Animal Physiology, 3rd Ed

985 Pages·2012·53.37 MB·English

by  Richard W. Hill, Gordon A. Wyse

Themes in the Study of Animal Physiology Theme An Example of the Theme in Action See Pages
Listed are 15 overarching themes that reappear throughout the study of animal in importance. To help explain each theme, an illustrative example is presented
physiology. Some of the listed themes overlap with, or even encompass, oth- in the second column of the table. Further examples are on the pages listed in The Dynamic State of Body Constituents: Averaged over the course of an ordinary 24-h day, an adult person is like- 11–12
ers; they are not intended to be mutually exclusive or, in all cases, equivalent the third column (italic listing elaborates the featured example). Great quantities of many of the key constituents ly to process more than 2 kg of adenosine triphosphate (ATP) each hour, 183–184
of the body are added and subtracted every day synthesizing that amount of ATP from adenosine diphosphate (ADP) 378
in many animals under many conditions. Thus the and, with only a short delay, breaking it back down to ADP. To synthe- 699
Theme An Example of the Theme in Action See Pages constituents of the body—far from being static—are size the ATP, the person—during each hour—will use about 20 liters of
continuously in a dynamic state of flux. This is true oxygen (O2) that he or she takes up from the atmosphere. During a 24-h 743–744
The Study of Function: When physiologists study muscle, one of their goals is to understand how 114 (Fig. 5.12) even though additions and subtractions are often day, the oxygen used will combine with almost 100 g (a fifth of a pound) (Fig. 28.21)
A
funnicmtiaoln p. hTyhsaito ilso,g iyt iiss tthhee ssttuuddyy ooff hhooww tahneiimr caelsll s tahre epmropteloinyse din i nm luosccolme coetlilosn a, rhee aabrtle c oton tdreavcteiloonp, more octhhaenri caaclt ifvoirtcies., which 652067 (Fig. 203.52)2) creolnactievnetlrya tbioanlasn (cae dp,h reensoumltienngo nin t reerlmateidve hlyom ceoonsstatasinst) . oabf ohuytd 8ro00g emni alltiolimtesrs t hoaf tw haatveer. bTeheins rweamteorv iesd a fdrdomed ftooo tdh me booledcyu lfelus,i dfosr. ming
and organs operate. 687 (Fig. 26.9)
Multiple Forms of Key Molecules: The cell membranes of all animals are composed principally of lipid mol- 34 (Fig. 2.3)
Animals have often evolved multiple molecular ecules. Physiologists have found, however, that the membranes of all ani- 242–243
Integration of the Sciences: To understand how animals employ odors to orient their movements, 6 (Fig. 1.2) forms (sometimes called isoforms) of particular mals are not composed of chemically identical lipid molecules. Instead, (Fig. 10.19)
Physiologists often find that they must integrate physiologists study the chemical structural differences between molecules 60 (Fig. 2.28) proteins or other sorts of molecules. Physiologists multiple molecular forms of lipids are employed by different animals 537
knowledge of mathematics, chemistry, or physics that attract or repel, and they mathematically describe the physics of how 165 (Fig. 7.3) hypothesize that when two species or two tissues living under different circumstances. Cold-water fish species, for instance, 620 (Fig. 24.2)
qwuitehs tkionnosw. Plehdygseio olof gbyio ilso goyn et oo fa nthsew mero ismt pinotretgarnatt ive wthien odlsf aocrt owryat oer gcaunrsr eonft sa ntrimanaslps.ort odor molecules from odor sources to 377 efoxrhmibsi ta dreif ofefrten ts mpeoclieacluizleadr ftor fmusn cotfi oan m ino ltehceu lsep, et-he cleosnss ltirkueclty tthoe ihra credlel nm aetm lobwra nteems puesirnagtu mreosl ethcuanla trh feo rmmosl eocfu lliapri dfos rtmhast s ayrne- 640 (Fig. 24.20)
branches of biology. 650 (Fig. 25.2) cific settings in which the animals live or the tissues thesized by warm-water species.
function.
