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Gene Review

emc  -  extra macrochaetae

Drosophila melanogaster

Synonyms: 0094/26, 0203/10, 0587/01, 0977/09, Ach, ...
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Psychiatry related information on emc

  • This indicates that the zebrafish gene described here is a genuine member of the Id family, and suggests that it may serve a function similar to that of the Drosophila gene emc and mammalian Ids during development [1].

High impact information on emc

  • While neither h nor emc is required for photoreceptor cell determination, in emc-h-clones the morphogenetic furrow and differentiated eye field advance up to eight ommatidial rows ahead of adjacent wild-type tissue [2].
  • We propose a model in which the emc protein negatively regulates sensory organ determination by forming heterodimers with the HLH proteins encoded by the AS-C and/or daughterless, thereby altering or interfering with their activity [3].
  • The emc protein shares the dimerization domain of other HLH proteins but lacks their DNA binding motif [3].
  • The tam mutation reduces the transcription of a repressor gene, extramacrochaetae, and causes enlargement of a proneural cluster where supernumerary precursor cells emerge, resulting in extra mechanosensory organs in the fly [4].
  • Spatial regulation of proneural gene activity: auto- and cross-activation of achaete is antagonized by extramacrochaetae [5].

Biological context of emc

  • As currently thought, this seems accomplished by heterodimerization via the HLH domain, because an amino acid substitution in this region abolishes the emc antagonistic effect both in vitro and in vivo [6].
  • We show that, during embryonic development, the extramacrochaetae gene is expressed in complex patterns that comprise derivatives of the three embryonic layers [7].
  • Neural fate specification in Drosophila is promoted by the products of the proneural genes, such as those of the achaete-scute complex, and antagonized by the products of the Enhancer of split [E(spl)] complex, hairy, and extramacrochaetae [8].
  • Mutations at some candidate loci (bb, emc, h, Dl, Hairless) showed strong interactions with selected chromosomes, whereas others interacted weakly (ASC, abd, Scr) or not at all (N, mab, E(spl)) [9].
  • In addition, cell proliferation behaviour of emc mutant cells varies depending on the mutant background [10].

Anatomical context of emc


Physical interactions of emc


Regulatory relationships of emc

  • We propose that at least during vein differentiation and wing margin formation, extramacrochaetae is regulated by Notch and collaborates with other Notch-downstream genes such as Enhancer of split-m(beta) [15].
  • The known proneural antagonists either titrate these proteins by forming inactive complexes (extramacrochaetae) or repress achaete/scute expression at specific sites (i.e., hairy) [16].

Other interactions of emc

  • When one of these spatial restrictions is eliminated, by ubiquitously expressing ac-sc, SMCs still emerge within minima of emc [17].
  • Within the ectodermal layer, its level is neither elevated (as in the case of AS-C genes) nor reduced (as in the case of emc product) in the proneural cluster [18].
  • These results suggest that negative regulation by emc may be a common feature of developmental processes involving da, the AS-C and possibly other helix-loop-helix proteins [19].
  • Synergistic interactions between pyd and Notch, Delta and extramacrochaetae mutations support this model [20].
  • Thus, loss-of-function conditions for ed give rise to the development of extra macrochaetae near the extant ones and increase the density of microchaetae [21].

Analytical, diagnostic and therapeutic context of emc

  • Our results strongly support the model of negative regulation of emc on ac and sc transcription through titration of their products [6].


