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12/5/2014 SignallingtheoryWikipedia,thefreeencyclopedia

http://en.wikipedia.org/wiki/Signalling_theory 1/19

"Stotting"or"pronking"byayoungspringbok(Antidorcasmarsupialis)maysignalhonestlytopredatorssuchascheetahsthatitisafitandfastindividual,andthereforenotworthchasing.

SignallingtheoryFromWikipedia,thefreeencyclopedia

Withinevolutionarybiology,signallingtheoryisabodyoftheoreticalworkexaminingcommunicationbetweenindividuals.Thecentralquestioniswhenorganismswithconflictinginterestsshouldbeexpectedtocommunicatehonestly(nopresumptionbeingmadeofconsciousintention)ratherthancheating.Mathematicalmodelsinwhichorganismssignaltheirconditiontootherindividualsaspartofanevolutionarilystablestrategyareimportantforresearchinthisfield.

Signalsaregivenincontextssuchasmateselectionbyfemales,whichsubjectsthemales'signalstoselectivepressure.Signalsthusevolvebecausetheymodifythebehaviourofthereceivertobenefitthesignaller.Signalsmaybehonest,conveyinginformationwhichusefullyincreasesthefitnessofthereceiver,ordishonest.Anindividualcancheatbygivingadishonestsignal,whichmightbrieflybenefitthatsignaller,attheriskofunderminingthesignallingsystemforthewholepopulation.

Thequestionofwhetherselectionofsignalsworksattheleveloftheindividualorganismorgene,oratthelevelofthegroup,hasbeendebatedbybiologistssuchasRichardDawkins,arguingthatindividualsevolvetosignalandtoreceivesignalsbetter,includingresistingmanipulation.AmotzZahavisuggestedthatcheatingcouldbecontrolledbythehandicapprinciple,wherethebesthorseinahandicapraceistheonecarryingthelargesthandicapweight.AccordingtoZahavi'stheory,signallerssuchasmalepeacockshave'tails'thataregenuinelyhandicaps,beingcostlytoproduce.Thesystemisevolutionarilystableasthelargeshowytailsarehonestsignals.Biologistshaveattemptedtoverifythehandicapprinciple,butwithinconsistentresults.ThemathematicalbiologistRonaldFisheranalysedthecontributionthathavingtwocopiesofeachgene(diploidy)wouldmaketohonestsignalling,demonstratingthatarunawayeffectcouldoccurinsexualselection,dependingsensitivelyonthebalanceofcostsandbenefits.

Thesamemechanismscanbeexpectedinhumans,whereresearchershavestudiedbehavioursincludingrisktakingbyyoungmen,huntingoflargegameanimals,andcostlyreligiousrituals,findingthattheseappeartoqualifyascostlyhonestsignals.

Contents

1Sexualselection2Honestsignals3Dishonestsignals4Thesportshandicappingmetaphor



Eurasianjay,Garrulusglandarius,giveshonestsignalsloudalarmcallsfromitstreeperchwhenitseesapredator.

5CostlysignallingandFisheriandiploiddynamics6Examples7Humanhonestsignals

7.1Costlysignallinginhunting7.2Physicalrisksasacostlysignal7.3Religionasacostlysignal

8Seealso9Notes10References11Sources12Externallinks

Sexualselection

Whenanimalschoosemates,traitssuchassignallingaresubjecttoevolutionarypressure.Forexample,themalegraytreefrog,Hylaversicolor,producesacalltoattractfemales.Onceafemalechoosesamate,thisselectsforaspecificstyleofmalecalling,thuspropagatingaspecificsignallingability.Thesignalcanbethecallitself,theintensityofacall,itsvariationstyle,itsrepetitionrate,andsoon.Varioushypothesesseektoexplainwhyfemaleswouldselectforonecallovertheother.Thesensoryexploitationhypothesisproposesthatpreexistingpreferencesinfemalereceiverscandrivetheevolutionofsignalinnovationinmalesenders,inasimilarwaytothehiddenpreferencehypothesiswhichproposesthatsuccessfulcallsarebetterabletomatchsome'hiddenpreference'inthefemale.[1]

