Eckert Animal Physiology: Mechanisms and Adaptations (Fourth Edition)

840 Pages·1997·59.63 MB·English

by  David J. Randall

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

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

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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

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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