Though the Price equation in itself is simply a statistical identity, biologists have often adopted a ‘causal interpretation’ of the equation, in the sense that its component terms have been supposed to correspond to distinct causal processes in evolution, such as natural selection and transmission bias. In this paper, we bring the issue of causal interpretation to the fore, by studying the conditions under which it is legitimate to read causal meaning into the Price equation. We argue that only if substantive assumptions about causal structure are made, which can be represented in the form of a causal model, can the component terms of the Price equation be interpreted as causally meaningful. We conclude with a reflection on the epistemic uses of the Price equation, emphasizing the difference between the description, explanation and prediction of evolutionary change. This article is part of the theme issue ‘Fifty years of the Price equation’.

We critically examine Denis Walsh's latest attack on the causalist view of fitness. Relying on Judea Pearl's Sure-Thing Principle and geneticist John Gillespie's model for fitness, Walsh has argued that the causal interpretation of fitness results in a reductio. We show that his conclusion only follows from misuse of the models, that is, (1) the disregard of the real biological bearing of the population-size parameter in Gillespie's model and (2) the confusion of the distinction between ordinary probability and Pearl's causal probability. Properly understood, the models used by Walsh do not threaten the causalist view of fitness.

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The causal nature of evolution is one of the central topics in the philosophy of biology. It has been discussed whether equations used in evolutionary genetics point to some causal processes or are purely phenomenological patterns. To address this question the present paper builds well-defined causal models that underlie standard equations in evolutionary genetics. These models are based on minimal and biologically-plausible hypotheses about selection and reproduction, and generate statistics to predict evolutionary changes. The causal reconstruction of the evolutionary principles shows adaptive evolution as a genuine causal process, where fitness and selection are both causes of evolution.

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