ANIMAL
Contents Preface PART 1 PRINCIPLES OF PHYSIOLOGY CHAPTER 1 Studying Animal Physiology THE SUBDISCIPLINES OF ANIMAL PHYSIOLOGY THE HISTORY OF ANIMAL PHYSIOLOGY WHY STUDY ANIMAL PHYSIOLOGY? Scientific Curiosity Commercial and Agricultural Applications Insights into Human Physiology CENTRAL THEMES IN ANIMAL PHYSIOLOGY Structure/Function Relationships Adaptation, Acclimatization, and Acclimation Homeostasis Feedback-Control Systems Conformity and Regulation THE LITERATURE OF THE PHYSIOLOGICAL SCIENCES Stiottif U: i -i The Concept of Feedback ANIMAL EXPERIMENTATION IN PHYSIOLOGY CHAPTER 2 Contemporary Experimental Methods for Exploring Physiology FORMULATING AND TESTING HYPOTHESES Animal Physiology and the August Krogh Principle Experimental Design and Levels of Biological Organization POPULAR MOLECULAR TECHNIQUES IN PHYSIOLOGY Tracing Molecules with Radioisotopes Visualization Techniques Using Antibodies Genetic Engineering CELLULAR TECHNIQUES Uses of Microelectrodes and Micropipettes Microscopic Techniques Cell Culture BIOCHEMICAL ANALYSIS Measuring Composition: What Is Present? Measuring Concentration: How Much Is Present? EXPERIMENTS WITH ISOLATED ORGANS AND ORGAN SYSTEMS OBSERVING AND MEASURING ANIMAL BEHAVIOR The Power of Behavioral Experiments Methods in Behavioral Research THE IMPORTANCE OF PHYSIOLOGICAL STATE IN RESEARCH XV I 3 4 4 6 7 8 9 9 10 ii 12 17 17 19 21 24 24 26 30 32 32 3 1 CHAPTER 3 Molecules, Energy, and Biosynthesis WATER: THE UNIQUE SOLVENT The Water Molecule Properties of Water Water as a Solvent PROPERTIES OF SOLUTIONS Concentration, Colligative Properties, and Activity Ionization of Water Acids and Bases The Biological Importance of ph The Henderson-Hasselbalch Equation Buffer Systems Electric Current in Aqueous Solutions Sp.olii.f-liir-\-i Electrical Terminology and Conventions Binding of Ions to Macromolecules BIOLOGICAL MOLECULES Lipids Carbohydrates Proteins Nucleic Acids ENERGETICS OF LIVING CELLS Energy: Concepts and Definitions Transfer of Chemical Energy by Coupled Reactions ATP: Energy Source for the Cell Temperature and Reaction Rates ENZYMES: GENERAL PROPERTIES Enzyme Specificity, and Active Sites Mechanism of Catalysis by Enzymes Effects of Temperature and ph on Enzymatic Reactions Cofactors Enzyme Kinetics Enzyme Inhibition REGULATION OF METABOLIC REACTIONS Control of Enzyme Synthesis Control of Enzyme Activity METABOLIC PRODUCTION OF ATP Oxidation, Phosphorylation, and Energy Transfer EFFICIENCY OF ENERGY METABOLISM CHAPTER 4 Membranes, Channels, and Transport MEMBRANE STRUCTURE AND ORGANIZATION 79 36 Membrane Composition 79 The Fluid Mosaic Model of Membranes 81 36 Variation in Membrane Form 83 36 CROSSING THE MEMBRANE: AN OVERVIEW 83 37 Diffusion 84 Membrane Flux 84 38 Osmosis 8 41 41 41 42 42 44 44 4 46 47 48 48 49 49 1 2 2 3 4 7 9 9 61 62 64 6 6 6 66 66 67 70 71 71 71 72 74 7 79
