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The Biological Nature of the State Author(s): Roger D. Masters Source: World Politics, Vol. 35, No. 2 (Jan., 1983), pp. 161-193Published by: Cambridge University PressStable URL: http://www.jstor.org/stable/2010269Accessed: 10-03-2015 04:29 UTC

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THE BIOLOGICAL NATURE OF THE STATE

By ROGER D. MASTERS*

W HY do human beings live in societies consisting of hundreds of V V thousands, if not millions, of individuals who owe obedience to

the state? This question, long at the center of political philosophy, has been given renewed importance by the findings of contemporary biology: from an evolutionary perspective, the very existence of legal and political institutions is far more problematic than is generally assumed. Para- doxically enough, however, the origin and nature of the state can be illuminated by the theory of natural selection.

For over three million years, our hominid ancestors lived in small bands, probably approximating a hundred in number.' Most other mammals-and notably our closest relatives, the primates also form groups that are several orders of magnitude smaller than contemporary societies. In such small, face-to-face bands, the extensive division of labor and social stratification connected with a formal state do not normally exist.

Recent models of natural selection indicate that sexually reproducing animals such as Homo sapiens are unlikely to form enormous, impersonal social systems in which individuals are called upon to sacrifice time, money, and sometimes even their lives for the benefit of the community.2 Because contemporary nation-states are so much larger than the groups observed in other mammals, it is appropriate to seek an evolutionary explanation for human legal and political institutions.

To avoid confusion, however, the use of biological concepts in social science must be based on careful specification on the level of analysis. Scientific comparisons, unlike organic metaphors, must distinguish care- fully between evolutionary processes at the species level, social behavior in groups, and individual physiological and neurological functions.3 As

* This article is a greatly expanded and revised version of a paper given at the Symposium on Law and Behavioral Research, Monterey Dunes, Calif., September 25-27, 5981.

C. Owen Lovejoy, "The Origin of Man," Science, Vol. 22I (January 23, i981), 34I-50; Richard E. Leakey and Roger Lewin, Origins (New York: Dutton, I977).

2Edward 0. Wilson, Sociobiology (Cambridge: Harvard University Press, 1975); David Barash, Sociobiology and Behavior (New York: Elsevier, I977).

3 Roger D. Masters, "Functional Approaches to Analogical Comparisons between Species," in Mario von Cranach, ed., Methods of Comparing Animal and Human Behavior (The Hague: Mouton, I976), 73-I02-

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162 WORLD POLITICS

used here, the concept of evolution concerns only the process of natural selection at the level of populations, and can refer to individually learned, culturally transmitted, or genetically programmed behavior.4 In this paper, therefore, no assumptions will be made concerning the extent- if any-of genetic constraints on human behavior.

Since social scientists have long assumed that humans are "unique," they have tended to ignore research on other species. It is becoming increasingly evident, however, that the separation of biology from the social sciences is unwise.5 Despite charges that "sociobiology" is ideo- logically biased and scientifically untenable,6 the neo-Darwinian theory of natural selection has been shown to be compatible with diverse political conclusions.7 Substantively, evolutionary biology has begun to enrich such diverse fields as anthropology,8 economics,9 sociology,bo and political science."

I will begin this essay by showing how evolutionary theory can be 4 Donald T. Campbell, "Variation and Selective Retention in Socio-cultural Evolution,"

in Herbert R. Barringer, George I. Blanksten, and Raymond W. Mack, eds., Social Change in Developing Areas (Cambridge, Mass.: Schenkman, I965), I9-48; Roger D. Masters, "Politics as a Biological Phenomenon," Social Science Information, xiv (April I975), 7-63; Richard D. Alexander, "Evolution, Human Behavior, and Determinism," Proceedings of the Biennial Meeting of the Philosophy of Science Association, 11 (977), 3-2I.

5Alexander Rosenberg, Sociobiology and the Preemption of Social Science (Baltimore: The Johns Hopkins Press, i980); Richard D. Alexander, "Evolution, Culture, and Human Be- havior: Some General Considerations," in Richard D. Alexander and Donald W. Tinkle, eds., Natural Selection and Social Behavior: Recent Research and Theory (New York: Chiron Press, i982), 504-20; John C. Wahlke, "Prebehavioralism in Political Science," American Political Science Review, Vol. 73 (March I979), 9-3I.

6 Marshall Sahlins, Use and Abuse of Biology (Ann Arbor: University of Michigan Press, I976); Arthur Caplan, ed., The Sociobiology Debate (New York: Harper & Row, I978).

7 Roger D. Masters, "Is Sociobiology Reactionary? The Political Implications of Inclusive Fitness Theory," Quarterly Review of Biology, Vol. 57 (September i982), 275-92; Marvin Bressler, "Biological Determinism and Ideological Indeterminacy," in Elliott White, ed., Sociobiology and Human Politics (Lexington, Mass.: Lexington Books, i98i), i8i-9i. For historical perspective, see also Loren Graham, "Science and Values: The Eugenics Movement in Germany and Russia in the I92os," American Historical Review, Vol. 82 (December I977), II33-64-

8 Robin Fox, ed., Biosocial Anthropology (London: Malaby, I975); Napoleon Chagnon and William Irons, eds., Evolutionary Biology and Human Social Behavior (N. Scituate, Mass.: Duxbury, I979); Joan S. Lockard, ed., The Evolution of Human Social Behavior (New York: Elsevier, i980); Pierre van den Berghe & David Barash, "Inclusive Fitness and Family Structure," American Anthropologist, Vol. 79 (No. 4, I977), 809-23.

9 Jack Hirshleifer, "Natural Economy vs. Political Economy," Journal of Social and Bio- logical Structures, I (October I978), 3I9-37; Ivan Chase, "Cooperative and Noncooperative Behavior in Animals," American Naturalist, Vol. II5 (June I980), 827-57; Howard Margolis, "A New Model of Rational Choice," Ethics, Vol. 9I (January I98I), 265-79.

10 Lionel Tiger, Men in Groups (New York: Random House, I969); Allan Mazur, "Bio- sociology," in James F. Short, Jr., ed., The State of Sociology (Beverly Hills, Calif.: Sage, I98I), I4I-60.

Albert Somit, ed., Biology and Politics (The Hague: Mouton, I976); Thomas C. Wiegele, Biopolitics (Boulder, Colo.: Westview, I979); White (fn. 7); Meredith Watts, ed., Biopolitics: Ethological and Physiological Approaches, New Directions for Methodology of Social and Behavioral Science, No. 7 (San Francisco: Jossey-Bass, I98I).



BIOLOGICAL NATURE OF THE STATE 163

extended to study human social institutions without engaging in genetic reductionism (Section I). Next, I will survey the relationship between cost-benefit models of natural selection and "rational actor" theories of human behavior (Section II), political philosophy (Section III), and empirical research on the origin of the state (Section IV). On this basis, I will reassess the potential contribution of evolutionary biology to the study of politics (Section V).

I. INCLUSIVE FITNESs THEORY AND HUMAN SOCIAL BEHAVIOR

Contemporary neo-Darwinian theory requires a fundamental shift in our understanding of social behavior. Political philosophers and social scientists usually treat the question of "sociability"-i.e., whether humans are naturally selfish or cooperative-as an attribute of "human nature." But biology now teaches us that the existence and structure of social groups varies, even within a single species, as a consequence of the interaction between animals and their environment; a species that is asocial in one setting may be social in others.12 From a scientific perspective, therefore, it is misleading to conceptualize social behavior as a constant natural trait (such as two eyes, one nose, and one mouth in the human face).

The conventional wisdom-according to which biology readily explains the existence of human society, but not the differences between human cultures-is thus inadequate: the emergence of large-scale human societies and formal legal systems must be treated as a biological problem.'3 The usual answers to this problem violate the premises of evolutionary theory because they assume that the state is beneficial to the group, results from chance, or is an act of human creation;'4 none of these conventional explanations is tenable without a careful consid- eration of the selective pressures against institutions like the state-in other animals, and during most of hominid evolution. The approach known as "inclusive fitness theory" suggests an alternative that is con-

Marc Beckoff and Michael C. Wells, "The Social Ecology of Coyotes," Scientific Amer- ican, Vol. 242 (April i980), I30-48; Richard Wrangham, "Review Essay: Sociobiology," Biological Journal of the Linnean Society, xiii (March i980), 171.

'3 Richard D. Alexander, "Natural Selection and the Analysis of Human Sociality," in C. E. Goulden, ed., Changing Scenes in Natural Sciences, I776-I976 (Special Publication I2, Philadelphia: Philadelphia Academy of Natural Sciences, I977), 283-337; Donald T. Camp- bell, "On the Genetics of Altruism and the Counter-hedonic Components in Human Culture," Journal of Social Issues, xxviii (No. 3, 1972), 21-37; Wilson (fn. 2), Part III.

