Animal Physiology, 3rd Ed

985 Pages·2012·53.374 MB·English

by  Richard W. Hill, Gordon A. Wyse, Margaret Anderson

Themes in the Study of Animal Physiology
Theme An Example of the Theme in Action See Pages
Listed are 15 overarching themes that reappear throughout the study of animal in importance. To help explain each theme, an illustrative example is presented
The Dynamic State of Body Constituents: Averaged over the course of an ordinary 24-h day, an adult person is like- 11–12
physiology. Some of the listed themes overlap with, or even encompass, oth- in the second column of the table. Further examples are on the pages listed in
ers; they are not intended to be mutually exclusive or, in all cases, equivalent the third column (italic listing elaborates the featured example). Great quantities of many of the key constituents ly to process more than 2 kg of adenosine triphosphate (ATP) each hour, 183–184
of the body are added and subtracted every day synthesizing that amount of ATP from adenosine diphosphate (ADP) 378
Theme An Example of the Theme in Action See Pages icno nmstaintuye anntsim ofa ltsh ue nbdoedry m—afnary fcroonmd ibteioinngs. sTtahtuics— thaer e asinzde, twheit Ah ToPn,l yth ae spheorrsto dne—ladyu, rbirnega keiancgh iht obuarc—k dwoiwll nu steo aAbDouPt. 2T0o lsityenrtsh oef- 699
continuously in a dynamic state of flux. This is true oxygen (O2) that he or she takes up from the atmosphere. During a 24-h 7 4 (3F–ig7.4 248 .21)
The Study of Function: When physiologists study muscle, one of their goals is to understand how 114 (Fig. 5.12) even though additions and subtractions are often day, the oxygen used will combine with almost 100 g (a fifth of a pound)
Animal physiology is the study of how animals the proteins in muscle cells are able to develop mechanical forces, which 527 (Fig. 20.5) relatively balanced, resulting in relatively constant of hydrogen atoms that have been removed from food molecules, forming
function. That is, it is the study of how their cells are employed in locomotion, heart contraction, or other activities. 606 (Fig. 23.22) concentrations (a phenomenon termed homeostasis). about 800 milliliters of water. This water is added to the body fluids.
and organs operate. 687 (Fig. 26.9)
Multiple Forms of Key Molecules: The cell membranes of all animals are composed principally of lipid mol- 34 (Fig. 2.3)
Animals have often evolved multiple molecular ecules. Physiologists have found, however, that the membranes of all ani- 242–243
Integration of the Sciences: To understand how animals employ odors to orient their movements, 6 (Fig. 1.2) forms (sometimes called isoforms) of particular mals are not composed of chemically identical lipid molecules. Instead, (Fig. 10.19)
Physiologists often find that they must integrate physiologists study the chemical structural differences between molecules 60 (Fig. 2.28) proteins or other sorts of molecules. Physiologists multiple molecular forms of lipids are employed by different animals 537
knowledge of mathematics, chemistry, or physics that attract or repel, and they mathematically describe the physics of how 165 (Fig. 7.3) hypothesize that when two species or two tissues living under different circumstances. Cold-water fish species, for instance, 620 (Fig. 24.2)
with knowledge of biology to answer important winds or water currents transport odor molecules from odor sources to 377 exhibit different molecular forms of a molecule, the construct their cell membranes using molecular forms of lipids that are 640 (Fig. 24.20)
questions. Physiology is one of the most integrative the olfactory organs of animals. forms are often specialized to function in the spe- less likely to harden at low temperatures than the molecular forms syn-
650 (Fig. 25.2)
branches of biology. cific settings in which the animals live or the tissues thesized by warm-water species.
function.
