Animal Physiology Pearson

Principles of
Animal Physiology
T HIRD EDIT ION
Christopher D. Moyes, Ph.D.

Queen’s University
Patricia M. Schulte, Ph.D.

University of British Columbia
Toronto
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Library and Archives Canada Cataloguing in Publication

Moyes, Christopher D., 1960-, author
Principles of animal physiology / Christopher D. Moyes, Ph.D.
(Queen’s University), Patricia M. Schulte, Ph.D. (University of British
Columbia). — Third edition.
Includes bibliographical references and index.
ISBN 978-0-321-83817-9 (pbk.)
1. Physiology—Textbooks. I. Schulte, Patricia M., 1965-, author II. Title.
QP31.2.M69 2014 571.1 C2014-906141-2
10 9 8 7 6 5 4 3 2 1 [WC]
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Brief Contents
Part One Part Two Part Three
Introduction to The Cellular Basis of Integrating Physiological
Physiology 2 Animal Physiology 98 Systems 256
Photo source: Scott Nielsen/Bruce Coleman Inc. Photo source: Francois Paquet-Durand / Science Source Photo source: Image Quest Marine
CHAPTER 1 CHAPTER 4 CHAPTER 7

Introduction to Physiological Cell Signaling and Endocrine Sensory Systems 256
Principles 2 Regulation 98
CHAPTER 8
CHAPTER 2 CHAPTER 5 Functional Organization of
Physiological Evolution of Neuron Structure and Nervous Systems 310
Animals 20 Function 154
CHAPTER 9
CHAPTER 3 CHAPTER 6 Circulatory Systems 356
Chemistry, Biochemistry, Cellular Movement and
and Cell Physiology 38 Muscles 208 CHAPTER 10
Immune Systems 414
CHAPTER 11
Respiratory Systems 442
CHAPTER 12
Locomotion 498
CHAPTER 13
Ion and Water Balance 542
CHAPTER 14
Digestion and Energy
Metabolism 592
CHAPTER 15
Thermal Physiology 634
CHAPTER 16
Reproductive Physiology 668
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About the Authors
Christopher D. Moyes, Ph.D. Patricia M. Schulte, Ph.D.
Queen’s University University of British Columbia
Chris Moyes received his Ph.D. Trish Schulte received her Ph.D. in
in Zoology from the University Biological Sciences from Stanford
of British Columbia in the area of University in the area of evolution-
comparative muscle physiology. ary physiology. After graduating,
After postdoctoral fellowships in she took a position as an assistant
molecular physiology at the U.S. professor in the Department of
National Institutes of Health and Biology at the University of

Simon Fraser University, he took Waterloo, and then moved to the
a position at Queen’s University, where he is a Full Profes- Department of Zoology at the University of British Columbia
sor in the Department of Biology, and Department Head in Vancouver where she is currently a Full Professor.
(2013–2015). He teaches a spectrum of courses in animal Research in her laboratory focuses on the mechanisms
physiology, comparative biochemistry, and cell biology. Us- that fish use to respond to environmental stressors such as
ing a wide range of comparative and traditional models, his high temperature, hypoxia, and altered salinity. She is particu-
research addresses questions in molecular physiology and larly interested in understanding how genetic variation among
metabolic biochemistry. One major theme of his research individuals contributes to variation in their stress response
is the study of the evolutionary and developmental origins across multiple levels of biological organization, and assess-
of variation in muscle structure and function. Another ma- ing the consequences of this variation for performance and
jor area of his research is the response of animals to envi- fitness in variable environments. Dr. Schulte’s research group
ronmental stress. In all of his research he emphasizes the also conducts applied research in fisheries, aquaculture, and
integration of physiological processes, from molecular to aquatic toxicology. She has published over 100 peer-reviewed
organismal levels. papers, including contributions to several books.
Dr. Moyes is a recipient of the Ontario Premier’s Re- Dr. Schulte was the President of the Canadian Society of
search Excellence Award. He is a member of the American Zoologists (2007–2008), and is a member of the Society for
Physiological Society and the Canadian Society of Zoologists Integrative and Comparative Biology, The Society for Exper-
and has served on research grant panels for the N atural Sci- imental Biology, and the American Physiological Society. She
ence and Engineering Research Council of Canada and the was the co-editor in chief of the journal Physiological and Bio-
U.S. National Science Foundation. He is also Editor-in-Chief chemical Zoology (2009–2014), and is a member of the editorial
of Comparative Biochemistry and Physiology B Biochemistry. board of the journal Comparative Biochemistry and Physiology.
He has published more than 100 peer-reviewed papers, Dr. Schulte has taught physiology courses at multiple
including contributions to four books. levels, including introductory physiology, comparative phys-
More of his research is detailed on his homepage at iology, and human physiology. She is a recipient of a several
http://post.queensu.ca/~cdm2/. teaching awards, including the UBC Science Undergraduate
Society Award for Excellence in Teaching and the Faculty of
Science Achievement Award for Teaching. She is currently
the departmental director for Life Sciences for the Carl Wie-
man Science Education Initiative at UBC, which is dedicated
to promoting the use of evidence-based approaches to sci-
ence education for undergraduates.
You can learn more about her research and teaching
activities on her homepage at http://www.zoology.ubc.ca/
person/pschulte.
Dedication
Thanks to our families, friends, colleagues, and students for their influence and support during the development of this textbook. We
dedicate this textbook to the memory of Peter Hochachka, an inspiration to comparative physiologists and valued mentor to both of us.
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Contents
Preface xix
Acknowledgments xxxiii
PART ONE INTRODUCTION TO PHYSIOLOGY 2
CHAPTER 1 CHAPTER 2
Introduction to Physiological Evolution
Physiological Principles 2 of Animals 20
OVERVIEW 4 INTRODUCTION 21
UNIFYING THEMES IN PHYSIOLOGY 5 ANIMAL EVOLUTION AND PHYSIOLOGY 22
Integration in Physiology 5 Multicellularity and the Invention of Tissues 23
Animal physiologists study phenomena at multiple levels Placozoans and sponges lack discrete tissues 24
of organization 6 Cnidarians possess true tissues 24
Animal physiologists address basic and applied Bilaterians are triploblastic with some degree of cephalization 25
questions 7 Protostomes and deuterostomes differ in the embryonic origins of
Physics and Chemistry: The Basis the mouth and anus 25
A coelom forms by enterocoely or schizocoely 25
of Physiology 7
Platyhelminthes include parasitic and free-living worms 26
The laws of diffusion help to explain the evolution of animal
Mollusks possess a calcareous shell 27
form and function 7
Annelids have segmented bodies 27
Mechanical theory helps us understand how organisms
work 8 Arthropods show metamerism and tagmatization 27
Electrical potentials are a fundamental physiological Deuterostomes include echinoderms and chordates 28
currency 9 Vertebrates 28
Temperature affects physiological processes 9
Different agnathan ancestors gave rise to modern agnathans
Biochemical and physiological patterns are influenced and fish 28
by body size 9
Cartilaginous fish evolved from placoderms 29
Form, Function, and Evolution 11 Several groups of bony fish evolved in the Devonian period 29
Form and function are the products of evolution 11 Sarcopterygians gave rise to tetrapods 31
Animals have many traits in common 12 Amphibians must return to water to breed 31
What is adaptation? 12 Reptiles and their ancestors have dominated land for 300 million
years 31
Not all differences are evolutionary adaptations 13
Mammals evolved from therapsid reptiles 32
Phenotypes may be homologous or analogous 14
Birds are modern reptiles 32
Regulation and Homeostasis 15
EVOLUTIONARY CONSERVATION AND
Animals can be physiological conformers or
regulators 15 CONVERGENCE IN ANIMAL PHYSIOLOGY 32
Homeostasis is the maintenance of internal Molecular Innovations 33
constancy 15 The myosin gene family divergence underlies much of animal
Feedback loops control physiological pathways 16 diversity 33
Negative feedback loops maintain homeostasis 17 Na+/K+ ATPase is essential for ion homeostasis and excitable
Positive feedback loops cause explosive responses 17 tissues 33
Acclimation and acclimatization result in reversible phenotypic The appearance of collagen coincided with tissue formation 35
changes 17 Hormones extended the range of cell-to-cell signaling 35
Animals can also irreversibly alter their phenotype 17
Integrative Processes 36
Summary 18 • Review Questions 19 • Synthesis The evolution of complexity was accompanied by an increase
Questions 19 in cephalization 36
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Terrestriality arose in multiple lineages 36 Gluconeogenesis builds glucose from noncarbohydrate

Metabolic pathways are broadly conserved, though metabolic precursors 62
rate varies widely 36 Glycolysis is a low-efficiency, high-velocity pathway 63
Terminal dehydrogenases oxidize NADH under anaerobic
Summary 37 • Review Questions 37 • Synthesis Questions 37 conditions 65
Lipids 66
Fatty acids are long aliphatic chains produced from acetyl CoA 66
Fatty acids are oxidized in mitochondrial b-oxidation 67
CHAPTER 3
Fatty acids can be converted to ketone bodies 68
Chemistry, Biochemistry, Triglyceride is the major form of lipid storage 68
Phospholipids predominate in biological membranes 69
and Cell Physiology 38 Steroids share a multiple ring structure 70
OVERVIEW 39 Mitochondrial Metabolism 71

