Hibbing Reviewed work(s): Source: Perspectives on Politics, Vol. 2, No. 4 (Dec., 2004), pp. 707-723 Published by: American Political Science Association Stable URL: http://www.jstor.org/stable/3688539 . Accessed: 31/10/2011 10:56
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Articles
The
Origin
of
Theory
hy do people hold the political attitudes they do? Despite the centrality of this question, the two guiding theoreticalorientationsof modern political science, rationalchoice and behavioralism,are surprisingly unhelpful.Rationalchoiceis contentto takepreferences as given and is not particularlymotivated to explore their origins or grounding in reality.Behavioralismis based on the untested assertion that preferencescan be understood it by an exclusivefocus on environmentalvariables; largely ignoresthe fundamentalissue of why people respondto the environmentas they do. It is quite common (and accurate) for adherentsof both positions to admit that their favored approachesare not theories at all. Our goal is not to criticizerationalchoice and behavioralism. Indeed, politicalscientistsworkingwithin both theoretical frameworkshave produced impressivefindings and insights, therebyrevealingthe value in identifying influential environmentalfactorsand in outlining the actions that a rationalindividualholding a given preference takes.Nonetheless, the fact remainsthat, as early proponents of rational choice correctlynoted, behavioralismis limited by its lack of a real theory to being a largely inductive mode of research.Rational choice, in turn, as many of its proponents have come to realize,is reaching the limits of what John R. Alfordis associate professor ofpolitical scienceat Rice John R. Hibbing is Foundation (jra@rice.edu). University Regents University Professor PoliticalScienceat the Univerof The sity of Nebraska-Lincoln (jhibbing@unl.edu). authors are gratefulto ChrisLarimer,LeventeLittvay,David Rapkin, KevinSmith,JeffSpinner-Halev, ElizabethTheissreviewers theirhelpffl comments Morse,and anonymous for and suggestions.
can be accomplished deductiveresearch basedon assumed by rather than actual preferences.To fully understand, connect, and transcendthe extant researchrequiresan empiriand of cally sustainabletheory of the source of preferences the reasons people respond as they do to environmental stimuli. Such a theory exists and is being employedwith increasing regularityin the social sciences, especiallyexperimental and economics, behavioralanthropology, socialpsychology. some political scientists are familiar with this Certainly, theory,but it has not been the basis for originalresearchin political science. Yet theories of the origins of preferences (and thereforebehaviors)may offer the ability to unite the individualsocial sciences-and even the social sciencesas a group-with the naturalsciences. Politicalscientistsmay find the theory we describewanting. But it should not be dismissedwithout due consideration of the evidence for and against it. Most important, the theory should not be dismissedbecauseof an unscientific aversionto its implications. The central tenet of this theory is that preferencesand behaviorsareat leastpartiallyshapedby evolutionaryforces and thereforeby genetic heritage.Just as evolutionarypressuresshapedgenes governingthe physicaltraitsof humans, they also shapedgenes governingbehavioraltraits.If this is true, researchin the social sciences will be furthered by taking into account evolutionarilyadvantageousbehaviors and by increased attentionto the connectionbetweenhuman and social behavior.This evolutionary-biological biology approachincorporatesvaluable insights from both behavioralism and rational choice. In fact, the most substantial currentapplicationsof this theory are seen in the two disciplines, psychology and economics, from which political scientists borrow most heavily-and from which we December 2004 i Vol. 2/No. 4 707
Articles
If and rationalchoice, respectively. borrowedbehavioralism a single theoretical approach can make headway in disciplines as disparateas economics and social psychology, it surely merits some attention in a discipline like ours, which has internalizedthe conflict betweenpsychologyand economics. In what follows,we providea generaldescriptionof modern evolutionarytheory and our own more sociopolitically focused behavioral theory, which we call "warycooperation."We then drawout the directimplicationsof this theory for interpretationsof politics and public policy. Finallywe directour attentionto evidencesupportingthe assertionthat is most likely to trouble political scientists: that political orientationsareto a significantextentgeneticallyinfluenced.
