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29/4/2016 CulturalEvolution(StanfordEncyclopediaofPhilosophy)

Stanford Encyclopedia of Philosophy


CulturalEvolution
FirstpublishedSunDec23,2007substantiverevisionWedFeb20,2013

Inthebroadestterms,theoriesofevolutionseektoexplainwhyspeciesarethewaystheyare.Formany
evolutionists,thismeansexplainingthepossessionbyspeciesofcharacteristicadaptations.Italsomeans
explainingdiversitywithinspecies.Thegeneralmarkofmoderntheoriesofculturalevolutionistheir
insistenceonthesignificanceofculturalinheritanceparticularlyvariousformsoflearningfromothers
forbothofthesequestions.Theprimafaciecaseforculturalevolutionarytheoriesisirresistible.
Membersofourownspeciesareabletosurviveandreproduceinpartbecauseofhabits,knowhowand
technologythatarenotonlymaintainedbylearningfromothers,theyareinitiallygeneratedaspartofa
cumulativeprojectthatbuildsondiscoveriesmadebyothers.Andourownspeciesalsocontainssub
groupswithdifferenthabits,knowhowandtechnologies,whichareonceagaingeneratedand
maintainedthroughsociallearning.Thequestionisnotsomuchwhetherculturalevolutionisimportant,
buthowtheoriesofculturalevolutionshouldbefashioned,andhowtheyshouldberelatedtomore
traditionalunderstandingsoforganicevolution.

1.WhatisCulturalEvolution?
2.NaturalSelectionandCulturalInheritance
3.HistoricalPedigree
4.Memes
5.ProblemswithMemes
6.CulturalEvolutionwithoutMemes
7.TheExplanatoryRoleofCulturalEvolutionaryTheories
8.PopulationThinking
9.Evolvability
10.CulturalPhylogenies
Bibliography
AcademicTools
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1.WhatisCulturalEvolution?
Theoriesofculturalevolutionneedtobedistinguishedfromtheorieswithinevolutionarypsychology,
eventhoughbothmayinvolveanapplicationofevolutionaryideastotheexplanationofcultural
phenomena.Theevolutionarypsychologist(e.g.ToobyandCosmides1992)tendstoassumethatthe
mostimportantinheritancemechanisminallspeciesourownincludedisgeneticinheritance.
Evolutionarypsychologyregardsthehumanmindasevolvingthroughaconventionalprocessofnatural
selectionactingongeneticallyinheritedvariation.Forexample,anevolutionarypsychologistmight
explainthewidespreadtasteamonghumansforfattyfoodsintermsoftheimportanceinourspecies'
distantpastofconsumingasmuchfataspossibleonthoserareoccasionswhenthecircumstances
presentedthemselves.Suchahypothesiscanalsohelptoexplainnovelculturaltrends:therecent
increaseinobesityisexplainedastheresultofanovelenvironmentalchangetheincreasedavailability
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ofcheap,highfatfoodsactinginconcertwithaonceadaptive,nowdangerous,gustatorypreference.

Darwinbelieved,asdobiologiststoday,thatnaturalselectioncanexplaintheoriginofmanycomplex
adaptivetraits.InDarwin'soriginalpresentationofnaturalselection,herequiresthatparentorganisms
differintheirabilitiestosurviveandreproduce,andthatoffspringresembletheirparentsintermsofthe
traitsthatpromoteorinhibittheseabilities(Darwin1859).Thisexplanatoryschemaislargelyneutral
regardingwhatmechanismaccountsforparentoffspringresemblance.Forexample,offspringmight
learnskillsfromtheirparents,andtherebycometoresemblethembehaviourally.Fromtheperspectiveof
naturalselectionexplanations,itdoesnotmatterwhyoffspringresembleparents,onlythattheydo
resemblethem.

Darwin'stheoryofnaturalselectionexplainsadaptationbyappealingtowhatwenowcallvertical
transmissiontheinheritanceofparentaltraitsbyoffspring.Aswehaveseen,culturalprocessessuchas
learningmight,inprinciple,underpinthisformofinheritance.Butwedonotlearnonlyfromourparents
wealsolearnfrompeers,authorityfiguresandsoforth.Thisisknownasobliquetransmission.Once
weacknowledgethepossibilitythatlearningcanunderpinnaturalselection,wealsoacknowledgethata
theoryofevolutionatheorywhichseekstoexplainchange,includingadaptivechangeinapopulation
mayalsoneedtobefurtherexpandedtoencompassobliquetransmission.Theadmittanceofoblique
transmissionintoevolutionarytheorynecessitatesfarmoreradicalrevisionstotraditionalDarwinian
modelsofevolution.Thisisbecauseobliquetransmissionopensupthepossibilitythatsometraitsmay
spreadthroughapopulationinspiteofthefactthattheyreducethefitnessoftheindividualswhobear
them.

2.NaturalSelectionandCulturalInheritance
Inaclassicearlyworkofculturalevolution,CavalliSforzaandFeldman(1981)ask(amongother
things)howwecanexplaindecliningbirthratesamongItalianwomeninthenineteenthcentury.These
womenwentfromhavingaroundfivechildrenonaveragetohavingonlytwo.Itwouldbeextremely
implausibletoarguethatthisoccurredasresultofnaturalselection(Sober1991,482).Itwouldbe
implausible,forexample,toarguethatthefitnessofwomenwithsmallerfamilieswasgreaterthanthe
fitnessofwomenwithlargerfamilies.Trueenough,anindividual'slongtermfitness(measuredinterms
ofnumbersofgrandchildren,orgreatgrandchildren)maysometimesbeaugmentedbyhavingafew
strongoffspringratherthanlotsofweakones(Lack1954).ButsurelyItalianwomencouldhaveraised
morethantwochildrentobehealthyadults.CavalliSforzaandFeldmaninsteadarguethatthepractice
ofhavingfewerchildrenspreadthroughItalybecausewomenacquiredthetraitbothfrompeersandfrom
individualsfromtheirmother'sgeneration,throughmodesofculturaltransmission.Formsofoblique
transmissionarerequiredtoexplainthistransition,becauseifculturaltransmissionwasalwaysvertical,
thenthetraitofhavinggreaternumbersofoffspringwouldbemaintainedinthepopulationbynatural
selection,albeitselectionactingviaculturalinheritance.

Onemightreacttothiswithconfusion:whyisabodyoftheoryneededtomaketheseclaims?Ofcourse
weacquiretraitsfromothersbylearning.Andofcoursethoseothersfromwhomwelearncaninclude
peersaswellasparents.Inpart,wecanrespondtothisbewildermentbypointingtothevirtuesof
clarifyingtheconditionsrequiredforculturalinheritancetoovercomenaturalselection.CavalliSforza
andFeldmanarguethatifwomensimplyacquiredwhicheverpreferenceforfamilysizewasthemost
widelyadoptedintheirlocalculturalenvironment,thenculturalinheritancewouldnothaveenoughofan
effecttoovercomenaturalselection.Womenmustbedisposedtoacquirethepreferenceforsmallfamily
sizeevenwhenitispresentinonlyasmallproportionoftheirculturalcircle,ifsmallfamilysizeisto
replacelargefamilysizeinthepopulationasawhole.Thisisanilluminatingclaim,andittakesa
quantitativemodeltoshowit.
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Thisquestionofwhatbenefitistobehadfromsettingthesesortsofclaimsinaquantitativetheorywill
beraisedinmoredetaillaterinthisarticle.Forthemoment,notethatonemayalsoaskwhyitshouldbe
thecasethatweareabletolearnfromnonparentsatall,giventheadaptivecostsofsuchadisposition.If
thetendencyofItalianwomentolearnfromtheirpeershasledthemtoreducetheirfitnessbyreducing
theirfamilysize,whydidnaturalselectionallowsuchlearningdispositionstobecomeestablishedinthe
firstplace?BoydandRicherson,twootherpioneersinculturalevolutionarytheory,claimthattheoverall
adaptivebenefitsoflearningfromnonparentsinfactoutweightheoveralladaptivecosts(Richersonand
Boyd2005,Ch.4).Theygiveseveralreasonsforthisview.Supposeaninventive(orlucky)individualis
abletodiscoversomebehaviour,ortechnique,whichaugmentsfitness.Ifotherindividualsinthe
populationcancopythatbehaviour,thentheirfitnesswillprobablybeaugmented,too.Itwilloftenbe
difficultforindividualstoascertainwhichbehavioursinfactaugmentfitness,hencewhichbehaviours
shouldbecopied.Theproblem,then,ishowtotunealearningmechanismsothatbeneficialbehaviours
arecopied,whilenonbeneficialbehavioursarenot.

