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background image   Themes in the Study of Animal Physiology Theme An Example of the Theme in Action See Pages The Study of Function:  Animal physiology is the study of how animals  function. That is, it is the study of how their cells  and organs operate. When physiologists study muscle, one of their goals is to understand how  the proteins in muscle cells are able to develop mechanical forces, which  are employed in locomotion, heart contraction, or other activities.  114 (Fig. 5.12)
527 (Fig. 20.5)
606 (Fig. 23.22)
687 (Fig. 26.9)
Integration of the Sciences:  Physiologists often find that they must integrate  knowledge of mathematics, chemistry, or physics  with knowledge of biology to answer important  questions. Physiology is one of the most integrative  branches of biology. To understand how animals employ odors to orient their movements,  physiologists study the chemical structural differences between molecules  that attract or repel, and they mathematically describe the physics of how  winds or water currents transport odor molecules from odor sources to  the olfactory organs of animals. 6 (Fig. 1.2)
60 (Fig. 2.28)
165 (Fig. 7.3)
377
650 (Fig. 25.2)
Emphasis on Quantitative Methods:  Physiologists quantify the properties of animals as  carefully as possible as they seek to test hypotheses  or make predictions. Starting in ancient Roman times, people thought that the dromedary camel  could carry enough water in its rumen to explain its unusual ability to live  without drinking. When physiologists quantified the amount of water in  the camel rumen rather than just speaking qualitatively about it, however,  they found that there was not nearly enough water in the rumen for the  old idea to make sense. Negating the old idea helped lead to understand- ing that camels do not store water to a greater degree than other mam- mals. Instead, they have excellent abilities to conserve water and endure  dehydration. 209 (Box 9.1)
212 (Fig. 9.7)
234–235  
   (Fig. 10.8)
799–800
The Tandem Goal of Understanding  Mechanism and Adaptation:  When physiologists study a process, they typically  emphasize a two-part goal: They try to understand  both the mechanism involved (i.e., how the process  is executed) and the potential adaptive significance of  the process (i.e., how, if at all, it enhances evolu- tionary fitness). A number of animals, including fireflies and certain fish, produce light.  When physiologists study light production in such animals, they try to  learn both how the animals make light and why they make it. 5–9
248–249  
   (Fig. 10.26)
455–456
735 (Fig. 28.12)
760 (Fig. 29.5)
The Comparative Method:  To understand the adaptive significance of animal  features, physiologists make extensive use of the  comparative method , which is the examination of how  particular functions are carried out by related and  unrelated species living in similar and dissimilar  environments. When physiologists compare animals as distantly related as mammals and  insects, they find that desert species tend consistently to have great abili- ties to concentrate their urine. Desert species of mammals are typically  able to make urine of higher concentration than nondesert mammals, and  desert insects are similarly superior to nondesert insects. These compara- tive observations provide evidence that the ability to make concentrated  urine is an advantage—favored by natural selection—in deserts. 26 (Fig. 1.18)
114 (Fig. 5.13)
675 (Box 25.3)
688 (Box 26.1)
743 (Fig. 28.20)
Phylogenetic Reconstruction: To understand the evolution of physiological  properties—and thus gain perspective on the evo- lutionary significance of modern-day properties— physiologists employ phylogenetic reconstructions, in  which genetic or other information on multiple spe- cies is used to reconstruct the paths of evolution. Although body temperature is the same as water temperature in most  species of fish, regardless of how big they are, certain species of fish  maintain elevated temperatures in some of their tissues. From phylo- genetic reconstructions, physiologists have found that the warm-tissue   condition evolved on at least four independent occasions. We know,  therefore, that today’s fish with warm tissues do not all simply inherit   the condition from a single common ancestor. 27
53 (Fig. 2.21)
71 (Fig. 3.5)
270 (Fig. 10.47)
732 (Box 28.4)
The Centrality of the Environment:  The specific environments in which animals have  evolved and live must be considered for the func- tional properties of the animals to make sense. Many specialists in high-altitude physiology argue that when lowland  people travel to high altitudes, some of their typical responses are more  harmful than helpful. These specialists emphasize that the human spe- cies did not evolve in high-altitude environments. Accordingly, there is  no reason to presume that all the human responses to such environments  would be beneficial. 53 (Fig. 2.22)
67
264 (Fig. 10.41)
639 (Box 24.5)
Body Size:  The physiological properties of related animal spe- cies typically scale in mathematically consistent  ways with their body sizes. These relations are  often nonproportional and thus termed allometric. The metabolic rate per gram of body weight is usually higher in small- bodied species than in related large-bodied ones. Because of this relation,  whenever two species of mammals of different body sizes—like mice and  horses—are compared, the smaller species typically needs more food per  gram of body weight than the larger one. 17 (Fig. 1.9)
173 (Fig. 7.6)
285 (Fig. 11.9)
741 (Fig. 28.18)
Listed are 15 overarching themes that reappear throughout the study of animal  physiology. Some of the listed themes overlap with, or even encompass, oth- ers; they are not intended to be mutually exclusive or, in all cases, equivalent  in importance. To help explain each theme, an illustrative example is presented  in the second column of the table. Further examples are on the pages listed in  the third column (italic listing elaborates the featured example). Theme An Example of the Theme in Action See Pages The Dynamic State of Body Constituents:  Great quantities of many of the key constituents  of the body are added and subtracted every day  in many animals under many conditions. Thus the  constituents of the body—far from being static—are  continuously in a dynamic state of flux. This is true  even though additions and subtractions are often  relatively balanced, resulting in relatively constant  concentrations (a phenomenon termed homeostasis). Averaged over the course of an ordinary 24-h day, an adult person is like- ly to process more than 2 kg of adenosine triphosphate (ATP) each hour,  synthesizing that amount of ATP from adenosine diphosphate (ADP)  and, with only a short delay, breaking it back down to ADP. To synthe- size the ATP, the person—during each hour—will use about 20 liters of  oxygen (O 2 ) that he or she takes up from the atmosphere. During a 24-h  day, the oxygen used will combine with almost 100 g (a fifth of a pound)  of hydrogen atoms that have been removed from food molecules, forming  about 800 milliliters of water. This water is added to the body fluids. 11–12
183–184
378
699
743–744  
   (Fig. 28.21) Multiple Forms of Key Molecules:  Animals have often evolved multiple molecular  forms (sometimes called isoforms) of particular  proteins or other sorts of molecules.  Physiologists  hypothesize that when two species or two tissues  exhibit different molecular forms of a molecule, the  forms are often specialized to function in the spe- cific settings in which the animals live or the tissues  function. The cell membranes of all animals are composed principally of lipid mol- ecules. Physiologists have found, however, that the membranes of all ani- mals are not composed of chemically identical lipid molecules. Instead,  multiple molecular forms of lipids are employed by different animals  living under different circumstances. Cold-water fish species, for instance,  construct their cell membranes using molecular forms of lipids that are  less likely to harden at low temperatures than the molecular forms syn- thesized by warm-water species. 34 (Fig. 2.3)
242–243  
   (Fig. 10.19)
537
620 (Fig. 24.2)
640 (Fig. 24.20)
Phenotypic Plasticity:  An individual animal is often able to change its  phenotype in response to changes in the particular  circumstances under which it is living (e.g., its par- ticular environment). This ability of an individual  animal to adopt two or more phenotypes despite  having a fixed genotype is termed phenotypic  plasticity . Animals that eat only occasionally, such as pythons, often alternate  between two intestinal phenotypes. When they have not had a meal for  weeks, their intestinal tract is physically small, and it has poorly devel- oped molecular mechanisms for absorbing food. After a meal, the tissues  of the intestinal tract enlarge greatly, and the intestinal tract expresses  well-developed absorption mechanisms. 15 (Fig. 1.5)
79 (Table 3.1)
90–92
157 (Box 6.2)
264 (Fig. 10.40)
555 (Fig. 21.7)
Interdependency of Function and Form: The function of a biological system typically cannot  be understood without knowledge of its structure and vice versa. The kidney tubules of mammals not only produce the most concentrated  urine observed in vertebrates but also differ from other vertebrate kidney  tubules in that they all have distinctive hairpin shapes. Physiologists have  shown that the functional ability to produce highly concentrated urine  depends on the hairpin structure, which guides the urine (as it is being  formed) to flow first in one direction and then in the opposite direction. 140 (Fig. 6.13)
259 (Fig. 10.35)
370 (Fig. 14.10)
649 (Fig. 25.1)
768 (Fig. 29.12)
Applicability of the Laws of Chemistry and  Physics:  Animals must adhere to the laws of chemistry and  physics. Sometimes chemistry and physics act as  constraints, but sometimes animals gain advantages  by evolving systems that capitalize on particular  chemical or physical principles. Heat transfer through air follows different physical laws when the air is  still rather than moving; heat tends to move much more slowly through  still air than moving air. Animals cannot change such laws of physics.  They sometimes can affect which law applies to them, however, as when  the ancestors of mammals evolved fur. The hairs of a furred mammal  keep the layer of air next to the body relatively motionless. Heat transfer  through that air is therefore slow, helping mammals retain internal heat  when living in cold environments.  230
493 (Fig. 18.8)
576 (Box 22.2)
694–695
The Interdependency of Levels of  Organization:  An animal’s overall functional properties depend on  how its tissues and organs function, and the function  of its tissues and organs depends on how its cells and  molecular systems  function. All these levels of organi- zation are interdependent. An important corollary  is that properties at one level of organization often  cannot be fully understood without exploring other  levels of organization. When your physician strikes a tendon near your knee with a mallet, your  leg straightens. For this response, electrical signals must travel along  nerve cells to the spinal cord and back. The rate of travel depends in part  on the molecular properties of ion-transporting proteins in the cell mem- branes of the nerve cells. It also depends in part on key cellular properties,  such as the spacing between the sections of each nerve cell membrane  that are fully exposed to the fluids bathing the cell. Molecular and cellular  properties of these sorts determine the overall properties of the process.  For instance, they determine the length of time that passes between the  moment the mallet strikes and the moment your leg muscles contract.  6 (Fig. 1.2)
116 (Box 5.2)
199 (Fig. 8.12)
306–307
508 (Fig. 19.4)
The Crucial Importance of Control  Mechanisms: In addition to mechanisms for reproducing, breath- ing, moving, and carrying out other overt functions,  animals require control mechanisms that orchestrate  the other mechanisms. The control mechanisms—so  diverse that they include controls of gene expres- sion as well as those exerted by the nervous  and endocrine systems—determine the relations  between inputs and outputs in physiological sys- tems. They thereby crucially affect the functional  properties of animals. Although sheep and reindeer are born at cold times of year, newborns  receive no heat from their parents and must keep warm on their own or  die. They possess a process for rapid heat production. Proper control of  this process requires that it be activated at birth, but not before birth when  it would tend needlessly to exhaust fetal energy supplies. The control  mechanism has two key properties: It activates heat production when  neural thermal sensors detect cold, but its capacity to activate heat produc- tion is turned off by chemical factors secreted by the placenta. The control  mechanism remains in a turned-off state until a newborn is separated from  the placenta at birth. The cold environment is then able to stimulate rapid  heat production. 50 (Fig. 2.19)
252 (Box 10.2)
283 (Fig. 11.8)
480 (Fig. 17.15)
background image   Themes in the Study of Animal Physiology Theme An Example of the Theme in Action See Pages The Study of Function:  Animal physiology is the study of how animals  function. That is, it is the study of how their cells  and organs operate. When physiologists study muscle, one of their goals is to understand how  the proteins in muscle cells are able to develop mechanical forces, which  are employed in locomotion, heart contraction, or other activities.  114 (Fig. 5.12)
527 (Fig. 20.5)
606 (Fig. 23.22)
687 (Fig. 26.9)
Integration of the Sciences:  Physiologists often find that they must integrate  knowledge of mathematics, chemistry, or physics  with knowledge of biology to answer important  questions. Physiology is one of the most integrative  branches of biology. To understand how animals employ odors to orient their movements,  physiologists study the chemical structural differences between molecules  that attract or repel, and they mathematically describe the physics of how  winds or water currents transport odor molecules from odor sources to  the olfactory organs of animals. 6 (Fig. 1.2)
