Gene Symbol: UBC9
Description: E2 SUMO-conjugating protein UBC9
Alias: E2 SUMO-conjugating protein UBC9
Species: Saccharomyces cerevisiae S288c
Products:     UBC9

Top Publications

  1. Takahashi Y, Kikuchi Y. Yeast PIAS-type Ull1/Siz1 is composed of SUMO ligase and regulatory domains. J Biol Chem. 2005;280:35822-8 pubmed
    ..Taken together, we conclude that the SP-RING and PINIT of Ull1/Siz1 are core domains of the SUMO ligase, and the other domains are regulatory for protein stability and subcellular localization. ..
  2. Windecker H, Ulrich H. Architecture and assembly of poly-SUMO chains on PCNA in Saccharomyces cerevisiae. J Mol Biol. 2008;376:221-31 pubmed
  3. 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...
  4. Yousef A, Fonseca G, Pelka P, Ablack J, Walsh C, Dick F, et al. Identification of a molecular recognition feature in the E1A oncoprotein that binds the SUMO conjugase UBC9 and likely interferes with polySUMOylation. Oncogene. 2010;29:4693-704 pubmed publisher
    ..This MoRF is necessary and sufficient to bind the N-terminal region of the SUMO conjugase UBC9, which also interacts with SUMO noncovalently and is involved in polySUMOylation...
  5. Johnson E, Blobel G. Ubc9p is the conjugating enzyme for the ubiquitin-like protein Smt3p. J Biol Chem. 1997;272:26799-802 pubmed
    ..cerevisiae. These results suggest that, like ubiquitination, Smt3p conjugation may be a critical modification in cell cycle regulation. ..
  6. Sollier J, Driscoll R, Castellucci F, Foiani M, Jackson S, Branzei D. The Saccharomyces cerevisiae Esc2 and Smc5-6 proteins promote sister chromatid junction-mediated intra-S repair. Mol Biol Cell. 2009;20:1671-82 pubmed publisher
    ..sister chromatid junctions at damaged replication forks, and its functionality seems to be regulated by Ubc9- and Mms21-dependent sumoylation...
  7. Hoege C, Pfander B, Moldovan G, Pyrowolakis G, Jentsch S. RAD6-dependent DNA repair is linked to modification of PCNA by ubiquitin and SUMO. Nature. 2002;419:135-41 pubmed
    ..Here we show that UBC9, a small ubiquitin-related modifier (SUMO)-conjugating enzyme, is also affiliated with this pathway and that ..
  8. Bencsath K, Podgorski M, Pagala V, Slaughter C, Schulman B. Identification of a multifunctional binding site on Ubc9p required for Smt3p conjugation. J Biol Chem. 2002;277:47938-45 pubmed
    ..b>Ubc9, the E2 for SUMO, synthesizes polymeric chains in the presence of its E1 and MgATP...
  9. Mullen J, Brill S. Activation of the Slx5-Slx8 ubiquitin ligase by poly-small ubiquitin-like modifier conjugates. J Biol Chem. 2008;283:19912-21 pubmed publisher
    ..Slx5-Slx8 primarily mono-ubiquitinated the N-terminal SUMO moiety of the chain. These data indicate that the Slx5-Slx8 Ub ligase is stimulated by poly-SUMO conjugates and that it can ubiquitinate a poly-SUMO chain. ..

More Information


  1. van Waardenburg R, Duda D, Lancaster C, Schulman B, Bjornsti M. Distinct functional domains of Ubc9 dictate cell survival and resistance to genotoxic stress. Mol Cell Biol. 2006;26:4958-69 pubmed
    ..Target recognition is determined, in part, by the SUMO E2 enzyme, Ubc9, while Siz/Pias E3 ligases may facilitate select interactions by acting as substrate adaptors...
  2. Jiang W, Koltin Y. Two-hybrid interaction of a human UBC9 homolog with centromere proteins of Saccharomyces cerevisiae. Mol Gen Genet. 1996;251:153-60 pubmed
    ..Overexpression of hUBC9 partially suppresses a S. cerevisiae ubc9 temperature-sensitive mutation, indicating that the UBC9 gene family is also functionally conserved...
