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About the Authors
Lisa A. Urry Lisa Urry is Professor of Biology and Chair of the Biology Department at Mills
College in Oakland, California, and a Visiting Scholar at the University of California,
Berkeley. After graduating from Tufts University with a double major in biology and French,
Lisa completed her Ph.D. in molecular and developmental biology at the Massachusetts
Institute of Technology (MIT) in the MIT/Woods Hole Oceanographic Institution Joint Pro-
gram. She has published a number of research papers, most of them focused on gene expres-
sion during embryonic and larval development in sea urchins. Lisa has taught a variety of
courses, from introductory biology to developmental biology and senior seminar. As a part
of her mission to increase understanding of evolution, Lisa also teaches a nonmajors course
called Evolution for Future Presidents and is on the Teacher Advisory Board for the Under-
standing Evolution website developed by the University of California Museum of Paleontol-
ogy. Lisa is also deeply committed to promoting opportunities in science for women and
underrepresented minorities. Lisa is also a co-author of Campbell Biology in Focus.

Michael L. Cain Michael Cain is an ecologist and evolutionary biologist who is now writ-
ing full-time. Michael earned a joint degree in biology and math at Bowdoin College,
an M.Sc. from Brown University, and a Ph.D. in ecology and evolutionary biology from
Cornell University. As a faculty member at New Mexico State University and Rose-Hulman
Institute of Technology, he taught a wide range of courses, including introductory biology,
ecology, evolution, botany, and conservation biology. Michael is the author of dozens of
scientific papers on topics that include foraging behaviour in insects and plants, long-
distance seed dispersal, and speciation in crickets. In addition to his work on Campbell
BIOLOGY and Campbell Biology in Focus, Michael is the lead author of an ecology textbook.

Steven A. Wasserman Steve Wasserman is Professor of Biology at the University of Cali-


fornia, San Diego (UCSD). He earned his A.B. in biology from Harvard University and
his Ph.D. in biological sciences from MIT. Through his research on regulatory pathway
mechanisms in the fruit fly Drosophila, Steve has contributed to the fields of develop-
mental biology, reproduction, and immunity. As a faculty member at the University of
Texas Southwestern Medical Center and UCSD, he has taught genetics, development,
and physiology to undergraduate, graduate, and medical students. He currently focuses
on teaching introductory biology. He has also served as the research mentor for more
than a dozen doctoral students and more than 50 aspiring scientists at the undergradu-
ate and high school levels. Steve has been the recipient of distinguished scholar awards
from both the Markey Charitable Trust and the David and Lucile Packard Foundation.
In 2007, he received UCSD’s Distinguished Teaching Award for undergraduate teaching.
Steve is also a co-author of Campbell Biology in Focus.

Peter V. Minorsky Peter Minorsky is Professor of Biology at Mercy College in New York,
where he teaches introductory biology, evolution, ecology, and botany. He received his
A.B. in biology from Vassar College and his Ph.D. in plant physiology from Cornell Uni-
versity. He is also the science writer for the journal Plant Physiology. After a postdoctoral
fellowship at the University of Wisconsin at Madison, Peter taught at Kenyon College,
Union College, Western Connecticut State University, and Vassar College. His research
interests concern how plants sense environmental change. Peter received the 2008
Award for Teaching Excellence at Mercy College. Peter is also a co-author of Campbell
Biology in Focus.

ABOUT THE AUTHORS v


Behaviour of Chromosome Sets in the Human Life Cycle 273 Abnormal Chromosome Number 325
The Variety of Sexual Life Cycles 274 Alterations of Chromosome Structure 326
CONCEPT 13.3 Meiosis reduces the number of chromosome Human Disorders Due to Chromosomal Alterations 327
sets from diploid to haploid 275 CONCEPT 15.5 Some inheritance patterns are exceptions to
The Stages of Meiosis 275 standard Mendelian inheritance 329
Crossing Over and Synapsis During Prophase I 278 Genomic Imprinting 329
A Comparison of Mitosis and Meiosis 278 Inheritance of Organelle Genes 330
CONCEPT 13.4 Genetic variation produced in sexual life
cycles contributes to evolution 281
Origins of Genetic Variation Among Offspring 281
The Evolutionary Significance of Genetic Variation Within
16 The Molecular Basis of
Populations 282 Inheritance 334
Life’s Operating Instructions 334
14 Mendel and the Gene Idea 285 CONCEPT 16.1 DNA is the genetic material 335
The Search for the Genetic Material: Scientific Inquiry 335
Drawing from the Deck of Genes 285 Building a Structural Model of DNA: Scientific Inquiry 337
CONCEPT 14.1 Mendel used the scientific approach to CONCEPT 16.2 Many proteins work together in DNA
identify two laws of inheritance 286 replication and repair 340
Mendel’s Experimental, Quantitative Approach 286 The Basic Principle: Base Pairing to a Template Strand 340
The Law of Segregation 287 DNA Replication: A Closer Look 342
The Law of Independent Assortment 290 Proofreading and Repairing DNA 346
CONCEPT 14.2 Probability laws govern Mendelian Evolutionary Significance of Altered DNA Nucleotides 348
inheritance 292 Replicating the Ends of DNA Molecules 348
The Multiplication and Addition Rules Applied to Monohybrid CONCEPT 16.3 A chromosome consists of a DNA molecule
Crosses 293 packed together with proteins 350
Solving Complex Genetics Problems with the Rules
of Probability 293
CONCEPT 14.3 Inheritance patterns are often more complex 17 Gene Expression: From Gene to
than predicted by simple Mendelian genetics 294
Extending Mendelian Genetics for a Single Gene 294
Protein 355
Extending Mendelian Genetics for Two or More Genes 297
The Flow of Genetic Information 355
Nature and Nurture: The Environmental Impact on
Phenotype 298 CONCEPT 17.1 Genes specify proteins via transcription
A Mendelian View of Heredity and Variation 299 and translation 356
Evidence from the Study of Metabolic Defects 356
CONCEPT 14.4 Many human traits follow Mendelian
Basic Principles of Transcription and Translation 357
patterns of inheritance 300
The Genetic Code 359
Pedigree Analysis 300
Recessively Inherited Disorders 301 CONCEPT 17.2 Transcription is the DNA-directed synthesis
Dominantly Inherited Disorders 303 of RNA: A closer look 362
Multifactorial Disorders 304 Molecular Components of Transcription 362
Genetic Testing and Counselling 304 Synthesis of an RNA Transcript 362
CONCEPT 17.3 Eukaryotic cells modify RNA after
transcription 365
15 The Chromosomal Basis Alteration of mRNA Ends 365

of Inheritance 312 Split Genes and RNA Splicing 365


CONCEPT 17.4 Translation is the RNA-directed synthesis
Locating Genes Along Chromosomes 312 of a polypeptide: A closer look 367
Molecular Components of Translation 368
CONCEPT 15.1 Morgan showed that Mendelian inheritance
Building a Polypeptide 370
has its physical basis in the behaviour of chromosomes:
Completing and Targeting the Functional Protein 372
Scientific Inquiry 314
Making Multiple Polypeptides in Bacteria and Eukaryotes 375
CONCEPT 15.2 Sex-linked genes exhibit unique patterns of
CONCEPT 17.5 Mutations of one or a few nucleotides can
inheritance 315
affect protein structure and function 377
The Chromosomal Basis of Sex 316
Types of Small-Scale Mutations 377
Inheritance of X-Linked Genes 317
New Mutations and Mutagens 378
X Inactivation in Female Mammals 318
What Is a Gene? Revisiting the Question 380
CONCEPT 15.3 Linked genes tend to be inherited together
because they are located near each other on the same

18
chromosome 319
How Linkage Affects Inheritance 319 Regulation of Gene Expression 385
Genetic Recombination and Linkage 320
Mapping the Distance Between Genes Using Recombination Data: Beauty in the Eye of the Beholder 385
Scientific Inquiry 323 CONCEPT 18.1 Bacteria often respond to environmental
CONCEPT 15.4 Alterations of chromosome number or change by regulating transcription 386
structure cause some genetic disorders 325 Operons: The Basic Concept 386

DETAILED CONTENTS xiii


Repressible and Inducible Operons: Two Types of Negative Gene CONCEPT 20.2 Biologists use DNA technology to study gene
Regulation 388 expression and function 446
Positive Gene Regulation 389 Analyzing Gene Expression 446
CONCEPT 18.2 Eukaryotic gene expression is regulated at Determining Gene Function 450
many stages 390 CONCEPT 20.3 Cloned organisms and stem cells are useful
Differential Gene Expression 390 for basic research and other applications 452
Regulation of Chromatin Structure 391 Cloning Plants: Single-Cell Cultures 453
Regulation of Transcription Initiation 392 Cloning Animals: Nuclear Transplantation 453
Mechanisms of Post-Transcriptional Regulation 397 Stem Cells of Animals 455
CONCEPT 18.3 Noncoding RNAs play multiple roles in CONCEPT 20.4 The practical applications of DNA-based
controlling gene expression 399 biotechnology affect our lives in many ways 458
Effects on mRNAs by MicroRNAs and Small Interfering RNAs 399 Medical Applications 458
Chromatin Remodelling and Effects on Transcription by ncRNAs 400 Forensic Evidence and Genetic Profiles 461
The Evolutionary Significance of Small ncRNAs 401 Environmental Cleanup 462
CONCEPT 18.4 A program of differential gene expression Agricultural Applications 462
leads to the different cell types in a multicellular Safety and Ethical Questions Raised by DNA Technology 463
organism 401
A Genetic Program for Embryonic Development 401
Cytoplasmic Determinants and Inductive Signals 402
21 Genomes and Their Evolution 468
Sequential Regulation of Gene Expression During Cellular
Combining Genomics with Traditional Ecological
Differentiation 403
Knowledge 468
Pattern Formation: Setting Up the Body Plan 404
CONCEPT 21.1 The Human Genome Project fostered
CONCEPT 18.5 Cancer results from genetic changes that
development of faster, less expensive sequencing
affect cell cycle control 408 techniques 469
Types of Genes Associated with Cancer 408
Interference with Normal Cell-Signalling Pathways 409
CONCEPT 21.2 Scientists use bioinformatics to analyze
The Multistep Model of Cancer Development 411
genomes and their functions 470
Centralized Resources for Analyzing Genome Sequences 470
Inherited Predisposition and Environmental Factors Contributing
Identifying Protein-Coding Genes and Understanding Their
to Cancer 414
Functions 471
The Role of Viruses in Cancer 415
Understanding Genes and Gene Expression at the Systems Level 472
CONCEPT 21.3 Genomes vary in size, number of genes, and
19 Viruses 419 gene density 474
Genome Size 474
Number of Genes 475
A Borrowed Life 419
Gene Density and Noncoding DNA 475
CONCEPT 19.1 A virus consists of a nucleic acid surrounded
CONCEPT 21.4 Multicellular eukaryotes have a lot of
by a protein coat 420
noncoding DNA and many multigene families 478
The Discovery of Viruses: Scientific Inquiry 420
Transposable Elements and Related Sequences 478
Structure of Viruses 420
Other Repetitive DNA, Including Simple Sequence DNA 480
CONCEPT 19.2 Viruses replicate only in host cells 422 Genes and Multigene Families 480
General Features of Viral Replicative Cycles 422
CONCEPT 21.5 Duplication, rearrangement, and mutation of
Replicative Cycles of Phages 423
DNA contribute to genome evolution 482
Replicative Cycles of Animal Viruses 426
Duplication of Entire Chromosome Sets 482
Evolution of Viruses 428
Alterations of Chromosome Structure 482
CONCEPT 19.3 Viruses and prions are formidable pathogens Duplication and Divergence of Gene-Sized Regions of DNA 483
in animals and plants 430 Rearrangements of Parts of Genes: Exon Duplication and Exon
Viral Diseases in Animals 430 Shuffling 484
Emerging Viruses 431 How Transposable Elements Contribute to
Viral Diseases in Plants 435 Genome Evolution 487
Prions: Proteins as Infectious Agents 435
CONCEPT 21.6 Comparing genome sequences provides clues
to evolution and development 487
20 DNA Tools and Biotechnology 438 Comparing Genomes 487

