Gene Symbol: usp
Description: ultraspiracle
Alias: CF1, CF1/USP, CF1/Usp, CG4380, Cf1, DmUSP, DmUsp, Dmel\CG4380, EG:22E5.1, NR2B4, USP, Usp, XR2C, cf1, dUSP, dmUSP, l(1)2Cf, l(1)usp, ultraspiracle, CG4380-PA, CG4380-PB, X receptor at 2C, chorion factor 1, chorion factor CF1, ultraspiricle, usp-PA, usp-PB
Species: fruit fly
Products:     usp

Top Publications

  1. Lee T, Luo L. Mosaic analysis with a repressible cell marker (MARCM) for Drosophila neural development. Trends Neurosci. 2001;24:251-4 pubmed
  2. Beckstead R, Ortiz J, Sanchez C, Prokopenko S, Chambon P, Losson R, et al. Bonus, a Drosophila homolog of TIF1 proteins, interacts with nuclear receptors and can inhibit betaFTZ-F1-dependent transcription. Mol Cell. 2001;7:753-65 pubmed
    ..Bon binds via an LxxLL motif to the AF-2 activation domain present in the ligand binding domain of betaFTZ-F1 and behaves as a transcriptional inhibitor in vivo. ..
  3. Ables E, Drummond Barbosa D. The steroid hormone ecdysone functions with intrinsic chromatin remodeling factors to control female germline stem cells in Drosophila. Cell Stem Cell. 2010;7:581-92 pubmed publisher
    ..Our findings support a link between systemic steroid hormones and the intrinsic chromatin remodeling machinery as a potential mechanism to promote broad transcriptional programs required for adult stem cell self-renewal. ..
  4. Awasaki T, Ito K. Engulfing action of glial cells is required for programmed axon pruning during Drosophila metamorphosis. Curr Biol. 2004;14:668-77 pubmed
    ..We therefore propose that neuron-glia interaction is essential for the precisely coordinated axon-pruning process during Drosophila metamorphosis. ..
  5. Dela Cruz F, Kirsch D, Heinrich J. Transcriptional activity of Drosophila melanogaster ecdysone receptor isoforms and ultraspiracle in Saccharomyces cerevisiae. J Mol Endocrinol. 2000;24:183-91 pubmed
    ..Upon co-expression with ultraspiracle (Usp), transcriptional activation was further increased with EcR-B1 or EcR-A, but was unchanged with EcR-B2 or ..
  6. Shea M, King D, Conboy M, Mariani B, Kafatos F. Proteins that bind to Drosophila chorion cis-regulatory elements: a new C2H2 zinc finger protein and a C2C2 steroid receptor-like component. Genes Dev. 1990;4:1128-40 pubmed
    ..The mRNAs for both of these components, CF1 and CF2, are differentially enriched in the follicles...
  7. Krusiński T, Wietrzych M, Grad I, Ozyhar A, Dobryszycki P. Equilibrium analysis of the DNA binding domain of the ultraspiracle protein interaction with the response element from the hsp27 gene promoter--the application of molecular beacon technology. J Fluoresc. 2008;18:1-10 pubmed
    ..The ecdysone receptor consists of two proteins: the ecdysone receptor (EcR) and the ultraspiracle (Usp). The EcR-Usp dimer conducts transcription through a hsp27(pal) response element...
  8. Bai J, Uehara Y, Montell D. Regulation of invasive cell behavior by taiman, a Drosophila protein related to AIB1, a steroid receptor coactivator amplified in breast cancer. Cell. 2000;103:1047-58 pubmed
    ..The finding of this type of coactivator required for cell motility suggests a novel role for steroid hormones, in stimulating invasive cell behavior, independent of effects on proliferation. ..
  9. . A unified nomenclature system for the nuclear receptor superfamily. Cell. 1999;97:161-3 pubmed

More Information


  1. Morris L, Spradling A. Steroid signaling within Drosophila ovarian epithelial cells sex-specifically modulates early germ cell development and meiotic entry. PLoS ONE. 2012;7:e46109 pubmed publisher
    ..Knocking down expression of the heterodimeric ecdysteroid receptor (EcR/Usp) or the E75 early response gene in escort cells disrupts 16-cell cyst production, meiotic entry and follicle ..
