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The Origin of Politics: An Evolutionary Theory of Political Behavior

Author(s): John R. Alford and John R. HibbingReviewed work(s):Source: Perspectives on Politics, Vol. 2, No. 4 (Dec., 2004), pp. 707-723Published by: American Political Science AssociationStable URL: http://www.jstor.org/stable/3688539 .

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Articles

T h e O r i g i n o f Politics: A n EvolutionaryTheory o f Political Behav i o rJohn R. Alford and John R. Hibbing

In this articlewe propose hatevolutionary iologycansupplypoliticalsciencewith a theoryof the ultimatecausesof humanpreferencesnd behaviors hat t otherwise acks.For he mostpart,political cientists reeitherunfamiliarwiththe socialsideofevolutionaryheoryormisidentifytskey eatures. ar rombeinggenetically eterministic rleading xclusivelyopredictionshatallhumanbehaviorwillbeselfish,modernevolutionaryheories tress hatadaptive ehaviorsfrequentlyharacterizedyaguardedsortof cooperation.Wedescribemodernbiological heory, fferour own versionof it, discussnewandpotentially sefulinterpre-tationsof politicalattitudesandpublicpolicies,andpresent cientific vidence,drawn rom research n autistic ndividuals ndmonozygotic nddizygoticwins,of thestartlinglymportant olegenetics laysnshaping oliticallyelevant ttitudes ndbehaviors.

W hy do people hold the politicalattitudesthey do?Despite the centralityof this question, the twoguiding theoreticalorientationsof modernpoliti-cal science, rational choice and behavioralism,aresurpris-ingly unhelpful.Rational hoice scontent to takepreferencesas given and is not particularlymotivatedto exploretheirorigins or groundingin reality.Behavioralisms basedonthe untested assertionthat preferencescan be understoodby an exclusivefocus on environmentalvariables;t largelyignoresthe fundamental ssue of why people respondto theenvironment as they do. It is quite common (andaccurate)for adherentsof both positions to admit that their favoredapproachesarenot theoriesat all.Our goal is not to criticizerationalchoiceand behavior-alism.Indeed,politicalscientistsworkingwithin both theo-reticalframeworkshave producedimpressive indingsandinsights,therebyrevealing he valuein identifyinginfluen-tial environmental actorsand in outlining the actionsthatarational ndividualholdingagiven preferenceakes.None-theless,the fact remainsthat, as earlyproponentsof ratio-nal choice correctlynoted, behavioralisms limited by itslack of a real theory to being a largely nductive mode ofresearch.Rational choice, in turn, as many of its propo-nents have come to realize, s reachingthe limits of what

JohnR.Alford s associateprofessorfpoliticalscienceat RiceUniversityjra@rice.edu).ohnR. Hibbingis FoundationRegentsUniversity rofessorfPoliticalScience t the Univer-sityofNebraska-Lincoln(jhibbing@unl.edu). heauthorsaregratefulto ChrisLarimer,LeventeLittvay,David Rap-kin, KevinSmith,Jeff Spinner-Halev, lizabethTheiss-Morse,and anonymous eviewersfor theirhelpffl commentsand suggestions.

canbeaccomplished ydeductiveresearch asedon assumedratherthan actual preferences.To fully understand,con-nect, and transcend he extantresearchrequiresan empiri-callysustainable heoryof the source of preferences nd ofthe reasonspeople respond as they do to environmentalstimuli.Sucha theoryexistsand is being employedwith increas-ing regularityn the socialsciences,especiallyexperimentaleconomics,behavioralanthropology, nd socialpsychology.Certainly,some political scientists are familiarwith thistheory,but it hasnot been the basisfor originalresearch npolitical science.Yet theoriesof the origins of preferences(and thereforebehaviors)may offerthe abilityto unite theindividualsocialsciences-and even the socialsciencesasa

group-with the naturalsciences.Politicalscientistsmayfind the theorywe describewant-ing. But it should not be dismissedwithout due consider-ation of the evidencefor and againstit. Most important,the theoryshould not be dismissedbecause of an unscien-tific aversion o its implications.The centraltenet of this theory is that preferencesandbehaviorsareat leastpartially hapedbyevolutionary orcesand thereforeby geneticheritage.Justasevolutionarypres-suresshapedgenesgoverning he physical raitsof humans,they alsoshapedgenesgoverningbehavioral raits.If this istrue, research n the social sciences will be furtheredbytakinginto account evolutionarilyadvantageousbehaviorsandbyincreased ttention o theconnectionbetweenhumanbiology and social behavior.This evolutionary-biologicalapproach ncorporatesvaluableinsights from both behav-ioralismand rationalchoice. In fact, the most substantialcurrentapplicationsof this theoryare seen in the two dis-ciplines, psychologyand economics, from which politicalscientists borrow most heavily-and from which we

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Articles I The Origin f Politicsborrowedbehavioralism nd rationalchoice, respectively.fa single theoreticalapproachcan make headwayin disci-plines as disparateas economics and social psychology,itsurely merits some attention in a discipline like ours,which hasinternalized he conflict betweenpsychologyandeconomics.Inwhatfollows,we provideageneraldescriptionof mod-ern evolutionary heoryand our own more sociopoliticallyfocused behavioraltheory, which we call "warycoopera-tion."We then drawout the directimplicationsof thistheoryfor interpretationsof politics and public policy.Finallywedirectourattentionto evidencesupporting he assertion hatis most likely to trouble political scientists:that politicalorientationsare o asignificant xtentgeneticallynfluenced.

An EvolutionaryTheoryof Political BehaviorWhile "survival f the fittest" s often automaticallyassoci-ated with Hobbesian self-interest,the Darwinianaccountof humanbehavior s not nearlyasdepressingas is typicallyimagined.As is often the case,adiscipletook a much harderline, which has coloredpopular nterpretations f the orig-inal message.In CharlesDarwin'scase,Thomas H. Huxleywasthe individuallargelyresponsible or creatinga storyofsocial evolution characterized y unceasingand often vio-lent self-interest.1Darwinhimselfrecognized he incrediblevarietyof adaptivebehaviorsand the value

and prevalenceof behaviordesigned,at least in the near term, to benefitother organisms.2To the extentthatevolutionary heoryhas influencedthethinkingof politicalscientists,it has been primarily n theequationof natural electionwithnarrow elf-interest.Devi-ationsfrom self-servingbehavioraretypicallyexplainedbyhumans'limited cognitive abilities,3or by humans'social-ization and ultimate internalizationof societal norms. AsLeda Cosmides and John Tooby put it, most economistsand social scientistsassume that "rationalbehavioris thestateof nature,requiringno explanation."4This sentimentis evident in the work of political scientists.For example,Elinor Ostrom assertsthat "ourevolutionaryheritagehashardwiredus to be boundedlyself-seekingat the sametimethat we arecapableof learningheuristicsand norms, suchas reciprocity."5From this perspective,self-servingbehav-ior is geneticallydrivenand can only be counteredby thedecidedlynongenetic teachingof societalniceties.In equating naturalselection with inherent selfishness,political scientistsreflectviews held by mainstreambiolo-gists from Huxley'stime through about a quartercenturyago.The overwhelming ocusof evolutionary heorists henwas largelycompatiblewith what came to be the rationalchoiceview of theappropriatetarting ssumptionor humanbehavior.In a well-known work, biologist RichardDaw-kins observed:"If we weretold thata man lived a long andprosperousife in theworldof Chicago gangsters,we would

be entitled to make some guessesas to the sort of man hewas.... Like successfulChicago gangsters,our genes havesurvived,in some cases for millions of years, in a highlycompetitiveworld .... Ifyou look at the waynaturalselec-tion works, t seemsto follow thatanythingthat has evolvedby naturalselection should be selfish."6In thisview,humanbeingsaremerely"survivalmachines,"constructedby the genes to help the genes continue intothe next generation.The interactionsof survivalmachinesarehardlyexpectedto be warmandfuzzy.Dawkinscontin-ues: "To a survivalmachine, another survivalmachine ispartof its environment, ike a rock or a riveror a lump offood. It is somethingthatgets in the wayor somethingthatcan be exploited."7One need not searchlong to find par-allel viewsin socialscience.In one of the foundingworks ofrationalchoice, JamesM. Buchanan and Gordon Tullockdeclaredthat when two people interact,each will alwaysseek "morerather hanless"and will "exclude rom consid-eration" he interestsof their "oppositenumber."8Evolutionary heoryhasundergone undamentalchangesin recentdecades,particularly s it relates o socialbehavior.Renewed enthusiasm for an earlier,but largely ignored,insight-that of multilevel selection-offers an ingeniousexplanationortheemergence f cooperationroma nuancedpatternof self-interest.Selectionpressuretselfis, afterall,alaw of nature,not an evolvedmechanism:overtime, thingsthat last longerand/or reproduce hemselvesmore success-fullywill come to dominatenumerically.This is not limitedto living things, and genetic evolution is only one mecha-nism that expresses his natural aw. Traditionalevolution-ary heoryhas focusedlargely n the roleof selectionpressureat the level of the gene, and this is what most politicalscientists think of when the topic of evolution comes up.9Multilevelselection begins by recognizingthe ubiquityof selectionpressure.Focusingat anysinglelevelwill high-light selection pressureand the associatednarrowlyself-interested modes of activity at that level. For example,focusingat the level of the gene highlightsthe fact that thebehaviorof human genes is ultimately a consequence ofself-interestat thatlevel. This hasled scholars uchasDaw-kins to think of organismsas nothing morethan machinesbuilt and utilizedby selfishgenes,geneswhich "care" oth-ing for the ultimatewelfareof the organism tself.'l

