臺灣博碩士論文加值系統

雜交後代的不孕或死亡的雜交不親合與行為上的隔離避免種族之間基因交流障礙是造成生殖隔離及種化的關鍵。黃果蠅與擬黃果蠅是僅在兩到三百萬年前左右分化，親緣關係很近的兩個種族，很少會有交配行為的發生。若黃果蠅的雌性與擬黃果蠅的雄性雜交只能產生存活的不孕雌性成蟲，雜交雄性則在幼蟲時期死亡。在相反方向的雜交，則只能產生不孕的雄性成蟲，雜交雌性則在胚胎時期死亡。根據附屬X染色體品系的研究，顯示雜交雌性的死亡是由於黃果蠅的X染色體與擬黃果蠅的母系產物不相容所造成。為了確認有參與雜交不親合的X染色體區段，我們系統性的利用黃果蠅Y染色體帶有多一段X染色體的基因重複 (gene duplication)並且帶有Zhr的品系來篩選。我篩選22個基因重複的品系涵蓋了97.22%的X染色體，結果顯示其中有三個X染色體重複的片段顯著造成雜交雄性的死亡。根據此結果指出在黃果蠅X染色體上至少有三個區域上有基因造成黃果蠅與擬黃果蠅雜交不親合。為了瞭解基因影響行為上隔離的機制，我利用EMS突變黃果蠅，試著找出影響雄果蠅改變求偶行為並讓雌性擬黃果蠅接受的基因。結果我篩選出的突變果蠅有增加對異族雌果蠅的求偶。

Hybrid incompatibility (HI) and behavioral isolation in the forms of hybrid lethality or sterility are important barriers to prevent gene flow among species that may encounter each other, causing reproductive isolation and thus lead to speciation. The closely related Drosophila species, D. melanogaster and D. simulans, rarely mate each in nature despite diverging 2-3 million years ago. The cross between D. melanogaster females and D. simulans males produces sterile F1 females and males died at larval stage. The reciprocal cross yields sterile F1 males and embryonic lethal females. Using attached-X chromosome strains, it is proposed that the hybrid female lethality is caused by the presence of D. melanogaster X chromosome (Xm) and maternal product from D. simulans. To identify loci which contribute to the hybrid lethality, I conducted a genetic screen by crossing D. simulans females to males of the D. melanogaster X chromosome duplication lines that contain an extra X fragment on Y chromosomes in Zhr background. I have screened 22 lines covering 97.22 % of the X chromosome, and the results show that three of the 22 duplications have significant effect on hybrid male lethality. My results indicated that there are at least three regions on the Xm chromosome contributing to HI between D. melanogaster and D. simulans. To identify the genes responsible for the behavioral isolation between these sibling species, I used EMS mutagenesis to mutagenize D. melanogaster males to identify genes affecting male behaviors responsive to the acceptance by D. simulans females. The results indicated that the mutant lines we chose spending more time courtship heterospecific females.

口試委員審定書………………………………………………………….............………………i
致謝…..………………………………………........….……………………........……………ii
Abstract (English)………………………...…….………………………………....…….iv
Abstract (Chinese)………….……..……………………………….……....………………vi
List of figures……………………………………………………..………….....…….…viii
List of tables…………………………………………...…………..……….....……….…ix
Introduction…..……………..……………………………………………………….......…..1
Materials and Methods……………………………………….……………………......……6
Results…………………………………………...……………………………….……...........10
Discussions...…………………………………..……………………...……….……....….16
Reproductive isolation between D. melanogaster and D. simulans………...….....................................16
X-linked hybrid incompatibility mapping by duplication lines………….….….......................................17
References..…………………………………………………………….…….………..........22
Appendix..…………………………………………………………………….………............30

Bokel, C. (2008). EMS Screens. Drosophila 420: 119-138.
Barbash, D. A., J. Roote., and Ashburner, M. (2000). The Drosophila melanogaster hybrid male rescue gene causes inviability in male and female species hybrids. Genetics 154: 1747-1771.
Bard, F., Casano, L., Mallabiabarrena, A., Wallace, E., Saito, K., Kitayama, H., Guizzunti, G., Hu, Y., Wendler, F., DasGupta, R., Perrimon, N., Malhotra, V. (2006). Functional genomics reveals genes involved in protein secretion and Golgi organization. Nature 439: 604--607.
Baudry, E., Derome, N., Huet, M., and Veuille, M. (2006). Contrasted polymorphism patterns in a large sample of populations from the evolutionary genetics model Drosophila simulans. Genetics 173: 759-767.
