具有費洛蒙作用的表皮碳氫化合物常在種辨識機制及生殖隔離中扮演重要角色。在黃果蠅的族群中，已發現雌性表皮碳氫化合物及不同的交配行為有明顯的高相關性。然而，雄性表皮碳氫化合物在生殖隔離中所扮演的角色至今仍是未解的問題。本研究發現不僅在Z及M這兩群間，存有不同的雄性表皮碳氫化合物，在Z群中又可分成ZA和ZB。ZA雌果蠅偏愛與帶有Z族雄性表皮碳氫化合物的雄果蠅交配，這代表著雄性表皮碳氫化合物在生殖隔離中極為重要。另一方面，不論Z雄果蠅帶有Z或M的雄性表皮碳氫化合物，ZB雌果蠅與兩者的交配率皆很相近。而透過染色體替換品系的遺傳分析，得知雄性費洛蒙訊號的產生最主要是由第三條染色體所控制的。綜合行為實驗及表皮碳氫化合物變異分析的結果，我找到五個對ZA雌果蠅而言可能被視為交配訊號的表皮碳氫化合物，包括7T、5T、25Br、7P和5P。我的研究提供了第一個証據，顯示雄性表皮碳氫化合物的改變是一種族群分化的關鍵。造成這種雌性交尾選擇的演化因素可在不久的將來進一步的釐清。

Pheromonal cuticular hydrocarbons have been shown to play important roles in species recognition and sexual isolation. A high correlation between female cuticular hydrocarbons and differentiation of mating behavior was observed in Drosophila melanogaster. However, the role of the male cuticular hydrocarbons in this sexual isolation remains elusive. In this study, male cuticular hydrocarbons not only diverged between Z and M races, but also within Z race (namely ZA and ZB groups). Females from ZA group prefer to mate with males carrying Z male cuticular hydrocarbons indicating that the importance of the male cuticular hydrocarbons in sexual isolation. On the other hand, females from ZB group prefer to mate with Z males, but mating success of Z males carrying Z or M male cuticular hydrocarbons is similar. Genetic mapping by using chromosome substitution lines showed that the production of the male pheromonal signal is mainly controlled by the third chromosome. Several cuticular hydrocarbon candidates for ZA females preference, including 7T, 5T, 25Br, 7P, and 5P, were suggested by combining the results of behavioral experiments and cuticular hydrocarbons variation analysis. My study provides the first evidence demonstrating that male cuticular hydrocarbon changes are one of the key switches in racial differentiation. Evolutionary forces shaping female preferences might uncover in the near feature.

ALIPAZ, J. A., S. FANG, N. OSADA and C.-I WU, 2005a Evolution of Sexual Isolation during Secondary Contact: Genotypic versus Phenotypic Changes in Laboratory Populations. Am. Nat. 165: 420-428.
ALIPAZ, J. A., T. L. KARR and C.-I WU, 2005b Evolution of Sexual Isolation in Laboratory Populations: Fitness Differences between Mating Types and the Associated Hybrid Incompatibilities. Am. Nat. 165: 429-438.
ALIPAZ, J. A., C.-I WU and T. L. KARR, 2001 Gametic incompatibilities between races of Drosophila melanogaster. Proc. R. Soc. Lond. B. Biol. Sci. 268: 789-795.
ANTONY, C., T. L. DAVIS, D. A. CARLSON, J.-M. PECHINE and J.-M. JALLOM, 1985 Compared behavioral responses of male Drosophila melanogaster (Canton-S) to natural and synthetic aphrodisiacs. J. Chem. Ecol. 11: 1617-1629.
ANTONY, C., and J.-M. JALLOM, 1982 The chemical basis for sex recognition in Drosophila melanogaster. J. Insect Physiol. 28: 873-880.
BARTELT, R. J., M. T. ARMOLD, A. M. SCHANER and L. L. JACKSON, 1986 Comparative analysis of cuticular hydrocarbons in the Drosophila virilis species group. Comp. Biochem. Physiol. B 83: 731-742.
BASTOCK, M., and A. MANNING, 1955 The courtship of Drosophila melanogaster. Behaviour 8: 85-111.
BEGUN, D. J., and C. F. AQUADRO, 1993 African and North American populations of Drosophila melanogaster are very different at the DNA level. Nature 365: 548-550.
BLOMQUIST, G. J., J. W. DILLWITH and T. S. ADAMS, 1987 Biosynthesis and endocrine regulation of sex pheromone production in Diptera, pp.217-250 in Pheromone Biochemistry, edited by G. D. Prestwich, G. J. Blomquist. Academic Press, New York.
BLOMQUIST, G. J., E. C. TOOLSON and D. R. NELSON, 1985 Epicuticular hydrocarbons of Drosophila pseudoobscura (Diptera: Drosophilidae). Insect Biochem. 15: 25-34.
