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Trustees of Princeton University
The Biological Nature of the State

Author(s): Roger D. Masters
Source: World Politics, Vol. 35, No. 2 (Jan., 1983), pp. 161-193
Published by: Cambridge University Press
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THE BIOLOGICAL NATURE OF

THE STATE
By ROGER D. MASTERS*
HY do humanbeingslivein societiesconsisting
ofhundredsof
V V thousands,if not millions,of individualswho owe obedience to

the state? This question, long at the centerof political philosophy,has
been given renewedimportancebythefindingsofcontemporarybiology:
froman evolutionaryperspective,theveryexistenceof legal and political
institutionsis far more problematicthan is generallyassumed. Paradoxically enough, however, the origin and nature of the state can be
illuminatedby the theoryof natural selection.
For over three million years, our hominid ancestorslived in small
bands,probablyapproximatinga hundredin number.'Most othermammals-and notablyour closestrelatives,the primates also formgroups
thatare severalordersof magnitudesmallerthancontemporary
societies.
In such small, face-to-facebands, the extensivedivision of labor and
social stratification
connectedwith a formalstatedo not normallyexist.
Recent models of natural selectionindicatethatsexuallyreproducing
animals such as Homo sapiensare unlikelyto formenormous,impersonal
social systemsin which individuals are called upon to sacrificetime,
money,and sometimeseven theirlives forthebenefitof thecommunity.2
Because contemporarynation-statesare so much largerthan the groups
observed in other mammals, it is appropriateto seek an evolutionary
explanation for human legal and political institutions.
To avoid confusion,however,the use of biological conceptsin social
science must be based on careful specificationon the level of analysis.
Scientificcomparisons,unlike organicmetaphors,mustdistinguishcarefullybetween evolutionaryprocessesat the species level, social behavior
in groups, and individual physiologicaland neurologicalfunctions.3
As
*
This articleis a greatlyexpanded and revisedversionof a papergivenat theSymposium
on Law and Behavioral Research,MontereyDunes, Calif., September25-27,
5981.
C. Owen Lovejoy, "The Origin of Man," Science,Vol. 22I (January23, i981), 34I-50;
Richard E. Leakey and Roger Lewin, Origins(New York: Dutton, I977).
2Edward 0. Wilson, Sociobiology(Cambridge: Harvard UniversityPress, 1975); David
Barash,Sociobiologyand Behavior(New York: Elsevier, I977).
3 RogerD. Masters,"FunctionalApproachesto AnalogicalComparisonsbetweenSpecies,"
in Mario von Cranach,ed., Methodsof ComparingAnimaland Human Behavior(The Hague:
Mouton, I976), 73-I02-
? i983 by the Trustees of PrincetonUniversity

WorldPolitics0043-887I/83/02016I-33$OI.65/I
For copyinginformation,see contributorpage.
WORLD POLITICS
162
used here, the conceptof evolutionconcernsonly the processof natural

selectionat thelevelofpopulations,and can referto individuallylearned,
culturally transmitted,or geneticallyprogrammed behavior.4In this
paper, therefore,no assumptionswill be made concerningthe extentif any-of genetic constraintson human behavior.
Since social scientistshave long assumed that humans are "unique,"
they have tended to ignore research on other species. It is becoming
increasinglyevident,however, that the separationof biology from the
social sciences is unwise.5 Despite charges that "sociobiology" is ideountenable,6the neo-Darwinian theory
logicallybiased and scientifically
of natural selectionhas been shown to be compatiblewith diverse political conclusions.7Substantively,evolutionarybiology has begun to
and
economics,9sociology,bo
enrich such diverse fieldsas anthropology,8
political science."
I will begin this essay by showing how evolutionarytheorycan be
4 Donald T. Campbell, "Variation and SelectiveRetentionin Socio-culturalEvolution,"
in HerbertR. Barringer,George I. Blanksten,and RaymondW. Mack, eds., Social Change
in DevelopingAreas(Cambridge,Mass.: Schenkman,I965), I9-48; RogerD. Masters,"Politics
xiv (April I975), 7-63; Richard D.
as a Biological Phenomenon,"Social ScienceInformation,
Alexander, "Evolution, Human Behavior,and Determinism,"Proceedingsof the Biennial
Meetingof thePhilosophyof ScienceAssociation,
11(977), 3-2I.
5Alexander Rosenberg,Sociobiology
and thePreemption
of Social Science(Baltimore:The
JohnsHopkins Press, i980); Richard D. Alexander,"Evolution,Culture,and Human Behavior: Some General Considerations,"in Richard D. Alexander and Donald W. Tinkle,
eds., Natural Selectionand Social Behavior:RecentResearchand Theory(New York: Chiron
Press, i982), 504-20; John C. Wahlke, "Prebehavioralismin Political Science," American
PoliticalScienceReview,Vol. 73 (March I979), 9-3I.
6 Marshall Sahlins, Use and Abuseof Biology(Ann Arbor: University
of Michigan Press,
I976); ArthurCaplan, ed., The Sociobiology
Debate (New York: Harper & Row, I978).
7 Roger D. Masters,"Is SociobiologyReactionary?The PoliticalImplicationsof Inclusive
Fitness Theory," QuarterlyReview of Biology,Vol. 57 (September i982), 275-92; Marvin
Bressler,"Biological Determinismand Ideological Indeterminacy,"in Elliott White, ed.,
Sociobiologyand Human Politics (Lexington,Mass.: Lexington Books, i98i), i8i-9i. For
historicalperspective,
see also Loren Graham,"Scienceand Values: The Eugenics Movement
in Germanyand Russia in the I92os," AmericanHistoricalReview,Vol. 82 (December I977),
II33-64-
8 Robin Fox, ed., BiosocialAnthropology

(London: Malaby, I975); Napoleon Chagnon and
William Irons, eds., EvolutionaryBiologyand Human Social Behavior(N. Scituate,Mass.:
Duxbury, I979); JoanS. Lockard, ed., The EvolutionofHuman Social Behavior(New York:
Elsevier, i980); Pierre van den Berghe & David Barash, "Inclusive Fitness and Family
Vol. 79 (No. 4, I977), 809-23.
Structure,"AmericanAnthropologist,
9 JackHirshleifer,"Natural Economy vs. Political Economy,"Journalof Social and BioI (October I978), 3I9-37; Ivan Chase, "Cooperativeand Noncooperative
logical Structures,
Behaviorin Animals,"AmericanNaturalist,Vol. II5 (JuneI980), 827-57; Howard Margolis,
"A New Model of Rational Choice," Ethics,Vol. 9I (JanuaryI98I), 265-79.
10Lionel Tiger, Men in Groups(New York: Random House, I969); Allan Mazur, "Biosociology,"in JamesF. Short,Jr.,ed., The State of Sociology(BeverlyHills, Calif.: Sage,
I98I), I4I-60.
AlbertSomit,ed., Biologyand Politics(The Hague: Mouton, I976); Thomas C. Wiegele,

Biopolitics(Boulder,Colo.: Westview,I979); White (fn.7); MeredithWatts,ed., Biopolitics:
Ethologicaland PhysiologicalApproaches,New Directions for Methodologyof Social and
Behavioral Science, No. 7 (San Francisco: Jossey-Bass,I98I).
BIOLOGICAL NATURE OF THE STATE
163
withoutengagingin genetic
extendedto studyhumansocialinstitutions
between
reductionism
(SectionI). Next, I will surveythe relationship
modelsof naturalselectionand "rationalactor"theoriesof
cost-benefit
humanbehavior(SectionII), politicalphilosophy
(SectionIII), and empiricalresearchon the originof the state(SectionIV). On thisbasis,I
of evolutionary
will reassessthe potentialcontribution
biologyto the
studyof politics(SectionV).
I. INCLUSIVE FITNESs THEORY AND HUMAN
SOCIAL BEHAVIOR
neo-Darwiniantheoryrequiresa fundamental
shiftin
Contemporary
of social behavior.Politicalphilosophers
our understanding
and social
scientists
usuallytreatthe questionof "sociability"-i.e.,whetherhumans are naturallyselfishor cooperative-as an attribute
of "human
nature."But biologynow teachesus thattheexistenceand structure
of
social groupsvaries,even withina singlespecies,as a consequenceof
the interaction
betweenanimalsand theirenvironment;
a speciesthat
is asocial in one settingmay be social in others.12 From a scientific
it is misleadingto conceptualizesocial behavior
perspective,
therefore,
as a constantnatural 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 differencesbetween
human cultures-is thus inadequate: the emergence of large-scale human societies and formal legal systemsmust be treated as a biological
problem.'3 The usual answers to this problem violate the premises of
evolutionarytheorybecause they assume that the state is beneficialto
the group, resultsfrom chance, or is an act of human creation;'4none
of these conventional explanations is tenable without a careful consideration of the selective pressures against institutionslike the state-in
other animals, and during most of hominid evolution. The approach
known as "inclusive fitnesstheory"suggestsan alternativethat is con-
Marc Beckoffand Michael C. Wells, "The Social Ecology of Coyotes,"Scientific

American, Vol. 242 (April i980), I30-48; Richard Wrangham, "Review Essay: Sociobiology,"
BiologicalJournalof theLinnean Society,xiii (March i980), 171.
'3 Richard D. Alexander, "Natural Selection and the Analysis of Human Sociality,"in
C. E. Goulden, ed., ChangingScenesin Natural Sciences,I776-I976 (Special Publication I2,
Philadelphia: Philadelphia Academy of Natural Sciences, I977), 283-337; Donald T. Campbell,"On theGeneticsof Altruismand theCounter-hedonic
Componentsin Human Culture,"
Journalof 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,
Discourseson Titus Liuy, ed. by Bernard Crick, I, ii and I, ix (Harmondsworth,England:
Penguin,1970),
104-II,
I3I-34-
164
WORLD POLITICS
sistentwith the widely observed tendenciesof animals to adjust their

behavior to maximize long-range reproductivesuccess.
Cooperation generallyentails some cost to the individual. To understand animal social behavior, biologiststhereforehave to discover how
individuals gain from cooperation and to show that these benefits
exceed the costs. For example, under conditionsof abundant and continuouslyavailable resources,animals not subject to predation tend to
be asocial. In such situations,there is virtuallyno benefitfrom cooperation that overbalances its costs in energyor time.'5Hence, in species
like the lepilemur or the orangutan, one finds asocial behavior that
resemblesRousseau's "state of nature" to a remarkabledegree.'6
According to the prevailinginterpretationof inclusivefitnesstheory,
natural selection operates primarily-though not necessarilysolely-at
the level of the individual.'7 If two organisms interact,whetheror not
theycooperate can usually be predictedby a cost-benefitcalculus 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 extraordinarynumber of species.
As in rationalactor models in economic theory-particularlyas applied
to questions of collective goods or public choice-all that need be assumed is that behavior will vary as if it were a strategychosen on the
basis of cost-benefitoptimization.,8
In his well-known articleintroducingthe conceptof inclusivefitness,
William Hamilton proposesfourgeneral "classes" or "typesof behavior"
definedin such cost-benefitterms.'9When organismsinteract,the actor
can derive eithera net benefit(+) or a net loss (-) fromhis behavior,
and "neighbors"-i.e., other organisms involved in the social interaction-can likewise derive eithera net benefitor a net loss. The resulting
four-foldtable, rewrittento stressbehavioral consequences ratherthan
1Barash (fn. 2), 57-62; Hans Kummer, "On the Value of Social Relationshipsto Nonhuman Primates:A HeuristicScheme," in Mario von Cranach and others,Human Ethology
(Cambridge,
England:CambridgeUniversity
Press,I979),
6
38I-95.
Jean-JacquesRousseau, Second Discourse,Part I and note j, in Roger D. Masters,ed.,
Rousseau's
FirstandSecondDiscourses
(New York:St. Martin'sPress,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-Jacquesis Alive and
Well: Rousseau and ContemporarySociobiology,"Daedalus, Vol. 107 (Summer 1978), 93I05.
'7George C. Williams,Adaptationand Natural Selection(Princeton:PrincetonUniversity

