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:
影响因子:
--
通讯作者:
I. Jamieson
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文献类型:
--
作者:
I. Jamieson
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