How development affects evolution.

Natural selection acts on developmentally constructed phenotypes, but how does development affect evolution? This question prompts a simultaneous consideration of development and evolution. However, there has been a lack of general mathematical frameworks mechanistically integrating the two, which may have inhibited progress on the question. Here, we use a new mathematical framework that mechanistically integrates development into evolution to analyse how development affects evolution. We show that, while selection pushes genotypic and phenotypic evolution up the fitness landscape, development determines the admissible evolutionary pathway, such that evolutionary outcomes occur at path peaks rather than landscape peaks. Changes in development can generate path peaks, triggering genotypic or phenotypic diversification, even on constant, single-peak landscapes. Phenotypic plasticity, niche construction, extra-genetic inheritance, and developmental bias alter the evolutionary path and hence the outcome. Thus, extra-genetic inheritance can have permanent evolutionary effects by changing the developmental constraints, even if extra-genetically acquired elements are not transmitted to future generations. Selective development, whereby phenotype construction points in the adaptive direction, may induce adaptive or maladaptive evolution depending on the developmental constraints. Moreover, developmental propagation of phenotypic effects over age enables the evolution of negative senescence. Overall, we find that development plays a major evolutionary role.