UBE2I

Summary

Gene Symbol: UBE2I
Description: ubiquitin conjugating enzyme E2 I
Alias: C358B7.1, P18, UBC9, SUMO-conjugating enzyme UBC9, RING-type E3 SUMO transferase UBC9, SUMO-1-protein ligase, SUMO-protein ligase, ubiquitin carrier protein 9, ubiquitin carrier protein I, ubiquitin conjugating enzyme 9, ubiquitin conjugating enzyme E2I, ubiquitin-conjugating enzyme E2I (UBC9 homolog, yeast), ubiquitin-conjugating enzyme E2I (homologous to yeast UBC9), ubiquitin-conjugating enzyme UbcE2A, ubiquitin-like protein SUMO-1 conjugating enzyme, ubiquitin-protein ligase E2I, ubiquitin-protein ligase I
Species: human
Products:     UBE2I

Top Publications

  1. Lu Z, Wu H, Mo Y. Regulation of bcl-2 expression by Ubc9. Exp Cell Res. 2006;312:1865-75 pubmed
    ..Sumoylation is a multiple-step process, involving maturation, activation, conjugation and ligation. Ubc9 is an E2 conjugating enzyme essential for sumoylation...
  2. Boggio R, Colombo R, Hay R, Draetta G, Chiocca S. A mechanism for inhibiting the SUMO pathway. Mol Cell. 2004;16:549-61 pubmed
    ..pathway with three discrete steps: activation involving the enzyme E1, conjugation involving the E2 enzyme UBC9, and substrate modification through the cooperative association of UBC9 and E3 ligases...
  3. Kahyo T, Nishida T, Yasuda H. Involvement of PIAS1 in the sumoylation of tumor suppressor p53. Mol Cell. 2001;8:713-8 pubmed
    ..In addition, PIAS1 bound p53 and Ubc9, the E2 for SUMO. PIAS1 that was mutated in the RING finger-like domain bound p53 and SUMO-1, but not Ubc9...
  4. Tashiro K, Pando M, Kanegae Y, Wamsley P, Inoue S, Verma I. Direct involvement of the ubiquitin-conjugating enzyme Ubc9/Hus5 in the degradation of IkappaBalpha. Proc Natl Acad Sci U S A. 1997;94:7862-7 pubmed
    ..mE2 is homologous to the yeast Ubc9/Hus5 ubiquitin-conjugating enzyme...
  5. Reverter D, Lima C. Insights into E3 ligase activity revealed by a SUMO-RanGAP1-Ubc9-Nup358 complex. Nature. 2005;435:687-92 pubmed
    ..to the cytoplasmic face of the nuclear pore complex by means of interactions in a complex that also includes Ubc9, the SUMO E2 conjugating protein. Here we describe the 3...
  6. Werner A, Flotho A, Melchior F. The RanBP2/RanGAP1*SUMO1/Ubc9 complex is a multisubunit SUMO E3 ligase. Mol Cell. 2012;46:287-98 pubmed publisher
    ..why RanBP2 functions in vivo has been unclear: throughout the cell cycle it stably interacts with RanGAP1*SUMO1 and Ubc9, whose binding sites overlap with the E3 ligase region...
  7. Zhang H, Saitoh H, Matunis M. Enzymes of the SUMO modification pathway localize to filaments of the nuclear pore complex. Mol Cell Biol. 2002;22:6498-508 pubmed
    ..We also investigated the localization of the SUMO conjugating enzyme, Ubc9. Using immunogold labeling of isolated nuclear envelopes, we found that Ubc9 localizes to both the cytoplasmic and ..
  8. Mo Y, Yu Y, Ee P, Beck W. Overexpression of a dominant-negative mutant Ubc9 is associated with increased sensitivity to anticancer drugs. Cancer Res. 2004;64:2793-8 pubmed
    b>Ubc9 is an E2-conjugating enzyme required for sumoylation and has been implicated in regulating several critical cellular pathways...
  9. Giorgino F, de Robertis O, Laviola L, Montrone C, Perrini S, McCowen K, et al. The sentrin-conjugating enzyme mUbc9 interacts with GLUT4 and GLUT1 glucose transporters and regulates transporter levels in skeletal muscle cells. Proc Natl Acad Sci U S A. 2000;97:1125-30 pubmed
    ..The mUbc9 sentrin-conjugating enzyme represents a novel regulator of GLUT1 and GLUT4 protein levels with potential importance as a determinant of basal and insulin-stimulated glucose uptake in normal and pathophysiological states. ..

More Information

Publications106 found, 100 shown here

  1. Seeler J, Marchio A, Losson R, Desterro J, Hay R, Chambon P, et al. Common properties of nuclear body protein SP100 and TIF1alpha chromatin factor: role of SUMO modification. Mol Cell Biol. 2001;21:3314-24 pubmed
  2. Yang S, Jaffray E, Hay R, Sharrocks A. Dynamic interplay of the SUMO and ERK pathways in regulating Elk-1 transcriptional activity. Mol Cell. 2003;12:63-74 pubmed
    ..Thus, the reciprocal regulation of the activation and repressive activities are coupled by MAP kinase modification of Elk-1. ..
  3. Dünnebier T, Bermejo J, Haas S, Fischer H, Pierl C, Justenhoven C, et al. Common variants in the UBC9 gene encoding the SUMO-conjugating enzyme are associated with breast tumor grade. Int J Cancer. 2009;125:596-602 pubmed publisher
    b>UBC9 encodes a protein that conjugates small ubiquitin-related modifier (SUMO) to target proteins resulting in a change of their localization, activity or stability...