Emphasis on Quantitative Methods: Starting in ancient Roman times, people thought that the dromedary camel 209 (Box 9.1)
Physiologists quantify the properties of animals as could carry enough water in its rumen to explain its unusual ability to live 212 (Fig. 9.7) Phenotypic Plasticity: Animals that eat only occasionally, such as pythons, often alternate 15 (Fig. 1.5)
carefully as possible as they seek to test hypotheses without drinking. When physiologists quantified the amount of water in 234–235 An individual animal is often able to change its between two intestinal phenotypes. When they have not had a meal for 79 (Table 3.1)
or make predictions. the camel rumen rather than just speaking qualitatively about it, however, (Fig. 10.8) phenotype in response to changes in the particular weeks, their intestinal tract is physically small, and it has poorly devel- 90–92
tohldey i dfoeua ntod mthaakt et hseernes ew. aNse ngoatt innega trhlye eonldo uidgeha w haetlepre idn ltehaed rtuom uennd eforsrt tahned - 799–800 ctiicruculamr setnavnicreosn umnednetr). wThhiics ha biti liist yli voifn agn (ien.gd.i,v iitds upaalr - oopf ethde m inotleesctuinlaarl tmraeccth eannliasrmges fgorre aatblyso, rabnidn gth feo oindt.e Astfitnearl at rmaceta el,x tphree stsisessu es 157 (Box 6.2)
ing that camels do not store water to a greater degree than other mam- animal to adopt two or more phenotypes despite well-developed absorption mechanisms. 264 (Fig. 10.40)
555 (Fig. 21.7)
mals. Instead, they have excellent abilities to conserve water and endure having a fixed genotype is termed phenotypic
dehydration. plasticity.
Interdependency of Function and Form: The kidney tubules of mammals not only produce the most concentrated 140 (Fig. 6.13)
The Tandem Goal of Understanding A number of animals, including fireflies and certain fish, produce light. 5–9
Mechanism and Adaptation: When physiologists study light production in such animals, they try to 248–249 The function of a biological system typically cannot urine observed in vertebrates but also differ from other vertebrate kidney 259 (Fig. 10.35)
When physiologists study a process, they typically learn both how the animals make light and why they make it. (Fig. 10.26) be understood without knowledge of its structure, tubules in that they all have distinctive hairpin shapes. Physiologists have 370 (Fig. 14.10)
emphasize a two-part goal: They try to understand 455–456 and vice versa. sdheopwennd tsh oant tthhee fhuanicrtpioinna slt arubciltiutyre ,t ow phriochd ugcuei dheigs hthlye cuorninceen (tarsa tiet dis ubreiinneg 649 (Fig. 25.1)
both the mechanism involved (i.e., how the process 735 (Fig. 28.12) formed) to flow first in one direction and then in the opposite direction. 768 (Fig. 29.12)
is executed) and the potential adaptive significance of
760 (Fig. 29.5)
the process (i.e., how, if at all, it enhances evolu-
Applicability of the Laws of Chemistry and Heat transfer through air follows different physical laws when the air is 230
tionary fitness).
Physics: still rather than moving; heat tends to move much more slowly through 493 (Fig. 18.8)
still air than moving air. Animals cannot change such laws of physics.
Animals must adhere to the laws of chemistry and 576 (Box 22.2)
The Comparative Method: When physiologists compare animals as distantly related as mammals and 26 (Fig. 1.18) They sometimes can affect which law applies to them, however, as when
physics. Sometimes chemistry and physics act as 694–695
To understand the adaptive significance of animal insects, they find that desert species tend consistently to have great abili- 114 (Fig. 5.13) constraints, but sometimes animals gain advantages the ancestors of mammals evolved fur. The hairs of a furred mammal
features, physiologists make extensive use of the ties to concentrate their urine. Desert species of mammals are typically 675 (Box 25.3) by evolving systems that capitalize on particular keep the layer of air next to the body relatively motionless. Heat transfer
comparative method, which is the examination of how able to make urine of higher concentration than nondesert mammals, and 688 (Box 26.1) chemical or physical principles. through that air is therefore slow, helping mammals retain internal heat
particular functions are carried out by related and desert insects are similarly superior to nondesert insects. These compara- when living in cold environments.