Honestsignals

Inbiology,signalsaretraits,includingstructuresandbehaviours,thathaveevolvedspecificallybecausetheychangethebehaviourofreceiversinwaysthatbenefitthesignaller.[2]Traitsoractionsthatbenefitthereceiverexclusivelyarecalledcues.Whenanalertbirddeliberatelygivesawarningcalltoastalkingpredatorandthepredatorgivesupthehunt,thesoundisasignal.Whenaforagingbirdinadvertentlymakesarustlingsoundintheleavesthatattractspredatorsandincreasestheriskofpredation,thesoundisa'cue'.[2]

Signallingsystemsareshapedbymutualinterestsbetweensignallersandreceivers.AnalertbirdsuchasaEurasianjaywarningoffastalkingpredatoriscommunicatingsomethingusefultothepredator:thatithasbeendetectedbythepreyitmightaswellquitwastingitstimestalkingthisalertedprey,whichitisunlikelytocatch.Whenthepredatorgivesup,thesignallercangetbackto



Malefiddlercrabsignalswithitsenlargedfightingclaw,butweakregrownclawsmaybedishonestsignals.

othertaskssuchasfeeding.Oncethestalkingpredatorisdetected,thesignallingpreyandreceivingpredatorthushaveamutualinterestterminatingthehunt.[3][4]

Withinspecies,mutualinterestsincreasewithkinship.[5]Kinshipiscentraltomodelsofsignallingbetweenrelatives,forinstancewhenbroodsofnestlingbirdsbegandcompeteforfoodfromtheirparents.[6][7]

Theconceptofhonestyinanimalcommunicationiscontroversialbecauseitisdifficulttodetermineintentandusethatasacriteriontodiscriminatedeceptionfromhonesty,aswedoinhumaninteractions.[8]Biologistsusethephrase"honestsignals"inastatisticalsense.Biologicalsignals,likewarningcallsorresplendenttailfeathers,arehonestiftheyarecorrelatedwithsomethingusefultothereceiver,thatis,thesignaltrait[a]iscorrelatedwiththeunobservablethingofvaluetothereceiver[b].Honestbiologicalsignalsdonotneedtobeperfectlyinformative,reducinguncertaintytozerotheyonlyneedtobecorrect"onaverage"tobeuseful.[9]Ultimatelythevalueofthesignalledinformationdependsontheextenttowhichitallowsthereceivertoincreaseitsfitness.[10]Hence,"honest"signalsareevolutionarilystable.

Dishonestsignals

Becausetherearebothmutualandconflictinginterestsinmostanimalsignallingsystems,acentralprobleminsignallingtheoryisdishonestyorcheating.Forexample,ifforagingbirdsaresaferwhentheygiveawarningcall,cheatscouldgivefalsealarmsatrandom,justincaseapredatorisnearby.Buttoomuchcheatingcouldcausethesignallingsystemtocollapse.Everydishonestsignalweakenstheintegrityofthesignallingsystem,andsoreducesthefitnessofthegroup.[11][12]AnexampleofdishonestsignallingcomesfromFiddlercrabssuchasUcalacteamjoebergi,whichhavebeenshowntobluff(noconsciousintentionbeingimplied)abouttheirfightingability.Whenaclawislost,acraboccasionallyregrowsaweakerclawthatneverthelessintimidatescrabswithsmallerbutstrongerclaws.[13][14]Theproportionofdishonestsignalsislowenoughforitnottobeworthwhileforcrabstotestthehonestyofeverysignalthroughcombat.[11]

