The Unselected Hypothesis for the Evolution of Helping Behavior: Too Much or too Little Emphasis on Natural Selection?

The Unselected Hypothesis for the Evolution of Helping Behavior: Too Much or too Little Emphasis on Natural Selection?
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DOI:
10.1086/285217
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发表时间:
1991-07
期刊:
The American Naturalist
影响因子:
--
通讯作者:
I. Jamieson
I. Jamieson
中科院分区:
其他
文献类型:
--
作者:
I. Jamieson

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贾米森和克雷格(1987)对被广泛引用的适应论假说提出了批评,这些假说认为,在共同/合作繁殖的鸟类中,帮助筑巢是进化的结果。批评部分集中在鸟类供给反应的控制和发展上,并认为辅助非繁殖者喂养幼崽的行为与父母喂养后代的行为几乎没有区别。这导致了这样的建议,即由辅助设备(即,帮助者)可能会保持在公共繁殖种群作为一个副产品的刺激-反应供应机制的演变,在鸟类中发现一般。当这一批评首次在第20届国际动物行为学会议(麦迪逊,威斯康星州,1987),随后的讨论清楚地表明,反对者认为该论点混淆了因果解释的近似/最终二分法。人们普遍认为,提供雏鸟的nonbreeding辅助控制下的相同的刺激-反应机制,参与父母的喂养。然而,有人认为,我们的讨论未能解决目前的适应性意义的帮助,处理其演变和维护的最终原因。为了澄清这种混乱,我对贾米森和克雷格(1987)提出的观点进行了更详细的讨论。这一次,我故意避免讨论供应行为的近端方面,而是集中在进化过程上(Jamieson 1989 a)。我再次提出,助人行为可能是父母关爱进化的副产品。它的表达在社区繁殖的背景下可能是不寻常的情况下,非分散的幼鸟获得乞讨雏鸟的结果。这是作为一个零假设进行测试的备择假设,选择直接作用于帮手,以自适应的方式修改基本的供应反应。这一条也遭到了反对。然而,这一次它不仅仅来自于像Emlen等人(1991)和Ligon和Stacey(1991)这样的适应主义者,他们声称进化论假说没有考虑一个重要的选择主义标准(即,当前的效用),但也来自“结构主义者”(白色等1991),他们认为结构主义假设过于强调选择。虽然我会强烈反对白色等人提出的“真正的替代方案”,在其他方面,他们的批评也颇受欢迎。他们的回答,与适应主义者的回答相比,
Jamieson and Craig (1987) presented a critique of the widely cited adaptationist hypotheses on the evolution of helping at the nest in communally/cooperatively breeding birds. The critique focused in part on the control and development of the provisioning response in birds and argued that the act of feeding young by auxiliary nonbreeders was virtually indistinguishable from the act of feeding offspring by parents. This led to the suggestion that the provisioning of nestlings by auxiliaries (i.e., helpers) might be maintained in communal breeding populations as a by-product of the evolution of the stimulus-response provisioning mechanism found in birds in general. When this critique was first presented at the 20th International Ethological Conference (Madison, Wis., 1987), the ensuing discussion indicated clearly that opponents felt the argument confused the proximate/ultimate dichotomy of causal explanations. It was generally agreed that the provisioning of nestlings by nonbreeding auxiliaries was under the control of the same stimulus-response mechanism that is involved in parental feedings. However, it was argued that our discussion failed to address the current adaptive significance of helping that deals with the ultimate reasons for its evolution and maintenance. In an attempt to clear up the confusion, I presented a more detailed discussion of the ideas presented in Jamieson and Craig (1987). This time, I deliberately avoided discussion of the proximate aspects of provisioning behavior and instead concentrated on evolutionary processes (Jamieson 1989a). Again, I suggested that helping behavior might be maintained as an unselected by-product of the evolution of parental care. Its expression in the context of communal breeding could be the result of the unusual circumstances in which nondispersing juvenile birds obtain access to begging nestlings. This was presented as a null hypothesis to be tested against the alternative hypothesis that selection has acted directly on helpers to modify the basic provisioning response in an adaptive manner. This article too has been met with opposition. However, this time it has not come only from adaptationists like Emlen et al. (1991) and Ligon and Stacey (1991), who claim the unselected hypothesis fails to consider an important selectionist criterion (i.e., current utility), but from "structuralists" as well (White et al. 1991), who argue that the unselected hypothesis places too mntlch emphasis on selection. Although I will argue strongly against the "genuine alternative" proposed by White et al., their criticisms are in other ways quite welcome. Their reply, when contrasted to that of the adaptationists, leaves the distinct impression