Affinage

UBE2I

SUMO-conjugating enzyme UBC9 · UniProt P63279

Length
158 aa
Mass
18.0 kDa
Annotated
2026-06-10
100 papers in source corpus 54 papers cited in narrative 54 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

UBE2I (UBC9) is the E2 SUMO-conjugating enzyme that drives protein sumoylation across eukaryotes, receiving activated SUMO from the SAE1/SAE2 E1 heterodimer via a Cys93 thioester intermediate and transferring SUMO-1, -2, or -3 to consensus ψKXE motifs in substrates using only E1, Ubc9, mature SUMO, and ATP (PMID:9435231, PMID:16275321). Ubc9 directly recognizes substrate sumoylation motifs and cooperates with E3 ligases—RanBP2/Nup358, which binds both SUMO and the Ubc9 β-sheet surface to orient the SUMO~E2 thioester for paralog-selective conjugation, and the yeast Siz/Pli/Mms21 ligases (PMID:15608651, PMID:15931224, PMID:16782883, PMID:21444718). A noncovalent "backside" interaction between Ubc9 and SUMO, structurally analogous to the Mms2–Ubc13 ubiquitin chain-forming mechanism, is dispensable for mono-SUMOylation but required for poly-SUMO chain assembly; engineered backside-binding inhibitors block heat-shock poly-SUMOylation and arsenic-induced PML degradation without affecting mono-SUMOylation (PMID:17466333, PMID:17491593, PMID:28784659). Ubc9 activity and substrate selectivity are tuned by modifications of the enzyme itself: autosumoylation at Lys14 redirects target discrimination, Lys65 acetylation (reversed by SIRT1 under hypoxia) selectively dampens conjugation of NDSM substrates, CDK1-dependent Ser71 phosphorylation and subsequent Pin1 isomerization enhance thioester formation, and yeast sumoylation at Lys153 converts Ubc9 into a chain-promoting cofactor (PMID:18691969, PMID:23395904, PMID:23644018, PMID:38167292). Beyond catalysis, Ubc9 acts as a SUMO-conjugation-independent coregulator and protein-stability factor—stimulating androgen-receptor and oocyte transcription, and stabilizing GLUT4, XBP1(S), and other clients via catalytically inactive (C93S/C93A) mutants that retain function (PMID:10383460, PMID:17536066, PMID:18703419, PMID:23470653). Functionally, Ubc9 is essential for cell viability, cytokinesis and G2/M progression, replication-fork repair, and chromosome segregation (PMID:14982631, PMID:17081974, PMID:17035631, PMID:12413887), and in vivo it maintains intestinal stem cells, oocyte and beta-cell development, cardiomyocyte protein quality control, and macrophage polarization through sumoylation of substrates including keratin 8, NRF2, lamin A/C, STAT4, and IRF4 (PMID:27142163, PMID:30744690, PMID:36626227, PMID:31767832, PMID:20951138, PMID:29299635, PMID:31704792).

Mechanistic history

Synthesis pass · year-by-year structured walk · 21 steps
  1. 1996 Medium

    Established the first physical and genetic links of human UBE2I to recombination/meiosis machinery and demonstrated functional conservation with yeast Ubc9, framing it as a cell-cycle-relevant factor before its enzymatic identity was known.

    Evidence Yeast two-hybrid against Rad51/Rad52/p53/UBL1, spermatocyte immunolocalization, and yeast ts-mutant complementation

    PMID:8610150 PMID:8668529 PMID:8921390

    Open questions at the time
    • Yeast two-hybrid interactions lacked biochemical validation
    • Did not define a molecular activity for the protein
    • Meiotic role inferred from localization only
  2. 1998 High

    Resolved what UBE2I actually does enzymatically: it is the SUMO E2, forming a thioester with SUMO/SMT3 and being required for SUMO conjugation in vivo, defining the entire pathway's E2 step.

    Evidence In vitro thioester formation assay plus in vivo complementation in a yeast ubc9 mutant

    PMID:9435231

    Open questions at the time
    • Did not resolve substrate-selection rules
    • E3 ligase requirements not yet defined
  3. 2001 High

    Showed Ubc9 with E1 can build polymeric SUMO-2/-3 chains and that SUMO-1 cannot self-polymerize, distinguishing mono- from poly-SUMOylation as separable outputs.