CHEMISTRY 39 The TCA cycle uses acetyl CoA to generate reducing
equivalents 71
Energy 39 The ETS generates a proton gradient, heat, and reactive oxygen
Food webs transfer energy 40 species 72
Energy is stored in electrochemical gradients 41 The F1Fo ATPase uses the proton motive force to generate
Chemical energy is transferred in chemical reactions 42 ATP 73
Covalent bonds involve shared electrons 42 Creatine phosphokinase enhances energy stores and
Weak bonds control macromolecular structure 43 transfer 74
Weak bonds are sensitive to temperature 44 Integration of Pathways of Energy Metabolism 74
Reaction rates are influenced by temperature 44 Oxygen and ATP control the balance between glycolysis
and oxidative phosphorylation 75
Properties of Water 45
Physical properties of fuels influence fuel selection 75
The properties of water are unique 45
Fuel selection can be calculated from the respiratory quotient 75
Solubility determines how much solute can dissolve in
water 46 Energetic intermediates regulate the balance between anabolism
and catabolism 76
Solutes influence the physical properties of water 46
Metabolic rate is the sum of all chemical reactions 76
Solutes move through water by diffusion 46
Solutes in biological systems impose osmotic pressure 47 CELL PHYSIOLOGY 77
Differences in osmolarity can alter cell volume 48 Membrane Structure and Transport 77
Acids and bases alter the pH of water 49 The lipid profile affects membrane properties 77
Both pH and temperature affect the ionization of biological Lipid membranes are heterogeneous 77
molecules 50
Environmental stress can alter membrane fluidity 78
Buffers limit changes in pH 50
Membranes possess integral and peripheral proteins 78
BIOCHEMISTRY 51 Many molecules must move across cellular membranes 78
Membrane proteins can facilitate the diffusion of impermeant
Enzymes 52 molecules 79
All chemical reactions are governed by the laws of Active transporters use energy to pump molecules against
thermodynamics 52 gradients 80
Enzymes accelerate reactions by reducing the reaction activation The movement of charged molecules depends on the
energy 53 electrochemical gradient 81
Enzyme kinetics describe enzymatic properties 53 The Nernst equation allows calculation of the equilibrium
The physicochemical environment alters enzyme kinetics 55 potential 82
Allosteric and covalent regulation control enzymatic rates 56 Cells maintain a resting membrane potential 82
Enzymes convert nutrients to reducing energy 57 The Na+/K+ATPase establishes concentration gradients 82
ATP is a carrier of free energy 57 The membrane potential represents a balance between
equilibrium potentials 82
Proteins 58
The Goldman equation can be used to calculate membrane
Proteins are polymers of amino acids 58 potential 83
Proteins are folded into three-dimensional shapes 59 Changes in membrane permeability alter membrane
Molecular chaperones help proteins fold 60 potential 83
Carbohydrates 60 Cellular Organization 84

Animals use monosaccharides for energy and biosynthesis 60 Mitochondria are the powerhouse of the cell 84
Complex carbohydrates perform many functional and structural The cytoskeleton controls cell shape and directs intracellular
roles 61 movement 85
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The endoplasmic reticulum and Golgi apparatus mediate DNA is organized into genomes 91
vesicular traffic 85 Transcriptional control acts at gene regulatory regions 91
The extracellular matrix mediates interactions between cells 86 RNA degradation influences RNA levels 92
Most tissues are composed of multiple cell types 88 Global changes in translation control many pathways 93
Epithelial tissues share four specialized properties that affect Cells rapidly reduce protein levels through protein
solute movements 88 degradation 93
Solutes move across epithelial tissues by paracellular and Protein variants arise through gene duplications and
transcellular transport 89 rearrangements 93
Ancient genome duplications contribute to physiological
Physiological Genetics and Genomics 90 diversity 95
Nucleic acids are polymers of nucleotides 90
DNA is a double-stranded a-helix packaged into Summary 96 • Review Questions 96 • Synthesis Questions 97
chromosomes 90 • Quantitative Questions 97
PART TWO THE CELLULAR BASIS OF ANIMAL PHYSIOLOGY 98
CHAPTER 4 Receptor type determines the cellular response 113

Receptors have several domains 113
Cell Signaling and A ligand may bind to more than one receptor 114
Endocrine Regulation 98
Ligand-receptor binding obeys the law of mass action 114
Receptor number can vary 114
Receptor affinity for a ligand can vary 115
OVERVIEW 99
Ligand signaling must be inactivated 115
THE BIOCHEMICAL BASIS OF CELL
• Box 4.2 MATH IN PHYSIOLOGY
SIGNALING 101
Ligand-Receptor Interactions 116
General Features of Cell Signaling 101
SIGNAL TRANSDUCTION PATHWAYS 118
Indirect signaling systems form a continuum 102
The structure of the messenger determines the type of signaling Intracellular Receptors 119
mechanism 103
Ligand-Gated Ion Channels 121
Peptide Messengers 104
Signal Transduction via Receptor-Enzymes 121
Peptide messengers are released by exocytosis 104
Receptor guanylate cyclases generate cyclic GMP 122
Peptide messengers dissolve in extracellular fluids 105
Receptor tyrosine kinases signal through Ras proteins 123
Peptides bind to transmembrane receptors 105
Receptor serine/threonine kinases directly activate
Steroid Messengers 106 phosphorylation cascades 124
Steroids bind to carrier proteins 107 Signal Transduction via G Protein–Coupled
• Box 4.1 CHALLENGES TO HOMEOSTASIS Receptors 124
Endocrine Disruptors 108 G protein–coupled receptors are extremely diverse 125
Steroids bind to intracellular receptors 109 G proteins can act through Ca2+-calmodulin 126
G proteins can interact with amplifier enzymes 126
Biogenic Amines 109
Amplifier enzymes alter the concentration of second
Thyroid hormones diffuse across the membrane 110 messengers 126
Thyroid hormones are hydrophobic messengers 110 Guanylate cyclase generates cGMP 127
Other Classes of Messenger 111 Phospholipase C generates phosphatidylinositol 127
Eicosanoids are lipid messengers 111 Cyclic AMP was the first second messenger discovered 128
There are three known gaseous chemical messengers 111 Signal transduction pathways can interact 130
Purines can act as neurotransmitters and paracrines 112 INTRODUCTION TO ENDOCRINE SYSTEMS 130
Communication of the Signal to the Target Cell 112 Characteristics of Endocrine Systems 130
Ligand-receptor interactions are specific 113 Hormone levels are regulated by feedback loops 133
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The actions of insulin illustrate the principle of negative CHAPTER 5