Takinga step back alters this picturesubstantially.Thegenesthemselvesare,afterall, merelysurvivalmachinesforthe complexproteinsthat makeup geneticmaterial.At thisdeeperlevel, it is the complexproteinsthat areselfish,andtheir survivalmachines-the genes-may behave in waysthat seemhighlyinconsistentwith selfishness.1 In termsofhumanbehavior, f we thinkof groupsas survivalmachinesfor collectionsof individuals,then selectionpressures hatlead individualsto behaveselfishly may well be in conflictwith selection pressures hat favor groups of individualsthat behave n concert.Wheneverworkingin groupsallowsfor advantagesstemming from specialization,division of

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labor,or even mere numerousness tself, groupscharacter-izedby cooperativebehaviorat the individual evel have thepotentialto out-competeand hence dominategroupschar-acterizedby selfish behavior at the individual level. Thusindividuals aresubjectto conflictingselectionpressure.Atthe individual level, pressure avorsentirelyself-regardingbehavior,but self-regardingndividualsmay have a betterchanceof survivalf theyaremembersof cooperative roups.The parameters hat characterize he equilibriumof theseconflictingpressureswill often yield behavior hat is selfish;however,under some realisticconditions the resultwill bethe evolution of mechanismsfor cooperation.The cooper-ation evidentamongproteins n creating ivingcells,amongcells in creatingtissues,among tissues in creating organs,andamongorgans n creatingcomplexlivingstructures ikeplantsandanimals,areall examplesof this balanceof com-petitionandcooperationwithin the evolutionaryprocess.12The important insightfrom continuing this Rus-sian doll analogyof compe-tition within cooperationto the level of families,ex-tended kin groups, clans,political parties,and nationstates-all of which arepotentiallysubjectto selec-tion pressureat the indi-vidual and collective levels.Environmental conditionsthat offersubstantial dvan-tages to clans that cooper-ateeasilyandefficientlywilltend to produceat equilib-

for human behavior comes

groupboundaries,but we are alsohighlysensitiveto selfishactions on the part of other group members.This sensi-tivity leads us to ceasecooperatingwhen that cooperationis not reciprocated, o avoid future interactionwith non-cooperators,and even to engage in personallycostly pun-ishment of individualswho fail to cooperate.

Perhaps he easiestwayto seewarycooperation n actionis to summarizefindingsfromthe ultimatumgame.In thiswidely used experimental cenariotwo anonymous playersarebrought togetherfora singleinteraction.13One is giventhe task of dividing a sum of money between the two ofthem. The other can either accept or rejectthe proposedallocation but if it is rejectedneither playerreceivesanymoney.Of course,froma rationalpoint of view, no playershouldever makea generousoffer to a receivingplayerandno receivingplayershould ever rejecta positive offer. Inpractice,however, he modal offeris 50 percentof the pot,and offers of less than one-third arerejectedby receiving

Humansrecooperative,utnotaltruistic;competitive,utnot xclusivelyo.Wehave ninnatenclinationocooperate,articularlyithindefinedroup oundaries,utwearealsohighly

sensitiveoselfish ctionsnthepartf othergroupmembers.

rium an environmentfilledlargelywith relatively oopera-tive clans. The selfish interest of the clan vis-a-vis otherclansthus acts to reduce the selfish behaviorof individualswithin a clan, but only to the degreethat that reduction nwithin-clanselfishnessimproves hesurvival hancesof indi-viduals n cooperative lans relative o lesscooperative lans.Thus it is not that evolution itself favorscooperationoroperates n anythingbut a selfishfashion;it is simply thatevolution is agnosticabout the methods (e.g., competitionor cooperation) by which overall survivaladvantagesareachieved.

OurTheory:WaryCooperationNew theories of politicalbehaviorcan be built on this cur-rent and subtle version of Darwinianselection theory.Inthat spiritwe offer our own theory of "warycooperation"drawn from the work of leading scholars in evolutionarypsychologyand experimentaleconomics. The theory maybesummarizedas follows.Humansarecooperative,but notaltruistic;competitive,but not exclusivelyso. We have aninnate inclination to cooperate,particularlywithin defined

playersmore than 50 per-cent of the time, meaningthata remarkably igh per-centage of people are will-ing to sacrifice their ownmonetaryrewards o sharewith another player or topunish an allocator whokept a largeportion of thepot.14The latterpractice ssometimescalled,for obvi-ous reasons,costlyor altru-istic punishment.15On the one hand, pureself-interestcannot explaineither our willingness to cooperateor our willingness topunishat our own expense n this scenario.16On the otherhand,purealtruismcannotexplainourwillingnessto pun-ishnoncooperators r,in multiple-play ames, o ceasecoop-eratingwithnoncooperators.Warycooperationdoesaccountfor these behaviors. Moreover, unlike short-term self-interestoraltruism,ourtheory s grounded n anevolution-ary view of human behavior, for it is likely that warycooperation s a productof innate and geneticallyheritablebehavioralpredispositions,and that these predispositionsare the resultof identifiableselectionpressures.17What arethe evolutionaryunderpinningsof our theory?Evolution s a slowprocess,and much of the environmental

pressure avoringhumancooperationhasexistedfor a longtime. Our genetic composition is to some extentthe prod-uct of conditionsfacedbyourhunter-gatherer redecessorsof perhaps 100,000 years ago. One of the keys to anindividual's urvivalwas being a respectedpartof a viablegroup.The centralinsight of a behavioraltheorybuilt onevolutionarybiology is that the desire for group life is afundamentalhuman preference.What kinds of behaviorsoptimally promotebelongingto aviablegroup?Thesetend

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Articles I The Origin f Politicsto be primitives s opposed o elaboratedehaviors incespecific, enetically etermined ehaviorsre ess ikely obe adaptive. paceprecludes complete ist;we mentionjustsix of the mostimportant efaultbehaviorsormem-bersof aviablegroup.Tosustaingroupmembership,ndi-vidualsmust

1. cooperatewith others n theirin-group;2. dislike hose n out-groups;3. punishor banishuncooperativen-groupmembers;4. encouragethersthrough orms,nstitutions,rmoralcodes o (1), (2), and(3);5. beever ensitiveostatus,payoffs, ndreputationel-ative o otherin-groupmembers;6. ceasecooperatingf thenoncooperationf othermem-bersgoesunpunished.Since thesebehaviorsostervaluedmembershipn viablegroups, nd sincebeinganacceptedmember f a properlyfunctioning roup acilitatesurvival,we believe hesesixbehaviors ill be common, houghperhaps ot universal,geneticendowments.Wary ooperationsthe mostsocialoftheories,ince t isbuiltaroundotherpeople.In this sense t affords nothercontrastwith existingapproaches. ccordingo rationalchoice(andold schoolevolutionaryheory), orexample,otherpeople get in the way.Accordingo the theoryofwarycooperation, therpeopleare heway.But this doesnot mean that behavior s altruistic.Social behavior soften taken o be goodand itspolaropposite-egocentricbehavior-bad.Butsocialbehaviors notsynonymous ithaltruistic ehavior.18ather,ocialbehaviors centered notherpeoplebutnotnecessarilyoncerned iththewelfareof otherpeople.In so saying,we arein completeaccordwithKristenRenwickMonroe's ecent mphasis n "inter-actionswith others,"on "sociability"nd on "an nnateneed for humanconnection" s well as MarilynBrewer'saccountof behaviorelevanto in- andout-groups.19In addition o expecting ooperative ehaviorn somecircumstances,ur heory lsoexpects-andempiricaltud-ies haveproven t to be the case-that peoplemindlesslyconform,20assivelybeyauthorityigures,21recompeti-tive to the pointof takingpleasuren the misfortunesfothers,22nitiate hostilities oward hose peoplein out-groups,23onstructut-groupsor hesake fhavinghem,24andaredisconcertinglynthusiasticboutpunishinghosenot perceived s livingup to the group's ehavioraltan-dards, specially henpersonallyictimized.25he theoryofwary ooperationredictsheconstructionf moral odesthatconditionallyaluecooperative ehavior;t doesnotpredictaltruisticbehavior.

