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NewHumanPhysiology

| P a u l e v Z u b i e t a 2 n d E d i t i o n

C h a p t e r 2 1 : T h e r m o R e g u l a t i o n , T e m p e r a t u r e a n d R a d i a t i o n
HO ME

P RE FA CE

TA B L E O F CO NTE NTS

S Y MB O L S

S E CTI O N I NFO

CO NTRI B UTO RS

L I NK S

CO NTA CT US

Hi ghl i ghts

Chapter
21

S tudy_Obj ecti ves


P ri nci pl es
Defi ni ti ons
E ssenti al s

P athophysi ol ogy

E quati ons
S el fAssessm ent
Answers

T his C hapter is w r ittenfollow ingdis c us s ions w ithmy c olleague,LeifVanggaar d,MD ,Ar c tic Ins titute,

C openhagen.

Further Readi ng

StudyObjectives
T odefinebody c or eandbody s hell,heatbalanc e,heatex c hange( c onduc tion,c onv er s ion,

ev apor ationandr adiation) ,hy per ther mia,hy pother mia,meanbody temper atur e,heatc apac ity ,
andther mals teady s tate.
T odes c r ibefev er ( py r ogens ) ,benignantandmalignanthy per ther mia,heatex haus tion,heat
s y nc ope,heats tr ok e,s uns tr ok e,andhy pother mia.
T odes c r iber adiations ic k nes s .
T oc alc ulateonether malv ar iable,w henr elev antv ar iables ar egiv en.
T oex plainthec onc epts heatex c hange,ther mogenes is by foodands hiv er ing,thehuman
temper atur ec ontr ols y s temandits func tionatdiffer entenv ir onmentaltemper atur es .
T ous etheabov ec onc epts inpr oblems olv ingandc as ehis tor ies .

Principles
N ew t o n slaw o f co o lin g :T hedr y heatlos s is pr opor tionaltothetemper atur ediffer enc e
betw eenthehumanbody ( s hell) andthes ur r oundings .
T h et o t alen erg yo f asyst em is c ons er v edinaninter ac tion,notthek inetic ener gy or themas s
( Eins tein) .Ifthemas s c hanges dur inganinter ac tion,ther eis ar es ultantc hangeink inetic ener gy ,
s othatthetotalener gy r emains c ons tant.H eatener gy is pr opor tionaltomolec ular mov ement
r ates heatener gy equals mov ement.
St ef an B o lt z m an n sru le:T hehigher thetemper atur eofanobjec t,themor eitr adiates .T he
ener gy r adiatedfr omanobjec tis pr opor tionaltothefour thpow er ofits Kelv intemper atur e.T he
ener gy r adiatingfr omanobjec tandr ec eiv edby thehumanbody is pr opor tionaltothetemper atur e
differ enc ebetw eentheobjec tandthes k in( s eeEq .21 4) .T his is bec aus ehumanlifeimplies
r elativ ely s malltemper atur egr adients .

Definitions
B o d yco rec ons is ts ofthether mor egulateddeeper par ts ofthebody andthepr ox imalex tr emity
por tions ofw ar m bloodedanimals inc ludingman.
B o d ysh ellr efer s tothos eouter par ts ofthebody ( s k inands ubc utaneous tis s ue) thatc hange
temper atur eatc oldex pos ur e.
C o n d u ct an cec hanges ofthes hellar eus edas ameas ur eofs k inbloodflow .

C o n d u ct iveh eat lo ssdes c r ibes adir ec ttr ans fer ofheatener gy by c ontac tbetw eentw obodies
ofdiffer enttemper atur e( eg,s k inandobjec ts ) .
C o n vect iveh eat lo ssis definedas theheatlos s by c ontac tbetw eenthes ur fac e( s k in) anda
mov ingmedium( air or w ater ) .
Evap o rat iveh eat lo ssis definedas theheatlos s by ev apor ationfr omthebody s ur fac eor
lungs .
F everoc c ur s w henthec or etemper atur eofthebody is r ais edabov enor mals teady s tatelev els .
T hebody r eac ts as ifitis tooc old.F ev er implies adis or der r es ultingins hiv er ingc ombinedw ith
v as oc ons tr ic tion,headac he,dedolation,andgener aldis c omfor t( eg,malar ia) .
H eat f lo w is definedas ener gy ex c hangedduetoatemper atur ediffer enc e.H eatflow is
tr ans mittedalongatemper atur egr adient.
H eat cap acit yis theamountofheatr equir edtopr oduc eatemper atur einc r eas efor agiv en
amountofs ubs tanc e.
H eat en erg yb alan ceinar es tingper s onis ac ondition,w her etheheatpr oduc tionis equaltothe
heatlos s .T hus thebody temper atur eis c ons tantandtheheats tor ageis z er o( ther mals teady
s tate) .U s ually ,ther eis nointer nalheatener gy flux betw eenbody c or eands hell.
H yp ert h erm iais aninc r eas einc or etemper atur eabov enor mal.
H yp o t h erm iar efer s toac linic alc onditionw ithalow er edc or etemper atur e( below 35 o C ) .
Mean b o d yt em p erat u reis definedac c or dingtoEq .21 1( s eeendofC hapter ) .
N o n sh iverin g t h erm o g en esisis ar is einmetabolis m,w hic his notr elatedtomus c ular ac tiv ity
( s hiv er ingor ex er c is e) .
In sen sib lep ersp irat io n ( leak ageofthes k in) is thes mallc utaneous ev apor ationlos s ,w hic his
unr elatedtos w eatglandfunc tion.
In su lat io n r efer s tor es is tanc etoheattr ans fer .
R ad iat iveh eat lo ssis atr ans fer ofheatener gy betw een2s epar ateobjec ts atdiffer ent
temper atur e.H eatener gy is tr ans fer r edv iaelec tr omagnetic w av es ( photons ) .T his heattr ans fer
does notr equir eamedium,andthetemper atur eofany inter v eningmediumis immater ial.
Sh ellt em p erat u reis thetemper atur eoftheouter par ts ofthebody ( meas ur edonthes k in
s ur fac e) andr elatedtoc oldenv ir onments .
Sh iverin g is ar eflex my ogenic r es pons etoc oldw ithas y nc hr onous or balanc edmus c le
c ontr ac tions per for mingnoex ter nalw or k .
Sp ecif ich eat cap acit yis ther elations hipbetw eenheatener gy ex c hangedper w eightunitofa
s ubs tanc eandthec or r es pondingtemper atur ec hange.T hes pec ific heatc apac ity ofw at eris 4.18
andofthehumanbody ( bloodandtis s ues ) 3.49k J k g 1 o C 1 ,r es pec tiv ely .T hes pec ific heat
c apac ity ofatmos pher ic air is 1.3k J ( m 3 ) 1 o C 1 .
T em p erat u reis themeas ur ementofheatener gy c ontent.

Essentials
T his par agr aphdeals w ith
1.T h et em p erat u reso f t h eb o d y,2.B o d yresp o n sest o co ld ,3.B o d yresp o n sest o h eat ,4.
Em o t io n alsw eat in g ,5.Met ab o licR at ean d en viro n m en t alt em p erat u re,6.T em p erat u reco n t ro l,
7.T h eh u m an t h erm o co n t ro lsyst em ,and8.T h erm o reg u lat o ryef f ect o rs.
1.T h et em p erat u reso f t h eb o d y
T hehumanbody c ons is ts ofaper ipher als hellandac entr alc or e( F ig .21 1) .T heheatc ontent( H or
enthalpy ) ofthehumanbody is r eflec tedby its temper atur e.By definitionather mometer only
meas ur es thetemper atur eofthether mometer ,s oits loc ationis es s ential.T hemeanc or e
temper atur eis 37 o C inhealthy adults atr es t,buts mallc hildr enhav elar ger diur nalv ar iations .

T hes k inis themainheatex c hanger ofthebody .T hes k intemper atur eis deter minedby thec or e
temper atur eandby theenv ir onment( temper atur e,humidity ,air v eloc ity ) .T hus thes helltemper atur e
is gov er nedby theneeds ofthebody toex c hangeheatener gy .

F ig .21 1: H eat t ran sf ers,b o d yco resan d sh ellst em p erat u reso f an aked p erso n st an d in g in
co ld an d w arm air,resp ect ively.
T hes helltemper atur eis meas ur edonthes k ins ur fac eandatthehands andfeettoappr oac hthe
r oomtemper atur eof19 o C inaper s ons tandinginac oldr oomfor hour s ( F ig.21 1,left) .T hes hell
temper atur eis s ev er aldegr ees low er thanthetemper atur einthec entr alc or e.T helimbs hav ebotha
longitudinalandar adialtemper atur egr adient.T hes helltemper atur eandthes iz eofthes hellv ar y
w iththeenv ir onmentaltemper atur eandtheter mals tateoftheper s on.Anak edper s on,s tandingona
c oldfloor in19 o C air has as mallc or eandathic k s hellc ompar edtothes ameper s oninaw ar m
env ir onment( F ig.21 1) .T hes helltemper atur eofthes k inanddis talex tr emities is diffic ultto
ev aluate.T hebes tes timateis meas ur ementoftheinfr ar edheatr adiationflux w ithar adiometer .
T hec or etemper atur eis ther ather c ons tanttemper atur einthedeeper par ts ofthebody andinthe
pr ox imalex tr emity por tions ( s eether eds tippledlines ofF ig.21 1) .H ow ev er ,thec or etemper atur e
may v ar y s ev er alC entigr ades betw eendiffer entr egions dependingonthec ellular ac tiv ity .T hebr ain
has ar adialtemper atur egr adientbetw eenits deepands uper fic ialpar ts .Inas ens e,thetemper atur e
ofthemix edv enous bloodr epr es ents anes s entialc or etemper atur e.
T her ec taltemper atur e
Ahighc or etemper atur eis foundtobec ons tantinther ec tumabout10 15c mfr omtheanus .When
meas ur ingther ec taltemper atur eas tandar ddepthof5 10c mis us edc linic ally .T hev enous plex us
ar oundther ec tumc ommunic atew iththec utaneous bloodintheanalr egion.T her ec taltemper atur e
falls w henthefeetar ec old,bec aus ec oldbloodpas s es ther ec tuminthev eins fr omthelegs for the
s amer eas on.T her ec taltemper atur er is es dur ingheav y w or k inv olv ingthelegs .
Par ents s houldbeadv is edtomeas ur ether ec taltemper atur eindis eas es us pec tc hildr en.T her ec tal
temper atur eis ar eliablees timateofthec or etemper atur einr es tingper s ons .
Sublingual( or al) or ax illar y temper atur es ar eunr eliablemeas ur es ofthec or etemper atur e often
mor ethanhalfadegr eelow er thanther ec taltemper atur e.
T hec r anialtemper atur e( ty mpanic andnas al)
T hemainc ontr oloftemper atur eis per for medby theanter ior hy pothalamus ,w hic hhas ahigh
bloodflow .Withinthec r aniumthehy pothalamus lies ov er theC ir c leofWillis ,w hic hs upplies itw ith
blood,andc los etothec av er nous s inus w hic hdr ains it.H y pothalamus elic its heatlos s r es pons es
w hens timulatedby heat.T hety mpanic membr aneandar eas inthenas alc av ity ( theanter ior
ethmoidalr egion,par tofthes phenoids inus ) ar es uppliedw ithbloodfr omtheinter nalc ar otidar ter y
jus tlik ethehy pothalamus .T hes ec r anialloc ations thens er v eas as ubs titutefor themeas ur ementof
theinac c es s iblehy pothalamic temper atur e.
Intak eof250gofic er eleas es anabr uptfallinthenas altemper atur einaw ar mper s on,w her eas the
c hangeinr ec taltemper atur eis s maller anddelay ed( F ig.21 2) .T hec r anialc or etemper atur eis mor e
dy namic thanther ec tal.

