W. D. Hamilton

William Donald Hamilton (1936–2000) was a British evolutionary biologist who developed inclusive fitness theory and explained how kin selection can favor altruistic behavior.

Inclusive fitness: An individual's genetic success through its own reproduction and effects on the reproduction of related individuals. Hamilton’s central framework counts effects on relatives alongside an organism’s own offspring.
Altruism (biology): A behavior that increases another individual’s reproductive success while reducing the actor’s. Hamilton explained how such costly behavior can still increase an actor’s inclusive fitness.
Eusociality: A social organization marked by cooperative brood care, overlapping generations, and reproductive division of labor. Relatedness-based explanations of reproductive helping made eusociality a central test case.
George R. Price: An American population geneticist whose equation describes changes in trait frequencies through selection and transmission. Price’s mathematical work clarified links between Hamilton’s social-selection ideas and evolutionary change.
The Genetical Evolution of Social Behaviour: Hamilton’s 1964 paper presenting a general genetic account of social behavior and inclusive fitness. This landmark paper established the theory that became most closely associated with Hamilton.
Kin selection: Natural selection favoring traits that increase the reproductive success of genetic relatives. Hamilton used this process to explain how altruism can evolve among relatives.
Direct fitness: An individual's contribution to future generations through its own reproduction. It is the reproductive route Hamilton’s framework combines with effects on relatives.
Haplodiploidy: A sex-determination system in which males develop from unfertilized eggs and females from fertilized eggs. Its unusual relatedness patterns prompted debate about its role in the evolution of social insects.
John Maynard Smith: A British evolutionary biologist who applied game theory to evolution and helped shape modern evolutionary biology. He engaged with Hamilton’s work and developed complementary models of social behavior.
The Selfish Gene: Richard Dawkins’s 1976 book presenting evolution from the perspective of genes and their effects on organisms. Dawkins popularized gene-centered interpretations built partly on Hamilton’s theoretical foundations.

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