Blumenstiel, J. P., Noll, A. C., Griffiths, J. A., Perera, A. G., Walton, K. N., Gilliland, W. D., Hawley, R. S., and Staehling-Hampton, K. (2009). Identification of EMS-induced mutations in Drosophila melanogaster by whole-genome sequencing. Genetics 182: 25-32.
Byrne, P. G. and Rice, W. R. (2006). Evidence for adaptive male mate choice in the fruit fly Drosophila melanogaster. Proc. Biol. Sci. 273: 917-922.
Cattani, M. V. and Presgraves, D. C (2012). Incompatibility between X chromosome factor and pericentric heterochromatic region causes lethality in hybrids between Drosophila melanogaster and its sibling species." Genetics 191: 549-559.
Chasan, R., Jin, Y., Anderson, K.V. (1992). Activation of the easter zymogen is regulated by five other genes to define dorsal-ventral polarity in the Drosophila embryo. Development 115(2): 607--616.
Cook, R. K., Deal, M. E., Deal, J. A., Garton, R. D., Brown, C. A., Ward, M. E., Andrade, R. S., Spana, E. P., Kaufman, T. C., and Cook, K. R. (2010). A new resource for characterizing X-linked genes in Drosophila melanogaster: systematic coverage and subdivision of the X chromosome with nested, Y-linked duplications. Genetics 186: 1095-1109.
Coyne, J. A. and Orr, H. A. (1989). Patterns of speciation in Drosophila. Evolution 43: 362-381.
Coyne, J. A. and Orr, H. A. (1997). ''Patterns of speciation in Drosophila'' revisited. Evolution 51: 295-303.
Coyne, J. A., C. Wicker-Thomas, and Jallon, J. M. (1999). A gene responsible for a cuticular hydrocarbon polymorphism in Drosophila melanogaster. Genet. Res. 73: 189-203.
Cully, M., Genevet, A., Warne, P., Treins, C., Liu, T., Bastien, J., Baum, B., Tapon, N., Leevers, S.J., Downward, J. (2010). A role for p38 stress-activated protein kinase in regulation of cell growth via TORC1. Mol. Cell. Biol. 30: 481--495.
Davis, A. W., Roote, J ., Morley, T., Sawamura, K., Herrmann, S., and Ashburner, M. (1996). Rescue of hybrid sterility in crosses between D. melanogaster and D. simulans." Nature 380: 157-159.
Dobzhansky, T. (1940). Speciation as a stage in evolutionary divergence. Am. Nat. 74: 312-321.
Galletti, M., Riccardo, S., Parisi, F., Lora, C., Saqcena, M.K., Rivas, L., Wong, B., Serra, A., Serras, F., Grifoni, D., Pelicci, P., Jiang, J., Bellosta, P. (2009). Identification of domains responsible for ubiquitin-dependent degradation of dMyc by glycogen synthase kinase 3beta and casein kinase 1 kinases. Mol. Cell. Biol. 29: 3424--3434.
Gavin-Smyth, J. and Matute, D. R. (2013). Embryonic lethality leads to hybrid male inviability in hybrids between Drosophila melanogaster and D. santomea. Ecol. Evol. 3: 1580-1589.
Gerard, P. R. and Presgraves, D. C. (2012). Abundant genetic variability in Drosophila simulans for hybrid female lethality in interspecific crosses to D. melanogaster. Genet. Res. 94: 1-7.
Hutter, P. and Ashburner, M. (1987). Genetic rescue of inviable hybrids between Drosophila melanogaster and its sibling species. Nature 327: 331-333.
Ivan, V., de Voer, G., Xanthakis, D., Spoorendonk, K.M., Kondylis, V., Rabouille, C. (2008). Drosophila Sec16 mediates the biogenesis of tER sites upstream of Sar1 through an arginine-rich motif. Mol. Biol. Cell 19: 4352--4365.
Orr, H. A. and Irving, S. (2000). Genetic analysis of the hybrid male rescue locus of Drosophila. Genetics 155: 225-231.
Kimura, M. (1957). Some Problems of Stochastic-Processes in Genetics. Annals of Mathematical Statistics 28: 882-901.
Kondylis, V., Goulding, S.E., Dunne, J.C., Rabouille, C., Frank, P.G., Zhu, L., Pestell, R.G., Lisanti, M.P. (2001). Biogenesis of Golgi stacks in imaginal discs of Drosophila melanogaster. Mol. Biol. Cell 12: 2308--2327.
Laturney, M. and Moehring, A. J. (2012). Fine-scale genetic analysis of species-specific female preference in Drosophila simulans. J. Evol. Biol. 25: 1718-1731.