  1. A zebrafish Id homologue and its pattern of expression during embryogenesis. Sawai, S., Campos-Ortega, J.A. Mech. Dev. (1997) [Pubmed]
  2. Hairy and emc negatively regulate morphogenetic furrow progression in the Drosophila eye. Brown, N.L., Sattler, C.A., Paddock, S.W., Carroll, S.B. Cell (1995) [Pubmed]
  3. extramacrochaetae, a negative regulator of sensory organ development in Drosophila, defines a new class of helix-loop-helix proteins. Ellis, H.M., Spann, D.R., Posakony, J.W. Cell (1990) [Pubmed]
  4. The Drosophila tamou gene, a component of the activating pathway of extramacrochaetae expression, encodes a protein homologous to mammalian cell-cell junction-associated protein ZO-1. Takahisa, M., Togashi, S., Suzuki, T., Kobayashi, M., Murayama, A., Kondo, K., Miyake, T., Ueda, R. Genes Dev. (1996) [Pubmed]
  5. Spatial regulation of proneural gene activity: auto- and cross-activation of achaete is antagonized by extramacrochaetae. Van Doren, M., Powell, P.A., Pasternak, D., Singson, A., Posakony, J.W. Genes Dev. (1992) [Pubmed]
  6. Regulation of the proneural gene achaete by helix-loop-helix proteins. Martínez, C., Modolell, J., Garrell, J. Mol. Cell. Biol. (1993) [Pubmed]
  7. The helix-loop-helix extramacrochaetae protein is required for proper specification of many cell types in the Drosophila embryo. Cubas, P., Modolell, J., Ruiz-Gómez, M. Development (1994) [Pubmed]
  8. A network of interacting transcriptional regulators involved in Drosophila neural fate specification revealed by the yeast two-hybrid system. Alifragis, P., Poortinga, G., Parkhurst, S.M., Delidakis, C. Proc. Natl. Acad. Sci. U.S.A. (1997) [Pubmed]
  9. Genetic interactions between naturally occurring alleles at quantitative trait loci and mutant alleles at candidate loci affecting bristle number in Drosophila melanogaster. Long, A.D., Mullaney, S.L., Mackay, T.F., Langley, C.H. Genetics (1996) [Pubmed]
  10. Dual role of extramacrochaetae in cell proliferation and cell differentiation during wing morphogenesis in Drosophila. Baonza, A., García-Bellido, A. Mech. Dev. (1999) [Pubmed]
  11. emc has a role in dorsal appendage fate formation in Drosophila oogenesis. Papadia, S., Tzolovsky, G., Zhao, D., Leaper, K., Clyde, D., Taylor, P., Asscher, E., Kirk, G., Bownes, M. Mech. Dev. (2005) [Pubmed]
  12. A gain-of-function screen for genes that affect the development of the Drosophila adult external sensory organ. Abdelilah-Seyfried, S., Chan, Y.M., Zeng, C., Justice, N.J., Younger-Shepherd, S., Sharp, L.E., Barbel, S., Meadows, S.A., Jan, L.Y., Jan, Y.N. Genetics (2000) [Pubmed]
  13. The Drosophila extramacrochaetae protein antagonizes sequence-specific DNA binding by daughterless/achaete-scute protein complexes. Van Doren, M., Ellis, H.M., Posakony, J.W. Development (1991) [Pubmed]
  14. Down regulation of extramacrochaetae mRNA by a Drosophila neural RNA binding protein Rbp9 which is homologous to human Hu proteins. Park, S.J., Yang, E.S., Kim-Ha, J., Kim, Y.J. Nucleic Acids Res. (1998) [Pubmed]
  15. Relationships between extramacrochaetae and Notch signalling in Drosophila wing development. Baonza, A., de Celis, J.F., García-Bellido, A. Development (2000) [Pubmed]
  16. Patterning of the adult peripheral nervous system of Drosophila. Modolell, J. Perspectives on developmental neurobiology. (1997) [Pubmed]
  17. The extramacrochaetae gene provides information for sensory organ patterning. Cubas, P., Modolell, J. EMBO J. (1992) [Pubmed]
  18. daughterless is essential for neuronal precursor differentiation but not for initiation of neuronal precursor formation in Drosophila embryo. Vaessin, H., Brand, M., Jan, L.Y., Jan, Y.N. Development (1994) [Pubmed]
  19. Embryonic expression and function of the Drosophila helix-loop-helix gene, extramacrochaetae. Ellis, H.M. Mech. Dev. (1994) [Pubmed]
  20. Polychaetoid is required to restrict segregation of sensory organ precursors from proneural clusters in Drosophila. Chen, C.M., Freedman, J.A., Bettler, D.R., Manning, S.D., Giep, S.N., Steiner, J., Ellis, H.M. Mech. Dev. (1996) [Pubmed]
  21. Echinoid synergizes with the Notch signaling pathway in Drosophila mesothorax bristle patterning. Escudero, L.M., Wei, S.Y., Chiu, W.H., Modolell, J., Hsu, J.C. Development (2003) [Pubmed]
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