Thushumanbehavioreems o beremarkablyonsistentwith the behaviorsxpected yourtheory-an issue uturescholarshipillcontinueoaddress.26erewe mustdefendthe claim that the behaviors isted above areindeed advan-

tageousfor humankind-or at least werein the past.Sinceno one has the abilityto recreate he Pleistocene,evolution-arypsychology s vulnerable o charges hat it merely spins"just-so"toriesby observingmodern human behaviorandthen offeringpost-hoc and perhapseven torturedaccountsof why those behaviorsmight have been valuable in ourspecies'distantpast.27These chargesshould be taken seri-ously because,as is the case with any theory,proponentshaveoccasionallymadeoverstatements.

Havingsaidthis,there s intriguing upport or the notionthat behaviorsand traitsconsistentwith the theoryof warycooperation are preciselythose that lead to success in aharsh and competitiveenvironment. Much of this supportcomesfromcomputersimulationsusedto testmodelsdevel-oped in evolutionarygametheory.These simulationsallowobservationof the relativesuccess of groupsof individualswho follow various behavioralrules.Some groupsin thesesimulationsarepopulatedby altruists, ome by egoists,andsome by warycooperators.When groupswith these differ-ent characteristics reallowedto competewith one another,those composedof warycooperators urvive he evolution-arytest. Groups composed mainlyof altruistsfail becausethey have no protection against the occasional egoist intheir midst. But egoist groups also fail because they aregroups in name only and lack the bonds that lead to thebenefits of group behavior.28We submit that it is not acoincidence that warycooperationwins in computersim-ulations of the evolutionaryprocess and is so frequentlyobservable n realhuman society.Evidencesuggests hat even humans'allegedmentalfrail-ties serve a purpose. Ralph Hertwig and Peter M. Todddemonstrate hatcognitive"limitationsn processing apac-ity, as well as in other resourcessuch as knowledge, canactuallyenablerather handisableimportantadaptive unc-tions."29Forexample, t is now thought thatchildren earnlanguagesmore quickly than adults in part due to theirlimitedmemories.Limitations onstrain olutionspace,allowa scaffolding o guide learning,and suggestpatterns.Neu-ral networksdesignedto simulatelanguagelearningactu-ally learn more quickly with less memory-more is notalwaysbetter.30The sameis trueof rationality.As statedbyCosmidesandTooby,"'[R]ational'decision-makingmeth-ods ... arecomputationallyveryweak; ncapableof solvingthe naturaladaptiveproblemsour ancestorshad to solvereliably n orderto reproduce."31hey concludethat, froman evolutionarypoint of view,humanmentalcapacities, arfrom preventingrationalthought,32actuallyallowus to be"better than rational."33Related evidence that there ismethod to human madness comes from identificationofthe tasks that humans do not do well. Humans are notparticularlyood atcalculation, torageand retrieval f facts,and pure logicalproblems.On the other hand, humans(evenveryyoung ones) areamazingly good at language recognition;intuitivelycom-prehendingthe physics of movement (e.g., throwing an

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objectaccuratelyat somethingwhile it and the throwerareboth moving); reading facial expressions for honest, asopposed to contrived, emotion; making inferencesaboutplantsand animals(asopposedto similarlydistinguishableman-made objects); and identifying others likely to benoncooperators,specially heaters-even thoughthesetasksareas, or more,demandingthanmanywe do poorly.34Forexample,when experimental ubjectsareshownpicturesofindividualsand told their names along with a single factabout them, subjectsare betterat remembering he namesof those who had been connected with a social fact (Sallyhelped a neighbor paint his house) than a nonsocial fact(Tom has an old refrigerator), nd they arebest at remem-beringthose who had been connected to a negativesocialfact (Harry did not return a CD he borrowed from hisfriend).35These very activities-accurately readingsocialsituations, intuiting the physics of the world around us,perceivingfood sources, and especiallyidentifying thosewho might harm us or our group status-are those thatwould have beenquiteuseful o ourancestors stheyroamedthe savannah n smallbands.As biologistWilliam Hamil-ton put it, "The tabulaof humannaturewasnever rasa."36The idea that human nature exists should not be con-fusedwith the belief that it is fixed. In fact, humansmustbe sensitive o environmentalsurroundingso achieveobjec-tives. For example,levels of trustingbehaviorvarywidelyaround the world and even within a country.37 f naturalselection over hundreds of thousands of generationshasweeded out the altruistsand the egoists, leavingonly warycooperators,why is cooperativebehaviorpresent in suchvastlydifferentproportionsaroundthe world?The answeris simplebut important:warycooperation s a conditionalbehaviorand thus fundamentallydifferent from strategiesof "alwaysact in your own interest"or "alwaysact in theinterestof the otherperson."The warycooperatoracts witha glanceoverthe shoulderand adjustsbehavioras needed.Thus the behavior of a warycooperator n Lombardywillbe quite unlike that of the same genotype transplanted oSicily, merely because these two individuals are likely toencounterdifferent evelsof cooperativebehavior.Most contributions of rationalchoice and behavioralistscholarship re n no waycompromisedbut rather nhancedby the acceptanceof evolutionary heories.The previouslycontent-free"preferencetructures" f rationalchoice canfinda sourceof clearandpotentially alsifiable ontent, andthe "incoherentenvironmentalism"38f behavioralism anbegiventheoretical oherence.The messagesnot thatnaturetrumpsnurturebut ratherthat nurture s in our nature-and that the precise nature of our nurturing tendenciesdependsuponenvironmental onditions.39Ourtheorysub-sumes a wide range of behavior-as any successfulsocialtheory must. Work that takesevolution seriouslyopens abroadrangeof cooperativeopportunitiesacrossmethodol-ogies and approaches hat in the past have created ines ofconflictwithin politicalscience; t drawson andencourages

more theory-drivenresearch n fields as diverseas gametheory, traditionalfield work in anthropology,and eco-nomic and social psychologicalexperiments,with world-wide human (and nonhuman)subjects.40Implicationsfor Politics andPublic PolicyBuchananandTullockareundeniablycorrect o assert hat"theonly final test of a model lies in its abilityto assistinunderstandingrealphenomena."41We believe our theoryof warycooperationhas practicalapplications n a varietyof areas n politicsand politicalscience.Death, taxes, and welfarePerhapsnot surprisingly, theorythathighlightsourinnatewillingnessto engagein costlypunishmentshedsconsider-able light on policies that relate to criminaljustice. In thedeath penaltydebate,the theoryof warycooperationsug-geststhat supportfor such draconianpunishmentsderivesmore from a desire for retributionthan from an effort todiscouragepotential murderers.This explains why, withresearchconsistentlyfindingthat the deathpenaltyhas nodeterrentvalue, the Americanpublic largelysupports it.Historically, xecutionshavebeen publicevents,and whiletheirpublicnature s certainlycompatiblewith the goal ofdeterrence, t is difficultto see why anyonewithout a gen-eralinterest n punishmentwould attend.Tax law providesanother example. Do we punish taxevadersbecausewe all wish to be tax evadersand must befrightened nto compliance,orarewewillingto payourfairshareonly assumingothers do the same and evadersfaceswiftand certainconsequences?Warycooperationsuggeststhe latterreason.