  4. Kawabe Y, Seki M, Seki T, Wang W, Imamura O, Furuichi Y, et al. Covalent modification of the Werner's syndrome gene product with the ubiquitin-related protein, SUMO-1. J Biol Chem. 2000;275:20963-6 pubmed
    ..One was a novel protein, and the other two were mouse Ubc9 and SUMO-1...
  5. Saltzman A, Searfoss G, Marcireau C, Stone M, Ressner R, Munro R, et al. hUBC9 associates with MEKK1 and type I TNF-alpha receptor and stimulates NFkappaB activity. FEBS Lett. 1998;425:431-5 pubmed
    ..hUBC9 also synergized with MEKK1 to activate NFkappaB reporter activity. Thus, hUBC9 modulates NFkappaB activity which, at least in part, can be attributed to its interaction with MEKK1 and the type I TNF-alpha receptor. ..
  6. Zhu S, Sachdeva M, Wu F, Lu Z, Mo Y. Ubc9 promotes breast cell invasion and metastasis in a sumoylation-independent manner. Oncogene. 2010;29:1763-72 pubmed publisher
    b>Ubc9 is an E2-conjugating enzyme that transfers the activated small ubiquitin-like modifier (SUMO) to protein substrates, and thus it has an important function in sumoylation-mediated cellular pathways...
  7. Pichler A, Knipscheer P, Saitoh H, Sixma T, Melchior F. The RanBP2 SUMO E3 ligase is neither HECT- nor RING-type. Nat Struct Mol Biol. 2004;11:984-91 pubmed
    ..modification with the ubiquitin-related protein SUMO1 requires the E1 enzyme Aos1-Uba2 and the E2 enzyme Ubc9. Distinct E3 ligases strongly enhance modification of specific targets...
  8. Martin S, Tatham M, Hay R, Samuel I. Quantitative analysis of multi-protein interactions using FRET: application to the SUMO pathway. Protein Sci. 2008;17:777-84 pubmed publisher
    ..These assays provide powerful tools for the study of competitive biochemical cascades and the extent to which drug candidates modify protein interactions. ..
  9. Lin D, Tatham M, Yu B, Kim S, Hay R, Chen Y. Identification of a substrate recognition site on Ubc9. J Biol Chem. 2002;277:21740-8 pubmed
    Human Ubc9 is homologous to ubiquitin-conjugating enzymes. However, instead of conjugating ubiquitin, it conjugates a ubiquitin homologue, small ubiquitin-like modifier 1 (SUMO-1), also known as UBL1, GMP1, SMTP3, PIC1, and sentrin...
  10. Sentis S, Le Romancer M, Bianchin C, Rostan M, Corbo L. Sumoylation of the estrogen receptor alpha hinge region regulates its transcriptional activity. Mol Endocrinol. 2005;19:2671-84 pubmed
    ..of transcription (PIAS)1 and PIAS3 as E3 ligases for ERalpha, we also found that PIAS1 and PIAS3, as well as Ubc9, modulated ERalpha-dependent transcription independently from their SUMO-1 conjugation activity...
  11. Rytinki M, Palvimo J. SUMOylation attenuates the function of PGC-1alpha. J Biol Chem. 2009;284:26184-93 pubmed publisher
    ..Thus, our findings showing that reversible SUMOylation can adjust the activity of PGC-1alpha add a novel layer to the regulation of the coactivator. ..
  12. Tatham M, Kim S, Jaffray E, Song J, Chen Y, Hay R. Unique binding interactions among Ubc9, SUMO and RanBP2 reveal a mechanism for SUMO paralog selection. Nat Struct Mol Biol. 2005;12:67-74 pubmed
    ..proteins requires the concerted action of the specific E1-activating enzyme SAE1/SAE2, the E2-conjugating enzyme Ubc9, and an E3-like SUMO ligase...
  13. Rajan S, Plant L, Rabin M, Butler M, Goldstein S. Sumoylation silences the plasma membrane leak K+ channel K2P1. Cell. 2005;121:37-47 pubmed
    ..Removal of the peptide adduct by SUMO protease reveals K2P1 to be a K+-selective, pH-sensitive, openly rectifying channel regulated by reversible peptide linkage. ..
  14. Chang Y, Huang C, Chan J, Liu P, Chang H, Huang S. Regulation of nuclear receptor and coactivator functions by the carboxyl terminus of ubiquitin-conjugating enzyme 9. Int J Biochem Cell Biol. 2007;39:1035-46 pubmed
    ..The SUMO E2 enzyme ubiquitin-conjugating enzyme 9 (Ubc9) is sufficient for substrate recognition and lysine modification of known SUMO targets...
  15. Kovalenko O, Plug A, Haaf T, Gonda D, Ashley T, Ward D, et al. Mammalian ubiquitin-conjugating enzyme Ubc9 interacts with Rad51 recombination protein and localizes in synaptonemal complexes. Proc Natl Acad Sci U S A. 1996;93:2958-63 pubmed
    ..The 18-kDa HsUbc9 protein is homologous to the ubiquitin-conjugating enzymes Hus5 of Schizosaccharomyces pombe and Ubc9 of Saccharomyces cerevisiae. The Hsubc9 gene complements a ubc9 mutation of S. cerevisiae...
