Nature-Nurture and the Debates Surrounding Ethology and ...

AMER. ZOOL., 31:286-296 (1991)

Nature-Nurture and the Debates SurroundingEthology and Sociobiology1

GEORGE W. BARLOW

Department of Integrative Biology and Museum of Vertebrate Zoology,University of California, Berkeley, California 94720

SYNOPSIS. The central problem in the history of animal behavior has been the inabilityto perceive the phenotype as the result of an interaction between genome and environ-ment, despite the considerable lip service paid to the interaction. In North America thecomparative study of animal behavior was overshadowed by the growth of an experimentalpsychology that produced the general-process view of learning, holding that the mecha-nisms underlying learning are much the same in all species. That made evolution irrelevant.During the same period ethology emerged in Europe as the study of naturally occurringbehavior in an evolutionary context. Because evolution is fundamental to ethology, thegenetic basis of behavior was a central precept. Ethology and psychology collided afterWorld War II. After a vigorous exchange on the issues, a synthesis by Robert Hindematerialized, one that advanced the study of behavior and produced a sophisticated under-standing of nature and nurture. A few decades later sociobiology appeared and wasimmediately assailed for making what were seen as unwarranted extensions from animalto human behavior, and for emphasizing genetic control of behavior. Much of the debatethat ensued was distractingly political and threw little light on the scientific merits of theissues although it moderated the stance of sociobiologists; on the other hand, the politicallyinspired debates may have harmed the field of animal behavior.

INTRODUCTION

"The yin-yang principle is not . . . whatwe would ordinarily call a dualism, butrather an explicit duality expressing animplicit unity" (Capra, 1983, p. 26).

To what extent is the development ofbehavior molded by its genetic substrate asopposed to the pervasive effects of theenvironment? That question has engagednatural historians and philosophers forcenturies (Richards, 1987), perhaps becauseof society's fascination with the inheritanceof behavior, whether cultural or genetic(e.g., Twain, 1894). Why does the issue sointrigue people, even in the seeminglyinnocuous framework of animal behavior?And why do people have such great diffi-culty admitting that nature and nurtureare inseparable, even though they can beanalyzed separately? Recent events in thestudy of animal behavior provide someinsight.

The nature-nurture issue is the domi-

1 From the Symposium on Animal Behavior: Past,Present, and Future presented at the Annual Meetingof the American Society of Zoologists, 27-30 Decem-ber 1989, at Boston, Massachusetts.

nant recurrent theme that has generatedacrimony and misunderstanding. Theethology debate was potentially highly per-sonal and political, but a rapprochementproduced a productive resolution (Hinde,1966). In contrast, the controversy sur-rounding sociobiology yielded a less posi-tive outcome because the scientific issueswere so clouded by the political nature ofthe debate.

I am not a historian. I merely offer aperspective, one appropriate for an audi-ence of biologists. It is a personal, and willbe to some a provocative view but one thatis meant to cause readers to examine bothsides of the issues.

EVENTS LEADING UP TO ETHOLOGY

Many date the start of the modern studyof animal behavior with Darwin's (1872)book, The Expression of the Emotions in Manand Animals. More relevant to this essay,however, was that Darwin (1871) treatedhuman and animal behavior on a contin-uum. Biologists long before him, and evensome noted theologians, had commonlyassumed such a continuity (Richards, 1987).But now, together with his persuasive pro-motion of the principles of natural selec-tion, biology impinged on the realm of

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NATURE-NURTURE: ETHOLOGY AND SOCIOBIOLOGY 287

morals and ethics in a way that alarmedtheologians and others. Many notable biol-ogists attacked Darwin because of theinference that even the moral sensitivitiesof humans resulted from organic evolution(Richards, 1987).

Less familiar to biologists was the largelyindependent formulation and advocacy byHerbert Spencer of what was to be calledsocial Darwinism. According to Kalikow(1980) and Stein (1988) the biologist ErnstHaeckel elaborated a particularly perni-cious expression of this doctrine, thoughRichards (1987) presented a gentler inter-pretation of Haeckel's world view. Haeck-el's conception, based on genetic deter-minism and natural selection, was seen byStein (1988) as advocating adherence tothe dictates of society and as good stuff fortotalitarian governments. Haeckel was saidto espouse eugenics, racism (Kalikow,1980), and the inherent inequality of indi-viduals and the sexes (Richards, 1987).Stein (1988) perceived Haeckel's mainthemes in Adolph Hitler's Mein Kampf.

