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Gene Review

usp  -  ultraspiracle

Drosophila melanogaster

Synonyms: 2C, CF1, CF1/USP, CF1/Usp, CG4380, ...
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Disease relevance of usp

  • We describe here the expression of the CF1/USP protein in an E. coli T7 RNA polymerase-driven expression system [1].

High impact information on usp

  • DHR38 also heterodimerizes with USP, and this complex responds to a distinct class of ecdysteroids in a manner that is independent of EcR [2].
  • This response is unusual in that it does not involve direct binding of ecdysteroids to either DHR38 or USP [2].
  • We now demonstrate that ultraspiracle (usp), a Drosophila RXR homolog, can substitute for RXR in stimulating the DNA binding of receptors for retinoic acid, T3, vitamin D, and peroxisome proliferator activators [3].
  • Replacement of USP with retinoid X receptor produces heterodimers with distinct pharmacological and functional properties [4].
  • Here we report that native EcR and USP are co-localized on ecdysone-responsive loci of polytene chromosomes [4].

Biological context of usp

  • Terminal filament morphogenesis is severely disrupted in homozygous usp clones [5].
  • We have identified the regions of Drosophila EcR and USP responsible for transcriptional activation of a semisynthetic Eip71CD promoter in Kc cells [6].
  • Mutagenesis reveals that AF2 function depends on EcR helix 12, but not on the cognate USP region [6].
  • Among other phenotypes, loss of functional USP in Drosophila eye development results in an accelerated morphogenetic furrow, although loss of ecdysone arrests the furrow [7].
  • A chimeric transgene (d/cusp), produced by replacing the ligand-binding domain (LBD) of Drosophila USP with the equivalent domain from another Diptera, Chironomus tentans, was tested for its ability to rescue Drosophila usp mutants from early larval lethality [8].

Anatomical context of usp


Associations of usp with chemical compounds

  • In vitro experiments with cultured wing discs showed that BRC-Z1 expression and early metamorphic development are rendered steroid-independent in the usp mutant clones [13].
  • Like, USP, it activates transcription in Drosophila Schneider S2 cells in a 20-hydroxyecdysone-dependent manner, via its interaction with the endogenous EcR protein [14].
  • EcRB1 is normally induced with a form of USP that is missing its DNA-binding domain (DBD), although potentiation by juvenile hormone (JH) III is reduced [15].
  • Substitution of E19, the most characteristic amino acid for estrogen receptor-like P-boxes, with alanine showed that the mutation did not appreciably alter the affinity of the wild-type Usp DNA-binding domain (UspDBD(WT)) for a probe containing natural ecdysone response element (hsp27(wt)) [16].
  • Toward this end, we have expressed the EcR DNA-binding domain and the Usp DNA-binding domain as proteins with an affinity tag consisting of six histidine residues (6xHis-EcRDBD and 6xHis-UspDBD, respectively) using the expression vector pQE-30 [17].

Physical interactions of usp

  • These results are consistent with a model in which these latter processes are induced by a signal arising during the middle of the last larval stage but suppressed by the unliganded EcR/USP complex [13].
  • In fluorescence-based binding assays, USP protein binds JH III and JH III acid with specificity, adopting for each ligand a different final conformational state [18].
  • Although members of the NGFI-B family are thought to function exclusively as monomers, we show that DHR38 and BHR38 in fact interact strongly with USP and that this interaction is evolutionarily conserved [19].
  • We have determined the optimal binding site of partially purified bacterially produced CF1/USP by an in vitro selection procedure and also have characterized its binding to the follicular-specific chorion s15 promoter [20].

Regulatory relationships of usp

  • JH III treatment of Saccharomyces cerevisiae expressing a LexA-USP fusion protein stabilizes an oligomeric association containing this protein, as detected by formation of a protein-DNA complex, and induces USP-dependent transcription in a reporter assay [18].
  • Seven-up inhibits ultraspiracle-based signaling pathways in vitro and in vivo [21].

