Abstract
SignificanceEvolution through natural selection is an overwhelmingly complex process, and it is not surprising that theoretical approaches are strongly simplifying it. For instance, population genetics considers mainly dynamics of gene allele frequencies. Here, we develop a complementary approach to evolutionary dynamics based on three elements-organism reproduction, variations, and selection-that are essential for any evolutionary theory. By considering such general dynamics as a stochastic thermodynamic process, we clarify the nature and action of the evolutionary forces. We show that some of the forces cannot be described solely in terms of fitness landscapes. We also find that one force contribution can make organism reproduction insensitive (robust) to variations.