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TheNatureofIntelligence:NovartisFoundationSymposium233.Volume233 EditedbyGregoryR.Bock,JamieA.GoodeandKateWebb CopyrightNovartisFoundation2000.ISBN:0-471-49434-8 THE NATURE OF INTELLIGENCE TheNovartisFoundationisaninternationalscienti¢candeducational charity(UKRegisteredCharityNo.313574).KnownuntilSeptember1997 astheCibaFoundation,itwasestablishedin1947bytheCIBAcompany ofBasle,whichmergedwithSandozin1996,toformNovartis.The FoundationoperatesindependentlyinLondonunderEnglishtrust law.Itwasformallyopenedon22June1949. TheFoundationpromotesthestudyandgeneralknowledgeof scienceandinparticularencouragesinternationalco-operationin scienti¢cresearch.Tothisend,itorganizesinternationally acclaimedmeetings(typicallyeightsymposiaandalliedopen meetingsand15^20discussionmeetings)andpublisheseightbooks peryearfeaturingthepresentedpapersanddiscussionsfromthesymposia. Althoughprimarilyanoperationalratherthanagrant-makingfoundation, itawardsbursariestoyoungscientiststoattendthesymposiaand afterwardsworkwithoneoftheotherparticipants. TheFoundation’sheadquartersat41PortlandPlace,LondonW1B1BN, providelibraryfacilities,opentograduatesinscienceandallieddisciplines. Mediarelationsarefosteredbyregularpressconferencesandbyarticles preparedbytheFoundation’sScienceWriterinResidence.TheFoundation o¡ersaccommodationandmeetingfacilitiestovisitingscientistsand theirsocieties. InformationonallFoundationactivitiescanbefoundat http://www.novartisfound.org.uk Novartis Foundation Symposium 233 TTHHEE NNAATTUURREE OOFF IINNTTEELLLLIIGGEENNCCEE 2000 JOHN WILEY & SONS, LTD Chichester · New York · Weinheim · Brisbane · Singapore · Toronto Copyright&NovartisFoundation2000 Publishedin2000byJohnWiley&SonsLtd, Ba⁄nsLane,Chichester, WestSussexPO191UD,England National 01243779777 International(+44)1243779777 e-mail(forordersandcustomerserviceenquiries):[email protected] VisitourHomePageonhttp://www.wiley.co.uk orhttp://www.wiley.com AllRightsReserved.Nopartofthisbookmaybereproduced,storedinaretrieval system,ortransmitted,inanyformorbyanymeans,electronic,mechanical,photocopying, recording,scanningorotherwise,exceptunderthetermsoftheCopyright,Designsand PatentsAct1988orunderthetermsofalicenceissuedbytheCopyrightLicensingAgency, 90TottenhamCourtRoad,London,W1P9HE,UK,withoutthepermissioninwriting ofthepublisher. OtherWileyEditorialO⁄ces JohnWiley&Sons,Inc.,605ThirdAvenue, NewYork,NY10158-0012,USA WILEY-VCHVerlagGmbH,Pappelallee3, D-69469Weinheim,Germany JacarandaWileyLtd,33ParkRoad,Milton, Queensland4064,Australia JohnWiley&Sons(Asia)PteLtd,2ClementiLoop#02-01, JinXingDistripark,Singapore129809 JohnWiley&Sons(Canada)Ltd,22WorcesterRoad, Rexdale,OntarioM9W1L1,Canada NovartisFoundationSymposium233 viii+300pages,24¢gures,24tables BritishLibraryCataloguinginPublicationData AcataloguerecordforthisbookisavailablefromtheBritishLibrary ISBN0471494348 Typesetin101(cid:1)2 on121(cid:1)2 ptGaramondbyDobbieTypesettingLimited,Tavistock,Devon. PrintedandboundinGreatBritainbyBiddlesLtd,GuildfordandKing’sLynn. Thisbookisprintedonacid-freepaperresponsiblymanufacturedfromsustainableforestry, inwhichatleasttwotreesareplantedforeachoneusedforpaperproduction. Contents SymposiumonThenatureofintelligence,heldattheNovartisFoundation,London,30Nov^2Dec 1999 Editors:GregoryR.Bock(Organizer),JamieA.GoodeandKateWebb ThissymposiumisbasedonaproposalmadebyGeo¡reyMiller MichaelRutter Introduction 1 DavidLubinski Intelligence:successand¢tness 6 Discussion 27 ArthurR.Jensen Thegfactor:psychometricsandbiology 37 Discussion 47 IanJ.Deary Psychometricintelligencedi¡erencesandbrainfunction 58 Discussion 73 BrittAnderson Thegfactorinnon-humananimals 79 Discussion 90 RandolphM.Nesse Naturalselection,mentalmodulesandintelligence 96 Discussion 105 GeneraldiscussionI 116 NathanBrody gandtheone^manyproblem:isoneenough? 122 Discussion 129 DouglasK.Detterman Generalintelligenceandthede¢nitionof phenotypes 136 Discussion 144 DavidHoule Isthereagfactorfor¢tness? 149 Discussion 159 J.MichaelBailey Howcanpsychologicaladaptationsbeheritable? 