Other interactions of usp


Analytical, diagnostic and therapeutic context of usp

  • AF2 function was studied using a two-hybrid assay in Kc cells, based on the observation that potent hormone-dependent activation by the EcR/USP ligand-binding domain heterodimer requires the participation of both partners [6].
  • We used in vivo, cell culture, and biochemical approaches to analyze the functions of two usp alleles, usp(3) and usp(4), which encode stable proteins with defective DNA-binding domains [25].
  • Molecular cloning and expression analysis of ultraspiracle (USP) from the rice stem borer Chilo suppressalis [26].
  • Biological activities have been determined in Drosophila melanogaster B(II) cell-based bioassay for ecdysteroid agonists and in an in vitro radioligand-displacement assay using bacterially-expressed D. melanogaster EcR/USP receptor proteins [27].
  • Using an antibody raised against DmUSP, Western blot analysis of proteins from epidermis and other tissues revealed five immunoreactive bands, corresponding to different phosphorylated forms of a unique polypeptide, as shown by lambda-phosphatase treatment [28].


  1. Binding affinity of the Drosophila melanogaster CF1/USP protein to the chorion s15 promoter. Christianson, A.M., Kafatos, F.C. Biochem. Biophys. Res. Commun. (1993) [Pubmed]
  2. The Drosophila orphan nuclear receptor DHR38 mediates an atypical ecdysteroid signaling pathway. Baker, K.D., Shewchuk, L.M., Kozlova, T., Makishima, M., Hassell, A., Wisely, B., Caravella, J.A., Lambert, M.H., Reinking, J.L., Krause, H., Thummel, C.S., Willson, T.M., Mangelsdorf, D.J. Cell (2003) [Pubmed]
  3. Drosophila ultraspiracle modulates ecdysone receptor function via heterodimer formation. Yao, T.P., Segraves, W.A., Oro, A.E., McKeown, M., Evans, R.M. Cell (1992) [Pubmed]
  4. Functional ecdysone receptor is the product of EcR and Ultraspiracle genes. Yao, T.P., Forman, B.M., Jiang, Z., Cherbas, L., Chen, J.D., McKeown, M., Cherbas, P., Evans, R.M. Nature (1993) [Pubmed]
  5. The ecdysone receptor and ultraspiracle regulate the timing and progression of ovarian morphogenesis during Drosophila metamorphosis. Hodin, J., Riddiford, L.M. Dev. Genes Evol. (1998) [Pubmed]
  6. Transcription activation by the ecdysone receptor (EcR/USP): identification of activation functions. Hu, X., Cherbas, L., Cherbas, P. Mol. Endocrinol. (2003) [Pubmed]
  7. Analyzing the repressive function of ultraspiracle, the Drosophila RXR, in Drosophila eye development. Ghbeish, N., McKeown, M. Mech. Dev. (2002) [Pubmed]
  8. Developmental effects of a chimeric ultraspiracle gene derived from Drosophila and Chironomus. Henrich, V.C., Vogtli, M.E., Antoniewski, C., Spindler-Barth, M., Przibilla, S., Noureddine, M., Lezzi, M. Genesis (2000) [Pubmed]
  9. Cell-autonomous requirement of the USP/EcR-B ecdysone receptor for mushroom body neuronal remodeling in Drosophila. Lee, T., Marticke, S., Sung, C., Robinow, S., Luo, L. Neuron (2000) [Pubmed]
  10. The RXR homolog ultraspiracle is an essential component of the Drosophila ecdysone receptor. Hall, B.L., Thummel, C.S. Development (1998) [Pubmed]
  11. Characterization of an EcR/USP heterodimer target site that mediates ecdysone responsiveness of the Drosophila Lsp-2 gene. Antoniewski, C., O'Grady, M.S., Edmondson, R.G., Lassieur, S.M., Benes, H. Mol. Gen. Genet. (1995) [Pubmed]
  12. Characterization and partial cloning of ecdysteroid receptor from a cotton boll weevil embryonic cell line. Dhadialla, T.S., Tzertzinis, G. Arch. Insect Biochem. Physiol. (1997) [Pubmed]
