Comparative Animal Physiology

Philip C. Withers
Senior Lecturer, Department of Zoology

The University of Western Australia

Technlsche Universitat
ft
FACHBEREICH 10 BIOLOGIE
Bibliothek

Schnittspahristralie 10
D'642 8 7 D a r m s t a d t

SAUNDERS COLLEGE PUBLISHING

Harcourt Brace jovanovich College Publishers

Fort Worth Philadelphia San Diego New York

Orlando Austin San Antonio Toronto
Montreal London Sydney Tokyo

Contents
About the Author vii
Preface ix
Acknowledgments xiii

Chapter l

Introduction 1

Comparative Animal Physiology 2

Objectives 3
Summary 5

Chapter 2 External and Internal

Environments 6
The External Environment 7
The Atmosphere 7
Aquatic Environments 7
Terrestrial Environments 9
The Internal Environment 10
The Extracellular Environment 11
The Intracellular Environment 13
Homeostasis and Regulation 14
Tolerance and Resistance 15
Acclimatization and Acclimation 16
Regulatory Mechanisms 17
Summary 27

First Law of Thermodynamics 31

Second Law of Thermodynamics 31
Free Energy 31
Diffusion Flux 33
Chemical Reactions 36
Free Energy Change 36
Reaction Kinetics 36
Activation Energy 38
Oxidative Metabolism 40
High-Energy Phosphate Compounds 40
Metabolic Substrates 42
Aerobic Metabolism 42
Anaerobic Metabolism 58
Anaerobic Metabolism Strategies 58
Invertebrates 61
Vertebrates 62
Energetics of Anaerobic Metabolism 65
Metabolic Fates of Anaerobic End Products 66
Bioluminescence 67
Thermogenesis 70
Leaky Membranes 70
Brown Adipose Tissue 71
Defense 71
Summary 73
Supplement 3-1 Diffusion in Various Geometries 74
Supplement 3-2 Mechanisms of Bioluminescence 76
Recommended Reading 81

Recommended Reading 28

Chapter 4 Animal Energetics 82

Chapter 3 Cellular Energetics 29
Thermodynamics 30
Work from Heat 30

Calorimetry 83
Energy Budgets 86
Production 91
xvii

Contents

Aerobic Metabolism 92
Body Size 93
Temporal and Geographic Effects 104
Food and Oxygen Availability 105
Temperature 106
Specific Dynamic Effect 108
Activity 110
Free-Living Metabolism 112
Anaerobic Metabolism 112
Invertebrates 112
Vertebrates 113
Scaling of Cellular Metabolism 115
Summary 117
Supplement 4-1 Stoichiometry for Carbohydrate,
Lipid, and Protein Metabolism 118
Supplement 42 Energetics of Nitrogenous
Waste Products 118
Supplement 4-3 Allometric Analysis
of Metabolic Rate 119
Recommended Reading 121 '

Chapter 5 Temperature 122

Physical Effects of Temperature 123
Temperature Scales 123
Reaction Rates 123
The Thermal Environment 125
Patterns of Temperature Regulation 127
Heat Exchange 128
Conduction 128
Convection 130
Radiation 131
Evaporation! Condensation 132
Heat Balance 132
Ectotherms 133
Aquatic Ectotherms 133
Terrestrial Ectotherms 135
Adaptations to Cold 143
Adaptations to Heat 150
Advantages of Ectothermy 152
Endotherms 154
Mammals and Birds 154
Reptiles 170

Fish 171
Insects 173
Plants 175
Evolution of Endothermy 175
Fever 177
Acclimation and Acclimatization 177
Biochemical Adaptations to Temperature 180
Summary 186
Supplement 5-1 Convective Heat Transfer 187
Supplement 5-2 Newtonian Model for
Thermoregulation in Ectothermic
Mammals and Birds 188
Supplement 5-3 Bioclimatic Rules 190
Recommended Reading 191

Chapter 6 Membrane Physiology 192

Membrane Structure 194
Membrane Permeation 196
Diffusion 196
Mediated Transport 198
Dynamics of Semipermeable Membranes 203
Resting Membrane Potentials 204
Diffusion Potentials 205
Equilibrium Potentials 207
Goldman-Hodgkin-Katz Potential 208
Conductance, Current, and Capacitance 210
Excitable Cell Membranes 212
Action Potentials 212
Properties of Action Potentials 220
Axonal Propagation 223
Synaptic Transmission 230
Electrical Synapses 230
Chemical Synapses 232
Neural Integration 243
Summary 247
Supplement 6-1 Hodgkin-Huxley Model
of Action Potentials 248
Supplement 6-2 Electrical Synapses 250
Supplement 6-3 Quantal Release
of Neurotransmitter Vesicles 251
Recommended Reading 252

