Principles of

Animal Physiology

Christopher D. Moyes, Ph.D.

Patricia M. Schulte, Ph.D.

Queens University

University of British Columbia

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ISBN 0-8053-7279-2
Copyright 2005 by Pearson Education, Inc., publishing as Benjamin Cummings, San Francisco, CA 94111.
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About the Authors

Christopher D. Moyes, Ph.D.
Queens University
Chris received his Ph.D. in Zoology from the University of British Columbia in the area of comparative
muscle physiology. After postdoctoral fellowships in molecular physiology at the U.S. National Institutes
of Health and Simon Fraser University, he took a position at Queens University, where he is an associate
professor in the Department of Biology. He teaches a spectrum of courses in cell biology and animal
physiology, while continuing to pursue his research interests in molecular physiology and biochemistry.
Chris is a recipient of the Premiers Research Excellence Award. He is a member of the American
Physiological Society and the Canadian Society of Zoologists and has served on the Natural Science and
Engineering Research Council of Canada grant panel for Animal Biology. He is also a member of the
Editorial Board of Comparative Biochemistry and Physiology.
He has published more than 60 peer-reviewed papers, including contributions to four books. Among his
recent papers are Moyes, C. D., and D. L. Hood (2003) Origins and consequences of mitochondrial
variation in vertebrate muscle, Annual Review of Physiology 65: 177201 and Moyes, C. D. (2003)
Controlling muscle mitochondrial content, Journal of Experimental Biology 206: 43854391.
More of Chriss research is detailed on his homepage at http://biology.queensu.ca/
moyesc.

Patricia M. Schulte, Ph.D.

University of British Columbia
Trish received her Ph.D. in Biological Sciences from Stanford University in the area of evolutionary
physiology focusing on the role that changes in gene expression play in evolution. She is currently an
assistant professor in the Department of Zoology at the University of British Columbia in Vancouver,
where she teaches animal physiology and evolutionary physiology and runs an active research program.
Using several species of fish as model systems, her research group is particularly focused on the
relationship between genetic variation, performance differences, and fitness in a changing environment.
She also conducts research into applied questions relating to fisheries, aquaculture, and aquatic
toxicology.
Trish is a recipient of the Premiers Research Excellence Award and several teaching awards, including
the UBC Science Undergraduate Society Award for Excellence in Teaching. Trish is a member of the
Canadian Society of Zoologists and the Society for Integrative and Comparative Biology. She is an
associate editor for the scientific journal Physiological and Biochemical Zoology.
She has published more than 30 peer-reviewed papers, including two book chapters. Among her recent
papers are DeKoning, A. B. L., D. J. Picard, S. R. Bond, and P. M. Schulte (2004) Stress and interpopulation variation in glycolytic enzyme expression in a teleost fish, Fundulus heteroclitus,
Physiological and Biochemical Zoology 77: 1826 and Richards, J. G., J. W. Semple, J. S. Bystriansky,
and P. M. Schulte (2003) Na
/K
-ATPase alpha-isoform switching in gills of rainbow trout
(Oncorhynchus mykiss) during salinity transfer, Journal of Experimental Biology 206: 44754486.
More of Trishs research is detailed on her homepage at www.zoology.ubc.ca/zoology/z/schulte.

iii

To the Student:
In this booklet are three preliminary chapters from Principles of Animal Physiology, a new animal
physiology text that we are publishing next year. Before we finalize the material and send the book to
press, we need your help in making the last round of edits.
We want to know what you think about the writing style, art, organization, boxes, and review questions
in these sample chapters.
Your instructor has agreed to try out these three sample chapters with you in a class test. Please read
these chapters in addition to, or in lieu of, the comparable chapters in your present book, and then pass
along your feedback by filling out the questionnaire located in the front of the book. Then, tear out the
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Your feedback will be used in the final editing process for this book. We want to make sure that this new
title is meeting your needs, so please be as clear and candid in your comments as possible. We truly
appreciate your time in helping us make this a spectacular textbook, and we hope you enjoy participating
in and contributing to the publishing process.
We look forward to hearing from you!

