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Biology 11e

Eldra P. Solomon
former affiliations
Hillsborough Community College, Tampa
University of South Florida

Charles E. Martin
professor emeritus, Rutgers University

Diana W. Martin
professor emeritus, Rutgers University

Linda R. Berg
former affiliations
University of Maryland, College Park
St. Petersburg College

Australia • Brazil • Mexico • Singapore • United Kingdom • United States

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Biology, Eleventh Edition © 2019, 2015, 2011 Cengage Learning, Inc.
Eldra P. Solomon, Charles E. Martin, WCN: 02-300
Diana W. Martin, Linda R. Berg Unless otherwise noted, all items are © Cengage.
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To our families, friends, and colleagues who gave
freely of their love, support, knowledge, and time as
we prepared this eleventh edition of Biology, and in
appreciation of all who teach and learn.
Especially to
My grandchildren and their generation
Margaret, Damian, and Ava
Alan, Jennifer, and Pat

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About the Authors

Eldra P. Solomon has writ- Charles E. Martin is professor Diana W. Martin is professor Linda R. Berg is an award-
ten several leading college emeritus of cell biology and emeritus and former direc- winning teacher and textbook
textbooks in biology and in neuroscience at Rutgers Uni- tor of general biology in the author. She received a B.S. in
human anatomy and physi- versity. He received his Ph.D. Division of Life Sciences at science education, an M.S. in
ology. Her books have been in genetics from Florida State Rutgers University. Dr. Martin botany, and a Ph.D. in plant
translated into more than ten University and engaged in received an M.S. from Florida physiology from the Univer-
languages. She earned an M.S. postdoctoral research in ge- State University, where she sity of Maryland. Her research
from the University of Florida netics and membrane biology studied the chromosomes of focused on the evolutionary
and an M.A. and Ph.D. from at the University of Texas at related plant species to under- implications of steroid bio-
the University of South Flor- Austin. He has taught general stand their evolutionary rela- synthetic pathways in various
ida. Dr. Solomon taught biol- biology as well as undergradu- tionships. She earned a Ph.D. organisms.
ogy and nursing students for ate and graduate level courses from the University of Texas Dr. Berg taught at the Uni-
more than 20 years. in genetics and molecular cell at Austin, where she studied versity of Maryland at College
In addition to being a bi- biology throughout his career the genetics of the fruit fly, Park for 17 years and at St. Pe-
ologist and science author, Dr. at Rutgers. An award-winning Drosophila melanogaster, and tersburg College in Florida for
Solomon is a biopsychologist teacher for more than 30 years, then conducted postdoc- 8 years. During her career, she
with a special interest in the in 2011 Dr. Martin was named toral research at Princeton taught introductory courses in
neurophysiology of traumatic Professor of the Year by the University. biology, botany, and environ-
experience. Her research has Molecular Biosciences Gradu- Dr. Martin taught general mental science to thousands
focused on the neurological, ate Student Association. biology and other courses at of students. At the University
endocrine, and psychological His research on gene regu- Rutgers for more than 30 years of Maryland, she received nu-
effects of trauma, including lation of membrane protein and has been involved in writ- merous teaching and service
complex post-traumatic stress enzyme systems in yeast and ing textbooks since 1988. She awards. Dr. Berg is also the
disorder and development of other fungi illustrates the in- is immensely grateful that her recipient of many national and
maladaptive coping strategies. terdisciplinary nature of the decision to study biology in regional awards, including
Dr. Solomon has pre- life sciences. He is most proud college has led to a career that the National Science Teach-
sented her research at numer- of the many generations of allows her many ways to share ers Association Award for In-
ous national and international undergraduate, graduate, and her excitement about all as- novations in College Science
conferences, and her work postdoctoral students who pects of biology. Teaching, the Nation’s Capital
has been published in lead- contributed to this research Area Disabled Student Ser-
ing professional journals. She and have gone on to produc- vices Award, and the Wash-
has been profiled more than tive careers. He continues to be ington Academy of Sciences
30 times in leading publica- committed to teaching and is Award in University Science
tions, including Who’s Who in grateful for the opportunities Teaching.
America, Who’s Who in Science to pursue a teaching and re- During her career as a
and Engineering, Who’s Who search career in what contin- professional science writer,
in Medicine and Healthcare, ues to be the most exciting era Dr. Berg has authored or coau-
Who’s Who in American Edu- of the biological sciences. thored several leading college
cation, Who’s Who of American science textbooks. Her writing
Women, and Who’s Who in the reflects her teaching style and
World. love of science.

iv

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Brief Contents

Preface xxiii 28 Seed Plants 584


To the Student xxx 29 The Fungi 603
part one: The Organization of Life 30 An Introduction to Animal Diversity 628
31 Sponges, Cnidarians, Ctenophores,
1 A View of Life 1 and Protostomes 641
2 Atoms and Molecules: 32 The Deuterostomes 676
The Chemical Basis of Life 26
part six: Structure and Life Processes in Plants
3 The Chemistry of Life: Organic Compounds 46
4 Organization of the Cell 73 33 Plant Structure, Growth, and Development 710

5 Biological Membranes 106 34 Leaf Structure and Function 729


6 Cell Communication 131 35 Stem Structure and Transport 745
36 Roots and Mineral Nutrition 762
part two: Energy Transfer through Living Systems
37 Reproduction in Flowering Plants 782
7 Energy and Metabolism 150
38 Plant Developmental Responses to ­External
8 How Cells Make ATP: and ­Internal Signals 803
Energy-Releasing Pathways 167
9 Photosynthesis: Capturing Light Energy 187 part seven: Structure and Life Processes in Animals

part three: The Continuity of Life: Genetics 39 Animal Structure and Function: An Introduction 821
40 Protection, Support, and Movement 842
10 Chromosomes, Mitosis, and Meiosis 206
41 Neural Signaling 860
11 The Basic Principles of Heredity 228
42 Neural Regulation 882
12 DNA: The Carrier of Genetic Information 253
43 Sensory Systems 911
13 Gene Expression 272
44 Internal Transport 936
14 Gene Regulation 297
45 The Immune System: Internal Defense 962
15 DNA Technology and Genomics 315
46 Gas Exchange 991
16 Human Genetics and the Human
Genome 340 47 Processing Food and Nutrition 1010
17 Developmental Genetics 362 48 Osmoregulation and Disposal of ­Metabolic
Wastes 1032
part four: The Continuity of Life: Evolution
49 Endocrine Regulation 1050
18 Introduction to Darwinian Evolution 385 50 Reproduction 1074
19 Evolutionary Change in Populations 406 51 Animal Development 1104
20 Speciation and Macroevolution 421 52 Animal Behavior 1124
21 The Origin and Evolutionary
History of Life 442 part eight: The Interactions of Life: Ecology
22 The Evolution of Primates 461 53 Introduction to Ecology: Population Ecology 1151
part five: The Diversity of Life 54 Community Ecology 1171
55 Ecosystems and the Biosphere 1194
23 Understanding Diversity: Systematics 478
56 Ecology and the Geography of Life 1216
24 Viruses and Subviral Agents 499
57 Biological Diversity and Conservation Biology 1241
25 Bacteria and Archaea 517
26 Protists 539 Glossary G-1
27 Seedless Plants 563 Index I-1

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Contents

part one: THE ORGANIZATION OF LIFE

1 A View of Life 1 2.2 Chemical Reactions 31


1.1 Major Themes of Biology 2 Atoms form compounds and molecules 31
1.2 Characteristics of Life 2 Simplest, molecular, and structural chemical formulas give
Organisms are composed of cells 3 different information 31
Organisms grow and develop 3 One mole of any substance contains the same number
Organisms regulate their metabolic processes 3 of units 31
Organisms respond to stimuli 4 Chemical equations describe chemical reactions 32
Organisms reproduce 5 2.3 Chemical Bonds 32
Populations evolve and become adapted to the environment 5 In covalent bonds electrons are shared 32
1.3 Levels of Biological Organization 6 The function of a molecule is related to its shape 34
Organisms have several levels of organization 6 Covalent bonds can be nonpolar or polar 34
Several levels of ecological organization can be identified 6 Ionic bonds form between cations and anions 34
1.4 Information Transfer 6 Hydrogen bonds are weak attractions 36
DNA transmits information from one generation to the next 8 van der Waals interactions are weak forces 37
Information is transmitted by chemical and electrical signals 8 2.4 Redox Reactions 37
Organisms also communicate information to one another 8 2.5 Water 38
1.5 The Energy of Life 9 Hydrogen bonds form between water molecules 38
1.6 Evolution: The Basic Unifying Concept of Biology 10 Water molecules interact with hydrophilic substances by
Biologists use a binomial system for naming organisms 11 hydrogen bonding 38
Taxonomic classification is hierarchical 11 Water helps maintain a stable temperature 39
Systematists classify organisms in three domains 11 2.6 Acids, Bases, and Salts 41
Species adapt in response to changes in their environment 14 pH is a convenient measure of acidity 41
Natural selection is an important mechanism by which Buffers minimize pH change 42
­evolution proceeds 14 An acid and a base react to form a salt 43
Populations evolve as a result of selective pressures from
changes in their environment 15
1.7 The Process of Science 15 3 The Chemistry of Life:
Science requires systematic thought processes 16 Organic Compounds 46
Scientists make careful observations and ask critical 3.1 Carbon Atoms and Organic Molecules 47
questions 16 Isomers have the same molecular formula but different
Chance often plays a role in scientific discovery 17 structures 48
A hypothesis is a testable statement 17 Functional groups change the properties of organic
Researchers must avoid bias 18 molecules 49
Scientists interpret the results of experiments and draw Many biological molecules are polymers 50
conclusions 18 3.2 Carbohydrates 51
A scientific theory is supported by tested hypotheses 20 Monosaccharides are simple sugars 51
Many hypotheses cannot be tested by direct experiment 21 Disaccharides consist of two monosaccharide units 52
Paradigm shifts accommodate new discoveries 21 Polysaccharides can store energy or provide structure 53
Systems biology integrates different levels of information 21 Some modified and complex carbohydrates have special
Science has ethical dimensions 21 roles 55
Science, technology, and society interact 22 3.3 Lipids 56
Triacylglycerol is formed from glycerol and three fatty
acids 56
2 Atoms and Molecules: Saturated and unsaturated fatty acids differ in physical
The Chemical Basis of Life 26 properties 57
2.1 Elements and Atoms 27 Phospholipids are components of cell membranes 57
An atom is uniquely identified by its number of protons 28 Carotenoids and many other pigments are derived from
Protons plus neutrons determine atomic mass 29 ­isoprene units 57
Isotopes of an element differ in number of neutrons 29 Steroids contain four rings of carbon atoms 58
Electrons move in orbitals corresponding to energy levels 30 Some chemical mediators are lipids 59

vi

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3.4 Proteins 59 Microfilaments consist of intertwined strings of actin 100
Amino acids are the subunits of proteins 60 Intermediate filaments help stabilize cell shape 102
Peptide bonds join amino acids 61 4.7 Cell Coverings 103
Proteins have four levels of organization 61
The amino acid sequence of a protein determines its
conformation 65 5 Biological Membranes 106
3.5 Nucleic Acids 68 5.1 The Structure of Biological Membranes 107
Some nucleotides are important in energy transfers and other Phospholipids form bilayers in water 107
cell functions 68 The fluid mosaic model explains membrane structure 108
3.6 Identifying Biological Molecules 69 Biological membranes are two-dimensional fluids 109
Biological membranes fuse and form closed vesicles 110
Membrane proteins include integral and peripheral
4 Organization of the Cell 73 proteins 111
4.1 The Cell: Basic Unit of Life 74 Proteins are oriented asymmetrically across the bilayer 111
The cell theory is a unifying concept in biology 74 5.2 Overview of Membrane Protein Functions 113
The organization and basic functions of all cells are 5.3 Cell Membrane Structure and Permeability 114
similar 74 Biological membranes present a barrier to polar
Cell size is limited 74 molecules 114
Cell size and shape are adapted to function 76 Transport proteins transfer molecules across membranes 115
4.2 Methods for Studying Cells 76 5.4 Passive Transport 115
Light microscopes are used to study stained or living cells 76 Diffusion occurs down a concentration gradient 115
Electron microscopes provide a high-resolution image that can Osmosis is diffusion of water across a selectively permeable
be greatly magnified 78 membrane 116
Biologists use biochemical and genetic methods to connect Facilitated diffusion occurs down a concentration gradient 118
cell structures with their functions 79 5.5 Active Transport 120
4.3 Prokaryotic and Eukaryotic Cells 82 Active transport systems “pump” substances against their
Organelles of prokaryotic cells are not surrounded by concentration gradients 120
membranes 82 Carrier proteins can transport one or two solutes 122
Membranes divide the eukaryotic cell into compartments 83 Cotransport systems indirectly provide energy for active
The unique properties of biological membranes allow transport 122
­eukaryotic cells to carry on many diverse functions 83 5.6 Exocytosis and Endocytosis 123
4.4 The Cell Nucleus 84 In exocytosis, vesicles export large molecules 123
Ribosomes manufacture proteins in the cytoplasm 87 In endocytosis, the cell imports materials 123
4.5 Membranous Organelles in the Cytoplasm 88 5.7 Cell Junctions 125
The endoplasmic reticulum is a multifunctional network of Anchoring junctions connect cells of an epithelial sheet 125
membranes 88 Tight junctions seal off intercellular spaces between some
The ER is the primary site of membrane assembly for animal cells 127
­components of the endomembrane system 91 Gap junctions allow the transfer of small molecules and ions 128
The Golgi complex processes, sorts, and routes proteins Plasmodesmata allow certain molecules and ions to move
from the ER to different parts of the endomembrane between plant cells 128
system 91
Lysosomes are compartments for digestion 93
Vacuoles are large, fluid-filled sacs with a variety of 6 Cell Communication 131
functions 94 6.1 Cell Communication: an Overview 132
Peroxisomes metabolize small organic compounds 94 6.2 Sending Signals 133
Mitochondria and chloroplasts are energy-converting 6.3 Reception 134
organelles 95 Cells regulate reception 135
Mitochondria make ATP through aerobic respiration 95 Three types of receptors occur on the cell surface 135
Chloroplasts convert light energy to chemical energy through Some receptors are located inside the cell 137
photosynthesis 97 6.4 Signal Transduction 138
4.6 The Cytoskeleton 98 Signaling molecules can act as molecular switches 138
Microtubules are hollow cylinders 98 Ion channel–linked receptors open or close channels 139
Centrosomes and centrioles function in cell division 99 G protein–linked receptors initiate signal transduction 139
Cilia and flagella are composed of microtubules 99 Second messengers are intracellular signaling agents 139

Contents / vii

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Many activated intracellular receptors are transcription Ras pathways involve tyrosine kinase receptors and
factors 142 G proteins 144
Scaffold proteins increase efficiency 143 The response to a signal is amplified 144
Signals can be transmitted in more than one direction 143 Signals must be terminated 145
6.5 Responses to Signals 143 6.6 Evolution of Cell Communication 146

part two: ENERGY Transfer THROUGH Living Systems

7 Energy and Metabolism 150 Pyruvate is converted to acetyl CoA 171


7.1 Biological Work 151 The citric acid cycle oxidizes acetyl groups derived from
Organisms carry out conversions between potential energy acetyl CoA 171
and kinetic energy 151 The electron transport chain is coupled to ATP
7.2 The Laws of Thermodynamics 151 synthesis 176
The total energy in the universe does not change 151 Aerobic respiration of one glucose yields a maximum of 36 to
The entropy of the universe is increasing 152 38 ATPs 180
7.3 Energy and Metabolism 152 Cells regulate aerobic respiration 181
Enthalpy is the total potential energy of a system 153 8.3 Energy Yield of Nutrients other than
Free energy is available to do cell work 153 Glucose 182
Chemical reactions involve changes in free energy 153 8.4 Anaerobic Respiration and Fermentation 182
Free energy decreases during an exergonic reaction 153 Alcohol fermentation and lactate fermentation are
Free energy increases during an endergonic reaction 154 inefficient 183
Diffusion is an exergonic process 154
Free-energy changes depend on the concentrations of 9 Photosynthesis: Capturing Light
­reactants and products 154 Energy 187
Cells drive endergonic reactions by coupling them to 9.1 Light and Photosynthesis 188
­exergonic reactions 154 9.2 Chloroplasts 189
7.4 Atp, the Energy Currency of the Cell 155 Chlorophyll is found in the thylakoid membrane 190
ATP donates energy through the transfer of a phosphate Chlorophyll is the main photosynthetic pigment 191
group 155 9.3 Overview of Photosynthesis 192
ATP links exergonic and endergonic reactions 156 ATP and NADPH are the products of the light-dependent
The cell maintains a very high ratio of ATP to ADP 156 reactions: An overview 192
7.5 Energy Transfer in Redox Reactions 157 Carbohydrates are produced during the carbon fixation
Most electron carriers transfer hydrogen atoms 157 ­reactions: An overview 193
7.6 Enzymes 158 9.4 The Light-Dependent Reactions 193
All reactions have a required energy of activation 158 Photosystems I and II each consist of a reaction center and
An enzyme lowers a reaction’s activation energy 159 multiple antenna complexes 194
An enzyme works by forming an enzyme–substrate Noncyclic electron transport produces ATP and
complex 159 NADPH 194
Enzymes are specific 160 Cyclic electron transport produces ATP but no
Many enzymes require cofactors 160 NADPH 196
Enzymes are most effective at optimal conditions 161 ATP synthesis occurs by chemiosmosis 196
Enzymes are organized into teams in metabolic pathways 162 9.5 The Carbon Fixation Reactions 198
The cell regulates enzymatic activity 162 Most plants use the Calvin cycle to fix
Enzymes are inhibited by certain chemical agents 163 carbon 198
Some drugs are enzyme inhibitors 164 Photorespiration reduces photosynthetic
efficiency 200
The initial carbon fixation step differs in C4 plants and in
8 How Cells Make ATP: CAM plants 200
Energy-Releasing Pathways 167 CAM plants fix CO2 at night 202
8.1 Redox Reactions 168 9.6 Metabolic Diversity 202
8.2 The Four Stages of Aerobic Respiration 168 9.7 Photosynthesis in Plants and in the
In glycolysis, glucose yields two pyruvates 170 Environment 203

viii / Contents

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part three: The Continuity of Life: Genetics

10 Chromosomes, Mitosis, and The rules of probability can be applied to a variety of


Meiosis 206 calculations 238
10.1 Eukaryotic Chromosomes 207 11.3 Inheritance and Chromosomes 240
DNA is organized into informational units called genes 207 Linked genes do not assort independently 240
DNA is packaged in a highly organized way in Calculating the frequency of crossing-over reveals the linear
chromosomes 207 order of linked genes on a chromosome 240
Chromosome number and informational content differ Sex is generally determined by sex chromosomes 241
among species 208 11.4 Extensions of Mendelian Genetics 246
10.2 The Cell Cycle and Mitosis 210 Dominance is not always complete 246
Chromosomes duplicate during interphase 210 Multiple alleles for a locus may exist in a population 248
During prophase, duplicated chromosomes become visible A single gene may affect multiple aspects of the phenotype 248
with the microscope 211 Alleles of different loci may interact to produce a
Prometaphase begins when the nuclear envelope breaks phenotype 248
down 211 In polygenic inheritance, the offspring exhibit a continuous
Duplicated chromosomes line up on the midplane during variation in phenotypes 249
metaphase 212 Genes interact with the environment to shape phenotype 250
During anaphase, chromosomes move toward the
poles 213
During telophase, two separate nuclei form 215 12 DNA: The Carrier of Genetic
Cytokinesis forms two separate daughter cells 215 Information 253
Mitosis produces two cells genetically identical to the 12.1 Evidence of Dna as the Hereditary Material 254
parent cell 215 DNA is the transforming factor in bacteria 254
Lacking nuclei, prokaryotes divide by binary fission 216 DNA is the genetic material in certain viruses 254
10.3 Regulation of the Cell Cycle 217 12.2 The Structure of Dna 257
10.4 Sexual Reproduction and Meiosis 219 Nucleotides can be covalently linked in any order to form long
Meiosis produces haploid cells with unique gene polymers 257
combinations 220 DNA is made of two polynucleotide chains intertwined to
Prophase I includes synapsis and crossing-over 221 form a double helix 258
During meiosis I, homologous chromosomes separate 221 In double-stranded DNA, hydrogen bonds form between
Chromatids separate in meiosis II 222 A and T and between G and C 261
Mitosis and meiosis lead to contrasting outcomes 223 12.3 DNA Replication 261
10.5 Sexual Life Cycles 224 Meselson and Stahl verified the mechanism of
­semiconservative replication 262
Semiconservative replication explains the perpetuation of
11 The Basic Principles of Heredity 228 mutations 262
11.1 Mendel’s Principles of Inheritance 229 DNA replication requires protein “machinery” 263
Alleles separate before gametes are formed: the principle Enzymes proofread and repair errors in DNA 268
of segregation 231 Telomeres cap eukaryotic chromosome ends 268
Alleles occupy corresponding loci on homologous
chromosomes 232
A monohybrid cross involves individuals with different alleles 13 Gene Expression 272
of a given locus 233 13.1 Discovery of the Gene–Protein Relationship 273
A dihybrid cross involves individuals that have different alleles Beadle and Tatum proposed the one-gene, one-enzyme
at two loci 235 hypothesis 273
Alleles on nonhomologous chromosomes are randomly 13.2 Information Flow from Dna to Protein:
distributed into gametes: the principle of independent An Overview 275
assortment 236 DNA is transcribed to form RNA 276
Recognition of Mendel’s work came during the early RNA is translated to form a polypeptide 276
20th century 236 Biologists cracked the genetic code in the 1960s 277
11.2 Using Probability to Predict Mendelian The genetic code is virtually universal 278
Inheritance 238 The genetic code is redundant 278

