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F O U R T H E D I T I O N E C K E R T ANIMAL PHYSIOLOGY M E C H A N I S M S A N D A D A P T A T I O N S D A V I D R A N D A L L U N I V E A S ~ T W O F B R I T I S H C O L U M B I A 1 W A R R E N B U R G G R E N K A T H L E E N F R E N C H U N I V E R S I T Y O F C A L I F O R N I A , S A N D L E G O W I T H C O N T R I B U T I O N S B Y R U S S E L L F E R N A L D S T A N F O R D U N I V E R S I T Y W. H. Freeman and Company New Ynrk ACQUISITIONS EDITOR: Deborah Allen DEVELOPMENT EDITOR: Kay Ueno PROJECT EDITOR: Kate Ahr PHOTO RESEARCH: Larry Marcus COVER DESIGNER: Michael Mendelsohn, Design 2000, Inc FRONT COVER PHOTOGRAPH: Arctic fox, Canada O Daniel J. Cox/Tony Stone Images BACK COVER ILLUSTRATION: Roberto Osti ' TEXT DESIGNERS: Michael Mendelsohn, Design 2000, Inc.; Victoria Tomaselli ILLUSTRATION COORDINATOR: Bill Page ILLUSTRATION: Fine Line Illustrations; Medical and Scientific Illustration, William C. Ober, M D and Claire Garrison, RN, BA PRODUCTION COORDINATOR: Maura Studley COMPOSITION: Progressive Information Technologies MANUFACTURING: RR Donnelley & Sons Company L~brary of Congress Cataloging-in-Publication Data Randall, Dav~d J., 1938- Eckert an~mal phys~ology: mechanisms and adaptations/Dav~d Randall, Warren Burggren, Kathleen French.-4th ed. P. cm. Includes blbl~ograph~cal references and ~ndex. ISBN 0-71 67-2414-6 (hardcover) 1. Physiology. I. Burggren, Warren. 11. French, Kathleen. 111. Tltle. QP31.2.R36 1997 591.1-dc20 96-31713 CIP Copyright O 1978, 1983, 1988, 1997 by W. H. Freeman and Company. All rights reserved. No part of this book may be reproduced by any mechanical, photographic, or electronic process, or in the form of a phonographic recording, nor may it be stored in a retrieval system, transmitted, or otherwise copied for public or private use, without written permission from the publisher. Printed in the United States of America Second printing, 1997, RRD ......... ... ,. . . . . . . . . . . . . . . , , , , ,,w~:~,;~,',;:;.,;~~,:2~~;:;;~~~::~q p;?, ,,$!,>,!;., f,: ::y:.;;;z$;@,&$,:.; :j!j?;:, .~i;~!~~~,,$$$~~!~~g~~;~~&~~~,~i~:~~t~:,,~~~~,~~;~~~!~,!~$~ ,:, ;m&.:,: ',:. :.L, :...'. ::.v -., .:::, '-:.. ,,:i.,,:+ A B O U T T H E A U T H O R S DAVID RANDALL A prominent fish physiologist and a leading expert in respiratory and circulatory physiology, David Randall collaborated with the late Roger Eckert on the earlier editions of Animal Physiology and continues his con- tribution in the fourth. A faculty member at the University of British Columbia in Vancouver, Canada, since 1963, and full professor since 1973, Randall was appointed Associate Dean of Graduate Studies in 1990. Elected a fellow of the Royal Society of Canada in 1981, Randall has been both a Guggenheim and a Killam fellow, and was awarded the prestigious Fry Medal for research contributions to zoology by the Canadian Soci- ety of Zoology in 1993. In 1995, he received the Award of Excellence from the American Fisheries Society for . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . WARREN BURGGREN Warren Burggren has taught in physiology for 23 years, and has been a professor of biological sciences at the Uni- versity of Nevada at Las Vegas since 1992. Courses he has taught at UNLV and at the University of Massachu- setts, where he was Professor of Zoology from 1987 through 1991, include Human Anatomy and Physiology, Bioenergetics, Introductory Zoology, and Comparative Physiology. Burggren's research interest include developmental physiology, comparative animal physiology, and environmental and ecological physiology. In particular, his research focuses on the ontogeny of respiratory and cardiovascular systems, and how the systems that regu- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . KATHLEEN FRENCH A neurobiologist at the University of California at San Diego since 1985, Kathleen French has for 10 years taught upper division courses in embryology, mammalian physiology for premedical students, and cellular neurobi- ology. In addition, at UCSD, French participates in a training program to instruct science teaching assistants in the techniques and philosophy of teaching. She also serves on the faculty of the Neuroscience and Behavior Course at the Marine Biological Laboratories in Woods Hole, Massachusetts, an intensive course designed primarily for graduate students and post-doctoral fellows. French . brings her expertise in-and love of-teaching to her contributions to the field of fish physiology. A frequent symposium lecturer on fish physiology and other sub- jects, most recently in Brazil, France, Germany, Italy, the People's Republic of China, Russia, and the United States. He has worked with both the World Health Or- ganization and the United States Environmental Protec- tion Agency in developing ammonia criteria. Widely published as author and co-author in leading journals, Randall is co-editor of the noted series Fish Physiolog?, (Academic Press), of which 15 volumes are in print. Vol- ume 16, subtitled "Deep-sea Fish," will appear in 1997. Along with his other duties, Randall co-teaches third year courses in vertebrate physiology and environmental physiology. His research interests concern the interactions between gas and ion exchange across fish gills. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . late them change over the course of development. Burggren has been actively involved in symposia, semi- nars, and formal extramural researchltraining activities in many countries. A co-author of The Evolution of Air Breathing in Vertebrates (Cambridge University Press, 1981), Burggren has been a frequent contributor since 1980 to edited collections of physiology, including Presser's Comparative Animal Physiology, Fourth Edi- tion (Wiley-Liss, 1991). Burggren co-edited Environ- mental Physiology of the Amphibia (University of Chicago Press, 1992), and more recently co-edited De- velopment of Cardiovascular Systems; Molecules to Or- ganisms (Cambridge University Press, 1997). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . role as co-author of the current edition of Animal Physi- ology, along with a lifelong interest in the nervous systems of organisms from a broad range of phyla. As an Associ- ate Project Scienrist at UCSD, French's research focuses on the control of neuronal development, a topic that she has studied in various invertebrate species. Her current research concerns the cellular events that control differenti- ation of identified neurons in the medicinal leech, with an emphasis on the cellular physiology of embryonic neurons and the effects of cell-cell contacts. She has been the author and co-author of numerous published research and review articles in journals including the Journal of Neuroscience and Journal of Neurophysiology. PART I PRINCIPLES OF PHYSIOLOGY 1 STUDYING ANIMAL PHYSIOLOGY 2 EXPERIMENTAL METHODS FOR EXPLORING PHYSIOLOGY 3 MOLECULES, ENERGY, AND BIOSYNTHESIS 4 MEMBRANES, CHANNELS, AND TRANSPORT PART II PHYSIOLOGICAL PROCESSES 5 THE PHYSICAL BASIS OF NEURONAL FUNCTION 6 COMMUNICATION ALONG AND BETWEEN NEURONS 7 SENSING THE ENVIRONMENT 8 GLANDS: MECHANISMS AND COSTS OF SECRETION 9 HORMONES: REGULATION AND ACTION 10 MUSCLES AND ANIMAL MOVEMENT 351 11 BEHAVIOR: INITIATION, PATTERNS, AND CONTROL 405 PART Ill INTEGRATION OF PHYSIOLOGICAL SYSTEMS 465 12 CIRCULATION 467 13 GAS EXCHANGE AND ACID-BASE BALANCE 517 14 IONIC AND OSMOTIC BALANCE 571 15 ACQUIRING ENERGY: FEEDING, DIGESTION. AND METABOLISM 627 16 USING ENERGY: MEETING ENVIRONMENTAL CHALLENGES 665 ,.::,:; $:::oi;, ,:, ,,,< :$,:',:,;;?:$ :{;, %<+,;>? :, , , , , , , , , ,, ,,*, :-,: >:,'.!I:, , , > , ; , J;:, ,,<,,, , ; ! < ) , ' i., 'if '?:'ii ,>,, 'a ,', ,."%q F": : ':;i.?'.;.;,', , . : , ., , , , ,, ,,, , , , , . . , , , , , , , ; .:-.: . . ,, ,,,,, .. 1 I i C O N T E N T S I I I Preface ix Acknowledgments xvii PART I PRINCIPLES OF PHYSIOLOGY CHAPTER 1 STUDYING ANIMAL PHYSIOLOGY 3 THE SUBDISCIPLINES OF ANIMAL PHYSIOLOGY 4 WHY STUDY ANIMAL PHYSIOLOGY? 4 Sc~ent~fic Curlos~ty 4 Commerc~aYAgr~cultural Appl~cat~ons 4 Inslghts into Human Physiology 4 CENTRAL THEMES IN ANIMAL PHYSIOLOGY 4 Structure-Funct~on Relat~onsh~ps 5 Adaptation, Accl~mat~zat~on, and Accl~mat~on 5 Homeostas~s 7 Feedback-Control Systems 8 Conform~t~ and Regulation 9 LITERATURE OF PHYSIOLOGICAL SCIENCES 10 SPOTLIGHT 1-1 THE CONCEPT OF FEEDBACK 12 ANIMAL EXPERIMENTATION IN PHYSIOLOGY 13 Summary 13 Rev~ew Quest~ons 14 Suggested Readlngs 14 CHAPTER 2 EXPERIMENTAL METHODS FOR EXPLORING PHYSIOLOGY FORMULATING AND TESTING HYPOTHESES The August Krogh Prlnc~ple Experimental Deslgn and Phys~olog~cal Level MOLECULAR TECHNIQUES Traclng Molecules w~th Radlolsotopes Traclng Molecules wlth Monoclonal Antlbod~es Genetic Englneerlng CELLULAR TECHNIQUES Uses of M~croelectrodes and Mlcroplpettes Structural Analys~s of Cells Cell Culture BIOCHEMICAL ANALYSIS Measurlng Composit~on: What Is Present Measurlng Concentration: How Much IS Present EXPERIMENTS WITH ISOLATED ORGANS AND ORGAN SYSTEMS OBSERVING AND MEASURING ANIMAL BEHAVIOR The Power of Behav~oral Experiments Methods In Behav~oral Research IMPORTANCE OF PHYSIOLOGICAL STATE IN RESEARCH Summary Revlew Quest~ons Suggested Readlngs CHAPTER 3 MOLECULES, ENERGY, AND BIOSYNTHESIS ORIGIN OF KEY BIOCHEMICAL MOLECULES ATOMS, BONDS, AND MOLECULES THE SPECIAL ROLES