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1. 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
3. 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
5. 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.
7. 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
8. 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
9. 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
10. 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
11. 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 protein–linked receptors initiate signal transduction 141 The cell regulates enzymatic activity 165 Second messengers are intracellular signaling agents 141 Enzymes are inhibited by certain chemical agents 166 Enzyme-linked receptors function directly 144 Some drugs are enzyme inhibitors 167 Many activated intracellular receptors are transcription factors 144 Scaffolding proteins increase efficiency 145 Signals can be transmitted in more than one direction 145 8 HOW CELLS MAKE ATP: ENERGY- Responses to Signals 145 RELEASING PATHWAYS 171 The response to a signal is amplified 146 Redox Reactions 172 Signals must be terminated 147 The Four Stages of Aerobic Respiration 173 Evolution of Cell Communication 147 In glycolysis, glucose yields two pyruvates 174 Pyruvate is converted to acetyl CoA 175 The citric acid cycle oxidizes acetyl CoA 178 The electron transport chain is coupled to ATP synthesis 179 Aerobic respiration of one glucose yields a maximumPart 2 ENERGY TRANSFER THROUGH of 36 to 38 ATPs 183 LIVING SYSTEMS 152 Cells regulate aerobic respiration 185 Energy Yield of Nutrients Other Than Glucose 185 7 ENERGY AND METABOLISM 152 Anaerobic Respiration and Fermentation 186 Biological Work 153 Alcohol fermentation and lactate fermentation are Organisms carry out conversions between potential energy inefficient 187 and kinetic energy 153 The Laws of Thermodynamics 154 The total energy in the universe does not change 154 9 PHOTOSYNTHESIS: The entropy of the universe is increasing 154 CAPTURING ENERGY 191 Energy and Metabolism 155 Light 192 Enthalpy is the total potential energy of a system 155 Chloroplasts 193 Free energy is available to do cell work 155 Chlorophyll is found in the thylakoid membrane 193 Chemical reactions involve changes in free energy 155 Chlorophyll is the main photosynthetic pigment 194 Contents xi
12. Overview of Photosynthesis 196 Prophase I includes synapsis and crossing-over 224 ATP and NADPH are the products of the light-dependent During meiosis I, homologous chromosomes separate 225 reactions: An overview 197 Chromatids separate in meiosis II 225 Carbohydrates are produced during the carbon fixation Mitosis and meiosis lead to contrasting outcomes 225 reactions: An overview 197 The timing of meiosis in the life cycle varies among The Light-Dependent Reactions 198 species 228 Photosystems I and II each consist of a reaction center and multiple antenna complexes 198 Noncyclic electron transport produces ATP and NADPH 198 Cyclic electron transport produces ATP but no NADPH 200 11 THE BASIC PRINCIPLES ATP synthesis occurs by chemiosmosis 200 OF HEREDITY 234 The Carbon Fixation Reactions 202 Mendel’s Principles of Inheritance 235 Most plants use the Calvin cycle to fix carbon 202 Alleles separate before gametes are formed: the principle Photorespiration reduces photosynthetic efficiency 204 of segregation 236 The initial carbon fixation step differs in C4 plants and in Alleles occupy corresponding loci on homologous CAM plants 204 chromosomes 238 Metabolic Diversity 206 A monohybrid cross involves individuals with different alleles of a given locus 238 Photosynthesis in Plants and in the Environment 207 A dihybrid cross involves individuals that have different alleles at two loci 240 Alleles on nonhomologous chromosomes are randomly distributed into gametes: the principle of independent assortment 241 Recognition of Mendel’s work came during the earlyPart 3 THE CONTINUITY OF LIFE: 20th century 242 GENETICS 211 Using Probability to Predict Mendelian Inheritance 243 The rules of probability can be applied to a variety 10 CHROMOSOMES, MITOSIS, of calculations 244 AND MEIOSIS 211 F o c u s O n : Solving Genetics Problems 245 Eukaryotic Chromosomes 212 Inheritance and Chromosomes 246 DNA is organized into informational units called genes 212 Linked genes do not assort independently 246 DNA is packaged in a highly organized way in Calculating the frequency of crossing-over reveals the linear chromosomes 212 order of linked genes on a chromosome 247 Chromosome number and informational content differ among Sex is generally determined by sex chromosomes 248 species 214 Extensions of Mendelian Genetics 252 The Cell Cycle and