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Fishphysiology
FromWikipedia,thefreeencyclopedia

Fishphysiologyisthescientificstudyofhowthecomponent
partsoffishfunctiontogetherinthelivingfish.[2]Itcanbe
contrastedwithfishanatomy,whichisthestudyoftheformor
morphologyoffishes.Inpractice,fishanatomyandphysiology
complementeachother,theformerdealingwiththestructure
ofafish,itsorgansorcomponentpartsandhowtheyareput
together,suchasmightbeobservedonthedissectingtableor
underthemicroscope,andthelaterdealingwithhowthose
componentsfunctiontogetherinthelivingfish.

Contents

Whenthreatened,thetoxicpufferfishfillsits
extremelyelasticstomachwithwater. [1]

1 Respiration
1.1 Bonyfish
1.2 Cartilaginousfish
1.3 Lampreysandhagfish
2 Circulation
3 Digestion
4 Endocrinesystem
5 Osmoregulation
6 Thermoregulation
7 Muscularsystem
8 Buoyancy
9 Sensorysystems
9.1 Vision
9.2 Hearing
9.3 Chemoreception
9.4 Magnetoception
9.5 Electroreception
9.6 Pain
10 Reproductiveprocesses
11 Socialbehaviour
12 Cognition
13 Seealso
14 References
15 Furtherreading
16 Externallinks

Respiration
Mostfishexchangegasesusinggillsoneithersideofthepharynx(throat).Gillsaretissueswhichconsistof
threadlikestructurescalledfilaments.Thesefilamentshavemanyfunctionsand"areinvolvedinionandwater
transferaswellasoxygen,carbondioxide,acidandammoniaexchange.[3][4]Eachfilamentcontainsacapillary
networkthatprovidesalargesurfaceareaforexchangingoxygenandcarbondioxide.Fishexchangegasesby
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pullingoxygenrichwaterthroughtheirmouthsandpumpingitovertheir
gills.Insomefish,capillarybloodflowsintheoppositedirectiontothe
water,causingcountercurrentexchange.Thegillspushtheoxygenpoor
wateroutthroughopeningsinthesidesofthepharynx.
Fishfrommultiplegroupscanliveoutofthewaterforextendedtime
periods.Amphibiousfishsuchasthemudskippercanliveandmoveabout
onlandforuptoseveraldays,orliveinstagnantorotherwiseoxygen
depletedwater.Manysuchfishcanbreatheairviaavarietyof
mechanisms.Theskinofanguillideelsmayabsorboxygendirectly.The
buccalcavityoftheelectriceelmaybreatheair.Catfishofthefamilies
Loricariidae,Callichthyidae,andScoloplacidaeabsorbairthroughtheir
digestivetracts.[5]Lungfish,withtheexceptionoftheAustralianlungfish,
andbichirshavepairedlungssimilartothoseoftetrapodsandmustsurface
togulpfreshairthroughthemouthandpassspentairoutthroughthegills.
Garandbowfinhaveavascularizedswimbladderthatfunctionsinthe
sameway.Loaches,trahiras,andmanycatfishbreathebypassingair
throughthegut.Mudskippersbreathebyabsorbingoxygenacrosstheskin
(similartofrogs).Anumberoffishhaveevolvedsocalledaccessory
breathingorgansthatextractoxygenfromtheair.Labyrinthfish(suchas
gouramisandbettas)havealabyrinthorganabovethegillsthatperforms
thisfunction.Afewotherfishhavestructuresresemblinglabyrinthorgans
informandfunction,mostnotablysnakeheads,pikeheads,andthe
Clariidaecatfishfamily.

Gillsinsidetheheadofatuna.The
headissnoutdown,withtheview
lookingtowardsthemouth.Onthe
rightarethedetachedgills.

Gillarchesbearinggillsinapike

Breathingairisprimarilyofusetofishthatinhabitshallow,seasonallyvariablewaterswherethewater'soxygen
concentrationmayseasonallydecline.Fishdependentsolelyondissolvedoxygen,suchasperchandcichlids,
quicklysuffocate,whileairbreatherssurviveformuchlonger,insomecasesinwaterthatislittlemorethanwet
mud.Atthemostextreme,someairbreathingfishareabletosurviveindampburrowsforweekswithoutwater,
enteringastateofaestivation(summertimehibernation)untilwaterreturns.
Airbreathingfishcanbedividedintoobligateairbreathersandfacultativeairbreathers.Obligateairbreathers,
suchastheAfricanlungfish,areobligatedtobreatheairperiodicallyortheysuffocate.Facultativeairbreathers,
suchasthecatfishHypostomusplecostomus,onlybreatheairiftheyneedtoandcanotherwiserelyontheirgills
foroxygen.Mostairbreathingfisharefacultativeairbreathersthatavoidtheenergeticcostofrisingtothesurface
andthefitnesscostofexposuretosurfacepredators.[5]
Allbasalvertebratesbreathewithgills.Thegillsarecarriedrightbehindthehead,borderingtheposteriormargins
ofaseriesofopeningsfromtheesophagustotheexterior.Eachgillissupportedbyacartilagenousorbonygill
arch.[6]Thegillsofvertebratestypicallydevelopinthewallsofthepharynx,alongaseriesofgillslitsopeningto
theexterior.Mostspeciesemployacountercurrentexchangesystemtoenhancethediffusionofsubstancesinand
outofthegill,withbloodandwaterflowinginoppositedirectionstoeachother.
Thegillsarecomposedofcomblikefilaments,thegilllamellae,whichhelpincreasetheirsurfaceareaforoxygen
exchange.[7]Whenafishbreathes,itdrawsinamouthfulofwateratregularintervals.Thenitdrawsthesidesof
itsthroattogether,forcingthewaterthroughthegillopenings,sothatitpassesoverthegillstotheoutside.The
bonyfishhavethreepairsofarches,cartilaginousfishhavefivetosevenpairs,whiletheprimitivejawlessfish
haveseven.Thevertebrateancestornodoubthadmorearches,assomeoftheirchordaterelativeshavemorethan
50pairsofgills.[8]

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Highervertebratesdonotdevelopgills,thegillarchesformduringfetaldevelopment,andlaythebasisofessential
structuressuchasjaws,thethyroidgland,thelarynx,thecolumella(correspondingtothestapesinmammals)and
inmammalsthemalleusandincus.[8]Fishgillslitsmaybetheevolutionaryancestorsofthetonsils,thymusgland,
andEustachiantubes,aswellasmanyotherstructuresderivedfromtheembryonicbranchialpouches.
Scientistshaveinvestigatedwhatpartofthebodyisresponsibleformaintainingtherespiratoryrhythm.They
foundthatneuronslocatedinthebrainstemoffishareresponsibleforthegenesisoftherespiratoryrhythm.[9]The
positionoftheseneuronsisslightlydifferentfromthecentersofrespiratorygenesisinmammalsbuttheyare
locatedinthesamebraincompartment,whichhascauseddebatesaboutthehomologyofrespiratorycenters
betweenaquaticandterrestrialspecies.Inbothaquaticandterrestrialrespiration,theexactmechanismsbywhich
neuronscangeneratethisinvoluntaryrhythmarestillnotcompletelyunderstood(seeInvoluntarycontrolof
respiration).
Anotherimportantfeatureoftherespiratoryrhythmisthatitismodulatedtoadapttotheoxygenconsumptionof
thebody.Asobservedinmammals,fish"breathe"fasterandheavierwhentheydophysicalexercise.The
mechanismsbywhichthesechangesoccurhavebeenstronglydebatedovermorethan100yearsbetween
scientists.[10]Theauthorscanbeclassifiedin2schools:
1.Thosewhothinkthatthemajorpartoftherespiratorychangesarepreprogrammedinthebrain,whichwould
implythatneuronsfromlocomotioncentersofthebrainconnecttorespiratorycentersinanticipationof
movements.
2.Thosewhothinkthatthemajorpartoftherespiratorychangesresultfromthedetectionofmusclecontraction,
andthatrespirationisadaptedasaconsequenceofmuscularcontractionandoxygenconsumption.Thiswould
implythatthebrainpossessessomekindofdetectionmechanismsthatwouldtriggerarespiratoryresponsewhen
muscularcontractionoccurs.
Manynowagreethatbothmechanismsareprobablypresentandcomplementary,orworkingalongsidea
mechanismthatcandetectchangesinoxygenand/orcarbondioxidebloodsaturation.

