The Evolution Of Cooperation Robert Axelrod; William D .

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The Evolution of CooperationRobert Axelrod; William D. HamiltonScience, New Series, Vol. 211, No. 4489. (Mar. 27, 1981), pp. 1390-1396.Stable URL:http://links.jstor.org/sici?sici C%3E2.0.CO%3B2-6Science is currently published by American Association for the Advancement of Science.Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available athttp://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtainedprior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content inthe JSTOR archive only for your personal, non-commercial use.Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained athttp://www.jstor.org/journals/aaas.html.Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printedpage of such transmission.The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academicjournals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers,and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community takeadvantage of advances in technology. For more information regarding JSTOR, please contact support@jstor.org.http://www.jstor.orgFri Jan 4 15:02:00 2008

The latest data for 1978 suggests that the situation may, in fact, be deteriorating. . . . we maybe losing the war on air pollution."3. For examples see House Subcommittee on theEnvironment and the Atmosphere, The Erzvironmentcrl Protection Agerzcy's Resecrrch Progrcrmwith Prirnury F,'rnphusis on the Communitylfecrlth und Environmenrul Surveillance Syatem(CHESS):An lnveatigative Report (GovernmentPrinting Office, Washington, D.C., 1976). especially chapters 4 to 6.4. Without trying to be entirely rigorous, we willuse an NSF definition: "Basic research is thattype of research which is directed toward increase of knowledge in science. It is researchwhere the primary aim of the investigator is afuller knowledge or understanding of the subjectunder study, rather than a practical applicationthereof." This was given by A. T. Waterman,then director of NSF, in Symposium on RuaicReaeurch, D. Wolfle, Ed. (American Association for the Advancement of Science, Washington, D.C., 1959). p. 20.5. For example, the EPA administrator, D. Costle,in a letter dated 12 June 1978 to Senator WilliamProxmire, chairman of the HUD-lndependentAgencies Subcommittee of the Senate Appropriatrons Committee, said concerning enviionmental research, "I've had to make too many billiondollar decisions over the last year without thecritical information this sort of investment,made five years ago, would have provided."6. U.S. Congress, Office of Technology Assessment, A Review i f the U . S . F,'nvironmrntulProtection Agency Environmentcrl ResecrrchOutlook FY I976 through 1980 (GovernmentPrinting Office, Washington, D.C., 1976).7. National Academy of Sciences, Commission onNatural Resources, Ar ul),ticcrlStudies for theU . S . Environmentul Protection Agency, vol. 3,Research and Development in the Environmentul Protection Agency (National Academy ofSciences, Washington, D.C., 1977).8. National Advisory Committee on Oceans andAtmosphere, A Report lo the President crnd theCongress, fifth annual report (GovernmentPrinting Ofice, Washington, D.C., 1976).9. ORD Progrcrm Guide (EPA-60019-79-038, Environmental Protection Agency, Washington,D.C. 1979).10. Kesecrrch r r t l o o k 1980,(EPA-60019-80-006, Environmental Protection Agency, Washington,D.C., 1980). This presents the agency's 5-yearenvironmental research plan in response to statutory requirement. The plan is updated and anew report issued annually.11. Many examples of the environmental problemsEPA faces are reported in Environmentcrl Oullook 1980 (EPA 60018-80-003, EnvironmentalProtection Agency, Washington, D.C., 1980).12. At the beginning of the 94th Congress (Janu-13.14.15.16.17.ary 1975) the Committee on Science and Technology received jurisdiction over envlronmental research" as a result of several changesin the rules of the House of Representatives.The Subcommittee on the Environment and theAtmosphere was formed to handle this jurisdiction and for 4 years (two Congresses), withCongressman Brown as chairman, had responsibility for ORD. In January 1979, as a result of reorganization within the Committee onScience and Technology, Congressman Brownmoved to the chair of the Subcommittee onScience, Research, and Technology. The Subcommittee on the Environment and Atmospherewas renamed Subcommittee on Natural Resources and Environment and given some additional jurisdiction.Eni ironmrnterlProtection Agency Reseurch undDevelopment laarres: 1978, hearings before theHouse Subcommittee on the Environment andthe Atmosphere, 19 July and 13 and 14 September 1978 (Government Printing Office, Washington, D.C., 1979).In making funding decisions, the agency uses azero-base budgeting (ZBB) process in whichprograms are approved by a consensusof theadministrator and the six assistant administrators. In the ZBB