Mitosis 215 Dominance is not always complete 252 Chromosomes duplicate during interphase 215 Multiple alleles for a locus may exist in a population 253 During prophase, duplicated chromosomes become visible with the microscope 216 A single gene may affect multiple aspects of the phenotype 253 Prometaphase begins when the nuclear envelope breaks down 218 Alleles of different loci may interact to produce a phenotype 254 Duplicated chromosomes line up on the midplane during metaphase 218 Polygenes act additively to produce a phenotype 255 During anaphase, chromosomes move toward the poles 219 Genes interact with the environment to shape phenotype 256 During telophase, two separate nuclei form 220 Cytokinesis forms two separate daughter cells 220 Mitosis produces two cells genetically identical to the parent cell 221 Lacking nuclei, prokaryotes divide by binary fission 221 12 DNA: THE CARRIER OF GENETIC Regulation of the Cell Cycle 221 INFORMATION 260 Sexual Reproduction and Meiosis 223 Evidence of DNA as the Hereditary Material 261 Meiosis produces haploid cells with unique gene DNA is the transforming principle in bacteria 261 combinations 224 DNA is the genetic material in certain viruses 263xii Contents www.thomsonedu.com/biology/solomon
13. The Structure of DNA 263 Some mutations involve larger DNA segments 298 Nucleotides can be covalently linked in any order Mutations have various causes 300 to form long polymers 263 DNA is made of two polynucleotide chains intertwined to form a double helix 264 In double-stranded DNA, hydrogen bonds form between A and T and between G and C 265 14 GENE REGULATION 304 DNA Replication 266 Gene Regulation in Bacteria and Eukaryotes: Meselson and Stahl verified the mechanism An Overview 305 of semiconservative replication 266 Gene Regulation in Bacteria 306 Semiconservative replication explains the perpetuation Operons in bacteria facilitate the coordinated control of mutations 268 of functionally related genes 306 DNA replication requires protein “machinery” 270 Some posttranscriptional regulation occurs in bacteria 309 Enzymes proofread and repair errors in DNA 274 Gene Regulation in Eukaryotic Cells 312 Telomeres cap eukaryotic chromosome ends 275 Eukaryotic transcription is controlled at many sites and by many different regulatory molecules 313 The mRNAs of eukaryotes have many types of posttranscriptional control 31713 GENE EXPRESSION 279 Posttranslational chemical modifications may alter the activity of eukaryotic proteins 318 Discovery of the Gene–Protein Relationship 280 Beadle and Tatum proposed the one-gene, one-enzyme hypothesis 280 Information Flow from DNA to Protein: An Overview 282 15 DNA TECHNOLOGY DNA is transcribed to form RNA 282 AND GENOMICS 322 RNA is translated to form a polypeptide 282 DNA Cloning 323 Biologists cracked the genetic code in the 1960s 283 Restriction enzymes are “molecular scissors” 323 Transcription 285 Recombinant DNA forms when DNA is spliced into The synthesis of mRNA includes initiation, elongation, a vector 324 and termination 286 DNA can be cloned inside cells 324 Messenger RNA contains base sequences that do not directly The polymerase chain reaction is a technique for amplifying code for protein 288 DNA in vitro 328 Translation 288 DNA Analysis 330 An amino acid is attached to tRNA before incorporation Gel electrophoresis is used for separating into a polypeptide 288 macromolecules 330 The components of the translational machinery come together DNA, RNA, and protein blots detect specific at the ribosomes 289 fragments 331 Variations in Gene Expression in Different Restriction fragment length polymorphisms are a measure Organisms 292 of genetic relationships 331 Transcription and translation are coupled in prokaryotes 293 One way to characterize DNA is to determine its sequence of nucleotides 331 Eukaryotic mRNA is modified after transcription and before translation 293 Genomics 333 Both noncoding and coding sequences are transcribed Identifying protein-coding genes is useful for research from eukaryotic genes 294 and for medical applications 334 Several kinds of eukaryotic RNA have a role in gene One way to study gene function is to silence genes one at expression 296 a time 335 The definition of a gene has evolved as biologists have learned DNA microarrays are a powerful tool for studying how genes more about genes 297 interact 335 The usual direction of information flow has exceptions 297 The Human Genome Project stimulated studies on the genome sequences of other species 337 Mutations 298 Base-substitution mutations result from the replacement Applications of DNA Technologies 338 of one base pair by another 298 DNA technology has revolutionized medicine Frameshift mutations result from the insertion or deletion and pharmacology 338 of base pairs 298 DNA fingerprinting has numerous applications 339 Contents xiii