Turningto the positiveside of government,welfarepro-gramsareoften poorly regarded, n largepart, apparently,because it is believed they provide benefits to unworthyrecipients.Why arepeopleso willingto helpthe downtrod-den, and why are they even more willing to stop helpingwhen they suspect even a handful of undeservingrecipi-ents?Our theoryprovidesa readyexplanation orboth ten-dencies.Peopleareinitiallyhelpfulandcooperative,evenatsome personalexpense,but they arehypersensitive o thepossibility that someone might take advantage of theirgenerosity.The warycooperationtheoryalso has somethingto sayabout the viabilityof strategies or dealingwith cheaters ngeneral.In the welfaredebate,for example,policy makerscommonlyargue hat increased ffort to eliminatecheatingis simply inefficient,as the amountspenton detectingeachadditional cheat will exceed, often substantially, he costthat the cheaterwas imposing. This rationalappealtypi-callyservesonly to furtheroutragethose whose taxcontri-butions finance welfarebenefits.But does anyone actuallythink that it would be a good idea to senda cheatto live in

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Articles I The Origin of Politicsluxuryin the Caribbean or what we would otherwisepayto bring him to justice for his noncompliant behavior?In this and relatedpolicy matters,we see clear evidencethat people are willing to support and engage in costlypunishment-and also to comply themselves-as long asthey remainconfident that noncontributorsare routinelypunished. The perception of a general failure to punishothers, ratherthan the perception that we are personallyunlikely to be punished, is the proximate cause ofnoncompliance.

WarThe phrase"millions or defense,not a penny for tribute,"popularizedin the United States during the days of theBarbary iratesbutjustas relevant o today'smood (if a fewzeroesareadded),nicelycaptures he rolewarycooperationhas played in the history of internationalaffairs.The factthat wary cooperation helps account for humans' innatecapacityto cooperateon a largesocialscale is by no meansequivalentto a notion that the socialside of human natureis benignin its aims and impacts.Perhaps he most strikingexampleof this is war.The most apparentcharacteristic fwar at the group level is conflict, but we should not missthe fact that at the individual level warfare s among themost cooperativeandself-sacrificing f all humanventures.In no other arenaare humans as likely to set aside theirpersonalhealth,well-being,comfort,and evenlife itself,infavor of group success. Selfless participation in warfareremains, even in modern societies, at the center of ournotionsof heroismandennoblingactions.Whilethesavageryof warclearlyderives rominnatehumanfeatures, t isclearthat the explanationof group-levelsavagery s not simplyan aggregationof individual-levelaggression.While theimportanceof cooperation orpeacemayseemself-evident,cooperationis no less criticalto war.This seemingcontra-diction is troublingto many.As KennethWaltznotes, forexample, "While human natureno doubt plays a role inbringingaboutwar, t cannotby itselfexplainboth warandpeace.Waris, of course,only an extremeexampleof the appar-ent contradictionwithin an explanationfor both coopera-tion and competition that focuses on human nature. Insome ways,avoidingthis contradictionmaybe the greatestvalue of the wary cooperationtheory.Though our theoryposits a simple baseline explanationof human nature, itdoes not assume that human naturemust be either com-petitiveor cooperative;rather, t explainsone sourceof therelativeimportanceof competitive and cooperativedriveswithin a givenbehavioralcontext.43Similarly, t freescom-petition from a necessary ie to self-interest ust as it freescooperationfrom any necessaryassociationwith altruism.We do not havea geneforcombativenessanymorethanwehave a gene for cooperation, or indeed a gene for warycooperation.What we have is a set of innate behavioral

repertoireshat allowswarycooperation o emergeand dom-inate in a varietyof fairlycommon social environments.Rational actor models would predictuniversalfree-ridingin the faceof the overwhelmingcosts to individualpartici-pationin warfare.44While some individualsdo indeedseekto avoid participationand sanctions againstdesertersarenecessary,he modal behavior s often voluntaryparticipa-tion, particularlywhen perceptionsof group membershipand expectationsof cooperationby other group membersare present. Behavioralists'explanationsfare no better,asneitherthe level of self-sacrificingooperationnor the levelof violentcompetitionfindanynaturalparallelsn the aver-ageindividual'speacetimebehavioralrepertoire.Bothcoop-erationand conflictreach heirzenithin the samebehavior.

What other features of war does an evolutionaryviewlike ourshighlight?Sincein this view,the conflict of war isagroup-levelphenomenon, group-level actorsbecomepar-ticularlysalient. Markersof in-group-out-group bound-aries,for example (e.g., borders, anguage,ethnicity,race,religion, citizenship) should assume exaggeratedimpor-tance in both the developmentand prosecutionof war.45Similarly,the intergrouppressureof competition shouldlead to an exaggerated ntragroupfocus on cooperation,groupsolidarity, ndpotentialinternalbetrayal witness hepublicmood in the United Statesafterthe terroristattacksof September11, 2001). Human behavior n war is a bun-dle of seemingly intractablecontradictions.Nevertheless,our theorycan accountfor the inherent humanwillingnessin wartimeto riskone's ife fora chanceto takethe life of anout-groupmember,or savethe life of an in-groupmemberwhile simultaneously supporting the killing of in-groupmemberswho areunwillingto kill out-groupmembers.

Political institutionsWhy do political nstitutionsexist?This questionhas bedev-iled politicalthinkersfor quite some time. If people preferthe kind of societal life that is made possible by politicalinstitutions,why do they not merelylead that life in thefirstplace?The infiniteregressproblem mmediately cuttlesmost attemptedexplanations,butwary cooperationoffersaway out: the existenceof politicalinstitutionsmay in largepartbe attributed o people's ntense desirefor sanctionstobebroughtagainstnoncooperators.Peoplemaybelievethat,if left to theirown devices,most of theircompatriotswouldbe cooperativegood citizens,but the possibilityof even averysmallnumberof badapples s enoughto drivethem tocreate nstitutions.Do people not realize hat those elevatedto positionsofpowerin institutionshavegreatpotentialto takeadvantageof their positions for personalgain?Indeed they do, andthusit shouldcome asno greatsurprise hatpublicopinionresearchindsthatAmericans'primary ourceof dissatisfac-tion with governmentis not that it makesbad decisions,but ratherthat it makes decisions for self-servingrather

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thancommon-good reasons.46Government s typicallydis-liked not because t is perceived o do thewrongthings,butbecause it is perceivedto do things for the wrong reasons.Ascribed motive is the main determinant of responsestothe actions of others in any societal interaction, but itbecomesacutelyimportantwhen we aredealingwith peo-ple possessingsubstantialclout.47In this sense, the theoryof warycooperationholds that those staffingpoliticalinsti-tutions should be subject to specialvigilancenot becausetheyarefundamentallydifferent ife forms butbecause heirambition for authorityraisessuspicions,and their posses-sion of authoritymakesthem potentiallydangerous.48What does the theory say about the specificmannerinwhichpolitical nstitutionsshould be constructed? orstart-ers,it points out the prescienceof the architectsof the U.S.political systemin making t difficultfordecisionmakers ofurther heir own ends.Separationof powersacross nstitu-tions, frequentelections,federalarrangements, nd astrongjudiciary-all facilitate he balancingof one group'spowersagainst another's,which is, not coincidentally,extremelypopularwith the masspublic.This does not mean, however, hat the Americanpoliti-cal systemis a perfectfit with the institutionalpreferencesflowingfromthe theoryof warycooperation. n thepeople'sview, fartoo many opportunitiesstill exist for electedoffi-cials to feathertheir own nests. Think of the ire evokedbymattersrelatingto congressional alary,pension plans,andperquisites,not to mention the perceivedunholyallianceofwell-heeledinterestgroupsand politicians.Warycoopera-tors like to cooperate,but they are also predisposed o bewaryof theactionsof others-especially thoseeager o makedecisions for the group.Reformerswould do well to realizethat people do not wish to be in control of the politicalsystem; heyonlywantthose who are n controlto be unableto take advantageof their positions. If people were confi-dent thatexistingconstraintsprohibitedsuchself-interestedactions, they would pay even lessattention to the politicalarenathan they do now. For most people, involvementinpolitics is drivennot by a desireto be heardbut by a desireto limit the power of others. CurrentAmerican foreignpolicy might be improved,for example,if decisionmakersrealizedthat, like Americans,people in AfghanistanandIraqdo not cravedemocraticprocedures.Kurdssimplydonot want to be dominatedby Sunnis;Sunnisdo not wantto be dominated by Shiites;Uzbekisby Tajiks;and Tajiksby Pashtuns.Peopleoften expressa desirefor participatorydemocracywhen they reallyjust want to avoid being vic-timizedby a more powerfulgroup.

The Genetic Inheritance ofPolitical OrientationsTothispoint,we haveasserted hatthemodalhumanbehav-ioral tendency is neitherself-interestednor altruistic,but,rather,warilycooperative.Further,we claim that the prev-

alenceof warycooperatorss the resultof evolutionarypres-suresthatrewardhumansskilled at creatingandexistinginviablesocialunits. Our theoryis only one of manypossibletheories hattakeseriously he connectionbetweenbiology,naturalselection,and human behavior.Evidencesupport-ing this connectioncan be found in the literature n genet-icsandhumanbehavior hathasflourished,asnew fields ofinquiryoften do, with the acceptanceof a new methodol-ogy. In this section we explore one such methodology-twin studies.