BoydandRichersonsuggestthatprestigebiascanovercomethisproblem:ifindividualscopytechniques
fromthosewhoareinprestigiouspositions,thenthisincreasesthechancesthattheywillcopytechniques
thatare,infact,beneficial.Astheyputit,Determiningwhoisasuccessismucheasierthandetermining
howtobeasuccess(RichersonandBoyd,2005,124).Moreover,evidencehasbeenaccumulatingfor
therealityofprestigebias.HenrichandBroesch(2011)haveargued,basedonfieldworkinFiji,thatan
individual'sperceivedsuccessinasingledomainofactivity(forexample,yamcultivation)predicts
whetherthatindividualwillbeaskedforadviceinotherdomains(forexample,fishing).Inotherwords,
theyclaimthatindividualsareaccordedabroadformofprestige,whichaffectstheirlikelihoodof
servingasaculturalmodel.Thevalueofprestigebiasreliesonthesuppositionthatthoseindividuals
whoareabletogetthemselvesintoprestigiouspositionshaveabetterthanaveragetendencytomakeuse
offitnessenhancingtechniques.Thisheuristicwillnotbefailsafe:afterall,noteverytechniquea
prestigiousindividualuseswillalsoaugmentfitness,andsomeindividualsmaybeaccordedprestige
withoutgoodcause.Butthequestionwhichsettlestheplausibilityofnaturalselectionexplainingprestige
biasisnotwhetherprestigebiaswillsometimesleadtothecopyingofmaladaptivetechniquesthe
question,rather,iswhetherindividualswholearnfromtheprestigiouswilltendtobefitteronaverage
thanindividualswhoeitherdonotlearnatall,orwhoareequallylikelytolearnfromanymemberofthe
population,regardlessoftheirsocialstatus.

RichersonandBoyd(2005,12022)suggestthatotherlearningheuristicsmaybeadaptive.Oneofthese
theycallconformistbias.TheyarguethatimitationofthecommontypetheWheninRomeruleis
morelikelythannotimitatingatall,andmorelikelythanimitationofarandomlychosenmemberofa
population,toprovideanindividualwithbehavioursthatareappropriatetonovelsituations.Thismay
meanacquiringbehavioursappropriatetoanewbiologicalenvironment:whenmovingintoanew
habitat,withunknownplantsandanimals,itisbesttoeatthefoodsthelocalseat,foronetherebyavoids
poisoning.Butitcanalsoleadtothegenerationofsociallyappropriatebehaviours,whichwillobviate
ostracismorattack.HarrisandCorriveau's(2011)empiricalworkconcludesthatwhileyoungchildren
areunselectivewithregardtowhattheylearn,theyarefarmoreselectiveregardingwhomtheylearn
from.Moreover,theyarguethatchildrentendtoseekoutculturalconformistsasindividualswhomthey
shouldtrust.Thesefindingsbroadlysupporttheexistenceofaformofconformistbias.

Theseexamplesshowthenatureoftheinteractionbetweenculturalevolutionarythinkingandmore
traditionalnaturalselectionthinking.Naturalselectionactingongeneticvariationcanestablish
dispositionstolearnfromnonkininspiteofthefactthatundersomecircumstancesthesedispositions
leadtotheproliferationofmaladaptivetraits.Itisworthnotingthatthisaspectofmuchcultural
evolutionarythinkingretainsastrongmethodologicalaffinitywiththeevolutionarypsychological
approachitissometimescontrastedwith.Learningdispositionsthemselvesareoftenunderstoodby
culturalevolutionistsasgeneticallyinheritedadaptations,producedinresponsetoadaptiveproblems
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facedbyourearlierancestors.Somerecentcriticsofculturalevolutionarythinking(e.g.Heyes2012)
consequentlyarguethatitisnotculturalenough,forittendstodownplaythepossibilitythatlearning
dispositionsthemselvesmightbeinheritedthroughformsoflearning.Allagree,though,thatoncethese
learningdispositionsareinplace,weshouldnotassumethateverytraitinapopulationmustbe
explainedbyreferencetothebiologicalfitnessbenefitithasconferredinthepast.Evolutionary
adaptationiststendtoask,ofanygiventrait,whateffectmighthavelednaturalselectiontofavourthat
trait.Evenifanadaptationiststanceofthissortisjustifiableforlearningmechanisms(andcultural
evolutioniststypicallyareadaptationistsinthisrespect)thisdoesnotmeanthatanadaptationiststanceis
justifiableforlearnedtraits.

3.HistoricalPedigree
Thenotionthatcultureitselfevolves,andthatDarwinianinsightscanbeappliedtounderstanding
culturalchange,isbynomeansnew.Averyearlyexampleofculturalevolutionarythinkingcomesfrom
WilliamJames:

Aremarkableparallel,whichtomymindhasneverbeennoticed,obtainsbetweenthefacts
ofsocialevolutionandthementalgrowthoftherace,ontheonehand,andofzoological
evolution,asexpoundedbyMrDarwin,ontheother.(James1880,441)

James'spaperisprimarilyconcernedwithusingwhatheregardsasaproperunderstandingofDarwinism
tounderminethesocalledevolutionaryphilosophyofMrHerbertSpencer(ibid.,422).Spencerhad
arguedthatgreatmenwereofsecondaryimportanceindeterminingthecourseofhistory,onthe
groundsthatBeforehecanremakehissociety,hissocietymustmakehim(fromSpencer'sStudyof
Sociology,quotedinibid.,449).Thegreatmanneedstobemade,andsocietydoesthis.Henceultimately
itissocietyitselfthatexplainssocialchange.

JamesarguesthatthecentralkeytoDarwin'snaturalselectionmechanismistodecouplethecausesof
variationfromthecausesofselection(seeLewens,2007,Ch.2).Variationsareproducedbyunknown
causes,andtheenvironmentselectsamongthem.Variationsthemselves(forJames'sDarwin)are
inexplicable.Thesameistrueofgreatmen:Thecausesofproductionofgreatmenlieinasphere
whollyinaccessibletothesocialphilosopher.Hemustsimplyacceptgeniusesasdata,justasDarwin
acceptshisspontaneousvariations(James,1880,445).Greatmen,likespontaneousvariations,are
essentialandinexplicableelementsoftheevolutionaryprocess.JustasDarwin'stheorycredits
environmentandvariationwithdistinctive,yetvital,roles,sobothgreatmenandthesocialenvironment
areimportantfortheexplanationofsocialchange:

Thissocialevolutionisaresultantoftheinteractionoftwowhollydistinctfactors:the
individual,derivinghispeculiargiftsfromtheplayofphysiologicalandinfrasocialforces,
butbearingallthepowerofinitiativeandoriginationinhisownhandsandsecond,the
socialenvironment,withitspowerofadoptingorrejectingbothhimandhisgifts.Both
factorsareessentialtochange.(Ibid.,448)

Thereareproblemsassociatedwithanyefforttotracethepedigreeofculturalevolutionarytheoriesback
toDarwinhimself.Oneofthereasonsforthisisthatculturalevolutionarytheoriesoftendefine
themselvesinoppositiontothosewhichclaimthatgeneticinheritanceistheonlysignificantinheritance
mechanism.ClearlyonecannotcastDarwinasaculturalevolutionistinthismanner,forhehadnonotion
ofgeneticinheritancetooppose.Havingsaidthis,Darwindidbelievethatwhatwaslearnedinone
generationcouldbeinheritedinlatergenerations.Butfarfromdistinguishingculturalinheritancefrom
organicinheritance,Darwinthoughtthatallinheritanceshouldbeexplainedbythetransmissionof

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gemmules.Thesewereunderstoodtobeparticlesproducedthroughoutthebody,ofacharacterspecific
tothebodypartthatproducesthem.Darwinbelievedthatgemmulesthentravelledtothegonads,where
theyweretransmittedtooffspringinthesexcells.Darwinclaimedthatgemmuleswereproduced
throughoutthebodyinordertoexplaintheinheritanceofacquiredcharacteristics.Soinonesense
Darwinisinalignmentwithmodernculturalevolutionistshebelievedthatcharacteristicslearned
duringthelifeofaparentcouldbetransmittedtooffspring.ButinanothersenseDarwinisopposedto
modernculturalevolutionists,forratherthandistinguishingbetweendifferentinteractinginheritance
systems(e.g.culturalandgeneticinheritance),Darwintendstousethetransmissionofgemmulesto
explaintheinheritanceofalltypesoftrait.