14. Transgenic organisms have incorporated foreign DNA The first cloned mammal was a sheep 371 into their cells 340 Stem cells divide and give rise to differentiated cells 372 DNA Technology Has Raised Safety The Genetic Control of Development 374 Concerns 342 A variety of model organisms provide insights into basic biological processes 374 Many examples of genes that control development have been identified in the fruit fly Drosophila 375 16 THE HUMAN GENOME 346 Caenorhabditis elegans has a relatively rigid developmental pattern 380 Studying Human Genetics 347 The mouse is a model for mammalian development 383 Human chromosomes are studied by karyotyping 347 Arabidopsis is a model for studying plant development, Family pedigrees help identify certain inherited including transcription factors 385 conditions 348 The Human Genome Project sequenced the DNA Cancer and Cell Development 386 on all human chromosomes 349 Comparative genomics has revealed several hundred DNA segments that are identical in both mouse and human genomes 350 Researchers use mouse models to study human genetic diseases 350 Part 4 THE CONTINUITY OF LIFE: Abnormalities in Chromosome Number and EVOLUTION 390 Structure 351 Down syndrome is usually caused by trisomy 21 353 Most sex chromosome aneuploidies are less severe than 18 INTRODUCTION TO DARWINIAN autosomal aneuploidies 354 EVOLUTION 390 Abnormalities in chromosome structure cause certain disorders 354 What Is Evolution? 391 Genetic Diseases Caused by Single-Gene Pre-Darwinian Ideas about Evolution 391 Mutations 356 Darwin and Evolution 392 Many genetic diseases are inherited as autosomal recessive Darwin proposed that evolution occurs by natural traits 356 selection 394 Some genetic diseases are inherited as autosomal dominant The modern synthesis combines Darwin’s theory traits 358 with genetics 394 Some genetic diseases are inherited as X-linked recessive Biologists study the effect of chance on evolution 395 traits 359 Evidence for Evolution 396 Gene Therapy 360 The fossil record provides strong evidence for evolution 396 Gene therapy programs are carefully scrutinized 360 Comparative anatomy of related species demonstrates Genetic Testing and Counseling 361 similarities in their structures 400 Prenatal diagnosis detects chromosome abnormalities The distribution of plants and animals supports and gene defects 361 evolution 402 Genetic screening searches for genotypes or karyotypes 362 Developmental biology helps unravel evolutionary patterns 404 Genetic counselors educate people about genetic diseases 363 Molecular comparisons among organisms provide evidence Human Genetics, Society, and Ethics 363 for evolution 405 Genetic discrimination provokes heated debate 364 Evolutionary hypotheses are tested experimentally 408 Many ethical issues related to human genetics must be addressed 364 19 EVOLUTIONARY CHANGE IN POPULATIONS 412 17 DEVELOPMENTAL GENETICS 368 Genotype, Phenotype, and Allele Cell Differentiation and Nuclear Equivalence 369 Frequencies 413 Most cell differences are due to differential gene The Hardy–Weinberg Principle 413 expression 369 Genetic equilibrium occurs if certain conditions are met 415 A totipotent nucleus contains all the instructions Human MN blood groups are a valuable illustration of the for development 370 Hardy–Weinberg principle 415xiv Contents www.thomsonedu.com/biology/solomon
15. Microevolution 416 Biological evolution began with the first cells 452 Nonrandom mating changes genotype frequencies 416 The first cells were probably heterotrophic 452 Mutation increases variation within a population 417 Aerobes appeared after oxygen increased in the In genetic drift, random events change allele frequencies 417 atmosphere 453 Gene flow generally increases variation within a Eukaryotic cells descended from prokaryotic cells 453 population 418 The History of Life 455 Natural selection changes allele frequencies in a way that Rocks from the Ediacaran period contain fossils of cells and increases adaptation 418 simple animals 455 Genetic Variation in Populations 421 A diversity of organisms evolved during the Paleozoic era 455 Genetic polymorphism exists among alleles and the proteins Dinosaurs and other reptiles dominated the Mesozoic era 459 for which they code 421 The Cenozoic era is the Age of Mammals 461 Balanced polymorphism exists for long periods 422 Neutral variation may give no selective advantage or F o c u s O n : The Origin of Flight in Birds 462 disadvantage 424 Populations in different geographic areas often exhibit genetic adaptations to local environments 425 22 THE EVOLUTION OF PRIMATES 466 Primate Adaptations 467 Primate Classification 46820 SPECIATION AND Suborder Anthropoidea includes monkeys, apes, and MACROEVOLUTION 428 humans 469 What Is a Species? 