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Gene Review

tra2  -  transformer 2

Drosophila melanogaster

Synonyms: CG10128, Dmel\CG10128, TRA 2, TRA-2, TRA2, ...
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Disease relevance of tra2


High impact information on tra2

  • We find that Tra2 interacts with itself, with the splicing regulator Transformer, and with the general splicing factor SF2 in vitro and in the yeast two-hybrid system [3].
  • These results demonstrate that both protein-RNA and protein-protein interactions are involved in tra2-dependent activation and repression of alternative splicing [3].
  • Characterization of the proteins associated with affinity-purified complex formed on dsx pre-mRNA reveals the presence of Tra, Tra2, SR proteins, and additional unidentified components [4].
  • We conclude that Tra, Tra2, and SR proteins are essential components of a splicing enhancer complex [4].
  • Sex-specific splicing and polyadenylation of dsx pre-mRNA requires a sequence that binds specifically to tra-2 protein in vitro [1].

Biological context of tra2

  • The transformer-2 (tra-2) gene of Drosophila melanogaster plays essential roles in both sexual differentiation in the female soma and spermatogenesis in the male germ line [5].
  • The translation of products from a tra-2-lacZ fusion gene in both sexes suggests that the female-specific functioning of tra-2 in somatic tissues is not attributable to a translational mechanism [5].
  • Temperature-shift experiments with temperature-sensitive tra-2 mutants demonstrate that within single cell lineages tra-2+ function is required at several times, and probably continuously, during development for the occurrence of a series of determinative decisions necessary for female sexual differentiation [6].
  • Four repeat elements identical to the D. melanogaster TRA/TRA-2 binding sites have been found in the untranslated region of the female-specific exon 4, predicting a common regulatory splicing mechanism in all studied species of Diptera [7].
  • Moreover, by altering the dosage of either the transgenes or the endogenous tra2(+) loci, one can vary the effect over a wide range of mutant phenotypes [8].

Anatomical context of tra2

  • We show that although two transcripts are expressed male specifically in the germ line, the tra-2 transcripts expressed in the soma are not sex-specific [5].
  • att, a target for regulation by tra2 in the testes of Drosophila melanogaster, encodes alternative RNAs and alternative proteins [9].
  • Chromosomal females homozygous for tra or tra-2 have no detectable hemolymph vitellogenins, while those homozygous for tra-2OTF exhibit appreciable levels of these proteins [10].
  • Female flies mutant for alleles of Sex-lethal, defective in sex determination, or null alleles of transformer or transformer-2 are converted into phenotypic males that formed male-specific abdominal muscles [11].
  • We find that Tra2 is expressed at higher levels in primary spermatocytes of males than in other cell types [12].

Associations of tra2 with chemical compounds

  • Independently of pre-mRNA, SRm160/300 specifically interacts with U2 snRNP and with a human homolog of the Drosophila alternative splicing regulator Transformer 2, which binds to purine-rich ESEs [13].
  • The products of the tra-2 gene are also required for continuous transcription of the yolk-protein genes, suggesting that the pathway inhibited by the cycloheximide is that of the sex-determination hierarchy [14].
  • 20-hydroxyecdysone can affect the transcription of these genes in males and females, as can mutations in the sex-determining genes tra, tra-2, ix and dsx [14].
  • Orcein-stained testis preparations from tra-2 males reveal a spermiogenic defect similar to that associated with dominant male sterile (X; autosome) translocations [15].
  • All isoforms of BmTRA-2 protein contain two arginine/serine-rich domains and one RNA recognition motif, showing striking organizational similarity to Drosophila TRA-2 proteins [16].

Regulatory relationships of tra2

  • These and other results support the model that the tra-2+ and tra+ loci act to control the expression of the bifunctional doublesex (dsx) locus [6].
  • The mechanisms of this process were investigated using an in vitro system in which recombinant TRA/TRA2 could activate the female-specific 5'-splice site of fru [17].
  • We suggest that tra-2 activity in somatic tissues is regulated through a post-translational sex-specific interaction with the product of the tra gene rather than through the expression of a female-specific tra-2 polypeptide [5].
  • Sex-specific processing of the Drosophila exuperantia transcript is regulated in male germ cells by the tra-2 gene [18].

Other interactions of tra2

  • We show that mutations in the Drosophila LAMMER kinase, Doa, alter sexual differentiation and interact synergistically with tra and tra2 mutations [19].
  • Furthermore, using a two-hybrid system, we show protein-protein interactions between RBP1 and itself and between RBP1 and TRA-2 [20].
  • We report here that tra-2 is required in male germ cells for efficient male-specific processing of exu RNA; in the absence of tra-2, X/Y males produce a new mRNA which is processed at its 3' end so that it contains sequences normally specific to the female 3' untranslated region [18].
  • Therefore, the wild-type products of the genes Sex-lethal, transformer and transformer-2 act to prevent the differentiation of male-specific muscles in female flies [11].
  • These arginine-rich regions contain stretches of arginine-serine dipeptides like those found in transformer, transformer-2, and suppressor-of-white-apricot proteins, all of which have been identified as regulators of mRNA splicing in Drosophila melanogaster [21].

