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Gene Review

ey  -  eyeless

Drosophila melanogaster

Synonyms: CG1464, DPax-6, Dmel\CG1464, EY, EYEL, ...
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Disease relevance of ey

  • The finding that ey of Drosophila, Small eye of the mouse, and human Aniridia are encoded by homologous genes suggests that eye morphogenesis is under similar genetic control in both vertebrates and insects, in spite of the large differences in eye morphology and mode of development [1].

Psychiatry related information on ey

  • Furthermore, severe defects in adult brain structures essential for vision, olfaction, and for the coordination of locomotion are provoked by two newly isolated mutations of Pax-6/eyeless that result in truncated proteins [2].
  • In addition, so1 (eyeless) mutant flies show the presence of a single evening locomotor activity peak during the whole circadian day, suggesting that in wild type flies the morning and evening activity peaks may be under separate control [3].
  • Brief heating of the oocytes and larvae of eyeless mutants during the critical periods of faceted-eye development caused an increase in the thermal sensitivity of the eye rudiments, which led to an increase in the number of one-eyed individuals among the flies that hatched [4].

High impact information on ey

  • These results imply that ey does not induce the entire eye morphogenetic program but rather modifies ato-dependent neuronal development [5].
  • The eyeless, dachshund, and eyes absent genes encode conserved, nuclear proteins that are essential for eye development in Drosophila [6].
  • These genes have been proposed to act with eyeless (Pax6) to regulate eye development in vertebrates and invertebrates. so encodes a highly diverged homeobox transcription factor and eya encodes a novel nuclear protein [7].
  • Pax-6 is a member of the Pax gene class and encodes a protein containing a paired domain and a homeodomain [8].
  • The molecular characterization of Pax-6 genes from species of different animal phyla and the analysis of Pax-6 function in the developing eyes and central nervous system of vertebrates, Drosophila melanogaster, and Caenorhabditis elegans suggest that Pax-6 homologues share conserved functions [8].

Biological context of ey

  • They are all conserved between vertebrates and insects and they interact in a combinatorial and hierarchical network to regulate each other expression. so has been shown to be directly regulated by ey through an eye-specific enhancer (so10) [9].
  • We further studied the regulation of this element and found that both Drosophila Pax6 proteins namely EY and TOY bind and positively regulate so10 expression through different binding sites [9].
  • Thus, eya-1 appears to play a partially redundant role with pax-6 during C. elegans embryogenesis [10].
  • However, this genetic network has been used in a novel developmental context, myogenesis rather than eye development, and has been expanded to include gene family members that are not directly homologous, for example Pax3 instead of Pax6 [11].
  • Using the emerging consensus sequence for SO-DNA binding we performed a genome-wide search and have thereby been able to identify eyeless as well as the signalling gene hedgehog as putative targets of so [12].

Anatomical context of ey

  • Mutations of ey completely disrupted the MB neuropils, and a null mutation of dac resulted in marked disruption and aberrant axonal projections [13].
  • Genetic control of development of the mushroom bodies, the associative learning centers in the Drosophila brain, by the eyeless, twin of eyeless, and Dachshund genes [13].
  • The tiny rudiments of the optic lobes in eyeless double mutants still contain tangential neurons of the medulla and of the lobula complex [14].
  • Therefore, in addition to performing a global photoreceptor-specific function, RCSI also appears to mediate the combined action of Pax6 and other factors and to contribute to rh regulation in subsets of photoreceptors [15].
  • It is required for development of the central nervous system, and is mutated in human aniridia, mouse and rat small eye and Drosophila eyeless [16].

Physical interactions of ey

  • To overcome the lack of high-quality consensus binding site sequences, phylogenetic shadowing of known Ey binding sites in sine oculis (so) was used to construct a position weight matrix (PWM) of the Ey protein [17].

Regulatory relationships of ey


Other interactions of ey


Analytical, diagnostic and therapeutic context of ey

  • 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 [17].
  • We used two different Drosophila full-genome DNA microarrays to compare gene expression in wild-type leg discs versus leg discs where eyeless, one of the two Drosophila Pax-6 genes, was ectopically expressed [24].
  • The results of protein, phospholipid and enzyme analyses were corroborated by analyses by 'genetic dissection' using an eyeless mutant line [25].
  • On a Western blot, the antibody recognizes a 41 kDa protein that is present in the heads of yellow white flies, but not in the heads of eyeless mutant flies, eyes absent [26].


