A fundamental tenet of modern evolutionary thought is that mechanisms must exist to promote the development of systems of interacting non-allelic genes, since few if any significant adaptations involve single gene effects. The preceding paper (Levitan, 1973) has demonstrated that the gene arrangements on the two arms of the X-chromosome of Drosophila robusta Sturtevant provide evidence for one such interaction system, inasmuch as the frequencies of certain combinations of these linked arrangements often exhibit cyclic seasonal variation not attributable to one or another of the constituent arrangements. The writer has shown earlier (Levitan, 1955, 1958a, 1961a, 1961b, 1964) that in several localities another manifestation of such an adaptive interaction system is the presence of linkage disequilibrium. A linkage disequilibrium or association exists when, contrary to theoretical expectations in the absence of selection (Robbins, 1916; Geiringer, 1944, 1948), certain combinations of linked genes or gene arrangements are significantly in excess of random proportions, other combinations deficient. A number of writers have since developed theoretical models outlining the conditions of recombination and selection for epistatic interaction that would be conducive to such disequilibriums (see Bodmer and Parsons (1962), Feldman and Crow (1970), and Sved (1971) for references to the work of Felsenstein, Karlin, Kimura, Lewontin, and others). The previously reported linkage disequilibriums were based on the frequencies of the arrangement combinations over a period of several years. They did not take into account the possibility that the total data were seasonally heterogeneous, as proved to be the case. This paper reanalyzes these data, and presents results from several additional localities, to determine whether the concept of linkage disequilibriums remains valid in the light of the variations in the component samples. It also examines the possibility that the seasonal samples themselves evidence disequilibriums and, if so, whether their nature are attributable to the fluctuations in the linkage combinations described by Levitan (1973). It turns out that the populations studied for temporal variation contain three different X-chromosomal disequilibrium (linkage association) systems. The data for each system will be discussed separately. Methods and notation are described in Levitan (1973).
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