In the last decade, the genetic basis of reproductive isolation has been shown to be surprisingly polygenic, and yet even the most efficient system currently in use could lend itself to molecular analysis only in highly selected cases. By extending the recent discovery of fertility rescue between Drosophila melanogaster and Drosophila simulans, we show that this hybridization can permit systematic and precise delineation of the genetic and molecular basis of speciation. In a region of 5% of the D. simulans genome introgressed into D. melanogaster, we discover at least six genes of hybrid male sterility and none for female sterility by deficiency mapping. A single case of hybrid inviability has been tracked down to a 3-Kb element that was inserted into the Cyclin E locus during species hybridization. The extent of interspecific genetic divergence underlying hybrid male sterility, especially in contrast with the low degree of inviability and female sterility, is far greater than expected from previous studies.O ne of the important subjects in speciation study is the genetic mechanism of reproductive isolation (1, 2). Drosophila melanogaster, with its wealth of resources, could allow the delineation of genes of reproductive isolation by genetic, cytological, and molecular means. Unfortunately, this species does not produce progeny beyond the F1 generation when crossed to any of its known sibling species, including Drosophila simulans (3-5). Since the time A. H. Sturtevant observed this sterility 80 years ago (3), evolutionary geneticists have relied on other species pairs. Th. Dobzhansky (6) was the first to do so, and many have since followed. Nevertheless, the progress has been slow (7), and not until recently has a gene of reproductive isolation been cloned from Drosophila (8). To move the field at a faster pace, a more efficient system involving D. melanogaster is highly desirable.Various attempts at bringing this species into speciation research have been made, including mating triploid D. melanogaster females to heavily irradiated D. simulans males to obtain the equivalent of F2s (9, 10), searching for sibling species that might be crosshybridizable (4, 5), examining far-flung populations that might be reproductively isolated (11, 12), and surveying for hybrid inviability rescue mutations (13-16). None has succeeded in making this ''model organism'' usable material for systematic studies of the genetics of speciation. The recent discovery of combinations of lines from D. melanogaster and D. simulans that yield fertile hybrid females has perhaps given us the best chance to tap into the resources of D. melanogaster (17).
Materials and MethodsNomenclature. We shall briefly explain the conventional Drosophila nomenclature used in this study. For example, Df(2L) J39 (31C-D; 32D-E) denotes a deletion on the left arm of the second chromosome with the two breakpoints determined to be at the locations of 31C (or 31D) and 32D (or 32E) of the polytene chromosome, respectively. This deletion is named J39. Following this ...