CharlesDarwin and AlfredRusselWallaceworkedoutthe detailsof natural electionat roughly he sametimeandwere in remarkableagreement-with one vital exception.Darwinwascompletelyconsistentandcontendedthatnat-uralselectionappliedto behavioral swell asphysical raits.Wallace,on the other hand, drew a bold line betweenthetwo, positingthat the mentalrealmwas immune to evolu-tion andwasinstead hepurviewof etherealreligiousuncer-tainties.Wallace's ositionisstillfavoredbygroupsasdiverseas religiousfundamentalistsand manyin the socialsciencecommunity who refuseto believethat behavioris innate.The naturalscienceshave sided almost unanimouslywithDarwin.Wallacespentthelastyearsof hislifelargelygnored,a religiousmysticwho thought it possibleto communicatewith the deadand who trieddesperately o hold on to thefictionthatourbrains,behaviors, ndbeingsconstitute ome-thing more than just anotherstep along the evolutionarypath.49If politicalscientistsbelieve,with Darwin, that geneticsinfluencessocialattitudesand behavior, t is not evident intheirresearch.Certainlyno recentarticle n aleadingpolit-ical science journal has used genetics as an independentvariable.50Perhapsmost believe that, while genetics mayplay some nebulous role in politics, environmentalcom-plexitiesandhumancognitivecapabilities verwhelmgeneticeffects, allowing the discipline'sresearchagenda to com-pletelyignorebiology. Contrary o popularbelief,however,the link betweengenesandpoliticalattitudesandbehaviorsis strong. In this section we hope to convince readers hatgeneticvariablesaremoreinfluentialthan is generallycon-cededby politicalscientists.To do so we summarizerecentresearch n twins,on autism,andon genetic-environmentalinteractions.

TwinsMonozygotic (MZ) twins have virtuallyidentical geneticcodes;dizygotic(DZ) twins,like otherfullsiblings,shareaslittle as 50 percent of the genetic materialthat varies inhumanbeings.51This makesthe studyof the adultcharac-teristicsof these two typesof twinsof specialvaluein com-putingthe relativenfluenceof heredityandenvironment-especiallywhen augmentedbya comparisonof twinsrearedseparatelyand those rearedtogether.Suchstudiestypicallyidentify a substantialamount of genetic influence and a

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Articles I The Origin f Politicssurprisinglypaltryamount of parental nfluenceoperatingthrough the environment.52To take a single example,thecorrelationn generalntelligencebetweenMZ twins s muchgreater han the correlation n general ntelligencebetweenDZ twins. While general ntelligenceappears o bethe mosthighlyheritablebehavioral raitdiscovered o date,53 hou-sands of twin studies have documented the heritabilityofnumerousothertraits, romsusceptibility o drinkingprob-lems to manners of talkingand from likelihood of divorceto religiosity.Those who believegeneticcompositionis irrel-evant to behaviormust offeran alternativeexplanationforthe persistentfindings from twin studies; otherwise, theunavoidableconclusionhas to be that an exclusive ocus onenvironmental actorsmissesa majorsource of the variancein attitudes and behavior.

As interestingand challengingas these findingsmay be,do they relate o the concernsof politicalscience?While, toour knowledge, there are no political scientists currentlyperformingtwin studies, much of what is being done inotherdisciplines s surprisingly elevant o avarietyof polit-ical science subfields.The most developedexamplescomefrom studies of the heritabilityof traits such as conserva-tism and altruism.These behaviorshave been studied indifferenttwin populationsin different countriesby differ-ent researchers verthe last twentyyears.All of the studiesreachthe sameconclusion: apredispositionoconservatismsgeneticallyheritable.This is an empirical assertionthat should startle andintrigueanypolitical cientist.Thus far hepertinentresearchhasbeen done largelyby psychologists,and there is consid-erableworkaheadin this field for politicalscientists.Socialpsychologists reatconservatismasa broadpersonality rait,roughlyequivalent o otherpersonality raits,such as intro-versionorauthoritarianism. or mostpoliticalscientists hisis not the first thing that comes to mind when someonementions a specific ideology like conservatism.Conserva-tism as a belief system,a bundle of policy preferences,or anascent group identificationare all examplesof commonpoliticalscienceconceptualizations f conservatismhat dif-fer,often substantially,rom the view of conservatismas apersonality trait. Fortunately,the details of at least onemethod by which social psychologistsmeasureconserva-tism allow us a glimpseat some component characteristicsthatare morecompatiblewith our conceptionsof it. Beforeproceedingto the detailsof conservatism,we must discussin more detail the methodologyof twin research.The value of twins to genetic researchdoes not comefrom the distinctivenessof their attitudes and behaviorscomparedto the population of nontwins. In fact, the lackof distinctivenessis a salient and empiricallyestablishedprerequisitefor generalizingtwin findings to the generalpopulation.54Nor is the valueof twinsthe obviousfact thatthey aregeneticallycloserthan nontwin siblings.The realresearchpower comes from the fact that the two distincttypes of twins differ geneticallyin known ways. For any

traitthat is at leastpartlyheritable he tendencyfor mono-zygotic (MZ) twins to sharethat characteristic hould bestronger hanthe tendencyfordizygotic(DZ) twins to shareit. In contrast,characteristicshat arisefrom the environ-ment, whethersharedby the twins, as would typicallybethe case for parentalsocialization,or not shared by thetwins, as would be the case for many adult experiences,shouldnot generateanysignificantly ifferentpatternswhenwe contrastMZ and DZ twins.55

This assertion-that the effectof geneticsis measurablydistinct for MZ and DZ twins while the effect of the envi-ronment is either equivalentor at least randomlydistrib-uted around equivalence-is crucial to everything thatfollows from twin research. t is important herefore o con-sider criticisms of this fundamental assumption, whichappear n two essentialvarieties.The first s thatMZ twins,genetics aside, experiencea similar environment becausethey are treatedmore alikethan areDZ twins. This couldbeparticularlyellingforchildhoodsocializationf, for exam-ple, parentsof MZ twins tended to treatthem lessas indi-viduals than do parents of DZ twins. The second basiccriticism is that MZ twins, genetics aside, interactwitheach other more throughoutlife than do DZ twins. Thiscould be particularlymportant or adultsocialization; loseadult contact betweenMZ twins might lead us to expectahigher degreeof environmentally nduced similaritythanwe would expectin DZ twins.Both criticismshave beensubjectto sustainedandvariedinvestigation,and neither hasbeen found to hold up underempirical crutiny.The argumentof more similar reatmentfails on several ronts.Parentsfrequentlymiscategorizeheirtwins (DZ twins are often believed by their parentsto beMZ twins) and the differentialcorrelationpersists n theseinstancesof miscategorization. n other words, the degreeof correspondencebetweenMZ twinssurpasseshat of DZtwins even in a subpopulationof twins all of whom werethoughtby theirparents o be MZ twins.56The speculationthat MZ twins have closeror more frequentcontact thanDZ twins turns out to be, at best, irrelevant.The correla-tion between interpaircontact, for example,and interpairdifferencesn conservatisms in factslightlynegative.57 hatis, the more contact, the less ideologicalsimilarity.But themost powerfulrefutationof both criticismscomes in recentstudiesthatcompareMZ and DZ twins raisedapart.Thesestudies uniformlyvalidateMZ and DZ differences oundin earlier tudiesof twins raisedtogether.Argumentsaboutthe relativedegreeof sharedenvironmentaleffectsbetweenMZ and DZ twins simplyoffer no credibleexplanation fthe twins in question have been raisedapart.58 n effect,this naturallyoccurring, f uncommon, condition providesprecisely he sort of laboratorycontrol that we want in anexperimental etting.59Empirical videncealsosuggests hat dentical winsrearedtogetherare often less ikely to sharebehavioral raits thanareidentical twins rearedapart,perhapsbecauseof parents'

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extra effortsto help the twins living togetherestablishdis-tinct identities.In addition,as adult MZ twins livingapartage, theytend to becomemore,not less,similar,60 findingdifficultto reconcilewith the belief that only the environ-ment matters.Interestingly, his preciseeffect is predictedin an early landmarkcriticism of behavioralismand theconditioned-responseesearch n animalbehavior t itscore.Over time, substantial anomalies accumulated in thisresearch,pointing towarda primacyfor some nonenviron-mental behaviors. As Keller and Marian Breland sum-marized this tendency, "Learned behavior drifts towardinstinctive behavior."61