F ig .21 2: In t akeo f icered u cest h et em p erat u rein aw arm p erso n rest in g at 45 o C .


Ins por ts andins ur gic alhy pother miady namic meas ur ements ofc or etemper atur ear ees s ential.T he
c r anialtemper atur eis oftenpr efer r ed.D ur ingfor c efulmov ements thether mis tor may bedis plac ed.In
s uc hs ituations anoes ophagealloc ationis appliedathear tlev el.T his is anappr ox imativ emeas ur eof
thetemper atur eofthemix edv enous bloodofther ighthear tloc atedc los etothether mis tor .
T hemeanbody temper atur eis definedac c or dingtoEq .21 1.T hes tor ageofheatener gy inthebody
c anbec alc ulatedac c or dingtoits heatc apac ity ( 3.49k J *k g 1 * o C 1 ) ,thebody w eight( k g) andthe
c hangeinmeanbody temper atur eintheper iod( Eq .21 2) .
Ac c or dingtothefir s tlaw ofther mody namic s ,thes tor ageofheatener gy equals themetabolic ener gy
c hangeminus theheatlos s ( Eq .21 3) .Q uantific ationofther mody namic s inhumans is pos s ibleus ing
equations 21 1to21 7( later inthis c hapter ) .
T hebody is inheatener gy balanc e,w henthes tor ageis z er o.H ow ev er ,thec or etemper atur emay
c hangew ithinter nalflux es ofheatener gy betw eenc or eands hellw ithouts tor ageor los s ofheat
ener gy atac ons tantac tiv ity .
Venous blooddr ainingac tiv emus c les andtheliv er is lik ely tobew ar mer thanpulmonar y v enous
blood,s inc ethis has under goneev apor ativ ec oolinginthealv eoli.Apatientw ithhighfev er c anbein
ther mals teady s tate,w ithahighc ons tantheatpr oduc tion,ifbothc or e ands hell temper atur es ar e
c ons tant,andnointer nalener gy flux oc c ur s .
War m bloodedanimals ,homeother ms s uc has humans ,c anc hangetheir metabolis minor der tok eep
their heatpr oduc tionequaltotheheatlos s .Suc hanimals hav eatemper atur ec ontr ols y s temand
ther eby maintainar ather c ons tantc or etemper atur e.War m bloodedanimals liv ew iththeadv antage
ofanunc hangedc ellac tiv ity andtemper atur eintheir c or e.H ow ev er ,thehumanc or etemper atur e
falls dur ingtheoes tr ogenphas eofthemens tr ualc y c leanddur ings leep( c ir c adianr hy thm) .T he
low es ttemper atur eis betw een18atnightand6oc loc k inthemor ning( F ig .21 3) .T hetemper atur e
c y c leis par tofthec ir c adianper iodic ity .O ur biologic alc loc k s eems tobes y nc hr onis edw iththe
r otationoftheglobe.Als omeals ,lightandtemper atur eplay s ar ole.
O v ulationr eleas es as har pr is einmor ningtemper atur e.Pr oges ter oneeffec ts s eemtoex plainthe
higher temper atur einthelas tphas eofthemens tr ualc y c le( F ig.21 3) .

F ig .21 3: Variat io n so f t h eco ret em p erat u red u rin g 24h o u r( ab o ve) ,an d variat io n srelat ed t o
p h aseso f t h em en st ru alcycle( b elo w ) .

C old bloodedanimals ( poik ilother ms ) liv ew ithabehav iour altemper atur er hy thm,buthav eno
autonomic temper atur ec ontr ol.T hec or e ands hell temper atur es v ar y w iththeenv ir onmentandthe
c ellular ac tiv ity .R eptiles ,pr ematur eandlow w eight pr ematur enew bor nbabies ar ec old blooded.
T hes ebabies hav enother mor egulation( s eelater ) .H ow ev er ,their c apac ity for heatpr oduc tionis 5
10times as gr eatper unitw eightas thatofadults .
H umans hav eaw ar m blooded( homeother mic ) c or eandac old blooded( poik ilother mic ) s hellinac old
env ir onment.
Per s ons ex pos edtogener alanaes thes ia,alc ohol,andc er taindr ugs los etheautonomic
ther mor egulation.C old bloodedanimals mus tliv ew ithv ar y ingc or eands helltemper atur e,w her eby
ther ateoftheir c ellular ac tiv ities v ar ies w iththes ur r oundingtemper atur e( F ig .21 4) .

F ig .21 4: T h eb o d yco ret em p erat u rean d t h een viro n m en t alb o d yt em p erat u ref o raw arm
b lo o d ed an im al( cat ) an d aco ld b lo o d ed an im al( liz ard ) .
a) C onv ec tion.T hec onv ec tiv eheatlos s is c alc ulatedby Eq .21 7.Ahealthy per s onins por ts
c lothes ex per ienc es ther malc omfor tatthr eetimes ther es tingmetabolic r ate( 3MET ) ,w henthe
s ur r oundingtemper atur eis 20 o C ,thehumidity is 50% andthew indv eloc ity is 0.5m*s 1.
D iv ing( w ater has ahighther malc onduc tiv ity ) illus tr ates theimpor tanc eofc onduc tionandc onv ec tion
inheatener gy tr ans fer .
T hedr y div ings uitex c ludes w ater fr omc ontac tw iththes k inandtr aps low c onduc tanc eair in
ins ulatingc lothingw or nins idethew ater tights ealing.
T hew ets uittr aps w ater nex ttothes k inbutpr ev ents its c ir c ulation.T hew ater is w ar medthr ough
c ontac tw iththes k in,andthehighins ulationofthefoamr ubber w etdiv ings uit,w ithits many poc k ets
oftr appedair ,minimis es ther ateofheatener gy los s tothes ur r oundingw ater .Air is apoor heat
c onduc tor andthus agoodins ulator .D ur ingdeepdiv inghighpr es s ur es c ompr es s thes eair poc k ets
andthus r educ etheins ulationpr oper ties ofw etdiv ings uits .
b) R adiationdes c r ibes atr ans fer ofener gy betw eenobjec ts inthefor mofelec tr omagnetic w av es
( photons ) .T his inc ludes ultr av ioletandv is ible( s unlight) r adiationfr omtheouts ideandfr omthe
body infr ar edor w ar mheatr adiation.
R adiativ eheattr ans fer c anbec alc ulatedfor anak edper s onac c or dingtoEq .21 4.
Whenthes k intemper atur e( T s k i n ) is les s thanthetemper atur eofthes ur r oundingobjec ts ,heatis
gainedby r adiation.
Atw inter time,heatc anbelos tthr oughaw indow glas s by r adiationfr omthebody tothec old
env ir onmentir r es pec tiv eofther oomtemper atur e.T his is bec aus ethes k intemper atur eis higher
thantheouts idetemper atur e.
c ) C onduc tion.Sittingonac olds toneis aty pic alex ampleofc onduc tionlos s ,jus tas s tandingona
c oldfloor ( F ig .21 1) .C onduc tionheatc anals obegained,althoughitis r eally pos s ibletow alk on
glow ingc oals w iths peedandathic k epider malhor nlay er .
d) Ev apor ativ eheatlos s s ees w eats ec r etionbelow .
2.B o d y resp o n sest o co ld
C utaneous v as oc ons tr ic tionlow er s s k intemper atur e,andther eby r educ es thec onduc tiv e c onv ec tiv e
heatlos s thatis deter minedby thetemper atur egr adientfr omthes k ins ur fac etotheenv ir onment.
C utaneous v as oc ons tr ic tiondir ec ts theper ipher alv enous bloodbac k tothebody c or ethr oughthe

deepv eins andthec ommitantv eins .T hes ev eins ar eloc atedar oundthear ter ies w ithw ar mblood,s o
thatthev enous bloodr ec eiv es par toftheheatener gy fr omthear ter ialblood s o c alledc ounter
c ur r entheatex c hange( F ig .21 5) .T hev as oc ons tr ic tionis s oeffec tiv e,thatthebloodflow thr ough
thear ter io v enous anas tomos es inthefinger s andtoes c anfalltobelow oneper c entoftheflow at
nor maltemper atur e.T hec oolingofthes hellis immediate,andthes iz eofthes hellinc r eas es ( F ig .
21 1) .O bv ious ly ,thes hellis lar gefor anak edper s oninc oldair .T her es is tanc ev es s els ofthe
hands may openper iodic ally tonour is hthetis s ues ,butthehighv is c os ity ofthec oldbloodc an
endanger thetis s uenutr itionandr es ultintr enc hfoot.
T hear ter io v enous s hunts ofthehands andfeetar ec los ed,s othebloodflow tothelimbs is a
nutr itiv eminimum.
T hedeepar ter ies andv eins ofthelimbs lieinpar allel,s othear ter ialbloodflow los es heattothe
inc omingv enous bloodpar tially s ur r oundingthear ter ies ( F ig.21 5) .T his is aty pic alc ounter c ur r ent
heatex c hange.Inac oldenv ir onment,w her ev as oc ons tr ic tionandheatex c hangepr oduc es c old
ex tr emities ,thetotalins ulationis inc r eas edattheex pens eofr educ edneur omus c ular effic ienc y .

