Fitness in time-dependent environments includes a geometric phase contribution.

Phenotypic evolution implies sequential rise in frequency of new genomic sequences. The speed of the rise depends, in part, on the relative fitness (selection coefficient) of the mutant versus the ancestor. Using a simple population dynamics model, we show that the relative fitness in dynamical environments is not equal to the geometric average of the fitness over individual environments. Instead, it includes a term that explicitly depends on the sequence of the environments. For slowly varying environments, this term depends only on the oriented area enclosed by the trajectory taken by the system in the environment state space. It is closely related to the well-studied geometric phases in classical and quantum physical systems. We discuss possible biological implications of these observations, focusing on evolution of novel metabolic or stress-resistant functions.