RichardDawkinsandJohnKrebsin1978consideredwhetherindividualsofthesamespecieswouldactasifattemptingtodeceiveeachother.Theyapplieda"selfishgene"viewofevolutiontoanimals'threatdisplaystoseeifitwouldbeintheirgenes'intereststogivedishonestsignals.Theycriticisedpreviousethologists,suchasNikolaasTinbergenandDesmondMorrisforsuggestingthatsuchdisplayswere"forthegoodofthespecies".Theyarguedthatsuchcommunicationoughttobeviewedasanevolutionaryarmsraceinwhichsignallersevolvetobecomebetteratmanipulatingreceivers,whilereceiversevolvetobecomemoreresistanttomanipulation.[11][12]Thegametheoreticalmodelofthewarofattritionsimilarlysuggeststhatthreatdisplaysoughtnottoconveyanyreliableinformationaboutintentions.[15]

Thesportshandicappingmetaphor



Thebesthorsesinahandicapracecarrythelargestweights,sothesizeofthehandicapisameasureoftheanimal'squality.

Peacocksignalsreproductivefitnesswithitslargecolourfultail,possiblybecauseitisahandicap.

In1975,AmotzZahaviproposedaverbalmodelforhowsignalcostscouldconstraincheatingandstabilizean"honest"correlationbetweenobservedsignalsandunobservablequalities,basedonananalogytosportshandicappingsystems.[16][17]Hecalledthisideathehandicapprinciple.Thepurposeofasportshandicappingsystemistoreducedisparitiesinperformance,makingthecontestmorecompetitive.Inahorsehandicaprace,intrinsicallyfasterhorsesaregivenheavierweightstocarryundertheirsaddles.Similarly,inamateurgolf,bettergolfershavefewerstrokessubtractedfromtheirrawscores.Thiscreatescorrelationsbetweenthehandicapandunhandicappedperformance,andifthehandicapsworkastheyaresupposedto,betweenthehandicapandhandicappedperformance.Ifyouknewnothingabouttworacehorsesortwoamateurgolfersexcepttheirhandicaps,youcouldinferwhichismostlikelytowin:thehorsewiththebiggerweighthandicap,andthegolferwiththesmallerstrokehandicap.Byanalogy,ifpeacock'tails'(largetailcovertfeathers)actasahandicappingsystem,andapeahenknewnothingabouttwopeacocksbutthesizesoftheirtails,shecould"infer"thatthepeacockwiththebiggertailhasgreaterunobservableintrinsicquality.Displaycostscanincludeextrinsicsocialcosts,intheformoftestingandpunishmentbyrivals,aswellasintrinsicproductioncosts.[18]AnotherexamplegivenintextbooksistheextinctIrishelk,Megalocerosgiganteus.ThemaleIrishelk'senormousantlerscouldperhapshaveevolvedasdisplaysofabilitytoovercomehandicap,thoughbiologistspointoutthatifthehandicapisinherited,itsgenesoughttobeselectedagainst.[19]

Theessentialideahereisintuitiveandprobablyqualifiesasfolkwisdom.ItwasarticulatedbyKurtVonnegutinhis1961shortstoryHarrisonBergeron.[20]InVonnegutsfuturisticdystopia,theHandicapperGeneralusesavarietyofhandicappingmechanismstoreduceinequalitiesinperformance.Aspectatorataballetcomments:"itwaseasytoseethatshewasthestrongestandmostgracefulofalldancers,forherhandicapbagswereasbigasthosewornbytwohundredpoundmen."Zahaviinterpretedthisanalogytomeanthathigherqualitypeacockswithbiggertailsaresignallingtheirabilityto"waste"moreofsomeresourcebytradingitoffforabiggertail.ThisresonateswithThorsteinVeblen'sideathatconspicuousconsumptionandextravagantstatussymbolscansignalwealth.[21]