F ig .21 5: C o u n t er cu rren t exch an g ein ah u m an arm co n servin g h eat en erg yin aco ld clim at e
( lef t ) .Su p erf icialven o u sco o lin g rib selim in at eh eat en erg yin aw arm clim at e( rig h t ) .
Inaw ar mc limatethehighbloodflow oftheex tr emities ens ur es anoptimaltemper atur eofthedeeper
s tr uc tur es ( eg,theneur omus c ular s y s tem) .T hetemper atur eofthear ter ialbloodis maintained( F ig.
21 5,r ight) andthear ter io v enous anas tomos es ar ew ideopenc onv ey ingw ar mbloodtothe
s uper fic ialv eins .T hes uper fic ialv eins als oac tas co o lin g rib sandtr ans fer lar geamounts ofheatto
thes k ins ur fac e,w her eitis eliminatedfr omthebody by c onv ec tion,c onduc tionandev apor ation
( F ig.21 5,r ight) .
Shiv er ingis ar eflex my ogenic r es pons etoc oldw ithas y nc hr onous or balanc edmus c lec ontr ac tions
elic itedfr omthehy pothalamus v iac utaneous r ec eptor s .T heac tiv ity inagonis tandantagonis t
mus c les balanc e,s other eis noex ter nalw or k .Withoutouts idew or k ,allener gy is liber atedas
metabolic heatener gy .H eatpr oduc tionis als oinc r eas edby thy r oidglandac tiv ity andby r eleas eof
c atec holamines fr omtheadr enalmedulla.
Ex ter nalw or k ,s uc has r unning,is helpfulinmaintainingbody temper atur ew henfeelingc old.C old
inc r eas es themotiv ationfor w ar m upex er c is es andillus tr ates thev oluntar y ,c or tic al( feedfor w ar d
influenc e) ontemper atur ehomeos tas is .T hec or etemper atur einc r eas es pr opor tionally tothew or k
intens ity dur ingpr olongeds teady s tatew or k ( F ig.21 6) .T hemeans k intemper atur efalls w ith
inc r eas ingw or k intens ity at20 o C ,bec aus ethes w eatev apor ationc ools thes k in.

F ig .21 6: Mu scu laran d o eso p h ag ealt em p erat u red u rin g st ead yst at eexercise.T h elevelso f
exerciseran g ef ro m z ero t o 100% o f t h em axim u m o xyg en u p t ake.

T hetemper atur eintheac tiv emus c les deter mines thelev elofther ec taltemper atur e.F ollow ing
mar athonr ec taltemper atur es ofmor ethan41 o C hav ebeenmeas ur edandheats tr ok es hav e
oc c ur r ed.Amar athonis ev enmor ediffic ulttoac c omplis hinw ar m,humidenv ir onments ands tr ong
s unmay c aus es uns tr ok e( s eelater ) .
Peoplemay adapttopr olongedex pos ur etoc oldby inc r eas ingtheir bas almetabolic r ateupto50%
higher thannor mal.T his metabolic adaptationis foundinInuits ( Es k imos ) andother people
c ontinuous ly s ubjec ttoc old.
T heenv ir onmentaltemper atur e,w her ew emaintainour autonomic temper atur ec ontr ol,is inther ange
ofz er oto45 o C .Below andabov ethis r angew eadapttotheenv ir onmentby behav iour ( addingor
r emov ingc lothing,w ar mor c oldbath,s unor s hadow ) .Ac or etemper atur eabov e44 o C s tar ts pr otein
denatur ationinallc ells andis inc ompatiblew ithlife.Below 32 o C humans los ec ons c ious nes s and
below 28 o C thefr equenc y ofmalignantc ar diac ar r hy thmias is inc r eas ing,endingw ithv entr ic ular
fibr illationanddeathatac or etemper atur ebelow 23 o C ( F ig.21 7) .

F ig .21 7: En viro n m en t alt em p erat u revariat io n san d t em p erat u reco n t ro l.L acko f vit alsig n sin
t h eclin ic( resp irat io n ,h eart rat e,EEG ) m u st n o t b et aken asd eat h .T reat m en t m u st b e
in st it u t ed u n t ild eat h sig n sared evelo p ed .
3.B o d y resp o n sest o h eat
Sw eat secret io n .T hr eemillions w eatglands pr oduc es w eatatar ateofupto2litr es per hour or
mor edur ingex er c is einex tr emew ar mc onditions .Ifnotc ompens atedby dr ink ing,s uc hhighs w eat
r ates leadtoc ir c ulator y failur eands hoc k .Sw eatr es embles adiluteultr afiltr ateofplas ma.H ealthy
humans c annotmaintaintheir body temper atur e,iftheenv ir onmentalair r eac hes body temper atur e
andtheair is s atur atedw ithw ater v apour .Pr imar y s w eatis s ec r etedas anis os motic fluidintothe
s w eatduc t,ands ubs equentN aC lr eabs or ptionr es ults inthefinalhy po os motic s w eat.T her mal
s w eatingis abolis hedby atr opine,pr ov ingthatthepos tganglionic fibr es ar ec holiner gic .C holiner gic
dr ugs pr ov ok es w eatingjus tas adr ener gic agonis ts do.Ev apor ationofw ater onthebody s ur fac e
eliminates 2428 2436J g 1 atmeans helltemper atur es of30 32 o C .Ev apor ationofalar gev olumeof
s w eatper timeunit( V s w e a t ) implies as ubs tantiallos s ofheatac c or dingtoEq .21 5.
N or mally ,thes k intemper atur efalls w ithinc r eas ingw or k intens ity ,bec aus ethes w eatev apor ation
c ools thes k in( F ig .21 6) .D anger oc c ur s w hentheav er ages k intemper atur eandthebody c or e
temper atur ec onv er getow ar ds thes amev alue.
C ondens ationofw ater onthes k ingains heatener gy ,w hic his s tor edinthebody .T his is w hat
happens inaSauna.
Vas odilatationofs k inv es s els inw ar menv ir onments r es ults ininc r eas edc ar diac output.T hear ter io
v enous anas tomos es inthehands andfeetar eopen,andthebloodflow c anr is euptoatleas t10
folds .T hes hellis minimal,w henanak edper s onis inw ar mair ( F ig .21 1,r ight) .T hes k inbloodflow ,
mainly intheex tr emities ,deter mines theamountofheatener gy ,w hic his c ar r iedfr omthebody c or e
tobelos tonthes ur fac e.T heheatener gy is tr ans por tedfr omthelar gebody c or etothes k inby
c onv ec tionintheblood.As ubs tantialpar toftheheatener gy is los tthr oughthes uper fic ialv eins of
theex tr emities ac tingas c oolingr ibs ( F ig .21 5) .T hebloodofthes uper fic ialv eins is thus

ar ter ializ ed,w hentheper s onis w ar m.


Apiec eofs teak has thes amec ompos itionas humans k inbutofc our s enobloodflow andnos w eat
ev apor ation.T hus thes teak w illbec ook edatanair temper atur ethathumans c ans ur v iv e.Aper s on
c ans tay inar oomw ithdr y air at128 o C for upto10mindur ingw hic htimethes teak is par tially
c ook ed.
4.Em o t io n alsw eat in g
T his is apar adox ic alr es pons einc ontr as ttothether mals w eatingofther mor egulation.Emotional
s tr es s elic its v as oc ons tr ic tioninthehands andfeetc ombinedw ithpr ofus es w eats ec r etiononthe
palmar andplantar s k ins ur fac es .
5.Met ab o licR at ean d en viro n m en t alt em p erat u re
T hetotalheatlos s c ons is ts oftheev apor ativ eheatlos s andthedr y heatlos s ( ie,thes umof
c onv ec tiv e,c onduc tiv eandr adiativ elos s ) .N ew tons law ofc oolings tates thatthedr y heatlos s is
pr opor tionaltothetemper atur ediffer enc ebetw eenthehumanbody ( s hell) andthes ur r oundings .
Letus look atahealthy ,lightly dr es s eds ittingper s oninther malbalanc e.H is heatlos s is plottedas
theor dinateandtheenv ir onmentaltemper atur eas theabs c is s a( F ig .21 8) .T hetw oty pes ofheat
los s ar eaddedinor der topr ov idethetotalheatener gy los s .Atar oomtemper atur eof37 o C ther eis
adr y heatlos s ofz er o,andbelow ther eis aninc r eas ingdr y heatlos s .Abov e37 o C theheatlos s
tur ns intoheatinput.O bv ious ly ,thedr y heatlos s als odepends uponc onduc tionandc onv ec tionof
heatins idethebody by c ontac tandby theper fus ion.Atex tr emely low env ir onmentaltemper atur ethe
dr y heatlos s bec omes lar ger thanthemetabolic heatliber ationandthebody is c ooleddow n.
T heper s onis inther mals teady s tate,andthemetabolic r ateis almos tc ons tantinthether moneutr al
z onebetw een20and30 o C ( F ig.21 8) .T helaw ofmetabolic r educ tionr eflec ts thetendenc y for heat
pr oduc tiontomatc hther ateofheatlos s .T hether moneutr alz one,w her eminimalc ompens ator y
ac tiv ity is r equir ed,is s epar atedinthelo w ervaso m o t o randtheupper s udomotor c ontr olz one.
Inthelow er ,c omfor tablez one( 20 26 o C ) thetotalheatdis s ipationis maintainedequaltothe
metabolic r ateby c utaneous ,v as omotor alter ations .T hes mallev apor ationlos s is ter medleak ageof
thes k inor ins ens ibleper s pir ation,w hic his unr elatedtos w eatglandfunc tionandr ather c ons tantat
thebas almetabolic r ate.