  1. Homology of the eyeless gene of Drosophila to the Small eye gene in mice and Aniridia in humans. Quiring, R., Walldorf, U., Kloter, U., Gehring, W.J. Science (1994) [Pubmed]
  2. Drosophila Pax-6/eyeless is essential for normal adult brain structure and function. Callaerts, P., Leng, S., Clements, J., Benassayag, C., Cribbs, D., Kang, Y.Y., Walldorf, U., Fischbach, K.F., Strauss, R. J. Neurobiol. (2001) [Pubmed]
  3. Extra ocular photic entrainment in Drosophila melanogaster. Zordan, M., Osterwalder, N., Rosato, E., Costa, R. J. Neurogenet. (2001) [Pubmed]
  4. Adaptive changes in degree of eye reduction in eyeless mutants of Drosophila melanogaster after thermal shock during the critical periods of faceted eye development. Svetlov, P.G., Korsakova, G.F. The Soviet journal of developmental biology. (1975) [Pubmed]
  5. A conserved developmental program for sensory organ formation in Drosophila melanogaster. Niwa, N., Hiromi, Y., Okabe, M. Nat. Genet. (2004) [Pubmed]
  6. Dachshund and eyes absent proteins form a complex and function synergistically to induce ectopic eye development in Drosophila. Chen, R., Amoui, M., Zhang, Z., Mardon, G. Cell (1997) [Pubmed]
  7. The eye-specification proteins So and Eya form a complex and regulate multiple steps in Drosophila eye development. Pignoni, F., Hu, B., Zavitz, K.H., Xiao, J., Garrity, P.A., Zipursky, S.L. Cell (1997) [Pubmed]
  8. PAX-6 in development and evolution. Callaerts, P., Halder, G., Gehring, W.J. Annu. Rev. Neurosci. (1997) [Pubmed]
  9. Differential interactions of eyeless and twin of eyeless with the sine oculis enhancer. Punzo, C., Seimiya, M., Flister, S., Gehring, W.J., Plaza, S. Development (2002) [Pubmed]
  10. The C. elegans eyes absent ortholog EYA-1 is required for tissue differentiation and plays partially redundant roles with PAX-6. Furuya, M., Qadota, H., Chisholm, A.D., Sugimoto, A. Dev. Biol. (2005) [Pubmed]
  11. Synergistic regulation of vertebrate muscle development by Dach2, Eya2, and Six1, homologs of genes required for Drosophila eye formation. Heanue, T.A., Reshef, R., Davis, R.J., Mardon, G., Oliver, G., Tomarev, S., Lassar, A.B., Tabin, C.J. Genes Dev. (1999) [Pubmed]
  12. Identification of functional sine oculis motifs in the autoregulatory element of its own gene, in the eyeless enhancer and in the signalling gene hedgehog. Pauli, T., Seimiya, M., Blanco, J., Gehring, W.J. Development (2005) [Pubmed]
  13. Genetic control of development of the mushroom bodies, the associative learning centers in the Drosophila brain, by the eyeless, twin of eyeless, and Dachshund genes. Kurusu, M., Nagao, T., Walldorf, U., Flister, S., Gehring, W.J., Furukubo-Tokunaga, K. Proc. Natl. Acad. Sci. U.S.A. (2000) [Pubmed]
  14. Cell degeneration in the developing optic lobes of the sine oculis and small-optic-lobes mutants of Drosophila melanogaster. Fischbach, K.F., Technau, G. Dev. Biol. (1984) [Pubmed]
  15. A conserved regulatory element present in all Drosophila rhodopsin genes mediates Pax6 functions and participates in the fine-tuning of cell-specific expression. Papatsenko, D., Nazina, A., Desplan, C. Mech. Dev. (2001) [Pubmed]
  16. Specification of sense-organ identity by a Caenorhabditis elegans Pax-6 homologue. Zhang, Y., Emmons, S.W. Nature (1995) [Pubmed]
  17. 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]
  18. The Drosophila homeobox gene optix is capable of inducing ectopic eyes by an eyeless-independent mechanism. Seimiya, M., Gehring, W.J. Development (2000) [Pubmed]
  19. Direct regulation of rhodopsin 1 by Pax-6/eyeless in Drosophila: evidence for a conserved function in photoreceptors. Sheng, G., Thouvenot, E., Schmucker, D., Wilson, D.S., Desplan, C. Genes Dev. (1997) [Pubmed]
  20. Notch signaling and the determination of appendage identity. Kurata, S., Go, M.J., Artavanis-Tsakonas, S., Gehring, W.J. Proc. Natl. Acad. Sci. U.S.A. (2000) [Pubmed]
  21. Combinatorial control of Drosophila eye development by eyeless, homothorax, and teashirt. Bessa, J., Gebelein, B., Pichaud, F., Casares, F., Mann, R.S. Genes Dev. (2002) [Pubmed]
  22. Evidence for a direct functional antagonism of the selector genes proboscipedia and eyeless in Drosophila head development. Benassayag, C., Plaza, S., Callaerts, P., Clements, J., Romeo, Y., Gehring, W.J., Cribbs, D.L. Development (2003) [Pubmed]
  23. Expression of evolutionarily conserved eye specification genes during Drosophila embryogenesis. Kumar, J.P., Moses, K. Dev. Genes Evol. (2001) [Pubmed]
  24. Analysis of the eye developmental pathway in Drosophila using DNA microarrays. Michaut, L., Flister, S., Neeb, M., White, K.P., Certa, U., Gehring, W.J. Proc. Natl. Acad. Sci. U.S.A. (2003) [Pubmed]
  25. Quantitative tissue isolation from Drosophila freeze-dried in acetone. Fujita, S.C., Inoue, H., Yoshioka, T., Hotta, Y. Biochem. J. (1987) [Pubmed]
  26. Identification of a retinal protein in Drosophila with antibody to the alpha subunit of bovine brain G(o) protein. Garen-Fazio, S., Neer, E.J., Schmidt, C.J. J. Comp. Neurol. (1991) [Pubmed]
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