Biology and the Social Sciences Author(s): Edward O. Wilson Source: Daedalus, Vol. 106, No. 4, Discoveries and Interpretations: Studies in Contemporary Scholarship, Volume II (Fall, 1977), pp. 127-140 Published by: The MIT Press on behalf of American Academy of Arts & Sciences Stable URL: http://www.jstor.org/stable/20024512 . Accessed: 01/06/2014 19:14

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This content downloaded from 130.92.9.57 on Sun, 1 Jun 2014 19:14:58 PM All use subject to JSTOR Terms and Conditions EDWARD O. WILSON

Biology and the Social Sciences

a correct statement a statement. But the of a The opposite of is false opposite truth.?Niels Bohr profound truth may well be another profound

an For every discipline in its early stages of development there exists antidisci for pline. For many-body physics, particle physics; chemistry, many-body molecular biol physics; for molecular biology, chemistry; for cellular biology, to ogy; and so forth. With the word antidiscipline I wish emphasize the special adversary relation that exists initially between the studies of adjacent levels of of a organization. This relationship is also creative, and with the passage great deal of time it becomes fully complementary. now to In this article I will argue that biology has moved close enough the social sciences to become their antidiscipline. Hitherto biology has affected the as social sciences largely through technological manifestations such the benefits of medicine, the mixed blessings of genetics, and the specter of population matters are growth. Although of great practical importance, these wholly trivial to foundation of the social sciences. The conven with reference the conceptual tional academic treatments of ''social biology" and "social issues of biology" are not present some formidable intellectual challenges, but they concerned with the core of social theory. Many scholars judge this core to be the deep structure of human an that is also the nature, essentially biological phenomenon primary focus of the humanities. If it is true that biology is the antidiscipline of the social sciences, the past a common failure of the social sciences to develop body of theory is under standable. The reason is that the relevant branches of biology?neurobiology now mature to a and sociobiology?are only becoming enough attain juncture to can with the social sciences. Although itwould be premature say that biology revolutionize the social sciences, at least to the extent that chemistry has revolu to cannot. tionized biology, it would be equally premature say that it This proposition is well worth examining. To that end I will first describe some precedents from the history of biology, then turn to developments within a to the individual social sciences that suggest growing susceptibility biological as a explanation. Finally, acknowledging my obligation student of the antidisci pline, I will attempt to define the obviously strong limitations of biological re ductionism. 127

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Disciplines andAntidisciplines a In general, the practitioners of given discipline in its early, natural-history are phase concerned with the discovery and classification of phenomena. They stress novelty and particularity. In terms of the classic thematic dualities of sci ence,1 their explanations are characteristically holistic, emphasizing pattern and are form over units of construction. In the early phase, specialists also likely are to be dualistic in philosophy, questioning whether their phenomena directly subject to the laws of the remainder of science.2 And in later stages, having been are more converted, they still concerned with what Victor Weisskopf has called as use to extensive opposed to intensive research, the of existing theory explain as to the widest possible range of phenomena opposed the search for fundamen tal laws.3 to Members of the antidiscipline are likely to be monistic with reference the own as discipline and dualistic with reference to their subject. Having chosen their primary subject the units of the lower of the paired levels of organization, can laws. they believe that the next discipline above be reformulated by their totem Their interest is relatively narrow, abstract, and exploitative, lacking the most ic attachment to phenomenology displayed by the devoted students of the discipline above. Thus P. A. M. Dirac, speaking of the theory of the hydrogen atom, could say that its consequences would unfold as mere chemistry, whereas to recent the biochemist Franz Hofmeister responded the great advances in cell more to en structure by recommending (in 1901) that biologists pay attention zymes. an A tense It is easy to see why each discipline is also antidiscipline. creative of are interplay is inevitable because the devotees of adjacent levels organization on committed to different methodologies when they focus the upper level. By a can as one a three today's standards broad scholar be defined who is student of to his subjects: his discipline, the lower antidiscipline, and the subject which as specialty stands antidiscipline. A well-rounded cellular neurophysiologist, of for example, is deeply involved in the microstructure and behavior single cells, but he also understands the molecular basis of electrical and chemical neuron to account transmission, and he hopes to explain enough of systems help more of animal behavior.4 for the elementary patterns

Cell Biology and Biochemistry at an In the late 1800s cell biology and biochemistry grew accelerating pace. was can Their relationship during this period very complicated, but it be broad as were ex ly characterized fitting the schema just described. The cytologists an cited by the mounting evidences of intricate cell architecture. They had also deduced the mysterious choreography of the chromosomes during cell division, setting the stage for the emergence of modern genetics and experimental devel on opmental biology. Many biochemists, the other hand, remained skeptical of the idea that so much structure exists. They emphasized the possibility of arti fact production by the chemical reactions used in cytological preparations and over stood apart from the debate then raging whether protoplasm is homoge or more neous, reticular, granular, foamlike. Their interest lay in the "funda