F ig .21 8: Met ab o licrat ean d en viro n m en t alt em p erat u rein af ast in g d ressed h u m an at rest .
T h ew et an d t h ed ryh eat lo ss,asw ellast h em et ab o lich eat an d t h eb asalm et ab o licrat e
( B MR ) ism easu red in Wat t s.
Intheupper s udomotor z oneabov e26 o C env ir onmentaltemper atur e,thebloodflow thr oughthes k in
r is es ,as does s w eats ec r etionandev apor ation.At37 o C ther is einener gy los s oc c ur s v ia
ev apor ation( F ig.21 8) .
Whentheenv ir onmentaltemper atur efalls themetabolic r ateinc r eas es fir s tby inc r eas ingmus c le
toneandthenby s hiv er ing.T hec hemic alor metabolic temper atur ec ontr olis intheenv ir onmental
r egionfr om20 o C andbelow ( F ig.21 8) ,w her es hiv er ing,dec r eas edbloodflow thr oughs k inandnon
my ogenic heatpr oduc tiontak eplac e.H er e,metabolis mc ontr ols thec or etemper atur eby inc r eas ing
metabolic r atew ithfallingtemper atur eintheenv ir onment.Abov e20 o C ,thephy s ic altemper atur e
c ontr oltak es ov er ,as anautonomic c apac ity for alter ations inheatlos s .Inthis ther moneutr alz one
thebody temper atur eis k eptc ons tantalmos tw ithouteither heat pr oduc ingmec hanis ms or s w eat
s ec r etion. T hether malc omfor tfor lightc lothed,s eatedper s ons is aboutanair temper atur eof

26 o C w henthehumidity is 50% ,30 o C for nudeper s ons ,andabout36 o C s ittinginw ater tothenec k .
Inthemetabolic z one,thetotalheatlos s r is es w ithfallingenv ir onmentaltemper atur e,butbelow 5 o C
intheenv ir onment,thedr y heatlos s ex c eeds themetabolic r ate,andthebody is c ooleddow n( F ig .
21 8) .T his is thez oneofhy pother mia,w her ec olddeathis inev itablew ithouttr eatment.
T hez oneofhy per ther miabegins atanenv ir onmentaltemper atur eof37 o C ,w her ehumans s oon
r eac hthemax imalc apac ity for ev apor ationandther eis anunbalanc edheatinflux tothebody ending
inheatdeath.
T hemetabolis mv ar ies w iththes hellandthec or etemper atur e.T hes er elations w er eeluc idatedby
s er ies ofs imilar ex per iments .Aper s onw as plac edins tir r edw ater tothenec k ,w her ethew ater
temper atur ethus definedthemeans helltemper atur e.By pr e tr eatmentinother baths anar r ay of
c or e ands hell temper atur ec ombinations w er eobtainedandmeas ur eds imultaneous ly w iththe
metabolic r ate( metabolic heatliber ation) .T hec or etemper atur ew as r educ edby intak eofupto2lof
c r us hedic einw ater .
Atas helltemper atur eof20 o C ,intak eofic ew ater r educ es thec or etemper atur ebelow 37.1 o C
( her ec alledthes etpoint) andthemetabolic r ateinc r eas es by s hiv er ing( F ig.21 9) w ithfallingc or e
temper atur e.

F ig .21 9: T h em et ab o licrat easaf u n ct io n o f t h eco rean d t h esh ellt em p erat u re.


War mer s k inmak es thes etpointfallandther is einmetabolic r ateper o C ( ofc or etemper atur efall) is
les s s teep.C or etemper atur es abov e37.1 o C allhav ealow metabolic r ate,r egar dles s ofthes hell
v alues .Atas helltemper atur eof30 o C thes etpointdec r eas es fr om37.1to36.7 o C .T he
hy pothalamus func tions as ather mos tatus ingther es tofthebody tos tabilis eits ow ntemper atur e.
Withr is ingc or etemper atur ethemetabolic r ateis maintainedlow andthebody tr ies toc ooldow n.
Whenthebody c or etemper atur efalls below s etpoint,themetabolic r ateinc r eas es by s hiv er ingand
theheatener gy s tor ageas w ell.T hec utaneous c oldr ec eptor s ar emax imally ac tiv eat20 o C and
they ar es ilentabov e33 o C they c anonly tr igger s hiv er ing,w henthec or etemper atur eis below s et
point.Shiv er ingc eas es immediately ,w henw etak eaw ar ms how er andthes k inis w ar med.T he
hy pothalamic ,pr eoptic heatr ec eptor s inhibits hiv er ing,ands hiv er ingis totally bloc k edabov ethes et
point.
Another s er ies ofex per iments w er edir ec tedtow ar ds heatlos s .T heheatlos s fr omev apor ateds w eat
w as r ec or deddur ingex er c is eatdiffer ents helltemper atur es .

F ig .21 10: Evap o rat iveh eat lo ssasaf u n ct io n o f t h eco rean d t h em ean skin t em p erat u re
d u rin g d if f eren t in t en sit ieso f exercise.

C olds timulates c utaneous c oldr ec eptor s w ithc onnec tions tothehy pothalamus ( F ig .21 12) .
Atmeans k intemper atur es of33 39 o C ,w her ethec utaneous c oldr ec eptor s ar es ilent,theper s onis
unabletos w eatbefor ethec or etemper atur eis abov e36.9 o C ( thes etpointofF ig.21 10) .Below the
s etpoint,theev apor ationis low ( per s pir atioins ens ibilis ) .Withinc r eas ingmus c leac tiv ity andc or e
temper atur etheev apor ativ eheatlos s may r is etow ar ds 20k J eac hmin.
Means k intemper atur es below 33 o C r educ etheev apor ativ eheatlos s .T her is inghy pothalamic
temper atur er eleas es thes w eats ec r etion,buttheloc als ec r etionis inhibitedby thec utaneous c old
r ec eptor s .
H eathas adir ec teffec tonpr eoptic heatr ec eptor s inthehy pothalamus .Atinc r eas edc or e
temper atur e,thepr eoptic heatr ec eptor s totally bloc k s hiv er ing,althoughthehy pothalamic c entr e
als or ec eiv es s hiv er ings ignals fr omc utaneous c oldr ec eptor s inc olds ur r oundings w ithmeans k in
temper atur es below 33 o C .T hefallingmeans k intemper atur ec anals oac tdir ec tly tolow er c utaneous
bloodflow ands w eats ec r etion.
6.T em p erat u reco n t ro l
H umans hav ear ather c ons tantc or etemper atur ealthoughthemetabolis mandenv ir onmental
temper atur emay v ar y c ons ider ably .T his implies thatc ontr olis ex er ted.

F ig .21 11: T h erm o reg u lat io n b yd yn am icg ain an d set p o in t syst em s.


Infor mationabouttheenv ir onmentaltemper atur eis pr ov idedby per ipher alther mo s ens or s ,w hic har e
loc atedinthes k in,abdominalor gans ,andmus c les .Inter nalor bloodtemper atur eis monitor edby
c entr alther mo s ens or s inthepr eoptic hy pothalamus andthemedulla.
Ar is einhy pothalamic temper atur ec aus es v as odilatationinthes k inandr educ es mus c ular tone.T he
per s onloos es motiv ationfor phy s ic alac tiv ity andr educ es c lothing.T henther mals w eatis obs er v ed,
andafter s ometime,r educ edac tiv ity oftheadr enalc or tex andofthethy r oidglandis als oobs er v ed.
Afallinhy pothalamic temper atur eby c oolingofthes hellandc or er eleas es c utaneous
v as oc ons tr ic tiontogether w ithinc r eas edmus c ular toneands hiv er ing.T her eis as y mpathetic
ac tiv ationw iths ec r etionofc atec holamines ,ox idationoffatty ac ids andgluc os e,andinc r eas ed
s ec r etionofthethy r oidandadr enalgland.T hemus c letoneis inc r eas ed,ands hiv er ingis tr igger ed
r eflex ly as as y nc hr onous mus c lec ontr ac tions w ithoutex ter nalw or k ( mov ements ) ,s oallthe
metabolic ener gy is r eleas edas heat.T hec apac ity for mus c ular ther mogenes is by s hiv er ingis high.
U ptofiv efolds bas almetabolic r ateis obs er v ed,w hic hc or r es ponds toheav y indus tr ialw or k .
Shiv er ingmay bes uppr es s edv oluntar ily atthebeginning.T r ans mis s ions ignals for s hiv er ingpas s es
ther ubr os pinalpathw ay s to and motor neur ons ofantagonis tmus c les .
D yn am icg ain an d set p o in t co n t ro l
A:Ady namic gains y s temr es ponds c ontinuous ly tofeedbac k s ignals r egar dles s ofthec or e
temper atur e.Withr is ingtis s uetemper atur e,theneur alac tiv ity ofheats ens or s inc r eas es
linear ly ,w her eas theac tiv ity ofc olds ens or s dec r eas es ( F ig .21 11A ) .
T his c ombineds ens or y inputtothehy pothalamus inc r eas es thec or etemper atur eandthus
inc r eas es theac tiv ity ofheatlos s effec tor s ,w hileinhibitingheatpr oduc tioneffec tor s ( F ig.
21 11A) .T his deter mines ther efer enc es ignal.
T hedy namic gains y s temhas afloatingr efer enc es ignalmov ingw iththec ontinuous heat
los s andheatpr oduc tion( F ig.21 11A) .

