Perturbation-reperturbation test of selection vs. hitchhiking of the two major alleles of Esterase-5 in Drosophila pseudoobscura.

A perturbation-reperturbation tests selective neutrality of 100/100/100/100/100 and 106/100/100/100/100, the two most common alleles at the highly polymorphic X-linked locus Esterase-5 in Drosophila pseudoobscura. A total of 22 replicate populations are set up in cages, 11 start at a high frequency of 76% (U) and 11 at a low frequency of 21% (N) of the 106 allele. Allele frequencies change directionally and decrease in both U and N populations as groups and reach equilibria of 60 and 14%, respectively, after 200-300 days. These changes suggest natural selection. A hypothesis of balancing selection accounts for the pattern and predicts a dynamic equilibrium. A rival neutral hypothesis accounts for the pattern equally well by postulating hitchhiking and breakup of linkage leaving the Est-5 variants to drift at neutral equilibria. A reperturbation of allele frequencies in each population, creating 22 additional reperturbed populations EN and EU, with the original populations as controls, directly addresses the question of balancing selection or hitchhiking and breakup of linkage effects. Allele frequencies do not change directionally among the reperturbed populations as a group. The hypothesis of balancing selection is rejected in favor of the hypothesis of initial hitchhiking and dissipated linkage effects. The power of the experimental design to detect selection is studied by simulation. Within the limits of power set by the design, it is concluded that the 100 and 106 are iso-fitness alleles of Est-5 under the environmental conditions of the laboratory populations. The requirements of a method of perturbation and reperturbation are discussed.