Contents / ix

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13.3 Transcription 279 15 DNA Technology and Genomics 315
The synthesis of mRNA includes initiation, elongation, 15.1 Dna Cloning 316
and termination 280 Restriction enzymes are “molecular scissors” used to
Messenger RNA contains base sequences that do not directly ­construct recombinant DNA molecules 316
code for protein 281 Recombinant DNA is formed when DNA is spliced into a
Eukaryotic mRNA is modified after transcription and before vector 317
translation 282 Scientists use restriction enzymes and gel electrophoresis to
Biologists debate the evolution of eukaryotic gene examine cloned DNA fragments 318
structure 282 The polymerase chain reaction amplifies DNA
13.4 Translation 284 in vitro 318
An amino acid is attached to tRNA before incorporation into a cDNA clones do not contain introns 319
polypeptide 284 15.2 CRISPR-Based Technologies 321
The components of the translational machinery come CRISPR-based technologies can be used to edit genes in
­together at the ribosomes 285 growing cells 321
Translation begins with the formation of an initiation CRISPR-based tools exploit host DNA repair systems to
complex 285 ­perform many types of recombinant DNA functions 322
During elongation, amino acids are added to the growing Engineered CRISPR systems are used for specialized research
polypeptide chain 286 applications 322
One of three stop codons signals the termination of 15.3 Tools for Studying Dna 323
translation 288 DNA, RNA, and protein blots detect differences in related
Transcription and translation are coupled in bacteria 288 molecules separated by gel electrophoresis 324
13.5 Mutations 290 Automated DNA sequencing methods have been
Base-pair substitution mutations result from the replacement developed 324
of one base pair by another 290 Gene databases are powerful research tools 325
Frameshift mutations result from the insertion or deletion of Reverse transcription of mRNA to cDNA is used to measure
base pairs 290 gene expression in numerous ways 326
Some mutations involve mobile genetic elements 290 15.4 Genomics 328
Mutations have various causes 292 Collaborative genome-wide association studies have radically
13.6 Variations in Gene Expression 292 changed our view of the human genome 328
Many eukaryotic genes produce “non-coding” RNAs with Comparative genomic databases are tools for uncovering
catalytic, regulatory, or other cellular functions 292 gene functions 328
The definition of a gene has evolved 293 RNA interference is used to study gene functions 329
The usual direction of information flow has exceptions 293 15.5 Applications of Dna Technologies 330
DNA technology has revolutionized medicine 330
DNA fingerprinting has numerous applications 331
14 Gene Regulation 297
Transgenic organisms have many research and technological
14.1 Gene Regulation in Bacteria and Eukaryotes:
applications 331
An Overview 298
15.6 CRISPR-Based Gene Drives 334
14.2 Gene Regulation in Bacteria 299
15.7 Dna Technology and Safety Concerns 336
Operons in bacteria facilitate the coordinated control of
­functionally related genes 299
Some posttranscriptional regulation occurs in 16 Human Genetics
bacteria 303 and the Human Genome 340
14.3 Gene Regulation in Eukaryotic Cells 304 16.1 Studying Human Genetics 341
Eukaryotic transcription is controlled at many sites and by Human chromosomes are studied by karyotyping 341
many regulatory molecules 305 Family pedigrees help identify certain inherited
Chromosome organization affects the expression of some conditions 342
genes 307 Human gene databases allow geneticists to map the locations
Long non-coding RNAs (lncRNAs) regulate transcription over of genes on chromosomes 342
long distances within the genome 309 16.2 Abnormalities in Chromosome Number
The mRNAs of eukaryotes are subject to many types of and Structure 344
­posttranscriptional control 309 Down syndrome is usually caused by trisomy 21 345
Posttranslational chemical modifications may alter the activity Most sex chromosome aneuploidies are less severe than
of eukaryotic proteins 312 autosomal aneuploidies 347

x / Contents

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Abnormalities in chromosome structure cause certain 17 Developmental Genetics 362
disorders 348 17.1 Cell Differentiation and Nuclear
Genomic imprinting may determine whether inheritance is Equivalence 363
from the male or female parent 349 Most cell differences are due to differential gene
16.3 Genetic Diseases Caused by Single-Gene expression 363
Mutations 351 A totipotent nucleus contains all the instructions for
Many genetic diseases are inherited as autosomal development 364
recessive traits 351 The first cloned mammal was a sheep 366
Some genetic diseases are inherited as autosomal Stem cells divide and give rise to differentiated
dominant traits 353 cells 367
Some genetic diseases are inherited as X-linked 17.2 The Genetic Control of Development 369
recessive traits 353 A variety of model organisms provide insights into basic
16.4 Gene Therapy 354 ­biological processes 369
Performing clinical trials on humans always has inherent Many genes that control development have been identified in
risks 354 the fruit fly 369
16.5 Genetic Testing and Counseling 355 Caenorhabditis elegans has a relatively rigid developmental
Prenatal diagnosis detects chromosome abnormalities and pattern 374
gene defects 355 The mouse is a model for mammalian development 377
Preimplanation genetic diagnosis is used to screen embryos Arabidopsis is a model for studying plant development,
produced by in vitro fertilization 356 ­including transcription factors 379
Genetic screening searches for genotypes or karyotypes 356 17.3 Cancer and Cell Development 380
Genetic counselors educate people about genetic Oncogenes are usually altered components of cell
diseases 357 ­signaling pathways that control growth and
16.6 Human Genetics, Society, and Ethics 357 differentiation 381
Genetic discrimination provokes heated debate 358 In many familial cancers, tumor suppressor genes must be
Many ethical issues related to human genetics must be inactivated before cells progress to cancer 382
addressed 358 Cancer cells evolve by accumulating new mutations 382

part four: The CONTINUITY of Life: Evolution

18 Introduction to Darwinian 19 Evolutionary Change in


Evolution 385 Populations 406
18.1 What Is Evolution? 386 19.1 Genotype, Phenotype, and Allele
18.2 Pre-Darwinian Ideas about Evolution 386 Frequencies 407
18.3 Darwin and Evolution 387 19.2 The Hardy–Weinberg Principle 407
Darwin proposed that evolution occurs by natural Genetic equilibrium occurs if certain conditions are met 409
selection 389 Human MN blood groups are a valuable illustration of the
The modern synthesis combines Darwin’s scientific theory of Hardy–Weinberg principle 409
evolution with genetics 390 19.3 Microevolution 410
Biologists study the effect of chance on evolution 390 Nonrandom mating changes genotype frequencies 410
18.4 Evidence for Evolution 391 Mutation increases variation within a population 410
The fossil record provides strong evidence for In genetic drift, random events change allele frequencies 411
evolution 391 Gene flow generally increases variation within a population 412
The distribution of plants and animals supports Natural selection changes allele frequencies in a way that
evolution 395 increases adaptation 412
Comparative anatomy of related species demonstrates 19.4 Genetic Variation in Populations 415
­similarities in their structures 396 Genetic polymorphism can be studied in several ways 415
Molecular comparisons among organisms provide evidence Balanced polymorphism exists for long periods 416
for evolution 399 Neutral variation may give no selective advantage or
Developmental biology helps unravel evolutionary disadvantage 418
patterns 400 Populations in different geographic areas often exhibit genetic
Evolutionary hypotheses are tested experimentally 401 adaptations to local environments 418

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20 Speciation and Macroevolution 421 Biological evolution began with the first cells 447
20.1 What is a Species? 422 Photosynthesis was a further step in the evolution of
The biological species concept is based on reproductive cells 448
isolation 422 Aerobes appeared after oxygen increased in the
The phylogenetic species concept defines species based on atmosphere 449
such evidence as molecular sequencing 422 Eukaryotic cells descended from prokaryotic cells 450
20.2 Reproductive Isolation 423 21.3 The History of Life 451
Prezygotic barriers interfere with fertilization 423 Rocks from the Ediacaran period contain fossils of cells and
Postzygotic barriers prevent gene flow when fertilization simple animals 451
occurs 425 A diversity of organisms evolved during the Paleozoic era 451
Biologists are discovering genes responsible for reproductive Dinosaurs and other reptiles dominated the Mesozoic era 454
isolating mechanisms 425 The Cenozoic era is the age of mammals 457
20.3 Speciation 425
Long physical isolation and different selective pressures result
in allopatric speciation 427 22 The Evolution of Primates 461
The evolutionary importance of hybridization is being 22.1 Primate Adaptations 462
re-evaluated 432 22.2 Primate Classification 462
20.4 The Rate of Evolutionary Change 434 Suborder Anthropoidea includes monkeys, apes, and
20.5 Macroevolution 435 humans 463
Evolutionary novelties originate through modifications of Apes are our closest living relatives 465
pre-existing structures 435 22.3 Hominin Evolution 467
Adaptive radiation is the diversification of an ancestral The earliest hominins may have lived 6 mya to 7 mya 468
­species into many species 436 Ardipithecus, Australopithecus, and Paranthropus are
Extinction is an important aspect of evolution 438 ­australopithecines, or “southern man apes” 468
Is microevolution related to speciation and Homo habilis is considered the oldest member of genus
macroevolution? 439 Homo 470
Homo ergaster may have arisen from H. habilis 471
Homo erectus probably evolved from H. ergaster 471
21 The Origin and Evolutionary Archaic humans date from about 1.2 mya to 200,000 years
History of Life 442 ago 471
21.1 Chemical Evolution on Early Earth 443 Neandertals appeared approximately 250,000 years ago 472
Organic molecules formed on primitive Earth 443 Scientists have reached a near consensus on the origin of
21.2 The First Cells 445 modern H. sapiens 473
The origin of a simple metabolism within a membrane 22.4 Cultural Change 474
­boundary may have occurred early in the evolution Development of agriculture resulted in a more dependable
of cells 445 food supply 475
Molecular reproduction was a crucial step in the origin Human culture has had a profound effect on the
of cells 445 biosphere 475

part five: The Diversity of Life

23 Understanding Diversity: Some biologists are moving away from Linnaean


Systematics 478 categories 482
23.1 Classifying Organisms 479 Phylogenetic trees show hypothesized evolutionary
Organisms are named using a binomial system 479 relationships 483
Each taxonomic level is more general than the one Systematists continue to consider other hypotheses 484
below it 480 23.3 Reconstructing Evolutionary History 485
23.2 Determining the Major Branches in the Tree Homologous structures are important in determining
of Life 480 ­evolutionary relationships 485
Systematics is an evolving science 480 Shared derived characters provide clues about phylogeny 486
The three domains form the three main branches of the tree Systematists base taxonomic decisions on recent shared
of life 482 ancestry 487

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Molecular homologies help clarify phylogeny 487 25.4 the phylogeny of the two prokaryote
Taxa are grouped based on their evolutionary domains 525
relationships 488 Key characters distinguish the three domains 526
23.4 Constructing Phylogenetic Trees 490 Taxonomy of archaea and bacteria continuously changes 526
Outgroup analysis is used in constructing and interpreting Most archaea live in marine and soil habitats, and many
cladograms 490 thrive in harsh environments 527
A cladogram is constructed by considering shared derived Bacteria are the most familiar prokaryotes 528
characters 491 25.5 Impact on Ecology, Technology, and
Each branch point represents a major evolutionary step 491 Commerce 528
Systematists use the principles of parsimony and maximum Prokaryotes form intimate relationships with other
likelihood to make decisions 494 organisms 529
23.5 Applying Phylogenetic Information 495 Prokaryotes play key ecological roles 529
Prokaryotes are important in many commercial processes and
in technology 532
24 Viruses and Subviral Agents 499 25.6 bacteria and disease 533
24.1 The Status and Structure of Viruses 500 Many scientists have contributed to our understanding of
Viruses are very small 500 infectious disease 533
A virus consists of nucleic acid surrounded by a Many adaptations contribute to pathogen success 533
protein coat 500 Antibiotic resistance is a major public health problem 535
The capsid is a protective protein coat 501
Some viruses are surrounded by an envelope 502
24.2 Classification of Viruses 502 26 Protists 539
24.3 Viral Replication 503 26.1 diversity in the protists 540
Bacteriophages infect bacteria 503 26.2 how did eukaryotes evolve? 541
Viruses replicate inside host cells 503 Mitochondria and chloroplasts probably originated
24.4 Viral Diseases 504 from endosymbionts 541
Viruses cause serious plant diseases 505 A consensus in eukaryote classification is beginning
Viruses cause serious diseases in animals 505 to emerge 541
24.5 Evolution of Viruses 511 26.3 excavates 544
24.6 Subviral Agents 512 Diplomonads are small, mostly parasitic flagellates 544
Satellites depend on helper viruses 512 Parabasilids are anaerobic endosymbionts that live in
Viroids are short, single strands of naked RNA 513 animals 544
Prions are protein particles 513 Euglenoids and trypanosomes include both free-living species
Defective interfering particles are virus mutants 514 and parasites 545
26.4 chromalveolates 546
Most dinoflagellates are a part of marine plankton 546
25 Bacteria and Archaea 517 Apicomplexans are spore-forming parasites of animals 547
25.1 The Structure of Bacteria and Archaea 518 Ciliates use cilia for locomotion 548
Prokaryotes have several common shapes 518 Water molds produce biflagellate reproductive cells 549
Prokaryotic cells do not have membrane-enclosed Diatoms are stramenopiles with shells composed of
organelles 518 two parts 551
A cell wall protects most prokaryotes 519 Brown algae are multicellular stramenopiles 551
Some bacteria produce capsules or slime layers 520 Most golden algae are unicellular biflagellates 552
Some prokaryotes have fimbriae or pili 520 26.5 rhizarians 553
Some bacteria survive unfavorable conditions by forming Forams extend cytoplasmic projections that form a threadlike,
endospores 520 interconnected net 553
Many types of prokaryotes are motile 521 Actinopods project slender axopods 553
25.2 Prokaryote Reproduction and Evolution 522 26.6 archaeplastids 554
Rapid reproduction contributes to prokaryote success 522 Red algae do not produce motile cells 554
Prokaryotes transfer genetic information 522 Green algae share many similarities with land plants 555
Evolution proceeds rapidly in prokaryote populations 523 26.7 unikonts 555
25.3 Nutritional and Metabolic Adaptations 524 Amoebozoa are unikonts with lobose pseudopodia 556
Most prokaryotes require oxygen 525 Choanoflagellates are opisthokonts closely related to
Some prokaryotes fix and metabolize nitrogen 525 animals 558

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27 Seedless Plants 563 29 The Fungi 603
27.1 adaptations of plants to life on land 564 29.1 Characteristics of Fungi 604
The plant life cycle alternates between haploid and diploid Fungi absorb food from the environment 604
generations 564 Fungi have cell walls that contain chitin 604
Four major groups of plants exist today 565 Most fungi consist of a network of filaments 604
27.2 bryophytes 568 29.2 Fungal Reproduction 605
Moss gametophytes are differentiated into “leaves” Many fungi reproduce asexually 605
and “stems” 568 Most fungi reproduce sexually 605
Liverwort gametophytes are either thalloid or leafy 571 29.3 Fungal Diversity 607
Hornwort gametophytes are inconspicuous thalloid Fungi are assigned to the opisthokont clade 607
plants 572 Diverse groups of fungi have evolved 607
Bryophytes are used for experimental studies 572 Chytrids have flagellate spores 608
Recap: details of bryophyte evolution are based on fossils and Zygomycetes reproduce sexually by forming
on structural and molecular evidence 573 zygospores 609
27.3 seedless vascular plants 574 Microsporidia have been a taxonomic mystery 610
Club mosses are small plants with rhizomes and short, Glomeromycetes have a symbiotic relationship with
erect branches 574 plant roots 611
Ferns are a diverse group of spore-forming vascular Ascomycetes reproduce sexually by forming
plants 575 ascospores 612
Whisk ferns are classified as reduced ferns 576 Basidiomycetes reproduce sexually by forming
Horsetails are an evolutionary line of ferns 576 basidiospores 615
Some ferns and club mosses are heterosporous 577 29.4 Ecological Importance of Fungi 618
Seedless vascular plants are used for experimental studies 578 Fungi form symbiotic relationships with some
Seedless vascular plants arose more than 420 mya 580 animals 618
Mycorrhizae are symbiotic associations between fungi
and plant roots 618
28 Seed Plants 584 A lichen consists of one or more fungi and a
28.1 an introduction to seed plants 585 photoautotroph 619
28.2 gymnosperms 586 29.5 Economic, Biological, and Medical Impact
Conifers are woody plants that produce seeds of Fungi 621
in cones 586 Fungi provide beverages and food 621
Pines represent a typical conifer life cycle 588 Fungi are important to modern biology and medicine 622
Cycads have seed cones and compound leaves 589 Fungi are used in bioremediation and to biologically
Ginkgo biloba is the only living species in its control pests 623
phylum 590 Some fungi cause diseases in humans and other
Gnetophytes include three unusual genera 591 animals 623
28.3 flowering plants 591 Fungi cause many important plant diseases 623
Monocots and eudicots are the two largest classes of
flowering plants 592
Sexual reproduction takes place in flowers 593 30 An Introduction to Animal
The life cycle of flowering plants includes double Diversity 628
fertilization 594 30.1 Animal Characteristics 629
Seeds and fruits develop after fertilization 596 30.2 Adaptations to Ocean, Freshwater, and
Flowering plants have many adaptations that account for ­Terrestrial Habitats 630
their success 596 Marine habitats offer many advantages 630
Floral structure provides insights into the evolutionary Some animals are adapted to freshwater
process 596 habitats 630
28.4 the evolution of seed plants 597 Terrestrial living requires major adaptations 630
Our understanding of the evolution of flowering plants has 30.3 Animal Evolution 631
made great progress in recent years 597 Molecular systematics helps biologists interpret the
The basal angiosperms comprise three clades 599 fossil record 631
The core angiosperms comprise magnoliids, monocots, Biologists develop hypotheses about the evolution
and eudicots 600 of development 631