Integration of Physiological Systems Osmolarity and Tonicity Electrical Influences on Ion Distribution Donnan Equilibrium OSMOTIC PROPERTIES OF CELLS Ionic Steady State Cell Volume MECHANISMS FOR TRANSMEMBRANE MOVEMENTS Passive (Simple) Diffusion Passive Transport (Facilitated Diffusion) Stio{:tt^4ft/(.-i Artificial Bilayers Active Transport The Na + /K + Pump as a Model of Primary Active Transport Cotransport Summary of Primary Active Transport ION GRADIENTS AS A SOURCE OF ENERGY Production of Electrical Signals Chemiosmotic Energy Transduction MEMBRANE SELECTIVITY Selectivity for Electrolytes Selectivity for Nonelectrolytes ENDOCYTOSIS AND EXOCYTOSIS Mechanisms of Endocytosis Mechanisms of Exocytosis JUNCTIONS BETWEEN CELLS Gap Junctions Tight Junctions EPITHELIAL TRANSPORT Active Salt Transport Across an Epithelium Transport of Water PART 2 PHYSIOLOGICAL PROCESSES CHAPTER The Physical Basis of Neuronal Function OVERVIEW OF NEURONAL STRUCTURE, FUNCTION, AND ORGANIZATION Transmission of Signals in a Single Neuron Transmission of Signals Between Neurons Organization of the Nervous System MEMBRANE EXCITATION Measuring Membrane Potentials Steottlefit >-' The Discovery of "Animal Electricity" Electrical Properties of Membranes The Role of Ion Channels PASSIVE ELECTRICAL PROPERTIES OF MEMBRANES Membrane Resistance and Conductance Membrane Capacitance Sfwitlg-U: <j-i of Different Cells Comparing Membrane Properties ELECTROCHEMICAL POTENTIALS The Nernst Equation: Calculating the Equilibrium Potential for Single Ions 86 86 87 88 88, 89 90 91 92 94 96 96 97 98 99 99 99 100 100 101 101 102 102 103 103 10 10 106 108 111 113 113 11 11 117 1 1 119 120 122 122 124 124 124 126 128 S-ta-otfijrM: -) A Quantitative Consideration of Charge Separation Across Membranes 128 The Goldman Equation: Calculating the Steady-State Potential for Multiple Ions THE RESTING POTENTIAL The Role of Ion Gradients and Channels The Role of Active Transport ACTION POTENTIALS General Properties of Action Potentials Ionic Basis of the Action Potential Sh..jti.itrUt -4 Pioneering Studies of Action Potentials in the Squid Giant Axon THE NATURE OF ION CHANNELS ShottiVfit - The Voltage-Clamping Method Voltage-Gated Na + Channels SHaifltAtk -6 Patch-Clamping Techniques Voltage-Gated K + Channels Absolute and Relative Refractory Periods Intracellular Ions and the Na + /K + Pump in Action Potentials 146 Molecular Structure of Voltage-Gated Ion Channels 146 Spo-tLlrUl: <y~y Ion Channels in Novel Locations: Artificial Membranes and Cellular Expression Systems 148 Other Electrically Excitable Channels 149 CHAPTER 6 Communication Along and Between Neurons TRANSMISSION OF SIGNALS IN THE NERVOUS SYSTEM: AN OVERVIEW TRANSMISSION OF INFORMATION WITHIN A SINGLE NEURON Passive Spread of Electrical Signals Propagation of Action Potentials Sbolllglit 6-1 Confirmation that Local Circuits Contribute to Action Potentials Speed of Propagation Sbutfi^fit 6-2. Extracellular Signs of Impulse Conduction Sfco-tkp-Iit 6-* I O ' Velocity Axon Diameter and Conduction 1 1 17 17 19 161 162 163 164 Rapid, Saltatory Conduction in Myelinated Axons 16 TRANSMISSION OF INFORMATION BETWEEN NEURONS 166 Synaptic Structure and Function: Electrical Synapses 167 Synaptic Structure and Function: Chemical Synapses 167 Fast Chemical Synapses 170 Sh.«rtkf M: &~\ Pharmacological Agents Useful in Neurophysiological Studies 173 StaotliVfii-6- Calculation of Reversal Potential 177 PRESYNAPTIC RELEASE OF NEUROTRANSMITTERS 1 Quantal Release of Neurotransmitters 1 Depolarization-Release Coupling 2
