Interpretation of X Chromosome Dose at Sex-lethalRequires Non-E-Box Sites for the Basic Helix-Loop-Helix Proteins SISB and Daughterless

REFERENCES

• Atchley, W. R., and W. M. Fitch. 1997. A natural classification of the basic helix-loop-helix class of transcription factors. Proc. Natl. Acad. Sci. USA 94:5172–5176.
   PubMed Web of Science ®Google Scholar
• Atchley, W. R., W. Terhalle, and A. Dress. 1999. Positional dependence, cliques, and predictive motifs in the bHLH protein domain. J. Mol. Evol. 48:501–516.
   PubMed Web of Science ®Google Scholar
• Barbash, D. A., and T. W. Cline. 1995. Genetic and molecular analysis of the autosomal component of the primary sex determination signal of Drosophila melanogaster. Genetics 141:1451–1471.
   PubMed Web of Science ®Google Scholar
• Blackwell, T. K., J. Huang, A. Ma, L. Kretzner, F. W. Alt, R. N. Eisenman, and H. Weintraub. 1993. Binding of myc proteins to canonical and noncanonical DNA sequences. Mol. Cell. Biol. 13:5216–5224.
   PubMed Web of Science ®Google Scholar
• Blackwell, T. K., L. Kretzner, E. M. Blackwood, R. N. Eisenman, and H. Weintraub. 1990. Sequence-specific DNA binding by the c-Myc protein. Science 250:1149–1151.
   PubMed Web of Science ®Google Scholar
• Blackwell, T. K., and H. Weintraub. 1990. Differences and similarities in DNA-binding preferences of MyoD and E2A protein complexes revealed by binding site selection. Science 250:1104–1110.
   PubMed Web of Science ®Google Scholar
• Burz, D. S., R. Rivera-Pomar, H. Jackle, and S. D. Hanes. 1998. Cooperative DNA-binding by Bicoid provides a mechanism for threshold-dependent gene activation in the Drosophila embryo. EMBO J. 17:5998–6009.
   PubMedGoogle Scholar
• Cabrera, C. V., and M. C. Alonso. 1991. Transcriptional activation by heterodimers of the achaete-scute and daughterless gene products of Drosophila. EMBO J. 10:2965–2973.
   PubMedGoogle Scholar
• Cai, H. N., D. N. Arnosti, and M. Levine. 1996. Long-range repression in the Drosophila embryo. Proc. Natl. Acad. Sci. USA 93:9309–9314.
   PubMedGoogle Scholar
• Carey, M.. 1998. The enhanceosome and transcriptional synergy. Cell 92:5–8.
   PubMed Web of Science ®Google Scholar
• Chen, H., A. Thiagalingam, H. Chopra, M. W. Borges, J. N. Feder, B. D. Nelkin, S. B. Baylin, and D. W. Ball. 1997. Conservation of the Drosophila lateral inhibition pathway in human lung cancer: a hairy-related protein (HES-1) directly represses achaete-scute homolog-1 expression. Proc. Natl. Acad. Sci. USA 94:5355–5360.
   PubMedGoogle Scholar
• Cline, T. W.. 1993. The Drosophila sex determination signal: how do flies count to two?. Trends Genet. 9:385–390.
   PubMedGoogle Scholar
• Cline, T. W.. 1988. Evidence that sisterless-a and sisterless-b are two of several discrete “numerator elements” of the X/A sex determination signal in Drosophila that switch Sxl between two alternative stable expression states. Genetics 119:829–862.
   PubMed Web of Science ®Google Scholar
• Cline, T. W., and B. J. Meyer. 1996. Vive la difference: males vs females in flies vs worms. Annu. Rev. Genet. 30:637–702.
   PubMed Web of Science ®Google Scholar
• Cronmiller, C., and C. A. Cummings. 1993. The daughterless gene product in Drosophila is a nuclear protein that is broadly expressed throughout the organism during development. Mech. Dev. 42:159–169.
   PubMedGoogle Scholar
• Dang, C. V., C. Dolde, M. L. Gillison, and G. J. Kato. 1992. Discrimination between related DNA sites by a single amino acid residue of Myc-related basic-helix-loop-helix proteins. Proc. Natl. Acad. Sci. USA 89:599–602.
   PubMedGoogle Scholar
• Deshpande, G., J. Stukey, and P. Schedl. 1995. scute (sis-b) function in Drosophila sex determination. Mol. Cell. Biol. 15:4430–4440.