'4 E.g., St. Thomas Aquinas, Summa Theologica, I-II, Q. 90-97, in Dino Bigongiari, ed., The Political Ideas of St. Thomas Aquinas (New York: Hafner, I957), 3-85; Machiavelli, Discourses on Titus Liuy, ed. by Bernard Crick, I, ii and I, ix (Harmondsworth, England: Penguin, 1970), 104-II, I3I-34-



164 WORLD POLITICS

sistent with the widely observed tendencies of animals to adjust their behavior to maximize long-range reproductive success.

Cooperation generally entails some cost to the individual. To under- stand animal social behavior, biologists therefore have to discover how individuals gain from cooperation and to show that these benefits exceed the costs. For example, under conditions of abundant and con- tinuously available resources, animals not subject to predation tend to be asocial. In such situations, there is virtually no benefit from cooperation that overbalances its costs in energy or time.'5 Hence, in species like the lepilemur or the orangutan, one finds asocial behavior that resembles Rousseau's "state of nature" to a remarkable degree.'6

According to the prevailing interpretation of inclusive fitness theory, natural selection operates primarily-though not necessarily solely-at the level of the individual.'7 If two organisms interact, whether or not they cooperate can usually be predicted by a cost-benefit calculus of the alternatives. This calculus need not be conscious; indeed, it provides an explanation of the natural selection (sometimes called the "ultimate causation") of animal behavior in an extraordinary number of species. As in rational actor models in economic theory-particularly as applied to questions of collective goods or public choice-all that need be assumed is that behavior will vary as if it were a strategy chosen on the basis of cost-benefit optimization.,8

In his well-known article introducing the concept of inclusive fitness, William Hamilton proposes four general "classes" or "types of behavior" defined in such cost-benefit terms.'9 When organisms interact, the actor can derive either a net benefit (+) or a net loss (-) from his behavior, and "neighbors"-i.e., other organisms involved in the social interaction-can likewise derive either a net benefit or a net loss. The resulting four-fold table, rewritten to stress behavioral consequences rather than

1Barash (fn. 2), 57-62; Hans Kummer, "On the Value of Social Relationships to Non- human Primates: A Heuristic Scheme," in Mario von Cranach and others, Human Ethology (Cambridge, England: Cambridge University Press, I979), 38I-95.

6 Jean-Jacques Rousseau, Second Discourse, Part I and note j, in Roger D. Masters, ed., Rousseau's First and Second Discourses (New York: St. Martin's Press, i964), I04-40, 203-I3; Birute Galdikas-Brindamour, "Orangutans, Indonesia's 'People of the Forest,'" National Geographic, Vol. I48 (October I975), 444-73; Roger D. Masters, "Jean-Jacques is Alive and Well: Rousseau and Contemporary Sociobiology," Daedalus, Vol. 107 (Summer 1978), 93- I05.

'7George C. Williams, Adaptation and Natural Selection (Princeton: Princeton University Press, i966).

,8 John Maynard-Smith, "The Evolution of Behavior," Scientific American, Vol. 239 (Sep- tember 1978), 176-92; Barash (fn. 2), 51-52; Hirshleifer (fn. 9); Chase (fn. 9). On the distinction between "ultimate" and "proximate" causation, see Barash (fn. 2), 37-39.

'9 Hamilton, "The Genetical Evolution of Social Behavior," Journal of Theoretical Biology, vii (No. I, i964), i-i6; reprinted in Caplan (fn. 6), I9I-209. Cf. Barash (fn. 2), 96.




subjective intention,20 has general relevance as a means of linking evolutionary biology to the study of human institutions (Figure I).

FIGURE I

HAMILTON S 'FOUR TYPES OF BEHAVIOR"

A GainI LossI

MUTUAL BENEFIT SOCIALITY OR l Gain - /VIRTUE

Loss NEPOTISM MUTUAL HARM

SOURCE: Modified from Hamilton (fn. I9), 207.

This approach analyzes social behavior in terms of each individual's benefits and costs, on the assumption that observable outcomes establish the net utility of the act. In economic theory, such cost-benefit analysis is usually measured in monetary or materialistic (economic) terms. Ev- olutionary biology substitutes long-range reproductive success-i.e., the proportion of an individual's genes transmitted to future generations of the species. Inclusive fitness thus differs from the traditional notion of "survival of the fittest" in two important respects: first, natural selection favors the ability of individuals to transmit their genes to posterity (rather than their "fitness" in terms of health, power, beauty, or other physical traits); second, an organism's reproductive success can sometimes be furthered by assisting others, instead of by mating.21

Social biologists call the lower left cell of Figure I nepotism, or kin selection, because the actor's positive payoff-a larger share of genes, in comparison to others, in future generations necessarily involves fa-

20 On the importance of this distinction, see Gunther Stent, "You Can Take the Ethics out of Altruism, But You Can't Take the Altruism out of Ethics," Hastings Center Report, VII (I977), 33-36; Roger D. Masters, "Beyond Reductionism," in von Cranach (fn. I5), 265- 84.

21 On the "coefficient of relatedness"-i.e., "the fraction of genes in two individuals that are identical by descent, averaged over all loci"-as a measure of reproductive success, see Wilson (fn. 2), 74. For examples of non-mating strategies, see also Barash (fn. 2), esp. 91- 93.



166 WORLD POLITICS

voring close kin as compared to more distant relatives or unrelated individuals. In contrast, the upper left cell-mutual benefit (often called mutualism or reciprocity)-refers to situations in which two presumably unrelated individuals both gain from an action; among animals, such an exchange of benefits often rests on a mechanism of recognition or reciprocity that discriminates against noncooperative "free riders."22 Mu- tual harm-the lower right cell of Figure i-can occur when organisms gain more from denying themselves the use of a beneficial resource than from running the risk that it be preempted by rivals;23 when such interactions predominate, a species will tend to be asocial.

Since nepotism, mutual benefit, and mutual harm all refer to self- interested or "egoistic" behavior, all can be readily explained-depending on circumstances-as results of natural selection.24 Situations in which an animal incurs a relative loss while potential competitors gain (the upper right cell in Figure i) seem harder to derive from evolutionary theory. Such group benefits or "giving" behaviors are instances of social cooperation between non-kin without personal reciprocity; anything that benefits the actor's kin is at least equally helpful to reproductive rivals and their offspring. Since the term "altruism," as used by many socio- biologists, confuses cost-benefit outcomes and motivations, it is preferable to call this category sociality or-to apply an old-fashioned term-virtue.

The cells in Figure i can also be described as four categories of social partners: kin (others benefiting from nepotistic behavior), allies (others sharing mutual benefits), non-kin (others benefiting by sociality or the mere fact of group membership), and enemies (others excluded from the group that is sharing benefits, even at the risk of mutual harm). Whether a human action falls in one or another of these four cells-and hence, whether an individual responds to the "other" as kin, as ally, as non- kin group member, or as enemy-depends on environmental conditions. At any time, moreover, humans may find a benefit in forgoing all social interaction, retreating into asocial isolation. According to the circumstances, the same behavior-be it eating or defense against a predator- can thus be entirely asocial or can generate shared benefits for kin, for allies, or for all group members.

22 Robert Trivers, "The Evolution of Reciprocal Altruism," Quarterly Review of Biology, XLVI (No. 4, I971), 35-57-

23Raymond Pierotti, "Spite and Altruism in Gulls," American Naturalist, Vol. I IS (Feb- ruary 1980), 290-300; Wilson (fn. 2), II9.

24 Richard D. Alexander, "Evolution of Social Behavior," Annual Review of Ecology and Systematics, v ('974), 25-83; Richard Dawkins, The Selfish Gene (New York: Oxford Uni- versity Press, I976).




Since contemporary theories of natural selection stress the priority of individual reproductive success,25 one can never presume that a behavior has been selected because it redounds to the benefit of the population or species. But it is equally erroneous to assume that, as a matter of principle, natural selection never generates behavior of this sort.26 Indeed, much recent theoretical work suggests a dialectical relationship between individual inclusive fitness and group interest,27 particularly in a complex social species like Homo sapiens.28

Many species have specialized in one or two of the five possible kinds of behavior (asociality, nepotism, mutual benefit, sociality, and mutual harm). But if environments change or are uncertain, a diverse and more plastic repertoire will be selected.29 Humans seem to have exploited a broader range of social alternatives than most other animals, thanks to our larger brains and complex linguistic systems. Moreover, the emergence of mutual benefit or sociality, not to mention occasions of mutual harm, in no way abolishes the adaptiveness of nepotism or- in appropriate conditions-asociality. In conventional terms, humans combine "selfishness" and "self-interested" cooperation with "altruism" and "spite" to an often bewildering degree.30

Social biology thus gives us a more precise understanding of the difference between human social behavior and that of other gregarious animals. Inclusive fitness theory leads to a prediction that the likely costs and benefits of alternative behaviors will vary, depending on ecological circumstances and the individual's social role, prior experience, age, or sex. Humans, especially because of their intelligence, are capable of exploiting many if not all of the possible alternatives in a single day, or even a single hour. Moreover, sociality or virtue among non-kin creates

25 Williams (fn. I7); Robert Trivers, "Sociobiology and Politics," in White (fn. 7), 4-7. 26 David Sloan Wilson, The Natural Selection of Populations and Communities (Menlo Park,

Calif.: Benjamin Cummings, I979); Roger D. Masters, "The Value-and Limitations-of Sociobiology," in White (fn. 7), I35-65.