    Evidence In vitro reconstitution of SUMO chain formation with in vivo immunoblot detection

    PMID:11451954

    Open questions at the time
    • Surface of Ubc9 mediating chain extension not yet mapped
    • Physiological triggers of chain formation undefined
  4. 2003 High

    Mapped distinct Ubc9 surfaces for SUMO transfer steps, separating an N-terminal noncovalent SUMO-binding region required for E1-to-E2 transfer from substrate recognition and E2-to-target transfer.

    Evidence Site-directed mutagenesis, ITC, NMR chemical shift perturbation, and in vitro conjugation

    PMID:12924945

    Open questions at the time
    • E3-binding surface not yet defined
    • In vivo consequence of these surface mutations untested
  5. 2004 High

    Defined how an E3 (RanBP2/Nup358) engages Ubc9 via its β-sheet surface to confer paralog-selective conjugation, showing SUMO E2–E3 logic differs from ubiquitin.

    Evidence NMR chemical shift perturbation, mutagenesis, and in vitro conjugation of Sp100/PML

    PMID:15608651

    Open questions at the time
    • Mechanism of thioester positioning not yet structurally resolved
    • Generality across other E3s unknown
  6. 2005 High

    Provided the structural mechanism of SUMO E3 catalysis: a Ubc9–RanBP2–SUMO~RanGAP1 complex showed the E3 binds both SUMO and Ubc9 to optimally orient the thioester, and parallel work reconstituted the minimal active E2.

    Evidence X-ray crystallography at 3.0 Å with kinetics; detailed in vitro single/multiple-turnover reconstitution

    PMID:15931224 PMID:16275321

    Open questions at the time
    • Catalytic chemistry of lysine attack not fully resolved
    • How modifications of Ubc9 alter this geometry untested here
  7. 2007 High

    Identified the noncovalent 'backside' Ubc9–SUMO interface as the determinant of poly-SUMO chain elongation, distinct from mono-SUMOylation and analogous to ubiquitin Mms2–Ubc13 chemistry.

    Evidence Crystal structures of noncovalent Ubc9–SUMO1, chain-formation assays, MS linkage analysis

    PMID:17466333 PMID:17491593

    Open questions at the time
    • In vivo importance of backside binding not yet probed
    • Substrate determinants directing chain vs. mono unclear
  8. 2008 High

    Showed autosumoylation at Lys14 is a self-encoded switch that retunes Ubc9 substrate discrimination, creating a SIM-dependent interface that favors some substrates over others.

    Evidence Crystal structure of sumoylated Ubc9, conjugation assays with multiple substrates, SIM mutagenesis

    PMID:18691969

    Open questions at the time
    • Cellular regulation of Ubc9 autosumoylation undefined
    • Breadth of SIM-dependent substrates not catalogued
  9. 2011 High

    Demonstrated that Ubc9 uses analogous noncovalent surfaces to engage SUMO and a SUMO-like domain (Rad60 SLD2), partitioning its activities between Nse2-dependent DNA repair and Pli1-dependent global sumoylation/chain formation.

    Evidence Crystallography, mutagenesis, and yeast genetic epistasis with in vivo sumoylation assays

    PMID:21444718

    Open questions at the time
    • Mammalian counterpart of SLD2 partitioning unconfirmed
    • How surfaces are selected in vivo unknown
  10. 2013 High

    Revealed multiple covalent modifications of Ubc9 (Lys65 acetylation reversed by SIRT1; Lys153 sumoylation in yeast) that switch substrate selectivity and convert Ubc9 between active enzyme and chain-promoting cofactor, linking sumoylation to hypoxia and meiosis.

    Evidence In vitro conjugation/chain assays, MS site mapping, mutagenesis, SIRT1 knockdown, yeast meiotic genetics

    PMID:23395904 PMID:23644018

    Open questions at the time
    • Enzymes installing Lys153 sumoylation in vivo not fully defined
    • Crosstalk among the various Ubc9 modifications unresolved
  11. 2016 High

    Engineered backside-binding inhibitors (SUBINs) provided a tool that selectively blocks poly-SUMO chains in cells, confirming the two activities are pharmacologically separable and required for stress-induced PML degradation.

    Evidence Phage-display protein engineering, in vitro chain assays, cellular heat-shock and arsenic treatment

    PMID:28784659

    Open questions at the time
    • Endogenous regulators that gate backside binding unknown
    • Range of chain-dependent processes in cells not mapped
  12. 2015 Medium

    Extended kinase control of Ubc9 by showing Akt phosphorylates Thr35 to boost thioester formation and global SUMOylation, integrating sumoylation with growth-factor signaling.

