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翻訳と辞書　辞書検索 [ 開発暫定版 ] スポンサード リンク inclusive fitness ： ウィキペディア英語版 inclusive fitness In evolutionary biology inclusive fitness theory is a model for the evolution of social behaviors (traits), first set forward by W. D. Hamilton in 1963 and 1964. Instead of a trait's frequency increase being thought of only via its average effects on an organism's ''direct reproduction'', Hamilton argued that its average effects on ''indirect reproduction'', via identical copies of the trait in other individuals, also need to be taken into account. Hamilton's theory, alongside reciprocal altruism, is considered one of the two primary mechanisms for the evolution of social behaviors in natural species. From the gene's point of view, evolutionary success ultimately depends on leaving behind the maximum number of copies of itself in the population. Until 1964, it was generally believed that genes only achieved this by causing the individual to leave the maximum number of viable direct offspring. However, in 1964 W. D. Hamilton showed mathematically that, because other members of a population may share identical genes, a gene can also increase its evolutionary success by indirectly promoting the reproduction and survival of such individuals. The most obvious category of such individuals is close genetic relatives, and where these are concerned, the application of inclusive fitness theory is often more straightforwardly treated via the narrower kin selection theory. Belding's ground squirrel provides an example. The ground squirrel gives an alarm call to warn its local group of the presence of a predator. By emitting the alarm, it gives its own location away, putting itself in more danger. In the process, however, the squirrel may protect its relatives within the local group (along with the rest of the group). Therefore, if the effect of the trait influencing the alarm call typically protects the other squirrels in the immediate area, it will lead to the passing on of more of copies of the alarm call trait in the next generation than the squirrel could leave by reproducing on its own. In such a case natural selection will increase the trait that influences giving the alarm call, provided that a sufficient fraction of the shared genes include the gene(s) predisposing to the alarm call.〔Mateo JM, 1996. (The development of alarm-call response behavior in free-living juvenile Belding's ground squirrels ). Animal Behaviour 52:489-505.〕 ''Synalpheus regalis'', a eusocial shrimp, also is an example of an organism whose social traits meet the inclusive fitness criterion. The larger defenders protect the young juveniles in the colony from outsiders. By ensuring the young's survival, the genes will continue to be passed on to future generations.〔 〕 Inclusive fitness is more generalized than strict kin selection, which requires that the shared genes are ''identical by descent''. Inclusive fitness is not limited to cases where "kin" ('close genetic relatives') are involved. == Hamilton's rule == In the context of sociobiology, which holds that some behavior can be influenced by genes and therefore can evolve by natural selection, Hamilton proposed that inclusive fitness offers a mechanism for the evolution of altruism. He claimed that this leads natural selection to favor organisms that behave in ways that correlate with maximizing their inclusive fitness. Hamilton's rule describes mathematically whether or not a gene for altruistic behavior will spread in a population: : where * $r\; \backslash$ is the probability, above the population average, of the individuals sharing an altruistic gene – commonly viewed as "degree of relatedness". * $b\; \backslash$ is the reproductive benefit to the recipient of the altruistic behavior, and * $c\; \backslash$ is the reproductive cost to the altruist, In a recent paper, Gardner ''et al.''(2007) suggest that Hamilton's rule can be applied to multi-locus models, but that it should be done at the point of interpreting theory, rather than the starting point of enquiry.〔Gardner A, West SA and Barton NH. The relation between multilocus population genetics and social evolution theory. ''Am Nat'' 169, 207–226.〕 They suggest that one should "use standard population genetics, game theory, or other methodologies to derive a condition for when the social trait of interest is favored by selection and then use Hamilton's rule as an aid for conceptualizing this result". A recent paper by Nowak et al. (2010) suggested that standard natural selection theory is superior to inclusive fitness theory, stating that the interactions between cost and benefit can not be explained only in terms of relatedness. This, Nowak said, makes Hamilton's rule at worst superfluous and at best ad hoc.〔Martin A. Nowak, Corina E. Tarnita & Edward O. Wilson The evolution of eusociality ''Nature'' 466 1057–1062〕 Gardner in turn was critical of the paper, describing it as "a really terrible article", and along with other co-authors has written a reply, submitted to ''Nature''.〔http://www.nytimes.com/2010/08/31/science/31social.html?pagewanted=1&_r=1〕 In work prior to Nowak, various authors derived different versions of a formula for $r$, all designed to preserve Hamilton's rule.〔Orlove, M. J. 1975 A Model of Kin Selection not Invoking Coefficients of Relationship J. Theor. Biol. v49 pp289-310〕〔Orlove, M. J. & Wood, C. L. 1978. "Coefficients of relationship and coefficients of relatedness in kin selection: A covariance form for the RHO formula". Journal of Theoretical Biology, Volume 73, Issue 4, 21 August 1978, Pages 679-686〕〔Michod, R. E. & Hamilton, W. D. 1980. "Coefficients of relatedness in sociobiology" Nature 288, 694 - 697 (18 December 1980)〕 Orlove noted that if a formula for $r$ is defined so as to ensure that Hamilton's rule is preserved, then the approach is by definition ad hoc. However, he published an unrelated derivation of the same formula for $r$ – a derivation designed to preserve two statements about the rate of selection – which on its own was similarly ad hoc. Orlove argued that the existence of two unrelated derivations of the formula for $r$ reduces or eliminates the ad hoc nature of the formula, and of inclusive fitness theory as well.〔Orlove, M. J. 1979 A Reconciliation of Inclusive Fitness and Personal Fitness Approaches: a Proposed Correcting Term for the Inclusive Fitness Formula, J. Theor. Biol. v81 pp 577–586〕 The derivations were demonstrated to be unrelated by corresponding parts of the two identical formulae for $r$ being derived from the genotypes of different individuals. The parts that were derived from the genotypes of different individuals were terms to the right of the minus sign in the covariances in the two versions of the formula for $r$. By contrast, the terms left of the minus sign in both derivations come from the same source. One study suggest the c/b ratio be considered as a continuum of this behavioral trait rather than discontinuous in nature. From this approach fitness transactions can be better observed because there is more to what is happening to affect an individual's fitness than just losing and gaining (Engles, W.R. 1982). 抄文引用元・出典: フリー百科事典『 ウィキペディア（Wikipedia）』 ■ウィキペディアで「inclusive fitness」の詳細全文を読む スポンサード リンク 翻訳と辞書 : 翻訳のためのインターネットリソース Copyright(C) kotoba.ne.jp 1997-2016. All Rights Reserved.