OF H, 0, N, AND C IN LIFE PROCESSES WATER: THE UNIQUE SOLVENT The Water Molecule Propert~es of Water Water as a Solvent PROPERTIES OF SOLUTIONS Concentration, Coll~gatlve Properties, and Activ~ty Ionizat~on of Water Ac~ds and Bases The B~olog~cal Importance of pH Henderson-Hasselbalch Equat~on Buffer Systems Electrlc Current ~n Aqueous Solutions SPOTLIGHT 3-1 ELECTRICAL TERMINOLOGY AND CONVENTIONS Blndlng of Ions to Macromolecules X CONTENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . BIOLOGICAL MOLECULES Llplds Carbohydrates Protelns Nucle~c Ac~ds ENERGETICS OF LIVING CELLS Energy: Concepts and Defin~t~ons Transfer of Chem~cal Energy by Coupled React~ons ATP: Energy Carrier of the Cell Temperature and React~on Rates ENZYMES: GENERAL PROPERTIES Enzyme Speclficlty and Actlve S~tes Mechanism of Catalys~s by Enzymes Effect of Temperature and pH on Enzymat~c React~ons Cofactors Enzyme Klnetlcs Enzyme Inhlb~t~on REGULATION OF METABOLIC REACTIONS Control of Enzyme Synthes~s Control of Enzyme Actlvlty METABOLIC PRODUCTION OF ATP Oxldat~on, Phosphorylatlon, and Energy Transfer Glycolys~s Cltr~c Ac~d Cycle Effic~ency of Energy Metabol~sm Oxygen Debt Summary Rev~ew Quest~ons Suggested Readlngs CHAPTER 4 MEMBRANES, CHANNELS, AND TRANSPORT MEMBRANE STRUCTURE AND ORGANIZATION Membrane Composlt~on Flu~d Mosa~c Membranes SPOTLIGHT 4-1 THE CASE FOR A LIPID BILAYER MEMBRANE Var~at~on In Membrane Form CROSSING THE MEMBRANE: AN OVERVIEW D~ffus~on Membrane Flux Osmos~s Osmolarlty and Tonlclty Electr~cal Influences on Ion D~str~but~on Donnan Equll~brlurn OSMOTIC PROPERTIES OF CELLS Ion~c Steady State Cell Volume PASSIVE TRANSMEMBRANE MOVEMENTS Slmple D~ffus~on through the Llp~d Bllayer D~ffus~on through Membrane Channels SPOTLIGHT 4-2 ARTIFICIAL BILAYERS Facllltated Transport across Membranes ACTIVE TRANSPORT The Na+lK+ Pump as a Model of Aave Transport . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ion Gradients as a Source of Cell Energy Coupled Transport MEMBRANE SELECTIVITY Selectlvlty for Electrolytes Selectlv~ty for Nonelectrolytes ENDOCYTOSIS AND EXOCYTOSIS Mechanlsms of Endocytosis Mechanlsms of Exocytosls JUNCTIONS BETWEEN CELLS Gap Junct~ons Tlght Junct~ons EPITHELIAL TRANSPORT Actlve Salt Transport across an Ep~thellum Transport of Water Summary Rev~ew Quest~ons Suggested Readlngs PART II PHYSIOLOGICAL PROCESSES CHAPTER 5 THE PHYSICAL BASIS OF NEURONAL FUNCTION OVERVIEW OF NEURONAL STRUCTURE, FUNCTION, AND ORGANIZATION Transmlss~on of S~gnals In a Slngle Neuron Transm~ss~on of Slgnals Between Neurons Organ~zat~on of the Nervous System MEMBRANE EXCITATION Measuring Membrane Potentials Dlstlngu~shlng Passlve and Actlve Membrane Electr~cal Propert~es SPOTLIGHT 5-1 THE DISCOVERY OF "ANIMAL ELECTRICITY" Role of Ion Channels PASSIVE ELECTRICAL PROPERTIES OF MEMBRANES Membrane Res~stance and Conductance Membrane Capac~tance ELECTROCHEMICAL POTENTIALS The Nernst Equat~on: Calculatlng the Equ~l~brlum Potent~al for Slngle Ions SPOTLIGHT 5-2 A QUANTITATIVE CONSIDERATION OF CHARGE SEPARATION ACROSS MEMBRANES The Goldman Equation: Calculatlng the Equll~brlum Potent~al for Mult~ple