Press, i966).
,8 JohnMaynard-Smith,
"The Evolution of Behavior,"Scientific
American,Vol. 239 (September1978), 176-92;Barash(fn.2), 51-52; Hirshleifer
(fn.9); Chase(fn.9).On thedistinction
between "ultimate" and "proximate"causation,see Barash (fn. 2), 37-39.
'9 Hamilton, "The GeneticalEvolution of Social Behavior,"Journalof Theoretical
Biology,
vii (No. I, i964), i-i6; reprintedin Caplan (fn. 6), I9I-209.
Cf. Barash (fn. 2), 96.
165
has general relevance as a means of linking evsubjective intention,20

olutionarybiology to the studyof human institutions(Figure I).
FIGURE
HAMILTON
S 'FOUR
TYPES
OF BEHAVIOR"
A
GainI
Gain
l
Loss
LossI
OR
MUTUAL
BENEFIT SOCIALITY
/VIRTUE
-
NEPOTISM
MUTUAL
HARM
SOURCE: ModifiedfromHamilton (fn. I9), 207.
This approach analyzes social behavior in termsof each individual's

benefitsand costs,on the assumptionthatobservableoutcomesestablish
analysis
the net utilityof the act. In economic theory,such cost-benefit
is usually measured in monetaryor materialistic(economic) terms.Evolutionarybiology substituteslong-rangereproductivesuccess-i.e., the
proportionof an individual's genes transmittedto futuregenerationsof
the species. Inclusive fitnessthus differsfromthe traditionalnotion of
"survivalof the fittest"in two importantrespects:first,naturalselection
favorstheabilityof individualsto transmittheirgenes to posterity(rather
than their"fitness"in termsof health,power, beauty,or otherphysical
traits); second, an organism's reproductivesuccess can sometimes be
furtheredby assistingothers,instead of by mating.21
Social biologistscall the lower leftcell of Figure I nepotism,or kin
selection,because the actor's positivepayoff-a largershare of genes, in
comparison to others, in futuregenerations necessarily involves faOn the importanceof this distinction,see Gunther Stent,"You Can Take the Ethics
out of Altruism,But You Can't Take the Altruismout of Ethics,"HastingsCenterReport,
VII (I977), 33-36; Roger D. Masters,"Beyond Reductionism,"in von Cranach (fn. I5), 26520
84.
On the "coefficient
of relatedness"-i.e., "the fractionof genes in two individualsthat
are identicalby descent,averaged over all loci"-as a measure of reproductivesuccess,see
Wilson (fn. 2), 74. For examples of non-matingstrategies,see also Barash (fn. 2), esp. 9193.
21
166
WORLD POLITICS
voring close kin as compared to more distant relatives or unrelated

individuals. In contrast,the upper leftcell-mutual benefit(oftencalled
mutualismor reciprocity)-refersto situationsin which two presumably
unrelated individuals both gain from an action; among animals, such
an exchange of benefitsoften restson a mechanism of recognitionor
reciprocitythatdiscriminatesagainstnoncooperative"freeriders."22 Mutual harm-the lower rightcell of Figure i-can occur when organisms
gain more fromdenyingthemselvesthe use of a beneficialresourcethan
from running the risk that it be preempted by rivals;23when such
interactionspredominate,a species will tend to be asocial.
Since nepotism,mutual benefit,and mutual harm all referto selfinterestedor "egoistic" behavior,all can be readilyexplained-depending on circumstances-as results of natural selection.24Situations in
which an animal incurs a relativeloss while potentialcompetitorsgain
(theupper rightcell in Figure i) seem harderto derivefromevolutionary
theory.Such group benefitsor "giving" behaviorsare instancesof social
cooperationbetweennon-kinwithoutpersonalreciprocity;
anythingthat
benefitsthe actor's kin is at least equally helpfulto reproductiverivals
and theiroffspring.Since the term "altruism,"as used by many sociobiologists,confusescost-benefit
outcomesand motivations,it is preferable
to call thiscategorysocialityor-to apply an old-fashionedterm-virtue.
The cells in Figure i can also be describedas fourcategoriesof social
partners:kin(others benefitingfromnepotisticbehavior),allies (others
sharing mutual benefits),non-kin(others benefitingby socialityor the
mere factof group membership),and enemies(othersexcluded fromthe
group thatis sharingbenefits,even at the riskof mutual harm). Whether
a human action falls in one or another of these four cells-and hence,
whetheran individual responds to the "other" as kin, as ally, as nonkin group member,or as enemy-depends on environmentalconditions.
At any time,moreover,humans may finda benefitin forgoingall social
interaction,retreatinginto asocial isolation. According to the circumstances,the same behavior-be it eating or defenseagainsta predatorcan thus be entirelyasocial or can generateshared benefitsfor kin, for
allies, or for all group members.
RobertTrivers,"The Evolution of ReciprocalAltruism,"QuarterlyReview of Biology,
(No. 4, I971), 35-5723Raymond Pierotti,"Spite and Altruismin Gulls," AmericanNaturalist,Vol. I IS (February1980), 290-300; Wilson (fn. 2), II9.
24 Richard D. Alexander,"Evolution of Social Behavior,"Annual Review of Ecologyand
v ('974), 25-83; Richard Dawkins, The SelfishGene (New York: Oxford UniSystematics,
versityPress, I976).
22
XLVI
167
Sincecontemporary
theoriesof naturalselectionstressthepriority
of
individual
reproductive
one can neverpresumethata behavior
success,25
has beenselectedbecauseit redoundsto thebenefit
of thepopulation
or
species.Butitis equallyerroneous
toassumethat,as a matter
ofprinciple,
naturalselectionnevergeneratesbehaviorof thissort.26Indeed,much
recenttheoretical
work suggestsa dialecticalrelationship
betweenindividualinclusivefitness
and groupinterest,27
in a complex
particularly
socialspecieslikeHomo sapiens.28
Manyspecieshavespecializedin one or twoofthefivepossiblekinds
of behavior(asociality,nepotism,mutual benefit,sociality,and mutual
harm). Butifenvironments
a diverseand more
changeor are uncertain,
plastic repertoirewill be selected.29Humans seem to have exploited a
broader range of social alternativesthan most other animals, thanks to
our larger brains and complex linguisticsystems.Moreover,the emergence of mutual benefitor sociality,not to mentionoccasions of mutual
harm, in no way abolishes the adaptivenessof nepotismor- in appropriate conditions-asociality. In conventionalterms,humans combine
"selfishness"
and "self-interested"
cooperationwith "altruism"and "spite"
to an oftenbewilderingdegree.30
Social biology thus gives us a more precise understandingof the
differencebetween human social behavior and thatof othergregarious
animals. Inclusivefitnesstheoryleads to a predictionthatthe likelycosts
and benefitsof alternativebehaviorswill vary,depending on ecological
circumstancesand the individual's social role, prior experience,age, or
sex. Humans, especially because of their intelligence,are capable of
exploitingmany if not all of the possible alternativesin a single day, or
even a single hour. Moreover,socialityor virtueamong non-kincreates
Williams (fn. I7); RobertTrivers,"Sociobiologyand Politics,"in White (fn. 7), 4-7.

David Sloan Wilson,The NaturalSelectionofPopulationsand Communities
(Menlo Park,
Calif.: Benjamin Cummings, I979); Roger D. Masters,"The Value-and Limitations-of
Sociobiology,"in White (fn. 7), I35-65.
Richard D. Alexander and Gerald Borgia, "Group Selection,Altruism,and the Levels
of the Organization of Life," Annual Review of Ecologyand Systematics,
IX (1978), 449-74;
David Layzer, "Altruismand Natural Selection,"JournalofSocial and BiologicalStructures,
I (JulyI978), 297-305;
Michael J.Wade, "Kin Selection:Its Components,"Science,Vol. 2i0
(November 7, i980), 665-67.
Martin L. Hoffman,"Is AltruismPart of Human Nature?" Journalof Personality
and
25
26
27
28
SocialPsychology,
XL (January
1981),
I2I-37;
Howard Margolis,Selfishness,
Altruism,
and
Rationality(Cambridge,England: Cambridge UniversityPress, 1982).

Leslie A. Real, "Fitness,Uncertainty,
and the Role of Diversificationin Evolution and
Behavior,"AmericanNaturalist,Vol. I I5 (May 1980), 623-38.
30 Roger D. Masters,"Of Marmotsand Men: Human Altruismand Animal Behavior,"
in Lauren Wispe, ed., Altruism,
Sympathy,
and Helping (New York: Academic Press, 1978),
59-77; Gunther Stent, ed., Moralityas a Biological Phenomenon(Berkeley: Universityof
29
California
Press,1979).
168
WORLD
POLITICS
a situationin whichcheating-i.e.,pretending
to cooperatewhile actuallyengagingin nepotism("selfish"behavior)-has greaterpayoffs
than reciprocity,
provided,of course,thatit goes undetected(Plato,
Republic,II.359b-36od).3'
Humansmanipulatethisrepertoire
ofsocialpossibilities
bymeansof
a dual transformation
of the cost-benefit
calculusobservedin animal
behavior.At the individuallevel,the assessmentof the interpersonal
situation(i.e., the categoriesin Figure i) is transformed-usually
in a
consciousway,but sometimesthroughrepression
or self-deceptionintoa subjective
motive.Thesepsychological
responses
arefurther
molded
intosociocultural
institutions
which,whilesuperficially
associatedwith
a corresponding
psychological
motive,actuallyuse diverseassessments
of theworld(Table i). Sinceeach biologicalsituationcan generateany
of thefivemotivational
responses-andeach of theindividualmotives
can producebehaviorsat anyof thefiveinstitutional
levels-a reductionistscienceof humanbehavioris highlyimprobable,
if not impossible.32
Languageis, of course,whatenableshumansto engagein thisdual