Bonyfish
Inbonyfish,thegillslieinabranchialchambercoveredbyabonyoperculum.Thegreatmajorityofbonyfish
specieshavefivepairsofgills,althoughafewhavelostsomeoverthecourseofevolution.Theoperculumcanbe
importantinadjustingthepressureofwaterinsideofthepharynxtoallowproperventilationofthegills,sothat
bonyfishdonothavetorelyonramventilation(andhencenearconstantmotion)tobreathe.Valvesinsidethe
mouthkeepthewaterfromescaping.[8]
Thegillarchesofbonyfishtypicallyhavenoseptum,sothatthegillsaloneprojectfromthearch,supportedby
individualgillrays.Somespeciesretaingillrakers.Thoughallbutthemostprimitivebonyfishlackaspiracle,the
pseudobranchassociatedwithitoftenremains,beinglocatedatthebaseoftheoperculum.Thisis,however,often
greatlyreduced,consistingofasmallmassofcellswithoutanyremaininggilllikestructure.[8]
Marineteleostsalsousegillstoexcreteelectrolytes.Thegills'largesurfaceareatendstocreateaproblemforfish
thatseektoregulatetheosmolarityoftheirinternalfluids.Saltwaterislessdilutethantheseinternalfluids,so
saltwaterfishloselargequantitiesofwaterosmoticallythroughtheirgills.Toregainthewater,theydrinklarge
amountsofseawaterandexcretethesalt.Freshwaterismoredilutethantheinternalfluidsoffish,however,so
freshwaterfishgainwaterosmoticallythroughtheirgills.[8]

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Insomeprimitivebonyfishesandamphibians,thelarvaebearexternalgills,branchingofffromthegillarches.[11]
Thesearereducedinadulthood,theirfunctiontakenoverbythegillsproperinfishesandbylungsinmost
amphibians.Someamphibiansretaintheexternallarvalgillsinadulthood,thecomplexinternalgillsystemasseen
infishapparentlybeingirrevocablylostveryearlyintheevolutionoftetrapods.[12]

Cartilaginousfish
Likeotherfish,sharksextractoxygenfromseawaterasitpassesovertheirgills.Unlikeotherfish,sharkgillslits
arenotcovered,butlieinarowbehindthehead.Amodifiedslitcalledaspiracleliesjustbehindtheeye,which
assiststhesharkwithtakinginwaterduringrespirationandplaysamajorroleinbottomdwellingsharks.
Spiraclesarereducedormissinginactivepelagicsharks.[13]Whilethesharkismoving,waterpassesthroughthe
mouthandoverthegillsinaprocessknownas"ramventilation".Whileatrest,mostsharkspumpwaterovertheir
gillstoensureaconstantsupplyofoxygenatedwater.Asmallnumberofspecieshavelosttheabilitytopump
waterthroughtheirgillsandmustswimwithoutrest.Thesespeciesareobligateramventilatorsandwould
presumablyasphyxiateifunabletomove.Obligateramventilationisalsotrueofsomepelagicbonyfish
species.[14]
Therespirationandcirculationprocessbeginswhendeoxygenatedbloodtravelstotheshark'stwochambered
heart.Herethesharkpumpsbloodtoitsgillsviatheventralaortaarterywhereitbranchesintoafferentbrachial
arteries.Reoxygenationtakesplaceinthegillsandthereoxygenatedbloodflowsintotheefferentbrachialarteries,
whichcometogethertoformthedorsalaorta.Thebloodflowsfromthedorsalaortathroughoutthebody.The
deoxygenatedbloodfromthebodythenflowsthroughtheposteriorcardinalveinsandenterstheposteriorcardinal
sinuses.Fromtherebloodenterstheheartventricleandthecyclerepeats.[15]
Sharksandraystypicallyhavefivepairsofgillslitsthatopendirectlytotheoutsideofthebody,thoughsome
moreprimitivesharkshavesixorsevenpairs.Adjacentslitsareseparatedbyacartilaginousgillarchfromwhich
projectsalongsheetlikeseptum,partlysupportedbyafurtherpieceofcartilagecalledthegillray.Theindividual
lamellaeofthegillslieoneithersideoftheseptum.Thebaseofthearchmayalsosupportgillrakers,small
projectingelementsthathelptofilterfoodfromthewater.[8]
Asmalleropening,thespiracle,liesinthebackofthefirstgillslit.Thisbearsasmallpseudobranchthatresembles
agillinstructure,butonlyreceivesbloodalreadyoxygenatedbythetruegills.[8]Thespiracleisthoughttobe
homologoustotheearopeninginhighervertebrates.[16]
Mostsharksrelyonramventilation,forcingwaterintothemouthandoverthegillsbyrapidlyswimmingforward.
Inslowmovingorbottomdwellingspecies,especiallyamongskatesandrays,thespiraclemaybeenlarged,and
thefishbreathesbysuckingwaterthroughthisopening,insteadofthroughthemouth.[8]
Chimaerasdifferfromothercartilagenousfish,havinglostboththespiracleandthefifthgillslit.Theremaining
slitsarecoveredbyanoperculum,developedfromtheseptumofthegillarchinfrontofthefirstgill.[8]

Lampreysandhagfish
Lampreysandhagfishdonothavegillslitsassuch.Instead,thegillsarecontainedinsphericalpouches,witha
circularopeningtotheoutside.Likethegillslitsofhigherfish,eachpouchcontainstwogills.Insomecases,the
openingsmaybefusedtogether,effectivelyforminganoperculum.Lampreyshavesevenpairsofpouches,while

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hagfishesmayhavesixtofourteen,dependingonthespecies.Inthehagfish,thepouchesconnectwiththepharynx
internally.Inadultlampreys,aseparaterespiratorytubedevelopsbeneaththepharynxproper,separatingfoodand
waterfromrespirationbyclosingavalveatitsanteriorend.[8]

Circulation
Thecirculatorysystemsofallvertebratesareclosed,justasinhumans.
Still,thesystemsoffish,amphibians,reptiles,andbirdsshowvarious
stagesoftheevolutionofthecirculatorysystem.Infish,thesystemhas
onlyonecircuit,withthebloodbeingpumpedthroughthecapillariesofthe
gillsandontothecapillariesofthebodytissues.Thisisknownassingle
cyclecirculation.Theheartoffishisthereforeonlyasinglepump
(consistingoftwochambers).Fishhaveaclosedloopcirculatorysystem.
Twochamberedheartofafish
Theheartpumpsthebloodinasingleloopthroughoutthebody.Inmost
fish,theheartconsistsoffourparts,includingtwochambersandan
entranceandexit.[17]Thefirstpartisthesinusvenosus,athinwalledsacthatcollectsbloodfromthefish'sveins
beforeallowingittoflowtothesecondpart,theatrium,whichisalargemuscularchamber.Theatriumservesasa
onewayantechamber,sendsbloodtothethirdpart,ventricle.Theventricleisanotherthickwalled,muscular
chamberanditpumpstheblood,firsttothefourthpart,bulbusarteriosus,alargetube,andthenoutoftheheart.
Thebulbusarteriosusconnectstotheaorta,throughwhichbloodflowstothegillsforoxygenation.
Inamphibiansandmostreptiles,adoublecirculatorysystemisused,buttheheartisnotalwayscompletely
separatedintotwopumps.Amphibianshaveathreechamberedheart.