The Physical Basis Of Neuronal Function Nonspiking Release THE CHEMICAL NATURE OF NEUROTRANSMITTERS Neurotransmitters of Fast, Direct Synaptic Transmission Neurotransmitters of Slow, Indirect Synaptic Transmission POSTSYNAPTIC MECHANISMS Receptors and Channels in Fast, Direct Neurotransmission Receptors in Slow, Indirect Neurotransmission Neuromodulation INTEGRATION AT SYNAPSES SYNAPTIC PLASTICITY Short-Term Homosynaptic Modulation: Facilitation, 6 6 6 9 190 190 194 196 198 202 i f Depression, and Potentiation 203 ( Heterosynaptic Modulation 20 ; Long-Term Potentiation and Depression 20 Sixoi[i^-(tt6-6 Neurotransmitters and Psychoactive : Drugs 208 I CHAPTER 7 Sensing the Environment 21 I GENERAL PROPERTIES OF SENSORY I RECEPTION 216 I Properties of Receptor Cells 216 ['. Common Mechanisms and Molecules of Sensory I Transduction 2 If From Transduction to Neuronal Output 220 f Encoding Stimulus Intensities 222 6 Control of Sensory Sensitivity 22 I THE CHEMICAL SENSES: TASTE AND I SMELL 230 Mechanisms of Taste Reception 231 I Mechanisms of Olfactory Reception 234 MECHANORECEPTION 238 j Insect Mechanoreceptors 239 [ Hair Cells 241 * Organs of Equilibrium 242 { The Vertebrate Ear 242 SholiiAdy-\ Mechanics of the Basilar Membrane 246 * An Insect Ear 249 I Sjiotdf M: y-2. Unusually Sensitive Sensory i Receptors 20 'i THERMORECEPTION 22 I VISION 22 Optic Mechanisms: Evolution and Function 23 Compound Eyes 24 I Sjj.oikjrk.t7- Subjective Correlates of Primary i Photoresponses 27 [ The Vertebrate Eye 28 I Photoreception: Converting Photons into Neuronal I Signals 262 Sfcoik.rk.t7-4 The Electroretinogram. 264 i Stiotiieki-y-'y Light, Paint, and Color Vision 270 LIMITATIONS ON SENSORY RECEPTION 271 CHAPTER 8 The Structural and Functional Organization of the Nervous System 277 THE FLOW OF INFORMATION IN THE NERVOUS SYSTEM 277 Sh.atkjrk.t8-i Sensing Effectors as Housekeepers 279 EVOLUTION OF NERVOUS SYSTEMS 280 ORGANIZATION OF THE VERTEBRATE NERVOUS SYSTEM 284 The Spinal Cord 28 The Brain 287 The Autonomic Nervous System 294 CHAPTER 9 Glands and Hormones 301 CELLULAR SECRETIONS 301 Sfe.otkjrk.t9-1 Substances with Similar Structures and Functions Secreted by Different Organisms 303 Surface Secretions: The Cell Coat, Collagen, and Mucus 303 Packaging and Transport of Secreted Material 304 Storage of Secreted Substances 307 SECRETORY MECHANISMS 307 Glandular Secretions 309 Types and General Properties of Glands 309 ENDOCRINE GLANDS 310 NEUROENDOCRINE SYSTEMS 314 v Hypothalamic Control of the Anterior Pituitary Gland 316 S-h.«jtk«rki cj-i Peptide Hormones 317 Glandular Hormones Released from the Anterior Pituitary Gland 317 Neurohormones Released from the Posterior Pituitary Gland 3 CELLULAR MECHANISMS OF HORMONE ACTION 320 Lipid-Soluble Hormones and Their Receptors 321 Lipid-Insoluble Hormones and Intracellular Signaling 321 PHYSIOLOGICAL EFFECTS OF HORMONES 333 Metabolic and Developmental Hormones 333 Hormones that Regulate Water and Electrolyte Balance 342 Reproductive Hormones 344 Prostaglandins, 348 HORMONAL ACTION IN INVERTEBRATES 349 EXOCRINE GLANDS 32 The Vertebrate Salivary Gland 32 Invertebrate Silk Glands 34 ENERGY COST OF GLANDULAR ACTIVITY 36 CHAPTER 10 Muscles and Animal Movement 361 ESSENTIALS OF SKELETAL MUSCLE CONTRACTION 361 Myofilament Substructure 364 Contraction of Sarcomeres: The Sliding-Filament Theory 366
