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Gene Review

ato  -  atonal

Drosophila melanogaster

Synonyms: ATO, At, Ato, CG7508, Dmel\CG7508, ...
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Disease relevance of ato

  • Math5, a mouse homolog of the Drosophila proneural bHLH transcription factor Atonal, is essential in the developing retina to establish retinal progenitor cell competence for a ganglion cell fate [1].

High impact information on ato

  • These results imply that ey does not induce the entire eye morphogenetic program but rather modifies ato-dependent neuronal development [2].
  • In D. melanogaster, a common proneural gene called atonal (ato) functions in the initial process of development of a number of segment-specific organs, including the compound eye, the auditory organ and the stretch receptor, suggesting that these organs share an evolutionary origin [2].
  • Like the ASC products, ato protein contains a basic-helix-loop-helix region and heterodimerizes with daughterless protein to bind to E boxes [3].
  • atonal is a proneural gene that directs chordotonal organ formation in the Drosophila peripheral nervous system [3].
  • The establishment and subsequent patterning of Atonal expression requires activity of the signalling transmembrane receptor Notch [4].

Biological context of ato

  • M8SD elicits neural hypoplasia in eye discs, elicits loss of phase-shifted Atonal-positive cells, i.e. the 'founding' R8 photoreceptors, and consequently leads to apoptosis [5].
  • We used ro(Dom) as a sensitive genetic background in which to identify mutations that affect hh signal transduction or regulation of ato expression [6].
  • The time required for N signaling to inhibit ato expression was at most 90 min, but changes in the Dl protein distribution in mutant genotypes arose more slowly [7].
  • The proneural proteins Atonal and Scute regulate neural target genes through different E-box binding sites [8].
  • First, tandem repeats of 20-bp sequences containing E(Ato) and E(Sc) sites are sufficient to confer Atonal- and Scute-specific expression patterns, respectively, on a reporter gene in vivo [8].

Anatomical context of ato

  • Expression of tll is normally confined to the optic lobe primordium, whereas ato appears in a subset of Bolwig's organ cells that we call Bolwig's organ founders [9].
  • Initiation of Drosophila peripheral nervous system (PNS) development requires the achaete-scute complex (AS-C) and the atonal (ato) genes [10].
  • atonal regulates neurite arborization but does not act as a proneural gene in the Drosophila brain [11].
  • Sensory neurons of the Atonal lineage pioneer the formation of glomeruli within the adult Drosophila olfactory lobe [12].
  • Previously, we showed that targeted deletion of the atonal (ato) homologue math5 blocked the differentiation of most retinal ganglion cells (RGCs), revealing an essential role for math5 in RGC differentiation [13].

Enzymatic interactions of ato

  • R8s in turn appear to signal through Rhomboid and Vein to create a patterned array of 'proneural clusters' which contain high levels of phosphorylated ERKA and the bHLH protein Atonal [14].

Regulatory relationships of ato

  • N and Dl were required to repress ato in the vicinity of R8 cells, whereas sca had effects over several cell diameters [15].

Other interactions of ato

  • E(spl)D, an allele of m8, encodes a truncated protein known as M8*, which, unlike wild type M8, displays exacerbated antagonism of Atonal via direct protein-protein interactions [5].
  • Neither Atonal expression nor Bolwig's organ formation occurred in the absence of hedgehog, eyes absent or sine oculis activity [16].
  • Normal ato or da activity is required for maintenance of MAPK activation [17].
  • The b domains of ato and sc differ in seven residues [18].
  • This supports a model of inhibitory interactions between proneural genes, whereby ato-like genes (amos and ato) must suppress sensory bristle fate as well as promote alternative sense organ subtypes [19].

Analytical, diagnostic and therapeutic context of ato

  • In parallel, microarray analysis of wild-type versus ectopic ey-expressing tissue, followed by microarray-based epistasis experiments in an atonal (ato) mutant background, identified 188 genes induced by ey [20].