The Zeitgeist of the nineteenth century,however, was that socioeconomic statusresulted from superior "blood"; racism andsexism were rampant, even in the writingsof Darwin (e.g., 1871). Lamentably, a num-ber of biologists in the early part of thetwentieth century favored eugenics, andsome of them worked in animal behaviorin North America (Dewsbury, 1989).

I mention this extension of Darwin's the-ory to political philosophy because thatphilosophy became anathema in NorthAmerica during the twentieth century,especially among experimental psycholo-gists. Consequently, it was a powerful com-ponent underlying the altercation thatdeveloped after World War II (WW II)between the ethologists and American psy-chologists. It also makes understandablesome of the heated rhetoric during thesociobiology quarrel, especially thatinvolving biological determinism. Now letus turn to the situation in the field of ani-mal behavior in North America before theadvent of ethology.

Dewsbury (1989) painstakingly docu-mented the important contributions of sev-eral biologists and comparative psycholo-

gists to the study of animal behavior in theperiod preceding and following the turn ofthe century. The picture that surfaces is arich mixture of field and laboratory inves-tigations on a remarkable variety of ani-mals, vertebrate and invertebrate. Oneleading group was the Allee school at theUniversity of Chicago; they becameincreasingly occupied with two issues,dominance hierarchies and cooperation(Banks, 1985); their conception of coop-eration, however, differed from our cur-rent image, was organized around groupselection, and died out.

Psychologists followed a different path.Some continued to be comparative in thebest sense of the word (Dewsbury, 1989).And some were homing in on concepts thatlater emerged as central tenets in ethology(Glickman, 1985; Thorpe, 1979).

Those contributions of comparative pys-chologists, however, were overshadowedby the proliferation of experimental psy-chologists pursuing learned behavior. Themain thrust of that school was that the basicprocesses of learning are common to allvertebrates. This became known as thegeneral-process model of learning, as pio-neered by Thorndike (1911; see Glickman,1985). Because learning is general, it wasreasoned, one need not consider phyleticrelationships, hence also not evolution.

This perspective underlies the publica-tions of a substantial literature involvingfamous experimental psychologists (seeLehrman, 1971), culminating in the per-suasive papers and books of B. F. Skinner(e.g., 1959): "Pigeon, rat, monkey, whichis which? It doesn't matter."

Psychologists were not entirely contentwith this view, however. Lehrman (1971)faulted the experimental psychologists forfeeling that the laboratory rat and humanswere fundamentally the same with regardto "learning, motivation, sensation, socialdominance, etc." (p. 462). Earlier, FrankBeach (1950, p. 119) had written:

"From its inception, American psy-chology has been strongly anthropocen-tric. Human behavior has been acceptedas the primary object of study and thereactions of other animals have been of

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288 GEORGE W. BARLOW

interest only insofar as they seemed tothrow light upon the psychology of ourown species. There has been no con-certed effort to establish a genuine com-parative psychology in this country forthe simple reason that with few excep-tions American psychologists have nointerest in animal behavior per se."

The overshadowed branch of compara-tive psychology most relevant to the ethol-ogy controversy was the one inspired byZ.-Y. Kuo. He emphasized the importanceof development, including embryogenesis.In his early papers (e.g., Kuo, 1922, 1929)he rejected any role for genetics in behav-ior, a view he later moderated (Kuo, 1967).Kuo (1967) also echoed the negative viewthat many experimental psychologists hadtoward field studies as compared with thetightly regulated circumstances of labora-tory experimentation. The school thatdeveloped around T. C. Schneirla, follow-ing Kuo, made development central to theircomparative psychology (e.g., Maier andSchneirla, 1935); although the evolution-ary perspective entered in, it was treatedgingerly and kept at arm's length (Rich-ards, 1987).

