Sociobiology: The New Synthesis
Since the discovery of the nature and mode of operation of the basic units of heredity, biologists in search of major new fields to conquer have increasingly turned their attention to what has conventionally been referred to as behavior, mentality, and society. Three of the most important students of animal behavior, Karl von Frisch, Konrad Lorenz, and Niko Tinbergen, who recently shared a Nobel Prize, have all recently published books about their work; but the most sustained attempt to synthesize the whole of our present-day knowledge is undoubtedly that of Professor Edward O. Wilson of Harvard. His book is a formidable work. If I had to find a single word to describe my feelings when I found on my desk this offering from the NYR—no fewer than 392 broad-columned galley proofs, with a promise of more to come, containing glossary, bibliography, and index, plus, as a small side dish, a work of a mere 200 pages of sophisticated neurobiology—I think the best I could do would be “dudgeon”—not high dudgeon, but certainly low dudgeon.
A first glance was not very appetizing. The first sentence of Sociobiology quotes Camus (“Camus said that the only serious philosophical question is suicide”), only to have it pointed out in the second that there Camus was—not unusually in my opinion—talking Gallic rhetoric through his hat (“That is wrong even in the strict sense intended”), and in the third sentence we find ourselves involved with the hypothalamus and the limbic system of the brain. H’m…. So I turned to the very last sentence, at the bottom of galley 392. Only to find Camus again. This time propounding one of his tautological aphorisms: “In a universe divested of illusions and lights, man feels an alien, a stranger.” Well, in such a universe this may be so. This time Wilson remarks, “This, unfortunately, is true. But we still have another 100 years.” So I applied what capacity for perseverance I possess to trying to discover what the intermediate 390 galleys told us to do in this 100 years.
It was quickly apparent that there is, as might be expected from Professor Wilson, a great deal of solid and interesting material behind the window dressing. He is well known as one of the most profound professional students of animal societies, particularly those of insects. He very early makes it. clear that his book has a well-defined and important aim, and it is rather closely organized around this coherent theme. By “Sociobiology” he means the systematic study of the biological basis of all aspects of social behavior, particularly of animal societies, but also encompassing the social behavior of early man, and the organization of the more primitive contemporary human societies. He foregoes the attempt to deal with more complex forms of human social organization.
He argues that societies have been produced by evolution, that is to say by the natural selection of genetic variants, and that the proper way to understand societies is to discover how natural selection could have brought them into being. He reminds us that biologists who study the evolution of anatomical and physiological characteristics have claimed to have achieved in recent years a New Synthesis, which provides a satisfactory explanation for that entire class of phenomena. Wilson sets himself nothing less than the task of producing an equally satisfying new synthesis about the evolution of the social behavior of animals.
He sees this synthesis as eventually having three major components. At what one might call the most dispersed level would be a theory of the integrated functioning of the cells of the brain (“integrative neurophysiology”), which would itself be related to general cellular biology. This would give rise to a second component, consisting of ethology and physiological psychology. Those fields in turn would form a bridge to the third major component, namely sociobiology and behavior related to the ecological situation of the population of organisms, which in its turn would be linked with general population biology.
This is clearly an extraordinarily ambitious aim. I think one should say immediately that Professor Wilson has been astonishingly successful in achieving it. This book will undoubtedly be for many years to come a major source of information about all aspects of our knowledge of social behavior in animals, from the most primitive types such as corals, through insects, fishes, birds, to the many varieties of mammals and primitive man. It has also some of the clearest discussions yet written of recent advances in general population biology and demography. Many large drawings illustrate the major aspects of social behavior in various groups of animals with exemplary clarity. Unfortunately, I feel in these drawings an exceptional lack of any aesthetic appeal whatever, but perhaps that is not important to their purpose.
There is no point in repeating that this is a very good and very important book. It is more interesting to get down to particular features in which it seems to me something less than perfect.
At the very beginning, when he is still warming up for what he intends to be a long game, Wilson defines “the central theoretical problem of sociobiology: how can altruism, which by definition reduces personal fitness, possibly evolve by natural selection?” This problem was raised, and the essential solution of it actually provided, more than forty years ago in one of the books which founded the theory usually called neo-Darwinism, in which evolution is explained in terms of genes. In The Causes of Evolution, published in 1932, J. B. S. Haldane wrote (p. 131): “Insofar as it makes for the survival of one’s descendants and near relations, altruistic behavior is a kind of Darwinian fitness, and may be expected to spread as a result of natural selection.”