feedback 133
The actions of oxytocin illustrate the principle of postive
feedback 133
Neuron Structure
Feedback loops can be complex 134 and Function 154
Pituitary hormones provide examples of several types of feedback
loops 135 OVERVIEW 155
Neurohormones from the posterior pituitary are involved in SIGNALING IN A VERTEBRATE MOTOR
first-order feedback loops 135
NEURON 157
The hypothalamus regulates the secretion of anterior pituitary
hormones 135 Electrical Signals in Neurons 157
Prolactin is an anterior pituitary hormone involved in Ionic concentration gradients and permeability establish
a second-order feedback loop 136 membrane potential 157
Many anterior pituitary hormones participate in third-order The Nernst equation can be used to calculate the equilibrium
pathways 136 potential of an ion 158
A single hormone can be regulated by multiple types of feedback The Goldman equation is used to calculate the membrane
pathways 137 potential 159
Antagonistic hormone pairs provide precise The Na+/K+ ATPase maintains the membrane potential 159
regulation 138
Hormones can demonstrate additivity and
Using the Nernst and Goldman Equations 160
synergism 139
Hormone levels are influenced by both synthesis and Changes in membrane permeability cause electrical signals 160
removal 140 Ion channel function can be studied using the voltage clamp 162
Endocrine pathologies occur when hormone levels are Signals in the Dendrites and Cell Body 162
disregulated 140
Graded potentials vary in magnitude 162
The responsiveness of the target cell can vary 140
Graded potentials are short-distance signals 163
• Box 4.3 APPLICATIONS Graded potentials can trigger action potentials at the axon
Cell-To-Cell Communication and Diabetes hillock 164
Mellitus 141 Graded potentials can be integrated across time and space 165
The nervous and endocrine systems interact in the stress
Signals in the Axon 166
response 142
Voltage-gated channels generate the action potential 167
The hypothalamo-pituitary axis is involved in the stress
response 143 Voltage-gated Na+ channels open at the threshold potential 167
A positive feedback loop drives the depolarization phase 168
Evolution of Endocrine Systems 144
Voltage-gated Na+ channels have two gates 168
Endocrine systems vary in complexity among animal Voltage-gated K+ channels open slowly 169
phyla 144
Hyperglycemic hormones are an example of an invertebrate • Box 5.2 APPLICATIONs
neurohormone 144 Voltage-Gated Na+ Channel Blockers 170
The major steroid hormones differ between vertebrates Both Na+ and K+ shape the action potential 171
and arthropods 145 Action potentials transmit signals across long distances 172
Arthropods also use other terpenoids as Vertebrate motor neurons are myelinated 174
hormones 146
Axons conduct action potentials unidirectionally 174
The structure and function of many hormones is highly
Action potential frequency carries information 176
conserved in vertebrates 146
Some hormonal pathways have evolved via gene Signals Across the Synapse 176
duplication 147 Intracellular Ca2+ regulates neurotransmitter release 176
Some hormones have acquired new functions during vertebrate Action potential frequency influences neurotransmitter
evolution 147 release 177
Some hormone pathways are reduced in Acetylcholine is the primary neurotransmitter at the vertebrate
humans 148 neuromuscular junction 178
The structure of endocrine glands varies among the Signaling is terminated by acetylcholinesterase 178
vertebrates 148
Postsynaptic cells express specific receptors 179
The structure of the pituitary differs among
Neurotransmitter amount and receptor activity influence signal
vertebrates 149
strength 179
The neurohormones of the posterior pituitary vary among
vertebrates 149 DIVERSITY OF NEURAL SIGNALING 180
Summary 151 • Review Questions 152 • Synthesis Questions 152 Structural Diversity of Neurons 180
• Quantitative Questions 153 Neurons can be classified based on their function 180
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Neurons can be classified based on their structure 180 • Box 6.1 APPLICATIONs
Neurons are associated with glial cells 182 Pharmaceutical Uses of Microtubule Disruptors 215
Diversity of Signal Conduction 184 Kinesin and dynein move along microtubules 215
Cilia and flagella are composed of microtubules 216
Voltage-gated ion channels are encoded by multiple genes 184
Voltage-gated Ca2+ channels can also be involved in action Microfilaments 217
potentials 185 Microfilaments are polymers of actin 217
Conduction speed varies among axons 185 Actin polymerization can generate movement 218
The cable properties of the axon influence current flow 185 Actin uses myosin as a motor protein 218
• Box 5.3 CHALLENGES TO HOMEOSTASIS The sliding filament model describes actino-myosin activity 219
RNA Editing of Potassium Channels as an Adaptation Myosin activity is influenced by unitary displacement and duty
to Cold Environments 186 cycle 222
Intracellular and membrane resistance influence conduction
MUSCLE 223
speed 188
Membrane capacitance influences the speed of General Features of Striated Muscles 223
conduction 189 Muscle cells possess thick and thin filaments 224
Giant axons have high conduction speed 191 Striated muscle thick and thin filaments are arranged into
Myelinated neurons evolved in the vertebrates 192 sarcomeres 224
Myelination increases conduction speed 193 Myosin II has a unique duty cycle and unitary displacement 226
Sarcomeric organization determines contractile properties of the
Diversity of Synaptic Transmission 194 muscle cell 226
Electrical and chemical synapses play different roles 194 Actino-myosin activity is activated by Ca2+ 228
Chemical synapses have diverse structures 195 Thick filaments also influence contractile properties 229
There are many types of neurotransmitters 196 Muscle contraction can generate force 230
Neurotransmitters can be excitatory or inhibitory 198 Cardiac and skeletal muscle cells differ in some structural
Neurotransmitter receptors can be ionotropic or properties 231
metabotropic 198
Acetylcholine receptors can be ionotropic or metabotropic 198 • Box 6.2 MATH IN PHYSIOLOGY
Factors Affecting Force, Work, and Power 232
The biogenic amines play diverse physiological roles 199
Neurons can synthesize more than one kind of Excitation in Vertebrate Skeletal and Cardiac
neurotransmitter 201 Muscles 234
Neurotransmitter release varies depending on physiological
Striated muscles are all activated by an action potential 234
state 201
Striated muscles differ in the time course of the action
Evolution of Neural Signaling 203 potential 235
Many organisms use electrical signaling 203 Cardiac and skeletal muscles differ in refractory periods 236
Action potentials in nonmetazoans involve Ca2+ 204 Skeletal muscle excitation is triggered by neurotransmitters 237
Animals have unique voltage-gated Na+ channels 204 T-tubules enhance action potential penetration into the myocyte 238
Neurotransmitters evolved from ancient signaling Cardiac muscle cells are stimulated by other muscle cells 238
molecules 204
Excitation-Contraction Coupling in Striated Muscles 239
Summary 205 • Review Questions 205 • Synthesis Questions 206 Depolarization leads to an increase in cytoplasmic [Ca2+] 239
• Quantitative Questions 207 DHPR activation induces Ca2+ release from the SR 240
Relaxation follows removal of Ca2+ from the cytoplasm 241
Many factors contribute to differences in properties of striated
muscles 242
CHAPTER 6 DIVERSITY IN MUSCLE STRUCTURE
Cellular Movement AND FUNCTION 243
and Muscles 208 Smooth Muscle 243

• Box 6.3 CHALLENGES TO HOMEOSTASIS
OVERVIEW 209 Remodeling Muscle in Response to Changing
Conditions 244
CYTOSKELETON AND MOTOR PROTEINS 210
Smooth muscle lacks organized sarcomeres 244
Microtubules 210
Smooth muscle contraction is regulated by both thick and thin
Tubulin is composed of α-tubulin and β-tubulin 212 filament proteins 246
Microtubules show dynamic instability 213 Latch cross-bridges maintain smooth muscle contraction for long
Microtubule polarity determines the direction of movement 214 periods 247
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Invertebrate Muscles 248 Specialized Muscles and Transdifferentiated

Many invertebrates possess obliquely striated muscle 248 Muscle 252
Invertebrate muscles contract in response to graded excitatory Sonic muscles produce rapid contractions but generate less
postsynaptic potentials 249 force 252
Asynchronous insect flight muscles do not use Ca2+ Heater organs and electric organs are modified muscles 253
transients 249
Mollusk catch muscles maintain contraction for long Summary 254 • Review Questions 254 • Synthesis Questions 255
periods 251 • Quantitative Questions 255
PART THREE INTEGRATING PHYSIOLOGICAL SYSTEMS 256
CHAPTER 7 Coding differs between the olfactory and gustatory systems 272
Taste reception differs between vertebrates and
Sensory Systems 256 invertebrates 272
Nociceptors detect noxious chemical stimuli 273
(a)
MECHANORECEPTION 273
OVERVIEW 257
Touch and Pressure Receptors 274
GENERAL PROPERTIES OF SENSORY Vertebrate tactile receptors are widely dispersed 274
RECEPTION 257 Vertebrate proprioceptors monitor body position 275
Classification of Sensory Receptors 259 Insects have several types of tactile and proprioceptors 275
(b)
Receptors can be classified based on stimulus location or Equilibrium and Hearing 276
modality 259
Statocysts are the organ of equilibrium for invertebrates 276
Receptors may detect more than one stimulus modality 260
Insects use a variety of organs for hearing 276
Stimulus Encoding in Sensory Systems 260 Vertebrate organs of hearing and equilibrium contain hair cells 277
Sensory pathways encode stimulus modality 260 Tip links are critical for mechanosensory transduction 280
Receptive fields provide information about stimulus location 261 Hair cells are found in the lateral line and ears of fish 280
Sensory receptors have a dynamic range 261 Vertebrate ears function in hearing and equilibrium 280
There is a trade-off between dynamic range and The vestibular apparatus is the organ of equilibrium in
discrimination 262 vertebrates 280
Range fractionation increases sensory discrimination 262 The inner ear detects sounds 282
Sense organs can have a very large dynamic range 263 In terrestrial vertebrates, hearing involves the inner, middle,
Many receptors encode signals logarithmically 263 and outer ears 284
Tonic and phasic receptors encode stimulus duration 264 Cetaceans have highly modified ears 285
The inner ear of mammals has specializations for sound
CHEMORECEPTION 265 detection 286
The Olfactory System 265 Outer hair cells amplify sounds 286
The ears can detect sound location 287
The vertebrate olfactory system can distinguish thousands of
odorants 265 PHOTORECEPTION 287
Odorant receptors are G protein coupled 265
Photoreceptors 287
An alternative chemosensory system detects pheromones 267
The structure of photoreceptor cells differs among animals 287
Invertebrate olfactory mechanisms differ from those
in vertebrates 268 Mammals have two types of photoreceptor cells 288
Chromophores allow photoreceptors to absorb light 290
• Box 7.1 APPLICATIONs The mechanisms of phototransduction differ among
Using Pheromones to Alter Behavior 269 organisms 291
The Gustatory System 270 The Structure and Function of Eyes 291
Taste buds are vertebrate gustatory receptors 270 There are two major types of compound eyes in arthropods 293
Vertebrate taste receptors use diverse signal transduction Structurally diverse eyes share underlying molecular
mechanisms 270 similarity 294
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The structure of the vertebrate eye relates to its function 294 Brain size varies among vertebrates 321
The lens focuses light on the retina 295 • Box 8.1 MATH IN PHYSIOLOGY
Vertebrate retinas have multiple layers 295 Brain Size and Brain Complexity 322
Information from rods and cones is processed differently 296
The relative sizes of brain regions vary among vertebrates 324
Signal processing in the retina enhances contrast 297
The hindbrain supports basic functions 325
The brain processes the visual signal 298
The midbrain is greatly reduced in mammals 325
Color vision requires multiple types of photoreceptors 300
The forebrain controls complex processes 325
Color vision evolved secondarily in primates 301
The corpus callosum allows communication between
Some photoreceptors are not involved in vision 302 hemispheres 325
Other Sensory Modalities 302 The hypothalamus maintains homeostasis 327
Thermoreceptors detect temperature 302 The limbic system influences emotions 327
The thalamus acts as a relay station 328
The cortex integrates and interprets information 328
Circadian Rhythms in the Modern World 303
The cortex exhibits topographic organization 330
Electroreceptors detect electrical fields 304 Mirror neurons fire in response to observed behaviors 330
Magnetoreceptors detect magnetic fields 305
THE PERIPHERAL NERVOUS SYSTEM
Communication in Weakly Electric Fish 306 OF VERTEBRATES 331
The Autonomic Nervous System 331
Summary 308 • Review Questions 308 • Synthesis Questions 309 The sympathetic and parasympathetic branches act together to
• Quantitative Questions 309 maintain homeostasis 332
The anatomy of the sympathetic and parasympathetic branches
differs 333
The neurotransmitters of the sympathetic and parasympathetic
systems differ 334
CHAPTER 8
Some effectors receive only sympathetic innervation 335
Functional Organization The central nervous system regulates the autonomic nervous
system 336
of Nervous Systems 310 The enteric nervous system regulates the gut 337
OVERVIEW 311 Somatic Motor Pathways 337
ORGANIZATION AND EVOLUTION OF NERVOUS INTEGRATIVE FUNCTIONS OF NERVOUS