27 Richard D. Alexander and Gerald Borgia, "Group Selection, Altruism, and the Levels of the Organization of Life," Annual Review of Ecology and Systematics, IX (1978), 449-74; David Layzer, "Altruism and Natural Selection," Journal of Social and Biological Structures, I (July I978), 297-305; Michael J. Wade, "Kin Selection: Its Components," Science, Vol. 2i0 (November 7, i980), 665-67.

28 Martin L. Hoffman, "Is Altruism Part of Human Nature?" Journal of Personality and Social Psychology, XL (January 1981), I2I-37; Howard Margolis, Selfishness, Altruism, and Rationality (Cambridge, England: Cambridge University Press, 1982).

29 Leslie A. Real, "Fitness, Uncertainty, and the Role of Diversification in Evolution and Behavior," American Naturalist, Vol. I I5 (May 1980), 623-38.

30 Roger D. Masters, "Of Marmots and Men: Human Altruism and Animal Behavior," in Lauren Wispe, ed., Altruism, Sympathy, and Helping (New York: Academic Press, 1978), 59-77; Gunther Stent, ed., Morality as a Biological Phenomenon (Berkeley: University of California Press, 1979).



168 WORLD POLITICS

a situation in which cheating-i.e., pretending to cooperate while actually engaging in nepotism ("selfish" behavior)-has greater payoffs than reciprocity, provided, of course, that it goes undetected (Plato, Republic, II.359b-36od).3'

Humans manipulate this repertoire of social possibilities by means of a dual transformation of the cost-benefit calculus observed in animal behavior. At the individual level, the assessment of the interpersonal situation (i.e., the categories in Figure i) is transformed-usually in a conscious way, but sometimes through repression or self-deception- into a subjective motive. These psychological responses are further molded into sociocultural institutions which, while superficially associated with a corresponding psychological motive, actually use diverse assessments of the world (Table i). Since each biological situation can generate any of the five motivational responses-and each of the individual motives can produce behaviors at any of the five institutional levels-a reduc- tionist science of human behavior is highly improbable, if not impossible.32

Language is, of course, what enables humans to engage in this dual transformation of natural potentialities. In other species, intention can be inferred from species-specific behaviors that have been ritualized into what Lorenz has called releasing mechanisms.33 Whatever the internal events in other animals, they do not seem to serve simultaneously as subjective states and as objective social signals or symbols. In contrast, human language permits this duality, since words can be used either in silent thought or in audible speech. Probably grounded in the unusual structure of phonemic utterances, speech can thus establish cultural rules and social institutions based on shared meaning.34 To put it more simply, language enables humans to establish cultures to a degree apparently impossible in other animals (Aristotle, Politics, I.ii.I253a; Hobbes, Levi- athan, I, iv).

We are now in a better position to link inclusive fitness theory with the conventional social sciences. Human institutions, in their bewildering

P Trivers (fn. 25), 9-39; Fred Willhoite, "Rank and Reciprocity: Speculations on Human Emotions and Political Life," in White (fn. 7), 239-58.

32 Glendon Schubert, "The Sociobiology of Political Behavior," in White (fn. 7), I93-238; Masters (fn. 20); Stent (fn. 30); P. W. Anderson, "More is Different," Science, Vol. I77 (August 4, I972), 393-96.

33 Konrad Lorenz, Studies in Animal and Human Behavior, 2 vols. (Cambridge: Harvard University Press, I970-I97I).

34 Roger D. Masters, "Genes, Language, and Evolution," Semiotica, ii (No. 4, I970), 295- 320; John Hurrell Crook, The Evolution of Human Consciousness (New York: Oxford Uni- versity Press, 198I).




TABLE 1

HUMAN TRANSFORMATIONS OF INCLUSIVE FITNESS STRATEGIES

Cultural Institution** (Sociology,

Natural Individual Anthropology, Situation* Motive** Economics, &

(Social Biology) (Psychology) Political Science)

Asociality Egoism Individuality ("Sauve qui peut")

Nepotism Love Family ("Be mine, I'm yours")

Mutual Benefit Reciprocity Trade: Marriage, ("I scratch your Barter, Markets back, you ...

Sociality Altruism Community: ("Do unto others .") State, Church

Mutual Harm Spite Feud: Economic ("If I can't have Competition, War it, no one can

* Situations defined in terms of Figure 1. ** Each human motive can result from any one of the five natural situations, and each human institution can rest on any one or a combination of the five individual motives.

variety and complexity, are constituted by the attributes defining who is included, who is excluded, and how participants are expected to behave. When social institutions are described, however, we usually distinguish several major categories or levels of analysis, such as the family, the business firm or market economy, and the state or community. Mediated by psychological assessments and motives, these familiar human institutions correspond-in a rough way-to categories of animal social interaction.

Among humans, situations that other animals usually-though not always35-"live" in an unreflective manner have been transformed into distinct, and often contradictory, phenomena of individual consciousness

35 Donald R. Griffin, The Question of Animal Awareness (New York: Rockefeller University Press, I976); John Tyler Bonner, The Evolution of Culture in Animals (Princeton: Princeton University Press, i980).



170 WORLD POLITICS

and cultural rules. This means that animal instinct and human learning are different ways of responding to similar problems created by environmental conditions and individual needs (cf. Aristotle, Politics, VIII.xvii.1337a). Hence the concepts of inclusive fitness theory are relevant to complex human institutions like the state, even though human social behavior often has different "proximate" causes than that of other species.

II. COST-BENEFIT THEORIES OF COOPERATION: FROM THE PRISONER S DILEMMA TO THE TRAGEDY OF THE COMMONS

As Brian Barry has argued,36 there are two broad traditions in the social sciences: economic models, in which aggregate phenomena are analyzed as if individuals rationally calculated the costs and benefits of alternatives, and sociological approaches, which treat entire systems as having properties that cannot be deduced from individual choices. A similar duality of methods can be discerned in the biological sciences.37 Since both inclusive fitness theory in biology and economic or rational actor models in the social sciences are based on a cost-benefit analysis, it has been argued that they can be integrated into a single approach.38 This prospect is particularly relevant to research concerning the state, because rational actor theories of public choice or collective goods, akin to those in economics, have increasingly been adopted by political scientists.39

To indicate how inclusive fitness theory is related to social science, it is convenient to begin with game theory, which has been used by many social biologists to explain the costs and benefits of alternative behavioral strategies.40 In particular, the well-known "Prisoner's Dilemma" is a fruitful way of examining the problem of cooperation: this class of games

36 Barry, Sociologists, Economists, and Democracy (London: Macmillan, I970). Cf. C. Wright Mills, The Sociological Imagination (New York: Oxford University Press, I959).

37 Masters (fns. 7 and 26); Williams (fn. 25); Alexander and Borgia (fn. 27). 38 Jack Hirshleifer, "Economics from a Biological Viewpoint," Journal of Law and Eco-

nomics, xx (April I977), I-52; Chase (fn. 9); Margolis (fn. 28). 39 E. g., Anthony Downs, An Economic Theory of Democracy (New York: Harper, I957);

Mancur Olson, Jr., The Logic of Collective Action (Cambridge: Harvard University Press, i965); Albert 0. Hirschman, Exit, Voice, and Loyalty (Cambridge: Harvard University Press, I974); Denis G. Sullivan, Robert T. Nakamura, and Richard F. Winters, How America Is Ruled (New York: Wiley, i980).

40 Trivers (fn. 22); Maynard-Smith (fn. i8); Donald L. McEachron and Darius Baer, "A Review of Selected Sociobiological Principles: Application to Hominid Evolution. II. The Effects of Intergroup Conflict," Journal of Social and Biological Structures, v (April i982), I2I-39.




has been widely used in the social sciences,41 and has also been applied in explanations of animal social life.42 For the sake of exposition, consider the following example of the Prisoner's Dilemma.