Emphasis on Quantitative Methods: Starting in ancient Roman times, people thought that the dromedary camel 209 (Box 9.1)
Physiologists quantify the properties of animals as could carry enough water in its rumen to explain its unusual ability to live 212 (Fig. 9.7) Phenotypic Plasticity: Animals that eat only occasionally, such as pythons, often alternate 15 (Fig. 1.5)
c
oar rmefauklley parse pdoicstsioibnlse. as they seek to test hypotheses wthiet hcoaumte dl rriunmkienng .r aWthheern t hpahny sjuioslto sgpisetask qinuagn qtiufiaeldi ttahteiv aemlyo aubnotu ot fi tw, hatoewr einv er, 2 3 (4F–i2g3. 51 0 .8) Aphne innodtyivpide uina lr aenspimonasl ei st oo fcthenan agbelse itno tchhea npgaret ictsu lar bweetewkese, nth tewiro i nintetesstitninaal lt rpahcet niso tpyhpyesi.c Wallhye snm thaelly, ahnadv eit nhoats h paodo arl ym deaelv feol-r 7909 –(9T2able 3.1)
othldey i dfoeua ntod mthaakt et hsernes ew. aNse ngoatt innega trhlye eonldo uidgeha w haetlepre idn ltehaed rtuom uennd eforsrt tahned - 799–800 ctiicruculamr setnavnicreosn umnednetr). wThics ha biti liist yli voifn agn (ien.gd.i,v iitds upaalr - opf ethde m inotleesctuinlarl tmraeccth eannliasrmges fgorre atblyso, rabnidn gth feo oindt.e Astfitnearl at rmaceta el,x tphree stsisessu es 157 (Box 6.2)
ing that camels do not store water to a greater degree than other mam- animal to adopt two or more phenotypes despite well-developed absorption mechanisms. 264 (Fig. 10.40)
555 (Fig. 21.7)
mals. Instead, they have excellent abilities to conserve water and endure having a fixed genotype is termed phenotypic
dehydration. plasticity.
The Tandem Goal of Understanding A number of animals, including fireflies and certain fish, produce light. 5–9 Interdependency of Function and Form: The kidney tubules of mammals not only produce the most concentrated 140 (Fig. 6.13)
Mechanism and Adaptation: When physiologists study light production in such animals, they try to 248–249 The function of a biological system typically cannot urine observed in vertebrates but also differ from other vertebrate kidney 259 (Fig. 10.35)
When physiologists study a process, they typically learn both how the animals make light and why they make it. (Fig. 10.26) be understood without knowledge of its structure, tubules in that they all have distinctive hairpin shapes. Physiologists have 370 (Fig. 14.10)
emphasize a two-part goal: They try to understand 455–456 and vice versa. s
dheopwennd tsh oant tthhee fhuanicrtpioina slt arubciltiutyre ,t ow phriochd ugcuei dheigs hthlye cuornincen (tarsa tiet dis ubreiinneg 649 (Fig. 25.1)
both the mechanism involved (i.e., how the process 735 (Fig. 28.12)
formed) to flow first in one direction and then in the opposite direction. 768 (Fig. 29.12)
is executed) and the potential adaptive significance of
760 (Fig. 29.5)
the process (i.e., how, if at all, it enhances evolu-
tionary fitness). Applicability of the Laws of Chemistry and Heat transfer through air follows different physical laws when the air is 230
Physics: still rather than moving; heat tends to move much more slowly through 493 (Fig. 18.8)
Animals must adhere to the laws of chemistry and still air than moving air. Animals cannot change such laws of physics. 576 (Box 22.2)
T
Toh eu nCdoermstpanadr athtiev ead Mapettivheo sdig: nificance of animal Winsheecnts p, thhyesyio fliongdi stthsa cto dmespearrte s apneicmieas ltse ansd d ciostnasnisttlyen rtellya teod h asv em garmeamt albsi lain- d 2161 4( F(Figig. .1 .51.81)3) pcohnysstircasi.n Stos,m beutti msoems cehtiemmeiss tarnyi manadls p ghayinsi casd avcatn atas ges Tthhee ayn scoemsteotrism oefs m canm amffaelcst ewvhoilcvhe dla fwur a. pTphlei ehsa tiors t hoef ma ,f uhrorwede vmera,m ams walh en 694–695
features, physiologists make extensive use of the ties to concentrate their urine. Desert species of mammals are typically 675 (Box 25.3) by evolving systems that capitalize on particular keep the layer of air next to the body relatively motionless. Heat transfer
comparative method, which is the examination of how able to make urine of higher concentration than nondesert mammals, and 688 (Box 26.1) chemical or physical principles. through that air is therefore slow, helping mammals retain internal heat
particular functions are carried out by related and desert insects are similarly superior to nondesert insects. These compara- when living in cold environments.