SYSTEMS 312 SYSTEMS 337
Evolution of Nervous Systems 312 Coordination of Behavior 338
Sponges lack a nervous system 313 Reflex arcs control many involuntary behaviors 338
Cnidarian nerve nets allow complex behaviors 314 Pattern generators initiate rhythmic behaviors 339
Nervous system complexity varies among flatworms 315 Pattern generators govern swimming behavior in the
Nematodes and annelids show substantial cephalization 315 leech 339
Pattern generators and reflexes are involved in tetrapod
THE ARTHROPOD NERVOUS SYSTEM CONTAINS locomotion 340
SEGMENTAL GANGLIA 315 The brain coordinates voluntary movements 341
Cephalopods have the largest and most complex brains among Communication is a complex behavior 341
invertebrates 316 • Box 8.2 CHALLENGES TO HOMEOSTASIS
The echinoderms lack an obvious brain 316 Ocean Acidification Affects Fish Behavior by Disturbing
Vertebrates have a hollow dorsal nerve cord 317 Brain Homeostasis 342
THE CNS OF VERTEBRATES 317 Learning and Memory 344

Anatomy of the Vertebrate CNS 317 Invertebrates show simple learning and memory 344
The vertebrate CNS is encased in cartilage or bone 317 The hippocampus is important for memory formation in
mammals 346
The meninges surround the CNS 317
The CNS is physiologically separated from the rest of the • Box 8.3 APPLICATIONs
body 318 Functional Magnetic Resonance Imaging and Brain
The CNS contains gray and white matter 319 Plasticity 347
The spinal cord mediates information flow between the brain Regulation and Homeostasis 349
and body 319
Multiple brain regions send inputs to the hypothalamus 349
The Vertebrate Brain 319 The hypothalamus helps maintain ion and water balance 349
Vertebrate brains have three main regions 320 The hypothalamus regulates body temperature 349
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The hypothalamus regulates food intake 349 STRUCTURE AND FUNCTION OF VERTEBRATE
The hypothalamus is involved in the stress response 349 HEARTS 376
The hypothalamus regulates circadian rhythms 350
Heart Anatomy 376
The hypothalamus regulates sleep-wake cycles 351
The myocardium can be spongy or compact 377
Sleep is divided into phases 353
Fish heart chambers are arranged in series 378
Summary 354 • Review Questions 354 • Synthesis Questions 355 Amphibian hearts have three chambers 379
• Quantitative Questions 355 Most reptiles have five heart chambers 379
Crocodilians have completely divided ventricles 380
Birds and mammals have four heart chambers 381
Cardiac anatomy is related to respiratory mode 382
CHAPTER 9 Cardiac anatomy changes during development 383
Circulatory Systems 356 The Cardiac Cycle 384

Fish hearts contract in series 384
The mammalian cardiac cycle is similar to that of fish 384
Some vertebrate hearts fill actively 385
OVERVIEW 357
The right and left ventricles develop different pressures 385
UNITY AND DIVERSITY OF CIRCULATORY
Control of Cardiac Contraction 386
SYSTEMS 358
Pacemaker cells initiate the heartbeat 387
General Characteristics of Circulatory Systems 358 The nervous and endocrine systems can modulate heart
Circulatory systems use diverse pumping structures 358 rate 387
Circulatory systems can be open or closed 359 Pacemaker depolarizations can spread via gap junctions 389
Circulatory systems pump several types of fluids 360 Cardiac action potentials have an extended depolarization
Blood and hemolymph contain proteins 361 phase 389
Blood and hemolymph contain cells 361 Conducting pathways spread the depolarization across
Vertebrate blood has three main components 361 the heart 390
The integrated electrical activity of the heart can be detected
Circulatory Plans of the Major Animal Phyla 362 with the EKG 390
Most annelids have closed circulatory systems 363 The heart functions as an integrated organ 391
Most mollusks have open circulatory systems 363 Cardiac output is the product of heart rate and stroke
Arthropod circulatory systems vary in complexity 363 volume 391
Crustacean hearts are both suction and pressure pumps 364 The nervous and endocrine systems can modulate stroke
Insects have simple open circulatory systems 365 volume 391
Chordates have both open and closed circulatory • Box 9.2 APPLICATIONs
systems 366 Using the EKG to Diagnose Heart Conditions 392
Closed circulatory systems evolved multiple times in End-diastolic volume modulates stroke volume 394
animals 366
REGULATION OF CIRCULATORY FUNCTION 395
The Circulatory Plan of Vertebrates 367
Vertebrate blood vessels have complex walls 368 Regulation of Flow 396
Wall thickness varies among blood vessels 368 The arterioles control blood distribution 396
Blood vessels undergo angiogenesis 370 Myogenic autoregulation maintains blood flow 396
Low oxygen levels can promote angiogenesis 370 Metabolic activity and paracrine signals influence blood
flow 397
Vertebrate circulatory systems contain one or more pumps
in series 371 The nervous and endocrine systems regulate arteriolar
diameter 398
Mammals and birds have completely separated pulmonary
and systemic circuits 371 Regulation of Blood Pressure 399
Many tetrapods have incompletely separated pulmonary The arteries dampen pressure fluctuations 399
and systemic circuits 372
Mean arterial pressure is determined by systolic and diastolic
The Physics of Circulatory Systems 372 pressures 400
The radius of a tube affects its resistance 373 The skeletal muscle and respiratory pumps aid venous return to
The resistance of a vessel determines the flow 373 the heart 400
The veins act as a volume reservoir 401
• Box 9.1 MATH IN PHYSIOLOGY Peripheral resistance influences pressure 401
Poiseuille’s Equation 374
The baroreceptor reflex is the primary means of regulating
Velocity of flow is determined by pressure and cross-sectional MAP 402
area 375 The kidneys also play a role in maintaining blood volume
Pressure exerts a force on the walls of blood vessels 375 and MAP 403
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• Box 9.3 CHALLENGES TO HOMEOSTASIS Cell-Mediated Immunity 430

Hypertension 404 Antigen-presenting cells display fragments of pathogens on the
Blood pressure can force fluid out of the capillaries 404 cell surface 431
The lymphatic system returns filtered fluids to the circulatory T cells recognize MHC; antigen complexes presented on the
system 406 surface of other immune cells 432
Changes in body position can alter blood pressure B cells produce antibodies 433
and flow 407
Changes in body position can cause orthostatic Elevating Antibody Titer 434
hypotension 408
Very tall animals must have specialized circulatory Immune cells move via the lymph 434
systems 408 Lymphocytes mature in lymph nodes 435
The Circulatory System During Exercise 410 Integration with Other Physiological Systems 435
The cardiovascular control center of the brain regulates the Allergic responses are stimulated by mast cells 435
circulatory system 410 Increases in body temperature impede pathogen replication 436
Cardiac output increases during exercise 410 The GI tract has immunological defenses 436
Patterns of blood flow change during exercise 411
Blood pressure changes only slightly during exercise 411 The Immune System and Thermoregulation 438
Higher brain centers are also involved 411
Some species can transfer immunity to offspring 438
Summary 412 • Review Questions 412 • Synthesis Questions 412 Steroid hormone levels affect the immune system 439
• Quantitative Questions 413
Summary 441 • Review Questions 441 • Synthesis Questions 441
CHAPTER 10
CHAPTER 11
Immune Systems 414
Respiratory Systems 442
OVERVIEW 415
INNATE IMMUNITY 418 OVERVIEW 443
Recognition of Pathogens 418 RESPIRATORY STRATEGIES 444
Pattern-recognition receptors detect pathogen-associated
The Physics of Respiratory Systems 444
molecular patterns 418
Gases exert a pressure 445
Toll-like receptors activate immune responses 419
Henry’s law describes how gases dissolve in liquids 445
Phagocytic Cells 420 Gases diffuse at different rates 446
Phagocytic cells engulf and digest foreign cells 420 Fluids flow from areas of high to low pressure 447
Opsonins promote phagocytosis 421 Resistance opposes flow 447
Complement molecules promote other immune processes 422
Types of Respiratory Systems 448
Executing Pathogens in the Innate Immune Very thin animals can rely on diffusion alone for gas exchange 449
System 423 Most animals use one of three major respiratory strategies 449
Granulocytes and natural killer cells secrete cytotoxic Gas-exchange surfaces are often ventilated 450
compounds 423 Perfusion of the respiratory surface affects gas exchange 450
Antimicrobial peptides can be secreted by many cell types in all
multicellular organisms 423 VENTILATION AND GAS EXCHANGE 452
Inflammation is an early response to pathogens and tissue Ventilation and Gas Exchange in Water 453
damage 423 Most mollusks ventilate their gills using cilia 453
ADAPTIVE IMMUNITY OF VERTEBRATES 425 Crustacean gills are located on the appendages 454
Echinoderms have diverse respiratory structures 454
• Box 10.1 APPLICATIONs
Transgenic Mosquitoes 426 Feeding lampreys ventilate their gills tidally 455
Elasmobranchs use a buccal pump for ventilation 455
Humoral Immunity 426 Teleost fishes use a buccal-opercular pump for ventilation 456
Antibodies are composed of variable and constant regions 427 Fish gills are arranged for countercurrent flow 458
Diversity in immunoglobulins arises through gene
recombination 428 Ventilation and Gas Exchange in Air 458
Antibody classes differ in the C regions 429 Arthropods use a variety of mechanisms for aerial gas
Complement molecules interact with immunoglobulins 430 exchange 459
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Tracheal systems are inefficient in water 460 Air breathers can experience high-altitude hypoxia 488
Some aquatic insects breathe through siphons 460 High altitude can cause pathological responses in lowland
Some aquatic insects carry bubbles of air 461 animals 490
Many insects actively ventilate the tracheae 461 Some human populations have colonized high altitudes 490
Air breathing has evolved multiple times in vertebrates 462 High-altitude mammals have various adaptations to function well
Amphibians ventilate their lungs using a buccal force pump 463 in hypoxia 490
Reptiles ventilate their lungs using a suction pump 465 Birds have a greater tolerance of high-altitude hypoxia than do
mammals 490
Birds unidirectionally ventilate their lungs 466
Metabolic suppression is a common response to hypoxia 491
The alveoli are the site of gas exchange in mammals 467
Mammals ventilate their lungs tidally 468 Diving 491
The work required for ventilation depends on lung compliance Anaerobic metabolism takes over at the aerobic dive limit 491
and resistance 469
Surfactants increase lung compliance 469
Adaptations to High Altitude in Bar-Headed Geese 492
Airway resistance affects the work required to breathe 470
Aspiration-based pulmonary systems have substantial dead Diving animals have increased body oxygen stores 494
space 470 Nitrogen narcosis is a problem at depth 494
Pulmonary function tests measure lung function and Decompression sickness can occur on ascent 495
volumes 470 Marine mammals decrease oxygen demand during a dive 495
Diving animals have modified responses to CO2 495
• Box 11.1 APPLICATIONs
Treating Respiratory Distress Syndrome in Premature Summary 496 • Review Questions 496 • Synthesis Questions 496
Infants 471
Ventilation-perfusion matching is important for gas
exchange 472
GAS TRANSPORT TO THE TISSUES 473