This content downloaded from 130.92.9.57 on Sun, 1 Jun 2014 19:14:58 PM All use subject to JSTOR Terms and Conditions BIOLOGY AND THE SOCIAL SCIENCES 129 mental" issues of the chemical nature of protoplasm, especially the newly formulated enzyme theory of life. too of to In general, biochemists judged the cytologists ignorant chemistry grasp the basic processes, whereas the cytologists considered the methods of the to structures that the cell. chemists inadequate characterize the diagnose living in The renewal of Mendelian genetics and subsequent progress chromosome at to a no immediate mapping did little first effect synthesis. Biochemists, seeing way to encompass classical genetics, by and large ignored it. now so Both sides were essentially correct. Biochemistry has explained own as most extrava much of the cellular machinery by its terms to justify its gant early claims. But in achieving this feat (mostly during the past thirty years) was new it partially transformed into the discipline of molecular biology?bio movements chemistry that entails particular spatial arrangements and of large a and molecules. Cytology forced the development of special kind of chemistry a new the employment of wide array of powerful methods, including electro phoresis, chromatography, density-gradient centrifugation, and X-ray crystal same cellular lography. At the time cytology metamorphosed into modern outlook biology. Aided by electron microscopy, it converged in language and toward molecular biology. Finally, classical genetics, by switching from Droso more and phila to the ultrafast-breeding and far simply constructed bacteria to viruses, has incorporated much of biochemistry become molecular genetics. a was Progress over large part of biology fueled by competition among the various attitudes and themata derived from biology and chemistry?the dis a cipline and its antidiscipline. Joseph Fruton, biochemist who has paid close com attention to this Hegelian interplay, has suggested that inevitably "such to petition is attended by tensions among the participants. I venture suggest that are source this competition and these tensions the principal of the vitality of biochemistry and are likely to lead to unexpected and exciting novelties in the as have in the future, they past."5

Ecology andPopulation Biology a as ten Modern ecology has had troubled history. As recently years ago it was painfully unfashionable in many American universities. An anecdote will illustrate how serious the situation had become. The Department of Biology at was Harvard University at that time increasingly dominated by molecular and new cellular biology, and there appeared to be little chance of adding faculty on or members specializing populations ecosystems. One afternoon I proposed a departmental membership for distinguished ecologist who had been appointed a new by another school within the university. One of the molecular geneticists, Nobel laureate, said, "Are they out of their minds?" When I asked what he an meant, he responded that anyone who hired ecologist must be out of his one mind. After the meeting of my senior colleagues in evolutionary biology not suggested that I refer explicitly to ecology in the future, because it had a was become "dirty word." The ecologist invited much later, after the molecu lar and cellular biologists had formed independent administrative units. at once Ecology is almost embarrassingly fashionable now, and I should add to are that the several ecologists subsequently added the Department of Biology

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on very cordial terms with all their colleagues. But the lesson learned at Har vard and at other universities with similar experiences was, I believe, that few scientists are to share resources with those whose research is more than willing one the central level of organization removed. Populations and communities, are an concern of ecology, separated from molecules and cells by entire level, an that of the organism. In environment ruled by competitive research, profit can be extracted only from the discipline and antidiscipline. was more of from The problem with ecology, however, than lack sympathy came molecular and cellular biologists. The difficulty ultimately from ecology's and the focus on the highest level of biological organization, the community, weakness of its connections to fundamental population biology. Without quite was an 1950s theoretical realizing it, ecology orphan discipline. By the popu one into a lation genetics, of the logical antidisciplines, had grown technically were formidable, even arcane subject. Its models derived from the distinctive chromosomal mechanics of Mendelian heredity, and it dealt almost exclusively with the of the Darwinian mutation interplay operators?selection, pressure, was reduced to the gene flow, genetic drift, and meiotic drive. Ecology single a parameter of the selection coefficient. By remarkable piece of bad luck, experi on are ments and field studies were concentrated Drosophila. Fruit flies superb insects for rapid genetic and demographic analysis in the laboratory, but they to to in habitats. Much of what are singularly hard find and study their natural was at food is known of their ecology deduced from their appearance traps. as a remained Partly result of this historical accident, population genetics apart from ecology.6 the fundamental Similarly, with the single exception of pure demography, not advanced much the theory of population ecology had beyond principles in 1920s and advanced by Alfred Lotka, Vito Volterra, and R. A. Fisher the at of entire commu early 1930s. Ecology, consisting largely of analysis the level as an ever more elaborate science. In some of its nities, developed descriptive of branches, such as biogeography and phytosociology, the systems classifica tion and reached extremes. quantitative description phantasmagoric a new The time was obviously ripe for effort that would bring population and cast them in their role as genetics and population ecology together proper 1950s 1960s a small antidisciplines to community ecology. In the late and early to a set about group of younger population biologists (I confess being member) an were held to discuss self-consciously to make such effort. Several meetings the matter and to some extent to divide the labor. The undoubted leader was the late Robert H. Mac Arthur, whose work on complex community phenomena discrimination of was exceptional in its originality and important problems.7 Mac Arthur's seminal contribution was his 1957 analysis of the population curve a bird ecolo abundance frequency of community of species.8 Descriptive a amount of information on this but had gista had accumulated large subject curves data to test a done little more than fit empirical to it.Mac Arthur used the on of various forms of interaction series of competing hypotheses based models a to that time had been diffi among species populations, process which up very nature. has had limited cult to study directly in Although his method only to it validated success when applied other biological communities, post can ulational-deductive theory and demonstrated that leaps of the imagination and them new lay open the most complex ecological processes give meaning.