unrelated species living in similar and dissimilar tive observations provide evidence that the ability to make concentrated 743 (Fig. 28.20)
environments. urine is an advantage—favored by natural selection—in deserts. The Interdependency of Levels of When your physician strikes a tendon near your knee with a mallet, your 6 (Fig. 1.2)
Organization: leg straightens. For this response, electrical signals must travel along 116 (Box 5.2)
nerve cells to the spinal cord and back. The rate of travel depends in part
Phylogenetic Reconstruction: Although body temperature is the same as water temperature in most 27 An animal’s overall functional properties depend on 199 (Fig. 8.12)
on the molecular properties of ion-transporting proteins in the cell mem-
To understand the evolution of physiological species of fish, regardless of how big they are, certain species of fish 53 (Fig. 2.21) how its tissues and organs function, and the function branes of the nerve cells. It also depends in part on key cellular properties, 306–307
properties—and thus gain perspective on the evo- maintain elevated temperatures in some of their tissues. From phylo- 71 (Fig. 3.5) of its tissues and organs depends on how its cells and such as the spacing between the sections of each nerve cell membrane 508 (Fig. 19.4)
lutionary significance of modern-day properties— genetic reconstructions, physiologists have found that the warm-tissue 270 (Fig. 10.47) molecular systems function. All these levels of organi- that are fully exposed to the fluids bathing the cell. Molecular and cellular
pwhhyicshio lgoegniesttisc e omr poltohye rp ihnyfloogrmenaettiioc nre oconn mstruulctitpiolne ss, pine- cthoenrdeiftoioren, ethvaotl vtoedd aoyn’s a fti sleha wsti tfhou wr ainrmde tpisesnudeesn dt oo cncoats iaolnl ssi. mWpel yk ninohwe,r it 732 (Box 28.4) zisa tthioant parreo pinetretiredse apte onndee nlet.v Aeln o ifm orpgoarntaiznat tcioonro ollfateryn pForor pinersttiaensc oef, tthheeyse d seotretrsm dientee rtmhei nleen tghteh o ovfe rtaimll ep rtohpate rptaiesss eosf btheetw pereonc etshse.
cies is used to reconstruct the paths of evolution. the condition from a single common ancestor. cannot be fully understood without exploring other moment the mallet strikes and the moment your leg muscles contract.
levels of organization.
The Centrality of the Environment: Many specialists in high-altitude physiology argue that when lowland 53 (Fig. 2.22)
The Crucial Importance of Control Although sheep and reindeer are born at cold times of year, newborns 50 (Fig. 2.19)
The specific environments in which animals have people travel to high altitudes, some of their typical responses are more 67 Mechanisms: receive no heat from their parents and must keep warm on their own or 252 (Box 10.2)
evolved and live must be considered for the func- harmful than helpful. These specialists emphasize that the human spe- 264 (Fig. 10.41) die. They possess a process for rapid heat production. Proper control of
tional properties of the animals to make sense. cnioe sr edaisdo nno tto e pvroelsvuem ien t hhaigt ha-lal ltthiteu hdue menavni rroenspmoennstess. Atoc scuocrdh ienngvlyir, othnemree nists 639 (Box 24.5) Iinng a, dmdoitvioinng t,o a mnde cchaarrnyisinmgs o fuotr orethperro douvceirnt gfu, bnrcetiaotnhs-, this process requires that it be activated at birth, but not before birth when 248830 ((FFiigg.. 1117..81)5)
it would tend needlessly to exhaust fetal energy supplies. The control
would be beneficial. animals require control mechanisms that orchestrate
mechanism has two key properties: It activates heat production when
the other mechanisms. The control mechanisms—so
neural thermal sensors detect cold, but its capacity to activate heat produc-
diverse that they include controls of gene expres-
Body Size: The metabolic rate per gram of body weight is usually higher in small- 17 (Fig. 1.9) sion as well as those exerted by the nervous tion is turned off by chemical factors secreted by the placenta. The control
The physiological properties of related animal spe- bodied species than in related large-bodied ones. Because of this relation, 173 (Fig. 7.6) and endocrine systems—determine the relations mechanism remains in a turned-off state until a newborn is separated from
cies typically scale in mathematically consistent whenever two species of mammals of different body sizes—like mice and 285 (Fig. 11.9) between inputs and outputs in physiological sys- the placenta at birth. The cold environment is then able to stimulate rapid
ways with their body sizes. These relations are horses—are compared, the smaller species typically needs more food per 741 (Fig. 28.18) tems. They thereby crucially affect the functional heat production.
often nonproportional and thus termed allometric. gram of body weight than the larger one. properties of animals.