The DNA Toolbox 438


UNIT 4 MECHANISMS OF EVOLUTION 495
CONCEPT 20.1 DNA sequencing and DNA cloning are
valuable tools for genetic engineering and biological Interview: Maydianne Andrade 495
inquiry 439
DNA Sequencing 439
Making Multiple Copies of a Gene or Other DNA Segment 441 22 Descent with Modification:
Using Restriction Enzymes to Make a Recombinant DNA A Darwinian View of Life 498
Plasmid 442
Amplifying DNA: The Polymerase Chain Reaction (PCR) and Its “Endless Forms Most Beautiful”—Charles Darwin 498
Use in DNA Cloning 443 CONCEPT 22.1 The Darwinian revolution challenged traditional
Expressing Cloned Eukaryotic Genes 445 views of a young Earth inhabited by unchanging species 499
xiv DETAILED CONTENTS
Scala Naturae and Classification of Species 500
Ideas About Change over Time 500 25 The History of Life on Earth 555
Lamarck’s Hypothesis of Evolution 500
CONCEPT 22.2 Descent with modification by natural Dinosaurs of a Feather 555
selection explains the adaptations of organisms and the CONCEPT 25.1 Conditions on early Earth made the origin
unity and diversity of life 501 of life possible 556
Darwin’s Research 501 Synthesis of Organic Compounds on Early Earth 556
CONCEPT 22.3 Evolution is supported by an overwhelming Abiotic Synthesis of Macromolecules 557
amount of scientific evidence 506 Protocells 557
Direct Observations of Evolutionary Change 507 Self-Replicating RNA 558
Homology 509 CONCEPT 25.2 The fossil record documents the history
The Fossil Record 511 of life 558
Biogeography 512 The Fossil Record 558
What Is Theoretical about Darwin’s View of Life? 513 How Rocks and Fossils Are Dated 560
The Origin of New Groups of Organisms 560

23
CONCEPT 25.3 Key events in life’s history include the
The Evolution of Populations 517 origins of unicellular and multicelled organisms and the
colonization of land 562
The Smallest Unit of Evolution 517 The First Single-Celled Organisms 564
CONCEPT 23.1 Genetic variation makes evolution The Origin of Multicellularity 565
possible 518 The Colonization of Land 566
Genetic Variation 518 CONCEPT 25.4 The rise and fall of groups of organisms
Sources of Genetic Variation 519 reflect differences in speciation and extinction rates 567
CONCEPT 23.2 The Hardy-Weinberg equation can be used Plate Tectonics 568
to test whether a population is evolving 520 Mass Extinctions 570
Gene Pools and Allele Frequencies 521 Adaptive Radiations 572
The Hardy-Weinberg Equation 521 CONCEPT 25.5 Major changes in body form can result from
CONCEPT 23.3 Natural selection, genetic drift, and gene changes in the sequences and regulation of developmental
flow can alter allele frequencies in a population 524 genes 574
Natural Selection 525 Effects of Developmental Genes 575
Genetic Drift 525 The Evolution of Development 576
Gene Flow 527 CONCEPT 25.6 Evolution is not goal oriented 578
CONCEPT 23.4 Natural selection is the only mechanism that Evolutionary Novelties 578
consistently causes adaptive evolution 528 Evolutionary Trends 579
Natural Selection: A Closer Look 528
The Key Role of Natural Selection in Adaptive Evolution 530
Sexual Selection 530 UNIT 5 THE EVOLUTIONARY HISTORY OF
Balancing Selection 531 BIOLOGICAL DIVERSITY 583
Frequency-Dependent Selection 531
Heterozygote Advantage 532 Interview: Laura Hug 583
Why Natural Selection Cannot Fashion Perfect Organisms 532
26 Phylogeny and the Tree of Life 586
24 The Origin of Species 536 Investigating the Tree of Life 586
CONCEPT 26.1 Phylogenies show evolutionary
That “Mystery of Mysteries” 536 relationships 587
CONCEPT 24.1 The biological species concept emphasizes Binomial Nomenclature 588
reproductive isolation 537 Hierarchical Classification 588
The Biological Species Concept 537 Linking Classification and Phylogeny 589
Other Definitions of Species 540 Visualizing Phylogenetic Relationships 589
CONCEPT 24.2 Speciation can take place with or without CONCEPT 26.2 Phylogenies are inferred from morphological
geographic separation 541 and molecular data 591
Allopatric (“Other Country”) Speciation 541 Morphological and Molecular Homologies 591
Sympatric (“Same Country”) Speciation 543 Sorting Homology from Analogy 591
Allopatric and Sympatric Speciation: A Review 546 Evaluating Molecular Homologies 591
CONCEPT 24.3 Hybrid zones reveal factors that cause CONCEPT 26.3 Shared characters are used to construct
reproductive isolation 546 phylogenetic trees 592
Patterns Within Hybrid Zones 546 Cladistics 592
Hybrid Zones and Environmental Change 547 Maximum Parsimony and Maximum Likelihood 594
Hybrid Zones over Time 547 Interpreting Phylogenetic Trees 595
CONCEPT 24.4 Speciation can occur rapidly or slowly and Applying Phylogenies 597
can result from changes in few or many genes 550 CONCEPT 26.4 An organism’s evolutionary history is
The Time Course of Speciation 550 documented in its genome 599
Studying the Genetics of Speciation 552 Gene Duplications and Gene Families 599
From Speciation to Macroevolution 553 Genome Evolution 600

DETAILED CONTENTS xv
CONCEPT 26.5 Molecular clocks help track evolutionary Alveolates 640
time 600 Rhizarians 641
Molecular Clocks 600 CONCEPT 28.4 Red algae and green algae are the closest
Applying a Molecular Clock: The Origin of HIV 601 relatives of land plants 645
CONCEPT 26.6 Our understanding of the tree of life Red Algae 645
continues to change based on new data 602 Green Algae 645
From Two Kingdoms to Three Domains 602 CONCEPT 28.5 Unikonts include protists that are closely
The Important Role of Horizontal Gene Transfer 603 related to fungi and animals 647
Amoebozoans 648

27
Opisthokonts 650
Bacteria and Archaea 607 CONCEPT 28.6 The relationships of some protists to other
eukaryotes is uncertain 650
Masters of Adaptation 607 Haptophytes 650
CONCEPT 27.1 Structural and functional adaptations Cryptomonads 651
contribute to prokaryotic success 608 CONCEPT 28.7 Protists play key roles in ecological
Cell-Surface Structures 608 communities 652
Endospores 609 Symbiotic Protists 652
Motility 610 Photosynthetic Protists 652
Internal Organization and DNA 611
Reproduction 612
CONCEPT 27.2 Rapid reproduction, mutation, and
genetic recombination promote genetic diversity in 29 Plant Diversity I: How Plants
prokaryotes 612 Colonized Land 657
Rapid Reproduction and Mutation 612
Genetic Recombination 612 The Greening of Earth 657
CONCEPT 27.3 Diverse nutritional and metabolic CONCEPT 29.1 Plants evolved from green algae 658
adaptations have evolved in prokaryotes 615 Morphological and Molecular Evidence 658
The Role of Oxygen in Metabolism 616 Adaptations Enabling the Move to Land 658
Nitrogen Metabolism 616 Derived Traits of Plants 659
Metabolic Cooperation 616 The Origin and Diversification of Plants 659
CONCEPT 27.4 Prokaryotes have radiated into a diverse set CONCEPT 29.2 Mosses and other nonvascular plants have
of lineages 617 life cycles dominated by gametophytes 663
An Overview of Prokaryotic Diversity 617 Bryophyte Gametophytes 664
Bacteria 620 Bryophyte Sporophytes 666
Archaea 620 The Ecological and Economic Importance of Mosses 666
CONCEPT 27.5 Prokaryotes play crucial roles in the CONCEPT 29.3 Ferns and other seedless vascular plants
biosphere 621 were the first plants to grow tall 668
Chemical Recycling 621 Origins and Traits of Vascular Plants 668
Ecological Interactions 622 Classification of Seedless Vascular Plants 672
CONCEPT 27.6 Prokaryotes have both beneficial The Significance of Seedless Vascular Plants 672
and harmful impacts on humans 623
Mutualistic Bacteria 623
Pathogenic Bacteria 623
Prokaryotes in Research and Technology 624 30 Plant Diversity II: The Evolution
of Seed Plants 678
28 Protists 628 Transforming the World 678
CONCEPT 30.1 Seeds and pollen grains are key adaptations
You Are What You Eat 628 for life on land 679
CONCEPT 28.1 Most eukaryotes are single-celled Advantages of Reduced Gametophytes 679
organisms 629 Heterospory: The Rule Among Seed Plants 680
Structural and Functional Diversity in Protists 629 Ovules and Production of Eggs 680
Four Supergroups of Eukaryotes 629 Pollen and Production of Sperm 680
Endosymbiosis in Eukaryotic Evolution 632 The Evolutionary Advantage of Seeds 681
Endosymbiosis and the Spread of Photosynthesis 632 Evolution of the Seed 682
How Does an Endosymbiont Evolve into an Organelle? 633 CONCEPT 30.2 Gymnosperms bear “naked” seeds, typically
CONCEPT 28.2 Excavates include protists with modified on cones 682
mitochondria and protists with unique flagella 635 The Life Cycle of a Pine 682
Diplomonads and Parabasalids 635 Evolution of Gymnosperms 686
Euglenozoans 635 Gymnosperm Diversity 686
CONCEPT 28.3 SAR is a highly diverse group of protists CONCEPT 30.3 The reproductive adaptations of angiosperms
defined by DNA similarities 637 include flowers and fruits 686
Stramenopiles 637 Characteristics of Angiosperms 686
Oomycetes (Water Moulds and Their Relatives) 639 Angiosperm Evolution 690