xii Integration of Physiological Systems Sh.otkjrk.tio~! The Geomehy of Muscle 368 Cross-Bridges and the Production of Force 371 Sh.otkirk.tio-1 Motor Molecules 372 Sbotkjrkt 10- Skinned Muscle Fibers 374 MECHANICS OF MUSCLE CONTRACTION 37 Relation Between Force and Shortening Velocity 376 Effect of Cross-Bridges on the Force-Velocity Relation 377 REGULATION OF MUSCLE CONTRACTION 379 The Role of Calcium in Cross-Bridge Attachment 379 Excitation-Contraction Coupling 380 The Contraction-Relaxation Cycle 387 THE TRANSIENT PRODUCTION OF FORCE 389 Series Elastic Components of Muscle 390 The Active State 391 Contractile States: Twitches and Tetanus 391 ENERGETICS OF MUSCLE CONTRACTION 393 ATP Consumption by the Myosin ATPase and Calcium Pumps 393 Regeneration of ATP During Muscle Activity 393 FIBER TYPES IN VERTEBRATE SKELETAL MUSCLE 394 Classification of Fiber Types 394 Functional Rationale for Different Fiber Types 396 ADAPTATION OF MUSCLES FOR VARIOUS ACTIVITIES 397 Adaptation for Power: Jumping Frogs 397 Adaptation for Contrasting Functions: Swimming Fish 400 Adaptation for Speed: Sound Production 40 High Power and High Frequency: Insect Asynchronous Flight Muscles 408 NEURONAL CONTROL OF MUSCLE CONTRACTION 411 Motor Control in Vertebrates 411 Motor Control in Arthropods 413 CARDIAC MUSCLE 414 SMOOTH MUSCLE 417 Vertebrate Single-Unit and Multi-Unit Smooth Muscles 417 Regulation of Smooth-Muscle Contraction 4 Unusual Features of Smooth-Muscle Contraction 420 CHAPTER 11 Behavior: Initiation, Patterns, and Control 42 Sb.otkirk.t I i -1 Behavior in Animals Without Nervous Systems 426 THE NEURONAL BASIS OF A SIMPLE BEHAVIOR 427 Simple Reflexes 427 Principles of Neuronal Control 430 THE STUDY OF BEHAVIOR 431 Basic Behavioral Concepts 431 Examples of Behavior 434 PROPERTIES OF NEURONAL CIRCUITS 439 Sensory Networks hottijfkt ii-i Motor Networks Sensory Tuning Curves taoik«-kt 11- Endocrine Control of Bird Song Neuroendocrine Control of Behavior PART 3 INTEGRATION OF PHYSIOLOGICAL SYSTEMS CHAPTER 12 Circulation GENERAL PLAN OF THE CIRCULATORY SYSTEM Open Circulations Sfe-<>tk<rk.t 12.-1 Pressure Measurement Closed Circulations THE HEART Electrical Properties of the Heart Mechanical Properties of the Heart Shotlivk.412.-2, The Frank-Starling Mechanism Vertebrate Hearts: Comparative Functional Morphology HEM0DYNAMICS Laminar and Turbulent Flow Relationship Between Pressure and Flow THE PERIPHERAL CIRCULATION Arterial System Venous System Capillaries and the Microcirculation The Lymphatic System CIRCULATION AND THE IMMUNE RESPONSE REGULATION OF CIRCULATION Control of the Central Cardiovascular System Control of the Microcirculation CARDIOVASCULAR RESPONSE TO EXTREME CONDITIONS Exercise Diving Hemorrhage CHAPTER 13 Gas Exchange and Acid-Base Balance OXYGEN AND CARBON DIOXIDE IN THE PHYSICAL ENVIRONMENT Sh.otk.rkt ij-i Exchange in Animals Early Experiments on Gas OXYGEN AND CARBON DIOXIDE IN LIVING SYSTEMS Sjxotkekt 1 j-2. Effects of Human Activity on Environmental Oxygen and Carbon Dioxide Levels Staotlwk.t }-} The Gas Laws OXYGEN AND CARBON DIOXIDE IN BLOOD Respiratory Pigments Oxygen Transport in Blood