  13. The RXR ortholog USP suppresses early metamorphic processes in Drosophila in the absence of ecdysteroids. Schubiger, M., Truman, J.W. Development (2000) [Pubmed]
  14. An rxr/usp homolog from the parasitic nematode, Dirofilaria immitis. Shea, C., Hough, D., Xiao, J., Tzertzinis, G., Maina, C.V. Gene (2004) [Pubmed]
  15. Analysis of transcriptional activity mediated by Drosophila melanogaster ecdysone receptor isoforms in a heterologous cell culture system. Beatty, J., Fauth, T., Callender, J.L., Spindler-Barth, M., Henrich, V.C. Insect Mol. Biol. (2006) [Pubmed]
  16. Functionality versus strength -- has functional selection taken place in the case of the ecdysteroid receptor response element?. Grad, I., Kochman, M., Ozyhar, A. Acta Biochim. Pol. (2002) [Pubmed]
  17. Expression and purification of 6xHis-tagged DNA binding domains of functional ecdysteroid receptor from drosophila melanogaster. Rusin, A., Niedziela-Majka, A., Rymarczyk, G., Ozyhar, A. Acta Biochim. Pol. (1996) [Pubmed]
  18. Ultraspiracle: an invertebrate nuclear receptor for juvenile hormones. Jones, G., Sharp, P.A. Proc. Natl. Acad. Sci. U.S.A. (1997) [Pubmed]
  19. Drosophila hormone receptor 38: a second partner for Drosophila USP suggests an unexpected role for nuclear receptors of the nerve growth factor-induced protein B type. Sutherland, J.D., Kozlova, T., Tzertzinis, G., Kafatos, F.C. Proc. Natl. Acad. Sci. U.S.A. (1995) [Pubmed]
  20. DNA binding and heteromerization of the Drosophila transcription factor chorion factor 1/ultraspiracle. Christianson, A.M., King, D.L., Hatzivassiliou, E., Casas, J.E., Hallenbeck, P.L., Nikodem, V.M., Mitsialis, S.A., Kafatos, F.C. Proc. Natl. Acad. Sci. U.S.A. (1992) [Pubmed]
  21. Seven-up inhibits ultraspiracle-based signaling pathways in vitro and in vivo. Zelhof, A.C., Yao, T.P., Chen, J.D., Evans, R.M., McKeown, M. Mol. Cell. Biol. (1995) [Pubmed]
  22. The ecdysone response enhancer of the Fbp1 gene of Drosophila melanogaster is a direct target for the EcR/USP nuclear receptor. Antoniewski, C., Laval, M., Dahan, A., Lepesant, J.A. Mol. Cell. Biol. (1994) [Pubmed]
  23. Cloning and characterization of CcEcR. An ecdysone receptor homolog from the mediterranean fruit fly ceratitis capitata. Verras, M., Mavroidis, M., Kokolakis, G., Gourzi, P., Zacharopoulou, A., Mintzas, A.C. Eur. J. Biochem. (1999) [Pubmed]
  24. Proteins that bind to Drosophila chorion cis-regulatory elements: a new C2H2 zinc finger protein and a C2C2 steroid receptor-like component. Shea, M.J., King, D.L., Conboy, M.J., Mariani, B.D., Kafatos, F.C. Genes Dev. (1990) [Pubmed]
  25. The dual role of ultraspiracle, the Drosophila retinoid X receptor, in the ecdysone response. Ghbeish, N., Tsai, C.C., Schubiger, M., Zhou, J.Y., Evans, R.M., McKeown, M. Proc. Natl. Acad. Sci. U.S.A. (2001) [Pubmed]
  26. Molecular cloning and expression analysis of ultraspiracle (USP) from the rice stem borer Chilo suppressalis. Minakuchi, C., Nakagawa, Y., Kiuchi, M., Seino, A., Tomita, S., Kamimura, M. Insect Biochem. Mol. Biol. (2003) [Pubmed]
  27. Synthesis and biological activities of turkesterone 11alpha-acyl derivatives. Dinan, L., Bourne, P., Whiting, P., Tsitsekli, A., Saatov, Z., Dhadialla, T.S., Hormann, R.E., Lafont, R., Coll, J. J. Insect Sci. (2003) [Pubmed]
  28. Molecular cloning and expression of Tenebrio molitor ultraspiracle during metamorphosis and in vivo induction of its phosphorylation by 20-hydroxyecdysone. Nicolaï, M., Bouhin, H., Quennedey, B., Delachambre, J. Insect Mol. Biol. (2000) [Pubmed]
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