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30.4 Reconstructing Animal Phylogeny 632 Many sea stars capture prey 678
Animals exhibit two main types of body symmetry 632 Basket stars and brittle stars make up the largest group
Animal body plans are linked to the level of tissue of echinoderms 680
development 633 Sea urchins and sand dollars have movable spines 680
Most bilateral animals have a body cavity lined with Sea cucumbers are elongated, sluggish animals 680
mesoderm 634 32.3 The Chordates: Major Characteristics 681
Bilateral animals form two main clades based on differences 32.4 Invertebrate Chordates 682
in development 635 Tunicates are common marine animals 682
Biologists have identified major animal clades based on Lancelets clearly exhibit chordate characteristics 682
­structure, development, and molecular data 635 Systematists debate chordate phylogeny 683
Segmentation apparently evolved three times 636 32.5 Introducing the Vertebrates 684
The vertebral column is a derived vertebrate character 684
Vertebrate taxonomy is a work in progress 686
31 Sponges, Cnidarians, Ctenophores, 32.6 Jawless Fishes 686
and Protostomes 641 32.7 Evolution of Jaws and Limbs: Jawed Fishes and
31.1 Sponges, Cnidarians, and Ctenophores 642 Tetrapods 688
Sponges have collar cells and other specialized cells 642 Most cartilaginous fishes inhabit marine environments 688
Cnidarians have unique stinging cells 644 Ray-finned fishes gave rise to modern bony fishes 690
Comb jellies have adhesive glue cells that trap prey 648 Tetrapods evolved from sarcopterygian ancestors 691
31.2 The Lophotrochozoa 649 Amphibians were the first successful land
Flatworms are bilateral acoelomates 649 vertebrates 693
Nemerteans are characterized by their proboscis 652 32.8 Amniotes: Terrestrial Vertebrates 694
Mollusks have a muscular foot, visceral mass, and mantle 653 Our understanding of amniote phylogeny is changing 695
Annelids are segmented worms 657 Reptiles have many terrestrial adaptations 695
The lophophorates are distinguished by a ciliated ring Biologists assign reptiles to two major lineages 696
of tentacles 659 Lizards and snakes are common modern reptiles 696
Rotifers have a crown of cilia 661 Tuataras superficially resemble lizards 698
31.3 The Ecdysozoa 662 Turtles have protective shells 698
Roundworms are of great ecological importance 662 Crocodilians have an elongated skull 699
Arthropods are characterized by jointed appendages and How do we know that birds are really dinosaurs? 699
an exoskeleton of chitin 662 Early birds were transitional forms 699
Modern birds are adapted for flight 700
Mammals (class Mammalia) have many unique
32 The Deuterostomes 676 characters 702
32.1 What are Deuterostomes? 677 New fossil discoveries are changing our understanding of
32.2 Echinoderms 677 the early evolution of mammals 702
Feather stars and sea lilies are suspension feeders 678 Modern mammals are assigned to three subclasses 703

part six: Structure and Life Processes in Plants

33 Plant Structure, Growth, and The orientation of cellulose microfibrils affects the direction
Development 710 of cell expansion 724
33.1 The Plant Body 711 Cell differentiation depends in part on a cell’s location 724
The plant body consists of cells and tissues 711 Morphogenesis occurs through pattern formation 725
The ground tissue system is composed of three simple tissues 711
The vascular tissue system consists of two complex tissues 716
The dermal tissue system consists of two complex tissues 718 34 Leaf Structure and Function 729
33.2 Plant Meristems 720 34.1 Leaf Form and Structure 730
Primary growth takes place at apical meristems 721 Leaf structure is adapted for maximum light absorption 730
Secondary growth takes place at lateral meristems 721 34.2 Stomatal Opening and Closing 736
33.3 Development of Form 722 Blue light triggers stomatal opening 736
The plane and symmetry of cell division affect plant form 723 Additional factors affect stomatal opening and closing 737

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34.3 Transpiration and Guttation 737 Many plants have mechanisms that prevent
Some plants exude liquid water 738 self-pollination 786
34.4 Leaf Abscission 739 Flowering plants and their animal pollinators have
In many leaves, abscission occurs at an abscission zone coevolved 786
near the base of the petiole 739 Some flowering plants depend on wind to disperse
34.5 Modified Leaves 740 pollen 788
Modified leaves of carnivorous plants capture insects 742 37.3 Fertilization and Seed and Fruit
Development 789
A unique double fertilization process occurs in flowering
35 Stem Structure and Transport 745 plants 790
35.1 Stem Growth and Structure 746 Embryonic development in seeds is orderly and
Herbaceous eudicot and monocot stems differ in internal predictable 790
structure 746 The mature seed contains an embryonic plant and storage
Woody plants have stems with secondary growth 748 materials 791
35.2 Water Transport 754 Fruits are mature, ripened ovaries 792
Water and minerals are transported in xylem 754 Seed dispersal is highly varied 794
Water movement can be explained by a difference in water 37.4 Germination and Early Growth 796
potential 755 Some seeds do not germinate immediately 797
According to the tension–cohesion model, water is pulled up Eudicots and monocots exhibit characteristic patterns of
a stem 755 early growth 797
Root pressure pushes water from the root up a stem 756 37.5 Asexual Reproduction in Flowering Plants 797
35.3 Translocation of Sugar in Solution 757 Apomixis is the production of seeds without the sexual
The pressure–flow model explains translocation in process 799
phloem 757 37.6 A Comparison of Sexual and Asexual
Reproduction 800
Sexual reproduction has some disadvantages 800
36 Roots and Mineral Nutrition 762
36.1 Root Structure and Function 763
Roots have root caps and root hairs 763 38 Plant Developmental Responses
The arrangement of vascular tissues distinguishes the roots of to ­External and Internal Signals 803
herbaceous eudicots and monocots 764 38.1 Tropisms 804
Woody plants have roots with secondary growth 767 38.2 Plant Hormones and Development 805
Some roots are specialized for unusual functions 768 Plant hormones act by signal transduction 805
36.2 Root Associations and Interactions 769 Auxins promote cell elongation 807
Mycorrhizae facilitate the uptake of essential minerals by Gibberellins promote stem elongation 809
roots 771 Cytokinins promote cell division 810
Rhizobial bacteria fix nitrogen in the roots of leguminous Ethylene promotes abscission and fruit ripening 811
plants 772 Abscisic acid promotes seed dormancy 812
36.3 The Soil Environment 773 Brassinosteroids are plant steroid hormones 812
Soil comprises inorganic minerals, organic matter, air, and Identification of a universal flower-promoting signal remains
water 773 elusive 813
About 50% of soil volume is composed of pore spaces 775 38.3 Light Signals and Plant Development 813
Soil organisms form a complex ecosystem 775 Phytochrome detects day length 814
Soil pH affects soil characteristics and plant growth 775 Competition for sunlight among shade-avoiding plants
Soil provides most of the minerals found in plants 776 involves phytochrome 815
Soil can be damaged by human mismanagement 778 Phytochrome is involved in other responses to light,
including germination 816
Phytochrome acts by signal transduction 816
37 Reproduction in Flowering Plants 782 Light influences circadian rhythms 816
37.1 The Flowering Plant Life Cycle 783 38.4 Responses to Herbivores and Pathogens 817
Flowers develop at apical meristems 783 Jasmonic acid activates several plant defenses 818
Each part of a flower has a specific function 783 Methyl salicylate may induce systemic acquired
37.2 Pollination 786 resistance 818

xvi / Contents

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part seven: Structure and Life Processes in Animals

39 Animal Structure and Function: 41 Neural Signaling 860


An Introduction 821 41.1 Neural Signaling: an Overview 861
39.1 Tissues, Organs, and Organ Systems 822 41.2 Neurons and Glial Cells 862
Epithelial tissues cover the body and line its Neurons receive stimuli and transmit neural signals 862
cavities 822 Certain regions of the CNS produce new neurons 862
Glands are made of epithelial cells 823 Axons aggregate to form nerves and tracts 863
Epithelial cells form membranes 823 Glial cells play critical roles in neural function 863
Connective tissues support other body 41.3 Transmitting Information along the Neuron 865
structures 823 Ion channels and pumps maintain the resting potential of the
Muscle tissue is specialized to contract 828 neuron 865
Nervous tissue controls muscles and glands 829 Ions cross the plasma membrane by diffusion through ion
Tissues and organs make up the organ systems of the channels 866
body 830 Ion pumping maintains the gradients that determine the
39.2 Regulating the Internal Environment 834 resting potential 867
Negative feedback systems restore homeostasis 834 Graded local signals vary in magnitude 867
A few positive feedback systems operate in the body 835 Axons transmit signals called action potentials 867
39.3 Regulating Body Temperature 836 An action potential is generated when the voltage reaches
Ectotherms absorb heat from their surroundings 836 threshold level 867
Endotherms derive heat from metabolic processes 836 The neuron repolarizes and returns to a resting state 868
Many animals adjust to challenging temperature changes 839 The action potential is an all-or-none response 869
An action potential is self-propagating 870
Several factors determine the velocity of an action
40 Protection, Support, and potential 871
Movement 842 41.4 Transmitting Information across Synapses 872
40.1 Epithelial Coverings 843 Signals across synapses can be electrical or chemical 872
Invertebrate epithelium may secrete a cuticle 843 Neurons use neurotransmitters to signal other cells 873
Vertebrate skin functions in protection and temperature Neurotransmitters bind with receptors on postsynaptic
regulation 843 cells 873
40.2 Skeletal Systems 844 Activated receptors can send excitatory or inhibitory
In hydrostatic skeletons body fluids transmit force 844 signals 874
Mollusks and arthropods have nonliving exoskeletons 845 41.5 Neural Integration 877
Internal skeletons are capable of growth 845 Postsynaptic potentials are summed over time and space 877
The vertebrate skeleton has two main divisions 846 Where does neural integration take place? 877
A typical long bone amplifies the motion generated by 41.6 Neural Circuits: Complex Information
muscles 846 Signaling 877
Bones are remodeled throughout life 847
Joints are junctions between bones 847
40.3 Muscle Contraction 848 42 Neural Regulation 882
Invertebrate muscle varies among groups 848 42.1 Invertebrate Nervous Systems: Trends in
Vertebrate skeletal muscles act antagonistically to one Evolution 883
another 849 42.2 Overview of the Vertebrate Nervous System 884
A vertebrate muscle may consist of thousands of muscle 42.3 Evolution of the Vertebrate Brain 885
fibers 849 The hindbrain develops into the medulla, pons, and
Contraction occurs when actin and myosin filaments move cerebellum 886
past one another 850 The midbrain is prominent in fishes and amphibians 886
ATP powers muscle contraction 853 The forebrain gives rise to the thalamus, hypothalamus, and
The type of muscle fibers determines strength and cerebrum 887
endurance 855 42.4 The Human Central Nervous System 888
Several factors influence the strength of muscle The spinal cord transmits impulses to and from the brain 888
contraction 855 The most prominent part of the human brain is the
Smooth muscle and cardiac muscle are involuntary 856 cerebrum 889
Contents / xvii

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Axons in the white matter of the cerebrum connect parts of 44.2 Vertebrate Blood 939
the brain 892 Plasma is the fluid component of blood 939
The body follows a circadian rhythm of sleep and Red blood cells transport oxygen 939
wakefulness 892 White blood cells defend the body against disease
The limbic system affects emotional aspects of behavior 896 organisms 940
Learning and memory involve long-term changes Platelets function in blood clotting 941
at synapses 897 44.3 Vertebrate Blood Vessels 942
Language involves comprehension and expression 901 44.4 Evolution of the Vertebrate Circulatory
42.5 The Peripheral Nervous System 901 System 944
The somatic division helps the body adjust to the external 44.5 The Human Heart 946
environment 901 Each heartbeat is initiated by a pacemaker 947
The autonomic division regulates the internal The cardiac cycle consists of alternating periods of contraction
environment 901 and relaxation 948
42.6 Effects of Drugs on the Nervous System 903 The nervous system regulates heart rate 949
Drug addiction is a serious issue 904 Stroke volume depends on venous return 950
Opioid overdose is an epidemic 904 Cardiac output varies with the body’s need 950
44.6 Blood Pressure 950
Blood pressure varies in different blood vessels 952
43 Sensory Systems 911 Blood pressure is carefully regulated 952
43.1 How Sensory Systems Work 912 44.7 The Pattern of Circulation 953
Sensory receptors receive information 912 The pulmonary circulation oxygenates the blood 954
Sensory receptors transduce energy 912 The systemic circulation delivers blood to the tissues 954
Sensory input is integrated at many levels 912 44.8 The Lymphatic System 955
We can classify sensory receptors based on location The lymphatic system consists of lymphatic vessels and lymph
of ­stimuli or on the type of energy they transduce 914 tissue 955
43.2 Thermoreceptors 915 The lymphatic system plays an important role in fluid
43.3 Electroreceptors and Magnetic Reception 916 homeostasis 956
43.4 Nociceptors 916 44.9 Cardiovascular Disease 956
43.5 Mechanoreceptors 916 Atherosclerosis develops progressively 957
Tactile receptors are located in the skin 917 Atherosclerosis has many effects 958
Proprioceptors help coordinate muscle movement 918 Cardiovascular disease can be treated 958
Many invertebrates have gravity receptors called statocysts 918 The risk of cardiovascular disease can be lowered 959
Hair cells are characterized by stereocilia 919
Lateral line organs supplement vision in fishes 919
The vestibular apparatus maintains equilibrium 919
45 The Immune System: Internal
Auditory receptors are located in the cochlea 921
Defense 962
45.1 Evolution of Immune Responses 963
43.6 Chemoreceptors 924
Invertebrates launch innate immune responses 963
Taste receptors detect dissolved food molecules 925
Vertebrates launch both innate and adaptive immune
The olfactory epithelium is responsible for the sense of smell 925
responses 964
Many animals communicate with pheromones 926
45.2 Innate Immune Responses in Vertebrates 965
43.7 Photoreceptors 926
Physical barriers and chemical weapons stop most
Invertebrates have several types of light-sensing organs 926
pathogens 965
Vertebrate eyes form sharp images 927
Cells of the innate immune system destroy pathogens 965
The retina contains light-sensitive rods and cones 929
Cytokines are important signaling molecules 966
Light activates rhodopsin 930
Complement promotes destruction of pathogens and
Color vision depends on three types of cones 931
­enhances inflammation 967
Integration of visual information begins in the retina 931
Inflammation is a protective response 967
45.3 Adaptive Immune Responses in Vertebrates 969
44 Internal Transport 936 Many types of cells are involved in adaptive immune
44.1 Types of Circulatory Systems 937 responses 969
Many invertebrates have an open circulatory system 937 The major histocompatibility complex is responsible for
Some invertebrates have a closed circulatory system 938 ­recognition of self 971
Vertebrates have a closed circulatory system 938 45.4 Cell-Mediated Immunity 972

xviii / Contents

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45.5 Antibody-Mediated Immunity 973 47 Processing Food and Nutrition 1010
A typical antibody consists of four polypeptide 47.1 Nutritional Styles and Adaptations 1011
chains 974 Animals are adapted to their mode of nutrition 1011
Antibodies are grouped in five classes 976 Some invertebrates have a digestive cavity with a single
Antigen–antibody binding activates other defenses 977 opening 1012
The immune system responds to millions of different Most animal digestive systems have two openings 1013
antigens 977 47.2 The Vertebrate Digestive System 1013
Monoclonal antibodies are highly specific 978 Food processing begins in the mouth 1015
Immunological memory is responsible for long-term The pharynx and esophagus conduct food to the
immunity 979 stomach 1016
45.6 Response to Disease, Immune Failures, and Food is mechanically and enzymatically digested in the
Harmful Reactions 980 stomach 1016
Cancer cells evade the immune system 981 Most enzymatic digestion takes place in the small
Immunodeficiency disease can be acquired or intestine 1017
inherited 982 The liver secretes bile 1019
HIV is the major cause of acquired immunodeficiency in The pancreas secretes digestive enzymes 1019
adults 982 Nutrients are digested as they move through the digestive
In an autoimmune disease, the body attacks its own tract 1019
tissues 984 Nerves and hormones regulate digestion 1020
Rh incompatibility can result in hypersensitivity 985 Absorption takes place mainly through the villi of the small
Allergic reactions are directed against ordinary environmental intestine 1021
antigens 985 The large intestine eliminates waste 1021
Graft rejection is an immune response against transplanted 47.3 Required Nutrients 1022
tissue 987 Carbohydrates provide energy 1022
Lipids provide energy and are used to make biological
molecules 1023
46 Gas Exchange 991
Proteins serve as enzymes and as structural components of
46.1 Adaptations for Gas Exchange in Air or
cells 1024
Water 992
Vitamins are organic compounds essential for normal
46.2 Types of Respiratory Surfaces 992
metabolism 1024
The body surface may be adapted for gas exchange 992
Minerals are inorganic nutrients 1026
Tracheal tube systems deliver air directly to the cells 992
Antioxidants inactivate reactive molecules 1026
Gills are the respiratory surfaces in many aquatic
Phytochemicals play important roles in maintaining
animals 994
health 1027
Terrestrial vertebrates exchange gases through
47.4 Energy Metabolism 1027
lungs 994
Energy metabolism is regulated by complex signaling 1028
46.3 The Mammalian Respiratory System 997
Obesity is a serious nutritional problem 1028
The airway conducts air into the lungs 997
Undernutrition can cause serious health problems 1029
Gas exchange occurs in the alveoli of the lungs 998
Ventilation is accomplished by breathing 998
The quantity of respired air can be measured 998 48 Osmoregulation and Disposal
Gas exchange takes place in the alveoli 998 of ­Metabolic Wastes 1032
Gas exchange takes place in the tissues 1001 48.1 Maintaining Fluid and Electrolyte Balance 1033
Respiratory pigments increase capacity for oxygen 48.2 Metabolic Waste Products 1033
transport 1001 48.3 Osmoregulation and Excretion in
Carbon dioxide is transported mainly as bicarbonate Invertebrates 1034
ions 1002 Nephridial organs are specialized for osmoregulation and/or
Breathing is regulated by respiratory centers in excretion 1034
the brain 1002 Malpighian tubules conserve water 1035
Hyperventilation reduces carbon dioxide 48.4 Osmoregulation and Excretion in
concentration 1004 Vertebrates 1036
High flying or deep diving can disrupt homeostasis 1004 Freshwater vertebrates must rid themselves of excess
Some mammals are adapted for diving 1004 water 1036
46.4 Breathing Polluted Air 1005 Marine vertebrates must replace lost fluid 1036
Contents / xix

