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CognitivePsychology61(2010)1–22 ContentslistsavailableatScienceDirect Cognitive Psychology journal homepage: www.elsevier.com/locate/cogpsych Adaptive memory: Ancestral priorities and the mnemonic value of survival processing James S. Nairnea,*, Josefa N.S. Pandeiradab aPurdueUniversity,UnitedStates bUniversityofAveiro,Portugal a r t i c l e i n f o a b s t r a c t Articlehistory: Evolutionary psychologists often propose that humans carry Accepted26January2010 around‘‘stone-age”brains,alongwithatoolkitofcognitiveadap- Availableonline6March2010 tationsdesignedoriginallytosolvehunter–gathererproblems.This perspective predicts that optimal cognitive performance might Keywords: sometimesbeinducedbyancestrally-basedproblems,thosepres- Memory entinancestralenvironments,ratherthanbyadaptiveproblems Evolution faced more commonly in modern environments. This prediction Survival was examinedin fourexperiments usingthe survival processing paradigm, in which retention is tested after participants process information in terms of its relevance to fitness-based scenarios. Ineachoftheexperiments,participantsrememberedinformation betterafter processingitsrelevanceinanancestralenvironment (the grasslands), compared to a modern urban environment (a city),despitethefactthatallscenariosdescribedsimilarfitness- relevantproblems. Thesedatasuggestthatourmemorysystems maybetunedtoancestralpriorities. !2010ElsevierInc.Allrightsreserved. 1. Introduction Thecapacitytorememberevolved,havingbeenshapedandsculptedbytheprocessesofnatural selection.Specificselectionpressures,relatedtosurvivalandreproduction,conferredselectionadvan- tagestoorganismscapableofusingthepastintheserviceofthepresent.Noscholarseriouslyques- tionstheseclaims,althoughthefunctionalpropertiesofmemoryarerarelyconsideredexplicitlyby modernmemorytheorists(Nairne,2005;Klein,Cosmides,Tooby,&Chance,2002).Instead,themain * Correspondingauthor.Address:DepartmentofPsychologicalSciences,PurdueUniversity,703ThirdStreet,WestLafayette, IN47907-2081,UnitedStates. E-mailaddress:[email protected](J.S.Nairne). 0010-0285/$-seefrontmatter!2010ElsevierInc.Allrightsreserved. doi:10.1016/j.cogpsych.2010.01.005 2 J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 empiricalandtheoreticaleffortoverthepastcenturyhasbeendirectedatunderstandingtheproxi- matemechanismsthatcontrolperformanceonspecifictasks—i.e.,the‘‘how”ratherthanthe‘‘why” ofremembering(seealso,Bruce,1985;Sherry&Schacter,1987). Yet,fromanevolutionaryperspective,itisreasonabletoaskwhethervestigesoftheselectionpres- suresthatledtomemory’sdevelopmentremainapparentinitsoperatingcharacteristics.Ifmemory systemsevolvedtoenhancefitness(survivalandreproduction),thenareorganismsespeciallygoodat encodingandretainingmaterialfromfitness-relevantsituations?Thisisultimatelyanempiricalques- tion, one that need not suffer from the criticisms usually directed at investigations in evolutionary psychology(e.g.,‘‘just-so”and‘‘Panglossian”reasoning;seeGould&Lewontin,1979).Indeed,there isnowsubstantialempiricalevidenceshowingthatmemoryisenhancedwhenpeopleprocessinfor- mationintermsofitsfitnessvalue(Kang,McDermott,&Cohen,2008;Nairne,Thompson,&Pandei- rada, 2007; Weinstein, Bugg, & Roediger, 2008). For example, Nairne, Pandeirada, and Thompson (2008)foundthatafewsecondsofsurvivalprocessing,duringwhichparticipantswereaskedtorate therelevanceofrandomwordstoasurvivalscenario,producedbetterlong-termretentionthanahost oftraditionallystrongencodingprocedures(e.g.,formingavisualimageorengaginginsemanticpro- cessing;seealsoNairne&Pandeirada,2008a). Still, establishing an empirical advantage for fitness-relevant processing tells us little about the cognitivemechanismsthatproducethoseadvantages,orabouttheirultimateorigins.Onecouldargue thatorganismsevolveddomain-generalmechanismsforretaininginformation,onesthatarematerial- andsituation-independent.Infact,memoryresearchersvirtuallyalwaysemploygeneralmechanisms toexplainsuperiormnemonicperformance,mechanismssuchaselaboration(connectinginformation tootherthingsinmemory),distinctiveprocessing(creatingunusualrepresentations),orprocessing thatistransfer-appropriate(encodingrepresentationsthatarelikelytobematchedinatargetedre- trievalenvironment).Certaintypesofmaterial,orsituations,maynaturallyaffordelaborate,distinc- tive, or appropriate encodings, but the mechanisms themselves are assumed to operate similarly acrossdomains(seeSurprenant&Neath,2009). Evolutionarypsychologiststendtoeschewdomain-generalmechanismsforavarietyofcompelling reasons.Natureusually‘‘selects”onephysicaldesignoveranotherwheniteffectivelysolvesaspecific