60 (Fig. 2.28)
165 (Fig. 7.3)
377
650 (Fig. 25.2)
Emphasis on Quantitative Methods:  Physiologists quantify the properties of animals as  carefully as possible as they seek to test hypotheses  or make predictions. Starting in ancient Roman times, people thought that the dromedary camel  could carry enough water in its rumen to explain its unusual ability to live  without drinking. When physiologists quantified the amount of water in  the camel rumen rather than just speaking qualitatively about it, however,  they found that there was not nearly enough water in the rumen for the  old idea to make sense. Negating the old idea helped lead to understand- ing that camels do not store water to a greater degree than other mam- mals. Instead, they have excellent abilities to conserve water and endure  dehydration. 209 (Box 9.1)
212 (Fig. 9.7)
234–235  
   (Fig. 10.8)
799–800
The Tandem Goal of Understanding  Mechanism and Adaptation:  When physiologists study a process, they typically  emphasize a two-part goal: They try to understand  both the mechanism involved (i.e., how the process  is executed) and the potential adaptive significance of  the process (i.e., how, if at all, it enhances evolu- tionary fitness). A number of animals, including fireflies and certain fish, produce light.  When physiologists study light production in such animals, they try to  learn both how the animals make light and why they make it. 5–9
248–249  
   (Fig. 10.26)
455–456
735 (Fig. 28.12)
760 (Fig. 29.5)
The Comparative Method:  To understand the adaptive significance of animal  features, physiologists make extensive use of the  comparative method , which is the examination of how  particular functions are carried out by related and  unrelated species living in similar and dissimilar  environments. When physiologists compare animals as distantly related as mammals and  insects, they find that desert species tend consistently to have great abili- ties to concentrate their urine. Desert species of mammals are typically  able to make urine of higher concentration than nondesert mammals, and  desert insects are similarly superior to nondesert insects. These compara- tive observations provide evidence that the ability to make concentrated  urine is an advantage—favored by natural selection—in deserts. 26 (Fig. 1.18)
114 (Fig. 5.13)
675 (Box 25.3)
688 (Box 26.1)
743 (Fig. 28.20)
Phylogenetic Reconstruction: To understand the evolution of physiological  properties—and thus gain perspective on the evo- lutionary significance of modern-day properties— physiologists employ phylogenetic reconstructions, in  which genetic or other information on multiple spe- cies is used to reconstruct the paths of evolution. Although body temperature is the same as water temperature in most  species of fish, regardless of how big they are, certain species of fish  maintain elevated temperatures in some of their tissues. From phylo- genetic reconstructions, physiologists have found that the warm-tissue   condition evolved on at least four independent occasions. We know,  therefore, that today’s fish with warm tissues do not all simply inherit   the condition from a single common ancestor. 27
53 (Fig. 2.21)
71 (Fig. 3.5)
270 (Fig. 10.47)
732 (Box 28.4)
The Centrality of the Environment:  The specific environments in which animals have  evolved and live must be considered for the func- tional properties of the animals to make sense. Many specialists in high-altitude physiology argue that when lowland  people travel to high altitudes, some of their typical responses are more  harmful than helpful. These specialists emphasize that the human spe- cies did not evolve in high-altitude environments. Accordingly, there is  no reason to presume that all the human responses to such environments  would be beneficial. 53 (Fig. 2.22)
67
264 (Fig. 10.41)
639 (Box 24.5)
Body Size:  The physiological properties of related animal spe- cies typically scale in mathematically consistent  ways with their body sizes. These relations are  often nonproportional and thus termed allometric. The metabolic rate per gram of body weight is usually higher in small- bodied species than in related large-bodied ones. Because of this relation,  whenever two species of mammals of different body sizes—like mice and  horses—are compared, the smaller species typically needs more food per  gram of body weight than the larger one. 17 (Fig. 1.9)
173 (Fig. 7.6)
285 (Fig. 11.9)
741 (Fig. 28.18)
Listed are 15 overarching themes that reappear throughout the study of animal  physiology. Some of the listed themes overlap with, or even encompass, oth- ers; they are not intended to be mutually exclusive or, in all cases, equivalent  in importance. To help explain each theme, an illustrative example is presented  in the second column of the table. Further examples are on the pages listed in  the third column (italic listing elaborates the featured example). Theme An Example of the Theme in Action See Pages The Dynamic State of Body Constituents:  Great quantities of many of the key constituents  of the body are added and subtracted every day  in many animals under many conditions. Thus the  constituents of the body—far from being static—are  continuously in a dynamic state of flux. This is true  even though additions and subtractions are often  relatively balanced, resulting in relatively constant  concentrations (a phenomenon termed homeostasis). Averaged over the course of an ordinary 24-h day, an adult person is like- ly to process more than 2 kg of adenosine triphosphate (ATP) each hour,  synthesizing that amount of ATP from adenosine diphosphate (ADP)  and, with only a short delay, breaking it back down to ADP. To synthe- size the ATP, the person—during each hour—will use about 20 liters of  oxygen (O 2 ) that he or she takes up from the atmosphere. During a 24-h  day, the oxygen used will combine with almost 100 g (a fifth of a pound)  of hydrogen atoms that have been removed from food molecules, forming  about 800 milliliters of water. This water is added to the body fluids. 11–12
183–184
378
699
743–744  
   (Fig. 28.21) Multiple Forms of Key Molecules:  Animals have often evolved multiple molecular  forms (sometimes called isoforms) of particular  proteins or other sorts of molecules.  Physiologists  hypothesize that when two species or two tissues  exhibit different molecular forms of a molecule, the  forms are often specialized to function in the spe- cific settings in which the animals live or the tissues  function. The cell membranes of all animals are composed principally of lipid mol- ecules. Physiologists have found, however, that the membranes of all ani- mals are not composed of chemically identical lipid molecules. Instead,  multiple molecular forms of lipids are employed by different animals  living under different circumstances. Cold-water fish species, for instance,  construct their cell membranes using molecular forms of lipids that are  less likely to harden at low temperatures than the molecular forms syn- thesized by warm-water species. 34 (Fig. 2.3)
242–243  
   (Fig. 10.19)
537
620 (Fig. 24.2)
640 (Fig. 24.20)
Phenotypic Plasticity:  An individual animal is often able to change its  phenotype in response to changes in the particular  circumstances under which it is living (e.g., its par- ticular environment). This ability of an individual  animal to adopt two or more phenotypes despite  having a fixed genotype is termed phenotypic  plasticity . Animals that eat only occasionally, such as pythons, often alternate  between two intestinal phenotypes. When they have not had a meal for  weeks, their intestinal tract is physically small, and it has poorly devel- oped molecular mechanisms for absorbing food. After a meal, the tissues  of the intestinal tract enlarge greatly, and the intestinal tract expresses  well-developed absorption mechanisms. 15 (Fig. 1.5)
79 (Table 3.1)
90–92
157 (Box 6.2)
264 (Fig. 10.40)
555 (Fig. 21.7)
Interdependency of Function and Form: The function of a biological system typically cannot  be understood without knowledge of its structure and vice versa. The kidney tubules of mammals not only produce the most concentrated  urine observed in vertebrates but also differ from other vertebrate kidney  tubules in that they all have distinctive hairpin shapes. Physiologists have  shown that the functional ability to produce highly concentrated urine  depends on the hairpin structure, which guides the urine (as it is being  formed) to flow first in one direction and then in the opposite direction. 140 (Fig. 6.13)
259 (Fig. 10.35)
370 (Fig. 14.10)
649 (Fig. 25.1)
768 (Fig. 29.12)
Applicability of the Laws of Chemistry and  Physics:  Animals must adhere to the laws of chemistry and  physics. Sometimes chemistry and physics act as  constraints, but sometimes animals gain advantages  by evolving systems that capitalize on particular  chemical or physical principles. Heat transfer through air follows different physical laws when the air is  still rather than moving; heat tends to move much more slowly through  still air than moving air. Animals cannot change such laws of physics.  They sometimes can affect which law applies to them, however, as when  the ancestors of mammals evolved fur. The hairs of a furred mammal  keep the layer of air next to the body relatively motionless. Heat transfer  through that air is therefore slow, helping mammals retain internal heat  when living in cold environments.  230
493 (Fig. 18.8)
576 (Box 22.2)
694–695
The Interdependency of Levels of  Organization:  An animal’s overall functional properties depend on  how its tissues and organs function, and the function  of its tissues and organs depends on how its cells and  molecular systems  function. All these levels of organi- zation are interdependent. An important corollary  is that properties at one level of organization often  cannot be fully understood without exploring other  levels of organization. When your physician strikes a tendon near your knee with a mallet, your  leg straightens. For this response, electrical signals must travel along  nerve cells to the spinal cord and back. The rate of travel depends in part  on the molecular properties of ion-transporting proteins in the cell mem- branes of the nerve cells. It also depends in part on key cellular properties,  such as the spacing between the sections of each nerve cell membrane  that are fully exposed to the fluids bathing the cell. Molecular and cellular  properties of these sorts determine the overall properties of the process.  For instance, they determine the length of time that passes between the  moment the mallet strikes and the moment your leg muscles contract.  6 (Fig. 1.2)
116 (Box 5.2)
199 (Fig. 8.12)
306–307
508 (Fig. 19.4)
The Crucial Importance of Control  Mechanisms: In addition to mechanisms for reproducing, breath- ing, moving, and carrying out other overt functions,  animals require control mechanisms that orchestrate  the other mechanisms. The control mechanisms—so  diverse that they include controls of gene expres- sion as well as those exerted by the nervous  and endocrine systems—determine the relations  between inputs and outputs in physiological sys- tems. They thereby crucially affect the functional  properties of animals. Although sheep and reindeer are born at cold times of year, newborns  receive no heat from their parents and must keep warm on their own or  die. They possess a process for rapid heat production. Proper control of  this process requires that it be activated at birth, but not before birth when  it would tend needlessly to exhaust fetal energy supplies. The control  mechanism has two key properties: It activates heat production when  neural thermal sensors detect cold, but its capacity to activate heat produc- tion is turned off by chemical factors secreted by the placenta. The control  mechanism remains in a turned-off state until a newborn is separated from  the placenta at birth. The cold environment is then able to stimulate rapid  heat production. 50 (Fig. 2.19)
252 (Box 10.2)
283 (Fig. 11.8)
480 (Fig. 17.15)
background image 00_Hill3e_FM.indd   ii 3/9/12   2:11 PM
background image ANIMAL PHYSIOLOGY THIRD EDITION 00_Hill3e_FM.indd   i 3/9/12   2:11 PM
background image
background image 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
background image 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: 413-549-1118
Email: publish@sinauer.comInternet: 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
background image 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
background image Thomas Kuhn wrote that a textbook is principally a means of com-
municating to students the paradigms of their time. We have had 
three principal goals in preparing the content of this book. One, 
in accord with Kuhn’s dictum, has been to articulate the central 
paradigms of contemporary animal physiology. A second content 
goal has been to provide our readers with a source of both lucid 
explanations of physiological concepts and accurate information 
about physiological systems. Our third content goal has been to 
draw attention to the cutting edges of physiological science, the 
places where the onward progress of research is challenging old 
paradigms and potentially creating footholds for new ones.
We have also had goals for presentation. Most visibly, we have  combined our words with an ambitious, informative art program. 
More fundamentally, we have strived to take advantage of all the 
assets of traditional bookmaking to achieve a book that—through 
constant integration of the full suite of pedgogically relevant ele-
ments—is a first-rate learning tool. Many sorts of professionals 
have important contributions to make for a book to be excellent. 
Thus many sorts of professionals have traditionally found personal 
fulfillment by engaging in the cooperative, synergistic production 
of books. The authors listed on the cover are just the tip of the 
iceberg. A book’s art program depends on scientific illustrators. 