Themes in the Study of Animal Physiology
Theme An Example of the Theme in Action See Pages
Listed are 15 overarching themes that reappear throughout the study of animal in importance. To help explain each theme, an illustrative example is presented
The Dynamic State of Body Constituents: Averaged over the course of an ordinary 24-h day, an adult person is like- 11–12
physiology. Some of the listed themes overlap with, or even encompass, oth- in the second column of the table. Further examples are on the pages listed in
ers; they are not intended to be mutually exclusive or, in all cases, equivalent the third column (italic listing elaborates the featured example). Great quantities of many of the key constituents ly to process more than 2 kg of adenosine triphosphate (ATP) each hour, 183–184
of the body are added and subtracted every day synthesizing that amount of ATP from adenosine diphosphate (ADP) 378
Theme An Example of the Theme in Action See Pages icno nmstaintuye anntsim ofa ltsh ue nbdoedry m—afnary fcroonmd ibteioinngs. sTtahtuics— thaer e asinzde, twheit Ah ToPn,l yth ae spheorrsto dne—ladyu, rbirnega keiancgh iht obuarc—k dwoiwll nu steo aAbDouPt. 2T0o lsityenrtsh oef- 699
continuously in a dynamic state of flux. This is true oxygen (O2) that he or she takes up from the atmosphere. During a 24-h 7 4 (3F–ig7.4 248 .21)
The Study of Function: When physiologists study muscle, one of their goals is to understand how 114 (Fig. 5.12) even though additions and subtractions are often day, the oxygen used will combine with almost 100 g (a fifth of a pound)
Animal physiology is the study of how animals the proteins in muscle cells are able to develop mechanical forces, which 527 (Fig. 20.5) relatively balanced, resulting in relatively constant of hydrogen atoms that have been removed from food molecules, forming
function. That is, it is the study of how their cells are employed in locomotion, heart contraction, or other activities. 606 (Fig. 23.22) concentrations (a phenomenon termed homeostasis). about 800 milliliters of water. This water is added to the body fluids.
and organs operate. 687 (Fig. 26.9)
Multiple Forms of Key Molecules: The cell membranes of all animals are composed principally of lipid mol- 34 (Fig. 2.3)
Animals have often evolved multiple molecular ecules. Physiologists have found, however, that the membranes of all ani- 242–243
Integration of the Sciences: To understand how animals employ odors to orient their movements, 6 (Fig. 1.2) forms (sometimes called isoforms) of particular mals are not composed of chemically identical lipid molecules. Instead, (Fig. 10.19)
Physiologists often find that they must integrate physiologists study the chemical structural differences between molecules 60 (Fig. 2.28) proteins or other sorts of molecules. Physiologists multiple molecular forms of lipids are employed by different animals 537
knowledge of mathematics, chemistry, or physics that attract or repel, and they mathematically describe the physics of how 165 (Fig. 7.3) hypothesize that when two species or two tissues living under different circumstances. Cold-water fish species, for instance, 620 (Fig. 24.2)
with knowledge of biology to answer important winds or water currents transport odor molecules from odor sources to 377 exhibit different molecular forms of a molecule, the construct their cell membranes using molecular forms of lipids that are 640 (Fig. 24.20)
questions. Physiology is one of the most integrative the olfactory organs of animals. forms are often specialized to function in the spe- less likely to harden at low temperatures than the molecular forms syn-
650 (Fig. 25.2)
branches of biology. cific settings in which the animals live or the tissues thesized by warm-water species.
function.