Zahavisconclusionsrestonhisverbalinterpretationofametaphor,andinitiallythehandicapprinciplewasnotwellreceivedbyevolutionarybiologists.[17]However,in1984,NurandHasson[22]usedlifehistorytheorytoshowhowdifferencesinsignallingcosts,intheformofsurvivalreproductiontradeoffs,couldstabilizeasignallingsystemroughlyasZahaviimagined.Geneticmodelsalsosuggestedthiswaspossible.[23]In1990AlanGrafenshowedthatahandicaplikesignallingsystemwasevolutionarilystableifhigherqualitysignallerspaidlowermarginalsurvivalcostsfortheirsignals.[24]



TheenormousantlersoftheextinctIrishelk,Megalocerosgiganteusmayhaveevolvedasdisplaysofabilitytoovercomehandicap.

In1982,W.D.Hamiltonproposedaspecificbutwidelyapplicablehandicapmechanism,parasitemediatedsexualselection.[25]Hearguedthatintheneverendingcoevolutionaryracebetweenhostsandtheirparasites,sexuallyselectedsignalsindicatehealth.Thisideawastestedin1994inbarnswallows,aspecieswheremaleshavelongtailstreamers.Mllerfoundthatthemaleswithlongertails,andtheiroffspring,didhavefewerbloodsuckingmites,whereasfosteredyoungdidnot.Theeffectwasthereforegenetic,confirmingHamilton'stheory.[26]

Anotherexampleistheideathatcarotenoidshavedualbutmutuallyincompatiblerolesinimmunefunctionandsignalling.[27]Giventhatanimalscannotsynthesizecarotenoidsdenovo,thesemustbeobtainedfromfood.Thehypothesisstatesthatanimalswithcarotenoiddependedsexualsignalsaredemonstratingtheirabilityto"waste"carotenoidsonsexualsignalsattheexpenseoftheirimmunesystem.[28][29]

Thehandicapprinciplehasprovenhardtotestempirically,partlybecauseofinconsistentinterpretationsofZahavismetaphorandGrafensmarginalfitnessmodel,andpartlybecauseofconflictingempiricalresults:insomestudiesindividualswithbiggersignalsseemtopayhighercosts,inotherstudiestheyseemtobepayinglowercosts.[30][31]ApossibleexplanationfortheinconsistentempiricalresultsisgiveninaseriesofpapersbyGetty,[32][33][34][35]whoshowsthatGrafensproofofthehandicapprincipleisbasedonthecriticalsimplifyingassumptionthatsignallerstradeoffcostsforbenefitsinanadditivefashion,thewayhumansinvestmoneytoincreaseincomeinthesamecurrency.[c]Buttheassumptionthatcostsandbenefitstradeoffinanadditivefashionisnotvalidforthesurvivalcostreproductionbenefittradeoffthatisassumedtomediatetheevolutionofsexuallyselectedsignals.Fitnessdependsonproducingoffspring,whichisamultiplicativefunctionofreproductivesuccessgivenanindividualisstillalivetimestheprobabilityofstillbeingalive,giveninvestmentinsignals.[22]

CostlysignallingandFisheriandiploiddynamics

Theefforttodiscoverhowcostscanconstrainan"honest"correlationbetweenobservablesignalsandunobservablequalitieswithinsignallersisbuiltonstrategicmodelsofsignallinggames,withmanysimplifyingassumptions.Thesemodelsaremostoftenappliedtosexuallyselectedsignallingindiploidanimals,buttheyrarelyincorporateafactaboutdiploidsexualreproductionnotedbythemathematicalbiologistRonaldFisherintheearly20thcentury:ifthereare"preferencegenes"correlatedwithchoosinessinfemalesaswellas"signalgenes"correlatedwithdisplaytraitsinmales,choosierfemalesshouldtendtomatewithshowiermales.Overgenerations,showiersonsshouldalsocarrygenesassociatedwithchoosierdaughters,andchoosierdaughtersshouldalsocarrygenesassociatedwithshowiersons.ThiscancausetheevolutionarydynamicknownasFisherianrunaway,inwhichmalesbecomeevershowier.RussellLandeexploredthiswithaquantitativegeneticmodel,[23]showingthatFisheriandiploiddynamicsaresensitivetosignallingandsearchcosts.OthermodelsincorporatebothcostlysignallingandFisherianrunaway.[37][38]Thesemodelsshowthatiffitnessdependsonbothsurvivalandreproduction,havingsexysonsandchoosydaughters(inthestereotypicalmodel)canbeadaptive,increasingfitnessjustasmuchashavinghealthysonsanddaughters.[37][38]