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Terrestrial vertebrates must conserve water 1037 The accessory glands produce the fluid portion of
48.5 The Urinary System of Mammals 1038 semen 1079
The nephron is the functional unit of the kidney 1040 The penis transfers sperm to the female 1080
Urine is produced by glomerular filtration, tubular Testosterone has multiple effects 1081
­reabsorption, and tubular secretion 1041 The hypothalamus, pituitary gland, and testes regulate
Glomerular filtration is not selective with regard to ions and male reproduction 1081
small molecules 1041 50.3 Human Reproduction: The Female 1082
Urine becomes concentrated as it passes through the renal The ovaries produce gametes and sex hormones 1083
tubule 1043 The oviducts transport the secondary oocyte 1084
Urine consists of water, nitrogenous wastes, and The uterus incubates the embryo 1084
salts 1044 The vagina receives sperm 1085
Hormones regulate kidney function 1044 The vulva are external genital structures 1085
Breasts function in lactation 1086
The hypothalamus, pituitary gland, and ovaries regulate
49 Endocrine Regulation 1050 ­female reproduction 1086
49.1 An Overview of Endocrine Regulation 1051 Menstrual cycles stop at menopause 1089
The endocrine system and nervous system interact to regulate Most mammals have estrous cycles 1091
the body 1051 50.4 Fertilization, Pregnancy, and Birth 1091
Negative feedback systems regulate endocrine activity 1051 Fertilization is the fusion of sperm and egg 1091
Hormones are assigned to four chemical groups 1052 Hormones are necessary to maintain pregnancy 1093
49.2 Types of Endocrine Signaling 1053 The birth process depends on a positive feedback
Neurohormones are transported in the blood 1053 system 1093
Some local regulators are considered hormones 1053 50.5 Human Sexual Response 1094
49.3 Mechanisms of Hormone Action 1055 50.6 Birth Control Methods and Abortion 1096
Lipid-soluble hormones enter target cells and activate Many birth control methods are available 1096
genes 1055 Most hormonal contraceptives prevent ovulation 1096
Water-soluble hormones bind to ­cell-surface receptors 1056 Intrauterine devices are widely used 1097
49.4 Neuroendocrine Regulation in Invertebrates 1058 Barrier methods of contraception include the diaphragm
49.5 Endocrine Regulation in Vertebrates 1058 and condom 1098
Homeostasis depends on normal concentrations of Emergency contraception is available 1098
hormones 1058 Sterilization renders an individual incapable of producing
The hypothalamus regulates the pituitary gland 1058 offspring 1098
The posterior pituitary gland releases hormones produced Future contraceptives may control regulatory
by the hypothalamus 1059 peptides 1099
The anterior pituitary gland regulates growth and other Abortions can be spontaneous or induced 1099
­endocrine glands 1059 50.7 Sexually Transmitted Infections 1099
Thyroid hormones increase metabolic rate 1062
Negative feedback systems regulate thyroid secretion 1064
The parathyroid glands regulate calcium concentration 1065 51 Animal Development 1104
The islets of the pancreas regulate blood glucose 51.1 Development of Form 1105
concentration 1065 51.2 Fertilization 1105
The adrenal glands help the body respond to stress 1068 The first step in fertilization involves contact and
Many other hormones help regulate life processes 1071 recognition 1105
Sperm entry is regulated 1106
Fertilization activates the egg 1107
50 Reproduction 1074 Sperm and egg pronuclei fuse, restoring the diploid
50.1 Asexual and Sexual Reproduction 1075 state 1107
Asexual reproduction is an efficient strategy 1075 51.3 Cleavage 1107
Most animals reproduce sexually 1075 The pattern of cleavage is affected by yolk 1107
Sexual reproduction increases genetic variability 1076 Cleavage may distribute developmental determinants 1109
50.2 Human Reproduction: the Male 1077 Cleavage provides building blocks for development 1110
The testes produce gametes and hormones 1077 51.4 Gastrulation 1110
A series of ducts store and transport sperm 1079 The amount of yolk affects the pattern of gastrulation 1111

xx / Contents

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51.5 Organogenesis 1113 Animal cognition is controversial 1130
51.6 Extraembryonic Membranes 1115 Play may be practice behavior 1131
51.7 Human Development 1115 52.3 Behavioral Responses to Environmental
The placenta is an organ of exchange 1116 Stimuli 1131
Organ development begins during the first trimester 1118 Biological rhythms regulate many behaviors 1131
Development continues during the second and third Environmental signals trigger physiological responses that
trimesters 1118 lead to migration 1132
More than one mechanism can lead to a multiple birth 1119 52.4 Foraging Behavior 1133
Environmental factors affect the embryo 1119 52.5 Costs and Benefits of Social Behavior 1134
The neonate must adapt to its new environment 1119 Communication is necessary for social behavior 1135
Aging is not a uniform process 1121 Dominance hierarchies establish social status 1136
Many animals defend a territory 1137
Some insect societies are highly organized 1138
52 Animal Behavior 1124 52.6 Sexual Selection 1140
52.1 Behavior and Adaptation 1125 Animals of the same sex compete for mates 1140
Behaviors have benefits and costs 1125 Animals select quality mates 1140
Genes interact with environment 1125 Sexual selection favors polygynous mating systems 1141
Behavior depends on physiological readiness 1126 Some animals care for their young 1142
Many behavior patterns depend on motor programs 1127 52.7 Helping Behavior 1143
52.2 Learning: Changing Behavior as a Result Altruistic behavior can be explained by inclusive fitness 1145
of Experience 1127 Helping behavior may have alternative explanations 1145
An animal habituates to irrelevant stimuli 1128 Some animals help nonrelatives 1145
Imprinting occurs during an early critical period 1129 52.8 Culture in Vertebrate Societies 1146
In classical conditioning, a reflex becomes associated with Some vertebrates transmit culture 1146
a new stimulus 1129 Sociobiology explains human social behavior in terms
In operant conditioning, spontaneous behavior is reinforced 1129 of adaptation 1147

part eight: The Interactions of Life: Ecology

53 Introduction to Ecology: 54 Community Ecology 1171


Population Ecology 1151 54.1 Community Structure and Functioning 1172
53.1 Features of Populations 1152 Community interactions are complex and often not readily
Density and dispersion are important features of populations 1152 apparent 1173
53.2 Changes in Population Size 1154 The niche is a species’ ecological role in the
Dispersal affects the growth rate in some populations 1154 community 1173
Each population has a characteristic intrinsic rate of Competition is intraspecific or interspecific 1175
increase 1154 Natural selection shapes the bodies and behaviors of both
No population can increase exponentially indefinitely 1155 predator and prey 1178
53.3 Factors Influencing Population Size 1156 Symbiosis involves a close association between
Density-dependent factors regulate population size 1156 species 1180
Density-independent factors are generally abiotic 1159 54.2 Strength and Direction of Community
53.4 Life History Traits 1160 Interactions 1183
Life tables and survivorship curves indicate mortality and Other species of a community depend on or are greatly
survival 1161 affected by keystone species 1183
53.5 Metapopulations 1163 Dominant species influence a community as a result of their
53.6 Human Populations 1164 greater size or abundance 1184
Not all countries have the same growth rate 1165 Ecosystem regulation occurs from the bottom up and top
The age structure of a country helps predict future population down 1184
growth 1166 54.3 Community Biodiversity 1185
Environmental degradation is related to population growth Ecologists seek to explain why some communities have more
and resource consumption 1167 species than others 1186

Contents / xxi

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Species richness may promote community stability 1187 Chaparral is a thicket of evergreen shrubs and small
54.4 Community Development 1189 trees 1221
Disturbance influences succession and species richness 1190 Deserts are arid ecosystems 1222
Ecologists continue to study community structure 1190 Savanna is a tropical grassland with scattered trees 1223
There are two basic types of tropical forests 1224
56.2 Aquatic Ecosystems 1226
55 Ecosystems and the Biosphere 1194 Freshwater ecosystems are linked to land and marine
55.1 Energy Flow through Ecosystems 1195 ecosystems 1226
Ecological pyramids illustrate how ecosystems Estuaries occur where fresh water and salt
work 1196 water meet 1230
Ecosystems vary in productivity 1197 Marine ecosystems dominate Earth’s surface 1231
Some toxins persist in the environment 1199 56.3 Ecotones 1235
55.2 Cycles of Matter in Ecosystems 1201 56.4 biogeography 1235
Carbon dioxide is the pivotal molecule in the carbon Land areas are divided into biogeographic realms 1236
cycle 1201
Bacteria and archaea are essential to the nitrogen
cycle 1202 57 Biological Diversity
The phosphorus cycle lacks a gaseous component 1204 and Conservation Biology 1241
Water moves among the ocean, land, and atmosphere in 57.1 The Biodiversity Crisis 1242
the hydrologic cycle 1205 Endangered species have certain characteristics in
55.3 Abiotic Factors in Ecosystems 1206 common 1243
The sun warms Earth 1206 Human activities contribute to declining biological
The atmosphere contains several gases essential to diversity 1245
organisms 1208 57.2 Conservation Biology 1248
The global ocean covers most of Earth’s surface 1209 In situ conservation is the best way to preserve biological
Climate profoundly affects organisms 1210 diversity 1249
Fires are a common disturbance in some ecosystems 1211 Ex situ conservation attempts to save species on the brink
55.4 Studying Ecosystem Processes 1212 of extinction 1252
The Endangered Species Act provides some legal protection
for species and habitats 1252
International agreements provide some protection for
56 Ecology and the Geography
­species and habitats 1253
of Life 1216
56.1 Biomes 1217 57.3 Deforestation 1253
Why are tropical rain forests continuing to disappear? 1254
Tundra is the cold, boggy plains of the far
Why are boreal forests disappearing? 1255
north 1217
Boreal forest is the evergreen forest of the
57.4 Climate Change 1255
Greenhouse gases cause climate change 1256
north 1219
What are the probable effects of climate change? 1257
Temperate rain forest has cool weather, dense fog, and
high precipitation 1219
The Future? 1260
Temperate deciduous forest has a canopy of broad-leaf
trees 1220
Glossary G-1
Temperate grasslands occur in areas of moderate
precipitation 1220 Index I-1

xxii / Contents

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Preface

This eleventh edition of Solomon, Martin, Martin, and Berg’s


Biology conveys our vision of the dynamic science of biology
The Solomon/Martin/Martin/Berg
and how it affects every aspect of our lives, from our own Learning System
health and behavior to the challenging global environmen-
tal issues that confront us. New discoveries in the biological In the eleventh edition, we have continued to refine our
sciences continue to increase our understanding of both the highly successful Learning System. This system provides the
unity and diversity of life’s processes and adaptations. With student with the learning strategies needed to integrate bio-
this understanding, we become ever more aware of our inter- logical concepts and demonstrate mastery of these concepts.
dependence with the vast diversity of organisms with which Learning biology is challenging because it requires learning
we share planet Earth. many new terms and facts that must then be integrated into
the framework of biological principles. To help students focus
on important principles and concepts, we provide Learning
­Outcomes for the course and Learning Objectives for each
Biology: The Student-Friendly major section of every chapter. At the end of each section, we
Biology Textbook provide Checkpoint questions based on the Learning ­Objectives
so that students can assess their level of understanding of the
We want beginning students to experience learning biology material presented in the section. At the end of each chap-
as an exciting journey of discovery. In the eleventh edition of ter, we include a Summary: Focus on Learning Objectives that
Biology, we explore Earth’s diverse organisms, their remark- is organized around the Learning Objectives and emphasizes
able adaptations to the environment, and their evolutionary key terms in context. The Summary is followed by Test Your
and ecological relationships. We present the workings of Understanding, a set of questions organized according to
science and the contributions of scientists whose discover- Bloom’s taxonomy. Questions include Know and Comprehend
ies not only expand our knowledge of biology but also help multiple-choice exercises as well as a variety of questions that
shape and protect the future of our planet. Biology provides encourage the student to Apply and Analyze and Evaluate and
insight into what science is, how scientists work, what scien- Synthesize the topics in the chapter.
tists have contributed, and how scientific knowledge affects Students are directed to www.cengagebrain.com, a pow-
daily life. erful online tool that offers access to MindTap and additional
Since the first edition of Biology, we have worked very study materials. See the Resources for Students section of the
hard to present the principles of biology in an integrated Preface for details.
way that is accurate, interesting, and conceptually accessible
to students. In this eleventh edition of Biology, we continue
this tradition. We also continue to present biology in an in- Pedagogical Features
quiry-based framework. Some professors interpret inquiry Our Learning System includes numerous learning strategies
as a learning method that takes place in the laboratory as that help students increase their success:
students perform experiments. While laboratory research is
certainly an integral part of inquiry-based learning, inquiry • Our updated art program reinforces concepts discussed
is also a way of learning in which students actively pursue in the text and presents complex processes in clear steps.
knowledge outside the laboratory. In Biology, we have al- Key Experiment figures emphasize the scientific pro-
ways presented the history of scientific advances, including cess in both classic and modern research. These figures
­scientific debates, to help students understand that science encourage students to evaluate investigative approaches
is a process—that is, a field of investigative inquiry—as that scientists have taken. Examples include Figures 4-12,
well as a body of knowledge, the product of inquiry. In the 8-9, and 52-1. Key Point figures state important concepts
­eleventh edition of Biology, we further integrate inquiry- in process diagrams of complex topics. Examples include
based ­learning into the textbook, as discussed in the follow- Figures 4-11, 4-15, 27-2, 31-32, and 42-6. Many of the Key
ing sections. Point figures have numbered parts that show sequences of
Throughout the text, we stimulate interest by relating con- events in biological processes or life cycles.
cepts to experiences within the student’s frame of reference. • Numerous photographs, both alone and combined with
By helping students make such connections, we facilitate their line art, help students grasp concepts by connecting the
mastery of general concepts. The combined effect of an engag- “real” to the “ideal.” The line art uses features such as
ing writing style, interesting and exciting features, and our ­orientation icons to help students put the detailed figures
well-tested Learning System provides the ingredients for stu- into the broader context. We use symbols and colors con-
dents to succeed in their study of biology. sistently throughout the book to help students connect

xxiii

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
concepts. For example, the same four colors and shapes of the material in the chapter. Know and Comprehend
are used throughout the book to identify guanine, cyto- multiple-choice questions reinforce important terms and
sine, adenine, and thymine. Similarly, the same colors concepts. Apply and Analyze questions challenge students
are used consistently in illustrations and tables to indi- to integrate their knowledge. Higher-level Evaluate and
cate specific clades of organisms. Research Method fig- Synthesize questions encourage students to apply the con-
ures describe why biologists use a particular method and cepts just learned to new situations or to make connec-
explain how the method is executed. Examples include tions among important concepts. Each chapter has one or
Figures 4-7 and 15-9. more Evolution Link questions, and many chapters con-
• Many questions carry special designations: Predict; tain one or more Interpret Data questions that require stu-
­Connect; Visualize; Evolution Link; Interpret Data; or dents to actively interpret experimental data presented in
­Science, Technology, and Society. These questions empha- the chapter. Also included are Predict, Connect, Visualize,
size that learning is enhanced by many diverse approaches. and Science, Technology, and Society questions. Answers
• Inquiring About boxes explore issues of special relevance to the Test Your Understanding questions are provided in
to students, such as the effects of smoking, how traumatic Appendix E online at www.cengagebrain.com.
experiences affect the body, and breast cancer. These • The Glossary at the end of the book, the most compre-
boxes also provide a forum for discussing some inter- hensive glossary found in any biology text, provides
esting topics in more detail, such as the smallest ancient precise definitions of terms. The Glossary is especially
humans, ancient plants and coal formation, hydrothermal useful because it is extensively cross-referenced and
vent communities, declining amphibian populations, and includes pronunciations for many terms. The vertical
stratospheric ozone depletion. ­yellow bar along the margin facilitates rapid access to the
• A list of Key Concepts at the beginning of each chapter ­Glossary. MindTap also includes glossary flash cards with
provides a chapter overview and helps the student focus pronunciations.
on important principles discussed in the chapter.
• Learning Objectives at the beginning of each major sec-
tion in the chapter indicate, in behavioral terms, what the Course Learning Outcomes
student must do to demonstrate mastery of the material At the end of a successful study of introductory biology,
in that section. the student can demonstrate mastery of biological concepts
• Each major section of the chapter is followed by a series of by responding accurately to the following Course Learning
Checkpoint questions that assess comprehension by ask- Outcomes:
ing the student to describe, explain, compare, contrast, or
illustrate important concepts. The Checkpoint questions • Design an experiment to test a given hypothesis, using the
are based on the section Learning Objectives. procedure and terminology of the scientific method.
• Concept Statement Subheads introduce sections, preview- • Cite the cell theory and relate the structure of organelles to
ing and summarizing the key idea or ideas to be discussed their functions in both prokaryotic and eukaryotic cells.
in that section. • Describe the mechanisms of evolution, explain why evo-
• Sequence Summaries within the text simplify and summa- lution is the principal unifying concept in biology, and
rize information presented in paragraph form. For exam- discuss natural selection as the primary agent of evolu-
ple, paragraphs describing blood circulation through the tionary change.
body or the steps by which cells take in certain materials • Explain the role of genetic information in all species and
are followed by a Sequence Summary listing the sequence discuss applications of genetics that affect society.
of structures or steps. • Describe several mechanisms by which cells and organ-
• Numerous tables, many illustrated, help the student orga- isms transfer information, including the use of nucleic
nize and summarize material presented in the text. Many acids in genetic transmission of information, signal trans-
tables are color-coded. duction, chemical signals (such as hormones and phero-
• A Summary: Focus on Learning Objectives at the end of mones), electrical signals (such as neural transmission),
each chapter is organized around the chapter Learning sounds, and visual displays.
Objectives. This summary provides a review of the mate- • Provide examples (at various levels of complexity) of
rial, and because selected key terms are boldfaced in interactions among biological systems that illustrate the
the summary, students learn vocabulary words within the interdependence of these systems.
context of related concepts. • Explain how any given structure is related to its function.
• Test Your Understanding end-of-chapter questions are • Argue for or against the classification of organisms in three
organized according to Bloom’s taxonomy, providing stu- domains and several kingdoms or supergroups, charac-
dents with the opportunity to evaluate their understanding terizing each of these clades; based on your knowledge

xxiv / Preface

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
of genetics and evolution, give specific examples of the Chapter 1 examines several fundamental concepts in biology
unity and diversity of organisms in different domains and and the nature of the scientific process, including a discus-
supergroups. sion of systems biology. Chapters 2 and 3, which focus on the
• Compare the structural adaptations, life processes, and life molecular level of organization, establish the foundations in
cycles of a prokaryote, protist, fungus, plant, and animal. chemistry necessary for understanding biological processes.
• Define homeostasis and give examples of regulatory Chapters 4, 5, and 6 focus on the cellular level of organiza-
mechanisms, including feedback systems. tion, including cell structure and function, cell membranes,
• Trace the flow of matter and energy through a photosyn- and cell signaling. We emphasize the interdisciplinary nature
thetic cell and a nonphotosynthetic cell and through the of cell research. Topics discussed include transport between
biosphere, comparing the roles of producers, consumers, the nucleus and cytoplasm, routing of proteins through the
and decomposers. endomembrane system, and cell communication.
• Describe the study of ecology at the levels of an individual
organism, a population, a community, and an ecosystem. Part 2 Energy Transfer through
Living Systems
What’s New: An Overview Because all living cells need energy for life processes, the flow
of energy through living systems—that is, capturing energy
of Biology, Eleventh Edition and converting it to usable forms—is a basic theme of Biology.
Chapter 7 examines how cells capture, transfer, store, and use
Five themes are interwoven throughout Biology: the evolu- energy. Chapters 8 and 9 discuss the metabolic adaptations
tion of life, the transmission of biological information, the by which organisms obtain and use energy through cellular
flow of energy through living systems, interactions among respiration and photosynthesis.
biological systems, and the inter-relationship of structure and
function. As we introduce the concepts of modern biology, we Part 3 The Continuity of Life: Genetics
explain how these themes are connected and how life depends
on them. We have updated and expanded the eight chapters of Part 3
Educators present the major topics of an introductory bi- for the eleventh edition. We begin this unit by discussing
ology course in a variety of orders. For this reason, we care- mitosis and meiosis in Chapter 10. Chapter 11 builds on this
fully designed the eight parts of this book so that they do not foundation as it considers Mendelian genetics and related
depend heavily on preceding chapters and parts. This flexible patterns of inheritance. We then turn our attention to the
organization means that an instructor can present the 57 chap- structure and replication of DNA in Chapter 12. The discus-
ters in any number of sequences with pedagogical success. sion of RNA and protein synthesis in Chapter 13 includes
Chapter 1, which introduces the student to the major prin- insights into how the small percentage of DNA that codes
ciples of biology, provides a comprehensive springboard for for polypeptides relates to the much larger percentage of the
future discussions, whether the professor prefers a “top-down” genome that is expressed. We provide information establish-
or “bottom-up” approach. ing that much of the genome encodes different classes of
In this edition as in previous editions, we examined every non-protein-coding RNAs, including microRNAs and long
line of every chapter for accuracy and currency. We carefully non-coding RNAs. The regulatory functions of these RNAs
updated each topic and verified new material. The following are further explored in Chapter 14, which also includes infor-
brief summary provides a general overview of the organiza- mation on eukaryotic promoters, enhancers, and silencers as
tion of Biology and some of the changes made to the eleventh well as on epigenetic inheritance. In Chapter 15 we focus
edition. on DNA technology and genomics, including a discussion
of rapid DNA sequencing and CRISPR-mediated gene edit-
ing. We discuss the importance of gene databases as tools for
Part 1 The Organization of Life understanding gene regulation, gene functions, and molecu-
The six chapters that make up Part 1 provide basic princi- lar evolution. We have also added a section on genetically
ples of biology and the concepts of chemistry and cell biol- engineered gene drive systems. These chapters build the
ogy that lay the foundation upon which the remaining parts necessary foundation for exploring human genetics and the
of the book build. We begin Chapter 1 with a discussion of human genome in Chapter 16, which includes sections on
the promise and challenges of CRISPR research. We then genomic imprinting and on genome-wide association stud-
introduce the main themes of the book: evolution, informa- ies. In Chapter 17 we introduce the role of genes in devel-
tion transfer, energy transfer, interactions in biological sys- opment, emphasizing studies on specific model organisms
tems, and the inter-relationship of structure and function. that have led to spectacular advances in this field. We discuss