Laturney, M. and Moehring, A. J. (2012). The genetic basis of female mate preference and species isolation in Drosophila. Int. J. Evol. Biol. 2012: 328392.
Maheshwari, S. and Barbash, D. A. (2011). The genetics of hybrid incompatibilities. Annu. Rev. Genet. 45: 331-355.
Matute, D. R. and Gavin-Smyth, J. (2014). Fine mapping of dominant x-linked incompatibility alleles in Drosophila hybrids. PLoS. Genet 10: e1004270.
Orr, H. A. (1993). Haldane''s rule has multiple genetic cause. Nature 361: 532-533.
Orr, H. A. and Presgraves, D. C. (2000). Speciation by postzygotic isolation: forces, genes and molecules. Bioessays 22: 1085-1094.
Sapiro, A. L., Ihry, R. J., Buhr, D. L., Konieczko, K. M., Ives, S. M., Engstrom, A. K., Wleklinski, N. P., Kopish, K. J., and Bashirullah, A. (2013). Rapid recombination mapping for high-throughput genetic screens in Drosophila. G3 (Bethesda) 3: 2313-2319.
Sawamura, K., Fujita, A., Yokoyama, R., Taira, T., Inoue, Y. H., Park, H. S., and Yamamoto, M. T. (1995). Molecular and genetic dissection of a reproductive isolation gene, zygotic hybrid rescue, of Drosophila melanogaster. Jpn. J. Genet. 70: 223-232.
Sawamura, K., Maehara, K., Mashino, S., Kagesawa, T., Kajiwara, M., Matsuno, K., Takahashi, A., and Takano-Shimizu, T. (2010). Introgression of Drosophila simulans nuclear pore protein 160 in Drosophila melanogaster alone does not cause inviability but does cause female sterility." Genetics 186: 669-676.
Sawamura, K., Taira, T., and Watanabe, T. K. (1993). Hybrid lethal systems in the Drosophila melanogaster species complex. I. The maternal hybrid rescue (mhr) gene of Drosophila simulans. Genetics 133: 299-305.
Sawamura, K. and Yamamoto, M. T. (1993). Cytogenetical localization of Zygotic hybrid rescue (Zhr), a Drosophila melanogaster gene that rescues interspecific hybrids from embryonic lethality. Mol. Gen. Genet. 239: 441-449.
Sawamura, K., Yamamoto, M. T., and Watanabe, T. K. (1993). Hybrid lethal systems in the Drosophila melanogaster species complex. II. The Zygotic hybrid rescue (Zhr) gene of D. melanogaster. Genetics 133: 307-313.
Siegel, R. W. and Hall, J. C. (1979). Conditioned responses in courtship behavior of normal and mutant Drosophila. PNAS 76: 3430-3434.
St Johnston, D. (2002). The art and design of genetic screens: Drosophila melanogaster. Nat. Rev. Genet. 3: 176-188.
Sturtevant, A. H. (1920). Genetics studies on Drosophila simulans. I. Introduction. hybrids with Drosophila melanogaster. Genetics 5: 488-500.
Takahashi, A., Fujiwara-Tsujii, N., Yamaoka, R., Itoh, M., Ozaki, M., and Takano-Shimizu, T. (2012). Cuticular hydrocarbon content that affects male mate preference of Drosophila melanogaster from west Africa. Int. J. Evol. Biol. 2012: 278903.
Tregenza, T. and Wedell, N. (2000). Genetic compatibility, mate choice and patterns of parentage: invited review. Mol. Ecol. 9: 1013-1027.
Tsai, I.-F. (2006). Bidirectional isolation between Zimbabwe (Z) and Cosmopolitan (M) races in Drosophila melanogaster. Institute of Molecular and Cellulor Biology, National Tsing Hwa University Master thesis.
Uenoyama, T. and Inoue, Y. (1995). Genetic studies on premating isolation in Drosophila simulans. I.AD. simulans line highly crossable with D. melanogaster. Jpn. J. Genet. 70: 365-371.
Watanabe, T. K. (1979). A gene that rescues the lethal hybrids between Drosophila melanogaster and D. simulans. Jpn. J. Genet. 54: 325-331.
Yamamoto, M. T. (1992). Inviability of hybrids between D. melanogaster and D. simulans results from the absence of simulans X not the presence of simulans Y chromosome. Genetica 87: 151-158.
Zhou, C., Huang, H., Kim, S. M., Lin, H., Meng, X., Han, K. A., Chiang, A. S., Wang, J. W., Jiao, R., and Rao, Y. (2012). Molecular genetic analysis of sexual rejection: roles of octopamine and its receptor OAMB in Drosophila courtship conditioning. J. Neurosci . 32: 14281-14287