在達爾文的天擇演化論得到認同後,動物行為就理所當然被視為天擇演化的產物。達爾文本人即有提到族群選擇的概念,認為這是利他行為的演化基礎[2];康拉德·洛倫茲在其書中也用「對物種整體有利」之類的詞語來解釋動物行為[1][3],但沒有真的去實際量測各種行為對適應度的影響。至1962年,動物學家V·C·韋恩-愛德華茲(V. C. Wynne-Edwards)觀察到大多數動物分佈在一個個小族群中,因此認定動物行為影響的是整個族群的生存和繁殖,並主張動物行為主要是源自作用在族群層級的天擇演化[4]。但J·M·史密斯(J. M. Smith)和G·C·威廉斯(G. C. Williams)等演化生物學隨即家用數學模型證明了生物不會為了族群而犧牲個體的適應度進行利他行為,因此主張天擇主要運作在個體上,只有在少數如蜜蜂和螞蟻等真社會性的動物中才會因為親屬選擇而出現利他行為[5][6][7]。
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^Richard Dawkins, "Replicators and Vehicles (页面存档备份,存于互联网档案馆)", King's College Sociobiology Group, eds., Current Problems in Sociobiology, Cambridge, Cambridge University Press, (1982), pp. 45-64