Digestion
Jawsallowfishtoeatawidevarietyoffood,includingplantsandotherorganisms.Fishingestfoodthroughthe
mouthandbreakitdownintheesophagus.Inthestomach,foodisfurtherdigestedand,inmanyfish,processedin
fingershapedpouchescalledpyloriccaeca,whichsecretedigestiveenzymesandabsorbnutrients.Organssuchas
theliverandpancreasaddenzymesandvariouschemicalsasthefoodmovesthroughthedigestivetract.The
intestinecompletestheprocessofdigestionandnutrientabsorption.
Inmostvertebrates,digestionisafourstageprocessinvolvingthemainstructuresofthedigestivetract,starting
withingestion,placingfoodintothemouth,andconcludingwiththeexcretionofundigestedmaterialthroughthe
anus.Fromthemouth,thefoodmovestothestomach,whereasbolusitisbrokendownchemically.Itthenmoves
totheintestine,wheretheprocessofbreakingthefooddownintosimplemoleculescontinuesandtheresultsare
absorbedasnutrientsintothecirculatoryandlymphaticsystem.
Althoughthepreciseshapeandsizeofthestomachvarieswidelyamongdifferentvertebrates,therelative
positionsoftheoesophagealandduodenalopeningsremainrelativelyconstant.Asaresult,theorganalways
curvessomewhattotheleftbeforecurvingbacktomeetthepyloricsphincter.However,lampreys,hagfishes,
chimaeras,lungfishes,andsometeleostfishhavenostomachatall,withtheoesophagusopeningdirectlyintothe
intestine.Theseanimalsallconsumedietsthateitherrequirelittlestorageoffood,ornopredigestionwithgastric
juices,orboth.[8]
Thesmallintestineisthepartofthedigestivetractfollowingthestomachandfollowedbythelargeintestine,and
iswheremuchofthedigestionandabsorptionoffoodtakesplace.Infish,thedivisionsofthesmallintestineare
notclear,andthetermsanteriororproximalintestinemaybeusedinsteadofduodenum.[18]Thesmallintestineis
foundinallteleosts,althoughitsformandlengthvaryenormouslybetweenspecies.Inteleosts,itisrelatively
short,typicallyaroundoneandahalftimesthelengthofthefish'sbody.Itcommonlyhasanumberofpyloric
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caeca,smallpouchlikestructuresalongitslengththathelptoincreasetheoverallsurfaceareaoftheorganfor
digestingfood.Thereisnoileocaecalvalveinteleosts,withtheboundarybetweenthesmallintestineandthe
rectumbeingmarkedonlybytheendofthedigestiveepithelium.[8]
Thereisnosmallintestineassuchinnonteleostfish,suchassharks,sturgeons,andlungfish.Instead,thedigestive
partofthegutformsaspiralintestine,connectingthestomachtotherectum.Inthistypeofgut,theintestineitself
isrelativelystraight,buthasalongfoldrunningalongtheinnersurfaceinaspiralfashion,sometimesfordozens
ofturns.Thisvalvegreatlyincreasesboththesurfaceareaandtheeffectivelengthoftheintestine.Theliningof
thespiralintestineissimilartothatofthesmallintestineinteleostsandnonmammaliantetrapods.[8]Inlampreys,
thespiralvalveisextremelysmall,possiblybecausetheirdietrequireslittledigestion.Hagfishhavenospiralvalve
atall,withdigestionoccurringforalmosttheentirelengthoftheintestine,whichisnotsubdividedintodifferent
regions.[8]
Thelargeintestineisthelastpartofthedigestivesystemnormallyfoundinvertebrateanimals.Itsfunctionisto
absorbwaterfromtheremainingindigestiblefoodmatter,andthentopassuselesswastematerialfromthe
body.[19]Infish,thereisnotruelargeintestine,butsimplyashortrectumconnectingtheendofthedigestivepart
oftheguttothecloaca.Insharks,thisincludesarectalglandthatsecretessalttohelptheanimalmaintainosmotic
balancewiththeseawater.Theglandsomewhatresemblesacaecuminstructure,butisnotahomologous
structure.[8]
Aswithmanyaquaticanimals,mostfishreleasetheirnitrogenouswastesasammonia.Someofthewastesdiffuse
throughthegills.Bloodwastesarefilteredbythekidneys.
Saltwaterfishtendtolosewaterbecauseofosmosis.Theirkidneysreturnwatertothebody.Thereversehappens
infreshwaterfish:theytendtogainwaterosmotically.Theirkidneysproducediluteurineforexcretion.Somefish
havespeciallyadaptedkidneysthatvaryinfunction,allowingthemtomovefromfreshwatertosaltwater.
Insharks,digestioncantakealongtime.ThefoodmovesfromthemouthtoaJshapedstomach,whereitisstored
andinitialdigestionoccurs.[20]Unwanteditemsmaynevergetpastthestomach,andinsteadthesharkeither
vomitsorturnsitsstomachsinsideoutandejectsunwanteditemsfromitsmouth.Oneofthebiggestdifferences
betweenthedigestivesystemsofsharksandmammalsisthatsharkshavemuchshorterintestines.Thisshort
lengthisachievedbythespiralvalvewithmultipleturnswithinasingleshortsectioninsteadofalongtubelike
intestine.Thevalveprovidesalongsurfacearea,requiringfoodtocirculateinsidetheshortgutuntilfully
digested,whenremainingwasteproductspassintothecloaca.[20]

Endocrinesystem
Osmoregulation
Twomajortypesofosmoregulationareosmoconformersandosmoregulators.Osmoconformersmatchtheirbody
osmolaritytotheirenvironmentactivelyorpassively.Mostmarineinvertebratesareosmoconformers,although
theirioniccompositionmaybedifferentfromthatofseawater.
Osmoregulatorstightlyregulatetheirbodyosmolarity,whichalwaysstaysconstant,andaremorecommoninthe
animalkingdom.Osmoregulatorsactivelycontrolsaltconcentrationsdespitethesaltconcentrationsinthe
environment.Anexampleisfreshwaterfish.Thegillsactivelyuptakesaltfromtheenvironmentbytheuseof
mitochondriarichcells.Waterwilldiffuseintothefish,soitexcretesaveryhypotonic(dilute)urinetoexpelall
theexcesswater.Amarinefishhasaninternalosmoticconcentrationlowerthanthatofthesurroundingseawater,
soittendstolosewaterandgainsalt.Itactivelyexcretessaltoutfromthegills.Mostfisharestenohaline,which
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meanstheyarerestrictedtoeithersaltorfreshwaterandcannot
surviveinwaterwithadifferentsaltconcentrationthantheyare
adaptedto.However,somefishshowatremendousabilityto
effectivelyosmoregulateacrossabroadrangeofsalinitiesfish
withthisabilityareknownaseuryhalinespecies,e.g.,salmon.
Salmonhasbeenobservedtoinhabittwoutterlydisparate
environmentsmarineandfreshwateranditisinherentto
adapttobothbybringinginbehavioralandphysiological
modifications.
Incontrasttobonyfish,withtheexceptionofthecoelacanth,[21]
thebloodandothertissueofsharksandChondrichthyesis
generallyisotonictotheirmarineenvironmentsbecauseofthe
highconcentrationofureaandtrimethylamineNoxide
(TMAO),allowingthemtobeinosmoticbalancewiththe
seawater.Thisadaptationpreventsmostsharksfromsurvivingin
freshwater,andtheyarethereforeconfinedtomarine
environments.Afewexceptionsexist,suchasthebullshark,
whichhasdevelopedawaytochangeitskidneyfunctionto
excretelargeamountsofurea.[22]Whenasharkdies,theureais
brokendowntoammoniabybacteria,causingthedeadbodyto
graduallysmellstronglyofammonia.[23][24]

Movementofwaterandionsinsaltwaterfish

Movementofwaterandionsinfreshwaterfish

Sharkshaveadoptedadifferent,efficientmechanismtoconservewater,i.e.,osmoregulation.Theyretainureain
theirbloodinrelativelyhigherconcentration.Ureaisdamagingtolivingtissueso,tocopewiththisproblem,some
fishretaintrimethylamineoxide.Thisprovidesabettersolutiontourea'stoxicity.Sharks,havingslightlyhigher
soluteconcentration(i.e.,above1000mOsmwhichisseasoluteconcentration),donotdrinkwaterlikefreshwater
fish.

