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Gene Review

exu  -  exuperantia

Drosophila melanogaster

Synonyms: CG8994, Dmel\CG8994, Exu, Maternal protein exuperantia, bs30h09.y1, ...
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Disease relevance of exu

  • In addition, we find that a smaller deletion which removes only sequence within the male 3'-UTR reduces the steady-state level of the mRNA and prevents an exu transgene from rescuing male sterility [1].

High impact information on exu

  • Drosophila bicoid mRNA is synthesized in the nurse cells and transported to the oocyte where microtubules and Exuperantia protein mediate localization to the anterior pole [2].
  • Here we express a chimaeric gene encoding a fusion between the Acquorea victoria green fluorescent protein (GFP) and the exu protein (Exu) in female germ cells, and find that the fusion protein fluoresces strongly in both live and fixed cells during Drosophila oogenesis [3].
  • Furthermore, both Exu/Yps particles and osk mRNA follow a similar temporal pattern of localization in which they transiently accumulate at the oocyte anterior and subsequently localize to the posterior pole [4].
  • One of these proteins was identified as the cold shock domain RNA-binding protein Ypsilon Schachtel (Yps), which we show binds directly to Exu and colocalizes with Exu in both the oocyte and nurse cells of the Drosophila egg chamber [4].
  • We discovered that exu-null mutants are defective in osk mRNA localization in both nurse cells and the oocyte [4].

Biological context of exu


Anatomical context of exu


Enzymatic interactions of exu


Co-localisations of exu

  • Notably, we find that exu protein is colocalized with bcd mRNA during the early phase of localization, when bcd mRNA is positioned at the apical regions of the nurse cells [10].

Regulatory relationships of exu


Other interactions of exu

  • In contrast, clearly nonadditive effects on the ftz pattern were seen when a mutation in a gene of the anterior class (exuperantia) was combined with mutations in posterior class genes [11].
  • Using forms of the 3' UTR lacking this RNA recognition redundancy, we reexamine the roles of the swallow, staufen, and exuperantia genes, which are all required for normal bicoid mRNA localization [12].
  • We report here that tra-2 is required in male germ cells for efficient male-specific processing of exu RNA; in the absence of tra-2, X/Y males produce a new mRNA which is processed at its 3' end so that it contains sequences normally specific to the female 3' untranslated region [9].

Analytical, diagnostic and therapeutic context of exu

  • To gain insight into the mechanism of anterior patterning, we have used time lapse laser scanning confocal microscopy to analyze transport of particles containing a Green Fluorescent Protein-Exu fusion (GFP-Exu), and to directly image microtubule organization in vivo [13].


  1. A male-specific 3'-UTR regulates the steady-state level of the exuperantia mRNA during spermatogenesis in Drosophila. Crowley, T.E., Hazelrigg, T. Mol. Gen. Genet. (1995) [Pubmed]
  2. In vivo analysis of Drosophila bicoid mRNA localization reveals a novel microtubule-dependent axis specification pathway. Cha, B.J., Koppetsch, B.S., Theurkauf, W.E. Cell (2001) [Pubmed]
  3. Implications for bcd mRNA localization from spatial distribution of exu protein in Drosophila oogenesis. Wang, S., Hazelrigg, T. Nature (1994) [Pubmed]
  4. Isolation of a ribonucleoprotein complex involved in mRNA localization in Drosophila oocytes. Wilhelm, J.E., Mansfield, J., Hom-Booher, N., Wang, S., Turck, C.W., Hazelrigg, T., Vale, R.D. J. Cell Biol. (2000) [Pubmed]
  5. The temporal and spatial distribution pattern of maternal exuperantia protein: evidence for a role in establishment but not maintenance of bicoid mRNA localization. Marcey, D., Watkins, W.S., Hazelrigg, T. EMBO J. (1991) [Pubmed]
  6. Par-1 regulates bicoid mRNA localisation by phosphorylating Exuperantia. Riechmann, V., Ephrussi, A. Development (2004) [Pubmed]
  7. The exuperantia gene is required for Drosophila spermatogenesis as well as anteroposterior polarity of the developing oocyte, and encodes overlapping sex-specific transcripts. Hazelrigg, T., Watkins, W.S., Marcey, D., Tu, C., Karow, M., Lin, X.R. Genetics (1990) [Pubmed]
  8. The molecular motor dynein is involved in targeting swallow and bicoid RNA to the anterior pole of Drosophila oocytes. Schnorrer, F., Bohmann, K., Nüsslein-Volhard, C. Nat. Cell Biol. (2000) [Pubmed]
  9. Sex-specific processing of the Drosophila exuperantia transcript is regulated in male germ cells by the tra-2 gene. Hazelrigg, T., Tu, C. Proc. Natl. Acad. Sci. U.S.A. (1994) [Pubmed]
  10. Protein encoded by the exuperantia gene is concentrated at sites of bicoid mRNA accumulation in Drosophila nurse cells but not in oocytes or embryos. Macdonald, P.M., Luk, S.K., Kilpatrick, M. Genes Dev. (1991) [Pubmed]
  11. Maternal-effect genes that alter the fate map of the Drosophila blastoderm embryo. Winslow, G.M., Carroll, S.B., Scott, M.P. Dev. Biol. (1988) [Pubmed]
  12. Redundant RNA recognition events in bicoid mRNA localization. Macdonald, P.M., Kerr, K. RNA (1997) [Pubmed]
  13. In vivo analyses of cytoplasmic transport and cytoskeletal organization during Drosophila oogenesis: characterization of a multi-step anterior localization pathway. Theurkauf, W.E., Hazelrigg, T.I. Development (1998) [Pubmed]
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