- recipient of many national and regional decision to study biology in college hasships among stress, emotions, and health awards, including the National Science led to a career that allows her many waysand on post-traumatic stress disorder. Teachers Association Award for Innova- to share her excitement about all aspects Dr. Solomon has presented her work in tions in College Science Teaching, the of biology.plenary sessions and scientific meetings at Nation’s Capital Area Disabled Studentmany national and international confer- Services Award, and the Washingtonences. She has been profiled more than Academy of Sciences Award in Univer-20 times in leading publications, includ- sity Science Teaching.ing Who’s Who in America, Who’s Who During her career as a professionalin Science and Engineering, Who’s Who in science writer, Dr. Berg has authored orMedicine and Healthcare, Who’s Who in co-authored several leading college sci-American Education, Who’s Who of Ameri- ence textbooks. Her writing reflects hercan Women, and Who’s Who in the World. teaching style and love of science.
Brief ContentsPreface xxvii 14 Gene Regulation 304To the Student xxxiii 15 DNA Technology and Genomics 322 16 The Human Genome 346 17 Developmental Genetics 368Part 1 THE ORGANIZATION OF LIFE 1 1 A View of Life 1 2 Atoms and Molecules: The Chemical Basis Part 4 THE CONTINUITY OF LIFE: of Life 25 EVOLUTION 390 3 The Chemistry of Life: Organic Compounds 45 18 Introduction to Darwinian Evolution 390 4 Organization of the Cell 73 19 Evolutionary Change in Populations 412 5 Biological Membranes 106 20 Speciation and Macroevolution 428 6 Cell Communication 134 21 The Origin and Evolutionary History of Life 447 22 The Evolution of Primates 466Part 2 ENERGY TRANSFER THROUGH LIVING SYSTEMS 152 7 Energy and Metabolism 152 Part 5 THE DIVERSITY OF LIFE 482 8 How Cells Make ATP: Energy-Releasing 23 Understanding Diversity: Systematics 482 Pathways 171 24 Viruses and Prokaryotes 500 9 Photosynthesis: Capturing Energy 191 25 Protists 530 26 Kingdom Fungi 555 27 The Plant Kingdom: Seedless Plants 581Part 3 THE CONTINUITY OF LIFE: 28 The Plant Kingdom: Seed Plants 600 GENETICS 211 29 The Animal Kingdom: An Introduction 10 Chromosomes, Mitosis, and Meiosis 211 to Animal Diversity 619 11 The Basic Principles of Heredity 234 30 The Animal Kingdom: The Protostomes 640 12 DNA: The Carrier of Genetic Information 260 31 The Animal Kingdom: The Deuterostomes 667 13 Gene Expression 279 vii
Part 6 STRUCTURE AND LIFE PROCESSES 48 Endocrine Regulation 1028 IN PLANTS 698 49 Reproduction 1051 32 Plant Structure, Growth, and 50 Animal Development 1080 Differentiation 698 51 Animal Behavior 1101 33 Leaf Structure and Function 715 34 Stems and Transport in Vascular Plants 731 35 Roots and Mineral Nutrition 748 Part 8 THE INTERACTIONS OF LIFE: 36 Reproduction in Flowering Plants 767 ECOLOGY 1126 37 Plant Growth and Development 789 52 Introduction to Ecology: Population Ecology 1126 53 Community Ecology 1146Part 7 STRUCTURE AND LIFE PROCESSES 54 Ecosystems and the Biosphere 1166 IN ANIMALS 807 55 Ecology and the Geography of Life 1189 38 Animal Structure and Function: 56 Global Environmental Issues 1212 An Introduction 807 39 Protection, Support, and Movement 827 Appendix A Periodic Table of the Elements A-1 40 Neural Signaling 845 Appendix B Classification of Organisms A-2 41 Neural Regulation 865 Appendix C Understanding Biological Terms A-6 42 Sensory Systems 893 Appendix D Abbreviations A-9 43 Internal Transport 919 Appendix E Answers to Test Your Understanding Questions A-11 44 The Immune System: Internal Defense 944 Glossary G-1 45 Gas Exchange 970 Index I-1 46 Processing Food and Nutrition 989 47 Osmoregulation and Disposal of Metabolic Wastes 1011viii Brief Contents www.thomsonedu.com/biology/solomon