171 Discussion 180 v vi CONTENTS AndrewWhiten Socialcomplexityandsocialintelligence 185 Discussion 196 JamesR.Flynn IQgains,WISCsubtestsand£uidg:gtheoryandtherelevanceof Spearman’shypothesistorace 202 Discussion 216 AndrewPomiankowski Mutation,selectionandtheheritabilityofcomplex traits 228 Discussion 237 PeterMcGu⁄n Thequantitativeandmoleculargeneticsofhuman intelligence 243 Discussion 255 Geo¡reyMiller Sexualselectionforindicatorsofintelligence 260 Discussion 270 Finalgeneraldiscussion 276 MichaelRutter Closingremarks 281 Indexofcontributors 288 Subjectindex 290 Participants B.Anderson Neurology(127),BirminghamVAMedicalCenter,700S19th Street,Birmingham,AL35233,USA J.M.Bailey DepartmentofPsychology,NorthwesternUniversity,2029 SheridanRoad,Evanston,IL60208-2710,USA N.Brody DepartmentofPsychology,WesleyanUniversity,Middletown, CT06459,USA I.J.Deary DepartmentofPsychology,UniversityofEdinburgh,7George Square,EdinburghEH89JZ,UK D.K.Detterman DepartmentofPsychology,CaseWesternReserveUniversity, 10900EuclidAvenue,Cleveland,OH44106-7123,USA R.Dunbar SchoolofBiologicalSciences,NicholsonBuilding,Universityof Liverpool,LiverpoolL693BX,UK J.R.Flynn DepartmentofPoliticalStudies,UniversityofOtago,POBox56, Dunedin,NewZealand S.Gangestad DepartmentofPsychology,UniversityofNewMexico, Albuquerque,NM87131,USA S.Harnad DepartmentofElectronicsandComputerScience,Universityof Southampton,High¢eld,SouthamptonSO171BJ,UK R.Hinde StJohn’sCollege,CambridgeCB21TP,UK D.Houle DepartmentofBiologicalScience,FloridaStateUniversity, Tallehassee,FL32306-1100,USA vii viii PARTICIPANTS N.Humphrey CentreforPhilosophyofNaturalandSocialScience(CPNSS), TymesCourtBuilding,LondonSchoolofEconomics,HoughtonStreet, LondonWC2A2AE,UK A.R.Jensen GraduateSchoolofEducation,UniversityofCalifornia,Berkeley, CA94720-1670,USA A.Karmilo¡-Smith NeurocognitiveDevelopmentUnit,InstituteofChild Health,30GuilfordStreet,LondonWC1N1EH,UK D.Lubinski DepartmentofPsychologyandHumanDevelopment,Vanderbilt University,Nashville,TN37203,USA N.J.Mackintosh DepartmentofExperimentalPsychology,Universityof Cambridge,DowningStreet,CambridgeCB23EB,UK J.MaynardSmith SchoolofBiologicalSciences,BiologyBuilding,University ofSussex,Falmer,BrightonBN19QG,UK P.McGu⁄n Social,GeneticandDevelopmentalPsychiatryResearchCentre, InstituteofPsychiatry,King’sCollegeLondon,DeCrespignyPark,Denmark Hill,LondonSE58AF,UK G.Miller CentreforEconomicLearningandSocialEvolution,University CollegeLondon,GowerStreet,LondonWC1E6BT,UK R.M.Nesse DepartmentofPsychiatryandInstituteforSocialResearch,The UniversityofMichigan,AnnArbor,MI48106-1248,USA A.Pomiankowski TheGaltonLaboratory,DepartmentofBiology,University CollegeLondon,LondonNW12HE,UK M.Rutter(Chair) InstituteofPsychiatry,DeCrespignyPark,DenmarkHill, LondonSE58AF,UK T.Suddendorf(Bursar) SchoolofPsychology,UniversityofQueensland, Brisbane,QLD4072,Australia A.Whiten SchoolofPsychology,UniversityofStAndrews,StAndrews,Fife KY169AJ,UK TheNatureofIntelligence:NovartisFoundationSymposium233.Volume233 EditedbyGregoryR.Bock,JamieA.GoodeandKateWebb CopyrightNovartisFoundation2000.ISBN:0-471-49434-8 Introduction MichaelRutter Social, Genetic and Developmental Psychiatry Research Centre, Institute of Psychiatry, DeCrespignyPark,DenmarkHill,LondonSE58AF,UK The main rationale put forwardfor this meeting was that there was an apparent clash between the expectations of evolutionary psychologists and the claims of behaviour geneticists with respect to genetic in£uences on variations in intelligence.Evolutionarypsychologists,itwassaid,wouldexpectthatatraitso strongly adaptive as general intelligence would have been intensely shaped by naturalselectionandthat,asaresult,intelligenceshouldshowonlyweakgenetic e¡ectsonindividualvariationswithinthepopulation(Tooby&Cosmides1990). By contrast, behaviour geneticists have argued that empirical research ¢ndings have been consistent in showing a high heritability for intelligence. The supposed quandary was how to explain why the high intelligence that distinguishes us from other species, and which must have been under strong selection, nevertheless still shows large individual di¡erences that are subject to stronggeneticin£uences. Intrying,overthecourseofthenextfewdays,tocomeupwithsomekindof resolution of this apparent paradox, we will need to consider carefully the assumptions that underlie this proposition. The starting point, I suppose, is the theoretical