labor, or even mere numerousnessitself, groups charactergroup boundaries,but we are also highly sensitiveto selfish ized by cooperativebehaviorat the individuallevel have the actions on the part of other group members. This sensito out-compete and hence dominate groupschartivity leads us to cease cooperatingwhen that cooperation potential acterized by selfish behavior at the individual level. Thus is not reciprocated,to avoid future interactionwith nonindividualsare subject to conflicting selection pressure.At cooperators,and even to engage in personallycostly punthe individual level, pressurefavors entirely self-regarding ishment of individualswho fail to cooperate. individuals may have a better behavior,but self-regarding Perhapsthe easiestway to see warycooperationin action chanceof survival they aremembersof cooperative if is to summarizefindings from the ultimatum game. In this groups. The parametersthat characterize equilibrium of these the widely used experimentalscenariotwo anonymous players will often yield behaviorthat is selfish; arebroughttogetherfor a single interaction.13 One is given conflicting pressures the task of dividing a sum of money between the two of however,under some realisticconditions the resultwill be the evolution of mechanismsfor cooperation.The cooperthem. The other can either accept or reject the proposed ation evident among proteinsin creatingliving cells, among allocation but if it is rejected neither player receives any cells in creating tissues, among tissues in creating organs, money. Of course, from a rationalpoint of view, no player and among organsin creatingcomplex living structures like should ever make a generousoffer to a receivingplayerand no receiving player should ever reject a positive offer. In plants and animals,areall examplesof this balanceof comand cooperationwithin the evolutionaryprocess.12 practice,however,the modal offer is 50 percentof the pot, petition The important insight for human behavior comes and offers of less than one-third are rejectedby receiving from continuing this Rusplayers more than 50 persian doll analogyof compecent of the time, meaning that a remarkably tition within cooperation Humans cooperative, notaltruistic; are but high perto the level of families, excentage of people are willtended kin groups, clans, but so. an competitive, notexclusively Wehave ing to sacrifice their own monetary rewardsto share politicalparties,and nation states-all of which are innate inclination tocooperate, within with another player or to particularly potentiallysubjectto selecpunish an allocator who tion pressure at the indidefined but boundaries, wearealsohighly kept a large portion of the group vidual and collective levels. pot.14The latterpracticeis Environmental conditions sometimes called, for obvito on sensitive selfish actions thepart other of that offersubstantial advanous reasons,costly or altruto clans that coopermembers. istic punishment.15 group tages ate easilyand efficientlywill On the one hand, pure tend to produce at equilibself-interestcannot explain rium an environmentfilled largelywith relativelycooperaeither our willingness to cooperate or our willingness to tive clans. The selfish interest of the clan vis-a-vis other On punish at our own expensein this scenario.16 the other clans thus acts to reduce the selfish behaviorof individuals altruismcannot explainour willingnessto punhand, pure within a clan, but only to the degreethat that reductionin ish noncooperators in multiple-play or, games,to ceasecoopwithin-clanselfishness the survivalchancesof indidoes account improves Warycooperation eratingwith noncooperators. vidualsin cooperativeclansrelativeto less cooperativeclans. for these behaviors. Moreover, unlike short-term selfThus it is not that evolution itself favors cooperation or interestor altruism,our theoryis groundedin an evolutionoperatesin anything but a selfish fashion; it is simply that ary view of human behavior, for it is likely that wary evolution is agnostic about the methods (e.g., competition cooperationis a product of innate and geneticallyheritable or cooperation) by which overall survival advantagesare behavioralpredispositions,and that these predispositions achieved. are the result of identifiableselection pressures.17 What are the evolutionaryunderpinningsof our theory? Evolutionis a slow process,and much of the environmental Our Theory: WaryCooperation pressurefavoringhuman cooperationhas existed for a long New theories of political behaviorcan be built on this curtime. Our genetic composition is to some extent the prodrent and subtle version of Darwinian selection theory. In uct of conditions faced by our hunter-gatherer predecessors that spirit we offer our own theory of "warycooperation" of perhaps 100,000 years ago. One of the keys to an drawn from the work of leading scholars in evolutionary individual'ssurvivalwas being a respectedpart of a viable and experimentaleconomics. The theory may psychology group. The central insight of a behavioraltheory built on be summarizedas follows. Humans arecooperative,but not evolutionary biology is that the desire for group life is a fundamentalhuman preference.What kinds of behaviors altruistic;competitive, but not exclusivelyso. We have an innate inclination to cooperate,particularly within defined optimallypromote belonging to a viable group?These tend
December 2004 I Vol. 2/No. 4 709
in fostervaluedmembership viable Sincethesebehaviors of member a properly andsincebeingan accepted groups, we survival, believethesesix functioning groupfacilitates not will behaviors be common,thoughperhaps universal, endowments. genetic is sinceit is Wary cooperation themostsocialof theories, In this senseit affords another builtaroundotherpeople. to contrastwith existingapproaches. According rational choice(andold schoolevolutionary theory),for example, to other people get in the way.According the theoryof otherpeoplearethe way.But this does warycooperation, not mean that behavioris altruistic.Social behavioris often takento be good and its polaropposite-egocentric with is behavior-bad.Butsocialbehavior notsynonymous is on socialbehavior centered behavior.18 altruistic Rather, with concerned thewelfare otherpeoplebutnot necessarily of otherpeople.In so saying,we are in completeaccord on Monroe's recent Renwick with Kristen emphasis "interand actionswith others,"on "sociability" on "aninnate as need for humanconnection" well as MarilynBrewer's to in- andout-groups.19 relevant accountof behavior in behavior some In additionto expecting cooperative studalso our circumstances, theory expects-and empirical ies haveprovenit to be the case-that peoplemindlessly are conform,20 obeyauthority passively figures,21 competiof in tive to the point of takingpleasure the misfortunes initiatehostilitiestowardthose people in outothers,22 of for construct them,24 out-groups thesake having groups,23 those aboutpunishing enthusiastic andaredisconcertingly stanbehavioral as not perceived living up to the group's The victimized.25 theory when personally dards, especially of codes the of wary predicts construction moral cooperation it valuecooperative that conditionally behavior; does not consistent seemsto be remarkably Thushumanbehavior with the behaviors by expected ourtheory-an issuefuture Here will to scholarship continue address.26 we mustdefend
the claim that the behaviorslisted above are indeed advan710 Perspectives on Politics predict altruisticbehavior.