Ions THE RESTING POTENTIAL Role of Ion Grad~ents and Channels Role of Actlve Transport ACTION POTENTIALS General Propert~es of Act~on Potent~als Ion~c Baas of the Act~on Potent~al SPOTLIGHT 5-3 THE VOLTAGE-CLAMP METHOD Changes In Ion Concentrat~on durlng Exc~tat~on OTHER ELECTRICALLY EXCITED CHANNELS Summary Review Questions Suggested Readings CHAPTER 6 COMMUNICATION ALONG AND BETWEEN NEURONS TRANSMISSION OF SIGNALS IN THE NERVOUS SYSTEM: AN OVERVIEW TRANSMISSION OF INFORMATION WITHIN A SINGLE NEURON Pass~ve Spread of Electr~cal S~gnals Propagatlon of Actlon Potentials Speed of Propagat~on Rapid, Saltatory Conduction In Myelmated Axons SPOTLIGHT 6-1 EXTRACELLULAR SIGNS OF IMPULSE CONDUCTION SPOTLIGHT 6-2 AXON DIAMETER AND CONDUCTION VELOCITY TRANSMISSION OF INFORMATION BETWEEN NEURONS: SYNAPSES / Synapt~c Structure and Function: Electr~cal Synapses Synaptlc Structure and Function: Chermcal Synapses Fast Chemical Synapses SPOTLIGHT 6-3 PHARMACOLOGICAL AGENTS USEFUL IN SYNAPTIC STUDIES SPOTLIGHT 6-4 CALCULATION OF REVERSAL POTENTIAL PRESYNAFTIC RELEASE OF NEUROTRANSMITTERS Quanta1 Release of Neurotransm~tters Depolarizat~on-Release Coupllng Nonsplking Release THE CHEMICAL NATURE OF NEUROTRANSMITTERS Fast, D~rect Neurotransm~ss~on Slow, Indirect Neurotransmission POSTSYNAPTIC MECHANISMS Receptors and Channels in Fast, Direct Neurotransmission Receptors m Slow, Ind~rect Neurotransm~ss~on Neuromodulat~on INTEGRATION AT SYNAPSES SYNAPTIC PLASTICITY Homosynaptlc Modulation: Facil~tatlon Homosynapt~c Modulat~on: Posttetan~c Potentlation Heterosynapt~c Modulation " Long Term Potent~at~on Summary Rev~ew Questions Suggested Read~ngs CHAPTER 7 SENSING THE ENVIRONMENT GENERAL PROPERTIES OF SENSORY RECEPTION Properties of Receptor Cells Common Mechanisms and Molecules of Sensory Transduction From Transduction to Neuronal Output C O N T E N T S xi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Encod~ng Stlmulus Intensities 224 Input-Output Relations 225 Range Fractionation 226 Control of Sensory Senslt~v~ty 226 THE CHEMICAL SENSES: TASTE AND SMELL 231 Mechan~sms of Taste Recept~on 232 Mechanisms of Olfactory Reception 235 MECHANORECEPTION 238 Ha~r Cells 238 Organs of Equ~librlum 241 The Vertebrate Ear 242 An Insect Ear 248 ELECTRORECEPTION 248 THERMORECEPTION 250 VISION 251 Optlc Mechan~sms: Evolut~on and Funct~on 252 Compound Eyes 253 SPOTLIGHT 7-1 SUBJECTIVE CORRELATES OF PRIMARY PHOTORESPONSES 256 The Vertebrate Eye 257 Photoreception: Convert~ng Photons into Neuronal Signals 261 SPOTLIGHT 7-2 THE ELECTRORETINOGRAM 263 SPOTLIGHT 7-3 LIGHT, PAINT, AND COLOR VISION 268 LIMITATIONS O N SENSORY RECEPTION 269 Summary 270 Revlew Quest~ons 271 Suggested Read~ngs 271 CHAPTER 8 GLANDS: MECHANISMS AND COSTS OF SECRETION CELLULAR SECRETION Types and Funct~ons of Secretions Surface Secretions: The Cell Coat and Mucus Packag~ng and Transport of Secreted Material SPOTLIGHT 8-1 SUBSTANCES WITH SIMILAR STRUCTURES AND FUNCTIONS SECRETED BY DIFFERENT ORGANISMS Storage of Secreted Substances Secretory Mechanlsms GLANDULAR SECRETIONS Types and General Properties of Glands Endocrine Glands