  3. Vigasová D, Sarangi P, Kolesar P, Vlasáková D, Slezakova Z, Altmannova V, et al. Lif1 SUMOylation and its role in non-homologous end-joining. Nucleic Acids Res. 2013;41:5341-53 pubmed publisher
    ..Taken together, these findings suggest that SUMOylation of Lif1 represents a new regulatory mechanism that downregulates NHEJ in a cell cycle phase-independent manner. ..
  4. Cremona C, Sarangi P, Yang Y, Hang L, Rahman S, Zhao X. Extensive DNA damage-induced sumoylation contributes to replication and repair and acts in addition to the mec1 checkpoint. Mol Cell. 2012;45:422-32 pubmed publisher
    ..We also show that DNA damage-induced sumoylation does not require Mec1 checkpoint signaling, and the presence of both enables optimal DNA damage resistance. ..
  5. Parker J, Bucceri A, Davies A, Heidrich K, Windecker H, Ulrich H. SUMO modification of PCNA is controlled by DNA. EMBO J. 2008;27:2422-31 pubmed publisher
    ..Instead, the stimulatory effect of DNA on conjugation is mainly attributable to DNA binding of PCNA itself. These findings imply a change in the properties of PCNA upon loading that enhances its capacity to be sumoylated. ..
  6. Meednu N, Hoops H, D Silva S, Pogorzala L, Wood S, Farkas D, et al. The spindle positioning protein Kar9p interacts with the sumoylation machinery in Saccharomyces cerevisiae. Genetics. 2008;180:2033-55 pubmed publisher
    ..We propose a model in which sumoylation regulates spindle positioning by restricting Kar9p to one SPB. These findings raise the possibility that sumoylation could regulate other microtubule-dependent processes. ..
  7. Johnson E, Gupta A. An E3-like factor that promotes SUMO conjugation to the yeast septins. Cell. 2001;106:735-44 pubmed
    ..Our results suggest that this family functions by promoting SUMO conjugation to specific substrates. ..
  8. Takahashi Y, Kahyo T, Toh e A, Yasuda H, Kikuchi Y. Yeast Ull1/Siz1 is a novel SUMO1/Smt3 ligase for septin components and functions as an adaptor between conjugating enzyme and substrates. J Biol Chem. 2001;276:48973-7 pubmed
    ..the SUMO1/Smt3 modification of septin components was abolished in the ull1 mutant and Ull1 associated with E2 (Ubc9) and the substrates (septin components) in immunoprecipitation experiments...
  9. Schwarz S, Matuschewski K, Liakopoulos D, Scheffner M, Jentsch S. The ubiquitin-like proteins SMT3 and SUMO-1 are conjugated by the UBC9 E2 enzyme. Proc Natl Acad Sci U S A. 1998;95:560-4 pubmed
    ..Herein we show that UBC9, one of the 13 different E2 enzymes from yeast, is required for SMT3 conjugation in vivo...
  10. Takahashi Y, Toh e A, Kikuchi Y. Comparative analysis of yeast PIAS-type SUMO ligases in vivo and in vitro. J Biochem. 2003;133:415-22 pubmed
    ..It interacted with Smt3, SUMO/Smt3 conjugating enzyme Ubc9 and a septin component Cdc3 in the two-hybrid system...
  11. Yunus A, Lima C. Structure of the Siz/PIAS SUMO E3 ligase Siz1 and determinants required for SUMO modification of PCNA. Mol Cell. 2009;35:669-82 pubmed publisher
    ..Mutational analysis of Siz1 and PCNA revealed surfaces on both proteins that are required for efficient SUMO modification of PCNA in vitro and in vivo. ..
  12. Duan X, Sarangi P, Liu X, Rangi G, Zhao X, Ye H. Structural and functional insights into the roles of the Mms21 subunit of the Smc5/6 complex. Mol Cell. 2009;35:657-68 pubmed publisher
    ..Through structure-based dissection of Mms21 functions, our studies establish a framework for understanding its roles in the Smc5/6 complex. ..