object accuratelyat something while it and the throwerare both moving); reading facial expressions for honest, as opposed to contrived, emotion; making inferences about plants and animals (as opposed to similarlydistinguishable man-made objects); and identifying others likely to be cheaters-even though thesetasks noncooperators, especially are as, or more, demanding than many we do poorly.34 For when experimentalsubjectsare shown picturesof example, individuals and told their names along with a single fact about them, subjectsare better at rememberingthe names of those who had been connected with a social fact (Sally helped a neighbor paint his house) than a nonsocial fact and (Tom has an old refrigerator), they are best at remembering those who had been connected to a negative social fact (Harry did not return a CD he borrowed from his friend).35These very activities-accurately reading social situations, intuiting the physics of the world around us, perceiving food sources, and especially identifying those who might harm us or our group status-are those that would havebeen quite usefulto our ancestors they roamed as the savannahin small bands. As biologist William Hamilton put it, "The tabulaof human naturewas neverrasa."36 The idea that human nature exists should not be confused with the belief that it is fixed. In fact, humans must be sensitiveto environmental to surroundings achieveobjectives. For example, levels of trusting behaviorvary widely around the world and even within a country.37If natural selection over hundreds of thousands of generations has weeded out the altruistsand the egoists, leaving only wary cooperators,why is cooperative behavior present in such vastly differentproportionsaround the world?The answer is simple but important:wary cooperation is a conditional behavior and thus fundamentallydifferent from strategies of "alwaysact in your own interest"or "alwaysact in the interestof the other person."The warycooperatoractswith a glance over the shoulder and adjustsbehavioras needed. Thus the behaviorof a wary cooperatorin Lombardywill be quite unlike that of the same genotype transplantedto Sicily, merely because these two individuals are likely to encounter differentlevels of cooperativebehavior. Most contributions of rational choice and behavioralist are scholarship in no way compromisedbut ratherenhanced the acceptanceof evolutionarytheories.The previously by content-free "preference structures" rational choice can of find a sourceof clearand potentiallyfalsifiable content, and the "incoherentenvironmentalism"38 behavioralismcan of be giventheoretical coherence. The messageis not thatnature trumps nurturebut ratherthat nurtureis in our natureand that the precise nature of our nurturing tendencies Our theorysubdependsupon environmentalconditions.39 sumes a wide range of behavior-as any successful social theory must. Work that takes evolution seriously opens a broad rangeof cooperativeopportunitiesacrossmethodologies and approachesthat in the past have createdlines of conflict within political science;it drawson and encourages
more theory-driven researchin fields as diverse as game theory, traditional field work in anthropology, and economic and social psychological experiments,with worldwide human (and nonhuman) subjects.40
Articles
luxury in the Caribbeanfor what we would otherwise pay to bring him to justice for his noncompliant behavior? In this and related policy matters, we see clear evidence that people are willing to support and engage in costly punishment-and also to comply themselves-as long as they remain confident that noncontributorsare routinely punished. The perception of a general failure to punish others, rather than the perception that we are personally unlikely to be punished, is the proximate cause of noncompliance. War The phrase"millionsfor defense, not a penny for tribute," popularized in the United States during the days of the Barbary piratesbut just as relevantto today'smood (if a few zeroesareadded), nicely capturesthe role warycooperation has played in the history of internationalaffairs.The fact that wary cooperation helps account for humans' innate capacityto cooperateon a large social scale is by no means equivalentto a notion that the social side of human nature is benign in its aims and impacts. Perhapsthe most striking of example of this is war.The most apparentcharacteristic war at the group level is conflict, but we should not miss the fact that at the individual level warfareis among the of most cooperativeand self-sacrificing all human ventures. In no other arena are humans as likely to set aside their personalhealth, well-being, comfort, and even life itself, in favor of group success. Selfless participation in warfare remains, even in modern societies, at the center of our notions of heroismand ennoblingactions.While the savagery of war clearlyderivesfrom innate human features,it is clear that the explanation of group-levelsavageryis not simply an aggregation of individual-level aggression. While the importanceof cooperationfor peace may seem self-evident, cooperation is no less critical to war.This seeming contradiction is troubling to many. As Kenneth Waltz notes, for example, "While human nature no doubt plays a role in bringingabout war,it cannot by itself explainboth war and peace. War is, of course, only an extremeexampleof the apparent contradictionwithin an explanationfor both cooperation and competition that focuses on human nature. In some ways, avoiding this contradictionmay be the greatest value of the wary cooperation theory. Though our theory posits a simple baseline explanation of human nature, it does not assume that human nature must be either competitive or cooperative;rather,it explainsone source of the relative importance of competitive and cooperativedrives within a given behavioralcontext.43Similarly,it freescompetition from a necessarytie to self-interestjust as it frees cooperation from any necessaryassociationwith altruism. We do not have a gene for combativenessany more than we have a gene for cooperation, or indeed a gene for wary cooperation. What we have is a set of innate behavioral 712 Perspectives on Politics