Thermoregulation
Homeothermyandpoikilothermyrefertohowstableanorganism'stemperatureis.Mostendothermicorganisms
arehomeothermic,likemammals.However,animalswithfacultativeendothermyareoftenpoikilothermic,
meaningtheirtemperaturecanvaryconsiderably.Similarly,mostfishareectotherms,asalloftheirheatcomes
fromthesurroundingwater.However,mostarehomeothermsbecausetheirtemperatureisverystable.
Mostorganismshaveapreferredtemperaturerange,howeversomecanbeacclimatedtotemperaturescolderor
warmerthanwhattheyaretypicallyusedto.Anorganism'spreferredtemperatureistypicallythetemperatureat
whichtheorganism'sphysiologicalprocessescanactatoptimalrates.Whenfishbecomeacclimatedtoother
temperatures,theefficiencyoftheirphysiologicalprocessesmaydecreasebutwillcontinuetofunction.Thisis
calledthethermalneutralzoneatwhichanorganismcansurviveindefinitely.[25]
H.M.Vernonhasdoneworkonthedeathtemperatureandparalysistemperature(temperatureofheatrigor)of
variousanimals.Hefoundthatspeciesofthesameclassshowedverysimilartemperaturevalues,thosefromthe
Amphibiaexaminedbeing38.5C,fish39C,Reptilia45C,andvariousMolluscs46C.
Tocopewithlowtemperatures,somefishhavedevelopedtheabilitytoremainfunctionalevenwhenthewater
temperatureisbelowfreezingsomeusenaturalantifreezeorantifreezeproteinstoresisticecrystalformationin
theirtissues.

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Mostsharksare"coldblooded"or,moreprecisely,poikilothermic,meaningthattheirinternalbodytemperature
matchesthatoftheirambientenvironment.MembersofthefamilyLamnidae(suchastheshortfinmakosharkand
thegreatwhiteshark)arehomeothermicandmaintainahigherbodytemperaturethanthesurroundingwater.In
thesesharks,astripofaerobicredmusclelocatednearthecenterofthebodygeneratestheheat,whichthebody
retainsviaacountercurrentexchangemechanismbyasystemofbloodvesselscalledtheretemirabile
("miraculousnet").Thecommonthreshersharkhasasimilarmechanismformaintaininganelevatedbody
temperature,whichisthoughttohaveevolvedindependently.[26]
Tunacanmaintainthetemperatureofcertainpartsoftheirbodyabovethetemperatureofambientseawater.For
example,bluefintunamaintainacorebodytemperatureof2533C(7791F),inwaterascoldas6C(43F).
However,unliketypicalendothermiccreaturessuchasmammalsandbirds,tunadonotmaintaintemperature
withinarelativelynarrowrange.[27][28]Tunaachieveendothermybyconservingtheheatgeneratedthroughnormal
metabolism.Theretemirabile("wonderfulnet"),theintertwiningofveinsandarteriesinthebody'speriphery,
transfersheatfromvenousbloodtoarterialbloodviaacountercurrentexchangesystem,thusmitigatingthe
effectsofsurfacecooling.Thisallowsthetunatoelevatethetemperaturesofthehighlyaerobictissuesofthe
skeletalmuscles,eyesandbrain,[27][29]whichsupportsfasterswimmingspeedsandreducedenergyexpenditure,
andwhichenablesthemtosurviveincoolerwatersoverawiderrangeofoceanenvironmentsthanthoseofother
fish.[28]Inalltunas,however,theheartoperatesatambienttemperature,asitreceivescooledblood,andcoronary
circulationisdirectlyfromthegills.[29]
Homeothermy:Althoughmostfishareexclusivelyectothermic,thereareexceptions.Certainspeciesoffish
maintainelevatedbodytemperatures.Endothermicteleosts(bonyfish)areallinthesuborderScombroidei
andincludethebillfishes,tunas,includinga"primitive"mackerelspecies,Gasterochismamelampus.All
sharksinthefamilyLamnidaeshortfinmako,longfinmako,white,porbeagle,andsalmonsharkare
endothermic,andevidencesuggeststhetraitexistsinfamilyAlopiidae(threshersharks).Thedegreeof
endothermyvariesfromthebillfish,whichwarmonlytheireyesandbrain,tobluefintunaandporbeagle
sharkswhomaintainbodytemperatureselevatedinexcessof20Caboveambientwatertemperatures.[30]
Seealsogigantothermy.Endothermy,thoughmetabolicallycostly,isthoughttoprovideadvantagessuchas
increasedmusclestrength,higherratesofcentralnervoussystemprocessing,andhigherratesofdigestion.
Insomefish,aretemirabileallowsforanincreaseinmuscletemperatureinregionswherethisnetworkofvein
andarteriesisfound.Thefishisabletothermoregulatecertainareasoftheirbody.Additionally,thisincreasein
temperatureleadstoanincreaseinbasalmetabolictemperature.ThefishisnowabletosplitATPatahigherrate
andultimatelycanswimfaster.
Theeyeofaswordfishcangenerateheattobettercopewithdetectingtheirpreyatdepthsof2000feet.[31]

Muscularsystem
Fishswimbycontractinglongitudinalredmuscleandobliquelyorientedwhitemuscles.Theredmuscleisaerobic
andneedsoxygenwhichissuppliedbymyoglobin.Thewhitemuscleisanaerobicanditdoesnotneedoxygen.
Redmusclesareusedforsustainedactivitysuchascruisingatslowspeedsonoceanmigrations.Whitemusclesare
usedforburstsofactivity,suchasjumpingorsuddenburstsofspeedforcatchingprey.[32]
Mostlyfishhavewhitemuscles,butthemusclesofsomefishes,suchasscombroidsandsalmonids,rangefrom
pinktodarkred.Theredmyotomalmusclesderivetheircolourfrommyoglobin,anoxygenbindingmolecule,
whichtunaexpressinquantitiesfarhigherthanmostotherfish.Theoxygenrichbloodfurtherenablesenergy
deliverytotheirmuscles.[27]

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Mostfishmovebyalternatelycontractingpairedsetsofmusclesoneither
sideofthebackbone.ThesecontractionsformSshapedcurvesthatmove
downthebody.Aseachcurvereachesthebackfin,backwardforceis
appliedtothewater,andinconjunctionwiththefins,movesthefish
forward.Thefish'sfinsfunctionlikeanairplane'sflaps.Finsalsoincrease
thetail'ssurfacearea,increasingspeed.Thestreamlinedbodyofthefish
decreasestheamountoffrictionfromthewater.
Atypicalcharacteristicofmanyanimalsthatutilizeundulatorylocomotion
isthattheyhavesegmentedmuscles,orblocksofmyomeres,runningfrom
theirheadtotailswhichareseparatedbyconnectivetissuecalled
myosepta.Inaddition,somesegmentedmusclegroups,suchthelateral
hypaxialmusculatureinthesalamanderareorientedatanangletothe
Crosssectionofateleost
longitudinaldirection.Fortheseobliquelyorientedfibersthestraininthe
longitudinaldirectionisgreaterthanthestraininthemusclefiberdirection
leadingtoanarchitecturalgearratiogreaterthan1.Ahigherinitialangleof
orientationandmoredorsoventralbulgingproducesafastermuscle
contractionbutresultsinaloweramountofforceproduction.[33]Itis
hypothesizedthatanimalsemployavariablegearingmechanismthat
allowsselfregulationofforceandvelocitytomeetthemechanical
demandsofthecontraction.[34]Whenapennatemuscleissubjectedtoa
Iridescentsharkfiletsshowing
lowforce,resistancetowidthchangesinthemusclecauseittorotatewhich
myomerestructure
consequentlyproduceahigherarchitecturalgearratio(AGR)(high
velocity).[34]However,whensubjecttoahighforce,theperpendicularfiber
forcecomponentovercomestheresistancetowidthchangesandthemusclecompressesproducingalowerAGR
(capableofmaintainingahigherforceoutput).[34]
Mostfishesbendasasimple,homogenousbeamduringswimmingviacontractionsoflongitudinalredmuscle
fibersandobliquelyorientedwhitemusclefiberswithinthesegmentedaxialmusculature.Thefiberstrain(f)
experiencedbythelongitudinalredmusclefibersisequivalenttothelongitudinalstrain(x).Thedeeperwhite
musclefibersfishesshowdiversityinarrangement.Thesefibersareorganizedintoconeshapedstructuresand
attachtoconnectivetissuesheetsknownasmyoseptaeachfibershowsacharacteristicdorsoventral()and
mediolateral()trajectory.Thesegmentedarchitecturetheorypredictsthat,x>f.Thisphenomenonresultsinan
architecturalgearratio,determinedaslongitudinalstraindividedbyfiberstrain(x/f),greaterthanoneand
longitudinalvelocityamplificationfurthermore,thisemergentvelocityamplificationmaybeaugmentedby
variablearchitecturalgearingviamesolateralanddorsoventralshapechanges,apatternseeninpennatemuscle
contractions.Aredtowhitegearingratio(redf/whitef)capturesthecombinedeffectofthelongitudinalred
musclefiberandobliquewhitemusclefiberstrains.[33][35]