Oxygen Transport 473 CHAPTER 12
There are three main types of respiratory pigments 473
Locomotion 498
Pulmonary Function Tests 474
Respiratory pigments have characteristic oxygen equilibrium OVERVIEW 499
curves 476
The shapes of oxygen equilibrium curves differ 478 LOCOMOTOR SYSTEMS 499
Blood pH and PCO2 can affect oxygen affinity 478 Muscle Fiber Types 500
Root-effect hemoglobins help to deliver oxygen to the swim Many invertebrates use simple circular and longitudinal muscles
bladder 479 to move 500
The rete mirabile maintains the low pH and high PO2 at the gas Fish have anatomically separated muscle fiber types 502
gland 480 The pattern of locomotor muscle contraction is controlled by
The Root effect may also assist in delivering oxygen to systemic motor neurons 503
tissues 481 Tetrapods have a multiplicity of fiber types 504
Temperature affects oxygen affinity 481 Locomotor muscles are organized into locomotor modules
Organic modulators can affect oxygen affinity 482 and functional groups 505
Carbon Dioxide Transport 482 Energy Metabolism 506

The carbon dioxide equilibrium curve quantifies carbon dioxide Glycolysis and mitochondria support different types of
transport 482 locomotion 507
Blood oxygenation affects CO2 transport 483 Mitochondrial content influences muscle aerobic
Vertebrate red blood cells play a role in CO2 transport 483 capacity 507
The respiratory system can regulate blood pH 484 Muscle must recover from high-intensity activity 508
Metabolic transitions accompany prolonged exercise 508
REGULATION OF VERTEBRATE RESPIRATORY Hormones control fuel oxidation in muscle 509
SYSTEMS 486
Perfusion and Oxygen Delivery to Muscle 511
Regulation of Ventilation 486 Capillary networks bring oxygen to vertebrate muscle
Chemosensory input influences ventilation 487 fibers 511
Other factors regulate breathing 488 • Box 12.1 APPLICATIONs
Environmental Hypoxia 488 Exercise and Type 2 Diabetes Mellitus 512
Fish respond to hypoxia in many ways 488 Vasoactive agents regulate blood vessel diameter 512
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Myoglobin aids in oxygen delivery and utilization 513 CHAPTER 13

Locomotor physiology is maximized in animal
athletes 514 Ion and Water Balance 542
Skeletal Systems 515
Hard skeletons are made from cellular secretions 515
OVERVIEW 543
Migration 516 IONIC AND OSMOTIC REGULATION 544
Vertebrate skeletons are composed of mineralized Strategies for Ionic and Osmotic Regulation 545
calcium 516
Animals may be regulators or conformers 545
Skeletal components act as mechanical levers 518
The environment provides water in many forms 546
Skeletons can store elastic energy 519
Solutes can be classified as perturbing, compatible, or
Work loops determine if a muscle is working as motor, brake, counteracting 547
or strut 521
Evolution, Environment, and Osmoregulation 548
MOVING IN THE ENVIRONMENT 522
Marine invertebrates are osmoconformers 548
Gravity and Buoyancy 522 Most ancient fish are osmoconformers 549
Body composition influences buoyant density 523 Bony fish are ionoregulators and osmoregulators 549
Lipid accumulations aid buoyancy in zooplankton and Some fish move between freshwater and seawater 550
chondrichthians 523 Terrestriality evolved multiple times in animals 550
Gas bladders aid buoyancy in bony fish 523 The integument is an osmotic barrier 551
Fluid Mechanics 524 Desert animals have water-conserving adaptations 553
Reynolds numbers determine turbulent or laminar • Box 13.1 CHALLENGES TO HOMEOSTASIS
flow 525 Life Without Water 554
The relative importance of viscous and inertial effects
determine Re 526 Nitrogen Excretion 555
Streamlining reduces drag 526 Ammonia is produced in amino acid metabolism 555
Ammonia can be excreted across epithelial tissues 556
Aerodynamics and Hydrodynamics 527
Birds, reptiles, and insects excrete uric acid 556
Aerofoils and hydrofoils generate lift 527
Urea is produced in the ornithine-urea cycle 557
Soaring uses lift from natural air currents to overcome
gravity 528 Each nitrogenous waste strategy has inherent costs 558
True flight arose at least four times 529 The mode of nitrogen excretion can change with development or
environment 558
Fluid movements can generate propulsion 530
Cartilaginous fish produce urea as an osmolyte 559
Fin and wing shapes influence fluid movements 531
THE KIDNEY 559
Terrestrial Life 532
Aquatic animals invaded the land several times 532 Kidney Structure and Function 560
Metamorphosis remodels anatomy and physiology for terrestrial The nephron is the functional unit of the kidney 561
locomotion 533 The kidney performs four main processes 562
Flightless birds evolved in the absence of terrestrial Filtration occurs at the glomerulus 562
predators 533 Glomerular filtration pressure is affected by hydrostatic pressure
Animals of similar geometry should be able to jump to the same and oncotic pressure 563
heights 534
Terrestrial animals require strong bones and postural
Calculating Glomerular Filtration Rate (GFR) and Renal
musculature 534
Clearance 564
Energetics of Movement 535 The kidney can maintain GFR across a range of blood
Energy demands of movement can be expressed as total costs pressures 565
or mass-specific costs 535 The primary urine is modified by reabsorption and secretion 567
• Box 12.3 MATH IN PHYSIOLOGY Cellular properties differ among regions of the tubule 568
COT in Cod 536 The proximal tubule reabsorbs salts and organic metabolites 569
Animals change style of movement to alter the costs The loop of Henle mediates sequential uptake of water, then
of locomotion 537 salt 570
Environment determines energetic costs 537 The distal tubule mediates K+ secretion, NaCl reabsorption,
and hormone-sensitive water recovery 571
Body size affects costs of locomotion 538
The collecting duct regulates ion and water flux 571
Summary 540 • Review Questions 540 • Synthesis Questions 541 The loop of Henle creates a countercurrent multiplier 572
• Quantitative Questions 541 Vasopressin alters the permeability of the collecting duct 573
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Recycling of urea helps to establish the osmotic gradient in the Digestion of specific nutrients requires specific enzymes 597
medulla 574 Many animals incorporate symbiotic organisms into their digestive
The vasa recta maintains the medullary osmotic gradient via physiology 597
a countercurrent exchanger 574 Enterosymbionts play multiple roles in digestive physiology 598
Micturition is regulated by reflex and higher pathways 575
Finding and Consuming Food 598
Roles of the Kidney in Homeostasis 575 Animals sense food using chemical, electrical, and thermal
Aldosterone regulates sodium and potassium balance 575 cues 598
The renin-angiotensin-aldosterone pathway regulates blood • Box 14.1 APPLICATIONs
pressure 575
The Human Microbiome 599
Natriuretic peptides also play a role in sodium balance 576
The respiratory system and excretory system contribute to Simple animals digest food within phagocytic vesicles 600
acid-base balance 576 Feeding structures are matched to diet 600
Bird beaks are composed of keratinized tissue 601
Water Intake and Excretion 577
Mammals have bony teeth 602
Hypothalamic factors regulate thirst 577
The excretory system interacts with the cardiovascular system to DIGESTION AND ASSIMILATION 603
regulate blood pressure 578 Digestive Systems 603
Evolutionary Variation in the Structure and Function of Gut complexity is linked to the appearance of the coelom 605
Excretory Systems 580 The digestive systems of complex animals maximize surface
Invertebrates have primitive kidneys called nephridia 580 area 605
Insects use Malpighian tubules and the hindgut for ion and water Specialized compartments increase the efficiency of
regulation 581 digestion 607
Chondrichthian kidneys produce hypoosmotic urine and retain Salivary glands secrete water and digestive enzymes 608
urea 582 The stomach secretes acid and mucus 608
The role of the fish kidney differs in freshwater and The intestine is where most nutrients are hydrolyzed and
seawater 583 absorbed 609
The amphibian kidney changes in metamorphosis 583 Assimilation 611
Terrestrial animals have kidneys that help conserve Carbohydrates are hydrolyzed in the lumen and transported
water 584 by multiple carriers 611
Fish gills transport ions into and out of the water 584 Proteins are broken down into amino acids by proteases
Digestive epithelia mediate ion and water transfers 586 and peptidases 612
Reptiles and birds possess salt glands 586 Lipids are transported in many forms 613
• Box 13.3 APPLICATIONs DIGESTION AND METABOLISM 615
Conservation Physiology of Salmon 587
Regulating Digestive Systems in Individuals 615
Elasmobranch rectal glands excrete Na+ and Cl−, while retaining
urea 588 Hormones control the desire to feed 615
Hormones and neurotransmitters control gastrointestinal
Summary 590 • Review Questions 590 • Synthesis Questions 590 secretions 617
• Quantitative Questions 591 Retention time affects the efficiency of nutrient uptake 618
Gut motility is regulated by nerves and hormones that act on
smooth muscle 618
Metabolic Transitions 619