More recently this subject became, for some rather mysterious reason, a fashionable topic for a rather foolish controversy. The argument was represented as turning on whether one cohesive and delimited population, whose members showed some degree of altruistic behavior among themselves, would in competition entirely overcome and eliminate some other population whose members were not so well disposed to their fellows. One has only to look at the results of the arrival of the white man in Tasmania, or in many parts of the United States, to realize that such differential population extinctions can occur. However, many people have argued that in theory they would happen only in rather exceptional circumstances.
In a very clear exposition, Wilson shows, first, that these doubts about the superior competitiveness of cohesive and internally altruistic populations are theoretically well founded, but he then has the common sense to demonstrate that the basic theory does not depend on population extinction at all. It only requires that the altruistic behavior should lead to a greater increase in numbers of individuals bearing hereditary factors tending in that direction than of other individuals not similarly genetically endowed. He shows that certain very plausible relations between populations would bring about such situations, and also that a similar differential multiplication might be produced by certain types of competition between individuals, rather than between groups.
This is an argument that has been advanced recently particularly by the English biologist Maynard Smith. I am personally not very convinced that anyone has yet provided a good explanation of how a tendency to altruistic behavior first takes hold in a population, but there certainly are, as Wilson so clearly expounds, several ways in which it might be expected to spread once it got started.
All this is an example of the admirable clarity of Wilson’s exposition. The point that remains most obscure is just why he should consider the existence of altruism to be the central theoretical problem of sociobiology. But he does not allow this point about altruism to limit his discussion. He puts under review most of the problems which have recently appealed to students of population biology (except for the peculiar phenomenon of protein polymorphism, which has interested many people, precisely for the paradoxical reason that it seems to have no interesting consequences for the effectiveness of the organisms, either as individuals or as social creatures).
Wilson’s discussion of modern selection theory and population dynamics is again a model of clarity. In my opinion he is somewhat too impressed by the distinction that has been made between r-selection and K-selection. These names are derived from coefficients that occur in certain algebraic theories about population growth recently developed by American authors. Many biologists on the other side of the Atlantic feel that the theories are rather schematic, and never fully apply to the complicated situations that arise in the actual ecological situations of nature. Most creatures are probably being subjected both to r-selection and K-selection, though maybe at different stages in their life histories.
In his discussion of modern evolutionary population biology, Wilson is broad-minded enough to bring out the fact, which most orthodox evolutionists try to keep in decent obscurity, that selection operates directly on phenotypes, that is to say on organisms after they have been affected during their development by the environment as well as by the genes they contain. Important effects on the fitness of an organism, and thus on the results of selection, may in some cases be brought about by the influence of stressful environments. It has been found experimentally that if individuals are selected over many generations for their ability to respond successfully to such environments, in the end the modifications may be exhibited even when the environment becomes no longer stressful. This is the process known as genetic assimilation. Wilson admits that “because behavior, and especially social behavior, has the greatest developmental plasticity of any category of phenotypic traits, it is also theoretically the most subject to evolution by genetic assimilation.”
However, this is in an early introductory part of his book, and when in the main text he gets down to cases the importance of phenotypic adaptation and genetic assimilation hardly gets a mention. Yet “phenotypic adaptation” is only another way of referring to learning, by a phrase broad enough to encompass modification of physical structure as well as of behavior. If one is going to discuss the evolution of social behavior with the aim of bringing it into connection with human social behavior, which is almost wholly learned, then “learning,” and the genetic basis for learning, must have an absolutely central position in the argument. I do not think that Wilson goes far enough to give it the importance it deserves.
This may be partly a defect of one of his great merits. Wilson is perhaps the leading authority in the world on the social life of insects, and insects are notoriously unteachable creatures. Most of them communicate with one another by primarily chemical means, though the bees may have gone slightly further by developing a system of bodily gestures. Neither mode would seem to have the potentiality for anything remotely resembling the semantic richness of the communication systems of apes, or even mammals, let alone the fantastic capacities of human language. It is, I think, a symptom of a certain insensitivity in Wilson to the importance of communication, particularly as a mode of teaching and learning, when he refers to Margaret Mead in this connection only as having employed the word “semiotic” to designate the analysis of communication in the broadest sense. He makes no mention of the fact that she has devoted a large and very illuminating book (Continuities in Cultural Evolution, Yale University Press, 1964) precisely to that analysis, covering many forms of transmission of social organization which do not depend on verbal language.
