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FishphysiologyWikipedia,thefreeencyclopedia
Fishphysiology
FromWikipedia,thefreeencyclopedia
Fishphysiologyisthescientificstudyofhowthecomponent
partsoffishfunctiontogetherinthelivingfish.[2]Itcanbe
contrastedwithfishanatomy,whichisthestudyoftheformor
[Link],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
[Link]"areinvolvedinionandwater
transferaswellasoxygen,carbondioxide,acidandammoniaexchange.[3][4]Eachfilamentcontainsacapillary
[Link]
[Link]
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pullingoxygenrichwaterthroughtheirmouthsandpumpingitovertheir
[Link],capillarybloodflowsintheoppositedirectiontothe
water,[Link]
wateroutthroughopeningsinthesidesofthepharynx.
Fishfrommultiplegroupscanliveoutofthewaterforextendedtime
[Link]
onlandforuptoseveraldays,orliveinstagnantorotherwiseoxygen
[Link]
[Link]
[Link]
Loricariidae,Callichthyidae,andScoloplacidaeabsorbairthroughtheir
digestivetracts.[5]Lungfish,withtheexceptionoftheAustralianlungfish,
andbichirshavepairedlungssimilartothoseoftetrapodsandmustsurface
togulpfreshairthroughthemouthandpassspentairoutthroughthegills.
Garandbowfinhaveavascularizedswimbladderthatfunctionsinthe
[Link],trahiras,andmanycatfishbreathebypassingair
[Link]
(similartofrogs).Anumberoffishhaveevolvedsocalledaccessory
[Link](suchas
gouramisandbettas)havealabyrinthorganabovethegillsthatperforms
[Link]
informandfunction,mostnotablysnakeheads,pikeheads,andthe
Clariidaecatfishfamily.
[Link]
headissnoutdown,withtheview
[Link]
rightarethedetachedgills.
Gillarchesbearinggillsinapike
Breathingairisprimarilyofusetofishthatinhabitshallow,seasonallyvariablewaterswherethewater'soxygen
[Link],suchasperchandcichlids,
quicklysuffocate,whileairbreatherssurviveformuchlonger,insomecasesinwaterthatislittlemorethanwet
[Link],someairbreathingfishareabletosurviveindampburrowsforweekswithoutwater,
enteringastateofaestivation(summertimehibernation)untilwaterreturns.
[Link],
suchastheAfricanlungfish,[Link],
suchasthecatfishHypostomusplecostomus,onlybreatheairiftheyneedtoandcanotherwiserelyontheirgills
[Link]
andthefitnesscostofexposuretosurfacepredators.[5]
[Link],borderingtheposteriormargins
[Link]
arch.[6]Thegillsofvertebratestypicallydevelopinthewallsofthepharynx,alongaseriesofgillslitsopeningto
[Link]
outofthegill,withbloodandwaterflowinginoppositedirectionstoeachother.
Thegillsarecomposedofcomblikefilaments,thegilllamellae,whichhelpincreasetheirsurfaceareaforoxygen
exchange.[7]Whenafishbreathes,[Link]
itsthroattogether,forcingthewaterthroughthegillopenings,[Link]
bonyfishhavethreepairsofarches,cartilaginousfishhavefivetosevenpairs,whiletheprimitivejawlessfish
[Link],assomeoftheirchordaterelativeshavemorethan
50pairsofgills.[8]
[Link]
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Highervertebratesdonotdevelopgills,thegillarchesformduringfetaldevelopment,andlaythebasisofessential
structuressuchasjaws,thethyroidgland,thelarynx,thecolumella(correspondingtothestapesinmammals)and
inmammalsthemalleusandincus.[8]Fishgillslitsmaybetheevolutionaryancestorsofthetonsils,thymusgland,
andEustachiantubes,aswellasmanyotherstructuresderivedfromtheembryonicbranchialpouches.
[Link]
foundthatneuronslocatedinthebrainstemoffishareresponsibleforthegenesisoftherespiratoryrhythm.[9]The
positionoftheseneuronsisslightlydifferentfromthecentersofrespiratorygenesisinmammalsbuttheyare
locatedinthesamebraincompartment,whichhascauseddebatesaboutthehomologyofrespiratorycenters
[Link],theexactmechanismsbywhich
neuronscangeneratethisinvoluntaryrhythmarestillnotcompletelyunderstood(seeInvoluntarycontrolof
respiration).
Anotherimportantfeatureoftherespiratoryrhythmisthatitismodulatedtoadapttotheoxygenconsumptionof
[Link],fish"breathe"[Link]
mechanismsbywhichthesechangesoccurhavebeenstronglydebatedovermorethan100yearsbetween
scientists.[10]Theauthorscanbeclassifiedin2schools:
[Link],whichwould
implythatneuronsfromlocomotioncentersofthebrainconnecttorespiratorycentersinanticipationof
movements.
[Link],
[Link]
implythatthebrainpossessessomekindofdetectionmechanismsthatwouldtriggerarespiratoryresponsewhen
muscularcontractionoccurs.
Manynowagreethatbothmechanismsareprobablypresentandcomplementary,orworkingalongsidea
mechanismthatcandetectchangesinoxygenand/orcarbondioxidebloodsaturation.