Exocr~ne Glands ENERGY COST OF GLANDULAR ACTIVITY Summary Rev~ew Questions Suggested Read~ngs CHAPTER 9 HORMONES: REGULATION AND ACTION 301 ENDOCRINE SYSTEMS: OVERVIEW 302 Chem~cal Types and General Functions of Hormones 3 02 Regulat~on of Hormone Secretion 303 NEUROENDOCRINE SYSTEMS 303 Hypothalmlc Control of the Anterlor Pitu~tary Gland 304 xii CONTENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glandular Hormones Released from the Anterlor P~tuitary Gland 305 Neurohormones Released from the Posterlor P~tu~tary Gland 308 SPOTLIGHT 9-1 PEPTIDE HORMONES 310 CELLULAR MECHANISMS OF HORMONE ACTION 311 Lipld-Soluble Hormones and Cytoplasmic Receptors 311 Lipld-Insoluble Hormones and Intracellular Slgnal~ng 312 SPOTLIGHT 9-2 AMPLIFICATION BY ENZYME CASCADES 322 PHYSIOLOGICAL EFFECTS OF HORMONES 328 Metabolic and Developmental Hormones 328 Hormones That Regulate Water and Electrolyte Balance 336 Reproduct~ve Hormones 338 Prostagland~ns 342 HORMONAL ACTION IN INVERTEBRATES 343 Summary 346 Revlew Quest~ons 348 Suggested Readings 349 CHAPTER 10 MUSCLES AND ANIMAL MOVEMENT STRUCTURAL BASIS OF MUSCLE CONTRACTION Myofilament Substructure Contraction of Sarcomeres: The Slldlng Fllament Theory Cross-Bridges and the Productlon of Force SPOTLIGHT 10-1 PARALLEL AND SERIES ARRANGEMENTS: THE GEOMETRY OF MUSCLE MECHANICS OF MUSCLE CONTRACTION Relatlon Between Force and Shortening Veloclty SPOTLIGHT 10-2 SKINNED MUSCLE FIBERS Effect of Cross-Brldges on Force-Veloclty Relatlon REGULATION OE CONTRACTION Role of Calclum In Cross-Brldge Attachment Excltatlon-Contraction Coupllng Contraction-Relaxat~on Cycle THE TRANSICNT PRODUCTION OF FORCE Serles Elast~c Component The Act~ve State Twitches and Tetanus ENERGETICS OF MUSCLE CONTRACTION ATP Usage by Myosln ATPase and Calclum Pumps Regeneration of ATP durlng Muscle Actlvlty FIBER TYPES IN VERTEBRATE SKELETAL MUSCLE Class~ficat~on of Flber Types Functional Rat~onale for Different Flber Types ADAPTATION OE MUSCLES FOR VARIOUS ACTIVITIES Adaptatlon for Power: Jumplng Frogs Diversity of Function: Swlmmmg Fish Adaptatlon for Speed: Sound Productlon Hlgh-Power, Hlgh Frequency Muscles:Asynchronous Fllght Muscles NEURONAL CONTROL OF MUSCLE CONTRACTION Motor Control in Vertebrates Motor Control ln Arthropods CARDIAC MUSCLE SMOOTH MUSCLE Summary Revlew Questions Suggested Readings CHAPTER 11 BEHAVIOR: INITIATION, PATERNS, AND CONTROL 405 SPOTLIGHT 1 1-1 BEHAVIOR IN ANIMALS THAT LACK A NERVOUS SYSTEM 403 EVOLUTION OF NERVOUS SYSTEMS 408 ORGANIZATION OF THE VERTEBRATE NERVOUS SYSTEM 412 Major Dlv~slons of the Central Nervous System 413 The Automatic Nervous System 420 ANIMAL BEHAVIOR 423 Bas~c Behavioral Concepts 423 Examples of Behavlor 426 PROPERTIES OF NEURONAL CIRCUITS 432 Pieces of the Neuronal Puzzle 433 Sensory Networks 434 $SPOTLIGHT 11-2 TUNING CURVES: THE RESPONSE OF A NEURON PLOllED AGAINST THE PARAMETERS OF A STIMULUS 436 SPOTLIGHT 11-3 SPECIFICITY OF NEURONAL CONNECTIONS AND INTERACTIONS 447 Motor Networks 453 Summary 461 Revlew Questions 462 Suggested Reahngs 462 PART Ill INTEGRATION OF PHYSIOLOGICAL SYSTEMS CHAPTER 12 CIRCULATION GENERAL PLAN