  2. Arbeitman M, Hogness D. Molecular chaperones activate the Drosophila ecdysone receptor, an RXR heterodimer. Cell. 2000;101:67-77 pubmed
    ..a nuclear receptor heterodimer consisting of the ecdysone receptor, EcR, and the Drosophila RXR receptor, USP. We show that EcR/USP DNA binding activity requires activation by a chaperone heterocomplex like that required for ..
  3. Niedziela Majka A, Kochman M, Ozyhar A. Polarity of the ecdysone receptor complex interaction with the palindromic response element from the hsp27 gene promoter. Eur J Biochem. 2000;267:507-19 pubmed
    ..of two members of the nuclear hormone receptors superfamily; the product of the EcR (EcR) and of the ultraspiracle (Usp) genes...
  4. Lee T, Marticke S, Sung C, Robinow S, Luo L. Cell-autonomous requirement of the USP/EcR-B ecdysone receptor for mushroom body neuronal remodeling in Drosophila. Neuron. 2000;28:807-18 pubmed
    ..To elucidate the underlying molecular mechanisms, we conducted a genetic mosaic screen and identified one ultraspiracle (usp) allele defective in larval process pruning...
  5. Rørth P. Initiating and guiding migration: lessons from border cells. Trends Cell Biol. 2002;12:325-31 pubmed
    ..This article discusses the implications of these studies and compares (invasive) migration through a tissue with what is known about cells crawling on a flat substratum. ..
  6. Lehmann M, Wattler F, Korge G. Two new regulatory elements controlling the Drosophila Sgs-3 gene are potential ecdysone receptor and fork head binding sites. Mech Dev. 1997;62:15-27 pubmed
    ..elements in the upstream region of the Drosophila Sgs-3 gene which are both able to bind the ecdysone receptor (EcR/USP) and the product of the fork head gene...
  7. Yao T, Segraves W, Oro A, McKeown M, Evans R. Drosophila ultraspiracle modulates ecdysone receptor function via heterodimer formation. Cell. 1992;71:63-72 pubmed
    ..We now demonstrate that ultraspiracle (usp), a Drosophila RXR homolog, can substitute for RXR in stimulating the DNA binding of receptors for ..
  8. Grueber W, Jan Y. Dendritic development: lessons from Drosophila and related branches. Curr Opin Neurobiol. 2004;14:74-82 pubmed
    ..Here, we review some recent advances in our understanding of dendritic development in insects, focusing primarily on insights that have been gained from studies of Drosophila. ..
  9. Escriva H, Safi R, Hanni C, Langlois M, Saumitou Laprade P, Stehelin D, et al. Ligand binding was acquired during evolution of nuclear receptors. Proc Natl Acad Sci U S A. 1997;94:6803-8 pubmed
    ..Strikingly, our results suggest that the ancestral NR was an orphan receptor that acquired ligand-binding ability during subsequent evolution. ..
  10. Dobryszycki P, Grad I, Krusiński T, Michaluk P, Sawicka D, Kowalska A, et al. The DNA-binding domain of the ultraspiracle drives deformation of the response element whereas the DNA-binding domain of the ecdysone receptor is responsible for a slight additional change of the preformed structure. Biochemistry. 2006;45:668-75 pubmed
    ..metamorphosis in insects via a heterodimeric complex of two nuclear receptors, the ecdysone receptor (EcR) and ultraspiracle protein (Usp)...
  11. Sedkov Y, Cho E, Petruk S, Cherbas L, Smith S, Jones R, et al. Methylation at lysine 4 of histone H3 in ecdysone-dependent development of Drosophila. Nature. 2003;426:78-83 pubmed by binding to a nuclear receptor, EcR, which heterodimerizes with the retinoid X receptor homologue Ultraspiracle. Both partners are required for binding to ligand or DNA...
  12. Palanker L, Necakov A, Sampson H, Ni R, Hu C, Thummel C, et al. Dynamic regulation of Drosophila nuclear receptor activity in vivo. Development. 2006;133:3549-62 pubmed
    ..identified the angular psoralen angelicin and the insect growth regulator fenoxycarb as activators of the Ultraspiracle (USP) ligand-binding domain...