ContentsPart 1 THE ORGANIZATION OF LIFE 1 Scientists interpret the results of experiments and make conclusions 18 A theory is supported by tested hypotheses 20 1 A VIEW OF LIFE 1 Many hypotheses cannot be tested by direct experiment 20 Paradigm shifts allow new discoveries 21 Three Basic Themes 2 Systems biology integrates different levels of information 21 Characteristics of Life 2 Science has ethical dimensions 21 Organisms are composed of cells 2 Organisms grow and develop 3 Organisms regulate their metabolic processes 3 Organisms respond to stimuli 4 Organisms reproduce 5 2 ATOMS AND MOLECULES: Populations evolve and become adapted THE CHEMICAL BASIS OF LIFE 25 to the environment 5 Elements and Atoms 26 Levels of Biological Organization 6 An atom is uniquely identiﬁed by its number of protons 27 Organisms have several levels of organization 6 Protons plus neutrons determine atomic mass 28 Several levels of ecological organization can be identiﬁed 6 Isotopes of an element differ in number of neutrons 28 Information Transfer 6 Electrons move in orbitals corresponding to energy levels 29 DNA transmits information from one generation Chemical Reactions 29 to the next 6 Atoms form compounds and molecules 29 Information is transmitted by chemical and electrical Simplest, molecular, and structural chemical formulas signals 8 give different information 30 Evolution: The Basic Unifying Concept One mole of any substance contains the same number of Biology 9 of units 31 Biologists use a binomial system for naming organisms 9 Chemical equations describe chemical reactions 31 Taxonomic classiﬁcation is hierarchical 9 Chemical Bonds 31 The tree of life includes three domains and six kingdoms 11 In covalent bonds electrons are shared 31 Species adapt in response to changes in their environment 11 Ionic bonds form between cations and anions 33 Natural selection is an important mechanism by which Hydrogen bonds are weak attractions 35 evolution proceeds 11 van der Waals interactions are weak forces 36 Populations evolve as a result of selective pressures from changes in their environment 13 Redox Reactions 36 The Energy for Life 14 Water 36 The Process of Science 15 Hydrogen bonds form between water molecules 37 Science requires systematic thought processes 16 Water molecules interact with hydrophilic substances by hydrogen bonding 37 Scientists make careful observations and ask critical questions 16 Water helps maintain a stable temperature 38 Chance often plays a role in scientiﬁc discovery 16 Acids, Bases, and Salts 39 A hypothesis is a testable statement 17 pH is a convenient measure of acidity 40 Many predictions can be tested by experiment 17 Buffers minimize pH change 40 Researchers must avoid bias 18 An acid and a base react to form a salt 41 ix
3 THE CHEMISTRY OF LIFE: F o c u s O n : Acetabularia and the Control of Cell ORGANIC COMPOUNDS 45 Activities 86 Carbon Atoms and Molecules 46 Organelles in the Cytoplasm 88 Isomers have the same molecular formula but different Ribosomes manufacture proteins 89 structures 47 The endoplasmic reticulum is a network of internal Functional groups change the properties of organic membranes 90 molecules 48 The Golgi complex processes, sorts, and modiﬁes proteins 91 Many biological molecules are polymers 50 Lysosomes are compartments for digestion 92 Vacuoles are large, ﬂuid-ﬁlled sacs with a variety Carbohydrates 50 of functions 93 Monosaccharides are simple sugars 51 Peroxisomes metabolize small organic compounds 94 Disaccharides consist of two monosaccharide units 52 Mitochondria and chloroplasts are energy-converting Polysaccharides can store energy or provide structure 52 organelles 94 Some modiﬁed and complex carbohydrates have special Mitochondria make ATP through cellular respiration 94 roles 54 Chloroplasts convert light energy to chemical energy Lipids 56 through photosynthesis 96 Triacylglycerol is formed from glycerol and three fatty The Cytoskeleton 97 acids 56 Microtubules are hollow cylinders 97 Saturated and unsaturated fatty acids differ in physical Microﬁlaments consist of intertwined