Coordination between the art and the text—a key to the success 
of any textbook—depends on the editorial expertise of the book’s 
editor. An attractive science text needs to be designed and physi-
cally executed by talented people who combine scientific acumen 
with artistic sensibility. To the degree that the presentation of 
the material in this book achieves success, the reason is that it is 
the creative product of a team of at least a dozen people playing 
diverse, mutually reinforcing roles. One of our goals has been to 
take advantage of this time-proven model to provide students with 
a superior text.
In these pages, we consistently and deliberately address animal  physiology as a discipline integrated with other disciplines in biol-
ogy—especially genetics, molecular biology, evolutionary biology, 
and ecology. We also consistently emphasize the roles of physiology 
throughout the life cycle of an animal by discussing physiologi-
cal development and by examining animal function during such 
important life-cycle processes as exercise, long-distance migration, 
seasonal rhythms, and accommodation to severe conditions (we 
generally omit pathology and parasitism, however). Although we give 
particular attention to mammals, we make a point of recognizing 
the other vertebrate groups and at least the arthropods and molluscs 
among invertebrates. We address all levels of organization that are 
germane, from the genome to the ecological context. 
We want to mention four specific strategies we have adopted to  add interest and breadth to the book. First, we start every chapter 
with a vivid example of the application of the chapter’s material to 
the lives of animals in their natural habitats. Second, we devote five 
entire chapters (our “At Work” chapters) to in-depth explorations 
of how physiologists do their work; in these chapters we break out 
of the usual textbook mold to discuss exciting topics—such as the 
diving physiology of marine mammals—with emphasis on experi-
ments, theory maturation, integration of physiological systems, and 
prospects for future research. Third, we include many photographs 
and drawings of animals throughout the book to remind readers of 
the animals we discuss. Fourth, entirely new to this edition, we have 
started a program of inviting specialists to contribute expert Guest 
Boxes on emerging topics that expand the book’s subject content.
With our aspirations being as numerous as we have described,  we have put a great deal of effort into balancing competing demands 
for space. The product is a complete physiology textbook that in 
one volume will meet the requirements of a diversity of one- or 
two-semester courses in animal function. Our intended audience 
is sophomores through beginning graduate students. To make 
the book accessible to as wide an audience as possible, we have 
included both a glossary of nearly 1200 terms and 11 appendices 
on important background concepts.
Our approach to the writing has been to work from the original  scientific literature and obtain extensive peer review. Another aspect 
of our approach is that we have opted for the pedagogical consis-
tency of a book written by just three principal authors. Margaret 
Anderson wrote Chapters 16, 20, and 21, and Gordon Wyse wrote 
Chapters 12–15, 18, and 19. Richard Hill wrote Chapters 1–11, 17, 
and 22–30. David S. Garbe, Scott A. Huettel, Matthew S. Kayser, 
Kenneth J. Lohmann, and Margaret McFall-Ngai wrote Guest 
Boxes. Matthew S. Kayser and Gordon Fain assisted with topic 
development in certain parts of the principal text.
NEW TO THIS EDITION: As in other editions, our two central goals 
for this edition were to update content and enhance pedagogical 
effectiveness. To these ends, we have reconsidered every sentence 
and every element of the art program. Of the 690 figures and tables 
in this edition, over 60 are either new or greatly enhanced. Chapter 
14, on sensory processes, has been entirely rewritten. We have also 
added a new chapter (Chapter 4) on physiological development and 
epigenetics. Other chapters that have received exceptional attention 
are: Chapter 5 (transport of solutes and water), Chapter 8 (aerobic 
and anaerobic metabolism), Chapter 9 (activity energetics), and 
Chapter 29 (kidney physiology, edited throughout to emphasize 
plasma regulation). The book now includes Guest Boxes on functional 
magnetic resonance imaging, magnetoreception, optogenetics, 
sleep, symbiosis in the bobtail squid-Vibrio system, and synaptic 
development. Treatment of topics in global warming has been 
tripled. Treatment of altitude physiology in Chapters 23 and 24 is 
entirely revised. The index is new and upgraded. A limited list of 
Preface 00_Hill3e_FM.indd   vi 3/9/12   2:11 PM
background image Preface  vii the many topics that have been added or substantially upgraded 
includes: aquaporins, basal ganglia function, bioluminescence, 
breathing in crocodilians, chemiosmosis, calcium metabolism, color 
change, daily rhythms in transcription, efficiency of ATP synthesis 
in oxidative phosphorylation, endothermy in plants, evolution of 
nervous systems, function of P-type ATPases, the gut microbiome, 
hibernation, hippocampal specialization in place learning, ion-
transport proteins in fish, kisspeptin neurons, metabolic scaling, 
neurotransmitter release mechanisms, photoperiodic control, 
reactive oxygen species, smooth muscle, thermal performance 
curves, and voltage-gated channels. 
The book is organized in modular fashion with the express  purpose of providing instructors and students with flexibility in 
choosing the order in which they move through the book. The 
first of the six parts (modules) consists of Chapters 1 to 5, which 
are background chapters for the book as a whole. Most instructors 
will want to assign those chapters at the beginning of the course 
of study (or, when students have exceptional preparation, skip the 
chapters in part or in whole). Each of the subsequent five parts of 
the book is written to be free-standing and self-contained, so that 
students who have mastered the material in Part I will be well 
prepared to work through any of the other five parts. Two of the 
final five parts begin with explicitly introductory chapters that 
present fundamentals. All five of these parts end with “At Work” 
chapters. Within a part, although chapters will probably be best 
read in order, most chapters are themselves written to be relatively 
self-contained, meaning that the order of reading chapters within 
a part is flexible. Three additional features promote flexibility in 
the order of reading: the glossary, the new index, and page cross-
references. Text is cross-referenced both forward and backward, 
so that instructors and students can link material across chapters.
We have tried to keep animals front and center. At the end of  our production, as the orchestra goes silent and the klieg lights dim, 
we hope that animals leading their lives in their natural habitats 
will be the enduring image and memory left by this work—animals 
now better understood, but still with much to attract the curiosity 
of upcoming generations of biologists.
Our peer reviewers are particularly important to the quality of  the book, even though at times—accepting full responsibility for 
the product—we have followed our own inclinations rather than 
theirs. We are thus happy to acknowledge our current peer reviewers 
as well as individuals who acted as reviewers for earlier editions 
and whose influence remains clearly evident: Doris Audet, Brian 
Bagatto, Jason Blank, Charles E. Booth, Eldon Braun, Warren Burg-
gren, Heather Caldwell, Jeffrey C. Carrier, Sheldon Cooper, Daniel 
Costa, Emma Creaser, David Crews, Stephanie Gardner, Stephen 
Gehnrich, Joseph Goy, Bernd Heinrich, Raymond Henry, James 
Hicks, Carl S. Hoegler, Richard Hoffman, Mark A. Holbrook, Jason 
Irwin, Steven H. Jury, William Karasov, Fred J. Karsch, Leonard 
Kirschner, Sharon Lynn, Megan M. Mahoney, Robert Malchow, 
Duane McPherson, Ulrike Muller, Barbara Musolf, Randy Nelson, 
Gilbert Pitts, Fernando Quintana, Matthew Rand, Susan Safford, 
Malcolm Shick, Bruce Sidell, Mark Slivkoff, Paul Small, George 
Somero, Frank van Breukelen, Itzick Vatnick, Curtis Walker, Zachary 
Weil, Alexander Werth, and Eric Widmaier.
Another group to whom we offer special thanks are the many  scientists who have provided us with photographs, drawings, 
or unpublished data for direct inclusion in this book: Jonathan 
Ashmore, William J. Baker, Lise Bankir, Jody M. Beers, Rudolf 
Billeter-Clark, Walter Bollenbacher, Richard T. Briggs, Klaus Bron, 
Marco Brugnoli, Jay Burnett, Christina Cheng, Daniel Costa, Mat-
thew Dalva, Hans-Ranier Duncker, Aaron M. Florn, Jamie Foster, 
Peter Gillespie, Greg Goss, Bernd Heinrich, Dave Hinds, Michael 
Hlastala, Hans Hoppeler, José Jalife, Kjell Johansen, Toyoji Kaneko, 
Matthew S. Kayser, Mary  B. Kennedy, Andor Kiss, Daniel Luchtel, 
David Mayntz, Margaret McFall-Ngai, Nathan Miller, Eric Montie, 
Michael Moore, Mikko Nikinmaa, Sami Noujaim, Dan Otte, Thomas 
Pannabecker, R. J. Paul, Steve Perry, Bob Robbins, Ralph Russell, 
Jr., Josh Sanes, Klaus Schulten, Stylianos Scordilis, Bruce Sidell, 
Helén Nilsson Sköld, Jake Socha, Kenneth Storey, Karel Svoboda, 
Emad Tajkhorshid, Irene Tieleman, Christian Tipsmark, Shinichi 
Tokishita, Walter S. Tyler, Tom Valente, Tobias Wang, Rüdiger 
Wehner, Ewald Weibel, Judith Wopereis, Eva Ziegelhoffer, and the 
Zoological Society of London.
Thanks are due too for encouragement, feedback, and other  help with writing that we have gratefully received from Richard 
T. Briggs, Michael Cook, John Dacey, Giles Duffield, Aaron M. 
Florn, Fritz Geiser, Loren Hayes, Gerhard Heldmaier, Richard L. 
Marsh, Steve Perry, George Somero, Mark Vermeij, Tobias Wang, 
and Joseph Williams.
Of course, no book of this scope emerges fully formed in a single  edition. Thus, we also thank the following who played important 
roles in earlier versions of this work: Simon Alford, Kellar Autumn, 
Robert Barlow, Al Bennett, Eric Bittman, Jeff Blaustein, Batrice Boily, 
Beth Brainerd, Richard C. Brusca, Gary Burness, Bruce Byers, John 
Cameron, Donald Christian, Barbara Christie-Pope, Corey Cleland, 
Randal Cohen, Joseph Crivello, Peter Daniel, Bill Dawson, Gregory 
Demas, Linda Farmer, Jane Feng, Milton Fingerman, Dale Forsyth, 
Christopher Gillen, Kathleen Gilmour, Judy Goodenough, Edward 
Griff, Jacob Gunn, James Harding, Jean Hardwick,  John Harley, Ian 
Henderson, David Hillis, Kay Holekamp, Charles Holliday, Henry 
John-Alder, Kelly Johnson, Alexander Kaiser, Reuben Kaufman, 
M. A. Q. Khan, William Kier, Peter King, Rosemary Knapp, 
Heather Koopman, Richard Lee, John Lepri, Robert Linsenmeier, 
Stephen Loomis, William Lutterschmidt, Steffen Madsen, Don 
Maynard, Grant McClelland, Kip McGilliard, Stephen McMann, 
00_Hill3e_FM.indd   vii 3/9/12   2:11 PM
background image viii  Preface Allen Mensinger, Tim Moerland, Thomas Moon, Thomas Moylan, 
Richard Nyhof, David O’Drobinak, Linda Ogren, Sanford Ostroy, 
Christine Oswald, Linda Peck, Sandra Petersen, Chuck Peterson, 
Richard Petriello, Nathan Pfost, Robert Rawding, Heinrich Reichert, 
Larry Renfro, David Richard, R. M. Robertson, Robert Roer, William 
Seddon, Brent Sinclair, Laura Smale, Amanda Southwood, Tony 
Stea, Philip Stephens, Georg Striedter, Rebekah Thomas, Heather 
Thompson, Irene Tieleman, Lars Tomanek, Terry Trier, Kay Ueno, 
Joshua Urio, Mark Wales, Winsor Watson, Leonard E. White, Susan 
Whittemore, Steve Wickler, Robert Winn, and Tom Zoeller.
Of the many colleagues who have made contributions, Richard  Hill would like in particular to thank Kjell Johansen, one of the 
greats, who way back at the beginning said without a moment’s 
hesitation, “This is good.” Energy still emanates from those words 
three decades later.