ThereareotherrespectsinwhichonemightchoosetoregardDarwinasaprotoculturalevolutionist.
Darwinsometimesintegratesdiscussionoftechnologicalevolutionintohisbroaderdiscussionsofnatural
selection.IntheDescentofMan,Darwinpausestodiscusstechnicalinnovation,arguingthatsuccessful
innovationswillusuallybeimitated,therebyincreasingthesuccessofagroupasawhole,increasingthe
sizeofthatgroup,andconsequentlyincreasingthechancesofinventivemembersbeingbornintoit
(Darwin1877).Thisexplanationcombinesnaturalselection'scentralconcernwithreproductiveoutput,
andculturalevolution'scentralconcernwithimitation.Darwinwrites:

ifsomeonemaninatribe,moresagaciousthantheothers,inventedanewsnareor
weapontheplainestselfinterest,withouttheassistanceofmuchreasoningpower,would
prompttheothermemberstoimitatehimandallwouldthusprofitIfthenewinvention
wereanimportantone,thetribewouldincreaseinnumber,spread,andsupplantother
tribesInatribethusrenderedmorenumeroustherewouldalwaysbearathergreater
chanceofthebirthofothersuperiorandinventivemembers.(Darwin1877,154)

Finally,Darwinendorsestheview,widelyfavouredthesedays,thatnaturalselectionneednotacton
organisms.Rather,naturalselectionissubstrateneutral.Anaturalselectionprocesscanoccurwhenever
certainabstractconditionsthesedaysoftenexpressedasdifferentialreproductionwithinheritanceare
met.Darwinexplicitlyendorsestheviewthatnaturalselectioncanactonentitiesotherthanorganismsin
thecontextoflanguagechange,aculturalphenomenon.ThispositionisbrieflyexploredintheOriginof
Species,andfurtherexpandedintheDescentofMan.Inthiswork,heendorsestheopinionofMax
Mller:

Astruggleforlifeisconstantlygoingonamongstthewordsandgrammaticalformsineach
language.Thebetter,theshorter,theeasierformsareconstantlygainingtheupperhand,and
theyowetheirsuccesstotheirowninherentvalue.(Darwin1877,113)

Darwinassertsthatthisisnomereanalogy:Thesurvivalorpreservationofcertainfavouredwordsin
thestruggleforexistenceisnaturalselection.Thisclaimthatculturalentitiesofvarioussortscan
undergonaturalselectionprocessesintheirownrightisnotanecessaryfeatureofatheoryofcultural
evolution.Culturalevolutionarytheoryingeneralrequiresonlyasystematicefforttomodeltheeffectsof
culturalinheritance,andonemightdecidethatthinkingintermsofnaturalselectionactingonunitsof
cultureisnotthebestwayofdoingthis.Wewillinvestigatetheseissuesinmoredetaillaterinthis
article.

WehavealreadymentionedHerbertSpencer,andSpencerissometimesregardedasakeyearlyadvocate
ofeffortstoapplyevolutionarythinkingtohumanculture(e.g.JablonkaandLamb2005,2122).As
earlyas1855,inhisPrinciplesofPsychology,Spencerproposedaformofevolutionaryepistemology,
arguingforathirdwaybetweenempiricism'semphasisonthenecessityofexperienceforknowledge,
andrationalism'sinsistenceontheimportanceofaprioriknowledge.Spencerreasonedthatifthe
experiencesofpastgenerationswereimprintedonhumanminds,thenitwouldbetrueboththatsome
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formsofknowledgeincurrentgenerationswereapriori,andalsothatthisknowledgehaditsoriginsin
experience,albeittheexperienceofourancestors.Darwinhimselfhadmadeabriefnotealongsimilar
linesinhisMnotebook:PlatosaysinPhaedothatournecessaryideasarisefromthepreexistenceof
thesoul,arenotderivablefromexperience.readmonkeysforpreexistence.(Barrettetal1987,551)
Muchlaterinthetwentiethcentury,KonradLorenzwouldargueforasimilarsetofviewsinhiseffortsto
seetheKantianapriorithroughthelensofevolutionarybiology,andnaturalselectionmorespecifically
(Lorenz1941).ThereisanimportantdifferencebetweenDarwinandLorenz,whichthesesuperficial
similaritiesmighthide.Darwin'scommentsdonotpresupposetheactionofnaturalselectionasthe
mechanismbywhichtheseadaptiveideasbecomecommoninthespeciesrather,themechanismofuse
inheritanceiswhatexplainsthepreservationofconceptsandtechniquesthatareseentoworkwell.

AlthoughSpencerissometimescreditedwithinitiatingtheapplicationofevolutionarythinkingto
culture,Spencer'scontributionsinthisdomainandothersareoftenregardedasscientificallyworthless
(althoughseeJablonkaandLamb2005,3723foranexception).ErnstMayr,forexample,claimedthat
ItwouldbequitejustifiabletoignoreSpencertotallyinahistoryofbiologicalideasbecausehispositive
contributionswerenil(Mayr1982,386).Spencerisusuallytreatedharshlyforhisadherencetothe
importanceofuseinheritance,accordingtowhichhabitsinitiallylearnedareeventuallyinherited
automaticallyinoffspring.ThisformofinheritancewouldbeclassedbymanyasLamarckian,in
contrasttotheDarwinianformsofinheritancethataretypicallyplacedintheforegroundin
presentationsofmodernevolutionarytheory.

SomerecentmoderntheoristshavearguedthatLamarckianinheritanceshouldnotbedismissedoutof
hand(e.g.JablonkaandLamb1995).Whateverwethinkofthismove,thetendencytopraiseDarwin
whiledamningSpenceroftenoverlooksthefactthatDarwin,too,believedinthebiologicalsignificance
ofuseinheritance,anditfiguredstronglyinhisownviewsofculturalevolution.Spencerisalso
criticisedforhissocialDarwinistbeliefs,butDarwin,too,wasasocialDarwinistofsorts,andheld
evolutionaryviewsregardingrace,socialdegenerationandothersuchtopicsthatmostwoulddismiss
today(seeLewens2007,chaptereight).Aswehaveseen,Darwin'stheoryofpangenesiswasdeveloped
partlyinordertoexplainwhathetooktobethephenomenaofuseinheritance,andageneralaccountof
useinheritanceplayedanimportantroleinDarwin'sculturalevolutionaryaccountofhumanmoral
progress.Indeed,atonepointintheDescentofMan,DarwinquotesSpenceratlengthandwithapproval:

Ourgreatphilosopher,HerbertSpencer,hasrecentlyexplainedhisviewsonthemoralsense.
Hesays,Ibelievethattheexperiencesofutilityorganisedandconsolidatedthroughallpast
generationsofthehumanrace,havebeenproducingcorrespondingmodifications,which,by
continuedtransmissionandaccumulation,havebecomeinuscertainfacultiesofmoral
intuitioncertainemotionscorrespondingtorightandwrongconduct,whichhaveno
apparentbasisintheindividualexperienceofutility.(Darwin1877,148)