Bonyfish
Inbonyfish,[Link]
specieshavefivepairsofgills,[Link]
importantinadjustingthepressureofwaterinsideofthepharynxtoallowproperventilationofthegills,sothat
bonyfishdonothavetorelyonramventilation(andhencenearconstantmotion)[Link]
mouthkeepthewaterfromescaping.[8]
Thegillarchesofbonyfishtypicallyhavenoseptum,sothatthegillsaloneprojectfromthearch,supportedby
[Link],the
pseudobranchassociatedwithitoftenremains,[Link],however,often
greatlyreduced,consistingofasmallmassofcellswithoutanyremaininggilllikestructure.[8]
[Link]'largesurfaceareatendstocreateaproblemforfish
[Link],so
[Link],theydrinklarge
[Link],however,so
freshwaterfishgainwaterosmoticallythroughtheirgills.[8]
[Link]
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Insomeprimitivebonyfishesandamphibians,thelarvaebearexternalgills,branchingofffromthegillarches.[11]
Thesearereducedinadulthood,theirfunctiontakenoverbythegillsproperinfishesandbylungsinmost
[Link],thecomplexinternalgillsystemasseen
infishapparentlybeingirrevocablylostveryearlyintheevolutionoftetrapods.[12]
Cartilaginousfish
Likeotherfish,[Link],sharkgillslits
arenotcovered,[Link],which
assiststhesharkwithtakinginwaterduringrespirationandplaysamajorroleinbottomdwellingsharks.
Spiraclesarereducedormissinginactivepelagicsharks.[13]Whilethesharkismoving,waterpassesthroughthe
mouthandoverthegillsinaprocessknownas"ramventilation".Whileatrest,mostsharkspumpwaterovertheir
[Link]
[Link]
[Link]
species.[14]
Therespirationandcirculationprocessbeginswhendeoxygenatedbloodtravelstotheshark'stwochambered
[Link]
[Link],
[Link]
deoxygenatedbloodfromthebodythenflowsthroughtheposteriorcardinalveinsandenterstheposteriorcardinal
[Link].[15]
Sharksandraystypicallyhavefivepairsofgillslitsthatopendirectlytotheoutsideofthebody,thoughsome
[Link]
projectsalongsheetlikeseptum,[Link]
[Link],small
projectingelementsthathelptofilterfoodfromthewater.[8]
Asmalleropening,thespiracle,[Link]
agillinstructure,butonlyreceivesbloodalreadyoxygenatedbythetruegills.[8]Thespiracleisthoughttobe
homologoustotheearopeninginhighervertebrates.[16]
Mostsharksrelyonramventilation,forcingwaterintothemouthandoverthegillsbyrapidlyswimmingforward.
Inslowmovingorbottomdwellingspecies,especiallyamongskatesandrays,thespiraclemaybeenlarged,and
thefishbreathesbysuckingwaterthroughthisopening,insteadofthroughthemouth.[8]
Chimaerasdifferfromothercartilagenousfish,[Link]
slitsarecoveredbyanoperculum,developedfromtheseptumofthegillarchinfrontofthefirstgill.[8]
Lampreysandhagfish
[Link],thegillsarecontainedinsphericalpouches,witha
[Link],[Link],the
openingsmaybefusedtogether,[Link],while
[Link]
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hagfishesmayhavesixtofourteen,[Link],thepouchesconnectwiththepharynx
[Link],aseparaterespiratorytubedevelopsbeneaththepharynxproper,separatingfoodand
waterfromrespirationbyclosingavalveatitsanteriorend.[8]
Circulation
Thecirculatorysystemsofallvertebratesareclosed,justasinhumans.
Still,thesystemsoffish,amphibians,reptiles,andbirdsshowvarious
[Link],thesystemhas
onlyonecircuit,withthebloodbeingpumpedthroughthecapillariesofthe
[Link]
[Link]
(consistingoftwochambers).Fishhaveaclosedloopcirculatorysystem.
Twochamberedheartofafish
[Link]
fish,theheartconsistsoffourparts,includingtwochambersandan
entranceandexit.[17]Thefirstpartisthesinusvenosus,athinwalledsacthatcollectsbloodfromthefish'sveins
beforeallowingittoflowtothesecondpart,theatrium,[Link]
onewayantechamber,sendsbloodtothethirdpart,[Link],muscular
chamberanditpumpstheblood,firsttothefourthpart,bulbusarteriosus,alargetube,andthenoutoftheheart.
Thebulbusarteriosusconnectstotheaorta,throughwhichbloodflowstothegillsforoxygenation.
Inamphibiansandmostreptiles,adoublecirculatorysystemisused,buttheheartisnotalwayscompletely
[Link].
Digestion
Jawsallowfishtoeatawidevarietyoffood,[Link]
[Link],foodisfurtherdigestedand,inmanyfish,processedin
fingershapedpouchescalledpyloriccaeca,[Link]
[Link]
intestinecompletestheprocessofdigestionandnutrientabsorption.
Inmostvertebrates,digestionisafourstageprocessinvolvingthemainstructuresofthedigestivetract,starting
withingestion,placingfoodintothemouth,andconcludingwiththeexcretionofundigestedmaterialthroughthe
[Link],thefoodmovestothestomach,[Link]
totheintestine,wheretheprocessofbreakingthefooddownintosimplemoleculescontinuesandtheresultsare
absorbedasnutrientsintothecirculatoryandlymphaticsystem.