Preface / xxv

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induced pluripotent stem cells and present a comprehensive Part 6 Structure and Life Processes
view of cancer and its relationship to cell signaling. We link in Plants
these advances to the application of genome-wide associa-
tion studies and whole genome sequencing. Part 6 introduces students to the fascinating plant world. Here
we stress relationships between structure and function in plant
cells, tissues, organs, and individual organisms. In Chapter 33
Part 4 The Continuity of Life: Evolution we consider plant structure, growth, and differentiation in the
Although we explore evolution as the cornerstone of biol- context of cell division, cell expansion, cell differentiation,
ogy throughout the book, in Part 4 we discuss evolutionary tissue culture, morphogenesis, pattern formation, positional
concepts in depth. We provide the history behind the dis- information, and Arabidopsis mutants. Chapters 34 through 36
covery of the scientific theory of evolution, the mechanisms discuss the structural and physiological adaptations of leaves,
by which it occurs, and the methods by which it is studied stems, and roots; these chapters include special consideration
and tested. Chapter 18 introduces the Darwinian concept of of plant transport systems. Chapter 37 describes reproduction
evolution and presents several kinds of evidence that support in flowering plants, including asexual reproduction, flowers,
the scientific theory of evolution. In Chapter 19 we examine fruits, and seeds. Chapter 38 focuses on growth responses and
evolution at the population level. Chapter 20 describes the regulation of growth, including the latest findings generated
evolution of new species and discusses aspects of macroevo- by the continuing explosion of knowledge in plant biology,
lution. Chapter 21 summarizes the evolutionary history of particularly at the molecular level.
life on Earth. In Chapter 22 we recount the evolution of pri-
mates, including humans. We explore recent molecular and Part 7 Structure and Life Processes
fossil findings, including those relating to human relatives
such as the Denisovans (a sister species to the Neandertals) in Animals
as well as Australopithecus sediba and Homo naledi, species In Part 7 we provide a strong emphasis on comparative animal
whose fossils exhibit unique mixtures of apelike and human- physiology, showing the structural, functional, and behavioral
like features. adaptations that help animals meet environmental challenges.
We use a comparative approach to examine how various ani-
mal groups have solved both similar and diverse problems. In
Part 5 The Diversity of Life
Chapter 39 we discuss the basic tissues and organ systems of
Emphasizing the cladistic approach, we use an evolutionary the animal body, homeostasis, and the ways that animals regu-
framework to discuss each group of organisms. We present late their body temperature. Chapter 40 focuses on different
current hypotheses of how groups of organisms are related. types of body coverings, skeletons, and muscles, and discusses
In Chapter 23 we discuss why organisms are classified and how they function. In Chapters 41 through 43, we discuss
provide insight into the scientific process of deciding how neural signaling, neural regulation, and sensory reception.
they are classified. New advances have enabled us to further In Chapters 44 through 51, we compare how different ani-
clarify the connection between evolutionary history and sys- mal groups carry on life processes, such as internal transport,
tematics in the eleventh edition. Chapter 24 focuses entirely internal defense, gas exchange, digestion, osmoregulation and
on viruses and subviral agents. Topics include giant viruses, disposal of metabolic wastes, endocrine regulation, reproduc-
viral origins, and the evolutionary importance of viruses. tion, and development. Each chapter in Part 7 considers the
Chapter 25 is devoted to the prokaryotes, both bacteria and adaptations for the life processes being discussed. Part 7 ends
archaea. Information about the evolution, structure, ecology, with a discussion of behavioral adaptations in Chapter 52.
and phylogeny of archaea has been updated and expanded. Reflecting recent research findings, we have updated or added
Implications of research on the human microbiome are new material on many topics, including neurotransmitters,
discussed and discussion of antibiotic resistance has been cardiovascular disease, immune function, nutrition, regula-
expanded. tion of appetite and energy metabolism, the mesentery, endo-
Chapter 26 describes the protists in the context of five crine function, contraception, sexually transmitted infections,
­“supergroups” of eukaryotes. Chapters 27 and 28 present the and transmission of culture in vertebrates.
members of the plant kingdom. Chapter 27 considers the evolu-
tion of land plants and the evolution of seedless vascular plants.
Discussion of the origin and early evolution of angiosperms
Part 8 The Interactions of Life: Ecology
is included in Chapter 28. Chapter 29 describes the fungi. In In Part 8 we discuss the ecology of populations, communities,
Chapters 30 through 32, we discuss the diversity of animals. We and ecosystems and the application of ecological principles to
have updated the discussions of phylogenetic relationships to disciplines such as conservation biology. Throughout Chap-
reflect recent research. ters 53 through 57, we focus on human interdependence with

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other organisms, how we have disrupted the biosphere, and Each MindTap product offers the full, mobile-ready t­extbook
what we can do to stop global climate change, decreases in combined with superior and proven learning tools at one
species diversity, and other negative human impacts. ­affordable price. Students who purchase digital access can add
In Chapter 53 we focus on population ecology. In Chapter 54 a print option at any time.
we focus on community ecology. We discuss research advances
that facilitate understanding of community structure, interac- A Problem-based Guide to Basic Genetics by Donald
tions, synergy, and potential medical benefits. For example, we Cronkite of Hope College. This brief guide provides students
discuss research on the keystone wolves of Yellowstone and eco- with a systematic approach to solving genetics problems along
system cascade effects. We have added a new box, Inquiring About: with numerous solved problems and practice problems.
The Human Microbiota: Our Many Symbionts, which introduces
students to the various relationships we have with our m
­ icrobiota, Spanish Glossary This glossary includes key terms and
including health, metabolism, and immune functions. definitions in Spanish.
In Chapter 55 we discuss ecosystems and the biosphere.
We present updated discussions on community composition
research, illustrating the potential for balance between species Additional Resources for Instructors
richness and agricultural productivity. Chapter 56 focuses on This edition includes a comprehensive package of supple-
ecology and the geography of life. We discuss the degradation ments, available to qualified adopters. Please ask your local
of permafrost and the resulting concerns for global climate sales representative for details.
change. We have added a discussion of research suggesting
­ecotones as potential global climate change indicators and Instructor Companion Site Everything you need for your
have expanded research information on coral reef bleaching course in one place! This collection of book-specific l­ecture
and ­recovery and the effects of global warming. and class tools is available online via www.cengage.com/login.­
In Chapter 57 we discuss biological diversity and conser- Access and download PowerPoint presentations, images, instruc-
vation biology. We have expanded information on human- tor’s manual, videos, and more.
caused extinctions and decreases in biodiversity. We include
updated information on biodiversity hotspots and conserva- Cengage Learning Testing Powered by Cognero A
tion partnerships. flexible, online system that allows you to import, edit, and
This chapter includes new conservation biology research ­manipulate test bank content from the test bank or elsewhere,
on ecosystem restoration partnerships. We have also expanded including your own favorite test questions; create multiple test
information linking global climate change with sea level rise versions in an instant; and deliver tests from your LMS, your
and the increasing diseases spread by mosquitos. classroom, or wherever you want.

MindTap A fully online, highly personalized learning


A Comprehensive Package ­experience built on Cengage Learning content. MindTap com-
for Learning and Teaching bines student learning tools—readings, multimedia, activities,
and ­assessments—into a singular Learning Path that guides
A carefully designed supplement package is available to fur- students through their course. Instructors personalize the
ther facilitate learning. In addition to print resources, we are ­experience by customizing authoritative Cengage Learning con-
pleased to present student multimedia tools that have been tent and learning tools, including the ability to add their own
developed in conjunction with the text. content in the Learning Path via apps that integrate into the
MindTap framework seamlessly with Learning Management
Systems.
Resources for Students MindTap for Biology is easy to use and saves instructors
time by allowing them to:
MindTap A fully online learning experience built on C ­ engage
Learning content. MindTap combines student learning tools— • Seamlessly deliver appropriate content from a number of
readings, study tools, animations, activities, and ­assessments— providers.
into a singular Learning Path that guides students through • Break course content down into movable objects to pro-
their course. New features for the eleventh edition include Data mote personalization, encourage interactivity, and ensure
Analysis activities and Mastery Training powered by Cerego. student engagement.
Mastery Training uses the science of how we learn to support • Customize the course—from tools to text—and make
students’ understanding of the key concepts in each chapter adjustments “on the fly,” making it possible to intertwine
and can be accessed through your ­computer or the mobile app. breaking news into their lessons.

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
• Bring interactivity into learning through the integration of our copy editor, Laura Specht Patchkofsky, who helped us
of multimedia assets. maintain consistency and improve the manuscript. We appre-
• Track students’ use, activities, and comprehension in real ciate the efforts of intellectual property project manager Erika
time, which provides opportunities for early intervention Mugavin, intellectual property analyst Christine Myaskovsky,
to influence progress and outcomes. Grades are visible and photo researcher Gopala Krishnan Sankar in helping us
and archived so that students and instructors always have acquire excellent images. We also appreciate the work of text
access to current standings in the class. researcher Rameshkumar P.M.
We appreciate the help, patience, and hard work of our
production team. Our schedule for this project was very de-
Acknowledgments manding. At times, it seemed like the team worked around the
clock. When we sent e-mails late at night or during weekends,
The development and production of the eleventh edition we often received immediate responses.
of Biology required extensive interaction and cooperation These dedicated professionals and many others on our
among the authors and many individuals in our families, Cengage team provided the skill, attention, patience, and good
social circles, and professional environments. We thank all of humor needed to produce Biology, Eleventh Edition. We thank
you—our editors, colleagues, students, families, and friends them for their help and support throughout this project.
for your help and support. We appreciate the help of obstetrician/gynecologist Dr.
Preparing a book of this complexity is challenging and Amy Solomon for her input regarding the most recent infor-
requires a cohesive, talented, and hardworking professional mation on pregnancy, childbirth, contraception, and sexually
team. We appreciate the contributions of everyone on the edi- transmitted infections. We are grateful to Mical Solomon for
torial and production staff at Cengage Learning who worked his computer help. We thank Dr. David Axelrod for insightful
on this eleventh edition of Biology. We thank our product discussions on the genetics and biology of cancer.
team manager, April Cognato, for her commitment to ­Biology We thank our families and friends for their understand-
and for working closely with us throughout the process of ing, support, and encouragement as we struggled through
­development and production. We greatly appreciate the help of many revisions and intense deadlines. We especially thank
­Suzannah Alexander, our very talented content developer, who Dr. Kathleen M. Heide, Freda Brod, Margaret and Damian
was a critical part of our team. Suzannah expertly coordinated Brown, Alan Berg, and Jennifer and Pat Roath for their sup-
many aspects of this challenging project, including the render- port and input.
ing of new art and selection of new photographs for this edi- Our colleagues and students who have used our book have
tion. She made herself available to advise and help us whenever provided valuable input by sharing their responses to past
we needed her, including late at night and during weekends. editions of Biology. We thank them and ask again for their
We thank Tom Ziolkowski, our market development man- comments and suggestions as they use this new edition. We
ager and brand manager, whose expertise ensured that you can be reached via our website at www.cengagebrain.com or
would know about our new edition. through our editors at Cengage Learning.
We appreciate the help of content project manager Hal We greatly appreciate the participation and help of our
Humphrey, who expertly guided overall production of Biology, contributors:
Eleventh Edition.
Lois A. Ball
We are grateful to product team assistants Marina Starkey
Meteorologist, Biologist, and Science Educator
and Vanessa Desiato for quickly providing us with resources
University of South Florida
whenever we needed them.
We thank Helen Bruno, senior designer, as well as cover Lois worked closely with us on the revision of several chap-
designer and text designer Jeanne Calabrese. ters, including the microbiology, immunology, and ecology
We appreciate the work of Lauren Oliveira, associate con- chapters, helping us to update them with new findings.
tent development manager, who coordinated and managed
Alexandra McMurray
the many print and high-tech components of our Learning
Biology Major and Premedical Student
­System. We also thank content developers Tyler Sally and John
University of Miami
­Anderson for coordinating the print and media supplements.
We are grateful to our production project manager, Phil Alexandra reviewed more than 20 chapters and gave us
Scott at SPi Global, for coordinating the many editors involved valuable feedback from a college student’s perspective. She also
in the preparation of this edition and bringing together the assisted with researching and with updating statistics.
thousands of complex pieces of the project that together pro- We express our thanks to the many biologists who have
duced Biology, Eleventh Edition. We value the careful work read the manuscript during various stages of its development

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
and provided us with valuable suggestions for improving it. Melissa Murray Reedy, University of Illinois
Eleventh edition reviewers include the following: at Urbana-Champaign
E.C. Nebel, Delgado Community College
Jason Adams, College of Dupage
F.E. Nelson, Temple University
Alicia Anzaldo, Wilbur Wright College
Maryann Page, Onondaga Community College
Josh R. Auld, West Chester University
Christopher Pantazis, Danville Community College
Kristin Y. Bridge, Motlow State Community College
Nancy Peterson, Mineral Area College
Mark A Buchheim, University of Tulsa
John Plunket, Horry Georgetown Technical College
Ray Canham, Richland College of the Dallas County
Robert H. Reavis, Glendale Community College, Arizona
­Community College District
George R. Robinson, SUNY Albany
Ethan A. Carver, University of Tennessee at Chattanooga
David J. Rosen, Lee College
G. Chung, Broward College
Jyotsna Sharma, University of Texas at San Antonio
Kari B. Clifton, University of West Florida
Mark Shotwell, Slippery Rock University
Bryan Coppedge, Tulsa Community College
Matthew T. Smith, North Dakota State University
Neta Dean, Stony Brook University
Ayodotun O. Sodipe, Texas Southern University
Susan Erster, Stony Brook University
Shannon Stevenson, University of Minnesota, Duluth
Robert F. Feissner, SUNY Geneseo
William Terzaghi, Wilkes University
Jillian C. Fesolovich, Keystone College
Neal J. Voelz, Saint Cloud State University
Cynthia M. Galloway, Texas A&M University, Kingsville
D. Alexander Wait, Missouri State University
Sharada Gollapudi, San Jacinto College South
Justin Walguarnery, University of Hawaii
Nan Ho, Las Positas College
Eileen Walsh, Westchester Community College
Kirsten Hokeness, Bryant University
Amy Wiles, Mercer University
Dianella Howarth, St John’s University
Beverly Wiltz, Delgado Community College
Nicole J. Huber, University of Colorado at Colorado Springs
Holly Woodruff, Central Piedmont Community College
Jason Jennings, Southwest Tennessee Community College
Carly N. Jordan, The George Washington University
Amy Kennedy, Central Carolina Community College We would also like to thank the hundreds of reviewers of
Brenda Leady, University of Toledo previous editions, both professors and students, who are too
Jonathan H. Lieman, South Texas College numerous to mention. They asked thoughtful questions, pro-
Cynthia Littlejohn, University of Southern Mississippi vided new perspectives, offered alternative wordings to clarify
Jason L. Locklin, Temple College difficult passages, and informed us of possible errors. We are
Melissa Masse, Tulsa Community College truly indebted to their excellent feedback. Their suggestions
Colleen Mikelson, University of Tennessee at Chattanooga have helped us improve each edition of Biology.

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To the Student

We have learned a great deal from tens of thousands of stu- material in your text. Do not copy the information; instead,
dents who have taken on the challenge of learning biology. process it and write out an explanation in your own words.
Although they have varied in their life goals and academic Read the entire chapter, including parts that are not covered
preparation, most have found that they needed to modify in lecture. This extra information will give you breadth of un-
their approach to learning to be successful. derstanding and will help you grasp key concepts. In addition,
You already know that memorization and cramming are you should make an effort to employ as many of the techniques
unsuccessful, and you probably also know that many students described in the next paragraphs as possible.
fall back on these methods as default strategies. So, what really If the assigned readings in the text are going to be tested,
works? you must use your text intensively. After reading the chapter
introduction, read the list of Learning Objectives for the first
Use the Wealth of Learning Aids section. These objectives are written in behavioral terms; that
is, they ask you to “do” something to demonstrate mastery. The
That Accompany Biology objectives give you a concrete set of goals for each section of
The Learning System we use in this book is described in the the chapter. At the end of each section, you will find Checkpoint
Preface. Using the strategies of the Learning System will help questions keyed to the Learning Objectives. Carefully examine
you master the language and concepts of biology. You will each figure, making certain that you understand what it is il-
also want to use the many online tools available to Biology lustrating. Answer the question at the end of each Key Point
students. These tools, described in the Resources for Students figure and at the end of each Key Experiment.
section of the Preface, are included in MindTap available at Read each chapter section actively. Highlighting and un-
www.cengagebrain.com. In addition to these learning strate- derlining are not always active learning techniques; sometimes
gies, you can make the task of learning biology easier by using they postpone learning. (“This part is important; I’ll learn it
approaches that have been successful for a broad range of our later.”) An active learner always has questions in mind and is
students over the years. constantly making connections. For example, there are many
processes that must be understood in biology. Don’t try to
blindly memorize them; instead, think about causes and ef-
Be Open to Many Learning Styles fects so that every process becomes a story. Eventually, you’ll
There is a popular belief that each person has an innate “learn- see that many processes are connected by common elements.
ing style” that is most successful for them. In fact, there is very To master the material, you will probably have to read each
little scientific evidence to support this view. What works will chapter more than once. Each time will be much easier than
depend on the nature of the material being learned, and in the previous time because you’ll be reinforcing concepts that
most cases a mix of activities and a variety of sensory inputs you have already partially learned.
will be most effective. Biology includes many kinds of questions Write a chapter outline and flesh out your outline by add-
to encourage you to think and learn in different ways. Make ing important concepts and boldface terms with definitions in
learning a part of your life as you think, listen, draw, write, your own words (not copied from the book or cut and pasted).
argue, describe, speak, observe, explain, and experiment. Use this outline when preparing for the exam.
Now it is time to test yourself. Answer the Test Your
­Understanding questions (Know and Comprehend, Apply and
Know Your Professor’s Expectations ­Analyze, and Evaluate and Synthesize) at the end of the c­ hapter.
Determine what your professor wants you to know and how You will sharpen your thinking if you take the time to type
your learning will be assessed. Some professors test almost or write out your answers. The answers are in Appendix E
exclusively on material covered in lecture. Others rely on their ­online at www.cengagebrain.com, but do not be too quick to
students’ learning most of, or even all, the content assigned check them. Think about them and discuss them with your
in chapters. Find out what your professor’s requirements are fellow students if possible. Consider each question as a kind of
because the way you study will vary accordingly. springboard that leads to other questions. Finally, review the
If lectures are the main source of examination questions, Learning ­Objectives in the Summary and try to answer them
make your lecture notes as complete and organized as possible. before reading the summary provided.
Before going to class, skim over the chapter, identifying key
terms and examining the main figures, so that you can take
effective lecture notes. Spend no more than 1 hour on this.
Learn the Vocabulary
Within 24 hours after class, type (or rewrite) your notes. Before One stumbling block for many students is learning the many
typing them, however, read the notes and make marginal notes terms that make up the language of biology. In fact, it would
about anything that is not clear. Then read the corresponding be much more difficult to learn and communicate if we did

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
not have this terminology because words are really tools for covered in Chapter 8. Before presenting all the details, we
thinking. Learning terminology generally becomes easier if provide an overview figure that emphasizes what the process
you realize that most biological terms are modular. They con- accomplishes.
sist of mostly Latin and Greek roots; once you learn many
of these roots, you will have a good idea of the meaning of a
new word even before it is defined. For this reason, we have Form a Study Group
included Appendix C, Understanding Biological Terms, online Active learning is facilitated if you do some of your studying
at www.cengagebrain.com. To be sure that you understand collaboratively in a small group. In a study group, the roles of
the precise definition of a term, use the Index and the ­Glossary. teacher and learner can be interchanged: a good way to learn
The more you use biological terms in speech and writing, the material is to teach, through a process that cognitive scien-
more comfortable you will be with the language of biology. tists describe as elaborative rehearsal (not to be confused with
memorization). A study group has other advantages: it can
Develop a Framework for Your make learning more fun, lets you meet challenges in a non-
threatening environment, and can provide some emotional
Learning support. When combined with individual study of text and
Always aim to get the big picture before adding details. lecture notes, study groups can be effective learning tools.
When attempting to learn a complex process, a struggling Our author team hopes that your study of biology will be
student will typically begin with the first part, try to learn an exciting journey for you, as it continues to be for us.
all the details, and then give up. Instead, begin by making
sure that you have a basic understanding of what is happen- Eldra P. Solomon
ing in the overall process. To encourage you in this way of Charles E. Martin
thinking, we have modeled this approach in Biology. As just Diana W. Martin
one example out of many, glycolysis is a multistep process Linda R. Berg