than common-good reasons.46 Governmentis typicallydisliked not becauseit is perceivedto do the wrong things, but because it is perceivedto do things for the wrong reasons. Ascribed motive is the main determinant of responsesto the actions of others in any societal interaction, but it becomes acutely importantwhen we are dealing with people possessingsubstantialclout.47In this sense, the theory of wary cooperationholds that those staffingpolitical institutions should be subject to special vigilance not because they arefundamentallydifferentlife formsbut becausetheir ambition for authority raisessuspicions, and their possession of authoritymakes them potentially dangerous.48 What does the theory say about the specific manner in which politicalinstitutionsshould be constructed? startFor ers, it points out the prescienceof the architectsof the U.S. political system in makingit difficultfor decision makersto furthertheir own ends. Separationof powersacrossinstituand tions, frequentelections, federalarrangements, a strong judiciary-all facilitatethe balancingof one group'spowers against another's,which is, not coincidentally, extremely popularwith the mass public. This does not mean, however,that the Americanpolitical system is a perfect fit with the institutional preferences flowing from the theoryof warycooperation.In the people's view, far too many opportunitiesstill exist for elected officials to feathertheir own nests. Think of the ire evoked by mattersrelatingto congressionalsalary,pension plans, and perquisites,not to mention the perceivedunholy allianceof well-heeled interest groups and politicians. Wary cooperators like to cooperate, but they are also predisposedto be waryof the actionsof others-especially those eagerto make decisions for the group. Reformerswould do well to realize that people do not wish to be in control of the political system;they only want those who arein controlto be unable to take advantageof their positions. If people were confident that existingconstraintsprohibitedsuch self-interested actions, they would pay even less attention to the political arena than they do now. For most people, involvement in politics is driven not by a desireto be heardbut by a desire to limit the power of others. Current American foreign policy might be improved, for example, if decision makers realized that, like Americans, people in Afghanistan and Iraqdo not cravedemocraticprocedures.Kurdssimply do not want to be dominated by Sunnis; Sunnis do not want to be dominated by Shiites; Uzbekis by Tajiks;and Tajiks by Pashtuns.People often expressa desirefor participatory democracywhen they reallyjust want to avoid being victimized by a more powerfulgroup.
alenceof warycooperatorsis the resultof evolutionarypressuresthat rewardhumans skilled at creatingand existing in viable social units. Our theory is only one of many possible theoriesthat takeseriouslythe connection betweenbiology, naturalselection, and human behavior.Evidence supporton ing this connection can be found in the literature genetics and human behaviorthat has flourished,as new fieldsof inquiry often do, with the acceptanceof a new methodology. In this section we explore one such methodologytwin studies. Charles Darwin and Alfred Russel Wallaceworked out the detailsof naturalselection at roughlythe same time and were in remarkableagreement-with one vital exception. Darwin was completelyconsistent and contended that naturalselection appliedto behavioralas well as physicaltraits. Wallace,on the other hand, drew a bold line between the two, positing that the mental realmwas immune to evolution and was insteadthe purviewof etherealreligiousuncertainties.Wallace's position is still favoredby groupsas diverse as religiousfundamentalistsand many in the social science community who refuse to believe that behavior is innate. The naturalsciences have sided almost unanimouslywith Darwin.Wallace spentthe lastyearsof his life largely ignored, a religiousmystic who thought it possible to communicate with the dead and who tried desperatelyto hold on to the fictionthatour brains,behaviors, beingsconstitutesomeand thing more than just another step along the evolutionary path.49 If political scientists believe, with Darwin, that genetics influencessocial attitudesand behavior,it is not evident in their research.Certainlyno recentarticlein a leadingpolitical science journal has used genetics as an independent variable.50 Perhapsmost believe that, while genetics may some nebulous role in politics, environmentalcomplay overwhelmgenetic plexitiesand humancognitivecapabilities effects, allowing the discipline'sresearchagenda to completely ignore biology. Contraryto popularbelief, however, the link betweengenes and political attitudesand behaviors is strong. In this section we hope to convince readersthat genetic variablesare more influentialthan is generallyconceded by political scientists.To do so we summarizerecent research twins, on autism,and on genetic-environmental on interactions.