Even Frank Beach, who rebuked theexperimental psychologists for not beingtruly comparative, did not appreciate theissue of adaptiveness of behavior. I recallwell that after his lecture on copulation indogs, which I heard a number of times,biologists present predictably asked aboutthe functional significance of the genitallock. Beach, just as predictably, became vis-ibly annoyed because he regarded thequestion as trivial. Ethologists, in contrast,were concerned with just such questions ofadaptive significance.

When you appreciate the stress of theSchneirla school, including Beach andLehrman, on development, and their sus-picion of evolutionary arguments, youbegin to understand some of the roots ofthe debate. Glickman (1985) observed thatfor psychologists in general, evolutionarytheory was (and to an extent remains) sus-pect because of three beliefs: (1) The the-ory is circular. (2) Humans are plastic, soevolution is irrelevant. And (3) social Dar-

winism has unpalatable ramifications (seealso Crawford, 1987; Hodos and Camp-bell, 1969).

THE RISE OF ETHOLOGY

Shortly after the close of WWII, KonradLorenz and Niko Tinbergen togetheradvanced an approach to thinking about,and studying, animal behavior, calledethology, that was new to North America.In part, ethology was a reaction against thesubjective interpretations of behavior thatcharacterized one school in Europe as wellas unorthodox interpretations of" Darwin-ian evolution (Richards, 1987). Lorenz andTinbergen held that ethology, in contrast,was the objective study of behavior, whichoffended the North American psycholo-gists who viewed their discipline as yet moreobjective than ethology.

But what truly distinguished ethologistswas that they dealt "with behavior patternsdrawn from the life cycle of the animalsdiscussed, rather than with the laboratorysituations most often found in Americancomparative psychology" (Lehrman, 1953).The Tinbergen school was particularlyactive in field studies (e.g., Tinbergen et al.,1962). In contrast, the Lorenz group wasnot. Typically, but not always, Lorenzianswent to pains to recreate a semblance ofthe natural habitat in captivity, or to employa seminatural situation, such as fish inaquaria, jackdaws nesting in an attic, andducks and geese on ponds. It was thisframework that permitted Lorenz to rec-ognize behavior that was inappropriatelytriggered in captivity (Lorenz, 1950, 1965).

The supporting framework for etholo-gists was evolution. Little attempt was madein the beginning, however, to show howbehavior is adaptive. The assumption ofadaptation was nonetheless importantbecause it guided them to questions aboutbehavior that had been all too oftenignored, as in their studies of communi-cation and displacement behavior.

The attention of ethologists in the begin-ning was drawn to naturally occurringbehavior of animals. That enterprise, how-ever, grew into an attempt to providereductionistic models for neurobiologists,probably because of Lorenz's early train-

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ing in medicine and his close associationwith Erich von Hoist; that lineage persistsas neuroethology. Lorenz and Tinbergenwere also concerned with learned behav-ior. Much of the early work of Tinbergenand his students, most notably GerardBaerends, was on learning in digger wasps,which led to the then controversial prop-osition of one-trial learning (Tinbergen,1951).

For his part, Lorenz wrote about learn-ing under a number of different cir-cumstances. His major contribution wasdrawing attention to the remarkable phe-nomenon of imprinting among ducks andgeese. Lorenz distinguished sharplybetween instinctive and learned compo-nents of behavior. In his theoreticalscheme, development was a process ofintercalating instincts and learning (Lorenz,1937, 1965).

THE CONFLICT BETWEENETHOLOGY AND PSYCHOLOGY

The field of ethology rapidly gainedattention, particularly for its stress oninnate behavior. It was not long after WWII before a critical review was published bythe comparative psychologist Daniel Lehr-man (1953). Reading the article today, itsessence does not seem so outrageous, butit did to ethologists at the time. Lehrman'sfrequent use of terms such as "gratuitous,""rash," "patently shallow," "facile," "pre-formationistic," and "preconceived andrigid ideas" were less vexing than the sub-stance of the penetrating critique. To fullyappreciate the undercurrents of the attack,you have to know more about the individ-ual players.