Buoyancy
Thebodyofafishisdenserthanwater,sofishmustcompensateforthe
differenceortheywillsink.Manybonyfisheshaveaninternalorgancalled
aswimbladder,orgasbladder,thatadjuststheirbuoyancythrough
manipulationofgases.Inthisway,fishcanstayatthecurrentwaterdepth,
orascendordescendwithouthavingtowasteenergyinswimming.The
Swimbladderofacommonrudd
bladderisonlyfoundinbonyfishes.Inthemoreprimitivegroupslike
someminnows,bichirsandlungfish,thebladderisopentotheesophagus
anddoubleasalung.Itisoftenabsentinfastswimmingfishessuchasthetunaandmackerelfamilies.The
conditionofabladderopentotheesophagusiscalledphysostome,theclosedconditionphysoclist.Inthelatter,the
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gascontentofthebladderiscontrolledthroughtheretemirabilis,anetwork
ofbloodvesselseffectinggasexchangebetweenthebladderandthe
blood.[36]
Insomefish,aretemirabilefillstheswimbladderwithoxygen.A
countercurrentexchangesystemisutilizedbetweenthevenousandarterial
capillaries.ByloweringthepHlevelsinthevenouscapillaries,oxygen
unbindsfrombloodhemoglobin.Thiscausesanincreaseinvenousblood
oxygenconcentration,allowingtheoxygentodiffusethroughthecapillary
membraneandintothearterialcapillaries,whereoxygenisstillsequestered
tohemoglobin.Thecycleofdiffusioncontinuesuntiltheconcentrationof
oxygeninthearterialcapillariesissupersaturated(largerthanthe
concentrationofoxygenintheswimbladder).Atthispoint,thefree
oxygeninthearterialcapillariesdiffusesintotheswimbladderviathegas
gland.[37]

Sharks,likethisthreetonnegreat
whiteshark,don'thaveswim
bladders.Mostsharksneedtokeep
swimmingtoavoidsinking.

Unlikebonyfish,sharksdonothavegasfilledswimbladdersforbuoyancy.Instead,sharksrelyonalargeliver
filledwithoilthatcontainssqualene,andtheircartilage,whichisabouthalfthenormaldensityofbone.[38]Their
liverconstitutesupto30%oftheirtotalbodymass.[22]Theliver'seffectivenessislimited,sosharksemploy
dynamiclifttomaintaindepthwhennotswimming.Sandtigersharksstoreairintheirstomachs,usingitasaform
ofswimbladder.Mostsharksneedtoconstantlyswiminordertobreatheandcannotsleepverylongwithout
sinking(ifatall).However,certainspecies,likethenurseshark,arecapableofpumpingwateracrosstheirgills,
allowingthemtorestontheoceanbottom.[39]

Sensorysystems
Mostfishpossesshighlydevelopedsenseorgans.Nearlyalldaylightfishhavecolorvisionthatisatleastasgood
asahuman's(seevisioninfishes).Manyfishalsohavechemoreceptorsthatareresponsibleforextraordinary
sensesoftasteandsmell.Althoughtheyhaveears,manyfishmaynothearverywell.Mostfishhavesensitive
receptorsthatformthelaterallinesystem,whichdetectsgentlecurrentsandvibrations,andsensesthemotionof
nearbyfishandprey.[40]Sharkscansensefrequenciesintherangeof25to50Hzthroughtheirlateralline.[41]
Fishorientthemselvesusinglandmarksandmayusementalmapsbasedonmultiplelandmarksorsymbols.Fish
behaviorinmazesrevealsthattheypossessspatialmemoryandvisualdiscrimination.[42]

Vision
Visionisanimportantsensorysystemformostspeciesoffish.Fisheyesaresimilartothoseofterrestrial
vertebrateslikebirdsandmammals,buthaveamoresphericallens.Theirretinasgenerallyhavebothrodcellsand
conecells(forscotopicandphotopicvision),andmostspecieshavecolourvision.Somefishcanseeultraviolet
andsomecanseepolarizedlight.Amongstjawlessfish,thelampreyhaswelldevelopedeyes,whilethehagfish
hasonlyprimitiveeyespots.[43]Fishvisionshowsadaptationtotheirvisualenvironment,forexampledeepsea
fisheshaveeyessuitedtothedarkenvironment.

Hearing
Hearingisanimportantsensorysystemformostspeciesoffish.Hearingthresholdandtheabilitytolocalizesound
sourcesarereducedunderwater,inwhichthespeedofsoundisfasterthaninair.Underwaterhearingisbybone
conduction,andlocalizationofsoundappearstodependondifferencesinamplitudedetectedbybone
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conduction.[44]Aquaticanimalssuchasfish,however,haveamorespecializedhearingapparatusthatiseffective
underwater.[45]
Fishcansensesoundthroughtheirlaterallinesandtheirotoliths(ears).Somefishes,suchassomespeciesofcarp
andherring,hearthroughtheirswimbladders,whichfunctionratherlikeahearingaid.[46]
Hearingiswelldevelopedincarp,whichhavetheWeberianorgan,threespecializedvertebralprocessesthat
transfervibrationsintheswimbladdertotheinnerear.
Althoughitishardtotestsharks'hearing,theymayhaveasharpsenseofhearingandcanpossiblyhearpreymany
milesaway.[47]Asmallopeningoneachsideoftheirheads(notthespiracle)leadsdirectlyintotheinnerear
throughathinchannel.Thelaterallineshowsasimilararrangement,andisopentotheenvironmentviaaseriesof
openingscalledlaterallinepores.Thisisareminderofthecommonoriginofthesetwovibrationandsound
detectingorgansthataregroupedtogetherastheacousticolateralissystem.Inbonyfishandtetrapodstheexternal
openingintotheinnerearhasbeenlost.

Chemoreception
Sharkshavekeenolfactorysenses,locatedintheshortduct(whichis
notfused,unlikebonyfish)betweentheanteriorandposteriornasal
openings,withsomespeciesabletodetectaslittleasonepartper
millionofbloodinseawater.[48]
Sharkshavetheabilitytodeterminethedirectionofagivenscentbased
onthetimingofscentdetectionineachnostril.[49]Thisissimilartothe
methodmammalsusetodeterminedirectionofsound.
Theyaremoreattractedtothechemicalsfoundintheintestinesofmany
species,andasaresultoftenlingernearorinsewageoutfalls.Some
species,suchasnursesharks,haveexternalbarbelsthatgreatlyincrease
theirabilitytosenseprey.

Theshapeofthehammerheadshark's
headmayenhanceolfactionbyspacing
thenostrilsfurtherapart.

Magnetoception
Electroreception
Somefish,suchascatfishandsharks,haveorgansthatdetectweakelectriccurrentsontheorderofmillivolt.[50]
Otherfish,liketheSouthAmericanelectricfishesGymnotiformes,canproduceweakelectriccurrents,whichthey
useinnavigationandsocialcommunication.Insharks,theampullaeofLorenziniareelectroreceptororgans.They
numberinthehundredstothousands.SharksusetheampullaeofLorenzinitodetecttheelectromagneticfieldsthat
alllivingthingsproduce.[51]Thishelpssharks(particularlythehammerheadshark)findprey.Thesharkhasthe
greatestelectricalsensitivityofanyanimal.Sharksfindpreyhiddeninsandbydetectingtheelectricfieldsthey
produce.OceancurrentsmovinginthemagneticfieldoftheEarthalsogenerateelectricfieldsthatsharkscanuse
fororientationandpossiblynavigation.[52]
TheampullaeofLorenziniallowsharkstosenseelectricaldischarges.
Electricfishareabletoproduceelectricfieldsbymodifiedmusclesintheirbody.