The physiology team at Benjamin Cummings Publishers,

Susan Malloy
Senior Project Editor

iv

Leslie Berriman
Executive Editor

Cheryl Cechvala
Market Development Manager

Table of Contents
PART ONE

The Cellular Basis of Animal Physiology

Chapter 1

Introduction to Physiological Principles

Chapter 2

Chemistry of Life

Chapter 3

Cell Metabolism and Physiology

Chapter 4

Hormones and Cell Signaling

Chapter 5

Neuron Structure and Function

Chapter 6

Cellular Movement and Muscles

PART TWO

Integrating Physiological Systems

Chapter 7

Sensory Systems

Chapter 8

Nervous Systems

Chapter 9

Circulatory Systems

Chapter 10

Respiratory Systems

Chapter 11

Ion and Water Balance

Chapter 12

Digestion

Chapter 13

Locomotion

Chapter 14

Thermal Biology

Chapter 15

Reproduction

Appendices
For Further Reading
Glossary
References
Index
v

Detailed Contents of the Three Sample Chapters

Chapter 5 Neuron Structure
and Function 2
Overview

Evolution of Neurons

Signaling in a Vertebrate Motor Neuron

Electrical Signals in Neurons

13

Voltage-gated channels shape the action potential 14

Voltage-gated Na
channels have two gates 15
Action potentials transmit signals across long
distances 17
Vertebrate motor neurons are myelinated 19
Axons conduct action potentials unidirectionally 19
Action potential frequency carries information 20

Signals Across the Synapse

21

Intracellular Ca2
regulates neurotransmitter release 21

Action potential frequency influences neurotransmitter
release 22
Acetylcholine is the primary neurotransmitter at the
vertebrate neuromuscular junction 22
Postsynaptic cells express specific receptors 22
Neurotransmitter amount and receptor activity influence
signal strength 23

Diversity of Neural Signaling

Structural Diversity of Neurons

24

25

Neurons can be classified based on their function 26

Neurons can be classified based on their structure 26
Neurons are associated with glial cells 27

Diversity of Signal Conduction

28

Voltage-gated ion channels are encoded by multiple

genes 28
Voltage-gated Ca2
channels can also be involved in action
potentials 30
Conduction speed varies among axons 30
The cable properties of the axon influence current flow 30
Intracellular and membrane resistance influence
conduction speed 32
Membrane capacitance influences the speed of
conduction 33
Giant axons have high conduction speed 35
Myelinated neurons evolved in the vertebrates 37
Myelination increases conduction speed 38

Diversity of Synaptic Transmission

38

Electrical and chemical synapses play different roles 40

Chemical synapses have diverse structures 41
There are many types of neurotransmitters 42
Neurotransmitters can be excitatory or inhibitory 44
Neurotransmitter receptors can be ionotropic or
metabotropic 44
Acetylcholine receptors can be ionotropic or
metabotropic 45
The biogenic amines play diverse physiological roles 45
Neurons can synthesize more than one kind of
neurotransmitter 47
Neurotransmitter release varies depending on
physiological state 47

vi

50

51

Synthesis Questions

51
52

Graded potentials vary in magnitude 7

Graded potentials are short-distance signals 10
Graded potentials are integrated to trigger action
potentials 11

Signals in the Axon

Summary

Review Questions

Gated ion channels allow neurons to alter their membrane

potentials 7

Signals in the Dendrites and Cell Body

49

Only animals have voltage-gated Na
channels

Most organisms use chemicals for cell-to-cell
communication 50

Chapter 6 Cellular Movement

and Muscles 54
Overview

56

Cytoskeleton and Motor Proteins

Microtubules

57

57

Microtubules are composed of -tubulin and -tubulin

Microtubules show dynamic instability 59
Microtubule polarity determines the direction of
movement 61
Kinesin and dynein move along microtubules 61
Cilia and flagella are composed of microtubules 62

Microfilaments

57

63

Microfilaments are polymers of actin 64

Actin polymerization can generate movement 65
Actin uses myosin as a motor protein 66
The sliding filament model describes actino-myosin
activity 68
Myosin activity is influenced by unitary displacement and
duty cycle 69

