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演化作为因果推断的过程

Evolution as a Process of Causal Inference

Department of Epidemiology and Data Science(流行病学与数据科学系);Fondazione IRCCS Istituto Nazionale dei Tumori di Milano(米兰国家肿瘤研究所IRCCS基金会)

Jacopo Iacovacci

AI总结:

本文提出自然选择应被理解为因果推断过程,利用Neyman-Rubin潜在结果框架形式化突变作为自然实验,并证明平均适应度的代际变化可分解为选择项和突变项。

摘要:

最近,复制子方程到贝叶斯定理的映射已被认识,导致了演化动力学与贝叶斯学习之间的类比。然而,这种类比仅适用于无限种群中的纯选择,当引入突变——演化的核心机制——时则失效。这里我提出,自然选择下的演化,至少在静态环境中的单倍体复制子种群中,最好不被理解为学习过程,而是因果推断过程。每个突变事件构成一个自然实验,其中亲本作为对照,突变后代作为处理单元。自然选择筛选突变对适应度的因果效应,保留非负效应的突变。我在Neyman-Rubin潜在结果框架内形式化了这一观点。我首先使用通用适应度结果发展了一般理论,并展示了因果推断中的核心识别假设(稳定单位处理值假设、一致性、无混杂性、积极性)如何映射到演化生物学。利用非归一化的准种方程,我证明了平均适应度的代际变化精确分解为一个选择项——恢复了费舍尔基本定理——加上一个突变项,该突变项对应于所有亲本基因型上所有突变的累积效应的适应度加权平均值。我展示了在适当假设下,这种分解扩展到广义复制子-突变子方程,并且匹配的亲本-后代群体的频率根据突变对适应度的平均因果效应成比例更新。

英文摘要:

Recently, the mapping of the replicator equation onto Bayes' theorem has been recognised, leading to an analogy between evolutionary dynamics and Bayesian learning. However, this analogy holds only for pure selection in infinite populations and breaks down when mutations -- a central mechanism of evolution -- are introduced. Here I propose that evolution by natural selection, at least for populations of haploid replicators in static environments, is best understood not as a learning process but as a process of causal inference. Each mutation event constitutes a natural experiment in which the parent serves as the control and the mutant offspring as the treated unit. Natural selection screens the causal effect of the mutation on fitness, retaining mutations with non-negative effects. I formalise this view within the Neyman-Rubin potential-outcomes framework. I first develop the general theory using a generic fitness outcome and show how the core identification assumptions in causal inference (Stable Unit Treatment Value Assumption, Consistency, Unconfoundedness, Positivity) map onto evolutionary biology. Using the unnormalised quasispecies equation, I prove that the intergenerational change in mean fitness decomposes exactly into a selection term -- recovering Fisher's Fundamental Theorem -- plus a mutation term that corresponds to a fitness-weighted average of the cumulated effect of all mutations over all parental genotypes. I show that this decomposition extends, under suitable assumptions, to the generalised replicator-mutator equation and that the frequencies of populations of matched parents-offspring update in proportion to the average causal effect of mutations on fitness.

机构、总结、中文摘要均由AI分析而来,可能存在错误,请以论文正文为准。

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