  13. Mullen J, Chen C, Brill S. Wss1 is a SUMO-dependent isopeptidase that interacts genetically with the Slx5-Slx8 SUMO-targeted ubiquitin ligase. Mol Cell Biol. 2010;30:3737-48 pubmed publisher
    ..The results suggest that Wss1 is a SUMO-dependent isopeptidase that acts on sumoylated substrates as they undergo proteasomal degradation. ..
  14. Burgess R, Rahman S, Lisby M, Rothstein R, Zhao X. The Slx5-Slx8 complex affects sumoylation of DNA repair proteins and negatively regulates recombination. Mol Cell Biol. 2007;27:6153-62 pubmed
    ..In addition, the complex inhibits Rad51-independent recombination via modulating the sumoylation of DNA repair proteins. ..
  15. Duda D, van Waardenburg R, Borg L, McGarity S, Nourse A, Waddell M, et al. Structure of a SUMO-binding-motif mimic bound to Smt3p-Ubc9p: conservation of a non-covalent ubiquitin-like protein-E2 complex as a platform for selective interactions within a SUMO pathway. J Mol Biol. 2007;369:619-30 pubmed
    ..In the complex, Smt3p binds a surface distal from Ubc9's catalytic cysteine...
  16. Branzei D, Sollier J, Liberi G, Zhao X, Maeda D, Seki M, et al. Ubc9- and mms21-mediated sumoylation counteracts recombinogenic events at damaged replication forks. Cell. 2006;127:509-22 pubmed
    The Ubc9 SUMO-conjugating enzyme and the Siz1 SUMO ligase sumoylate several repair and recombination proteins, including PCNA. Sumoylated PCNA binds Srs2, a helicase counteracting certain recombination events...
  17. Xie Y, Rubenstein E, Matt T, Hochstrasser M. SUMO-independent in vivo activity of a SUMO-targeted ubiquitin ligase toward a short-lived transcription factor. Genes Dev. 2010;24:893-903 pubmed publisher
    ..We suggest that alpha2, and presumably other proteins, have surface features that mimic SUMO, and therefore can directly recruit STUbLs without prior SUMO conjugation. ..
  18. Zhao X, Blobel G. A SUMO ligase is part of a nuclear multiprotein complex that affects DNA repair and chromosomal organization. Proc Natl Acad Sci U S A. 2005;102:4777-82 pubmed
    ..We propose that Mms21 sumoylates proteins involved in these diverse processes and that the other members of the complex, particularly Smc5/6, facilitate proper substrate sumoylation by localizing Mms21 to specific chromosomal regions. ..
  19. Pfander B, Moldovan G, Sacher M, Hoege C, Jentsch S. SUMO-modified PCNA recruits Srs2 to prevent recombination during S phase. Nature. 2005;436:428-33 pubmed
    ..Our finding suggests a model in which SUMO-modified PCNA recruits Srs2 in S phase in order to prevent unwanted recombination events of replicating chromosomes. ..
  20. Psakhye I, Jentsch S. Protein group modification and synergy in the SUMO pathway as exemplified in DNA repair. Cell. 2012;151:807-820 pubmed publisher
    ..We propose that SUMOylation may thus often target a protein group rather than individual proteins, whereas localized modification enzymes and highly specific triggers ensure specificity. ..
  21. Montpetit B, Hazbun T, Fields S, Hieter P. Sumoylation of the budding yeast kinetochore protein Ndc10 is required for Ndc10 spindle localization and regulation of anaphase spindle elongation. J Cell Biol. 2006;174:653-63 pubmed
    ..These data suggest that sumoylation of Ndc10 and other kinetochore proteins play a critical role during the mitotic process. ..
