The evolution of social behavior

For several years the study of social behavior has been undergoing a revolution with far-reaching consequences for the social and biological sciences. Partly responsible are three recent changes in the attitudes of evolutionary biologists. First was growing acceptance of the evidence that the potency of natural selection is overwhelmingly concentrated at levels no higher than that of the individual. Second was revival of the comparative method, especially as applied to behavior and life histories. Third was spread of the realization that not only are all aspects of structure and function of organisms to be understood solely as products of selection, but because of their peculiarly direct relationship to the forces of selection, behavior and life history phenomena, long neglected by the evolutionists, may be among the most predictable of all phenotypic attributes. These ideas have been appreciated by a few biologists for a long time, but they have only recently begun to characterize the science as a whole. Darwin’s discussion of sterility between species as an incidental effect of evolutionary adaptation (41, p. 260) and his refusal to deal with sex ratio selection (42, p. 399) suggest awareness of the difficult problem of determining the levels at which selection is most powerful. Yet significant clarification of this basic issue did not really commence until publication of Wynne-Edwards’ massive volume (179) championing group selection and inadvertently exposing its unlikelihood. As late as 1958, Fisher felt constrained to add to the revised edition of his 1929 classic, The Genetical Theory of Natural Selection, the admonishment (53, p. 49) that his fundamental theorem and its associated considerations, already misused then by decades of population geneticists dealing (as they saw it) with the fitness of populations, refer strictly to "the progressive modification of structure or function only in so far as variations in these are ofadvantage to the individual... [and afford] no corresponding explanation for any properties of animals and plants.., supposed to be of service to the species to which they belong." Williams’ critique (171) provided a significant turning point. Nevertheless, one has only to pick up any biological journal or attend any biological meeting to realize that this question has not yet been settled for all