429 Apes are our closest living relatives 469 Reproductive Isolation 430 Hominid Evolution 471 Prezygotic barriers interfere with fertilization 430 The earliest hominids may have lived 6 mya to 7 mya 473 Postzygotic barriers prevent gene flow when fertilization Australopithecines are the immediate ancestors of genus occurs 431 Homo 474 Biologists are discovering the genetic basis of isolating Homo habilis is the oldest member of genus Homo 474 mechanisms 432 Homo erectus apparently evolved from Homo habilis 475 Speciation 432 Archaic Homo sapiens appeared between 400,000 and 200,000 years ago 475 Long physical isolation and different selective pressures result in allopatric speciation 433 Neandertals appeared approximately 230,000 years ago 475 Two populations diverge in the same physical location by F o c u s O n : The Smallest Humans 476 sympatric speciation 434 Biologists debate the origin of modern Homo sapiens 477 Reproductive isolation breaks down in hybrid zones 437 Cultural Change 478 The Rate of Evolutionary Change 438 Development of agriculture resulted in a more dependable Macroevolution 439 food supply 478 Evolutionary novelties originate through modifications of Cultural evolution has had a profound impact on the pre-existing structures 439 biosphere 479 Adaptive radiation is the diversification of an ancestral species into many species 440 Extinction is an important aspect of evolution 442 Is microevolution related to speciation and macroevolution? 444 Part 5 THE DIVERSITY OF LIFE 48221 THE ORIGIN AND EVOLUTIONARY 23 UNDERSTANDING DIVERSITY: HISTORY OF LIFE 447 SYSTEMATICS 482 Chemical Evolution on Early Earth 448 Classifying Organisms 483 Organic molecules formed on primitive Earth 449 Organisms are named using a binomial system 483 The First Cells 450 Each taxonomic level is more general than the one Molecular reproduction was a crucial step in the origin below it 484 of cells 450 Biologists are moving away from Linnaean categories 484 Contents xv
16. Determining the Major Branches in the Tree The Two Prokaryote Domains 518 of Life 484 Some archaea survive in harsh environments 519 Reconstructing Phylogeny 487 Bacteria are the most familiar prokaryotes 520 Homologous structures are important in determining Impact of Prokaryotes 520 evolutionary relationships 489 Some prokaryotes cause disease 523 Shared derived characters provide clues about phylogeny 489 Prokaryotes are used in many commercial processes 526 Biologists carefully choose taxonomic criteria 490 Molecular homologies help clarify phylogeny 491 Taxa are grouped based on their evolutionary relationships 492 25 PROTISTS 530 Constructing Phylogenetic Trees 493 Introduction to the Protists 531 Outgroup analysis is used in constructing and interpreting Evolution of the Eukaryotes 531 cladograms 493 Mitochondria and chloroplasts probably originated A cladogram is constructed by considering shared derived from endosymbionts 532 characters 494 A consensus is emerging in eukaryote classification 532 In a cladogram each branch point represents a major evolutionary step 496 Representative Protists 534 Systematists use the principle of parsimony to make Excavates are anaerobic zooflagellates 534 decisions 497 Discicristates include euglenoids and trypanosomes 536 Alveolates have flattened vesicles under the plasma membrane 537 Motile cells of heterokonts are biflagellate 540 24 VIRUSES AND PROKARYOTES 500 Red algae, green algae, and land plants are collectively classified Viruses 501 as plants 545 A virus consists of nucleic acid surrounded by a protein Cercozoa are amoeboid cells enclosed in shells 546 coat 501 Amoebozoa have lobose pseudopodia 547 Viruses may have evolved from cells 502 Opisthokonts include choanoflagellates, fungi, The International Committee on Taxonomy of Viruses and animals 551 classifies viruses 502 Bacteriophages are viruses that attack bacteria 503 Viruses reproduce only inside host cells 503 26 KINGDOM FUNGI 555 Lytic reproductive cycles destroy host cells 503 Characteristics of Fungi 556 Temperate viruses integrate their DNA into the host DNA 503 Fungi absorb food from the environment 556 Many viruses infect vertebrates 505 Fungi have cell walls that contain chitin 556 Most fungi have a filamentous body plan 556F o c u s O n : Influenza and Other Emerging and Fungi reproduce by spores 556 Re-emerging Diseases 506 Fungal Diversity 558 Some viruses infect plant cells 510 Fungi are assigned to the opisthokont clade 