transformation
of naturalpotentialities.
In otherspecies,intention
can
be inferred
fromspecies-specific
behaviorsthathavebeenritualizedinto
whatLorenz has called releasingmechanisms.33
Whatevertheinternal
eventsin otheranimals,theydo not seem to servesimultaneously
as
subjectivestatesand as objectivesocialsignalsor symbols.In contrast,
humanlanguagepermitsthisduality,sincewordscan be usedeitherin
silentthoughtor in audiblespeech.Probablygroundedin theunusual
structure
ofphonemicutterances,
speechcan thusestablish
culturalrules
and socialinstitutions
basedon sharedmeaning.34
To putitmoresimply,
languageenableshumansto establishculturesto a degreeapparently
impossiblein otheranimals(Aristotle,
Politics,
I.ii.I253a; Hobbes,Leviathan,I, iv).
We are now in a betterpositionto linkinclusivefitness
theorywith
theconventional
socialsciences.Humaninstitutions,
intheirbewildering
P Trivers (fn. 25), 9-39; Fred Willhoite,"Rank and Reciprocity:
Speculationson Human
Emotions and Political Life," in White (fn. 7), 239-58.
32 Glendon Schubert,"The Sociobiologyof PoliticalBehavior,"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, Studiesin Animaland Human Behavior,2 vols. (Cambridge: Harvard
University
Press,I970-I97I).
34 Roger D. Masters,"Genes, Language, and Evolution,"Semiotica,ii (No. 4, I970), 295320; JohnHurrell Crook, The Evolutionof Human Consciousness
(New York: Oxford UniversityPress, 198I).
BIOLOGICAL
NATURE
TABLE
HUMAN
TRANSFORMATIONS
Natural
Situation*
(Social Biology)
OF THE STATE
169
OF INCLUSIVE
FITNESS
STRATEGIES
Cultural
Institution**
(Sociology,
Anthropology,
Economics,&
PoliticalScience)
Individual
Motive**
(Psychology)
Asociality
Egoism
("Sauve qui peut")
Individuality
Nepotism
Love
("Be mine,I'm
yours")
Family
Mutual Benefit
Reciprocity
("I scratchyour
back, you ...
Trade: Marriage,
Barter,Markets
Sociality
Altruism
("Do unto others
Mutual Harm
Spite
("If I can't have
it, no one can
.")
Community:
State,Church
Feud: Economic
Competition,War
Situationsdefinedin termsof Figure 1.
** Each human motive can result from any one of the five natural situations,and each
human institutioncan reston any one or a combinationof the fiveindividualmotives.
varietyand complexity,are constitutedby the attributesdefiningwho

is included, who is excluded, and how participantsare expected to
behave. When social institutionsare described, however, we usually
distinguishseveral major categories or levels of analysis, such as the
family,the businessfirmor marketeconomy,and thestateor community.
Mediated by psychologicalassessmentsand motives,these familiarhuman institutionscorrespond-in a rough way-to categoriesof animal
social interaction.
Among humans, situationsthat other animals usually-though not
always35-"live" in an unreflectivemanner have been transformedinto
distinct,and oftencontradictory,
phenomenaof individualconsciousness
35 Donald R. Griffin,
The QuestionofAnimalAwareness(New York: RockefellerUniversity
Press, I976); JohnTyler Bonner,The Evolutionof Culturein Animals(Princeton:Princeton
UniversityPress, i980).
170
WORLD POLITICS
and culturalrules. This means thatanimal instinctand human learning

are differentways of respondingto similar problems created by environmental conditions and individual needs (cf. Aristotle, Politics,
VIII.xvii.1337a). Hence the conceptsof inclusivefitnesstheoryare relevant to complex human institutionslike the state,even thoughhuman
social behavioroftenhas different
"proximate"causes than thatof other
species.
II. COST-BENEFIT
FROM THE PRISONER
THEORIES OF COOPERATION:
S DILEMMA
TO THE TRAGEDY OF THE COMMONS
As Brian Barry has argued,36there are two broad traditionsin the

social sciences: economic models, in which aggregate phenomena are
analyzed as if individuals rationallycalculated the costs and benefitsof
alternatives,and sociologicalapproaches, which treatentire systemsas
having propertiesthat cannot be deduced from individual choices. A
similar dualityof methods can be discernedin the biological sciences.37
Since both inclusive fitnesstheoryin biology and economic or rational
actor models in the social sciences are based on a cost-benefitanalysis,
it has been argued that theycan be integratedinto a single approach.38
This prospect is particularlyrelevantto researchconcerningthe state,
because rationalactor theoriesof public choice or collectivegoods, akin
to those in economics, have increasinglybeen adopted by political scientists.39
To indicate how inclusivefitnesstheoryis related to social science,it
is convenientto begin with game theory,which has been used by many
social biologiststo explain the costsand benefitsof alternativebehavioral
In particular,the well-known "Prisoner's Dilemma" is a
strategies.40
fruitfulway of examiningtheproblemof cooperation:thisclass of games
36 Barry,Sociologists,
Economists,
and Democracy(London: Macmillan,I970). Cf. C. Wright
Mills, The SociologicalImagination(New York: Oxford UniversityPress, I959).
37 Masters
(fns.7 and 26); Williams(fn.25); Alexander
and Borgia(fn.27).
38 Jack Hirshleifer,"Economics froma Biological Viewpoint,"Journalof Law and Economics,
xx (AprilI977), I-52; Chase(fn.9); Margolis(fn.28).
39 E. g., AnthonyDowns, An EconomicTheoryof Democracy(New York: Harper, I957);
Mancur Olson, Jr.,The Logic of CollectiveAction(Cambridge: Harvard UniversityPress,
i965); Albert0. Hirschman,Exit,Voice,and Loyalty(Cambridge:Harvard University
Press,
I974); Denis G. Sullivan, RobertT. Nakamura, and Richard F. Winters,How AmericaIs
Ruled (New York: Wiley, i980).
40 Trivers (fn. 22);
Maynard-Smith(fn. i8); Donald L. McEachron and Darius Baer, "A
Review of Selected SociobiologicalPrinciples:Applicationto Hominid Evolution. II. The
Effectsof IntergroupConflict,"Journalof Social and BiologicalStructures,
v (April i982),
I2I-39.
171
has been widelyused in thesocialsciences,41and has also beenapplied

consider
inexplanations
ofanimalsociallife.42
For thesakeofexposition,
thefollowingexampleof thePrisoner'sDilemma.
$20, whichthey
split
Two men,A andB, havecollaborated
in stealing
jail cells.The
fifty-fifty.
Theyhavebeencaughtand placedin separate
fora full
he lacksevidence
jailertellsthemthatif bothremainsilent,
bothA
prosecution
andwillreleasethemwitha fineof$i each(leaving
andB witha netgainof$9 each).Ifoneofthetwotalks,buttheother
doesnot,theonewhotalks("defector")
willgo scotfree(netgain:$io),
toreturn
thestolen
whilethesilentindividual
willbe forced
("sucker")
to tendaysin jail or a fineof$io (netcost:
moneyand be sentenced
toninedays
-$io). Ifbothtalk,bothforfeit
themoney
andaresentenced
thislastresult
injailor$9 fines(netcost:-$9 each).ForA andB jointly,
and
is theworstpossible
outcome,
sinceall thestolenmoneyis forfeited
botharepunished
(forthetwo
(jailor fine).Yetitis thisworstoutcome
on
"rationally"
collectively)
thatis chosenifA andB decidetheiractions
becauseeachindividual
thebasisofa cost-benefit
calculation.
Thisfollows
action.
musttrytogainthemost(orlosetheleast)regardless
oftheother's
A gainsmore(orlosesless)bytalking
thanbysilenceifB is silentor if
B talks;B, deciding
separately,
makesthesamecalculation.
The game matrixforthisPrisoner'sDilemmais setforthin Figure

2a. This game has at leastthreeessentialconditions:first,the payoffs
must be transitive(in the example, + io
+9
-9
) -
io); second,
each actorindependently
choosesa strategy
thatwillmaximizehisgains
or behavioralcoercion
(or minimizehis losses);third,communication
as game
betweentheactorsis notallowed.Underthesecircumstances,
theorists
are fondof pointingout,it is "rational"forbothA and B to
choosethe outcomein whichbothlose, sinceforeach individualthe
of talking(defecting)
dominatesthatof silence(cooperating).
strategy
On a moment'sreflection,
Figure2a willbe seento containthesame
basiccategories
as Figurei (derivedfrominclusivefitness
theory).
The
of course,is thatthe Prisoner'sDilemma specifiesa set of
difference,
that
environmental
constraints
and thenseeks to predictthe strategy
willbe chosen,whereasHamilton'smoregeneralmodeldefinesthebasic
optionsin any set of environmental
conditions.Hence, the Prisoner's
Dilemmacan be takenas a specialcase of thegeneraltheory,
albeitone
41MortonA. Kaplan, Systemand Processin International
Politics(New York: Wiley,1957),
Part IV; RobertAxelrod, "EffectiveChoice in the Prisoner'sDilemma, Journalof Conflict
Resolution,xxiv (March i980), 3-25, and "More EffectiveChoice inthePrisoner'sDilemma,"
ibid. (Septemberi980), 379-403.
42 Robert Axelrod and William D. Hamilton, "The Evolution of Cooperation,"Science,
Vol.2I I (March 27, i98i), I390-96; RobertAxelrod,"The Emergenceof Cooperationamong
Egoists,"AmericanPoliticalScienceReview,Vol. 75 (Junei980), 306-i8.
172
WORLD
POLITICS
FIGURE 2A
PRISONER'S
DILEMMA:
NON-KIN
A's Strategy
Silent
Silent
_
A=-+9
A- +I0
B:+9
B=-10
BENEFIT
MMUTUAL
al
Talk
B =+IO
IALITY OR VIRTUE|
NEPOTISMM
B -9
MUTUALHARM
ASSUMPTIONS: 2-persongame in which payoffsare transitive(+ IO > +9 > -9 > -I o)

and commensurable;communicationor behavioralcoercionbetweenA and B impossible.
* Nepotism(privateself-interest)
and Sociality(virtue)definedin termsof A's payoffs.
situations.
thatis particularlyusefulin explainingmany counterintuitive
One major consequence of thissimilaritydeservesemphasis.The labels
in thesefiguresconcernthefour outcomesas defined
fromtheperspective
of
A. In the Prisoner'sDilemma, what is nepotism("selfishness")fromthe
perspectiveof A will be socialityor virtue("altruism") fromB's point of
view. One man's meat is another man's poison. Is it any wonder that
humans oftensuspect the motivationsof others-and proclaim the innocence of their own intentions(even if, or ratherespecially if, these
As Trivers
intentionsare consistentwith theirlong-rangeself-interest)?
has shown,43humans seem to specialize in deceit in order to "cheat" on
theirpartners,in moralisticaggressionagainst those who cheat, and in
deceitfulimitationof compliance and moralityin order to avoid punishment.
Even if the Prisoner'sDilemma is usefulin analyzing such problems
of social cooperation,what is gained by followingthose biologistswho
have extended this model to evolutionarytheory?To illustratethe unexpected consequences of linkinggame theoryand inclusivefitnesstheory,consider a Prisoner'sDilemma in which the two prisonersare full
siblings,or fatherand son. If the coefficientof relatednessbetween A
and B is I/2, A's payoffforeach cell of the matrixmust be rewrittento
include one-halfof B's payoff(which also accrues to A), and vice versa.
When we recalculate the game matrix in this way (Figure 2b), the
strategyof cooperatingnow dominates that of defectingfor each pris43
Trivers (fns.22 and 25).
BIOLOGICAL
NATURE
OF THE STATE
173
oner, regardlessof the response of the other. Rather than resultingin