Onetheoryisthatautumnalcoloursareasignalfromtreestoaphidsofpowerfulchemicaldefences.

AmalehunterandafemalegathereroftheKali'napeopleofGuyana,drawnbyPierreBarrrein1743.Generoussharingbymalehuntersmayserveasa"costlysignal",helpingthemtoacquiremates.

Examples

SamBrownandW.D.Hamilton[39]andMarcoArchetti[40]proposedthatautumnleafcolourisasignalfromtreestoaphidsandotherpestspeciesthatmigrateinautumntothetrees.Intheirtheory,brightautumncolorationwithpinksandyellowsiscostlytotreesbecausepigmentsrequireenergytosynthesize,buttheinvestmentmayhelpthemtoreducetheirparasiteload.[39][40]

Stotting,forexampleinThomson'sGazelle,iscitedasanexampleofsignalling:thegazellesjumpclosetoapredatorinsteadofescaping,inwhatcouldbeasignalofstrength.[41]

Humanhonestsignals

Humanbehaviourmayalsoprovideexamplesofcostlysignals.Ingeneral,thesesignalsprovideinformationaboutapersonsphenotypicqualityorcooperativetendencies.Evidenceforcostlysignallinghasbeenfoundinmanyareasofhumaninteractionincludingrisktaking,hunting,andreligion,[42]whicharediscussedinturnbelow.

Costlysignallinginhunting

Largegamehuntinghasbeenstudiedextensivelyasasignalofmenswillingnesstotakephysicalrisks,aswellasshowcasestrengthandcoordination.[42][43][44][45]Costlysignallingtheoryisausefultoolforunderstandingfoodsharingamonghuntergatherersbecauseitcanbeappliedtosituationsinwhichdelayedreciprocityisnotaviableexplanation.[46][47][48]Instancesthatareparticularlyinconsistentwiththedelayedreciprocityhypothesisarethoseinwhichahuntershareshiskillindiscriminatelywithallmembersofalargegroup.[49]Inthesesituations,theindividualssharingmeathavenocontroloverwhetherornottheirgenerositywillbereciprocated,andfreeridingbecomesanattractivestrategyforthosereceivingmeat.Freeridersarepeoplewhoreapthebenefitsofgrouplivingwithoutcontributingtoitsmaintenance.[50]Fortunately,costlysignallingtheorycanfillsomeofthegapsleftbythedelayedreciprocityhypothesis.[51][52]Hawkeshassuggestedthatmentargetlargegameandpubliclysharemeattodrawsocialattentionorto

showoff.[53]Suchdisplayandtheresultingfavorableattentioncanimproveahuntersreputationbyprovidinginformationabouthisphenotypicquality.Highqualitysignallersaremoresuccessfulinacquiringmatesandallies.Thus,costlysignallingtheorycanexplainapparentlywastefulandaltruisticbehaviour.[24][52][54][55][56]



Inordertobeeffective,costlysignalsmustfulfillspecificcriteria.[16][42][57]Firstly,signallersmustincurdifferentlevelsofcostandbenefitforsignallingbehaviour.Secondly,costsandbenefitsmustreflectthesignallersphenotypicquality.Thirdly,theinformationprovidedbyasignalshouldbedirectedatandaccessibletoanaudience.Areceivercanbeanyonewhostandstobenefitfrominformationthesignallerissending,suchaspotentialmates,allies,orcompetitors.Honestyisguaranteedwhenonlyindividualsofhighqualitycanpaythe(high)costsofsignalling.Hence,honestsignallingmakeitimpossibleforlowqualityindividualstofakeasignalandfoolareceiver.[16][42][57]