Thanks to our students, who have challenged us, encouraged  us, taught us, and—if nothing else—listened to us over our many 
years of classroom teaching. Our classes with our students have 
been our proving ground for teaching physiology and our most 
fundamental source of reinforcement to take on a project of this 
magnitude. We are grateful to work and teach at institutions—
Michigan State University, the University of Massachusetts, and 
Smith College—at which efforts of this sort are possible.
Special thanks to Andy Sinauer, who has helped us to think  big and provided the resources to realize ambitious goals for three 
editions. We have all worked with many editors and publishers in 
our careers, and Andy is tops: an entrepreneur dedicated to put-
ting the life of ideas on the printed page. We also extend special 
thanks to our editor, Laura Green, who has brought expertise and 
sound judgment to our work on every aspect of the book, includ-
ing text, art, and pedagogy. Warm thanks, too, to Chris Small, 
head of production, David McIntyre, photo editor, Joan Gemme, 
production specialist, and the others at Sinauer Associates whose 
talents and dedication have been indispensable. We feel privileged 
to have had Elizabeth Morales execute the art, which makes such 
a contribution to our pages.
We each have particular thanks to offer to the people in our  personal lives whose support and patience have been indispensable. 
Richard Hill thanks Sue, Dave, and Chrissie, who have always been 
there even though the hours of writing have often meant long waits 
between sightings of their husband and father. Sue in particular 
has been a major contributor by repeatedly offering the benefits of 
her knowledge and judgment as a biologist. Gordon Wyse thanks 
Mary for her editorial talents, support, and willingness to keep 
planning around this long project, and Jeff, Karen, and Nancy for 
inspiration. Likewise, Margaret Anderson expresses gratitude to her 
family, especially Andy and Anita, and to her friends and students, 
whose boundless enthusiasm and idealism provide great inspiration.
While acknowledging the many ways others have helped, we  of course accept full responsibility for the finished product and 
invite readers’ opinions on how we could do better. Please contact 
us with your observations.
One of the gratifications of writing a book like this is the  opportunity to participate in the raw enthusiasm of scientists for 
science. On countless occasions, many colleagues have performed 
great favors on short notice without the slightest hint of wanting pay 
for their professional expertise. Pure science must be one of the last 
redoubts of this ethic in today’s professional world. We are honored 
to play the role of synthesizing and communicating the insights 
and questions that arise from the exciting search for knowledge.
R ICHARD  W. H ILL East Lansing, Michigan G ORDON  A. W YSE Amherst, Massachusetts M ARGARET  A NDERSON Northampton, Massachusetts February 2012 00_Hill3e_FM.indd   viii 3/9/12   2:11 PM
background image If you’ve ever been to a show and one of the producers stepped 
out on stage before the curtain went up to offer remarks about 
the upcoming event, you will understand the nature of these two 
pages. We, your authors, want to say a few words about the way 
we approached writing this book. We would also like to mention 
how we have handled several challenging issues.
One of our primary goals has been to create a book in which you  will find the fascination of physiology as well as its content. Thus 
we have started each of the 30 chapters with an intriguing example 
that illustrates the application of the chapter to understanding the 
lives of animals. Collectively, these examples highlight the many 
ways in which the study of physiology relates to biology at large.
Besides our desire to emphasize the fascination of physiology, we  have also wanted to stress the importance of integrating knowledge 
across physiological disciplines—and the importance of integrating 
physiology with ecology, behavior, molecular biology, genetics, and 
other fields. We have wanted, in addition, to discuss how concepts 
are tested and revised during research in physiology and to focus 
on the cutting edges in physiological research today. To help meet 
these goals, we have included five “At Work” chapters, which ap-
pear at the ends of five of the book’s six parts. You will find that the 
“At Work” chapters are written in a somewhat different style than 
the other chapters because they give extra emphasis to the process
of discovery. For the topics of the “At Work” chapters, we chose 
subjects that are especially intriguing and important: diving by 
seals and whales, animal navigation, muscle in states of use (e.g., 
athletic training) and disuse, mammals in the Arctic, and desert 
animals. Each “At Work” chapter uses concepts introduced in the 
chapters preceding it. We hope you will find these chapters to be 
something to look forward to: enjoyable to read and informative.
One of the thrills of science today, besides the extraordinary  pace at which new knowledge is being generated worldwide, is the 
revolution in how readily each of us can access information. The 
first step in learning more about a field of knowledge is to gather 
references. Even as recently as 15 years ago, the reference-gathering 
stage could easily require days or weeks. Today, however, the search 
engines available to find references in the scientific literature enable 
rapid review and assembly of information sources. Specialized 
search engines such as the Web of Science
® —which are similar  to internet search engines but far more effective for exploring the 
scientific literature—will permit you to glean references rapidly from 
the thousands of scientific journals in which research is reported. 
Such search engines will then enable you to read the abstracts of 
dozens of papers in a few hours of time, so you can identify the 
research reports and other papers you want to read in full. Today 
is the information era. And indeed, knowledge is power. We en-
courage you to place a priority on mastering the tools available for 
information-gathering from the scientific literature.
You might wonder, if information is so easy to find, why should  I take the course in which I am enrolled and why should I read this 
book? The answer in a few words is that extraordinary quantities of 
information create extraordinary challenges for synthesis. The more 
information each of us can locate, the more we need frameworks 
for organizing knowledge. Scientists, philosophers, and historians 
who comment on the practice of science are of one mind that the 
mere accumulation of facts leads quite literally nowhere. The suc-
cessful pursuit of scientific knowledge requires testable concepts 
that organize facts. Scientists create concepts that organize raw 
information. Then, in science, it is these concepts that we test for 
their accuracy and utility.
A good course taught with a good textbook provides a concep- tual framework into which raw information can be fitted so that it 
becomes part of the life of ideas and concepts. We hope we have 
provided you not simply with a conceptual framework, but one that 
is “good for the future.” By this we mean we have not tried merely 
to organize the knowledge already available. We have tried in equal 
measure to articulate a conceptual framework that is poised to grow 
and mature as new knowledge becomes available.
Just briefly we want to comment on four particular topics. First,  our Box design. Boxes that start on the pages of this book often 
continue on the web. To find the web content, go to the book’s 
website that is mentioned prominently at the end of each chapter. 
The part of a Box that you will read online is called a Box Exten-
sion. All the Box Extensions are fully integrated with the rest of the 
book in terms of concepts, terminology, and artistic conventions. 
Moreover, many of the Box Extensions are extensive and include 
informative figures. Thus, we urge that you keep reading when a 
Box directs you to a Box Extension.
Second, units of measure. For 30 years there has been a revolu- tion underway focused on bringing all human endeavor into line 
with a single system of units called the Système International (SI). 
Different countries have responded differently, as have different 
fields of activity. Thus, if you purchase a box of cereal in much of 
the world, the cereal’s energy value will be quoted on the box in 
kilojoules, but elsewhere it will be reported in kilocalories. If you 
go to a physician in the United States and have your blood pressure 
measured, you will have it reported in millimeters of mercury, but 
if you read a recent scientific paper on blood pressures, the pres-
sures will be in kilopascals. The current state of transition in units 
of measure presents challenges for authors just as it does for you. 
We have tried, in our treatment of each physiological discipline, to 
familiarize you with the pertinent units of measure you are most 
likely to encounter
 (SI or not). Moreover, you will find in Appendix 
A an extensive discussion of the Système International and its 
relations to other systems of units.
To Our Readers 00_Hill3e_FM.indd   ix 3/9/12   2:11 PM
background image x   To Our Readers A third specific matter we want to mention is the classifica- tion of birds. Systematists now agree that birds and crocodilians 
(alligators and crocodiles) are more closely related to each other 
than crocodilians are related to lizards, snakes, and turtles. This 
means that, logically, when we speak of reptiles, the birds belong 
with them. Probably your textbook in general biology has already 
presented this new classification of the vertebrates. In this book, we 
treat birds as being reptiles, but we also make a point of speaking 
of the groups in ways that keep the traditional distinctions clear.
The fourth and final specific matter on our minds is to mention  our referencing system. For each chapter, there are three reference 
lists: (1) a brief list of particularly important or thought-provoking 
references at the end of the chapter, (2) a longer list of references 
in the section titled Additional References at the back of the book, 
and (3) a list of all the references cited as sources of information for 
figures or tables in the chapter. The final list appears in the Figure 
and Table Citations at the back of the book; highly detailed or specific 
references that we used to prepare figures or tables often appear 
only in the Figure and Table Citations. In terms of the formats used 
in citations to research reports in the scientific journals, our most 
common citation format is to provide the journal volume number 
and inclusive page numbers where a report is found. However, 
online journals often do not employ inclusive page numbers; in 
those cases, our citation provides the volume number and an index 
to the location of the report in that volume. In unusual cases, our 
citation format is to employ a doi (digital object identifier) number. 
You can enter the doi number into a search engine to find the 
pertinent research report; the most reliable search engine for use 
of doi numbers is found at www.doi.org. 
All three of us who wrote this book have been dedicated teach- ers throughout our careers. In addition, we have been fortunate to 
develop professional relationships and friendships with many of 
our students. This book is a product of that two-way interaction. 
In the big universities today, there are many forces at work that 
encourage passivity and anonymity. We urge the opposite. We 
encourage you to talk science as much as possible with each other 
and with your instructors, whether in classroom discussions, study 
groups, office hours, or other contexts. Active learning of this sort 
will contribute in a unique way to your enjoyment and mastery of 
the subjects you study. We have tried, deliberately, to write a book 
that will give you a lot to talk about.
R ICHARD  W. H ILL   G ORDON  A. W YSE   M ARGARET  A NDERSON 00_Hill3e_FM.indd   x 3/9/12   2:11 PM
background image eBook (ISBN 978-0-87893-879-7) www.coursesmart.com Animal Physiology, Third Edition is available as an eBook via CourseSmart, at a substan-
tial discount off the price of the printed textbook. The CourseSmart eBook reproduces 
the look of the printed book exactly, and includes convenient tools for searching the text, 
highlighting, and note-taking. The eBook is viewable in any Web browser, and via free 
apps for iPhone/iPad, Android, and Kindle Fire.
Companion Website sites.sinauer.com/animalphys3e New for the Third Edition, the Animal Physiology Companion Website includes content 
that expands on the coverage in the textbook as well as study and review tools. The site 
includes Chapter Outlines & Summaries to provide quick overviews of each chapter; 
Box Extensions, which expand on topics introduced in the textbook and cover important 
additional conceptual material; Online Quizzes, which cover all the key material in each 
chapter; Flashcards and Key Terms which allow the student to master the many new 
terms introduced in the textbook; and a complete Glossary.
Instructor’s Resource Library (ISBN 978-0-87893-880-3) Available to qualified adopters, the Animal Physiology Instructor’s Resource Library in-
cludes all of the figures (including photos) and tables from the textbook in a variety of 
formats, making it easy to incorporate images from the book into your lecture presenta-
tions and other course materials. The Resource Library includes both labeled and un-
labeled versions of all figures in JPEG format (both high- and low-resolution versions) 
and in PowerPoint
®  format. New to this edition are a Test Bank, answers to the Online Quiz questions, and  answers to the end-of-chapter Study Questions. The Test Bank consists of a broad range 
of questions covering key facts and concepts in each chapter. Both multiple-choice and 
short-answer questions are provided. The Test Bank also includes the Companion Web-
site Online Quiz questions. All questions are ranked according to Bloom’s Taxonomy 
and referenced to specific textbook sections and page numbers. The entire Test Bank is 
provided in Wimba’s Diploma software, making it easy to assemble quizzes and exams 
from any combination of publisher-provided questions and instructor-created questions. 
Answers to the end-of-chapter Study Questions are provided as Word documents.