At that time, Lorenz's critics suspectedhe had been a Nazi. He rose rapidly, forsuch a young academician, to Kant's chairat Konigsberg. His writings, especially anarticle in 1940 (Lorenz, 1940), could beconsidered (e.g., Kalikow, 1980; Klopfer,1977; Richards, 1987) to have been instru-mental in the promulgation of a suprem-acist racist doctrine, even eugenics. Kali-kow (1980) argued that Lorenz was greatlyinfluenced in his youth by the world viewof Haeckel, and that that perspective madehim receptive to Nazi philosophy; the con-

nection to Haeckel, however, was tenuousat best (Richards, 1987).

Long after the stinging critique by Lehr-man, Kalikow (1980) documented thatLorenz joined the Nazi party eight weeksafter Anschluss with Austria (Kalikow,1980). It is nonetheless unclear whetherLorenz was opportunistic or simply accom-modated to a difficult situation. As Rich-ards (1987) pointed out, Lorenz's views onthe application of the "laws" of nature tohuman society, and on the degenerativeeffects of modern society on humans, werenot unique to Germany.

"At this point in Lorenz's career, certainwell-entrenched evolutionary ideas hap-pened to intersect with despicable Nazidogma. Certainly he fostered the unionof biology and propaganda, but I doubtthat his main concerns would have beenmarkedly different had the WeimarRepublic survived. Nonetheless, just thissort of public association of Nazism withhuman evolutionary biopsychology frozeany enthusiasm for the discipline (ofethology) immediately after the war, andcontinues to chill its development withincontemporary biology of behavior as wellas within the social sciences" (Richards,1987, p. 536).

For the rest of his life Lorenz was trou-bled by his experiences in the Third Reichand turned away from political issues. Butas a measure of how he was accepted afterWW II, the friendly collaboration betweenLorenz and Tinbergen, started in 1936,resumed. Bear in mind that Tinbergen wasimprisoned by the Germans for protestingagainst the dismissal of three Jewish facultymembers (Dewsbury, 1990).

The biting nature of Lehrman's critiquemust therefore be taken in the context ofa recent war in which genocide was directedat Jews and others and was said to be jus-tified in Haeckelian biopolitics. Lorenz wasperceived as a vestige of that tradition. Theoriginal, submitted version of Lehrman'scritique was scathing and personal; fortu-nately, the editor insisted the article berecast and focussed on the scientific issues(Eckhard Hess, personal communication).

The content of the critique reflected the

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thinking of Kuo and the Schneirla schooland was fairly representative of compara-tive psychology in North America at thetime. Lehrman (1953) objected to the useof terms such as innate and instinctive toexplain behavior. So naming behavior, heargued, substituted for an explanation ofhow behavior works. Lorenz was also saidto be insensitive to the interactive natureof development and to its underlying pro-cesses in producing the behavioral phe-notype, as reflected in the use by etholo-gists of maturation as opposed to learning.Lehrman faulted the ethologists for whathe felt was pseudo-physiological explana-tions. Finally, Lehrman took exception towhat he perceived as careless drawing ofparallels between the behavior of animalsand humans.

The criticism was accurate with regardto the dichotomization into innate andlearned behavior but was nonetheless anoverstatement because Lorenz wroteabundantly about ontogeny and imprint-ing. Yet it was true that ethologists wereinsufficiently concerned at that time withhow behavior was produced throughdevelopment. And they had been carelessin a number of ways.

The dispute was polarized and exagger-ated into the environmentalist doctrine ofthe American psychologists versus thegenetic determinism of the Europeanethologists. Nonetheless, Lehrman's cri-tique had the desired effect of bringing thetwo camps together. Much of the ensuingdebate took place at small meetings ratherthan in the literature, as in the first Etho-logical Conference in Freiburg in 1953.But passions ran high for several years,especially in the interactions with the Ger-man ethologists. For instance, in 1959 inCambridge, Lehrman met one evening ina small room to discuss some of the issueswith a group of German ethologists thatincluded Lorenz and von Hoist. At onepoint, von Hoist referred to Lehrman's1953 article as the Vergewaltigung (rape)of ethology (Lehrman, who understoodsome German, asked with a disarming smilefor the meaning of the word and chuckledwhen told).