BliegeBirdetal.observedturtlehuntingandspearfishingpatternsinaMeriamcommunityintheTorresStraitofAustralia,publishingtheirfindingsin2001.Here,onlysomeMeriammenwereabletoaccumulatehighcaloricgainsfortheamountoftimespentturtlehuntingorspearfishing(reachingathresholdmeasuredinkcal/h).Sinceadailycatchoffishiscarriedhomebyhandandturtlesarefrequentlyservedatlargefeasts,membersofthecommunityknowwhichmenmostreliablybroughtthemturtlemeatandfish.Thus,turtlehuntingqualifiesasacostlysignal.Furthermore,turtlehuntingandspearfishingareactuallylessproductive(inkcal/h)thanforagingforshellfish,wheresuccessdependsonlyontheamountoftimededicatedtosearching,soshellfishforagingisapoorsignalofskillorstrength.Thissuggeststhatenergeticgainsarenottheprimaryreasonmentakepartinturtlehuntingandspearfishing.[42]AfollowupstudyfoundthatsuccessfulMeriamhuntersdoexperiencegreatersocialbenefitsandreproductivesuccessthanlessskilledhunters.[58]

TheHadzapeopleofTanzaniaalsosharefood,possiblytogaininreputation.[59]Hunterscannotbesharingmeatmainlytoprovisiontheirfamiliesortogainreciprocalbenefits,asteenageboysoftengiveawaytheirmeateventhoughtheydonotyethavewivesorchildren,socostlysignallingoftheirqualitiesisthelikelyexplanation.[60]Thesequalitiesincludegoodeyesight,coordination,strength,knowledge,endurance,orbravery.Hadzahuntersmoreoftenpairwithhighlyfertile,hardworkingwivesthannonhunters.[61]Awomanbenefitsfrommatingwithamanwhopossessessuchqualitiesasherchildrenwillmostlikelyinheritqualitiesthatincreasefitnessandsurvivorship.Shemayalsobenefitfromherhusbandshighsocialstatus.Thus,huntingisanhonestandcostlysignalofphenotypicquality.[62]

AmongthemenofIfalukatoll,costlysignallingtheorycanalsoexplainwhymentorchfish.[51]TorchfishingisaritualizedmethodoffishingonIfalukwherebymenusetorchesmadefromdriedcoconutfrondstocatchlargedogtoothedtuna.Preparationfortorchfishingrequiressignificanttimeinvestmentsandinvolvesagreatdealoforganization.Duetothetimeandenergeticcostsofpreparation,torchfishingresultsinnetcaloriclossesforfishers.Therefore,torchfishingisahandicapthatservestosignalmensproductivity.[51]TorchfishingisthemostadvertisedfishingoccupationonIfaluk.Womenandothersusuallyspendtimeobservingthecanoesastheysailbeyondthereef.Also,localritualshelptobroadcastinformationaboutwhichfishersaresuccessfulandenhancefishersreputationsduringthetorchfishingseason.Severalritualbehaviouralanddietaryconstraintsclearlydistinguishtorchfishersfromothermen.First,malesareonlypermittedtotorchfishiftheyparticipatedonthefirstdayofthefishingseason.Thecommunityiswellinformedastowhoparticipatesonthisday,andcaneasilyidentifythetorchfishers.Second,torchfishersreceivealloftheirmealsatthecanoehouseandareprohibitedfromeatingcertainfoods.Peoplefrequentlydiscussthequalitiesoftorchfishermen.OnIfaluk,womenclaimthattheyarelookingforhardworkingmates.[63]WiththedistinctsexualdivisionoflaboronIfaluk,industriousnessisahighlyvaluedcharacteristicinmales.[64]Torchfishingthusprovideswomenwithreliableinformationontheworkethicofprospectivemates,whichmakesitanhonestcostlysignal.[52]



Youngmenmaytakepartinriskysportslikemotorcycleracingtosignaltheirstrengthandskill.