  1. A gene network downstream of transcription factor Math5 regulates retinal progenitor cell competence and ganglion cell fate. Mu, X., Fu, X., Sun, H., Beremand, P.D., Thomas, T.L., Klein, W.H. Dev. Biol. (2005) [Pubmed]
  2. A conserved developmental program for sensory organ formation in Drosophila melanogaster. Niwa, N., Hiromi, Y., Okabe, M. Nat. Genet. (2004) [Pubmed]
  3. atonal is a proneural gene that directs chordotonal organ formation in the Drosophila peripheral nervous system. Jarman, A.P., Grau, Y., Jan, L.Y., Jan, Y.N. Cell (1993) [Pubmed]
  4. Scabrous complexes with Notch to mediate boundary formation. Powell, P.A., Wesley, C., Spencer, S., Cagan, R.L. Nature (2001) [Pubmed]
  5. Drosophila CK2 regulates eye morphogenesis via phosphorylation of E(spl)M8. Karandikar, U.C., Trott, R.L., Yin, J., Bishop, C.P., Bidwai, A.P. Mech. Dev. (2004) [Pubmed]
  6. A screen for dominant modifiers of ro(Dom), a mutation that disrupts morphogenetic furrow progression in Drosophila, identifies groucho and hairless as regulators of atonal expression. Chanut, F., Luk, A., Heberlein, U. Genetics (2000) [Pubmed]
  7. The R8-photoreceptor equivalence group in Drosophila: fate choice precedes regulated Delta transcription and is independent of Notch gene dose. Baker, N.E., Yu, S.Y. Mech. Dev. (1998) [Pubmed]
  8. The proneural proteins Atonal and Scute regulate neural target genes through different E-box binding sites. Powell, L.M., Zur Lage, P.I., Prentice, D.R., Senthinathan, B., Jarman, A.P. Mol. Cell. Biol. (2004) [Pubmed]
  9. The control of cell fate in the embryonic visual system by atonal, tailless and EGFR signaling. Daniel, A., Dumstrei, K., Lengyel, J.A., Hartenstein, V. Development (1999) [Pubmed]
  10. daughterless is required for Drosophila photoreceptor cell determination, eye morphogenesis, and cell cycle progression. Brown, N.L., Paddock, S.W., Sattler, C.A., Cronmiller, C., Thomas, B.J., Carroll, S.B. Dev. Biol. (1996) [Pubmed]
  11. atonal regulates neurite arborization but does not act as a proneural gene in the Drosophila brain. Hassan, B.A., Bermingham, N.A., He, Y., Sun, Y., Jan, Y.N., Zoghbi, H.Y., Bellen, H.J. Neuron (2000) [Pubmed]
  12. Sensory neurons of the Atonal lineage pioneer the formation of glomeruli within the adult Drosophila olfactory lobe. Jhaveri, D., Rodrigues, V. Development (2002) [Pubmed]
  13. Math5 determines the competence state of retinal ganglion cell progenitors. Yang, Z., Ding, K., Pan, L., Deng, M., Gan, L. Dev. Biol. (2003) [Pubmed]
  14. Regulation of EGF receptor signaling establishes pattern across the developing Drosophila retina. Spencer, S.A., Powell, P.A., Miller, D.T., Cagan, R.L. Development (1998) [Pubmed]
  15. The scabrous gene encodes a secreted glycoprotein dimer and regulates proneural development in Drosophila eyes. Lee, E.C., Hu, X., Yu, S.Y., Baker, N.E. Mol. Cell. Biol. (1996) [Pubmed]
  16. Transcriptional regulation of atonal required for Drosophila larval eye development by concerted action of eyes absent, sine oculis and hedgehog signaling independent of fused kinase and cubitus interruptus. Suzuki, T., Saigo, K. Development (2000) [Pubmed]
  17. Negative regulation of atonal in proneural cluster formation of Drosophila R8 photoreceptors. Chen, C.K., Chien, C.T. Proc. Natl. Acad. Sci. U.S.A. (1999) [Pubmed]
  18. Neuronal type information encoded in the basic-helix-loop-helix domain of proneural genes. Chien, C.T., Hsiao, C.D., Jan, L.Y., Jan, Y.N. Proc. Natl. Acad. Sci. U.S.A. (1996) [Pubmed]
  19. The Drosophila proneural gene amos promotes olfactory sensillum formation and suppresses bristle formation. zur Lage, P.I., Prentice, D.R., Holohan, E.E., Jarman, A.P. Development (2003) [Pubmed]
  20. Genome-wide identification of direct targets of the Drosophila retinal determination protein Eyeless. Ostrin, E.J., Li, Y., Hoffman, K., Liu, J., Wang, K., Zhang, L., Mardon, G., Chen, R. Genome Res. (2006) [Pubmed]
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