adaptiveproblemrelatedtofitness—thus,wehaveheartsuniquelydesignedtopumpblood,kidneys tofilterimpurities,andsoforth.Fromasocialandcognitiveperspective,therearetoomanycritical problemstobesolved,suchasavoidingpredators,locatingnourishment,orselectinganappropriate mate,torelyonthewhimsofgenerallearningmechanismsthataresimplydesignedtoextractinfor- mationfromindividualexperiences(e.g.,Cosmides&Tooby,2005).Relevantexperiencesmayoccur naturally with too low a probability, and problems of combinatorial explosion arise quickly with a cognitivesystemthatisdesignedtostoreeverything.Selectivestorage,basedoninherentcognitive ‘‘biases”or‘‘tunings,”wouldappeartobenecessaryforspeciestosurviveandreproduceeffectively. Of course, these are not airtight arguments (see Buller, 2005), but they can serve as the impetus andfoundationforempiricalinvestigation. 1.1. Ancestralpriorities Tothe extentthat our brainswere sculptedprimarilyduring thePleistocene, theperiod usually considered as the environment of evolutionary adaptation (e.g., Symons, 1992), natural selection wouldhavebeendrivenbyPleistocenepressures—thatis,theuniquesetofadaptiveproblemsfaced byour hunter–gathererancestors. Evolutionarypsychologists commonlyarguethatwecontinueto carry around these ‘‘stone-age” brains, along with a toolkit of cognitive adaptations that were de- signed originally to solve hunter–gatherer problems (Tooby & Cosmides, 1992). One might predict, therefore,thatoptimalcognitiveperformanceshouldbeinducedbyancestrally-basedproblems,par- ticularlythosepresentinancestralforagingenvironments,ratherthanbyproblemsfacedinmodern environments.If,infact,ourcognitivesystemsevolvedtosolveparticularadaptiveproblemsthenthe fitbetweenproblemandprocessshouldaccount,atleastinpart,forprocessingefficiency.Thisisnota strongpredictionofevolutionarytheory,becauseadaptationscanbeeffectivelyexaptedtoperform rolesthatareunrelatedtotheoriginalselectionenvironment(seeAndrews,Gangestad,&Matthews, 2002),butitrepresentsaninterestingempiricaltest. J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 3 Thereissomesupportforthenotionofancestralprioritiesincognitiveprocessing.Forexample,it iswidelyacceptedthatorganisms,includinghumans,haveaninbornattentionalbiasforthreatening stimuli—particularly stimuli that were relevant in ancestral environments (see Öhman & Mineka, 2001,forareview).Bothadultsandveryyoungchildrendetectevolutionarily-relevantstimuli,such assnakesandspiders,morequicklythannon-threateningstimuli(LoBue&DeLoache,2008).Even7- to9-month-oldinfants, whopresumablyhavelimitedexperiencewithsnakes,showan attentional biastosnakescomparedtootherunfamiliaranimals(DeLoache&LoBue,2009).Inaddition,New,Cos- mides,andTooby(2007)foundfasterandmoreaccuratedetectionforanimals,bothhumanandnon- human,thanforinanimateobjectsusingthechange-detectionparadigm,aprocedureinwhichpeople areaskedtodetectdifferencesbetweentworapidly-alternatingimages.Changedetectionfornonhu- mananimalswasfasterthanforvehicles,eventhoughthelatterarecommonplaceineverydayexpe- rienceandcanbesurvival-related.Avarietyofalternativeexplanationsforthedatawerediscounted, includingtheinterestvalueandlow-levelvisualcharacteristicsofthestimuli,leadingtheauthorsto concludethatpeoplepossessananimatemonitoringsystemthatis‘‘bettertunedtoancestralthanto modernpriorities”(Newetal.,2007,p.16603). Inthelearningdomain,evolutionarily-significantstimulicanbeeasiertoassociatewithaversive stimuli (e.g., shock or loud noise) than comparable modern controls (Öhman & Mineka, 2003). For example, some studies have found that snakes and/or spiders are easier to associate with aversive stimuli,orshowmoreresistancetosubsequentextinction,thanontogeneticfear-relevantstimulisuch asgunsorelectricaloutlets(seeÖhman&Mineka,2001,forareview).Peoplearealsomorelikelyto perceiveillusorycorrelationsbetweenpicturesofsnakesandshockthanbetweenpicturesofdamaged electricalequipmentandshock(Tomarken,Sutton,&Mineka,1995).Inaddition,specificphobiasare more likely to develop to ancestral stimuli (e.g., predators) than to aversive stimuli exclusively encounteredinmodernenvironments(e.g.,weapons;seeDeSilva,Rachman,&Seligman,1977).Col- lectively,thesedatasuggestacertainamountof‘‘preparedness”forassociatingancestrally-relevant stimuliwithparticularoutcomes. Demonstratingthatourcognitivesystemsshowancestralprioritiesinprocessingisanimportant stepinfurtheringevolutionaryaccounts.Asnotedbymany(e.g.,Buller,2005;Richardson,2007),evo- lutionaryaccountscanbedifficulttodefend,largelybecausecriticaldataareeithermissingorimpos- sibletoobtain(e.g.,cognitiveprocessescannotbefossilized).Toestablishanevolutionarylocus—that