    Evidence In vitro kinase and thioester assays, T35A mutagenesis, co-IP, reporter assays

    PMID:25867063

    Open questions at the time
    • Single lab with pleiotropic claims spanning STAT1/CREB and SUMO1 stabilization
    • In vivo significance of Thr35 phosphorylation untested
  13. 2024 Medium

    Connected CDK1 phosphorylation of Ubc9 to Pin1 prolyl isomerization as a sequential activation module driving hypersumoylation in glioma stem cells.

    Evidence Co-IP, in vitro thioester assays, phosphosite mutagenesis, inhibitor studies, in vivo tumor model

    PMID:38167292

    Open questions at the time
    • Single lab; generality beyond glioma stem cells unknown
    • Direct structural effect of isomerization on the active site not resolved
  14. 2002 Medium

    Established Ubc9 as essential in vertebrate cells with a cytokinesis defect distinct from the yeast G2/M block, showing SUMO conjugation has organism-specific essential roles.

    Evidence Tetracycline-repressible conditional knockout in DT40 cells with FACS and nuclear morphology analysis

    PMID:12413887

    Open questions at the time
    • Critical SUMO substrates for cytokinesis not identified
    • Single cell-type system
  15. 2006 High

    Placed Ubc9/SUMO in a defined replication-repair pathway, acting with Mms21 and Sgs1 to resolve damaged-fork cruciforms separately from the Siz1/PCNA/Srs2 route, and in vertebrate G2/M control.

    Evidence Yeast genetics with 2D-gel cruciform readout and epistasis; zebrafish dominant-negative/morpholino with cell-cycle FACS

    PMID:17035631 PMID:17081974

    Open questions at the time
    • Direct SUMO substrates at the fork not pinpointed
    • Mechanistic basis of zebrafish G2/M arrest unresolved
  16. 1999 Medium

    Uncovered a SUMO-conjugation-independent coregulatory function: a catalytic-dead Ubc9 still potentiates androgen-receptor transactivation, separating enzymatic from scaffolding roles.

    Evidence Yeast two-hybrid, co-IP, reporter assays with C93S mutant

    PMID:10383460

    Open questions at the time
    • Structural basis of conjugation-independent coactivation unknown
    • Breadth across nuclear receptors not tested here
  17. 2007 Medium

    Generalized the conjugation-independent role to membrane-protein trafficking, with catalytically inactive Ubc9 (C93A) regulating GLUT4 stability and storage-compartment targeting.

    Evidence Adenoviral overexpression/siRNA in adipocytes, glucose transport, fractionation, C93A mutant

    PMID:17536066

    Open questions at the time
    • Partner proteins mediating GLUT4 effect undefined
    • Whether SUMO of a third party is required not excluded
  18. 2010 High

    Demonstrated an essential in vivo tissue-maintenance role: inducible intestinal knockout depletes stem cells and disrupts epithelial polarity, with keratin 8 identified as a SUMO target.

    Evidence Inducible conditional knockout mouse, histology/IF, in vitro keratin-8 sumoylation

    PMID:20951138

    Open questions at the time
    • Causal substrate(s) for stem-cell loss not established
    • Link between keratin-8 sumoylation and phenotype indirect
  19. 2019 Medium

    Defined specific substrate-level mechanisms in vivo across tissues—lamin A/C for nucleophagy, STAT4 and IRF4 for macrophage polarization, NRF2 for beta-cell ROS defense, plus oocyte fertility—anchoring Ubc9's physiology to named substrates.

    Evidence Cell-type-specific knockouts, SUMO-site mutagenesis (e.g., STAT4 K350R), co-IP, fractionation, in vivo disease/tumor models

    PMID:29299635 PMID:30744690 PMID:31704792 PMID:31767832 PMID:36626227

    Open questions at the time
    • Each substrate axis is single-lab
    • Whether SUMOylation is sufficient versus necessary not fully isolated for all targets
  20. 2018 Medium

    Showed therapeutically tractable substrate engagement: Ubc9 binds CRMP2 at low-micromolar affinity, and peptide disruption reduces NaV1.7 trafficking and reverses neuropathic pain, illustrating substrate-targeted modulation of Ubc9.