Pain
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ExperimentsdonebyWilliamTavolgaprovideevidencethat
fishhavepainandfearresponses.Forinstance,inTavolgas
experiments,toadfishgruntedwhenelectricallyshockedand
overtimetheycametogruntatthemeresightofan
electrode.[53]
In2003,ScottishscientistsattheUniversityofEdinburghand
theRoslinInstituteconcludedthatrainbowtroutexhibit
behaviorsoftenassociatedwithpaininotheranimals.Bee
venomandaceticacidinjectedintothelipsresultedinfish
rockingtheirbodiesandrubbingtheirlipsalongthesidesand
floorsoftheirtanks,whichtheresearchersconcludedwere
attemptstorelievepain,similartowhatmammalswould
do.[54][55][56]Neuronsfiredinapatternresemblinghuman
neuronalpatterns.[56]

Electromagneticfieldreceptors(ampullaeof
Lorenzini)andmotiondetectingcanalsinthehead
ofashark

ProfessorJamesD.RoseoftheUniversityofWyomingclaimedthestudywasflawedsinceitdidnotprovide
proofthatfishpossess"consciousawareness,particularlyakindofawarenessthatismeaningfullylikeours".[57]
Rosearguesthatsincefishbrainsaresodifferentfromhumanbrains,fishareprobablynotconsciousinthe
mannerhumansare,sothatreactionssimilartohumanreactionstopaininsteadhaveothercauses.Rosehad
publishedastudyayearearlierarguingthatfishcannotfeelpainbecausetheirbrainslackaneocortex.[58]
However,animalbehavioristTempleGrandinarguesthatfishcouldstillhaveconsciousnesswithoutaneocortex
because"differentspeciescanusedifferentbrainstructuresandsystemstohandlethesamefunctions."[56]
Animalwelfareadvocatesraiseconcernsaboutthepossiblesufferingoffishcausedbyangling.Somecountries,
suchasGermanyhavebannedspecifictypesoffishing,andtheBritishRSPCAnowformallyprosecutes
individualswhoarecrueltofish.[59]

Reproductiveprocesses
Seealso:Fishreproduction,Spawning,Ichthyoplankton
Oogoniadevelopmentinteleostsfishvariesaccordingtothegroup,andthe
determinationofoogenesisdynamicsallowstheunderstandingof
maturationandfertilisationprocesses.Changesinthenucleus,ooplasm,
andthesurroundinglayerscharacterizetheoocytematurationprocess.[60]
Postovulatoryfolliclesarestructuresformedafteroocytereleasetheydo
nothaveendocrinefunction,presentawideirregularlumen,andarerapidly
reabsorbedinaprocessinvolvingtheapoptosisoffollicularcells.A
degenerativeprocesscalledfollicularatresiareabsorbsvitellogenicoocytes
notspawned.Thisprocesscanalsooccur,butlessfrequently,inoocytesin
otherdevelopmentstages.[60]
Somefisharehermaphrodites,havingbothtestesandovarieseitherat
differentphasesintheirlifecycleor,asinhamlets,havethem
simultaneously.

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Typicaleggofabonyfish,about1
mmindiameter.Fisheggsareoften
releasedintothewatercolumn,where
theydriftaszooplankton

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Over97%ofallknownfishareoviparous,[61]thatis,theeggsdevelopoutsidethemother'sbody.Examplesof
oviparousfishincludesalmon,goldfish,cichlids,tuna,andeels.Inthemajorityofthesespecies,fertilisationtakes
placeoutsidethemother'sbody,withthemaleandfemalefishsheddingtheirgametesintothesurroundingwater.
However,afewoviparousfishpracticeinternalfertilisation,withthemaleusingsomesortofintromittentorganto
deliverspermintothegenitalopeningofthefemale,mostnotablytheoviparoussharks,suchasthehornshark,
andoviparousrays,suchasskates.Inthesecases,themaleisequippedwithapairofmodifiedpelvicfinsknown
asclaspers.
Marinefishcanproducehighnumbersofeggswhichareoftenreleasedintotheopenwatercolumn.Theeggshave
anaveragediameterof1millimetre(0.039in).Theeggsaregenerallysurroundedbytheextraembryonic
membranesbutdonotdevelopashell,hardorsoft,aroundthesemembranes.Somefishhavethick,leatherycoats,
especiallyiftheymustwithstandphysicalforceordesiccation.Thesetypeofeggscanalsobeverysmalland
fragile.

Eggoflamprey

Eggofcatshark
(mermaids'purse)

Eggofbullheadshark

Eggofchimaera

Thenewlyhatchedyoungofoviparousfisharecalledlarvae.Theyareusuallypoorlyformed,carryalargeyolk
sac(fornourishment)andareverydifferentinappearancefromjuvenileandadultspecimens.Thelarvalperiodin
oviparousfishisrelativelyshort(usuallyonlyseveralweeks),andlarvaerapidlygrowandchangeappearanceand
structure(aprocesstermedmetamorphosis)tobecomejuveniles.Duringthistransitionlarvaemustswitchfrom
theiryolksactofeedingonzooplanktonprey,aprocesswhichdependsontypicallyinadequatezooplankton
density,starvingmanylarvae.
Inovoviviparousfishtheeggsdevelopinsidethemother'sbodyafterinternalfertilisationbutreceivelittleorno
nourishmentdirectlyfromthemother,dependinginsteadontheyolk.Eachembryodevelopsinitsownegg.
Familiarexamplesofovoviviparousfishincludeguppies,angelsharks,andcoelacanths.
Somespeciesoffishareviviparous.Insuchspeciesthemotherretainstheeggsandnourishestheembryos.
Typically,viviparousfishhaveastructureanalogoustotheplacentaseeninmammalsconnectingthemother's
bloodsupplywiththatoftheembryo.Examplesofviviparousfishincludethesurfperches,splitfins,andlemon
shark.Someviviparousfishexhibitoophagy,inwhichthedevelopingembryoseatothereggsproducedbythe
mother.Thishasbeenobservedprimarilyamongsharks,suchastheshortfinmakoandporbeagle,butisknownfor
afewbonyfishaswell,suchasthehalfbeakNomorhamphusebrardtii.[62]Intrauterinecannibalismisaneven
moreunusualmodeofvivipary,inwhichthelargestembryoseatweakerandsmallersiblings.Thisbehaviorisalso
mostcommonlyfoundamongsharks,suchasthegreynurseshark,buthasalsobeenreportedforNomorhamphus
ebrardtii.[62]
Inmanyspeciesoffish,finshavebeenmodifiedtoallowInternalfertilisation.
Aquaristscommonlyrefertoovoviviparousandviviparousfishaslivebearers.
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Manyfishspeciesarehermaphrodites.Synchronoushermaphroditespossessbothovariesandtestesatthe
sametime.Sequentialhermaphroditeshavebothtypesoftissueintheirgonads,withonetypebeing
predominantwhilethefishbelongstothecorrespondinggender.