Inmanyhumancases,astrongreputationbuiltthroughcostlysignallingenhancesamanssocialstatusoverthestatusesofmenwhosignallesssuccessfully.[49][65][66]AmongnorthernKalahariforaginggroups,traditionalhuntersusuallycaptureamaximumoftwoorthreeantelopesperyear.[67]Itwassaidofaparticularlysuccessfulhunter:[68]

"Itwassaidofhimthatheneverreturnedfromahuntwithouthavingkilledatleastawildebeest,ifnotsomethinglarger.Hencethepeopleconnectedwithhimateagreatdealofmeatandhis

popularitygrew."[68]

Althoughthishunterwassharingmeat,hewasnotdoingsointheframeworkofreciprocity.[68]Thegeneralmodelofcostlysignallingisnotreciprocalrather,individualswhoshareacquiremorematesandallies.[16][42]CostlysignallingappliestosituationsinKalahariforaginggroupswheregivingoftengoestorecipientswhohavelittletoofferinreturn.Ayounghunterismotivatedtoimpresscommunitymemberswithdaughterssothathecanobtainhisfirstwife.Olderhuntersmaywishtoattractwomeninterestedinanextramaritalrelationship,ortobeacowife.[69][70]InthesenorthernKalaharigroups,thekillingofalargeanimalindicatesamanwhohasmasteredtheartofhuntingandcansupportafamily.[71]Generally,manywomenseekamanwhoisagoodhunter,hasanagreeablecharacter,isgenerous,andhasadvantageoussocialties.[67][70][71]Sincehuntingabilityisaprerequisiteformarriage,menwhoaregoodhuntersenterthemarriagemarketearliest.Costlysignallingtheoryexplainsseeminglywastefulforagingdisplays.[61]

Physicalrisksasacostlysignal

Costlysignallingcanbeappliedtosituationsinvolvingphysicalstrainandriskofphysicalinjuryordeath.[72][73][74]Researchonphysicalrisktakingisimportantbecauseinformationregardingwhypeople,especiallyyoungmen,takepartinhighriskactivitiescanhelpinthedevelopmentofpreventionprograms.[73][74]Recklessdrivingisalethalproblemamongyoungmeninwesternsocieties.[75]Amalewhotakesaphysicalriskissendingthemessagethathehasenoughstrengthandskilltosurviveextremelydangerousactivities.Thissignalisdirectedatpeersandpotentialmates.[16]

Inastudyofrisktaking,sometypesofrisk,suchasphysicalorheroicriskforothers'benefit,areviewedmorefavorablythanothertypesofrisk,suchastakingdrugs.Malesandfemalesvalueddifferentdegreesofheroicriskformatesandsamesexfriends.Malesvaluedheroicrisktakingbymalefriends,butpreferredlessofitinfemalemates.Femalesvaluedheroicrisktakinginmalematesandlessofitinfemalefriends.Femalesmaybeattractedtomalesinclinedtophysicallydefendthemandtheirchildren.Malesmaypreferheroicrisktakingbymalefriendsastheycouldbegoodallies.[74]

Inwesternsocieties,voluntaryblooddonationisacommon,yetlessextreme,formofrisktaking.Costsassociatedwiththesedonationsincludepainandriskofinfection.[76]Ifblooddonationisanopportunitytosendcostlysignals,thendonorswillbeperceivedbyothersasgenerousandphysicallyhealthy.[16][77]Ina



Religiousritualssuchassnakehandlingmaybeexplainableascostlysignals.

