British evolutionary biologist who derived the mathematical theory of kin selection — Hamilton's rule — solving the problem of altruism that had troubled Darwin, and who was arguably the greatest evolutionary biologist since Darwin.
Hamilton's rule (rb > c): altruism evolves when the relatedness-weighted benefit to the recipient exceeds the cost to the actor. Inclusive fitness: an organism's fitness includes its effect on the fitness of relatives. Kin selection: the mechanism by which altruism evolves. This solved Darwin's problem of sterile worker insects and altruism generally. Evolutionarily Stable Strategy (with Maynard Smith): game theory applied to evolution. The evolution of sex (the red queen).
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