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“隔离机制的发展”(以及其特殊形式“强化”)存在什么问题?

[What's wrong with "development of isolating mechanisms" (and its special case in form of "reinforcement")?].

作者信息

Fridman V S

出版信息

Zh Obshch Biol. 2014 Jul-Aug;75(4):261-86.

Abstract

Examined are the conceptual grounds of the "reinforcement" model that play an important role in explanatory schemes of the theory of evolution. According to the current views, reinforcement (of obstacles for crossing under natural selection when there occurs a repeated contact with hybridization between two substantially divergent forms, up to the level where started hybridization is terminated and the forms become unable to cross) should be a common or even frequent phenomenon. In fact, reinforcement turns out to be a rare event needed to be specially searched for, all such events are dubious, etc. Here, empirical data on reinforcement pro and contra are analyzed, using mostly ornithological (and some other zoological) materials. It is shown that in most cases where the theory predicts reinforcement to occur, it does not happen. On the contrary, something quite opposite is observed--selection for non-crossing enhancement does not stop up the leaks in gene pools of the forms that are due to their hybridization within the zone of a repeated contact, but instead the stable hybrid zone is established which serve as a conduit for gene exchange between the forms. Through hybridization, those forms can adopt strange genes, remaining, at that, their isolation at the level of separate species. This is achieved through the system of self-non-self recognition in population system of both forms being in contact. Within the zone of a repeated contact, conditions are usually such that individuals repeatedly make mistakes forming mixed pairs and producing hybrids, i.e., at the level of individuals, self-non-self recognition is often inefficient, especially at the range edges, in zone of repeated contact and such. Efficient recognition, that ensures species separateness even under conditions of continuing hybridization, takes place upon inclusion of individuals of different origin--phenotypically 'pure' ones, hybrids, and backcrosses--into spatio-ethologic population structure of both forms, upon acquisition of territories by established groups, their inclusion in flocks and other intra-population associations of resident individuals. At that, groups of both forms predominantly incorporate their 'own' ('pure') phenotypes and reject 'strange' ones, including hybrids of intermediate phenotype. If the forms have attained the species level of separateness, hybrids do not form their own stable groups, but try to individually inset themselves into the habitation of both parents, which turns out to be not so successful as compared with phenotypically 'pure' individuals of the given form. Thereby, hybrids happen to be directly excluded from the population system of both forms even if hybridization goes on and produces them in sufficient quantity while hybrids and backcrosses themselves are not inferior to 'pure' individuals either in survivorship or reproductive success (those ones who became residents have occupied a territory and are reproducing). It is shown that usual examples of reinforcement are not only sparse but could be explained differently. Particularly, 'splitting up' of a population system which gives rise to two forms of different ecology with following strengthening/enhancement of the divergence through sexual selection is better explained by our model of self-non-self recognition than by reinforcement. Relying on this concept, the conclusion is drawn that the model of non-crossing 'reinforcement' in zones of a repeated contact with hybridization is not supported by the data, as well as the model of character displacement that follows from it. Apparently, the reasons behind its popularity consist in conformity with the 'ideology' of a more general theory that is agreed upon by most researchers, and corresponds, in whole, to known facts (in this case, the theory of modern evolutionary synthesis).

摘要

本文探讨了“强化”模型的概念基础,该模型在进化理论的解释框架中起着重要作用。根据当前观点,强化(当两个差异较大的形态之间反复发生杂交接触时,自然选择会增加杂交的障碍,直至杂交停止,两个形态无法再杂交)应该是一种常见甚至频繁出现的现象。但实际上,强化是一种罕见的事件,需要专门去寻找,而且所有这类事件都存在疑问等。在此,我们主要利用鸟类学(以及其他一些动物学)资料,分析了支持和反对强化的实证数据。结果表明,在大多数理论预测会发生强化的情况下,强化并未出现。相反,观察到的情况恰恰相反——选择增强杂交障碍并没有阻止由于在反复接触区域内杂交而导致的形态基因库中的基因渗漏,反而形成了稳定的杂交区域,成为形态之间基因交换的通道。通过杂交,这些形态可以获得奇特的基因,同时在物种层面保持隔离。这是通过两个接触形态的种群系统中的自我 - 非自我识别系统实现的。在反复接触区域内,通常情况下个体在形成混合配对和产生杂种时会反复犯错,即在个体层面,自我 - 非自我识别往往效率低下,尤其是在分布范围边缘、反复接触区域等地方。当不同来源的个体——表型上“纯”的个体、杂种和回交个体——被纳入两个形态的空间 - 行为种群结构中,当已建立的群体获得领地、被纳入鸟群以及其他常驻个体的种群内关联时,就会发生高效的识别,这种识别即使在持续杂交的条件下也能确保物种的分离。此时,两个形态的群体主要纳入它们“自己的”(“纯”的)表型个体,并排斥“陌生的”个体,包括中间表型的杂种。如果这些形态已经达到物种层面的分离,杂种不会形成自己稳定的群体,而是试图个体融入双亲的栖息地,但与给定形态的表型“纯”个体相比,这并不那么成功。因此,即使杂交持续进行并产生大量杂种,杂种也会直接被排除在两个形态的种群系统之外,而杂种和回交个体在生存能力或繁殖成功率方面并不比“纯”个体差(那些成为常驻个体的已经占据了领地并正在繁殖)。研究表明,通常的强化例子不仅稀少,而且可以有不同的解释。特别是,一个种群系统“分裂”产生两种不同生态形态,随后通过性选择加强/增强分化,用我们的自我 - 非自我识别模型比用强化来解释更好。基于这一概念,得出的结论是,在与杂交的反复接触区域中,非杂交“强化”模型以及由此得出的特征取代模型都没有得到数据支持。显然,其广受欢迎的原因在于与大多数研究人员认同的更一般理论的“意识形态”相符,并且总体上与已知事实(在这种情况下是现代进化综合理论)一致。

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