unrelated species living in similar and dissimilar tive observations provide evidence that the ability to make concentrated 743 (Fig. 28.20)
environments. urine is an advantage—favored by natural selection—in deserts. The Interdependency of Levels of When your physician strikes a tendon near your knee with a mallet, your 6 (Fig. 1.2)
Organization: leg straightens. For this response, electrical signals must travel along 116 (Box 5.2)
Phylogenetic Reconstruction: Although body temperature is the same as water temperature in most 27 An animal’s overall functional properties depend on nerve cells to the spinal cord and back. The rate of travel depends in part 199 (Fig. 8.12)
wp
cTluihroehto ysiiuop csinehisodr a ulgtreoisyergen ssidt—esia gttnisaonc dne iorf mdteirch c ptaoeohnlt nouehcsyvset o rgpo ulihaufnc yitmntfli o topghrnmde nore apserftpt naipioect-h dcnrytsae i oscyvooin eofnp lemosrovtongruop uitlcecthuaitreptiltoi iloenevns ss—,o. pi-ne- msgcthpoeaeneni rcdecneitoietfitaoncsiior dnorenei f,t e cietflohivesnavohs talf, tv rtreoeuedmgc atato iyearonmd’ ns lai spefnt,is e gsplsrehl haeoa tyw sfuc tsohir itfoemohoslwu o mwirg nb oai sinsnrgtomd s amt e hnhtpeaicese evyosnse ufatd oeftreohsren. u,ed ticn roed trcn itctsaohasitsuan iate o slstnlph. seFei.c m rWwioepmsael r yo kmp fni h-nfotiyhiswslheos,r u- ite 57273173 02( F(FBigiog. x.23 1.225081).4)47)) ohmc
lziesafo vo ntwilhietenoslac son iut tt iolsp asab fsrtr eiueo os repsfyigunsse aeltralesntynm iraie dzusnsae n dofatpudir toeogn eonrnacrgnd.stnaiteosno n lnsodet .df.ve uAAp ewnleln lcnoi tifdhihm oosoen rpuso,geo tna rle nethxadviopzne watltloh tsc ire ioit nsnfrfu og coonl erloflcaglttsterhai ynoane nri- d mstpFobhunroroaocr tmnh ph iae neeasrs nse tmto it atef fohntu slhtec elhoel ecysef u,pm ltenahxaercepl riplysnveoe rgetds o se secpbodtteerere litrtlrkwsostmi e. de etsIihse tn a etnoeaen f lfrtdts lhmihuo oete iinhd d nsl-eset prnc mbtateghainonotteshmhdn po isoenvo fnioerg nrtf ai i tnpymelhlagoa eepcr u p hctrtr eorh o nolpanleet.teg er Mkrp ivtmneaieoyess ul s cseic seneoceclsl uflt euhblmtashle arect rcwmoa epnelbrldteo ormn acpc encetsthelmr.ls teu.i - elasr, 53086– (3F0ig7. 19.4)
The Centrality of the Environment: Many specialists in high-altitude physiology argue that when lowland 53 (Fig. 2.22) The Crucial Importance of Control Although sheep and reindeer are born at cold times of year, newborns 50 (Fig. 2.19)
The specific environments in which animals have people travel to high altitudes, some of their typical responses are more 67 Mechanisms: receive no heat from their parents and must keep warm on their own or 252 (Box 10.2)
teivoonlavle pdr oanpder ltivese omf uthste baen cimonaslsid teor emda fkoer stehnes feu.nc-
whcniaoeor surm edldafiusd bol nenth o btatoe ne np vehrofeilsclvupieamf uli.enl .t hTaihgte has-leal lstthpiteu chdiauel miesntasvn ie rrmoensppmhoaensnsitzeses. tAthoc asctuo tcrhdhe i enhnguvlmyir,a otnhn emsrpee ni-sts 266394 (FBiogx. 2140.54)1) iI
tanhnnge iam ,o dmtadhloisetv irroi enmnqg eut,o ciahr mnea dnce oiccsnhamtarrrnsoy.li siTmnmhgese co fhcuooatrn ortiesrthompelr srom dtoheuvcacehtir anotn grfciu,s hbnmercsestti—arotanhstseo-, mditthi weisc. ohpTuarhlonedciys et msepsno hdrseas nqes seutsewi rdaeol sep kstrseholyacy et p tsiorsto befpoexer har ratcaiutepisvit:da Ift teeh tadeac alat tiet v npbaeritoregtdshy uh, csbetuauioptt n pn.lr oiPoetdr sbo.u epTcfehtoirero e cnc ob owninrthtrtrhoeo lnlw o hf en 24803 (Fig. 117.81)5)
diverse that they include controls of gene expres- neural thermal sensors detect cold, but its capacity to activate heat produc-
Body Size: The metabolic rate per gram of body weight is usually higher in small- 17 (Fig. 1.9) sion as well as those exerted by the nervous tion is turned off by chemical factors secreted by the placenta. The control
The physiological properties of related animal spe- bodied species than in related large-bodied ones. Because of this relation, 173 (Fig. 7.6) and endocrine systems—determine the relations mechanism remains in a turned-off state until a newborn is separated from