OF THE CIRCULATORY SYSTEM Open C~rculat~ons Closed C~rculatlons THE HEART Electrical Act~v~ty of the Heart Coronary Clrculat~on Mechanical Properties of the Heart SPOTLIGHT 12-1 THE FRANK-STARLING MECHANISM The Pericardlum Vertebrate Hearts: Comparative Functional Morphology HEMODYNAMICS Laminar and Turbulent Flow Relat~onship between Pressure and Flow THE PERIPHERAL CIRCULATION Arterlal System Venous System Capillaries and the M~croc~rculat~on THE LYMPHATIC SYSTEM . . . CONTENTS xi11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . CIRCULATION AND THE IMMUNE RESPONSE REGULATION OF CIRCULATION Control of the Central Cardiovascular System Control of the M~croc~rculat~on CARDIOVASCULAR RESPONSE T O EXTREME CONDITIONS Exerc~se D~vmg Hemorrhage Summary Rev~ew Questions Suggested Readings CHAPTER 13 GAS EXCHANGE AND ACID-BASE BALANCE GENERAL CONSIDERATIONS SPOTLIGHT 13-1 EARLY EXPERIMENTS ON GAS EXCHANGE IN ANIMALS OXYGEN AND CARBON DIOXIDE IN BLOOD Resp~ratory P~gments SPOTLIGHT 13-2 THE GAS LAWS Oxygen Transport In Blood Carbon D~ox~de Transport m Blood Transfer of Gases to and from the Blood REGULATION OF BODY pH Hydrogen Ion Production and Excret~on Hydrogen Ion D~str~but~on between Compartments Factors Influenc~ng Intracellular pH Factors Influenc~ng Body pH GAS TRANSFER IN AIR: LUNGS AND OTHER SYSTEMS Funct~onal Anatomy of the Lung Pulmonary C~rculat~on SPOTLIGHT 13-3 LUNG VOLUMES Vent~lation of the Lung Pulmonary Surfactants Heat and Water Loss across the Lung Gas Transfer In B~rd Eggs Insect Tracheal Systems GAS TRANSFER IN WATER: GILLS Flow and Gas Exchange across G~lls Funct~onal Anatomy of the G~ll REGULATION OF GAS TRANSFER AND RESPIRATION Ventdat~on-to-Perfus~on Rat~os Neural Regulat~on of Breath~ng RESPIRATORY RESPONSES T O EXTREME CONDITIONS Reduced Oxygen Levels (Hypox~a) Increased Carbon D~ox~de Levels (Hypercapma) D~v~ng by Air-Breathmg An~mals Exerc~se SWIMBLADDERS: OXYGEN ACCUMULATION AGAINST LARGE GRADIENTS The Rete M~rab~le Oxygen Secret~on Summary Review Questions Suggested Readings CHAPTER 14 IONIC AND OSMOTIC BALANCE 571 PROBLEMS OF OSMOREGULATION 571 OBLIGATORY EXCHANGE OF IONS AND WATER 574 Grad~ents Between the An~mal and the Env~ronment 574 Surface-to-Volume Ratlo 574 Permeab~l~ty of the Integument 575 Feedmg, Metabol~c Factors, and Excret~on 577 Temperature, Exerc~se, and Resp~rat~on 5 78 OSMOREGULATORS AND OSMOCONFORMERS 580 OSMOREGULATION IN AQUEOUS AN TERRESTRIAL ENVIRONMENTS 581 Water-Breathmg An~mals 581 hr-breathmg Animals 584 OSMOREGULATORY ORGANS 587 MAMMALIAN KIDNEY 587 Anatomy of the Mammakn ldney 588 Ur~ne Product~on 590 SPOTLIGHT 14-1 RENAL CLEARANCE 595 Regulat~on of pH by the K~dney 601 Ur~ne-Concentratmg Mechan~sm 603 SPOTLIGHT 14-2 COUNTERCURRENT SYSTEMS 604 Control of Water Reabsorpt~on 606 NONMAMMALIAN VERTEBRATE KIDNEYS 608 EXTRARENAL OSMOREGULATORY ORGANS IN VERTEBRATES 608 Salt Glands 608 F~sh G~lls 613 INVERTEBRATE OSMOREGULATORY ORGANS 616 F~ltrat~on-Reabsorpt~on Systems 616 Secretory-Reabsorpt~on Systems 617 EXCRETION OF NITROGENOUS WASTES 620 Ammon~a-Excretmg (Ammonotel~c) An~mals 621 Urea-Excret~ng (Ureotehc) An~mals 