This content downloaded from 130.92.9.57 on Sun, 1 Jun 2014 19:14:58 PM All use subject to JSTOR Terms and Conditions BIOLOGY AND THE SOCIAL SCIENCES 131 at The response of other ecologists to such renewed model building the was to population level predictably mixed. Many of those devoted painstaking so descriptive work refused to believe that general laws could be cheaply bought. The patterns they discerned seemed too elaborate, the variables too numerous, and the mark of history too deep and idiosyncratic to make general more a a models anything than clever illusion. A community of organisms is use tangled bank, to Darwin's famous phrase in the closing paragraph of the or a woven one Origin of Species, uniquely tapestry, in the metaphor of of the recent critics. Prominent ecological schools still exist, comprised of ecosystems so analysts, theoretical population ecologists, physiological ecologists, and versus over forth. It is cytology biochemistry all again. But the history of this now subject, if I have interpreted it correctly, is entering the middle phase of the classic Broad areas of are and model ontogeny. agreement already apparent, at building the population level has become routine.9

and The Social Sciences Biology us now return Let to the original proposition that biology is the antidisci to pline of the social sciences. This assessment is not congenial the prevailing view in the social sciences and humanities that human social life is the nearly exclusive product of cultural determinism, constrained only by the most ele a mentary and unstructured biological drives. There is strong tendency to think our own as of species entirely plastic and hence all but equipotent in the design of its social institutions. However, this conception will not stand close scrutiny. on A comparison of the literally tens of thousands of other highly social species Earth, from colonial coelenterates through the social insects to the most social of the birds and mammals, reveals that the sum of all the varieties of human social a behavior occupies only small envelope in the space of realized social arrange ments.10 a Anthropocentrism is disabling vice of the intellect. I am reminded of the clever Robert Nozick deflated our sense of to other animal way superiority spe cies in order to make his principal argument for vegetarianism. If visitors from to more another planet happened be far intelligent and sensitive than ourselves, our own and applied criteria of relativity, they could proceed in good conscience to eat us.11 the same and to our considerable scientists By token, chagrin, them find us as a social another cultur among might uninteresting species?just variant on the basic mammalian theme?and instead turn to al-linguistic study more the theoretically challenging societies of ants and termites. It is this quality of specificity and restriction that biologists have in mind to a when they speak of genetic determinism. In order define genetic trait pre it is to two or more states same cisely necessary compare of the character. To that are not say blue eyes inherited, without further qualification, is meaning are ful, because everyone knows that blue eyes the product of the interaction of and genes the largely physiological environment that brought final coloration to the irides. But to say that the difference between blue and brown eyes is based or a partly wholly upon differences in genes is meaningful statement by virtue of being testable and translatable into the laws of genetics. can same Human social behavior be evaluated in the way, first by com parison with the systems of other species and then, with far greater difficulty

