Genetic draft and valley crossing

Living systems are characterized by complex adaptations which require multiple coordinated mutations in order to function. Empirical studies of fitness landscapes that result from the many possible mutations in a gene region reveal many fitness peaks and valleys that connect them. Thus, it is possible that some complex adaptations have arisen by evolutionary paths whose intermediate states are neutral or even deleterious. When intermediates are deleterious, traversing such an evolutionary path is known as \"crossing a fitness valley\". Previous efforts at studying this problem have rigorously characterized the rate at which such complex adaptations evolve in populations of roughly equally fit individuals. However, populations that are very large or have broad fitness distributions, such as many microbial populations, adapt quickly, which substantially alters the fate and dynamics of individual mutations due to the action of genetic draft. We investigate the rate at which complex adaptations evolve in these rapidly adapting populations in regions without recombination. We confirm that rapid adaptation overall increases the time required to cross a valley; however, rapid adaptation can make it easier for deeper valleys to be crossed relative to the time required for single beneficial mutations to sweep to fixation.