    Evidence Microscale thermophoresis, AlphaLISA, peptide inhibitor, patch clamp, in vivo nerve-injury model

    PMID:29847471

    Open questions at the time
    • Selectivity of the peptide for the Ubc9–CRMP2 interface incompletely defined
    • Single lab
  21. 2022 High

    Demonstrated Ubc9 active-site plasticity by engineering a chimeric E1 that loads native ubiquitin onto Ubc9, enabling tag-directed ubiquitylation, showing the catalytic cleft can accommodate ubiquitin.

    Evidence Directed evolution of chimeric E1, in vitro ubiquitylation, E. coli reconstitution

    PMID:35237717

    Open questions at the time
    • Native cellular relevance of ubiquitin loading unknown
    • Engineered system only

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the diverse Ubc9 inputs—covalent modifications, backside/SLD surfaces, E3 partners, and conjugation-independent scaffolding—are integrated to select among hundreds of substrates in a given cell state remains unresolved.
  • No unified model linking Ubc9 PTMs to substrate choice in vivo
  • Conjugation-independent functions lack defined structural mechanism
  • Relative in vivo contribution of mono- vs poly-SUMOylation per process unmapped

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140096 catalytic activity, acting on a protein 6 GO:0016740 transferase activity 4 GO:0098772 molecular function regulator activity 4 GO:0060090 molecular adaptor activity 3 GO:0016874 ligase activity 2
Localization
GO:0005634 nucleus 4 GO:0005654 nucleoplasm 1 GO:0005829 cytosol 1
Pathway
R-HSA-1640170 Cell Cycle 3 R-HSA-392499 Metabolism of proteins 3 R-HSA-9612973 Autophagy 3 R-HSA-168256 Immune System 2 R-HSA-73894 DNA Repair 2
Partners
CRMP2MEL-18PIAS/SIZ/MMS21 (NSE2)RAD51RAD52RANBP2SAE2SUMO1
Complex memberships
SAE1/SAE2–Ubc9 SUMO E1–E2 systemUbc9–RanBP2/Nup358–SUMO1–RanGAP1 E3 complex