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and ambiguity, by studies of variation within the species. For example, certain are most general traits shared with other Old World primates, including size of on intimate social groups the order of 10-100; males larger than females, prob in to a ably relation polygyny; long period of socialization in the young, shifting in to focus from the mother age- and sex-peer groups; and social play strongly on developed, with emphasis role practice, mock aggression, and exploration. It so is virtually inconceivable that primates, including human beings, could be cialized into the radically different repertories of insects, fish, birds, or ante or reverse lopes; that the could be accomplished. Human beings, by conscious design, might well imitate such arrangements; but it would be a fiction played out on a counter to emotional and with no chance stage, running deep responses as as a of persistence through much single generation. Homo sapiens is distinct from other primate species in ways that can be ex as a plained only the result of unique human genotype. Universal or near-uni versal traits include the facial expressions that denote basic emotions, and some other forms of paralinguistic communication; elaborate kinship rules that in a clude incest avoidance; semantic, symbolical language that develops in the a young through relatively strict timetable; close sexual and parent-offspring a bonding; and others. Again, to socialize human being out of such species specific traits would be very difficult if not impossible, and almost certainly destructive to mental development. People might imitate the distinctive social a arrangements of white-handed gibbon or hamadryas baboon, but it seems extremely unlikely that human social systems could be stably reconstructed by such an effort. a It is significant that not only do human beings develop species-character set are istic of social behaviors, but that these behaviors generally mammalian, and most specifically Old-World primate in character. Furthermore, even the are cases species-specific traits logically derivable in some from the inferred a ancestral modes still displayed by few related species. For example, the facial some can expressions and nonlinguistic vocalizations be plausibly derived in phylogenetic reconstructions.12 This is precisely the pattern to be expected if the human species was derived from Old-World primate ancestors (a fact) and in still retains genetic constraints the development of social behavior (a hy pothesis). a An important quality of genetic determinism is that it seldom entails the a a control of single phenotype by single gene. Poly genie inheritance is the rule, a and the entity determined is not one trait but rather range of possible pheno com types. For example, diabetes and schizophrenia possess moderate genetic a to ponents. The multiple genes underlying them produce stronger tendency develop the traits; they also prescribe the range of possible manifestations that are a probable under specified environmental conditions. In parallel way basic as as human social behaviors?including those structured male-group bonding, as outermost territoriality, and kinship rules?emerge phenotypes following be havioral development, the range and scope of which is constrained by the inter action of poly genes with the environment. With reference to this interaction, no reason as there is to regard most forms of human social behavior qualitatively different from physiological and nonsocial psychological traits. our Whatever the present social arrangements of species, the biological foun

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dation of human nature arose in populations that adapted to special environ were ments. The prevailing hypothesis, which holds that the basic qualities fashioned as an to a more existence in adaptation predatory open habitats, may con or may not be correct in detail. The important point is that the emotional are to trols and the developmental pathways considered be structured in idio can syncratic ways that be wholly understood only by retracing the ecological a history of the species.13 That such relationship exists in other social species can be readily demonstrated.14 This is why paleoanthropology, by reconstruct an to ing the Pleistocene African environment, has important role play in behav ioral biology. extent to An unavoidable question is the which social behavior varies geneti a cally within the human species. This is subject entirely removed from the distinctive properties of human behavior vis-?-vis that of other species. The evidence is strong that almost but probably not quite all differences among cul are on on same tures based learning and socialization rather than genes.15 At the time variation within populations is evidently great enough to create the poten tial for further human social evolution by population-wide genetic change. same-sex a Studies comparing monozygotic twins with dizygotic twins suggests a an genetic component in the variation of large array of traits having influence on the development of social behavior. These traits include verbal and number ability, word fluency, perceptual speed, memory, the timing of language acqui and certain sition, psychomotor skill, extroversion-introversion, homosexuality, forms of neuroticism and psychosis.16 Although not conclusive in themselves, are such studies strongly suggestive. Behavioral heritability is also enhanced by the undoubted existence of single, identifiable point mutations and chromosome such as those the and Turner's aberrations, causing Lesch-Nyhan syndromes, that alter various components of behavior differentially. Some geneticists have so as once gone far to suggest that the conditions make their appearance in spite of medical precautions, their study can permit the indirect genetic dissection of behavioral traits, in analogy to the technique used for nematodes and fruit flies. a My overall impression of the available information is that Homo sapiens is to typically animal species with reference the quality and magnitude of the genetic diversity affecting behavior. If the comparison is correct, the psychic a a unity of humankind has been reduced in status from dogma to testable hypothesis. This is not an easy thing to say in the present political ambience of as the United States, and it is regarded punishable heresy in some sectors of the to academic community. But the idea needs be faced squarely if the social sci ences are to cannot as con be entirely honest. I regard it dangerous. Quite the trary: the political consequences of its objective examination will be determined our not reverse. by value system, the It will be better for scientists to study the to a subject of genetic behavioral diversity than maintain conspiracy of silence out to of good intentions and thereby default ideologues. Following this elementary excursion into genetic determinism, let us now consider how the various social sciences might be influenced by biological theo ry. In the brief sections to follow I have little more than scraps of information to and impressions offer. It is hoped that they will nevertheless suffice to pro vide a view of the reverse side of the as a social sciences it has been glimpsed by biologist approaching in that direction.