To the Student / xxxi

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-202
Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s).
Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
1 A View of Life

This is a very exciting time to study biology, the science of


life. Biologists are making remarkable new discoveries that
affect every aspect of our lives, including our health, food, key concepts
safety, relationships with humans and other organisms, and 1.1 Basic themes of biology include evolution, interactions
the environment of our planet. New knowledge provides of biological systems, inter-relationships of structure and
new insights into the human species and the millions of other function, information transfer, and energy transfer.
organisms with which we share planet Earth. Biology affects 1.2 Characteristics of life include cellular structure, growth
our personal, governmental, and societal decisions. and development, self-regulated metabolism, response to
One of the most exciting areas of current research is genetic stimuli, and reproduction.
engineering, specifically CRISPR, a breakthrough technology 1.3 Biological organization is hierarchical and includes
chemical, cell, tissue, organ, organ system, and organism
that allows researchers to edit genes. CRISPR (clustered regularly
levels; ecological organization includes population,
interspaced short palindromic repeats) are sections of prokaryotic
community, ecosystem, and biosphere levels.
(archaeal and bacterial) DNA that have short repeating base 1.4 Information transfer includes DNA transfer of information
sequences. Near each CRISPR there are groups of cas genes. The from one generation to the next, chemical and electrical
CRISPR/Cas system protects prokaryotes from foreign genetic signals within and among the cells of every organism,
material (which is carried and inserted into their genetic material and sensory receptors and response systems that allow
by plasmids and phages). RNA in nearby spacer sequences works organisms to communicate with one another and interact
with Cas proteins to recognize and remove the foreign DNA with their environment.
or RNA. 1.5 Individual organisms and entire ecosystems depend on a
continuous input of energy. Energy is transferred within
cells and from one organism to another.
photo: CRISPR editing a genome (genetic material in a cell). 1.6 Evolution is the process by which populations of organisms
This molecular model of gene editing using CRISPR (clustered change over time, adapting to changes in their environment;
regularly interspaced short palindromic repeats) shows a Cas9 protein the tree of life includes three major branches, or domains.
(purple) attached to the DNA of a cell using a guide RNA (orange) 1.7 Biologists ask questions, develop hypotheses, make
that matches a target DNA sequence (light blue). The Cas9 protein predictions, and collect data by careful observation and
separates the DNA strands of the double helix. The green area of the by performing experiments; based on their results, they
DNA identifies the location where the Cas9 protein is attached and come to conclusions and then share their work with other
“cuts” the target DNA, changing the gene sequence. Evan Oto/Science Source scientists and with the public.
1

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Researchers are now using CRISPR as a tool for editing genes blood cells, which function to transport oxygen. Similarly,
rapidly and inexpensively (see image). By “knocking out” specific on the level of organisms, the canine teeth of carnivorous
genes, scientists discover the function of these genes. CRISPR mammals are adapted for stabbing their prey and ripping
identifies the specific region of DNA that has been targeted. It flesh. In contrast, horses and other herbivorous mammals
then cuts out and removes the gene of interest. You will learn have teeth adapted for cutting off bits of vegetation and
more about CRISPR when you study genetics later in this book. grinding plant material. In each case, structure and func-
CRISPR/Cas9 technology has been used to successfully tion are inter-related.
deactivate specific human genes, modify yeasts for biofuel 3. Information must be transmitted within organisms and
production, and modify agricultural crops. Scientists are among organisms. Each organism must be able to receive
researching how CRISPR technology could produce mutant information from the surrounding environment. The
model stem cell lines for studying disease, eliminate proteins survival and function of every cell and every organism
that cause rejection in organ transplants, and selectively cut out depend on the orderly transmission of information. As
or alter any targeted human gene. CRISPR research will almost we will learn, evolution depends on the transmission of
certainly lead to more effective ways of preventing and treating genetic information from one generation to another.
cancer and HIV, and of dramatically decreasing the spread of 4. Life depends on a continuous input of energy from the
mosquito-transmitted diseases, such as Zika and malaria. sun because every activity of a living cell or organism
CRISPR technology is just one of hundreds of exciting areas of requires energy. Energy from the sun flows through indi-
biological research that bring together science, technology, and vidual organisms and through ecosystems. Within living
society. Whatever your college major or career goals, knowledge of cells energy is continuously transferred from one chemi-
biological concepts is a vital tool for understanding our world and cal compound to another.
for meeting many of the personal, societal, and global challenges 5. Evolution is the process by which populations of organ-
that confront us. Among these challenges are the expanding isms change over time. Scientists have accumulated a
human population, decreasing biological diversity, diminishing wealth of evidence showing that the diverse life-forms on
natural resources, global climate change, and prevention and cure this planet are related and that populations have evolved—
of diseases, such as heart disease, cancer, diabetes, and Alzheimer’s that is, have changed over time—from earlier forms of life.
disease. Meeting these challenges will require the combined efforts The process of evolution is the framework for the science
of biologists and other scientists, health professionals, educators, of biology and is a major theme of this book.
politicians, and biologically informed citizens.
This book is a starting point for your exploration of biology.
The interaction of biological systems, the inter-relationship of
It will provide you with the basic knowledge and the tools to
structure and function, information transfer, energy transfer,
become a part of this fascinating science as well as a more
and the process of evolution are forces that give life its unique
informed member of society.
characteristics. You will find reference to one or more of these
unifying themes in every chapter of Biology. We begin our
study of biology by developing a more precise understanding
1.1 Major Themes of Biology of the fundamental characteristics of living systems and of the
levels of biological organization. We then take a closer look at
learning objective some of the major themes of biology. We end Chapter 1 with
a discussion of the process of science.
1 Describe five basic themes of biology.

In this first chapter we introduce five major themes of biology.


These themes are interconnected with one another and with checkpoint 1.1
almost every concept that we discuss in this book. • Why are information transmission, energy transfer, and
evolution considered basic to life?
1. Biological systems interact. Every organism is a biological
system made up of millions of other biological systems. Each • connect What are some ways in which an organism
of its cells is a biological system, as is each organ (e.g., heart is dependent on other biological systems?
and liver) and body system (e.g., cardiovascular system and
digestive system). Each of the multitude of microorgan-
isms (e.g., bacteria) that inhabit an organism is also a bio-
logical system. Making this concept even more interesting, 1.2 Characteristics of Life
an organism cannot survive on its own. Every organism is
learning objective
a biological system that is interdependent with many other
biological systems. Clearly, scientists can study biological 2 Distinguish between living systems and nonliving things by
systems and their interactions at many different levels. describing the features that characterize living organisms.
2. Structure and function are interrelated in all biological
systems. The structure of neurons that function to trans- We easily recognize that a pine tree, a butterfly, and a horse are
mit information is very different from the structure of red living systems, whereas a rock is not. Despite their diversity,
2 / Chapter 1

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the organisms that inhabit our planet share a common set of
characteristics that distinguish them from nonliving things.
These features include a precise kind of organization, growth
and development, self-regulated metabolism, the ability to

Wim van Egmond/Visuals Unlimited/Getty Images


respond to stimuli, reproduction, and adaptation to environ-
mental change.

Organisms are composed of cells


Although they vary greatly in size and appearance, all organ-
isms consist of basic units called cells. New cells are formed
only by the division of previously existing cells. As will be dis-
cussed in Chapter 4, these concepts are expressed in the cell
theory, another fundamental unifying concept of biology. 250 mm
Some of the simplest life-forms, such as protozoa, are (a) Unicellular organisms consist of one cell that performs all the
unicellular organisms, meaning that each consists of a single functions essential to life. Ciliates, such as this Paramecium, move
cell (FIG. 1-1a). In contrast, the body of a maple tree or a buf- about by beating their hairlike cilia.
falo is made of billions of cells (FIG. 1-1b). In such complex
multicellular organisms, life processes depend on the coordi-
nated functions of component cells that are organized to form
tissues, organs, and organ systems.
Every cell is enveloped by a protective plasma membrane
that separates it from the surrounding external environment.
The plasma membrane regulates passage of materials between
the cell and its environment. Cells have specialized molecules—
deoxyribonucleic acid, more simply known as DNA—that
contain genetic instructions and transmit genetic information.
Cells typically have internal structures called organelles that
are specialized to perform specific functions. For example,
mitochondria, which we can think of as the power plants of the
cell, convert energy in food molecules into energy forms that
can be more conveniently used by the cell.
There are two fundamentally different types of cells: pro-
karyotic and eukaryotic. Prokaryotic cells are microscopic

McMurray Photography
organisms classified in two large groups (called domains):
domain Bacteria and domain Archaea. Prokaryotic cells do
not have a nucleus or other membrane-enclosed organelles.
All other organisms are characterized by their eukaryotic
(b) Multicellular organisms, such as this African buffalo (Syncerus
cells. These cells typically contain a variety of organelles caffer) and the plants on which it grazes, may consist of billions of
enclosed by membranes, including a nucleus, which contains cells specialized to perform specific functions.
DNA, the genetic information.
Figure 1-1 Unicellular and multicellular life-forms
Organisms grow and develop
Biological growth involves an increase in the size of individ- to the functions the organism must perform. Like many other
ual cells of an organism, in the number of cells, or in both. organisms, every human begins life as a fertilized egg that
Growth may be uniform in the various parts of an organism, or then grows and develops.
it may be greater in some parts than in others, causing the body
proportions to change as growth occurs. Some organisms— Organisms regulate their metabolic
most trees, for example—continue to grow throughout their
lives. Many animals have a defined growth period that termi-
processes
nates when a characteristic adult size is reached. An intriguing Within all organisms, chemical reactions and energy transforma-
aspect of the growth process is that each part of the organism tions occur that are essential to nutrition, the growth and repair of
typically continues to function as it grows. cells, and the conversion of energy into usable forms. The sum
Organisms develop as well as grow. Development includes of all the chemical activities of the organism is its metabolism.
all the changes that take place during an organism’s life. The Metabolic processes occur continuously in every organism,
structures and body form that develop are exquisitely adapted and they must be carefully regulated to maintain homeostasis,
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an appropriate, balanced internal environment. The term
homeostasis also refers to the automatic tendency of the organ-
ism to maintain a steady state. For example, when a particu-
lar substance is required, cell processes that produce it must
be turned on. When enough of a cell product has been made,
its manufacture must be decreased or turned off. These
homeostatic mechanisms are self-regulating control systems
that are remarkably sensitive and efficient.
The regulation of glucose (a simple sugar) concentration Flagella
in the blood of complex animals is a good example of a homeo-
static mechanism. Your cells require a constant supply of glu-
cose molecules, which they break down to obtain energy. The
circulatory system delivers glucose and other nutrients to all
the cells. When the concentration of glucose in the blood rises

CAMR/A. Barry Dowsett/Science Source


above normal limits, glucose is stored in the liver and in mus-
cle cells. When you have not eaten for a few hours, the glucose
concentration begins to fall. Your body mobilizes stored glu-
cose. If necessary, the body converts other stored nutrients to
glucose, bringing the glucose concentration in the blood back
to normal levels. When the glucose concentration decreases,
you also feel hungry and can restore nutrients by eating.
1 mm
Figure 1-2 Biological movement
Organisms respond to stimuli These bacteria (Helicobacter pylori), equipped with flagella for
locomotion, have been linked to stomach ulcers. The photograph
All forms of life respond to stimuli, physical or chemical was taken using a scanning electron microscope. The bacteria are
changes in their internal or external environment. Stimuli that not really red and blue. Their color has been artificially enhanced.
evoke a response in most organisms are changes in the color,
intensity, or direction of light; changes in temperature, pres-
sure, or sound; and changes in the chemical composition of specialized cells that respond to specific types of stimuli. For
the surrounding soil, air, or water. Responding to stimuli often example, cells in the retina of the vertebrate eye respond to light.
involves movement, although not always locomotion (moving Although their responses may not be as obvious as those
from one place to another). of animals, plants do respond to light, gravity, water, touch,
In simple organisms, the entire individual may be sensitive and other stimuli. For example, plants orient their leaves to
to stimuli. Certain unicellular organisms, for example, respond to the sun and grow toward light. Many plant responses involve
bright light by retreating. In some organisms, locomotion is different growth rates of various parts of the plant body. A
achieved by the slow oozing of the cell, the process of a­ moeboid few plants, such as the Venus flytrap of the Carolina swamps,
movement. Other organisms move by beating tiny, hairlike are very sensitive to touch and catch insects (FIG. 1-3). Their
extensions of the cell called cilia or longer structures known as leaves are hinged along the midrib, and they have a scent
flagella (FIG. 1-2). Some bacteria
move by rotating their flagella.
Most animals move very
obviously. They wiggle, crawl,
swim, run, or fly by contracting
muscles. Sponges, corals, and
oysters have free-swimming lar-
val stages, but most are sessile as
adults, meaning that they do not
Linas T/Shutterstock.com
Linas T/Shutterstock.com

move from place to place. In fact,


they may remain firmly attached
to a surface, such as the sea bot-
tom or a rock. Many sessile
organisms have cilia or flagella
(b) The edges of the leaf come together and
that beat rhythmically, bring- (a) When hairs on the leaf surface of the Venus interlock, preventing the insect’s escape. The
flytrap (Dionaea muscipula) detect the touch
ing them food and oxygen in of an insect, the leaf responds by folding. leaf then secretes enzymes that kill and digest
the surrounding water. Complex the insect.
animals, such as grasshoppers, Figure 1-3 Plants respond to stimuli
lizards, and humans, have highly
4 / Chapter 1

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that attracts insects. Trigger hairs on the leaf surface detect
the arrival of an insect and stimulate the leaf to fold. When the
edges come together, they interlock, preventing the insect’s
escape. The leaf then secretes enzymes that kill and digest the
insect. The Venus flytrap usually grows in nitrogen-deficient
soil. The plant obtains part of the nitrogen required for its
growth from the insects it “eats.”

Cabisco/Visuals Unlimited, Inc.


Organisms reproduce
At one time, people thought worms arose spontaneously from
horsehair in a water trough, maggots from decaying meat,
and frogs from the mud of the Nile. Thanks to the work of 100 mm
a great many scientists, beginning with pioneering studies
by Italian physician Francesco Redi in the 17th century and (a) Asexual reproduction. One individual gives rise to two
French chemist Louis Pasteur in the 19th century, we know or more offspring that are similar to the parent. Difflugia, a
unicellular amoeba, is shown dividing to form two amoebas.
that organisms arise only from previously existing organisms.
Simple organisms, such as amoebas, perpetuate them-
selves by asexual reproduction (FIG. 1-4a). When an amoeba
has grown to a certain size, it reproduces by splitting to form
two new amoebas. Before an amoeba divides, its hereditary
material (set of genes) is duplicated, and one complete set is
distributed to each new cell. Except for size, each new amoeba
is similar to the parent cell. The only way that variation
occurs among asexually reproducing organisms is by genetic
mutation, a permanent change in the genes.
In most plants and animals, sexual reproduction is car-
ried out by the fusion of an egg and a sperm cell to form a

iStock.com/PK6289
fertilized egg (FIG. 1-4b). The new organism develops from
the fertilized egg. Offspring produced by sexual reproduc-
tion are the product of the interaction of various genes con-
tributed by the mother and the father. Genetic variation is (b) Sexual reproduction. Typically, each of two parents
important in the vital processes of evolution and adaptation. contributes a gamete (sperm or egg). Gametes fuse to produce
the offspring, which has a combination of the traits of both
parents. A pair of cucumber beetles is shown mating.
Populations evolve and become
Figure 1-4 Asexual and sexual reproduction
adapted to the environment
The ability of a population to evolve over many generations
and adapt to its environment equips it to survive in a chang-
ing world. Adaptations are inherited characteristics that
enhance an organism’s ability to survive in a particular envi-
ronment. The long, flexible tongue of the frog is an adapta-
tion for catching insects. The feathers and lightweight bones
of birds are adaptations for flying, and their thick fur coats
allow polar bears to survive in frigid temperatures. Adapta-

McMurray Photography
tions may be structural, physiological, biochemical, behav-
ioral, or a combination of all four (FIG. 1-5). Every biologically
successful organism is a complex collection of coordinated
adaptations produced through evolutionary processes.
Figure 1-5 Adaptations
These Burchell’s zebras (Equus burchelli), photographed in Tanzania,
are behaviorally adapted to position themselves to watch for lions
checkpoint 1.2 and other predators. Stripes are thought to be an adaptation for
visual protection against predators. They serve as camouflage or to
• What characteristics distinguish a living organism from a rock? break up form when spotted from a distance. The zebra stomach is
• predict What would be the consequences to an organism adapted for feeding on coarse grass passed over by other grazers,
if its homeostatic mechanisms failed? Explain your answer. an adaptation that helps the animal survive when food is scarce.

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1.3 L
 evels of Biological roots and leaves in plants. In animals, each major group of
biological functions is performed by a coordinated group
Organization of tissues and organs called an organ system. The circulatory
and digestive systems are examples of organ systems. Func-
learning objective tioning together with great precision, organ systems make up
a complex, multicellular organism. Again, emergent proper-
3 Construct a hierarchy of biological organization, including
levels characteristic of individual organisms and levels ties are evident. An organism is much more than its compo-
characteristic of ecological systems. nent organ systems.

Whether we study a single organism or the world of life as a Several levels of ecological
whole, we can identify a hierarchy of biological organization
organization can be identified
(FIG. 1-6). At every level, structure and function are precisely
coordinated. One way to study a particular level is by look- Organisms interact to form still more complex levels of bio-
ing at its components. Biologists can gain insights about cells logical organization. All the members of one species living
by studying atoms and molecules. Learning about a struc- in the same geographic area at the same time make up a
ture by studying its parts is called reductionism. However, population. The populations of various types of organisms
the whole is more than the sum of its parts. Each level has that inhabit a particular area and interact with one another
emergent properties, characteristics not found at lower form a community. A community can consist of hundreds of
levels. For example, populations of organisms have emergent different types of organisms. The human body, for example,
properties such as population density, age structure, and birth harbors a community of hundreds of species of microorgan-
and death rates. The individuals that make up a population isms, its microbiota, that critically affect human health.
do not have these characteristics. Consider also the human A community together with its nonliving environment is
brain. The brain is composed of billions of neurons (nerve an ecosystem. An ecosystem can be as small as a pond (or even
cells). However, we could study every one of these individual a puddle) or as vast as the Great Plains of North America or
neurons and have no clue about the functional capacities of the Arctic tundra. All Earth’s ecosystems together are known
the brain. Only when the neurons interact are the emergent as the biosphere. The biosphere includes all systems of Earth
properties, such as the capacity for thought, judgment, and that are inhabited by living organisms: the atmosphere, the
motor coordination, evident. hydrosphere (water in any form), and the lithosphere (Earth’s
crust). The study of how organisms relate to one another and
Organisms have several levels to their physical environment is called ecology (derived from
the Greek oikos, meaning “house”).
of organization
The chemical level, the most basic level of organization,
includes atoms and molecules. An atom is the smallest unit
of a chemical element that retains the characteristic proper- checkpoint 1.3
ties of that element. For example, an atom of hydrogen is the • What are the levels of organization within an organism?
smallest possible amount of hydrogen. Atoms combine chem- • predict At which level do you think more biological
ically to form molecules. Two atoms of hydrogen combine systems would be interacting: organism, population, or
with one atom of oxygen to form a single molecule of water. ecosystem? Justify your answer.
Although composed of two types of atoms that are gases under
conditions found on Earth, water can exist as a gas, l­iquid, or
solid. The properties of water are very different from those of
its hydrogen and oxygen components, an example of emer- 1.4 Information Transfer
gent properties.
At the cellular level, many types of atoms and molecules learning objective
associate with one another to form cells. However, a cell is 4 Summarize the importance of information transfer within
much more than a heap of atoms and molecules. Its emergent and between living systems, giving specific examples.
properties make it the basic structural and functional unit of
life, the simplest component of living matter that can carry on Biological systems receive and respond to information. They
all the activities necessary for life. also store information in the form of DNA (or RNA). An
During the evolution of multicellular organisms, cells organism inherits the information it needs to grow, develop,
associated to form tissues. For example, most animals have carry on self-regulated metabolism, respond to stimuli, and
muscle tissue and nervous tissue. Plants have epidermis, a tis- reproduce. Each organism must also have precise instructions
sue that serves as a protective covering, and vascular tissues for making the molecules necessary for its cells to communi-
that move materials throughout the plant body. In most com- cate. The information an organism requires to carry on these
plex organisms, tissues organize into functional structures life processes is coded and transmitted in the form of chemi-
called organs, such as the heart and stomach in animals and cal substances and electrical impulses.
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Organism
Organ systems
work together
in a functional
organism.
Population
A population
consists of
organisms of the
Organism same species.