Emphasis on Quantitative Methods: Starting in ancient Roman times, people thought that the dromedary camel 209 (Box 9.1)
Physiologists quantify the properties of animals as could carry enough water in its rumen to explain its unusual ability to live 212 (Fig. 9.7) Phenotypic Plasticity: Animals that eat only occasionally, such as pythons, often alternate 15 (Fig. 1.5)
carefully as possible as they seek to test hypotheses without drinking. When physiologists quantified the amount of water in 234–235 An individual animal is often able to change its between two intestinal phenotypes. When they have not had a meal for 79 (Table 3.1)
or make predictions. the camel rumen rather than just speaking qualitatively about it, however, (Fig. 10.8) phenotype in response to changes in the particular weeks, their intestinal tract is physically small, and it has poorly devel- 90–92
tohldey i dfoeua ntod mthaakt et hseernes ew. aNse ngoatt innega trhlye eonldo uidgeha w haetlepre idn ltehaed rtuom uennd eforsrt tahned - 799–800 ctiicruculamr setnavnicreosn umnednetr). wThhiics ha biti liist yli voifn agn (ien.gd.i,v iitds upaalr - oopf ethde m inotleesctuinlaarl tmraeccth eannliasrmges fgorre aatblyso, rabnidn gth feo oindt.e Astfitnearl at rmaceta el,x tphree stsisessu es 157 (Box 6.2)
ing that camels do not store water to a greater degree than other mam- animal to adopt two or more phenotypes despite well-developed absorption mechanisms. 264 (Fig. 10.40)
555 (Fig. 21.7)
mals. Instead, they have excellent abilities to conserve water and endure having a fixed genotype is termed phenotypic
dehydration. plasticity.
Interdependency of Function and Form: The kidney tubules of mammals not only produce the most concentrated 140 (Fig. 6.13)
The Tandem Goal of Understanding A number of animals, including fireflies and certain fish, produce light. 5–9
Mechanism and Adaptation: When physiologists study light production in such animals, they try to 248–249 The function of a biological system typically cannot urine observed in vertebrates but also differ from other vertebrate kidney 259 (Fig. 10.35)
When physiologists study a process, they typically learn both how the animals make light and why they make it. (Fig. 10.26) be understood without knowledge of its structure, tubules in that they all have distinctive hairpin shapes. Physiologists have 370 (Fig. 14.10)
emphasize a two-part goal: They try to understand 455–456 and vice versa. sdheopwennd tsh oant tthhee fhuanicrtpioinna slt arubciltiutyre ,t ow phriochd ugcuei dheigs hthlye cuorninceen (tarsa tiet dis ubreiinneg 649 (Fig. 25.1)
both the mechanism involved (i.e., how the process 735 (Fig. 28.12) formed) to flow first in one direction and then in the opposite direction. 768 (Fig. 29.12)
is executed) and the potential adaptive significance of
760 (Fig. 29.5)
the process (i.e., how, if at all, it enhances evolu-
Applicability of the Laws of Chemistry and Heat transfer through air follows different physical laws when the air is 230
tionary fitness).
Physics: still rather than moving; heat tends to move much more slowly through 493 (Fig. 18.8)
still air than moving air. Animals cannot change such laws of physics.
Animals must adhere to the laws of chemistry and 576 (Box 22.2)
The Comparative Method: When physiologists compare animals as distantly related as mammals and 26 (Fig. 1.18) They sometimes can affect which law applies to them, however, as when
physics. Sometimes chemistry and physics act as 694–695
To understand the adaptive significance of animal insects, they find that desert species tend consistently to have great abili- 114 (Fig. 5.13) constraints, but sometimes animals gain advantages the ancestors of mammals evolved fur. The hairs of a furred mammal
features, physiologists make extensive use of the ties to concentrate their urine. Desert species of mammals are typically 675 (Box 25.3) by evolving systems that capitalize on particular keep the layer of air next to the body relatively motionless. Heat transfer
comparative method, which is the examination of how able to make urine of higher concentration than nondesert mammals, and 688 (Box 26.1) chemical or physical principles. through that air is therefore slow, helping mammals retain internal heat
particular functions are carried out by related and desert insects are similarly superior to nondesert insects. These compara- when living in cold environments.