Contents

Chapter 7 Sensory Physiology 254

Chapter 8 Nervous Systems 329

Structural and Functional Classification 255

Sensory Coding 257
Transduction 258

Evolution 330
Nervous Tissue 332
Neurons 332

Relationship between Stimulus Intensity

and Response 259
Adaptation 261
Sensory Threshold 263
Central Control of Sensory Reception 265
Epithelial Modification of Receptor
Current Circuit 266
Chemoreception 267
Gustation and Olfaction 267
Humidity 274
Temperature 274
Infrared Reception 276
Mechanoreception 278
Mechanotransduction 278
Invertebrate Mechanoreceptors 278
Vertebrate Mechanoreceptors 282
Electroreception 293
Magnetoreception 297
Photoreception 300
Light 300
Photoreceptors 301
Image Formation 303
Structure and Evolution of Photoreceptors 309
Photochemistry 309
Electrophysiology 313
Color Vision 317
Polarized Light 319
Photopic and Scotopic Vision 319
Summary 322
Supplement 7-1 Electrophysiologkal
Recording from Sensory Receptors 323
Supplement 7-2 Lenses, Image Formation,
and Visual Resolution 325
Supplement 7-3 Visual Sensitivity 327
Recommended Reading 328

Glia 334
Integrative Neurophysiology 335
Interneurons 335
Neural Circuits 337
Nerve Networks 339
Primitive Nervous Systems 339
Nerve Nets 340
Nerve Cords and Brains 342
Chordate Nervous System 351
Visual Processing 365
Retinal Processing 367
Central Nervous System Processing 368
Binocular Processing 371
Memory and Learning 373
Biological Clocks 380
Circadian Rhythms 380
Lunar Rhythms 384
Cireannual Rhythms 385
Summary 385
Supplement 8-1 Inhibition by Mauthner
Interneurons 387
Supplement 8-2 The Nervous System
of the Leech 388
Supplement 8-3 The Giant Fiber
System of Cephalopods 390
Supplement 8^4 Development of the Cerebral
Cortex in Vertebrates 392
Recommended Reading 393

Chapter 9 Cell Movement 394

The Cytoskeleton 395
Microtubules 395
Microfilaments 397
Intermediate Filaments 398
Principles of Cellular Movement 399

Contents

Microtubular Movement 400

Cytoplasmic Streaming 401
Cell Division 402
jCilia and Flagella 405
Microfilament Movement 415

Invertebrate Endocrine Systems 509

Lower Invertebrates 509

Amoeboid Movement 415

Muscle 415
Summary 442
Supplement 9-1 Dynamics of Actin 443
Supplement 92 Muscle Force, Velocity,
Power, and Work Done 445
Supplement 9-3 Neurogenic WeaklyElectric Fish 446

Insects 521
Echinoderms 529
Chordate Endocrine Systems 529
Hypothalamo-Hypophyseal Axis 532
Pineal Gland 540
Urophysis 540

Recommended Reading 447

Chapter 10 Support and Locomotion 448

Support 450
Material Properties 450
Skeletons 452
Joints 457
Locomotion 460
Terrestrial Locomotion 460
Aquatic Locomotion 468
Aerial Locomotion 479
Comparison of Locomotory Costs 484
Summary 486 '
Supplement 10-1 Analysis of Bending Forces 487
Supplement 10-2 Mechanics of Fusiform
and Pennate Muscles 489
Supplement 10-3 Aerodynamic Forces 490
Recommended Reading 492

Annelids 511
Mollusks 512
Crustaceans 515

Thyroid Gland 540

Parathyroid Gland, the Ultimobranchial Gland,
and Corpuscles of Stannius 542
Gastroenteropancreatic Cells 545
Chromaffin Tissues 548
Steroid Hormones 550
Evolution of Endocrine Systems 557
Summary 559
Supplement 11-1 Radioimmunoassay: A Technique
Using Hormone-Binding Specificity 560
Recommended Reading 563

Chapter 12 Aquatic Respiration 564

Respiratory Gases 565
Composition of Air 565
Gases Dissolved in Water 568
Diffusion 570
Bulk Flow 572
Respiratory Systems 573
> Generalized Respiratory Systems 573
Invertebrates 574

Chapter 11 Endocrinology 493

Vertebrates 585
Regulation of Respiration 599

Endocrine Systems 495

Physiological Implications of Aquatic Respiration 601

Summary 602

Neurosecretion 496
Classical Endocrine Glands 499
Mechanisms of Hormone Action 501
Hormone Binding 503 "

