Hamilton's rule

Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even at a cost to the organism's own survival and reproduction. Kin altruism is altruistic behaviour whose evolution is driven by kin selection. Kin selection is an instance of inclusive fitness, which combines the number of offspring produced with the number an individual can produce by supporting others, such as siblings.

Charles Darwin discussed the concept of kin selection in his 1859 book, The Origin of Species, where he reflected on the puzzle of sterile social insects, such as honey bees, which leave reproduction to their mothers, arguing that a selection benefit to related organisms (the same "stock") would allow the evolution of a trait that confers the benefit but destroys an individual at the same time. R.A. Fisher in 1930 and J.B.S. Haldane in 1932 set out the mathematics of kin selection, with Haldane famously joking that he would willingly die for two brothers or eight cousins. In 1964, W.D. Hamilton popularised the concept and the major advance in the mathematical treatment of the phenomenon by George R. Price which has become known as "Hamilton's rule". In the same year John Maynard Smith used the actual term kin selection for the first time.

According to Hamilton's rule, kin selection causes genes to increase in frequency when the genetic relatedness of a recipient to an actor multiplied by the benefit to the recipient is greater than the reproductive cost to the actor. The rule is difficult to test but studies of red squirrels appear to confirm it. Hamilton proposed two mechanisms for kin selection: kin recognition, where individuals are able to identify their relatives, and viscous populations, where dispersal is rare enough for populations to be closely related. The viscous population mechanism makes kin selection and social cooperation possible in the absence of kin recognition. Nurture kinship, the treatment of individuals as kin when they live together, is sufficient for kin selection, given reasonable assumptions about dispersal rates. Kin selection is not the same thing as group selection, where natural selection acts on the group as a whole.

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