While the twin-studymethodologyhasyet to be appliedby politicalscientists,a varietyof socialpsychologystudiesusethismethodologyto assessconservatismas apersonalitytrait.The measureof most interest to politicalscientists isthe Wilson-PattersonAttitude Inventory.This inventory sadministeredby presenting subjectswith a short stimulusterm,such as deathpenalty,and elicitinga simpleresponseof agree,disagree,or uncertain.The broadestversionof theW-P inventoryincludes50 items, half of which contributepositivelyto the conservatism core and halfof which con-tributenegatively.While some of the items relateheavilytoconceptions of social conservatism-for example,pajamaparties,casualliving, nudist camps, computer music, andhoroscopes-others aremore directly political-for exam-ple disarmament, egalizedabortion, socialism,patriotism,and deathpenalty.Studiestypicallyuse reducedsets of W-P items or tailorindividual items to the country in which the studies arebeing administered.Forpolitical scientists,this is frustrat-ing on two counts. The list of politicallyrelevant tems istantalizingbut limitedandunfocused;moreover, he resultsareoften presentedonly for the entirecombinedscale,mak-ing it difficult o assess he contributionof thedirectlypolit-ical items to the overall index of heritability.Heritabilityestimatesfor overallconservatismas measuredby the W-Pinventory are quite high, usually around 0.60. Further-more,the modelstypicallyshowlittleor no effect forsharedenvironment(the restis likelythe resultof nonsharedenvi-ronmental factors, including the prenatalenvironment).These findings come from studies of twins in settings asdisparateasAustralia,Virginia,and Minnesota. The Min-nesotafindings,basedon twins rearedapart,conformwellto the otherstudies,basedon twins raisedtogether.62Our concern at thispoint is not to quibbleoverwhetherin factgeneticsaccountsfor more thanhalfof the variationin individual evels of conservatism.The importantpoint isthat our discipline currentlybelievesthat the state of theworldwouldyieldanenvironmentali.e.,socialization) om-ponent greater han 50 percentand a genetic componentofessentiallyzero.63Whateverthe precisenumbers,the dura-ble and starkresultsof twin studies arecompletelyat oddswith currentthinking in our discipline.At the very least,political science can no longer dismiss genetics as some

minor or distant precursor o important political orienta-tions. Taken at full face value, these results suggest thatvirtuallyeverythingwe havesupposedto be true about theoriginsof broadpoliticalorientations s false.

As strongas this assertionmayseem,it does not indicatethat most of the extantempiricalresultsconcerningpoliti-cal socializationareincorrect,only that the interpretationscommonly appliedto those resultsare incorrect.In fact, itstrongly reinforces the most salient empirical findings.64We know,for example,that if both parents harea politicalorientation and find politics to be important, their off-springwill likely have that same political orientation andalso valuepolitics.65Nothing in thisdescriptioncontradictsthe twin-based contention that political orientations arelargely he resultof genetic inheritance.In fact, this is pre-ciselythe patternwe would expectto find if the twin stud-ies are correct.The challengeof the twin researchs only toour assumptionthat, unlike sharedphysicaltraits,sharedbehavioralraitsmust be purelya function ofenculturation.At this point, the carefulreader s undoubtedlythinkingthat we are getting ahead of ourselves.As we mentionedearlier, he 50-itemW-P inventorycontains both items thatwe would recognizeas havingsubstantivepoliticalcontentand others about which we would entertain substantialdoubt. If the overalldegreeof heritability s largelya func-tion of attitudestowardmodernartandpajamaparties, hedirectrelevance o politicalsciencemay be minimal. Whatthen do we find in the limited studiesthat reportheritabil-ity for individual items?A 1986 studyby Martin and col-leagues of over 3,800 Australianand British twin pairsreportedthe following estimatesof heritability on a scaleof 0 to 1.0) for the following items: death penalty,0.51;white superiority,0.40; royalty,0.44; apartheid,0.43; dis-armament,0.38; censorship,0.41. The heritability stimateforpajamaparties,on the otherhand,was a mere 0.08. Thecomparableestimates for the influenceof shared environ-ment were: death penalty, 0.00; white superiority,0.09;royalty,0.14; apartheid,0.05; disarmament,0.00; censor-ship, 0.03 (but pajamaparties,0.44).66These findingsarepowerfulyet incomplete.As politicalscientists,we could allprovideourown list of items thatwewould want to see includedin this sortof inventory.Partyidentification,for example,is probablyan obvious item.67In anycase,the implicationfor politicalscience is clear:wemust takeseriously he possibilitythatat least somepartofwhat we haveattributed o activesocialization s a functionof genetic inheritance.Equally mportant-and even moreurgent-is the need to adapt twin-studymethodology toour specificdisciplinaryresearchagendas.Weinvitedubiousreaderso indulgeus in athoughtexper-iment.Suspenddisbeliefandthinkseriouslyaboutwhatthismight mean for our entireconceptionof politicalattitudesandorientations.Daniel Elazar haracterizedheAmericanSouth as havingwhat he calleda traditionalistorientationtowardpolitics, and arguedfurther that these presumably

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that differences n methodology within and acrossdisci-plines may deriveat least in partfrom heritabledifferencesin brainphysiology.When MorrisFiorinawrites that he is"rational hoice down to [his]DNA," he could be literallycorrect.71Likewise,seemingly ncompatibleconceptionsofbasic human nature may in fact be equallycorrectwhenappliedto the propersubset of the population:fundamen-tal behavioral differencesmay also derive from heritabledifferences n brainphysiology.

GenesFor a varietyof reasons,skepticism persistsregardingesti-matesofheritabilityderived rom twin studies. But the casefor genetics playinga role in shapingbehavior s based onmuch more than twin research.Since much of the resis-tance among social scientists to genetic independentvari-ables stems from unfamiliaritywith modern biology, it isessentialthat we offer some indication of the manner inwhich geneticsconnects to behavior.The most interesting and numerous genes in humanbeings are not structural blue eyes or brown),but regula-tory. Regulatorgenes allow an organismto respondto itsenvironment;they are the genes that turn on and off thetranscriptionof other genes (or themselves).They are thebody's thermostats. Consider, for example, biologicalresponses o fearand threat.The amygdala,a primitivepartof the lower brain,sounds an alarm,adrenalineand stresshormones are released,extrablood is sent to the muscles,breathingquickens,glucoseis disgorgedby the liver,bloodpressure kyrockets,and nonessentialbodilyfunctionsstop.While this generalpattern s always n evidence,sensitivity,rapidity,and depth of responsevarywidely. Some peopleexperiencea full-fledgedpanic attack at the slightestprov-ocation;othersdisplay quite muted physiologicalreactionsto evenlife-threatening vents.Why?Justaspeople displayphysicaldifferencespartlybecauseof differences n struc-tural genes, they displaybehavioraldifferencespartlybe-cause of differencein regulator genes. People'sregulatorgenes encourage them to react to environmental condi-tions in certain ways, whether those conditions involvedanger, lactose, alcohol, an uncooperativeindividual, orsexual stimulation. These on-off switches shape much ofour personalities.72

Regulator genes are not merely theoreticalconstructs.They havebeenseen,andbehaviorhas beenaccuratelypre-dicted merely by knowing the shapeof an individual'skeyregulatorgenes. Considera gene calledD4DR, which canbe found on the short arm of chromosome 11. This geneinfluences the receptionof dopamine, a chemical neededby portionsof the brain.Individualswith a long version ofD4DR are ess efficientatcollecting dopamine,so theyneedto takemore risksandseekmore thrills n orderto generateas much dopamineas othersacquirefrom more sedentarylifestyles.73