survey,bothdonorsandnondonorsexpressedperceptionsofthehealth,generosity,andabilityofblooddonorstooperateinstressfulsituations.[77]

Religionasacostlysignal

Costlyreligiousritualssuchasmalecircumcision,foodandwaterdeprivation,andsnakehandlinglookparadoxicalinevolutionaryterms.Devoutreligiousbeliefswhereinsuchtraditionsarepracticedthereforeappearmaladaptive.[78]Religionmayhavearisentoincreaseandmaintainintragroupcooperation.[79]Cooperationleadstoaltruisticbehaviour,[80]andcostlysignallingcouldexplainthis.[16]Allreligionsmayinvolvecostlyandelaboraterituals,performedpublicly,todemonstrateloyaltytothereligiousgroup.[81]Inthisway,groupmembersincreasetheirallegiancetothegroupbysignallingtheirinvestmentingroupinterests.However,asgroupsizeincreasesamonghumans,thethreatoffreeridersgrows.[50]Costlysignallingtheoryaccountsforthisbyproposingthatthesereligiousritualsarecostlyenoughtodeterfreeriders.[82]

Ironsproposedthatcostlysignallingtheorycouldexplaincostlyreligiousbehaviour.Hearguedthathardtofakereligiousdisplaysenhancedtrustand

solidarityinacommunity,producingemotionalandeconomicbenefits.HeshowedthatdisplaysignalsamongtheYomutTurkmenofnorthernIranhelpedtosecuretradeagreements.These"ostentatious"displayssignalledcommitmenttoIslamtostrangersandgroupmembers.[83]Sosisdemonstratedthatpeopleinreligiouscommunitiesarefourtimesmorelikelytolivelongerthantheirsecularcounterparts,[51][80]andthattheselongerlifespanswerepositivelycorrelatedwiththenumberofcostlyrequirementsdemandedfromreligiouscommunitymembers.[84]However,confoundingvariablesmaynothavebeenexcluded.[85]Iannacconestudiedtheeffectsofcostlysignalsonreligiouscommunities.Inaselfreportedsurvey,asthestrictnessofachurchincreased,theattendanceandcontributionstothatchurchincreasedproportionally.Ineffect,peopleweremorewillingtoparticipateinachurchthathasmorestringentdemandsonitsmembers.[86]

Despitetheexperimentalsupportforthishypothesis,itremainscontroversial.Acommoncritiqueisthatdevoutnessiseasytofake,suchassimplybyattendingareligiousservice.[87]However,thehypothesispredictsthatpeoplearemorelikelytojoinandcontributetoareligiousgroupwhenitsritualsarecostly.[86]Anothercritiquespecificallyasks:whyreligion?Thereisnoevolutionaryadvantagetoevolvingreligionoverothersignalsofcommitmentsuchasnationality,asIronsadmits.[80]However,thereinforcementofreligiousritesaswellastheintrinsicrewardandpunishmentsystemfoundinreligionmakesitanidealcandidateforincreasingintragroupcooperation.Finally,thereisinsufficientevidenceforincreaseinfitnessasaresultofreligiouscooperation.[80]However,Sosisarguesforbenefitsfromreligionitself,suchasincreasedlongevity,improvedhealth,assistanceduringcrises,andgreaterpsychologicalwellbeing.[88]

Seealso

Animalcommunication



AutumnleafcolorBaculumCheaptalkConspicuousconsumptionDramaturgy(sociology)HandicapprincipleMultiplesexualornamentsOriginoflanguageSignalling(economics)Signallinggame

Notes

a. Êconomistscallwhatisavailabletothereceiver"publicinformation".b. Êconomistscalltheunobservablethingthatwouldbeofvaluetothereceiver"privateinformation"biologists

oftencallit"quality"c. ^GrafensproofisformallysimilartoaclassicmonographoneconomicmarketsignallingbyNobellaureate

MichaelSpence.[36]

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