unrelated species living in similar and dissimilar tive observations provide evidence that the ability to make concentrated 743 (Fig. 28.20)
environments. urine is an advantage—favored by natural selection—in deserts. The Interdependency of Levels of When your physician strikes a tendon near your knee with a mallet, your 6 (Fig. 1.2)
Organization: leg straightens. For this response, electrical signals must travel along 116 (Box 5.2)
nerve cells to the spinal cord and back. The rate of travel depends in part
Phylogenetic Reconstruction: Although body temperature is the same as water temperature in most 27 An animal’s overall functional properties depend on 199 (Fig. 8.12)
on the molecular properties of ion-transporting proteins in the cell mem-
To understand the evolution of physiological species of fish, regardless of how big they are, certain species of fish 53 (Fig. 2.21) how its tissues and organs function, and the function branes of the nerve cells. It also depends in part on key cellular properties, 306–307
properties—and thus gain perspective on the evo- maintain elevated temperatures in some of their tissues. From phylo- 71 (Fig. 3.5) of its tissues and organs depends on how its cells and such as the spacing between the sections of each nerve cell membrane 508 (Fig. 19.4)
lutionary significance of modern-day properties— genetic reconstructions, physiologists have found that the warm-tissue 270 (Fig. 10.47) molecular systems function. All these levels of organi- that are fully exposed to the fluids bathing the cell. Molecular and cellular
pwhhyicshio lgoegniesttisc e omr poltohye rp ihnyfloogrmenaettiioc nre oconn mstruulctitpiolne ss, pine- cthoenrdeiftoioren, ethvaotl vtoedd aoyn’s a fti sleha wsti tfhou wr ainrmde tpisesnudeesn dt oo cncoats iaolnl ssi. mWpel yk ninohwe,r it 732 (Box 28.4) zisa tthioant parreo pinetretiredse apte onndee nlet.v Aeln o ifm orpgoarntaiznat tcioonro ollfateryn pForor pinersttiaensc oef, tthheeyse d seotretrsm dientee rtmhei nleen tghteh o ovfe rtaimll ep rtohpate rptaiesss eosf btheetw pereonc etshse.
cies is used to reconstruct the paths of evolution. the condition from a single common ancestor. cannot be fully understood without exploring other moment the mallet strikes and the moment your leg muscles contract.
levels of organization.
The Centrality of the Environment: Many specialists in high-altitude physiology argue that when lowland 53 (Fig. 2.22)
The Crucial Importance of Control Although sheep and reindeer are born at cold times of year, newborns 50 (Fig. 2.19)
The specific environments in which animals have people travel to high altitudes, some of their typical responses are more 67 Mechanisms: receive no heat from their parents and must keep warm on their own or 252 (Box 10.2)
evolved and live must be considered for the func- harmful than helpful. These specialists emphasize that the human spe- 264 (Fig. 10.41) die. They possess a process for rapid heat production. Proper control of
tional properties of the animals to make sense. cnioe sr edaisdo nno tto e pvroelsvuem ien t hhaigt ha-lal ltthiteu hdue menavni rroenspmoennstess. Atoc scuocrdh ienngvlyir, othnemree nists 639 (Box 24.5) Iinng a, dmdoitvioinng t,o a mnde cchaarrnyisinmgs o fuotr orethperro douvceirnt gfu, bnrcetiaotnhs-, this process requires that it be activated at birth, but not before birth when 248830 ((FFiigg.. 1117..81)5)
it would tend needlessly to exhaust fetal energy supplies. The control
would be beneficial. animals require control mechanisms that orchestrate
mechanism has two key properties: It activates heat production when
the other mechanisms. The control mechanisms—so
neural thermal sensors detect cold, but its capacity to activate heat produc-
diverse that they include controls of gene expres-
Body Size: The metabolic rate per gram of body weight is usually higher in small- 17 (Fig. 1.9) sion as well as those exerted by the nervous tion is turned off by chemical factors secreted by the placenta. The control
The physiological properties of related animal spe- bodied species than in related large-bodied ones. Because of this relation, 173 (Fig. 7.6) and endocrine systems—determine the relations mechanism remains in a turned-off state until a newborn is separated from
cies typically scale in mathematically consistent whenever two species of mammals of different body sizes—like mice and 285 (Fig. 11.9) between inputs and outputs in physiological sys- the placenta at birth. The cold environment is then able to stimulate rapid
ways with their body sizes. These relations are horses—are compared, the smaller species typically needs more food per 741 (Fig. 28.18) tems. They thereby crucially affect the functional heat production.