xvi DETAILED CONTENTS


Evolutionary Links Between Angiosperms and Animals 692 The Diversification of Animals 726
Angiosperm Diversity 692 Future Directions in Animal Systematics 728
CONCEPT 30.4 Human welfare depends greatly on seed
plants 695
Products from Seed Plants 695 33 An Introduction to Invertebrates 731
Threats to Plant Diversity 695
Life Without a Backbone 731

31
CONCEPT 33.1 Sponges are basal animals that lack true
Fungi 698 tissues 735
CONCEPT 33.2 Cnidarians are an ancient phylum of
Brewer’s Yeast and Climate Change 698 eumetazoans 736
CONCEPT 31.1 Fungi are heterotrophs that feed Medusozoans 737
by absorption 699 Anthozoans 737
Nutrition and Ecology 699 CONCEPT 33.3 Lophotrochozoans, a clade identified by
Body Structure 699 molecular data, have the widest range of animal body
Specialized Hyphae in Mycorrhizal Fungi 700 forms 739
CONCEPT 31.2 Fungi produce spores through sexual or Flatworms 739
asexual life cycles 702 Rotifers 742
Sexual Reproduction 702 Lophophorates: Ectoprocts and Brachiopods 743
Asexual Reproduction 702 Molluscs 744
CONCEPT 31.3 The ancestor of fungi was an aquatic, single- Annelids 747
celled, flagellated protist 703 CONCEPT 33.4 Ecdysozoans are the most species-rich animal
The Origin of Fungi 703 group 750
Basal Fungal Groups 703 Nematodes 750
The Move to Land 704 Arthropods 751
CONCEPT 31.4 Fungi have radiated into a diverse set of CONCEPT 33.5 Echinoderms and chordates are
lineages 704 deuterostomes 759
Chytrids 704 Echinoderms 759
Zygomycetes 706 Chordates 761
Glomeromycetes 707
Ascomycetes 707
Basidiomycetes 709
CONCEPT 31.5 Fungi play key roles in nutrient cycling, 34 The Origin and Evolution
ecological interactions, and human welfare 711 of Vertebrates 765
Fungi as Decomposers (Saprotrophs) 711
Fungi as Mutualists 711 Half a Billion Years of Backbones 765
Fungi as Parasites 712
CONCEPT 34.1 Chordates have a notochord and a dorsal,
Practical Uses of Fungi 714
hollow nerve cord 766
Derived Characters of Chordates 766
32 An Overview of Animal Lancelets 767
Tunicates 768
Diversity 717 Early Chordate Evolution 769
CONCEPT 34.2 Vertebrates are chordates that have a
Welcome to Your Kingdom 717 backbone 769
CONCEPT 32.1 Animals are multicellular, heterotrophic Derived Characters of Vertebrates 769
eukaryotes with tissues that develop from embryonic layers 718 Hagfishes and Lampreys 770
Nutritional Mode 718 Hagfishes 770
Cell Structure and Specialization 718 Lampreys 770
Reproduction and Development 718 Early Vertebrate Evolution 771
CONCEPT 32.2 The history of animals spans more than half CONCEPT 34.3 Gnathostomes are vertebrates that have
a billion years 719 jaws 772
Steps in the Origin of Multicellular Animals 719 Derived Characters of Gnathostomes 772
Neoproterozoic Era (1 Billion–542 Million Years Ago) 720 Fossil Gnathostomes 773
Paleozoic Era (542–251 Million Years Ago) 720 Chondrichthyans (Sharks, Rays, and Their Relatives) 773
Mesozoic Era (251–65.5 Million Years Ago) 722 Ray-Finned Fishes and Lobe-Fins 775
Cenozoic Era (65.5 Million Years Ago to the Present) 722
CONCEPT 34.4 Tetrapods are gnathostomes that have
CONCEPT 32.3 Animals can be characterized by “body limbs 777
plans” 723 Derived Characters of Tetrapods 777
Symmetry 724 The Origin of Tetrapods 777
Tissues 724 Amphibians 779
Body Cavities 724 Salamanders 779
Protostome and Deuterostome Development 725 Frogs 779
CONCEPT 32.4 Views of animal phylogeny continue to be Caecilians 779
shaped by new molecular and morphological data 726 Lifestyle and Ecology of Amphibians 779

DETAILED CONTENTS xvii


CONCEPT 34.5 Amniotes are tetrapods that have CONCEPT 36.2 Different mechanisms transport substances
a terrestrially adapted egg 782 over short or long distances 837
Derived Characters of Amniotes 782 The Apoplast and Symplast: Transport Continuums 837
Early Amniotes 783 Short-Distance Transport of Solutes across Plasma
Reptiles 783 Membranes 838
CONCEPT 34.6 Mammals are amniotes that have hair and Short-Distance Transport of Water Across Plasma
produce milk 788 Membranes 838
Derived Characters of Mammals 789 Long-Distance Transport: The Role of Bulk Flow 841
Early Evolution of Mammals 789 CONCEPT 36.3 Transpiration drives the transport
Monotremes 790 of water and minerals from roots to shoots via the
Marsupials 790 xylem 842
Eutherians (Placental Mammals) 791 Absorption of Water and Minerals by Root Epidermal
CONCEPT 34.7 Humans are mammals that have a large brain Cells 842
and bipedal locomotion 796 Transport of Water and Minerals into the Xylem 842
Derived Characters of Humans 796 Bulk Flow Transport via the Xylem 842
The Earliest Hominins 797 Xylem Sap Ascent by Bulk Flow: A Review 846
Australopiths 797 CONCEPT 36.4 The rate of transpiration is regulated by
Bipedalism 798 stomata 846
Tool Use 798 Stomata: Major Pathways for Water Loss 847
Early Homo 799 Mechanisms of Stomatal Opening and Closing 847
Neanderthals 800 Stimuli for Stomatal Opening and Closing 848
Effects of Transpiration on Wilting and Leaf Temperature 848
Adaptations That Reduce Evaporative Water Loss 848
UNIT 6 PLANT FORM AND FUNCTION 805 CONCEPT 36.5 Sugars are transported from sources to sinks
via the phloem 849
Interview: Jacqueline Monaghan 805 Movement from Sugar Sources to Sugar Sinks 849
Bulk Flow by Positive Pressure: The Mechanism of Translocation
35 Plant Structure, Growth, and in Angiosperms 850
CONCEPT 36.6 The symplast is highly dynamic 852
Development 808 Changes in Plasmodesmatal Number and Pore Size 852
Phloem: An Information Superhighway 852
Why Did the Palm Tree Cross the Road? 808 Electrical Signalling in the Phloem 852
CONCEPT 35.1 Plants have a hierarchical organization
consisting of organs, tissues, and cells 809
Basic Vascular Plant Organs: Roots, Stems, and Leaves 809
Dermal, Vascular, and Ground Plant Tissues 812
Common Types of Plant Cells 813
37 Soil and Plant Nutrition 855
CONCEPT 35.2 Different meristems generate cells for The Corkscrew Carnivore 855
primary and secondary growth 816 CONCEPT 37.1 Soil contains a living, complex
CONCEPT 35.3 Primary growth lengthens roots and shoots 818 ecosystem 856
Primary Growth of Roots 818 Soil Texture 856
Primary Growth of Shoots 818 Topsoil Composition 856
CONCEPT 35.4 Secondary growth increases the diameter of Soil Conservation and Sustainable Agriculture 857
stems and roots in woody plants 822 CONCEPT 37.2 Plant roots absorb essential elements from
The Vascular Cambium and Secondary Vascular Tissue 823 the soil 860
The Cork Cambium and the Production of Periderm 824 Essential Elements 860
Evolution of Secondary Growth 824 Symptoms of Mineral Deficiency 861
CONCEPT 35.5 Growth, morphogenesis, and cell Improving Plant Nutrition by Genetic Modification 862
differentiation produce the plant body 825 CONCEPT 37.3 Plant nutrition often involves relationships
Model Organisms: Revolutionizing the Study of Plants 826 with other organisms 863
Growth: Cell Division and Cell Expansion 826 Bacteria and Plant Nutrition 863
Morphogenesis and Pattern Formation 828 Fungi and Plant Nutrition 867
Gene Expression and Control of Cell Differentiation 828 Vertebrates and Plant Nutrition 868
Shifts in Development: Phase Changes 829 Epiphytes, Parasitic Plants, and Carnivorous Plants 869
Genetic Control of Flowering 830