4.Memes
SeriouseffortstoconstructculturalevolutionarytheoriescanbetracedtotheworkofLumsdenand
Wilson(1981),CavalliSforzaandFeldman(1981),andBoydandRicherson(1985).Alloftheseauthors
haveattempted,inonewayoranother,toproduceformalmodelsthatcanintegratetheeffectsofcultural
inheritanceintomorestandardbiologicalmodelsofevolution.Wehavealreadylookedatsomeofthe
claimsofthesetheorists,butbeforelookingattheirworkinmoredetail,letuslookatthetheoryof
memetics.Thistheory,originallyputforwardbyRichardDawkins(1976),isperhapsthebestknown
attemptstoapplyevolutionarythinkingtoculturethatsaid,whileithasenjoyedconsiderablepopular
attention,ithasnotbecomewellestablishedinscientificcircles.Instead,theschoolofBoydand
Richersonhasbeenfarmoresuccessful,forreasonsthatareexplainedbelow.
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Thememetheoryseekstodrawaverystronganalogybetweenevolutionattheculturallevel,and
biologicalevolution.Itbeginswithanabstractcharacterisationofselectionasaprocessrequiringentities
thatreproduce,suchthatparentsresembleoffspring.Memeticstakestheview,popularisedbyDawkins,
thatentitieswhichhavetheabilitytomakefaithfulcopiesofthemselvessocalledreplicatorsare
requiredtoexplainthistransgenerationalresemblance.Instandardbiologicalmodelsofevolutionitis
assumedthatgenesaretherelevantreplicators.Genesmakecopiesofthemselves,andthisability
explainswhyoffspringorganismsresembletheirparents.Ifcultureistoevolve,itthereforebecomes
necessarytofindsomeformofculturalreplicatorthatexplainsculturalinheritance.Memesplaythisrole.
Dawkinsgivesalistofsomeexemplarymemes:tunes,ideas,catchphrases,clothesfashions,waysof
makingpotsorofbuildingarches.Notethatwhileitissometimesassumedthatallmemesareideas(and
viceversa)Dawkins'slistincludesothertypesofthing,suchaswaysofmakingpots,whichare
techniques.

Dawkins'sclaimisthatideas,forexample,canbeconceptualisedasentitiesthathopfrommindtomind,
makingcopiesofthemselvesastheygo.Onthefaceofthings,thisseemsanattractiveproposition.Just
asgenesmakecopiesofthemselvesatdifferentratesaccordingtotheireffectsontheorganismsthatbear
themandontheirlocalenvironments,soideasmakecopiesofthemselvesatdifferentratesaccordingto
theireffectsontheorganismsthatbearthemandontheirlocalenvironments.Inacommunityof
scientists,forexample,differenthypothesesareentertained,andsomecometobebelievedmorewidely
thanothers.Ahypothesisthatbeginsinthemindofoneortwoscientiststherebyspreads,untilitis
widelyheldintheresearchcommunity.Anotherhypothesisquicklydies.Wecanperhapscharacterisethe
featuresthatmakesomehypotheseslikelytospread,andotherslikelytoperish.Fithypothesesmay
havepredictivepower,orsimplicity,ortheymayintegratewellwithexistingbodiesoftheory.Notethat
whatthisexampleshowsisthattakingthememe'seyeperspectivedoesnotliterallyshowthatweare
beingmanipulatedbyselfishculturalreplicators.Onecandescribescientificchangeasastruggle
betweenselfishmemes,butonecanalsodescribejustthesameprocessintermsofscientistschoosingto
accept,ortoreject,theoriesbyreferencetofamiliarcriteriaofexplanatorypower,theoreticalelegance
andsoforth.Itisonlyanincidentalfeatureofthemetaphorofmemeticselfishnessthatappearsto
deprivehumansofcontroloverwhichideastheydo,anddonot,accept.

5.ProblemswithMemes
Therearevariousproblemsassociatedwithmemeticviews,mostofwhichfocusonlimitationsofthe
gene/memeanalogy.Theseworriesaresometimesraisedbytheoristsfromthesocialscienceswhoare
hostiletoevolutionarytheoriesofculture.Buttheyarealsoraisedquitefrequentlybycultural
evolutionistswhoarguethatthememeconceptisnottherightwaytogroundatheoryofcultural
evolution:itisessentialtobearinmind,then,thatculturalevolutionarytheoriesingeneraldonotrequire
thememetheorytobetrue(seeHenrichetal2008).Herearesomeofthemostfrequentlyencountered
argumentsagainstthememeconcept(theremainderofthissectiondrawsonLewens,2007,Ch.7):

Culturalunitsarenotreplicators:Replicators,remember,aresupposedtobeunitsthatmakecopiesof
themselves.Somecriticsofthememeconceptarguethatthereisnoknownmechanismthatcould
explainhowmemesarecopied.Butthisisamistake.Anideacanbecopiedsimplythroughobservation
andinference:agentBcanobservethebehaviourofagentA,inferthatAholdssomebeliefX,and
therebycometoholdthesamebeliefasA.Ideascanalsobecopiedusinglinguisticcommunication.
AgentAmightbeconvincedofbeliefX,shetellsBaboutit,andBcomestobelieveXtoo.Inbothcases
onecansaythatbeliefXmakesacopyofitself,albeitvialanguage,inference,andsoforth.Amore
pressingworryformemeticsisthatimitationisusuallytooerrorpronetounderpinreplication.IfImake
acakeonthebasisofasecretfamilyrecipe,youeatthecake,andyouthenattempttomakeanotherone,
thenthechancesarethattherecipeyouhituponwillnot,infact,beexactlythesameastheoneIused,
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evenifyouareabletomakeasimilartastingcake.Anothersignificantworryformemeticsisthatwhen
thesameideasdospreadthroughapopulation,itisrarelybecausetheyareliterallycopiedfromeach
other.Sperberarguesthatculturalreproductionisrarelymemelike,butinsteadmakesuseofwhathe
callsattractorsculturallysharedpatternsofthought,whichenablerepresentationstospreadthrougha
populationwithoutliteralcopying.Stickingwiththecakeexample,perhapsyoueatasliceofmy
Victoriasponge,youlikeit,andyoudecidetomakeoneforyourself.Perhapstherecipeyouuseisvery
similartomine.Butyouhavenotfiguredoutbytastingmycakewhichingredientsneededtogoinand
inwhatorder.Rather,youalreadyknewhowtomakeaVictoriasponge.Eatingmycakesimplytriggered
theuseofarecipethatwasalreadyinyourrepertoire.Inthiscase,therecipehasappearedinmyhead,
andbecauseofthisithasappearedinyours,butnotbecauseyourrecipeisacopyofmine.Sperber
arguesthatmemeticsismistakenbecausemostcasesofthespreadofideasarelikethis:

mostculturalitemsarereproducedinthesensethattheyareproducedagainandagain
with,ofcourse,acausallinkbetweenalltheseproductionsbutarenotreproducedinthe
senseofbeingcopiedfromoneanotherHencetheyarenotmemes,evenwhentheyare
closecopiesofoneanother(inaloosesenseofcopy,ofcourse).(Sperber2000,16465)

Bothoftheseconcernsraiseseriousproblemsforthegeneralityofmemetics:notallideasarereplicators,
hencenotallideasarememes.Whetherthisshowsthememeconcepttobeuselessdependsonwhether
thereareinsightstobehadbydistinguishingculturalinheritancethatismemelikefromcultural
inheritancethatisnot(Sterelny2006a).