Althoughthepreciseshapeandsizeofthestomachvarieswidelyamongdifferentvertebrates,therelative
[Link],theorganalways
[Link],lampreys,hagfishes,
chimaeras,lungfishes,andsometeleostfishhavenostomachatall,withtheoesophagusopeningdirectlyintothe
[Link],ornopredigestionwithgastric
juices,orboth.[8]
Thesmallintestineisthepartofthedigestivetractfollowingthestomachandfollowedbythelargeintestine,and
[Link],thedivisionsofthesmallintestineare
notclear,andthetermsanteriororproximalintestinemaybeusedinsteadofduodenum.[18]Thesmallintestineis
foundinallteleosts,[Link],itisrelatively
short,typicallyaroundoneandahalftimesthelengthofthefish'[Link]
[Link]
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caeca,smallpouchlikestructuresalongitslengththathelptoincreasetheoverallsurfaceareaoftheorganfor
[Link],withtheboundarybetweenthesmallintestineandthe
rectumbeingmarkedonlybytheendofthedigestiveepithelium.[8]
Thereisnosmallintestineassuchinnonteleostfish,suchassharks,sturgeons,[Link],thedigestive
partofthegutformsaspiralintestine,[Link],theintestineitself
isrelativelystraight,buthasalongfoldrunningalongtheinnersurfaceinaspiralfashion,sometimesfordozens
[Link]
thespiralintestineissimilartothatofthesmallintestineinteleostsandnonmammaliantetrapods.[8]Inlampreys,
thespiralvalveisextremelysmall,[Link]
atall,withdigestionoccurringforalmosttheentirelengthoftheintestine,whichisnotsubdividedintodifferent
regions.[8]
[Link]
absorbwaterfromtheremainingindigestiblefoodmatter,andthentopassuselesswastematerialfromthe
body.[19]Infish,thereisnotruelargeintestine,butsimplyashortrectumconnectingtheendofthedigestivepart
[Link],thisincludesarectalglandthatsecretessalttohelptheanimalmaintainosmotic
[Link],butisnotahomologous
structure.[8]
Aswithmanyaquaticanimals,[Link]
[Link].
[Link]
infreshwaterfish:[Link]
havespeciallyadaptedkidneysthatvaryinfunction,allowingthemtomovefromfreshwatertosaltwater.
Insharks,[Link],whereitisstored
andinitialdigestionoccurs.[20]Unwanteditemsmaynevergetpastthestomach,andinsteadthesharkeither
[Link]
[Link]
lengthisachievedbythespiralvalvewithmultipleturnswithinasingleshortsectioninsteadofalongtubelike
[Link],requiringfoodtocirculateinsidetheshortgutuntilfully
digested,whenremainingwasteproductspassintothecloaca.[20]
Endocrinesystem
Osmoregulation
[Link]
[Link],although
theirioniccompositionmaybedifferentfromthatofseawater.
Osmoregulatorstightlyregulatetheirbodyosmolarity,whichalwaysstaysconstant,andaremorecommoninthe
[Link]
[Link]
[Link],soitexcretesaveryhypotonic(dilute)urinetoexpelall
[Link],
[Link],which
[Link]
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meanstheyarerestrictedtoeithersaltorfreshwaterandcannot
surviveinwaterwithadifferentsaltconcentrationthantheyare
[Link],somefishshowatremendousabilityto
effectivelyosmoregulateacrossabroadrangeofsalinitiesfish
withthisabilityareknownaseuryhalinespecies,e.g.,salmon.
Salmonhasbeenobservedtoinhabittwoutterlydisparate
environmentsmarineandfreshwateranditisinherentto
adapttobothbybringinginbehavioralandphysiological
modifications.
Incontrasttobonyfish,withtheexceptionofthecoelacanth,[21]
thebloodandothertissueofsharksandChondrichthyesis
generallyisotonictotheirmarineenvironmentsbecauseofthe
highconcentrationofureaandtrimethylamineNoxide
(TMAO),allowingthemtobeinosmoticbalancewiththe
[Link]
freshwater,andtheyarethereforeconfinedtomarine
[Link],suchasthebullshark,
whichhasdevelopedawaytochangeitskidneyfunctionto
excretelargeamountsofurea.[22]Whenasharkdies,theureais
brokendowntoammoniabybacteria,causingthedeadbodyto
graduallysmellstronglyofammonia.[23][24]
Movementofwaterandionsinsaltwaterfish
Movementofwaterandionsinfreshwaterfish
Sharkshaveadoptedadifferent,efficientmechanismtoconservewater,i.e.,[Link]
[Link],tocopewiththisproblem,some
[Link]'[Link],havingslightlyhigher
soluteconcentration(i.e.,above1000mOsmwhichisseasoluteconcentration),donotdrinkwaterlikefreshwater
fish.
Thermoregulation
Homeothermyandpoikilothermyrefertohowstableanorganism'[Link]
arehomeothermic,[Link],animalswithfacultativeendothermyareoftenpoikilothermic,
[Link],mostfishareectotherms,asalloftheirheatcomes
[Link],mostarehomeothermsbecausetheirtemperatureisverystable.