Two men, A and B, have collaborated in stealing $20, which they split fifty-fifty. They have been caught and placed in separate jail cells. The jailer tells them that if both remain silent, he lacks evidence for a full prosecution and will release them with a fine of $i each (leaving both A and B with a net gain of $9 each). If one of the two talks, but the other does not, the one who talks ("defector") will go scot free (net gain: $io), while the silent individual ("sucker") will be forced to return the stolen money and be sentenced to ten days in jail or a fine of $io (net cost: -$io). If both talk, both forfeit the money and are sentenced to nine days in jail or $9 fines (net cost: -$9 each). For A and B jointly, this last result is the worst possible outcome, since all the stolen money is forfeited and both are punished (jail or fine). Yet it is this worst outcome (for the two collectively) that is chosen if A and B decide their actions "rationally" on the basis of a cost-benefit calculation. This follows because each individual must try to gain the most (or lose the least) regardless of the other's action. A gains more (or loses less) by talking than by silence if B is silent or if B talks; B, deciding separately, makes the same calculation.

The game matrix for this Prisoner's Dilemma is set forth in Figure 2a. This game has at least three essential conditions: first, the payoffs must be transitive (in the example, + io ) +9 ) -9 ) - io); second, each actor independently chooses a strategy that will maximize his gains (or minimize his losses); third, communication or behavioral coercion between the actors is not allowed. Under these circumstances, as game theorists are fond of pointing out, it is "rational" for both A and B to choose the outcome in which both lose, since for each individual the strategy of talking (defecting) dominates that of silence (cooperating).

On a moment's reflection, Figure 2a will be seen to contain the same basic categories as Figure i (derived from inclusive fitness theory). The difference, of course, is that the Prisoner's Dilemma specifies a set of environmental constraints and then seeks to predict the strategy that will be chosen, whereas Hamilton's more general model defines the basic options in any set of environmental conditions. Hence, the Prisoner's Dilemma can be taken as a special case of the general theory, albeit one

41Morton A. Kaplan, System and Process in International Politics (New York: Wiley, 1957), Part IV; Robert Axelrod, "Effective Choice in the Prisoner's Dilemma, Journal of Conflict Resolution, xxiv (March i980), 3-25, and "More Effective Choice in the Prisoner's Dilemma," ibid. (September i980), 379-403.

42 Robert Axelrod and William D. Hamilton, "The Evolution of Cooperation," Science, Vol. 2I I (March 27, i98i), I390-96; Robert Axelrod, "The Emergence of Cooperation among Egoists," American Political Science Review, Vol. 75 (June i980), 306-i8.



172 WORLD POLITICS

FIGURE 2A PRISONER'S DILEMMA: NON-KIN

A's Strategy Silent Talk

A =-+9 A- +I0 Silent B:+9 B=-10

_ MMUTUAL BENEFIT NEPOTISMM

al B =+IO B -9

IALITY OR VIRTUE| MUTUAL HARM

ASSUMPTIONS: 2-person game in which payoffs are transitive (+ IO > +9 > -9 > -I o) and commensurable; communication or behavioral coercion between A and B impossible. * Nepotism (private self-interest) and Sociality (virtue) defined in terms of A's payoffs.

that is particularly useful in explaining many counterintuitive situations. One major consequence of this similarity deserves emphasis. The labels

in these figures concern the four outcomes as defined from the perspective of A. In the Prisoner's Dilemma, what is nepotism ("selfishness") from the perspective of A will be sociality or virtue ("altruism") from B's point of view. One man's meat is another man's poison. Is it any wonder that humans often suspect the motivations of others-and proclaim the in- nocence of their own intentions (even if, or rather especially if, these intentions are consistent with their long-range self-interest)? As Trivers has shown,43 humans seem to specialize in deceit in order to "cheat" on their partners, in moralistic aggression against those who cheat, and in deceitful imitation of compliance and morality in order to avoid punishment.

Even if the Prisoner's Dilemma is useful in analyzing such problems of social cooperation, what is gained by following those biologists who have extended this model to evolutionary theory? To illustrate the un- expected consequences of linking game theory and inclusive fitness theory, consider a Prisoner's Dilemma in which the two prisoners are full siblings, or father and son. If the coefficient of relatedness between A and B is I/2, A's payoff for each cell of the matrix must be rewritten to include one-half of B's payoff (which also accrues to A), and vice versa. When we recalculate the game matrix in this way (Figure 2b), the strategy of cooperating now dominates that of defecting for each pris-

43 Trivers (fns. 22 and 25).




oner, regardless of the response of the other. Rather than resulting in mutual harm, the same payoffs lead to cooperation and mutual benefit. It should be little wonder that the Mafia is organized in "families" based on pseudo-kinship if not real kin relations-or that states often use symbols of kinship ("fatherland," "motherland," 'fraternite") to legitimate obedience.

FIGURE 2B PRISONER'S DILEMMA: KIN**

A's Strateg y Silent I Talk

A =+l13.5 A=+ 5 >t Slent B+35B- B A- + 13.5 B - 5 a MMUTUAL BENEFIT NEPOTISM

%q) ~~A =-5 A= -13.5 ~ ak B =+5 B= -13.5 SOCIALITY OR VIRTUE MUTUAL HARM

ASSUMPTIONS: as in Figure 2A. * Nepotism (private self-interest) and Sociality (virtue) defined in terms of A's payoffs. ** Kin are either full siblings or parent-offspring (i.e., "coefficient of relatedness" or r .5); hence, in each cell, A adds .5 of B's pay-off in Figure 3A to his own (and vice-versa).

As this example suggests, inclusive fitness theory leads to the prediction that social cooperation is most likely when the partners are kin or can gain mutual and reciprocal benefits. But, as Axelrod and Hamilton show,44 the Prisoner's Dilemma also illustrates the natural constraints that inhibit self-sacrifice for others (sociality or virtue): when confronted with multiple plays of Prisoner's Dilemma, the most reliable way to avoid mutual harm is called a "TIT-FOR-TAT" strategy-i.e., respond to the other player or organism as it behaved in the last round. The virtuous response cannot simply be established as the "best" policy in all cases, because such a strategy will be vulnerable to cheating by a free rider who gets the benefits of the virtue of others without paying any of the costs.

From the perspective of game theory, social cooperation usually occurs only in cases of mutual benefit. To be sure, the decision rule need not be "TIT-FOR-TAT" (using the other's last action as a cue for the appropriate

44 Axelrod and Hamilton (fn. 42).



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response). Maynard-Smith has shown that the location of an interaction can also serve as a behavioral rule, leading to a mutually beneficial "bourgeois" strategy of deferring to the other individual when on the other's "turf."45 Other formalizations include Hirshleifer's "tender trap"- a game in which any rule of behavior that permits an actor to predict what others will do is preferable to the absence of rules.46 All these models can be used to develop Trivers's original insight that reciprocity can produce mutually beneficial social cooperation among non-kin, even in circumstances of short-term loss to the cooperator (Aristotle, Ethics, V.v-vi.i I32b-I i34b).

Although this approach can readily explain nepotism and mutual benefit, it can also account for virtue or-to use the language of rational actor theory-the choice of collective goods. Admittedly, this choice is rare in other species, but humans occasionally act in the group interest even when reciprocity is unlikely or is deferred beyond the temporal framework of the alternatives. Game theory is a useful way to illustrate the origins of such sociality or virtue, especially because it reveals the critical role of coercive restraints imposed by an impersonal and bu- reaucratic state.

Although the problem of collective goods can be analyzed in terms of an n-person Prisoner's Dilemma,47 for heuristic purposes it is clearer to consider a similar model that Garrett Hardin popularized as the "Tragedy of the Commons."48 In these situations, one individual can be considered to play a game against each of many neighbors.

Picture a pasture open to all. It is to be expected that each herdsman will try to keep as many cattle as possible on the commons.... As a rational being, each herdsman seeks to maximize his gain.... Adding together the component partial utilities, the rational herdsman concludes that the only sensible course for him to pursue is to add another animal to his herd. And another; and another.... But this is the conclusion reached by each and every rational herdsman sharing a commons. Therein is the tragedy. Each man is locked into a system that compels him to increase his herd without limit-in a world that is limited.49

45 Maynard-Smith (fn. i8). 46 Jack Hirshleifer, "Evolutionary Models in Economics and Law: Cooperation versus

Conflict Strategies," Research in Law and Economics (Greenwich, Conn.: JAI Press, i982), IV, i-6o.

47 Russell Hardin, "Collective Action as an Agreeable n-Prisoners' Dilemma," Behavioral Science, xvi (September I971), 472-8i; Iain McLean, "The Social Contract in Leviathan and the Prisoner's Dilemma Supergame," Political Studies, xxix (September i98i), 339-5I.

48 Garrett Hardin, "The Tragedy of the Commons," Science, Vol. i62 (December I3, i968), I243-48, reprinted in Herman E. Daley, ed., Toward a Steady-State Economy (San Francisco: Freeman, I973), chap. 6.