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Anthropology nature The central question of biological anthropology is the and strength of the coupling between cultural and biological evolution. Cultural evolution is Lamarckist and usually very fast; biological evolution is Darwinist and slower at an most by least order of magnitude. Because the rapid cultural changes track too to environmental fluctuations brief in duration influence directional genetic can to selection, cultural fitness be expected diverge frequently from genetic must fitness.17 But the divergence be limited in degree and duration, because ultimately the newly created social environment will be tracked and the gap narrowed by natural selection. If this modification of basic sociobiological theo ry is correct, there should be two detectable consequences. First, it will be are learning rules rather than specific cultural forms that inherited.18 And are most as where the rules directly concerned with survival and reproduction, case in the of sexual and parental bonding, incest avoidance, and xenophobia, are most they likely to be the rigid and structured. Marshall Sahlins has recently argued that the lack of clear correspondence between human kinship rules and the details of genetic kin selection theory disproves basic sociobiological theory are too as far as human beings concerned.19 But this goes much far. Kinship rules are central to social in most and in organization societies, aggregate they appear to enhance inclusive genetic fitness. Richerson and Boyd20 have sug can gested that four competing hypotheses be posed for the explanation of kin ship rules: detailed genetic control, rational strategizing, complete cultural determinism, and coupled cultural and genetic control. They conclude that the are ethnographic facts consistent only with the model of coupled cultural and control. genetic con The second expected consequence of coupling is that the genetic fitness ferred by particular cultural traits should be strongest in the oldest, culturally most it correct to focus attention on stable societies. Thus appears hunter-gatherer and persistent, preliterate herding and agricultural societies. This circum stance explains why anthropology has already become the social science closest are to sociobiology. Explicit tests of sociobiological theory being made in studies on societal and warfare.21 polygyny, status, fissioning, territory, to ac It is probable that population biology will be simultaneously altered commodate the special problems of anthropology. In addition to the solution of a the dual inheritance problem, there is need for advances in the theory of group to transac and kin selection distinguish unilateral, "hard core" altruism from tional, "soft-core" altruism. Also, the complexity of human population struc to are tures presents unique challenges biology. Population boundaries seldom sharp, often being confused by discordant linguistic, cultural, and historical political patterns. Groups also shift rapidly in their loyalties, forming alliances one next. in year and dividing into quarreling factions the The present theory of to com population genetics and ecology is entirely inadequate handle such plications.22

Psychology and Psychoanalytic Theory as com Just anthropology has been burdened in the past by the doctrine of plete cultural determinism, conventional psychology has been burdened by

This content downloaded from 130.92.9.57 on Sun, 1 Jun 2014 19:14:58 PM All use subject to JSTOR Terms and Conditions BIOLOGY AND THE SOCIAL SCIENCES 135 was general process learning theory. It natural for Thorndike, Watson, and to mam other pioneering psychologists choose large animals, that is, birds and mals, for their study objects, rather than insects and opisthobranch mollusks. came as Partly as a result of this choice, learning to be accepted the central was as essen process of behavior. Moreover, the learning process viewed being same were to to tially equipotential: the laws theorized apply whatever learning are process and organism chosen. Thus Skinner could say in 1938, "The general not most topography of op?rant behavior is important, because if not all specific are op?rants conditioned. I suggest that the dynamic properties of op?rant be a was havior may be studied with single reflex."23 It believed that by placing can animals in simplified laboratory environments, where stimulation be rigidly the most laws would controlled, general emerge. a This is powerful idea, with seductive precedents in the physical sciences, and it has resulted in substantial advances in the study of animal and human to behavior. Nevertheless, general process learning theory has started crumble. a In its place is appearing the description of melange of specialized learning that conform to no law evolution natu phenomena general except, perhaps, by ral selection.24 The full range of learning potential of each species appears to be to separately programmed. According the species, each animal is "prepared" to to learn certain stimuli, counterprepared learn others, and unprepared (neutral) to with respect still others. For example, adult herring gulls quickly learn to never own distinguish their newly hatched chicks but their eggs, which are as nevertheless visually diversified. Indigo buntings are prepared to learn the circumpolar constellations by which they orient their nocturnal migrations, but are counterprepared to learn other constellations. When chicks are shocked at water a their beaks while drinking and simultaneously given visual stimulus, they thereafter avoid the visual stimulus, but they do not learn to avoid an same auditory stimulus presented the way. The reverse is true when the shock to are is administered the feet; that is, the chicks prepared to learn sound but to counterprepared learn visual cues.25 The timing of preparedness in the life are cycle is also programmed and species-specific in ways that readily inter as to preted adaptations the particular environments experienced by the species their recent during evolutionary pasts. as The hypothesis of learning rules idiosyncratic evolutionary adaptations to seems has seldom been examined with reference the human species. It signifi cant that some phobias, which share properties with imprinting in lower ani are and other mals, readily acquired against snakes, spiders, rats, potentially in dangerous objects mankind's ancient natural environment, but only rarely such as against dangerous artifacts knives and electrical outlets.26 Language is a acquired by small children through progression of closely timed steps, in distinctive volving vocalizations and phrase forms later replaced by adult lan It is difficult to believe that the rules of this most guage.27 human of all learning events not have been shaped by natural selection. to Psychoanalytic theory appears be exceptionally compatible with socio a fact some biological theory, already appreciated by of the psychoanalysts themselves.28 If the essence of the Freudian revolution was that it gave structure to the unconscious, the logical role of sociobiology is to reconstruct the evolu tionary history ofthat structure. When Freud speculated in Totem and Taboo on the primal father, primal horde, and the origins of the incest taboo, he created a