  22. Kolesar P, Sarangi P, Altmannova V, Zhao X, Krejci L. Dual roles of the SUMO-interacting motif in the regulation of Srs2 sumoylation. Nucleic Acids Res. 2012;40:7831-43 pubmed publisher
    ..The SIM region is required for Srs2 sumoylation, likely by binding to SUMO-charged Ubc9. Srs2's SIM also cooperates with an adjacent PCNA-specific interaction site in binding to sumoylated PCNA to ensure ..
  23. Takahashi Y, Toh e A, Kikuchi Y. A novel factor required for the SUMO1/Smt3 conjugation of yeast septins. Gene. 2001;275:223-31 pubmed
    ..A conserved cysteine residue in the domain was essential for this conjugation. Furthermore, Siz1 was localized at the mother-bud neck in the M-phase and physically bound to both E2 and the target proteins. ..
  24. Moldovan G, Dejsuphong D, Petalcorin M, Hofmann K, Takeda S, Boulton S, et al. Inhibition of homologous recombination by the PCNA-interacting protein PARI. Mol Cell. 2012;45:75-86 pubmed publisher
    ..Thus, we propose that PARI is a long sought-after factor that suppresses inappropriate recombination events at mammalian replication forks...
  25. Seufert W, Futcher B, Jentsch S. Role of a ubiquitin-conjugating enzyme in degradation of S- and M-phase cyclins. Nature. 1995;373:78-81 pubmed
    ..Here we show that B-type-cyclin degradation in yeast involves an essential nuclear ubiquitin-conjugating enzyme, UBC9. Repression of UBC9 synthesis prevents cell cycle progression at the G2 or early M phase, causing the accumulation ..
  26. Ho C, Chen H, Hwang J. UBC9 autosumoylation negatively regulates sumoylation of septins in Saccharomyces cerevisiae. J Biol Chem. 2011;286:21826-34 pubmed publisher
    ..that two lysine residues (Lys-153 and Lys-157) in the C-terminal region of the yeast E2-conjugating enzyme Ubc9 are the major and minor autosumoylation sites, respectively...
  27. Klug H, Xaver M, Chaugule V, Koidl S, Mittler G, Klein F, et al. Ubc9 sumoylation controls SUMO chain formation and meiotic synapsis in Saccharomyces cerevisiae. Mol Cell. 2013;50:625-36 pubmed publisher
    ..Here, we describe modification of the budding yeast E2 enzyme Ubc9 by sumoylation (Ubc9(*)SUMO)...
  28. Böhm S, Mihalevic M, Casal M, Bernstein K. Disruption of SUMO-targeted ubiquitin ligases Slx5-Slx8/RNF4 alters RecQ-like helicase Sgs1/BLM localization in yeast and human cells. DNA Repair (Amst). 2015;26:1-14 pubmed publisher
    ..Our results point to a model where RecQ-like helicase subcellular localization is regulated by STUbLs in response to DNA damage, presumably to prevent illegitimate recombination events. ..
  29. Takahashi S, Araki Y, Ohya Y, Sakuno T, Hoshino S, Kontani K, et al. Upf1 potentially serves as a RING-related E3 ubiquitin ligase via its association with Upf3 in yeast. RNA. 2008;14:1950-8 pubmed publisher
    ..These results suggest that Upf1 may serve as an E3 ubiquitin ligase upon its association with Upf3 and play an important role in signaling to the NMD pathway. ..
  30. Silva S, Altmannova V, Eckert Boulet N, Kolesar P, Gallina I, Hang L, et al. SUMOylation of Rad52-Rad59 synergistically change the outcome of mitotic recombination. DNA Repair (Amst). 2016;42:11-25 pubmed publisher
    ..In some genetic assays, Rad52 and Rad59 SUMOylation act synergistically. Collectively, our data indicate that the described SUMO modifications affect the balance between conservative and non-conservative mechanisms of HR. ..