  13. Oro A, McKeown M, Evans R. Relationship between the product of the Drosophila ultraspiracle locus and the vertebrate retinoid X receptor. Nature. 1990;347:298-301 pubmed
    ..This receptor-like protein is encoded by ultraspiracle (usp), a locus required both maternally and zygotically for pattern formation...
  14. Tsai C, Kao H, Yao T, McKeown M, Evans R. SMRTER, a Drosophila nuclear receptor coregulator, reveals that EcR-mediated repression is critical for development. Mol Cell. 1999;4:175-86 pubmed
    The Drosophila ecdysone receptor (EcR)/ultraspiracle (USP) heterodimer is a key regulator in molting and metamorphoric processes, activating and repressing transcription in a sequence-specific manner...
  15. Jang A, Chang Y, Bai J, Montell D. Border-cell migration requires integration of spatial and temporal signals by the BTB protein Abrupt. Nat Cell Biol. 2009;11:569-79 pubmed publisher
    ..Taken together, these findings provide a molecular mechanism by which spatial and temporal cues are integrated. ..
  16. Kuo C, Jan L, Jan Y. Dendrite-specific remodeling of Drosophila sensory neurons requires matrix metalloproteases, ubiquitin-proteasome, and ecdysone signaling. Proc Natl Acad Sci U S A. 2005;102:15230-5 pubmed
    ..These results demonstrate that, in response to environmental changes, certain neurons have cell-intrinsic abilities to completely lose their dendrites but keep their axons and subsequently regrow their dendritic arbors. ..
  17. Lehmann M, Korge G. Ecdysone regulation of the Drosophila Sgs-4 gene is mediated by the synergistic action of ecdysone receptor and SEBP 3. EMBO J. 1995;14:716-26 pubmed
    ..Thus, the available data suggest that induction of Sgs-4, and possibly other 'intermolt genes', is a combination of a primary and a secondary response to the hormone. ..
  18. Huet F, Ruiz C, Richards G. Sequential gene activation by ecdysone in Drosophila melanogaster: the hierarchical equivalence of early and early late genes. Development. 1995;121:1195-204 pubmed
    ..sequential activation of members of the superfamily of nuclear receptors (DHR3, DHR39, EcR, E75, E78, FTZ-F1, usp)...
  19. Baehrecke E. Steroid regulation of programmed cell death during Drosophila development. Cell Death Differ. 2000;7:1057-62 pubmed
    ..This article reviews the current knowledge of steroid signaling and the regulation of programmed cell death during development of Drosophila. ..
  20. Laudet V, Hanni C, Coll J, Catzeflis F, Stehelin D. Evolution of the nuclear receptor gene superfamily. EMBO J. 1992;11:1003-13 pubmed
    ..These data suggest a complex evolutionary history for nuclear receptor genes in which gene duplication events and swapping between domains of different origins took place. ..
  21. Elke C, Vogtli M, Rauch P, Spindler Barth M, Lezzi M. Expression of EcR and USP in Escherichia coli: purification and functional studies. Arch Insect Biochem Physiol. 1997;35:59-69 pubmed
    ..ecdysteroid receptor complex consists of a nuclear receptor heterodimer of ecdysteroid receptor (EcR) and ultraspiracle (USP)...
  22. Henrich V, Sliter T, Lubahn D, MacIntyre A, Gilbert L. A steroid/thyroid hormone receptor superfamily member in Drosophila melanogaster that shares extensive sequence similarity with a mammalian homologue. Nucleic Acids Res. 1990;18:4143-8 pubmed
    ..The high degree of similarity in the hormone-binding domain between the 2C protein and the H2RII binding protein outside regions II-IV suggests specific functional roles which are not shared by other members of the superfamily. ..
  23. Ghbeish N, McKeown M. Analyzing the repressive function of ultraspiracle, the Drosophila RXR, in Drosophila eye development. Mech Dev. 2002;111:89-98 pubmed
    Response to the insect hormone ecdysone is mediated by a nuclear receptor complex containing Ultraspiracle (USP) and the Ecdysone Receptor (EcR)...
  24. King Jones K, Thummel C. Nuclear receptors--a perspective from Drosophila. Nat Rev Genet. 2005;6:311-23 pubmed
    ..Here, we review recent breakthroughs in our understanding of D. melanogaster nuclear receptors, and interpret these results in light of findings from their evolutionarily conserved vertebrate homologues. ..