Blomquist, G. J., and R. G. Vogt, 2003 Genetic studies on pheromone production in Drosophila, pp. 254-281, in Insect
pheromone biochemistry and Molecular biology, edited by J.-M. Jallon, C. Wicker-Thomas. Academic Press, California.
BONAVITA-COUGOURDAN, A., J. L. CL´EMENT and C. LANGE, 1987 Nestmate recognition: The role of cuticular hydrocarbons in the ant Camponotus vagus Scop. J. Entomol. Sci. 22: 1-10.
BRAY, S., and H. AMREIN, 2003 A putative Drosophila pheromone receptor expressed in male-specific taste neurons is required for efficient courtship. Neuron 39: 1019-1029.
BURG, J. G., and R. C. AXTELL, 1984 Monitoring house fly, Musca domestica (Diptera: Muscidae), populations in caged-layer poultry houses using a baited jug-trap. Environ. Entomol. 13: 1083-1090.
BUTLIN, R. K., and M. G. RITCHIE, 1989 Genetic coupling in mate recognition systems: what is the evidence? Biol. J. Linn. Soc. 37: 237-246.
CARLSON, D. A., and M. BEROZA, 1973 Field evaluation of (Z)-9- tricosene, a sex pheromone of the house fly. Environ. Entomol. 2: 555-559.
COLEGRAVE, N., H. HOLLOCHER, K. HINTON and G. RITCHIE, 2000 The courtship song of African Drosophila melanogaster. J. Evol. Biol. 13: 145-150.
COYNE, J. A., 1992 Genetics and speciation. Nature 355: 511-515.
COYNE, J. A., 1996a Genetics of a difference in male cuticular hydrocarbons between two sibling species, Drosophila simulans and D. sechellia. Genetics 143: 1689-1698.
COYNE, J. A., and B. CHARLESWORTH, 1997 Genetics of a pheromonal difference affecting sexual isolation between Drosophila mauritiana and D. sechellia. Genetics 145: 1015-1030.
COYNE, J. A., A. P. CRITTENDEN and K. MAH, 1994 Genetics of a pheromonal difference contributing to reproductive isolation in Drosohpila. Science 265: 1461-1464.
COYNE, J. A., C. WICKER-THOMAS and J.-M. JALLON, 1999 A gene responsible for a cuticular hydrocarbon polymorphism in Drosophila melanogaster. Genet. Res. 73: 189-203.
DALLERAC, R., C. LABEUR, J. M. JALLON, D. C. KNIPPLE, W. L. ROELOFS et al., 2000 A delta 9 desaturase gene with a different substrate specificity is responsible for the cuticular diene hydrocarbon polymorphism in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA 97: 9449-9454.
DANI, F. R., G. R. JONES, S. DESTRI, S. H. SPENCER and S. TURILLAZZI, 2001 Deciphering the recognition signature within the cuticular chemical profile of paper wasps. Anim. Behav. 62: 165-171.
DE RENOBALES, M., C. CRIPPS, D. W. STANLEY-SAMUELSON, R. A. JURENKA and G. J. BLOMQUIST, 1987 Biosynthesis of linoleic acid in insects. Trends Biochem. Sci. 12: 364-366.
DIETEMANN, V., C. PEETERS, J. LIEBIG, V. THIVET and B. HOLLDOBLER, 2003 Cuticular hydrocarbons mediate discrimination of reproductives and nonreproductives in the ant Myrmecia gulosa. Proc. Natl. Acad. Sci. USA 100: 10341-10346.
EDNEY, E. B., 1967 Water balance in desert arthropods. Despite their small size, arthropods may be highly adapted for life in xeric conditions. Science 156: 1059-1066.
FANG, S., A. TAKAHASHI and C.-I WU, 2002 A mutation in the promoter of desaturase 2 is correlated with sexual isolation between Drosophila behavioral races. Genetics 162: 781-784.
FERVEUR, J.-F., 1991 Genetic control of pheromones in Drosophila simulans. I. Ngbo, a locus on the second chromosome. Genetics 128: 293-301.
FERVEUR, J.-F., 2005 Cuticular hydrocarbons: their evolution and roles in Drosophila pheromonal communication. Behav. Genet. 35: 279-295.
FERVEUR, J.-F., and J.-M. JALLON, 1993a Genetic control of pheromones in Drosophila simulans. II. kete, a locus on the X chromosome. Genetics 133: 561-567.
FERVEUR, J.-F., and J.-M. JALLON, 1993b Nerd, a locus on chromosome III, affects male reproductive behavior in Drosophila melanogaster. Naturwissenschaften 80: 474-475.
FERVEUR, J.-F., and J.-M. JALLON, 1996 Genetic control of male cuticular hydrocarbons in Drosophila melanogaster. Genet. Res. 67: 211-218.