Culturalunitsdonotformlineages:Acloselyrelatedcriticismofmemeticsdrawsonthefactthatwhile
ingeneticreplicationwecantraceanewcopyofagenebacktoasingleparent,ideasarerarelycopied
fromasinglesourceinawaythatallowsustotraceclearlineages(BoydandRicherson2000).
Memeticistsarefondofanalysingreligiousbeliefintermsofthespreadofmemes.Butwhilereligious
beliefsmaywellspreadthroughpopulationsofhumans,itseemsunlikelythatwewillalwaysbeableto
tracetokeninstancesoffaithbacktoonesource.Instead,individualsoftenacquirebeliefinGodthrough
exposuretoseveralbelieversintheirlocalcommunity.Inthesecircumstances,beliefinGodisnot
causedbyoneidentifiableearliertokenofthesametype.Withintherealmofbiologicalevolution,an
understandingofMendel'slawshasbeenimportantinexplainingsomeaspectsofevolutionarydynamics.
Mendel'slawsrelyonanunderstandingofgenesasdiscrete,transmittedunits.Butiftokenideascan
appearinanindividualinvirtueofthatindividual'sexposuretoseveralsources,thenthismakesit
unlikelythatanythingclosetoMendel'slawswillbediscoveredwithinculturalevolution.Thissuggests
apracticallimitationonenquirythatmayresultfromthisdifferencebetweenideasandgenes.Criticisms
ofthisformhavebeenputespeciallyforcefullybyWilliamWimsatt(1999),whoarguesthatthecreative
andinferentialabilitiesofhumanusersmakeitthecasethatanygivenidea,oritemoftechnology,can
havefluctuatingnumbersofculturalparentsovertime.Thisisbecausethecausalsourcesofits
reproductionmayvary.BeliefinGodmaysometimesbecausedbyexposuretoasinglecharismatic
evangelist,itmaysometimesbecausedbythejointinculcationoftwobiologicalparents,anditmay
sometimesbecausedbyimmersioninadiffusecommunityoftheists.Ideasanditemsoftechnologyalso
havenostableanaloguetothegenome,orgermline,becausedifferentelementswithincyclesof
technologicalreproduction,includingideas,behavioursofartisans,andmaterialelementsoftechnologies
themselves,canalltemporarilyacquirethestatusofreplicatorsdependingontheattentionthathuman
agentshappentobepayingtothem.Accidentalvariationsinone'smentalplanforconstructingapot,or
inone'sactionsinproducingthepot,orinthemadepotitself,canallconceivablybereproducedwhen
anotherartisancomestomakearesemblingitem.Wimsattusesthesedisanalogiestohighlightthe
formidableproblemsfacinganyefforttousepopulationgeneticmodelsintheexplanationofcultural
change.

Culturecannotbeatomisedintodiscreteunits:Ideasstandinlogicalrelationstoeachother.Whetheran
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individualisabletoacquiresomebelief,forexample,dependsontheirrelatedconceptualcompetencies.
Itisimpossibletobelieveinthetheoryofrelativitywithoutunderstandingit,andonecannotunderstand
itwithoutholdingmanyadditionalbeliefsrelatingtophysics.Thesameistruefornontechnicalbeliefs.
Dependingonwhichreligiononeistalkingabout,beliefinGodislikelytoberelatedtovariousother
beliefsconcerningforgiveness,retribution,loveandsoforth.Thishasledsomecriticstoarguethatitisa
mistaketotakeaviewofculturewhichatomisesitintodiscreteunits,assigningreplicativepowerto
themindividually.TheanthropologistAdamKupercomplainsthatUnlikegenes,culturaltraitsarenot
particulate.AnideaaboutGodcannotbeseparatedfromotherideaswithwhichitisindissolublylinked
inaparticularreligion(Kuper2000,180).Memeticistsarelikelytorespondbysayingthatalthough
ideasareinterlinked,thisdoesnotunderminethememegeneanalogy:O'Brienetal.(2010)haveargued
thatamorematureviewoftheroleofgenesinevolutionanddevelopmentreinstatesthememegene
parallel.Forthereisasenseinwhichgenes,too,needtobestudiedinacontextthattakesothergenes,
andtheirbroaderdevelopmentalandenvironmentalsettings,intoaccount.ADNAsequencecanhave
differenteffectsindifferentorganisms,dependingonthenetworkofrelationsitentersintowithother
geneticanddevelopmentalresources.JustasthesignificanceofbeliefinGodcanvarywithsocial
context,withtheresultthatitcanmakelittlesensetothinkofbeliefinGodasameme,sothefunction
ofsomeDNAsequencecanvarywithorganiccontext,withtheresultthatitmakeslittlesensetoidentify
somesequencetypeasageneforthepurposesofevolutionaryanalysis.

Thesecriticismsfocusonwhethertheretrulyarememes.Buttherearealsocriticismsoftheusefulness
ofthememeconcept,regardlessofwhethermemesexist.Aswasalreadyindicatedabove,onemight
worrythatmemeticsmerelyoffersacosmeticrepackagingofafamiliarsetofstoriesaboutcultural
change.Perhapsscienceiscomposedofreplicatingentitiesstrugglingagainstvariousselectionpressures,
butwhatinsightdoesthisofferus,ifintheenditpresentsuswithnothingmorethananalternativeidiom
inwhichtodescribethevariousfactorsthataffecttheevaluationofscientifichypotheses?Perhaps
clothesfashionsarememes,butevenifthatisthecase,onestillneedstoexplainwhatmakesone
clothingmemefitterthananother,andthefearisthatoncespelledoutthiswillquicklyboildowntoa
wellknownappealtoconsumerpsychology.

6.CulturalEvolutionwithoutMemes
Themostseriousandmostrespectedeffortstoapplyevolutionarythinkingtoculturebeginfroma
differentstartingpointtomemetics(althoughseeShennan2008,2011forsignificantworkthattakesthe
meme'seyeview).Memetheoriststendtobeginwithageneralcharacterisationofevolutionbynatural
selection,namelyasaprocessthatrequiresdifferentialcompetitionbetweenreplicators.Hencethememe
theoristlooksforsomestrictanaloguetothegeneintheculturalrealm,whichcanplaythereplicator
role.Dawkinsimpliesthatitisonlybecausehumansaresubjecttocolonisationbyreplicatorsotherthan
genesthathumanevolutionescapesthetyrannyofthegene.Onthisview,memesarerequiredinorderto
freeourspeciesfromaformofbiologicaldeterminism.

Thealternativetothisviewbeginswiththeobservationthatculturalinheritanceisimportant,anditseeks
tointegrateculturalinheritanceintotraditionalevolutionarymodels.Butthisgeneralmotivationleaves
opentheissueofwhetherculturalevolutionrequirestheexistenceofculturalreplicators.Clearlyonecan
acceptmanyofthecriticismsofthememeconcept,andstillattempttomodeltheeffectsofcultural
inheritance.Ratherthanseekingtoshowthatthereareculturalreplicators,onecaninsteadseektobuild
modelsthatallowforerrorpronelearning,andthatacknowledgethatanindividual'sbeliefsareoftenthe
resultofexposuretomanysources,ratherthancopyingfromjustonesource.Theinterestofcultural
evolutionarymodelsinthistraditionissometimessimplytoshowhowculturalchangeofvarioussorts
notnecessarilyadaptiveculturalchangecansubsequentlyaffectgeneticevolution,andviceversa.This
isthegeneralgoalofmodelsofgeneculturecoevolution.Butculturalevolutionarymodelsalsoaimto
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assesstheroleofculturalinheritanceintheconstructionofadaptation:here,culturalevolutionary
theoristsarenotmerelyseekingtoexplaindistributionsoftraitsinpopulations,theyareseekingto
explaintheappearanceofvaluableculturalnovelties(GodfreySmith2012).

Onemightthinkthatevenifculturalchangedoesnotrequireculturalreplicators,atleastadaptive
culturalchangedoes.ThegeneralDarwinianschemeforexplainingadaptationdemandsreliable
inheritanceitdemandsthatonceafitnessaugmentingmutationarises,itcanberetainedinfuture
generations.Ifculturallearningiserrorprone,orifindividualsacquireculturaltraitsbytakingan
averageofmanydifferentmodels,thenonemightthinkthatifsomeindividualisabletodiscovera
fitnessenhancingbehaviour,thattraitwillbelosttofuturegenerationseitherbecauseitismiscopied,or
becauseitiscombinedwithotherlessadaptivetraitstoproduceanaveragedmishmashofabehaviour.