  16. Saitoh H, Pizzi M, Wang J. Perturbation of SUMOlation enzyme Ubc9 by distinct domain within nucleoporin RanBP2/Nup358. J Biol Chem. 2002;277:4755-63 pubmed
    b>Ubc9, a conjugation enzyme for the ubiquitin-related modifier SUMO, is present predominantly in the nucleus and at the nuclear pore complex...
  17. Yang S, Galanis A, Witty J, Sharrocks A. An extended consensus motif enhances the specificity of substrate modification by SUMO. EMBO J. 2006;25:5083-93 pubmed
    ..Mechanistically, the acidic residues are important in enhancing the efficiency of Elk-1 sumoylation by Ubc9. Similar mechanisms operate in other transcription factors and phosphorylation sites can functionally substitute ..
  18. Moschos S, Jukic D, Athanassiou C, Bhargava R, Dacic S, Wang X, et al. Expression analysis of Ubc9, the single small ubiquitin-like modifier (SUMO) E2 conjugating enzyme, in normal and malignant tissues. Hum Pathol. 2010;41:1286-98 pubmed publisher
    ..multiple E2 enzymes required for ubiquitination, there is only one E2-conjugating enzyme for sumoylation, which is Ubc9. In line with increasing evidence that sumoylation plays an important role in tumorigenesis, we recently ..
  19. Qin Y, Xu J, Aysola K, Begum N, Reddy V, Chai Y, et al. Ubc9 mediates nuclear localization and growth suppression of BRCA1 and BRCA1a proteins. J Cell Physiol. 2011;226:3355-67 pubmed publisher
    ..We have previously reported BRCA1 proteins unlike K109R and cancer-predisposing mutant C61G to bind Ubc9 and modulate ER-? turnover...
  20. Desterro J, Rodriguez M, Hay R. SUMO-1 modification of IkappaBalpha inhibits NF-kappaB activation. Mol Cell. 1998;2:233-9 pubmed
    ..Thus, while ubiquitination targets proteins for rapid degradation, SUMO-1 modification acts antagonistically to generate proteins resistant to degradation. ..
  21. Yunus A, Lima C. Lysine activation and functional analysis of E2-mediated conjugation in the SUMO pathway. Nat Struct Mol Biol. 2006;13:491-9 pubmed
    ..Genetic complementation revealed three side chains of the E2 Ubc9 that were crucial for normal growth...
  22. Boyer Guittaut M, Birsoy K, Potel C, Elliott G, Jaffray E, Desterro J, et al. SUMO-1 modification of human transcription factor (TF) IID complex subunits: inhibition of TFIID promoter-binding activity through SUMO-1 modification of hsTAF5. J Biol Chem. 2005;280:9937-45 pubmed
    ..Our observations suggest that reversible SUMO modification at hsTAF5 contributes to the dynamic regulation of TFIID promoter-binding activity in human cells. ..
  23. Pichler A, Knipscheer P, Oberhofer E, van Dijk W, Körner R, Olsen J, et al. SUMO modification of the ubiquitin-conjugating enzyme E2-25K. Nat Struct Mol Biol. 2005;12:264-9 pubmed
    ..In contrast, adjacent SUMO consensus sites are modified only when in unstructured peptides. The demonstration that secondary structure elements are part of SUMO attachment signals could contribute to a better prediction of SUMO targets. ..
  24. Kuo F, Bentsi Barnes I, Barlow G, Bae J, Pisarska M. Sumoylation of forkhead L2 by Ubc9 is required for its activity as a transcriptional repressor of the Steroidogenic Acute Regulatory gene. Cell Signal. 2009;21:1935-44 pubmed publisher
    ..In this study, we found that FOXL2 interacts with Ubc9, an E2-conjugating enzyme that mediates sumoylation, a key mechanism in transcriptional regulation...
  25. Lin X, Liang M, Liang Y, Brunicardi F, Feng X. SUMO-1/Ubc9 promotes nuclear accumulation and metabolic stability of tumor suppressor Smad4. J Biol Chem. 2003;278:31043-8 pubmed
    ..Smad4 levels, while inhibition of SUMOylation by small interfering RNA (siRNA)-mediated knockdown of the E2 enzyme Ubc9 reduces endogenous Smad4 levels...
  26. Argasinska J, Zhou K, Donnelly R, Hay R, Lee C. A functional interaction between RHA and Ubc9, an E2-like enzyme specific for Sumo-1. J Mol Biol. 2004;341:15-25 pubmed
    ..We previously reported that Ubc9, an E2-like enzyme specific for small ubiquitin-like modifier 1 (Sumo-1), is required for the interaction between ..
  27. Matic I, van Hagen M, Schimmel J, Macek B, Ogg S, Tatham M, et al. In vivo identification of human small ubiquitin-like modifier polymerization sites by high accuracy mass spectrometry and an in vitro to in vivo strategy. Mol Cell Proteomics. 2008;7:132-44 pubmed publisher
    ..The developed methodology is generic and can be adapted for the identification of other sumoylation sites in complex samples. ..
  28. Hsieh Y, Kuo H, Chang C, Naik M, Liao P, Ho C, et al. Ubc9 acetylation modulates distinct SUMO target modification and hypoxia response. EMBO J. 2013;32:791-804 pubmed publisher
    ..Here, we show that acetylation of SUMO E2 conjugase Ubc9 selectively downregulates the sumoylation of substrates with negatively charged amino acid-dependent sumoylation ..