process, the ORD has onlyabout one of six votes, and thus research programs are vulnerable to a great deal of influencefrom the program offices. Because they playsuch a substantial role in defining the program ofresearch ultimately conducted by ORD, theadministrator and all assistant administratorswere asked to testify on what they expect fromthat office.Speciul Urban Air Pollution Problema: Denverund Houaton, hearings before the House Subcommittee on the Environment and the Atmosphere, 19 and 21 November 1977 (GovernmentPrinting Office, Washington, D.C., 1978).Long-Term Environmental Research in the Eni ironmenterlProtection Agency, hearings beforethe House Subcommittee on the Environmentand the Atmosphere, 30 June 1977 (GovernmentPrinting Office, Washington, D.C., 1978). Seethe testimony of R. L . Sansom, especially hissupplemental statement, p. 52.H. Kissinger, The Reporter, 5 March 1959, p.10.For example, the agency has instituted a newsystem of research grants putatively aimed atbringing new work of high quality into its program. Despite this aim the published solicitationfor grant proposals does not explicitly andunambiguously state that funding decisions willbe based on scientific quality. Instead the following appears: "Scientific merit and relevanceof proposals will be significant and balancedfactors in the evaluation procedures since allprojects must be in concert with the Agency's-18. The Evolution of CooperationRobert Axelrod and William D. HamiltonThe theory of evolution is based on thestruggle for life and the survival of thefittest. Yet cooperation is common between members of the same species andeven between members of different species. Before about 1960, accounts of theevolutionary process largely dismissedcooperative phenomena as not requiringspecial attention. This position followedfrom a misreading of theory that assigned most adaptation to selection at1390the level of populations or whole species. As a result of such misreading,cooperation was always consideredadaptive. Recent reviews of the evolutionary process, however, have shownno sound basis for a pervasive groupbenefit view of selection; at the level of aspecies or a population, the processes ofselection are weak. The original individualistic emphasis of Darwin's theory ismore valid (1, 2).0036-807518110327-1390 01 .SO10 Copyright O 1981 AAASbudget appropriations." In other words, itseems that work on highly relevant mattersmight be funded even if of poor quality. [SeeEPA and the Academic Community (EPA-6001880-010, Environmental Protection Agency, Cincinnati, Ohio, 19801, p. 2.119. National Academy of Sciences, Materials Advisory Board, Repor1 of the Ad Hoc C o m m i t t eonPrinciples i?fReseurch-Engineerinx Interuc'tion(National Academy of Sciences, Washington,D.C., 1966), p. 16.20. W. 0 . Baker, in House Committee on Scienceand Technology, Serninur on Reaeurch, Productivity, und the Natiorzul Economy, 18 Jrrne I980(Government Printing Office, Washington,D.C., 1980).21. Testimony of J. N. Pitts, in 1980 Aulhorizcltion,for the Office o f Resecrrch und Developnzenl,Eni ironmrntulProtection Ayency, hearings before the House Subcommittee on Science andTechnology, 13 and 15 February 1979 (Government Printing Office, Washington, D.C., 1979).22. H. W. Bode, in nusic Resecrrch und NrrtioncrlGorrls, a report to the House Committee onScience and Astronautics (National Academy ofSciences, Washington, D.C., 1965), p. 74.23. At present the program offices guide EPA'sresearch not only through the ZBB process butalso through the mechanism of 13 research committees. These committees translate programofice needs into "research strategy documents" which guide all EPA research (10).24. This provision is contained in section 6 of PublicLaw 95-155, the FY 1978 authorization act forORD. For explanation of congressional intentsee Conference Report to Accompcrny H.R.5101, 95th Congress, Report No. 95-722 (Government Printing Office, Washington, D.C.,1977). ,.25. This provision is contained in section 11 ofPublic Law 95-155. For explanation see thereport cited in (10), and also Report lo Acc'ompuny H . R . 5101, 95th Congress, Report No. 95157 (Government Printing Office, Washington,D.C. 1977).26. This was cbntained in section 4(a) of H.R. 7099,the House version of the FY 1981 authorizationbill. The provision was deleted from the finalversion of the bill at least in part because theagency strenuously (if informally) opposed itand succeeded in having it removed from theSenate-passed version of the bill. For cxplanation of intent, see Report to Accompan) H . K .7099, 96th Congress, Report No. 96-959 (Government Printing Office, Washington, D.C.,1980).27. J. Bronowski, The Common Senae of Science(Harvard Univ. Press, Cambridge, Mass., 19781,p. 143.28. We thank A. V. Applegate for substantial assistance in the preparation of this paper.To account for the manifest existenceof cooperation and related group behavior, such as