B:As et points y s temdoes notr es pondtoar is eintemper atur ebefor eac er tains etpointis
r eac hed.T hes etpointis thec or etemper atur eatw hic hneither heatlos s mec hanis ms or
heatpr oduc tionmec hanis ms ar eac tiv e.
Whenather maldis or der r eac hes ac er tains et pointinthehy pothalamus ,s ignals pas s es to
theeffec tor s .T hedes ir edc or etemper atur e( t s e t =s etpointtemper atur e) is c ompar edtothe
ac tualv alue( t c o r e inF ig .21 11B ) .T hec audalhy pothalamus w or k s as ather mos tat.Er r or
s ignals adev iationfr omt s e t ev ok er es pons es thattendtor es tor ec or eandhy pothalamic
temper atur etow ar dthes etpoint.Whentheac tualc or eandhy pothalamic temper atur er is es
abov ethedes ir eds etpoints uc has 37 o C ,effec tor s ar etur nedon,andthec ompens ator y
heatener gy los s is almos tlinear ( F ig.21 11B) .T hes ec ompens ator y mec hanis ms
( v as odilatation,s w eat,r educ edmus c letone) donottur noffuntilthetemper atur edr ops to
thes etpoint( ie,anall or nonr es pons e) .
Whentheac tualc or eandhy pothalamic temper atur eis jus tbelow thes etpoint,the
c ompens ator y mec hanis ms ( v as oc ons tr ic tionands hiv er ing) ar er elativ ely inac tiv e.
T hehy pothalamic s etpointc hangew iththephy s iologic alc onditions andis elev atedinfev er
by py r ogens fr ommic r oor ganis ms .T her is einmetabolis mis mainly ac c omplis hedby
s hiv er ing.
7.T h eh u m an t h erm o co n t ro lsyst em
H umantemper atur ec ontr olex hibits bothdy namic gainands etpointc har ac ter is tic s .T hec ontr ol
s y s temimplies w ides pr eadc utaneous anddeeps ens or s .T heir affer ents c onv er getow ar ds the
hy pothalamic integr ator ,w hic hac ts as ather mos tat.T hehy pothalamus als oc ontains ther mos ens or s
inthepr eoptic r egion,andinhibitor y neur ons per for mc r os s inginhibition( F ig .21 12) .T hec entr al
heatdr iv efr omthepr eoptic hy pothalamus is maintained( F ig.21 12) .T hes tability ofthec or e
temper atur eis maintainedby thelar geheatc apac ity ofthebody mas s ,andby thedeep
ther mos ens or s ,w hic har edominant.
Shiv er ingis r eleas edfr omc utaneus c olds ens or s fir ingmax imally at20 o C .T hes ec olds ens or s ar e
s ilentabov e33 o C .T hec olds hell( F ig.21 12) ac tiv ates deepc olds ens or s inthepr eoptic
hy pothalamus .T his inc r eas es heatpr oduc tionby s hiv er ing.T hepr eoptic ther mos tats imultaneous ly
r educ es heatlos s by c r os s inginhibition.

F ig .21 12: T h eh yp o t h alam ict h erm o st at an d it sco n n ect io n s.


Sw eats ec r etionis r eleas edby pr eoptic w ar ms ens or s as s oonas their temper atur eis 37 o C or
abov e( t s e t ) .C utaneous c olds ens or s inhibits w eats ec r etionats helltemper atur es below 33 o C ,
s inc ethey ar es ilentabov ethis temper atur e.
Inc onc lus ion,pr eoptic w ar ms ens or s s how s etpointc har ac ter is tic s below thes etpoint,andpr eoptic
c olds ens or s s how s etpointc har ac ter is tic s abov ethes etpoint.
Apar tfr omthat,pr eoptic s ens or s s how dy namic gain:Withr is ingtis s uetemper atur e,theneur al
ac tiv ity ofheats ens or s inc r eas es linear ly ,w her eas thes ignalfr equenc y ofc olds ens or s inc r eas es
w ithfallingtemper atur e( F ig .21 11A ) .
C utaneous s y mpathetic v as odilatationis pr obably als or eleas edby pr eoptic w ar ms ens or s abov es et
point( F ig.21 12) .Afallins k intemper atur ebelow 33 o C w illr educ es k inbloodflow by c r os s ing
inhibition( F ig.21 12) .

Alc ohols eems tooffs etthether moc ontr olmainly by inhibitionofthehy pothalamic ther mos tat( F ig.
21 12) .
N ew bor ns ,dow ntopr ematur ebabies abov e1000g,pos s es s c er tainther mor egulator y func tions .T he
new bor nc aninc r eas ether mogenes is by afac tor ofthr eew ithouts hiv er ings how s v as omotor
r eac tions ,s w eats ec r etion,andr educ es thes ur fac ear eainc oldair .H ow ev er thebaby has s pec ial
pr oblems :T hes ur fac e v olumer atiois 3 foldhigher thanthatofanadult.T hebaby has athins hell
duetothethins ubc utaneous fatlay er ,s oev enamax imalv as oc ons tr ic tionc annotlimitheatlos s w ith
ac apac ity c ompar abletothatofanadult.N ew bor ns ar es pec ially equippedtoper for mnon s hiv er ing
ther mogenes is ( c hemic alther mogenes is ) .N on s hiv er ingther mogenes is is any r is einmetabolis m,
w hic his notr elatedtos hiv er ing.Inbabies this for mofther mogenes is is par tic ular ly lar geintheir
br ow nadipos etis s ue.T his tis s ueis abundantar oundv italor gans ,innec k andmedias tinum,
betw eens c apulaeandinthear mpits .Br ow nadipos etis s uec ells c ontains multiloc ular dr oplets inthe
fatphas eandmany mitoc hondr ia.T hetis s uer ec eiv es s y mpathetic inner v ationandis s timulatedby
c atec holamines andthy r oidhor mones .C oolinginc r eas es thebloodflow andtemper atur eofbr ow n
adipos etis s ue.N or adr enalineinjec tions c aus ev as odilatationv ia r ec eptor s ,andinc r eas ethe
metabolis mtothes ameex tent.T hec aus eoftheinc r eas edmetabolis mis aninc r eas edc ell
membr aneper meability for N a + andK + ions ,w her eby theN a + K + pump( highAT Pdemand) is
ac tiv ated.

F ig .21 13: T h erm o co n t ro lin t h en ew b o rn .


N ew bor ns ar einther malbalanc eataminimalmetabolis monly w henthes ur r oundingtemper atur eis
high( 32 34 o C ) .Withother w or ds ,new bor ns hav eanex tr emely highlow er thr es holdfor the
ther moneutr alz one,namely 32 o C .T hethr es holdfor max imalus eofs hiv er ingis appr ox imately 23
o C c ompar edtoanak edadultabout5 o C .Ingener al,thes ethr es holdv alues for new bor ninc r eas e

w ithfallingbody w eightinpr ematur e.Pr ematur ebelow 1000ghas har dly any ther moneutr alz one
( F ig.21 13) .T hey ar eac tually c old bloodedandtheir temper atur ec ontr olis maintainedw itha
c ouv eus e.
8.T h erm o reg u lat o ryef f ect o rs
T hes y mpathetic andthes omatomotor ner v ous s y s tempar tic ipateinther mor egulation( F ig .21 14) .
N or adr ener gic s y mpathetic neur ons c ontr olthebloodflow thr oughfinger s ,hands ,ear s ,lips andnos e.
Ar ter ioles c ontr ac tandar ter iov enous anas tomos es c los e( ther malins ulation) follow inganinc r eas ein
s y mpathetic tone,anddilatatefollow ingadec r eas eintone.Whenar ter ioles andar ter iov enous
anas tomos es open,thebloodflow is mar k edly inc r eas edandthus thec onv ec tiv eheatlos s fr omthe
s k inis inc r eas ed.

F ig .21 14: T h et h erm o reg u lat o ryf eed b acksyst em .


C holiner gic s y mpathetic fibr es c ontr ols w eats ec r etion.T hev as odilatator br ady k ininis liber atedin
thes k in.T hus ,pr ofus es w eats ec r etionis alw ay s ac c ompaniedby v as odilatation.
Sy mpathetic ac tiv ationr eleas es thy r oidhor mones fr omthethy r oidglandandc atec holamines fr omthe
adr enalmedulla.T hes ehor mones liber atefatty ac ids andgluc os efor c ombus tion.Ar educ ed
s y mpathetic toneals or educ es theac tiv ity oftheadr enalandthethy r oidgland.
T hether mogenic r es pons etoc oldals oinv olv es anon my ogenic or non s hiv er ingc omponent
pr obably inadipoc y tes .N on s hiv er ingheatpr oduc tionis c ontr olledby thes y mpathetic ner v ous
s y s temv iaadr ener gic r ec eptor s .T henor adr enaline( N AinF ig.21 14) r eleas edatthener v e
ter minals c los etotheadipoc y tes ,s timulates theliber ationoffr eefatty ac ids andtheir s ubs equent
ox idation.N on my ogenic heatpr oduc tioninc ludes ac ontr ibutionfr omthebr ow nfatofbabies ,butis
ins ignific antinadults .
Shiv er ingis induc edby w ay ofthemotor s y s tem.T hec entr als hiv er ingpathw ay pas s es fr omthe
hy pothalamus tothemotor neur ons inthes pinalc or d.Shiv er ingis abolis hedby bloc k adeofthe
neur omus c ular endplatew ithc ur ar e.
T her mor egulator y behav iour s uc has fanningandaddingor r emov ingc lothingis effec tiv einc hanging
thether malins ulation.Sev er allay er s ofc lothingw ithtr appedair ac tas agoodins ulator .
Inhealthy per s ons ,heatener gy is liber atedfr omc ellular metabolis mandtr ans fer r edtothe
env ir onmentthr oughthes k inby s w eatingandv as odilatation.Sw eatingoc c ur s dur ingex er c is e,and
its ev apor ationis themos timpor tantmec hanis minmaintainingthec or etemper atur eas c los eto37 o
C as pos s ible.T hether mor egulator y c entr einthehy pothalamus c ontr ols allpr oc es s es .

F ig .21 15: H eat an d co ld ad ap t at io n


C oldadaptationis foundamongAus tr alianabor igines andInuits inG r eenland.Inuits hav er elativ ely
mor es w eatglands inthefac eandles s onthebody .
Abor igines c ans leepnak edonthegr oundev enatlow temper atur e.Inuits hav eabas almetabolic r ate
50% higher thanper s ons liv inginatemper atec limatedo.T hethr es holdfor s hiv er ingis s hifted
tow ar ds theleftinc old adaptedper s ons ,butthey maintainnor malfunc tionatthenew s etpoint.Ver y
oldpeoplemay s how thes amephenomenon,andliv ew ithac or etemper atur eof35 o C w ithout
s hiv er ing.O bv ious ly ,c oldadaptationimplies non s hiv er ingther mogenes is ,w hic his oec onomic
metabolic heatliber ation( F ig .21 15) .
H eatac c limatiz ationis ac tually as w eatglandadaptation,anda2 w eek pr oc es s follow ingar r iv altoa
hotc limates uc has thetr opic s or ades er t.G r adually s w eat ev apor ationis inc r eas edandtheN aC l
los s is r educ ed.T hes w eats ec r etionc apac ity may r eac h4leac hhour w ithathins w eat.T he
adaptationis c aus edby inc r eas edaldos ter ones ec r etionfr omtheadr enalc or tex .Aldos ter one
inc r eas es ther eabs or ptionofN aC landthes ec r etionofK + fr omthes w eat,dur ingits pas s ageofthe
s w eatglandtubules .T helar ger s w eatlos s thethir s tier onefeels .T his is bec aus ethelar ges w eat
s ec r etionr educ es thetimeper iodfor N aC l r eabs or ptioninthes w eatglandtubules .T her es ulting
highN aC lc onc entr ationintheplas maimplies thir s t,s oheatadaptedper s ons hav etodr ink alot.
T hir s tis anex tr emely lateindic ator ofdehy dr ationdur ingw or k inaw ar mc limate.
T r opic inhabitants ar eofc our s eheat adapted.T hey hav eaninc r eas edc or etemper atur e( s etpoint)
andtheir thr es holdfor s w eatandv as odilatationis ty pic ally 0.5 o C higher thanthatofaper s onliv ing
inthetemper atez one( F ig.21 15) .