Evidence

Reading pass · 54 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1998 UBC9/UBE2I functions as the E2 SUMO-conjugating enzyme, forming thioester intermediates with SMT3 (yeast) and SUMO-1 (mammalian) analogous to ubiquitin-conjugating enzymes; yeast UBC9 is required for SMT3 conjugation in vivo. In vitro thioester formation assay, in vivo complementation in yeast ubc9 mutant Proceedings of the National Academy of Sciences of the United States of America High 9435231
2001 SAE1/SAE2 (E1) and Ubc9 (E2) catalyze formation of polymeric SUMO-2 and SUMO-3 chains on protein substrates in vitro; SUMO-2 chains are also detected in vivo. SUMO-1 lacks the consensus ψKXE site and cannot form such chains. In vitro reconstitution of SUMO chain formation; in vivo detection by immunoblot The Journal of biological chemistry High 11451954
2003 An N-terminal region of Ubc9 (residues including R13, K14, R17, K18) mediates noncovalent interaction with SUMO-1; mutations at this site (R13A/K14A, R17A/K18A) reduce transfer of SUMO-1 from E1 to E2 without abolishing substrate recognition or SUMO transfer from E2 to target. Site-directed mutagenesis, isothermal titration calorimetry, in vitro SUMO conjugation assay, NMR chemical shift perturbation Biochemistry High 12924945
2004 NMR chemical shift perturbation identified the beta-sheet surface of Ubc9 as the binding site for the RanBP2/Nup358 E3 SUMO ligase; this interaction is distinct from classical ubiquitin E2–E3 interactions and is required for SUMO-2 (but not SUMO-1) conjugation to Sp100 and PML. RanBP2 contains a SUMO-1-specific binding site, enabling paralog-selective conjugation. NMR chemical shift perturbation, site-directed mutagenesis, in vitro SUMO conjugation assays Nature structural & molecular biology High 15608651
2005 Crystal structure of a four-protein complex (Ubc9, Nup358/RanBP2 E3 ligase domain IR1-M, SUMO-1 conjugated to RanGAP1 C-terminal domain) at 3.0 Å reveals that Nup358/RanBP2 acts as an E3 by binding both SUMO and Ubc9 to position the SUMO–E2 thioester for optimal orientation during conjugation. X-ray crystallography (3.0 Å), biochemical and kinetic assays with additional substrates Nature High 15931224
2005 Ubc9 directly purifies as an active SUMO E2: it forms E2~SUMO thioester intermediates and catalyzes SUMO conjugation to consensus ψKXE motifs in substrate proteins in vitro using only E1, Ubc9, mature SUMO, and ATP. In vitro reconstitution, E2 thioester formation assay, single-turnover and multiple-turnover conjugation assays Methods in enzymology High 16275321
2006 In budding yeast, distinct functional domains of Ubc9 are required for cell viability versus resistance to DNA damage: swapping domains affecting substrate binding, RanBP2 E3 interaction, and E1/SUMO binding had distinct effects; the activities of E3 ligases Siz1 and Siz2 in genotoxic stress responses were distinguished. Crystal structure of yeast Ubc9 at 1.75 Å resolved these domain functions. X-ray crystallography (1.75 Å), human/yeast chimera construction, yeast genetic complementation, DNA damage sensitivity assays Molecular and cellular biology High 16782883
2007 Ubc9 forms a noncovalent complex with SUMO-1 through a surface far from the E2 active site; crystal structure at 2.4 Å shows this interface is conserved with ubiquitin pathway E2–Ubl interactions. Biochemically, this noncovalent interface is less important for E1 activation or di-SUMO-2 formation but important for E3 interactions and for poly-SUMO chain formation beyond two SUMO units. X-ray crystallography (2.4 Å), in vitro SUMO chain formation assays, E1 activation assays Journal of molecular biology High 17466333
2007 The noncovalent interaction between Ubc9 and SUMO (via the 'backside' binding site far from the active site) promotes formation of short SUMO chains on substrates Sp100 and HDAC4; crystal structure of the noncovalent Ubc9–SUMO1 complex reveals structural analogy to the Mms2–Ubc13 ubiquitin chain-forming mechanism. X-ray crystallography, in vitro SUMO chain formation assays, MS identification of chain linkages The EMBO journal High 17491593
2008 Auto-sumoylation of Ubc9 at Lys14 regulates target discrimination: sumoylated Ubc9 impairs activity toward RanGAP1 and strongly activates sumoylation of Sp100 via a SUMO-interacting motif (SIM) in Sp100 that creates an additional interface with the SUMO conjugated to the E2. Crystal structure of sumoylated Ubc9 demonstrates the new binding interface. Crystal structure of sumoylated Ubc9, in vitro SUMO conjugation assays with multiple substrates, SIM mutagenesis Molecular cell High 18691969
