Trivers R L, Hare H
Science. 1976 Jan 23;191(4224):249-63. doi: 10.1126/science.1108197.
Halminton (1) was apparently the first to appreciate that the synthesis of Mendelian genetics with Darwin's theory of natural selection had profound implications for social theory. In particular, insofar as almost all social behavior is either selfish or altruistic (or has such effects), genetical reasoning suggests that an individual's social behavior should be adjusted to his or her degree of relatedness, r, to all individuals affected by the behavior. We call this theory kinship theory. The social insects provide a critical test of Hamilton's kinship theory. When such theory is combined with the sex ratio theory of Fisher (9), a body of consistent predictions emerges regarding the haplodiploid Hymenoptera. The evolution of female workers helping their mother reproduce is more likely in the Hymenoptera than in diploid groups, provided that such workers lay some of the male-producing eggs or bias the ratio of investment toward reproductive females. Once eusocial colonies appear, certain biases by sex in these colonies are expected to evolve. In general, but especially in eusocial ants, the ratio of investment should be biased in favor of females, and in it is expected to equilibrate at 1 : 3 (male to female). We present evidence from 20 species that the ratio of investment in monogynous ants is, indeed, about 1 : 3, and we subject this discovery to a series of tests. As expected, the slave-making ants produce a ratio of investment of 1 : 1, polygynoys ants produce many more males than expected on the basis of relative dry weight alone, solitary bees and wasps produce a ratio of investment near 1 : 1 (and no greater than 1 : 2), and the social bumblebees produce ratios of investment between 1 : 1 and 1 : 3. In addition, sex ratios in monogynous ants and in trapnested wasps are, as predicted by Fisher, inversely related to the relative cost in these species of producing a male instead of a female. Taken together, these data provide quantitative evidence in support of kinship theory, sex ratio theory, the assumption that the offspring is capable of acting counter to its parents' best interests, and the supposition that haplodiploidy has played a unique role in the evolution of the social insects. Finally, we outline a theory for the evolution of worker-queen conflict, a theory which explains the queen's advantage in competition over male-producing workers and the workers' advantage regarding the ratio of investment. The theory uses the asymmetries of haplodiploidy to explain how the evolved outcome of parent-offspring conflict in the social Hymenoptera is expected to be a function of certain social and life history parameters.
哈明顿（1）显然是第一个认识到孟德尔遗传学与达尔文自然选择理论的结合对社会理论具有深远影响的人。特别是，由于几乎所有的社会行为要么是自私的，要么是利他的（或有这样的影响），遗传学推理表明，个体的社会行为应该根据其与受该行为影响的所有个体的亲缘关系程度r进行调整。我们将这一理论称为亲缘关系理论。群居昆虫为汉密尔顿的亲缘关系理论提供了关键检验。当这一理论与费希尔（9）的性别比例理论相结合时，就会出现一系列关于单倍二倍体膜翅目的一致预测。在膜翅目中，雌性工蜂帮助其母亲繁殖的进化比在二倍体群体中更有可能发生，前提是这些工蜂产下一些产生雄性的卵，或者将投资比例偏向于繁殖雌性。一旦出现了真社会性群体，这些群体中按性别划分的某些偏差就有望进化。一般来说，尤其是在真社会性蚂蚁中，投资比例应该偏向雌性，预计会在1:3（雄性与雌性）达到平衡。我们提供了来自20个物种的证据，表明单蚁后的蚂蚁的投资比例确实约为1:3，并且我们对这一发现进行了一系列测试。正如预期的那样，奴役蚁产生的投资比例为1:1，多蚁后的蚂蚁产生的雄性比仅根据相对干重预期的要多得多，独居蜜蜂和黄蜂产生的投资比例接近1:1（且不大于1:2），而社会性大黄蜂产生的投资比例在1:1和1:3之间。此外，正如费希尔所预测的，单蚁后的蚂蚁和筑巢黄蜂中的性别比例与这些物种中产生雄性而非雌性的相对成本呈负相关。综合起来，这些数据提供了定量证据，支持亲缘关系理论、性别比例理论、后代能够违背其父母最佳利益行事的假设，以及单倍二倍体在群居昆虫进化中发挥了独特作用的假设。最后，我们概述了一个关于工蜂与蚁后冲突进化的理论，该理论解释了蚁后在与产生雄性的工蜂竞争中的优势以及工蜂在投资比例方面的优势。该理论利用单倍二倍体的不对称性来解释群居膜翅目中亲子冲突的进化结果如何预期是某些社会和生活史参数的函数。
