进化理论对我们对世界的理解和管理产生了巨大的影响,部分原因是它能够成功地整合新数据和新见解。尽管如此,目前,某些生物学现象与主流进化论之间存在紧张关系。例如,分子表观遗传变化的遗传如何适应主流进化理论?生态位构建是一个进化过程吗?通过栖息地选择的局部适应也是适应性进化吗?这些例子表明有范围(甚至可能是需要)扩大我们对进化的看法。我们确定了三个方面,这些方面纳入单一框架将使理解和研究适应性进化成为一种更普遍的方法:(i)扩展表型的扩展观点;(ii)性状可以相互响应;(iii)遗传可以是非遗传的。我们使用因果模型将这三个方面与驱动和允许自适应进化的变量和机制的既定观点相结合。我们的因果模型将适应性父母反应的自然选择和非遗传遗传鉴定为适应性进化的两个互补但不同和独立的驱动因素。这两个驱动因素都符合价格方程;具体来说,父母反应的非遗传遗传被Price方程中经常被忽视的部分捕获。我们的因果模型是通用的和简化的,但可以在变量和因果联系方面灵活调整,取决于研究问题和/或生物系统。通过回顾上面给出的三个例子,我们展示了如何使用它作为一个启发式工具来澄清概念问题,并帮助设计实证研究。与仅从遗传变化角度定义进化的基因中心观点相反,我们的广义方法使我们能够将进化视为整个因果结构的变化,不仅包括遗传,还包括表型和环境变量。
Evolutionary theory has made large impacts on our understanding and management of the world, in part because it has been able to incorporate new data and new insights successfully. Nonetheless, there is currently a tension between certain biological phenomena and mainstream evolutionary theory. For example, how does the inheritance of molecular epigenetic changes fit into mainstream evolutionary theory? Is niche construction an evolutionary process? Is local adaptation via habitat choice also adaptive evolution? These examples suggest there is scope (and perhaps even a need) to broaden our views on evolution. We identify three aspects whose incorporation into a single framework would enable a more generalised approach to the understanding and
study of adaptive evolution: (i) a broadened view of extended phenotypes; (ii) that traits can respond to each other; and (iii) that inheritance can be non-genetic. We use causal modelling to integrate these three aspects with established views on the variables and mechanisms that drive and allow for adaptive evolution. Our causal model identifies natural selection and non-genetic inheritance of adaptive parental responses as two complementary yet distinct and independent drivers of adaptive evolution. Both drivers are compatible with the Price equation; specifically, non-genetic inheritance of parental responses is captured by an often-neglected component of the Price equation. Our causal model is general and simplified, but can be adjusted flexibly in terms of variables and causal connections, depending on the research question and/or biological system. By revisiting the three examples given above, we show how to use it as a heuristic tool to clarify conceptual issues and to help design empirical research. In contrast to a gene-centric view defining evolution only in terms of genetic change, our generalised approach allows us to see evolution as a change in the whole causal structure, consisting not just of genetic but also of phenotypic and environmental variables.