Mutation bias can shape adaptation in large asexual populations experiencing clonal interference

The Extended Evolutionary Synthesis invokes a role for development in shaping adaptive evolution, which in population genetics terms corresponds to mutation-biased adaptation. Critics have claimed that clonal interference makes mutation-biased adaptation rare. We consider the behavior of two simultaneously adapting traits, one with larger mutation rate U, the other with larger selection coefficient s, using asexual traveling wave models. We find that adaptation is dominated by whichever trait has the faster rate of adaptation v in isolation, with the other trait subject to evolutionary stalling. Reviewing empirical claims for mutation-biased adaptation, we find that not all occur in the "origin-fixation" regime of population genetics where v is only twice as sensitive to s as to U. In some cases, differences in U are at least ten to twelve times larger than differences in s, as needed to cause mutation-biased adaptation even in the "multiple mutations" regime. Surprisingly, when U > s in the "diffusive-mutation" regime, the required sensitivity ratio is also only two, despite pervasive clonal interference. Given two traits with identical v, the benefit of having higher s is surprisingly small, occurring largely when one trait is at the boundary between the origin-fixation and multiple mutations regimes.