  25. Antoniewski C, Laval M, Dahan A, Lepesant J. The ecdysone response enhancer of the Fbp1 gene of Drosophila melanogaster is a direct target for the EcR/USP nuclear receptor. Mol Cell Biol. 1994;14:4465-74 pubmed
    ..antibodies, one of the complexes was identified as a heterodimer consisting of the ecdysone receptor (EcR) and Ultraspiracle (USP) proteins...
  26. Lehmann M. Drosophila Sgs genes: stage and tissue specificity of hormone responsiveness. Bioessays. 1996;18:47-54 pubmed
    ..g. secretion enhancer binding proteins, which act together in a synergistic manner with the 20E receptor and early gene products to establish a stage- and tissue-specific expression pattern. ..
  27. Gates J, Lam G, Ortiz J, Losson R, Thummel C. rigor mortis encodes a novel nuclear receptor interacting protein required for ecdysone signaling during Drosophila larval development. Development. 2004;131:25-36 pubmed
    ..The ecdysone signal is transduced by the EcR/USP nuclear receptor heterodimer that binds to specific response elements in the genome and directly regulates target ..
  28. Kozlova T, Thummel C. Spatial patterns of ecdysteroid receptor activation during the onset of Drosophila metamorphosis. Development. 2002;129:1739-50 pubmed
    Ecdysteroid signaling in insects is transduced by a heterodimer of the EcR and USP nuclear receptors...
  29. Koelle M, Talbot W, Segraves W, Bender M, Cherbas P, Hogness D. The Drosophila EcR gene encodes an ecdysone receptor, a new member of the steroid receptor superfamily. Cell. 1991;67:59-77 pubmed
    ..As expected, EcR is nuclear and found in all ecdysone target tissues examined. Furthermore, the EcR gene is expressed at each developmental stage marked by a pulse of ecdysone. ..
  30. Christianson A, King D, Hatzivassiliou E, Casas J, Hallenbeck P, Nikodem V, et al. DNA binding and heteromerization of the Drosophila transcription factor chorion factor 1/ultraspiracle. Proc Natl Acad Sci U S A. 1992;89:11503-7 pubmed
    The Drosophila chorion factor 1/ultraspiracle (CF1/USP) transcription factor, a homologue of the retinoid X receptor, is a developmentally important member of the family of nuclear (steroid) hormone receptors...
  31. Brodu V, Mugat B, Roignant J, Lepesant J, Antoniewski C. Dual requirement for the EcR/USP nuclear receptor and the dGATAb factor in an ecdysone response in Drosophila melanogaster. Mol Cell Biol. 1999;19:5732-42 pubmed
    The EcR/USP nuclear receptor controls Drosophila metamorphosis by activating complex cascades of gene transcription in response to pulses of the steroid hormone ecdysone at the end of larval development...
  32. Grad I, Niedziela Majka A, Kochman M, Ozyhar A. Analysis of Usp DNA binding domain targeting reveals critical determinants of the ecdysone receptor complex interaction with the response element. Eur J Biochem. 2001;268:3751-8 pubmed
    ..formed by two members of the nuclear hormone receptors superfamily, the product of the EcR (EcR) and of the ultraspiracle (Usp) genes. Our previous study [Niedziela-Majka, A., Kochman, M., Ozyhar, A. (2000) Eur. J. Biochem...
  33. Hall B, Thummel C. The RXR homolog ultraspiracle is an essential component of the Drosophila ecdysone receptor. Development. 1998;125:4709-17 pubmed
    ..that the EcR ecdysone receptor requires an RXR heterodimer partner for its activity, encoded by the ultraspiracle (usp) locus...
  34. Jiang C, Lamblin A, Steller H, Thummel C. A steroid-triggered transcriptional hierarchy controls salivary gland cell death during Drosophila metamorphosis. Mol Cell. 2000;5:445-55 pubmed
    ..This study identifies transcriptional regulators of programmed cell death in Drosophila and provides a direct link between a steroid signal and a programmed cell death response. ..
  35. Niedziela Majka A, Rymarczyk G, Kochman M, Ozyhar A. GST-Induced dimerization of DNA-binding domains alters characteristics of their interaction with DNA. Protein Expr Purif. 1998;14:208-20 pubmed
    ..Two members of the nuclear receptor superfamily, the product of the EcR (EcR) and of the ultraspiracle genes (Usp), heterodimerize to form its functional receptor...
