The Evolution of Social Behavior
Encyclopedia
The Genetical Evolution of Social Behaviour is a 1964 scientific paper by the British evolutionary biologist W.D. Hamilton in which he mathematically lays out the basis for inclusive fitness
Inclusive fitness
In evolutionary biology and evolutionary psychology, the inclusive fitness of an organism is the sum of its classical fitness and the number of equivalents of its own offspring it can add to the population by supporting others...

.

Hamilton, then only a PhD student, completed his work in London. It was based on Haldane
J. B. S. Haldane
John Burdon Sanderson Haldane FRS , known as Jack , was a British-born geneticist and evolutionary biologist. A staunch Marxist, he was critical of Britain's role in the Suez Crisis, and chose to leave Oxford and moved to India and became an Indian citizen...

's idea, but Hamilton showed that it applied to all gene frequencies. Although initially obscure, it is now highly cited in biology books, and has gone on to reach such common currency that citations are now often unnecessary as it is assumed that the reader is so familiar with kin selection and inclusive fitness that he need not use the reference to obtain further information.

The paper's peer review
Peer review
Peer review is a process of self-regulation by a profession or a process of evaluation involving qualified individuals within the relevant field. Peer review methods are employed to maintain standards, improve performance and provide credibility...

 process led to disharmony between one of the reviewers, John Maynard Smith
John Maynard Smith
John Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....

 and Hamilton. Hamilton thought that Maynard Smith had deliberately kept the paper, which has difficult mathematics, from publication so that Maynard Smith could claim credit for the concept of kin selection in his own paper. Indeed such was the time taken for peer review that Hamilton published a magazine essay in American Naturalist
American Naturalist
The American Naturalist is a monthly peer-reviewed scientific journal that was established in 1867. It is published by the University of Chicago Press on behalf of the American Society of Naturalists. The journal covers research in ecology, evolutionary biology, population, and integrative biology....

in 1963.

The American George R. Price
George R. Price
George Robert Price was an American population geneticist. Originally a physical chemist and later a science journalist, he moved to London in 1967, where he worked in theoretical biology at the Galton Laboratory, making three important contributions: first, rederiving W.D...

 found Hamilton's paper, and finding trouble in its implications for sociobiology
Sociobiology
Sociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology,...

, tried to disprove it but ended up rederiving his work through the Price equation
Price equation
The Price equation is a covariance equation which is a mathematical description of evolution and natural selection. The Price equation was derived by George R. Price, working in London to re-derive W.D. Hamilton's work on kin selection...

.

The paper has been reprinted in books twice, firstly in George C. Williams
George C. Williams
Professor George Christopher Williams was an American evolutionary biologist.Williams was a professor emeritus of biology at the State University of New York at Stony Brook. He was best known for his vigorous critique of group selection. The work of Williams in this area, along with W. D...

's Group Selection, and secondly in the first volume of Hamilton's collected papers Narrow Roads of Gene Land. The latter includes a background essay by Hamilton.

Hamilton had previously written a short note explaining the background in 1988 when ISI
Institute for Scientific Information
The Institute for Scientific Information was founded by Eugene Garfield in 1960. It was acquired by Thomson Scientific & Healthcare in 1992, became known as Thomson ISI and now is part of the Healthcare & Science business of the multi-billion dollar Thomson Reuters Corporation.ISI offered...

 recorded it as a citation classic.

See also

  • Group Selection
    Group Selection (book)
    Group Selection is a 1971 book edited by George C. Williams, containing papers written by biologists arguing against the view of group selection as a major force in evolution...

    (book by G. C. Williams which contains this paper)

External links

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