altruism and restraint incompetition, evolutionary theory has recently acquired two kinds of extension.These extensions are, broadly, geneticalkinship theory (3) and reciprocation theory (4, 5). Most of the recent activity,both in field work and in further developments of theory, has been on the side ofkinship. Formal approaches have varied,but kinship theory has increasingly takena gene's-eye view of natural selection(6). A gene, in effect, looks beyond itsmortal bearer to interests of the potentially immortal set of its replicas existingin other related individuals. If interactants are sufficiently closely related, alDr. Axelrod is a professor of political science andresearch scientist at the Institute ol' Public PolicyStudies, University of Michigan, Ann Arbor 48109.Dr. Hamilton is a professor of evolutionary biologyin the Museum of Zoology and the Division ofBiological Sciences, University of Michigan.SCIENCE, VOL. 21 1, 27 MARCH 1981

truism can benefit reproduction of theset, despite losses to the individual altruist. In accord with this theory's predictions, apart from the human species,almost all clear cases of altruism, andmost observed cooperation, occur incontexts of high relatedness, usually between immediate family members. Theevolution of the suicidal barbed sting ofthe honeybee worker could be taken asparadigm for this line of theory (7).Conspicuous examples of cooperation(although almost never of ultimate selfsacrifice) also occur where relatedness islow or absent. Mutualistic symbiosesoffer striking examples such as these: thefungus and alga that compose a lichen;the ants and ant-acacias, where the treeshouse and feed the ants which, in turn,protect the trees (8); and the fig waspsand fig tree, where wasps, which areobligate parasites of fig flowers, serve asthe tree's sole means of pollination andseed set ( 9 ) . Usually the course of cooperation in such symbioses is smooth, butsometimes the partners show signs ofantagonism, either spontaneous or elicited by particular treatments (10). Although kinship may be involved, as willbe discussed later, symbioses mainly illustrate the other recent extension ofevolutionary theory, the theory of reciprocation.Cooperation per se has received comparatively little attention from biologistssince the pioneer account of Trivers (5);but an associated issue, concerning restraint in conflict situations, has beendeveloped theoretically. In this connection, a new concept, that of an evolutionarily stable strategy, has been formallydeveloped (6, 11). Cooperation in themore normal sense has remained clouded by certain difficulties, particularlythose concerning initiation of cooperation from a previously asocial state (12)and its stable maintenance once established. A formal theory of cooperation isincreasingly needed. The renewed emphasis on individualism has focused onthe frequent ease of cheating in reciprocatory arrangements. This makes thestability of even mutualistic symbiosesappear more questionable than under theold view of adaptation for species benefit. At the same time other cases thatonce appeared firmly in the domain ofkinship theory now begin to reveal relatednesses of interactants that are too lowfor much nepotistic altruism to be expected. This applies both to cooperativebreeding in birds (13) and to cooperativeacts more generally in primate groups(14). Here either the appearances of cooperation are deceptive-they are casesof part-kin altruism and part cheat27 MARCH 1981ing-or a larger part of the behavior isattributable to stable reciprocity. Previous accounts that already invoke reciprocity, however, underemphasize thestringency of its conditions (15).Our contribution in this area is new inthree ways.1) In a biological context, our model isnovel in its probabilistic treatment of thepossibility that two individuals may interact again. This allows us to shed newPrisoner's Dilemma game in particular,allow a formalization of the strategicpossibilities inherent in such situations.The Prisoner's Dilemma game is anelegant embodiment of the problem ofachieving mutual cooperation (16), andtherefore provides the basis for our analysis. To keep the analysis tractable, wefocus on the two-player version of thegame, which describes situations thatinvolve interactions between pairs ofSummary. Cooperation in organisms, whether bacteria or primates, has been adifficulty for evolutionary theory since Darwin. On the assumption that interactionsbetween pairs of individuals occur on a probabilistic basis, a model is developedbased on the concept of an evolutionarily stable strategy in the context of thePrisoner's Dilemma game. Deductions from the model, and the results of a computertournament show how cooperation based on reciprocity can get started in an asocialworld, can thrive while interacting with a wide range of other strategies, and can resistinvasion once fully established. Potential applications include specific aspects