mutual harm, the same payoffslead to cooperationand mutual benefit.
It should be littlewonder thatthe Mafia is organized in "families"based
on pseudo-kinship if not real kin relations-or that states often use
symbolsof kinship ("fatherland,""motherland,"'fraternite") to legitimate obedience.
FIGURE
PRISONER'S
2B
DILEMMA:
A's
Strategy
Silent I
A=+l13.5
B+35BSlent B
>t
A- + 13.5
MMUTUAL
BENEFIT
a
%q)
~ ak
KIN**
~~A=-5
B =+5
SOCIALITY
OR VIRTUE
Talk
A=+ 5
B - 5
NEPOTISM
A= -13.5
B= -13.5
MUTUALHARM
as in Figure 2A.
Nepotism(privateself-interest)
and Sociality(virtue)definedin termsof A's payoffs.
** Kin are eitherfull siblingsor parent-offspring
of relatedness"or r
(i.e., "coefficient
.5); hence, in each cell,A adds .5 of B's pay-offin Figure 3A to his own (and vice-versa).
ASSUMPTIONS:
*
As this example suggests,inclusivefitnesstheoryleads to the prediction that social cooperationis most likelywhen the partnersare kin or
can gain mutual and reciprocalbenefits.But, as Axelrod and Hamilton
show,44the Prisoner's Dilemma also illustratesthe natural constraints
thatinhibitself-sacrifice
forothers(socialityor 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
theotherplayeror organismas it behaved in thelast round.The virtuous
response cannot simply be establishedas the "best" policy in all cases,
because such a strategywill be vulnerable to cheating by a free rider
who gets the benefitsof the virtueof otherswithoutpaying any of the
costs.
From theperspectiveof game theory,social cooperationusuallyoccurs
only in cases of mutual benefit.To be sure, the decision rule need not
be "TIT-FOR-TAT" (using theother'slastactionas a cue fortheappropriate
44Axelrod and Hamilton (fn. 42).
WORLD POLITICS
174
response).Maynard-Smithhas shown thatthe locationof an interaction

can also serve as a behavioral rule, leading to a mutually beneficial
"bourgeois" strategyof deferringto the other individual when on the
other's"turf."45
Other formalizations
includeHirshleifer's"tendertrap"a game in which any rule of behavior that permitsan actor to predict
what others will do is preferableto the absence of rules.46All these
models can be used to develop Trivers's originalinsightthatreciprocity
can produce mutuallybeneficialsocial cooperationamong non-kin,even
in circumstancesof short-termloss 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 forvirtueor-to use the language of rational
actor theory-the choice of collectivegoods. Admittedly,this choice is
rare in other species,but humans occasionallyact in the group interest
even when reciprocityis unlikely or is deferredbeyond the temporal
frameworkof the alternatives.Game theoryis a usefulway to illustrate
the origins of such socialityor virtue,especially because it reveals the
critical role of coercive restraintsimposed by an impersonal and bureaucraticstate.
Although the problem of collectivegoods can be analyzed in terms
of an n-personPrisoner'sDilemma,47forheuristicpurposes it is clearer
to consider a similar model that Garrett Hardin popularized as the
"Tragedy of the Commons."48In these situations,one individual can be
considered to play a game against each of many neighbors.
Picturea pastureopen to all. It is to be expectedthateach herdsmanwill
tryto keep as manycattleas possibleon thecommons.... As a rational
being,each herdsmanseeks to maximizehis gain.... Adding together
thecomponentpartialutilities,
the rationalherdsmanconcludesthatthe
onlysensiblecourseforhim to pursueis to add anotheranimalto his
herd.And another;and another.... But thisis theconclusionreachedby
each and every rational herdsman sharing a commons. Therein is the
tragedy.Each man is locked into a systemthat compels him to increase
his herdwithoutlimit-in a worldthatis limited.49
45 Maynard-Smith(fn. i8).
46Jack Hirshleifer,"EvolutionaryModels in Economics and Law: Cooperation versus

ConflictStrategies,"Researchin Law and Economics(Greenwich,Conn.: JAI Press, i982),
IV, i-6o.
47Russell Hardin, "CollectiveActionas an Agreeablen-Prisoners'Dilemma," Behavioral
Science,xvi (SeptemberI971), 472-8i; Iain McLean, "The Social Contractin Leviathan and
the Prisoner'sDilemma Supergame,"PoliticalStudies,xxix (Septemberi98i), 339-5I.
48GarrettHardin, "The Tragedy of the Commons," Science, Vol. i62 (December I3,
Economy(San
i968), I243-48, reprintedin Herman E. Daley, ed., Toward a Steady-State
Francisco:Freeman,I973), chap.6.
49 Ibid., I04.
BIOLOGICAL
NATURE
OF THE STATE
175
Many ecologistshave agreed with Hardin's argumentthatthissituation

approximatesa wide range of phenomena,fromtheexhaustionof scarce
5
natural resourcesto the pollution of the environment.
To show how game theoryand inclusivefitnesstheorycan be combined in the analysis of social institutions,consider the Tragedy of the
Commons as a game withmanyplayerschoosingindependentlywhether
or not to add to theirherds. Since each knows that his decision cannot
influencethe choice of the others,it is inevitablethat some herdsmen
will adopt conflictingstrategies(here called StrategyA and StrategyB).
Assume that,ultimately,the commons will supportonly thexth cow of
one set of strategists(eitherA's or B's, but not both). Nobody knows,
however, either the value of x (i.e., the number of cows at which the
carryingcapacity is exceeded) or the number of playersadopting each
strategy.
The game has two outcomes,one in the shortrun (the addition of a
single cow; Figure 3a), the otherover a longer term(Figure 3b). In the
short run, everyone will add a cow, for the same reason that both
prisonerstalk in the simple Prisoner'sDilemma: to do otherwisewould
be a gratuitousact of sociality,incurringloss forno conceivablebenefit.
Unlike the Prisoner'sDilemma, however,thisstrategyis mutuallybeneficial in the short run (Figure 3a); it is only later that losses are encountered.
FIGURE
TRAGEDY
3A
OF THE COMMONS
ShortRun-Add i Cow
StrategyA
Don'tAddCow
Don't A:-I
AddCow
m%
Br-I
|I
MUTUAL
HARM
+ I
Add
u~~~i
Cow ~BBA+I
SOCIALITY*
jAddCowI
A=+I
NEPOTISM*
A +I
+ I
MUTUAL
BENEFIT
An n-persongame in whicheach actorknows thatonlytwo strategiesexistand thatothersare freeto choose either.All cows have a marginalutility= + i.
* Nepotism(privateself-interest)
and Sociality(virtue)are definedin termsof A's payoffs.
ASSUMPTIONS:
50 E.g., JohnN. Cole, "Learning a Hard Lesson in AroostookCounty,"Country

Journal
(November i980), 42-46.
176
WORLD POLITICS
If thecarrying
capacityof thecommonsis exceededin thelong run
(Figure3b),themutualharmtoall participants
willbe incommensurably
large.Virtue(notaddingcows) entailsa loss,particularly
comparedto
the strategy
of nepotismor privateself-interest;
but the dangerthat
everyonewill seeksucha selfishoutcomemakesself-restraint
relatively
beneficial
as soonas thereis a highenoughprobability
thatmutualharm
can be prevented.
The problemis thatno one knowswhichwillbe the
xthcow. Hence,withoutsomeformof organization,
it is impossibleto
focuson the long-termgame; the playerscontinueto calculateonly
immediatebenefits
and costs;and thetragedyseemsinevitable.
FIGURE
TRAGEDY
3B
OF THE COMMONS
Long Run-Add
Xth Cow
Strategy
Don'tAddCow]
Don't A= X-I
AddCow
B=X-I
>%
MUTUALBENEFIT
AddCow
A =X -I
B X+I
SOCIALITY
[AddCowl
A=X+I
B=X-I
NEPOTISM
A=X-1OOO
BXX-O000
MUTUALHARM
ASSUMPTIONS: as in Figure 3A.

Commons can supportan xth cow if added by eitherA strategistsor B strategists,
but not
by both; value of x is unknown.
the playersinto two groupsis a way out. If those

Transforming
adoptingstrategy
B couldcoercetheA's intolimiting
theirherds,while
theB's themselves
add cows,thelattercould gain thebenefits
of nepotism;but forthelosers,virtuestillhas itsrewards,sincetheoutcome
ofbeingcoercedis preferable
to mutualharm.By notaddingcows,the
A strategists
avoidcatastrophe-eventhoughtheyknowthattheirrivals
maybe less thanperfectly
virtuous.And if thecommonsor collective
goods can be expanded-forexample,by jointworkor byconquesteventhegroupwiththerelatively
inferior
strategy
can gainthemutual
benefits
of extendingthe short-run
game beyondtheoriginallimitsof
theresourcebase.
From thisperspective,
citizenscan benefitfromthe politicalcommunityonly by acceptingthe risk that thosewith power will gain
177
fromtheirrole as enforcers
disproportionately
of the socialcontract.5'
Governments
thus can be describedas providingthe serviceof cosocialbehavior,exactinga "side-payment"
forprotecting
ordinating
the
collectivegoods.52Once begun, such institutionscan createconsiderable
mutualbenefitsforall membersas long as obedienceto social rules

prevents
a Tragedyof theCommons.Undertheseconditions,
it can be
rationalforself-interested
individualsto forma politicalsociety,even
if it will occasionallyentailself-sacrifice
or "virtue"-includingpaying
taxes,servingin thearmy,evendyingin defenseof thecommunity.53
It is relatively
straightforward
to movefromsuchsimplified
models
ofgame theoryto morepreciseversionsof rationalchoicetheory.54
Not
onlydoes thisapproachsuggestthattheeconomists'
distinction
between
selectivebenefitsand collectivegoods can be foundedon evolutionary
biology,butit explainstheoriginofthestate-and itsultimatefragility
and disintegration.55
Cooperationcan readilyarisein smallgroupsofextendedkin(Figure
2b),and can evenexpandtomutually
beneficial
exchangesbetweennonkin (Figure3a). But as soon as collectivegoods come intoquestionforinstance,when groupscompetefornondivisible
resources,
or the
ecologicalcarryingcapacityis approached-theriskof mutualharm
increasesthebenefits
of enforceable
norms;freeriderscan be punished
by a coercivegovernment,
and well-organized
groupsoftenhave the
optionof conqueringweakeror less cohesivegroups.
havelongdebatedwhether
Anthropologists
thestatearosefromeither
intragroupcooperationor intergroup
conflict;56
the presentapproach
suggeststhatbothhypothesesare likelyto be correct.Indeed, both
processeswere probablynecessaryto overcomethegreatselectivedisadvantageof extendingsocialityto a societyof hundredsof thousands
of non-kin(in whichmutualbenefitcannotbe based on personalrecor individualrecognition
iprocity
of thefreeriders).Once established,
however,socialcooperationwithina statecan createextensivemutual
as symbolized
benefits,
bytheawesomegrowthin thehumanpopulation
51Margolis(fn. 28), chap. 9.
52 Michael Laver, The PoliticsofPrivateDesire(Harmondsworth,
England: Penguin,i98i),
esp. chap. 3.
53 Richard D. Alexander,"Natural Selectionand Societal Laws," in H. TristramEngelhardtand 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).
55Donald T. Campbell, "Legal and Primary-GroupSocial ControlsCurbing Selfishand