cies typically scale in mathematically consistent whenever two species of mammals of different body sizes—like mice and 285 (Fig. 11.9) between inputs and outputs in physiological sys- the placenta at birth. The cold environment is then able to stimulate rapid
ways with their body sizes. These relations are horses—are compared, the smaller species typically needs more food per 741 (Fig. 28.18) tems. They thereby crucially affect the functional heat production.
often nonproportional and thus termed allometric. gram of body weight than the larger one. properties of animals.
Themes in the Study of Animal Physiology Theme An Example of the Theme in Action See Pages
Listed are 15 overarching themes that reappear throughout the study of animal in importance. To help explain each theme, an illustrative example is presented
physiology. Some of the listed themes overlap with, or even encompass, oth- in the second column of the table. Further examples are on the pages listed in The Dynamic State of Body Constituents: Averaged over the course of an ordinary 24-h day, an adult person is like- 11–12
ers; they are not intended to be mutually exclusive or, in all cases, equivalent the third column (italic listing elaborates the featured example). Great quantities of many of the key constituents ly to process more than 2 kg of adenosine triphosphate (ATP) each hour, 183–184
of the body are added and subtracted every day synthesizing that amount of ATP from adenosine diphosphate (ADP) 378
in many animals under many conditions. Thus the and, with only a short delay, breaking it back down to ADP. To synthe- 699
Theme An Example of the Theme in Action See Pages constituents of the body—far from being static—are size the ATP, the person—during each hour—will use about 20 liters of
continuously in a dynamic state of flux. This is true oxygen (O2) that he or she takes up from the atmosphere. During a 24-h 743–744
The Study of Function: When physiologists study muscle, one of their goals is to understand how 114 (Fig. 5.12) even though additions and subtractions are often day, the oxygen used will combine with almost 100 g (a fifth of a pound) (Fig. 28.21)
A
funnicmtiaoln p. hTyhsaito ilso,g iyt iiss tthhee ssttuuddyy ooff hhooww tahneiimr caelsll s tahre epmropteloinyse din i nm luosccolme coetlilosn a, rhee aabrtle c oton tdreavcteiloonp, more octhhaenri caaclt ifvoirtcies., which 652067 (Fig. 203.52)2) creolnactievnetlrya tbioanlasn (cae dp,h reensoumltienngo nin t reerlmateidve hlyom ceoonsstatasinst) . oabf ohuytd 8ro00g emni alltiolimtesrs t hoaf tw haatveer. bTeheins rweamteorv iesd a fdrdomed ftooo tdh me booledcyu lfelus,i dfosr. ming
and organs operate. 687 (Fig. 26.9)
Multiple Forms of Key Molecules: The cell membranes of all animals are composed principally of lipid mol- 34 (Fig. 2.3)
Animals have often evolved multiple molecular ecules. Physiologists have found, however, that the membranes of all ani- 242–243
Integration of the Sciences: To understand how animals employ odors to orient their movements, 6 (Fig. 1.2) forms (sometimes called isoforms) of particular mals are not composed of chemically identical lipid molecules. Instead, (Fig. 10.19)
Physiologists often find that they must integrate physiologists study the chemical structural differences between molecules 60 (Fig. 2.28) proteins or other sorts of molecules. Physiologists multiple molecular forms of lipids are employed by different animals 537
knowledge of mathematics, chemistry, or physics that attract or repel, and they mathematically describe the physics of how 165 (Fig. 7.3) hypothesize that when two species or two tissues living under different circumstances. Cold-water fish species, for instance, 620 (Fig. 24.2)
qwuitehs tkionnosw. Plehdygseio olof gbyio ilso goyn et oo fa nthsew mero ismt pinotretgarnatt ive wthien odlsf aocrt owryat oer gcaunrsr eonft sa ntrimanaslps.ort odor molecules from odor sources to 377 efoxrhmibsi ta dreif ofefrten ts mpeoclieacluizleadr ftor fmusn cotfi oan m ino ltehceu lsep, et-he cleosnss ltirkueclty tthoe ihra credlel nm aetm lobwra nteems puesirnagtu mreosl ethcuanla trh feo rmmosl eocfu lliapri dfos rtmhast s ayrne- 640 (Fig. 24.20)
branches of biology. 650 (Fig. 25.2) cific settings in which the animals live or the tissues thesized by warm-water species.