36 Resource Acquisition and Transport 38 Angiosperm Reproduction and


in Vascular Plants 834 Biotechnology 873
Overview 834 Canola (Canadian Oil Low Acid): A Canadian Invention 873
CONCEPT 36.1 Adaptations for acquiring resources were key CONCEPT 38.1 Flowers, double fertilization, and fruits are
steps in the evolution of vascular plants 835 unique features of the angiosperm life cycle 874
Shoot Architecture and Light Capture 835 Flower Structure and Function 874
Root Architecture and Acquisition of Water and Minerals 837 Methods of Pollination 875

xviii DETAILED CONTENTS


The Angiosperm Life Cycle: An Overview 875 CONCEPT 40.2 Feedback control maintains the internal
Seed Development and Structure: A Closer Look 878 environment in many animals 936
Sporophyte Development from Seed to Mature Plant 881 Regulating and Conforming 936
Fruit Structure and Function 882 Homeostasis 936
CONCEPT 38.2 Flowering plants reproduce sexually, CONCEPT 40.3 Homeostatic processes for thermoregulation
asexually, or both 883 involve form, function, and behaviour 938
Mechanisms of Asexual Reproduction 883 Endothermy and Ectothermy 939
Advantages and Disadvantages of Asexual Versus Sexual Variation in Body Temperature 939
Reproduction 885 Balancing Heat Loss and Gain 940
Mechanisms That Prevent Self-Fertilization 886 Acclimation and Acclimatization 943
Totipotency, Vegetative Reproduction, and Tissue Culture 887 Physiological Thermostats and Fever 944
CONCEPT 38.3 Humans modify crops by breeding and CONCEPT 40.4 Energy requirements are related to animal
genetic engineering 888 size, activity, and environment 945
Plant Breeding 889 Energy Allocation and Use 945
Plant Biotechnology and Genetic Engineering 889 Quantifying Energy Use 945
The Debate over Plant Biotechnology 891 Minimum Metabolic Rate and Thermoregulation 946
Influences on Metabolic Rate 946
Torpor and Energy Conservation 948
39 Plant Responses to Internal
and External Signals 895 41 Animal Nutrition 951
Stimuli and a Stationary Life 895 The Need to Feed 951
CONCEPT 39.1 Signal transduction pathways link signal CONCEPT 41.1 An animal’s diet must supply chemical energy
reception to response 896 and building blocks 952
Reception 897 Essential Nutrients 952
Transduction 897 Dietary Deficiencies 954
Response 898 Assessing Nutritional Needs 956
CONCEPT 39.2 Plant hormones help coordinate growth, CONCEPT 41.2 Food processing involves ingestion,
development, and responses to stimuli 899 digestion, absorption, and elimination 956
A Survey of Plant Hormones 900 Digestive Compartments 958
CONCEPT 39.3 Responses to light are critical for plant CONCEPT 41.3 Organs specialized for sequential stages
success 908 of food processing form the mammalian digestive
Blue-Light Photoreceptors 908 system 959
Phytochromes as Photoreceptors 909 The Oral Cavity, Pharynx, and Esophagus 960
Biological Clocks and Circadian Rhythms 910 Digestion in the Stomach 960
The Effect of Light on the Biological Clock 911 Digestion in the Small Intestine 962
Photoperiodism and Responses to Seasons 912 Absorption in the Small Intestine 963
CONCEPT 39.4 Plants respond to a wide variety of stimuli Absorption in the Large Intestine 964
other than light 914 CONCEPT 41.4 Evolutionary adaptations of vertebrate
Gravity 914 digestive systems correlate with diet 965
Mechanical Stimuli 914 Dental Adaptations 965
Environmental Stresses 915 Stomach and Intestinal Adaptations 965
CONCEPT 39.5 Plants respond to attacks by herbivores Mutualistic Adaptations 966
and pathogens 919 Mutualistic Adaptations in Herbivores 968
Defences against Pathogens 919 CONCEPT 41.5 Feedback circuits regulate digestion, energy
Defences Against Herbivores 921 storage, and appetite 969
Regulation of Digestion 969
Regulation of Energy Storage 969
UNIT 7 ANIMAL FORM AND FUNCTION 925 Regulation of Appetite and Consumption 971
Obesity and Evolution 972
Interview: Matt Vijayan 925

42 Circulation and Gas Exchange 975


40 Basic Principles of Animal Form
and Function 928 Trading Places 975
CONCEPT 42.1 Circulatory systems link exchange surfaces
Diverse Forms, Common Challenges 928 with cells throughout the body 976
General Properties of Circulatory Systems 976
CONCEPT 40.1 Animal form and function are correlated at
Open and Closed Circulatory Systems 977
all levels of organization 929
Evolution of Vertebrate Circulatory Systems 978
Evolution of Animal Size and Shape 929
Exchange with the Environment 929 CONCEPT 42.2 Coordinated cycles of heart contraction drive
Hierarchical Organization of Body Plans 931 double circulation in mammals 981
Coordination and Control 935 Mammalian Circulation 981

DETAILED CONTENTS xix


The Mammalian Heart: A Closer Look 981
Maintaining the Heart’s Rhythmic Beat 982 44 Osmoregulation and Excretion 1035
CONCEPT 42.3 Patterns of blood pressure and flow reflect
the structure and arrangement of blood vessels 983 A Balancing Act 1035
Blood Vessel Structure and Function 984 CONCEPT 44.1 Osmoregulation balances the uptake and loss
Blood Flow Velocity 984 of water and solutes 1036
Blood Pressure 985 Osmosis and Osmolarity 1036
Capillary Function 987 Osmotic Challenges 1036
Fluid Return by the Lymphatic System 988 Energetics of Osmoregulation 1039
CONCEPT 42.4 Blood components function in exchange, Transport Epithelia in Osmoregulation 1039
transport, and defence 989 CONCEPT 44.2 An animal’s nitrogenous wastes reflect its
Blood Composition and Function 989 phylogeny and habitat 1040
Cardiovascular Disease 992 Forms of Nitrogenous Waste 1040
CONCEPT 42.5 Gas exchange occurs across specialized The Influence of Evolution and Environment on Nitrogenous
respiratory surfaces 994 Wastes 1041
Partial Pressure Gradients in Gas Exchange 994 CONCEPT 44.3 Diverse excretory systems are variations
Respiratory Media 994 on a tubular theme 1042
Respiratory Surfaces 994 Excretory Processes 1042
Gills in Aquatic Animals 995 Survey of Excretory Systems 1042
Tracheal Systems in Insects 996 CONCEPT 44.4 The nephron is organized for stepwise
Lungs 997 processing of blood filtrate 1046
CONCEPT 42.6 Breathing ventilates the lungs 999 From Blood Filtrate to Urine: A Closer Look 1046
How an Amphibian Breathes 999 Solute Gradients and Water Conservation 1047
How a Bird Breathes 999 Adaptations of the Vertebrate Kidney to Diverse Environments 1049
How a Mammal Breathes 999 CONCEPT 44.5 Hormonal circuits link kidney function, water
Control of Breathing in Humans 1000 balance, and blood pressure 1052
CONCEPT 42.7 Adaptations for gas exchange include Antidiuretic Hormone 1052
pigments that bind and transport gases 1001 The Renin-Angiotensin-Aldosterone System 1053
Coordination of Circulation and Gas Exchange 1001 Coordinated Regulation of Salt and Water Balance 1054
Respiratory Pigments 1002

45
Respiratory Adaptations of Diving Mammals 1003
Hormones and the Endocrine
System 1057
43 The Immune System 1008
The Body’s Long-Distance Regulators 1057
Recognition and Response 1008 CONCEPT 45.1 Hormones and other signalling molecules
CONCEPT 43.1 In innate immunity, recognition and response bind to target receptors, triggering specific response
rely on traits common to groups of pathogens 1009 pathways 1058
Innate Immunity of Invertebrates 1009 Intercellular Communication 1058
Innate Immunity of Vertebrates 1011 Chemical Classes of Intercellular Signalling Factors 1059
Evasion of Innate Immunity by Pathogens 1014 Cellular Response Pathways 1060
Multiple Effects of Hormones 1062
CONCEPT 43.2 In adaptive immunity, receptors provide
pathogen-specific recognition 1014 CONCEPT 45.2 Feedback regulation and coordination with
Antigen Recognition by B Cells and Antibodies 1014 the nervous system are common in endocrine systems 1064
Antigen Recognition by T Cells 1015 Simple Endocrine Pathways 1064
B Cell and T Cell Development 1016 Simple Neuroendocrine Pathways 1064
Feedback Regulation 1064
CONCEPT 43.3 Adaptive immunity defends against infection
Coordination of the Endocrine and Nervous Systems 1065
of body fluids and body cells 1019
Thyroid Regulation: A Hormone Cascade Pathway 1068
Helper T Cells: A Response to Nearly All Antigens 1020
Hormonal Regulation of Growth 1069
B Cells and Antibodies: A Response to Extracellular Pathogens 1020
Cytotoxic T Cells: A Response to Infected Cells 1023 CONCEPT 45.3 Endocrine glands respond to diverse stimuli
Summary of the Humoral and Cell-Mediated Immune in regulating homeostasis, development, and behaviour 1070
Responses 1023 Parathyroid Hormone and Vitamin D: Control of Blood Calcium 1070
Immunization 1024 Adrenal Hormones: Response to Stress 1070
Active and Passive Immunity 1025 Sex Hormones 1072
Antibodies as Tools 1025 Hormones and Biological Rhythms 1074
Immune Rejection 1026
CONCEPT 43.4 Disruptions in immune system function can
elicit or exacerbate disease 1027 46 Animal Reproduction 1078
Exaggerated, Self-Directed, and Diminished Immune
Responses 1027 Pairing Up for Sexual Reproduction 1078
Evolutionary Adaptations of Pathogens That Underlie Immune CONCEPT 46.1 Both asexual and sexual reproduction occur
System Avoidance 1029 in the animal kingdom 1079
Cancer and Immunity 1032 Mechanisms of Asexual Reproduction 1079

xx DETAILED CONTENTS
Variation in Patterns of Sexual Reproduction 1079 CONCEPT 48.4 Neurons communicate with other cells
Reproductive Cycles 1080 at synapses 1139
Sexual Reproduction: An Evolutionary Enigma 1081 Generation of Postsynaptic Potentials 1139
CONCEPT 46.2 Fertilization depends on mechanisms that Summation of Postsynaptic Potentials 1141
bring together sperm and eggs of the same species 1082 Termination of Neurotransmitter Signalling 1142
Ensuring the Survival of Offspring 1082 Modulated Signalling at Synapses 1142
Gamete Production and Delivery 1083 Neurotransmitters 1142
CONCEPT 46.3 Reproductive organs produce and transport
gametes 1085
Human Male Reproductive Anatomy 1085
Human Female Reproductive Anatomy 1086
49 Nervous Systems 1147
Gametogenesis 1087 Command and Control Centre 1147
CONCEPT 46.4 The interplay of tropic and sex hormones CONCEPT 49.1 Nervous systems consist of circuits
regulates mammalian reproduction 1090 of neurons and supporting cells 1148
Hormonal Control of the Male Reproductive System 1091
Glia 1149
Hormonal Control of Female Reproductive Cycles 1091
Organization of the Vertebrate Nervous System 1150
Human Sexual Response 1094
The Peripheral Nervous System 1151
CONCEPT 46.5 In placental mammals, an embryo develops
CONCEPT 49.2 The vertebrate brain is regionally
fully within the mother’s uterus 1094
specialized 1153
Conception, Embryonic Development, and Birth 1094
Arousal and Sleep 1153
Maternal Immune Tolerance of the Embryo and Fetus 1098
Biological Clock Regulation 1156
Contraception and Abortion 1098
Emotions 1156
Modern Reproductive Technologies 1099
Functional Imaging of the Brain 1158

