The traveling-wave approach to asexual evolution: Mullerʹs ratchet and speed of adaptation

We use traveling-wave theory to derive expressions for the rate of accumulation of deleterious mutations under Mullerʹs ratchet and the speed of adaptation under positive selection in asexual populations. Traveling-wave theory is a semi-deterministic description of an evolving population, where the bulk of the population is modeled using deterministic equations, but the class of the highest-fitness genotypes, whose evolution over time determines loss or gain of fitness in the population, is given proper stochastic treatment. We derive improved methods to model the highest-fitness class (the stochastic edge) for both Mullerʹs ratchet and adaptive evolution, and calculate analytic correction terms that compensate for inaccuracies which arise when treating discrete fitness classes as a continuum. We show that traveling-wave theory makes excellent predictions for the rate of mutation accumulation in the case of Mullerʹs ratchet, and makes good predictions for the speed of adaptation in a very broad parameter range. We predict the adaptation rate to grow logarithmically in the population size until the population size is extremely large