Alloftheseinferenceshavebeenchallengedbyrecentculturalevolutionarytheory.Culturalevolutionists
agreethatatthelevelofthepopulation,cumulativeevolutionrequiresthatfitnessenhancingcultural
traitsarepreservedintheoffspringgeneration.However,theydenythatthisrequiresfaithful
transmissionbetweenindividuals.AformalmodelfromHenrichandBoyd(2002)showshowsocalled
conformistbiascanovercometheeffectsoferrorpronelearningtoproducereliableinheritanceatthe
populationlevel.Conformistbiasisthetendencyofindividualstoadoptwhattheybelieveisthemost
commonrepresentationinapopulation.HenrichandBoydciteevidencethatconformistbiasisareal
phenomenon.HenrichandBoyd'stheoreticalmodelassumesthatindividualsarepooratinferringthe
representationsofothers.Evenso,theyarguethatwhenwelooktothepopulationlevel,conformistbias
helpstocorrecttheeffectsofsucherrors,producingapopulationwidedistributionofrepresentationsin
theoffspringgenerationthatisclosetothepopulationwidedistributionofrepresentationsintheparent
generation.HenrichandBoydexplainthereasonforthis.Ingeneral,errorpronetransmissionhasa
tendencytoproduceamixtureofdifferentrepresentations.Inapopulationthatalreadycontainsseveral
differentrepresentationsatsignificantfrequencies,theeffectoferroronapopulationwidedistributionof
representationsisthereforelow.Inapopulationinwhichonerepresentationiscommon,theeffectsof
erroraremuchmoresignificant.Butifweaddconformistbias,weincreasethechancesofacommonly
heldrepresentationremainingcommonlyheldinfuturegenerations,evenwitherrorproneimitation.

BoydandHenrichacknowledgethatthisdoesnotmakepopulationleveldistributionsperfectlyreliably
inherited.Butthisdoesnotshowthatcumulativeevolutionactingonculturalinheritanceisimpossible.
Atthegeneticlevel,highlyfaithfulcopyingprocessesallowevenverysmallselectiveforcestopreserve
adaptivevariation.Lessfaithfulcopyingdemandsstrongerselectiveforcesifadaptivevariationisnotto
belost.BoydandHenrichareconfidentthatselectiveforcesintheculturalrealmarestrongerthan
selectiveforcesinthegeneticrealm.Themoral,onceagain,isthatitisimportantnottofocustooclosely
ongeneticevolutionasamodelforculturalevolution.

BoydandHenrichalsoarguethataculturalevolutionarytheorycanaccommodateSperber'sclaimthat
culturalreproductionrarelyworksthroughgenuinecopying.Evenifthereisasmallnumberof
attractorswaysofthinkingthatweareparticularlylikelytoadoptgivensomeexternalstimulusit
doesnotfollowthatevolutionarymodelshavenoroletoplay.Mostobviously,onecanstillarguethat
variousselectiveforcesaffectwhichofanumberofattractorscomestopredominateinapopulation.
Returningonceagaintothecakeexample,theremaybeattractorscorrespondingtoVictoriaSponges,
GingerCakesandBananaBread.Evenso,onecanseektounderstandwhyatagivenmomentintime
moreVictoriaSpongesarebeingproducedthanGingerCakes,andtheframeworkofcultural
evolutionarytheory,whichlookstothefactorsthatmakeindividualslikelytobeusedasmodelsfor
imitation,andthefactorsthatmakerepresentations(recipes,inthiscase)likelytobeemulatedoncea
modelispicked,canbeusedtoexplainthisdifferentialsuccesswithoutstrictcopying.

Memeticstendstobedrivenbyadesiretoseeculturalanalogiestogeneticevolution.Cultural
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evolutionarymodelsinthemannerofBoydandHenricharedriveninsteadbyadesiretofindwaysof
understandinghowculturalinheritanceaffectsevolutionaryprocesses.Thesesortsofcultural
evolutionarymodelsdonotassumethatculturalinheritanceworksinthesamewayasgenetic
inheritance.Indeed,theyarefreetomodelculturalinheritanceinwaysthatdepartquitemarkedlyfrom
geneticinheritance.Yettheyremainrecognisablyevolutionaryinstyle,primarilybecausetheyseekto
explainthechangesintraitfrequenciesinapopulationovertime.Theydothisbymakingbroad
assumptionsabouthowindividualsacquireculturaltraitsforexample,theymayassumethatan
individual'srepresentationsaretheproductoflearningfromavarietyofpeers,orthattheyarisefrom
attendingparticularlytoauthorityfiguresandbyassessinghowsuchruleswillplayoutatthe
populationlevel.Moreover,theserulesforculturalacquisitionarenotmerelyconjectured,theyaregiven
experimentalsupport.Culturalevolutionarytheoristsseektodocumenttheeffectsofvariousempirically
confirmedformsofbias,suchasconformistbiasandprestigebias.JustasDarwin'sowntheoryof
evolutionbynaturalselectionremainedlargelyconjecturaluntilsupplementedbyempiricalwork
showinghowinheritanceworked,andbystatisticalworkfocusingonthepopulationlevelconsequences
ofinheritance,selection,mutationandotherforces,soculturalevolutionarytheoryhasgaineditsinsights
fromasimilarcombinationofempiricalandmathematicalapproaches.

7.TheExplanatoryRoleofCulturalEvolutionaryTheories
Atthebeginningofthisentryitwasclaimedthatthecaseforculturalevolutionwasirresistible.Noone
candenythatculturalinheritanceisanimportantfactorinexplaininghowourspecieshaschangedover
time.Culturalinheritanceisnotmerelyaprocessthatactsinparalleltogeneticevolution,itis
intertwinedwithgeneticevolution.Culturalchangesbringaboutalterationstotheenvironment,whichin
turnaffectbothhowgenesactindevelopment,andwhatselectionpressuresactongenes.Inspiteofall
this,onemightstillworrythatitisamistaketounderstandtheimportanceofcultureusingthetoolsof
evolutionarytheory.Thisisbecauseonemaybescepticaloftheexistenceofatheorythatisbothgeneral
enoughtocoverallformsofculturalchange,andinformativeenoughtobeenlightening.

ThereisnodoubtthatitisoftenimportanttoremindoverlyenthusiasticorthodoxDarwiniansofthe
importanceofculture.Forexample,itseemsthattheincreasedincidenceoflactosetoleranceamong
humanpopulationshasarisenasaconsequenceofaculturalinnovationnamelydairyfarming.The
relativelyrecentappearanceofthisgeneticallycontrolledadaptationdemonstratesthathuman
physiologicalnatureissomethingthatcontinuestochange,anditalsodemonstratesthecausalimpactof
cultureongenes(RichersonandBoyd2005,19192).Suchexamplesbythemselvesshowtherashness
ofanyviewthatclaimseitherthathumannaturehasremainedfixedsincetheStoneAge,orthatgenes
aresomehowintheevolutionarydrivingseat.Yetnoneofthisshowsthatwecandevelopageneral,
informativetheoryofculturalevolution.Onemightfearthatintheendculturalchange,andtheinfluence
ofculturalchangeonotheraspectsofthehumanspecies,arebestunderstoodthroughaseriesof
individualnarratives.Ourbriefexaminationofmemeticsillustratedthisconcern.Wegainnoreal
explanatoryinsightifwearetoldthatideasspreadthroughpopulations,somemoresuccessfullythan
others.Wewanttoknowwhatmakessomeideasfitterthanothers.Anditisnotclearthattherewillbe
anygeneralrulesthatcanhelpustoanswerthisquestion.Inthebiologicalrealmweneeddetailed
accountsoflocalenvironmentalcircumstances,speciesspecificphysiologyandanatomy,andsoforth,to
telluswhatmakesoneorganicvariantfitterthananother.Similarly,intheculturalrealmwewillneedto
lookatlocalpsychologicaldispositionstoexplainwhysomeideasaremorelikelytospreadthanothers.
Soanyexplanatoryvaluetobehadfrommemeticsisparasiticonconventionalworkdoneinpsychology.
Andifindividualpreferencesaresubjecttochangeovertime,thentheremaybenogeneraland
informativetheoryofculturalevolutiontobehadrather,wewillhavetosettleforlocalexplanationsthat
looktoshiftingpreferences.Ratherthanprovideanewscientificframeworkforanunderstandingof
culture,memeticswilltendtodegenerateintoconventionalnarrativeculturalhistory.
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Therearethreebroadsetsofresponsestothislineofargument,eachofwhichpicksuponadifferent
explanatoryelementofmainstreamevolutionarytheory(Lewens2012).First,BoydandRichersonargue
thatinformativeinsightsariseoutofculturalevolution'spopulationthinking(RichersonandBoyd
2005).Second,Sterelny(2001,2003,2006a,2006b,2007,2012)andWimsatt(1999)arguethat
illuminatinginsightsregardingthegeneralconditionsrequiredforevolvabilityalsoapplyinthecultural
realm.Third,manyhavearguedthatculturalevolutionarytheoriescanprovideinsightsintothehistorical
patternofculturalchange,inmuchthesamewaythatevolutionarybiologicaltoolshaveenabledusto
reconstructthebranching(orperhapsreticulated)historyoflife.Wewilllookateachlineofdefencein
turn.Foramuchfulleraccountofthevalueofculturalevolutionarythinking,andindeedforathorough
andaccessibleintroductiontothetheoryasawhole,AlexMesoudi's(2011)bookisanexcellentplaceto
look.