CHAPTER 14
Digestion and Energy Gut Reactor Theory 620
Metabolism 592 Nutrient stores are regulated between meals 622

OVERVIEW 593 Obesity 624
THE NATURE AND ACQUISITION OF Prolonged food deprivation can trigger a starvation
NUTRIENTS 594 response 626
Pythons may rebuild the digestive tract for each meal 627
Nutrients 594
Diets provide energy for activity, growth, maintenance, Evolutionary Variation in Metabolic Rate 628
and reproduction 594 Metabolic rate can be measured by direct or indirect
Vitamins and minerals participate in catalysis 595 calorimetry 628
Inadequate supply of essential amino acids compromises Field metabolic rate relies upon doubly labeled water 629
growth 596 Maximal sustained MR is about five times greater than
Animals require linoleic and linolenic acid in the diet 597 RMR 629
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Body size influences metabolic rate 629 MAINTAINING A CONSTANT BODY

Body temperature affects metabolic rate 630 TEMPERATURE 655
The metabolic theory of ecology links animal metabolism to
Thermogenesis 656
ecological relationships 630
Shivering thermogenesis results from unsynchronized muscle
Summary 631 • Review Questions 632 • Synthesis Questions 632 contractions 656
• Quantitative Questions 632 Some insects use metabolic futile cycles to warm flight
muscle 657
Membrane leakiness enhances thermogenesis 657
Thermogenin enhances mitochondrial proton leak 658
CHAPTER 15 • Box 15.3 CHALLENGES TO HOMEOSTASIS
Thermal Physiology 634 Evolution and Development of Thermogenin 659
Regulating Body Temperature 660

A central thermostat integrates central and peripheral
OVERVIEW 635 thermosensory information 660
Piloerection reduces heat losses 661
HEAT EXCHANGE AND THERMAL
Changes in blood flow affect thermal exchange 661
STRATEGIES 636
Countercurrent exchangers in the vasculature help retain
Controlling Heat Fluxes 636 heat 662
Water has a higher thermal conductivity than air 637 Sweating reduces body temperature by evaporative cooling 663
Convective heat exchange depends on fluid movements 638 Panting increases heat loss across the respiratory surface 663
Radiant energy warms some animals 638 Relaxed endothermy results in hypometabolic states 664
Evaporation induces heat losses 639
Summary 666 • Review Questions 666 • Synthesis Questions 666
Ratio of surface area to volume affects heat flux 639
Insulation reduces thermal exchange 640
Thermal Strategies 640
Poikilotherms and homeotherms differ in the stability of TB 641
Ectotherms and endotherms differ in the source of body thermal CHAPTER 16
energy 641
Heterotherms exhibit temporal or regional endothermy 641 Reproductive
Animals have a characteristic degree of thermotolerance 642
Temperature affects metabolic scope 644
Physiology 668
The evolution of thermal tolerance has complex origins 644 OVERVIEW 669
COPING WITH A CHANGING BODY REPRODUCTION 670
TEMPERATURE 645 REPRODUCTIVE ENDOCRINOLOGY 671
Macromolecular Structure and Metabolism 645 Reproductive Hormones 671
Animals can remodel membrane fluidity 645 Vertebrates rely on progesterone, androgens, and
• Box 15.1 APPLICATIONs estrogens 671
Thermal Tolerance and Conservation Biology of Atlantic Gonadotropins control steroid hormone levels 672
Cod 646 JH and 20HE control development and reproductive physiology
Animals remodel membranes to maintain near-constant of arthropods 672
fluidity 647 • Box 16.1 APPLICATIONs
Temperature changes enzyme kinetics 648 Pesticides Targeting Insect-Specific Hormonal
Evolution may lead to changes in enzyme kinetics 649 Pathways 674
Ectotherms can remodel tissues in response to long-term
changes in temperature 649 Sex Determination 674
Clonal reproduction is asexual 675
• Box 15.2 MATH IN PHYSIOLOGY Parthenogenesis is a short-circuit of sexual reproduction 675
Evaluating Thermal Effects on Physiological Processes
Using Q10 and Arrhenius Plots 650 Animals may be simultaneous or serial hermaphrodites 675
Sex is determined in some species by environmental
Life at High and Low Body Temperatures 650 conditions 676
Some enzymes display cold adaptation 651
GAMETOGENESIS AND FERTILIZATION 677
Stress proteins are induced at thermal extremes 652
Ice nucleators control ice crystal growth in freeze-tolerant Gametogenesis 677
animals 653 Ova are produced within follicles of somatic tissue 677
Antifreeze proteins can prevent intracellular ice formation 654 The yolk provides building blocks and metabolic precursors 678
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Insect eggs are surrounded by a chorion 679 REGULATING REPRODUCTION AND DEVELOPMENT
Egg structure differs in aquatic and terrestrial IN MAMMALS 688
vertebrates 680
Coordinating the Ovarian and Uterine Cycles 688
Spermatogenesis requires production of motile
gametes 680 Hormones control the ovarian and uterine cycles 688
Leydig cells and Sertoli cells control spermatogenesis 681 The follicular phase of ovulation is driven by FSH 689
Reproductive hormones interact with other hormones 682 Ovulation and the luteal phase follow an LH surge 689
The endometrial cycle parallels the ovulatory cycle 691
Mating, Fertilization, and Embryonic
A placenta forms after a fertilized ovum implants in the uterine
Development 682 wall 691
• Box 16.2 CHALLENGES TO HOMEOSTASIS Maternal changes in physiology accompany pregnancy 692
Reproduction and Stress 683 Contractions of uterine smooth muscle induce parturition 693
Mating is under physiological control 684 Postnatal Growth and Development 693
Male copulatory organs increase the efficiency of sperm Prolactin also controls parental care of offspring 693
transfer 684 Milk is a secretory product of mammary glands 694
Erection is controlled by vascular changes in the penis 685
Mammary gland secretions include two novel products, casein
Sperm alter activity in response to chemokinetic and chemotaxic and lactose 695
molecules 686
Females use sperm storage to ensure uninterrupted • Box 16.3 MATH IN PHYSIOLOGY
reproduction 686 Scaling of Milk Production 696
Individual sperm can compete for the opportunity to fertilize Milk energy output influences infant growth rate 697
the egg 686 Early postnatal development requires remodeling of each
Some animals delay embryonic development 686 physiological system 698
Postfertilization development relies on maternal factors 687
Amniotes produce four extraembryonic membranes early Summary 698 • Review Questions 699 • Synthesis Questions 699
in development 687 • Quantitative Question 699
Glossary 700
Animal Index 723
Subject Index 729
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Preface
The 21st century is an incredibly exciting time to be a biolo- • Physiological processes are based in the laws of chemis-
gist. Animal biologists now have access to data from a range of try and physics.
complete animal genomes covering a broad spectrum of the • Physiological diversity among animals is the result of
diversity of animals. At the time of writing this preface, com- evolutionary processes.
plete genomes already exist for several hundred species of in-
• Physiological processes are homeostatically regulated.
vertebrates and over two hundred species of vertebrates; in the
next few years, we expect that genome sequences will be avail- Every chapter revisits these key themes, providing a uni-
able for thousands of species of animals. But the fundamental fying thread that ties together our concept of animal
questions about how the genes in these genomes work to- physiology.
gether to allow animals to perform their diverse physiological
Orientation Around Learning. To promote comprehension,
functions and to go about their daily lives are still largely unan-
each chapter begins with Learning Objectives that connect
swered. Animal physiologists are at the forefront of integrating
directly with the headings in the chapter and with the Review
this new genome sequence information into a functional and
Questions at the end of the chapter. To assist with the integration
evolutionary framework as part of their efforts to understand
of material across chapters, many chapters feature a new
how animals work. Our goal in writing this textbook is to con-
Looking Back section that identifies the critical background
vey a sense of this excitement to students who are approaching
material found in earlier chapters.
the study of animal physiology for the first time.
One of the challenges that students face when they ap- An Emphasis on Animal Diversity and Evolution. We
proach their first course in physiology is the great breadth are strongly committed to the importance of teaching
and diversity of the subject matter. Physiology is among the about the physiological diversity of animals, because we
most integrative of the life sciences, drawing on ideas from feel that this diversity is a fundamental property of the
chemistry, physics, mathematics, molecular biology, and cell natural world. We also believe that books focusing only
biology for its conceptual underpinnings. In addition, to fully on humans can cause students to form the erroneous
appreciate the physiological diversity of animals, students impression that physiological processes in humans are
must have a working knowledge of environmental biology, typical of those in all animals, and thus we provide diverse
ecology, systematics, and evolutionary biology. We have writ- examples in their evolutionary context. As a result, we
ten this book to give students a well-organized and engaging include extensive discussion of physiological processes in
treatment of the fundamental principles of animal physiology. both vertebrates and invertebrates throughout the book
Throughout the book, we integrate concepts from all levels of and attempt to interweave evolutionary thinking into these
biological organization to explore the nature of diversity in discussions. Our new Chapter 2 discusses the major events
biological molecules, cells, physiological systems, and whole in the evolution of animals, with a focus on the evolution of
animals. We hope that this approach will spark the interest of physiologically significant traits and how they contributed to
all students, whatever their background preparation. the evolutionary diversification of the major animal groups.
Attention to the Integrative Nature of Animal