Media and Supplements  to accompany Animal Physiology,  Third Edition 00_Hill3e_FM.indd   xi 3/9/12   2:11 PM
background image 00_Hill3e_FM.indd   xii 3/9/12   2:11 PM
background image PART I Fundamentals of Physiology  1 1  Animals and Environments: Function on the  Ecological Stage  3 2  Molecules and Cells in Animal Physiology  31
3  Genomics, Proteomics, and Related 
Approaches to Physiology  67 4  Physiological Development and  Epigenetics 85  5  Transport of Solutes and Water  99  PART II Food, Energy, and Temperature  125 6  Nutrition, Feeding, and Digestion  127
7  Energy Metabolism  161
8  Aerobic and Anaerobic Forms of 
Metabolism 183 9  The Energetics of Aerobic Activity  207 10  Thermal Relations  225
11  Food, Energy, and Temperature 
at Work:   The Lives of Mammals in Frigid Places  277 PART III Integrating Systems  293 12 Neurons  295
13 Synapses  327 
14  Sensory Processes  359 
15  Nervous System Organization and Biological 
Clocks 397  16  Endocrine and Neuroendocrine  Physiology 419  17 Reproduction  455 
18  Integrating Systems 
at Work: Animal Navigation  485  PART IV Movement and Muscle  501 19  Control of Movement: The Motor Bases of  Animal Behavior  503  20 Muscle  523 
21  Movement and Muscle 
at Work:  Plasticity in  Response to Use and Disuse  549  PART V Oxygen, Carbon Dioxide, and Internal  Transport 567 22  Introduction to Oxygen and Carbon Dioxide  Physiology 569 23  External Respiration: The Physiology of  Breathing 583 24  Transport of Oxygen and Carbon Dioxide in  Body Fluids (with an Introduction to Acid-Base  Physiology) 617 25 Circulation  647
26  Oxygen, Carbon Dioxide, and Internal Transport 
at Work:  Diving by Marine Mammals  679  PART VI Water, Salts, and Excretion  697  27  Water and Salt Physiology: Introduction and  Mechanisms 699  28  Water and Salt Physiology of Animals in Their  Environments 717  29  Kidneys and Excretion (with Notes on Nitrogen  Excretion) 753  30  Water, Salts, and Excretion  at Work:  Mammals  of Deserts and Dry Savannas  787 Brief Contents 00_Hill3e_FM.indd   xiii 3/9/12   2:11 PM
background image CHAPTER 1 Animals and Environments:  Function on the Ecological Stage  3 The Importance of Physiology  4
Mechanism and Origin: Physiology’s Two Central 
Questions 5 The study of mechanism: How do modern-day animals carry out  their functions?  5 The study of origin: Why do modern-day animals possess the  mechanisms they do?  7 Natural selection is a key process of evolutionary origin  8 Mechanism and adaptive significance are distinct concepts that  do not imply each other  8 This Book’s Approach to Physiology  10
Animals 11
The structural property of an animal that persists through time is  its organization  11 Most cells of an animal are exposed to the internal environment,  not the external environment  11 The internal environment may be permitted to vary when the  external environment changes, or it may be kept constant  12 Homeostasis in the lives of animals: Internal constancy is often  critical for proper function  12 BOX 1.1  Negative Feedback  13 Time in the lives of animals: Physiology changes in five time  frames 14 BOX 1.2    The Evolution of Phenotypic Plasticity  16 Size in the lives of animals: Body size is one of an animal’s most  important traits  16 Environments 18 Earth’s major physical and chemical environments  18 The environment an animal occupies is often a  microenvironment or microclimate  22 Animals often modify their own environments  23 Evolutionary Processes  24 Some processes of evolution are adaptive, others are not  24 A trait is not an adaptation merely because it exists  25 Adaptation is studied as an empirical science  25 Evolutionary potential can be high or low, depending on  available genetic variation  27 CHAPTER 2 Molecules and Cells in Animal  Physiology 31 Cell Membranes and Intracellular Membranes  32 The lipids of membranes are structured, diverse, fluid, and  responsive to some environmental factors  33 Proteins endow membranes with numerous  functional capacities  35 BOX 2.1    Protein Structure and the Bonds That  Maintain It  35 Carbohydrates play important roles in membranes  36 Epithelia 37
Elements of Metabolism  40
Enzyme Fundamentals  40
Enzyme-catalyzed reactions exhibit hyperbolic or sigmoid  kinetics 42 Contents PART I • Fundamentals of Physiology 00_Hill3e_FM.indd   xiv 3/9/12   2:11 PM
background image Contents  xv Maximum reaction velocity is determined by the amount and  catalytic effectiveness of an enzyme  43 Enzyme–substrate affinity affects reaction velocity at the  substrate concentrations that are usual in cells  43 Enzymes undergo changes in molecular conformation and have  specific binding sites that interact  44 Enzymes catalyze reversible reactions in both directions  45 Multiple molecular forms of enzymes occur at all levels of animal  organization 46 Regulation of Cell Function by Enzymes  47 The types and amounts of enzymes present depend on gene  expression and enzyme degradation  48 Modulation of existing enzyme molecules  permits fast regulation of cell function  48 Evolution of Enzymes  52
Enzymes Are Instruments of Change in All Time 
Frames 54 The Life and Death of Proteins  54
Light and Color  55
BOX 2.2    Squid and Bioluminescent Bacteria, a Study  in Cross-Phylum Coordination: The Euprymna  scolopes–Vibrio fischeri Symbiosis  Margaret McFall-Ngai  57 Reception and Use of Signals by Cells  58 Extracellular signals initiate their effects by binding to receptor  proteins 58 Cell signal transduction often entails sequences of amplifying  effects 61 Several second-messenger systems participate in cell signal  transduction 63 CHAPTER 3 Genomics, Proteomics, and Related  Approaches to Physiology  67 Genomics 72 Genomics is inextricably linked with advanced methods of  information processing  72 One overarching goal of genomics is to elucidate the evolution  of genes and genomes  73 A second overarching goal of genomics is to elucidate the  current functioning of genes and genomes  73 Genomes must ultimately be related empirically  to phenotypes  74 Top-down versus Bottom-up Approaches  to the Study of Physiology  75 Screening or Profiling as a Research Strategy  76
The Study of Gene Transcription: Transcriptomics  76
Transcription profiling often identifies large numbers of genes  that exhibit altered transcription in response to environmental 
or other conditions  78
Transcription profiling reveals that many genes routinely  undergo daily cycles of transcription  78 Manipulations of protein synthesis can be used to clarify gene  function 79 Proteomics 80
Metabolomics 82
CHAPTER 4 Physiological Development and  Epigenetics 85 The Physiology of Immature Animals Always Differs  from That of Adults  86 Phenotypic Plasticity during Development  90 Environmental effects during development may arise from  programmed responses to the environment or may be forced 
by chemical or physical necessity  91
Insect polyphenic development underlies some of the most  dramatic cases of phenotypic plasticity  91 Epigenetics 93 Two major mechanisms of epigenetic marking are DNA  methylation and covalent modification of histones  93 Epigenetic marking during an animal’s early development affects  the animal’s lifelong phenotype  94 Epigenetic marks on paternal and maternal copies of genes set  the stage in mammals and insects for the two copies to exert 
nonequivalent effects  95
CHAPTER 5 Transport of Solutes and Water  99 Passive Solute Transport by Simple Diffusion  101 Concentration gradients give rise to the most elementary form of  simple solute diffusion  102 Electrical gradients often influence the diffusion of charged  solutes at membranes  103 Biological aspects of diffusion across membranes: Some solutes  dissolve in the membrane; others require channels  104 Diffusion of ions across cell membranes is determined by  simultaneous concentration and electrical effects  105 Diffusion often creates challenges for cells and animals  105 Concentration gradients can create electrical gradients that alter  concentration gradients  107 Passive Solute Transport by Facilitated Diffusion  108
Active Transport  108
Active transport and facilitated diffusion  are types of carrier-mediated transport  109 Basic properties of active-transport mechanisms  109 Recognition of active transport completes  our overview of a single animal cell  109 Primary and secondary active transport differ in their cellular- molecular mechanisms  110 00_Hill3e_FM.indd   xv 3/9/12   2:11 PM
background image xvi  Contents BOX 5.1    Energy Coupling via the Potential Energy of  Electrochemical Gradients  113 Active transport across an epithelium does not imply a specific  transport mechanism  114 Two epithelial ion-pumping mechanisms help freshwater fish  maintain their blood composition  114 BOX 5.2    Cellular Mechanisms of Ion Pumping in  Fresh-water Fish Gills  116 Diversity and Modulation of Channels  and Transporters  116 Osmotic Pressure and Other Colligative Properties of  Aqueous Solutions  117 Physiologists usually express osmotic pressure  in osmolar units  118 Osmotic pressures can be measured in several ways  118 Osmosis 120 Quantification and terminology  120 Hydrostatic pressures develop from osmotic pressures only  when two or more solutions interact  121 Water may dissolve in membranes or pass through aquaporin  water channels during osmosis  121 Aquaporins 121 Osmosis and solute physiology often interact  122 CHAPTER 6 Nutrition, Feeding, and Digestion  127 Nutrition 129 Proteins are “foremost”  129 Lipids are required for all membranes and are the principal  storage compounds of animals  132 Carbohydrates are low in abundance in many animals but highly  abundant when they play structural roles  133 Vitamins are essential organic compounds required in small  amounts 134 Elemental nutrition: Many minerals are essential nutrients  134 Feeding 136 Many animals feed on organisms that are individually attacked  and ingested  137 Suspension feeding is common in aquatic animals  139 Symbioses with microbes often play key roles in animal feeding  and nutrition  141 BOX 6.1    Types of Meal Processing Systems  146 Digestion and Absorption  148 Vertebrates, arthropods, and molluscs represent three important  digestive–absorptive plans  148 Digestion is carried out by specific enzymes operating in three  spatial contexts  151 Absorption occurs by different mechanisms for hydrophilic and  hydrophobic molecules  153 Responses to Eating  155
Nutritional Physiology in Additional Time Frames  157
Nutritional physiology is responsive to the environment  157 BOX 6.2    Long-term Natural Fasting, Emphasizing  Pythons 157 The nutritional physiology of individuals is often endogenously  programmed to change over time  158 CHAPTER 7 Energy Metabolism  161 Why Animals Need Energy: The Second Law of  Thermodynamics 161 Fundamentals of Animal Energetics  163 The forms of energy vary in their capacity for physiological  work 163 Transformations of high-grade energy are always inefficient  163 Animals use energy to perform three major functions  164 BOX 7.1    Views on Animal Heat Production  165 Metabolic Rate: Meaning and Measurement  166 BOX 7.2    Units of Measure for Energy and Metabolic  Rates 166 Direct calorimetry: The metabolic rate of an animal can be  measured directly  167 Indirect calorimetry: Animal metabolic rates are usually  measured indirectly  167 BOX 7.3    Direct Measurement versus Indirect  Measurement 168 PART II • Food, Energy, and Temperature 00_Hill3e_FM.indd   xvi 3/9/12   2:11 PM
background image Contents  xvii BOX 7.4    Respirometry 170 Factors That Affect Metabolic Rates  170 Ingestion of food causes metabolic rate to rise  170 Basal Metabolic Rate and Standard Metabolic Rate  172
Metabolic Scaling: The Relation between Metabolic 
Rate and Body Size  172 Resting metabolic rate is an allometric function of body weight  in related species  173 The metabolic rate of active animals is often also an allometric  function of body weight  175 The metabolism–size relation has important physiological and  ecological implications  176 BOX 7.5    Scaling of Heart Function  177 The explanation for allometric metabolism–size relations  remains unknown  178 Energetics of Food and Growth  180
Conclusion: Energy as the Common Currency of 