Pathophysiology
T his par agr aphdeals w ith1.H eat cram p s,2.H eat exh au st io n andheats tr ok e,3.Malig n an t
h yp ert h erm ia,4.H yp o t h erm ia,5.F ro st b it e,and6.F everan d h yp ert h erm ia.
Apar agr aphc onc er ningnuc lear ener gy r adiationis giv enattheend.
1.H eat C ram p s
Painfulc r amps inthelegmus c les oc c ur follow ingex er c is e,w henathletes r untoofas tinahot
c limate.H eatc r amps r es pondtos altandw ater r eplenis hmentinanor maldiet,andthec r amps ar e
pr obably c aus edby hy ponatr aemia.D ur ingpr olongeds w eating,ther unner is los ings altandw ater .If
only thew ater los s is r eplenis hed,ther es ultis w ater induc edhy ponatr aemia apar alleltothe
c las s ic alminer s c r amps .
2.H eat Exh au st io n an d H eat St ro ke
Whenthew ater s altbalanc eis atr is k inahotc limate,itis alw ay s athr eattothec ir c ulation.Pr ofus e
s w eats ec r etion,inas ubjec tw hois notac c limatiz ed,r es ults ins alt andw ater depletion,w ithadaily
los s ofmor ethanonemolofN aC landmor ethan6lofw ater .Withinaper iodofone hour s tr enuous
w or k ingendur anc eathletes hav elos tupto8lofw ater .
T hefallingex tr ac ellular fluidv olumeandinc r eas ingbody andbr aintemper atur etoabov e40 o C elic it
s ev er es y mptoms ands igns .As thev olumeands altdepletiondev elops ,thes w eatpr oduc tiongoes
dow nins piteofex tr emev as odilatation.T hefallingbloodpr es s ur es timulates thehigh pr es s ur e
bar or ec eptor s r es ultinginar is inghear tr ate.T hedehy dr ationw ithimminents hoc k fr equently r es ults
inc er ebr al,r enalandhepatic failur e.T helow br ainbloodflow thr oughanov er heatedbr ainleads to
fatigue( heatex haus tion) ,c onfus ionandunc ons c ious nes s or s y nc ope( heats y nc ope) .T hec onfus ion
may dev elopintoav er itabledelir ium( anac uteimpair mentofc ons c ious nes s ) w ithbr ainoedema.
H eats tr ok e inthes unitis c alleds uns tr ok e is heatc ollaps ethatoc c ur s s uddenly ,her eby
c r eatingalife thr eateningc ondition.T his heatc ollaps eoftenoc c ur s inw ar m,humidenv ir onments ,
w henanunac c limatiz eds ubjec tex er c is es .T hes ubjec t w ithouts w eating( hy pothalamic failur e)
s uddenly falls intoc oma,ifnotpr ec ededby as hor tper iodofc onfus ionanddelir ium.T hec onditionis
fatal,ifnotr eliev edby r apidc ooling.
3.Malig n an t h yp ert h erm ia
Malignanthy per py r ex iais oftenc aus edby agenetic defec t( autos omaldominant) inthes ar c oplas mic
r etic ulumofs k eletalmus c les .G ener alanaes thes ia( oftenhalogen s ubs titutedethane) tr igger s an
aller gic r eac tionw iths uddenopeningofC a 2 + c hannels inthemus c lec ells .T hefollow inginflux of
C a 2 + elic its gener aliz edandmaintainedmus c lec ontr ac tion( r igidity ) ,w hic hliber ates enor mous
quantities ofheatener gy .T his c onditionis lifethr eateningandoftenr es ults ins uddendeathdur ingor
jus tafter anaes thes ia.
4.H yp o t h erm ia
H y pother miais afallinc or etemper atur etov alues below 35 o C .H y pother mic s ubjec ts los e
c ons c ious nes s ,w henthec or etemper atur efalls below 32 o C apotentially lethalc onditionc alled
s ev er ehy pother mia.
D ur inganaes thes iaands ur ger y thec or etemper atur eofthepatientfalls ands tabilis ear ound34 35
o C after 4 5hour s .Suc has ur ger y hy pother miainc r eas es ther is k ofc ar diac c omplic ations ,bleeding

tendenc y andpr olongedw oundhealing.


T hec onditionis apr ominentc aus eofdeathinc limber s ands k ier s ,as w ellas inper s ons being
immer s edinc oldw ater or liv ingintheAntar c tis .T hec limber s ands k ier s ar eoftenex pos edtoac old,
w etandw indy env ir onmentc ar r y ingins uffic ientc lothing.As theneur omus c ular func tions uffer s ,they
c annolonger mov e.G ener alhy pother miadev elops andthey die.
Inmildhy pother miathes ubjec tc ans tilltak eac tiontor ew ar mby ex er c is eandc lothing,butas
c ons c ious nes s is los tthec or etemper atur efalls fur ther ,bec aus es hiv er ingis abolis hed.T hepatient
feels c oldtotouc h,is inadev elopingc oma,thec ir c ulationandr es pir ationfall,as does themetabolic
r ate.T hedis s oc iationc ur v eofox y haemoglobinis mov edtotheleft,andthes olubility ofgas s es
inc r eas es as thebloodtemper atur efalls .

T hehear tis thetar getor ganinhy pother mia.Below 30 o C ,s pontaneous r emis s ionis pr ac tic ally
impos s ible,anddeathens ues fr omv entr ic ular fibr illationoc c ur r ingar ound28 o C .
C ar efulmonitor ingofallv italfunc tions is r equir eddur ingr ew ar ming,w hic his per for medeither
pas s iv ely ( r ew ar mingby thepatients ow nmetabolis m) w ithins ulationands pac eblank ets or ac tiv ely
by aw ar mw ater bath,w hilemonitor ingthepatient.Agoods tr ategy intr eatinghy pother mic v ic tims is
thes logan: T hey ar enotdeaduntilthey ar ew ar manddead.
Anar ter ialbloods amplefr omahy pother mic patientis r outinely analy s edat37 o C .T heP a O 2 andpH a
is fals ely higher ,theP a C O 2 is fals ely low er ,thaninthec ir c ulatingbloodofthehy pother mic patient.
T his is bec aus emor eC O 2 is boundals oas c ar bamino haemoglobin,andles s O 2 is boundinthec old
blood.Bas eEx c es s is definedat37 o C andthus atr uemetabolic v ar iable( s eeC h ap t er17) .
Ar tific ially induc edhy pother miais us edinbr ain andhear t s ur ger y ,w her etheus ualther moc ontr olis
inac tiv atedby gener alanaes thes ia.T hepr oc edur ebec omes danger ous ,w henitelic its v entr ic ular
fibr illation.
5.F ro st b it e
F r os tbiteis aloc alc oldinjur y duetothefor mationofex tr ac ellular ic ec r y s tals inthes k inandother
tis s ues .T his leads toex tr ac ellular dehy dr ationandhy per os mos is ,w her eby thec ells los ew ater until
they die.As thes k intemper atur efalls below +7 o C thes ubjec ts hav elos ttheir s ens or y func tions ,
andthus donotr ec ognis ethedev elopingfr os tbite.
T hemos teffec tiv etr eatmentis w ar mingofthes tillfr oz enar eaby immer s ionin40 42 o C hotw ater
follow inghos pitalis ation.T hepatientex per ienc es s ev er epain( abov e+7 o C ) andmor phinemus tbe
adminis ter ed.T is s uew hic his nolonger fr oz enmus tbetr eateds ter ile.
6.F everan d h yp ert h erm ia
F ev er oc c ur s w henthec or etemper atur eofthebody is r ais edabov enor mals teady s tatelev els .T he
body r eac ts as ifitis tooc old,w hilethetemper atur er is es uptothenew higher s etpoint.F ev er
attac k s imply s hiv er ingc ombinedw ithv as oc ons tr ic tion,headac he,dedolationpains ,andgener al
dis c omfor t.F ev er is ther es ultofoneoftw ophenomena:thes etpointmay bes ettoahigher lev el,or
theeffic ac y ofthetemper atur ec ontr ols y s temmay beimpair ed.F ev er implies hy per ther miahow ev er
many c as es ofhy per ther miadonotc ons titutefev er .F ev er r es ults fr omtheac tionofendogenous
py r ogens onthehy pothalamic heatc ontr olc entr e( they inc r eas ethes etpointfor thec or e
temper atur ev iapr os taglandins ) .Ex ogenous py r ogens fr ommic r obes c aus ethes eendogenous
poly peptides tober eleas edfr omthedefenc ec ells ofthebody ( ie,ther etic uloendothelials y s tem,
R ES) .Antipy r etic dr ugs inhibitc y c lo ox y genas eac tiv ity ,her eby inter fer ingw iththes y nthes is of
pr os taglandins andthr ombox anes .
F ollow inganattac k offev er ,v as odilatationands w eatev apor ationr educ es thec or etemper atur e.
Phy s iologic hy per ther miais aninc r eas einc or etemper atur ec aus edby ex tr emeheats tr es s or
ex er c is e.D ur ingw or k thebody temper atur er is es upto39 o C w ithoutc linic alc ons equenc es .H er eby ,
theheatlos s c apac ity is ex c eeded.D ur inghy per ther miatheheatlos s effec tor s ar es tr ainedtothe
utmos t.T hehighbody temper atur es ofex er c is eac tiv atec oolingmec hanis ms andelic its w eatlos s ,
w hic hs tr iv etor etur nthec or etemper atur etoits nor mallev el.Inex tr emehy per ther miathec or e
temper atur emay r is etomor ethan41 o C ( heats tr ok e) .Ir r ev er s iblepr oteindenatur ationoc c ur s
abov e44 o C w ithbr ainoedemaanddes tr oy edther moneur ons inthehy pothalamus .C linic ally ,the
br aindamageis s how nw ithdis or ientation,lac k ofs w eats ec r etion,delir iumanduniv er s alc r amps
befor edeath.
Io n iz in g R ad iat io n H az ard s
Ioniz ingr adiationimplies des tr uc tionoftis s uemolec ules .
D os ages ofabs or bedr adiationis meas ur edinJ oules per k g( 1J k g 1 is k now nas ag ray,G y) .O ne
G y is equiv alentto100r ads .R adioac tiv ity is meas ur edas thenumber ofdegr adations per s ec ondin
bequer els or Bq.O nedegr adationper s fr omr adioac tiv emater ialequals oneBq.
T heabs or beddos eofr adiationis balanc edfor damagingioniz ingtis s ueeffec t,s inc ediffer entty pes
ofr adiationhav ediffer entdens ity ofioniz ation.Ados eequiv alentter meds iev er t( Sv ) c aus es a