558 Viroids and Prions 510 Diverse groups of fungi have evolved 560 Viroids are the smallest known pathogens 510 Chytrids have flagellate spores 560 Prions are protein particles 511 Zygomycetes reproduce sexually by forming zygospores 561 Prokaryotes 512 Glomeromycetes are symbionts with plant roots 565 Prokaryotes have several common shapes 512 Ascomycetes reproduce sexually by forming ascospores 566 Prokaryotic cells lack membrane-enclosed organelles 512 Basidiomycetes reproduce sexually by forming A cell wall typically covers the cell surface 512 basidiospores 568 Many types of prokaryotes are motile 513 Ecological Importance of Fungi 570 Prokaryotes have a circular DNA molecule 514 Fungi form symbiotic relationships with some animals 570 Most prokaryotes reproduce by binary fission 514 Mycorrhizae are symbiotic relationships between fungi Bacteria transfer genetic information 514 and plant roots 570 Evolution proceeds rapidly in bacterial populations 516 Lichens are symbiotic relationships between a fungus Some bacteria form endospores 516 and a photoautotroph 572 Many bacteria form biofilms 516 Economic, Biological, and Medical Impact Metabolic diversity has evolved among prokaryotes 517 of Fungi 574 Most prokaryotes require oxygen 517 Fungi provide beverages and food 574xvi Contents www.thomsonedu.com/biology/solomon
17. Fungi are important to modern biology and medicine 574 The Evolution of Seed Plants 614 Some fungi cause animal diseases 576 Our understanding of the evolution of flowering plants has Fungi cause many important plant diseases 576 made great progress in recent years 614 29 THE ANIMAL KINGDOM: 27 THE PLANT KINGDOM: AN INTRODUCTION TO SEEDLESS PLANTS 581 ANIMAL DIVERSITY 619 Adaptations of Plants 582 Animal Characters 620 The plant life cycle alternates haploid and diploid Adaptations to Habitats 620 generations 582 Marine habitats offer many advantages 620 Four major groups of plants evolved 583 Some animals are adapted to freshwater habitats 621 Bryophytes 584 Terrestrial living requires major adaptations 621 Moss gametophytes are differentiated into “leaves” and “stems” 585 Animal Origins 621 Liverwort gametophytes are either thalloid or leafy 587 Molecular systematics helps biologists interpret the fossil record 622 Hornwort gametophytes are inconspicuous thalloid plants 587 Biologists develop hypotheses about the evolution of development 622 Bryophytes are used for experimental studies 588 Details of bryophyte evolution are based on fossils Reconstructing Animal Phylogeny 622 and on structural and molecular evidence 589 Animals exhibit two main types of body symmetry 622 Animal body plans are linked to the level of tissue Seedless Vascular Plants 589 development 624F o c u s O n : Ancient Plants and Coal Formation 590 Biologists group animals according to type of body cavity 624 Club mosses are small plants with rhizomes and short, erect Bilateral animals form two main groups based on differences branches 591 in development 625 Ferns are a diverse group of spore-forming vascular Biologists have identified major animal groups based plants 591 on structure 625 Some ferns and club mosses are heterosporous 595 Molecular data contribute to our understanding Seedless vascular plants are used for experimental studies 595 of animal relationships 626 Seedless vascular plants arose more than 420 mya 595 The Parazoa: Sponges 630 Collar cells characterize sponges 630 The Radiata: Animals with Radial Symmetry and Two Cell Layers 631 28 THE PLANT KINGDOM: Cnidarians have unique stinging cells 631 SEED PLANTS 600 Comb jellies have adhesive glue cells that trap prey 636 An Introduction to Seed Plants 601 Gymnosperms 601 Conifers are woody plants that produce seeds 30 THE ANIMAL KINGDOM: in cones 602 THE PROTOSTOMES 640 Cycads have seed cones and compound leaves 605 Importance of the Coelom 641 Ginkgo biloba is the only living species in its phylum 606 Gnetophytes include three unusual genera 606 The Lophotrochozoa 641 Flatworms are bilateral acoelomates 641 Flowering Plants 607 Phylum Nemertea is characterized by the proboscis 643 Monocots and eudicots are the two largest classes Mollusks have a muscular foot, visceral mass, and mantle 644 of flowering plants 608 Annelids are segmented worms 649 Flowers are involved in sexual reproduction 608 The lophophorate phyla are distinguished by a ciliated ring of The life cycle of flowering plants includes double tentacles 652 fertilization 610 Rotifers have a crown of cilia 653 Seeds and fruits develop after fertilization 611 Flowering plants have many adaptations that account The Ecdysozoa 654 for their success 611 Roundworms are of great ecological importance 654 Studying how flowers evolved provides insights into the Arthropods are characterized by jointed appendages and an evolutionary process 613 exoskeleton of chitin 655 Contents xvii