  29. Kagey M, Melhuish T, Wotton D. The polycomb protein Pc2 is a SUMO E3. Cell. 2003;113:127-37 pubmed
    ..In vitro, CtBP sumoylation minimally requires the SUMO E1 and E2 (Ubc9) and SUMO-1. However, Pc2 dramatically enhances CtBP sumoylation...
  30. Mishra R, Jatiani S, Kumar A, Simhadri V, Hosur R, Mittal R. Dynamin interacts with members of the sumoylation machinery. J Biol Chem. 2004;279:31445-54 pubmed
    ..Here, we show that dynamin interacts with Sumo-1, Ubc9, and PIAS-1, all of which are members of the sumoylation machinery...
  31. Xu J, Watkins T, Reddy A, Reddy E, Rao V. A novel mechanism whereby BRCA1/1a/1b fine tunes the dynamic complex interplay between SUMO-dependent/independent activities of Ubc9 on E2-induced ERalpha activation/repression and degradation in breast cancer cells. Int J Oncol. 2009;34:939-49 pubmed
    ..Furthermore, we have found SUMO E2-conjugating enzyme Ubc9 to bind BRCA1 proteins...
  32. Zamborlini A, Coiffic A, Beauclair G, Delelis O, Paris J, Koh Y, et al. Impairment of human immunodeficiency virus type-1 integrase SUMOylation correlates with an early replication defect. J Biol Chem. 2011;286:21013-22 pubmed publisher
    ..Because SUMOylation-defective IN mutants retained WT catalytic activity, we hypothesize that SUMOylation might regulate the affinity of IN for co-factors, contributing to efficient HIV-1 replication. ..
  33. Lin D, Huang Y, Jeng J, Kuo H, Chang C, Chao T, et al. Role of SUMO-interacting motif in Daxx SUMO modification, subnuclear localization, and repression of sumoylated transcription factors. Mol Cell. 2006;24:341-54 pubmed
    ..Our results provide mechanistic insights into Daxx in SUMO-dependent transcriptional control and subnuclear compartmentalization. ..
  34. Meulmeester E, Kunze M, Hsiao H, Urlaub H, Melchior F. Mechanism and consequences for paralog-specific sumoylation of ubiquitin-specific protease 25. Mol Cell. 2008;30:610-9 pubmed publisher
    ..One mechanism for paralog-specific sumoylation may, thus, involve SIM-dependent recruitment of SUMO1 or SUMO2/3 thioester-charged Ubc9 to targets.
  35. Brunet Simioni M, de Thonel A, Hammann A, Joly A, Bossis G, Fourmaux E, et al. Heat shock protein 27 is involved in SUMO-2/3 modification of heat shock factor 1 and thereby modulates the transcription factor activity. Oncogene. 2009;28:3332-44 pubmed publisher
    ..As we also show that HSP27 binds to the SUMO-E2-conjugating enzyme, Ubc9, our study raises the possibility that HSP27 may act as a SUMO-E3 ligase specific for SUMO-2/3.
  36. Okuma T, Honda R, Ichikawa G, Tsumagari N, Yasuda H. In vitro SUMO-1 modification requires two enzymatic steps, E1 and E2. Biochem Biophys Res Commun. 1999;254:693-8 pubmed
    ..The conjugation steps are thought to be similar to those of ubiquitination; and human Ubc9, which is homologous to the E2 enzyme for the ubiquitin conjugation step, was identified and shown to be necessary ..
  37. Carbia Nagashima A, Gerez J, Perez Castro C, Paez Pereda M, Silberstein S, Stalla G, et al. RSUME, a small RWD-containing protein, enhances SUMO conjugation and stabilizes HIF-1alpha during hypoxia. Cell. 2007;131:309-23 pubmed
    ..enhancer (RSUME), that enhances overall SUMO-1, -2, and -3 conjugation by interacting with the SUMO conjugase Ubc9. RSUME increases noncovalent binding of SUMO-1 to Ubc9 and enhances Ubc9 thioester formation and SUMO ..
  38. Tong H, Hateboer G, Perrakis A, Bernards R, Sixma T. Crystal structure of murine/human Ubc9 provides insight into the variability of the ubiquitin-conjugating system. J Biol Chem. 1997;272:21381-7 pubmed
    Murine/human ubiquitin-conjugating enzyme Ubc9 is a functional homolog of Saccharomyces cerevisiae Ubc9 that is essential for the viability of yeast cells with a specific role in the G2-M transition of the cell cycle...
  39. Joanisse D, Inaguma Y, Tanguay R. Cloning and developmental expression of a nuclear ubiquitin-conjugating enzyme (DmUbc9) that interacts with small heat shock proteins in Drosophila melanogaster. Biochem Biophys Res Commun. 1998;244:102-9 pubmed
    ..that interact with small heat shock proteins (sHsps), a Drosophila melanogaster homologue of yeast and human ubc9 (Dmubc9) was found to interact with Drosophila Hsp23...
  40. Minty A, Dumont X, Kaghad M, Caput D. Covalent modification of p73alpha by SUMO-1. Two-hybrid screening with p73 identifies novel SUMO-1-interacting proteins and a SUMO-1 interaction motif. J Biol Chem. 2000;275:36316-23 pubmed
    ..Alternatively, it may modulate the interaction of p73 with other proteins that are substrates for SUMO-1 modification or which interact with SUMO-1, such as those identified here. ..