49 Ibid., I04.




Many ecologists have agreed with Hardin's argument that this situation approximates a wide range of phenomena, from the exhaustion of scarce natural resources to the pollution of the environment. 5

To show how game theory and inclusive fitness theory can be combined in the analysis of social institutions, consider the Tragedy of the Commons as a game with many players choosing independently whether or not to add to their herds. Since each knows that his decision cannot influence the choice of the others, it is inevitable that some herdsmen will adopt conflicting strategies (here called Strategy A and Strategy B). Assume that, ultimately, the commons will support only the xth cow of one set of strategists (either A's or B's, but not both). Nobody knows, however, either the value of x (i.e., the number of cows at which the carrying capacity is exceeded) or the number of players adopting each strategy.

The game has two outcomes, one in the short run (the addition of a single cow; Figure 3a), the other over a longer term (Figure 3b). In the short run, everyone will add a cow, for the same reason that both prisoners talk in the simple Prisoner's Dilemma: to do otherwise would be a gratuitous act of sociality, incurring loss for no conceivable benefit. Unlike the Prisoner's Dilemma, however, this strategy is mutually beneficial in the short run (Figure 3a); it is only later that losses are en- countered.

FIGURE 3A TRAGEDY OF THE COMMONS Short Run-Add i Cow

Strategy A Don't Add Cow jAdd CowI

Don't A:-I A=+I Add Cow

m% B |I Br-I MUTUAL HARM NEPOTISM*

Add Cow BA+I A + I u~~~i ~B + I A - + I

SOCIALITY* MUTUAL BENEFIT

ASSUMPTIONS: An n-person game in which each actor knows that only two strategies exist- and that others are free to choose either. All cows have a marginal utility = + i. * Nepotism (private self-interest) and Sociality (virtue) are defined in terms of A's payoffs.

50 E.g., John N. Cole, "Learning a Hard Lesson in Aroostook County," Country Journal (November i980), 42-46.



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If the carrying capacity of the commons is exceeded in the long run (Figure 3b), the mutual harm to all participants will be incommensurably large. Virtue (not adding cows) entails a loss, particularly compared to the strategy of nepotism or private self-interest; but the danger that everyone will seek such a selfish outcome makes self-restraint relatively beneficial as soon as there is a high enough probability that mutual harm can be prevented. The problem is that no one knows which will be the xth cow. Hence, without some form of organization, it is impossible to focus on the long-term game; the players continue to calculate only immediate benefits and costs; and the tragedy seems inevitable.

FIGURE 3B TRAGEDY OF THE COMMONS Long Run-Add Xth Cow

Strategy A

Don't Add Cow] [Add Cowl

Don't A= X-I A=X+I Add Cow

>% B= X-I B=X-I MUTUAL BENEFIT NEPOTISM

Add Cow A =X -I A=X-1OOO B X+I B XX -O000

SOCIALITY MUTUAL HARM

ASSUMPTIONS: as in Figure 3A. Commons can support an xth cow if added by either A strategists or B strategists, but not by both; value of x is unknown.

Transforming the players into two groups is a way out. If those adopting strategy B could coerce the A's into limiting their herds, while the B's themselves add cows, the latter could gain the benefits of nepotism; but for the losers, virtue still has its rewards, since the outcome of being coerced is preferable to mutual harm. By not adding cows, the A strategists avoid catastrophe-even though they know that their rivals may be less than perfectly virtuous. And if the commons or collective goods can be expanded-for example, by joint work or by conquest- even the group with the relatively inferior strategy can gain the mutual benefits of extending the short-run game beyond the original limits of the resource base.

From this perspective, citizens can benefit from the political community only by accepting the risk that those with power will gain




disproportionately from their role as enforcers of the social contract.5' Governments thus can be described as providing the service of co- ordinating social behavior, exacting a "side-payment" for protecting the collective goods.52 Once begun, such institutions can create considerable mutual benefits for all members as long as obedience to social rules prevents a Tragedy of the Commons. Under these conditions, it can be rational for self-interested individuals to form a political society, even if it will occasionally entail self-sacrifice or "virtue"-including paying taxes, serving in the army, even dying in defense of the community.53

It is relatively straightforward to move from such simplified models of game theory to more precise versions of rational choice theory.54 Not only does this approach suggest that the economists' distinction between selective benefits and collective goods can be founded on evolutionary biology, but it explains the origin of the state-and its ultimate fragility and disintegration.55

Cooperation can readily arise in small groups of extended kin (Figure 2b), and can even expand to mutually beneficial exchanges between non- kin (Figure 3a). But as soon as collective goods come into question- for instance, when groups compete for nondivisible resources, or the ecological carrying capacity is approached-the risk of mutual harm increases the benefits of enforceable norms; free riders can be punished by a coercive government, and well-organized groups often have the option of conquering weaker or less cohesive groups.

Anthropologists have long debated whether the state arose from either intragroup cooperation or intergroup conflict;56 the present approach suggests that both hypotheses are likely to be correct. Indeed, both processes were probably necessary to overcome the great selective dis- advantage of extending sociality to a society of hundreds of thousands of non-kin (in which mutual benefit cannot be based on personal reciprocity or individual recognition of the free riders). Once established, however, social cooperation within a state can create extensive mutual benefits, as symbolized by the awesome growth in the human population

51Margolis (fn. 28), chap. 9. 52 Michael Laver, The Politics of Private Desire (Harmondsworth, England: Penguin, i98i),

esp. chap. 3. 53 Richard D. Alexander, "Natural Selection and Societal Laws," in H. Tristram Engel-

hardt and Daniel Callahan, eds., Morals, Science, and Society, Vol. III (Hastings-on-Hudson, N.Y.: Hastings Center, I978), chap. 7; Hirshleifer (fn. 46); R. Hardin (fn. 47).

54 Olson (fn. 39); Laver (fn. 52). 55 Donald T. Campbell, "Legal and Primary-Group Social Controls Curbing Selfish and

Nepotistic Human Nature," in Margaret Gruter and Paul Bohannon, eds., Biology, Culture, and Law. (Special Issue of Journal of Social and Biological Structures, in press.)

56 Ronald Cohen and Elman R. Service, eds., Origins of the State (Philadelphia: Institute for the Study of Human Issues, I978).



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that is associated with the spread of centralized political institutions over the last ten thousand years.

Paradoxically, it is the very success of the state that creates its vul- nerability. The legal rules enforced by a government constitute collective goods that are subject to violation in order to reap selfish benefits. Empirical evidence seems to confirm that the decline of prior civilizations is related to, if not caused by, overexploitation of environmental resources- i.e., lack of self-restraint and virtue; with good reason, Garrett Hardin reminds us that the mere existence of the state does not guarantee immunity from the Tragedy of the Commons.57

III. INCLUSIVE FITNESS THEORY AND POLITICAL PHILOSOPHY

It should not be surprising that models of rational decision making can be so easily related to inclusive fitness theory; both approaches deduce social behavior from a calculus of individual costs and benefits. But recent evolutionary theory can be linked to more traditional approaches in political science as well. In Western political philosophy, for example, definitions of human nature and the origin of the state frequently parallel the concepts of evolutionary biology. This proposition can be readily illustrated by considering the thought of some of the best-known modern political theorists.

(A) HOBBES

Hobbes's political theory is easily translated into the terms of inclusive fitness and rational actor models. The Hobbesian "natural condition of mankind" is marked primarily by nepotism or selfishness, since cooperation is essentially limited to the kin-group based on "natural lust." As a result, social interaction readily degenerates into mutual harm ("war of all against all")-unless, for reasons of mutual benefit ("natural right"), individuals are induced to agree ("social contract") to form a political community or state ("commonwealth"). Hobbes thus not only denies that sociality, or virtue, is natural; he is unable to justify any case of self- sacrifice on the basis of natural right alone. Individuals can-and if rational, always will-reclaim their natural independence whenever they fear for their continued safety and self-interest.58

Hobbes's state of nature is like a Prisoner's Dilemma or Tragedy of

57 G. Hardin (fn. 48); Marvin Harris, Cannibals and Kings (New York: Random House, I977). Cf. Machiavelli, Discourses on Titus Livy, I, ii and III, i; Plato, Republic, VIII.546a- 547c; Aristotle, Politics, V.ii.I3o2a-b and V.xii.13i6a-b.

58 Thomas Hobbes, Leviathan, ed. by Michael Oakeshott, I, xiii-xiv (London: Colliers- Macmillan, I 962), 98- I I 2.




the Commons,59 and can be escaped only through a freely accepted arbiter or a sovereign capable of imposing restraints on short-term self-interest. In other words, the Hobbesian solution can be viewed as an n-person game like Figure 3, in which each A strategist agrees with the other A's to accept the rule of a B strategist or strategists who, as "sovereign," is not party to the social contract. Even though such a sovereign may reap a selfish benefit compared to each individual subject, it is rational to form a commonwealth because the result permits the A strategists to avoid the large negative payoffs of mutual harm ("war of all against all") while preserving short-term mutual benefits.