NepotisticHuman Nature," in MargaretGruterand Paul Bohannon,eds.,Biology,Culture,
and Law. (Special Issue ofJournalof Social and BiologicalStructures,
in press.)
56 Ronald Cohen and Elman R. Service,eds., Originsof the State (Philadelphia: Institute
forthe Study of Human Issues, I978).
178
WORLD POLITICS
thatis associatedwiththespreadofcentralized
politicalinstitutions
over
thelasttenthousandyears.
Paradoxically,
it is the verysuccessof the statethatcreatesits vulnerability.
The legalrulesenforced
bya government
constitute
collective
goods that are subjectto violationin orderto reap selfishbenefits.
Empiricalevidenceseemstoconfirm
thatthedeclineofpriorcivilizations
is relatedto, if not caused by,overexploitation
of environmental
resources-i.e., lackofself-restraint
and virtue;withgood reason,Garrett
Hardinremindsus thatthemereexistence
ofthestatedoesnotguarantee
immunity
fromtheTragedyof theCommons.57
III.
INCLUSIVE
FITNESS
THEORY
AND POLITICAL
PHILOSOPHY
It shouldnot be surprising
thatmodelsof rationaldecisionmaking
canbe so easilyrelatedtoinclusive
fitness
theory;
bothapproaches
deduce
social behaviorfroma calculusof individualcostsand benefits.But
recentevolutionary
theorycan be linkedto moretraditional
approaches
in politicalscienceas well.In Westernpoliticalphilosophy,
forexample,
definitions
ofhumannatureand theoriginofthestatefrequently
parallel
the conceptsof evolutionary
can be readily
biology.This proposition
illustrated
byconsidering
thethought
ofsomeofthebest-known
modern
politicaltheorists.
(A)
HOBBES
Hobbes'spoliticaltheory
is easilytranslated
intothetermsofinclusive
and rationalactormodels.The Hobbesian"naturalconditionof
fitness
or selfishness,
since comankind"is markedprimarilyby nepotism
limitedto thekin-group
basedon "naturallust."
operationis essentially
intomutualharm("war
As a result,socialinteraction
readilydegenerates
ofall againstall")-unless, forreasonsofmutualbenefit
("naturalright"),
individualsare inducedto agree ("socialcontract")to forma political
or state("commonwealth").
Hobbes thusnot onlydenies
community
thatsociality,
or virtue,
is natural;he is unableto justify
anycase ofselfon
sacrifice the basis of naturalrightalone. Individualscan-and if
whenever
rational,
alwayswill-reclaimtheirnaturalindependence
they
fearfortheircontinuedsafetyand self-interest.58
Hobbes'sstateof natureis like a Prisoner'sDilemmaor Tragedyof
57
G. Hardin (fn. 48); Marvin Harris, Cannibalsand Kings (New York: Random House,
I977). Cf. Machiavelli,Discourseson TitusLivy, 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: ColliersMacmillan,I 962), 98-I I 2.
BIOLOGICAL
NATURE
OF THE STATE
179
theCommons,59and can be escaped onlythrougha freelyacceptedarbiter

or a sovereigncapable of imposingrestraintson short-termself-interest.
In other words, the Hobbesian solution can be viewed as an n-person
game like Figure 3, in which each A strategistagrees with the otherA's
to accept the rule of a B strategistor strategistswho, as "sovereign,"is
not partyto the social contract.Even thoughsuch a sovereignmay reap
a selfishbenefitcompared to each individual subject, it is rational to
form a commonwealth because the result permitsthe A strategiststo
avoid the large negative payoffsof mutual harm ("war of all against
all") while preservingshort-termmutual benefits.
The same cost-benefitcalculus persistswithin societyas in the state
of nature. Since "no man can transfer,or lay down his rightto save
himselffromdeath, wounds, and imprisonment,"even convictedcriminals can be expected to tryto escape legitimateand just punishment:
For man by naturechooseththelesserevil,whichis dangerof deathin
resisting;ratherthanthe greater,whichis certainand presentdeathin
not resisting.6o
Pure socialityor virtuebenefitingthe group as a whole cannot be relied
upon, since men are "not bound to hurtthemselves,"nor are theybound
"to warfare,unless theyvoluntarilyundertakeit."61
Neither mutual benefitnor sociality(virtue)can exist withouta centralized state to induce self-sacrifice
throughfear.
Thereforebeforethenamesofjust,and unjustcan haveplace,theremust
of
be some coercivepower,to compelmen equallyto the performance
theircovenants,
bytheterrorofsomepunishment,
greaterthanthebenefit
theyexpectby the breachof theircovenant;and to make good that
of the
propriety,
whichby mutualcontractmen acquire,in recompense
universalrighttheyabandon;and such powerthereis none beforethe
erectionof 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 contractis only binding if there
is a sovereigncapable of enforcingit.
(B)
ROUSSEAU
Rousseau criticizes Hobbes in two differentways, each of which

becomes clearer if formulatedin terms consistentwith evolutionary
59WilliamOphuls, "Leviathan or Oblivion?" in Herman E. Daly, ed., Towarda Steady2I5-30; McLean(fn.47).
IOHobbes (fn. 58), I, xiv, p. IIO.
6, Ibid.,I, xxi,p. i65.
62Ibid., I, xv, p. II3-
StateEconomy
(San Francisco:Freeman,I973),
WORLD
180
POLITICS
biology.First,Rousseau attacksHobbes forpresumingthatthe familyand therewitha war-like state of nature between kin-groups-is natural.63Since asocial or purely egoistic behavior is a conceivable means
of survival in some species, institutionsbased on the perpetuationof a
bond between parentscannot be taken forgranted.And once the family
is analyzed as the result of a prior evolution away fromasociality,the
original state of nature must be presumed to have been "peaceful" and
"solitary,"at least as a heuristicmodel or a hypothesis.
Let us concludethatwanderingin theforests,
withoutindustry,
without
speech,withoutdomicile,withoutwar and withoutliaisons,withno need
forhis fellow-men,
likewisewithno desireto harmthem,perhapsnever
evenrecognizing
savageman,subjectto fewpassions
anyoneindividually,
and self-sufficient,
had only the sentiments
and intellectsuitedto that
.. . .64
state
According to Rousseau, therefore,Hobbes makes the mistakeof taking

as "natural" a condition that is the resultof historicalchange.65
Rousseau's second major criticismof Hobbes is diametricallyopposed
to the first:having argued thatthe original conditionof the species was
less social than Hobbes assumed, Rousseau also concludes that cultural
institutionscan be more social or virtuous than the Hobbesian commonwealth. Rousseau's Social Contractcan be read as a denunciationof
the Hobbesian theorybecause it focuses on mutual benefitratherthan
on a patrioticcommunityfor which individuals would virtuouslysacrificetheir lives. Whereas Hobbes limits reciprocityor mutualism to
Rousthosecommon institutionsthatare in the individual'sself-interest,
seau seeks a logic of reciprocity("social contract")that could generate
binding obligationsconstitutingthe group's collectiveinterest("general
will").
Like Hobbes, Rousseau explicitlytreats the agreement to found a
societyas a calculation of cost and benefit:
Let us reducethe pros and cons to easilycomparedterms.What man
losesbythesocialcontractis his naturalfreedomand an unlimitedright
to everything
thattemptshim and thathe can get; whathe gainsis civil
of everything
he possesses.66
freedomand theproprietorship
But unlike Hobbes, Rousseau insiststhat consent to enter the political
communitygenerates a duty to sacrificefor it i.e., the essential condition of socialitythat Rousseau explicitlycalls the "virtue" of the "cit63 Rousseau,
PartI
SecondDiscourse,
64Ibid.,I37-
(fn.i6), esp. I07,
I28-30.
ofRousseau(Princeton:PrincetonUniversity
65 Roger D. Masters,The PoliticalPhilosophy
Press, i968), chap. 3.
Rousseau, Social Contract,I, viii,in Roger D. Masters,ed., Social Contract,
66Jean-Jacques
withGenevaManuscriptand PoliticalEconomy(New York: St. Martin'sPress, I978), 56.
BIOLOGICAL
NATURE
OF THE STATE
181
izen."67 The willingness to die in defense of the state, dismissed by

Hobbes as not binding without a specificcontractualcommitment,becomes a moral obligation of citizenship for Rousseau.68
Rousseau thus seeks to reconstructan evolutionary theory of the
origins of the civilized community,accepting Hobbes's critique of theories that assume natural sociality-but nonethelessachieving a level of
communal unity that Hobbes rejects as naturallyimpossible. Rousseau
agrees that one must adopt a social contractmodel of the origin of the
state,but disagrees with Hobbes's conceptionof human nature both as
an original condition and as a limit of political possibility.The paradoxical mixture of individualism and collectivism,which has led many
can thereforebe traced to
readers to accuse Rousseau of inconsistency,69
a single view of human evolution: preciselybecause humans were not
competitivein the original state of nature,theycould develop virtuein
a freelyconstitutedpolitical community.70
(C) KANT
Kantian ethics,in thisview, transformRousseau's conceptionof virtue
by seeking to remove its limits. Whereas Rousseau's social contract
cannot bind anyone who does not share the patrioticspiritsymbolized
by ancient Sparta and Rome,7' Kant seeks a principleof social obligation
that is pure-i.e., free of all empiricallydeterminedor emotional limitations.72Although Kant is oftenregarded as an abstractmetaphysician
whose thoughtis unrelated to natural science (and especiallyto biology),
this interpretationis highlyquestionable;73rather,Kantian ethicscan be
viewed as a scientificresponse to the "natural" limitationson social
cooperation adduced by both Hobbes and Rousseau.
Kant's ethical teaching is based on the observationthat the "disposition" or motive of "duty" is never observed unambiguously:
if we attendto the experienceof men's conduct,we meetfrequentand,
as we admit ourselves,just complaintsthatthereis not to be founda
singlecertainexampleof thedispositionto act frompureduty.Although
67 Jean-Jacques
Rousseau,FirstDiscourse (fn.i6), 4I-42, 49, 5I-52; Rousseau,Emile, ed.
byAllan Bloom,I (New York: Basic Books,I979), 39-40.
68
Rousseau, Social Contract,II, iv (fn. 66), 64.
69E.g., JohnCharvet, The Social Problemin the Philosophyof Rousseau (London: CambridgeUniversity
Press,I974).
7? Roger D. Masters,"Nothing Fails Like Success: Developmentand Historyin Rousseau's

Political Teaching," in Jim MacAdam and others,eds., TrentRousseau Papers (Ottawa:
of OttawaPress,i980), 99-Ii8; Masters(fn.i6, I978); Masters(fn.65).
University
7' Rousseau, Geneva Manuscript,
I, ii, and PoliticalEconomy(fn. 66), I57-63, 2i8-20.
72 Immanuel Kant, MetaphysicalFoundationsof Morals, in Carl J. Friedrich, ed., The
Philosophyof Kant (New York:ModernLibrary,i964), 140-208.
73 JohanHjort, The Human Value ofBiology(Cambridge: Harvard University
Press, I938);
Stent(fn.30).
WORLD POLITICS
182
it is neverto whatdutyprescribes,
manythingsare done in conformity
outofduty.74
thelessalwaysdoubtfulwhethertheyare done strictly
Kant sees thatpure "altruism"would require a motiveuncontaminated
by self-interest-andthe existenceof such a motiveis "always doubtful."
As contemporarysocial biologyindicates,thereis reasonenough to raise
this question with regard to humans as well as to other species.75Kant
answersbydenyingthatmoralitycould ever be deduced from"particular
attributesof human nature,"insistingthatit is only in abstractionfrom
empirical interestsand factsthat a man-made, rational"imperative"to
is possible: "every empirical element is not only quite inself-sacrifice
capable of aiding the principleof morality,but is even highlyprejudicial
to the purityof morals."76Hence, Kant's categoricalimperativeis "unconditional" and "not grounded in any interest"-and therebysharply
differentfromconsiderationsof "market price" or mutual benefit(i.e.,
fromconsiderationsthatcould apply to the behaviorof otheranimals).77
Kant thus maintains that there is no naturalisticground for virtue
other than an act of the free human will, imposing on itselfa rational
obligation as an end in itself.The individual, generalizing his rule of
choice ("maxim") into a law governingall rationalbeings,engages in a
logical transformationof the situation that is not unlike Rousseau's
"general will"-except that for Kant, the categoricalimperativeis addressed to all humans as rational beings, not to all fellow-citizensas
This
membersof thesame (admittedlycontingent)politicalcommunity.78
means that,for Kant, moral obligationis a pure case of sociality,since
the individual acts morallywithoutany expectationof personal gain in
the shortor long run. Whereas Trivers's "reciprocalaltruism"can include elementsof mutuallybeneficialreciprocityand even of nepotism,
Kant thusprovidesa logic of freelywilled virtuewithoutany conceivable
admixtureof self-interest.
(D) HEGEL
is an extraordinaryattemptto transcend
Hegel's Philosophyof Right79
the divergencesbetween the theoreticalperspectivesof Hobbes, Rousseau, and Kant. The First Part, devoted to "AbstractRight," could be
74Kant (fn.72), 154 (emphases
in original).
(fn.30), esp. chap. 4; Hans Kummer,"Analogs of MoralityAmong Nonhuman
75 Wispe
Primates,"
in Stent(fn.30),
76 Kant(fn.72), 174-
31-47.
Ibid., i8o, I82.

