HAMILTON'S RULE

HAMILTON, W. D.

The Genetical Evolution of Social Behaviour, I (+) The Genetical Evolution of Social Behaviour, II.

London, Academic Press, 1964. 8vo. In the original grey printed wrappers. In "Journal of Theoretical Biology", Volume 7, Number 1, July 1964. Entire issue offered. A very fine and clean copy. Pp. 1-16; Pp. 17-52. [Entire volume: 170, (2) pp.].

First printing of Hamilton's two seminal publications, perhaps the most important in evolutionary biology in the 20th century, on altruism in relation to kin selection. Hamilton is, primarily because of the present publication, widely regarded as being one of the most influential theoretical biologists of the twentieth century. "Hamilton's principal achievement was so thoroughly to revise the language of evolutionary biology that it has become nearly impossible to speak in evolutionary explanations except in terms of the self-interest of the organism or gene." (DSB)

Hamilton's rule: k> 1/r, a gene causing an organism to benefit relatives at the expense of its own reproduction will be selected and increase in a population if the benefit to the "altruist" outweighs the discounted relationship, or as Hamilton himself described it: "a gene causing altruistic behavior towards brothers and sisters will be selected only if the behavior and the circumstances are generally such that the gain is more than twice the loss; for half-brothers it must be more than four times the loss; and so on. To put the matter more vividly, an animal acting on this principle would sacrifice its life if it could thereby save more than two brothers, but not for less." (DSB).

Due to the complexity and advanced mathematics the paper was rejected twice until it was accepted by the reviewer's an it was not until the mid 1970ies that his theory became widely know and cited: "Hamilton wrote up the theory of inclusive fitness in two versions. One was a lengthy, fully mathematical treatment that unified understanding of a considerable body of case studies of altruistic behaviors that Hamilton drew from the scientific literature, the fruit of his graduate research. The second was a short, mostly verbal abstract of the whole, containing only the mathematical relation of Hamilton's rule and some general, theoretical remarks on its applicability. He met difficulty in publishing both. The first he submitted to the Journal of Theoretical Biology, where it spent considerable time in the reviewing process; ultimately the referee (John Maynard Smith, a mathematical biologist of similar interests) asked that it be split into two parts.
After the revisions and splitting called for by the referee for the Journal of Theoretical Biology, that journal published "The Genetical Evolution of Social Behaviour," parts 1 and 2, in 1964. The first part of the paper contained the mathematical arguments culminating in the derivation of Hamilton's rule; its arguments were almost exclusively cast in the language and methodology of modern population genetics. The second part hearkened back in its methodology to Darwin's, as Hamilton used the theory of inclusive fitness to explain a diverse array of social traits recorded in the biological literature, including alarm calling, mutual grooming, the fusion of colony organisms, and postreproductive behavior in cryptic (camouflaged) moth species compared with that of aposematic species (bad-tasting with vivid warning colors). In each case, Hamilton argued that his theory of inclusive fitness could coherently explain the evolution of phenomena that had been disparate in the literature as aspects of a single principle at work, Fisher's fundamental theorem of natural selection, mandating the maximization of favorable genes under selection.

Hamilton's influence began to grow among evolutionary biologists as the few who had read and understood the import of his papers worked to bring him from his initial scientific and social isolation into the networks of scientists interested in evolution and behavior. Wilson, for example, invited Hamilton to lecture at Harvard University in 1969, en route to a Smithsonian Institution conference on "Man and Beast" that brought together specialists from various fields to discuss the impact of recent biological work on understandings of human nature

"From about 1974, citations of Hamilton's 1964 papers in the scientific literature began an exponential rise, reaching some four thousand total in the Institute for Scientific Information (ISI) Web of Science database by 2007, making "The Genetical Evolution of Social Behaviour" the most-cited paper ever published in the Journal of Theoretical Biology. Hamilton's principal achievement was so thoroughly to revise the language of evolutionary biology that it has become nearly impossible to speak in evolutionary explanations except in terms of the self-interest of the organism or gene." (DSB)

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