Muscle Structure and Regulation

of Contraction 71
Structure of the Vertebrate Striated Muscle Contractile
Apparatus 72
Muscles are composed of thick and thin filaments 72
Thick and thin filaments are arranged into
sarcomeres 73
Myosin II has a unique duty cycle and unitary
displacement 73
Sarcomeric organization determines contractile properties
of the muscle cell 75

Regulation of Contraction in Vertebrate Striated

Muscle 76

Thin filament proteins confer Ca2
sensitivity 77

The troponin-tropomyosin complex influences contraction
kinetics 78
Thick filaments also influence contractile properties 81

Excitation

81

Muscles are excited by an action potential 83

Myogenic muscle cells spontaneously depolarize 85
Neurogenic muscle is excited by neurotransmitters 85
T-tubules enhance action potential penetration into the
myocyte 85
Ca2
for contraction comes from intracellular or
extracellular stores 86
DHPR activation induces Ca2
release from the SR 87
Relaxation follows removal of Ca2
from the
cytoplasm 88

Muscle Diversity in Vertebrates and

Invertebrates 89

Specific combinations of proteins result in muscle fiber

types 90
Individuals alter fiber type in response to changing
conditions 91
Invertebrate muscles contract in response to graded
excitatory postsynaptic potentials 92
Asynchronous insect flight muscles do not use Ca2

Translating Contraction into Movement

transients 93
Heater organs and electric organs are modified
muscles 94
Smooth muscle lacks organized sarcomeres 95
Smooth muscle contraction is regulated by both thick and
thin filament proteins 98
Latch cross-bridges maintain smooth muscle contraction
for long periods 99

Gravity and Buoyancy

Summary

100

Review Questions

101

Synthesis Questions

Chapter 13
Overview

101

Locomotion

104

Locomotor Systems
Muscle Fiber Types

104

111

Glycolysis and mitochondria support different types of

locomotion 111
Mitochondrial content determines muscle aerobic
capacity 112
Muscle must recover from high-intensity activity 113
Metabolic transitions accompany prolonged exercise 114
Hormones control fuel oxidation in muscle 115

Perfusion and Oxygen Delivery to Muscle

116

Capillary networks bring oxygen to the vertebrate muscle

fibers 117
Vasoactive agents regulate blood vessel diameter 118
Myoglobin aids in oxygen delivery and utilization 119

Skeletal Systems

127

127

Body composition influences buoyant density 127

Lipid stores increase the buoyancy of zooplankton and
sharks 129
Swim bladders are gas-filled sacs that increase
buoyancy 130

Fluid Mechanics

131

Reynolds numbers determine turbulent or laminar

flow 131
The relative importance of viscous and inertial effects
determine Re 132
Streamlining reduces drag 133

120

Hard skeletons are made from cellular secretions 120

Vertebrate skeletons are composed of mineralized
calcium 121
Skeletal components act as mechanical levers 122
Skeletons can store elastic energy 123

134

Aerofoils and hydrofoils generate lift 134

Soaring uses lift from natural air currents to overcome
gravity 135
Fluid movements can generate propulsion 136
Fin and wing shapes influence fluid movements 137

Terrestrial Life

104

Most invertebrates use simple circular and longitudinal

muscles to move 105
Fish use two or three fiber types to swim 106
The pattern of locomotor muscle contraction is controlled
by motor neurons 107
Tetrapods have a multiplicity of fiber types 109
Locomotor muscles are organized into locomotor modules
and functional groups 109

Energy Metabolism

Moving in the Environment

Aerodynamics and Hydrodynamics

102

124

Muscles are specialized for force generation or power

generation 125
Work loops show the balance between positive and
negative work 126

138

Aquatic animals invaded the land several times 140

Metamorphosis remodels anatomy and physiology for
terrestrial locomotion 140
Flightless birds evolved in the absence of terrestrial
predators 141
Animals of similar geometry should be able to jump to the
same heights 141
Terrestrial animals require strong bones and postural
musculature 142

Energetics of Movement

143

Energy demands of movement can be expressed as total

costs or mass-specific costs 143
Velocity of movement affects locomotor costs 144
Animals change style of movement to alter the costs of
locomotion 146
Environment determines energetic costs 146
Body size affects costs of locomotion 148

Integrating Systems: Migration

Summary

149

151

Review Questions
Synthesis Questions

153
153

For Further Reading

155

vii