  41. Tatham M, Jaffray E, Vaughan O, Desterro J, Botting C, Naismith J, et al. Polymeric chains of SUMO-2 and SUMO-3 are conjugated to protein substrates by SAE1/SAE2 and Ubc9. J Biol Chem. 2001;276:35368-74 pubmed
    ..modifier SUMO-1/SMT3C/Sentrin-1 to proteins in vitro is dependent on a heterodimeric E1 (SAE1/SAE2) and an E2 (Ubc9)...
  42. Shiio Y, Eisenman R. Histone sumoylation is associated with transcriptional repression. Proc Natl Acad Sci U S A. 2003;100:13225-30 pubmed
    ..H4 binds to the SUMO-conjugating enzyme (E2), UBC9, and can be sumoylated in an E1 (SUMO-activating enzyme)- and E2-dependent manner...
  43. Lois L, Lima C. Structures of the SUMO E1 provide mechanistic insights into SUMO activation and E2 recruitment to E1. EMBO J. 2005;24:439-51 pubmed
    ..A mechanism for E2 recruitment to E1 is suggested by biochemical and genetic data, each of which supports a direct role for the E1 C-terminal ubiquitin-like domain for E2 recruitment during conjugation. ..
  44. Hecker C, Rabiller M, Haglund K, Bayer P, Dikic I. Specification of SUMO1- and SUMO2-interacting motifs. J Biol Chem. 2006;281:16117-27 pubmed
  45. Pan X, Li H, Zhang P, Jin B, Man J, Tian L, et al. Ubc9 interacts with SOX4 and represses its transcriptional activity. Biochem Biophys Res Commun. 2006;344:727-34 pubmed
    ..Deletion mapping demonstrated that HMG-box domain of SOX4 is required to mediate the interaction with Ubc9 in yeast. Furthermore, confocal microscopy showed that Ubc9 co-localized with SOX4 in the nucleus...
  46. Morris J, Boutell C, Keppler M, Densham R, Weekes D, Alamshah A, et al. The SUMO modification pathway is involved in the BRCA1 response to genotoxic stress. Nature. 2009;462:886-90 pubmed publisher
    ..to genotoxic stress, and co-localizes at sites of DNA damage with SUMO1, SUMO2/3 and the SUMO-conjugating enzyme Ubc9. PIAS SUMO E3 ligases co-localize with and modulate SUMO modification of BRCA1, and are required for BRCA1 ..
  47. Synowiec E, Krupa R, Morawiec Z, Wasylecka M, Dziki L, Morawiec J, et al. Efficacy of DNA double-strand breaks repair in breast cancer is decreased in carriers of the variant allele of the UBC9 gene c.73G>A polymorphism. Mutat Res. 2010;694:31-8 pubmed publisher
    b>UBC9 (E2) SUMO conjugating enzyme plays an important role in the maintenance of genome stability and integrity. In the present work we examined the association between the c...
  48. Gostissa M, Hengstermann A, Fogal V, Sandy P, Schwarz S, Scheffner M, et al. Activation of p53 by conjugation to the ubiquitin-like protein SUMO-1. EMBO J. 1999;18:6462-71 pubmed
    ..We suggest that posttranslational modification of p53 by SUMO-1 conjugation provides a novel mechanism to regulate p53 activity. ..
  49. Muller S, Berger M, Lehembre F, Seeler J, Haupt Y, Dejean A. c-Jun and p53 activity is modulated by SUMO-1 modification. J Biol Chem. 2000;275:13321-9 pubmed
    ..Taken together, these data provide a first link between the SUMO-1 conjugation pathway and the regulation of transcription factors. ..
  50. Mingot J, Kostka S, Kraft R, Hartmann E, Gorlich D. Importin 13: a novel mediator of nuclear import and export. EMBO J. 2001;20:3685-94 pubmed
  51. Kaul S, Blackford J, Cho S, Simons S. Ubc9 is a novel modulator of the induction properties of glucocorticoid receptors. J Biol Chem. 2002;277:12541-9 pubmed
    ..We now present evidence that the GMEBs contact Ubc9, which is the mammalian homolog of a yeast E2 ubiquitin-conjugating enzyme...
  52. Bernier Villamor V, Sampson D, Matunis M, Lima C. Structural basis for E2-mediated SUMO conjugation revealed by a complex between ubiquitin-conjugating enzyme Ubc9 and RanGAP1. Cell. 2002;108:345-56 pubmed
    ..In contrast to most ubiquitin conjugation, the SUMO E2 enzyme Ubc9 is sufficient for substrate recognition and lysine modification of known SUMO targets...
  53. Girdwood D, Bumpass D, Vaughan O, Thain A, Anderson L, Snowden A, et al. P300 transcriptional repression is mediated by SUMO modification. Mol Cell. 2003;11:1043-54 pubmed
    ..p300-mediated transcriptional activity and expression of a SUMO-specific protease or catalytically inactive Ubc9 relieved repression, demonstrating that p300 repression was mediated by SUMO conjugation...
  54. Tatham M, Kim S, Yu B, Jaffray E, Song J, Zheng J, et al. Role of an N-terminal site of Ubc9 in SUMO-1, -2, and -3 binding and conjugation. Biochemistry. 2003;42:9959-69 pubmed
    ..ubiquitin-like proteins SUMO-1, -2, and -3 interact with the same N-terminal region of the E2 conjugating enzyme Ubc9 with similar affinities...
