Disentangling evolutionary, plastic and demographic processes underlying trait dynamics: a review of four frameworks

Biologists are increasingly interested in decomposing trait dynamics into underlying processes, such as evolution, plasticity and demography. Four important frameworks that allow for such a decomposition are the quantitative genetic animal model (AM), the Geber' method (GM), the age-structured Price equation(APE) and the integral projection model (IPM). However, as these frameworks have largely been developed independently, they differ in the assumptions they make, the data they require, as well as their outcomes and interpretation. Here, we evaluate how each framework decomposes trait dynamics into underlying processes. To do so, we apply them to simulated data for a hypothetical animal population. Individual body size was affected by, among others, genes, maternal effects and food intake. We simulated scenarios with and without selection on body size and with high and low heritability. The APE and IPM provided similar results, as did the AM and GM, with important differences between the former and the latter. All frameworks detected positive contributions of selection in the high but not in the low selection scenarios. However, only the AM and GM distinguished between the high and low heritability scenarios. Furthermore, the AM and GM revealed a high contribution of plasticity. The APE and IPM attributed most of the change in body size to ontogenetic growth and inheritance, where the latter captures the combined effects of plasticity, maternal effects and heritability. We show how these apparent discrepancies are mostly due to differences in aims and definitions. For example, the APE and IPM capture selection, whereas the AM and GM focus on the response to selection. Furthermore, the frameworks differ in the processes that are ascribed to plasticity and in how they take into account demography. We conclude that no single framework provides the true' contributions of evolution, plasticity and demography. Instead, different research questions require different frameworks. A thorough understanding of the different definitions of their components is necessary for selecting the most appropriate framework for the question at hand and for making biologically meaningful inferences. This work thus supports both future analysis and the careful interpretation of existing work.