This content downloaded from 130.92.9.57 on Sun, 1 Jun 2014 19:14:58 PM All use subject to JSTOR Terms and Conditions 136 EDWARD O. WILSON a one. same true sociobiological hypothesis, but poor The is of his insights into the conflict of self and society presented in Civilization and Its Discontents. can to restructure Whether population biology and evolutionary theory be used some to and objectively test of psychoanalytic theory remains be seen.29

Economics

can Classic economic theory restricted itself to the goods and services that be measured and market In recent less by money pricing. years new, easily quan tified parameters have entered the equations, including time, human capital, and environmental quality.30 A closer scrutiny is also being made of what Lei benstein has termed the X-efficiency factors, which include motivation, esprit, made In a effort, persistence, and other psychological variables mensurable.31 now word, microeconomics has begun to incorporate social psychology. It is not as economic widely appreciated that human beings do behave rationalizing machines. Macroeconomic predictions of the future will almost certainly be in nature. based on the wiser perception of irrational elements human are To the extent that the new parameters of human irrationality interpreted as an evolutionary product, the methods of economics will converge toward borrowed those of biology. Already, models in ecology and sociobiology have and decision heavily from the graphical methods of economics. Optimization measure theory are routinely used. The utility of biology is genetic fitness, and are I that once the enabling devices anatomy, physiology, and behavior. expect a method is developed for assessing the coupling of genetic and cultural evolu will come to tion, the utility measures of economics and evolutionary biology overlap broadly. An economics is also a recent interspecific comparative possibility. During research on the evolution of division of labor in insect societies, George Oster and I have written a short book that resembles a textbook in microeconomics.32

Insect economics differs in several respects: the transactions among colony members are almost exclusively instead of merely partially instinctive (that is, are and and of the "irrational"), the societies mostly sterile female, (because far than haplodiploid genetic bias) unilateral altruism has greater genetic utility are in human societies. The broad forms of the analyses otherwise much the is not but same. The point is that human economics really general economics, in one mammalian with a rather the description of economic behavior species state limited range of the biological variables.

Sociology of By virtue of its loftier perch in the hierarchy subject matter, sociology most should be the queen of the social sciences. Yet I personally find it the alien cause statement and least interesting. Part of the is revealed in the following by same Durkheim: "In a word, there is between psychology and sociology the Con break in continuity as between biology and the physicochemical sciences. a is a sequently, every time that social phenomenon directly explained by psy we be sure that the is false."33 I chological phenomenon, may explanation as as it to suspect that this statement is completely wrong for sociology proved

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would ac be for biology. Although few contemporary sociologists uncritically on cept The Rules of Sociological Method, Durkheim's dualism lives by tradition. Sociological analysis seldom utilizes the known facts of social psychology to any depth, and evolutionary biology remains all but taboo. The specters of biolo are gism and social Darwinism still feared, entirely without justification.34 The situation is so extreme that I suspect that progress in the near future will be measured by the connections sociology makes with its antidisciplines. To the an extent that it does not make these connections, itwill remain ad hoc, descrip tive science. not to Yet sociology is destined be cannibalized by the antidisciplines, any more was reason than cytology absorbed by biochemistry. The is that sociology is truly the subject most remote from the fundamental principles of individual behavior. Advanced literate the main concern of are the societies, sociology, most removed in character from the kinds of social and economic systems in which the genetic basis of human social behavior evolved. Having been jer on are ry built the Pleistocene human biogram, they the least stable, probably have the greatest discrepancies between genetic and cultural fitness, and hence are most to not a likely display emergent properties predictable from knowledge of individual psychology alone.

The Limits ofReductionism

Karl von Frisch once made a remarkable statement about his research. He a more said that the honeybee is like magic well?the you draw from it, the more there is to draw. Other students of social insects share this sense of seem ingly infinite richness in the phenomena of colonial life. They have learned that a can a great deal of evolutionary novelty at the social level be generated by only at small amount of genetic change the level of the individual. A slight modifica one can a tion in parameter of allometric pupal growth, for example, produce new of whereas an altered to a can create a array castes; response pheromone new mode of communication. cannot a The full phenomenology of social life be predicted from knowledge of the genetic programs of the individuals alone. When the observer shifts his one attention from level of organization to the next, he expects to find obedience to all of the laws of the levels below. But upper levels of organization require specification of the arrangement of the lower units, which in turn generates new richness and the basis of and unexpected principles. The specification can be classified into three categories: combinatoric, spatial, and historical. Thus the ammonia molecule neutralizes its electric dipole moment and conserves the laws of nuclear physics by inverting the negatively charged nitrogen back and forth at a X through the triangle of positively charged hydrogens frequency of 3 1010 per second. But this symmetry is broken in the case of sugar and other are too larger organic molecules, which large and complexly structured to invert not themselves. They break but do repeal the symmetry laws of physics.35 This not specification may be of great interest to particle physicists, but its effects redound throughout organic chemistry and biology. Primitive wasps, comprising early members of the order Hymenoptera, evolved sex the determination mechanism of haplodiploidy, whereby unferti

