Change in gene frequency under kin selection is the sum of two components, namely, [See equation in the PDF file], a change in gene frequency caused by individual selection, and [See equation in the PDF file], a change caused by group selection. For the evolution of altruistic traits by kin selection, [See equation in the PDF file] is always negative-that is, individual selection operates against altruism-and [See equation in the PDF file] is always positive, so that selection between groups favors altruism. Hamilton's rule specifies the conditions under which [See equation in the PDF file]-that is, the conditions necessary for intergroup selection to override individual selection.
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Michael J. Wade (1980) studied this question.
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