r ather lar gedamage.T heannualbac k gr oundr adiationis 2.5milli Sv ( 2.5mSv ) .N on penetr ating
r adiationoc c ur s fr omalpha andbetapar tic les .T hes epar tic les ar es toppedby paper ,butw henthey
enter atis s ue s uc has thebonemar r ow they s tay ther eands poilev er y thing.
Penetr atingr adiationc ons is ts ofeither gammar ay s ( neutr on) or X r ay s .Sur v iv or s fr omnuc lear
pow er plantac c idents ,w ithw holebody abs or ptiongr eater than100r ads ,ar ethr eatenedby ac ute
andc hr onic r adiations ic k nes s .
Ac uter adiations ic k nes s appear s as v omitingandmalais efollow ingex pos ur eto1G y ( 100r ad) or
mor e.Ly mphoc y tepr oduc tionis r educ edimmediately ,s oonfollow edby leuc openiaand
thr omboc y topeniaw ithbleeding.T hev illiofthegas tr ointes tinaltr ac tar edes tr oy ed,abs or ptionof
nutr ients is impair ed,andnew v illous c ells ar enotpr oduc ed.D iar r hoea,oftenw ithbloodlos s ,r es ults
indehy dr ationandanaemia.T hes k inis r edandblis ter ing,andthehair is loos ed.T he
immunodefic ienc y s y s temis des tr oy ed,ands ec ondar y infec tions hav eahighmor tality es pec ially
pulmonar y infec tions .C er ebr aloedemamay k illthev ic timw ithinhour s w henex pos edto35G y or
mor e.
C ontaminationw ithr adioac tiv eiodineis tr eatedby immediateintak eofpotas s iumiodide,w hic hbloc k
themajor par tofthethy r oidabs or ption.T hetr eatmentofs er ious ly ex pos edr adiationv ic tims is
s uppor tiv eandfr us tr ane.
C hr onic r adiations ic k nes s or later adiationdamageimplies aninc r eas edr ateofmutagenes is ,w hic h
inc ludes ahighfr equenc y ofleuc aemia,c anc er ofthebr ain,thethy r oidandthes aliv ar y glands ,
infer tility andc atar ac t( ie,aney edis eas ew her ethev is ionis blur r edby anopaquelens ) .T he
r adiationliber ates lar geamounts ofhighly r eac tiv eions inthec ells .T her eac tiv eions r uptur eD N A
s tr ands andc aus emutations w ithpr oduc tionofc anc er c ells .C anc er c ells multiply ex ponentially and
their ener gy demandappr oac hthetotalnutr itiv eener gy av ailable,c aus ingmalnutr itionanddeath.
T henuc lear pow er plantac c identinT jer noby l,U k r aine,hadmany v ic tims fr omr adiationex pos ur e.
T henear by tow n,Pr ipatja,pr ev ious ly w ith40,000r es idents is now abandoned.
T hefr equenc y ofthy r oidc anc er amongc hildr eninU k r aineis mor ethan100times theex pec ted.
Mos tofthec hildr enandy oungper s ons ex pos edhav edev elopedleuc aemiaor c anc er .T heir air w ay
epitheliumandr es pir ationis s er ious ly affec ted,andthey hav eanunus ualhighfr equenc y ofc er ebr al
haemor r hage.

Equations
T hemeanbody temper atur e( T b o d y ) is c alc ulatedw iththefollow ingequation:
Eq .21 1: T b o d y =( 0.7T c o r e +0.3T s h e l l ) ,
w her ethemeans helltemper atur e( T s h e l l ) is es timatedfr omas er ies ofr epr es entativ es k in
temper atur es .T heobv ious as s umptioninthis equationis that70% ofthebody w eightis c or e( T c o r e ) ,
andthebalanc eis s hell,butthes iz eofthes hellv ar ies w iththeenv ir onmentaltemper atur e.
T hes tor ageofheatener gy inthebody is c alc ulatedas follow s :
Eq .21 2:H S T O R E =3.49*BO D YWEIG H T *( T 2 T 1 )
w iththeunitk J per hour .
Ac c or dingtothefir s tlaw ofther mody namic s ,thefollow ingr elationis v alid:
Eq .21 3:H S T O R E =H M E T A B O L I C ( H R A D I A T I O N +H C O N +H E V A P O R A T I O N )
w her eH C O N is thec hangeinheatlos s by c onv ec tionandc onduc tion.
R adiativ eheatlos s fr omaw ar mobjec t( T o b j ) c anbec alc ulatedfor anak edper s onw ithak now n
s k intemper atur e( T s k i n ) :
Eq .21 4:H R A D I A T I O N =0.5A( T s k i n T o b j )
w her e0.5is k J min 1 m 2 K 1 ,Ais thear eaofthehumanbody ( r adiatingor r ec eiv ingr adiation) ,and
T o b j is thetemper atur eoftheobjec tex c hangingener gy .T his equationis anappr ox imationofStefan

Boltz manns r ule.


Ev apor ationofw ater onthebody s ur fac eeliminates 2430J g 1 .Ev apor ationoflar gev olume
r ates ofs w eat( V s w e a t ) implies as ubs tantiallos s ofev apor ativ eener gy ( J /min) ac c or dingtothe
equation:
Eq .21 5: H E V A P O R A T I O N ( J min 1 ) =2430( J g 1 ) V s w e a t ( gmin 1 ) .
C hanges inheatc onduc tionthr oughthes hell( c onduc tanc e s h e l l ) r eflec tc hanges ins k in
bloodflow .T hec onduc tanc ec anbec alc ulatedby thefor mula:
Eq .21 6:C onduc tanc e s h e l l =Metabolic R ate/( T c o r e T s h e l l ) .
Alar getemper atur ediffer enc eimplies aneffec tiv eis olation,w her eas as malldiffer enc eimplies alow
is olationc apac ity .
T heheatlos s ofanak edper s onr es tinginquietair by c onv ec tion( inc ludingaminor par tby
c onduc tionw ithoutmov ement) is giv enby thefollow ingequation:
Eq .21 7: H C O N =0.5*( T s h e l l T a i r ) ink J per min.
T his equationis anappr ox imationofN ew tons law ofc ooling.

SelfAssessment
Mu lt ip leC h o iceQ u est io n s
Each o f t h ef o llo w in g f ivest at em en t sh aveT ru e/F alseo p t io n s:
A .H umans c ans ur v iv eatatemper atur ethatw ouldc ook apiec eofs teak ,bec aus ethes teak
c annotdis s ipatec or eheatener gy .

B .H eatc onduc tanc eofair athighpr es s ur eex c eeds thatatlow pr es s ur e,w her eby mor eheat
ener gy is los tfr omthediv er s body by c onduc tionthr oughair ins ideadiv ingbelldeepunder
w ater thanatoneatmos pher eofpr es s ur e.

C .T emper atur ehomeos tas is is pr es entw henheatener gy pr oduc tionequals heatlos s .


D .Bas almetabolic r ate( BMR ) is low er befor ethanafter ameal.

E. T heneutr alenv ir onmentaltemper atur edefines thelev el,w her ether es tingmetabolic r ateis
minimal.

II.Each o f t h ef o llo w in g f ivest at em en t sh aveT ru e/F alseo p t io n s.


A .Water induc edhy ponatr aemiais c alledminer s c r amps .

B .Subs tantialinflux ofC a 2 + tothec ells is pr obably inv olv edins omec as es ofmalignant
hy per ther mia.

C .H y pother miais afallinc or etemper atur ebelow 32 o C .

D .O nedegr adationper s ec ondfr omr adioac tiv emater ialequals onebequer els ( 1Bq) ,w hic his
als oequaltoonec ur ie.

E. Ac uter adiationr educ es leuc oc y teandthr omboc y tepr oduc tion.

C aseH ist o ryA


Amalew ithabody w eightof70k gs tops his malar iapr ophy lax is w ithpr imaquinew henleav ingthe
endemic ar ea.T hr eew eek s later ,as uddenattac k offev er inc r eas es his c or etemper atur efr om37to
40 o C w ithin30min.T heheatc apac ity ofthehumanbody is 3.47k J /k gandper o C .T hemetabolic
r ateofthes ubjec tinc r eas eds ubs tantially dur ingther is eintemper atur e.F ollow ingthec olds tage
w ithunc ontr ollables hiv er ing,thepatientdev elops adelir ious c onditionw iths ev er eheadac he.T w o
hour s later thepatientdev elops apr ofus es w eating.Par tially ev apor ationofthew ater in32ml
s w eat/min( 25% ev apor ates ) oc c ur s fr omthebody s ur fac e( eliminating2,436J /g) ,dur ingw hic hbody
temper atur edr ops .

C alc ulatetheex tr aheatener gy s tor edinhis body after 30min.