The same cost-benefit calculus persists within society as in the state of nature. Since "no man can transfer, or lay down his right to save himself from death, wounds, and imprisonment," even convicted crim- inals can be expected to try to escape legitimate and just punishment:

For man by nature chooseth the lesser evil, which is danger of death in resisting; rather than the greater, which is certain and present death in not resisting.6o

Pure sociality or virtue benefiting the group as a whole cannot be relied upon, since men are "not bound to hurt themselves," nor are they bound "to warfare, unless they voluntarily undertake it."61

Neither mutual benefit nor sociality (virtue) can exist without a centralized state to induce self-sacrifice through fear.

Therefore before the names of just, and unjust can have place, there must be some coercive power, to compel men equally to the performance of their covenants, by the terror of some punishment, greater than the benefit they expect by the breach of their covenant; and to make good that propriety, which by mutual contract men acquire, in recompense of the universal right they abandon; and such power there is none before the erection of a commonwealth.62

Contract-and hence, presumed mutual benefit-is the only reliable ground on which humans will risk their lives to defend the state and the non-kin who live in it, but even a contract is only binding if there is a sovereign capable of enforcing it.

(B) ROUSSEAU

Rousseau criticizes Hobbes in two different ways, each of which becomes clearer if formulated in terms consistent with evolutionary

59William Ophuls, "Leviathan or Oblivion?" in Herman E. Daly, ed., Toward a Steady- State Economy (San Francisco: Freeman, I973), 2I5-30; McLean (fn. 47).

IOHobbes (fn. 58), I, xiv, p. IIO. 6, Ibid., I, xxi, p. i65. 62 Ibid., I, xv, p. II3-



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biology. First, Rousseau attacks Hobbes for presuming that the family- and therewith a war-like state of nature between kin-groups-is natural.63 Since asocial or purely egoistic behavior is a conceivable means of survival in some species, institutions based on the perpetuation of a bond between parents cannot be taken for granted. And once the family is analyzed as the result of a prior evolution away from asociality, the original state of nature must be presumed to have been "peaceful" and "solitary," at least as a heuristic model or a hypothesis.

Let us conclude that wandering in the forests, without industry, without speech, without domicile, without war and without liaisons, with no need for his fellow-men, likewise with no desire to harm them, perhaps never even recognizing anyone individually, savage man, subject to few passions and self-sufficient, had only the sentiments and intellect suited to that state .. . .64

According to Rousseau, therefore, Hobbes makes the mistake of taking as "natural" a condition that is the result of historical change.65

Rousseau's second major criticism of Hobbes is diametrically opposed to the first: having argued that the original condition of the species was less social than Hobbes assumed, Rousseau also concludes that cultural institutions can be more social or virtuous than the Hobbesian commonwealth. Rousseau's Social Contract can be read as a denunciation of the Hobbesian theory because it focuses on mutual benefit rather than on a patriotic community for which individuals would virtuously sacrifice their lives. Whereas Hobbes limits reciprocity or mutualism to those common institutions that are in the individual's self-interest, Rous- seau seeks a logic of reciprocity ("social contract") that could generate binding obligations constituting the group's collective interest ("general will").

Like Hobbes, Rousseau explicitly treats the agreement to found a society as a calculation of cost and benefit:

Let us reduce the pros and cons to easily compared terms. What man loses by the social contract is his natural freedom and an unlimited right to everything that tempts him and that he can get; what he gains is civil freedom and the proprietorship of everything he possesses.66

But unlike Hobbes, Rousseau insists that consent to enter the political community generates a duty to sacrifice for it i.e., the essential condition of sociality that Rousseau explicitly calls the "virtue" of the "cit-

63 Rousseau, Second Discourse, Part I (fn. i6), esp. I07, I28-30. 64Ibid., I37- 65 Roger D. Masters, The Political Philosophy of Rousseau (Princeton: Princeton University

Press, i968), chap. 3. 66 Jean-Jacques Rousseau, Social Contract, I, viii, in Roger D. Masters, ed., Social Contract,

with Geneva Manuscript and Political Economy (New York: St. Martin's Press, I978), 56.




izen."67 The willingness to die in defense of the state, dismissed by Hobbes as not binding without a specific contractual commitment, becomes a moral obligation of citizenship for Rousseau.68

Rousseau thus seeks to reconstruct an evolutionary theory of the origins of the civilized community, accepting Hobbes's critique of theories that assume natural sociality-but nonetheless achieving a level of communal unity that Hobbes rejects as naturally impossible. Rousseau agrees that one must adopt a social contract model of the origin of the state, but disagrees with Hobbes's conception of human nature both as an original condition and as a limit of political possibility. The para- doxical mixture of individualism and collectivism, which has led many readers to accuse Rousseau of inconsistency,69 can therefore be traced to a single view of human evolution: precisely because humans were not competitive in the original state of nature, they could develop virtue in a freely constituted political community.70

(C) KANT

Kantian ethics, in this view, transform Rousseau's conception of virtue by seeking to remove its limits. Whereas Rousseau's social contract cannot bind anyone who does not share the patriotic spirit symbolized by ancient Sparta and Rome,7' Kant seeks a principle of social obligation that is pure-i.e., free of all empirically determined or emotional limitations.72 Although Kant is often regarded as an abstract metaphysician whose thought is unrelated to natural science (and especially to biology), this interpretation is highly questionable;73 rather, Kantian ethics can be viewed as a scientific response to the "natural" limitations on social cooperation adduced by both Hobbes and Rousseau.

Kant's ethical teaching is based on the observation that the "disposition" or motive of "duty" is never observed unambiguously:

if we attend to the experience of men's conduct, we meet frequent and, as we admit ourselves, just complaints that there is not to be found a single certain example of the disposition to act from pure duty. Although

67 Jean-Jacques Rousseau, First Discourse (fn. i6), 4I-42, 49, 5I-52; Rousseau, Emile, ed. by Allan Bloom, I (New York: Basic Books, I979), 39-40.

68 Rousseau, Social Contract, II, iv (fn. 66), 64. 69 E.g., John Charvet, The Social Problem in the Philosophy of Rousseau (London: Cam-

bridge University Press, I974). 7? Roger D. Masters, "Nothing Fails Like Success: Development and History in Rousseau's

Political Teaching," in Jim MacAdam and others, eds., Trent Rousseau Papers (Ottawa: University of Ottawa Press, i980), 99-Ii8; Masters (fn. i6, I978); Masters (fn. 65).

7' Rousseau, Geneva Manuscript, I, ii, and Political Economy (fn. 66), I57-63, 2i8-20. 72 Immanuel Kant, Metaphysical Foundations of Morals, in Carl J. Friedrich, ed., The

Philosophy of Kant (New York: Modern Library, i964), 140-208. 73 Johan Hjort, The Human Value of Biology (Cambridge: Harvard University Press, I938);

Stent (fn. 30).



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many things are done in conformity to what duty prescribes, it is never- theless always doubtful whether they are done strictly out of duty.74

Kant sees that pure "altruism" would require a motive uncontaminated by self-interest-and the existence of such a motive is "always doubtful." As contemporary social biology indicates, there is reason enough to raise this question with regard to humans as well as to other species.75 Kant answers by denying that morality could ever be deduced from "particular attributes of human nature," insisting that it is only in abstraction from empirical interests and facts that a man-made, rational "imperative" to self-sacrifice is possible: "every empirical element is not only quite incapable of aiding the principle of morality, but is even highly prejudicial to the purity of morals."76 Hence, Kant's categorical imperative is "un- conditional" and "not grounded in any interest" -and thereby sharply different from considerations of "market price" or mutual benefit (i.e., from considerations that could apply to the behavior of other animals).77

Kant thus maintains that there is no naturalistic ground for virtue other than an act of the free human will, imposing on itself a rational obligation as an end in itself. The individual, generalizing his rule of choice ("maxim") into a law governing all rational beings, engages in a logical transformation of the situation that is not unlike Rousseau's "general will"-except that for Kant, the categorical imperative is ad- dressed to all humans as rational beings, not to all fellow-citizens as members of the same (admittedly contingent) political community.78 This means that, for Kant, moral obligation is a pure case of sociality, since the individual acts morally without any expectation of personal gain in the short or long run. Whereas Trivers's "reciprocal altruism" can include elements of mutually beneficial reciprocity and even of nepotism, Kant thus provides a logic of freely willed virtue without any conceivable admixture of self-interest.