Twins Monozygotic (MZ) twins have virtually identical genetic codes;dizygotic (DZ) twins, like other full siblings,shareas little as 50 percent of the genetic material that varies in human beings.51 This makes the study of the adult characteristicsof these two types of twins of specialvalue in computing the relativeinfluenceof heredityand environmentespeciallywhen augmentedby a comparisonof twins reared separatelyand those rearedtogether.Such studies typically identify a substantial amount of genetic influence and a December 2004 1Vol. 2/No. 4 713
extraefforts to help the twins living together establishdistinct identities. In addition, as adult MZ twins living apart a age, they tend to become more, not less, similar,60 finding difficult to reconcilewith the belief that only the environment matters. Interestingly,this precise effect is predicted in an early landmark criticism of behavioralismand the on research animalbehaviorat its core. conditioned-response Over time, substantial anomalies accumulated in this research,pointing towarda primacyfor some nonenvironmental behaviors. As Keller and Marian Breland summarized this tendency, "Learnedbehavior drifts toward instinctive behavior."61 While the twin-studymethodology has yet to be applied by political scientists, a varietyof social psychologystudies use this methodology to assessconservatismas a personality trait. The measureof most interest to political scientists is Attitude Inventory.This inventoryis the Wilson-Patterson administeredby presentingsubjectswith a short stimulus term, such as death penalty,and eliciting a simple response of agree,disagree,or uncertain.The broadestversionof the W-P inventory includes 50 items, half of which contribute positively to the conservatismscore and half of which conWhile some of the items relateheavilyto tribute negatively. of social conservatism-for example, pajama conceptions parties, casual living, nudist camps, computer music, and horoscopes-others are more directlypolitical-for example disarmament,legalizedabortion, socialism, patriotism, and death penalty. Studies typicallyuse reducedsets of W-P items or tailor individual items to the country in which the studies are being administered.For political scientists, this is frustrating on two counts. The list of politically relevantitems is tantalizingbut limited and unfocused;moreover,the results areoften presentedonly for the entirecombined scale,making it difficultto assessthe contributionof the directlypolitical items to the overall index of heritability.Heritability estimatesfor overallconservatismas measuredby the W-P inventory are quite high, usually around 0.60. Furthermore, the models typicallyshow little or no effectfor shared environment (the rest is likely the resultof nonsharedenvironmental factors, including the prenatal environment). These findings come from studies of twins in settings as disparateas Australia,Virginia, and Minnesota. The Minnesota findings, based on twins rearedapart,conform well to the other studies, based on twins raisedtogether.62 Our concern at this point is not to quibble over whether in fact genetics accounts for more than half of the variation in individuallevels of conservatism. The importantpoint is that our discipline currentlybelieves that the state of the worldwould yield an environmental com(i.e., socialization) ponent greaterthan 50 percentand a genetic component of Whateverthe precisenumbers,the duraessentiallyzero.63 ble and starkresultsof twin studies are completely at odds with current thinking in our discipline. At the very least, political science can no longer dismiss genetics as some
minor or distant precursorto important political orientations. Taken at full face value, these results suggest that virtuallyeverythingwe have supposed to be true about the originsof broad political orientationsis false. As strong as this assertionmay seem, it does not indicate that most of the extant empiricalresultsconcerningpolitical socializationare incorrect,only that the interpretations commonly applied to those resultsare incorrect.In fact, it strongly reinforces the most salient empirical findings.64 We know, for example,that if both parentssharea political orientation and find politics to be important, their offspring will likely have that same political orientation and also valuepolitics.65Nothing in this descriptioncontradicts the twin-based contention that political orientations are largelythe resultof genetic inheritance.In fact, this is precisely the patternwe would expect to find if the twin studis ies are correct.The challengeof the twin research only to our assumption that, unlike shared physical traits, shared traitsmust be purelya function ofenculturation. behavioral At this point, the carefulreaderis undoubtedlythinking that we are getting ahead of ourselves.As we mentioned earlier,the 50-item W-P inventorycontains both items that we would recognizeas having substantivepolitical content and others about which we would entertain substantial doubt. If the overalldegreeof heritabilityis largelya function of attitudestowardmodern art and pajamaparties,the direct relevanceto political science may be minimal. What then do we find in the limited studies that reportheritability for individual items?A 1986 study by Martin and colleagues of over 3,800 Australian and British twin pairs reportedthe following estimatesof heritability(on a scale of 0 to 1.0) for the following items: death penalty, 0.51; white superiority,0.40; royalty,0.44; apartheid,0.43; disarmament,0.38; censorship,0.41. The heritabilityestimate for pajamaparties,on the other hand, was a mere0.08. The comparableestimates for the influence of sharedenvironment were: death penalty, 0.00; white