Chromosome replacement in mixed populations of compound-2L; free-2R and standard strains of Drosophila melanogaster : An example of unstable genetic isolation.

Crosses between compound-2L; free-2R (free-arm) and standard strains of Drosophila melanogaster produce two classes of inviable aneuploid hybrids in equal proportions: monosomic 2L and trisomic 2L. The lethal period for monosomics occurs during embryogenesis while the trisomics survive to late pupae. Since the hybrids are inviable, standard and free-arm strains within a mixed population remain genetically isolated. Genetic isolation in the absence of mating isolation offers an extreme example of unstable equilibrium. Relative fitness data indicate that an unstable equilibrium will be established between free-arm and standard strains at a ratio of 2.5∶1. Indeed, in three cage experiments established at initial ratios of 3∶1, free arms to standards, laboratory (Oregon R) or native (Okanagan S) standard strains were completely replaced in approximately 100 days by free-arm lines derived either from laboratory or from native genetic background. In contrast, one cage established at an initial ratio of 4∶1 failed to show replacement and for 92 days remained at approximately the initial ratio. Subsequent genetic analysis of flies removed from this cage identified the presence of an anomalous strain through which genetic information was transferred reciprocally between the free-arm and standard lines. The second chromosomes carried by this strain consisted of a free-2R and a standard second on the right arm of which was attached a duplication for all of 2L. While the origin of the 2L·2R+2L chromosome was uncertain, genetic and cytological examinations revealed that it represented the reciprocal crossover product expected from an exchange that generated a F(2R). Additional crosses disclosed that the transmission frequency of the asymmetrical pair of second chromosomes, as well as their right-arm crossover products, was disproportionately in favor of the short arm. Since unequal transmission was invariably greater from female parents, this phenomenon was viewed as further evidence in support of the drag hypothesis.