   PubMedGoogle Scholar
• Erickson, J. W., and T. W. Cline. 1993. A bZIP protein, SISTERLESS-A, collaborates with bHLH transcription factors early in Drosophila development to determine sex. Genes Dev. 7:1688–1702.
   PubMed Web of Science ®Google Scholar
• Erickson, J. W., and T. W. Cline. 1998. Key aspects of the primary sex determination mechanism are conserved across the genus Drosophila. Development 125:3259–3268.
   PubMedGoogle Scholar
• Erickson, J. W., and T. W. Cline. 1991. Molecular nature of the Drosophila sex determination signal and its link to neurogenesis. Science 251:1071–1074.
   PubMedGoogle Scholar
• Estes, P. A., L. N. Keyes, and P. Schedl. 1995. Multiple response elements in the Sex-lethal early promoter ensure its female-specific expression pattern. Mol. Cell. Biol. 15:904–917.
   PubMedGoogle Scholar
• Falvo, J. V., B. S. Parekh, C. H. Lin, E. Fraenkel, and T. Maniatis. 2000. Assembly of a functional beta interferon enhanceosome is dependent on ATF-2-c-jun heterodimer orientation. Mol. Cell. Biol. 20:4814–4825.
   PubMedGoogle Scholar
• Gonzalez-Crespo, S., and M. Levine. 1993. Interactions between dorsal and helix-loop-helix proteins initiate the differentiation of the embryonic mesoderm and neuroectoderm in Drosophila. Genes Dev. 7:1703–1713.
   PubMedGoogle Scholar
• Grandori, C., and R. N. Eisenman. 1997. Myc target genes. Trends Biochem. Sci. 22:177–181.
   PubMed Web of Science ®Google Scholar
• Grandori, C., J. Mac, F. Siebelt, D. E. Ayer, and R. N. Eisenman. 1996. Myc-Max heterodimers activate a DEAD box gene and interact with multiple E box-related sites in vivo. EMBO J. 15:4344–4357.
   PubMed Web of Science ®Google Scholar
• Grosschedl, R.. 1995. Higher-order nucleoprotein complexes in transcription: analogies with site-specific recombination. Curr. Opin. Cell Biol. 7:362–370.
   PubMedGoogle Scholar
• Han, K., M. S. Levine, and J. L. Manley. 1989. Synergistic activation and repression of transcription by Drosophila homeobox proteins. Cell 56:573–583.
   PubMedGoogle Scholar
• Hoshijima, K., A. Kohyama, I. Watakabe, K. Inoue, H. Sakamoto, and Y. Shimura. 1995. Transcriptional regulation of the Sex-lethal gene by helix-loop-helix proteins. Nucleic Acids Res. 23:3441–3448.
   PubMedGoogle Scholar
• Ip, Y. T., R. E. Park, D. Kosman, E. Bier, and M. Levine. 1992. The dorsal gradient morphogen regulates stripes of rhomboid expression in the presumptive neuroectoderm of the Drosophila embryo. Genes Dev. 6:1728–1739.
   PubMed Web of Science ®Google Scholar
• Ip, Y. T., R. E. Park, D. Kosman, K. Yazdanbakhsh, and M. Levine. 1992. dorsal-twist interactions establish snail expression in the presumptive mesoderm of the Drosophila embryo. Genes Dev. 6:1518–1530.
   PubMed Web of Science ®Google Scholar
• Jan, Y. N., and L. Y. Jan. 1993. Functional gene cassettes in development. Proc. Natl. Acad. Sci. USA 90:8305–8307.
   PubMed Web of Science ®Google Scholar
• Jan, Y. N., and L. Y. Jan. 1993. HLH proteins, fly neurogenesis, and vertebrate myogenesis. Cell 75:827–830.
   PubMedGoogle Scholar
• Jiang, J., and M. Levine. 1993. Binding affinities and cooperative interactions with bHLH activators delimit threshold responses to the dorsal gradient morphogen. Cell 72:741–752.
   PubMedGoogle Scholar
• Jinks, T. M., A. D. Polydorides, G. Calhoun, and P. Schedl. 2000. The JAK/STAT signalling pathway is required for the initial choice of sexual identity in Drosophila melanogaster. Mol. Cell 5:581–587.
   PubMedGoogle Scholar
• Keyes, L. N., T. W. Cline, and P. Schedl. 1992. The primary sex determination signal of Drosophila acts at the level of transcription. Cell 68:933–943 (Corrigendum, 69:following 572).
