Table Of Content

F O U R T H E D I T I O N E C K E R T A N I M A L P H Y S I O L O G Y 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 UNIVEAS~OTFW B RITISHC OLUMBIA 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 UNIVERSITOYF CALIFORNIAS, AN DLEGO WITHC ONTRIBUTIONBSY R U S S E L L F E R N A L D STANFORDU NIVERSITY 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 FRONTC OVER PHOTOGRAPAHr: ctic 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, MDa nd Claire Garrison, RN, BA PRODUCTION COORDINATOR: Maura Studley COMPOSITION: Progressive Information Technologies MANUFACTURING: RR Donnelley & Sons Company L~braryo f Congress Cataloging-in-PublicationD ata Randall, Dav~dJ. , 1938- Eckert an~malp hys~ology:m echanisms and adaptations/Dav~d Randall, Warren Burggren, Kathleen French.-4th ed. P. cm. Includes blbl~ograph~carel ferences and ~ndex. ISBN 0-716 7-2414-6 (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 p;,,$?!,>,!;., ,f ,@ ;z$:;.;,& :;,$ ,;.y::: , :j!j?;:, .~i;~!~~~,,$$$~~!~~g~~;~~&~~~,~,:, i..~..:..~..;~.m t~:&,,...~.:',:. ,~:~: .L~, ,:..~.'. ~::;.v~ -.~,, .~ .::!:, ~'-:,.. !~,$,~:, i.,. .,.,.:..+... ..,.,..w ~:~,;~,',;:;.,;~~,:2~~;:;;~~~::~q A B O U T T H E A U T H O R S DAVID RANDALL contributions to the field of fish physiology. A frequent A prominent fish physiologist and a leading expert in symposium lecturer on fish physiology and other sub- respiratory and circulatory physiology, David Randall jects, most recently in Brazil, France, Germany, Italy, the collaborated with the late Roger Eckert on the earlier People's Republic of China, Russia, and the United editions of Animal Physiology and continues his con- States. He has worked with both the World Health Or- tribution in the fourth. A faculty member at the ganization and the United States Environmental Protec- University of British Columbia in Vancouver, Canada, tion Agency in developing ammonia criteria. Widely since 1963, and full professor since 1973, Randall was published as author and co-author in leading journals, appointed Associate Dean of Graduate Studies in 1990. Randall is co-editor of the noted series Fish Physiolog?, Elected a fellow of the Royal Society of Canada in 1981, (AcademicP ress), of which 15 volumes are in print. Vol- Randall has been both a Guggenheim and a Killam fel- ume 16, subtitled "Deep-sea Fish," will appear in 1997. low, and was awarded the prestigious Fry Medal for re- Along with his other duties, Randall co-teaches third search contributions to zoology by the Canadian Soci- year courses in vertebrate physiology and environmental ety of Zoology in 1993. In 1995, he received the Award physiology. His research interests concern the interac- of Excellence from the American Fisheries Society for tions between gas and ion exchange across fish gills. ......................................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . WARREN BURGGREN late them change over the course of development. Warren Burggren has taught in physiology for 23 years, Burggren has been actively involved in symposia, semi- and has been a professor of biological sciences at the Uni- nars, and formal extramural researchltraining activities versity of Nevada at Las Vegas since 1992. Courses he in many countries. A co-author of The Evolution of Air has taught at UNLV and at the University of Massachu- Breathing in Vertebrates (Cambridge University Press, setts, where he was Professor of Zoology from 1987 1981),B urggren has been a frequent contributor since through 1991, include Human Anatomy and Physiology, 1980 to edited collections of physiology, including Bioenergetics, Introductory Zoology, and Comparative Presser's Comparative Animal Physiology, Fourth Edi- Physiology. Burggren's research interest include develop- tion (Wiley-Liss, 1991). Burggren co-edited Environ- mental physiology, comparative animal physiology, and mental Physiology of the Amphibia (University of environmental and ecological physiology. In particular, Chicago Press, 1992), and more recently co-edited De- his research focuses on the ontogeny of respiratory and velopment of Cardiovascular Systems; Molecules to Or- cardiovascular systems, and how the systems that regu- ganisms (CambridgeU niversity Press, 1997). ........................................ ....................................... KATHLEEN FRENCH role as co-author of the current edition of Animal Physi- A neurobiologist at the University of California at San ology, along with a lifelong interest in the nervous systems Diego since 1985, Kathleen French has for 10 years of organisms from a broad range of phyla. As an Associ- taught upper division courses in embryology, mammalian ate Project Scienrist at UCSD, French's research focuses physiology for premedical students, and cellular neurobi- on the control of neuronal development, a topic that she ology. In addition, at UCSD, French participates in a has studied in various invertebrate species. Her current re- training program to instruct science teaching assistants in search concerns the cellular events that control differenti- the techniques and philosophy of teaching. She also serves ation of identified neurons in the medicinal leech, with an on the faculty of the Neuroscience and Behavior Course emphasis on the cellular physiology of embryonic neu- at the Marine Biological Laboratories in Woods Hole, rons and the effects of cell-cell contacts. She has been the Massachusetts, an intensive course designed primarily for author and co-author of numerous published research graduate students and post-doctoral fellows. French and review articles in journals including the Journal of . brings her expertise in-and love of-teaching to her 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 F": ,."%q : ':;i.?'.;.;,', , . ,.::,::; , $.,: :,: oi;, , ,:,, ,,,<:, $, ,,,:, ',:,, ;,; ?:{;:,, $ % <, +,;:,> ? . , , . , , , , , , , ,, , ,,*,, :-, : >:,'.!I,:,, , ,J:; , ,,<, ,,, > , ; , , ;! ,< ); , ' .:-.: i., . '.if , , '?:'ii, >,, 'a ,', ,,,,, .. 1 I C O N T E N T S i I I I Preface ix Structural Analys~so f Cells Acknowledgments xvii Cell Culture BIOCHEMICAL ANALYSIS PART I PRINCIPLES OF Measurlng Composit~onW: hat Is Present PHYSIOLOGY Measurlng Concentration: How Much IS Present EXPERIMENTS WITH ISOLATED ORGANS CHAPTER 1 STUDYING ANIMAL AND ORGAN SYSTEMS PHYSIOLOGY 3 OBSERVING AND MEASURING ANIMAL THE SUBDISCIPLINES OF ANIMAL PHYSIOLOGY 4 BEHAVIOR WHY STUDY ANIMAL PHYSIOLOGY? 4 The Power of Behav~oraEl xperiments Sc~ent~fCicu rlos~ty 4 Methods In Behav~oralR esearch Commerc~aYAgr~culturAalp pl~cat~ons 4 IMPORTANCE OF PHYSIOLOGICAL STATE Inslghts into Human Physiology 4 IN RESEARCH CENTRAL THEMES IN ANIMAL Summary PHYSIOLOGY 4 Revlew Quest~ons Structure-Funct~onR elat~onsh~ps 5 Suggested Readlngs Adaptation, Accl~mat~zat~aonnd, Accl~mat~on 5 CHAPTER 3 MOLECULES, ENERGY, Homeostas~s 7 AND BIOSYNTHESIS Feedback-Control Systems 8 Conform~atn~d Regulation 9 ORIGIN OF KEY BIOCHEMICAL MOLECULES LITERATURE OF PHYSIOLOGICAL SCIENCES 10 ATOMS, BONDS, AND MOLECULES SPOTLIGHT 1-1 THE CONCEPT OF FEEDBACK 12 THE SPECIAL ROLES OF H, 0, N, AND C ANIMAL EXPERIMENTATION IN PHYSIOLOGY 13 IN LIFE PROCESSES Summary 13 WATER: THE UNIQUE SOLVENT Rev~ewQ uest~ons 14 The Water Molecule Suggested Readlngs 14 Propert~eso f Water Water as a Solvent PROPERTIES OF SOLUTIONS CHAPTER 2 EXPERIMENTAL METHODS Concentration, Coll~gatlveP roperties, FOR EXPLORING PHYSIOLOGY and Activ~ty FORMULATING AND TESTING HYPOTHESES Ionizat~ono f Water The August Krogh Prlnc~ple Ac~dsa nd Bases Experimental Deslgn and Phys~olog~cLaelv el The B~olog~cIaml portance of pH MOLECULAR TECHNIQUES Henderson-Hasselbalch Equat~on Traclng Molecules w~thR adlolsotopes Buffer Systems Traclng Molecules wlth Monoclonal Antlbod~es Electrlc Current ~nA queous Solutions Genetic Englneerlng SPOTLIGHT 3-1 ELECTRICAL TERMINOLOGY CELLULAR TECHNIQUES AND CONVENTIONS Uses of M~croelectrodesa nd Mlcroplpettes Blndlng of Ions to Macromolecules

Description:

Eckert Animal Physiology is the essential text for courses exploring the structure, function and evolution of animals. The new Fourth Edition , revised, redesigned and updated, builds on Roger Eckert's earlier success in establishing this book as a classic. Comparing examples and experimental data a