Mostorganismshaveapreferredtemperaturerange,howeversomecanbeacclimatedtotemperaturescolderor
[Link]'spreferredtemperatureistypicallythetemperatureat
whichtheorganism'[Link]
temperatures,[Link]
calledthethermalneutralzoneatwhichanorganismcansurviveindefinitely.[25]
[Link](temperatureofheatrigor)of
[Link],thosefromthe
Amphibiaexaminedbeing38.5C,fish39C,Reptilia45C,andvariousMolluscs46C.
Tocopewithlowtemperatures,somefishhavedevelopedtheabilitytoremainfunctionalevenwhenthewater
temperatureisbelowfreezingsomeusenaturalantifreezeorantifreezeproteinstoresisticecrystalformationin
theirtissues.
[Link]
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Mostsharksare"coldblooded"or,moreprecisely,poikilothermic,meaningthattheirinternalbodytemperature
[Link](suchastheshortfinmakosharkand
thegreatwhiteshark)[Link]
thesesharks,astripofaerobicredmusclelocatednearthecenterofthebodygeneratestheheat,whichthebody
retainsviaacountercurrentexchangemechanismbyasystemofbloodvesselscalledtheretemirabile
("miraculousnet").Thecommonthreshersharkhasasimilarmechanismformaintaininganelevatedbody
temperature,whichisthoughttohaveevolvedindependently.[26]
[Link]
example,bluefintunamaintainacorebodytemperatureof2533C(7791F),inwaterascoldas6C(43F).
However,unliketypicalendothermiccreaturessuchasmammalsandbirds,tunadonotmaintaintemperature
withinarelativelynarrowrange.[27][28]Tunaachieveendothermybyconservingtheheatgeneratedthroughnormal
[Link]("wonderfulnet"),theintertwiningofveinsandarteriesinthebody'speriphery,
transfersheatfromvenousbloodtoarterialbloodviaacountercurrentexchangesystem,thusmitigatingthe
[Link]
skeletalmuscles,eyesandbrain,[27][29]whichsupportsfasterswimmingspeedsandreducedenergyexpenditure,
andwhichenablesthemtosurviveincoolerwatersoverawiderrangeofoceanenvironmentsthanthoseofother
fish.[28]Inalltunas,however,theheartoperatesatambienttemperature,asitreceivescooledblood,andcoronary
circulationisdirectlyfromthegills.[29]
Homeothermy:Althoughmostfishareexclusivelyectothermic,[Link]
[Link](bonyfish)areallinthesuborderScombroidei
andincludethebillfishes,tunas,includinga"primitive"mackerelspecies,[Link]
sharksinthefamilyLamnidaeshortfinmako,longfinmako,white,porbeagle,andsalmonsharkare
endothermic,andevidencesuggeststhetraitexistsinfamilyAlopiidae(threshersharks).Thedegreeof
endothermyvariesfromthebillfish,whichwarmonlytheireyesandbrain,tobluefintunaandporbeagle
sharkswhomaintainbodytemperatureselevatedinexcessof20Caboveambientwatertemperatures.[30]
[Link],thoughmetabolicallycostly,isthoughttoprovideadvantagessuchas
increasedmusclestrength,higherratesofcentralnervoussystemprocessing,andhigherratesofdigestion.
Insomefish,aretemirabileallowsforanincreaseinmuscletemperatureinregionswherethisnetworkofvein
[Link],thisincreasein
[Link]
andultimatelycanswimfaster.
Theeyeofaswordfishcangenerateheattobettercopewithdetectingtheirpreyatdepthsof2000feet.[31]
Muscularsystem
[Link]
[Link].
[Link]
usedforburstsofactivity,suchasjumpingorsuddenburstsofspeedforcatchingprey.[32]
Mostlyfishhavewhitemuscles,butthemusclesofsomefishes,suchasscombroidsandsalmonids,rangefrom
[Link],anoxygenbindingmolecule,
[Link]
deliverytotheirmuscles.[27]
[Link]
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Mostfishmovebyalternatelycontractingpairedsetsofmusclesoneither
[Link]
[Link],backwardforceis
appliedtothewater,andinconjunctionwiththefins,movesthefish
[Link]'sfinsfunctionlikeanairplane'[Link]
thetail'ssurfacearea,[Link]
decreasestheamountoffrictionfromthewater.
