A fundamental task of evolutionary biology is to explain the pervasive impression of organismal design in nature, including traits benefiting kin. Inclusive fitness is considered by many to be a crucial piece in this puzzle, despite ongoing discussion about its scope and limitations. Here, we use individual-based simulations to study what quantity (if any) individual organisms become adapted to maximize when genetic architectures are more or less suitable for the presumed main driver of biological adaptation, namely cumulative multi-locus evolution. As an expository device, we focus on a hypothetical situation called Charlesworth's paradox, in which altruism is seemingly predicted to evolve, yet altruists immediately perish along with their altruistic genes. Our results support a recently proposed re-definition of inclusive fitness, which is concerned with the adaptive design of whole organisms as shaped by multi-locus evolution, rather than with selection for any focal gene. They also illustrate how our conceptual understanding of adaptation at the phenotypic level should inform our choice of genetic assumptions in abstract simplified models.

中文翻译：

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现实的遗传结构使有机体适应成为根据包容性适应的民间定义所预测的。

进化生物学的一项基本任务是解释自然界中有机体设计的普遍印象，包括有益于亲属的特征。尽管一直在讨论其范围和局限性，但许多人认为包容性健身是这个难题中的关键部分。在这里，我们使用基于个体的模拟来研究当遗传结构或多或少适合假定的生物适应的主要驱动因素时，个体生物体适应的数量（如果有的话）最大化，即累积多位点进化。作为一种说明手段，我们关注一种称为查尔斯沃思悖论的假设情况，其中似乎预测利他主义会发展，但利他主义者立即与他们的利他基因一起灭亡。我们的结果支持最近提出的对包容性健身的重新定义，它与由多位点进化形成的整个生物体的适应性设计有关，而不是与对任何焦点基因的选择有关。它们还说明了我们对表型水平适应性的概念理解应该如何为我们在抽象简化模型中选择遗传假设提供信息。