The Physical Basis Of Neuronal Function xiii Carbon Dioxide Transport in Blood 34 Transfer of Gases to and from the Blood 3 REGULATION OF BODY ph 39 Hydrogen Ion Production and Excretion 40 Hydrogen Ion Distribution Between Compartments 41 Factors Influencing Intracellular ph 43 Factors Influencing Body ph 44 GAS TRANSFER IN AIR: LUNGS AND OTHER SYSTEMS 4 Functional Anatomy of the Lung 4 Lung Ventilation 47 Sfaotkjrkt i}~4 Lung Volumes 48 Pulmonary Circulation 0 Mechanisms for Ventilation of the Lung 1 Pulmonary Surfactants Heat and Water Loss Across the Lung Gas Transfer in Bird Eggs 6 Insect Tracheal Systems 7 GAS TRANSFER IN WATER: GILLS 9 Functional Anatomy of the Gill 62 REGULATION OF GAS TRANSFER 62 Ventilation-to-Perfusion Ratios 62 Neural Regulation of Breathing 64 RESPIRATORY RESPONSES TO EXTREME CONDITIONS 68 Reduced Oxygen Levels (Hypoxia) 68 Increased Carbon Dioxide Levels (Hypercapnia) 70 Diving by Air-Breathing Animals 70 Exercise 71 SYMMORPHOSIS _^- 72 SWIMBLADDERS: OXYGEN ACCUMULATION AGAINST LARGE GRADIENTS 73 The Rete Mirabile 74 Oxygen Secretion 74 CHAPTER 14 Ionic and Osmotic 79 Balance PROBLEMS OF OSMOREGULATION 79 OSMOREGULATORS AND OSMOCONFORMERS 82 OBLIGATORY EXCHANGES OF IONS AND WATER 83 Gradients Between the Animal and the Environment 83 Surface-to-Volume Ratio 83 Permeability of the Integument 83 Feeding, Metabolic Factors, and Excretion 8 Water Loss via Respiration 86 OSMOREGULATION IN AQUEOUS AND TERRESTRIAL ENVIRONMENTS 88 Water-Breathing Animals 88 Air-Breathing Animals 90 OSMOREGULATORY ORGANS 93 THE MAMMALIAN KIDNEY 96 Anatomy of the Mammalian Kidney 96 Urine Production 98 Shotkekti4.-i Renal Clearance 603 Regulation of ph by the Kidney 608 The Urine-Concentrating Mechanism 610 Sfe.oikjrkt 14.-2, Countercurrent Multiplier Systems 611 Control of Water Reabsorption 613 NON-MAMMALIAN VERTEBRATE KIDNEYS 614 EXTRARENAL OSMOREGULATORY ORGANS IN VERTEBRATES 61 Salt Glands 61 Fish Gills 6 INVERTEBRATE OSMOREGULATORY ORGANS 621 Filtration-Reabsorption Systems 621 Secretion-Reabsorption Systems 622 EXCRETION OF NITROGENOUS WASTES 624 Ammonia-Excreting (Ammonotelic) Animals 62 Urea-Excreting (Ureotelic) Animals 626 Uric Acid-Excreting (Uricotelic) Animals 627 CHAPTER 1 Acquiring Energy: Feeding, Digestion, and Metabolism 631 FEEDING METHODS 631 Food Absorption Through Exterior Body Surfaces 631 Endocytosis 631 Filter Feeding 632 Fluid Feeding 633 Seizing Prey 634 Herbivory and Grazing 637 OVERVIEW OF ALIMENTARY SYSTEMS 637 Headgut: Food Reception 640 Foregut: Food Conduction, Storage, and Digestion 642 Midgut: Chemical Digestion and Absorption 643 Hindgut: Water and Ion Absorption and Defecation 646 Dynamics of Gut Structure and the Influence of Diet ; 647 MOTILITY OF THE ALIMENTARY CANAL 648 Muscular and Ciliary Motility 648 Peristalsis 648 Control of Motility 649 GASTROINTESTINAL SECRETIONS 61 Exocrine Secretions of the Alimentary Canal 61 Control of Gastrointestinal Secretions 6 Sfeottijrkt 1-1 Behavioral Conditioning in Feeding and Digestion 66 ABSORPTION 69 Nutrient Uptake in the Intestine 660 Transport of Nutrients in the Blood 661 Water and Electrolyte Balance in the Gut 662 NUTRITIONAL REQUIREMENTS 663 Nutrient Molecules 663 CHAPTER 16 Energy Expenditure: Body Size, Locomotion, and Reproduction 667 THE CONCEPT OF ENERGY METABOLISM 667 DESCRIBING AND MEASURING METABOLIC RATE 667 Basal and Standard Metabolic Rates 668