often nonproportional and thus termed allometric. gram of body weight than the larger one. properties of animals.
00_Hill3e_FM.indd ii 3/9/12 2:11 PM
ANIMAL PHYSIOLOGY
THIRD EDITION
00_Hill3e_FM.indd i 3/9/12 2:11 PM

ANIMAL
PHYSIOLOGY
THIRD EDITION
Richard W. Hill
Michigan State University
Gordon A. Wyse
University of Massachusetts, Amherst
Margaret Anderson
Smith College
Sinauer Associates, Inc. Publishers • Sunderland, Massachusetts
00_Hill3e_FM.indd iii 3/13/12 2:26 PM
About the Cover
One of the central themes of this book is the intimate relationship between animals
and their environments. The gemsbok oryx (Oryx gazella) provides an iconic example.
Gemsboks succeed in one of Earth’s most demanding settings—the hot, dry deserts of
Africa—because of their evolution of a variety of specialized behavioral, morphological,
and physiological attributes. Among the truly wild large mammals, the three species of
oryxes that are scientifically well known probably represent the pinnacle of evolution
in their ability to survive in such deserts. Oryxes are discussed in depth in Chapter 30.
Animal Physiology, Third Edition
Copyright © 2012. All rights reserved.
This book may not be reproduced in whole or in part without permission from the publisher.
Address editorial correspondence and orders to:
Sinauer Associates, 23 Plumtree Road, Sunderland, MA 01375 U.S.A.
FAX: [phone]
Email: [email protected]
Internet: www.sinauer.com
Library of Congress Cataloging-in-Publication Data
Hill, Richard W.
Animal physiology / Richard W. Hill, Gordon A. Wyse, Margaret Anderson. -- 3rd ed.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-87893-559-8 (casebound)
1. Physiology, Comparative. I. Wyse, Gordon A. II. Anderson, Margaret, 1941- III. Title.
QP33.H54 2012
571.8’1--dc23
2012005574
Printed in U.S.A.
7 6 5 4 3 2 1
00_Hill3e_FM.indd iv 3/9/12 2:11 PM
To Sue, Dave, and Chrissie, from RWH
To Mary, from GAW
To Anita and Andy, from MA
00_Hill3e_FM.indd v 3/9/12 2:11 PM
Preface
Thomas Kuhn wrote that a textbook is principally a means of com- the lives of animals in their natural habitats. Second, we devote five
municating to students the paradigms of their time. We have had entire chapters (our “At Work” chapters) to in-depth explorations
three principal goals in preparing the content of this book. One, of how physiologists do their work; in these chapters we break out
in accord with Kuhn’s dictum, has been to articulate the central of the usual textbook mold to discuss exciting topics—such as the
paradigms of contemporary animal physiology. A second content diving physiology of marine mammals—with emphasis on experi-
goal has been to provide our readers with a source of both lucid ments, theory maturation, integration of physiological systems, and
explanations of physiological concepts and accurate information prospects for future research. Third, we include many photographs
about physiological systems. Our third content goal has been to and drawings of animals throughout the book to remind readers of
draw attention to the cutting edges of physiological science, the the animals we discuss. Fourth, entirely new to this edition, we have
places where the onward progress of research is challenging old started a program of inviting specialists to contribute expert Guest
paradigms and potentially creating footholds for new ones. Boxes on emerging topics that expand the book’s subject content.