function.
Emphasis on Quantitative Methods: Starting in ancient Roman times, people thought that the dromedary camel 209 (Box 9.1)
Physiologists quantify the properties of animals as could carry enough water in its rumen to explain its unusual ability to live 212 (Fig. 9.7) Phenotypic Plasticity: Animals that eat only occasionally, such as pythons, often alternate 15 (Fig. 1.5)
c
oar rmefauklley parse pdoicstsioibnlse. as they seek to test hypotheses wthiet hcoaumte dl rriunmkienng .r aWthheern t hpahny sjuioslto sgpisetask qinuagn qtiufiaeldi ttahteiv aemlyo aubnotu ot fi tw, hatoewr einv er, 2 3 (4F–i2g3. 51 0 .8) Aphne innodtyivpide uina lr aenspimonasl ei st oo fcthenan agbelse itno tchhea npgaret ictsu lar bweetewkese, nth tewiro i nintetesstitninaal lt rpahcet niso tpyhpyesi.c Wallhye snm thaelly, ahnadv eit nhoats h paodo arl ym deaelv feol-r 7909 –(9T2able 3.1)
othldey i dfoeua ntod mthaakt et hsernes ew. aNse ngoatt innega trhlye eonldo uidgeha w haetlepre idn ltehaed rtuom uennd eforsrt tahned - 799–800 ctiicruculamr setnavnicreosn umnednetr). wThics ha biti liist yli voifn agn (ien.gd.i,v iitds upaalr - opf ethde m inotleesctuinlarl tmraeccth eannliasrmges fgorre atblyso, rabnidn gth feo oindt.e Astfitnearl at rmaceta el,x tphree stsisessu es 157 (Box 6.2)
ing that camels do not store water to a greater degree than other mam- animal to adopt two or more phenotypes despite well-developed absorption mechanisms. 264 (Fig. 10.40)
555 (Fig. 21.7)
mals. Instead, they have excellent abilities to conserve water and endure having a fixed genotype is termed phenotypic
dehydration. plasticity.
The Tandem Goal of Understanding A number of animals, including fireflies and certain fish, produce light. 5–9 Interdependency of Function and Form: The kidney tubules of mammals not only produce the most concentrated 140 (Fig. 6.13)
Mechanism and Adaptation: When physiologists study light production in such animals, they try to 248–249 The function of a biological system typically cannot urine observed in vertebrates but also differ from other vertebrate kidney 259 (Fig. 10.35)
When physiologists study a process, they typically learn both how the animals make light and why they make it. (Fig. 10.26) be understood without knowledge of its structure, tubules in that they all have distinctive hairpin shapes. Physiologists have 370 (Fig. 14.10)
emphasize a two-part goal: They try to understand 455–456 and vice versa. s
dheopwennd tsh oant tthhee fhuanicrtpioina slt arubciltiutyre ,t ow phriochd ugcuei dheigs hthlye cuornincen (tarsa tiet dis ubreiinneg 649 (Fig. 25.1)
both the mechanism involved (i.e., how the process 735 (Fig. 28.12)
formed) to flow first in one direction and then in the opposite direction. 768 (Fig. 29.12)
is executed) and the potential adaptive significance of
760 (Fig. 29.5)
the process (i.e., how, if at all, it enhances evolu-
tionary fitness). Applicability of the Laws of Chemistry and Heat transfer through air follows different physical laws when the air is 230
Physics: still rather than moving; heat tends to move much more slowly through 493 (Fig. 18.8)
Animals must adhere to the laws of chemistry and still air than moving air. Animals cannot change such laws of physics. 576 (Box 22.2)
T
Toh eu nCdoermstpanadr athtiev ead Mapettivheo sdig: nificance of animal Winsheecnts p, thhyesyio fliongdi stthsa cto dmespearrte s apneicmieas ltse ansd d ciostnasnisttlyen rtellya teod h asv em garmeamt albsi lain- d 2161 4( F(Figig. .1 .51.81)3) pcohnysstircasi.n Stos,m beutti msoems cehtiemmeiss tarnyi manadls p ghayinsi casd avcatn atas ges Tthhee ayn scoemsteotrism oefs m canm amffaelcst ewvhoilcvhe dla fwur a. pTphlei ehsa tiors t hoef ma ,f uhrorwede vmera,m ams walh en 694–695
features, physiologists make extensive use of the ties to concentrate their urine. Desert species of mammals are typically 675 (Box 25.3) by evolving systems that capitalize on particular keep the layer of air next to the body relatively motionless. Heat transfer