The whole topic of communication among individuals and all the many forms in which it may be carried out seems to me a much more central issue in sociobiology than the problem of altruism, which Wilson singles out as the dominant theme. One can, I think, conceive of a society in which no individual shows any altruism toward any other: anthropologists such as Fortune on the Mundugumor have described human societies that come close to this. But to conceive of a society in which there is no communication among individuals would seem to be a contradiction in terms.
Another field which Wilson explores less deeply than one might hope is the reciprocal interaction between behavior and selection. To a major extent, animals lose out in selection, not so much for doing the wrong thing, but for their inefficiency in performing whatever it is they are doing. The nature of the selection is to a very large degree dictated by the nature of their behavior. The accounts of social behavior in animals which Wilson provides very often imply this, but he does not, so far as I can discover, explicitly state it.
That brings one to what is, to my mind, the weakest feature in the whole grand structure which Wilson has built up. Is it not surprising that in a book of 700 large pages about social behavior there is no explicit mention whatever of mentality? In the index, covering more than thirty pages of three columns each, there is no mention of mind, mentality, purpose, goal, aim, or any word of similar connotation. Of course, something very similar to mind or purpose is very often implied. In fact, in searching the index for the latter word, the nearest you get to it is the entry “pursuit, invitation,” which refers to the peculiar behavior of certain types of gazelle, apparently “for the purpose” of luring a predator into giving its presence away by starting a pursuit which the gazelle can dispose of by escaping.
Or consider the following account on pages 122 and 123:
A distraction display is any distinctive behavior used to attract the attention of an enemy and to draw it away from an object that the animal is trying to protect. In the great majority of instances the display directs a predator away from the eggs or young. Bird species belonging to many different families have evolved their own particular bag of tricks…. New Zealand pied stilts are among the great actors of the animal world. Guthrie-Smith (1925) has described their response to intrusion in the vicinity of the nest as follows:
“Dancing, prancing, galumphing over one spot of ground, the stricken bird seems simultaneously to jerk both legs and wings, as strange toy beasts can be agitated by elastic wires, the extreme length of the bird’s legs producing extraordinary effects. It gradually becomes less and less able to maintain an upright attitude. Lassitude, fatigue, weariness, faintings—lackadaisical and fine ladyish—supervene. The end comes slowly, surely, a miserable flurry and scraping, the dying Stilt, however, even in articulo mortis, contriving to avoid inconvenient stones and to select a pleasant sandy spot upon which decently to expire. When on some shingle bank well removed from eggs and nests half a dozen Stilts—for they often die in companies—go through their performances, agonizing and fainting, the sight is quaint indeed.”
Can there be any point in renouncing the use of words like mind, or goal-seeking, in the face of phenomena of this kind? Is not Wilson guilty of at the very least a certain incoherence, in indulging himself in a delicious mental frisson of Camusian alienation, while denying the poor New Zealand Stilt, working away so very hard to confuse the sense of reality, even a hint of mentality? I think one is bound to come to the conclusion that the socio-biologists are just “running scared” of ferocious philosophers. A few years ago it may have been tactically wise for quiet behavioral scientists to practice their own distraction procedures against the threat of predatory positivists, but I doubt if there is any longer any need for such super-caution.
Perhaps one can get some insight into what is bringing it about by a glance at the introductory portions of the book by Laughlin and d’Aquili. Most of their discussion involves more technical detail of the anatomical structures and functional intrarelations of parts of the brain than is likely to appeal to those who are not specialists. This is a topic which Wilson suggested would eventually form one of the two major branches of a well-developed sociobiology. But so far we have extremely little understanding of it, and that of a very technical nature, and Wilson does not attempt to go into it in any depth in the main body of his text. However, in their first chapter Laughlin and d’Aquili have sufficient consideration for their readers to explain what they mean by their somewhat daunting title Biogenetic Structuralism. They expound the meaning of structuralism in the words of an article originally published in the Scientific American in 1972 by the molecular biologist Gunther Stent:
It is only in the past 20 years or so, more or less contemporaneously with the growth of molecular biology, that a resolution of the age-old epistemological conflict of materialism vs. idealism was found in the form of what has come to be known as structuralism.
