Adaptive peak shifts in a heterogenous environment.

The problem of moving from one coadapted gene complex to a better one can be divided into two steps: first the advantageous combination has to appear and then it has to take over the population. Selection can have contrasting effects on the two stages. When selection is weak intermediate forms are frequent, and the advantageous combination appears easily. Spreading of that advantageous combination, on the other hand, tends to be hard, as recombination acts to break it. The opposite is true when selection is strong. Spreading is easier, but if selection is also strong against the intermediate forms, the appearance of the beneficial combination becomes an extremely rare event. This inherent contrast in the optimal conditions for the two stages raises the possibility that proximity of areas differing in the intensity of selection may significantly shorten the expected waiting time for a peak shift. We studied this phenomenon in a haploid two-locus diallelic model of two neighboring subpopulations. Our results show that limited migration between the two areas might shorten the waiting time for a peak shift by orders of magnitude in comparison with either complete isolation or complete mixing. Implications for peripheral evolution and speciation are discussed.