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lized eggs yield males and fertilized eggs yield females. This mechanism may a have been specific adaptation that permits females to choose the sex of the to nature are offspring according the of the separate prey items they able to subdue. But whatever the initial cause, haplodiploidy represented a historical accident that these insects to predisposed develop advanced forms of sociality. reason causes to more The is that it sisters be closely related genetically than mothers are to so a daughters, and they find genetic profit in becoming sterile caste an specialized for the rearing of sisters. As apparent result, social life to among insects is almost limited the phylogenetically advanced hymenopter the social social ants. most cases ans, namely wasps, bees, and Furthermore, of insect social life can be classified either as matriarchies, in which queens control or as colonies of daughters, sisterhoods, in which sterile daughters control the egg-laying mothers. Many other strange effects flow from this genetic asymme try. In addition, the hymenopterous societies have proved so successful that they dominate and alter much of the terrestrial ecosystems of the Earth.36 Who a could have guessed all of this from knowledge of haplodiploidy? The urge to be a reductionist is an understandable human trait. Ernst Mach as a captured it in the following definition: "Science may be regarded minimal problem consisting of the completest presentment of facts with the least possible a expenditure of thought."37 This is the sentiment of member of the antidisci to set on pline, impatient aside complexity and get with the search for more are fundamental ideas. The laws of his subject necessary to the discipline above, a more they challenge and force mentally efficient restructuring; but they are not sufficient for its purposes. Biology is the key to human nature, and social cannot to scientists afford ignore its emerging principles. But the social sciences are content. potentially far richer in Eventually they will absorb the relevant on ideas of biology and go to beggar them by comparison.

References to 1G. Holton, Thematic Origins of Scientific Thought: Kepler Einstein (Cambridge, Mass.: Harvard University Press, 1973). 2 A useful ontogeny of disciplines, starting with the natural history phase, has been outlined by F. S. C. Northrop, The Logic of the Sciences and Humanities (New York: Macmillan, 1947). 20 3V. F. Weisskopf, "Nuclear Structure and Modern Research,"Physics Today, (1967): 23-26. to 4An admirable example of this approach neurobiology is provided by E. R. Kandel, Cellular Basis of Behavior (San Francisco: Freeman, 1976). 192 recent 5J. S. Fruton, "The Emergence of Biochemistry, "Science, (1976): 327-334. Another one product of this interplay is molecular immunobiology. G. M. Edelman, of the leading contrib was was neces utors, has noted that "the idea of specificity and molecular recognition prior to, and at sary but not sufficient for, the idea of clonal selection which operated the level of cells. In turn, at the particular picture of variability and constancy which emerged the level of the molecular to at it analysis of antibodies could be applied refine the idea of clonal selection those points where seemed most dubious." See "The Problem of Molecular Recognition by Systems," in F. J. Ayala in of and T. Dobzhansky (eds.), Studies the Philosophy of Biology (Berkeley: University California Press, 1974), pp. 45-56. 6This imbalance in experimental population genetics is well illustrated in the work of T. Dob zhansky and his associates; see T. Dobzhansky, Genetics of the Evolutionary Process (New York: Co lumbia University Press, 1970). They made brilliant progress in elucidating the genetic processes of unaware natural selection and speciation but remained largely of the central issues of ecology. In recent more on years someDrosophila geneticists have concentrated deliberately natural history, and more now information useful for ecological analysis is accumulating rapidly. 7See E. O. Wilson, "Eminent Ecologist, 1973: Robert Helmer MacArthur," Bulletin of the Eco 11-12. logical Society of America, 54 (1973):