Despite such heated conversations, the

two sides had been moving toward under-standing one another, especially in Britain.Changes were taking place there that facil-itated resolution of the controversy(Thorpe, 1979; Durant, 1986). With DavidLack's support, Oxford University hiredNiko Tinbergen. Shortly thereafter,Oxford and Cambridge, as is their way,divided up the world of ethology. Oxfordwas to pursue behavior in the context ofevolution and field biology. Cambridge,under the guidance of William Thorpe andlater Robert Hinde, was to concentrate onmechanisms of behavior.

Right from the beginning, the Englishethologists took to heart Lehrman's cri-tique. Hinde and Lehrman developed aclose friendship that modified the way eachviewed animal behavior. And Tinbergenand Lehrman progressed to a warm andcordial relationship. One of Lehrman's lastessays (1971), shortly before his untimelydeath, reveals how far he traveled towardthe perspective of ethologists.

The culmination of the rapprochementwas the appearance of Hinde's (1966, 1970)tome subtitled, A Synthesis of Ethology andComparative Psychology. The book effec-tively demonstrated the intricacies ofbehavioral explanations, particularly thedelicate interplay of genome and environ-ment that results in the behavioral phe-notype. Lorenz (e.g., 1965) softened hisposition, now writing of the genome andenvironment as sources of informationduring ontogeny. But in many respectsLorenz seemed to be merely attempting tomake his original ideas more palatablewithout making fundamental changes(Lehrman, 1970). Nonetheless, ethologygradually entered a less turbulent periodcharacterized by more refined analyses.

For many years after the peak of thecontroversy, ethology was often used in thepejorative sense. Oddly, as the term becameless frequently heard among biologists itbecame increasingly accepted in allied areassuch as medicine (e.g., McGuire and Fair-banks, 1977) and political science (e.g.,Schubert, 1975).

The controversy had been intense but inpublications, at least, done withoutimpugning one another's motives. The

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issues were those of science, and the debateaccelerated understanding. Further, start-ing in the late fifties, many universitiesacross North America created new posi-tions in animal behavior.

THE ADVENT OF SOCIOBIOLOGY

Meanwhile at Oxford, Tinbergen'sgroup had started asking how specificbehavioral differences affect the survival,hence fitness, of individuals (e.g., Tinber-gen et at., 1962; Patterson, 1965). JohnCrook (1965), under David Lack's super-vision, showed how avian mating systemsand displays ultimately derive from feed-ing habits, predation, and physical envi-ronment. Other major contributors wereW. D. Hamilton (e.g., 1964) on kinship andaltruism, and John Maynard Smith (e.g.,1972) on the theoretical substrates of socialbehavior. In the Netherlands, Lukas Tin-bergen, Niko's brother, helped set the stagefor optimal foraging in his pioneering studyof search image (L. Tinbergen, 1960).

Things were also stirring in NorthAmerica. The Scottish biologist Wynne-Edwards' (1962) advocacy of group selec-tion stimulated George Williams (1966) towrite his classic "Adaptation and NaturalSelection"; he was between jobs in Berke-ley and had time to write (personal com-munication). Williams' book was foreshad-owed by a seldom cited article (Williamsand Williams, 1957) on the evolution ofaltruism among social insects. Alexander(1977) credited Williams for triggering arevolution in our thinking about evolution.Alexander himself was contributingimportantly, especially on the costs andbenefits of social aggregations (Alexander,1974). Three pivotal articles by Trivers(1971, 1972, 1974) finished setting thestage for the climax by E. O. Wilson (1975),Sociobiology. The New Synthesis.

THE STORM SURROUNDINGSOCIOBIOLOGY

Wilson's book was acclaimed by manybiologists. It cut a scholarly swath throughanimal behavior, ecology, population biol-ogy and evolution. Although Wilsontouched on a wide range of topics, the unit-ing theme was the modulation of social

behavior by kinship and its consequences,as articulated by Hamilton (1964), Wil-liams (1966) and Trivers (1971).