is,thepresenceofanadaptationsculptedbynaturalselection—requiresbuildingamultiprongedcase based partly on the systematic ruling out of alternative accounts (see Andrews, Gangestad, & Mat- thews,2002;Nairne,inpress).Establishingancestralprioritiesisrelevantpartlybecauseitisdifficult toseehowgenerallearningmechanisms,thoseessentiallytunedbyexperiencesthroughoutdevelop- ment,couldyieldsuperiorperformanceforsituationsandstimulithatarerootedinthedistantpast ratherthaninthepresent.Atthesametime,eachofthestudiesconductedtodatesuffersfroman inherent confound—ancestral versus modern comparisons require one to compare across different stimuli(e.g.,snakesversusguns)andsuchstimulicandifferinmanyuncontrolledways(Blanchette, 2006;Fox,Griggs,&Mouchlianitis,2007). 1.2. Thecurrentresearch Thepresentexperimentsweredesignedtoinvestigateancestralprioritiesinthesurvivalprocess- ingparadigm(Nairneetal.,2007).Asnotedearlier,processinginformationintermsofitsrelevanceto fitness can improve retention dramatically, at least compared to traditional encoding procedures (Nairneetal.,2008).Thisparadigmpresentssomemethodologicaladvantagesovertheworkjust-re- viewed,mainlybecauseallparticipantsareaskedtorememberexactlythesamematerial.Whatmat- ters to retention is how people process the information—i.e., in terms of survival or some other relevant control. This means that fitness-based retention differences cannot easily be attributed to the characteristics of thestimuli, or toassumptionsabout whatthosestimulinaturallyafford (e.g., naturalfearversuslearnedfear). Researchbasedonthesurvivalprocessingparadigmhasfirmlyestablishedthemnemonicpowerof fitness-relevantprocessing,butthelocusoftheprocessingadvantageremainscontroversial.Thereis someevidencethatsurvivalprocessingmaybespecial,atleastcomparedtootherformsofencoding, 4 J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 andnoteasilyinterpretedviastandardtheoreticalframeworks(seeNairne,inpress;Nairne&Pandei- rada,2008a).However,thefactthatfitness-relevantprocessingmaybepowerfulor‘‘special”doesnot meanthatitsmnemonicadvantagesaccruefromaspecificevolutionarytuning—moreandvariedevi- denceisneededtomakesuchacase.Asnotedearlier,establishinganevolutionarylocusforanycog- nitivephenomenonisdifficultandnosingleempiricalattackislikelytobedefinitive.Demonstrating thatsurvivalprocessingissensitivetoancestralprioritieswouldexpandtheexistingdatabaseonsur- vivalprocessingconsiderablyandstronglybolstertheevolutionaryaccount. ArecentexperimentreportedbyWeinsteinetal.(2008;Experiment2)providesinitialsupportfor thecontentionthatsurvivalprocessingmaybesensitivetoancestralpriorities.Thoseauthorscom- pared retention after people processed the relevance of words to a survival situation, but varied whether the scenario invoked an ancestral or a modern context. In one situation, following Nairne etal.(2007),peoplewereaskedtoimaginethemselvesstrandedinthegrasslandsofaforeignland withoutbasicsurvivalmaterials.Overthenextfewmonths,theinstructionsexplained,theywould needtofindsteadysuppliesoffoodandwaterandprotectthemselvesfrompredators.Theexperimen- taltaskwastoratetherelevanceofwordstothissurvivalsituation.Inasecondcondition,exactlythe samescenariowasusedbuttwocriticalwordswerechanged:citywassubstitutedforgrasslandsand predatorswasreplacedbyattackers.Escapingfrompredatorsinthegrasslands,theauthorsreasoned, is a closer fit to the problems faced in the environment of evolutionary adaptation; as a result, it shouldproducebettermemorythanprocessinginamoderncontext,eventhoughthelatterisargu- ablymorefamiliarandlikelytoleadtogreateramountsofelaboration.Consistentwiththeirhypoth- esis,betterretentionfortheratedwordswasobtainedinthegroupprocessingtheancestralscenario. Theexperimentsreportedherehadthreemaingoals:first,giventheirpotentialimportancetothe evolutionaryhypothesis,wesoughtinitiallytoreplicatethefindingsofWeinsteinetal.(2008).Will ratingtherelevanceofwordstoanancestralscenarioleadtobetterretentionthanprocessingthose samewordsinamatchedmoderncontext?Second,tohelpestablishthegeneralityofthephenome- nonweexploredtheroleofancestralprioritiesintwonewdomains,healingadangerousinfectionand collectingfoodtoeat.Bothdescribedrecurrentadaptiveproblemsthatwerelikelytobepresentinthe environmentofevolutionaryadaptation,butthenewdomainsweredesignedtoactivate‘‘gathering” strategiesratherthanpredatoravoidancestrategiessuchasthoseusedintheWeinsteinetal.exper- iment.Onceagain,allparticipantswereaskedtoprocesstherelevanceofwordstoasurvivalscenario. Whatdifferedacrossconditionswastheancestralnatureofthescenario:participantswerepresented withthesamegeneralsurvivalproblembutineitheranancestral(grasslands)oramoderncontext (city).Totheextentthatourmemorysystemsare‘‘prepared”toprocessinformationinancestralenvi- ronments(foragingashunter–gatherersinthegrasslands),weanticipatedbetterretentionafterpro- cessingwiththeancestralscenario.Finally,inanefforttoprovidegreaterinsightintotheproximate mechanismsthatmightunderlieancestraladvantages,andtogainadditionalsupportfortheevolu- tionary account, in the last two experiments we also asked participants to evaluate the ancestral andmodernscenariosalonganumberofmnemonically-relevantdimensions(e.g.,interest,familiarity, etc.). 