47 Animal Development 1103


CONCEPT 49.3 The cerebral cortex controls voluntary
movement and cognitive functions 1158
Information Processing 1158
A Body-Building Plan 1103 Language and Speech 1160
CONCEPT 47.1 Fertilization and cleavage initiate embryonic Lateralization of Cortical Function 1160
development 1104 Frontal Lobe Function 1160
Fertilization 1104 Evolution of Cognition in Vertebrates 1160
Cleavage 1107 CONCEPT 49.4 Changes in synaptic connections underlie
CONCEPT 47.2 Morphogenesis in animals involves specific memory and learning 1161
changes in cell shape, position, and survival 1109 Neural Plasticity 1162
Gastrulation 1109 Memory and Learning 1162
Developmental Adaptations of Amniotes 1113 Long-Term Potentiation 1163
Organogenesis 1114 CONCEPT 49.5 Many nervous system disorders can be
Cellular Mechanisms in Morphogenesis 1115 explained in molecular terms 1164
CONCEPT 47.3 Cytoplasmic determinants and inductive Schizophrenia 1165
signals contribute to cell fate specification 1117 Depression 1165
Fate Mapping 1117 The Brain’s Reward System and Drug Addiction 1166
Axis Formation 1119 Alzheimer’s Disease 1166
Cell Fate Determination and Pattern Formation by Inductive Parkinson’s Disease 1167
Signals 1121 Future Directions 1167
Climate Change and Development 1124

48 Neurons, Synapses, 50 Sensory and Motor


and Signalling 1129 Mechanisms 1170
Lines of Communication 1129 Sense and Sensibility 1170
CONCEPT 48.1 Neuron organization and structure reflect CONCEPT 50.1 Sensory receptors transduce stimulus
function in information transfer 1130 energy and transmit signals to the central nervous
Neuron Structure and Function 1130 system 1171
Introduction to Information Processing 1130 Sensory Reception and Transduction 1171
CONCEPT 48.2 Ion pumps and ion channels establish the Transmission 1172
resting potential of a neuron 1132 Perception 1172
The Resting Membrane Potential 1132 Amplification and Adaptation 1172
Modelling the Resting Membrane Potential 1133 Types of Sensory Receptors 1173
CONCEPT 48.3 Action potentials are the signals conducted CONCEPT 50.2 The mechanoreceptors responsible for
by axons 1134 hearing and equilibrium detect moving fluid or settling
Hyperpolarization and Depolarization 1135 particles 1175
Graded Potentials and Action Potentials 1135 Sensing of Gravity and Sound in Invertebrates 1175
Generation of Action Potentials: A Closer Look 1136 Hearing and Equilibrium in Mammals 1175
Conduction of Action Potentials 1137 Hearing and Equilibrium in Other Vertebrates 1179

DETAILED CONTENTS xxi


CONCEPT 50.3 Visual receptors in animals depend on General Features of Terrestrial Biomes 1239
light-absorbing pigments 1180 Disturbance and Terrestrial Biomes 1239
Evolution of Visual Perception 1180 CONCEPT 52.3 Aquatic biomes are diverse and dynamic
The Vertebrate Visual System 1181 systems that cover most of Earth 1244
CONCEPT 50.4 The senses of taste and smell rely on similar Zonation in Aquatic Biomes 1244
sets of sensory receptors 1186 CONCEPT 52.4 Interactions between organisms and the
Taste in Mammals 1187 environment limit the distribution of species 1245
Smell in Humans 1188 Dispersal and Distribution 1250
CONCEPT 50.5 The physical interaction of protein filaments Abiotic Factors 1251
is required for muscle function 1189 Biotic Factors 1252
Vertebrate Skeletal Muscle 1189 CONCEPT 52.5 Ecological change and evolution affect one
Other Types of Muscle 1195 another over time 1254
CONCEPT 50.6 Skeletal systems transform muscle
contraction into locomotion 1196
Types of Skeletal Systems 1196
Types of Locomotion 1197
53 Population Ecology 1258
Sable Island, a Natural Laboratory 1258
51 Animal Behaviour 1203 CONCEPT 53.1 Dynamic biological processes
influence population density, dispersion, and
demographics 1259
The How and Why of Animal Activity 1203
Density and Dispersion 1259
CONCEPT 51.1 Discrete sensory inputs can stimulate both Demographics 1261
simple and complex behaviours 1204
CONCEPT 53.2 The exponential model describes
Fixed Action Patterns 1204
population growth in an idealized, unlimited
Migration 1204
environment 1264
Behavioural Rhythms 1205
Change in Population Size 1264
Animal Signals and Communication 1205
Exponential Growth 1264
CONCEPT 51.2 Learning establishes specific links between
CONCEPT 53.3 The logistic model describes how a
experience and behaviour 1207
population grows more slowly as it nears its carrying
Experience and Behaviour 1208
capacity 1265
Learning 1208
The Logistic Growth Model 1266
CONCEPT 51.3 Selection for individual survival and The Logistic Model and Real Populations 1267
reproductive success can explain most behaviours 1213
CONCEPT 53.4 Life history traits are products of natural
Foraging Behaviour 1213
selection 1268
Mating Behaviour and Mate Choice 1214
Evolution and Life History Diversity 1268
CONCEPT 51.4 Genetic analyses and the concept of inclusive “Trade-offs” and Life Histories 1269
fitness provide a basis for studying the evolution of
CONCEPT 53.5 Density-dependent factors regulate
behaviour 1219
population growth 1271
Genetic Basis of Behaviour 1219
Population Dynamics 1272
Genetic Variation and the Evolution of Behaviour 1220
Altruism 1221 CONCEPT 53.6 The human population is no longer growing
Inclusive Fitness 1222 exponentially but is still increasing rapidly 1275
Evolution and Human Culture 1223 The Global Human Population 1275
Global Carrying Capacity 1277

UNIT 8 ECOLOGY 1227


54 Community Ecology 1282
Interview: Erin Bertrand 1227
Dynamic Communities 1282
52 An Introduction to Ecology CONCEPT 54.1 Community interactions are classified by
whether they help, harm, or have no effect on the species
and the Biosphere 1230 involved 1283
Competition 1283
Life on and in the Ice 1230 Exploitation 1285
CONCEPT 52.1 Earth’s climate varies by latitude and season Positive Interactions 1288
and is changing rapidly 1231 Facilitation 1290
Global Climate Patterns 1231 CONCEPT 54.2 Diversity and trophic structure characterize
Regional and Local Effects on Climate 1231 biological communities 1290
Microclimate 1235 Species Diversity 1290
Global Climate Change 1236 Diversity and Community Stability 1291
CONCEPT 52.2 The structure and distribution of terrestrial Trophic Structure 1291
biomes are controlled by climate and disturbance 1237 Species with a Large Impact 1293
Climate and Terrestrial Biomes 1238 Bottom-Up and Top-Down Controls 1294

xxii DETAILED CONTENTS


CONCEPT 54.3 Disturbance influences species diversity
and composition 1296 56 Conservation Biology and Global
Characterizing Disturbance 1297 Change 1330
Ecological Succession 1298
Human Disturbance 1300 What Is Going Wrong with the Right Whales? 1330
CONCEPT 54.4 Biogeographic factors affect community CONCEPT 56.1 Human activities threaten Earth’s
diversity 1300 biodiversity 1331
Latitudinal Gradients 1300 Three Levels of Biodiversity 1331
Area Effects 1301 Biodiversity and Human Welfare 1333
Island Equilibrium Model 1301 Threats to Biodiversity 1334
CONCEPT 54.5 Pathogens alter community structure locally CONCEPT 56.2 Population conservation focuses on
and globally 1302 population size, genetic diversity, and critical habitat 1336
Pathogens and Communities 1303 Small-Population Approach 1336
Community Ecology and Zoonotic Diseases 1303 Field Study: The Greater Prairie Chicken and the Extinction Vortex 1337
Field Study: Analysis of Grizzly Bear Populations 1338
Declining-Population Approach 1338
Field Study: Decline of the Rufa Red Knot 1339
55 Ecosystems and Restoration Weighing Conflicting Demands 1340
CONCEPT 56.3 Landscape and regional conservation help
Ecology 1308 sustain biodiversity 1340
Landscape Structure and Biodiversity 1340
Engineering Salmon 1308 Establishing Protected Areas 1341
CONCEPT 55.1 Physical laws govern energy flow and Urban Ecology 1344
chemical cycling in ecosystems 1309 CONCEPT 56.4 Earth is changing rapidly as a result of
Conservation of Energy 1309 human actions 1344
Conservation of Mass 1310 Acid Precipitation 1344
Energy, Mass, and Trophic Levels 1310 Nutrient Enrichment 1345
CONCEPT 55.2 Energy and other limiting factors control Toxins in the Environment 1346
primary production in ecosystems 1311 Greenhouse Gases and Climate Change 1347
Ecosystem Energy Budgets 1311 Depletion of Atmospheric Ozone 1347
Primary Production in Aquatic Ecosystems 1313 CONCEPT 56.5 Sustainable development can improve
Primary Production in Terrestrial Ecosystems 1315 human lives while conserving biodiversity 1351
CONCEPT 55.3 Energy transfer between trophic levels is Sustainable Development 1351
typically only 10% efficient 1317 Field Study: Sustainable Development in Costa Rica 1352
Production Efficiency 1317 The Future of the Biosphere 1353
Trophic Efficiency and Ecological Pyramids 1317
APPENDIX A Answers A-1
CONCEPT 55.4 Biological and geochemical processes cycle
APPENDIX B Periodic Table of the Elements B-1
nutrients and water in ecosystems 1319
Biogeochemical Cycles 1319 APPENDIX C A Comparison of the Light Microscope
Decomposition and Nutrient Cycling Rates 1322 and the Electron Microscope C-1
CONCEPT 55.5 Restoration ecologists help return degraded APPENDIX D Classification of Life D-1
ecosystems to a more natural state 1323 APPENDIX E Scientific Skills Review E - 1
Bioremediation 1324
Biological Augmentation 1325 GLOSSARY G-1
Restoration Projects Worldwide 1325 INDEX I-1