623 Ur~c Ac~d-Excretmng (Ur~cotel~c) An~mals 624 Summary 624 Rev~ew Quest~ons 625 Suggested Read~ngs 625 CHAPTER 15 ACQUIRING ENERGY: FEEDING, DIGESTION, AND METABOLISM FEEDING METHODS Food Absorpt~on through Extenor Body Surfaces Endocytos~s Filter Feeding Fluid Feed~ng Se~z~ng of Prey Herb~vory and Graz~ng To Collect Food OVERVIEW OF ALIMENTARY SYSTEMS Headgut: Food Recept~on Foregut: Food Conduct~on, Storage, and Digestion M~dgut: Chemical D~gest~on and Absorpt~on xiv CONTENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hindgut: Water and Ion Absorption and Defecation 643 Dynamics of Gut Structure-Influence of Diet 644 MOTILITY OF THE ALIMENTARY CANAL 644 Muscular and Ciliary Mot~lity 645 Peristalsis 645 Control of Motility 646 GASTROINTESTINAL SECRETIONS 649 Exocr~ne Secret~ons of the Ahrnentary Canal 650 Control of Dlgestlve Secret~ons 653 SPOTLIGHT 15-1 BEHAVIORAL CONDITIONING IN FEEDING AND DIGESTION 654 ABSORPTION 657 Nutnent Uptake in the Intestme 657 Blood Transport of Nutr~ents 658 Water and Electrolyte Balance tn the Gut 659 NUTRITIONAL REQUIREMENTS 66 1 Energy Balance 66 1 Nutr~ent Molecules 66 1 Summary 663 Review Questions 664 Suggested Readlngs 664 CHAPTER 16 USING ENERGY MEETING ENVIRONMENTAL CHALLENGES 665 THE CONCEPT OF ENERGY METABOLISM 665 MEASURING METABOLIC RATE 666 Basal and Standard Metabol~c Rate 666 Metabol~c Scope 667 D~rect Calor~rnetry 668 SPOTLIGHT 16-1 ENERGY UNITS (OR WHEN IS A CALORIE NOTA CALORIE?) 668 Indlrect Calorimetry-Measurement from Food Intake and Waste Excretion 668 Ind~rect Measures of Metabol~c Rate 669 Resp~ratory Quohent 670 Energy Storage 671 Spec~fic Dynam~c Act~on 672 BODY SIZE AND METABOLIC RATE 672 SPOTLIGHT 16-2 THE REYNOLDS NUMBER: IMPLICATIONS FOR BIG AND SMALL ANIMALS 676 TEMPERATURE AND ANIMAL ENERGETICS 677 Temperature Dependence of Metabohc Rate 677 Determ~nants of Body Heat and Temperature 680 Temperature Classlficat~ons of Anlmals 682 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . TEMPERATURE RELATIONS OF ECTOTHERMS Ectotherms In Freezlng and Cold Env~ronments , Ectotherms ln Water and Hot Envlronments Costs and Benefits of Ectothermy: A Compar~son w~th Endothermy TEMPERATURE RELATIONS OF HETEROTHERMS TEMPERATURE RELATIONS OF ENDOTHERMS Mechan~sms for Body Temperature Regulation Thermostatic Regulat~on of Body Temperature Fever DORMANCY: SPECIALIZED METABOLIC STATES Sleep Torpor H~bernat~on and Wlnter Sleep Estlvat~on ENERGETICS OF LOCOMOTION An~mal Sue, Velocity, and Cost of Locomohon Physlcal Factors Affecting Locomotlon Aquat~c, Aer~al, and Terrestrial Locomotlon BODY RHYTHMS AND ENERGETICS C~rcadlan Rhythms Nonclrcad~an Endogenous Rhythms Temperature Regulat~on, Metabohsm, and B~olog~cal Rhythms ENERGETICS OF REPRODUCTION Patterns of Energet~c Investment In Reproduct~on The "Cost" of Gamete Product~on Parental Care as an Energy Cost of Reproduct~on ENERGY, ENVIRONMENT, AND EVOLUTION Summary Rev~ew Quest~ons Suggested Readings Appendzx 1: SI Units Appendlx 2: Logs and Exponentials Appendrx 3: Conversions, Formulas, Physzcal and Chemrcal Constants, Definitions References Czted Glossary Index