Fixation of advantageous mutations in an off-equilibrium population

We investigate the evolution of asexual populations subject to a large supply of deleterious mutations such that Muller's ratchet operates. In this regime, the accumulation of deleterious mutations takes place continuously with the resulting loss of the least-loaded class of individuals. In the current work, we study the effect of the supply of beneficial mutations on the ratchet's speed. We also examine how the rate of substitution of favorable mutations as well as the mean selective effect of favorable mutations that reach fixation is compared to those assuming a population at equilibrium. We observe that under Muller's ratchet, the rate of fixation of advantageous mutations is higher than that predicted for an equilibrium population. The difference between the rate supposing an equilibrium regime and that for the non-equilibrium case becomes larger as we increase the rate of deleterious mutations. On the other hand, the mean selective effect of beneficial mutations that reach fixation is smaller than the expected value for the equilibrium situation. Copyright EDP Sciences/Società Italiana di Fisica/Springer-Verlag 2006