  31. Tan W, Wang Z, Prelich G. Physical and Genetic Interactions Between Uls1 and the Slx5-Slx8 SUMO-Targeted Ubiquitin Ligase. G3 (Bethesda). 2013;3:771-780 pubmed publisher
  32. Parker J, Ulrich H. SIM-dependent enhancement of substrate-specific SUMOylation by a ubiquitin ligase in vitro. Biochem J. 2014;457:435-40 pubmed publisher
    ..of enhancing PCNA SUMOylation in a SIM-dependent manner in vitro, most probably by means of directing SUMO-loaded Ubc9 (ubiquitin-conjugating enzyme 9) towards the substrate...
  33. Klenk C, Humrich J, Quitterer U, Lohse M. SUMO-1 controls the protein stability and the biological function of phosducin. J Biol Chem. 2006;281:8357-64 pubmed
    ..Together, these findings show that phosducin is a previously unrecognized target of SUMO modification and that SUMOylation controls phosducin stability in cells as well as its functional properties. ..
  34. Jacquiau H, van Waardenburg R, Reid R, Woo M, Guo H, Johnson E, et al. Defects in SUMO (small ubiquitin-related modifier) conjugation and deconjugation alter cell sensitivity to DNA topoisomerase I-induced DNA damage. J Biol Chem. 2005;280:23566-75 pubmed
    ..Benedetti, P., and Bjornsti, M. A. (2004) J. Biol. Chem. 279, 21271-21281). Here we report that tah mutant ubc9-10 harbors a hypomorphic allele of UBC9, which encodes the essential SUMO (small ubiquitin-related modifier) E2-..
  35. Chen Y, Chuang Y, Chuang C, Cheng Y, Chang C, Leng C, et al. S. cerevisiae Mre11 recruits conjugated SUMO moieties to facilitate the assembly and function of the Mre11-Rad50-Xrs2 complex. Nucleic Acids Res. 2016;44:2199-213 pubmed publisher
    ..Mre11(SIM1) is indispensable for MRX assembly. Mre11(SIM2) non-covalently links MRX with the SUMO enzymes (E2/Ubc9 and E3/Siz2) to promote global SUMOylation of DNA repair proteins...
  36. Streich F, Lima C. Capturing a substrate in an activated RING E3/E2-SUMO complex. Nature. 2016;536:304-8 pubmed
  37. Bermúdez López M, Pociño Merino I, Sánchez H, Bueno A, Guasch C, Almedawar S, et al. ATPase-dependent control of the Mms21 SUMO ligase during DNA repair. PLoS Biol. 2015;13:e1002089 pubmed publisher
    ..sumoylation requires binding of ATP to Smc5, a step that is part of the ligase mechanism that assists Ubc9 function...
  38. Cheng C, Lo Y, Liang S, Ti S, Lin F, Yeh C, et al. SUMO modifications control assembly of synaptonemal complex and polycomplex in meiosis of Saccharomyces cerevisiae. Genes Dev. 2006;20:2067-81 pubmed
    ..At and after mid-prophase, the Zip1 protein begins to associate with Zip3-dependent Smt3 conjugates (e.g., Red1) along meiotic chromosomes in the wild-type cell to form SCs and with Smt3 polymeric chains in the zip3 mutant to form PCs...
  39. Mullen J, Das M, Brill S. Genetic evidence that polysumoylation bypasses the need for a SUMO-targeted Ub ligase. Genetics. 2011;187:73-87 pubmed publisher
    ..In support of this latter possibility we find that the WSS1 isopeptidase is required for suppression by ulp2?. ..
  40. Felberbaum R, Wilson N, Cheng D, Peng J, Hochstrasser M. Desumoylation of the endoplasmic reticulum membrane VAP family protein Scs2 by Ulp1 and SUMO regulation of the inositol synthesis pathway. Mol Cell Biol. 2012;32:64-75 pubmed publisher
    ..These results provide the first evidence of cross-regulation between the SUMO and inositol pathways, including the sumoylation of an ER membrane protein central to phospholipid synthesis and phosphoinositide signaling...
