238646392 - Animal - Physiology - Eckert

238646392 - Animal - Physiology - Eckert

(Parte 1 de 9)

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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. 1. French, Kathleen. 11. 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.

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Printed in the United States of America Second printing, 1997, RRD

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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 fel- low, and was awarded the prestigious Fry Medal for re- search 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 develop- mental 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-

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 interac- tions 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 re- search concerns the cellular events that control differenti- ation of identified neurons in the medicinal leech, with an emphasis on the cellular physiology of embryonic neu- rons 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.

1 STUDYING ANIMAL PHYSIOLOGY 2 EXPERIMENTAL METHODS FOR EXPLORING PHYSIOLOGY 3 MOLECULES, ENERGY, AND BIOSYNTHESIS 4 MEMBRANES, CHANNELS, AND TRANSPORT

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 1 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

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i CONTENTS

Preface ix Acknowledgments xvii

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

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

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

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~s~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

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

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~s~on Slow, Indirect Neurotransmission POSTSYNAPTIC MECHANISMS

Receptors and Channels in Fast, Direct Neurotransmission Receptors m Slow, Ind~rect Neurotransm~s~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

CONTENTS 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 ON 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

(Parte 1 de 9)

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