…Both materialism and idealism take it for granted that all the information gathered by our senses actually reaches our mind; materialism envisions that thanks to this information reality is mirrored in the mind, whereas idealism envisions that thanks to this information reality is constructed by the mind. Structuralism, on the other hand, has provided the insight that knowledge about the world enters the mind not as raw data but in some already highly abstracted form, namely as structures. In the preconscious process of converting the primary data of our experience step by step into structure, information is necessarily lost, because the creation of structures, or the recognition of patterns, is nothing less than the selective destruction of information.
Thus since the mind does not gain access to the full set of data about the world, it can neither mirror nor construct reality. Instead for the mind reality is a set of structural transforms of primary data taken from the world. This transformation process is hierarchical, in that “stronger” structures are formed from “weaker” structures through selective destruction of information. Any set of primary data becomes meaningful only after a series of such operations has so transformed it that it has become congruent with a stronger structure preexisting in the mind. Neurophysiological studies carried out in recent years on the process of visual perception in higher mammals have not only shown directly that the brain operates according to the tenets of structuralism but also offer an easily understood illustration of those tenets.
Laughlin and d’Aquili first make the point that structuralism is not really related to molecular biology: in fact for them “it has been the worldwide interest in the work of Lévi-Strauss, and later Noam Chomsky, that seems to have made structuralism a watchword for many.” In fact, I think this bit of dialogue signifies no more than the passage of generations. Gunther Stent began to develop into a mature scientist at the time of the flowering of molecular biology, and it was then that he came across philosophical views called structuralism. Laughlin and d’Aquili belong to a more recent generation, and they came across them first at the time when Lévi-Strauss and Noam Chomsky held the limelight. I belong to an even older generation than Gunther Stent’s, and like him I first met these ideas at the time when I was becoming a mature scientist; but that was in the late 1920s; and I met them not in connection with any particular kind of science, but straight from the mouth of a philosopher, Alfred North Whitehead.
Whitehead used another terminology, almost as unexpected as that of structuralism, but he meant essentially the same thing. Recent authors use the word “structure,” which to me suggests the steel frame of a building or something similar, as a technical word to refer to the form in which “knowledge about the world enters the mind.” Whitehead used the word “object,” which might seem to suggest an external independent thing, like a jug or a table, to refer to the same sort of entity as a “structure”; namely, a something—a concept if you like—which is formed in the mind as a result of its interaction with nature.
Whitehead defined several different classes of “objects,” but one of these was “scientific objects,” and that comprised all the entities used in scientific theorizing, such as atoms, cells, nerves, and so on. Many of these have been conventionally regarded, under the influence of simple materialist philosophy, as “things” existing quite independently of people or mind. Whitehead argued that we can, of course, experience the world, but we cannot know anything until it has entered the mind and become what he called an object and what the structuralists call a structure.
Among these objects there are hierarchies, as Stent points out, but these are connected only with the inclusiveness of the object or structure, and not with its reality. This is the point which Laughlin and d’Aquili do not seem to have grasped. A few pages after quoting Stent’s clear account of the essential meaning of structuralism, they retreat back to a more old-fashioned kind of realist philosophy, and on that basis reject mind as a possible object or structure. “The major ontological assumption,” they state, “upon which biogenetic structuralism is founded is that there exists no reality intervening between the central nervous system and the environment” (their italics). This is to forget the basic insight of structuralism, namely that both the “central nervous system” and the “environment” are themselves not “reality,” but are structures or, as Whitehead would say, scientific objects. The concept of “mind” also would be a scientific object. It is probably a tricky one, particularly because it is difficult to keep it apart from the very difficult notion of consciousness or self-awareness; but if mind is defined primarily in terms of goal-seeking behavior, it is a scientific object which is almost universally present by implication in all discussions of animal behavior.
I personally think it preferable to have the question of mind out in the daylight and put under inspection, rather than to try to hide it away like a guilty secret. That is why, in the final analysis, I think the most profound discussions of the evolution of behavior, both individual and social, are those which begin from an undismayed recognition of the total nature of human experience; such as, for instance, Susanne Langer’s book Mind: A Study in Human Feeling, a work which rather astonishingly is never referred to even in Wilson’s very extensive bibliography.
Against “Sociobiology” November 13, 1975