  36. Francis V, Zorzano A, Teleman A. dDOR is an EcR coactivator that forms a feed-forward loop connecting insulin and ecdysone signaling. Curr Biol. 2010;20:1799-808 pubmed publisher
    ..Furthermore, because ecdysone signaling inhibits insulin signaling in the fat body, this also uncovers a feed-forward mechanism whereby ecdysone potentiates its own signaling via dDOR. ..
  37. Kozlova T, Thummel C. Essential roles for ecdysone signaling during Drosophila mid-embryonic development. Science. 2003;301:1911-4 pubmed
    ..We show that the EcR-USP ecdysone receptor is first activated in the extraembryonic amnioserosa, implicating this tissue as a source of ..
  38. Schubiger M, Carré C, Antoniewski C, Truman J. Ligand-dependent de-repression via EcR/USP acts as a gate to coordinate the differentiation of sensory neurons in the Drosophila wing. Development. 2005;132:5239-48 pubmed
    Loss of function of either the ecdysone receptor (EcR) or Ultraspiracle (USP), the two components of the ecdysone receptor, causes precocious differentiation of the sensory neurons on the wing of Drosophila...
  39. Shi L, Lin S, Grinberg Y, Beck Y, Grozinger C, Robinson G, et al. Roles of Drosophila Kruppel-homolog 1 in neuronal morphogenesis. Dev Neurobiol. 2007;67:1614-26 pubmed
    ..addition, Kr-h1 expression remains strongly linked to ecdysone-regulated pathways: Kr-h1 expression is regulated by usp, which dimerizes to the ecdysone receptor, and Kr-h1 expression is essential for proper patterning of the ecdysone ..
  40. Biyasheva A, Do T, Lu Y, Vaskova M, Andres A. Glue secretion in the Drosophila salivary gland: a model for steroid-regulated exocytosis. Dev Biol. 2001;231:234-51 pubmed
    ..We show that 20E mediates secretion through the EcR/USP receptor, and two early-gene products, the rbp(+) function of BR-C and the Ca2+ binding protein E63-1, are involved...
  41. Kucherenko M, Barth J, Fiala A, Shcherbata H. Steroid-induced microRNA let-7 acts as a spatio-temporal code for neuronal cell fate in the developing Drosophila brain. EMBO J. 2012;31:4511-23 pubmed publisher
    ..Our data propose a novel role for miRNAs as transducers between chronologically regulated developmental signalling and physical cell adhesion. ..
  42. Yao T, Forman B, Jiang Z, Cherbas L, Chen J, McKeown M, et al. Functional ecdysone receptor is the product of EcR and Ultraspiracle genes. Nature. 1993;366:476-9 pubmed
    ..Rather, these activities are dependent on heterodimer formation with Ultraspiracle (USP) the insect homologue of vertebrate retinoid X receptor...
  43. Dressel U, Thormeyer D, Altincicek B, Paululat A, Eggert M, Schneider S, et al. Alien, a highly conserved protein with characteristics of a corepressor for members of the nuclear hormone receptor superfamily. Mol Cell Biol. 1999;19:3383-94 pubmed
    ..the ecdysone receptor and Seven-up, the Drosophila homologue of COUP-TF1, but not with retinoic acid receptor, RXR/USP, DHR 3, DHR 38, DHR 78, or DHR 96...
  44. Thomas H, Stunnenberg H, Stewart A. Heterodimerization of the Drosophila ecdysone receptor with retinoid X receptor and ultraspiracle. Nature. 1993;362:471-5 pubmed that EcR needs to heterodimerize with either the retinoid X receptor (RXR) or its Drosophila homologue, ultraspiracle (USP), for DNA binding and transactivation...
  45. Zelhof A, Ghbeish N, Tsai C, Evans R, McKeown M. A role for ultraspiracle, the Drosophila RXR, in morphogenetic furrow movement and photoreceptor cluster formation. Development. 1997;124:2499-506 pubmed
    ..We report that ultraspiracle, the gene encoding the Drosophila cognate of the Retinoid X Receptor, is required for normal morphogenetic ..