"Sociobiology" also evoked vigorous butconstructive criticism (e.g., reviews in Ani-mal Behaviour, 1976). One group of sci-entists, however, ripped the book as a polit-ical tract. Those attacks were unequalledin recent times in vituperation and seemedaimed at Wilson as much as at the contentof the book, obliquely questioning hismotives. The most extreme assaults werewritten by fellow academicians, some ofwhom professed a fondness for Wilson(Wade, 1976). The group behaved con-trary to our expectations of academiciansas responsible critics by flagrantly distort-ing Wilson's position (Wade, 1976).

During that period, a network of activistscientists triggered graduate seminars inuniversities across North America. Oddly,the participants were seldom those whostudied behavior (see below); typically, theywere molecular biologists, geneticists andthe likes. One spin-off was a group thatcalled itself something like the CommitteeAgainst Racism; in 1979 they mounted thespeakers' platform at a symposium onsociobiology in Washington, D.C., andphysically assaulted E. O. Wilson. Prior toand during that symposium other anti-sociobiologists tried unsuccessfully to takeover and subvert the symposium to theirown agenda.

But why should "Sociobiology" havedrawn such organized, passionate and sus-tained attack at the biopolitical level? Jer-ram Brown's (1975) textbook coveredmuch the same material, interpreted sim-ilarly, and appeared the same year as Wil-son's book. Brown, however, did notextrapolate to human society. Brown wasalso more sensitive to the issue of behav-ioral processes and development. The yearbefore "Sociobiology" came out, MichaelGhiselin (1974) published a book thataddressed much the same issues in whatmight be considered an even more pro-vocative manner with regard to humans;other than some uncharitable reviews,nothing more followed.

The strong response to Wilson's bookwas said to have two roots, possibly three.

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The first and most frequently mentionedreason was the content of Wilson's lastchapter. There he speculated incautiouslyon the genetic basis of human social behav-ior, and often with regard to highly com-plex, situation-sensitive behavior. That wasthe target of most of the blistering reviews(see below). Unfortunately, Wilson's lastchapter resulted in the nature-nurturedebate regressing to a polarized state suchas prevailed when Lehrman (1953) tookthe ethologists to task.

The second reason for the negativeresponse is found in the first chapter. ThereWilson presented the infamous "dumb-bell" model of the future. The two largespheres represented population biology onthe one side and physiology on the other.The thin connecting link was all that wasto remain of animal behavior in the con-ventional sense. Further, sociobiology (thepopulation level) would replace the socialsciences, such as anthropology. Thatshocked those in the fields ordained to dis-appear, and it galvanized them into action(e.g., Sahlins, 1976; Washburn, 1978).

The last reason was inferred by Wade(1976), and the evidence is at best circum-stantial; it might be called local competi-tion. First a little background. Science forthe People, consisting of academicians andother scientists, championed progressiveto radical social issues at that time. Fromthem came the Sociobiology Study Group.The most active segment of that group wasat Harvard University, Wilson's homeinstitution. Most notably, three of them,Steven J. Gould, Richard Levins and Rich-ard Lewontin, were in Wilson's adminis-trative unit and interacted with him.

The first critique was the most scathing.Allen et al. (1975, in Wilson, 1976) statedthat sociobiology in an earlier form pro-vided "an important basis" for sterilizationlaws and restrictive immigration policiesduring the period between 1910 and 1930,and also "for the eugenics policies whichled to the establishment of gas chambersin Nazi Germany. . . . Wilson joins the longparade of biological determinists whosework has served to buttress the institutionsof our society by exonerating them fromresponsibility for social problems." The

issue of responsibility was echoed in suc-ceeding critiques in a form that appearedat times to advocate prior censorship (e.g.,Kitcher, 1985).

Later, the Sociobiology Study Group(1976) characterized genetic determinismas follows: Present societies are a conse-quence of a combination of inherent prop-erties of humans and of "biological forces."The status quo is justified by the homeo-static nature of societies; attempts to alterthe situation are fruitless because societywill return to its natural arrangement.Genetic determinism is merely an out-growth of the author's particular socioeco-nomic prejudices.