Atypicalcharacteristicofmanyanimalsthatutilizeundulatorylocomotion
isthattheyhavesegmentedmuscles,orblocksofmyomeres,runningfrom
theirheadtotailswhichareseparatedbyconnectivetissuecalled
[Link],somesegmentedmusclegroups,suchthelateral
hypaxialmusculatureinthesalamanderareorientedatanangletothe
Crosssectionofateleost
[Link]
longitudinaldirectionisgreaterthanthestraininthemusclefiberdirection
[Link]
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
[Link](f)
experiencedbythelongitudinalredmusclefibersisequivalenttothelongitudinalstrain(x).Thedeeperwhite
[Link]
attachtoconnectivetissuesheetsknownasmyoseptaeachfibershowsacharacteristicdorsoventral()and
mediolateral()[Link],x>[Link]
architecturalgearratio,determinedaslongitudinalstraindividedbyfiberstrain(x/f),greaterthanoneand
longitudinalvelocityamplificationfurthermore,thisemergentvelocityamplificationmaybeaugmentedby
variablearchitecturalgearingviamesolateralanddorsoventralshapechanges,apatternseeninpennatemuscle
[Link](redf/whitef)capturesthecombinedeffectofthelongitudinalred
musclefiberandobliquewhitemusclefiberstrains.[33][35]
Buoyancy
Thebodyofafishisdenserthanwater,sofishmustcompensateforthe
[Link]
aswimbladder,orgasbladder,thatadjuststheirbuoyancythrough
[Link],fishcanstayatthecurrentwaterdepth,
[Link]
Swimbladderofacommonrudd
[Link]
someminnows,bichirsandlungfish,thebladderisopentotheesophagus
[Link]
conditionofabladderopentotheesophagusiscalledphysostome,[Link],the
[Link]
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gascontentofthebladderiscontrolledthroughtheretemirabilis,anetwork
ofbloodvesselseffectinggasexchangebetweenthebladderandthe
blood.[36]
Insomefish,aretemirabilefillstheswimbladderwithoxygen.A
countercurrentexchangesystemisutilizedbetweenthevenousandarterial
[Link],oxygen
[Link]
oxygenconcentration,allowingtheoxygentodiffusethroughthecapillary
membraneandintothearterialcapillaries,whereoxygenisstillsequestered
[Link]
oxygeninthearterialcapillariesissupersaturated(largerthanthe
concentrationofoxygenintheswimbladder).Atthispoint,thefree
oxygeninthearterialcapillariesdiffusesintotheswimbladderviathegas
gland.[37]
Sharks,likethisthreetonnegreat
whiteshark,don'thaveswim
[Link]
swimmingtoavoidsinking.
Unlikebonyfish,[Link],sharksrelyonalargeliver
filledwithoilthatcontainssqualene,andtheircartilage,whichisabouthalfthenormaldensityofbone.[38]Their
liverconstitutesupto30%oftheirtotalbodymass.[22]Theliver'seffectivenessislimited,sosharksemploy
[Link],usingitasaform
[Link]
sinking(ifatall).However,certainspecies,likethenurseshark,arecapableofpumpingwateracrosstheirgills,
allowingthemtorestontheoceanbottom.[39]
Sensorysystems
[Link]
asahuman's(seevisioninfishes).Manyfishalsohavechemoreceptorsthatareresponsibleforextraordinary
[Link],[Link]
receptorsthatformthelaterallinesystem,whichdetectsgentlecurrentsandvibrations,andsensesthemotionof
nearbyfishandprey.[40]Sharkscansensefrequenciesintherangeof25to50Hzthroughtheirlateralline.[41]
[Link]
behaviorinmazesrevealsthattheypossessspatialmemoryandvisualdiscrimination.[42]
Vision
[Link]
vertebrateslikebirdsandmammals,[Link]
conecells(forscotopicandphotopicvision),[Link]
[Link],thelampreyhaswelldevelopedeyes,whilethehagfish
hasonlyprimitiveeyespots.[43]Fishvisionshowsadaptationtotheirvisualenvironment,forexampledeepsea
fisheshaveeyessuitedtothedarkenvironment.
Hearing
[Link]
sourcesarereducedunderwater,[Link]
conduction,andlocalizationofsoundappearstodependondifferencesinamplitudedetectedbybone
[Link]
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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
[Link],andisopentotheenvironmentviaaseriesof
[Link]
[Link]
openingintotheinnerearhasbeenlost.
Chemoreception
Sharkshavekeenolfactorysenses,locatedintheshortduct(whichis
notfused,unlikebonyfish)betweentheanteriorandposteriornasal
openings,withsomespeciesabletodetectaslittleasonepartper
millionofbloodinseawater.[48]
Sharkshavetheabilitytodeterminethedirectionofagivenscentbased
onthetimingofscentdetectionineachnostril.[49]Thisissimilartothe
methodmammalsusetodeterminedirectionofsound.
Theyaremoreattractedtothechemicalsfoundintheintestinesofmany
species,[Link]
species,suchasnursesharks,haveexternalbarbelsthatgreatlyincrease
theirabilitytosenseprey.
Theshapeofthehammerheadshark's
headmayenhanceolfactionbyspacing
thenostrilsfurtherapart.
Magnetoception
Electroreception
Somefish,suchascatfishandsharks,haveorgansthatdetectweakelectriccurrentsontheorderofmillivolt.[50]
Otherfish,liketheSouthAmericanelectricfishesGymnotiformes,canproduceweakelectriccurrents,whichthey
[Link],[Link]
[Link]
alllivingthingsproduce.[51]Thishelpssharks(particularlythehammerheadshark)[Link]
[Link]
[Link]
fororientationandpossiblynavigation.[52]
TheampullaeofLorenziniallowsharkstosenseelectricaldischarges.
Electricfishareabletoproduceelectricfieldsbymodifiedmusclesintheirbody.