strings of actin 99 properties 56 Intermediate ﬁlaments help stabilize cell shape 101 Phospholipids are components of cell membranes 57 Carotenoids and many other pigments are derived Cell Coverings 102 from isoprene units 57 Steroids contain four rings of carbon atoms 57 Some chemical mediators are lipids 58 5 BIOLOGICAL MEMBRANES 106 Proteins 59 Amino acids are the subunits of proteins 62 The Structure of Biological Membranes 107 Proteins have four levels of organization 63 Phospholipids form bilayers in water 107 The amino acid sequence of a protein determines its Current data support a ﬂuid mosaic model of membrane conformation 66 structure 108 Biological membranes are two-dimensional ﬂuids 109 Nucleic Acids 67 Biological membranes fuse and form closed vesicles 110 Some nucleotides are important in energy transfers and other Membrane proteins include integral and peripheral cell functions 68 proteins 110 Identifying Biological Molecules 70 Proteins are oriented asymmetrically across the bilayer 111 Membrane proteins function in transport, in information transfer, and as enzymes 113 Passage of Materials through Cell 4 ORGANIZATION OF THE CELL 73 Membranes 114 The Cell Theory 74 Biological membranes present a barrier to polar molecules 115 Cell Organization and Size 74 Transport proteins transfer molecules across The organization of all cells is basically similar 74 membranes 115 Cell size is limited 74 Cell size and shape are related to function 76 Passive Transport 115 Diffusion occurs down a concentration gradient 116 Methods for Studying Cells 76 Osmosis is diffusion of water across a selectively permeable Light microscopes are used to study stained membrane 116 or living cells 76 Facilitated diffusion occurs down a concentration Electron microscopes provide a high-resolution image that can gradient 118 be greatly magniﬁed 78 Biologists use biochemical techniques to study cell Active Transport 120 components 78 Active transport systems “pump” substances against their concentration gradients 120 Prokaryotic and Eukaryotic Cells 80 Carrier proteins can transport one or two solutes 122 Cell Membranes 81 Cotransport systems indirectly provide energy The Cell Nucleus 84 for active transport 122x Contents www.thomsonedu.com/biology/solomon
Exocytosis and Endocytosis 123 Free energy decreases during an exergonic reaction 155 In exocytosis, vesicles export large molecules 123 Free energy increases during an endergonic reaction 156 In endocytosis, the cell imports materials 123 Diffusion is an exergonic process 156 Cell Junctions 127 Free-energy changes depend on the concentrations of reactants and products 156 Anchoring junctions connect cells of an epithelial sheet 127 Cells drive endergonic reactions by coupling them Tight junctions seal off intercellular spaces between some to exergonic reactions 157 animal cells 127 Gap junctions allow the transfer of small molecules ATP, the Energy Currency of the Cell 157 and ions 128 ATP donates energy through the transfer Plasmodesmata allow certain molecules and ions to move of a phosphate group 158 between plant cells 129 ATP links exergonic and endergonic reactions 158 The cell maintains a very high ratio of ATP to ADP 159 Energy Transfer in Redox Reactions 159 6 CELL COMMUNICATION 134 Most electron carriers transfer hydrogen atoms 159 Cell Signaling: An Overview 135 Enzymes 160 Sending Signals 136 All reactions have a required energy of activation 161 Reception 137 An enzyme lowers a reaction’s activation energy 162 Cells regulate reception 138 An enzyme works by forming an enzyme–substrate Three types of receptors occur on the cell surface 138 complex 162 Some receptors are intracellular 140 Enzymes are speciﬁc 163 Many enzymes require cofactors 163 Signal Transduction 140 Enzymes are most effective at optimal conditions 163 Ion channel–linked receptors open or close channels 140 Enzymes are organized into teams in metabolic pathways 164 G protei