Nine palearctic and one North American Drosophila species of the obscura group have been compared with respect to electrophoretically detectable differences at 24 enzyme loci. The genetic distances between the species have been calculated. The constructed tree is divided into two evolutionary lineages.
The genetic polymorphism of a population of Pinus halepensis was studied using 12 electrophoretically detectable enzyme systems encoded by 7 monomorphic and 10 polymorphic loci. An estimation of population parameters revealed a relatively high average heterozygosity (H = 0.179). The total variation present in the population was partitioned into a high amount of genetic variation within groups, with only 1.3-4.4% of the total variation resulting from genic differences among groups.
Gametic frequencies were obtained in four natural populations of D. subobscura by extracting wild chromosomes and subsequently analyzing them for inversions and allozymes. The genes Lap and Pept-1, both located within the same inversions of chromosome 0, were found in striking nonrandom associations with them of the same kind and degree in all populations studied. On the contrary, the gene Acph, also located within the previously mentioned inversions, was found in linkage disequilibrium with them only in two populations and of opposite directions. This is also the case for the genes Est-9 and Hk, both located within chromosome E inversions. While the gene Est-9 was in strong linkage disequilibrium with the inversions, of the same kind and degree in all populations studied, Hk was found to be in linkage equilibrium.
Allele frequencies for the 29 genes studied do not show geographical variation except for the genes Lap, Pept-1 and Est-9, the ones found in linkage disequilibria with the geographically varying gene arrangements. Although mechanical or historical explanations for these equilibria cannot be ruled out, these data cannot be explained satisfactorily by the "middle gene explanation," which states that loci displaying such linkage disequilibria are the ones located near the break points of inversions, while the ones displaying linkage equilibria with them are located in the middle of them.
There is no evidence for consistent linkage disequilibria between pairs of loci, except for the closely linked genes of the complex locus, Est-9. This would imply, if it is not a peculiarity of the Est-9 complex, that the linkage disequilibria aye found only between very closely linked loci or that, far less closely linked genes, the associations are too weak to be detected by the usual samples sizes.
Urea denaturation of allozymes was used to provide finer resolution of allelic states within classes of different electrophoretic mobility. This method gives perfectly repeatable results. About 170 isogenic strains for the O chromosome of Drosophila subobscura, derived from two natural populations, were constructed. Their gene arrangements were studied, as well as eight polymorphic genes located on the O chromosome (Est-5, Odh, Ao, ME, Xdh, Lap, Pept-1 and Acph). Crosses performed indicate that differences in urea sensitivity are genetically controlled by the same genes that control electrophoretic mobility. Twice as many alleles have been detected in comparison to the usual electrophoretic method. However, the effective number of alleles did not increase considerably.
Studies of linkage disequilibria, by taking into account the finer resolution of allelic states, gave results nearly identical with those obtained in studies where the usual electrophoretic method was used. Although the power of the test is diminished, the absence of genic associations seems to indicate that there are no hidden linkage disequilibria in electrophoretic studies (because of consolidation effects of real alleles into few electromorph classes). The paucity of linkage disequilibria would indicate that there are no epistatic interactions such as those suggested in the model of Franklin and Lewontin (1970).
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