DETAILED CONTENTS xxiii


Preface
W e are honoured to present
the Third Canadian Edition
of Campbell BIOLOGY. For the last
exercises are designed to engage
students through compelling case
studies and provide practice with
three decades, Campbell BIOLOGY data analysis skills. Problem-Solving
has been the leading university text Exercises have assignable versions
in the biological sciences. It has been in Mastering Biology. Some also
translated into 19 languages and has have more extensive “Solve It”
provided millions of students with a investigations to further explore a
solid foundation in university-level given topic.
biology. This success is a testament not Ready-to-Go Teaching
only to Neil Campbell’s original vision Modules on key topics provide
but also to the dedication of thousands instructors with assignments to use
of reviewers, who, together with before and after class, as well as in-
editors, artists, and contributors, have class activities that use clickers or
shaped and inspired this work. Learning Catalytics™ for assessment.
Our goals for the Third Canadian Integrated text and media:
Edition include: Media references in the printed
book direct students to the wealth of
increasing visual literacy through new figures,
online self-study resources available
questions, and exercises that build students’ skills in
to them in the Study Area section of Mastering Biology. In
understanding and creating visual representations of
the eText, these resources are integrated directly into the
biological structures and processes
eText. The new online learning tools include:
asking students to practise scientific skills by applying
Figure Walkthroughs that guide students through
scientific skills to real-world problems
key figures with narrated explanations, figure markups,
supporting instructors by providing teaching modules
and questions that reinforce important points. Additional
with tools and materials for introducing, teaching, and
questions can be assigned in Mastering Biology.
assessing important and often challenging topics
Animations and videos that bring biology to life.
integrating text and media to engage, guide, and
These include resources from HHMI BioInteractive
inform students in an active process of inquiry and learning
that engage students in topics from the discovery of the
Our starting point, as always, is our commitment to craft- double helix to evolution.
ing text and visuals that are accurate, current, and reflect our Interviews from the First Edition through the Third
passion for teaching biology. Canadian Edition of Campbell BIOLOGY are referenced
in the chapter where they are most relevant. The
interviews show students the human side of science
New to This Edition by featuring diverse scientists talking about how they
Here we provide an overview of the new features that we have became interested in what they study, how they began,
developed for the Third Canadian Edition; we invite you to
and what inspires them.
explore pages xxvi–xxix for more information and examples.
The impact of climate change at all levels of the biological
Visualizing Figures and Visual Skills Questions hierarchy is explored throughout the text, starting with a new
give students practice in interpreting and creating visual figure (Figure 1.12) and discussion in Chapter 1 and concluding
representations in biology. The Visualizing Figures have with a new Unit 8 Make Connections Figure and expanded
embedded questions that guide students in exploring coverage on causes and effects of climate change in Chapter 56.
how diagrams, photographs, and models represent and As in each new edition of Campbell BIOLOGY, the
reflect biological systems and processes. Assignable Third Canadian Edition incorporates new content and
questions are also available in Mastering Biology to give pedagogical improvements. These are summarized on
students practice with the visual skills addressed in the pp. vi–viii, following this Preface. Content updates
figures. reflect rapid, ongoing changes in technology and
Numeracy Questions and Problem-Solving knowledge in the fields of genomics, gene editing
Exercises challenge students to apply scientific skills technology (CRISPR), evolutionary biology,
and interpret data in solving real-world problems. These microbiology, and more. In addition, significant

xxiv PREFACE
revisions to Unit 8, Ecology, improve the conceptual Your Knowledge and Interpret the Data Questions. The answers
framework for core ecological topics (such as to most of these questions require students to write as well as
population growth, species interactions, and think and thus help develop the core competency of commu-
community dynamics) and more deeply integrate nicating science.
evolutionary principles. Finally, Campbell BIOLOGY has always featured scientific
inquiry, an essential component of any biology course.
Complementing stories of scientific discovery in the text
Our Hallmark Features narrative, the unit-opening interviews, and our standard-
setting Inquiry Figures all deepen the ability of students to
Teachers of general biology face a daunting challenge: to help understand how we know what we know. Scientific Inquiry
students acquire a conceptual framework for organizing an ever- Questions give students opportunities to practise scientific
expanding amount of information. The hallmark features of thinking, along with the Problem-Solving Exercises, ­Scientific
Campbell BIOLOGY provide such a framework while promoting Skills Exercises, and Interpret the Data Questions. Together,
a deeper understanding of biology and the process of science. these activities provide students practice both in applying the
Chief among the themes of Campbell BIOLOGY is evolution. process of science and in using quantitative reasoning.
Chapters throughout the text include at least one Evolution
section that explicitly focuses on evolutionary aspects of the
chapter material, and chapters end with an Evolution Connec-
Mastering Biology®
tion Question and a Write about a Theme Question. Mastering Biology, the most widely used online assessment
To help students distinguish the “forest from the trees,” and tutorial program for biology, provides an extensive library
each chapter is organized around a framework of three to of homework assignments that are graded automatically. In
seven carefully chosen Key Concepts. The text, Concept addition to the new Figure Walkthroughs, Problem-
Check Questions, Summary of Key Concepts, and Mastering Solving Exercises, and Visualizing Figures, Mastering
Biology all reinforce these main ideas and essential facts. Biology offers Dynamic Study Modules, Adaptive Follow-Up
In an effort to act on the Calls to Action from the Truth Assignments, Scientific Skills Exercises, Interpret the Data
and Reconciliation Commissioners’ Report (2012) , the Cana- Questions, Solve It Tutorials, HHMI Bio-Interactive Short Films,
dian authors were committed to including more Indigenous BioFlix. Tutorials with 3-D Animations, Experimental Inquiry
content in this text. Pearson’s fist step in this third Canadian Tutorials, Interpreting Data Tutorials, BLAST Tutorials, Make
edition is to acknowledge and highlight terminologies that Connections Tutorials, Get Ready for Biology, Activities, Read-
come from Indigenous origins and include references to ing Quiz Questions, Student Misconception Questions, and
how Traditional Ecological Knowledge (TEK) is being used in 4500 Test Bank Questions. Mastering Biology also includes the
Chapter 21 and 54. The authors recognize this is just the be- Campbell BIOLOGY eText, Study Area, and Instructor Resources.
ginning in a long process of responding to Truth and Recon- Go to www.masteringbiology.com for more details.
ciliation with Indigenous Peoples of this land.
Because text and illustrations are equally important for learn-
Our Partnership with Instructors
ing biology, integration of text and figures has been a hall-
mark of this text since the first edition. In addition to the new
and Students
Visualizing Figures, our popular Exploring Figures and Make A core value underlying our work is our belief in the importance
Connections Figures epitomize this approach. Each Exploring of a partnership with instructors and students. One primary way
Figure is a learning unit of core content that brings together of serving instructors and students, of course, is providing a text
related illustrations and text. Make ­Connections Figures reinforce that teaches biology well. In addition, Pearson Education offers a
fundamental conceptual connections throughout biology, rich variety of instructor and student resources, in both print and
helping students overcome tendencies to compartmentalize in- electronic form. In our continuing efforts to improve the book
formation. The Third Canadian Edition features two new Make and its supplements, we benefit tremendously from instructor
Connections Figures. There are also Guided Tour Figures that and student feedback, not only in formal reviews from hundreds
walk students through complex figures as an instructor would. of scientists, but also via e-mail and other forms of informal
To encourage active reading of the text, Campbell communication.
BIOLOGY includes numerous opportunities for students to The real test of any textbook is how well it helps instructors
stop and think about what they are reading, often by putting teach and students learn. We welcome comments from both
pencil to paper to draw a sketch, annotate a figure, or graph students and instructors. Please address your suggestions to
data. Active reading questions include Make Connections Fiona Rawle, Lead Author, at fiona.rawle@utoronto.ca, and
Questions, What If? Questions, Figure Legend Questions, Draw Cathleen Sullivan, Executive Acquisitions Editor, cathleen.
It Questions, Summary Questions, and the new Synthesize sullivan@pearsoned.com

PREFACE xxv
New and Featured Content
T his section highlights selected new and featured content blocking quorum sensing. In Chapter 12, the mechanism of
and organizational changes in Campbell BIOLOGY, Third chromosome movement in bacteria has been updated and
more cell cycle control checkpoints have been added, includ-
Canadian Edition.
ing one recently proposed by researchers. Unit 2 also features
the identification of LHON mutations by Eric Shoubridge at
CHAPTER 1 Evolution, the Themes of Biology, McGill University; the International Cancer Genome Consor-
and Scientific Inquiry tium, co-founded by Thomas Hudson, Scientific Director of
To help students focus on the big ideas of biology, we continue the Ontario Institute of Cancer Research; and work on mem-
to emphasize five themes: Organization, Information, Energy brane proteins by Frances Sharom at the University of Guelph.
and Matter, Interactions, and the core theme of Evolution. A new interview featuring Jason Treberg, from the University
Chapter 1 opens with a new introduction to a case study about of Manitoba, is also included.
the Canadian Yew tree.