(D) HEGEL

Hegel's Philosophy of Right79 is an extraordinary attempt to transcend the divergences between the theoretical perspectives of Hobbes, Rous- seau, and Kant. The First Part, devoted to "Abstract Right," could be

74Kant (fn. 72), 154 (emphases in original). 75 Wispe (fn. 30), esp. chap. 4; Hans Kummer, "Analogs of Morality Among Nonhuman

Primates," in Stent (fn. 30), 31-47. 76 Kant (fn. 72), 174- 77 Ibid., i8o, I82. 78 Rousseau, Political Economy (fn. 66), 2i8-20. 79 Georg Wilhelm Friedrich Hegel, The Philosophy of Right, ed. by T. M. Knox (New

York: Oxford University Press, i964).




viewed as the perspective of Hobbes;80 the Second, dealing with "Mo- rality," focuses on the dimension that Rousseau and especially Kant found lacking in Hobbes;8" and the Third, "Ethical Life," presents Hegel's mode of transcending the contradictions in prior political thought. The Third Part is itself organized in three divisions, corresponding to the main institutional forms described in Table I: "The Family;82 Civil Society, or the market economy;83 and The State.84 Thus the Hegelian dialectic of "right," "morality," and "ethical life" moves from the perspective of the individual's calculus of costs and benefits (rights or claims) to a Kantian notion of pure duty or virtue as uniquely human (morality), and then to human institutions as social systems integrating all consti- tutive levels (ethical life).

Although detailed analysis of Hegel is impossible here, we should note that he explicitly treats both marriage and economic activity-i.e., "needs and the means of satisfying them"-as transformations of "natural" or "animal" functions.85 In contrast, the state, for Hegel, is "a self- dependent organism"; its Constitution is the manifestation, in "the objective world," of the "self-development of the Idea."86 In more familiar usage, the state is a cultural, or man-made, phenomenon; rather than being a transformation of a natural or animal need, the state is thus a purely human construction dependent on the self-conscious activity of its citizens and the evolution of human thought (Hegel's "world mind").

(E) THE RELEVANCE OF POLITICAL PHILOSOPHY

This summary of political philosophy in terms consistent with contemporary social biology could easily be expanded to include the ancients. For example, the pre-Socratics' analysis of individual costs and benefits leads to a social contract theory parallel to that of Hobbes, whereas Plato's Republic offers a model of group selection for sociality by de- stroying both the family and the market economy; Aristotle, like Hegel, develops a complex balance between institutions at the level of family, economic system, and polity.87 Although the specific vocabulary differs, the issues posed by contemporary biological theory are thus fundamentally similar to the traditional questions in Western political thought.

The analysis of human nature, long approached by observation and

80 Ibid., 37-74. 8i Ibid., 75-105. 82 Ibid., 110-22. 83Ibid., 122-55- 84Ibid., 155-223- 85 Ibid., Para. i6i, p. III; Para. 190, p. 127. 86Ibid., Para. 259, p. i6o; Para. 267, p. i63. 87 Roger D. Masters, "Classical Political Philosophy and Contemporary Biology," paper

presented to Conference for the Study of Political Thought, Chicago, Ill., April, 1978.



184 WORLD POLITICS

insight, can thus be illuminated by scientific research. As Trivers has indicated, the logic of inclusive fitness explains such supposedly "selfish" behaviors as parent-offspring conflict, sibling rivalry, and a host of other frequently observed human traits.88 But, in recognizing the ubiquity of egoistic motives and behaviors, evolutionary biology also points to the countervailing tendencies toward cooperation, particularly in a species that experiences prolonged helplessness in infancy, predators in its environment, and scarcity of food requiring cooperative hunting and food sharing. From the earliest epochs, therefore, we can assume that human nature exhibited a complex of both cooperation and competition.89

In theoretical terms, this means that Hobbes's definition of the state of nature as a mutually harmful "war of all" among selfish individuals, like Rousseau's assertion of human asociality or Kant's formulation of a pure and universal duty of virtue, cannot fully describe the origins of the state. That is tantamount to saying that the accounts of human nature and political institutions found in Hobbes, Rousseau, or Kant are only partially correct heuristic models: each is true as a formal description of human behavior under some circumstances; none is a global definition of the unchanging basis of human nature.

The fact that the major political thinkers of the past have elaborated systems that are incomplete or partial should not be taken to indicate that traditional political theory is irrelevant or useless. Each of the major theorists discussed above analyzes a particular issue or transformation with archetypical clarity. Of the five conceivable modes of behavior- asociality, nepotism, mutual benefit, sociality, and mutual harm-different philosophers have tended to isolate specific situations and to ex- plore with great depth how they relate to political experience. For example, Hobbes shows how nepotism, under circumstances producing mutual harm, can be transformed into mutual benefit only by the construction of a man-made "commonwealth." Rousseau spells out how humans could have evolved from an asocial condition (the "pure state of nature") into a Hobbesian condition of mutual harm, as well as how a political community could go beyond the mutually beneficial contracts of Hobbes to a patriotic community united by feeling and civic virtue. And Kant indicates how even such a form of virtue would fall short of the pure demands of a moral obligation, thus suggesting an inevitable tension between private self-interest and group interests. From the perspective of evolutionary biology, it is therefore evident that, contrary to

88 Trivers (fn. 25)- 89 Roger D. Masters, "Nature, Human Nature, and Political Thought," in J. Roland

Pennock and John Chapman, eds., Human Nature in Politics (New York: New York Uni- versity Press, 1977), 69-iio; Edgar Morin, Le Paradigme Perdu (Paris: Seuil, 1973).




the claim of "historicists," the great political theorists address the per- ennial issues of human social life.90

IV. THE EMERGENCE OF THE STATE Political philosophers usually treat the emergence of the state as a

central problem, fundamentally different from anything in the animal world. Contemporary evolutionary biologists agree, pointing to the limitations in the scale of social cooperation in other sexually reproducing mammals. The sheer size of political communities, and their inclusion of heterogeneous populations of different ethnic groups and classes, suggests that something unique was necessary to produce what is di- versely called human "civilization," "the state," or "euculture."9'

For most of the last 3.5 million years our ancestors lived in relatively small bands, presumably structured in groups of extended kin (like many primate species and human hunter-gatherer tribes). Exogamy, reinforced by the incest taboo, probably resulted in an exchange of mates between lineages or groups, thereby encouraging conscious norms of reciprocity in social interactions. Hominids seem to have evolved in groups favoring nepotism or kin selection-a structure that greatly increases the probability of cooperation within the group-reinforced by "altruistic" motives toward group members and hostility to outsiders.92 But since cooperation among millions of non-kin cannot be explained by "human nature" alone, the origin of the state poses the following scientific question: What circumstances in the last ten thousand years permitted the formation of very large groups-especially since the establishment and maintenance of such societies (let alone the formation of political systems to run them) were previously impossible or unnecessary?

In the anthropological literature on the origins of the state, two main hypotheses have been presented.93 In the first view, the state-organizing and controlling human populations-arose as the result of internal processes within prestate societies; in the second, it is conflict between groups that generated states. Is new light cast on the problem by considering the work of evolutionary biologists?

According to contemporary evolutionary theory, so-called "group se-

90 Leo Strauss, Natural Right and History (Chicago: University of Chicago Press, 1953). 91 A. L. Kroeber, The Nature of Culture (Chicago: University of Chicago Press, 1952);

Samuel M. Hines, Jr., "The Origins of the State," in Heiner Flohr and Wolfgang Tom- mesmann, eds., Politik und Biologie (Berlin: Verlag Paul Parey, in press); Charles J. Lumsden and Edward 0. Wilson, Genes, Mind, and Culture (Cambridge: Harvard University Press, i98i).

92 Robin Fox, "Primate Kin and Human Kinship," in Fox (fn. 8), 9-35; Leakey and Lewin (fn. i), chap. 5; Alexander (fn. I3); Masters (fn. 30); Morin (fn. 89).

93 Cohen and Service (fn. 56).



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lection" is not typically expected; in general, social cooperation will develop only when the actor's own kin ultimately benefit through reciprocity-and this factor can be expected to limit the size of groups to that in which individuals have a reasonable probability of recognition or role reversal in future encounters.94

There are, however, some restrictive circumstances in which this individualistic calculus does not hold. Where a number of small populations of extended kin are relatively isolated from each other, conflicts between groups can and will occur if resources are insufficient to support all of them. In that case, a behavior that benefits the group-even at a cost to the actor-can be favored by natural selection whenever competition within the group is harmful to both the individual's "inclusive fitness" and the group's "collective interest." Under these circumstances (which Alexander describes as a "balance of power" between competing bands of extended kin), group selection would expand the sphere of social cooperation.s5 As Benjamin Franklin put it, "We must all hang together, or assuredly we shall all hang separately."

