Loss of Least-Loaded Class in Asexual Populations Due to Drift and Epistasis

We consider the dynamics of a nonrecombining haploid population of finite size that accumulates deleterious mutations irreversibly. This ratchet-like process occurs at a finite speed in the absence of epistasis, but it has been suggested that synergistic epistasis can halt the ratchet. Using a diffusion theory, we find explicit analytical expressions for the typical time between successive clicks of the ratchet for both nonepistatic and epistatic fitness functions. Our calculations show that the interclick time is of a scaling form that in the absence of epistasis gives a speed that is determined by size of the least-loaded class and the selection coefficient. With synergistic interactions, the ratchet speed is found to approach zero rapidly for arbitrary epistasis. Our analytical results are in good agreement with the numerical simulations.