Singular effect of linkage on long-term genetic gain in Fisher's infinitesimal model

Elise Tourrette, Olivier C Martin
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Abstract

During the founding of the field of quantitative genetics, Fisher formulated in 1918 his “infinitesimal model” that provided a novel mathematical framework to describe the Mendelian transmission of quantitative traits. If the infinitely many genes in that model are assumed to segregate independently during reproduction, corresponding to having no linkage, directional selection asymptotically leads to a constant genetic gain at each generation. In reality, genes are subject to strong linkage because they lie on chromosomes and thus segregate in a correlated way. Various approximations have been used in the past to study that more realistic case of the infinitesimal model with the expectation that the asymptotic gain per generation is modestly decreased. To treat this system even in the strong linkage limit, we take the genes to lie on continuous chromosomes. Surprisingly, the consequences of genetic linkage are in fact rather singular, changing the nature of the long-term gain per generation: the asymptotic gain vanishes rather than being simply decreased. Nevertheless, the per-generation gain tends to zero sufficiently slowly for the total gain, accumulated over generations, to be unbounded.
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费雪无穷小模型中连锁对长期遗传增益的奇异效应
在数量遗传学领域的创立过程中,费雪于 1918 年提出了 "无限小模型",为描述数量性状的孟德尔传递提供了一个新颖的数学框架。如果假定该模型中的无限多基因在繁殖过程中独立分离,相当于没有连锁,那么定向选择就会近似地导致每一代的遗传增益不变。在现实中,由于基因位于染色体上,因此会以相关的方式发生分离,因此基因会有很强的连锁性。过去曾使用过各种近似方法来研究无穷小模型中更现实的情况,期望每一代的渐进增益会适度降低。即使在强联系极限下,为了处理这个系统,我们也要把基因放在连续的染色体上。令人惊讶的是,遗传连锁的后果实际上相当奇特,它改变了每代长期收益的性质:渐近收益消失了,而不仅仅是减少了。尽管如此,每代收益趋向于零的速度非常缓慢,因此世代累积的总收益是没有界限的。
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