2. Experiment1 Inthesurvivalprocessingparadigm,participantsareaskedtoratetherelevanceofwordstoasur- vivalscenario.Theratingtaskisthenfollowedbyasurpriseretentiontest,usuallyfreerecall,andper- formanceiscomparedtoanon-fitness-basedcontrol.FollowingNairneetal.(2007),Weinsteinetal. (2008)usedaschematiccontrol,movingtoaforeignland,astheircomparison.The‘‘moving”control iscloselymatchedtothesurvivalscenario,butisnotfitness-relevant(thatis,itdoesnotrequiredeal- ingwithissuesdirectlyrelatedtosurvivalorreproduction).Betterrecallforthewordsratedwiththe survivalscenario,comparedtothemovingcondition,definesthebasicsurvivalprocessingadvantage. FollowingtheWeinsteinetal.(2008)study(Experiment2)wecomparedtwoversionsofthesur- vival scenario in Experiment 1—one rooted in an ancestral environment (avoiding predators in the grasslands) and one in a modern context (avoiding attackers in a city). The critical question asked whetherthesizeofthesurvivalprocessingadvantagewouldvaryacrossthesetwosurvivalscenarios. J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 5 Weinsteinetal.foundthatthesurvivalprocessingadvantagewaslargerwhenpeopleratedtherele- vanceofwordstotheancestral(grasslands)scenario,consistentwithanevolutionaryaccount. 2.1. Method 2.1.1. Participantsandapparatus EightyPurdueundergraduatesparticipatedforcreditinanintroductorypsychologycourse.Stimuli werepresentedandcontrolledbypersonalcomputersinsessionslastingapproximately30min.Ses- sionswereconductedingroupsofuptofourparticipants. 2.1.2. Materialsanddesign Thirty-twounrelatedconcretenouns,drawnlargelyfromtheupdatedBattigandMontaguenorms (Van Overschelde, Rawson, & Dunlosky, 2004) or from the extended Paivio norms (Clark & Paivio, 2004), were used as target words in the experiment (six additional concrete nouns were used in a practice phase). The stimuli are presented in the Appendix A. Everyone rated and recalled exactly the same 32words, presented in the same order. Withina session participantsrated the relevance of 16 target words to the survival (S) scenario and 16 to the moving (M) control. Rating condition wasblockedintrialsofeightwordsintheformSMSMorMSMS;halfoftheparticipantsreceivedeach version,ensuringthateachwordwasratedequallyoftenunderbothscenarios.Thecontentofthesur- vivalscenariowasmanipulatedbetween-subjects:onegroupofparticipantswasgiventheancestral (grasslands) scenario (N=40); the other group received the modern (city) scenario (N=40). Both groups received the same ‘‘moving” instructions. Immediately following the last block of ratings, everyonecompletedashortdigit-distractortaskfollowedbyasurprisefreerecalltest. 2.1.3. Procedure On arrival in the laboratory, participants were assigned randomly to either the ancestral or the modern group. The instructions for the two survival scenarios are presented below, along with the instructionsforthemovingcontrol. 2.1.3.1. Ancestral. Inthistask,pleaseimaginethatyouarestrandedinthegrasslandsofaforeignland, withoutanybasicsurvivalmaterials.Youhaverecentlyreceivedwordthatadangerouspredatorhas beenseeninthearea.Youwillneedtoavoidand/orescapefromthepredatortoensureyoursurvival. Wearegoingtoshowyoualistofwords,andwewouldlikeyoutoratehowrelevanteachofthese wordswouldbeinyourattempttoavoidthepredator. 2.1.3.2. Modern. Inthistask,pleaseimaginethatyouarestrandedinthecityofaforeignland,without anybasicsurvivalmaterials.Youhaverecentlyreceivedwordthatadangerousattackerhasbeenseen inyourarea.Youwillneedtoavoidand/orescapefromtheattackertoensureyoursurvival.Weare goingtoshowyoualistofwords,andwewouldlikeyoutoratehowrelevanteachofthesewords wouldbeinyourattempttoavoidtheattacker. 