Influence of plasticity and learning on evolution under directional selection.

Phenotypic plasticity and related processes (learning, developmental noise) have been proposed to both accelerate and slow down genetically based evolutionary change. While both views have been supported by various mathematical models and simulations, no general predictions have been offered as to when these alternative outcomes should occur. Here we propose a general framework to study the effects of plasticity on the rate of evolution under directional selection. It is formulated in terms of the fitness gain gradient, which measures the effect of a marginal change in the degree of plasticity on the slope of the relationship between the genotypic value of the focal trait and log fitness. If the gain gradient has the same sign as the direction of selection, an increase in plasticity will magnify the response to selection; if the two signs are opposite, greater plasticity will lead to slower response. We use this general result to derive conditions for the acceleration/deceleration under several simple forms of plasticity, including developmental noise. We also show that our approach explains the results of several specific models from the literature and thus provides a unifying framework.