78 Rousseau, PoliticalEconomy(fn. 66), 2i8-20.
77
79Georg Wilhelm Friedrich Hegel, The Philosophy

of Right,ed. by T. M. Knox (New
York: Oxford UniversityPress, i964).
183
viewed as the perspectiveof Hobbes;80the Second, dealing with "Morality,"focuses on the dimension that Rousseau and especially Kant
found lacking in Hobbes;8" and the Third, "Ethical Life," presents
in priorpoliticalthought.
the contradictions
Hegel's mode of transcending
The Third Part is itselforganized in threedivisions,correspondingto
the main institutionalformsdescribedin Table I: "The Family;82Civil
Society,or the market economy;83and The State.84Thus the Hegelian
dialectic of "right,""morality,"and "ethical life" moves fromthe perspectiveof theindividual'scalculus of costsand benefits(rightsor claims)
to a Kantian notionof pure dutyor virtueas uniquelyhuman (morality),
and then to human institutionsas social systemsintegratingall constitutivelevels (ethical life).
Although detailed analysis of Hegel is impossible here, we should
note that he explicitlytreatsboth marriageand economic activity-i.e.,
of "nat"needs and the means of satisfyingthem"-as transformations
In contrast,the state,forHegel, is "a selfural" or "animal" functions.85
dependent organism"; its Constitutionis the manifestation,in "the objectiveworld," of the "self-developmentof the Idea."86In more familiar
usage, the state is a cultural,or man-made, phenomenon; ratherthan
of a natural or animal need, the state is thus a
being a transformation
purely human constructiondependent on the self-consciousactivityof
its citizensand the evolutionof human thought(Hegel's "world mind").
(E)
THE RELEVANCE
OF POLITICAL
PHILOSOPHY
This summaryof political philosophyin termsconsistentwith contemporarysocial biologycould easilybe expanded to includetheancients.
For example, the pre-Socratics'analysisof individual costsand benefits
leads to a social contracttheoryparallel to that of Hobbes, whereas
Plato's Republic offersa model of group selection for socialityby destroyingboth the familyand the marketeconomy;Aristotle,like Hegel,
develops a complex balance between institutionsat the level of family,
economic system,and polity.87
Although the specificvocabularydiffers,
the issues posed by contemporarybiological theoryare thus fundamentallysimilar to the traditionalquestions in Western political thought.
The analysis of human nature,long approached by observationand
80
Ibid., 37-74.
82 Ibid., 110-22.
8i Ibid., 75-105.
83Ibid.,122-55-
84Ibid.,155-22385Ibid.,Para. i6i, p. III;

Para. 190, p. 127.
86Ibid.,
Para. 259, p. i6o; Para. 267, p. i63.
87 Roger D. Masters,"Classical Political Philosophyand ContemporaryBiology,"paper
presentedto Conferenceforthe Study of Political Thought,Chicago, Ill., April, 1978.
184
WORLD POLITICS
insight,can thus be illuminated by scientificresearch.As Trivers has

indicated,the logic of inclusivefitnessexplainssuch supposedly"selfish"
behaviorsas parent-offspring
conflict,siblingrivalry,and a hostof other
frequentlyobserved human traits.88
But, in recognizingthe ubiquityof
egoistic motives and behaviors,evolutionarybiology also points to the
countervailingtendenciestoward cooperation,particularlyin a species
that experiencesprolonged helplessnessin infancy,predatorsin its environment,and scarcityof food requiringcooperativehuntingand food
sharing.From the earliestepochs,therefore,we can assume thathuman
nature exhibiteda complex of both cooperationand competition.89
In theoreticalterms,this means that Hobbes's definitionof the state
of nature as a mutuallyharmful"war of all" among selfishindividuals,
like Rousseau's assertionof human asocialityor Kant's formulationof
a pure and universaldutyof virtue,cannot fullydescribethe originsof
the state. That is tantamount to saying that the accounts of human
nature and political institutionsfound in Hobbes, Rousseau, or Kant
are only partially correct heuristicmodels: each is true as a formal
descriptionof human behavior under some circumstances;none is a
global definitionof the unchangingbasis of human nature.
The factthat the major politicalthinkersof the past have elaborated
systemsthat are incompleteor partial should not be taken to indicate
thattraditionalpoliticaltheoryis irrelevantor useless.Each of the major
theoristsdiscussed above analyzes a particularissue or transformation
with archetypicalclarity.Of the five conceivable modes of behaviorasociality,nepotism,mutual benefit,sociality,and mutual harm-differentphilosophershave tended to isolate specificsituationsand to explore with great depth how they relate to political experience. For
example, Hobbes shows how nepotism,under circumstancesproducing
mutual harm, can be transformedinto mutual benefitonly by the constructionof a man-made "commonwealth." Rousseau spells out how
humans could have evolved froman asocial condition(the "pure state
of nature") into a Hobbesian conditionof mutual harm,as well as how
a politicalcommunitycould go beyond the mutuallybeneficialcontracts
of Hobbes to a patrioticcommunityunited by feelingand civic virtue.
And Kant indicateshow even such a formof virtuewould fall shortof
the pure demands of a moral obligation,thus suggestingan inevitable
tensionbetween privateself-interest
and group interests.From the perspectiveof evolutionarybiology,it is thereforeevidentthat,contraryto
Trivers (fn. 25)89Roger D. Masters, "Nature, Human Nature, and Political Thought," in J. Roland
88
Pennock and JohnChapman, eds., Human Naturein Politics(New York: New York UniversityPress, 1977), 69-iio; Edgar Morin,Le ParadigmePerdu (Paris: Seuil, 1973).
185
thegreatpoliticaltheorists
addressthe pertheclaimof "historicists,"
ennial issues of human social life.90
IV. THE
EMERGENCE
OF THE STATE
usuallytreatthe emergenceof the stateas a

Politicalphilosophers
fromanything
in theanimal
different
centralproblem,fundamentally
evolutionary
biologists
agree,pointingto thelimworld.Contemporary
in othersexuallyreproducing
itationsin thescale of socialcooperation
and theirinclusion
mammals.The sheersize of politicalcommunities,
ethnicgroupsand classes,
of heterogeneous
populationsof different
suggeststhatsomethingunique was necessaryto producewhat is diverselycalled human "civilization,""the state,"or "euculture."9'
For most of the last 3.5 million yearsour ancestorslived in relatively
small bands, presumablystructuredin groupsof extendedkin (like many
tribes).Exogamy,reinforced
primatespeciesand human hunter-gatherer
by the incesttaboo, probablyresultedin an exchange of mates between
lineages or groups, therebyencouragingconscious norms of reciprocity
in social interactions.Hominids seem to have evolved in groups favoring
nepotismor kin selection-a structurethat greatlyincreasesthe probabilityof cooperationwithin the group-reinforced by "altruistic"motives toward group members and hostilityto outsiders.92But since cooperation among millions of non-kin cannot be explained by "human
nature" alone, the originof the stateposes the followingscientificquestion: What circumstancesin the last ten thousand years permittedthe
formationof verylarge groups-especially since the establishmentand
maintenanceof such societies(let alone the formationof politicalsystems
to run them) were previouslyimpossibleor unnecessary?
In the anthropologicalliteratureon the originsof the state,two main
In thefirstview,thestate-organizing
hypotheseshave been presented.93
and controllinghuman populations-arose as the resultof internalprocesses withinprestate societies;in the second,it is conflictbetweengroups
that generated states. Is new light cast on the problem by considering
the work of evolutionarybiologists?
According to contemporaryevolutionarytheory,so-called "group se90Leo Strauss,Natural Rightand History(Chicago: Universityof Chicago Press, 1953).
91A. L. Kroeber, The Nature of Culture(Chicago: Universityof Chicago Press, 1952);
Samuel M. Hines, Jr.,"The Origins of the State," in Heiner Flohr and WolfgangTommesmann,eds.,PolitikundBiologie(Berlin:Verlag Paul Parey,in press);CharlesJ.Lumsden
and Edward 0. Wilson, Genes,Mind, and Culture(Cambridge: Harvard UniversityPress,
i98i).
92
Robin Fox, "PrimateKin 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).

93Cohen and Service (fn. 56).
186
WORLD POLITICS
lection" is not typicallyexpected; in general, social cooperation will

develop only when the actor's own kin ultimatelybenefitthroughreciprocity-and this factorcan be expected to limit the size of groups to
that in which individuals have a reasonable probabilityof recognition
or role reversalin futureencounters.94
There are, however, some restrictivecircumstancesin which this
individualisticcalculus does not hold. Where a number of small populationsof extended kin are relativelyisolated fromeach other,conflicts
betweengroups can and will occur ifresourcesare insufficient
to support
all of them. In that case, a behavior that benefitsthe group-even at a
cost to the actor-can be favored by natural selectionwhenever competitionwithin the group is harmfulto both the individual's "inclusive
fitness"and the group's "collectiveinterest."Under these circumstances
(which Alexander describesas a "balance of power" betweencompeting
bands of extended kin), group selection would expand the sphere of
social cooperation.s5As Benjamin Franklin put it, "We must all hang
together,or assuredlywe shall all hang separately."
Fred Willhoite has shown how this "balance of power" hypothesis
can explain the progressiveevolutionof hominidsthroughthe stages of
the band (exemplifiedby the !Kung bushmen), the sovereign village
ranked society
(e.g., Yanomomo), the chiefdomas a multi-community,
based on extendedkin-groupsor lineages (like so manyof the preliterate
societiesstudied by anthropologists),and finallythe early state (ranging
historicallyfrom ancient Egypt to i9th-centuryAfrica). At each level,
as communities expanded to include more distant kin or nonrelated
kin-groups,the role of coerciveinstitutionsalso expanded. And clearly,
thereis some evidence thattheearliestcivilizationsevolved out of rivalry
and conflict.96
An alternativemodel of the origins of the state focuseson the new
environmentalor productiveresourcescreated by large-scale cooperation. This model only works, however,if some new "collectivegoods"
are available from which individuals can benefitrelativelyquicklyindeed, more rapidlythan the incrementsto inclusivefitnessthatwould
follow fromeithernepotism (refusalto cooperate with non-kin) or an
94 Wilson (fn. 2), chap. 5; Philip R. Thompson, "And Who is My Neighbor? An Answer
fromEvolutionaryGenetics,"Social ScienceInformation,
xix (No. 2, ig80), 341-84.
95Alexander (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 presentedat 76th
Annual Meeting,AmericanPoliticalScience Association,Washington,D.C., ig80; Willhoite
(fn.3i); AlexanderRustow,Freedomand Domination(Princeton:PrincetonUniversityPress,
1980).
187
insistenceon individual reciprocity.The developmentof extensiveirrigationsystems,formingthe basis of what Wittfogelcalls "hydraulic"