ofterritoriality, mating, and disease.light on certain specific biological processes such as aging and territoriality.2) Our analysis of the evolution ofcooperation considers not just the finalstability of a given strategy, but also theinitial viability of a strategy in an environment dominated by noncooperatingindividuals, as well as the robustness of astrategy in a variegated environmentcomposed of other individuals using avariety of more or less sophisticatedstrategies. This allows a richer understanding of the full chronology of theevolution of cooperation than has previously been possible.3) Our applications include behavioralinteraction at the microbial level. Thisleads us to some speculative suggestionsof rationales able to account for theexistence of both chronic and acutephases in many diseases, and for a certain class of chromosomal nondisjunction, exemplified by Down's syndrome.Strategies in the Prisoner's DilemmaMany of the benefits sought by livingthings are disproportionally available tocooperating groups. While there are considerable differences in what is meant bythe terms "benefits" and "sought," thisstatement, insofar as it is true, lays downa fundamental basis for all social life.The problem is that while an individualcan benefit from mutual cooperation,each one can also do even better byexploiting the cooperative efforts of others. Over a period of time, the sameindividuals may interact again, allowingfor complex patterns of strategic interactions. Game theory in general, and theindividuals. In the Prisoner's Dilemmagame, two individuals can each eithercooperate or defect. The payoff to aplayer is in terms of the effect on itsfitness (survival and fecundity). No matter what the other does, the selfishchoice of defection yields a higher payoffthan cooperation. But if both defect,both do worse than if both had cooperated.Figure 1 shows the payoff matrix ofthe Prisoner's Dilemma. If the otherplayer cooperates, there is a choice between cooperation which yields R (thereward for mutual cooperation) or defection which yields T (the temptation toaefect). By assumption, T K ,so that itpays to defect if the other player cooperates. On the other hand, if the otherplayer defects, there is a choice betweencooperation which yields S (the sucker'spayof) or defection which yields P (thepunishment for mutual defection). Byassumption P S , so it pays to defect ifthe other player defects. Thus, no matterwhat the other player does, it pays todefect. But, if both defect, both get Prather than the larger value of R that theyboth could have gotten had both cooperated. Hence the dilemma (17).With two individuals destined never tomeet again, the only strategy that can becalled a solution to the game is to defectalways despite the seemingly paradoxical outcome that both do worse thanthey could have had they cooperated.Apart from being the solution in gametheory, defection is also the solution inbiological evolution (18). It is the outcome of inevitable evolutionary trendsthrough mutation and natural selection:if the payofs are in terms of fitness, and

Player Bplay"*ICDCooperationDefection7CR 3Reward forCooperationmutual cooperationI11Sucker's payoffC--------------i--------------- DDefection!T 5'Temptation toidefectIP l/Punishment for1mutual defectionthe interactions between pairs of individuals are random and not repeated, thenany population with a mixture of heritable strategies evolves to a state where allindividuals are defectors. Moreover, nosingle differing mutant strategy can dobetter than others when the population isusing this strategy. In these respects thestrategy of defection is stable.This concept of stability is essential tothe discussion of what follows and it isuseful to state it more formally. A strategy is evolutionarily stable if a populationof individuals using that strategy cannotbe invaded by a rare mutant adopting adifferent strategy (11). In the case of thePrisoner's Dilemma played only once,no strategy can invade the strategy ofpure defection. This is because no otherstrategy can do better with the defectingindividuals than the P achieved by thedefecting players who interact with eachother. So in the single-shot Prisoner'sDilemma, to defect always is an evolutionarily stable strategy.In many biological settings, the sametwo individuals may meet more thanonce. If an individual can recognize aprevious interactant and remember someaspects of the prior outcomes, then thestrategic situation becomes an iteratedPrisoner's Dilemma with a much richerset of possibilities. A strategy would takethe form of a decision rule which determined the probability of cooperation ordefection as a function of

Robert Axelrod and William D. Hamilton The theory of evolution is based on the struggle for life and the survival of the fittest. Yet cooperation is common be-tween members of the same species and even between members of diff

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