superiority,0.09; royalty,0.14; apartheid,0.05; disarmament,0.00; censorship, 0.03 (but pajamaparties,0.44).66 These findings are powerfulyet incomplete. As political scientists,we could all provideour own list of items that we would want to see included in this sort of inventory.Party identification,for example, is probablyan obvious item.67 In any case, the implicationfor political science is clear:we must take seriouslythe possibilitythat at least some part of what we have attributedto activesocializationis a function of genetic inheritance.Equallyimportant-and even more urgent-is the need to adapt twin-study methodology to our specific disciplinaryresearchagendas. We invitedubiousreaders indulgeus in a thoughtexperto iment. Suspenddisbeliefand think seriouslyaboutwhat this might mean for our entire conception of political attitudes and orientations.Daniel Elazarcharacterized American the South as having what he called a traditionalistorientation toward politics, and argued further that these presumably December 2004 1Vol. 2/No. 4 715
that differencesin methodology within and across disciplines may deriveat least in part from heritabledifferences in brain physiology.When Morris Fiorinawrites that he is choice down to [his] DNA," he could be literally "rational correct.71 Likewise,seeminglyincompatibleconceptions of basic human nature may in fact be equally correct when applied to the propersubset of the population:fundamental behavioral differences may also derive from heritable differencesin brain physiology. Genes For a varietyof reasons,skepticism persistsregardingestimates ofheritability derivedfrom twin studies. But the case for genetics playing a role in shaping behavioris based on much more than twin research.Since much of the resistance among social scientists to genetic independent variables stems from unfamiliaritywith modern biology, it is essential that we offer some indication of the manner in which genetics connects to behavior. The most interesting and numerous genes in human beings are not structural(blue eyes or brown), but regulatory. Regulatorgenes allow an organism to respond to its environment; they are the genes that turn on and off the transcriptionof other genes (or themselves).They are the body's thermostats. Consider, for example, biological responsesto fearand threat.The amygdala,a primitivepart of the lower brain, sounds an alarm, adrenalineand stress hormones are released,extra blood is sent to the muscles, breathingquickens,glucose is disgorgedby the liver,blood pressureskyrockets,and nonessentialbodily functions stop. While this generalpatternis alwaysin evidence, sensitivity, rapidity,and depth of responsevary widely. Some people experiencea full-fledgedpanic attack at the slightest provocation; others displayquite muted physiologicalreactions to even life-threateningevents.Why?Just as people display physical differencespartly because of differencesin structural genes, they display behavioraldifferencespartly because of difference in regulator genes. People's regulator genes encourage them to react to environmental conditions in certain ways, whether those conditions involve danger, lactose, alcohol, an uncooperative individual, or sexual stimulation. These on-off switches shape much of our personalities.72 Regulator genes are not merely theoretical constructs. They have been seen, and behaviorhas been accurately predicted merely by knowing the shape of an individual'skey regulatorgenes. Consider a gene called D4DR, which can be found on the short arm of chromosome 11. This gene influences the reception of dopamine, a chemical needed by portions of the brain. Individualswith a long version of D4DR areless efficientat collectingdopamine,so they need to take more risksand seek more thrillsin orderto generate as much dopamine as others acquirefrom more sedentary lifestyles.73
None of this is to say that environmental factors are In irrelevant.74 fact, surprisingly,and perhaps for many, currentbiological thinking unites genetic and reassuringly, environmentalinfluences in a complicatedbut compelling interactivefashion. For example, recent work on violence calledmonohas found a connection to a blood transmitter amine oxidase A (MAOA), but the connection is not simply that those with genetic deficienciesin MAOA production are more violent. Rather, those individuals who have low MAOA production and experiencedviolence as children are much more likely to be violent as adults.75 Acting alone, MAOA deficiencies or a violent childhood have little predictivepower, but the interactionof genetic and environmental factors is disconcertinglypowerful. A similar pattern has been found in studies attempting to predictthose individualslikelyto sufferfrom clinicaldepression. There is a gene on chromosome 17 called 5-HTT. This gene also comes in a long form and a short form. An extensive study of nearly 1,000 New Zealanders found that clinical depression was associated not just with the presence of the short form of 5-HTT and not just with the presence of many high-stressevents in a subject'slife, but with a combination of high-stressevents and the shortform of 5-HTT.76 If we generalizethis patternto predispositionsmore central to political concerns,such as conservatismand cooperation, it suggeststhat the eventualgenetic-levelexplanation may fit surprisinglywell with much of what we already know and much of what we havesuggestedherewith regard to genetics and evolution. Perhapsfor a substantialportion of the population the short form of a yet-to-be-identified to gene allows their degree of cooperation/retaliation vary with regardboth to the specificcurrentcontext and to their broaderexperienceover time. We might call these individuals