comparative method, which is the examination of how able to make urine of higher concentration than nondesert mammals, and 688 (Box 26.1) chemical or physical principles. through that air is therefore slow, helping mammals retain internal heat
particular functions are carried out by related and desert insects are similarly superior to nondesert insects. These compara- when living in cold environments.
unrelated species living in similar and dissimilar tive observations provide evidence that the ability to make concentrated 743 (Fig. 28.20)
environments. urine is an advantage—favored by natural selection—in deserts. The Interdependency of Levels of When your physician strikes a tendon near your knee with a mallet, your 6 (Fig. 1.2)
Organization: leg straightens. For this response, electrical signals must travel along 116 (Box 5.2)
Phylogenetic Reconstruction: Although body temperature is the same as water temperature in most 27 An animal’s overall functional properties depend on nerve cells to the spinal cord and back. The rate of travel depends in part 199 (Fig. 8.12)
wp
cTluihroehto ysiiuop csinehisodr a ulgtreoisyergen ssidt—esia gttnisaonc dne iorf mdteirch c ptaoeohnlt nouehcsyvset o rgpo ulihaufnc yitmntfli o topghrnmde nore apserftpt naipioect-h dcnrytsae i oscyvooin eofnp lemosrovtongruop uitlcecthuaitreptiltoi iloenevns ss—,o. pi-ne- msgcthpoeaeneni rcdecneitoietfitaoncsiior dnorenei f,t e cietflohivesnavohs talf, tv rtreoeuedmgc atato iyearonmd’ ns lai spefnt,is e gsplsrehl haeoa tyw sfuc tsohir itfoemohoslwu o mwirg nb oai sinsnrgtomd s amt e hnhtpeaicese evyosnse ufatd oeftreohsren. u,ed ticn roed trcn itctsaohasitsuan iate o slstnlph. seFei.c m rWwioepmsael r yo kmp fni h-nfotiyhiswslheos,r u- ite 57273173 02( F(FBigiog. x.23 1.225081).4)47)) ohmc
lziesafo vo ntwilhietenoslac son iut tt iolsp asab fsrtr eiueo os repsfyigunsse aeltralesntynm iraie dzusnsae n dofatpudir toeogn eonrnacrgnd.stnaiteosno n lnsodet .df.ve uAAp ewnleln lcnoi tifdhihm oosoen rpuso,geo tna rle nethxadviopzne watltloh tsc ire ioit nsnfrfu og coonl erloflcaglttsterhai ynoane nri- d mstpFobhunroroaocr tmnh ph iae neeasrs nse tmto it atef fohntu slhtec elhoel ecysef u,pm ltenahxaercepl riplysnveoe rgetds o se secpbodtteerere litrtlrkwsostmi e. de etsIihse tn a etnoeaen f lfrtdts lhmihuo oete iinhd d nsl-eset prnc mbtateghainonotteshmhdn po isoenvo fnioerg nrtf ai i tnpymelhlagoa eepcr u p hctrtr eorh o nolpanleet.teg er Mkrp ivtmneaieoyess ul s cseic seneoceclsl uflt euhblmtashle arect rcwmoa epnelbrldteo ormn acpc encetsthelmr.ls teu.i - elasr, 53086– (3F0ig7. 19.4)
The Centrality of the Environment: Many specialists in high-altitude physiology argue that when lowland 53 (Fig. 2.22) The Crucial Importance of Control Although sheep and reindeer are born at cold times of year, newborns 50 (Fig. 2.19)
The specific environments in which animals have people travel to high altitudes, some of their typical responses are more 67 Mechanisms: receive no heat from their parents and must keep warm on their own or 252 (Box 10.2)
teivoonlavle pdr oanpder ltivese omf uthste baen cimonaslsid teor emda fkoer stehnes feu.nc-
whcniaoeor surm edldafiusd bol nenth o btatoe ne np vehrofeilsclvupieamf uli.enl .t hTaihgte has-leal lstthpiteu chdiauel miesntasvn ie rrmoensppmhoaensnsitzeses. tAthoc asctuo tcrhdhe i enhnguvlmyir,a otnhn emsrpee ni-sts 266394 (FBiogx. 2140.54)1) iI
tanhnnge iam ,o dmtadhloisetv irroi enmnqg eut,o ciahr mnea dnce oiccsnhamtarrrnsoy.li siTmnmhgese co fhcuooatrn ortiesrthompelr srom dtoheuvcacehtir anotn grfciu,s hbnmercsestti—arotanhstseo-, mditthi weisc. ohpTuarhlonedciys et msepsno hdrseas nqes seutsewi rdaeol sep kstrseholyacy et p tsiorsto befpoexer har ratcaiutepisvit:da Ift teeh tadeac alat tiet v npbaeritoregtdshy uh, csbetuauioptt n pn.lr oiPoetdr sbo.u epTcfehtoirero e cnc ob owninrthtrtrhoeo lnlw o hf en 24803 (Fig. 117.81)5)