In a number of essays, sociobiologists ingeneral were accused of being geneticdeterminists but, except for Wilson, with-out naming them (e.g., Gould, 1978).Dawkins (1985), taking note of the anon-ymous nature of the repeated accusation,flatly asserted that no legitimate sociobiol-ogist advocates genetic determinism oranything approaching it. Probably true, butthe views of David Barash and Pierre vanden Berghe came close (Kitcher, 1985).Wilson is on record as rejecting geneticdeterminism. He declared that the envi-ronment is more important than geneticsto human behavior. "When any genetic biasis demonstrated, it cannot be used to justifya continuing practice in present and futuresocieties . . . it would invite disaster" (Wil-son, 1976).

One of the curious things about theattack on Wilson and "Sociobiology" wasthe absence, among the outspoken critics,of any authoritative student of animalbehavior. This sharp contrast to the ethol-ogy controversy has gone without com-ment. I was therefore intrigued, during thatturbulent period, when individuals fromthe Sociobiology Study Group directed myattention to one example, a paper by aneminent student of animal behavior, PeterKlopfer (1977). The brief abstract of thatpaper stated that, "Sociobiology has madea resurgence in recent years, but hasbecome enmeshed in political controversy.Indeed, much of the work on sociobiologyhas been used to justify repressive or racistmeasures." To my surprise, the so-called

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sociobiologist he attacked was KonradLorenz, who abhorred sociobiology. Klop-fer referred to Wilson's book in but onesentence: "The recent monumental studyby E. O. Wilson, through the impetus ithas given biosociology, no less through thecontroversy it has spawned, makes this aparticularly topical issue" (emphasis added).Otherwise, Klopfer said only a few cau-tiously approving words about the writingsof sociobiologists sensu stricto.

The main thrust of the attack on "Socio-biology" by the Sociobiology Study Groupwas that Wilson was politically motivated,even if he were himself unaware of it. Iwould be remiss, therefore, if I did notmention the political views of the leadingscientists of the Study Group. It is not outof the ordinary for scientists to distort viewsof the opposing camp (Hull, 1980). It isunusual, however, for the distorting groupsto have such openly political aims.

Richard Levins and Richard Lewontinhave professed to being Marxists (Lums-den and Wilson, 1981a; Joravsky, 1970;Wade, 1976), and Steven J. Gould has pub-licly proclaimed that he learned Marxismon his father's knee. (I am compelled toadd that their personal politics are of noconcern to me.) Lumsden and Wilson(198 la) provided the following quotes fromThe Radicalization of Science by Lewontin andLevins: "As working scientists in the fieldof evolutionary genetics and ecology, wehave been attempting with some success toguide our own research by a consciousapplication of Marxist philosophy. . . .There is nothing in Marx, Lenin or Maothat is or that can be in contradiction withthe particular physical facts and processesof a particular set of phenomena in theobjective world."

This type of thinking led Lewontin(1977) to advocate stopping all studies onIQ, Allen et al. (1975) to question whetherthe genetics of human behavior in any formis warranted, and Gould (1978) to assertthat human nature is simply too complexand difficult to understand, so any attemptto do so is bound to fail. Kitcher (1985, p.9) seemed to oppose even hypothesizingabout human behavior. I understand andappreciate their concerns, given the sexist

attitudes and racial discrimination we allsee. Still, their injunctions are a chillingreminder of Lysenko's damaging proscrip-tion of all studies of genetics in the SovietUnion (see Caspari and Marshak, 1965;Joravsky, 1970).,

To return to the starting point, thenature-nurture issue, I quote a passage fromthe Sociobiology Study Group (1976) thatillustrates their environmentalistic view ofthe dichotomy: ". . . the truth is that theindividual's social activity is to be under-stood only by first understanding socialinstitutions" (emphasis added). Extremeenvironmentalism can result in a doctrineas dangerous and loathsome as geneticdeterminism, as the Minister of Propa-ganda in the Third Reich well appreciated.

Anti-sociobiology scientists obviouslyhad, and have, a sophisticated understand-ing of the contributions of environmentand genome to the development of thephenotype (Kitcher, 1985). Yet theyseemed to move in a different world wherebehavior was concerned, one blind to thatunderstanding. On different occasionsshortly after the publication of "Socio-biology" Richard Lewontin and LawrenceSlobodkin gave lectures in Berkeley inwhich they bluntly asserted that animalbehavior, unlike other biological traits, hasno genetic basis.