FERVEUR, J.-F., F. SAVARIT, C. J. O'KANE, G. SUREAU, R. J. GREENSPAN et al., 1997 Genetic feminization of pheromones and its behavioral consequences in Drosophila males. Science 276: 1555-1558.
FERVEUR, J.-F., and G. SUREAU, 1996 Simultaneous influence on male courtship of stimulatory and inhibitory pheromones
produced by live sex-mosaic Drosophila melanogaster. Proc. R. Soc. Lond. B. Biol. Sci. 263: 967-973.
FITZPATRICK, S., 1994 Colourful migratory birds: evidence for a mechanism other than parasite resistance for the maintenance of "good genes" sexual selection. Proc. R. Soc. Lond. Ser. B. Biol. Sci. 257: 155-160.
GIBBS, A. G., 1998 Water-proofing properties of cuticular lipids. Amer. Zool. 38: 471-482.
GREENBERG, A. J., J. R. MORAN, J. A. COYNE and C.-I WU, 2003 Ecological adaptation during incipient speciation revealed by precise gene replacement. Science 302: 1754-1757.
GREENSPAN, R. J., and J.-F. FERVEUR, 2000 Courtship in Drosophila. Annu. Rev. Genet. 34: 205-232.
GROSSFIELD, J., 1971 Behavioral differentiation of three races of Drosophila auraria. J. Hered. 62: 117-118.
HOLLOCHER, H., C.-T. TING, F. POLLACK and C.-I WU, 1997a Incipient speciation by sexual isolation in Drosophila melanogaster: variation in mating preference and correlation between sexs. Evolution 51: 1175-1181.
HOLLOCHER, H., C.-T. TING, M.-L. WU and C.-I WU, 1997b Incipient speciation by sexual isolation in Drosophila melanogaster: extensive genetic divergence without reinforcement. Genetics 147: 1191-1201.
IYENGAR, V. K., and T. EISNER, 1999 Female choice increases offspring fitness in an arctiid moth (Utetheisa ornatrix). Proc. Natl. Acad. Sci. USA 96: 15013-15016.
JALLON, J.-M., 1984 A few chemical words exchanged by Drosophila during courtship and mating. Behav. Genet. 14: 441-478.
JOHNSTONE, R. A., 1995 Sexual selection, honest advertisement and the handicap priniciple reviewing the evidence. Biol. Rev. Camb. Philos. Soc. 70: 1-65.
KAVALIERS, M., and D. D. COLWELL, 1995 Discrimination by female mice between the odours of parasitized and non-parasitized males. Proc. R. Soc. Lond. Ser. B. Biol. Sci. 261: 31-35.
KLEINBAUM, D. G., L. L. KUPPER, K. E. MULLER and A. NIZAM, 1998 Applied Regression Analysis and Other Multivariable Methods. Duxbury Press, Pacific Grove, CA.
KUO, T.-H., 2004 Behavioral and genetics of incipient speciation in Drosophila melanogaster. Master's thesis. National Tsing Hua University, Hsinchu, Taiwan.
LOCKEY, K. H., 1976 Cuticular hydrocarbons of Locusta, Schistocerca and Periplaneta. Insect Biochem. 6: 457-472.
MARCILLAC, F., F. BOUSQUET, J. ALABOUVETTE, F. SAVARIT and J.-F. FERVEUR, 2004 Genetic and molecular characterization of a mutation that largely affects the production of sex pheromones in Drosophila melanogaster. Genetics: doi: 10.1534/genetics.1104.033159.
MARKOW, T. A., and P. M. O'GRADY, 2005 Evolutionary genetics of reproductive behavior in Drosophila: connecting the dots. Annu. Rev. Genet. 39: 263-291.
MAYR, E., 1963 Animal Species and Evolution. Harvard Univ. Press, Cambridge, MA.
MCROBERT, S. P., and L. TOMPKINS, 1987 The effect of light on the sexual behavior of Drosophila affinis. Behav. Neural. Biol. 47: 151-157.
MEUNIER, N., J. F. FERVEUR and F. MARION-POLL, 2000 Sex-specific non-pheromonal taste receptors in Drosophila. Curr. Biol. 10: 1583-1586.
MONTOOTH, K. L., and A. G. GIBBS, 2003 Cuticular pheromones and water balance in the house fly, Musca domestica. Comp. Biochem. Physiol. A 135: 457-465.
MORGAN, P. B., I. H. GILBERT and R. L. FYE, 1974 Evaluation of (Z)-9-tricosene for attractancy for Musca domestica in the field. Fla. Entomol. 57: 136-140.
NAYAK, S. V., and R. N. SINGH, 1983 Sensilla on the tarsal segments and the mouthparts of adult Drosophila melanogaster Meigen (Diptera: Drosophilidae). Int. J. Insect Morphol. Embryol. 12: 273-291.