2011 Crystal structure of Ubc9 in complex with Rad60's SUMO-like domain 2 (SLD2) reveals that Ubc9:SLD2 and Ubc9:SUMO noncovalent complexes are structurally analogous. Disrupting Ubc9:SLD2 selectively impairs Nse2 E3 ligase-dependent DNA repair, while disrupting Ubc9:SUMO noncovalent interaction impairs global sumoylation and SUMO chain formation via Pli1 E3 ligase. X-ray crystallography, site-directed mutagenesis, yeast genetic epistasis, in vivo sumoylation assays Molecular and cellular biology High 21444718
2012 CDK1/cyclin B phosphorylates Ubc9 at Ser71 in vitro; phosphorylated Ubc9 shows increased SUMOylation activity and elevated Ubc9–SUMO1 thioester accumulation. CDK2/cyclin E and other cell cycle kinases tested did not show this activity. In vitro kinase assay, in vitro SUMO conjugation assay, site-directed mutagenesis (S71A) PloS one Medium 22509284
2013 Acetylation of Ubc9 at Lys65 selectively downregulates sumoylation of substrates with negatively charged amino acid-dependent sumoylation motifs (NDSM), such as CBP and Elk-1, by attenuating Ubc9 binding to NDSM substrates, without affecting substrates with core ψKXE motif alone. SIRT1 deacetylates Ubc9 K65 under hypoxia, linking this modification to the hypoxia response. In vitro SUMO conjugation assays, mass spectrometry identification of acetylation site, mutagenesis, siRNA knockdown of SIRT1, reporter assays The EMBO journal High 23395904
2013 In S. cerevisiae, sumoylation of Ubc9 at Lys153 (Ubc9*SUMO) converts it from an active enzyme into a catalytic cofactor: Ubc9*SUMO is severely impaired in classical SUMO conjugation activity but promotes SUMO chain assembly by cooperating with charged Ubc9 (Ubc9~SUMO) through noncovalent backside SUMO binding, positioning donor SUMO for optimal transfer. A sumoylation-deficient mutant shows reduced meiotic SUMO conjugates and abrogated synaptonemal complex formation. In vitro SUMO conjugation/chain formation assays, site-directed mutagenesis, yeast genetics (meiotic phenotype), biochemical reconstitution Molecular cell High 23644018
2015 Akt directly phosphorylates Ubc9 at Thr35; this phosphorylation promotes Ubc9 thioester bond formation with SUMO1, increasing global SUMOylation and substrate-specific SUMOylation (e.g., STAT1, CREB). Akt also phosphorylates SUMO1 at Thr76, stabilizing SUMO1 protein. In vitro kinase assay, thioester formation assay, site-directed mutagenesis (T35A), co-immunoprecipitation, reporter assays Oncogene Medium 25867063
2017 High-affinity SUMO2 variants (SUBINs) engineered to bind the backside binding site of Ubc9 selectively inhibit poly-SUMO chain formation without impairing mono-SUMOylation, demonstrating that these two activities use distinct surfaces of Ubc9. In cells, SUBINs largely prevent heat shock-triggered poly-SUMOylation and abrogate arsenic-induced PML degradation. Phage display protein engineering, in vitro SUMO chain formation assays, cellular heat shock and arsenic treatment assays The Journal of biological chemistry High 28784659
2024 Pin1 isomerizes Ubc9 in a CDK1-phosphorylation-dependent manner (CDK1-mediated phosphorylation of Ubc9 is required), upregulating Ubc9 thioester formation with SUMO1 and driving SUMO1-modified hypersumoylation in glioma stem cells. Pin1 stability is maintained by USP34-mediated deubiquitination, facilitated by Plk1-mediated phosphorylation of Pin1. Co-immunoprecipitation, in vitro thioester formation assay, site-directed mutagenesis (CDK1 phosphosite), inhibitor studies (sulfopin, RO3306), in vivo tumor model Nature communications Medium 38167292
1996 Human UBE2I (HsUbc9) interacts with Rad51 recombination protein in a yeast two-hybrid assay, and mouse MmUbc9 protein localizes to synaptonemal complexes in spermatocytes, suggesting a regulatory role in meiosis. Yeast two-hybrid, immunolocalization in mouse spermatocytes Proceedings of the National Academy of Sciences of the United States of America Low 8610150
1996 UBE2I interacts with RAD52, RAD51, p53, and UBL1 (SUMO-1) in yeast two-hybrid assays; these interactions are UBE2I-specific (RAD6/UBC2 does not interact). The RAD52 interaction is mediated by RAD52's self-association region. Yeast two-hybrid system Genomics Low 8921390
1996 Human UBC9 (UBE2I) functionally complements a S. cerevisiae ubc9 temperature-sensitive mutant, rescuing cell cycle progression defects, demonstrating structural and functional conservation. Yeast genetic complementation Nucleic acids research Medium 8668529
1999 Ubc9 interacts with the androgen receptor (AR) hinge region containing the nuclear localization signal and enhances AR-dependent transcription; a C93S catalytic mutant of Ubc9 that cannot form SUMO-1 thioester still stimulates AR-dependent transactivation, indicating a SUMO-conjugation-independent coregulatory function. Yeast two-hybrid, co-immunoprecipitation in COS-1 cells, transient transfection reporter assays, C93S mutagenesis The Journal of biological chemistry Medium 10383460