  46. Oro A, McKeown M, Evans R. The Drosophila retinoid X receptor homolog ultraspiracle functions in both female reproduction and eye morphogenesis. Development. 1992;115:449-62 pubmed
    b>Ultraspiracle (usp) encodes the Drosophila cognate of RXR, the human retinoid X receptor. To examine how RXR subfamily members function in development, we have undertaken a phenotypic analysis of usp mutants...
  47. Fahrbach S, Smagghe G, Velarde R. Insect nuclear receptors. Annu Rev Entomol. 2012;57:83-106 pubmed publisher
    ..An exciting new technology allows EcR to be used in chimeric, ligand-inducible gene-switch systems with applications in pest management and medicine. ..
  48. Marin E, Watts R, Tanaka N, Ito K, Luo L. Developmentally programmed remodeling of the Drosophila olfactory circuit. Development. 2005;132:725-37 pubmed
    ..As with MB gamma neurons, PN pruning requires cell-autonomous reception of the nuclear hormone ecdysone. Thus, these synaptic partners are independently programmed to prune their dendrites and axons. ..
  49. Cakouros D, Daish T, Kumar S. Ecdysone receptor directly binds the promoter of the Drosophila caspase dronc, regulating its expression in specific tissues. J Cell Biol. 2004;165:631-40 pubmed
    ..mediates its biological effects by either direct activation of gene transcription after binding to its receptor EcR-Usp or via hierarchical transcriptional regulation of several primary transcription factors...
  50. Zhu J, Chen L, Sun G, Raikhel A. The competence factor beta Ftz-F1 potentiates ecdysone receptor activity via recruiting a p160/SRC coactivator. Mol Cell Biol. 2006;26:9402-12 pubmed
    ..Thus, our results reveal the molecular basis of competence for the stage-specific 20E response...
  51. Hodin J, Riddiford L. The ecdysone receptor and ultraspiracle regulate the timing and progression of ovarian morphogenesis during Drosophila metamorphosis. Dev Genes Evol. 1998;208:304-17 pubmed
    ..receptor complex, a heterodimeric nuclear receptor consisting of the ecdysone receptor (EcR) and its partner ultraspiracle (USP)...
  52. Ghbeish N, Tsai C, Schubiger M, Zhou J, Evans R, McKeown M. The dual role of ultraspiracle, the Drosophila retinoid X receptor, in the ecdysone response. Proc Natl Acad Sci U S A. 2001;98:3867-72 pubmed
    The Drosophila homolog of the retinoid X receptor, ultraspiracle (USP), heterodimerizes with the ecdysone receptor (EcR) to form a functional complex that mediates the effects of the steroid molting hormone ecdysone by activating and ..
  53. Gancz D, Lengil T, Gilboa L. Coordinated regulation of niche and stem cell precursors by hormonal signaling. PLoS Biol. 2011;9:e1001202 pubmed publisher
    ..Similar principles might govern the formation of additional stem cell units during organogenesis. ..
  54. Sutherland J, Kozlova T, Tzertzinis G, Kafatos F. 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. Proc Natl Acad Sci U S A. 1995;92:7966-70 pubmed
    In Drosophila the response to the hormone ecdysone is mediated in part by Ultraspiracle (USP) and ecdysone receptor (EcR), which are members of the nuclear receptor superfamily...
  55. Mouillet J, Henrich V, Lezzi M, Vogtli M. Differential control of gene activity by isoforms A, B1 and B2 of the Drosophila ecdysone receptor. Eur J Biochem. 2001;268:1811-9 pubmed
    ..molting and metamorphosis in Drosophila via a heterodimeric receptor consisting of EcR that binds hormone, and USP, a homolog of the vertebrate RXR receptor...
  56. König A, Yatsenko A, WEISS M, Shcherbata H. Ecdysteroids affect Drosophila ovarian stem cell niche formation and early germline differentiation. EMBO J. 2011;30:1549-62 pubmed publisher
    ..Additionally, when ecdysone signalling is perturbed during the process of somatic stem cell niche establishment enlarged functional niches able to host additional stem cells are formed. ..
  57. Schubiger M, Truman J. The RXR ortholog USP suppresses early metamorphic processes in Drosophila in the absence of ecdysteroids. Development. 2000;127:1151-9 pubmed
    ..insects by signaling through the ecdysone receptor complex, a heterodimer of the ecdysone receptor (EcR) and ultraspiracle (USP)...