wary cooperators.For another portion of the population, those with the long form of the gene, context seems to make little difference.While we believe our theory of wary describesthe behavcooperationdiscussedearlieraccurately ior of the majorityof the population, it needs to and does leave room for a minority that behaves in a predictably differentfashion.This minoritymay be fixedin eithercooperativeor noncooperativemodes, or may alternatebetween these modes with no apparentregardfor the cooperationor lack of cooperation evidenced by those in their environment. This could provide a rough genetic sketch of the biologicalunderpinningsfor the starklydifferentsocialabilities and orientations discussed above in the section on autism. And that discussion in turn providesthe cognitive underpinningsfor the theory of wary cooperationthat was the startingpoint for this piece. Interestingly, computer the simulationsthat we discussedabovehavedemonstratedthat (under reasonableassumptions)a population consisting of two types roughly compatiblewith mildly autistic individuals and wary cooperators,respectively, can reach a stable with the larger part of the final population equilibrium December 2004 1Vol. 2/No. 4 717
Notes
1 Huxley 1989. For an intriguing counterpoint to Huxley, see Kropotkin 1972. 2 Darwin 1968; Darwin 1981. 3 Simon 1957; Simon 1985; Simon 1997. 4 Cosmides and Tooby 1994, 327. 5 Ostrom 1998, 2. See also Jones 2001. 6 Dawkins 1976, 2-4. 7 Ibid., 66. At a later point, Dawkins clearlyrecognizes that genes occasionallyengage in cooperativebehavior. 8 Buchananand Tullock 1962, 18. 9 See, for example, Dawkins 1976; Dawkins 1982. 10 Such "selfish" genes have been used to explain the selfdestructivebehaviorof organismssuch as salmon, which leavethe comfortsof the ocean to swim upstream, spawn, and die, and male mantises,which sometimes celebrate end of copulationby throwingthemthe selves back into the open, waiting jaws of the much largerfemales. 11 For a descriptionof the biology of cooperationat this level, see Mark Ridley 2003. 12 Frank 1988; Sober and Wilson 1998. The claim that selection pressurescan be profitablyviewed at levels other than the gene, while increasinglyacceptedby students of evolutionarypsychology,has had a checkered past, and many scholars remain unpersuaded. 1962 account of rooks volunWynne-Edwards's tarily curtailingtheir clutch size for the welfareof the flock was roundly criticizedby the biological community and gave relatednotions much to overcome. enviso-calledgroupselectionists Moreover, occasionally sioned an ethereal,almost inexplicable,and somewhat magicaldevotion to the group. As emphasizedin our descriptionabove, mysticism is not needed, and modern thinking is firmlyembeddedin the logical consequencesof selection pressuresat differentlevels. The switch in common phraseologyfrom group selection to multilevel selection is more than semantic. In addition, continuing advancesin genetics cast more and more doubt on genes as stand-alone, clearly distinct actors in the evolutionarydrama.Instead, we now know from Gibbs 2003 and others that many traitsare epigenetic, that is, influenced by chemical marksoutside the DNA sequence.Today,much of the dispute centers on the relativeimportanceof multilevel selection ratherthan the fact of its existence. For some sense of the opposing arguments, see the and Brain exchangein the 1994 issue of Behavioral in Sciences, which Wilson and Sober make the case for multilevel selection, and which is commented on by numerous scholars.See also Sober and Wilson 1998. 13 For a good summary,see Guth and Tietz 1990. 14 See Nowak, Page, and Sigmund 2000. Lest it be is of thoughtthatthe generosity allocators merelystrategic, when the game is changed to a dictatorgame, in
Conclusion
We began by noting the usefulnessto the social sciences of greaterattention to biological evolution. Part of the problem may be that the last time biology came to the attention of political scientists (in the 1970s, afterthe publication of E. O. Wilson's Sociobiology), they believed that advocates were saying that behaviorwas determined by biology.78If that was ever the position of biology proponents, it is no longer. A researchprogram focused on genetic factors to the exclusion of environmental factors would be deeply misguided-perhaps as misguided as a researchprogram factorsto the exclusionof genetics, focusedon environmental We believethat modernbiologicaltheory,properlyunderstood, holdsthe promiseof unifyingpoliticalscienceandintegratingpoliticalsciencewith othersocialsciences-and even with the naturalsciences. Concepts borrowedfrom evolutionary biology can provide the theoreticalguidance both need to becomeeven more rationalchoice and behavioralism usefulthan they have been to date. Scholarsoperatingin the traditioncan certainlycontinueto identifyaspects behavioral of the environment that seem more influentialthan others and the conditions underwhich these environmentalinfluences vary.Though we are convinced that it is necessaryto would be useunderstand originalcauses,this understanding did less if behavioralists not link these causesto more proxin Scholars imateenvironmental factors. operating the rational choice tradition are similarlyadvantagedby the incorporacan tion of biologicaltheory,sincepreferences be derivedfrom an understandingof evolutionarypressuresacting on individuals and groups. Baselinesof behaviorno longer have to be questionableassumptionsthat people pursueshort-term tangible gains or largelytautologicalassumptionsthat people must preferwhateverthey pursue. We are not suggestingthat all political scientistsbecome evolutionary theorists or adopt a certain methodological approach.We need a healthy