PAGE, M., L. J. NELSON, M. I. HAVERTY and G. J. BLOMQUIST, 1990 Cuticular hydrocarbons of eight species of North American cone beetles, Conophthorus Hopkins. J. Chem. Ecol. 16: 1173-1198.
PATTERSON, H., 1985 The recognition concept of species. In Species and Speciation, ed. es vrba. Transvaal Mus. Monogr. 4:21-29. Pretoria: Transvall Mus.
POSSIDENTE, D. R., and R. K. MURPHEY, 1989 Genetic control of sexually dimorphic axon morphology in Drosophila sensory
neurons. Dev. Biol. 132: 448-457.
ROUAULT, J., P. CAPY and J.-M. JALLON, 2001 Variations of male cuticular hydrocarbons with geoclimatic variables: an
adaptative mechanism in Drosophila melanogaster? Genetica 110: 117-130.
ROUAULT, J. D., C. MARICAN, C. WICKER-THOMAS and J.-M. JALLON, 2004 Relations between cuticular hydrocarbon (HC)
polymorphism, resistance against desiccation and breeding temperature; a model for HC evolution in D. melanogaster and D. simulans. Genetica 120: 195-212.
ROURKE, B. C., and A. G. GIBBS, 1999 Effects of lipid phase transitions on cuticular permeability: model membrane and in situ studies. J. Exp. Biol. 202: 3255-3262.
SAVARIT, F., and J. F. FERVEUR, 2002 Genetic study of the production of sexually dimorphic cuticular hydrocarbons in relation with the sex-determination gene transformer in Drosophila melanogaster. Genet. Res. 79: 23-40.
SCOTT, D., 1986 Sexual mimicry regulates the attractiveness of mated Drosophila melanogaster females. Proc. Natl. Acad. Sci. USA 83: 8429-8433.
SHANBHAG, S. R., and R. N. SINGH, 1992 Functional implications of the projections of neurons from individual labellar sensillum of Drosophila melanogaster as revealed by neuronal-marker horseradish peroxidase. Cell Tissue Res. 267: 273-282.
SPIETH, H. T., 1974 Courtship behavior in Drosophila. Annu. Rev. Entomol. 19: 385-405.
SPIETH, H. T., and T. C. HSU, 1950 The influence of light on the mating behavior of seven species of the Drosophila
melanogaster species group. Evolution 4: 316-325.
STOCKER, R. F., 1994 The organization of the chemosensory system in Drosophila melanogaster: a review. Cell Tissue Res. 275: 3-26.
STURTEVANT, A. H., 1915 Experiments on sexual recognition and the problems of sexual selection in Drosophila. J. Anim. Behav. 5: 351-366.
SUREAU, G., and J.-F. FERVEUR, 1999 Co-adaptation of pheromone production and behavioural responses in Drosophila

melanogaster males. Genet. Res. 74: 129-137.
TAKAHASHI, A., S. C. TSAUR, J. A. COYNE and C.-I WU, 2001 The nucleotide changes governing cuticular hydrocarbon variation and their evolution in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA 98: 3920-3925.
TAKAHASHI, S., and A. Gassa, 1995 Roles of cuticular hydrocarbons in intra- and interspecific recognition behavior of two Rhinotermitidae species. J. Chem. Ecol. 21: 1837-1845.
TING, C.-T., A. Takahashi and C.-I WU, 2001 Incipient speciation by sexual isolation in Drosophila: concurrent evolution at multiple loci. Proc. Natl. Acad. Sci. USA 98: 6709-6713.
TOMPKINs, L., 1984 Genetic analysis of sex appeal in Drosophila. Behav. Genet. 14: 411-440.
VENARD, R., C. Antony and J.-M. Jallon, 1989 Drosophila chemoreceptors, pp 377-385 in: Neurobiology of sensory systems, edited by R. N. Singh and J. S. Strausfeld. Plenum, New York.
WAGNER, D., M. Tissot, W. Cuevas and D. M. Gordon, 2000 Harvester ants utilize cuticular hydrocarbons in nestmate recognition. J. Chem. Ecol. 26: 2245-2257.
WU, C.-I, H. Hollocher, D. J. Begun, C. F. Aquadro, Y. Xu et al., 1995 Sexual isolation in Drosophila melanogaster: a
possible case of incipient speciation. Proc. Natl. Acad. Sci. USA 92: 2519-2523.
WU, C.-I, and M. F. Palopoli, 1994 Genetics of post-mating reproductive isolation in animals. Annu. Rev. Genet. 28: 283-308.
WU, C.-I, and C.-T. Ting, 2004 Genes and speciation. Nat. Rev. Genet. 5: 114-122.
ZAHAVI, A., 1991 On the definition of sexual selection, Fisher’s model, and the evolution of waste and of signals in general. Anim. Behav. 42: 501-503.