2.1.3.3. Moving. Inthistask,pleaseimaginethatyouareplanningtomovetoanewhomeinaforeign land.Overthenextfewmonths,you’llneedtolocateandpurchaseanewhomeandtransportyour belongings.Wearegoingtoshowyoualistofwords,andwewouldlikeyoutoratehowrelevanteach ofthesewordswouldbeforyouinaccomplishingthistask. Theto-be-ratedwordswerepresentedindividually,centeredonthescreen,for5sapiece.People wereaskedtoratethewordsona5-pointscale,with1indicatingtotallyirrelevanttothescenario and5signifyingextremelyrelevant.Theratingscalewasshownjustbeloweachwordandpeoplere- spondedbyclickingontheirvalueofchoice.Theywerecautionedtorespondquickly,withinthe5-s presentationwindow,andthelaterretentiontestwasnotmentioned.Ashortpracticesessionpre- cededeachofthefirsttwoblocks(oneSandoneM). Afterthelastwordwasrated,instructionsappearedforthedigitrecalltask.Forthistask,sevendig- its,rangingbetweenzeroandnine,werepresentedsequentiallyfor1sapieceandparticipantswere requiredtorecallthedigitsinorderbytypingresponsesintoatextbox.Thedigitrecalltasklastedfor 6 J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 approximately2min.Recallinstructionsthenappeared.Participantswereaskedtowritedownthe earlier-ratedwords,inanyorder,onaresponsesheet.Thefinalrecallphaselastedfor10minandpar- ticipantswereaskedtodrawalineontherecallsheet,underthelastrecalledword,aftereachminof recall.Aclockwasdisplayedonthecomputermonitoranda‘‘beep”soundedeveryminsignalingthe participantstodrawtheline.Usingthisprocedureallowsonetoconstructcumulativerecallcurves, buttheyarenotreportedhere. 2.2. Resultsanddiscussion Thelevelofstatisticalsignificance,unlessotherwisenoted,wassetatp<.05forallcomparisons. Participantshadlittledifficultyproducingtherelevanceratingswithintheallottedtime,andnosig- nificantdifferencesincompletionrateswerefoundacrossgroupsorconditions. The data of main interest are shown in Fig. 1. These data show proportion correct recall for the rated target words for each of the main comparisons. The left-hand side of the figure shows recall for thoseparticipantsreceiving the ancestralscenario; the moderndata are shownto the right. An overall analysis of variance (ANOVA) on these data, with group (ancestral versus modern) as a be- tween-subjectvariable,andratingcondition(survivalversusmoving)asawithin-subjectvariable,re- vealedonlyasignificantinteractionbetweengroupandratingcondition,[F(1,78)=6.27,MSe=.016, g2 :074].Plannedt-testsrevealedasignificantsurvivaladvantageintheancestralgroup(ancestral p¼ versusmoving;t(39)=2.91,p<.01);thecomparablecomparisoninthemoderngroup(modernversus moving) failed to approach significance [t(39)<1.0]. Turning to the direct planned comparison be- tween the ancestral and modern scenarios, participants recalled more words after rating with the ancestralscenariobutthedifferencewasonlymarginallyreliable[t(78)=1.65,p=.052].Additional analysesusingitemsastheanalyticunit,ratherthanparticipants,revealedthesamepatternsofsig- nificance—however,therecalldifferencebetweentheancestralandthemodernscenariosdidreach conventionallevelsofstatisticalsignificanceintheitemanalysis[t(31)=1.93,p=.031]. These results confirm the main predictions of the evolutionary account. There was a significant interaction between group and rating condition, indicating a larger survival effect for the ances- trally-basedgrasslandsscenario.Infact,althoughtherewasarobustfitnesseffectfortheancestral scenario,therewasnoindicationthatsurvival-basedprocessingproducedanymnemonicadvantages in the modern scenario. Interestingly, Weinstein et al. (2008) found essentially the same pattern, althoughtheinteractionofgroupandratingconditionisnotreportedintheirarticle.Theyreportonly theoverallmeansfortheancestral(.38),modern(.31),andmovingconditions(.28),butthepatternis clearlysimilartotheonereportedhere.Itissomewhatsurprisingthatthecityscenariofailedtopro- ducearetentionadvantageoverthemovingcontrol.Survivinginacityandescapingfromattackers 0.60 0.55 call 0.50 e R ct 0.45 e orr 0.40 C n ortio 0.35 op 0.30 Pr 0.25 0.20 Ancestral Moving Modern Moving Fig.1. ProportioncorrectrecallperformanceforeachoftheconditionsinExperiment1.Thedataontheleftarefromthegroup receivingtheancestral(predator)scenario;datafromthegroupreceivingthemodern(attacker)scenarioareshownonthe right.Errorbarsrepresentwithin-subject95%confidenceintervals(asperMassonandLoftus(2003)). J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 7 Table1 Ratingandresponsetime(ms)averagesforthevariousconditionsinExperiment1. Ancestral Modern Predator Moving Attacker Moving Rating 3.30 2.63 3.07 2.74 Responsetime 2341 2486 2362 2322 areclearlyfitness-relevant,yettherewasnomemoryenhancementforsurvivalprocessinginthiscon- text.WereturntothisissueinExperiments2and3. Itisalsoofinteresttoexaminetheaverageratingsandresponsetimesforeachoftheconditions; thesedataareshowninTable1.Turningfirsttotheratingdata,anoverallANOVArevealedamain effectofratingcondition[F(1,78)=95.77,MSe=.104,g2 :551];thetargetwordswereratedasmore p¼ relevanttothesurvivalscenariosthantothemovingscenario.Therewasalsoareliableinteraction betweengroupandratingcondition[F(1,78)=11.22,MSe=.104,g2 :126],reflectingalargerdiffer- p¼ enceinratingbetweenthesurvivalscenarioandmovingfortheancestralgroup.Individualpost-hoc comparisonsdeterminedthatthedifferencesbetweenthesurvivalandmovingconditionswerereli- ableforboththeancestralandthemoderngroups(botht-testvaluesweregreaterthan4.5)andthere wasasignificantratingadvantagefortheancestralscenariooverthemodernscenario[t(78)=2.29, p<.03].Thispatternisalittletroublingbecauseitmirrorsthepatternfoundinrecall,suggestingthat therecalldifferencesmayhavebeendeterminedbytheratingdifferences.However,thereareseveral reasonstodiscountthispossibility.First,priorresearchinthesurvivalprocessingparadigmhascon- sistently shown that rating differences cannot account for observed differences in recall (Nairne & Pandeirada,2008a;Nairneetal.,2007).Second,supportingthisconclusion,thecurrentdatashowa highlyreliableratingadvantageforthemodernscenariooverthemovingcontrol,yetthereisnoevi- denceforacomparabledifferenceinrecall.Finally,therecallresultsofExperiment1essentiallyrep- licatethoseofWeinsteinetal.(2008),whofoundnosignificantratingdifferencesacrossconditions. Analysisoftheresponsetimedatashowednomaineffectsofconditionorgroup,butdidshowa reliablegroup conditioninteraction[F(1,78)=7.27,MSe=46782.17,g2 :085].Post-hoctestsre- " p¼ vealed that participants in the ancestral group took significantly longer to make relevance ratings tothemovingscenariothantheydidtothesurvivalscenario[t(39)= 2.97,p<.01];therewereno # similarsignificantdifferencesfortheparticipantsinthemoderngroup.Asaresult,differencesinre- sponsetimescannoteasilybeusedtoexplaintherecalldifferencesthatwereobtained. The results of Experiment 1 replicate the general patterns found previously by Weinstein et al. (2008).Therecalldifferencebetweentheancestralandmodernscenarioswassmallerthanreported by Weinstein et al. (2008), and statistically significant only in the item analysis, but the important interaction of group by condition was demonstrated unequivocally. The survival processing advan- tage,definedastheadvantageofsurvivalprocessingoveranon-fitness-basedmovingcontrol,waslar- ger in the ancestral than in the modern group. This data pattern is broadly consistent with the evolutionaryaccount,suggestingthatourmemorysystemsaretunedtofitness-relevantprocessing, butespeciallyincontextsthatmirrorthoselikelytohavebeenpresentintheenvironmentofevolu- tionaryadaptation. 3. Experiment2 Critics of evolutionary accounts often focus on our lack of knowledge about ancestral environ- ments,whichsomescholarsbelieveareessentiallyunknowable(e.g.,Buller,2005;Reeve&Sherman, 2007). Moreover, the idea that there is a single ancestral environment,whichuniquely defined the selectionpressuresthatledtothedevelopmentofhumancognition,isdebatable.Atthesametime, ourancestorslikelyfacedrecurrentadaptiveproblems,onesthatremainedconstantacrosssituations. Ourancestorscertainlyneededtofindfood,potentialmates,trackandescapefrompredators,andfind remedies for injuries and disease (see Tooby & Cosmides, 2005). Cognitive adaptations may have 8 J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 developedtodealwiththesespecificproblems(seeBarrett,2008)and,moreimportantly,thoseadap- tationsmaycontinuetobearthegeneralimprintofourspecificforagingpast. TheancestralscenariousedinExperiment1wasdesignedtotaponesuchrecurrentproblemfaced byourforagingancestors—survivingandescapingfrompredatorsinagrasslandsenvironment.Exper- iment 2 explores another ancestral problem,one designed to activate a ‘‘gathering” strategy rather thanonedealingexclusivelywithpredatoravoidance.Onceagain,twoversionsofthesurvivalsce- narioweredeveloped,onerootedinagrasslandsenvironment(ancestral)andoneinthecity(mod- ern). In both cases participants were told they had been hurt and a dangerous infection might be developing. In the ancestral scenario, the task was to search and find relevant medicinal plants in anattempttocuretheinfection;inthemodernscenario,thetaskwastofindrelevantantibiotics.Sce- nariotypewasmanipulatedbetween-subjectsandeachgroupreceivedanon-fitness-basedcontrolas well.RatherthanusingmovingasourcontrolinExperiment2,however,participantswereaskedto makepleasantness ratingsabout half of thewords. Pleasantnessratings have been used frequently as a control in the survival processing paradigm (e.g., Nairne et al., 2008; Weinstein et al., 2008), andaregenerallyconsideredtobeaneffectiveformofdeepprocessing(e.g.,Packman&Battig,1978). 3.1. Method 3.1.1. Participantsandapparatus OnehundredandtwentyPurdueundergraduatesparticipatedforcoursecredit.Sixtypeoplewere assignedtoeachscenariogroup.Stimuliwerepresentedandcontrolledbypersonalcomputers.Ses- sionswereconductedingroupsofuptofourparticipants. 