diverse that they include controls of gene expres- neural thermal sensors detect cold, but its capacity to activate heat produc-
Body Size: The metabolic rate per gram of body weight is usually higher in small- 17 (Fig. 1.9) sion as well as those exerted by the nervous tion is turned off by chemical factors secreted by the placenta. The control
The physiological properties of related animal spe- bodied species than in related large-bodied ones. Because of this relation, 173 (Fig. 7.6) and endocrine systems—determine the relations mechanism remains in a turned-off state until a newborn is separated from
cies typically scale in mathematically consistent whenever two species of mammals of different body sizes—like mice and 285 (Fig. 11.9) between inputs and outputs in physiological sys- the placenta at birth. The cold environment is then able to stimulate rapid
ways with their body sizes. These relations are horses—are compared, the smaller species typically needs more food per 741 (Fig. 28.18) tems. They thereby crucially affect the functional heat production.
often nonproportional and thus termed allometric. gram of body weight than the larger one. properties of animals.
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ANIMAL PHYSIOLOGY
THIRD EDITION
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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
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About the Cover
One of the central themes of this book is the intimate relationship between animals
and their environments. The gemsbok oryx (Oryx gazella) provides an iconic example.
Gemsboks succeed in one of Earth’s most demanding settings—the hot, dry deserts of
Africa—because of their evolution of a variety of specialized behavioral, morphological,
and physiological attributes. Among the truly wild large mammals, the three species of
oryxes that are scientifically well known probably represent the pinnacle of evolution
in their ability to survive in such deserts. Oryxes are discussed in depth in Chapter 30.
Animal Physiology, Third Edition
Copyright © 2012. All rights reserved.
This book may not be reproduced in whole or in part without permission from the publisher.
Address editorial correspondence and orders to:
Sinauer Associates, 23 Plumtree Road, Sunderland, MA 01375 U.S.A.
FAX: [phone]
Email:
To Sue, Dave, and Chrissie, from RWH
To Mary, from GAW
To Anita and Andy, from MA
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Preface
Thomas Kuhn wrote that a textbook is principally a means of com- the lives of animals in their natural habitats. Second, we devote five
municating to students the paradigms of their time. We have had entire chapters (our “At Work” chapters) to in-depth explorations
three principal goals in preparing the content of this book. One, of how physiologists do their work; in these chapters we break out
in accord with Kuhn’s dictum, has been to articulate the central of the usual textbook mold to discuss exciting topics—such as the
paradigms of contemporary animal physiology. A second content diving physiology of marine mammals—with emphasis on experi-
goal has been to provide our readers with a source of both lucid ments, theory maturation, integration of physiological systems, and
explanations of physiological concepts and accurate information prospects for future research. Third, we include many photographs
about physiological systems. Our third content goal has been to and drawings of animals throughout the book to remind readers of
draw attention to the cutting edges of physiological science, the the animals we discuss. Fourth, entirely new to this edition, we have
places where the onward progress of research is challenging old started a program of inviting specialists to contribute expert Guest
paradigms and potentially creating footholds for new ones. Boxes on emerging topics that expand the book’s subject content.