xiv Integration of Physiological Systems The Cellular and Biochemical Components of Basal Metabolic Rate 669 Metabolic^Scope 669 Direct Calorimetry: Measuring Metabolism from Heat Production 670 StaotliVkt i6-i Energy Units (or When Is a Calorie Not a Calorie?) 671 Indirect Calorimetry: Measuring Metabolism from Food Intake and Waste Excretion 671 Radioisotopes: Measuring Metabolism by Tracking Atom Movements 671 Respirometry: Measuring Metabolism from Gas Exchange 672 Respiratory Quotient 673 Energy Storage 674 Specific Dynamic Action 67 BODY SIZE AND METABOLIC RATE 67 Isometric and Allometric Scaling of Metabolism 67 Interspecific versus Intraspecific Allometry 679 ENERGETICS OF LOCOMOTION 681 Animal Size, Velocity, and Cost of Locomotion 681 Physical Factors Affecting Locomotion 683 Aquatic, Aerial, and Terrestrial Locomotion 683 Sbotk^-kt 16-2. The Reynolds Number: Implications for Large and Small Animals 684 Locomotory Energetics of Ectotherms versus Endotherms 688 BODY RHYTHMS AND ENERGETICS 688 Circadian Rhythms 689 Noncircadian Endogenous Rhythms 690 Biological Rhythms, Temperature Regulation, and Metabolism 690 ENERGETICS OF REPRODUCTION 693 Patterns of Energetic Investment in Reproduction 693 The Cost of Egg and Sperm Production 694 Parental Care as an Energy Cost of Reproduction 69 DETERMINANTS OF BODY HEAT AND TEMPERATURE 704 The Physics of Heat Gain, Loss, and Storage 704 General Biological Mechanisms for Regulating Heat Transfer 706 PHYSIOLOGICAL CLASSIFICATION USING THERMAL BIOLOGY 707 Homeothermy and Poildlothermy 707 Endothermy and Ectothermy 708 Thermal Classification of Real versus Ideal Animals 709 THERMAL BIOLOGY OF ECTOTHERMS 710 Ectotherms in Cold and Freezing Environments 710 Ectotherms in Warm and Hot Environments 712 Costs and Benefits of Ectothermy Relative to Endothermy 714 THERMAL BIOLOGY OF HETEROTHERMS 71 THERMAL BIOLOGY OF ENDOTHERMS 717 The Concept of the Thermal Neutral Zone 7 Endothermy in Cold Environments: Producing and Retaining Body Heat 719 Endothermy in Hot Environments: Dissipating Body Heat 722 NEURONAL MECHANISMS OF TEMPERATURE CONTROL 72 Thermostatic Regulation of Body Temperature 72 Thermoregulation During Activity 728 THERMOREGULATION AND SPECIALIZED METABOLIC STATES 730 Sleep 730 Torpor 730 Hibernation and Winter Sleep 731 Estivation 732 Pyrogens and Fever 732 ENERGY, ENVIRONMENT, AND EVOLUTION 733 CHAPTER 1 7 Energetic Costs of Meeting Environmental Challenges 699 TEMPERATURE DEPENDENCE OF METABOLIC RATE 699 The Biochemical and Molecular Basis for Thermal Influences on Metabolism 699 Appendix 1: Units, Conversions, Constants, and Definitions Appendix 2: Logs and Exponentials References Cited Glossary Index Rl Gl II