Socialbehaviour
Oxytocinisagroupofneuropeptidesfoundinmostvertebrate.Oneformofoxytocinfunctionsasahormone
whichisassociatedwithhumanlove.In2012,researchersinjectedcichlidsfromthesocialspecies
Neolamprologuspulcher,eitherwiththisformofisotocinorwithacontrolsalinesolution.Theyfoundisotocin
increased"responsivenesstosocialinformation",whichsuggests"itisakeyregulatorofsocialbehaviorthathas
evolvedandenduredsinceancienttimes".[63][64]

Cognition
Seealso
Anatomicaltermsoflocation
DigitalFishLibrary
Evolutionoffish
Fishanatomy
Fishdevelopment
Fishmeasurement

Fishmeasurement
Ichthyologyterms
Panderichthysdigits

References
1.Schwab,IvanR(2002)."Morethanjustcoolshades".BritishJournalofOphthalmology86:1075.
doi:10.1136/bjo.86.10.1075.
2.Prosser,C.Ladd(1991).ComparativeAnimalPhysiology,EnvironmentalandMetabolicAnimalPhysiology(4thed.).
Hoboken,NJ:WileyLiss.pp.112.ISBN047185767X.
3.HoarWSandRandallDJ(1984)FishPhysiology:Gills:PartAAnatomy,gastransferandacidbaseregulation(htt
p://books.google.co.nz/books?id=yINDnV4mWi8C&pg=PA263&dq=%22FISH+PHYSIOLOGY+V10A%22&hl=en&sa=
X&ei=BPwUpeILMbOkAWXl4DAAQ&ved=0CCwQ6AEwAA#v=onepage&q=%22FISH%20PHYSIOLOGY%20V10
A%22&f=false)AcademicPress.ISBN9780080585314.
4.HoarWSandRandallDJ(1984)FishPhysiology:Gills:PartBIonandwatertransfer(http://books.google.co.nz/boo
ks?id=8fuEB7O3IqIC&pg=PA380&lpg=PA380&dq=%22Fish+physiology%22&source=bl&ots=MeKDBPDZIi&sig=2K
FLDRsgMXpQU0i_Ufj75aC0hzw&hl=en&sa=X&ei=RRHwUs7EFcnGlAWIz4GgCw&ved=0CLkBEOgBMBU#v=onep
age&q=%22Fish%20physiology%22&f=false)AcademicPress.ISBN9780080585321.
5."ModificationsoftheDigestiveTractforHoldingAirinLoricariidandScoloplacidCatfishes"(PDF).Copeia(3):663
675.1998.doi:10.2307/1447796.Retrieved25June2009.
6.Scott,Thomas(1996).Conciseencyclopediabiology.WalterdeGruyter.p.542.ISBN9783110106619.
7.Andrews,ChrisAdrianExellNevilleCarrington(2003).ManualOfFishHealth.FireflyBooks.
8.Romer,AlfredSherwoodParsons,ThomasS.(1977).TheVertebrateBody.Philadelphia,PA:HoltSaunders
International.pp.316327.ISBN003910284X.
9.Rovainen,1985(http://www.springerlink.com/content/j1q45g379420w127/)
10.Waldrop,IwamotoandHaouzi,2006(http://jap.physiology.org/cgi/content/full/100/3/1077)
11."TheOriginoftheLarvaandMetamorphosisinAmphibia".TheAmericanNaturalist(EssexInstitute)91:287.1957.
doi:10.1086/281990.JSTOR2458911.
12.Clack,J.A.(2002):Gainingground:theoriginandevolutionoftetrapods.IndianaUniversityPress,Bloomington,
Indiana.369pp
13.Gilbertson,Lance(1999).ZoologyLaboratoryManual.NewYork:McGrawHillCompanies,Inc.ISBN007237716X.
14.WilliamJ.Bennetta(1996)."DeepBreathing".Retrieved20070828.
15."SHARKS&RAYS,SeaWorld/BuschGardensANIMALS,CIRCULATORYSYSTEM".BuschEntertainment
Corporation.Retrieved20090903.
https://en.wikipedia.org/wiki/Fish_physiology

14/17

7/3/2016

FishphysiologyWikipedia,thefreeencyclopedia

16.LaurinM.(1998):Theimportanceofglobalparsimonyandhistoricalbiasinunderstandingtetrapodevolution.PartI
systematics,middleearevolution,andjawsuspension.AnnalesdesSciencesNaturelles,Zoologie,Paris,13eSrie19:
pp142.
17.Setaro,JohnF.(1999).CirculatorySystem.MicrosoftEncarta99.
18.Guillaume,JeanPraxisPublishingSadasivamKaushikPierreBergotRobertMetailler(2001).NutritionandFeeding
ofFishandCrustaceans.Springer.p.31.ISBN1852332417.ISBN9781852332419.Retrieved20090109.
19.NationalCancerInstitute."NCIDictionaryofCancerTermslargeintestine".Retrieved20120916.
20.Martin,R.Aidan."NoGuts,NoGlory".ReefQuestCentreforSharkResearch.Retrieved20090822.
21.Chemistryofthebodyfluidsofthecoelacanth,Latimeriachalumnae(http://www.jstor.org/pss/35431)
22.Compagno,LeonardDando,MarcFowler,Sarah(2005).SharksoftheWorld.CollinsFieldGuides.ISBN000
7136102.OCLC183136093.
23.JohnA.Musick(2005)."Managementtechniquesforelasmobranchfisheries:14.SharkUtilization".FAO:Fisheriesand
AquacultureDepartment.Retrieved20080316.
24.ThomasBatten."MAKOSHARKIsurusoxyrinchus".DelawareSeaGrant,UniversityofDelaware.Retrieved
20080316.
25.Wilmer,PatStone,GrahamJohnston,Ian(2009).EnvironmentalPhysiologyofAnimals.Wiley.pp.198200.
ISBN9781405107242.
26.Martin,R.Aidan(April1992)."FireintheBellyoftheBeast".ReefQuestCentreforSharkResearch.Retrieved
20090821.
27.Sepulveda,C.A.Dickson,K.A.Bernal,D.Graham,J.B.(1July2008)."Elevatedredmyotomalmuscletemperatures
inthemostbasaltunaspecies,Allothunnusfallai"(PDF).JournalofFishBiology73(1):241249.doi:10.1111/j.1095
8649.2008.01931.x.Retrieved2November2012.
28."TunaBiologyOfTuna".RetrievedSeptember12,2009.
29.LandeiraFernandez,A.M.Morrissette,J.M.Blank,J.M.Block,B.A.(16October2003)."Temperaturedependenceof
theCa2+ATPase(SERCA2)intheventriclesoftunaandmackerel".AJP:Regulatory,IntegrativeandComparative
Physiology286(2):398R404.doi:10.1152/ajpregu.00392.2003.
30.Block,BAFinnerty,JR(1993)."Endothermyinfishes:aphylogeneticanalysisofconstraints,predispositions,and
selectionpressures"(PDF).EnvironmentalBiologyofFishes40(3):283302.doi:10.1007/BF00002518.
31.DavidFleshler(10152012)SouthFloridaSunSentinel(http://www.sunsentinel.com/news/local/breakingnews/flgiante
yeballmystery20121015,0,2019024.story),
Swordfishheattheireyes(http://www.newscientist.com/article/dn6861swordfishheattheireyesforthehunt.htm
l)
32.Kapoor2004
33.Brainerd,E.L.Azizi,E.(2005)."MuscleFiberAngle,SegmentBulgingandArchitecturalGearRatioinSegmented
Musculature".JournalofExperimentalBiology208(17):32493261.doi:10.1242/jeb.01770.
34.Azizi,E.Brainerd,E.L.Roberts,T.J.(2008)."VariableGearinginPennateMuscles".PNAS105(5):17451750.
doi:10.1073/pnas.0709212105.PMC2234215.PMID18230734.
35.Brainerd,E.L.Azizi,E.(2007)."ArchitecturalGearRatioandMuscleFiberStrainHomogeneityinSegmented
Musculature".JournalofExperimentalZoology307(A):145155.doi:10.1002/jez.a.358.
36.Kardong,K.(2008).Vertebrates:Comparativeanatomy,function,evolution(5thed.).Boston:McGrawHill.ISBN007
3040584.
37.Kardong,K.(2008).Vertebrates:Comparativeanatomy,function,evolution,(5thed.).Boston:McGrawHill.
38.Martin,R.Aidan."TheImportanceofBeingCartilaginous".ReefQuestCentreforSharkResearch.Retrieved
20090829.
39."Dosharkssleep".Flmnh.ufl.edu.Retrieved20100923.
40.Orr,James(1999).Fish.MicrosoftEncarta99.ISBN0811423468.
41.Popper,A.N.C.Platt(1993)."Innerearandlateralline".ThePhysiologyofFishes(CRCPress)(1sted.).
42.JournalofUndergraduateLifeSciences."Appropriatemazemethodologytostudylearninginfish"(PDF).Retrieved
28May2009.
43.N.A.CampbellandJ.B.Reece(2005).Biology,SeventhEdition.BenjaminCummings,SanFrancisco,California.
44.ShupakA.SharoniZ.YanirY.KeynanY.AlfieY.HalpernP.(January2005)."UnderwaterHearingandSound
LocalizationwithandwithoutanAirInterface".Otology&Neurotology26(1):127130.doi:10.1097/00129492
20050100000023.
45.Graham,Michael(1941)."SenseofHearinginFishes".Nature147(3738):779.doi:10.1038/147779b0.
46.B,WILLIAMSC."SenseofHearinginFishes."Nature147.3731(n.d.):543.Print.
47.Martin,R.Aidan."HearingandVibrationDetection".Retrieved20080601.
48.Martin,R.Aidan."SmellandTaste".ReefQuestCentreforSharkResearch.Retrieved20090821.
https://en.wikipedia.org/wiki/Fish_physiology