18. 31 THE ANIMAL KINGDOM: The vascular tissue system consists of two complex THE DEUTEROSTOMES 667 tissues 706 The dermal tissue system consists of two complex tissues 708 What Are Deuterostomes? 668 Plant Meristems 710 Echinoderms 668 Primary growth takes place at apical meristems 711 Members of class Crinoidea are suspension feeders 668 Secondary growth takes place at lateral meristems 712 Many members of class Asteroidea capture prey 669 Class Ophiuroidea is the largest class of echinoderms 670 Members of class Echinoidea have movable spines 670 Members of class Holothuroidea are elongated, sluggish 33 LEAF STRUCTURE AND FUNCTION 715 animals 671 Leaf Form and Structure 716 Chordate Characters 671 Leaf structure consists of an epidermis, photosynthetic ground Invertebrate Chordates 672 tissue, and vascular tissue 716 Tunicates are common marine animals 672 Leaf structure is related to function 720 Lancelets may be closely related to vertebrates 673 F o c u s O n : Air Pollution and Leaves 721 Systematists are making progress in understanding chordate phylogeny 674 Stomatal Opening and Closing 722 Blue light triggers stomatal opening 722 Introducing the Vertebrates 674 Additional factors affect stomatal opening and closing 724 The vertebral column is a key vertebrate character 674 Vertebrate taxonomy is a work in progress 676 Transpiration and Guttation 724 Some plants exude liquid water 725 Jawless Fishes 677 Leaf Abscission 725 Evolution of Jaws and Limbs: Jawed Fishes and Amphibians 678 In many leaves, abscission occurs at an abscission zone near the base of the petiole 726 Members of class Chondrichthyes are cartilaginous fishes 678 The ray-finned fishes gave rise to modern bony fishes 680 Modified Leaves 726 Descendants of the lungfishes moved onto the land 680 Modified leaves of carnivorous plants capture insects 728 Amphibians were the first successful land vertebrates 682 Amniotes 683 Our understanding of amniote phylogeny is changing 684 34 STEMS AND TRANSPORT Reptiles have many terrestrial adaptations 686 IN VASCULAR PLANTS 731 We can assign extant reptiles to four groups 686 External Stem Structure in Woody Twigs 732 Are birds really dinosaurs? 687 Stem Growth and Structure 732 Some dinosaurs had feathers 688 Herbaceous eudicot and monocot stems differ in internal Modern birds are adapted for flight 689 structure 733 Mammals are characterized by hair and mammary glands 690 Woody plants have stems with secondary growth 734 F o c u s O n : Tree-Ring Analysis 738 Transport in the Plant Body 739 Water and minerals are transported in xylem 740 Sugar in solution is translocated in phloem 742 Part 6 STRUCTURE AND LIFE PROCESSES IN PLANTS 698 35 ROOTS AND MINERAL NUTRITION 748 Root Structure and Function 749 32 PLANT STRUCTURE, GROWTH, Roots have root caps and root hairs 749 AND DIFFERENTIATION 698 The arrangement of vascular tissues distinguishes the roots of Plant Structure and Life Span 699 herbaceous eudicots and monocots 749 Plants have different life history strategies 700 Woody plants have roots with secondary growth 753 The Plant Body 700 Some roots are specialized for unusual functions 754 The plant body consists of cells and tissues 700 Root Associations with Fungi and Bacteria 756 The ground tissue system is composed of three simple Mycorrhizae facilitate the uptake of essential minerals by tissues 702 roots 756i xviii Contents www.thomsonedu.com/biology/solomon
19. Rhizobial bacteria fix nitrogen in the roots of leguminous Gibberellins promote stem elongation 794 plants 756 Cytokinins promote cell division 795 The Soil Environment 757 F o c u s O n : Cell and Tissue Culture 796 Soil is composed of inorganic minerals, organic matter, air, Ethylene promotes abscission and fruit ripening 797 and water 758 Abscisic acid promotes seed dormancy 797 The organisms living in the soil form a complex ecosystem 760 Additional signaling molecules affect growth and development, including plant defenses 798 Soil pH affects soil characteristics and plant growth 760 Progress is being made in identifying the elusive flower- Soil provides most of the minerals found in plants 761 promoting signal 799 Soil can be damaged by human mismanagement 762 Light Signals and Plant Development 800 Phytochrome detects day length 801 Competition for sunlight among shade-avoiding plants36 REPRODUCTION IN FLOWERING involves phytochrome 802 PLANTS 767 Phytochrome is involved in other responses to light, including germination 802 The Flowering Plant Life Cycle 768 Phytochrome acts by signal transduction 