empires,97 would be an example of such a situation.
Eisenstadt's analysis of the emergenceof early empires,for instance,
associates the rise of a centralized bureaucracy-necessary to establish
a lastingpolitical tie by enforcingthe commands of the state-with the
At
existenceof productivesurplusesand socioeconomicdifferentiation.s8
least one careful study of the emergence of a "primarystate" in high
antiquityfinds that the population explosion took place afterthe first
developmentof the state's institutions-not before,as mighthave been
Since historyis littered
predictedfroma balance-of-powerhypothesis.99
with tales of Pyrrhicvictoriesand futileconquests (like those of Alexander the Great), it is likely that both intergroupcompetitionand intragroupcooperation played a role in the origin of the state.
In an interactivemodel, intergroupcompetitionplaces a premiumon
within-groupcooperation-and within-group
cooperationleads to greater
intergroupcompetition;only in this way could the free-ridereffectsof
nepotism,which have generallyprecluded the emergenceof large societiesand states,be overcome.In particular,it is the interactionbetween
the existenceof tangiblebenefits(increased quantitiesand reliabilityof
resources,made possible by compulsoryeconomic cooperation)and the
threatof outsiders seeking to gain these benefits,that permitsthe use
of symbolicappeals to unityand virtue withinthecommunity.The result
is a processof "moralizing aggression,"which reinforcescooperationby
creatingfeelingsof guilt in the potentialcheaterand of "righteousness"
in the cooperators,as well as by metingout legitimatepunishmentfor
the guiltywho are caught.Ioo
In this model, since the institutionalizedstate can only occur under
a narrowsetof conditions,timewould be a criticalfactor.If thecombined
effectsof intergroupcompetitionand within-groupcooperationfavored
the formationof a state only during a shortperiod, these unique "occasions" (to use Machiavelli's term)may have givenplayto theleadership
qualities of particularindividuals.Hence, in an interactivemodel of the
origin of the state,there is even a role for the "great man" in history.
The "critical period" hypothesisis an importantpart of the model,
97Karl Wittfogel,OrientalDespotism(New Haven: Yale UniversityPress, 1957). For an
example in the New World, see B. L. Turner, II, and P. D. Harrison,"PrehistoricRaisedField Agriculturein the Maya Lowlands," Science,Vol. 213 (July24, i98i), 399-405.
98 S. N. Eisenstadt,The PoliticalSystems
ofEmpires(London: Free Pressof Glencoe, i963).
99Cohen and Service (fn. 56), 6i.
.ooTrivers(fn. 25); Willhoite(fn.31); Peter Corning,The Synergism
Hypothesis:
A Theory
of Progressive
Evolution(New York: McGraw-Hill, in press).
188
WORLD POLITICS
becauseit helpsto explainhow the free-rider

problemwas overcome.
Althoughundetectednepotismpayswithinthe state,at its formation
cheatingbehaviorwould be reducedin frequency
becauseit would be
visibleat the decisivemoment.When thebattlestarts,thosewho stay
in theirtentsratherthanfightare usuallyobvious(Achillesin theIliad);
thosewho builtthe pyramidsof ancientEgypt-the epitomeof cooperativeendeavorwithinan earlystate-engraved"tallymarkson the
casingstones"in orderto "givethetitlesof theindividualworkteams
who wereto be creditedwiththesupply."IoI
Governments
and lawsthusseemto havemadeaccessiblethebenefits
ofcooperation
amongnon-kinin larger-scale
societies.
At thesametime,
thosewho providetheirfellowswiththe"services"oforganizingsocial
cooperationand catchingcheatersare presumably
as subjectto evolutionaryprinciplesas thosewho obey(or disobey)thelaws. Governing
officials
fortheirpositions-prestige,
everywhere
reap benefits
power,
as well as accessto femalesand thelikelihoodthatoffspring
will gain
privilegedtreatment.
Indeed,inclusivefitness
theoryleadsus to predict
thatenvyor jealouslywill be widespread,and will ultimately
destroy
anypoliticaleliteincapableof providingthemanywithbenefits
which
exceedthe relativecostsof supporting
thepoliticalsystem.
Unliketraditional
theevolutionary
theories,
approachsketchedhere
does not presumeeitherthatthestateoriginateddirectly
froma priori
traitsof humannatureor thatit resultedsolelyfromhistorical
circumstances.Rather,the humancapacityto institutionalize
new modes of
improvinginclusivefitness,
whencombinedwithappropriate
external
conditions-andperhapsonlyin thepresenceof individualscapableof
seizingtheopportunity-would
explainthespecifictimesand placesin
whichstateshave arisen.For example,the jointrole of international
is particularly
rivalryand economicdevelopment
evidentin theorigins
of the modernstatesystemin the I7th and i8th centuries-aswell as
in thecity-states
of classicalantiquityand RenaissanceItaly.
Once theyhave been established,
however,statesremainsubjectto
the dialecticalrelationship
betweenindividualand groupinterests.
As
theTragedyoftheCommonsindicates(Figure3b),itis alwaystempting
to seek to combinethebenefits
of collectivegoods(basedon thevirtue
ofothers)
withtheselectivebenefits
ofnepotismor selfishness.
For every
law thereis alwaysa new loophole;foreverynew abuse,a possible
... Homer, Iliad, Book IX; Kurt Mendelssohn,The Riddle of the Pyramids(New York:
Praeger, 1975), 148. For a strikingillustration,see also Exod. 33:21-29. On the origin of
writingas a systemfor ensuringcreditfor tax payments,see Denise Schmandt-Besserat,
"Deciphermentof the Earliest Tablets," Science,Vol. 22i (Januaryi6, i98i), 283-85.
189
legislativeresponse.Periodically,it is necessaryto reinvigoratethe state;

even a cursoryglance at recorded historyreveals the extraordinaryinstabilityof political systems.
Paradoxically,the state is vulnerableover the long run preciselybecause it is so successful in generatingmutual benefitsand collective
goods in the short time horizon of an individual's conscious decision
making. The very outcomes that representthe strengthof political
communitiescan thus be the seeds of theirdecay. Especially if further
researchconfirmsthe soundness of this approach, evolutionarybiology
can providea frameworkthatgreatlyilluminatesthe futureof industrial
societyas well as the origin of the state.'02
V.
CONCLUSION:
THREE
REASONS
FOR LINKING
EVOLUTIONARY
POLITICAL
THEORY
AND
BIOLOGY
There are threebroad reasonsfordevelopinglinkagesbetweenbiology

and political science along the lines suggested above. First,the establishment of a unifyingfoundation for the varied perspectiveswithin
contemporarysocial science will do much to improve research.Second,
in empiricalstudiesof politics,variablesand approachessimilarto those
in evolutionarybiology could reveal hithertounknown relationshipsof
great importance.Third,and most important,evolutionarybiology has
implicationsfor public policy and ethical judgment.
I. Unfyingthedisciplineofpoliticalscienceand relating
it to thenatural
sciences,which has long been a dream,103
now seems possible. Although
cost-benefittheories(whethergame theory,conventionaleconomics,or
decision-makingmodels) need not be modifiedfundamentallyin order
to be linked with inclusive fitnesstheory,by so doing one gains a
heightenedawareness of the domain within which collectivedecisions
can be derived from the aggregationof individual choices based on a
cost-benefit
Politicalphilosophy,long divorcedfromthe "hard"
calculus.104
sciences,can onlybe strengthenedbytherealizationthatfamousthinkers
of
describe,with archetypicalclarity,the categoriesand transformations
behaviorthatare centralto a scientificunderstandingof human political
102 Roger D. Masters,"Social Biology and the Welfare State," in Richard F. Tomasson,
ed., ComparativeSocial Research,Vol. VI (Greenwich,Conn.: JAI Press, i983), in press.
03 Karl Marx, Economicand Philosophic
of i844, ed. by Dirk J. Struik(New
Manuscripts
York: International,1964), 143.
04For a concreteexample, compare Hirschman (fn. 39) with Roger D. Masters,"Exit,
(DeVoice, and Loyaltyin Animal and Human Behavior,"Social ScienceInformation,.xv
calculus,see Masters(fn. 26);
cember 1976), 78-85. On the limitsof individualcost-benefit
Alexander and Borgia (fn. 27).
190
WORLD POLITICS
experience.105 And the characteristicempirical theoriesof the state,os-
cillatingbetween models of conflictand of functionalintegration,can

now be relatedand combined in a more comprehensiveapproach to the
studyof political and social history.
2. New and moresignificant
empiricalresearchshould, of course, be a
goal of anyone claiming to be engaged in a scientificendeavor. William
Durham has presentedthe firstcompellingcase of a culturalprohibition
whose distributionand timingwere the resultof naturalselection.o6 In
West Africa,many tribestraditionallyhad a taboo on the consumption
of newly ripe yams before a specificfestival(even though other foods
were in shortsupplyat thisseason). It has been claimed thatconsumption
of theseyams could counteractthe sicklingeffectsin hemoglobin,making individualswith the sickle cell gene bothmorevulnerableto malaria
and less vulnerable to sickling crises; this hypothesisexplains both the
taboo (sicklingwould have providedan adaptive advantageduringrainy
periods because it confersrelativeimmunityto malaria) and the festival
(as soon as the climate became dryer,the reduced incidenceof malarial
mosquitoes would have made it beneficialforthosewith sicklinghemoglobin to consume large quantitiesof yams).
Although there is controversyabout the precise biochemical mechanisms involved, the timingof yam consumptionclearlyhad biological
implications.Durham shows that the New Yam festivalsoccurredonly
in settingswith a "short dry season" in the summer,and were timed
to coincide with changes in rainfall.Under these environmentalconditions,the taboo postponed yam consumptionuntil the rains stopped,
and thus reduced the incidence of malaria (if only by keeping people
out of the yam fields,where pools of water tend to increase the risk of
exposure to mosquitoes). But this means that the taboo had different
selectiveconsequences foreach of thecharacteristic
genotypes:regardless
of the biochemical effectsof yam consumption,individuals with sickle
cell trait-and especially homozygotes-had to be affecteddifferently
than those with normal hemoglobin.Hence thistaboo, thoughcontrary
to the short-range"selfish" interestsof hungryindividuals, seems to
have been adaptive both for carriersof the sickle-cellgene and for the
population as a whole. While convincingas an example of the natural
selectionof a legal customthatbenefitstheentiregroup(especiallyinsofar
as geneticdiversityis in the group's interest),thiscase is obviouslyhighly
"5 Masters
(fn.I6, 1978; fn. 87; fn. 89); Masters,"The Duties of Humanity: Legal and
Moral Obligation in Rousseau's Thought," in Fred Eidlin, ed., Constitutional
Democracy:
Essaysin ComparativePolitics(forthcoming).
io6 Durham, "Coevolution and Law: The New Yam Festivals of West Africa," paper
presentedto Symposiumon Law and Behavioral Research,MontereyDunes, Calif., Sep-
tember25-27,
i98i.
191
localized and verycomplex. Are theremore generallessonsto be learned

