Kin Selection: Its Components

Abstract
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 negativethat is, individual selection operates against altruismand [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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