UNIT 3 GENETICS
UNIT 1 THE CHEMISTRY OF LIFE
In Chapters 13–17, we have incorporated changes that help
In Unit 1, new content engages students in learning this students grasp the more abstract concepts of genetics and their
foundational material. The new Figure 3.7 shows organisms chromosomal and molecular underpinnings. For example, a
affected by loss of Arctic sea ice. Chapter 5 has updates new Visual Skills Question with Figure 13.6 asks students to
on lactose intolerance, trans fats, the effects of diet on blood identify where in the three life cycles haploid cells undergo
cholesterol, protein sequences and structures, and intrinsically mitosis, and what type of cells are formed. Chapter 14 includes
disordered proteins. New Visualizing Figure 5.16 helps students new information from a recent genomic study on the number
understand various ways proteins are depicted. A new Problem- of genes and genetic variants contributing to height. Chapters
Solving Exercise engages students by having them compare 14 and 15 are more inclusive, clarifying the meaning of the
DNA sequences in a case of possible fish fraud. Unit 1 also term “normal” in genetics and explaining that sex is no longer
highlights research by the Department of Fisheries and Oceans, thought to be simply binary. Other updates in Chapter 15
discussion of include new research in sex determination and a technique be-
Figure 3.7 Effects of climate change on
Frances Old- ing developed to avoid passing on mitochondrial diseases. New
the Arctic.
ham Kelsey and Visualizing Figure 16.7 shows students various ways that DNA
thalidomide, as Species that are benefiting from loss of ice:
is illustrated. To help students understand the Beadle and Ta-
well as work by More light and warmer Some fish species, tum experiment, new Figure 17.2 explains how they obtained
waters result in more
Edward Fon and phytoplankton, which
such as capelin,
benefit from nutritional mutants. A new Problem-Solving Exercise asks
are eaten by having more
Kalle Gehring other
isms.
organ- plankton to
eat.
students to identify mutations in the insulin gene and predict
from McGill Species being harmed
by loss of ice: their effect on the protein. Chapters 18–21 are extensively
University on Loss of ice has
Russia
Arctic updated, driven by exciting new discoveries based on DNA se-
reduced feeding
the structure opportunities for
polar bears, who
ocean
Extent of sea ice in Sept. 2014 quencing and gene-editing technology. Chapter 18 has updates
of the parkin hunt from the ice.
Bering
Extent of sea ice in Sept. 1979
on histone modifications, nuclear location and the persistence
protein. A new The Pacific walrus depends
Strait
North Pole of transcription factories, chromatin remodelling by ncRNAs,
on the ice to rest; its
interview featur- fate is uncertain. Greenland long noncoding RNAs (­lncRNAs), the role of ­master regulatory
ing Roberta Black guillemots in Alaska
cannot fly from their nests genes in modifying chromatin structure, and the possible role
Alaska
Hamme, from on land to their fishing
grounds at the edge of p53 in the low incidence of cancer in elephants. Make Con-
of the ice, which is
the University of now too far from land; Canada nections Figure 18.27, “Genomics, Cell Signalling, and Can-
young birds are starving.
Victoria, is also Sea ice in Sept. 2014 cer,” has been expanded to include more information on cell
included. Ice lost from Sept.
1979 to Sept. 2014 signaling. Chapter 19 expands the section that covers bacterial
defences against bacteriophages and describes the CRISPRCas9
system (Figure 19.8); updates include the Ebola, Chikungunya,
UNIT 2 THE CELL and Zika viruses (Figure 19.12) and discovery of the largest
Our main goal for this unit was to make the material more virus known to date. A discussion has been added of mosquito
accessible and inviting to students. New Visualizing Figure transmission of diseases and concerns about the effects of
6.32 shows the profusion of molecules and structures in a cell, global climate change on disease transmission. In Chapter 21,
all drawn to scale. In Chapter 7, a new figure illustrates levels in addition to the usual updates of sequence-related data (speed
of LDL receptors in people with and without familial hyper- of sequencing, number of species’ genomes sequenced, etc.),
cholesterolemia. Chapter 8 includes a beautiful new photo of there are several research updates, including some early results
a geyser with thermophilic bacteria in Figure 8.17, bringing from the new Roadmap Epigenomics Project and results from a
to life the graphs of optimal temperatures for enzyme func- 2015 study focusing on 414 important yeast genes.
tion. Chapter 10 discusses current research trying to genetically Unit 3 also features the work of Stephen Scherer, who pro-
modify rice (a C3 crop) so that it is capable of carrying out duced a detailed annotated map and DNA sequence of human
C4 photosynthesis to increase yields. Chapter 11 includes a chromosome 7; Calvin Harley and the discovery of telomeres;
new Problem-Solving Exercise that guides students through Michael Houghton, whose research team recently developed a
assessing possible new treatments for bacterial infections by vaccine for the hepatitis C virus at the University of Alberta; the

xxvi NEW AND FEATURED CONTENT


Michael Smith Genome Sciences Centre in Vancouver, which
generated the first genome sequence of SARS; Frank Plummer UNIT 5 THE EVOLUTIONARY HISTORY
at the National Microbiology Laboratory in Winnipeg, whose OF BIOLOGICAL DIVERSITY
team sequenced the full genome of H1N1 flu samples; James Till
In keeping with our goal of improving how students inter-
and Ernest McCulloch, the Canadian scientists who discovered
pret and create visual representations in biology, we have
stem cells; and Michael Rudnicki, who led the team that discov-
added a new figure (Figure 26.6, “Visualizing Phylogenetic
ered adult muscle stem cells at the Sprott Centre for Stem Cell
Relationships”) that introduces the visual conventions used
Research in Ottawa. In addition, a range of genomics research
in phylogenetic trees and helps students understand what
in Canada is featured in the updated Exploring Figure 21.6. A
such trees do and don’t convey. Students are also provided
new interview featuring Julie Claycomb, from the University of
many opportunities to practise their visual skills, with more
Toronto, is also included.
than ten new Visual Skills Questions on topics ranging from
interpreting phylogenetic trees to predicting which regions of
a bacterial flagellum are hydrophobic. The unit also contains
UNIT 4 MECHANISMS OF EVOLUTION new content on tree thinking, emphasizing such key points
A major goal for this revision was to strengthen how we help as how sister groups provide a clear way to describe evolution-
students understand and interpret visual representations of ary relationships and how trees do not show a “direction”
evolutionary data and concepts. Toward this end, we have in evolution. Other major content changes include new text
added a new figure (Figure 25.8), “Visualizing the Scale of in Concepts 26.6, 27.4, and 28.1 on the 2015 discovery of
Geologic Time,” and a new figure (Figure 23.11) on gene flow. the Lokiarchaeota, a group of archaea that may represent
Several figures have been revised to improve the presentation the sister group of the eukaryotes, new text and a new figure
of data, including Figure 24.6 (on reproductive isolation in (Figure 26.22) on horizontal gene transfer from prokaryotes to
mosquitofish), Figure 24.10 (on allopolyploid speciation), eukaryotes, new text in Concept 27.6 describing the CRISPR-
and Figure 25.26 (on the origin of the insect body plan). A Cas9 system and a new figure (Figure 27.22) that illustrates
new Problem-Solving Exercise is included in Chapter 24 on one example of how CRISPR-Cas 9 technology has opened
how hybridization may have led to the spread of insecticide new avenues of research on HIV, and new material in Concept
resistance genes in mosquitoes that transmit malaria. The 29.3 describing how early forests contributed to global climate
unit also includes new chapter opening stories in Chapter change (in this case, global cooling). A new Problem-Solving
22 (expanding on the evolution of the bombardier beetle Exercise in Chapter 34 engages students in interpreting data
defence mechanism) and Chapter 23 (on the Vancouver from a study investigating whether frogs can acquire resis-
Island marmot and population change over time). Additional tance to a fungal pathogen through controlled exposure to it.
changes include new text in Concept 22.3 emphasizing how Other updates include the revision of many phylogenies to
populations can evolve over short periods of time, a new table reflect recent phylogenomic data, new chapter-opening sto-
(Table 23.1) highlighting the five conditions required for a ries in Chapter 28 (on the role of heterotrophy in establishing
population to be in Hardy-­Weinberg equilibrium, and new endosymbioses), Chapter 31 (on how mycorrhizae link trees
material in Concept 25.1 describing how researchers recently of different species on the importance of yeast in creating
succeeded for the first time in constructing a “protocell” ethanol, an important biofuel.) and Chapter 33 (on the visual
in which replication of a template strand of RNA could perception by the eyes of the blue-eyed scallop). There is also
occur. Unit 4 includes new text and a new figure (Figure 34.38) on the adaptations of
updated data on MRSA Figure 23.11 Gene flow and local the kangaroo rat to its arid environment, and new material in
incidence at Canadian adaptation in the Lake Erie water Concept 34.7, including a new figure (Figure 34.52) describing
hospitals, and profiles snake (Nerodia sipedon). fossil and DNA evidence indicating that humans and Nean-
the research of Darla derthals interbred, producing viable offspring. The discussion
Unbanded
Zelenitsky at the ONTARIO
N. sipedon of human evolution also includes new text and a new figure
(Pattern D)
University of Calgary Detroit (Figure 34.54) on Homo naledi, the most recently discov-
on the discovery of a LAKE ERIE
Pelee ered member of the human evolutionary lineage. This unit
Island
winged dinosaur with also highlights research on mycorrhizal networks by Suzanne
Middle
feathers in the Badlands OHIO Cleveland Island Simard at the University of British Columbia; research on early
of Alberta, the research BASS eukaryotic evolution by Patrick Keeling at the University of
ISLANDS
of Hans Larsson from LAKE ERIE British Columbia; data from COSEWIC (Committee on the
McGill University on Status of Endangered Wildlife in Canada), a profile of the Banff
phenotype plasticity Kelleys
Island spring snail,
in tetrapods, and the OHIO and endangered Figure 34.54 Fossils of hand bones and
research of Charles Banded N. sipedon
5 km
species; the foot bones (top and side views) of Homo
(Pattern C)
Henderson and others Hydrocarbon naledi.
who pinpointed the 100 Metagenome
Percentage of individuals

end-Permian mass 80 projects run out


extinction. A new 60 of the University
interview featuring 40 of Calgary and
Maydianne Andrade, 20 the University
from the University of 0
A B C D A B C D A B C D
of Alberta, and
Toronto Scarborough, Ohio mainland Islands Ontario mainland the Wildlife
is also included. Banding patterns in snake populations DNA Forensic