Pain
[Link]
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ExperimentsdonebyWilliamTavolgaprovideevidencethat
[Link],inTavolgas
experiments,toadfishgruntedwhenelectricallyshockedand
overtimetheycametogruntatthemeresightofan
electrode.[53]
In2003,ScottishscientistsattheUniversityofEdinburghand
theRoslinInstituteconcludedthatrainbowtroutexhibit
[Link]
venomandaceticacidinjectedintothelipsresultedinfish
rockingtheirbodiesandrubbingtheirlipsalongthesidesand
floorsoftheirtanks,whichtheresearchersconcludedwere
attemptstorelievepain,similartowhatmammalswould
do.[54][55][56]Neuronsfiredinapatternresemblinghuman
neuronalpatterns.[56]
Electromagneticfieldreceptors(ampullaeof
Lorenzini)andmotiondetectingcanalsinthehead
ofashark
[Link]
proofthatfishpossess"consciousawareness,particularlyakindofawarenessthatismeaningfullylikeours".[57]
Rosearguesthatsincefishbrainsaresodifferentfromhumanbrains,fishareprobablynotconsciousinthe
mannerhumansare,[Link]
publishedastudyayearearlierarguingthatfishcannotfeelpainbecausetheirbrainslackaneocortex.[58]
However,animalbehavioristTempleGrandinarguesthatfishcouldstillhaveconsciousnesswithoutaneocortex
because"differentspeciescanusedifferentbrainstructuresandsystemstohandlethesamefunctions."[56]
[Link],
suchasGermanyhavebannedspecifictypesoffishing,andtheBritishRSPCAnowformallyprosecutes
individualswhoarecrueltofish.[59]
Reproductiveprocesses
Seealso:Fishreproduction,Spawning,Ichthyoplankton
Oogoniadevelopmentinteleostsfishvariesaccordingtothegroup,andthe
determinationofoogenesisdynamicsallowstheunderstandingof
[Link],ooplasm,
andthesurroundinglayerscharacterizetheoocytematurationprocess.[60]
Postovulatoryfolliclesarestructuresformedafteroocytereleasetheydo
nothaveendocrinefunction,presentawideirregularlumen,andarerapidly
reabsorbedinaprocessinvolvingtheapoptosisoffollicularcells.A
degenerativeprocesscalledfollicularatresiareabsorbsvitellogenicoocytes
[Link],butlessfrequently,inoocytesin
otherdevelopmentstages.[60]
Somefisharehermaphrodites,havingbothtestesandovarieseitherat
differentphasesintheirlifecycleor,asinhamlets,havethem
simultaneously.
[Link]
Typicaleggofabonyfish,about1
[Link]
releasedintothewatercolumn,where
theydriftaszooplankton
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Over97%ofallknownfishareoviparous,[61]thatis,theeggsdevelopoutsidethemother'[Link]
oviparousfishincludesalmon,goldfish,cichlids,tuna,[Link],fertilisationtakes
placeoutsidethemother'sbody,withthemaleandfemalefishsheddingtheirgametesintothesurroundingwater.
However,afewoviparousfishpracticeinternalfertilisation,withthemaleusingsomesortofintromittentorganto
deliverspermintothegenitalopeningofthefemale,mostnotablytheoviparoussharks,suchasthehornshark,
andoviparousrays,[Link],themaleisequippedwithapairofmodifiedpelvicfinsknown
asclaspers.
[Link]
anaveragediameterof1millimetre(0.039in).Theeggsaregenerallysurroundedbytheextraembryonic
membranesbutdonotdevelopashell,hardorsoft,[Link],leatherycoats,
[Link]
fragile.
Eggoflamprey
Eggofcatshark
(mermaids'purse)
Eggofbullheadshark
Eggofchimaera
[Link],carryalargeyolk
sac(fornourishment)[Link]
oviparousfishisrelativelyshort(usuallyonlyseveralweeks),andlarvaerapidlygrowandchangeappearanceand
structure(aprocesstermedmetamorphosis)[Link]
theiryolksactofeedingonzooplanktonprey,aprocesswhichdependsontypicallyinadequatezooplankton
density,starvingmanylarvae.
Inovoviviparousfishtheeggsdevelopinsidethemother'sbodyafterinternalfertilisationbutreceivelittleorno
nourishmentdirectlyfromthemother,[Link].
Familiarexamplesofovoviviparousfishincludeguppies,angelsharks,andcoelacanths.
[Link].
Typically,viviparousfishhaveastructureanalogoustotheplacentaseeninmammalsconnectingthemother's
[Link],splitfins,andlemon
[Link],inwhichthedevelopingembryoseatothereggsproducedbythe
[Link],suchastheshortfinmakoandporbeagle,butisknownfor
afewbonyfishaswell,suchasthehalfbeakNomorhamphusebrardtii.[62]Intrauterinecannibalismisaneven
moreunusualmodeofvivipary,[Link]
mostcommonlyfoundamongsharks,suchasthegreynurseshark,buthasalsobeenreportedforNomorhamphus
ebrardtii.[62]
Inmanyspeciesoffish,finshavebeenmodifiedtoallowInternalfertilisation.
Aquaristscommonlyrefertoovoviviparousandviviparousfishaslivebearers.
[Link]
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[Link]
[Link],withonetypebeing
predominantwhilethefishbelongstothecorrespondinggender.
Socialbehaviour
[Link]
whichisassociatedwithhumanlove.In2012,researchersinjectedcichlidsfromthesocialspecies
Neolamprologuspulcher,[Link]
increased"responsivenesstosocialinformation",whichsuggests"itisakeyregulatorofsocialbehaviorthathas
evolvedandenduredsinceancienttimes".[63][64]
Cognition
Seealso
Anatomicaltermsoflocation
DigitalFishLibrary
Evolutionoffish
Fishanatomy
Fishdevelopment
Fishmeasurement
Fishmeasurement
Ichthyologyterms
Panderichthysdigits
References
[Link],IvanR(2002)."Morethanjustcoolshades".BritishJournalofOphthalmology86:1075.
doi:10.1136/bjo.86.10.1075.