None of this is to say that environmental factors areirrelevant.74 n fact, surprisingly,and perhapsfor many,reassuringly,urrentbiologicalthinkingunitesgeneticandenvironmental nfluences n a complicatedbut compellinginteractivefashion. For example,recentwork on violencehas found a connectionto a blood transmitter alledmono-amine oxidase A (MAOA), but the connection is notsimplythat those with genetic deficiencies n MAOA pro-duction are more violent. Rather,those individualswhohave low MAOA production and experiencedviolence aschildren are much more likely to be violent as adults.75Acting alone, MAOA deficienciesor a violent childhoodhave little predictivepower,but the interaction of geneticand environmentalfactors is disconcertinglypowerful.Asimilar pattern has been found in studies attempting topredict hose individualslikelyto suffer romclinicaldepres-sion. There is a gene on chromosome 17 called 5-HTT.This gene also comes in a long form and a short form.Anextensive study of nearly 1,000 New Zealanders foundthat clinical depressionwas associatednot just with thepresenceof the short form of 5-HTT and not just withthe presenceof many high-stressevents in a subject's ife,but with a combinationof high-stress ventsand the short-form of 5-HTT.76If we generalize hispattern o predispositionsmorecen-tral to politicalconcerns,suchas conservatismand cooper-ation, it suggests hat the eventualgenetic-levelexplanationmay fit surprisinglywell with much of what we alreadyknowandmuch of what we havesuggestedherewith regardto geneticsand evolution. Perhaps or a substantialportionof the population the short form of a yet-to-be-identifiedgene allowstheir degreeof cooperation/retaliationo varywith regardboth to the specificcurrentcontext and to theirbroaderexperienceover time. We might call these individ-ualswarycooperators.Foranotherportion of the popula-tion, those with the long form of the gene, contextseems tomake little difference.While we believeour theoryof warycooperationdiscussedearlieraccurately escribes he behav-ior of the majorityof the population,it needs to and doesleave room for a minority that behaves in a predictablydifferent ashion.This minoritymaybe fixed n eithercoop-erativeor noncooperativemodes,or mayalternatebetweenthese modes with no apparent egard or the cooperationorlack of cooperation evidenced by those in their environ-ment. This could provide a rough genetic sketch of thebiologicalunderpinningsor the starklydifferent ocial abil-ities and orientations discussed above in the section onautism.And that discussionin turn providesthe cognitiveunderpinnings or the theoryof warycooperationthatwasthe startingpoint for thispiece. Interestingly,he computersimulations hatwe discussedabovehavedemonstratedhat(underreasonableassumptions)a populationconsistingoftwo typesroughly compatiblewith mildlyautisticindivid-uals and wary cooperators,respectively, an reach a stableequilibriumwith the larger part of the final population

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Articles I The Origin f Politicscomposed of wary cooperatorsand the smallerremainderbehavingmore like the mildly autistic.77

ConclusionWe beganby noting the usefulness o the social sciences ofgreaterattention to biologicalevolution. Part of the prob-lem maybe thatthe last time biologycame to the attentionof politicalscientists(in the 1970s, after the publicationofE. O. Wilson'sSociobiology),hey believedthat advocateswere sayingthat behaviorwas determinedby biology.78 fthat was ever the position of biology proponents, it is nolonger.A researchprogramfocused on genetic factorstothe exclusion of environmental factors would be deeplymisguided-perhaps as misguided as a researchprogramfocusedon environmentalactors o theexclusionof genetics,

We believe hat modernbiological heory,properlyunder-stood,holds thepromiseof unifyingpolitical cienceand nte-gratingpoliticalsciencewith other socialsciences-and evenwith the naturalsciences.Concepts borrowedfrom evolu-tionarybiology can providethe theoreticalguidancebothrational hoice and behavioralism eed to becomeeven moreuseful thantheyhave been to date.Scholarsoperating n thebehavioralraditioncancertainly ontinueto identifyaspectsof the environment that seem more influential than othersand the conditions under which these environmental nflu-encesvary.Though we are convincedthat it is necessary ounderstandoriginalcauses, hisunderstandingwouldbe use-lessif behavioralists id not link thesecausesto moreprox-imateenvironmentalactors.Scholarsoperatingn the rationalchoice tradition aresimilarlyadvantagedby the incorpora-tion ofbiological heory, incepreferencesanbe derived roman understandingof evolutionarypressuresactingon indi-viduals and groups.Baselinesof behaviorno longerhave tobe questionableassumptions hatpeoplepursueshort-termtangible gainsor largely autologicalassumptions hatpeo-ple mustpreferwhatevertheypursue.We arenot suggestingthat all politicalscientists becomeevolutionarytheorists or adopt a certain methodologicalapproach.We need a healthydisciplinarydivisionof laborin serviceof a genuinetheoryof politicaland social behav-ior. To date in the social science disciplines that haveembracedevolutionary heory,everything romparticipant-observation to pure mathematicalgame theory to labora-tory(and field)experimentshas beenemployedwith fruitfulresults.As the old sawgoes, "Ifyou arelooking in the rightplacethere s no wrongwayto look."79Evolutionaryheoryhas the potential to render obsolete our intradisciplinaryconflicts over approach,method, and theory. It is morethan a replacementfor existing political science theories,sincerationalchoice andbehavioralism resimplynot capa-ble of supplyingan account of ultimatecauses.It is a truetheory of the origins of behavior and as such providesabasisfor bringingtogetherthe remarkably iverseand use-ful ongoing researchn politicalscience and beyond.

Notes1 Huxley 1989. For an intriguingcounterpointto Hux-ley,see Kropotkin1972.2 Darwin 1968; Darwin 1981.3 Simon 1957; Simon 1985; Simon 1997.4 Cosmides andTooby 1994, 327.5 Ostrom 1998, 2. See alsoJones2001.6 Dawkins 1976, 2-4.7 Ibid., 66. At a laterpoint, Dawkinsclearlyrecognizesthat genesoccasionallyengagein cooperativebehavior.8 Buchananand Tullock 1962, 18.9 See, for example,Dawkins 1976; Dawkins 1982.10 Such "selfish"geneshave been used to explainthe self-destructivebehaviorof organismssuch as salmon,which leave he comfortsof the oceanto swimupstream,

spawn,and die, and male mantises,which some-timescelebratehe end of copulationbythrowing hem-selves back into the open, waitingjawsof the muchlarger emales.11 Fora descriptionof the biology of cooperationat thislevel, see MarkRidley2003.12 Frank1988; SoberandWilson 1998. The claim thatselectionpressures an be profitablyviewed at levelsother than the gene,while increasingly cceptedby stu-dents of evolutionarypsychology,has had a check-ered past, and many scholarsremain unpersuaded.Wynne-Edwards's962 accountof rooks volun-tarilycurtailingtheir clutch size for the welfareof theflock was roundlycriticizedby the biologicalcom-munity and gave relatednotions much to overcome.Moreover,o-calledgroupselectionists ccasionally nvi-sioned an ethereal,almost inexplicable,and some-what magicaldevotion to the group.As emphasized nour descriptionabove,mysticismis not needed,andmodernthinkingis firmlyembedded n the logicalcon-sequencesof selectionpressuresat different evels.The switch in common phraseology rom groupselec-tion to multilevelselection is more than semantic.

In addition, continuing advances n geneticscastmore and more doubt on genes as stand-alone,clearlydistinct actors n the evolutionarydrama.Instead,we now know from Gibbs 2003 and othersthat manytraits areepigenetic,that is, influencedby chemicalmarksoutside the DNA sequence.Today,much of thedisputecenters on the relativeimportanceof multi-level selectionrather han the factof its existence.Forsome sense of the opposing arguments,see theexchange n the 1994 issueof Behavioral nd BrainSciences,n which Wilson and Sobermake the case formultilevelselection,and which is commentedon bynumerous scholars.See also Sober and Wilson 1998.

13 For a good summary, ee Guth andTietz 1990.14 See Nowak, Page, and Sigmund 2000. Lest it bethoughtthat thegenerosity f allocatorssmerely tra-tegic, when the game is changedto a dictatorgame, in718 Perspectives on Politics

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which thereceivingplayerhasno chanceto veto thepro-posedallocation,anonymousallocators till makegen-erousoffers,albeittypically essgenerousthan inultimatumgames.See Fehr and Fischbacher 003 fora usefulsummary.See Van Dijk andVermunt2000for researchsuggestingplayers n the dictatorgame areactuallymore generousthan those in the ultimatumgame.15 Boyd and Richerson1992; Gintis 2000; HenrichandBoyd 2001; Price,Cosmides,andTooby 2002.16 Thus the discipline'sheoreticalbasis orexplaining elf-interestedand non-self-interestedbehavior s much asOstrom, Gardner,andWalker(1994) describe twith regard o the specificinstanceof abusingor notabusing common pool resources. "No existingtheoryprovidesa consistentexplanation or how andwhy many appropriators xtricate hemselves romcommon pool resourcedilemmas[and]why this is notuniversallyhe case"(p. 18).The theoryof warycoop-eration,in contrast,does explainthesepuzzles.17 The main reason for opening up social scienceto therole of biology is not that it can betterexplainvaria-tions of human behavioraround the mean but ratherthat it can help us to betterunderstand he meanitself.We do not, however,believeall individualspos-sess in the samedegreethe biologicalmaterials hatpushus towardwarycooperativebehavior. nfact,asdis-cussedbelow,we believe that a minorityof humansaregeneticallypredisposed owardinsensitivityto thecooperation evels of people aroundthem.18 Fora thoroughtreatmentof altruisticbehavior, eeMonroe 1996.