Population

Organ system Organ system


(e.g., skeletal
system) Tissues
and organs make Community
up organ systems. The populations
of different
species that
populate the
Organ Organ
same area make
(e.g., bone) up a community.
Tissues form
organs.

Tissue Community
(e.g., bone tissue)
Cells associate Tissue Bone cells
to form tissues.

Nucleus
Cellular level Cell
Ecosystem
Atoms and molecules
A community
make up the cytoplasm
together with
and form organelles,
the nonliving
such as the nucleus
environment
and mitochondria (the
forms an
site of many energy
ecosystem.
transformations). Organelle
Organelles perform Ecosystem
various functions
of the cell.

Chemical level Macromolecule Biosphere


Atoms join to Earth and all
form molecules. its communities
Biosphere
Macromolecules are constitute the
large molecules such biosphere.
as proteins and DNA. Oxygen atom
Hydrogen atoms
Molecule

Water

Figure 1-6 The hierarchy of biological organization

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DNA transmits information from
one generation to the next
Humans give birth only to human babies, not to giraffes or rose-
bushes. In organisms that reproduce sexually, each offspring is
a combination of the traits of its parents. In 1953, James Watson
and Francis Crick worked out the structure of DNA, the large
molecule that makes up the genes, units of hereditary infor-
mation (FIG. 1-7). A DNA molecule consists of two chains of
atoms twisted into a helix. As will be described in Chapter 3,
each chain is made up of a sequence of chemical subunits called
nucleotides. There are four types of nucleotides in DNA, and
each sequence of three nucleotides is part of the genetic code.
Watson and Crick’s work, building on the earlier findings of
many researchers, led to the understanding of the genetic code.
The information coded in sequences of nucleotides in DNA
transmits genetic information from generation to generation. The
code works somewhat like an alphabet of 4 letters (A, G, C, T).
The nucleotides can “spell” an amazing variety of instructions
for making organisms as diverse as bacteria, frogs, and redwood
trees. The genetic code is universal—that is, virtually identical
in all organisms—and is a dramatic example of the unity of life.

Information is transmitted by chemical


and electrical signals
Genes control the development and functioning of every
organism. As you will learn in later chapters, the information
carried by the DNA that makes up the genes has many func-
tions, including providing the “recipes” for making all the pro-
teins required by the organism. Proteins are large molecules
important in determining the structure and function of cells
and tissues. For example, brain cells differ from muscle cells in
large part because they have different types of proteins. Some
proteins are important in communication within and among
cells. Certain proteins on the surface of a cell serve as markers
so that other cells “recognize” them. Other cell-surface proteins Figure 1-7 DNA
serve as receptors that combine with chemical messengers. DNA is the hereditary material that transmits information from
Cells use proteins and many other types of molecules one generation to the next. As shown in this model, DNA is a
to communicate with one another. In a multicellular organ- macromolecule that consists of two chains of atoms twisted into
a helix. Each chain consists of subunits called nucleotides. The
ism, cells produce chemical compounds, such as hormones, sequence of nucleotides makes up the genetic code.
that signal other cells. Hormones and other chemical mes-
sengers can signal cells in distant organs to secrete a particu-
lar required substance or change some metabolic activity. In compounds known as neurotransmitters. Information trans-
this way chemical signals help regulate growth, development, mitted from one part of the body to another is important in reg-
and metabolic processes. The mechanisms involved in cell ulating life processes. In complex animals, the nervous system
signaling often involve complex biochemical processes. gives the animal information about its outside environment by
Cell signaling is currently an area of intense research. transmitting signals from sensory receptors such as the eyes
A major focus has been the transfer of information among cells and ears to the brain. Neurons in the brain make appropriate
of the immune system. A better understanding of how cells decisions and transmit signals to specific muscles or glands.
communicate promises new insights into how the body pro-
tects itself against disease organisms. Learning to manipulate Organisms also communicate
cell signaling may lead to new methods of delivering drugs
into cells and new treatments for cancer and other diseases.
information to one another
Many organisms use electrical signals to transmit infor- Organisms communicate information to other organisms by
mation. Most animals have nervous systems that transmit releasing chemicals, sounds, and visual displays. Typically, organ-
information by way of both electrical impulses and chemical isms use a combination of several types of communication
8 / Chapter 1

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Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
signals. A dog may signal aggression by growling, using a par-
ticular facial expression, and laying its ears back. Many animals

Russ Witherington/Shutterstock.com
Light
perform complex courtship rituals in which they display parts of energy
their bodies, often elaborately decorated, to attract a mate.
Seaweed algae compete with coral for light and space.
Marine biologists studying endangered coral reefs have discov-
ered that certain seaweed algae secrete chemical compounds
that kill coral. Researchers have reported that some coral can
Photosynthesis
fight back. When they come into contact with toxic seaweed, captures light energy. Oxygen
the coral release chemical compounds that signal certain spe-
cies of goby fish. In response to this chemical signal, the fish eat
the seaweed. This action helps preserve their coral reef habitat.

Energy stored in glucose


and other nutrients
checkpoint 1.4 Carbon dioxide
and water
• What is the function of DNA? Oxygen
• How does a nervous system transmit information?
Energy

1.5 The Energy of Life


Cellular respiration in
learning objective plants and animals
releases energy stored
in glucose molecules.
5 Summarize the flow of energy through ecosystems
and contrast the roles of producers, consumers, and
decomposers.
Synthesis Life Activities
of needed • Homeostasis
The sun provides most of the energy that powers life on molecules • Growth and development

BMJ/Shutterstock.com
Earth. All life processes, including thousands of chemi- and structures • Reproduction
cal transactions that maintain life’s organization, require • Movement of materials
a continuous input of energy. Organisms can neither in and out of cells
• Movement of body
create energy nor use it with complete efficiency. During
every energy transaction, some energy is converted to Figure 1-8 Energy flow within and among organisms
heat and dispersed into the environment. Energy flows Algae and certain plant cells carry on photosynthesis, a process that uses light
through individual organisms and through ecosystems. energy to produce glucose from carbon dioxide and water. Energy is stored in the
A self-sufficient ecosystem consists of a physical chemical bonds of glucose and other nutrients produced from glucose. Through
environment inhabited by three types of organisms: the process of cellular respiration, cells of all organisms, including algae and plant
cells, then break down glucose and other nutrients. The energy released can be
producers, consumers, and decomposers. These organ- used to produce needed molecules and to fuel other life activities.
isms depend on one another and on the environment
for nutrients, energy, oxygen, and carbon dioxide.
Plants, algae, and certain bacteria and archaea are producers, Recall that all of the energy transformations and chemical
or autotrophs, organisms that use simple raw materials (inor- processes that occur within an organism are referred to as its
ganic compounds) to produce organic compounds (which metabolism. Energy is necessary to carry on the metabolic activi-
may be used for food). Most of these organisms carry on ties essential for growth, repair, and maintenance. Each cell of an
photosynthesis, the process during which autotrophs use organism requires nutrients that contain energy. During cellular
carbon dioxide, water, and light energy to synthesize complex respiration, cells capture energy stored in glucose and other
molecules such as glucose and other sugars (FIG. 1-8): nutrient molecules through a series of carefully regulated chemi-
cal reactions. We can summarize these reactions as follows:
carbon dioxide + water + light energy
glucose + oxygen glucose + oxygen carbon dioxide + water + energy

The light energy is transformed into chemical energy, which When chemical bonds are broken during cellular respira-
is stored within the chemical bonds of the glucose and other tion, their stored energy is made available for life processes.
food molecules produced. Oxygen, which is required by the Cells use this energy to do work, including the synthesis of
cells of most organisms including plant cells, is produced as a required materials, such as new cell components. Virtually all
byproduct of photosynthesis. cells carry on cellular respiration.
A View of Life / 9

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Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s).
Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it.
Key Point (Note that producers also carry on cellular respiration.) The
metabolism of consumers and producers helps maintain the
Energy flows from the sun to producers life-sustaining mixture of gases in the atmosphere.
Most bacteria, many archaea, and fungi are decomposers,
and then to consumers and decomposers.
heterotrophs that obtain nutrients by breaking down nonliv-
ing organic material such as wastes, dead leaves and branches,
and the bodies of dead organisms. In their process of obtaining
Light energy, decomposers make the components of these materi-
Heat energy
als available for reuse. If decomposers did not exist, nutrients
Heat Heat would remain locked up in wastes and dead bodies, and the
supply of elements required by living systems would soon be
exhausted.

Food

checkpoint 1.5
• predict What components do you think a forest
ecosystem might include?
Producer Primary
• connect In what ways do consumers depend
(plant) consumer on producers? on decomposers? Include energy
(caterpillar) considerations in your answer.
Secondary
consumer Dr. Morley Read/Shutterstock.com/Robert McGouey/Getty Images
(bird)
1.6 Evolution: The Basic
Unifying Concept of Biology
learning objectives
Plant litter, Dead
wastes bodies
6 Demonstrate the binomial system of nomenclature by
using specific examples and classify an organism (such as
Decomposers a human) in its domain, kingdom, phylum, class, order,
Soil (bacteria, fungi) family, genus, and species.
7 Identify the three domains and the kingdoms of living
organisms, and give examples of organisms assigned to
Figure 1-9 Energy flow through the biosphere each group.
Continuous energy input from the sun operates the biosphere. 8 Give a brief overview of the scientific theory of evolution and
During photosynthesis, producers use the energy from sunlight to explain why it is the principal unifying concept in biology.
make complex molecules from carbon dioxide and water. Primary
consumers, such as the caterpillar shown here, obtain energy, 9 Apply the concept of natural selection to any given
nutrients, and other required materials when they eat producers. adaptation and suggest a logical explanation of how
Secondary consumers, such as the bird, obtain energy, nutrients, the adaptation may have evolved.
and other required materials when they eat primary consumers
that have eaten producers. Decomposers obtain their energy and Evolution is the process by which populations of organ-
nutrients by breaking down wastes and dead organic material. isms change over time. The scientific theory of evolution
During every energy transaction, some energy is lost to biological has become the most important unifying concept of biol-
systems, dispersing into the environment as heat.
ogy. As we will discuss, evolution involves passing genes for
predict How does air pollution caused by human activity affect new traits from one generation to another, leading to differ-
the balance of energy flow through the biosphere? ences in populations. The evolutionary perspective is impor-
tant in every specialized field within biology. Biologists try to
understand the structure, function, and behavior of organ-
Animals are consumers, or heterotrophs—that is, they isms and their interactions with one another by considering
are organisms that depend on producers for food, energy, them in the context of the long, continuing process of evolu-
and oxygen (FIG. 1-9). Primary consumers eat producers. tion. Although we discuss evolution in depth in Chapters 18
Secondary consumers eat primary consumers. Consumers through 22, we present a brief overview here to give you the
obtain energy by breaking down sugars and other nutrients background necessary to understand other aspects of biology.
originally produced during photosynthesis. Consumers also First, we examine how biologists organize the millions of
contribute to the balance of the ecosystem. For example, organisms that have evolved, and then we summarize some
consumers produce carbon dioxide required by producers. of the mechanisms that drive evolution.
10 / Chapter 1

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doubtless instructed by the cook, that he received unceremonious
dismissal. The road could not be hard to find in any case, so we set
out, leaving the knight of the toasting-fork to follow at his leisure.
Soon he thought better of it, and took his place again at the head of
the procession. In the lower reaches of the valley we saw further
evidence of Circassian industry, in the rich crops that waved by the
wayside. A short distance east of our route lies a small village, which
takes its name from the saint whose tomb stands there—Neby Hûd,
renowned in ancient Arab story.
We took the more easterly of two possible roads—the longer, but
also the easier for the animals. The greater variety and beauty of the
scenery repay the extra travel. The descent into the Jabbok valley
winds down a narrow ravine, turning sharply round jutting crags, and,
in parts, almost precipitous. Oaks and thorns clung to the steeps;
luxuriant vegetation covered the ground. The fertile soil of the valley
supported a fine crop of wheat. The line of the river could be traced
by a winding glory of oleander bloom, overtopped by tall, gracefully-
bending papyrus reeds, whose heavy heads swayed in the breeze.
Reaching the “brook,” we found the bed more than half dry, but even
thus the water took the horses above the saddle-girths. With a short
struggle, we all landed safely on the other side. During the winter
months this must be a perfectly impassable torrent.
On the farther bank we sat to rest and lunch. The horses too
refreshed themselves before facing the steep mountain in front. We
gathered bunches of papyrus heads—an operation requiring both
care and skill, as we found the undergrowth bound together with
trailing brambles, furnished with the sharpest of prickles. Two square
towers stand one at each end of the meadow in which we halted.
They have not the appearance of great antiquity. There is no
entrance to their interior, and their use we were unable to discover.
Climbing the mountains south of the Jabbok, or Zerka—“the blue”
river, as it is now called—was the hardest work our horses had to
face. The track was narrow, and the foothold often extremely
precarious, especially over rocky parts where a slip would have
meant a fall of hundreds of feet. What a tremendous gorge that
Jabbok is! It literally cleaves the country in twain.
GORGE OF THE JABBOK
Now we were within the borders of the modern province of el-Belkâ,
of which es-Salt is the principal—indeed, the only—city. This lies in
the land of the ancient Ammonites. These cool, breezy uplands,
beautifully diversified with wooded knoll and pleasant vale, in which
may be heard the murmur of flowing water most of the year, offer a
rich return to the hand of the enterprising and diligent cultivator. But
whence is he to come? Numerous are the flocks and herds that
browse on the grassy slopes, find shelter in the shady woods, and
drink from the oleander-fringed streams in the vales. But no one who
sees it can for a moment suppose that this rich soil is designed
simply for the support of sheep and oxen. Those who hope for the
return of Israel to the land of their fathers should turn their eyes
rather to this rich and empty land than to the more populous and less
kindly country west of Jordan.
CHAPTER XII
“Time is money”—Rumamain—Priestly hospitality—Fair
mountain groves—Es-Salt—The springs—Relation to
Arabs—Raisins—Descent to the Jordan—Distant view of
Jerusalem—View of the river, the plains of Jordan, the
Dead Sea, and the mountains beyond—The bridge—The
“publican’s” shed—The men from Kerâk.
Causes for delay are never far to seek among Orientals. “Time is
money” is a phrase void of meaning in Arab ears. Money is precisely
the thing he lacks most, while of time he has more than abundance.
An Eastern in a hurry is one of the rarest sights. We were still on the
uplands, far from our destination, when the sun began to throw his
evening glories over the western hills. Our cook thought fit to profess
that he had lost the way—this doubtless to pay us out for our refusal
of a guide. His manner, however, was much too cool and collected,
so we were not deceived. But it was annoying, as the whole caravan
drew up, to see him comfortably seated among the bushes, on the
top of a huge precipice, enjoying a cigarette. We moved rapidly
forward, and fortunately found a wandering Bedawy who, for a
consideration, agreed to conduct us to es-Salt. He led us by a steep
pathway to the bottom of a sweet valley. Then suddenly we plunged
into a romantic ravine, down which dashed a brawling stream,
sprinkling rock and bush with sparkling diamonds. A stiff climb up the
farther bank brought us to the little village of Rumamain, just as the
light departed. Our tents were pitched by candle-light. The villagers,
who are Christians, were most cordial in their welcome, and
hastened to furnish whatever we required, as far as it was in their
power. The priest invited our party to take refreshments with him,
and those who could be spared from the camp gladly accepted his
hospitality. He well maintained the eastern traditions in the
entertainment of strangers, although some of his beverages were
stronger than the desert law prescribes!
RUMAMAIN
The village stands on the edge of the gorge through which we had
passed. The mountains tower aloft on either side. The valley narrows
southward, but to the north it opens out into a broad, fertile expanse,
bounded by the mountains, torn with water-courses, which form the
southern bank of the gorge of the Jabbok.
Before daybreak we were all astir. Waiting only for a supply of
beautiful milk, which was brought us by the Arabs in the mountains,
we got to horse again. Our guide shouldered his club and marched
off towards the thickets that hung, shaggy and dark, on the sides of
the valley. Our path wound among these delightful groves almost to
the confines of es-Salt. About three hours sufficed to bring us to the
city, but the ride was one not soon to be forgotten. It was one of the
most enjoyable parts of our whole journey. Thick oaks and thorns
gathered in the bottom of the hollows; honeysuckle, entwining their
gnarled limbs, shed perfume on the air; the hillsides were clad with
trees of richly varied foliage, while tall pines swayed gracefully high
over all. The morning was fresh and beautiful. Even the horses
seemed to feel the inspiration of surroundings, and footed it merrily
along. It seemed all too soon when we reached the edge of the
forest, and looked forth on the treeless hills beyond. Many of these
are, however, covered with vineyards, whose sprouting green
relieved the dull monotony. Tree-clad they too once were, as we can
see from the numerous stumps in the fields. A few of the forest
patriarchs, left here and there in the cultivated ground, would have
done much to beautify the land, and would have yielded grateful
shade to the labourers. But it is difficult to restrain the axe when it is
once set in motion. Here we found evidences of genuine industry.
Wherever it is possible, vines are planted and carefully tended, so
that the face of the country assumes quite a cheerful and prosperous
aspect.
The city of es-Salt, lying as it does on the steep slope of a valley, is
not seen until one is fairly upon it. The name es-Salt is evidently
derived from salton hieraticon—“the sacred forest.” The inhabitants
may number in all some seven thousand, of whom the great majority
are Moslems. With the Christians, however, they continue to dwell
together in harmony. There may be about two thousand Christians,
taking Greeks, Latins, and Protestants all together. The last belong
to the church founded by the Church Missionary Society. It was our
good fortune to meet with the Syrian clergyman of this congregation,
Kassîs Khalîl Jamal—a gentleman whose praise is in all the
churches. With his counsel, we were persuaded to stay here for the
day, instead of pushing straight on to Jordan, as we had originally
intended. In the valley under the city are olive groves, where the
company, dismounting, sat down for lunch, under shadow of the
trees, with the sound of running water in their ears. Our tents we
pitched on the top of the hill overlooking the town, separated by a
narrow valley from that on which the ruins of the old castle stand.
Having seen all right about the camp, some of us set out, gun in
hand, tempted by the numerous partridges, and enjoyed an excellent
opportunity to see the surroundings of es-Salt. There are few
remains of antiquity, and these not of great interest. Traces of old
graves, found along the hill-faces, and the bare ruins of the old
castle are the chief. The springs, to which the town owes so much of
its life, are, of course, highly prized. The town itself is interesting as
being the chief mercantile centre in all the district east of Jordan
through which we travelled. The market is frequented by the Beduw
from far and near, and everything necessary for their poor life is
found exposed for sale in the streets. Hither the “housewives” bring
their samn, jibn (clarified butter and cheese), skins, and other
products of the wilds, and carry off in return the cloth of which their
scanty clothing is made, coffee, tobacco, etc. Es-Salt thus forms an
excellent basis from which to reach the Arab tribes in these parts.
The advantages it offers are utilised, as far as possible, by the
missionaries there, and, with the help of the medical department of
the mission, they have found considerable entrance; but, in order to
overtake the work in a manner at all satisfactory, men would have to
be set apart to devote all their time to evangelising the Arabs. With a
well-manned station here and another at Bozrah, nearly all, if indeed
not all, the tribes that touch the eastern borders of Palestine might
be reached; and in due time evangelists from among themselves
would go forth with the glad tidings into the inhospitable wastes
beyond.
We were assured that the grapes grown in the district are unsuitable
for the making of wine. However that may be, wine is not made, but
a great business is carried on in raisins, those of es-Salt being
famed throughout the whole country. The Jewish merchants of
Tiberias buy large quantities of them and use them to produce ’arak
—a distilled spirit which is working havoc among the youth of
western Palestine, Moslems as well as others falling a prey to its
seductive influences, although all use of intoxicants is for them under
religious ban.
ES-SALT, THE FOUNTAIN
Our stores were replenished from the market, two days’ provision
only being required, as in that time we hoped to reach Jerusalem.
Early next morning all was packed up and ready for the descent to
Jericho. Several of our party were already in the saddle, when the
horse of one who had been assisting the ladies to mount thought fit
to bolt, and, in their wild efforts to catch him, the European horsemen
may have given the Arabs some new ideas in horsemanship. In any
case, we gained a more intimate acquaintance with the nearer
surroundings of the city than would have been possible otherwise.
When at last the runaway was captured, the main part of our
caravan had already disappeared some distance in front; and it may
give some idea of the crookedness and irregularity of the road to say
that we did not catch a glimpse of them again until we were almost
upon them, on reaching the plain to the north of the Dead Sea. It is a
descent of over four thousand feet in about fifteen miles. The road
turns abruptly now to one hand, now to the other, adapting itself to
the possibilities of the rough, rocky surface, plunging into ravines,
and anon emerging on grassy tracts; but downward, ever downward,
is its course. The wild birds here are evidently little used to be
disturbed by man. Even the timid partridge sat quite close, or nimbly
ran along the rocks on either side of the path. We had not left es-Salt
half-an-hour when, from an eminence commanding a wide prospect,
we saw the high tower that crowns the Mount of Olives in the far
distance, and thus caught the first glimpse of the environs of
Jerusalem. This is one of the most tantalising sights. It seems so
near, and yet hours of toil in the hot sun seem to bring the traveller
no nearer. And when, as the sun sinks, he descends into the valley,
and it is lost to view, it seems as if he had been following some
strange kind of “Will o’ the wisp.” Ere long, too, we obtained a view of
the Dead Sea, lying under a blue haze away below us to the left; and
soon we could trace the course of Jordan through the sandy plain by
the winding breadth of deep green that fills the valley within the
valley in which the river is confined.