expectation that traits under strong selection tend to have low heritabilityandthatevolutionbynaturalselectiondestroysthegeneticvariation on which it feeds (Dawkins 1999, citing Falconer 1960 and Lewontin 1979). Probably, most of us willbe inclined to acceptthis asa reasonable presumption from an understanding of how evolution operates, but questions may be raised on the extent to which the postulate is actually supported by empirical research ¢ndings. Which multifactorial traits can be contrasted with respect to the likelihood that they were or were not subject to strong selection? Is it the case thattheheritabilitiesoftheformerareconsistentlylowerthanthoseofthelatter? Is intelligence truly a rather unusual exception to a well-validated, empirically supported,generalrule,oristherealackofresearchdataontheissue? Thesecondassumptionisthatintelligencehasbeensubjecttostrongselection. On the face of it, it would seem obvious that this must have been the case. In support, Pinker (1997) pointed to the tripling in brain size during the time periodbetweensome4millionyearsagoandsome100000yearsago,andtothe 1 2 RUTTER factthatthecognitiveskillsofhumansfarexceedthoseofotherspecies.Clearly, there are huge interspecies di¡erences and it may be accepted that it is virtually certain that there have been major increases in cognitive capacity over time. Whatisnotquitesoself-evidentisthatitwillhavetohavetakenplaceasaresult ofevolutionaryadaptationofasinglecognitivetrait(cid:1)generalintelligence. Thornhill (1997) argued that a Darwinian adaptation was a feature of an organism that had been functionally designed through the process of evolution operatingbymeansof naturalselectionin thepast. Notethat theproposition is thatthetraitwasadaptiveinthepastduringtheperiodwhenitevolved;itdoes not necessarily have to be adaptive currently. Nevertheless, we may need to consider the extent to which intelligence is adaptive today. It has often been assumed that it is obvious that it must be so on the grounds that measured IQ correlates so consistently with measures of social success such as occupation, earnings, and parenting and crime. Jensen (1969) presented data on occupation 30 years ago, and Herrnstein & Murray (1994) outlined the ¢ndings more recently on a broader range of variables. There can be no doubt that IQ does indeed correlate with most measures of social success but, apart from scholastic achievement,thecorrelationsaremostlymoderateratherthanstrong(seeCawley et al 1997, on income, and Manolakes 1997, Rutter et al 1998, on crime). It is clear that IQ is only one of many factors, both in the individual and in society, that predisposeto social success.The ¢ndings on scholastic achievement do not help much because IQ tests were designed to predict it. We may conclude that theclaimsontheoverridingimportanceofIQhavebeenoverstatedbut,equally, itdoeshaveconsiderable predictivevalue. These ¢ndings, however, are of very limited relevance to the question of the adaptivevalueofIQduringthecourseofpastevolution.Pinker(1997)posedthe questionintermsofwhyitwashumans(ratherthanotheranimals)whodeveloped high intelligence in the way that they did. He argued that vision, group living, precision-grasp hands and hunting all played a part. All of these, like language (Pinker 1994), represent characteristics that are universally present, with individualdi¡erencesoflessimportance.Iftheseconstitutethekeyadaptations, it may be that the current individual di¡erences in intelligence are less of a paradox.Also,weneedtonotethatthekeyfeatureswithrespecttoevolutionary adaptationconcernreproductive¢tnessandnotsocialsuccess. That brings us to the next assumption, namely that intelligence constitutes a singletraitthatfunctionsinaunitaryfashioninadaptation.Wewillhearpapers thatargue forcefullyforthevalidityoftheconceptof‘g’(cid:1)generalintelligence. There can be no doubt that there are very substantial intercorrelations among cognitive tests and that, as an abstraction, g has a greater predictive power than more speci¢c cognitive tests (Carroll 1997). Whether that implies a biological unity, rather than a psychometric construct, is perhaps less certain. Also, to an

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