802 Flowers develop at apical meristems 769 Light influences circadian rhythms 803 Each part of a flower has a specific function 769 Female gametophytes are produced in the ovary, male gametophytes in the anther 769 Pollination 771 Many plants have mechanisms to prevent self-pollination 771 Flowering plants and their animal pollinators have Part 7 STRUCTURE AND LIFE PROCESSES coevolved 771 IN ANIMALS 807 Some flowering plants depend on wind to disperse pollen 774 Fertilization and Seed/Fruit Development 775 38 ANIMAL STRUCTURE AND FUNCTION: A unique double fertilization process occurs in flowering plants 776 AN INTRODUCTION 807 Embryonic development in seeds is orderly and Tissues 808 predictable 776 Epithelial tissues cover the body and line its cavities 808 The mature seed contains an embryonic plant and storage Connective tissues support other body structures 809 materials 777 Muscle tissue is specialized to contract 815 Fruits are mature, ripened ovaries 777 Seed dispersal is highly varied 780 F o c u s O n : Unwelcome Tissues: Cancers 816 Germination and Early Growth 782 Nervous tissue controls muscles and glands 817 Some seeds do not germinate immediately 782 Organs and Organ Systems 817 Eudicots and monocots exhibit characteristic patterns of early The body maintains homeostasis 817 growth 782 Regulating Body Temperature 822 Asexual Reproduction in Flowering Plants 783 Ectotherms absorb heat from their surroundings 823 Apomixis is the production of seeds without the sexual Endotherms derive heat from metabolic processes 823 process 784 Many animals adjust to challenging temperature changes 824 A Comparison of Sexual and Asexual Reproduction 785 Sexual reproduction has some disadvantages 785 39 PROTECTION, SUPPORT, AND MOVEMENT 827 Epithelial Coverings 82837 PLANT GROWTH Invertebrate epithelium may function in secretion AND DEVELOPMENT 789 or gas exchange 828 Tropisms 790 Vertebrate skin functions in protection and temperature Plant Hormones and Development 791 regulation 828 Plant hormones act by signal transduction 791 Skeletal Systems 829 Auxins promote cell elongation 792 In hydrostatic skeletons, body fluids transmit force 829 Contents xix
20. Mollusks and arthropods have nonliving exoskeletons 830 The midbrain is prominent in fishes and amphibians 869 Internal skeletons are capable of growth 830 The forebrain gives rise to the thalamus, hypothalamus, The vertebrate skeleton has two main divisions 831 and cerebrum 869 Muscle Contraction 833 The Human Central Nervous System 871 Invertebrate muscle varies among groups 834 The spinal cord transmits impulses to and from the Insect flight muscles are adapted for rapid contraction 834 brain 871 Vertebrate skeletal muscles act antagonistically The most prominent part of the human brain is the to one another 834 cerebrum 872 A vertebrate muscle may consist of thousands Brain activity cycles in a sleep–wake pattern 873 of muscle fibers 835 The limbic system affects emotional aspects of behavior 876 Contraction occurs when actin and myosin filaments F o c u s O n : The Neurobiology of Traumatic slide past one another 837 Experience 879 ATP powers muscle contraction 840 The strength of muscle contraction varies 840 Information Processing 880 Muscle fibers may be specialized for slow or quick Learning involves the storage of information and its responses 841 retrieval 881 Smooth muscle and cardiac muscle are involuntary 842 Language involves comprehension and expression 883 The Peripheral Nervous System 883 The somatic division helps the body adjust to the external environment 884 40 NEURAL SIGNALING 845 The autonomic division regulates the internal Information Flow through the Nervous environment 884 System 846 Effects of Drugs on the Nervous System 886 Neurons and Glial Cells 847 F o c u s O n : Alcohol: The Most Abused Drug 887 A typical neuron consists of a cell body, dendrites, and an axon 847 Glial cells provide metabolic and structural support 848 Transmitting Information along the Neuron 849 The neuron membrane has a resting potential 849 42 SENSORY SYSTEMS 893 Graded local signals vary in magnitude 851 How Sensory Systems Work 894 An action potential is generated by an influx of Naϩ and an Sensory receptors receive information 894 efflux of Kϩ 851 Sensory receptors transduce energy 894 Neural Signaling across Synapses 855 Sensory input is integrated at many levels 894 Signals across synapses can be electrical or chemical 855 Types of Sensory Receptors 896 Neurons use neurotransmitters to signal other cells 855 Thermoreceptors 897F o c u s O n : Alzheimer’s Disease 857 Electroreceptors and Electromagnetic Neurotransmitters bind with receptors on postsynaptic Receptors 898 cells 