1. C alc ulatethes malles tpos s iblemetabolic r ateinthe30 minper iod.
2. Whatc aus es theex tr aheatener gy s tor edinthebody ?
3. C alc ulatether educ tioninbody temper atur eby inges tionofoneLofic ew ater ,w henthe
fev er is atits highes tlev el.
4. C alc ulatethetimeittak es tolos etheac c umulatedheatener gy by ev apor ation.
C aseH ist o ryB
Amales edentar y per s on,w eight70k g,has adaily foodintak eof400gc ar bohy dr ate( 17.5k J /g) ,
100gfat( 39k J /g) ,and100gpr otein( 17k J /g) .T her eis ametabolic w ater for mationof32mg/k J .
T hemanex c r etes 2100mlofw ater ( i.e.,1200mlintheur ine,100mlinfaec es ,and800mlthr ough
lungs ands k in) .
1.C alc ulatethemetabolic r ate( ink J /day or MJ /day ) .
2.T hemanis inw ater balanc eby aw ater intak eof1,700mlas atotal.Ex plainthis appar ent
imbalanc e.
C aseH ist o ryC
A79 y ear oldmaleis foundappar ently deadinthes now follow ingaw inter s tor m,w her ealltr affic
w as ar r es tedby s now .H is mus c les ar es tiff,andthehear tr ateis notpalpable.T hetendonr eflex es
ar edepr es s ed,andthepupillar y andother br ains temr eflex es ar elos t.
T hebody is plac edinac hapelatthehos pitaluntilthefuner al.T henex tday theper s onnelar e
dis tur bedby nois es fr omthec hapel.O bv ious ly ,themanis aliv e.
1.Whathas aw ak enedtheman?
2.Sugges talik ely c or etemper atur e,atthetimew her ethemanw as admittedtothehos pital.
C aseH ist o ryD
A20 y ear oldper s on,w ithabody s ur fac ear eaof1.8m 2 ,is atr es tw ithametabolic r ate( MR ,or
heatener gy pr oduc tionandtr ans fer ) of80Watts .H is r ec taltemper atur eis 37 o C andhis means k in
temper atur eis 33 o C . Suddenly ,amalar iaattac k dev elops w itha7 foldr is einMR ,andthepatient
r eac hatemper atur eplateauof40 o C ,w ithameans k intemper atur eof34 o C .
T hec onduc tanc eofthes hell( C s h e l l ) is ameas ur eofs k inbloodflow .T his c anbec alc ulatedby the
for mula:C s h e l l =MR /( T c o r e T s h e l l ) .
1C alc ulatethec onduc tanc eofthes hellatr es t.
2.C alc ulatethec onduc tanc eofthes hellatthefev er plateau.
3.H ow does thebody ac c omplis hthis inc r eas e?
T r y tos olv ethepr oblems befor elook ingupthean sw ers..

Highlights
T h eco ret em p erat u reis 37 o C inhealthy adults atr es t,buts mallc hildr enhav elar ger diur nal
v ar iations .
Paren t ss houldbeadv is edtomeas ur ether ec taltemper atur eindis eas es us pec tc hildr en.T he
r ec taltemper atur eis ar eliablees timateofthec or etemper atur einr es tingper s ons .
A co n st an t co ret em p erat u redoes notguar anteeheatener gy balanc e,bec aus etheener gy
c ontentofthes hellmay c hangew iththes ur r oundings .
T h eh u m an co ret em p erat u refalls dur ingtheoes tr ogenphas eofthemens tr ualc y c leand
dur ings leep.T helow es ttemper atur eis betw een18atnightand6oc loc k inthemor ning.
Pr oges ter oneeffec ts s eemtoex plainthehigher temper atur einthelas tphas eofthemens tr ual
c y c le

T h et em p erat u recycleis par tofthec ir c adianper iodic ity .O ur biologic alc loc k s eems tobe
s y nc hr onis edw ithther otationoftheglobe.Als omeals ,lightandtemper atur eplay s ar ole.
T h et h erm alco m f o rt p o in t for light c lothed,s eatedper s ons is about26 o C w henthehumidity is
50% ,30 o C for nudeper s ons ,andabout36 o C s ittinginw ater tothenec k .
Vaso d ilat at io ninw ar menv ir onments per mits inc r eas eds k inbloodflow as thec ar diac output
als obegins tor is e.T hear ter io v enous anas tomos es ar edilatated,andthebloodflow c anr is etoat
leas t10folds inhands andfeet.
Inthem et ab o licz o n e,thetotalheatlos s r is es w ithfallingtemper atur e,butbelow 5 o C thedr y
heatlos s ex c eeds themetabolic r ate,andthebody is c ooleddow n.T his is thez oneof
hy pother mia,w her ec olddeathis inev itablew ithoutther apy .
Sw eat secret io n .T hr eemillions w eatglands pr oduc es w eatatar ateofupto2litr es per hour
or mor edur ingex tr emec onditions .Suc hhighs w eatr ates leadtoc ir c ulator y failur eands hoc k .
Sw eatr es embles adiluteultr afiltr ateofplas ma.
T h ez o n eo f h yp ert h erm iabegins atanenv ir onmentaltemper atur eof37 o C inhumidair ,w her e
w es oonr eac hthemax imalc apac ity for s w eats ec r etion,andther eis aheatinflux tothebody
endinginheatdeath.
T h eh yp o t h alam u sfunc tions as ather mos tatus ingther es tofthebody tos tabilis eits ow n
temper atur e.
A d yn am icg ain syst em r es ponds c ontinuous ly tofeedbac k s ignals ,r egar dles s ofthes iz eof
thec or etemper atur e.Withr is ingtis s uetemper atur e,theneur alac tiv ity ofheats ens or s
inc r eas es linear ly ,w her eas theac tiv ity ofc olds ens or s dec r eas es
A set p o in t syst em does notr es pondtoar is eintemper atur ebefor eac er tains etpointis
r eac hed.T hes etpointis thec or etemper atur eatw hic hneither heatlos s mec hanis ms or heat
pr oduc tionmec hanis ms ar eac tiv e.
H u m an t em p erat u reco n t ro lex hibits bothdy namic gainands etpointc har ac ter is tic s .T he
c ontr ols y s temimplies w ides pr eadc utaneous anddeeps ens or s .T heir affer ents c onv er ge
tow ar ds thehy pothalamic integr ator andther mos tat.
T h et h erm o g en icresp o n setoc oldals oinv olv es anon my ogenic or non s hiv er ingc omponent
pr obably inadipoc y tes .N on s hiv er ingheatpr oduc tionis c ontr olledby thes y mpathetic ner v ous
s y s temv iaadr ener gic r ec eptor s .
N o n sh iverin g t h erm o g en esisis any r is einmetabolis m,w hic his notr elatedtomus c ular
ac tiv ity .Inbabies this for mofther mogenes is is par tic ular ly lar geintheir br ow nadipos etis s ue.
T hebr ow nadipos etis s ueofbabies is abundantar oundv italor gans ,innec k andmedias tinum,
betw eens c apulaeandinthear mpits .
N ew b o rn sar einther malbalanc eataminimalmetabolis monly w henthes ur r ounding
temper atur eis high( 32 34 o C ) .
T h erm o reg u lat o ryb eh avio u rs uc has fanningandaddingor r emov ingc lothingis effec tiv ein
c hangingthether malins ulation.Sev er allay er s ofc lothingw ithtr appedair ar egoodins ulator s .
H eat acclim at iz at io n is ac tually as w eatglandadaptation,anda2 w eek pr oc es s follow ing
ar r iv altoahotc limate.G r adually s w eat ev apor ationis inc r eas edandtheN aC llos s is r educ ed.
T headaptationis c aus edby inc r eas edaldos ter ones ec r etionfr omtheadr enalc or tex .
H yp o vo laem iaw ithlow br ainbloodflow thr oughanov er heatedbr ainleads tofatigue( heat
ex haus tion) ,c onfus ionandunc ons c ious nes s or s y nc ope( heats y nc ope) .T hec onfus ionmay
dev elopintoav er itabledelir iumw ithbr ainoedema.
H eat st ro ke inthes unc alleds uns tr ok e is heatc ollaps ew ithhighbr aintemper atur ethat
oc c ur s s uddenly ,her eby c r eatingalife thr eateningc ondition.
H yp o t h erm iais afallinc or etemper atur etov alues below 35 o C .H y pother mic s ubjec ts los e
c ons c ious nes s ,w henthec or etemper atur efalls below 32 o C apotentially lethalc onditionc alled
s ev er ehy pother mia.

A rt if iciallyin d u ced h yp o t h erm iais us edinbr ain andhear t s ur ger y ,w her etheus ual
ther moc ontr olis inac tiv atedby gener alanaes thes ia.
Pen et rat in g rad iat io n c ons is ts ofeither gammar ay s ( neutr ons ) or X r ay s .Sur v iv or s fr om
nuc lear pow er plantac c idents w ithw holebody abs or ptiongr eater than100r ads ar ethr eatenedby
ac uteandc hr onic r adiations ic k nes s .
A cu t erad iat io n sickn essappear s as v omitingandmalais efollow ingex pos ur eto1G y ( 100
r ad) or mor e.Ly mphoc y tepr oduc tionis r educ edimmediately ,s oonfollow edby leuc openiaand
thr omboc y topeniaw ithbleeding.T hev illiofthegas tr ointes tinaltr ac tar edes tr oy ed,abs or ptionof
nutr ients is impair ed,andnew v illous c ells ar enotpr oduc ed.D iar r hoea,oftenw ithbloodlos s ,
r es ults indehy dr ation.T hes k inis r edandblis ter ing,andthehair is loos ed.
C h ro n icrad iat io n sickn essor later adiationdamageimplies aninc r eas edr ateofmutagenes is ,
w hic hinc ludes ahighfr equenc y ofleuc aemia,c anc er ofthebr ain,thethy r oidandthes aliv ar y
glands ,infer tility andc atar ac t.

FurtherReading
Sc ientific Amer ic an.Monthly jour nalpublis hedby Sc ientific Amer ic anInc .,415Madis onAv enue,N .Y.,
U SA.
Benz inger ,T .H ." T hehumanther mos tat." Sc i.Am.204:134 147,1961.
H ong,S.K.N ew s inPhy s iol.Sc i.2:79,1987.
Was s er man,D .H .andA.D .C her r ington." H epatic fuelmetabolis mdur ingmus c ular w or k :R oleand
r egulation." Am.J .Phy s iol.260:E811,1991.

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