JORDAN, SHOWING TERRACES


In that clear, dry atmosphere distances are most deceptive. Seen
through the openings in the hills, one would think that on touching
the plain we should immediately reach the river, but there are miles
of flat, sandy ground to cover ere we pass under the shadow of the
embowering foliage and hear the rush of the waters of Jordan.
Leaving behind us the mountains of Ammon, which form the eastern
boundary of these deep plains, their scarred sides stretching away
into the beetling heights that rise darkly over the Salt Sea, the
prospect before us was one of enchanting interest. We were already
on the borders of the “circle” of Sodom, which charmed the worldly
eye and heart of Abraham’s nephew, when viewed from yonder
mountains to the west, and which was so generously given up by the
aged uncle to the youthful Lot. How strikingly these barren plains
enforce the lessons of that old-world history! How vain the choice of
beauteous pastures, one day ere long to be o’erwhelmed in desert
sand! How infinitely wise the choice of Abraham, the portion of
whose inheritance was the unchanging God Himself! There,
stretching away southward until lost in a blue haze between her
guardian mountains, are the waters of the great sea, which still, in
name, is associated with the unhappy Lot; for the natives call it only
Bahr Lût—“the sea of Lot.” Before us flowed the river whose tide
rolled backward, and over which, while the ark of the covenant stood
in the midst, the great “congregation” of Israel crossed dry-shod.
Yonder lies the site of Gilgal, whence the conquering hosts went
forth under the gallant Joshua. Towards the western border of the
plain we saw great patches of green, over which rose a curl of blue
smoke, marking the position of Erîha, the village which now
represents the city of Jericho, whose walls fell down at the shout of
the armies of Israel. Beyond rose the dark, frowning crags of
Karantal, by tradition identified with the wild scenes of our Lord’s
Temptation.
The Jordan Valley stretched away northward between its mountain
walls almost as far as the eye could reach; the high cone of Karn
Sartabeh rising full two thousand feet above the plain. This last has
been by some identified with the great altar raised by the Eastern
tribes on their return from the conquest of the West. It is in reality an
“altar” of Nature’s raising, and is interesting as one of the signal
stations from which, by means of great fires, intelligence was flashed
over the land when the new moon had been seen in Jerusalem. The
wooden bridge by which we crossed the Jordan was entirely
concealed by the groves around until we were almost upon it. A
substantial structure it looked, made of strong beams, supported by
great posts, all securely fastened together; it seemed as if it would
outlive any ordinary flood, and so, doubtless, it would. But the floods
of the winter 1890-91 were not ordinary. For some months the rains
were excessive. The river rose far above its usual level, submerging
large tracts, and carrying off much that was valuable—among the
rest the bridge, which had done us and others such good service.
The river here is deep and strong, sweeping with great rapidity round
its swift curves. The water is of a thick, brown colour, charged with
the soil over which it passes. Care must be taken in approaching the
treacherous banks of sand and clay. Toll was collected by an
enterprising Syrian, who had erected a wooden shed at the Jericho
end of the bridge. One or two native huts also stood on the little
level, almost surrounded by a bend of the river, and protected to
westward by high sand bluffs.
It was already past mid-day when we arrived, and, as the heat was
terrible, we were thankful to take refuge in the “publican’s” shed. We
were received with every token of welcome by the owner, who at
once busied himself to provide for our refreshment. We found it an
excellent plan to carry with us a supply of tea. It is often easy to get
boiling water when it is hard to get other things. Tea is swiftly made,
and, on a hot journey, is most refreshing. Leban, also, there was in
plenty. After our meal, the more weary of the party, stretched in the
shade, enjoyed a delightful nap,—only for a little, however; for this
being the chief thoroughfare between east and west of Jordan, quiet
could not be secured for long at a time. Shepherds passed over with
their flocks, and guided them down to the water’s edge to drink.
Negotiating with some of the more tractable of these half-wild men,
we secured draughts of delicious milk. Then came Arabs from the
uplands of Moab; strong, stalwart, sallow-featured men; some armed
with the spear, others carrying rifles, with belts stuck full of battered
cartridges. They sat down sociably around the shed, and conversed
freely. They were greatly impressed with the strength of the men of
Kerâk, the ancient Kir Moab. The district was not even nominally
subject to the Ottoman Government. The spirit of wild independence
was abroad among them. While yielding ready obedience to their
own sheyûkh, they resent and battle to the death against any
interference with their tribal liberty. Holding themselves absolute
masters of the soil, they consider themselves entitled to levy
blackmail on all who pass through their territory. This varies in
amount, according to the prudence or the want of it displayed by the
traveller. One who goes with tents, a large retinue of servants, and
luxurious appointments may have to pay some hundreds of pounds
before he escapes their hands; another may shoulder his camera,
ride in on a mule, and with a few rotls of coffee and a judicious
distribution of a few pounds of tobacco and snuff, may march about
with freedom, photograph all of interest in the district, and carry off
his work in safety. In these regions one must avoid every
appearance of wealth. The Turkish Government claims a supremacy
which for long it was unable to assert. Often we heard that the Hajj
guard, returning from Mecca, was to attack and subdue the
Kerâkers. The report that there were four thousand trained men,
armed with repeating rifles and no lack of ammunition, no doubt
restrained the valour of the gallant guard. At length the Turk has
established a certain shadowy authority in the town of Kerâk itself;
but over the Arabs, who occupy the town only a few months in the
year, it is difficult, if not impossible, to exercise any effective control.
The stronghold, the thought of attacking which gave cold shivers to
the Turkish soldiery, could not daunt the high-hearted soldiers of the
Cross. With admirable courage, in the true spirit of Christian
heroism, a missionary and his wife braved all the dangers in the way,
and made for themselves a home in the midst of these people. That
they had dangers not a few to face, and many privations to endure,
needs hardly to be said. But these “things did not move them”; and
the bold warrior Arab learned to love the man of peace, and prize
him as a friend. Who knows whether the Cross may not soon
triumph where the crescent so long struggled for supremacy in vain!
CHAPTER XIII
The banks and thickets of the Jordan—Bathing-place—The
Greek convent—A night of adventures in the plains of
Jericho—The modern village—Ancient fertility—Possible
restoration—Elisha’s fountain—Wady Kelt—The Mountain
of Temptation—The path to Zion.
The Jordan, in the lower reaches, is shaded by overhanging willows,
and the path along the bank is lined with tall oleanders. In the
brushwood, which grows thickly over the little peninsulas formed by
the circlings of the river, we were assured that the nimr (small
leopard) found a lurking-place; and, further, that he and his grim
neighbour, the hyena, haunted the bushy hollows between the sand-
dunes which stretch away towards the sea.
We shouldered our guns, and, armed with ball cartridge, set out to
beat the brushwood in the hope of starting game of this class.
Perhaps it was as well for ourselves that we were disappointed; but
a gun in a man’s hand adds marvellously to his powers of endurance
in walking, so we were able to explore the shady banks of the river,
and attain a fuller knowledge of its windings. The bridge was
photographed with a group in front as a souvenir of our visit; then,
sending the muleteers straight to Jericho with tents and baggage, we
prepared to ride towards the sea, with the intention of returning to
Jericho for the night. Our road wound among the sand-dunes for
some distance. Here we were charged to keep close together. The
advice was necessary. Had one lingered behind, and by any
mischance lost the way, he might have wandered long enough in the
labyrinth formed by these little sand-hills, which resemble each other
so closely as to be distinguished only by the trained eye. They are
the haunts of robbers too, who, in the multitudinous winding hollows,
may easily escape pursuit.
We did not go so far down as the fords and famous bathing-place;
but on a subsequent occasion I saw something of the extraordinary
scenes enacted there—a great company of Russian pilgrims, men,
women, and children, plunging promiscuously into the sacred river.
Most wore a thin linen garment as they went down into the water.
This is afterwards carefully preserved, and is worn again only as a
shroud. Certain men standing in the stream saw that each one went
at least three times over the head.

FORDS OF JORDAN. PILGRIMS BATHING


Leaving the labyrinth, we emerged on a wide sterile plain, over which
grew only a few stunted desert bushes. Here and there we could see
the shimmer of the sunlight on a thin crust of salt. The Greek
convent, standing in the midst of the waste, served as a landmark,
and thither we directed our course, leaving the winding road. This
cross-country riding in these parts has its dangers. The horses are
apt to sink through the soft surface, into holes burrowed by the
rodents; and deep ruts, worn by winter torrents, are often difficult to
pass. At length we stood before the gate of the convent, and the
hospitable monks regaled us with refreshing draughts of pure, cool
well-water. It seemed only a short distance farther to the sea; but
some of our company were growing fatigued, and, at our rate of
riding, it would take almost till sunset to reach our camp by the “City
of Palms”; so, reluctantly, we turned our horses’ heads northward,
comforting ourselves with the hope that in a more convenient season
we should stand on the shores of the Dead Sea and plunge in its
sullen waters.
On arriving, we found the mukaries busy with the tents, which soon
were ready for our reception. But we were disturbed somewhat to
hear that one of our attendants was missing. He ought to have come
with us toward the sea; but when we found he had not come, we
concluded that he had accompanied the muleteers to Jericho. Now,
however, he was nowhere to be found, and the mukaries assured us
he had started immediately behind us. The evil reputation of the
district made us rather anxious; but the reckless character of the
man, and his habit of starting off on the wildest projects without a
moment’s warning, led us to believe that Saʿid would turn up again,
as he had often done before. We set up lights, however, on the most
conspicuous points, when darkness fell, so as to guide him, if
possible, over the plain. As the night advanced, and we still had no
news of him, our anxiety increased, our main hope being that, as he
followed us, he had arrived late at the convent and taken shelter for
the night, or that haply he had found his way back to the bridge.
Even with fifty men it would have been futile to search that
wilderness in the dark. A hunt after a number of wild swine that came
to the neighbourhood of the camp served only as a temporary
diversion from the graver subject occupying our thoughts. The
“garrison” of the town consisted of one soldier, whose services we
secured without much difficulty. He was despatched, with the
breaking light, to go towards Jordan bridge, and southwards to the
convent, while some of our party prepared to scour the plain, the rest
getting in order for the ascent to Jerusalem.
All were very early astir. Just before mounting, the plain was swept
with the telescope, and in the distance a coming horseman was
descried, who seemed to resemble the lost man. His appearance
caused no little excitement in camp; for, on a nearer view, there was
no doubt of his identity. Both man and horse were utterly wearied,
and Saʿid’s garments were covered with blood. The tale he told of
his experiences was terrible enough. On our departure, he lingered a
moment to see the muleteers ready and on the road; then, as he
thought, he followed us, but missing the way, he pushed straight
southward, guided in part by the river, arriving on the Dead Sea
shore just as the sun set. As he had ridden rapidly, he thought he
might have passed us on the way; but, after waiting and careful
search along the lonely beach, he despaired of finding us. He turned
away in the twilight, hoping by instinct to hit upon some way leading
to Jericho. Soon the night was filled with the horrid howlings of the
jackals, all over that weary waste, and here and there the shrill laugh
of the hyena fell on his ear. Pushing forward, he was speedily
entangled among the sand-dunes. Climbing one after another, he
sought to hit upon some landmark to guide him; but ever, on
descending, his bearings were lost, and he wandered almost
hopelessly. As the night closed in, he could see shadowy forms
moving around, and the howlings came nearer and nearer.
Suddenly, in the track before him, he saw a hyena glaring upon him.
The natives believe that the hyena will not attack a man save when
famishing or grievously provoked. But in such moments one does
not reflect much on these things.
Fortunately, Saʿid had my gun and ball cartridge with him. With
nerves steadied by something like despair, he aimed at the brute’s
head, and, the ball crashing through the brain, he rolled over with a
groan. A momentary silence followed the report, and then the dismal
noises broke out again. Riding onward he saw, and not a moment
too soon, a leopard crouching for a spring. Aiming swiftly, he fired,
and the ball took effect in the leopard’s neck; but, withal, the brute
was able to spring, and almost to reach his mark. “Then,” said Saʿid,
his lip quivering and an unaccustomed tear trembling in his eye
—“then I must have had strength given me from heaven; for, as the
brute sprang, I reached forward, caught him by the neck, and hurled
him back violently to the ground, where he lay stunned, and a
second ball finished him.” He attempted to carry the leopard on the
pommel of his saddle, and this accounted for the blood-stains on his
garments. But weariness and anxiety soon overcame his purpose,
and the leopard was left to decay among its native wilds.
Finally, baffled by the darkness and the intricacies of the labyrinthine
windings, he climbed a little knoll, and tying the horse’s halter to his
arm, stretched himself on the ground to sleep; but ever and anon, as
the denizens of the desert drew near, the trembling creature tugged
at the halter to awaken his master, and seemed to long for waking
fellowship. At last he was overcome with sleep, and was only
aroused when the horse had apparently come close up and whinnied
in his ear. Then, rising bolt upright, it seemed to him as if the gloomy
waste were all alive with moving shades, and vocal with dismal
howlings. How he came out of that pandemonium he never could
say; but ere long the dawn stole into the sky, and the wild beasts
crept away into their dens. He found a poor shepherd, walking out
with his flocks, who, seeing his forlorn condition, sustained the best
traditions of desert hospitality by offering what he had for his day’s
food to the weary traveller. Then he was able to make out the
direction of Jericho, and, before the morning was well born, he was
again with his companions, uttering perhaps the most sincere
hamdulillahs of his life. After hearing this tale from the lips of that
worn-out man, on the borders of the desert plain of Jericho, one
could better realise what it meant for the Son of Man when, driven of
the Spirit into the neighbouring wilderness, through the long nights
He was “with the wild beasts.”
JERICHO. ELISHA’S FOUNTAIN
Poor Saʿid has now “joined the majority.” He escaped the beasts of
the Dead Sea plains to fall a victim to a more subtle and deadly foe.
While at work on the new railway near the Hummeh, the hot springs
in Wady Yarmuk, he and many others were mortally stricken with
typhoid. A heavy toll in human lives these valleys have exacted on
the passage of that road of iron. Saʿid was a typical light-hearted
Syrian, prone to err, who yet through years of service proved faithful
according to his light, with a larger infusion of courage in him than is
common among his countrymen. In steadfast loyalty at his master’s
side, neither fatigue nor peril daunted him.
A burden was lifted from all our minds, and, moving forward that
morning, even higher spirits than usual prevailed. We rode into the
village, and stopped a little at the hotel, where souvenirs of the Dead
Sea plain, staffs made from the oaks of Bashan, Bedawy clubs, and
such-like articles, are exposed for sale to travellers. Then we rode
along lanes bounded by hedges of prickly pear and thorns towards
the spring of Elisha, called now ʿAin es-Sultan—“Fountain of the
Sultan,” where man and beast drank from the cool stream and were
refreshed.
The modern Erîha is a miserable representative of the famous cities
that rose one after another in the neighbourhood of this copious
spring. The land immediately around is a marvel of fertility, bearing,
when under cultivation, with tropical luxuriance. Near by the spring
stood the ancient city which was attacked and overthrown by the
Israelites—the first stronghold gained on this side of the Jordan.
Near the same spot rose the city of Heil, who dared the curse and
endured it—the curse pronounced by Joshua—in order to restore the
crown of splendour which his ancestors had crushed so ruthlessly.
This latter was the city known to Elijah and Elisha, not yet ancient in
their time, for the inhabitants of which the waters of this lovely spring
were miraculously healed. One of the “schools of the prophets”
existed here; and here Elijah had his last interview with the youth,
the hope of Israel, ere he went forth to yonder lonely tracts eastward,
to be parted from the faithful Elisha by the chariot of fire, and caught
away to heaven in the whirlwind. The fertility of the plains of Jericho
was almost proverbial in later days, when its rich revenues were
farmed by Herod from Cleopatra.
Then the magnificent balsam gardens and the groves of stately
palms lent added beauty to a scene in the midst of which the luxury-
loving Herod had his winter quarters. In the days of our Saviour the
sycamore tree was not wanting, as we learn from the story of
Zaccheus. The remains of old sugar-mills testify to the ancient
culture of the sugar-cane; and the ruins of ancient aqueducts, dating
from high antiquity, which brought the contributions of distant springs
to the grounds around the city, show with what care the paradise of
Jericho was watered. The balsam has now utterly disappeared; and
the stranger, coming unprepared upon the scene, might well-
exclaim, “How could Jericho ever be called ‘the city of palm trees’?”
But great fruitful vines may yet be seen, in the badly-kept gardens,
yielding with prodigality in spite of indifferent husbandry; and heavily-
laden bananas, bending over the hedges, offer of their sweetness to
the hand of the passer-by. The modern village boasts a hotel, a
Greek hospice for the accommodation of Russian pilgrims, and the
ruins of an old castle, which frown out upon the wilderness to
eastward. Those who are willing to be pleasantly deceived by
monkish tradition may also have the house of Zaccheus pointed out
to them. There are some three hundred inhabitants in the village,
creatures of a miserable physique, and with a most undesirable
reputation for laziness and thievishness.
That the plains of Jericho might be once again what they were of old
—a very garden of delights, wherein is enough and to spare for all—
needs hardly to be said. Were proof required, it would be found in
the surroundings of Elisha’s fountain. Wherever the waters of the
fountain come, the desert sands are transformed into fruitful fields,
and all its banks are clothed with emerald. No small supply of water
would serve to waken life over all the plains; but is not the vast
volume of the Jordan only waiting to be caught in the higher reaches,
and taught to run in fertilising streams all over the broad lands? For
long generations it has flowed idly past, only a few yards on either
side of the rushing flood tasting its refreshing power. What untold
wealth is rolling there, in these rich brown waves! What hand shall
arrest the flow, and turn its powers to noblest uses, so that all the
valley may be filled with the music of rustling grain and with the
beauty of ripening fruit?—that the wilderness may rejoice and
blossom as the rose. Ere this is possible, the reins of government
must be held by stronger and more righteous hands, the
husbandman must be secured in the enjoyment of the fruit of his toil.
But surely now the appointed season for favour cannot be long
delayed. Happy the eyes that shall behold the awaking of all the land
to fresh life after its death-like slumber! In those sweet days of
reviving, no fairer scenes will greet the eye than these broad
stretches, proudly in the midst of which will rise once more the
beautiful palm-girt city.

MOUTH OF WADY KELT


We could have lingered long beside that delightful spring, pouring its
sparkling waters forth in blessing over the plain; but the sun rose
higher in the heavens, and we had a hard ascent before us. We rode

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