2001 The IR1+2 domain of RanBP2/Nup358 binds Ubc9 with high affinity in vitro and in vivo; when overexpressed in COS-7 cells, IR1+2 sequesters ~90% of nuclear Ubc9 to the cytoplasm, causing mislocalization of SUMO-1, SUMO-2/3, PML nuclear body enlargement, and cytoplasmic mislocalization of Rad51 with failure to form Rad51 DNA-damage foci, implicating nuclear Ubc9 in Rad51-mediated homologous recombination. GST pulldown, co-immunoprecipitation, GFP-IR1+2 overexpression with immunofluorescence, DNA damage assays The Journal of biological chemistry Medium 11709548
2001 UBC9 interacts with STRA13 in mammalian cells and promotes its ubiquitin-dependent proteasomal degradation; co-expression of STRA13 and UBC9 leads to increased pSTRA13 ubiquitination, and proteasome inhibitor blocks this degradation. Yeast two-hybrid, co-immunoprecipitation in mammalian cells, proteasome inhibitor treatment, ubiquitination assay The Journal of biological chemistry Medium 11278694
2003 siRNA-mediated knockdown of Ubc9 in HeLa cells reduces endogenous Smad4 levels and intranuclear Smad4 accumulation; SUMO-1 overexpression protects Smad4 from ubiquitin-dependent degradation by competing for the same modification site, leading to enhanced TGF-β transcriptional and growth-inhibitory responses. siRNA knockdown, SUMO-1 overexpression, subcellular fractionation, half-life analysis of Smad4 mutants, reporter assays The Journal of biological chemistry Medium 12813045
2004 In budding yeast, Ubc9 and Smt3 (SUMO) are required for efficient APC/C-mediated proteolysis: temperature-sensitive ubc9-2 and smt3-331 mutants show delayed degradation of securin Pds1 and cyclin Clb2 during mitosis, and are defective in chromosome segregation, while proteolysis of non-APC/C substrates is unaffected. Yeast genetics (temperature-sensitive mutants), cyclin/securin degradation assays, cell cycle analysis, chromosome segregation assays Molecular microbiology Medium 14982631
2006 In S. cerevisiae, Ubc9 and Mms21 SUMO ligase act in concert with Sgs1 helicase to resolve X-shaped cruciform structures at damaged replication forks; ubc9 mutants show Rad51-dependent accumulation of cruciform structures during replication of damaged templates. This SUMOylation function is distinct from the Siz1/PCNA/Srs2 pathway. Yeast genetics (ubc9 mutant), 2D gel electrophoresis to detect cruciform structures, epistasis analysis with siz1, srs2, pcna, sgs1, top3 mutants Cell High 17081974
2006 In zebrafish, reduction of Ubc9 activity (dominant-negative expression or antisense knockdown) causes defects in G2/M transition and mitotic progression, resulting in cells with 4n or 8n DNA content and fewer cells in mitosis in cartilage/eye tissues, without equivalently increasing apoptosis. Dominant-negative expression, antisense morpholino knockdown, FACS analysis, BrdU incorporation, mitotic marker staining in zebrafish embryos Molecular biology of the cell Medium 17035631
2007 Ubc9 regulates GLUT4 stability and targeting to the insulin-responsive GLUT4 storage compartment (GSC) in 3T3-L1 adipocytes; overexpression inhibits GLUT4 degradation and promotes GSC targeting, increasing insulin-responsive glucose transport. A catalytically inactive mutant Ubc9-C93A produces the same effects, indicating SUMO-conjugation-independent regulation of GLUT4. Adenoviral overexpression, siRNA knockdown, glucose transport assay, subcellular fractionation, catalytic mutant (C93A) Diabetes Medium 17536066
2008 UBE2I localizes to nuclear speckles (colocalizing with SFRS2/SC35) in mouse oocytes; overexpression of either wild-type or catalytically inactive UBE2I increases BrUTP incorporation (transcription), demonstrating that transcriptional activation by UBE2I in oocytes is independent of its SUMO-conjugating activity. Immunocytochemistry, microinjection/overexpression of WT and catalytic mutant, BrUTP incorporation assay Biology of reproduction Medium 18703419
2009 Ubc9 interacts with HIV-1 Gag protein and colocalizes at perinuclear puncta; Ubc9 knockdown reduces virion infectivity 8–10-fold, associated with decreased cell-associated Env glycoprotein stability and altered Env incorporation into virions, without affecting Gag assembly or processing. The catalytic mutant Ubc9-C93A suggests SUMO-conjugating activity may not be required for this function. RNAi knockdown, colocalization imaging, virion infectivity assay, Env stability assay, catalytic mutant expression Journal of virology Medium 19640976