  41. Labedzka K, Tian C, Nussbaumer U, Timmermann S, Walther P, Müller J, et al. Sho1p connects the plasma membrane with proteins of the cytokinesis network through multiple isomeric interaction states. J Cell Sci. 2012;125:4103-13 pubmed publisher
    ..Owing to the overlapping binding specificities of its members the HICS complex is best described as ensembles of isomeric interaction states that precisely coordinate the different functions of the interactors during cytokinesis. ..
  42. Novoselova T, Rose R, Marks H, Sullivan J. SUMOylation regulates the homologous to E6-AP carboxyl terminus (HECT) ubiquitin ligase Rsp5p. J Biol Chem. 2013;288:10308-17 pubmed publisher
    ..This reciprocal regulation of these highly conserved ligases represents an exciting and previously unidentified system of cross talk between the ubiquitin and SUMO systems. ..
  43. Eichinger C, Jentsch S. Synaptonemal complex formation and meiotic checkpoint signaling are linked to the lateral element protein Red1. Proc Natl Acad Sci U S A. 2010;107:11370-5 pubmed publisher
    ..Thus, Red1, in addition to its structural role in the SC, is a crucial coordinator of meiosis by coupling checkpoint signaling to SC formation...
  44. Hooker G, Roeder G. A Role for SUMO in meiotic chromosome synapsis. Curr Biol. 2006;16:1238-43 pubmed
    ..A non-null allele of the UBC9 gene, which encodes the SUMO-conjugating enzyme, impairs Zip1 polymerization along chromosomes...
  45. Sarangi P, Bartosova Z, Altmannova V, Holland C, Chavdarova M, Lee S, et al. Sumoylation of the Rad1 nuclease promotes DNA repair and regulates its DNA association. Nucleic Acids Res. 2014;42:6393-404 pubmed publisher
    ..These findings suggest a model whereby sumoylation of Rad1 promotes its disengagement from DNA after nuclease cleavage, allowing it to efficiently attend to large numbers of DNA lesions. ..
  46. Oling D, Eisele F, Kvint K, Nystrom T. Opposing roles of Ubp3-dependent deubiquitination regulate replicative life span and heat resistance. EMBO J. 2014;33:747-61 pubmed publisher
    ..with this, Ubp3 destabilized, deubiquitinated, and diminished the toxicity of the JUNQ-associated misfolded protein Ubc9(ts) in a proteasome-dependent manner...
  47. Alonso A, D Silva S, Rahman M, Meluh P, Keeling J, Meednu N, et al. The yeast homologue of the microtubule-associated protein Lis1 interacts with the sumoylation machinery and a SUMO-targeted ubiquitin ligase. Mol Biol Cell. 2012;23:4552-66 pubmed publisher
    ..Pac1p modification is also altered by Kar9p and the dynein regulator She1p. This work has implications for the regulation of dynein's interaction with various cargoes, including its off-loading to the cortex. ..
  48. Rothenbusch U, Sawatzki M, Chang Y, Caesar S, Schlenstedt G. Sumoylation regulates Kap114-mediated nuclear transport. EMBO J. 2012;31:2461-72 pubmed publisher
    ..Remarkably, sumoylation of Kap114 greatly stimulates cargo dissociation in vitro. We propose that sumoylation occurs at the site of Kap114 cargo function and that SUMO is a cargo release factor involved in intranuclear targeting. ..
  49. Parker J, Ulrich H. A SUMO-interacting motif activates budding yeast ubiquitin ligase Rad18 towards SUMO-modified PCNA. Nucleic Acids Res. 2012;40:11380-8 pubmed publisher
  50. Hiraishi H, Mochizuki M, Takagi H. Enhancement of stress tolerance in Saccharomyces cerevisiae by overexpression of ubiquitin ligase Rsp5 and ubiquitin-conjugating enzymes. Biosci Biotechnol Biochem. 2006;70:2762-5 pubmed
    ..Co-overexpression of Rsp5 and Ubc1, Ubc2, Ubc3, Ubc5, Ubc6, Ubc9, Ubc10, Ubc11, Ubc12, or Ubc13 further enhanced stress tolerance...
