No description available.

Many species of Drosophila have an X chromosome inversion polymorphism: the Standard arrangement and the Sex-Ratio (SR) arrangement. Male carriers of SR transmit predominantly X-bearing sperm and hence produce nearly all-female progeny. In the absence of strong counteracting selection, SR would be meiotically driven to fixation, causing population extinction. In PART I, the role of virility selection in maintaining the SR polymorphism in D. pseudoobscura is examined by partitioning it into two components. The SR males are found to suffer substantial virility reduction which would not be detected if the differences between the two components are not heeded. The significance of this finding is discussed in light of two specific observations usually associated with the Sex-Ratio polymorphism in this species. One is the absence of suppressor modifiers of "sex-ratio" expression and the other is the temperature-dependent distribution of the sex-ratio trait. In PART II, a model is proposed to describe the behavior of the autosomal suppressor modifiers of "sex-ratio" meiotic drive. These modifiers, if neutral in other respects, should increase because they tend to be associated with the rare sex (males). However, selection operating on the sex-linked drive locus will sometimes act against autosomal modifiers as well because the two loci are always in gametic phase disequilibrium. Conditions under which modifiers will not increase are presented in terms of the fitnesses of different genotypes at the sex-linked drive locus. To prevent an increase in modifier alleles, the fitness of Sex-Ratio males relative to Standard males has to be no greater than 0.3. This result is in good agreement with that of PART I. PART III deals with "sex-ratio" genes, tightly linked within the Sex-Ratio inversion. By taking advantage of the fact that the Sex-Ratio chromosome of D. persimilis (SR(B)) is homosequential to the Standard chromosome of D. pseudoobscura (ST(A)), two reciprocal introgression experiments were carried out. Individual segments of SR(B) or ST(A) were introgressed into the genome of D. pseudoobscura or D. persimilis, respectively. Males possessing a hybrid SR(B)-ST(A) X chromosome and a genetic background derived from either of the two species were tested for fertility and "sex-ratio" expression. It was found that, in terms of the meiotic drive genes, the Sex-Ratio chromosome differs extensively from the Standard chromosome. Because recombinations of these genes result in a complete loss of sex-ratio expression, this finding lends strong support to the hypothesis of gene coadaptation. Coadaptation, in this context, is the advantage of being transmitted preferentially. In light of this finding, evolution of the sex-ratio system in these two sibling species is discussed. Introgression experiments also yielded information about hybrid sterility. Four types of sterility interactions were identified; one of them involved at least three genetic elements. With reciprocal introgression, sterility interactions were found to be "asymmetric". The asymmetry is fully expected from the viewpoint of evolution of postmating reproductive isolation. / Medicine, Faculty of / Medical Genetics, Department of / Graduate

No description available.

<p>Biodiversity is generated by the process of speciation. Because biological species are defined as populations that are unable to exchange genes with one another, the study of the evolution of reproductive isolation occupies the center of speciation research. A key to deciphering how reproductive isolation evolves is to understand the genetic changes that underlie these barriers to gene flow. Intrinsic postzygotic barriers, such as hybrid sterility or inviability, are known to impede gene flow and especially lend themselves to genetic analysis because of their ease of study in a laboratory setting. Because hybrid sterility likely evolves before hybrid inviability, it potentially plays an important role in the cessation of gene flow. Yet, while their X-linked counterparts have been precisely localized, we remain ignorant of the numbers of and interactions among dominant autosomal loci that are predicted to contribute to F1 hybrid male sterility. </p><p> To address this conceptual void, I examine the genetic architecture of hybrid male sterility between the allopatric sister species Drosophila persimilis and D. pseudoobscura bogotana. First, using a large-scale backcross analysis, I fine-map autosomal QTL from D. persimilis that confer sterility in male hybrids. This fine-mapping shows that loci contributing to hybrid male sterility reside outside chromosomal rearrangements (i.e., regions of reduced recombination) in this allopatric species pairs. In contrast, these QTL do not contribute to hybrid male sterility in the comparable sympatric hybridizing species D. persimilis and D. pseudoobscura, as predicted by models that suggest that hybridizing species persist because of broad regions of reduced recombination. Next, I use a serial backcross design to introgress these sterility-conferring QTL from D. persimilis into a D. p. bogatana genetic background devoid of other alleles from D. persimilis. This introgression study tested a prediction of the dominance theory proposed to explain Haldane's rule: dominant-acting autosomal loci should interact with recessive-acting X-linked loci to produce sterile hybrid males. Surprisingly, the results demonstrated that the "composite" dominance of the autosomal QTL is more important than the dominance of individual QTL for producing Haldane's rule: epistasis among loci elevated their dominant effects on sterility such that individually-recessive-acting autosomal QTL can contribute to F1 male infertility. Finally, using recombination to generate independent lines bearing only small segments of the identified QTL regions, I examine whether single or multiple loci within these regions contribute to the overall effect of hybrid sterility. While the effect of one QTL depends on epistasis between several loci within that small region, the effect of the other QTL appears to derive from a single genetic factor. These results suggest that estimates of the number of genes that contribute to reproductive isolation are at best, likely too low and, at worst, unattainable with the mapping resolution attainable by standard backcross and introgression approaches.</p><p> This dissertation addresses both evolutionary and genetic hypotheses of intrinsic postzygotic isolation. Hybrid male sterility between D. persimilis and D. p. bogotana clearly involves highly specific and complex interactions between homoospecific loci. The mapping results presented here also lay the foundation for the identification and cloning of multiple autosomal sterility-conferring "speciation genes."</p> / Dissertation

Patterns of heterospecific matings are important to understand in order to learn about speciation. I proposed a classical signal detection model as a representation of heterospecific mating patterns in both sexes. From the model I proposed that males would be able to achieve successive heterospecific mating success, as those males were more similar to conspecifics. I also proposed that restrictive females that rejected heterospecific matings would also be more likely to reject conspecific matings. I used the sibling species Drosophila pseudoobscura and D. persimilis to examine heterospecific mating patterns in males and females. I found that males previously accepted as heterospecific mates were more likely to achieve heterospecific matings than males that had previously been rejected as heterospecific mates. This was most likely due to decreased courtship by rejected males. Males rejected heterospecifically but mated conspecifically also were less likely to achieve heterospecific matings than males that had initially been accepted as heterospecific mates. This indicated that the same males were able to achieve successive heterospecific matings, despite rejected males also having had initial mating success. This was again likely influenced by decreased courtship in rejected males. Furthermore, I found that females that rejected heterospecific males were also more likely to reject conspecific males than naive females. There was also a non-significant trend of females that previously accepted heterospecific males were more likely to again accept heterospecific males than females that had previously rejected heterospecific males. / Thesis / Master of Science (MSc)