NEW AND FEATURED CONTENT  xxvii


Laboratory at Trent University. A new interview featuring Laura animals (Concept 42.7), the sensation of thirst (Concept 44.4)
Hug, from the University of Waterloo, is also included. and the locomotion of kangaroos and jellies (Concept 50.6).
Furthermore, new text and figures introduce students to
cutting-edge technology relating to such topics as RNA-based
UNIT 6 PLANT FORM AND FUNCTION antiviral defence in invertebrates (Figure 43.4) and rapid, com-
A major aim in revising Chapter 35 was to help students better prehensive characterization of viral exposure (Figure 43.24), as
understand how primary and secondary growth are related. well as recent discoveries regarding brown fat in adult humans
New Visualizing Figure 35.11 enables students to picture growth (Figure 40.14), the microbiome (Figure 41.18), parthenogen-
at the cellular level. Also, the terms protoderm, procambium, esis (Concept 46.1), and magnetoreception (Concept 50.1). As
and ground meristem have been introduced to underscore the always, there is fine-tuning of pedagogy, as in discussions of the
transition of meristematic to mature tissues. A new flowchart complementary roles of inactivation and voltage gating of ion
(Figure 35.24) summarizes growth in a woody shoot. New text channels during action potential formation (Concept 48.3) and
and a figure (Figure 35.26) focus on genome analysis of Ara- of the experimental characterization of genetic determinants in
bidopsis ecotypes, relating plant morphology to ecology and bird migration (Figure 51.24). Additional research highlighted
evolution. In Chapter 36, new Figure 36.8 illustrates the fine in this unit includes Janet Rossant at the University of Toronto
branching of leaf veins, and information on phloem-xylem on cell fate determination; Naweed Syed at the University of
water transfer has been updated. New Make Connections Calgary on synaptic repair; University of British Columbia
Figure 37.10 highlights mutualism across kingdoms and do- researchers exploring the impact of global warming trends on
mains. Figure 37.13 and the related text include new findings on salmon; University of Manitoba explores the evolution of the
how some soil nitrogen derives from weathering of rocks. New thermogenin gene in mammals; Karen Kidd at the University
Figure 38.3 clarifies how the terms carpel and pistil are related. of New Brunswick on environmental estrogens; Barrie Frost of
The text on flower structure and the angiosperm life cycle figure Queen’s University, who explored the navigational mechanisms
identify carpels as megasporophylls and stamens as microsporo- used by monarch butterflies; and Suzie Currie of Mount Allison
phylls, correlating with the plant evolution discussion in Unit 5. University on phenotypic plasticity and environmental stress.
In Concept 38.3, the current problem of glyphosate-resistant A new interview featuring Matt Vijayan, from the University of
crops is discussed in detail. A revised Figure 39.7 helps students Calgary, is also included.
visualize how cells elongate. Figure 39.8 now addresses apical
dominance in a Guided Tour format. Information about the role Figure 41.18 Variation in human gut microbiome at
of sugars in controlling apical dominance has been added. In different life stages.
Concept 39.4, a new Problem-Solving Exercise highlights how
global climate change affects crop productivity. Figure 39.26 on
defence responses against pathogens has been simplified and
On breastmilk Healthy Healthy 65–80
improved. years old

Amongst others, we highlight the work of Rob Guy at the


University of British Columbia on balsam poplar trees; Doug On formula On antibiotic Obese >100

Larson at the University of Guelph on cedars growing out of the treatment years old

rock face of the Niagara Escarpment; R. Keith Downey at the Key


Firmicutes
Ministry of Agriculture in Saskatoon and Baldur Stefansson at On solid food Malnourished Bacteroidetes
Actinobacteria
the University of Manitoba in Winnipeg on canola oil, and Mark Proteobacteria

Belmonte at the University of Manitoba on disease resistance in Other bacterial


phyla
Infant Toddler Adult Elderly person
plants. An Inquiry Figure features the work of Bruce Greenberg
and Bernie Glick at the University of Waterloo on the possible
effects of soil bacteria. A new interview featuring Jacqueline
Monaghan, from Queen’s University, is also included. UNIT 8 ECOLOGY
The Ecology Unit has been extensively revised for the Third
Canadian Edition. We have reorganized and improved the con-
UNIT 7 ANIMAL FORM AND FUNCTION ceptual framework with which students are introduced to the
A major goal of the Unit 7 revision was to transform how stu- following core ecological topics: life tables, per capita popula-
dents interact with and learn from representations of anatomy tion growth, intrinsic rate of increase (“r”), exponential popu-
and physiology. For example, gastrulation is now introduced lation growth, logistic population growth, density dependence,
with a Visualizing Figure (Figure 47.8) that provides a clear and species interactions (in particular, parasitism, commensalism,
carefully paced introduction to three-dimensional processes and mutualism), and MacArthur and Wilson’s island biogeog-
that may be difficult for students to grasp. In addition, a number raphy model. The revision also includes a deeper integration of
of the new and revised figures help students explore spatial evolutionary principles, including a new Key Concept (52.5)
relationships in anatomical contexts, such as the interplay of and two new figures (Figures 52.20 and 52.21) on the reciprocal
lymphatic and cardiovascular circulation (Figure 42.15) and effects of ecology and evolution, new material in Concept 52.4
the relationship of the limbic system to overall brain structure on how the geographic distributions of species are shaped by
(Figure 49.14). A new Problem-Solving Exercise in Chapter 45 a combination of evolutionary history and ecological factors,
taps into student interest in medical mysteries through a case and five new Make Connections Questions that ask students
study that explores the science behind laboratory testing and to examine how ecological and evolutionary mechanisms inter-
diagnosis. Content updates help students appreciate the con- act. In keeping with our goal of expanding and strengthening
tinued evolution of our understanding of even familiar phe- our coverage of climate change, we have added a new discus-
nomena, such as the evolution of hemoglobin in high altitude sion and a new figure (Figure 52.20) on how climate change has

xxviii NEW AND FEATURED CONTENT


affected the distribution of a keystone species, a new section of
Figure 55.9 Climate change,
text in Concept 55.2 on how climate change affects NPP, a new wildfires, and insect outbreaks.
figure (Figure 55.9) on how climate change has caused an in-
crease in wildfires and insect outbreaks, a new Problem-Solving
Exercise in Chapter 55 that explores how insect outbreaks
induced by climate change can cause an ecosystem to switch
from a carbon sink to a carbon source, a new figure (Figure
56.29) on the greenhouse effect, new text in Concept 56.4 on
biological effects of climate change, and a new Unit 8 Make 104

Area burned by wildfires


Connections Figure on how climate change affects all levels of 2

Drought-stress index:
biological organization. Additional updates include a new figure 103

(km2, log scale):


(Figure 53.26) on per capita ecological footprints, a new chapter-
opening story in Chapter 54 on a seemingly unlikely mutualism 102
0
between a shrimp and a much larger predatory fish, new text
in Concept 54.1 emphasizing that each partner in a mutualism 10
experiences both benefits and costs, new text in Concept 54.1
1
describing how the outcome of an ecological interaction can 22
change over time, two new figures (Figures 54.29 and 54.30) on
the island equilibrium model, a new figure (Figure 54.31) docu-
104

Drought-stress index:
menting two shrew species as unexpected hosts of the Lyme

Area affected by
1

(km2, log scale):


disease, new text in Concept 56.1 comparing extinction rates

bark beetles
today with those typically seen in the fossil record, and a new 103
discussion and figure (Figure 56.20) on the restoration of a de- 0
graded urban stream. Unit 8 also profiles the research of David 102
Schindler from the University of Alberta, and Verena Tunnicliffe –1
from the University of Victoria. A new interview featuring Erin
Bertrand, from Dalhousie University, is also included. The book 1997 2000 2005 2010 2014
ends on a hopeful note, charging students to use biological Year
knowledge to help solve problems and improve life on Earth.

NEW AND FEATURED CONTENT xxix


See the Big Picture
KEY CONCEPTS
Each chapter is organized around a framework of 3 to 7 Key Concepts that focus on
the big picture and provide a context for the supporting details.

Every chapter opens


with a visually
Osmoregulation and Excretion
44
dynamic photo
accompanied by
an intriguing
question that
invites students
into the chapter.

David Wall/Alamy Stock Photo


Figure 44.1 How does an albatross drink salt water without ill effect?

The List of Key KEY CONCEPTS A Balancing Act


Seabirds, such as the wandering albatross (Diomedea exulans) (Figure 44.1), spend
Concepts 44.1 Osmoregulation balances the
uptake and loss of water and most of their lives living near the ocean, eating marine organisms, and drinking
seawater. Birds and reptiles have evolved unique adaptations that permit them to
solutes

introduces the 44.2 An animal’s nitrogenous wastes


reflect its phylogeny and habitat
tolerate a high-salt diet and maintain the osmolarity of the fluids in a range similar
to your own. In addition, ions that are abundant in seawater, such as sodium and

big ideas covered 44.3 Diverse excretory systems are


variations on a tubular theme
calcium, must be eliminated to maintain their internal levels within a range that per-
mits normal function of muscles, neurons, and other cells of the body. Homeostasis
thus requires osmoregulation, the general term for the processes by which ani-
in the chapter. 44.4 The nephron is organized for
stepwise processing of blood
mals control solute concentrations and balance water gain and loss.
A number of mechanisms for water and solute control have arisen during evo-
filtrate
lution, reflecting the varied and often severe osmoregulatory challenges presented
44.5 Hormonal circuits link kidney by an animal’s surroundings. The arid environment of a desert, for instance, can
function, water balance, and
quickly deplete an animal of body water. Despite a quite different environment, ma-
blood pressure
rine animals also face potential dehydration. The success of animals in an ocean en-
vironment depends critically on conserving water and, for marine birds and fishes,
eliminating excess salts. In contrast, freshwater animals live in an environment that
Peter van Dam/Shutterstock
threatens to flood and dilute their body fluids. These organisms survive by
conserving solutes and absorbing salts from their surroundings.
In safeguarding their internal fluid environment, animals
must also deal with a hazardous metabolite produced by the

After reading a Key Concept section, students


can check their understanding using the
Concept Check Questions.
Questions throughout
the chapter encourage
students to read the CONCEPT CHECK 44.5
text actively. 1. How does alcohol affect regulation of water balance in
the body?
2. Why could it be dangerous to drink a very large amount
What if? Questions of water in a short period of time?
ask students to apply 3. WHAT IF? Conn’s syndrome is a condition caused by
tumours of the adrenal cortex that secrete high amounts
what they’ve learned. of aldosterone in an unregulated manner. What would
you expect to be the major symptom of this disorder?

Make Connections For suggested answers, see Appendix A.

Questions ask
students to relate
content in the chapter
to material presented
earlier in the course.

xxx SEE THE BIG PICTURE


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no related content on Scribd:
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