Analytical, diagnostic and therapeutic context of tra2

  • Sequence analysis of the tra-2 gene and 10 tra-2 cDNA clones coupled with nuclease protection analysis reveals a variety of alternatively spliced tra-2 mRNAs that each encode one of four distinct but overlapping polypeptides [5].
  • Molecular cloning of a mouse homologue for the Drosophila splicing regulator Tra2 [22].


  1. Sex-specific splicing and polyadenylation of dsx pre-mRNA requires a sequence that binds specifically to tra-2 protein in vitro. Hedley, M.L., Maniatis, T. Cell (1991) [Pubmed]
  2. Sex determination in the Drosophila germline is dictated by the sexual identity of the surrounding soma. Waterbury, J.A., Horabin, J.I., Bopp, D., Schedl, P. Genetics (2000) [Pubmed]
  3. The role of specific protein-RNA and protein-protein interactions in positive and negative control of pre-mRNA splicing by Transformer 2. Amrein, H., Hedley, M.L., Maniatis, T. Cell (1994) [Pubmed]
  4. A splicing enhancer complex controls alternative splicing of doublesex pre-mRNA. Tian, M., Maniatis, T. Cell (1993) [Pubmed]
  5. Alternative splicing of the sex determination gene transformer-2 is sex-specific in the germ line but not in the soma. Mattox, W., Palmer, M.J., Baker, B.S. Genes Dev. (1990) [Pubmed]
  6. Sex determination in Drosophila melanogaster: analysis of transformer-2, a sex-transforming locus. Belote, J.M., Baker, B.S. Proc. Natl. Acad. Sci. U.S.A. (1982) [Pubmed]
  7. Isolation and characterization of the Bactrocera oleae genes orthologous to the sex determining Sex-lethal and doublesex genes of Drosophila melanogaster. Lagos, D., Ruiz, M.F., Sánchez, L., Komitopoulou, K. Gene (2005) [Pubmed]
  8. Temperature-dependent gene silencing by an expressed inverted repeat in Drosophila. Fortier, E., Belote, J.M. Genesis (2000) [Pubmed]
  9. att, a target for regulation by tra2 in the testes of Drosophila melanogaster, encodes alternative RNAs and alternative proteins. Madigan, S.J., Edeen, P., Esnayra, J., McKeown, M. Mol. Cell. Biol. (1996) [Pubmed]
  10. Interactions between sex-transformation mutants of Drosophila melanogaster. I. Hemolymph vitellogenins and gonad morphology. Ota, T., Fukunaga, A., Kawabe, M., Oishi, K. Genetics (1981) [Pubmed]
  11. Differentiation of a male-specific muscle in Drosophila melanogaster does not require the sex-determining genes doublesex or intersex. Taylor, B.J. Genetics (1992) [Pubmed]
  12. Concentration dependent selection of targets by an SR splicing regulator results in tissue-specific RNA processing. Qi, J., Su, S., McGuffin, M.E., Mattox, W. Nucleic Acids Res. (2006) [Pubmed]
  13. The SRm160/300 splicing coactivator is required for exon-enhancer function. Eldridge, A.G., Li, Y., Sharp, P.A., Blencowe, B.J. Proc. Natl. Acad. Sci. U.S.A. (1999) [Pubmed]
  14. The use of an inhibitor of protein synthesis to investigate the roles of ecdysteroids and sex-determination genes on the expression of the genes encoding the Drosophila yolk proteins. Bownes, M., Scott, A., Blair, M. Development (1987) [Pubmed]
  15. The dual functions of a sex determination gene in Drosophila melanogaster. Belote, J.M., Baker, B.S. Dev. Biol. (1983) [Pubmed]
  16. Cloning and alternative splicing analysis of Bombyx mori transformer-2 gene using silkworm EST database. Niu, B.L., Meng, Z.Q., Tao, Y.Z., Lu, S.L., Weng, H.B., He, L.H., Shen, W.F. Acta Biochim. Biophys. Sin. (Shanghai) (2005) [Pubmed]
  17. Enhancer-dependent 5'-splice site control of fruitless pre-mRNA splicing. Lam, B.J., Bakshi, A., Ekinci, F.Y., Webb, J., Graveley, B.R., Hertel, K.J. J. Biol. Chem. (2003) [Pubmed]
  18. Sex-specific processing of the Drosophila exuperantia transcript is regulated in male germ cells by the tra-2 gene. Hazelrigg, T., Tu, C. Proc. Natl. Acad. Sci. U.S.A. (1994) [Pubmed]
  19. Protein phosphorylation plays an essential role in the regulation of alternative splicing and sex determination in Drosophila. Du, C., McGuffin, M.E., Dauwalder, B., Rabinow, L., Mattox, W. Mol. Cell (1998) [Pubmed]
  20. In vivo analysis of the functional domains of the Drosophila splicing regulator RBP1. Heinrichs, V., Baker, B.S. Proc. Natl. Acad. Sci. U.S.A. (1997) [Pubmed]
  21. Structure and expression of the Drosophila melanogaster gene for the U1 small nuclear ribonucleoprotein particle 70K protein. Mancebo, R., Lo, P.C., Mount, S.M. Mol. Cell. Biol. (1990) [Pubmed]
  22. Molecular cloning of a mouse homologue for the Drosophila splicing regulator Tra2. Segade, F., Hurlé, B., Claudio, E., Ramos, S., Lazo, P.S. FEBS Lett. (1996) [Pubmed]
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