In the space of a few years after "Socio-biology" much of the heat went out of thedebate. Workers started calling themselvesbehavioral ecologists instead of sociobiol-ogists. Wilson wrote further books in whichhe took greater pains to make explicit hisviews on human behavior (Wilson, 1978;Lumsden and Wilson, 19816). Had thesebooks appeared without the prior publi-cation of "Sociobiology" they still wouldhave received hostile reviews, but I believethe debate would have been on a higher,more objective plane. Problems remained,however. Wilson carefully explained thecontributions of both genetic and environ-mental substrates, and their intricate inter-actions. But he then proceeded to discussspecific aspects of human behavior in aselectionist framework that requiresgenetic control (Kitcher, 1985).

Critical reviews and commentaries con-

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tinued to appear, though for the most partthey became progressively more cautiousand constructive (e.g., Gould, 1978, 1980;but see Lumsden and Wilson, 1981a).Recently, Kitcher (1985) stressed the needto apply a higher standard of evidence whenwriting about human behavior because ofthe potential for misuse; in his view, a lowerstandard has prevailed in the publicationsof sociobiologists.

Kitcher's injunction can be applied toboth sides in the argument, not just to thesociobiologists. Kitcher (1985:292) illus-trated this point when he accused socio-biologists of the " . . . parading of gossip asobservational data," but then did the sameto support his own arguments. The worstexample of this double standard came whenhe sought to refute the notion of unitarydrive by describing a lascivious banquetfrom the cinema "Tom Jones," but manyother lapses occurred. Kitcher also faultedsociobiologists for not entertaining alter-native hypotheses. Yet, when he claimedthat sociobiological conclusions abouthuman behavior merely coincided with folkpsychology, he never considered the myr-iad of alternative interpretations of humanbehavior available in folk psychology.

Science for the People repeatedly heldup the claim that sociobiologists were irre-sponsible (see also Kitcher, 1985). On theirside, their attack so exceeded the usualbounds of scientific debate that they wereopen to the charge of irresponsibility. Thedebate did advance our appreciation ofmany sensitive issues, and the complexinterrelationship of science and politics;that must be counted as a plus. On theother hand, their attack contributed littleto clarifying purely scientific issues. Theirmain accomplishment seems to have beenthe creation of an image of sociobiology asan evil doctrine, an image that extendedto those studying animal behavior. Thatimage cost the field, and precipitated ascramble to call oneself a behavioral ecol-ogist.

During and after the sociobiology tumultI traveled to a number of meetings anduniversities. Old colleagues, not reading theprimary literature nor working the field ofbehavior, responded to the mention of

sociobiology as anathema. When askedabout the workings of the field, they hadno understanding whatsoever. They knewit had to do with animal behavior, and thatit was bad, it was sexist and racist or some-thing like that, but little more. I believethe aftermath, from which we still suffer,is that such faculty often argued againsthiring in the area of animal behavior. Atthe heart of all this is the nature-nurtureissue.

The issue will not die. It is all too easyto demonstrate the effects of environment,often reversible, on the behavior of indi-viduals. It is difficult, but not impossible,to tease out the contribution of the geneticsubstrate in animal behavior. Where theproblem becomes acute is when the exten-sion is made to humans, with implicationsfor ethical, hence political, behavior.Genetics of human behavior is pretty muchlimited to family histories and adoptionstudies. The argument will continue to bepropelled by our growing understandingof the evolution of behavior among ani-mals, the perceived continuity between thebehavior of animals and humans, and thetemptation to resort to correlations.

ACKNOWLEDGMENTS

I am grateful to Donald A. Dewsbury,Stephen E. Glickman and Judy Stamps fortheir constructive comments on an earlydraft of this article, and to Colin G. Beer,Patricia A. Gowaty and an anonymousreviewer for invaluable criticism of thepenultimate draft. Not only did theirremarks improve the manuscript, they ledme to a deeper exploration of the issues. Ialso thank Zuleyma Halpin for inviting meto present this essay.

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