We have also had goals for presentation. Most visibly, we have With our aspirations being as numerous as we have described,
combined our words with an ambitious, informative art program. we have put a great deal of effort into balancing competing demands
More fundamentally, we have strived to take advantage of all the for space. The product is a complete physiology textbook that in
assets of traditional bookmaking to achieve a book that—through one volume will meet the requirements of a diversity of one- or
constant integration of the full suite of pedgogically relevant ele- two-semester courses in animal function. Our intended audience
ments—is a first-rate learning tool. Many sorts of professionals is sophomores through beginning graduate students. To make
have important contributions to make for a book to be excellent. the book accessible to as wide an audience as possible, we have
Thus many sorts of professionals have traditionally found personal included both a glossary of nearly 1200 terms and 11 appendices
fulfillment by engaging in the cooperative, synergistic production on important background concepts.
of books. The authors listed on the cover are just the tip of the Our approach to the writing has been to work from the original
iceberg. A book’s art program depends on scientific illustrators. scientific literature and obtain extensive peer review. Another aspect
Coordination between the art and the text—a key to the success of our approach is that we have opted for the pedagogical consis-
of any textbook—depends on the editorial expertise of the book’s tency of a book written by just three principal authors. Margaret
editor. An attractive science text needs to be designed and physi- Anderson wrote Chapters 16, 20, and 21, and Gordon Wyse wrote
cally executed by talented people who combine scientific acumen Chapters 12–15, 18, and 19. Richard Hill wrote Chapters 1–11, 17,
with artistic sensibility. To the degree that the presentation of and 22–30. David S. Garbe, Scott A. Huettel, Matthew S. Kayser,
the material in this book achieves success, the reason is that it is Kenneth J. Lohmann, and Margaret McFall-Ngai wrote Guest
the creative product of a team of at least a dozen people playing Boxes. Matthew S. Kayser and Gordon Fain assisted with topic
diverse, mutually reinforcing roles. One of our goals has been to development in certain parts of the principal text.
take advantage of this time-proven model to provide students with
a superior text. NEW TO THIS EDITION: As in other editions, our two central goals
In these pages, we consistently and deliberately address animal for this edition were to update content and enhance pedagogical
physiology as a discipline integrated with other disciplines in biol- effectiveness. To these ends, we have reconsidered every sentence
ogy—especially genetics, molecular biology, evolutionary biology, and every element of the art program. Of the 690 figures and tables
and ecology. We also consistently emphasize the roles of physiology in this edition, over 60 are either new or greatly enhanced. Chapter
throughout the life cycle of an animal by discussing physiologi- 14, on sensory processes, has been entirely rewritten. We have also
cal development and by examining animal function during such added a new chapter (Chapter 4) on physiological development and
important life-cycle processes as exercise, long-distance migration, epigenetics. Other chapters that have received exceptional attention
seasonal rhythms, and accommodation to severe conditions (we are: Chapter 5 (transport of solutes and water), Chapter 8 (aerobic
generally omit pathology and parasitism, however). Although we give and anaerobic metabolism), Chapter 9 (activity energetics), and
particular attention to mammals, we make a point of recognizing Chapter 29 (kidney physiology, edited throughout to emphasize
the other vertebrate groups and at least the arthropods and molluscs plasma regulation). The book now includes Guest Boxes on functional
among invertebrates. We address all levels of organization that are magnetic resonance imaging, magnetoreception, optogenetics,
germane, from the genome to the ecological context. sleep, symbiosis in the bobtail squid-Vibrio system, and synaptic
We want to mention four specific strategies we have adopted to development. Treatment of topics in global warming has been
add interest and breadth to the book. First, we start every chapter tripled. Treatment of altitude physiology in Chapters 23 and 24 is
with a vivid example of the application of the chapter’s material to entirely revised. The index is new and upgraded. A limited list of
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