3.1.2. Materialsanddesign TargetwordsweredrawnfromthesamepoolusedinExperiment1.Onceagain,everyonerated andrecalledexactlythesamesetofwordspresentedinthesameorder.However,unlikeinExperi- ment1,theratingtask(survivalscenarioversuspleasantness)wasnotblocked,butwasdistributed randomlythroughoutthesession.(Intermixingcanhelptoreducevariabilitycausedbyhavingdiffer- entconditionsblockedatthebeginningandendofasession.)Presentationorderwasconstrainedas follows:nomorethantworatingsofagiventypecouldoccurinarowandeachratingtypehadto occurequallyofteninthefirstandsecondhalfofthelist.Ratingtaskwasalsocounterbalancedacross participantstoensurethateachtargetwordwasratedforsurvivalandpleasantnessanequalnumber oftimes.AsinExperiment1,theratingtaskwasfollowedbyashortdistractortaskandthenasurprise freerecalltest. 3.1.3. Procedure On arrival in the laboratory, participants were assigned randomly to either the ancestral or the modern group.The instructionsfor the two survival scenarios are presented below, along with the instructionsforthepleasantnesscontrol. 3.1.3.1. Ancestral. Forthesurvivalsituation,pleaseimaginethatyouarestrandedinthegrasslandsof a foreign land, without any basic survival materials. You have recently been hurt and a dangerous infectionmightbedeveloping.Youwillneedtosearchandfindrelevantmedicinalplantstoensure your survival. We would like you to rate how relevant the word would be in your attempt to cure theinfection.Thescaleofrelevancerangesfromonetofive,withone(1)indicatingtotallyirrelevant andfive(5)signifyingextremelyrelevant.Someofthewordsmayberelevantandothersmaynot—it’s uptoyoutodecide. 3.1.3.2. Modern. Forthesurvivalsituation,pleaseimaginethatyouarestrandedinthecityofaforeign land, without any basic survival materials. You have recently been hurt and a dangerous infection might be developing. You will need to search and find relevant antibiotics to ensure your survival. We would like you to rate how relevant the word would be in your attempt to cure the infection. Thescaleofrelevancerangesfromonetofive,withone(1)indicatingtotallyirrelevantandfive(5) J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 9 signifyingextremelyrelevant.Someofthewordsmayberelevantandothersmaynot—it’suptoyou todecide. 3.1.3.3. Pleasantness. Fortheotherdimension,wewouldlikeyoutoratethepleasantnessoftheword. Thescaleofrelevancerangesfromonetofive,withone(1)indicatingtotallyunpleasantandfive(5) signifyingextremelypleasant.Someofthewordsmaybepleasantandothersmaynot—it’suptoyou todecide. AsinExperiment1,theto-be-ratedwordswerepresentedindividually,centeredonthescreen,for 5sapiece.Theratingscalewasshownjustbeloweachwordandpeoplerespondedbyclickingontheir valueofchoice.Boththesurvivalscenario(ancestralormodern)andthepleasantnesstaskwerefully explainedtoeachparticipantatthebeginningofthesession.Inthesessionitself,eachwordwaspre- sentedalongwithaquestionthatexplainedtheratingtasktobeperformed(Howrelevantisthisword to the survival situation?; How pleasant is this word?). Participants were cautioned to respond quickly,withinthe5-spresentationwindow,andthelaterretentiontestwasnotmentioned.Thepro- cedurefortheshortdistractorperiodandtherecalltestmatchedExperiment1. 3.2. Resultsanddiscussion AsinExperiment1,participantshadnotroublecompletingtherespectiveratingdecisionswithin the5-spresentationwindow.ThedataofmaininterestareshowninFig.2,whichshowsproportion correctrecallperformanceforthewordsratedinthesurvivalandpleasantnesstasks.Theleft-hand side of the figure shows performance for the group receiving the ancestral scenario; the data for themoderngroupareshownontheright. AmixedANOVAwithgroup(ancestralversusmodern)asabetween-subjectvariableandcondition (survival versus pleasantness) as a within-subject variable revealed a highly reliable main effect of condition [F(1,118)=45.61, MSe=.010, g2 :279] as well as a reliable interaction between group p¼ andcondition[F(1,118)=5.31,MSe=.010,g2 :043].Pairedt-testsconfirmedasurvivaladvantage p¼ (compared to pleasantness) for both the ancestral [t(59)=6.14, p<.001] and the modern groups [t(59)=3.30, p<.002]. Mostimportantly, rating wordswithrespecttothe ancestralscenarioled to better recall performance than did rating words about the modern scenario [t(118)=2.19, p=.03]. Additional item analyses revealed the same patterns, including the significant advantage for the ancestraloverthemodernscenario[t(31)=2.04,p<.05],butthedifferencebetweensurvivalprocess- ingandpleasantnessdidnotquitereachstatisticalsignificanceinthemoderncondition[t(31)=1.88, p<.07]. 