19 Monroe 2002, 203; Brewer2000.20 Asch 1951.21 Milgram1974.22 Frank1999.23 Sherifet al. 1961.24 Barash1994.25 Price,Cosmides,andTooby2002.26 Extantresearchroughlyconsistent with our themeincludes KahnemanandTversky1984; Kahneman,Knetsch,andThaler1986;Orbelland Dawes1991;Tha-ler 1992; Kahnand Murnighan1993; Komoritaand Parks1996; Ostrom, Gardner,and Walker1994;Hoffman, McCabe,and Smith 1996; Wedekindand Milinski 1996; Thompson, Kray,and Lind 1998;Smith 2000; Lubelland Scholz2001; and FehrandFischbacher 003.27 See Gould 1981; Gould 2000.28 See MaynardSmith 1982; Axelrod 1984; Brewer2000; Hammond 2000; Fehrand Gachter2002;Rauch2002. See also Putnaml993 and Putnam2000for a real-worlddescriptionof this problem.29 HertwigandTodd 2003. See also GigerenzerandSelten 2001.

30 Elman 1993.31 CosmidesandTooby 1994, 329.32 As positedby Simon 1957; Simon 1985; Simon1997.33 CosmidesandTooby 1994, 327.34 SeePinker1997;Toobyand Cosmides1992; CosmidesandTooby 1992; Frank1988.35 See Chiappe,Brown,and Rodriquez2002. See alsoBaron and Burnstein2002.36 Quoted in Sanderson2001,120; elaborated n Pinker2002.37 See, for example,Putnam 1993.38 Toobyand Cosmides 1992, 37.39 Matt Ridley2003.40 For recentcreativeevidenceinvolvingnonhumanpri-mates,see Brosnanand de Waal2003.41 Buchananand Tullock 1962, 21. See also Selten1999.42 Waltz2001, 29. Fora more positivetakeon the valueof an evolutionaryapproach o understandingwar,seeStrate1983 and 1985, andAlexander1987.43 This is a point that critics such as Waltzseem to miss.As Wrangham 1999) points out, it is a gross"miscon-ception" o believethat "theonly behavioralpat-ternsexplainableby biologyare nstincts, i.e., behaviorsthat areobligatoryand/or inevitable .... [T]hiserrorseems remarkablebecause behavioralecologistshavelong stressed hat psychologicaladaptationsareexpectedto respondin a contingentway to appropri-ate contexts"(p. 21).44 See Olson 1965.45 Proponentsof rationalchoice such as Rabushkaand

Shepsle(1972) admit they are"unable o explain...the preeminenceof ethnicity" p. 64). They do,however,attemptto do so by claimingthis preemi-nence is entirelythe result of elitespushingtheirfol-lowersto view the world throughethnic lenses,acontention that flies in the face of even casualobserva-tion of rank-and-filepeople.Accordingto Varshney(2003), the abilityof rationalchoice to explainwhy somuch conflict is organizedalong ethnic, religious,and linguistic ratherthan economic lines has notimprovedin the decades since Rabushkaand Shepslewrote.46 Hibbing andTheiss-Morse2002.47 Hibbing and Alford2004.48 Froman evolutionarypoint of view,what could beworsethan being a partof a groupin which leadersare bent on personalgain at the expenseof the welfareof othermembers of the group?49 Raby2001 and Pinker2002.50 Sullivan ndMasters1988maybe theclosest o anexcep-tion to this statement.Severalother essayshavecalledfororiginalresearchusingbiologicalandgeneticprin-ciples.Examples ncludeWahlke1979; Masters1993;

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Articles I The Origin f PoliticsSomit and Peterson1996; Ostrom 1998; and Somitand Peterson1999.

51 Approximately98 percentof all human genetic mate-rial is estimatedto be sharedby all membersof thespecies.52 See Eaves,Eysenck,and Martin 1989; Plomin 1990;Plomin, Owen, and McGuffin 1994; Bouchard1998;and Plomin et al. 2000. Adoption studies dealwithcasesin which the varyingportion of the geno-type sharedby siblings approaches ero but theportion of the environmentsharedby them is sub-stantial.These studiescome to the same conclusionas twin studies.For a good summary, ee Plomin et al.2000.

53 See Plomin 1990; Plomin et al. 2000.54 See Bouchardand McGue 2003.55 See Plomin et al. 2000 for a thoroughdiscussionofthe relevantstatisticaltechniques.56 Plomin 1990; Bouchardet al. 1990; and Bouchardand McGue 2003.57 See Martinet al. 1986.58 See Bouchardet al. 1990; Bouchardand McGue2003.59 To explainthis finding, opponentswould need to

arguethatadoptionagenciesare morelikelyto placeMZ twins in similarhomes than they areto placeDZtwins in similarhomes. In fact, informationon twinzygosityis typicallyunavailable o those makingplace-ment decisions. Even if it were, it seems highlyunlikelythat it would factorinto theirdecisions,evenif agenciesweregenerallymore likelyto placetwinsthan nontwin siblingsin similarhomes.60 See Bouchardand McGue 2003.

61 Brelandand Breland1961, 684.62 Conservatism s not unusualin this regard.Rushton,Littlefield,and Lumsden(1986), find that approxi-mately 50 percent of the variancein altruismis theresultof"directgenetic inheritance"p. 7340), withfamilyenvironmentresponsible or 0 percent.63 To takejust one example,a leadingtext on politicalbehaviorstateswithout equivocationthat "attitudes,beliefs,and valuesare learnedpredispositions,and thevariousforms of politicalparticipationarelearnedbehaviors." eeConway2000,61. The twin studiessug-gest this assertion s quite wrong.64 Though clearlythe phrase"political ocialization"s amisnomer incemuchof thecorrelation etween heatti-tudes of parentsand childrenis not due to a social-izationprocessat all.65 See, for example,Jenningsand Niemi 1968; Tedin1974.66 A listingof the heritabilityof all items can be foundin Martin et al. 1986. Our own analysisof twin pairsin the United Statesproducessimilar results.SeeAlford, Funk, and Hibbing 2004.

67 Twinresearchers ave not focuseddirectlyon partyaffil-iation. The closesttheyhave come maybe some unpub-lished researchby Lykken,referencedby Pinker2002. Our expectation s that just as affiliationwith aparticular eligiousdenomination is heavilyenviron-mental,while the tendencyto be religiousat all is heav-ily inherited(Bouchardand McGue2003), particularpolitical identifications, ike partyidentification,aremuch more environmental han the tendencytobe interested n and involvedwith politics in any fash-ion. The tendencyof traits ike religiousaffiliationto not be sharedby MZ twinsanymore than DZ twinsis furtherevidenceagainstthe argumentthat differ-ences in correlationbetween the two typesof twins aredue to environmental actors.If this alternativeexpla-nationwere rue,religiosity houldproduce he samepatternasreligionandthis simplyis not the case.68 Elazar1984.

69 Baron-Cohen2003; see also Baron-Cohen,Tager-Flusberg,and Cohen 2000.70 Baron-Cohen2003 furthersuggests hat the meanplace-ment for males and femaleson such a spectrumis dis-tinct, with males located closerto the autisticpole.This conception could account for the fact that fourout of five individualsclinicallydiagnosedwith autismaremales.

71 Fiorina1996, 85.72 Formore, see Plomin, Owen, and McGuffin 1994.73 See Hamer and Copeland 1998; Ridley 1999.74 At the broadest evel the interactionbetweengenesand the environment includesnot only the naturalenvironment but the man-made environmentas well.

With the recognitionthat culture,a key compo-nent of the manmadeenvironment,can also developalong lines quite similar to evolution and naturalselection,scholarshaverecentlybegunattemptsto con-ceptualizeand empirically tudywhat is commonlytermed"gene-culture oevolution."Foran early appli-cation see Boyd and Richerson1985; for a morerecentapplication,see Fehrand Fischbacher 003.75 Caspiet al. 2002.76 Caspiet al. 2003.77 See Fehr and Fischbacher 003; Boyd et al. 2003.78 Wilson 1975.79 Einsteinsaid, "Withouttheoryobservation s impossi-ble,"to which we would only add that with theoryallmethodsof observationareuseful.

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