AnimalSentience2016.X:CommentaryonKeyonFishPain Paininparallel CommentaryonKeyonFishPain PeterGodfrey-Smith Philosophy CUNYGraduateCenter&UniversityofSydney Abstract: Brian Key's arguments against the view that fish feel pain can be shown to be fallacious by considering some damage-related behaviors in invertebrates. Pain may have different neural bases in different organisms, so the absence in fish of the cortical structures that might underlie pain in mammalsdoesnotsettlethequestionoffishpain. Peter Godfrey-Smith [email protected] is Distinguished Professor of Philosophy, Graduate Center,CUNY,andProfessorofHistoryandPhilosophy of Science, University of Sydney. His main research interests are philosophy of biology and philosophy of mind. His books include Complexity and the Function of Mind in Nature (Cambridge, 1996), Darwinian Populations and Natural Selection (Oxford, 2009) and Philosophy of Biology(Princeton,2014)http://petergodfreysmith.com/biographical BrianKeyholdsthatonceweworkouttheneuralbasisforpaininmammals,some questionsaboutpaininotheranimalscanbefairlyreadilyanswered: “If“fishpain”existeditwouldneedtobeconsciouslyprocessedusingat leasttheminimalneuralarchitecturedescribedabove.Thus,intheabsence ofthisarchitecture,fishcannotfeelanysortofpain”(Key2016). Iwillaccept,forthesakeofargument,alltheclaimsKeymakesaboutmammals.To assesshowtheseclaimsbearonfish,I'llputonthetablesomeotheranimalsfrom furtherafield.Ina2012studyofhermitcrabs,RobertElwoodfoundthattheycouldbe inducedtoleavetheirshellsbyelectricshock,butweresurprisinglyparticularinhow theydidthis,tradingoffthequalityoftheirpresentshellwiththeintensityofshock, alongwithwhetherolfactorycuessuggestedthatpredatorswereabout:"[I]tisclear thathermitcrabstrade-offcompetingdemandsintheirresponsestoelectricshockina waythatcannotbeexplainedbyanociceptivereflexresponse."Healsoreports "prolongedabdominalgroomingatthesiteofashockinhermitcrabsthatevacuate theirshells"–groomingofakindnotseenwhencrabsareremovedfromtheirshellby otherprocesses.ArecentstudybyJeanAlupayandherco-authors(2014)showedthata crushofanarmintheoctopusAbdopusaculeatusledtoafamilyofwound-directed AnimalSentience2016.X:CommentaryonKeyonFishPain responses,someofwhicharefamiliar–groomingandprotectingthedamagedsite, sensitization,long-termdecreasedthresholdsforescaperesponses–andsomemore unusual:amputationofthearm. Didtheseexperiencesfeellikeanythingfortheoctopusesandcrabs?It'svery hardtoknow.ElwoodandAlupayetal.arecautious.Theynotethatinstudiesofother animals,long-termtendingandgroomingbehaviorsareoftentakentobesomewhat indicativeofpain.Setthatissueasideforamoment,though,assomeconclusionscan bedrawnfromthisworkwithoutconsideringquestionsaboutthefeelofpain. Thenervoussystemsofoctopuses,crabs,andmammalsallhavedifferent architectures.Themostrecentcommonancestorofallthreeofthese(the protostome/deuterostomecommonancestor)probablylivedover600millionyearsago (Petersonetal.2008).Thisanimalisunknown,butprobablyhadaworm-likebody-plan andasimplenervoussystem.Complexnervoussystemshaveseenparallelevolutionin severallinessincethen.Whatevertheneuralarchitectureunderlyingwound-tending andflexibleprotectionofinjuredareasinmammalsmightbe,itisalmostcertainlynot homologoustothearchitecturesubservingthatfunctioninoctopuses.(Alupay'spaper doesnotofferaviewonwhattheneuralbasismightbe,andnotesthataplausible candidate,theverticallobe,doesnotseemtobetherelevantplace).Neurons themselvesarepresentinalltheseanimals,andalmostcertainlyexistedintheir commonancestor,buttheorganizationoftheseneuronsintocomplexnervoussystems occurredindependently.Theseanimalsshareflexiblewound-tendingbehaviors,butdo notsharetheneuralorganizationthatmakesthosebehaviorspossible.