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8R. H. Mac Arthur, "On the Relative Abundance of Bird Species," Proceedings of the National Academy of Sciences, United States, 43 (1957): 293-295. 9See, for example, R. M. May (ed.), Theoretical Ecology: Principles and Applications (Philadelphia: Saunders, 1976). 10E. O. Wilson, Sociobiology: The New Synthesis (Cambridge, Mass.: Belknap Press of Harvard University Press, 1975). 1 Robert Nozick, Anarchy, State, and Utopia (New York: Basic Books, 1974). A. van e to 12J. A. R. M. Hooff, "A Comparar Approach the Phylogeny of Laughter and Smiling," in R. A. Hinde (ed.), Non-verbal Comr nication (Cambridge: University Press, 1972), pp. 209-241. i 13G. E. King, "Socioterritorial Units and erspecific Competition: Modern Carnivores and Early Hominids," journal of Anthropological Research (1977), in press. 14E. O. Wilson, Sociobiology. motor 15D. G. Freedman has provided evidence of average differences in activity and excitabili in neonates ty between Navajo, Japanese, and Caucasian populations. Sec Human Infancy: An Evolu to at tionary Perspective (New York: Wiley, 1974). Such differences could lead least slight variations in the ways infants are handled and carried. 16L. Ehrman and P. A. Parsons, The Genetics of Behavior (Sunderland, Mass.: Sinauer, 1976); G. E. McClearn and J. C. DeFries, Introduction to Behavioral Genetics (San Francisco: Freeman, L. L. Heston and in a 1973); J. Shields, "Homosexuality Twins: A Family Study and Registry Archives General 18 E. H. Study," of Psychiatry, (1968): 149-160; Lenneberg, "On Explaining Lan 164 B. D. guage," Science, (1969): 635-643; Davis and Patricia Flaherty (eds.), Human Diversity: Its Causes and Social Significance (Cambridge, Mass.: Ballinger, 1976). 17P. "A J. Richerson and R. Boyd, Dual Inheritance Model of the Human Evolutionary Proc ess, "Journal of Theoretical Biology (1977), in press. 18See M. W. Feldman and L. L. Cavalli-Sforza, "Cultural and Biological Evolutionary Process es: Selection for a Trait under 9 Complex Transmission," Theoretical Population Biology, (1976): 238? 259. These authors an model for the selection of that alter provide explicit genes the learning rules of cultural transmission. 19M. The Use and Abuse Arbor: Sahlins, of Biology (Ann University of Michigan Press, 1976). 20Richerson and Boyd, "A Dual Inheritance Model of the Human Evolutionary Process." an 21See for example N. A. Chagnon, "Fission in Amazonian Tribe," The Sciences, 16 (1976): 14 18; and W. H. Durham, "Resource Competition and Human Aggression, Part I: A Review of Primitive War," Quarterly Review of Biology, 51 (1976): 385-415. 22E. O. "Some Central Problems of Social Wilson, Sociobiology," Sciences Information, Biology and Social Life, 14(1975): 5-18. 23B. F. Skinner, The Behavior of Organisms: An Experimental Analysis (New York: Appleton, 1938). to Skinner has since broadened his view incorporate genetically programmed behavior patterns as the of learned behavior in see analogs lower animals; "The Phylogeny and Ontogeny of Behavior," not Science, 153 (1966): 1205-1213. He differs from most biologists in his view of the basic mecha nisms amount occurs of learning and instinct, but in the of structuring that in the human brain. are most Matters of degree generally the easily resolvable in science. 24Summarized in M. E. P. and L. Seligman J. Hager (eds.), Biological Boundaries of Learning (Englewood Cliffs, N.J.: Prentice-Hall, 1972); and R. A. Hinde and J. Stevenson-Hinde (eds.), on Constraints Learning: Limitations and Predispositions (New York: Academic Press, 1973). A contrary view has been M. E. can expressed by Bitterman, who concludes that underlying principles still be drawn, at least for the vertebrates (Science, 188 (1975): 699-709). to 25S. J. Shettleworth, "Conditioning of Domestic Chicks Visual and Auditory Stimuli," in Seligman and Hager (eds.), Biological Boundaries of Learning, pp. 228-236. 26M. E. P. "Phobias and in Seligman, Preparedness," Seligman and Hager (eds.), Biological Boundaries of Learning, pp. 451-462. 27R. A First Brown, Language (Cambridge, Mass.: Harvard University Press, 1973). 28For R. The the a example, J. Lifton, Life of Self: Toward New Psychology (New York: Simon and Schuster, 1976). 29The same is true of which is to a considerable Marxism, degree sociobiology without biology. 30P. A. Samuelson, Economics, 10th ed. (New York: McGraw-Hill, 1976). 31H. Economic Man: A New Foundation Leibenstein, Beyond for Economics (Cambridge, Mass.: Har vard University Press, 1976). 32G. F. Oster and E. O. Caste and Wilson, Ecology in the Social Insects, in preparation. 33E. The Rules 8th Durkheim, of Sociological Method, ed., translated by S. A. Solovay and J. H. Mueller (New York: Free Press, 1938). 34The last remnants of social Darwinism died with the advent of sociobiology, which delineated the roles of and altruism in societies and cooperation rendered them consistent with population for the R. genetics. See, example, exposition by Dawkins, The Selfish Gene (New York: Oxford

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van University Press, 1976). A frankly biological critique of sociology has been provided by P. L. den Berghe inMan in Society: A Biosocial View (New York: Elsevier, 1975). 35P. W. Anderson, "More is Different," Science, 177 (1972): 393-396. 36E. O. Wilson, The Insect Societies (Cambridge, Mass.: Belknap Press of Harvard University Press, 1971). 37E. Mach, The Science ofMechanics, 9th ed. (LaSalle, 111.: Open Court, 1942).

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