[Link],[Link](1991).ComparativeAnimalPhysiology,EnvironmentalandMetabolicAnimalPhysiology(4thed.).
Hoboken,NJ:[Link].112.ISBN047185767X.
[Link](1984)FishPhysiology:Gills:PartAAnatomy,gastransferandacidbaseregulation(htt
p://[Link]/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.
[Link](1984)FishPhysiology:Gills:PartBIonandwatertransfer([Link]
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
[Link].2307/1447796.Retrieved25June2009.
[Link],Thomas(1996).[Link].p.542.ISBN9783110106619.
[Link],ChrisAdrianExellNevilleCarrington(2003).[Link].
[Link],AlfredSherwoodParsons,ThomasS.(1977).[Link],PA:HoltSaunders
[Link].316327.ISBN003910284X.
[Link],1985([Link]
[Link],IwamotoandHaouzi,2006([Link]
11."TheOriginoftheLarvaandMetamorphosisinAmphibia".TheAmericanNaturalist(EssexInstitute)91:287.1957.
doi:10.1086/281990.JSTOR2458911.
[Link],J.A.(2002):Gainingground:[Link],Bloomington,
Indiana.369pp
[Link],Lance(1999).[Link]:McGrawHillCompanies,Inc.ISBN007237716X.
[Link](1996)."DeepBreathing".Retrieved20070828.
15."SHARKS&RAYS,SeaWorld/BuschGardensANIMALS,CIRCULATORYSYSTEM".BuschEntertainment
Corporation.Retrieved20090903.
[Link]
14/17
7/3/2016
FishphysiologyWikipedia,thefreeencyclopedia
[Link].(1998):[Link]
systematics,middleearevolution,[Link],Zoologie,Paris,13eSrie19:
pp142.
[Link],JohnF.(1999).CirculatorySystem.MicrosoftEncarta99.
[Link],JeanPraxisPublishingSadasivamKaushikPierreBergotRobertMetailler(2001).NutritionandFeeding
[Link].p.31.ISBN1852332417.ISBN9781852332419.Retrieved20090109.
[Link]."NCIDictionaryofCancerTermslargeintestine".Retrieved20120916.
[Link],[Link]."NoGuts,NoGlory".ReefQuestCentreforSharkResearch.Retrieved20090822.
[Link],Latimeriachalumnae([Link]
[Link],LeonardDando,MarcFowler,Sarah(2005).[Link].ISBN000
7136102.OCLC183136093.
[Link](2005)."Managementtechniquesforelasmobranchfisheries:[Link]".FAO:Fisheriesand
AquacultureDepartment.Retrieved20080316.
[Link]."MAKOSHARKIsurusoxyrinchus".DelawareSeaGrant,[Link]
20080316.
[Link],PatStone,GrahamJohnston,Ian(2009).[Link].198200.
ISBN9781405107242.
[Link],[Link](April1992)."FireintheBellyoftheBeast".[Link]
20090821.
[Link],[Link],[Link],[Link],J.B.(1July2008)."Elevatedredmyotomalmuscletemperatures
inthemostbasaltunaspecies,Allothunnusfallai"(PDF).JournalofFishBiology73(1):[Link].1111/j.1095
8649.2008.01931.x.Retrieved2November2012.
28."TunaBiologyOfTuna".RetrievedSeptember12,2009.
[Link],[Link],[Link],[Link],B.A.(16October2003)."Temperaturedependenceof
theCa2+ATPase(SERCA2)intheventriclesoftunaandmackerel".AJP:Regulatory,IntegrativeandComparative
Physiology286(2):[Link].1152/ajpregu.00392.2003.
[Link],BAFinnerty,JR(1993)."Endothermyinfishes:aphylogeneticanalysisofconstraints,predispositions,and
selectionpressures"(PDF).EnvironmentalBiologyofFishes40(3):[Link].1007/BF00002518.
[Link](10152012)SouthFloridaSunSentinel([Link]
yeballmystery20121015,0,[Link]),
Swordfishheattheireyes([Link]
l)
32.Kapoor2004
[Link],[Link],E.(2005)."MuscleFiberAngle,SegmentBulgingandArchitecturalGearRatioinSegmented
Musculature".JournalofExperimentalBiology208(17):[Link].1242/jeb.01770.
[Link],[Link],[Link],T.J.(2008)."VariableGearinginPennateMuscles".PNAS105(5):17451750.
doi:10.1073/pnas.0709212105.PMC2234215.PMID18230734.
[Link],[Link],E.(2007)."ArchitecturalGearRatioandMuscleFiberStrainHomogeneityinSegmented
Musculature".JournalofExperimentalZoology307(A):[Link].1002/jez.a.358.
[Link],K.(2008).Vertebrates:Comparativeanatomy,function,evolution(5thed.).Boston:McGrawHill.ISBN007
3040584.
[Link],K.(2008).Vertebrates:Comparativeanatomy,function,evolution,(5thed.).Boston:McGrawHill.