Life 181 POSTSCRIPT:  The Energy Cost of Mental Effort  181 CHAPTER 8 Aerobic and Anaerobic Forms  of Metabolism  183 Mechanisms of ATP Production and Their  Implications 184 Aerobic catabolism consists of four major sets of reactions  184 BOX 8.1    Reactive Oxygen Species (ROS)  189 O 2  deficiency poses two biochemical challenges: Impaired ATP  synthesis and potential redox imbalance  189 Certain tissues possess anaerobic catabolic pathways that  synthesize ATP  190 Anaerobic glycolysis is the principal anaerobic catabolic pathway  of vertebrates  190 What happens to catabolic end products?  190 The functional roles of ATP-producing mechanisms depend on  whether they operate in steady state or nonsteady state  191 Phosphagens provide an additional mechanism of ATP  production without O 192 Internal O 2  stores may be used to make ATP  192 Comparative Properties of Mechanisms of ATP  Production 193 Question 1: What is each mechanism’s total possible ATP yield  per episode of use?  193 Question 2: How rapidly can ATP production be  accelerated? 193 BOX 8.2   Genetic Engineering as a Tool to Test  Hypotheses of Muscle Function and  Fatigue 194 Question 3: What is each mechanism’s peak rate of ATP  production (peak power)?  194 Question 4: How rapidly can each mechanism  be reinitialized?  194 Conclusion: All mechanisms have pros and cons  194 Two Themes in Exercise Physiology: Fatigue and  Muscle Fiber Types  194 Fatigue has many, context-dependent causes  194 The muscle fibers in the muscles used for locomotion are  heterogeneous in functional properties  195 The Interplay of Aerobic and Anaerobic Catabolism  during Exercise  196 Metabolic transitions occur at the start and end of vertebrate  exercise 196 The ATP source for all-out exercise varies in a regular manner  with exercise duration  198 Related species and individuals within one species are often  poised very differently for use of aerobic and anaerobic 
catabolism 200
Responses to Impaired O 2  Influx from the  Environment 201 Air-breathing vertebrates during diving: Preserving the brain  presents special challenges  201 Animals faced with reduced O 2  availability in their usual  environments may show conformity or regulation of aerobic 
ATP synthesis  202
Water-breathing anaerobes: Some aquatic animals are capable of  protracted life in water devoid of O 2  202 BOX 8.3    Human Peak O 2  Consumption and Physical  Performance at High Altitudes  204 CHAPTER 9 The Energetics of Aerobic Activity  207 How Active Animals Are Studied  208 BOX 9.1    The Cost of Carrying Massive Loads  209 The Energy Costs of Defined Exercise  210 The most advantageous speed depends on the function of  exercise 211 The minimal cost of transport depends in regular ways on mode  of locomotion and body size  213 The Maximal Rate of Oxygen Consumption  215 BOX 9.2    Finding Power for Human-Powered  Aircraft 215 V O2max  differs among phyletic groups and often from species to  species within a phyletic group  216 V O 2 max  varies among individuals within a species  217 V O 2 max  responds to training and selection  217 The Energetics of Routine and Extreme Daily Life  218
Long-Distance Migration  219
Ecological Energetics  220
BOX 9.3    Eel Migration and Energetics: A 2300-Year  Detective Story  221 00_Hill3e_FM.indd   xvii 3/9/12   2:11 PM
background image xviii  Contents CHAPTER 10 Thermal Relations  225 Temperature and Heat  227
Heat Transfer between Animals and Their 
Environments 227 BOX 10.1  Global Warming  228 Conduction and convection: Convection is intrinsically  faster 230 Evaporation: The change of water from liquid to gas carries  much heat away  230 Thermal radiation permits widely spaced objects to exchange  heat at the speed of light  231 Poikilothermy (Ectothermy)  233 Poikilotherms often exert behavioral control over their body  temperatures 234 Poikilotherms must be able to function over a range of body  temperatures 234 Poikilotherms respond physiologically to their environments in  all three major time frames  234 Acute responses: Metabolic rate is an approximately exponential  function of body temperature  235 Chronic responses: Acclimation often blunts metabolic  responses to temperature  236 The rate–temperature relations and thermal limits of individuals:  Ecological decline occurs at milder temperatures than acute 
stress 239
Evolutionary changes: Species are often specialized to live at  their respective body temperatures  241 Temperature and heat matter to animals because they affect the  rates of processes and the functional states of molecules  242 Poikilotherms threatened with freezing: They may survive by  preventing freezing or by tolerating it  246 Homeothermy in Mammals and Birds  250 Metabolic rate rises in cold and hot environments because of the  costs of homeothermy  251 BOX 10.2    Thermoregulatory Control, Fever, and  Behavioral Fever  252 The shape of the metabolism–temperature curve depends on  fundamental heat-exchange principles  252 Homeothermy is metabolically expensive  255 Insulation is modulated by adjustments of the pelage or  plumage, blood flow, and posture  256 Heat production is increased below thermoneutrality by  shivering and nonshivering thermogenesis  256 Regional heterothermy: In cold environments, allowing some  tissues to cool can have advantages  257 Countercurrent heat exchange permits selective restriction of  heat flow to appendages  258 Mammals and birds in hot environments: Their first lines of  defense are often not evaporative  260 Active evaporative cooling is the ultimate line of defense against  overheating 261 Mammals and birds acclimatize to winter and summer  263 Evolutionary changes: Species are often specialized to live in  their respective climates  264 Mammals and birds sometimes escape the demands of  homeothermy by hibernation, torpor, or related processes  265 Warm-Bodied Fish  268
Endothermy and Homeothermy in Insects  270
The insects that thermoregulate during flight require certain  flight-muscle temperatures to fly  271 Solitary insects employ diverse mechanisms of  thermoregulation 272 Colonies of social bees and wasps often display sophisticated  thermoregulation 273 Coda 273 BOX 10.3  Warm Flowers  273 CHAPTER 11 Food, Energy, and Temperature  at  Work:  The Lives of Mammals in Frigid  Places 277 Food, Nutrition, Energy Metabolism, and  Thermoregulation in the Lives of Adult  Reindeer 277 Newborn Reindeer  280 BOX 11.1    Knockout Mice Clarify the Function of  Brown Fat  281 BOX 11.2    Genomics Confirms That Piglets Lack  Brown Fat  282 The Future of Reindeer: Timing and Ice  283
Thermoregulatory Development: Small Mammals 
Compared with Large  283 The Effect of Body Size on Mammals’ Lives in Cold  Environments: An Overview  284 Hibernation as a Winter Strategy:  New Directions and Discoveries  285 Arctic ground squirrels supercool during hibernation and arouse  periodically throughout their hibernation season  286 The composition of the lipids consumed before hibernation  affects the dynamics of hibernation  286 Although periodic arousals detract from the energy savings of  hibernation, their function is unknown  288 The intersection of sociobiology and hibernation physiology  289 00_Hill3e_FM.indd   xviii 3/9/12   2:11 PM
background image Contents  xix PART III • Integrating Systems CHAPTER 12 Neurons 295 The Physiology of Control: Neurons and Endocrine  Cells Compared  295 Neurons transmit electrical signals to target cells  296 Endocrine cells broadcast hormones  297 Nervous systems and endocrine systems tend to control different  processes 298 Neurons Are Organized into Functional Circuits in  Nervous Systems  298 The Cellular Organization of Neural Tissue  299 Neurons are structurally adapted to transmit action  potentials 299 Glial cells support neurons physically and metabolically  300 The Ionic Basis of Membrane Potentials  301 Cell membranes have passive electrical properties: Resistance  and capacitance  302 Resting membrane potentials depend on selective permeability  to ions: The Nernst equation  305 Ion concentration differences result from active ion transport  and from passive diffusion  306 Membrane potentials depend on the permeabilities to and  concentration gradients of several ion species: The Goldman 
equation 308
Electrogenic pumps also have a small direct effect on V m   308 The Action Potential  309 Action potentials are voltage-dependent, all-or-none electrical  signals 309 Action potentials result from changes in membrane  permeabilities to ions  310 The molecular structure of the voltage-dependent ion channels  reveals their functional properties  315 There are variations in the ionic mechanisms of excitable  cells 316 BOX 12.1    Evolution and Molecular Function of Voltage- Gated Channels  317 BOX 12.2    Optogenetics: Controlling Cells with Light Matthew S. Kayser  318 The Propagation of Action Potentials  320 Local circuits of current propagate an action potential  320 Membrane refractory periods prevent bidirectional  propagation 320 The conduction velocity of an action potential depends on axon  diameter, myelination, and temperature  322 BOX 12.3    Giant Axons  322 CHAPTER 13 Synapses 327 Synaptic Transmission Is Usually Chemical but Can  Be Electrical  328 Electrical synapses transmit signals instantaneously  329 Chemical synapses can modify and amplify signals  329 Synaptic Potentials Control Neuronal Excitability  332 Synapses onto a spinal motor neuron exemplify functions of fast  synaptic potentials  332 Synapses excite or inhibit a neuron by depolarization or  hyperpolarization at the site of impulse initiation  332 Fast Chemical Synaptic Actions Are Exemplified by  the Vertebrate Neuromuscular Junction  333 Chemical synapses work by releasing and  responding to neurotransmitters  335 Postsynaptic potentials result from permeability changes that are  neurotransmitter-dependent and voltage-independent  335 EPSPs between neurons resemble neuromuscular EPSPs but are  smaller 336 Fast IPSPs can result from an increase in permeability to  chloride 337 Presynaptic Neurons Release Neurotransmitter  Molecules in Quantal Packets  337 Acetylcholine is synthesized and stored in the presynaptic  terminal 338 Neurotransmitter release requires voltage-dependent Ca 2+   influx 338 Neurotransmitter release is quantal and vesicular  338 Synaptic vesicles are cycled at nerve terminals in distinct  steps 339 Several proteins play roles in vesicular release and recycling  340 00_Hill3e_FM.indd   xix 3/9/12   2:11 PM
background image xx  Contents Neurotransmitters Are of Two General Kinds  341 Neurons have one or more characteristic neurotransmitters  342 An agent is identified as a neurotransmitter  if it meets several criteria  342 Vertebrate neurotransmitters have several general modes  of action  343 Neurotransmitter systems have been conserved in evolution  344 Postsynaptic Receptors for Fast Ionotropic Actions:  Ligand-Gated Channels  345 ACh receptors are ligand-gated channels that  function as ionotropic receptors  345 Many, but not all, ligand-gated channel receptors  have evolved from a common ancestor  347 Postsynaptic Receptors for Slow, Metabotropic  Actions: G Protein–Coupled Receptors  347 G protein–coupled receptors initiate signal transduction  cascades 347 Metabotropic receptors act via second messengers  347 Other mechanisms of G protein–mediated activity  349 G protein–coupled receptors mediate permeability-decrease  synaptic potentials and presynaptic inhibition  350 Synaptic Plasticity: Synapses Change Properties with  Time and Activity  350 Neurotransmitter metabolism is regulated homeostatically  351 Learning and memory may be based on synaptic plasticity  351 Habituation and sensitization in Aplysia 351 Long-term potentiation in the hippocampus  353 BOX 13.1    Synapse Formation: Competing Philosophies   Matthew S. Kayser  356 Long-term potentiation is a necessary component of  learning 356 CHAPTER 14 Sensory Processes  359 Organization of Sensory Systems  360 Sensory receptor cells can be classified in four different  ways 360 Sensory receptor cells transduce and encode  sensory information  361 Mechanoreception and Touch  362 Insect bristle sensilla exemplify mechanoreceptor responses  362 Touch receptors in the skin of mammals have specialized  endings 364 Proprioceptors monitor internal mechanical stimuli  365 Vestibular Organs and Hearing  366 Insects hear with tympanal organs  366 Vertebrate hair cells are used in hearing  and vestibular sense  366 Vertebrate vestibular organs sense acceleration and gravity  368 Sound stimuli create movements in the vertebrate cochlea that  excite auditory hair cells  369 The localization of sound is determined  by analysis of auditory signals in the CNS  372 BOX 14.1  Echolocation 373 Chemoreception and Taste  373 Insect taste is localized at chemoreceptive sensilla  373 Taste in mammals is mediated by receptor cells in taste buds  374 Olfaction 377 The mammalian olfactory epithelium contains odor generalist  receptor cells  378 The vomeronasal organ of mammals detects pheromones  380 Photoreception 381 Photoreceptor cells and eyes of different groups have evolved  similarities and differences  382 Rhodopsin consists of retinal conjugated to opsin,  a G protein–coupled receptor  382 Phototransduction in Drosophila leads to a depolarizing receptor  potential 382 The vertebrate eye focuses light onto retinal rods and cones  385 Rods and cones of the retina transduce light  into a hyperpolarizing receptor potential  386 Enzymatic regeneration of rhodopsin is slow  388 Visual Sensory Processing  389 Retinal neurons respond to contrast  389 The vertebrate brain integrates visual information through  parallel pathways  392 BOX 14.2    What roles do individual neurons play in  higher visual integration?  394 Color vision is accomplished by populations of photoreceptors  that contain different photopigments  394 00_Hill3e_FM.indd   xx 3/9/12   2:11 PM