Supplement 12-1 Boundary Layers 603

Supplement 122 Countercurrent Exchange 605
Recommended Reading 607

Contents

Chapter 13 Aerial Respiration 608

Vertebrates 609
The Vertebrate Transition from Water to Land 609
Principles of Pulmonary Respiration 625
Invertebrates 631
Invertebrate Transition from Water to Land 632
Chelicerates 636
Millipedes, Centipedes, and Insects 639
Physical Gills and Plastrons 644
Regulation of Respiration 646
Vertebrates 646
Invertebrates 650
Effects of Diving and Altitude 651
Water Loss from Gas Exchange Surfaces 654
Gas Exchange in Terrestrial Eggs 655
Physiological Implications of Air Breathing 659
Summary 660
Supplement 13-1 Branching Pattern of the
Mammalian Respiratory System 662
Recommended Reading 664

Chapter 14 Circulation 665

Design of Circulatory Systems 666
Rheology 669
Viscosity 669
Poiseuille-Hagen Flow Formula 671
Laminar and Turbulent Flow 672
Resistance 672
Distensibility and Compliance 673
Pressure, Velocity, and Gravity 675
Invertebrate Circulatory Systems 677
Coelenterates 677
Nemertean Worms 677
Aschelminths 677
Annelids 678
Mollusks 680
Arthropods 685
Echinoderms 690
Hemichordates 692

Chordate Circulatory Systems 692

Vertebrates 693
Circulatory Patterns 702
Regulation of the Cardiovascular System 715
The Heart 715
Cardiac Output 717
Blood Pressure 719
Selective Distribution of Blood Flow 721
Summary 723
Supplement 14-1 Laminar, Turbulent,
and Pulsatile Flow 724
Recommended Reading 726

Chapter 15 Blood 727

Blood 728
Composition 728
Proteins 728
Cells 729
Respiratory Pigments 732
Hemoglobin 733
Chlorocruorin 737
Hemerythrin 738
Hemocyanin 738
Oxygen-Binding Properties 740
Gas Transport 745
Oxygen 745
Carbon Dioxide 756
Transport of Other Gases 759
Acid-Base Balance 759
Defense 765
Hemostasis 765
Immunology 766
Physical Deterrence 771
Summary 772
Supplement 15-1 Oxygen Equilibrium 773
Recommended Reading 775

xxi

Contents

Chapter 16 Water and Solute Balance 777

Chapter 18 Digestion 892

Body Fluid Composition 778

'Water 778

Feeding Patterns 893

Solutes 780
Intracellular Volume Regulation 783
Water and Ion Budgets 788
Aquatic Environments 791
Protozoans 791
Invertebrates 791
Vertebrates 798

Large-Particle Feeding 896

Fluid Feeding 897
Surface Nutrient Absorption 899
Generalized Structure and Function
of the Digestive Tract 902
Why a Digestive Tract? 902

Metabolic Cost of Ionoregulation 803

Terrestrial Environments 806
Water Vapor in Air 807
Invertebrates 819
Vertebrates 822
Summary 827
Supplement 16-1 Benefits and Costs
of Ureo-osmoconforming 829
Recommended Reading 830

Chapter 17 Excretion 831

Excretory Organs 832
Epithelial Exchange 832
Tubular Excretion 838
Water and Solute Excretion 845
"Lower" Invertebrates 845
Annelids 848
Mollusks 849
Arthropods 851
Cephalochordates and Urochordates 855"
'3
Vertebrates 855
Nitrogen Metabolism 874
Amino N Metabolism 874
Nucleic Acid Metabolism 875
Other Nitrogenous Waste Products 875
Patterns of Nitrogen Excretion 878
Summary 886

Supplement 17-1 Urea and Uric Acid Synthesis

Suspension Feeding 893

Regions of the Gut 902

Digestion 907
Digestive Systems 921
Protozoans 922
Bivalve Mollusks 923
Insects 924
Vertebrates 924
Absorptive Adaptations of the Gut 928
Specialized Digestive Systems 928
Cellulose 928
Chitin 936
Wax 936
Algal farming 936
Symbiont Transfer 937
Nutrition 937
Amino-acids 937
Fatty Acids and Sterols 937
Vitamins 938
Regulation of Nutritional Intake 942
Summary 943
Supplement 18-1 Adaptations of the Gut
to Metabolic Rate and Diet 946
Recommended Reading 948
APPENDIX A

Classification of Living Animals A-l

APPENDIX B

Units A-5

APPENDIX C Constants and Coefficients A-7

References R-l
Glossary G-l

Recommended Reading 890

Genus Index 1-1
Subject Index 1-17