  51. Panse V, Kressler D, Pauli A, Petfalski E, Gnädig M, Tollervey D, et al. Formation and nuclear export of preribosomes are functionally linked to the small-ubiquitin-related modifier pathway. Traffic. 2006;7:1311-21 pubmed
    ..Mutations in UBC9 (E2 enzyme), ULP1 [small-ubiquitin-related modifier (SUMO) isopeptidase] and SMT3 (SUMO-1) caused 60S export ..
  52. Burkovics P, Sebesta M, Sisakova A, Plault N, Szukacsov V, Robert T, et al. Srs2 mediates PCNA-SUMO-dependent inhibition of DNA repair synthesis. EMBO J. 2013;32:742-55 pubmed publisher
    ..Our results suggest that cycling cells use the Siz1-dependent SUMOylation of PCNA to limit the extension of repair synthesis during template switch or HR and attenuate reciprocal DNA strand exchanges to maintain genome stability. ..
  53. Wang Z, Jones G, Prelich G. Genetic analysis connects SLX5 and SLX8 to the SUMO pathway in Saccharomyces cerevisiae. Genetics. 2006;172:1499-509 pubmed
  54. Ii T, Mullen J, Slagle C, Brill S. Stimulation of in vitro sumoylation by Slx5-Slx8: evidence for a functional interaction with the SUMO pathway. DNA Repair (Amst). 2007;6:1679-91 pubmed
    ..Physical interactions between the Slx5-8 complex and both Ubc9 and Smt3 were identified and characterized...
  55. Wang J, Taherbhoy A, Hunt H, Seyedin S, Miller D, Miller D, et al. Crystal structure of UBA2(ufd)-Ubc9: insights into E1-E2 interactions in Sumo pathways. PLoS ONE. 2010;5:e15805 pubmed publisher
    ..Aos1-Uba2 E1 enzyme (in humans called Sae1-Uba2), which activates Sumo's C-terminus, binds the dedicated E2 enzyme Ubc9, and promotes Sumo C-terminal transfer between the Uba2 and Ubc9 catalytic cysteines...
  56. Bustard D, Ball L, Cobb J. Non-Smc element 5 (Nse5) of the Smc5/6 complex interacts with SUMO pathway components. Biol Open. 2016;5:777-85 pubmed publisher
    ..Nse5 physically associates with the E2 conjugating enzyme, Ubc9, where contacts are stabilized by non-covalent interactions with SUMO...
  57. Takahashi Y, Yong Gonzalez V, Kikuchi Y, Strunnikov A. SIZ1/SIZ2 control of chromosome transmission fidelity is mediated by the sumoylation of topoisomerase II. Genetics. 2006;172:783-94 pubmed
    ..We propose a model for the positive control of the centromeric pool of Top2p, required for high segregation fidelity, by Smt3p modification. ..
  58. Bergink S, Ammon T, Kern M, Schermelleh L, Leonhardt H, Jentsch S. Role of Cdc48/p97 as a SUMO-targeted segregase curbing Rad51-Rad52 interaction. Nat Cell Biol. 2013;15:526-32 pubmed publisher
    ..We propose that Cdc48, through its ability to associate with co-factors that have affinities for ubiquitin and SUMO, connects the two modification pathways for protein degradation or other regulatory purposes. ..
  59. Hochstrasser M. Ubiquitin-dependent protein degradation. Annu Rev Genet. 1996;30:405-39 pubmed
    ..This diversity underlies both the high substrate specificity of the ubiquitin system and the variety of regulatory mechanisms that it serves. ..
  60. Ramsey K, Smith J, Dasgupta A, Maqani N, Grant P, Auble D. The NEF4 complex regulates Rad4 levels and utilizes Snf2/Swi2-related ATPase activity for nucleotide excision repair. Mol Cell Biol. 2004;24:6362-78 pubmed
    ..Elc1 is the yeast homologue of a mammalian E3 subunit, and it is a novel component of NEF4. Moreover, the E2s Ubc9 and Ubc13 were linked to the NEF4 repair pathway by genetic criteria...