fromusing evolutionarybiologyto studypoliticaland legal institutions?
In other species, it is well known that mating patternsvary widely
depending not only on the degree of physicaldifferencebetween males
and females(sexual dimorphism),but on such otherfactorsas predation,
the scarcityof resources,or the costsand benefitsof parentalinvestment
in the young. In analyzing these relationships,Alexander and others
have advanced the hypothesisthatthe establishmentof the stategreatly
increases the selective advantage of monogamous mating systems.'07
Whatever the extent to which early hominids establishedlasting pair
bonds-and on this question thereis considerabledisagreementso the
centralizationof political power would thus change the environmental
constraintson the kinship systemselaborated by various cultures.109
Whereas most social scientistsstresseconomic variablesin theirstudy
of politics,theoriesof the stateshould also make explicitthe relationship
between familial (or kinship) institutionsand the political system.For
example,Dickemann has shown how the variablesinfluencingsex ratios
in other species operate in a class- or caste-basedsocietythat is subject
to unpredictable disasters which can wipe out accumulated material
wealth; epitomized by traditionalNorthernIndia, these circumstances
seem to generate a "hypergamous"mating system,in which the upper
classesdiscriminateagainstinvestmentin femaleswhile thelower classes
discriminateagainstinvestmentin males.II0 Such findingsare, moreover,
of contemporaryrelevance:Dickemann's model pointsto theempirically
testablehypothesisthat relativeequality between the sexes is to be expected in a market economy of widespread abundance and social mobility.Thus, by providingsocial sciencewithan evolutionaryfoundation,
it may be possible to explain the incidence of socially variable norms
(like those toward gender) which have hithertooftenbeen treated as
preferencesor choices withouta rational foundation."'
3. Policyproposalsand ethicaljudgmentscan, in this perspective,also
107 Richard D. Alexander and others,"Sexual Dimorphisms and Breeding Systemsin
Pinnipeds,Ungulates,Primates,and Humans," in Chagnon and Irons (fn. 8), 402-35.
io8 Lovejoy (fn. i);
Sarah BlafferHrdy and William Bennett,"Lucy's Husband: What
Did He Stand For?" Harvard Magazine, Vol. 83 (July-Augusti981), 7-9, 46.
'09 Pierre L. van den Berghe,Human FamilySystems
(New York: Elsevier, I979).
0Mildred Dickemann, "The Ecology of Mating Systemsin HypergynousDowry Societies,"Social ScienceInformation,
xviii (No. 2, 1979), i63-95; Dickemann, "Female InfanA PreliminaryModel," in Chagnon
ticide,ReproductiveStrategies,and Social Stratification:
and Irons (fn. 8), 321-67; Robert L. Trivers and Daniel E. Willard, "Natural Selectionof
Parental Abilityto Vary the Sex Ratio of Offspring,"Science,Vol. I79 (January1973), 9092.
III Roger D. Masters, "Explaining 'Male Chauvinism' and 'Feminism': Differencesin

Male and Female ReproductiveStrategies,"Womenand Politics,in press; Ralph Pettman,
Biopoliticsand International
Norms(New York: Pergamon,i98i).
192
WORLD POLITICS
be informedby an evolutionary
approachto humanlife.To be sure,
we are constantly
toldthatthe"naturalistic
fallacy"is committed
wheneverscientific
"facts"are relatedto decisionsabout"values."But inclusivefitness
theory(unliketheconventional
imageofbiology)establishes
differences
of judgmentas naturaland inevitable.Recallhow thecategoriesin Figure i relateto thevarioussituations
set forthin Figures
2a, 2b,3a, and 3b; in everycase,thedescription
ofan outcomedepends
on theindividualwho is makingthejudgment:whatis virtue
decisively
foractorA is nepotism
forB (and viceversa).Accordingto evolutionary
biology,as forAristotle,what is in "accordancewithnature"is not
givenor rigidlyfixed,but changesdependingon circumstances;
a biologicalperspective
on humanethicstakesthegroundsof opposedconceptsof justiceseriously,
if onlybecausealternative
viewsoftenreflect
predictable
responsesto veryrealsocialproblems(e.g.,Politics,I.I255a;
III.I280a-I 282b).
Far frombeinga logicalfallacy,
"naturalism"
thuspromisesa return
toan ethicaltradition
witha longand noblehistory.
It doesnot,however,
resurrect
older "naturallaw" teachingswhichpresumably
enforceabsolutestandardsregardless
oftimeand place.Rather,naturalism
would
seemto pointtowarda biologicaland culturalequivalentof thetheory
of relativity
in physics.112
Indeed,one could argue thatsocial biology
promisesan understanding
of both"specialrelativity"
(i.e., whyindividualswithinanysocietyor culturehave ethicaldisagreements,
based
on theirdiffering
perspectives)
and "generalrelativity"
(i.e., whydifferentsocietiesand cultureshave different
laws and customs,based on
theirenvironment
and history).
Yet naturalism
wouldaccountforsuch
culturalvariations
withoutfallingintotheoppositeexcessofmere"relativism,"sinceevolutionary
theoryprovidesa standardagainstwhich
humanbehaviorcan be measured.
Lest thisview be dismissedas a returnto thediscredited
view that
"whateveris,is right,"one shouldrecallAristotle's
emphasison nature
as a criterion
ofethicaljudgment(Politics,
II.I I03a;
I.1252a-I253a; Ethics,
V. I I34b-i135a). In medicine, no one would maintain that,because a

rupturedappendix can occur in the "natural course of events,"people
oughtto die of appendicitis.Those who have confusedbad biologywith
a so-called "naturalisticfallacy"should-especially in an age dominated
by the threatof nuclear war, geneticengineering,and pervasiveterrorism-make it clear what they propose as the foundation of ethical
judgment.The fact-valuedichotomymay sound innocuous and reason112
FrijhofCapra, The Tao of Physics(Berkeley,Calif.: Shambala, 1975).
193
able in calm and tranquiltimes,but-as Leo Strausshas pointedout

so effectively-inour day it amountsto "retailsanityand wholesale
madness.""13
These are, however,complexand controversial

matters.They have
of an evolutionary
been raisedhere to indicatethatthe introduction
approachto the studyof politicsand social behaviorhas implications
thatare farfromtrivial.It maybe thattheprojectoflinkingthenatural
and socialsciencesis merelyan intellectual
luxuryof no lastingimportance,and thatnew empiricaldiscoveries
concerning
socialand political
changewillfailtomaterialize
fromit.Butevenifevolutionary
reasoning
couldilluminate
nothingbutourethicalor policyjudgments,
surelythat
wouldbe reasonenoughtoexplorethebiologicalfoundations
ofpolitical
life.If legal institutions
and governments
are not merelyman-made
conventions
or arbitrary
manifestations
ofhuman"will"(Plato,Republic,
IV.428a-444e),it behoovesus to followtheSocratictradition
ofseeking
standardsof thatwhichis rightor just,"accordingto nature."
II3Strauss(fn.90), 4. Cf. Masters(fns.26 and 105).

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Trustees of Princeton University

The Biological Nature of the State

THE BIOLOGICAL NATURE OF

V V thousands,if not millions,of individualswho owe obedience to

? i983 by the Trustees of PrincetonUniversity

used here, the conceptof evolutionconcernsonly the processof natural

8 Robin Fox, ed., BiosocialAnthropology

AlbertSomit,ed., Biologyand Politics(The Hague: Mouton, I976); Thomas C. Wiegele,

BIOLOGICAL NATURE OF THE STATE

Marc Beckoffand Michael C. Wells, "The Social Ecology of Coyotes,"Scientific

sistentwith the widely observed tendenciesof animals to adjust their

Jean-JacquesRousseau, Second Discourse,Part I and note j, in Roger D. Masters,ed.,

Birute Galdikas-Brindamour,"Orangutans, Indonesia's 'People of the Forest,'" National

'7George C. Williams,Adaptationand Natural Selection(Princeton:PrincetonUniversity

BIOLOGICAL NATURE OF THE STATE

has general relevance as a means of linking evsubjective intention,20

SOURCE: ModifiedfromHamilton (fn. I9), 207.

This approach analyzes social behavior in termsof each individual's

voring close kin as compared to more distant relatives or unrelated

BIOLOGICAL NATURE OF THE STATE

Williams (fn. I7); RobertTrivers,"Sociobiologyand Politics,"in White (fn. 7), 4-7.

Rationality(Cambridge,England: Cambridge UniversityPress, 1982).

Languageis, of course,whatenableshumansto engagein thisdual

Situationsdefinedin termsof Figure 1.

human institutioncan reston any one or a combinationof the fiveindividualmotives.

varietyand complexity,are constitutedby the attributesdefiningwho

and culturalrules. This means thatanimal instinctand human learning

TO THE TRAGEDY OF THE COMMONS

As Brian Barry has argued,36there are two broad traditionsin the

BIOLOGICAL NATURE OF THE STATE

has been widelyused in thesocialsciences,41and has also beenapplied

The game matrixforthisPrisoner'sDilemmais setforthin Figure

must be transitive(in the example, + io

ASSUMPTIONS: 2-persongame in which payoffsare transitive(+ IO > +9 > -9 > -I o)

Trivers (fns.22 and 25).

oner, regardlessof the response of the other. Rather than resultingin

response).Maynard-Smithhas shown thatthe locationof an interaction

Although this approach can readily explain nepotism and mutual

his herdwithoutlimit-in a worldthatis limited.49

46Jack Hirshleifer,"EvolutionaryModels in Economics and Law: Cooperation versus

Many ecologistshave agreed with Hardin's argumentthatthissituation

50 E.g., JohnN. Cole, "Learning a Hard Lesson in AroostookCounty,"Country

ASSUMPTIONS: as in Figure 3A.

the playersinto two groupsis a way out. If those

BIOLOGICAL NATURE OF THE STATE

mutualbenefitsforall membersas long as obedienceto social rules

55Donald T. Campbell, "Legal and Primary-GroupSocial ControlsCurbing Selfishand

I977). Cf. Machiavelli,Discourseson TitusLivy, I, ii and III, i; Plato, Republic,VIII.546a-

547c; Aristotle,Politics,V.ii.I3o2a-b and V.xii.13i6a-b.

theCommons,59and can be escaped onlythrougha freelyacceptedarbiter

Rousseau criticizes Hobbes in two differentways, each of which

According to Rousseau, therefore,Hobbes makes the mistakeof taking

(fn.i6), esp. I07,

izen."67 The willingness to die in defense of the state, dismissed by

Rousseau, Social Contract,II, iv (fn. 66), 64.

7? Roger D. Masters,"Nothing Fails Like Success: Developmentand Historyin Rousseau's

Ibid., i8o, I82.

79Georg Wilhelm Friedrich Hegel, The Philosophy

BIOLOGICAL NATURE OF THE STATE

84Ibid.,155-22385Ibid.,Para. i6i, p. III;

insight,can thus be illuminated by scientificresearch.As Trivers has

BIOLOGICAL NATURE OF THE STATE