Key Themes Physiology. Throughout the book, we emphasize the
integrative nature of physiology in a number of ways. Each
Students are sometimes so focused on remembering the
chapter begins with an opening essay that provides a short,
“facts” of physiology that they are unable to place these facts
engaging vignette that places the system under discussion
into a well-developed conceptual framework. To help stu-
into its environmental or evolutionary framework. Together,
dents get past this difficult barrier, we organized this book
these features help to build student understanding of how
around several key themes and fundamental principles that
physiological systems interrelate and depend on each other.
are highlighted in each chapter and strove to present this ma-
terial in an accessible fashion that engages student learning. Integration of Physiology with Cell and Molecular
Biology. We divided this book into three main sections. In
A Focus on Unifying Principles. In Chapter 1, we
Part One, we provide an overview of the basic principles of
introduce four unifying themes in animal physiology:
animal physiology, identifying the common themes in the
• Physiology integrates across levels of biological organi- discipline and emphasizing the role of evolution in animal
zation from molecules to populations. diversity.
xix
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xx Pr eface
In Part Two, we discuss the cellular basis of animal phys- for the 3rd Edition, you will find the following in each
iology. The goal of Part Two is to provide students with a chapter:
general context for understanding animal physiology and to
• A short and engaging chapter-opening essay that intro-
show how, at a cellular level, animals are both similar to and
duces an animal or scenario that epitomizes the impor-
different from other organisms. We hope that this treatment
tance of the physiological system discussed in the chapter.
will help students begin to see how the somewhat abstract
processes that they study in other courses have direct rele- • Learning Objectives that organize ideas into major
vance to the understanding of animal physiology. themes for students.
Providing a strong foundation in cellular and molecular • Looking Back sections that direct students to specific
physiology is critical for students because our understanding material earlier in the text.
of animal physiology has changed dramatically in the last 10 • More succinct chapter summaries that focus on the ma-
years due to advances in fields such as genomics, transcrip- jor points.
tomics, proteomics, and cell biology, and a solid understand-
ing of these disciplines is central to the modern concept of From Chapter 4 onward, each chapter showcases these
physiology. feature boxes:
In Part Three, we discuss how cells and tissues interact • Math in Physiology takes a quantitative approach to
to form the integrative physiological systems of animals. We physiological principles.
consider each of the major physiological systems in turn, • Challenges to Homeostasis discusses how animals re-
building on the twin themes of conservation and diversity to spond to physiological challenges.
address the question: How do different animals use funda-
• Applications addresses how physiology can be used or
mentally similar building blocks to construct unique phys-
studied to solve real-world problems.
iological systems to meet the challenges imposed by the
environment? Throughout the third part of this book, we In addition, we revised the narrative and the figures exten-
integrated the discussion of the cellular and molecular pro- sively with the goal of helping students to master some of the
cesses that underpin physiological processes, at a depth that most difficult concepts in physiology. The highlights of these
will encourage students to understand the relevance of these changes in the 3rd Edition include:
disciplines to animal physiology.
Chapter 1, Introduction to Physiological Principles
Integrated Treatment of Endocrine Regulation. The
• A new opening feature on Porcelain crabs to emphasize
treatment of endocrine systems is one unique element in the
environmental physiology and the applications of phys-
book’s organization. Rather than relegating these systems to
iology to conservation biology.
a single isolated chapter, we discuss endocrinology in Part
Two in the context of the various means of cellular signaling • A new focus on exploring the unifying themes that tie
and communication, and then integrate the presentation of together both the basic and applied aspects of the disci-
its various physiological roles throughout the chapters in Part pline of animal physiology.
Two. We find that students better understand how hormones • An expanded discussion of the relationship between
control systems once they have been introduced to all the diverse form and function, the concepts of homology and anal-
ways in which cells send and receive signals. By establishing ogy, and scaling as a unifying principle in physiology,
the foundation of cellular control early in the text, we are able including several new Figures
to discuss the impact of specific hormones and glands in the
Chapter 2, Physiological Evolution of Animals
context of each physiological system, increasing the integrative
nature of the discussion. This approach places the endocrine • New to the 3rd Edition! This chapter provides a survey
system in its appropriate evolutionary framework—as one of animal diversity, focusing on the origins of physiolog-
of several means of intercellular communication that are ical traits and the significance of phylogenies.
available to multicellular organisms—and clearly demonstrates • This chapter introduces the critical events in animal
how communication and coordination are critical for the evolution and the role of environment in the selective
functioning of essentially every organ system. process.
Chapter 3, Chemistry, Biochemisty, and Cell

New for the 3rd Edition Physiology
For the 3rd Edition, we expanded the pedagogical features • A more refined discussion of energetics, including an ex-
throughout the text to facilitate students’ learning. New planation of chemical energy transfers, bonds, solubility,
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Preface xxi
and thermal effects, clearing up ambiguity about these • A reorganization of the discussion of EC coupling that
topics. more clearly distinguishes between cardiac and skeletal
• A reorganized and expanded discussion of metabolic muscle.
rate determinants, collecting information from dispa- • A new feature on muscle remodeling in exercise, combin-
rate 2nd Edition chapters into a single section. ing the themes of structural changes and cellular regulation.
• A more complete discussion of the membrane potential/ Chapter 7, Sensory Systems
Nernst equation/Goldman equation, with this impor-
tant information in the body of the chapter, rather than • New sections on topics including nociception, hearing
in a boxed feature. in whales and dolphins, and the photoreceptors involved
in circadian rhythms.
• A discussion of tissue types and the roles and regu-
lation of epithelial tissues, including transport and • A new boxed feature on using pheromones to alter behavior.
transporters. • An expanded discussion of electroreception, including a
new Figure.
Chapter 4, Cell Signaling and Endocrine Regulation • An updated discussion of magnetoreception.
• A substantial reorganization of the second half of the Chapter 8, Functional Organization
chapter to provide a more focused discussion of the fun- of Nervous Systems
damental shared principles of endocrine regulation, us-
ing selected examples from vertebrates to illustrate these • An expanded treatment of the organization and evolu-
principles. tion of nervous systems.
• An expanded section discussing endocrine systems and • Increased coverage of the general anatomy of the central
how they evolved, including a new Figure showing the nervous system, with more information about the spinal
major endocrine glands of mammals. cord.
• A new section on the evolution of the vertebrate pitu- • New boxed features examining (1) the scaling of brain
itary gland. size, neuron number, and behavioral complexity; (2) how
ocean acidification affects fish behavior by disturbing
Chapter 5, Neuron Structure and Function brain homeostasis; and (3) functional magnetic reso-
nance imaging and brain plasticity.
• A more comprehensive explanation of the Nernst and
• New sections on the corpus callosum, mirror neurons,
Goldman equations, including a new Figure and boxed
and language acquisition in birds.
feature.
• An expanded discussion of the enteric nervous system.
• A revised discussion of saltatory conduction, including
a new Figure. • A new section focusing on the role of the hypothalamus
in regulating bodily functions such as circadian rhythms
• An expanded discussion of molecular events at the
and sleep-wake cycles.
synapse.
• An updated discussion of the evolution of neurons that Chapter 9, Circulatory Systems
reflects the recent cloning of bacterial voltage-gated Na+ • New discussions of orthostatic hypotension and space
channels. flight, physiology of dinosaur circulatory systems, the
development of the human heart, and the coevolution of
Chapter 6, Cellular Movement and Muscles circulatory and respiratory systems.
• New Figures to illustrate topics including (1) skeletal • Revised and clarified discussion of the evolution of the
muscle structure, explaining how all of the muscles fit lymphatic system, amphibian circulatory systems, ion
together; (2) the impact of arrangement (series ver- channels and pacemaker currents, and the cardiovascu-
sus parallel) on muscle structure; and (3) muscle fiber lar physiology of giraffes.
mosaics. • A new boxed feature dealing with the use of EKG tech-
• An expanded feature on force and work, which consoli- nology to diagnose heart conditions.
dates the force/work/power material in a single location. • New Figures to illustrate the evolution of vertebrate cir-
• New and revised Figures that help distinguish between culatory systems and cardiac anatomy, the development
muscle fiber types, expanding the discussion of smooth of the mammalian heart, and the effect of elevated blood
muscle. pressure on risk of cardiovascular disease.
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xxii Pre fac e
Chapter 10, Immune Systems • A new feature, “Conservation Physiology of Salmon,”