  55. Kagey M, Melhuish T, Powers S, Wotton D. Multiple activities contribute to Pc2 E3 function. EMBO J. 2005;24:108-19 pubmed
    ..However, a carboxyl-terminal fragment of Pc2 that recruits both Ubc9 and CtBP lacks E3 activity...
  56. Kobayashi S, Shibata H, Yokota K, Suda N, Murai A, Kurihara I, et al. FHL2, UBC9, and PIAS1 are novel estrogen receptor alpha-interacting proteins. Endocr Res. 2004;30:617-21 pubmed
    ..We have also detected strong interactions of ERalpha with Ubc9 and PIAS1 in yeast...
  57. Weger S, Hammer E, Heilbronn R. Topors acts as a SUMO-1 E3 ligase for p53 in vitro and in vivo. FEBS Lett. 2005;579:5007-12 pubmed
    ..In HeLa cells, Topors induced p53 sumoylation was accompanied by an increase in endogenous p53 protein levels. Furthermore, Topors enhances the sumoylation of a variety of other, yet unidentified, cellular proteins. ..
  58. Benson M, Li Q, Kieckhafer K, Dudek D, Whorton M, Sunahara R, et al. SUMO modification regulates inactivation of the voltage-gated potassium channel Kv1.5. Proc Natl Acad Sci U S A. 2007;104:1805-10 pubmed
    ..We find that Kv1.5 interacts specifically with the SUMO-conjugating enzyme Ubc9 and is a target for modification by SUMO-1, -2, and -3 in vivo. In addition, purified recombinant Kv1...
  59. Riising E, Boggio R, Chiocca S, Helin K, Pasini D. The polycomb repressive complex 2 is a potential target of SUMO modifications. PLoS ONE. 2008;3:e2704 pubmed publisher
    ..Furthermore, we show that SUZ12 interacts with the E2-conjugating enzyme UBC9 both in vitro and in vivo and that mutation of the SUZ12 sumoylation site does not abolish this binding...
  60. Martinez N, Xue X, Berro R, Kreitzer G, Resh M. Kinesin KIF4 regulates intracellular trafficking and stability of the human immunodeficiency virus type 1 Gag polyprotein. J Virol. 2008;82:9937-50 pubmed publisher
    ..transiently accumulated in discrete, perinuclear, nonendocytic clusters that colocalized with endogenous KIF4, with Ubc9, an E2 SUMO-1 conjugating enzyme, and with SUMO...
  61. Knipscheer P, Flotho A, Klug H, Olsen J, van Dijk W, Fish A, et al. Ubc9 sumoylation regulates SUMO target discrimination. Mol Cell. 2008;31:371-82 pubmed publisher
    ..Here we show that autosumoylation of the mammalian E2-conjugating enzyme Ubc9 at Lys14 regulates target discrimination...
  62. Jaber T, Bohl C, Lewis G, Wood C, West J, Weldon R. Human Ubc9 contributes to production of fully infectious human immunodeficiency virus type 1 virions. J Virol. 2009;83:10448-59 pubmed publisher
    b>Ubc9 was identified as a cellular protein that interacts with the Gag protein of Mason-Pfizer monkey virus...
  63. Driscoll J, Pelluru D, Lefkimmiatis K, Fulciniti M, Prabhala R, Greipp P, et al. The sumoylation pathway is dysregulated in multiple myeloma and is associated with adverse patient outcome. Blood. 2010;115:2827-34 pubmed publisher
    ..The Sumo-conjugating enzyme Ube2I, the Sumo-ligase PIAS1, and the Sumo-inducer ARF were elevated in MM patient samples and cell lines...
  64. Chu Y, Yang X. SUMO E3 ligase activity of TRIM proteins. Oncogene. 2011;30:1108-16 pubmed publisher
    ..These TRIM proteins bind both the SUMO-conjugating enzyme Ubc9 and substrates and strongly enhance transfer of SUMOs from Ubc9 to these substrates...
  65. Boutell C, Cuchet Louren o D, Vanni E, Orr A, Glass M, McFarlane S, et al. A viral ubiquitin ligase has substrate preferential SUMO targeted ubiquitin ligase activity that counteracts intrinsic antiviral defence. PLoS Pathog. 2011;7:e1002245 pubmed publisher
    ..We conclude that ICP0 has dual targeting mechanisms involving both SUMO- and substrate-dependent targeting specificities in order to counteract intrinsic antiviral resistance to HSV-1 infection...
  66. Poukka H, Aarnisalo P, Karvonen U, Palvimo J, Janne O. Ubc9 interacts with the androgen receptor and activates receptor-dependent transcription. J Biol Chem. 1999;274:19441-6 pubmed
    b>Ubc9, a homologue of the class E2 ubiquitin-conjugating enzymes, has recently been shown to catalyze conjugation of a small ubiquitin-like molecule-1 (SUMO-1) to a variety of target proteins...
  67. Rodriguez M, Desterro J, Lain S, Midgley C, Lane D, Hay R. SUMO-1 modification activates the transcriptional response of p53. EMBO J. 1999;18:6455-61 pubmed
    ..Modification in vitro requires only SUMO-1, the SUMO-1 activating enzyme and ubc9. SUMO-1 and ubiquitin modification do not compete for the same lysine acceptor sites in p53...
  68. Bies J, Markus J, Wolff L. Covalent attachment of the SUMO-1 protein to the negative regulatory domain of the c-Myb transcription factor modifies its stability and transactivation capacity. J Biol Chem. 2002;277:8999-9009 pubmed
    ..In accordance with this observation, we found that the SUMO-1-conjugating enzyme Ubc9 interacted only with a region surrounding Lys(523) (also called the PEST/EVES motif)...
