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Gene Review

so  -  sine oculis

Drosophila melanogaster

Synonyms: CG11121, Dmel\CG11121, Drl, Mdu, Protein sine oculis, ...
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Disease relevance of so


High impact information on so

  • We demonstrate that So and Eya (1) regulate common steps in eye development including cell proliferation, patterning, and neuronal development; (2) synergize in inducing ectopic eyes; and (3) interact in yeast and in vitro through evolutionarily conserved domains [3].
  • The homeobox gene sine oculis (so) is essential for visual system formation in Drosophila [4].
  • Moreover, Eya2 can also synergize with Six1 (a vertebrate homolog of the Drosophila gene sine oculis) to regulate myogenesis [5].
  • Molecular and developmental analyses reveals that the sine oculis (so) locus encodes a homeodomain-containing protein that is expressed and required in the unpatterned epithelium prior to morphogenesis [6].
  • We find that the conserved C-terminal EYA domain negatively regulates EYA transactivation potential, and that GROUCHO-SINE OCULIS (SO) interactions provide another mechanism for negative regulation of EYA-SO target genes [7].

Biological context of so


Anatomical context of so

  • Regulatory genes that are required for eye/optic lobe fate, including sine oculis (so) and eyes absent (eya), are turned on in their respective domains by Dpp [13].
  • A somatic role for eyes absent (eya) and sine oculis (so) in Drosophila spermatocyte development [14].
  • In this report, we find that the transcription factors eyes absent (eya) and sine oculis (so), previously shown to play major roles during eye development [Cell 91 (1997), 881] are each required in the somatic cyst cells of the testis for proper Drosophila spermatocyte development. eya mutant testes exhibit degenerating young spermatocytes [14].
  • In this study, we report the identification of two novel factors that participate in specific partnerships with Sine oculis and Optix during photoreceptor neurons formation and in eye progenitor cells [15].
  • Six1 and Six4 are the mammalian homologs of Drosophila sine oculis, and they are coexpressed in the nephrogenic mesenchyme [16].

Associations of so with chemical compounds

  • We provide evidence that Sine oculis and Glass are the two major activators of Lz expression during eye development [17].
  • Our findings are as follows: (a) Large amounts of histamine are synthesized by wild-type heads, whereas heads from the eye-deficient mutants, eyes absent and sine oculis, show reduced histamine synthesis [18].

Regulatory relationships of so

  • Previous work in our laboratory identified an eye/ocellus specific enhancer of the sine oculis gene that is directly regulated by eyeless and twin of eyeless [19].
  • Podocoryne Twist is expressed in the early embryo until the myoepithelial cells of the larva differentiate and then again during medusa development [20].

Other interactions of so

  • Eya and So, which are induced by Dpp, are epistatic to the Hh signal [13].
  • 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 [21].
  • Neither Atonal expression nor Bolwig's organ formation occurred in the absence of hedgehog, eyes absent or sine oculis activity [21].
  • These targets, which share homology with the fly genes sine oculis, eyes absent and dachshund, exist in mice and humans as the Six, Eya and Dach gene families [22].
  • Partner specificity is essential for proper function of the SIX-type homeodomain proteins Sine oculis and Optix during fly eye development [23].


  1. The homeobox gene Otx of the jellyfish Podocoryne carnea: role of a head gene in striated muscle and evolution. Müller, P., Yanze, N., Schmid, V., Spring, J. Dev. Biol. (1999) [Pubmed]
  2. Functional analysis of the homeodomain protein SIX5. Harris, S.E., Winchester, C.L., Johnson, K.J. Nucleic Acids Res. (2000) [Pubmed]
  3. 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]
  4. Six3 overexpression initiates the formation of ectopic retina. Loosli, F., Winkler, S., Wittbrodt, J. Genes Dev. (1999) [Pubmed]
  5. 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]
  6. The Drosophila sine oculis locus encodes a homeodomain-containing protein required for the development of the entire visual system. Cheyette, B.N., Green, P.J., Martin, K., Garren, H., Hartenstein, V., Zipursky, S.L. Neuron (1994) [Pubmed]
  7. Functional dissection of eyes absent reveals new modes of regulation within the retinal determination gene network. Silver, S.J., Davies, E.L., Doyon, L., Rebay, I. Mol. Cell. Biol. (2003) [Pubmed]
  8. 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]
  9. 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]
  10. Embryonic development of the Drosophila corpus cardiacum, a neuroendocrine gland with similarity to the vertebrate pituitary, is controlled by sine oculis and glass. De Velasco, B., Shen, J., Go, S., Hartenstein, V. Dev. Biol. (2004) [Pubmed]
  11. Molecular analysis of Drosophila eyes absent mutants reveals features of the conserved Eya domain. Bui, Q.T., Zimmerman, J.E., Liu, H., Bonini, N.M. Genetics (2000) [Pubmed]
  12. Six3, a medaka homologue of the Drosophila homeobox gene sine oculis is expressed in the anterior embryonic shield and the developing eye. Loosli, F., Köster, R.W., Carl, M., Krone, A., Wittbrodt, J. Mech. Dev. (1998) [Pubmed]
  13. Dpp and Hh signaling in the Drosophila embryonic eye field. Chang, T., Mazotta, J., Dumstrei, K., Dumitrescu, A., Hartenstein, V. Development (2001) [Pubmed]
  14. A somatic role for eyes absent (eya) and sine oculis (so) in Drosophila spermatocyte development. Fabrizio, J.J., Boyle, M., DiNardo, S. Dev. Biol. (2003) [Pubmed]
  15. Fly SIX-type homeodomain proteins Sine oculis and Optix partner with different cofactors during eye development. Kenyon, K.L., Li, D.J., Clouser, C., Tran, S., Pignoni, F. Dev. Dyn. (2005) [Pubmed]
  16. Six1 and Six4 are essential for Gdnf expression in the metanephric mesenchyme and ureteric bud formation, while Six1 deficiency alone causes mesonephric-tubule defects. Kobayashi, H., Kawakami, K., Asashima, M., Nishinakamura, R. Mech. Dev. (2007) [Pubmed]
  17. A transcriptional chain linking eye specification to terminal determination of cone cells in the Drosophila eye. Yan, H., Canon, J., Banerjee, U. Dev. Biol. (2003) [Pubmed]
  18. Histamine: a neurotransmitter candidate for Drosophila photoreceptors. Sarthy, P.V. J. Neurochem. (1991) [Pubmed]
  19. 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]
  20. The mesoderm specification factor twist in the life cycle of jellyfish. Spring, J., Yanze, N., Middel, A.M., Stierwald, M., Gröger, H., Schmid, V. Dev. Biol. (2000) [Pubmed]
  21. 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]
  22. Vertebrate eye development as modeled in Drosophila. Wawersik, S., Maas, R.L. Hum. Mol. Genet. (2000) [Pubmed]
  23. Partner specificity is essential for proper function of the SIX-type homeodomain proteins Sine oculis and Optix during fly eye development. Kenyon, K.L., Yang-Zhou, D., Cai, C.Q., Tran, S., Clouser, C., Decene, G., Ranade, S., Pignoni, F. Dev. Biol. (2005) [Pubmed]
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