background image Contents  xxi CHAPTER 15 Nervous System Organization and  Biological Clocks  397 The Organization and Evolution of Nervous  Systems 398 Nervous systems consist of neurons organized  into functional circuits  398 Many types of animals have evolved complex  nervous systems  398 BOX 15.1    Evolution of Nervous Systems  399 The Vertebrate Nervous System: A Guide to the  General Organizational Features of Nervous  Systems 401 Nervous systems have central and peripheral divisions  401 The central nervous system controls physiology  and behavior  401 Five principles of functional organization apply  to all mammalian and most vertebrate brains  402 BOX 15.2    Functional Magnetic Resonance  Imaging  Scott A. Huettel  405 The peripheral nervous system has somatic  and autonomic divisions that control different 
parts of the body  405
The autonomic nervous system has three divisions  406 Biological Clocks  410 Organisms have endogenous rhythms  410 BOX 15.3  Sleep  David S. Garbe  411 Biological clocks generate endogenous rhythms  412 Control by biological clocks has adaptive advantages  412 Endogenous clocks correlate with natural history  and compensate for temperature  413 Clock mechanisms are based on rhythms of gene expression  414 The loci of biological clock functions vary among animals  415 Circannual and circatidal clocks: Some endogenous clocks time  annual or tidal rhythms  416 Interval, or “hourglass,” timers can time shorter intervals  416 CHAPTER 16 Endocrine and Neuroendocrine  Physiology 419 Introduction to Endocrine Principles  420 Hormones bind to receptor molecules expressed  by target cells  421 Concentrations of hormones in the blood vary  421 Most hormones fall into three chemical classes  421 Hormone molecules exert their effects by producing biochemical  changes in target cells  423 Synthesis, Storage, and Release of Hormones  425 Peptide hormones are synthesized at ribosomes, stored in  vesicles, and secreted on demand  425 Steroid hormones are synthesized on demand prior to secretion,  and are released into the blood by diffusion  426 Types of Endocrine Glands and Cells  426
Control of Endocrine Secretion: The Vertebrate 
Pituitary Gland  427 The posterior pituitary illustrates neural control  of neurosecretory cells  427 The anterior pituitary illustrates neurosecretory control of  endocrine cells  428 Hormones and neural input modulate endocrine control  pathways 430 The Mammalian Stress Response  432 The autonomic nervous system and HPA axis coordinate the  stress response to an acute threat  433 The HPA axis modulates the immune system  434 Chronic stress causes deleterious effects  435 Plasma glucocorticoid concentrations  show seasonal variations  436 Endocrine Control of Nutrient Metabolism in  Mammals 436 Insulin regulates short-term changes in nutrient availability  436 Glucagon works together with insulin to ensure  stable levels of glucose in the blood  437 Other hormones contribute to the regulation  of nutrient metabolism  439 Endocrine Control of Salt and Water Balance in  Vertebrates 439 Antidiuretic hormones conserve water  439 The renin–angiotensin–aldosterone system  conserves sodium  440 Atrial natriuretic peptide promotes excretion  of sodium and water  442 Endocrine Control of Calcium Metabolism in  Mammals 442 Parathyroid hormone increases Ca 2+  in the blood  442 Active vitamin D increases Ca 2+  and phosphate  in the blood  442 Calcitonin opposes bone resorption and decreases Ca 2+  and  phosphate in the blood  443 Endocrine Principles in Review  444
Chemical Signals along a Distance Continuum  444
BOX 16.1    Can Mating Cause True Commitment?  445 Paracrines and autocrines are local chemical signals distributed  by diffusion  446 BOX 16.2    Hormones and Neuromodulators Influence  Behavior 447 00_Hill3e_FM.indd   xxi 3/9/12   2:11 PM
background image xxii  Contents CHAPTER 19 Control of Movement: The Motor Bases of  Animal Behavior  503 Neural Control of Skeletal Muscle Is the Basis of  Animal Behavior  503 Invertebrate neural circuits involve fewer neurons than  vertebrate circuits  504 Vertebrate spinal reflexes compensate for circumstances, as well  as initiate movements  504 BOX 19.1    Muscle Spindles  505 Motor neurons are activated primarily by central input rather  than by spinal reflexes  507 Pheromones and kairomones are used as chemical signals  between animals  447 Insect Metamorphosis  448 Insect metamorphosis may be gradual or dramatic  448 BOX 16.3  Insects in Forensics and Medicine  449 Hormones and neurohormones control insect  metamorphosis 450 CHAPTER 17 Reproduction 455 What Aspects of Reproduction Do Physiologists  Study? 457 Reproduce Once or More Than Once?—Semelparity  versus Iteroparity  459 BOX 17.1    Semelparity in a Mammal  460 Eggs, Provisioning, and Parental Care  460
External or Internal Fertilization?  461
The Environment as a Player in Reproduction  462
The Timing of Reproductive Cycles  463
Sperm storage permits flexible timing between copulation and  fertilization 463 Embryonic diapause permits flexible timing between fertilization  and the completion of embryonic development  463 The timing of reproductive events is often rigorously controlled  in seasonal environments  464 Sex Change  467
Reproductive Endocrinology of Placental 
Mammals 468 Females ovulate periodically and exhibit menstrual  or estrous cycles  468 Males produce sperm continually during the reproductive  season 473 BOX 17.2    Sex Determination and Differentiation,  Emphasizing Mammals  476 Pregnancy and birth are orchestrated by specialized endocrine  controls 477 Lactation is governed by neuroendocrine reflexes  480 CHAPTER 18 Integrating Systems  at Work:   Animal Navigation  485 The Adaptive Significance of Animal Navigation  486 Navigational abilities promote reproductive success  486 Navigational abilities facilitate food acquisition  487 Migrating animals need navigation  487 Navigational Strategies  487 Trail following is the most rudimentary form of animal  navigation 488 Piloting animals follow a discontinuous series of learned  cues 488 Path integration is a form of dead reckoning  489 Animals can derive compass information from environmental  cues 489 Some animals appear to possess a map sense  494 BOX 18.1    Magnetoreceptors and Magnetoreception  Kenneth J. Lohmann  495 Sea turtles exemplify the degree of our understanding of  navigation 496 Innate and Learned Components of Navigation  497 Some forms of navigation have strong innate aspects  497 The hippocampus is a critical brain area for vertebrate spatial  learning and memory  497 PART IV • Movement and Muscle 00_Hill3e_FM.indd   xxii 3/9/12   2:11 PM
background image Contents  xxiii Neural Generation of Rhythmic Behavior  509 Locust flight results from an interplay of central  and peripheral control  509 There are different mechanisms of central pattern  generation 510 Central pattern generators can underlie relatively complex  behavior 513 Control and Coordination of Vertebrate  Movement 514 Locomotion in cats involves a spinal central pattern  generator 515 Central pattern generators are distributed and interacting  515 The generation of movement involves several areas in the  vertebrate brain  516 BOX 19.2    Basal Ganglia and Neurodegenerative  Diseases 521 CHAPTER 20 Muscle 523 Vertebrate Skeletal Muscle Cells  524 Thick and thin filaments are polarized polymers  of individual protein molecules  526 Muscles require ATP to contract  527 Calcium and the regulatory proteins tropomyosin  and troponin control contractions  528 Excitation–Contraction Coupling  529
Whole Skeletal Muscles  531
Muscle contraction is the force generated by a muscle during  cross-bridge activity  531 A twitch is the mechanical response of a muscle to a single  action potential  532 The velocity of shortening decreases as the load increases  532 The frequency of action potentials determines the tension  developed by a muscle  532 A sustained high calcium concentration in the cytoplasm permits  summation and tetanus  533 The amount of tension developed by a muscle depends on the  length of the muscle at the time it is stimulated  534 In general, the amount of work a muscle can do depends on its  volume 535 BOX 20.1    Electric Fish Exploit Modified Skeletal  Muscles to Generate Electric Shocks  536 Muscle Energetics  536 ATP is the immediate source of energy for powering muscle  contraction 536 Vertebrate muscle fibers are classified into different types  537 BOX 20.2  Insect Flight  539 Neural Control of Skeletal Muscle  540 The vertebrate plan is based on muscles organized into motor  units 540 The innervation of vertebrate tonic muscle is intermediate  between the general vertebrate and arthropod plans  540 The arthropod plan is based on multiterminal innervation of  each muscle fiber by more than one neuron  540 Vertebrate Smooth (Unstriated) Muscle  542 Smooth muscle cells are broadly classified  542 Ca 2+  availability controls smooth muscle contraction by myosin- linked regulation   543 Most smooth muscles are innervated by the autonomic nervous  system 545 Vertebrate Cardiac Muscle  545 CHAPTER 21 Movement and Muscle  at Work:  Plasticity  in Response to Use and Disuse  549 Muscle Phenotypes  550 Power output determines a muscle’s contractile performance,  and changes in response to use and disuse  551 Endurance training elicits changes in fiber type, increased  capillary density, and increased mitochondrial density  551 Resistance training causes hypertrophy and changes in fiber  type 555 Hypertrophy also occurs in cardiac muscles  557 Atrophy 559 Humans experience atrophy in microgravity  559 Disuse influences the fiber-type composition of muscles  560 Muscles atrophy with age  560 Some animals experience little or no disuse atrophy  561 BOX 21.1    No Time to Lose  562 Regulating Muscle Mass  563 Myostatin 563 The PI3-K–Akt1 pathway  564 Summary 565 00_Hill3e_FM.indd   xxiii 3/9/12   2:11 PM
background image xxiv  Contents CHAPTER 22 Introduction to Oxygen and Carbon Dioxide Physiology  569 The Properties of Gases in Gas Mixtures and Aqueous  Solutions 570 Gases in the gas phase  570 Gases in aqueous solution  571 Diffusion of Gases  572 Gases diffuse far more readily through gas phases than through  aqueous solutions  574 Gas molecules that combine chemically with other molecules  cease to contribute to the gas partial pressure  574 BOX 22.1    Diffusion through Tissues Can Meet O 2   Requirements over Distances of Only 1  Millimeter or Less  575 Convective Transport of Gases: Bulk Flow  575 BOX 22.2    Induction of Internal Flow by Ambient  Currents 576 Gas transport in animals often occurs by alternating convection  and diffusion  576 The Oxygen Cascade  577
Expressing the Amounts and Partial Pressures of 
Gases in Other Units  578 The Contrasting Physical Properties of Air and  Water 579 Respiratory Environments  580 CHAPTER 23 External Respiration: The Physiology of Breathing  583 Fundamental Concepts of External Respiration  584
Principles of Gas Exchange by Active Ventilation  585
The O 2  partial pressure in blood leaving a breathing organ  depends on the spatial relation between the flow of the blood 
and the flow of the air or water  585
The relative changes in the partial pressures of O 2  and CO 2   depend dramatically on whether air or water is breathed  587 Introduction to Vertebrate Breathing  588
Breathing by Fish  590