Fred Willhoite has shown how this "balance of power" hypothesis can explain the progressive evolution of hominids through the stages of the band (exemplified by the !Kung bushmen), the sovereign village (e.g., Yanomomo), the chiefdom as a multi-community, ranked society based on extended kin-groups or lineages (like so many of the preliterate societies studied by anthropologists), and finally the early state (ranging historically from ancient Egypt to i9th-century Africa). At each level, as communities expanded to include more distant kin or nonrelated kin-groups, the role of coercive institutions also expanded. And clearly, there is some evidence that the earliest civilizations evolved out of rivalry and conflict.96

An alternative model of the origins of the state focuses on the new environmental or productive resources created by large-scale cooperation. This model only works, however, if some new "collective goods" are available from which individuals can benefit relatively quickly- indeed, more rapidly than the increments to inclusive fitness that would follow from either nepotism (refusal to cooperate with non-kin) or an

94 Wilson (fn. 2), chap. 5; Philip R. Thompson, "And Who is My Neighbor? An Answer from Evolutionary Genetics," Social Science Information, xix (No. 2, ig80), 341-84.

95 Alexander (fn. 53). Cf. Robert L. Carneiro, "A Theory of the Origin of the State," Science, Vol. i69 (August 2i, 1970), 733-38.

96 Willhoite, "Reciprocity, Political Origins, and Legitimacy," paper presented at 76th Annual Meeting, American Political Science Association, Washington, D.C., ig80; Willhoite (fn. 3i); Alexander Rustow, Freedom and Domination (Princeton: Princeton University Press, 1980).




insistence on individual reciprocity. The development of extensive ir- rigation systems, forming the basis of what Wittfogel calls "hydraulic" empires,97 would be an example of such a situation.

Eisenstadt's analysis of the emergence of early empires, for instance, associates the rise of a centralized bureaucracy-necessary to establish a lasting political tie by enforcing the commands of the state-with the existence of productive surpluses and socioeconomic differentiation.s8 At least one careful study of the emergence of a "primary state" in high antiquity finds that the population explosion took place after the first development of the state's institutions-not before, as might have been predicted from a balance-of-power hypothesis.99 Since history is littered with tales of Pyrrhic victories and futile conquests (like those of Alex- ander the Great), it is likely that both intergroup competition and intragroup cooperation played a role in the origin of the state.

In an interactive model, intergroup competition places a premium on within-group cooperation-and within-group cooperation leads to greater intergroup competition; only in this way could the free-rider effects of nepotism, which have generally precluded the emergence of large societies and states, be overcome. In particular, it is the interaction between the existence of tangible benefits (increased quantities and reliability of resources, made possible by compulsory economic cooperation) and the threat of outsiders seeking to gain these benefits, that permits the use of symbolic appeals to unity and virtue within the community. The result is a process of "moralizing aggression," which reinforces cooperation by creating feelings of guilt in the potential cheater and of "righteousness" in the cooperators, as well as by meting out legitimate punishment for the guilty who are caught.Ioo

In this model, since the institutionalized state can only occur under a narrow set of conditions, time would be a critical factor. If the combined effects of intergroup competition and within-group cooperation favored the formation of a state only during a short period, these unique "occasions" (to use Machiavelli's term) may have given play to the leadership qualities of particular individuals. Hence, in an interactive model of the origin of the state, there is even a role for the "great man" in history.

The "critical period" hypothesis is an important part of the model,

97 Karl Wittfogel, Oriental Despotism (New Haven: Yale University Press, 1957). For an example in the New World, see B. L. Turner, II, and P. D. Harrison, "Prehistoric Raised- Field Agriculture in the Maya Lowlands," Science, Vol. 213 (July 24, i98i), 399-405.

98 S. N. Eisenstadt, The Political Systems of Empires (London: Free Press of Glencoe, i963). 99 Cohen and Service (fn. 56), 6i. .oo Trivers (fn. 25); Willhoite (fn. 31); Peter Corning, The Synergism Hypothesis: A Theory

of Progressive Evolution (New York: McGraw-Hill, in press).



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because it helps to explain how the free-rider problem was overcome. Although undetected nepotism pays within the state, at its formation cheating behavior would be reduced in frequency because it would be visible at the decisive moment. When the battle starts, those who stay in their tents rather than fight are usually obvious (Achilles in the Iliad); those who built the pyramids of ancient Egypt-the epitome of cooperative endeavor within an early state-engraved "tally marks on the casing stones" in order to "give the titles of the individual work teams who were to be credited with the supply."IoI

Governments and laws thus seem to have made accessible the benefits of cooperation among non-kin in larger-scale societies. At the same time, those who provide their fellows with the "services" of organizing social cooperation and catching cheaters are presumably as subject to evolutionary principles as those who obey (or disobey) the laws. Governing officials everywhere reap benefits for their positions-prestige, power, as well as access to females and the likelihood that offspring will gain privileged treatment. Indeed, inclusive fitness theory leads us to predict that envy or jealously will be widespread, and will ultimately destroy any political elite incapable of providing the many with benefits which exceed the relative costs of supporting the political system.

Unlike traditional theories, the evolutionary approach sketched here does not presume either that the state originated directly from a priori traits of human nature or that it resulted solely from historical circumstances. Rather, the human capacity to institutionalize new modes of improving inclusive fitness, when combined with appropriate external conditions-and perhaps only in the presence of individuals capable of seizing the opportunity-would explain the specific times and places in which states have arisen. For example, the joint role of international rivalry and economic development is particularly evident in the origins of the modern state system in the I7th and i8th centuries-as well as in the city-states of classical antiquity and Renaissance Italy.

Once they have been established, however, states remain subject to the dialectical relationship between individual and group interests. As the Tragedy of the Commons indicates (Figure 3b), it is always tempting to seek to combine the benefits of collective goods (based on the virtue of others) with the selective benefits of nepotism or selfishness. For every law there is always a new loophole; for every new abuse, a possible

... Homer, Iliad, Book IX; Kurt Mendelssohn, The Riddle of the Pyramids (New York: Praeger, 1975), 148. For a striking illustration, see also Exod. 33:21-29. On the origin of writing as a system for ensuring credit for tax payments, see Denise Schmandt-Besserat, "Decipherment of the Earliest Tablets," Science, Vol. 22i (January i6, i98i), 283-85.




legislative response. Periodically, it is necessary to reinvigorate the state; even a cursory glance at recorded history reveals the extraordinary in- stability of political systems.

Paradoxically, the state is vulnerable over the long run precisely because it is so successful in generating mutual benefits and collective goods in the short time horizon of an individual's conscious decision making. The very outcomes that represent the strength of political communities can thus be the seeds of their decay. Especially if further research confirms the soundness of this approach, evolutionary biology can provide a framework that greatly illuminates the future of industrial society as well as the origin of the state.'02

V. CONCLUSION: THREE REASONS FOR LINKING POLITICAL THEORY AND

EVOLUTIONARY BIOLOGY

There are three broad reasons for developing linkages between biology and political science along the lines suggested above. First, the establishment of a unifying foundation for the varied perspectives within contemporary social science will do much to improve research. Second, in empirical studies of politics, variables and approaches similar to those in evolutionary biology could reveal hitherto unknown relationships of great importance. Third, and most important, evolutionary biology has implications for public policy and ethical judgment.

I. Unfying the discipline of political science and relating it to the natural sciences, which has long been a dream,103 now seems possible. Although cost-benefit theories (whether game theory, conventional economics, or decision-making models) need not be modified fundamentally in order to be linked with inclusive fitness theory, by so doing one gains a heightened awareness of the domain within which collective decisions can be derived from the aggregation of individual choices based on a cost-benefit calculus.104 Political philosophy, long divorced from the "hard" sciences, can only be strengthened by the realization that famous thinkers describe, with archetypical clarity, the categories and transformations of behavior that are central to a scientific understanding of human political

102 Roger D. Masters, "Social Biology and the Welfare State," in Richard F. Tomasson, ed., Comparative Social Research, Vol. VI (Greenwich, Conn.: JAI Press, i983), in press.

03 Karl Marx, Economic and Philosophic Manuscripts of i844, ed. by Dirk J. Struik (New York: International, 1964), 143.

04 For a concrete example, compare Hirschman (fn. 39) with Roger D. Masters, "Exit, Voice, and Loyalty in Animal and Human Behavior," Social Science Information,.xv (De- cember 1976), 78-85. On the limits of individual cost-benefit calculus, see Masters (fn. 26); Alexander and Borgia (fn. 27).



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experience.105 And the characteristic empirical theories of the state, os- cillating between models of conflict and of functional integration, can now be related and combined in a more comprehensive approach to the study of political and social history.

2. New and more significant empirical research should, of course, be a goal of anyone claiming to be engaged in a scientific endeavor. William Durham has presented the first compelling case of a cultural prohibition whose distribution and timing were the result of natural selection. o6 In West Africa, many tribes traditionally had a taboo on the consumption of newly ripe yams before a specific festival (even though other foods were in short supply at this season). It has been claimed that consumption of these yams could counteract the sickling effects in hemoglobin, making individuals with the sickle cell gene both more vulnerable to malaria and less vulnerable to sickling crises; this hypothesis explains both the t

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