859 Nociceptors 898 Activated receptors can send excitatory or inhibitory Mechanoreceptors 898 signals 859 Touch receptors are located in the skin 899 Neural Integration 860 Proprioceptors help coordinate muscle movement 899 Neural Circuits 861 Many invertebrates have gravity receptors called statocysts 900 Hair cells are characterized by stereocilia 901 Lateral line organs supplement vision in fishes 901 41 NEURAL REGULATION 865 The vestibular apparatus maintains equilibrium 901 Invertebrate Nervous Systems 866 Auditory receptors are located in the cochlea 903 Organization of the Vertebrate Nervous Chemoreceptors 906 System 867 Taste receptors detect dissolved food molecules 906 Evolution of the Vertebrate Brain 868 The olfactory epithelium is responsible for the sense The hindbrain develops into the medulla, pons, of smell 908 and cerebellum 868 Many animals communicate with pheromones 909xx Contents www.thomsonedu.com/biology/solomon
21. Photoreceptors 909 Cytokines and complement mediate immune responses 948 Invertebrate photoreceptors include eyespots, simple eyes, Inflammation is a protective response 949 and compound eyes 909 Specific Immune Responses 950 Vertebrate eyes form sharp images 910 Many types of cells are involved in specific immune The retina contains light-sensitive rods and cones 912 responses 950 The major histocompatibility complex is responsible for recognition of self 952 43 INTERNAL TRANSPORT 919 Cell-Mediated Immunity 952 Types of Circulatory Systems 920 Antibody-Mediated Immunity 952 Many invertebrates have an open circulatory system 920 A typical antibody consists of four polypeptide chains 953 Some invertebrates have a closed circulatory system 921 Antibodies are grouped in five classes 954 Vertebrates have a closed circulatory system 922 Antigen–antibody binding activates other defenses 955 Vertebrate Blood 922 The immune system responds to millions of different antigens 956 Plasma is the fluid component of blood 922 Monoclonal antibodies are highly specific 958 Red blood cells transport oxygen 923 White blood cells defend the body against disease Immunological Memory 958 organisms 924 A secondary immune response is more effective than a primary Platelets function in blood clotting 925 response 958 Immunization induces active immunity 959 Vertebrate Blood Vessels 925 Passive immunity is borrowed immunity 960 Evolution of the Vertebrate Cardiovascular System 927 The Immune System and Disease 960 The Human Heart 928 Cancer cells evade the immune system 960 Each heartbeat is initiated by a pacemaker 930 Immunodeficiency disease can be inherited or acquired 961 The nervous system regulates heart rate 932 HIV is the major cause of acquired immunodeficiency in adults 961 Stroke volume depends on venous return 932 Cardiac output varies with the body’s need 932 Harmful Immune Responses 964 Graft rejection is an immune response against transplanted Blood Pressure 933 tissue 964F o c u s O n : Cardiovascular Disease 934 Rh incompatibility can result in hypersensitivity 965 Blood pressure varies in different blood vessels 934 Allergic reactions are directed against ordinary environmental Blood pressure is carefully regulated 935 antigens 965 In an autoimmune disease, the body attacks its own The Pattern of Circulation 937 tissues 966 The pulmonary circulation oxygenates the blood 937 The systemic circulation delivers blood to the tissues 937 The Lymphatic System 938 45 GAS EXCHANGE 970 The lymphatic system consists of lymphatic vessels and lymph Adaptations for Gas Exchange in Air tissue 939 or Water 971 The lymphatic system plays an important role in fluid Types of Respiratory Surfaces 971 homeostasis 940 The body surface may be adapted for gas exchange 971 Tracheal tube systems deliver air directly to the cells 971 Gills are the respiratory surfaces in many aquatic animals 972 44 THE IMMUNE SYSTEM: Terrestrial vertebrates exchange gases through lungs 973 INTERNAL DEFENSE 944 The Mammalian Respiratory System 975 Nonspecific and Specific Immunity: The airway conducts air into the lungs 975 An Overview 945 Gas exchange occurs in the alveoli of the lungs 976 The immune system responds to danger signals 945 Ventilation is accomplished by breathing 976 Invertebrates launch nonspecific immune responses 946 The quantity of respired air can be measured 978 Vertebrates launch nonspecific and specific immune Gas exchange takes place in the alveoli 978 responses 946 Gas exchange takes place in the tissues 979 Nonspecific Immune Responses 946 Respiratory pigments increase capacity for oxygen Phagocytes and natural killer cells destroy pathogens 947 transport 979 Contents xxi
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