highlighting recent work showing how ionoregulatory
• New to the 3rd Edition! This chapter discusses compar-
physiology influences the survival of animals in nature.
ative immunology, with a focus on evolutionary diver-
sity of the innate and adaptive immune systems. Chapter 14, Digestion and Energy Metabolism
• It includes discussion of the molecular mechanisms that
• A change in the scope of the chapter to also include en-
organisms use to detect foreign molecules and the roles of
ergy metabolism and its regulation.
the various immune cells, particularly B cells and T cells.
• A reorganization of the section on regulation of diges-
• The addition of a chapter on immunology provides con-
tion to discuss processes along a linear timeline.
text for the interaction between immunity and other
physiological processes, particularly the circulatory, • A new Applications feature focusing on the gut micro-
thermal, and digestive systems. biome, with appropriate cross-referencing to the new
Immune Systems chapter.
Chapter 11, Respiratory Systems • A new feature focusing on obesity as a homeostatic challenge.
• A new discussion of the potential for unidirectional ven- • A more consistent treatment of the many hormones that
tilation in crocodile lungs. regulate digestion and metabolic rate.
• A revised discussion of Root effect hemoglobins, em-
Chapter 15, Thermal Physiology
phasizing recent research suggesting a role for these he-
moglobins in delivery of oxygen to systemic tissues in • An expanded discussion of thermal biology to better con-
fish. sider physiological ecology, including a new Applications
• A new section on the evolution of myoglobin in diving feature on thermal tolerance and conservation biology
mammals. of Atlantic cod, an expanded discussion of the impact of
temperature on metabolism, and new material on ther-
• New boxed features dealing with (1) the treatment
mal effects on aerobic scope and the OCLTT hypothesis.
of respiratory distress syndrome in premature infants,
(2) pulmonary function tests, and (3) adaptations to • Revised discussion of the evolution of uncoupling
high altitude in bar-headed geese. proteins, including introduction of a Challenges to
Homeostasis feature on the evolution and development
Chapter 12, Locomotion of thermogenin and brown adipose tissue.
• An expanded discussion of the importance of animal • Revised treatment of the ectotherm/endotherm,
athletes as models for understanding physiological poikilotherm/homeotherm distinctions, with a revised
evolution. Figure.
• New features on type II diabetes and migration. • A new summary Figure on the diversity in futile cycles.
• An expanded discussion of the regulation of homeosta- • A modified discussion of Arrhenius plots to include
sis in muscle. more student-driven calculations as part of a Math in
• A new feature on the cost of transport and a revised dis- Physiology feature.
cussion of work loops that deconstruct positive and neg- Chapter 16, Reproductive Physiology
ative work to help students understand the biophysical
basis of locomotion. • A new feature on pesticides targeting insect-specific
pathways addresses how pesticides can be used to target
Chapter 13, Ion and Water Balance insect development and reproduction.
• An expanded discussion of osmotic strategies used by • A Math in Physiology feature combines the concepts of
animals, highlighting the important transitions that allometry with the constraints on milk production.
arose in the context of animal evolution. We hope that you enjoy using this textbook. Please feel free to
• A reorganized section on kidney function and regula- contact us at the email addresses below if you have any com-
tion, focusing on the four main homeostatic functions: ments or suggestions on how we could make this book an
ion balance, osmotic balance, pH balance, and blood even better tool to help you learn or teach animal physiology.
pressure regulation.
• A new feature that delves into the quantitative analysis Chris Moyes Trish Schulte
of renal clearance, including an explicit discussion of the Queen’s University University of British Columbia
concept of a “virtual volume.” [email protected] [email protected]
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Supplements
Companion Website need at a lower cost for students. And even as students save
money, you can save time and hassle with a digital eTextbook
This student resource features answers to the Review Ques- that allows you to search for the most relevant content at the
tions and Concept Checks that appear in the text, chap- very moment you need it. Whether it’s evaluating textbooks or
ter-specific quizzes, links to physiology labs and other creating lecture notes to help students with difficult concepts,
relevant websites, an interactive glossary, and more. Please CourseSmart can make life a little easier. See how when you
visit www.pearsoncanada.ca/animalphysiology. visit www.coursesmart.com/instructors.
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xxiii
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Acknowledgments
Preparing the 3rd Edition of our textbook required a team of Reviewers
dedicated people to bring it to fruition. The Pearson Canada
We would also like to thank the instructors who reviewed the
team conducted the extensive research needed to identify
3rd Edition manuscript:
the strengths and weaknesses of our 2nd Edition. They also
helped us revise the chapters, developing the manuscript into Eli Asem, Purdue University
the finished product. A team of three editors managed the Adam Oliver Brown, University of Ottawa
three-year revision process. Maurice Esses, Lise Dupont, and Julian Christians, Simon Fraser University
Lisa Rahn managed to keep us on track, successfully main- Reyniel Cruz-Aguado, Douglas College
taining the delicate balance between motivation and persis- Rosa da Silva, McMaster University
tence. It is impossible to imagine how the 3rd Edition could Jeff W. Dawson, Carleton University
have been completed without their support. Heidi Englehardt, University of Waterloo
The 3rd Edition continues to benefit from the work David H. Evans, University of Florida
done by the developmental team in place for the 1st and Corey Flynn, University of Calgary
2nd Editions, notably Catherine Murphy, the developmental Wei Ge, Chinese University of Hong Kong
editor, Laura Southworth, our art development editor, and Kathleen Gilmour, University of Ottawa
Susan Malloy and Marie Beaugureau, our project editors at Helga Guderley, Université Laval
Benjamin Cummings. Raymond P. Henry, Auburn University
We would particularly like to thank various friends Kelly S. Johnson, Ohio University
and colleagues who have provided input and suggestions Kevin S. Kinney, Depauw University
on specific chapters. CDM would like to thank his col- Heather Koopman, University of North Carolina
leagues at Queen’s for their contributions to the new chap- at Wilmington
ters on evolution of physiological systems (Steve Lougheed, Roswitha Marx, University of Victoria
Vicki Freisen) and comparative immunology (Bill Bendena, Joanne Nash, University of Toronto at Scarborough
Virginia Walker), and to thank Doug Symes (University of Linda Ogren, University of California, Santa Cruz
Calgary) for his advice on the Cost of Transport feature. PMS Robert J. Omeljaniuk, Lakehead University
would like to thank her colleagues in the comparative phys- Sushama Pavi-Denver, University of Michigan
iology group at UBC for their advice and support through- Scott D. Reid, University of British Columbia
out this project, with particular thanks to Tony Farrell for Gregory Schmaltz, University of the Fraser Valley
suggestions regarding cardiac physiology, Bill Milsom for Jason Schreer, State University of New York at Potsdam
his advice on respiratory physiology, and Doug Altshuler James Staples, University of Western Ontario
for helpful discussions regarding ways to approach teaching Jonathon Stillman, San Francisco State University
neurophysiology and the nervous system. Marva Sweeney-Nixon, University of Prince Edward Island
Steven Swoap, Williams College
Keith Tierney, University of Alberta
Helene Volkoff, Memorial University
Tracy L. E. Wagner, Washburn University
xxiv
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Principles of
Animal Physiology
T HIRD EDIT ION
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Principles of

Christopher D. Moyes, Ph.D.

Patricia M. Schulte, Ph.D.

Library and Archives Canada Cataloguing in Publication

CHAPTER 1 CHAPTER 4 CHAPTER 7

PART ONE INTRODUCTION TO PHYSIOLOGY 2

Terrestriality arose in multiple lineages 36 Gluconeogenesis builds glucose from noncarbohydrate

OVERVIEW 39 Mitochondrial Metabolism 71

Carbohydrates 60 Cellular Organization 84

PART TWO THE CELLULAR BASIS OF ANIMAL PHYSIOLOGY 98

CHAPTER 4 Receptor type determines the cellular response 113

The actions of insulin illustrate the principle of negative CHAPTER 5

and Muscles 208 Smooth Muscle 243

Invertebrate Muscles 248 Specialized Muscles and Transdifferentiated

PART THREE INTEGRATING PHYSIOLOGICAL SYSTEMS 256

OVERVIEW 311 Somatic Motor Pathways 337

ORGANIZATION AND EVOLUTION OF NERVOUS INTEGRATIVE FUNCTIONS OF NERVOUS

THE CNS OF VERTEBRATES 317 Learning and Memory 344

Circulatory Systems 356 The Cardiac Cycle 384

• Box 9.3 CHALLENGES TO HOMEOSTASIS Cell-Mediated Immunity 430

GAS TRANSPORT TO THE TISSUES 473

Carbon Dioxide Transport 482 Energy Metabolism 506

Myoglobin aids in oxygen delivery and utilization 513 CHAPTER 13

Metabolic Transitions 619

Metabolism 592 Nutrient stores are regulated between meals 622

Body size influences metabolic rate 629 MAINTAINING A CONSTANT BODY

Regulating Body Temperature 660

Attention to the Integrative Nature of Animal

Chapter 3, Chemistry, Biochemisty, and Cell

Chapter 10, Immune Systems • A new feature, “Conservation Physiology of Salmon,”

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