   PubMedGoogle Scholar
• Kramer, S. G., T. M. Jinks, P. Schedl, and J. P. Gergen. 1999. Direct activation of Sex-lethal transcription by the Drosophila runt protein. Development 126:191–200.
   PubMedGoogle Scholar
• Littlewood, T., and G. I. Evan. 1998. Helix-loop-helix transcription factors. Oxford University Press, Oxford, United Kingdom
   Google Scholar
• Martinez, C., J. Modolell, and J. Garrell. 1993. Regulation of the proneural gene achaete by helix-loop-helix proteins. Mol. Cell. Biol. 13:3514–3521.
   PubMedGoogle Scholar
• Massari, M. E., and C. Murre. 2000. Helix-loop-helix proteins: regulators of transcription in eucaryotic organisms. Mol. Cell. Biol. 20:429–440.
   PubMed Web of Science ®Google Scholar
• Ohsako, S., J. Hyer, G. Panganiban, I. Oliver, and M. Caudy. 1994. Hairy functions as a DNA-binding helix-loop-helix repressor of Drosophila sensory organ formation. Genes Dev. 8:2743–2755.
   PubMedGoogle Scholar
• Pan, D., S. A. Valentine, and A. J. Courey. 1994. The bipartite D. melanogaster twist promoter is reorganized in D. virilis. Mech. Dev. 46:41–53.
   PubMedGoogle Scholar
• Parkhurst, S. M., D. Bopp, and D. Ish-Horowicz. 1990. X:A ratio, the primary sex-determining signal in Drosophila, is transduced by helix-loop-helix proteins. Cell 63:1179–1191 (Corrigendum, 64:following 1046).
   PubMedGoogle Scholar
• Paroush, Z., R. L. Finley Jr., T. Kidd, S. M. Wainwright, P. W. Ingham, R. Brent, and D. Ish-Horowicz. 1994. Groucho is required for Drosophila neurogenesis, segmentation, and sex determination and interacts directly with hairy-related bHLH proteins. Cell 79:805–815.
   PubMed Web of Science ®Google Scholar
• Penalva, L. O., H. Sakamoto, A. Navarro-Sabate, E. Sakashita, B. Granadino, C. Segarra, and L. Sanchez. 1996. Regulation of the gene Sex-lethal: a comparative analysis of Drosophila melanogaster and Drosophila subobscura. Genetics 144:1653–1664.
   PubMedGoogle Scholar
• Pultz, M. A., and B. S. Baker. 1995. The dual role of hermaphrodite in the Drosophila sex determination regulatory hierarchy. Development 121:99–111.
   PubMedGoogle Scholar
• Sefton, L., J. R. Timmer, Y. Zhang, F. Beranger, and T. W. Cline. 2000. An extracellular activator of the Drosophila JAK/STAT pathway is a sex-determination signal element. Nature 405:970–973.
   PubMedGoogle Scholar
• Swanson, H. I., W. K. Chan, and C. A. Bradfield. 1995. DNA binding specificities and pairing rules of the Ah receptor, ARNT, and SIM proteins. J. Biol. Chem. 270:26292–26302.
   PubMed Web of Science ®Google Scholar
• Szymanski, P., and M. Levine. 1995. Multiple modes of dorsal-bHLH transcriptional synergy in the Drosophila embryo. EMBO J. 14:2229–2238.
   PubMedGoogle Scholar
• Thanos, D.. 1996. Mechanisms of transcriptional synergism of eukaryotic genes. The interferon-beta paradigm. Hypertension 27:1025–1029.
   PubMedGoogle Scholar
• Torres, M., and L. Sanchez. 1989. The scute (T4) gene acts as a numerator element of the X:A signal that determines the state of activity of Sex-lethal in Drosophila. EMBO J. 8:3079–3086.
   PubMed Web of Science ®Google Scholar
• Van Doren, M., A. M. Bailey, J. Esnayra, K. Ede, and J. W. Posakony. 1994. Negative regulation of proneural gene activity: hairy is a direct transcriptional repressor of achaete. Genes Dev. 8:2729–2742.
   PubMedGoogle Scholar
• Verma-Kurvari, S., T. Savage, K. Gowan, and J. E. Johnson. 1996. Lineage-specific regulation of the neural differentiation gene MASH1. Dev. Biol. 180:605–617.
   PubMedGoogle Scholar
• Younger-Shepherd, S., H. Vaessin, E. Bier, L. Y. Jan, and Y. N. Jan. 1992. deadpan, an essential pan-neural gene encoding an HLH protein, acts as a denominator in Drosophila sex determination. Cell 70:911–922.
   PubMedGoogle Scholar