disciplinarydivision of labor in serviceof a genuine theory of political and social behavior. To date in the social science disciplines that have embracedevolutionarytheory,everythingfrom participantobservation to pure mathematicalgame theory to laboratory (and field) experimentshas been employedwith fruitful results.As the old saw goes, "Ifyou are looking in the right place thereis no wrong way to look."79Evolutionarytheory has the potential to render obsolete our intradisciplinary conflicts over approach, method, and theory. It is more than a replacementfor existing political science theories, are since rationalchoice and behavioralism simply not capable of supplying an account of ultimate causes. It is a true theory of the origins of behavior and as such provides a diverseand usebasis for bringing together the remarkably ful ongoing researchin political science and beyond. 718 Perspectives on Politics
15 16
17
18 19 20 21 22 23 24 25 26
27 28
29
which the receivingplayerhas no chanceto veto the prostill posed allocation,anonymousallocators makegenerous offers, albeit typicallyless generous than in ultimatum games. See Fehr and Fischbacher 2003 for a useful summary.See Van Dijk and Vermunt2000 for researchsuggestingplayersin the dictatorgame are actuallymore generous than those in the ultimatum game. Boyd and Richerson 1992; Gintis 2000; Henrich and Boyd 2001; Price, Cosmides, and Tooby 2002. Thus the discipline's theoreticalbasisfor explainingselfinterestedand non-self-interestedbehavioris much as Ostrom, Gardner,and Walker(1994) describeit with regardto the specific instance of abusing or not abusing common pool resources. "No existing theory providesa consistent explanationfor how and extricatethemselvesfrom why many appropriators common pool resourcedilemmas [and] why this is not the universally case"(p. 18). The theoryof warycooperation, in contrast,does explain these puzzles. The main reasonfor opening up social science to the role of biology is not that it can better explain variations of human behavior around the mean but rather that it can help us to better understandthe mean itself. We do not, however,believe all individualspossess in the same degree the biological materialsthat In behavior. fact,as dispush us towardwarycooperative cussed below, we believe that a minority of humans are geneticallypredisposedtowardinsensitivityto the cooperationlevels of people around them. For a thorough treatmentof altruisticbehavior,see Monroe 1996. Monroe 2002, 203; Brewer2000. Asch 1951. Milgram 1974. Frank 1999. Sherifet al. 1961. Barash1994. Price, Cosmides, and Tooby 2002. Extant researchroughly consistent with our theme includes Kahnemanand Tversky 1984; Kahneman, Knetsch,andThaler1986; Orbelland Dawes 1991;Thaler 1992; Kahn and Murnighan 1993; Komorita and Parks1996; Ostrom, Gardner,and Walker 1994; Hoffman, McCabe, and Smith 1996; Wedekind and Milinski 1996; Thompson, Kray,and Lind 1998; Smith 2000; Lubell and Scholz 2001; and Fehr and Fischbacher2003. See Gould 1981; Gould 2000. See MaynardSmith 1982; Axelrod 1984; Brewer 2000; Hammond 2000; Fehr and Gachter2002; Rauch 2002. See also Putnaml993 and Putnam 2000 for a real-worlddescriptionof this problem. Hertwig and Todd 2003. See also Gigerenzerand Selten 2001.
30 Elman 1993. 31 Cosmides and Tooby 1994, 329. 32 As posited by Simon 1957; Simon 1985; Simon 1997. 33 Cosmides and Tooby 1994, 327. 34 See Pinker 1997; Tooby and Cosmides 1992; Cosmides and Tooby 1992; Frank 1988. 35 See Chiappe, Brown, and Rodriquez2002. See also Baron and Burnstein2002. 36 Quoted in Sanderson2001,120; elaboratedin Pinker 2002. 37 See, for example, Putnam 1993. 38 Tooby and Cosmides 1992, 37. 39 Matt Ridley 2003. 40 For recent creativeevidence involving nonhuman primates, see Brosnanand de Waal 2003. 41 Buchananand Tullock 1962, 21. See also Selten 1999. 42 Waltz 2001, 29. For a more positive take on the value of an evolutionaryapproachto understandingwar, see Strate 1983 and 1985, and Alexander 1987. 43 This is a point that critics such as Waltz seem to miss. As Wrangham(1999) points out, it is a gross "misconception"to believe that "theonly behavioralpatternsexplainableby biology areinstincts, i.e., behaviors that areobligatoryand/or inevitable .... [T]his error seems remarkablebecause behavioralecologists have long stressedthat psychologicaladaptationsare expected to respond in a contingent way to appropriate contexts"(p. 21). 44 See Olson 1965. 45 Proponentsof rationalchoice such as Rabushkaand Shepsle (1972) admit they are "unableto explain ... the preeminenceof ethnicity"(p. 64). They do, however,attempt to do so by claiming this preeminence is entirely the result of elites pushing their followers to view the world through ethnic lenses, a contention that flies in the face of even casualobservation of rank-and-filepeople. Accordingto Varshney (2003), the ability of rationalchoice to explainwhy so much conflict is organizedalong ethnic, religious, and linguistic rather than economic lines has not improved in the decades since Rabushkaand Shepsle wrote. 46 Hibbing and Theiss-Morse2002. 47 Hibbing and Alford 2004. 48 From an evolutionarypoint of view, what could be worse than being a part of a group in which leaders are bent on personalgain at the expense of the welfare of other membersof the group? 49 Raby 2001 and Pinker2002. 50 Sullivanand Masters1988 maybe the closestto an exception to this statement. Severalother essayshave called for originalresearch usingbiologicaland geneticprinciples. Examplesinclude Wahlke 1979; Masters 1993; December 2004 1Vol. 2/No. 4 719
51
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53 54 55 56 57 58 59
60 61 62
63
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