15/17

7/3/2016

FishphysiologyWikipedia,thefreeencyclopedia

49.TheFunctionofBilateralOdorArrivalTimeDifferencesinOlfactoryOrientationofSharks(http://shell.cas.usf.edu/mott
a/Gardiner%20and%20Atema%202010.pdf),JayneM.Gardiner,JelleAtema,CurrentBiology13July2010(Vol.20,
Issue13,pp.11871191)
50.Albert,J.S.,andW.G.R.Crampton.2005.Electroreceptionandelectrogenesis.pp.431472inThePhysiologyofFishes,
3rdEdition.D.H.EvansandJ.B.Claiborne(eds.).CRCPress.
51.KalmijnAJ(1982)."Electricandmagneticfielddetectioninelasmobranchfishes".Science218(4575):9168.
doi:10.1126/science.7134985.PMID7134985.
52.MeyerCG,HollandKN,PapastamatiouYP(2005)."Sharkscandetectchangesinthegeomagneticfield".Journalofthe
RoyalSociety,Interface2(2):12930.doi:10.1098/rsif.2004.0021.PMC1578252.PMID16849172.
53.Dunayer,Joan,"Fish:SensitivityBeyondtheCaptor'sGrasp,"TheAnimals'Agenda,July/August1991,pp.1218
54.VantressaBrown,"FishFeelPain,BritishResearchersSay,"AgenceFrancePresse,1May2003(http://www.buzzle.co
m/editorials/430200339769.asp)
55.Kirby,Alex(30April2003)."Fishdofeelpain,scientistssay".BBCNews.Retrieved4January2010.
56.Grandin,TempleJohnson,Catherine(2005).AnimalsinTranslation.NewYork,NewYork:Scribner.pp.183184.
ISBN0743247698.
57."Rose,J.D.2003.ACritiqueofthepaper:"Dofishhavenociceptors:Evidencefortheevolutionofavertebratesensory
system" "(PDF).Retrieved21May2011.
58.JamesD.Rose,DoFishFeelPain?(http://www.coloradotu.org/dofishfeelpain/),2002.Retrieved27September2007.
59.Leake,J."AnglerstoFaceRSPCACheck,"TheSundayTimesBritain,14March2004(http://www.timesonline.co.uk/
newspaper/0,,1761037515,00.html)
60.GuimaraesCruz,RodrigoJ.,RodrigoJ.Santos,JosE.dosSantos,GilmarB.(JulySeptember2005)."Gonadal
structureandgametogenesisofLoricarialentiginosaIsbrcker(Pisces,Teleostei,Siluriformes)".Rev.Bras.Zool.22
(3):556564.doi:10.1590/S010181752005000300005.ISSN01018175.
61.PeterScott:LivebearingFishes,p.13.TetraPress1997.ISBN1564651932
62.Meisner,A&Burns,J:ViviparityintheHalfbeakGeneraDermogenysandNomorhamphus(Teleostei:Hemiramphidae)"
JournalofMorphology234,pp.295317,1997
63.Reddon,ARO'Connor,CMMarshRollo,SEBalcshine,S(2012)."Effectsofisotocinonsocialresponsesina
cooperativelybreedingfish".AnimalBehaviour84(4):753760.doi:10.1016/j.anbehav.2012.07.021.
64.FishVersionOfOxytocinDrivesTheirSocialBehavior(http://www.science20.com/news_articles/fish_version_oxytocin_
drives_their_social_behavior_says_study95041)Science2.0,10October2012.

Furtherreading
BernierNJ,VanDerKraakG,FarrellAPandBraunerCJ(2009)FishPhysiology:FishNeuroendocrinology
(http://books.google.com/books?id=vIOm8QXb7JEC&printsec=frontcover&dq=%22Fish+physiology%22&
hl=en&sa=X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CE8Q6AEwBg)AcademicPress.ISBN
9780080877983.
EddyFBandHandyRD(2012)EcologicalandEnvironmentalPhysiologyofFishes(http://books.google.co
m/books?id=VpWllLJ8INYC&printsec=frontcover&dq=%22Ecological+and+Environmental+Physiology+o
f+Fishes%22&hl=en&sa=X&ei=QPgBUfSoOMWHlAXK3IDoCA&ved=0CC8Q6AEwAA)Oxford
UniversityPress.ISBN9780199540952.
EvansDH,JBClaiborneandSCurrie(Eds)(2013)ThePhysiologyofFishes(https://books.google.com/boo
ks?id=KHtcAgAAQBAJ&printsec=frontcover&dq=%22The+Physiology+of+Fishes%22&hl=en&sa=X&ei
=2XoaVfiwO8yB8QX_soGwBw&ved=0CCUQ6AEwAA#v=onepage&q=%22The%20Physiology%20of%
20Fishes%22&f=false)4thedition,CRCPress.ISBN9781439880302.
GrosellM,FarrellAPandBraunerCJ(2010)FishPhysiology:TheMultifunctionalGutofFish(http://book
s.google.com/books?id=eddWw_nyRlYC&printsec=frontcover&dq=%22Fish+physiology%22&hl=en&sa=
X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CEsQ6AEwBQ)AcademicPress.ISBN9780080961361.
HaraTJandZielinskiB(2006)FishPhysiology:SensorySystemsNeuroscience(http://books.google.com/bo
oks?id=KpXBu4y4XNIC&printsec=frontcover&dq=%22Fish+physiology%22&hl=en&sa=X&ei=454PUeG
DLsGimQXCpYCIBA&ved=0CIMBEOgBMBA#v=onepage&q=%22Fish%20physiology%22&f=false)
AcademicPress.ISBN9780080469614.
KapoorBGandKhannaB(2004)"Ichthyologyhandbook"(http://books.google.com/books?id=I7WhoPBdA
ooC&printsec=frontcover&dq=%22Ichthyology+handbook%22&hl=en&sa=X&ei=mB_zTo7JBouWmQXY
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v6iXAg&ved=0CC8Q6AEwAA#v=onepage&q=%22Ichthyology%20handbook%22&f=false)Pages137
140,Springer.ISBN9783540428541.
McKenzieDJ,FarrellAPandBraunerCJ(2007)FishPhysiology:PrimitiveFishes(http://books.google.co
m/books?id=gfBc_omOIeAC&printsec=frontcover&dq=%22Fish+physiology%22&hl=en&sa=X&ei=454P
UeGDLsGimQXCpYCIBA&ved=0CF4Q6AEwCQ#v=onepage&q=%22Fish%20physiology%22&f=false)
AcademicPress.ISBN9780080549521.
SlomanKA,WilsonRWandBalshineS(2006)BehaviourAndPhysiologyofFish(http://books.google.com/
books?id=CBE0T2ADFoC&printsec=frontcover&dq=%22Fish+physiology%22&hl=en&sa=X&ei=454PU
eGDLsGimQXCpYCIBA&ved=0CGMQ6AEwCg#v=onepage&q=%22Fish%20physiology%22&f=false)
GulfProfessionalPublishing.ISBN9780123504487.
WoodCM,FarrellAPandBraunerCJ(2011)FishPhysiology:HomeostasisandToxicologyofNon
EssentialMetals(http://books.google.com/books?id=Nnh1WpHZbysC&printsec=frontcover&dq=%22Fish+
physiology%22&hl=en&sa=X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CFcQ6AEwCA#v=onepage&q
=%22Fish%20physiology%22&f=false)AcademicPress.ISBN9780123786340.

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