  69. Kurihara I, Shibata H, Kobayashi S, Suda N, Ikeda Y, Yokota K, et al. Ubc9 and Protein Inhibitor of Activated STAT 1 Activate Chicken Ovalbumin Upstream Promoter-Transcription Factor I-mediated Human CYP11B2 Gene Transcription. J Biol Chem. 2005;280:6721-30 pubmed
    ..for COUP-TFI-interacting proteins from a human adrenal cDNA library using a yeast two-hybrid system and identified Ubc9 and PIAS1, which have small ubiquitin-related modifier-1 (SUMO-1) conjugase and ligase activities, respectively...
  70. Bossis G, Malnou C, Farras R, Andermarcher E, Hipskind R, Rodriguez M, et al. Down-regulation of c-Fos/c-Jun AP-1 dimer activity by sumoylation. Mol Cell Biol. 2005;25:6964-79 pubmed
    ..This supports the idea that this modification does not constitute a final inactivation step that necessarily precedes protein degradation. ..
  71. Moschos S, Smith A, Mandic M, Athanassiou C, Watson Hurst K, Jukic D, et al. SAGE and antibody array analysis of melanoma-infiltrated lymph nodes: identification of Ubc9 as an important molecule in advanced-stage melanomas. Oncogene. 2007;26:4216-25 pubmed
    ..Equally important, we provide evidence that the ubiquitin-conjugating enzyme, Ubc9, is expressed at high levels in melanoma-positive lymph nodes, and that it plays a crucial role in preventing ..
  72. Guo Y, Yang M, Weissler J, Yang Y. Modulation of PLAGL2 transactivation activity by Ubc9 co-activation not SUMOylation. Biochem Biophys Res Commun. 2008;374:570-5 pubmed publisher
    ..Unexpectedly, the SUMO-conjugating enzyme Ubc9 enhanced, rather than reduced, PLAGL2 activated promoter activity but did not affect TTF-1 activation of the ..
  73. Matic I, Schimmel J, Hendriks I, van Santen M, van de Rijke F, van Dam H, et al. Site-specific identification of SUMO-2 targets in cells reveals an inverted SUMOylation motif and a hydrophobic cluster SUMOylation motif. Mol Cell. 2010;39:641-52 pubmed publisher
    ..In 16 proteins we identified a hydrophobic cluster SUMOylation motif (HCSM). SUMO conjugation of RanGAP1 and ZBTB1 via HCSMs is remarkably efficient. ..
  74. Chen S, Gong C, Luo M, Yao H, Zeng Y, Su F. Ubc9 expression predicts chemoresistance in breast cancer. Chin J Cancer. 2011;30:638-44 pubmed publisher
    Ubiquitin-conjugating enzyme 9 (Ubc9), the sole conjugating enzyme for sumoylation, regulates protein function and plays an important role in tumorigenesis. Whether Ubc9 is involved in the chemoresistance of breast cancer remains unknown...
  75. Pichler A, Gast A, Seeler J, Dejean A, Melchior F. The nucleoporin RanBP2 has SUMO1 E3 ligase activity. Cell. 2002;108:109-20 pubmed
    ..SUMOylation requires the E1 enzyme Aos1/Uba2 and the E2 enzyme Ubc9. A family of E3-like factors, PIAS proteins, was discovered recently...
  76. Schmidt D, Muller S. Members of the PIAS family act as SUMO ligases for c-Jun and p53 and repress p53 activity. Proc Natl Acad Sci U S A. 2002;99:2872-7 pubmed
    ..pathway that is mechanistically analogous to ubiquitination, but requires a different E1-activating enzyme and Ubc9, a SUMO-specific E2-conjugating enzyme...
  77. Sapetschnig A, Rischitor G, Braun H, Doll A, Schergaut M, Melchior F, et al. Transcription factor Sp3 is silenced through SUMO modification by PIAS1. EMBO J. 2002;21:5206-15 pubmed
    ..We identified the protein inhibitor of activated STAT1 (PIAS1) as an interaction partner of Sp3 and Ubc9. Moreover, PIAS1 strongly stimulated SUMO conjugation to Sp3, thus acting as an E3 ligase for SUMO conjugation to ..
  78. Ross S, Best J, Zon L, Gill G. SUMO-1 modification represses Sp3 transcriptional activation and modulates its subnuclear localization. Mol Cell. 2002;10:831-42 pubmed
    ..These studies reveal a direct effect of SUMO-1 modification on activity of a dual function transcription factor and provide a mechanism for functional specificity within the Sp transcription factor family. ..
  79. Tatham M, Chen Y, Hay R. Role of two residues proximal to the active site of Ubc9 in substrate recognition by the Ubc9.SUMO-1 thiolester complex. Biochemistry. 2003;42:3168-79 pubmed
    ..residues on target proteins by a specific conjugation pathway involving the E1 enzyme SAE1/SAE2 and the E2 enzyme Ubc9. In an ATP-dependent manner, the C-terminus of SUMO-1 forms consecutive thiolester bonds with cysteine residues in ..