We have also had goals for presentation. Most visibly, we have With our aspirations being as numerous as we have described,
combined our words with an ambitious, informative art program. we have put a great deal of effort into balancing competing demands
More fundamentally, we have strived to take advantage of all the for space. The product is a complete physiology textbook that in
assets of traditional bookmaking to achieve a book that—through one volume will meet the requirements of a diversity of one- or
constant integration of the full suite of pedgogically relevant ele- two-semester courses in animal function. Our intended audience
ments—is a first-rate learning tool. Many sorts of professionals is sophomores through beginning graduate students. To make
have important contributions to make for a book to be excellent. the book accessible to as wide an audience as possible, we have
Thus many sorts of professionals have traditionally found personal included both a glossary of nearly 1200 terms and 11 appendices
fulfillment by engaging in the cooperative, synergistic production on important background concepts.
of books. The authors listed on the cover are just the tip of the Our approach to the writing has been to work from the original
iceberg. A book’s art program depends on scientific illustrators. scientific literature and obtain extensive peer review. Another aspect
Coordination between the art and the text—a key to the success of our approach is that we have opted for the pedagogical consis-
of any textbook—depends on the editorial expertise of the book’s tency of a book written by just three principal authors. Margaret
editor. An attractive science text needs to be designed and physi- Anderson wrote Chapters 16, 20, and 21, and Gordon Wyse wrote
cally executed by talented people who combine scientific acumen Chapters 12–15, 18, and 19. Richard Hill wrote Chapters 1–11, 17,
with artistic sensibility. To the degree that the presentation of and 22–30. David S. Garbe, Scott A. Huettel, Matthew S. Kayser,
the material in this book achieves success, the reason is that it is Kenneth J. Lohmann, and Margaret McFall-Ngai wrote Guest
the creative product of a team of at least a dozen people playing Boxes. Matthew S. Kayser and Gordon Fain assisted with topic
diverse, mutually reinforcing roles. One of our goals has been to development in certain parts of the principal text.
take advantage of this time-proven model to provide students with
a superior text. NEW TO THIS EDITION: As in other editions, our two central goals
In these pages, we consistently and deliberately address animal for this edition were to update content and enhance pedagogical
physiology as a discipline integrated with other disciplines in biol- effectiveness. To these ends, we have reconsidered every sentence
ogy—especially genetics, molecular biology, evolutionary biology, and every element of the art program. Of the 690 figures and tables
and ecology. We also consistently emphasize the roles of physiology in this edition, over 60 are either new or greatly enhanced. Chapter
throughout the life cycle of an animal by discussing physiologi- 14, on sensory processes, has been entirely rewritten. We have also
cal development and by examining animal function during such added a new chapter (Chapter 4) on physiological development and
important life-cycle processes as exercise, long-distance migration, epigenetics. Other chapters that have received exceptional attention
seasonal rhythms, and accommodation to severe conditions (we are: Chapter 5 (transport of solutes and water), Chapter 8 (aerobic
generally omit pathology and parasitism, however). Although we give and anaerobic metabolism), Chapter 9 (activity energetics), and
particular attention to mammals, we make a point of recognizing Chapter 29 (kidney physiology, edited throughout to emphasize
the other vertebrate groups and at least the arthropods and molluscs plasma regulation). The book now includes Guest Boxes on functional
among invertebrates. We address all levels of organization that are magnetic resonance imaging, magnetoreception, optogenetics,
germane, from the genome to the ecological context. sleep, symbiosis in the bobtail squid-Vibrio system, and synaptic
We want to mention four specific strategies we have adopted to development. Treatment of topics in global warming has been
add interest and breadth to the book. First, we start every chapter tripled. Treatment of altitude physiology in Chapters 23 and 24 is
with a vivid example of the application of the chapter’s material to entirely revised. The index is new and upgraded. A limited list of
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