0.60 0.55 all c 0.50 e R ct 0.45 e Corr 0.40 n ortio 0.35 p o 0.30 Pr 0.25 0.20 Ancestral Pleasantness Modern Pleasantness Fig.2. ProportioncorrectrecallperformanceforeachoftheconditionsinExperiment2.Thedataontheleftarefromthegroup receivingtheancestral(medicinalplants)scenario;datafromthegroupreceivingthemodern(antibiotics)scenarioareshown ontheright.Errorbarsrepresentwithin-subject95%confidenceintervals(asperMassonandLoftus(2003)). 10 J.S.Nairne,J.N.S.Pandeirada/CognitivePsychology61(2010)1–22 Table2 Ratingandresponsetime(ms)averagesforthevariousconditionsinExperiment2. Ancestral Modern Medicinalplants Pleasantness Antibiotics Pleasantness Rating 3.50 2.91 3.38 2.92 Responsetime 3104 2864 3101 2849 Average rating and response time data are shown in Table 2 for each group and condition. The mixedANOVAontheratingdatashowedthatthesurvivalratingsweresignificantlyhigherthanpleas- antnessratings[F(1,118)=114.67,MSe=.145,g2 :493];neitherthemaineffectofscenarionorthe p¼ group ratingconditioninteractionwassignificantinthisanalysis.TheresultsofamixedANOVAon " responsetimerevealedonlyamaineffectofratingcondition:participantstooksignificantlylongerto make survival ratings than pleasantness ratings [F(1,118)=81.15, MSe=44685.18, g2 :407]. The p¼ samepatternsofresultswereobtainedintheitem-basedanalysis. Generally,theresultsofExperiment2replicatethemainfindingsofExperiment1.Onceagain,the criticalgroup conditioninteractionwassignificant—therewasalargersurvivalprocessingeffectfor " the group receiving the ancestral (grasslands) scenario. Moreover, individual planned comparisons confirmedthatprocessing withtheancestralscenarioproducedhigher levels ofrecallthan did the modernscenario—and,importantly,neithertheancestralnorthemodernscenariosproducedsignif- icant differences on either the rating or the response time measures. Consequently, the ancestral advantage reported here provides additional supporting evidence for the evolutionary account: fit- ness-based processing is particularly effective in contexts that presumably mirror those found in theenvironmentsofevolutionaryadaptation. Itisinterestingtonotethatsurvivalprocessingproducedenhancedrecallforboththeancestral andthemodernscenariosinExperiment2,relativetothepleasantnesscontrol(althoughthemod- ern-pleasantness advantage was only marginally significant in the item analysis). In Experiment 1, there was a robust survival advantage only for the ancestral scenario. The pattern seen here could bedue,inpart,toouruseofthe‘‘pleasantness”ratherthan‘‘moving”controlcondition.Ratingitems forpleasantnessisahighlyeffectiveformofdeepprocessing,butittendstoinduceprimarilyitem- specificratherthanrelational(oracross-item)processing.Itiswell-knownthatitem-specificprocess- ingismosteffectivewhento-be-recalleditemsarerelated,suchaswhenalistiscategorized,because ithelpsonediscriminatebetweenitemsthatdidanddidnotoccur(seeHunt&Einstein,1981;Hunt& McDaniel,1993).Giventhatourto-be-ratedwordswereunrelated(seeAppendixA),itispossiblethat both survival scenarios induced greater amounts of relationalprocessing than did the pleasantness tasktherebyaccounting,atleastinpart,forthesurvivaladvantagesseeninthisexperiment.However, Nairne and Pandeirada (2008a) have shown that survival processing produces better recall perfor- mancethanpleasantnessprocessingevenwhenwordlistsarecategorized,soitisunlikelythaten- hanced relational processing can completely explain the survival advantages seen here, or in other experimentsusingthesurvivalprocessingparadigm.Moreover,importantly,thecriticalcomparison betweentheancestralandmodernscenarioswasreliableinboththesubjectandtheitemanalyses anddoesnotdependontheuseofthepleasantnesscontrol(seealsoExperiment4). 4. Experiment3 TheancestralscenariosofExperiments1and2weredesignedtoevokerecurrentadaptiveprob- lems faced by our foraging ancestors—escaping from predators and treating a dangerous infection. Experiment3investigatesyetanotherbasicrequirement—searchingfornourishment.Onceagain,par- ticipantswererequiredtoratetherelevanceofwordseithertoasurvivalscenario,onerootedinthe grasslandsandoneinacity,orforpleasantness.Thesurvivaltaskaskedparticipantstoimaginegain- ingneedednourishmenteitherbysearchingforandbuyingfoodinacity,orbysearchingforandgath- ering edible plants in the grasslands. Consistent with the evolutionary account, and the results of Experiments 1 and 2, we anticipated the best final free recall for words rated with respect to the ancestralscenario.

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modern priorities” (New et al., 2007, p. 16603). In the learning domain, evolutionarily-significant stimuli can be easier to associate with aversive.
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