  80. Weger S, Hammer E, Engstler M. The DNA topoisomerase I binding protein topors as a novel cellular target for SUMO-1 modification: characterization of domains necessary for subcellular localization and sumolation. Exp Cell Res. 2003;290:13-27 pubmed
    ..In a yeast two-hybrid system, Topors interacted with both SUMO-1 and the SUMO-1 conjugating enzyme UBC9. Multiple SUMO-1 modified forms of Topors could be detected after cotransfection of exogenous SUMO-1 and Topors ..
  81. Gurer C, Berthoux L, Luban J. Covalent modification of human immunodeficiency virus type 1 p6 by SUMO-1. J Virol. 2005;79:910-7 pubmed
    ..p6 was found to interact with small ubiquitin-like modifier 1 (SUMO-1) as well as the E2 SUMO-1 transfer enzyme, Ubc9. Interaction with p6 was also detected with Daxx, a cellular protein to which SUMO-1 is sometimes covalently ..
  82. Eladad S, Ye T, Hu P, Leversha M, Beresten S, Matunis M, et al. Intra-nuclear trafficking of the BLM helicase to DNA damage-induced foci is regulated by SUMO modification. Hum Mol Genet. 2005;14:1351-65 pubmed
    ..These results constitute evidence that BLM is a DNA damage sensor that signals the formation of DDI, and they establish SUMO modification as a negative regulator of BLM's signaling function. ..
  83. Bossis G, Melchior F. Regulation of SUMOylation by reversible oxidation of SUMO conjugating enzymes. Mol Cell. 2006;21:349-57 pubmed
    ..a) disulfide bond(s) involving the catalytic cysteines of the SUMO E1 subunit Uba2 and the E2-conjugating enzyme Ubc9. The same phenomenon is also observed in a physiological scenario of endogenous ROS production, the respiratory ..
  84. Ivanov A, Peng H, Yurchenko V, Yap K, Negorev D, Schultz D, et al. PHD domain-mediated E3 ligase activity directs intramolecular sumoylation of an adjacent bromodomain required for gene silencing. Mol Cell. 2007;28:823-37 pubmed
    ..The RING finger-like structure of the PHD domain is required for both Ubc9 binding and sumoylation and directs modification to specific lysine residues in the bromodomain...
  85. Dünnebier T, Bermejo J, Haas S, Fischer H, Pierl C, Justenhoven C, et al. Polymorphisms in the UBC9 and PIAS3 genes of the SUMO-conjugating system and breast cancer risk. Breast Cancer Res Treat. 2010;121:185-94 pubmed publisher
    ..We recently reported that variation in the UBC9 gene encoding the SUMO-conjugating enzyme may affect the grade of breast tumors...
  86. Chang C, Naik M, Huang Y, Jeng J, Liao P, Kuo H, et al. Structural and functional roles of Daxx SIM phosphorylation in SUMO paralog-selective binding and apoptosis modulation. Mol Cell. 2011;42:62-74 pubmed publisher
    ..Our findings provide structural insights into the Daxx-SIM:SUMO-1 complex, a model of SIM phosphorylation-enhanced SUMO paralog-selective modification and interaction, and phosphorylation-regulated Daxx function in apoptosis. ..
  87. Tomasi M, Tomasi I, Ramani K, Pascale R, Xu J, Giordano P, et al. S-adenosyl methionine regulates ubiquitin-conjugating enzyme 9 protein expression and sumoylation in murine liver and human cancers. Hepatology. 2012;56:982-93 pubmed publisher
    Ubiquitin-conjugating enzyme 9 (Ubc9) is required for sumoylation and is overexpressed in several malignancies, but its expression in hepatocellular carcinoma (HCC) is unknown...
  88. Desterro J, Thomson J, Hay R. Ubch9 conjugates SUMO but not ubiquitin. FEBS Lett. 1997;417:297-300 pubmed
    Ubiquitin conjugating enzymes participate in the thioester cascade that leads to protein ubiquitination. Although Ubc9 is homologous to E2 ubiquitin conjugating enzymes we have shown that it is unable to form a thioester with ubiquitin, ..
  89. Vertegaal A, Ogg S, Jaffray E, Rodriguez M, Hay R, Andersen J, et al. A proteomic study of SUMO-2 target proteins. J Biol Chem. 2004;279:33791-8 pubmed
    ..SART1 and heterogeneous nuclear RNP M were both shown to be genuine SUMO targets, confirming the validity of the approach. ..
  90. Mo Y, Yu Y, Theodosiou E, Ee P, Beck W. A role for Ubc9 in tumorigenesis. Oncogene. 2005;24:2677-83 pubmed
    ..b>Ubc9 is a sole E2-conjugating enzyme essential for sumoylation...
  91. Cheng T, Chang L, Howng S, Lu P, Lee C, Hong Y. SUMO-1 modification of centrosomal protein hNinein promotes hNinein nuclear localization. Life Sci. 2006;78:1114-20 pubmed
    ..of human ninein in centrosome, we performed yeast two-hybrid screen and isolated the SUMO-conjugating E2 enzyme, Ubc9, and SUMOylation enhancing enzymes, including PIAS1 and PIASxalpha, as binding partners of hNinein...
  92. Tomoiu A, Gravel A, Tanguay R, Flamand L. Functional interaction between human herpesvirus 6 immediate-early 2 protein and ubiquitin-conjugating enzyme 9 in the absence of sumoylation. J Virol. 2006;80:10218-28 pubmed publisher
    ..The most frequently isolated IE2-interacting protein was the human ubiquitin-conjugating enzyme 9 (Ubc9), a protein involved in the small ubiquitin-like modifier (SUMO) conjugation pathway...