[Link],[Link]."TheImportanceofBeingCartilaginous".[Link]
20090829.
39."Dosharkssleep".[Link].Retrieved20100923.
[Link],James(1999).Fish.MicrosoftEncarta99.ISBN0811423468.
[Link],[Link](1993)."Innerearandlateralline".ThePhysiologyofFishes(CRCPress)(1sted.).
[Link]."Appropriatemazemethodologytostudylearninginfish"(PDF).Retrieved
28May2009.
[Link](2005).Biology,[Link],SanFrancisco,California.
[Link].(January2005)."UnderwaterHearingandSound
LocalizationwithandwithoutanAirInterface".Otology&Neurotology26(1):[Link].1097/00129492
20050100000023.
[Link],Michael(1941)."SenseofHearinginFishes".Nature147(3738):[Link].1038/147779b0.
46.B,WILLIAMSC."SenseofHearinginFishes."Nature147.3731(n.d.):[Link].
[Link],[Link]."HearingandVibrationDetection".Retrieved20080601.
[Link],[Link]."SmellandTaste".ReefQuestCentreforSharkResearch.Retrieved20090821.
[Link]
15/17
7/3/2016
FishphysiologyWikipedia,thefreeencyclopedia
[Link]([Link]
a/Gardiner%20and%20Atema%[Link]),[Link],JelleAtema,CurrentBiology13July2010(Vol.20,
Issue13,pp.11871191)
[Link],J.S.,[Link].431472inThePhysiologyofFishes,
[Link](eds.).CRCPress.
[Link](1982)."Electricandmagneticfielddetectioninelasmobranchfishes".Science218(4575):9168.
doi:10.1126/science.7134985.PMID7134985.
[Link],HollandKN,PapastamatiouYP(2005)."Sharkscandetectchangesinthegeomagneticfield".Journalofthe
RoyalSociety,Interface2(2):[Link].1098/rsif.2004.0021.PMC1578252.PMID16849172.
[Link],Joan,"Fish:SensitivityBeyondtheCaptor'sGrasp,"TheAnimals'Agenda,July/August1991,pp.1218
[Link],"FishFeelPain,BritishResearchersSay,"AgenceFrancePresse,1May2003([Link]
m/editorials/[Link])
[Link],Alex(30April2003)."Fishdofeelpain,scientistssay".BBCNews.Retrieved4January2010.
[Link],TempleJohnson,Catherine(2005).[Link],NewYork:[Link].183184.
ISBN0743247698.
57."Rose,[Link]:"Dofishhavenociceptors:Evidencefortheevolutionofavertebratesensory
system" "(PDF).Retrieved21May2011.
[Link],DoFishFeelPain?([Link]
[Link],J."AnglerstoFaceRSPCACheck,"TheSundayTimesBritain,14March2004([Link]
newspaper/0,,1761037515,[Link])
[Link],RodrigoJ.,[Link],[Link],GilmarB.(JulySeptember2005)."Gonadal
structureandgametogenesisofLoricarialentiginosaIsbrcker(Pisces,Teleostei,Siluriformes)".[Link].22
(3):[Link].1590/S010181752005000300005.ISSN01018175.
[Link]:LivebearingFishes,p.13.TetraPress1997.ISBN1564651932
[Link],A&Burns,J:ViviparityintheHalfbeakGeneraDermogenysandNomorhamphus(Teleostei:Hemiramphidae)"
JournalofMorphology234,pp.295317,1997
[Link],ARO'Connor,CMMarshRollo,SEBalcshine,S(2012)."Effectsofisotocinonsocialresponsesina
cooperativelybreedingfish".AnimalBehaviour84(4):[Link].1016/[Link].2012.07.021.
[Link]([Link]
drives_their_social_behavior_says_study95041)Science2.0,10October2012.
Furtherreading
BernierNJ,VanDerKraakG,FarrellAPandBraunerCJ(2009)FishPhysiology:FishNeuroendocrinology
([Link]
hl=en&sa=X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CE8Q6AEwBg)[Link]
9780080877983.
EddyFBandHandyRD(2012)EcologicalandEnvironmentalPhysiologyofFishes([Link]
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([Link]
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([Link]
[Link]/books?id=eddWw_nyRlYC&printsec=frontcover&dq=%22Fish+physiology%22&hl=en&sa=
X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CEsQ6AEwBQ)AcademicPress.ISBN9780080961361.
HaraTJandZielinskiB(2006)FishPhysiology:SensorySystemsNeuroscience([Link]
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"([Link]
ooC&printsec=frontcover&dq=%22Ichthyology+handbook%22&hl=en&sa=X&ei=mB_zTo7JBouWmQXY
[Link]
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v6iXAg&ved=0CC8Q6AEwAA#v=onepage&q=%22Ichthyology%20handbook%22&f=false)Pages137
140,Springer.ISBN9783540428541.
McKenzieDJ,FarrellAPandBraunerCJ(2007)FishPhysiology:PrimitiveFishes([Link]
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([Link]
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([Link]
physiology%22&hl=en&sa=X&ei=454PUeGDLsGimQXCpYCIBA&ved=0CFcQ6AEwCA#v=onepage&q
=%22Fish%20physiology%22&f=false)AcademicPress.ISBN9780123786340.
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