(Thecommon ancestorprobablyhadnothingresemblinglimbs,either.)Soevenbeforeweconsider subjectiveexperience,wehavelearnedthatflexible,non-reflexive,andadaptive responsestodamageandothernoxiousstimulidonotrequirethe"neuralarchitecture" (inKey'swords)foundinmammals.Theydonotrequireabrainwithamammalian architectureatall. Let'snowturntothemoredifficultquestionsaboutsubjectiveexperience.How doesthisrelatetoKey'sargumentthatfishdonotfeelpain?LikeElwoodandAlupayet al.,Idon'tprofesstoknowwhethercrabsandoctopusesfeelpain.Asacandidate hypothesis,though,supposethatthecapacitytofeelpainispresentwhenananimal hasaneuralorganizationthatmediatesadaptiveresponsestodamage-relatedstimuliin aflexibleandnon-reflexiveway,includinglong-termmodificationofbehaviorinthe lightofthestimulus,perhapsalsowithsomeoftheintegrationofinformation exemplifiedbythetrade-offsinElwood'scrabs.Thattheoryisaone-sentenceplaceholder(seealsoKlein2015),butitwilldoformypurposeshere.Weknowthatthissort ofcapacitycanbeachievedbymeansofdifferentneuralarchitectures.Itis"multiply realizable,"asphilosopherssay.Ifso,thenfindingthatthearchitecturethatsubserves thesefunctionsandhenceunderliespaininanimalXisnotpresentinanimalYdoesnot tellyouthatanimalYdoesnotfeelpain.Itmighthavedifferentmechanismssubserving thatfunction,andpainmightresultbothways.PerhapsanimalYdoesnotfeelpain, becauseofsubtledifferences.Thatquestionwillbehardtoanswer.Butitisnot answeredbysimplynotingthattheneuralbasisforpaininanimalXisnotfoundin animalY.Key,however,wantstomakeaninferenceofthatsimpleforminthecaseof AnimalSentience2016.X:CommentaryonKeyonFishPain fish.Eveniftherightstoryaboutpainisquitedifferentfromtheplace-holderIused above,thesamequestionswillariseandtheinvertebrateworkshowsthattheneural basesforquitecomplicatedresponsestobodilydamagecanexistwithindifferent architectures. SomeonelikeKeymightobjecttoallthisasfollows:"Fisharevertebrates,as mammalsare.Evenifsomearthropodsandmolluscshaveadifferentneuralbasisfor pain,fishlackthestructuresthatarerelevantinvertebrates.Ifyouareavertebrate, theninordertofeelpainyouneedtohavethesamesortofthingthatmammalshave. Clearlyyouwon'thavetheneuralbasisforpainseeninarthropods."Butthose considerationsshownothingaboutwhethertheremightbeasomewhatdifferent neuralarchitectureunderlyingpaininfish.Thecategory"vertebrate"isnotsacred. Mightfishhavepainviadifferentmechanisms?ThatisaninterestingquestionandI don'tknowtheanswer.ButthisisthesortofquestionthatKeyrefusestoevenaddress; hesimplydismissesit. IemphasizeagainthatKey'sclaimsaboutthecorticalbasisofmammalianpain wereacceptedhereonlyforthesakeofargument;othercommentatorsaremore qualifiedthanItoassessthisissue.Lastly,Keydrawsananalogybetweenerringonthe sideofgenerosityinourviewsaboutfishpainandanerrormadeinadifferentcontext. "[A]scientificresearcharticlewaspublishedthatpurportedlylinkedmeaslesmumps-rubella(MMR)vaccinationcausallytoautism.Althoughthislinkwas subsequentlydisproven,manypeoplecontinuedtoacceptat‘facevalue’the causalassociationbetweenMMRvaccinationandthedevelopmentofautismin children....Thiscausedparentsnottohavetheirchildrenvaccinatedand subsequentlyledtoapublichealthcrisis.Thus,whileinitiallyacceptingtheidea thatMMRvaccinationcausesautismmaybeconsideredasafewaytoproceed (evenifitisnottrue),itcancausecatastrophiceffects"(Key2016). Theirrelevanceofthiscomparisonistotal.OnecouldaswellsaythatKey'spositionis akintotheviewsofthosewhoweretardyinrecognizingthedangersofthalidomide(a drugthatcausesbirthdefectswhentakenbypregnantwomen)asittookawhileforits dangerstobecomecompletelyclear.Keymightreplythatanycomparisonbetweenhis stanceandthatofadilatorydrugcompanyisoutrageous,andsoitis,buthisanalogy withtheMMRvaccinecaseisentirelyonapar. References Alupay,J.,Hadjisolomou,S.,andCrook,R.J.(2014).Arminjuryproduceslong-term behavioralandneuralhypersensitivityinoctopus.NeuroscienceLetters558:137– 142. Elwood,R.(2012).Evidenceforpainindecapodcrustaceans.AnimalWelfare21(S2): 23-27.doi:10.7120/096272812X13353700593365 AnimalSentience2016.X:CommentaryonKeyonFishPain Key,B.(2016).Whyfishdonotfeelpain.AnimalSentience2016.3 Klein,C.(2015).WhattheBodyCommands:TheImperativeTheoryofPain.Cambridge MA:MITPress. Peterson,K.,Cotton,J.,Gehling,J.andPisani,D.(2008).TheEdiacaranEmergenceof Bilaterians:CongruenceBetweentheGeneticandtheGeologicalFossilRecords. PhilosophicalTransactionsoftheRoyalSocietyofLondonB363:1435–1443.
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