8.PopulationThinking
Populationthinkingmeansmanythingstomanypeople.ForBoydandRichersonitdenotesanyeffort
toabstractfromacharacterisationofindividualpsychologicalprofiles,inawaythatallowsan
explorationoftheconsequencesoftheseindividualleveldispositionsforpopulationlevelproperties.We
havealreadyseenanexampleofthissortofpopulationthinkinginaction.Itisfarfromobviousthat
conformistbiasamongindividualscanenablepopulationlevelinheritanceinspiteofindividuallevel
errorsincopying.Toshowthatthesepropertiesofindividualpsychology(conformistbiasanderror
pronelearning)combinetoyieldpopulationlevelinheritancerequiressomeabstractmathematical
modelling.Andtheestablishmentofthispopulationlevelconsequenceisimportant,foritenablesthe
investigatortorevisetheconstraintsonemightnaivelythinkmustbearonculturalinheritanceif
cumulativeculturalevolutionistooccur.

Inausefularticle,ElliottSober(1991)suggeststhattheoriesofculturalevolutionmayhavelimitedvalue
fortheworkofsocialscientists,onthegroundsthatsocialscientistsareprimarilyinterestedinexplaining
whatmakesindividualslikelytoadoptoneidea,ratherthananother.Theywanttoknow,forexample,
whynineteenthcenturyItalianwomendecidedthattheywouldratherhavetwochildrenthanfive,not
whatthepopulationlevelconsequencesoftheirdecisionsmightbe.RichersonandBoydrespondby
sayingthatSober'sargumentassumes,erroneously,thatweareallgoodintuitivepopulationthinkers
(RichersonandBoyd2005,97).InSober'soriginalarticlehepointsoutthatpopulationthinkingmight
saveculturalevolutionarymodelsfromvacuityinjustthisway:

Sothequestionabouttheusefulnessofthesemodelsofculturalevolutiontothedaytoday
researchofsocialscientistscomestothis:Aresocialscientistsgoodatintuitivepopulation
thinking?Iftheyare,thentheirexplanationswillnotbeunderminedbyprecisemodelsof
culturalevolution.Iftheyarenot,thensocialscientistsshouldcorrecttheirexplanations(and
theintuitionsonwhichtheyrely)bystudyingthesemodels.(Sober1991,492)

ManyofRichersonandBoyd'smodelsareenlightening.Aswehaveseen,ittakesworktoshowthat
cumulativeculturaladaptationdoesnotrequirereplication.Note,however,infavourofSober's
scepticism,thatthemostinterestingculturalevolutionarymodelsareoftenthosewhichshowthegeneral
circumstancesunderwhichitispossibleforculturalinheritancetobeeffectiveinproducingadaptation.
BoydandRicherson'sclaiminfavouroftheimportanceofprestigebiasisprimarilyanefforttoshow
hownaturalselectionmighthavefavouredculturallearning.Sober'sconcerniswithwhethermodels
suchasthesewillalsoaffectthedaytodayresearchofsocialscientists,whoarenotsointerestedin
establishingsuchgeneralconditionsforcumulativeculturalevolution,butwhoareinsteadinterestedin
understandingparticularepisodesofsocialandculturalchange.Evenhere,RichersonandBoyd's
populationthinkingmayhavesomebite.Theyseek,forexample,toexplainthedisappearanceof
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importanttechnologiesonTasmania.DrawingontheworkofJosephHenrich,theysuggest(Richerson
andBoyd2005,138)thatthemaintenanceoftechnologiesandtheassociatedbehavioursrequiredto
produceandoperatethemmayrequireapopulationthatislargeenoughfortherateofinnovationto
offsetthedegradationthatresultsfromerrorproneimitation.IfBoydandRichersonarerightaboutthis
episodeinthehistoryofTasmania,thenwemaybeabletoexplainthedifferencesintheabilitiesofthe
Tasmanians,comparedwithotherpeoples,tomaintainasetoftechnologies,merelybycitingpopulation
size,ratherthanotherformsofsocialorculturaldifference.Note,finally,thatHenrich'smodel,likeany
populationalmodel,mustproceedbymakinghighlysimplifiedassumptionsaboutthepropertiesofthe
individualentitiesthatmakeupthelargerpopulation.Thisinvitesagenericseriesofcriticismsaimedat
anyeffortatabstractmodelling:Henrich'smodelhasbeencriticizedbythosewhodoubttherobustness
ofitsassumptionsanditsmatchwithempiricaldata(seeRead2006forsuchcriticism,KlineandBoyd
2010foraresponse,andHoukes2012forausefulphilosophicaloverview).Atthesametime,this
abstractionconstitutesapotentialstrengthofthepopulationalapproach,foritoffersusthepossibilityof
understandingacomplexsystemwithoutneedingcomprehensiveinformationaboutallofitsparts.

9.Evolvability
Asecond,related,waytovindicatemodelsofculturalevolutionlookstothequestionofthegeneral
featuresofinheritancesystemsthatmakeforevolvabilityinalineage.Thisprojecthasbeenpioneeredin
recentyearsbyKimSterelny(e.g.2001,2003,2006a,2006b,2007,2012).Onceagain,letusillustrate
thegeneralnatureoftheseissuesbybeginningintheorganicrealm.Darwin'stheoryisintendedto
explainadaptation.Thebasicconditionsfornaturalselectiondonot,inspiteofappearances,sufficefor
theappearanceoffunctionaltraits.Asysteminwhichoffspringresembleparentswithrespecttofitness
enhancingtraitsmaynotdevelopcomplexadaptations.Theenvironmentneedstocooperate:ifselective
pressureschangeveryquicklythentherewillbenosustainedenvironmentaldemandsofthesortthat
mightbuildcomplexadaptationsovertime.Developmentalsomatters.Ifontogenyissetupinsucha
waythatchangestoanyonetraittendtobeaccompaniedbychangestoallothertraits,thenthechances
arethatcumulativeadaptationwillbeparticularlyhardtocomeby.Foreveninthosecaseswherea
mutationcontributespositivelytothefunctionofonetrait,thechancesarethatitwillcontribute
negativelytooverallfitnessinvirtueofitsdisruptionofthefunctioningofothertraits.Developmentalso
needstomakeawiderangeofvariationavailable.Ifitishighlyconstrained,sothatonlyasmallnumber
offormsarepossible,thenselectionisnotpresentedwithabroadenoughrangeofrawmaterialsfrom
whichtofashioncomplextraits.Italsoseemsthatcumulativeadaptationreliesonthesuppressionof
outlaws(Sterelny2001,2006b).Groupselection,forexample,isoftenheldtobeanineffectiveagentof
groupleveladaptation,onthegroundsthatitisvulnerabletosubversionfromwithin.Thisoccurswhen
individualorganismsgoitalone,sabotagingcomplexfeaturesofgrouporganisationinfavouroftheir
ownfitness.Individuallevelselection,incontrast,canbuildindividualleveladaptations.Thisis
because,byandlarge,genesinagivenhumanorganismshareacommonfatetheydonotbehaveas
thoughtheywereindirectcompetition,strugglingforrepresentationinfuturegenerations.Whengenes
genuinelygoitalone,forexamplebysabotagingmeiosissothatsomehavegreaterchancesof
appearinginfuturegenerationsthanothers,thentheoverallintegrityoftheorganismcanbe
compromised,andindividualleveladaptationisundermined.