Gill ventilation is usually driven by buccal–opercular  pumping 592 Many fish use ram ventilation on occasion, and some use it all  the time  593 Decreased O 2  and exercise are the major stimuli for increased  ventilation in fish  593 Several hundred species of bony fish are able to breathe air  593 Breathing by Amphibians  594 Gills, lungs, and skin are used in various combinations to  achieve gas exchange  595 Breathing by Reptiles Other than Birds  596
Breathing by Mammals  597
The total lung volume is employed in different ways in different  sorts of breathing  598 The gas in the final airways differs from atmospheric air in  composition and is motionless  599 The power for ventilation is developed by the diaphragm and  the intercostal and abdominal muscles  599 The control of ventilation  600 BOX 23.1  Low O 2 : Detection and Response  601 BOX 23.2     Mammals at High Altitude (with Notes on  High-Flying Birds)  602 In species of different sizes, lung volume tends to be a constant  proportion of body size, but breathing frequency varies 
allometrically 604
Pulmonary surfactant keeps the alveoli from collapsing  604 Breathing by Birds  605 Ventilation is by bellows action  606 Air flows unidirectionally through the parabronchi  606 The gas-exchange system is cross-current  608 BOX 23.3    Bird Development: Filling the  Lungs with Air Before Hatching  608 PART V • Oxygen, Carbon Dioxide,  and Internal Transport 00_Hill3e_FM.indd   xxiv 3/9/12   2:11 PM
background image Contents  xxv Breathing by Aquatic Invertebrates and Allied  Groups 608 Molluscs exemplify an exceptional diversity of breathing organs  built on a common plan  608 Decapod crustaceans include many important water breathers  and some air breathers  610 Breathing by Insects and Other Tracheate  Arthropods 611 BOX 23.4    The Book Lungs of Arachnids  612 Diffusion is a key mechanism of gas transport through the  tracheal system  612 Some insects employ conspicuous ventilation  613 Microscopic ventilation is far more common than believed even  a decade ago  614 Control of breathing  614 Aquatic insects breathe sometimes from the water, sometimes  from the atmosphere, and sometimes from both  615 CHAPTER 24 Transport of Oxygen and Carbon Dioxide  in Body Fluids (with an Introduction to  Acid-Base Physiology)  617 The Chemical Properties and Distributions of the  Respiratory Pigments  618 BOX 24.1    Absorption Spectra of Respiratory  Pigments 619 Hemoglobins contain heme and are the most widespread  respiratory pigments  619 BOX 24.2    Blood Cells and Their Production  622 Copper-based hemocyanins occur in many arthropods and  molluscs 622 Chlorocruorins resemble hemoglobins and occur in certain  annelids 623 Iron-based hemerythrins do not contain heme and occur in  three or four phyla  623 The O 2 -Binding Characteristics of Respiratory  Pigments 623 Human O 2  transport provides an instructive case study  624 A set of general principles helps elucidate O 2  transport by  respiratory pigments  627 The shape of the oxygen equilibrium curve depends on O 2 - binding site cooperativity  627 Respiratory pigments exhibit a wide range of affinities for O 2  628 The Bohr effect: Oxygen affinity depends on the partial pressure  of CO 2  and the pH  629 The Root effect: In unusual cases, CO 2  and pH dramatically affect  the oxygen-carrying capacity of the respiratory pigment  631 Thermal effects: Oxygen affinity depends on tissue  temperature 631 Organic modulators often exert chronic effects on oxygen  affinity 631 BOX 24.3    The Challenges of Regional Hypothermia  and the Resurrection of Mammoth  Hemoglobin 632 Inorganic ions may also act as modulators of respiratory  pigments 633 The Functions of Respiratory Pigments in Animals  633 BOX 24.4    Heme-Containing Globins in Intracellular  Function: Myoglobin Regulatory and  Protective Roles, Neuroglobins, and  Cytoglobins 634 Patterns of circulatory O 2  transport: The mammalian model is  common but not universal  635 Respiratory pigments within a single individual often display  differences in O 2  affinity that aid successful O 2  transport  636 Evolutionary adaptation: Respiratory pigments are molecules  positioned directly at the interface between animal and 
environment 636
The respiratory-pigment physiology of individuals undergoes  acclimation and acclimatization  637 Icefish live without hemoglobin  638 Carbon Dioxide Transport  638 BOX 24.5    Blood and Circulation in Mammals at High  Altitude 639 The extent of bicarbonate formation depends on blood  buffers 640 Carbon dioxide transport is interpreted by use of carbon dioxide  equilibrium curves  640 The Haldane effect: The carbon dioxide equilibrium curve  depends on blood oxygenation  641 Critical details of vertebrate CO 2  transport depend on carbonic  anhydrase and anion transporters  642 Acid–Base Physiology  643 Acid–base regulation involves excretion or retention of chemical  forms affecting H +  concentration  644 Disturbances of acid–base regulation fall into respiratory and  metabolic categories  644 CHAPTER 25 Circulation 647 Hearts 648 The heart as a pump: The action of a heart can be analyzed in  terms of the physics of pumping  649 The circulation must deliver O 2  to the myocardium  649 The electrical impulses for heart contraction may originate in  muscle cells or neurons  650 A heart produces an electrical signature, the  electrocardiogram 653 Heart action is modulated by hormonal, nervous, and intrinsic  controls 653 Principles of Pressure, Resistance, and Flow in  Vascular Systems  655 00_Hill3e_FM.indd   xxv 3/9/12   2:11 PM
background image xxvi  Contents The rate of blood flow depends on differences in blood pressure  and on vascular resistance  656 The dissipation of energy: Pressure and flow turn to heat during  circulation of the blood  657 Circulation in Mammals and Birds  658 The circulatory system is closed  658 Each part of the systemic vascular system has distinctive  anatomical and functional features  658 Mammals and birds have a high-pressure systemic circuit  660 Fluid undergoes complex patterns of exchange across the walls  of systemic capillaries  662 The pulmonary circuit is a comparatively low-pressure system  that helps keep the lungs “dry”  662 During exercise, blood flow is increased by orchestrated changes  in cardiac output and vascular resistance  663 Species have evolved differences in their circulatory  physiology 663 Circulation in Fish  664 The circulatory plans of fish with air-breathing organs (ABOs)  pose unresolved questions  666 Lungfish have specializations to promote separation of  oxygenated and deoxygenated blood  666 Circulation in Amphibians and in Reptiles Other than  Birds 668 BOX 25.1    An Incompletely Divided Central Circulation  Can Potentially Be an Advantage for  Intermittent Breathers  669 Concluding Comments on Vertebrates  670
Invertebrates with Closed Circulatory Systems  670
BOX 25.2    Bearing the Burden of Athleticism, Sort of:  A Synthesis of Cephalopod O 2   Transport 672 Invertebrates with Open Circulatory Systems  672 The crustacean circulatory system provides an example of an  open system  673 Open systems are functionally different from closed systems but  may be equal in critical ways  674 BOX 25.3    Circulation and O 2 : Lessons from the  Insect World  675 CHAPTER 26 Oxygen, Carbon Dioxide, and  Internal Transport  at Work:   Diving by Marine Mammals  679 Diving Feats and Behavior  679
Types of Dives and the Importance of Method  682
Physiology: The Big Picture  682
The Oxygen Stores of Divers  683
The blood O 2  store tends to be large in diving mammals  683 Diving mammals have high myoglobin concentrations and large  myoglobin-bound O 2  stores  683 Diving mammals vary in their use of the lungs as an O 2 store 684 Total O 2  stores never permit dives of maximum duration to be  fully aerobic  685 Circulatory Adjustments during Dives  685 Regional vasoconstriction: Much of a diving mammal’s  body is cut off from blood flow during forced or protracted 
dives 686
Diving bradycardia matches cardiac output to the circulatory  task 687 Cardiovascular responses are graded in freely diving  animals 687 BOX 26.1    The Evolution of Vertebrate Cardiac and  Vascular Responses to Asphyxia  688 Red blood cells are removed from the blood between dive  sequences in some seals  689 Metabolism during Dives  689 The body becomes metabolically subdivided  during forced or protracted dives  689 Metabolic limits on dive duration are determined by O 2  supplies,  by rates of metabolic O 2  use and lactic acid production, and by  tissue tolerances  690 The Aerobic Dive Limit: One of Physiology’s  Key Benchmarks for Understanding Diving  Behavior 691 Marine mammals exploit multiple means of reducing their  metabolic costs while under water  693 Decompression Sickness  694 Human decompression sickness is usually caused  by N 2  absorption from a compressed-air source  694 Breath-hold dives must be repeated many times to cause  decompression sickness in humans  694 Marine mammals have been thought—perhaps erroneously— to avoid decompression sickness during deep dives by alveolar 
collapse 694
Decompression sickness is an unresolved phenomenon  695 A Possible Advantage for Pulmonary O 2 Sequestration in Deep Dives  695 00_Hill3e_FM.indd   xxvi 3/9/12   2:11 PM
background image Contents  xxvii CHAPTER 27 Water and Salt Physiology: Introduction and Mechanisms  699 The Importance of Animal Body Fluids  700
The Relationships among Body Fluids  701
Types of Regulation and Conformity  701
Natural Aquatic Environments  703
Natural Terrestrial Environments  705
Organs of Blood Regulation  707
The osmotic U/P ratio is an index of the action of the kidneys in  osmotic regulation  707 The effects of kidney function on volume regulation depend on  the amount of urine produced  708 The effects of kidney function on ionic regulation depend on  ionic U/P ratios  709 Food and Drinking Water  709 Salty drinking water may not provide H 2 O 709 Plants and algae with salty tissue fluids pose challenges for  herbivores 710 Air-dried foods contain water  710 Protein-rich foods can be dehydrating for terrestrial animals  710 Metabolic Water  710 Metabolic water matters most in animals that conserve water  effectively 711 BOX 27.1    Net Metabolic Water Gain in Kangaroo  Rats 711 Cell-Volume Regulation  712
From Osmolytes to Compatible Solutes: Terms and 
Concepts 714 CHAPTER 28 Water and Salt Physiology  of Animals in Their Environments  717 Animals in Freshwater  717 Passive water and ion exchanges: Freshwater animals tend to  gain water by osmosis and lose major ions by diffusion  718 Most types of freshwater animals share similar regulatory  mechanisms 719 BOX 28.1    Fish Mitochondria-Rich Cells and Their  Diversity 723 A few types of freshwater animals exhibit exceptional patterns  of regulation  723 Why do most freshwater animals make dilute urine?  724 Animals in the Ocean  724 Most marine invertebrates are isosmotic to seawater  725 Hagfish are the only vertebrates with blood inorganic ion  concentrations that make them isosmotic to seawater  725 The marine teleost fish are markedly hyposmotic to  seawater 725 BOX 28.2    Where Were Vertebrates at Their  Start? 726 BOX 28.3    Epithelial NaCl Secretion in Gills, Salt  Glands, and Rectal Glands  728 Some arthropods of saline waters are hyposmotic regulators  729 Marine reptiles (including birds) and mammals are also  hyposmotic regulators  729 Marine elasmobranch fish are hyperosmotic but hypoionic to  seawater 731 BOX 28.4    The Evolution of Urea Synthesis in  Vertebrates 732 Animals That Face Changes in Salinity  733 Migratory fish and other euryhaline fish are dramatic and  scientifically important examples of hyper-hyposmotic 
regulators 734
Animals undergo change in all time frames in their relations to  ambient salinity  735 Responses to Drying of the Habitat in Aquatic  Animals 736 Animals on Land: Fundamental Physiological  Principles 737 BOX 28.5    Anhydrobiosis: Life as Nothing More  than a Morphological State  737 A low integumentary permeability to water is a key to reducing  evaporative water loss on land  738 Respiratory evaporative water loss depends on the function of  the breathing organs and the rate of metabolism  739 PART VI • Water, Salts, and Excretion 00_Hill3e_FM.indd   xxvii 3/9/12   2:11 PM

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School: University of Miami
Department: Communications
Course: Comparative Physiology
Professor: Dana Krempels
Term: Fall 2016
Tags:
Name: Comp. Phys. Textbook
Description: This is the textbook for the class.
Uploaded: 09/19/2016
985 Pages 181 Views 144 Unlocks
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