Byapplyingthesesortsofconsiderationstotheculturalrealmwecanattempttounderstandthelikely
costsandbenefitsassociatedwithvariousdifferentformsofculturalinheritance(vertical,oblique,
memelikeandsoforth).Wecanalsoperhapscometoanunderstandingofthedifferentevolutionary
forcesthatmightbringthesedifferentformsofculturalinheritanceintoexistence.And,inturn,these
insightsmayfacilitatecomparativeworkthatseekstodocumentthegeneralconditionsthatarerequired
foraspeciestomakeuseofculturalinheritanceinordertobuildcomplexadaptationssuchastools.This
wayofthinkingoffersthepromise,forexample,ofexplainingwhyfew,ifany,nonhumanspeciesare
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abletobuildprogressivelymoreandmorecomplexculturalfeaturesinacumulativemanner(Richerson
andBoyd2005,107).Theexplorationofthesignificanceoftheseconditionsintheculturalrealmis
contentious,partlybecausetheconditionsforevolvabilitythemselvesaredisputed(seeGodfreySmith
2009).BoydandHenrich'sworkbringsoutthefactthatalthoughpopulationlevelinheritanceis
importantforadaptation,parentoffspringresemblanceisnot,infact,necessary.Questionsrelatingto
evolvabilityarealsotiedupwithdifficultissuesrelatingtotheunitsofselectiondebate(Okasha2006).
Aswehaveseen,naturalselectionatahigherleveloforganisationmayberequiredtogenerate
mechanismsthatsuppresstheabilityofdisruptiveoutlawstogoitaloneatlowerlevelsoforganisation.
Doessomethinglikethisoccurintheculturalrealm?Doesselectiononhumangroupsactsoastolimit
theabilityofindividualhumanstogoitalone?Inwhatwaysmightculturalinheritancebeinvolvedin
theseprocesses?BoydandRicherson(2009),forexample,havearguedthatculturalinheritancepromotes
theexistenceofbetweengroupdifferences,andtherebyfacilitiesgroupselection.Thesequestionsare
complex,bothintermsofhowtheyshouldbeposedandhowtheyshouldbeanswered.Butsomeofthe
mostinterestingworkinculturalevolutionarytheorymaycomefromeffortstoanswerthem.

Issuesrelatingtoevolvabilityaresometimesframedintermsofsystemsofinformationtransfer.Onthis
view,ifoffspringaretoresembleparents,developmentalinformationmustbetransmittedfromone
generationtothenext.Thequestioniswhatformsofinformationtransmissionsystemdothisjob.This
modeofframingtheissueiscontentious,foritisnotalwaysclearhowwearetounderstandtheconcept
ofinformation,andwhatitmeansforsomecausalcontributortodevelopmenttocountasaninformation
bearer,ratherthansomeotherkindofdevelopmentalparticipant,suchasaninformationreader,say,ora
backgroundconditionforinformationtransfer(seeOyama2000andGriffiths2001fordiscussionof
theseissues).Thisgeneralwayofthinkingaboutinheritancehas,however,beeninfluentialin
characterisingsocalledmajorevolutionarytransitions(MaynardSmithandSzathmary1995).Key
transitionsinthehistoryoflifearesaidtoincludethedevelopmentofDNAbasedinheritance,the
emergenceofchromosomes,theadventofthegeneticcode,andeventssuchasthearrivalofsociality
andlanguage.MaynardSmithandSzathmaryproposethatwecanthinkoftheseeventsasmodifications
tothemechanismsofintergenerationalinformationtransmission.

JablonkaandLamb(2005)arguethatthinkingintermsofinformationtransmissionsystemsalsoallows
ustopointoutsalientdifferencesintheformsofsocialtransmissionunderlyingculturalevolution.They
claimthatonlysomeformsofsocialtransmissionmakeuseofasystemofsymbols.Consider,for
example,thattosaythatsomebirdsinherittheirsongbysocialtransmissionisnottosaythatbirdsongis
asymbolicsystem.Humans,ontheotherhand,tradeinpubliclyaccessiblesymbols.Moreover,
repositoriesofsymbols,mostobviouslyintheformoflibrariesandcomputerdatabases,arevital
inheritancesystemsforhumans,allowingthepreservationandaccumulationofknowledgeacross
generations.Note,also,thattherearedifferenttypesofsymbolsystem.Insomecasestherelationship
betweenasymbolandwhatisrepresentedisarbitrary.Theisthecaseforawordlikeman,whichdoes
notlookorsoundlikeahumanmale.Inothercasesoficonicsymbolism,therelationshipisoneof
resemblance:asignforthegents'toiletlookslikeaman.

JablonkaandLambusethecharacteristicdifferencesbetweentypicalmodesofsocialinheritancein
animalsandhumanstoilluminatetheimpactourownsymbolictransmissionsystemshaveonhuman
culturalevolution(seealsoDeacon1997).Althoughtheyarguethattherecanbenonlinguisticsymbolic
systems(2005,224),languageexemplifiesnicelythewayinwhichsystemsofsymbolscontainelements
thatcanberecombinedincountlesswaystoyieldavastarrayofdifferentmeaningfulmessages.
Repositoriesofsymbolicallystoredinformation,suchasbooks,canalsobesearched,annotated,edited
andsoforth,inwaysthataddtotheirpowerandversatility.Thismannerofthinkingopensupanumber
ofchallengingissues.Thequestionofthedegreetowhichsymbolicsystemsresembleotherinheritance
systemsisanilluminatingone.Consider,bywayofexample,Stegmann's(2004)discussionofthesense
inwhichthegeneticcodeisarbitrary.Onequicklyrealisesthatanyattempttosaypreciselywhat
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makessomeinheritancesystemasymbolicsystem,andanyattempttodifferentiatebetweentypesof
symbolicsystems(linguistic,nonlinguisticandsoforth),willbeexceptionallyphilosophically
demanding.

10.CulturalPhylogenies
Manyevolutionistshavearguedthatbiologicaltoolscanhavegreatvaluewhenwewishtodevelopa
historicalviewofthepatternofculturalchange(see,forexample,Grayatal2007,MaceandHolden
2005:thissectionitselfdrawsinLewens2012).Avarietyofbiologicalmethodshavebeendeveloped
thathelpustouncoverthestructureofevolutionarytrees:theyhelpustounderstandwhichtaxasplit
fromwhichothersandwhen.Itseemsclearthatculturalitemsofmanykinds(mostobviouslylanguages,
butalsotoolsandtechniques)alsostandinrecognizablegenealogicalrelationships,andthishasled
manybiologicalanthropologiststousephylogeneticmethodsborrowedoradaptedfromthebiological
sciencesinordertoreconstructthehistoryofborrowingsintheculturalrealm.Criticshavesometimes
followedGould(1988)inarguingthatthesebiologicalmethodscannotbeproperlyappliedtothecultural
realm,becauseculturalgenealogiestaketheformofreticulatednetworks,ratherthanbranchingtrees.
Culturalchangeisindeedoftenhighlyreticulated:itisobviousthatacomplexobjectlikeacarisa
confluenceofnumeroustechnicallineages,whichcometogethertoformthehifisystem,theengine,the
safetydevices,andsoforth.Moreover,asimprovementsaremadetocarsthesenewdevelopmentsmay
beborrowedbyinnovatorsofbicycles,furniture,toysandothershiftingconstellationsofartifacts.

Theseimportantobservationsneednotunderminetheprojectofculturalphylogeny.Muchofbiological
evolutionisalsoreticulated.Bacteria,forexample,donotformgenealogicallyisolatedlineages,
hybridizationisrifeamongplants,andthereisalsoconsiderableborrowingofelementsofthegenome
betweenapparentlyisolatedmammalianspecies.Ofcoursethismightshowsimplythatphylogenetic
modesofinferencearedoublyimperilled:theydon'tworkformuchofthebiologicalworldeither.But
culturalevolutionists(e.g.Grayetal2007)areencouragedbyinferentialdevelopmentswithinbiology
itself,whichaimtoreconstructpartiallyreticulatedtreesbyproposingsocalledreconciliationsofthe
conflictingtreesthattraditionalmethodsoftenproposeforspeciesandgenes.

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TheauthorwouldliketothankBethHannonforhelpinpreparingrevisionstothisentry,andthe
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ChristophervonBlowforcarefullyreadingtheentryandidentifyingagoodnumberoftypographical
errorsforcorrection.

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