2013 UBC9 specifically binds the leucine zipper motif of spliced XBP1 (pXBP1(S)) and increases its protein stability; UBC9 knockdown reduces pXBP1(S) levels and ER stress-induced transcription. A SUMO-conjugating-inactive UBC9 mutant equally stabilizes pXBP1(S), indicating SUMOylation-independent stabilization. Co-immunoprecipitation, siRNA knockdown, reporter assay, catalytic mutant (SUMO-inactive UBC9) Cell structure and function Medium 23470653
2014 Synaptic diffusion of Ubc9 in hippocampal neurons is regulated by mGlu5R-dependent signaling: activation of mGlu5R increases Ubc9 synaptic residency time via a Gαq/PLC/Ca2+/PKC cascade, promoting transient PKC phosphorylation-dependent synaptic trapping of Ubc9 and consequent modulation of synaptic sumoylation. Restricted photobleaching/photoconversion (FRAP/photoconversion) of individual hippocampal spines, pharmacological receptor activation, PKC inhibitor studies Nature communications High 25311713
2016 UBC9 overexpression in cardiomyocytes increases SUMOylation and upregulates autophagic flux; in a proteotoxic model (CryAB-R120G), UBC9 overexpression reduces aggregate formation, decreases fibrosis and hypertrophy, and improves cardiac function and survival, demonstrating that UBC9-mediated SUMOylation promotes cardiac autophagy and protein quality control. Adenoviral overexpression and siRNA knockdown in neonatal rat cardiomyocytes, transgenic mouse model, autophagic flux assays, cardiac function measurements Circulation research Medium 27142163
2019 DNA damage induces nuclear accumulation of UBC9, which SUMOylates lamin A/C; this SUMOylation is required for the interaction between autophagy protein LC3 and lamin A/C, which is required for nucleophagy (degradation of nuclear lamin A/C and leaked nuclear DNA). UBC9 knockdown prevents lamin A/C SUMOylation and attenuates LC3–lamin A/C interaction and nucleophagy. siRNA knockdown of UBC9, co-immunoprecipitation, immunofluorescence, subcellular fractionation Journal of experimental & clinical cancer research Medium 30744690
2019 UBC9 SUMOylates STAT4 at Lys350 in macrophages; mutation K350R enhances STAT4 nuclear translocation and stability, facilitating proinflammatory macrophage activation. Macrophage-specific Ubc9 deficiency augments CD8+ T cell-mediated antitumor response in prostate cancer. Biochemical/molecular analysis of SUMO modification site (site-directed mutagenesis K350R), co-immunoprecipitation, macrophage-specific knockout mouse model, tumor growth assays The Journal of clinical investigation Medium 36626227
2019 Ubc9-mediated SUMOylation of IRF4 enhances its nuclear localization and stability, thereby transcribing IL-4 and arginase 1 to promote macrophage M2 polarization; macrophage-specific Ubc9 knockout impairs M2 polarization and exacerbates streptozotocin-induced diabetes. Macrophage-specific knockout mouse (LyzM-Cre-Ubc9fl/fl), co-immunoprecipitation, subcellular fractionation, gene expression analysis Cell death & disease Medium 31767832
2019 Ubc9/SUMO-conjugase overexpression induces SUMO1-dependent DAT SUMOylation, reduces DAT ubiquitination and lysosomal degradation, enhances DAT plasma membrane steady-state level, and increases dopamine uptake capacity; Ubc9 knockdown has the opposite effects. Confocal microscopy, FRET, Western blot, RNAi knockdown, dopamine uptake assay Frontiers in cellular neuroscience Medium 30828290
2019 UBC9 interacts with Nedd4-2 E3 ubiquitin ligase (by co-immunoprecipitation) and promotes ubiquitination and proteasomal degradation of Nav1.5 (cardiac sodium channel); UBC9 overexpression decreases Nav1.5 expression and reduces sodium current densities, while UBC9 knockdown has the opposite effect. The C93S catalytic mutant equally regulates Nav1.5, indicating a SUMO-conjugating-independent mechanism. Co-immunoprecipitation, overexpression/knockdown in HEK293 and cardiomyocytes, patch clamp electrophysiology, proteasome inhibitor (MG132), catalytic mutant (C93S) Journal of molecular and cellular cardiology Medium 30772377
2000 Ubc9 interacts with bovine papillomavirus E1 replication protein both in vitro and in vivo; Ubc9 catalyzes covalent SUMO-1 conjugation to E1. An E1 mutant unable to bind Ubc9 shows impaired intranuclear distribution but normal stability, suggesting sumoylation is important for E1 nuclear subdomain localization. Yeast two-hybrid, in vitro binding, in vivo co-immunoprecipitation, in vitro SUMO-1 conjugation assay, mutational mapping The Journal of biological chemistry Medium 10871618
2002 Loss of Ubc9 in chicken DT40 cells (conditional knockout) causes cell death with increased multinucleated cells (defect in cytokinesis) and apoptosis, rather than the G2/M block seen in yeast ubc9 mutants, demonstrating distinct essential roles of SUMO conjugation in higher eukaryotes. Conditional gene knockout (tetracycline-repressible transgene), FACS cell cycle analysis, microscopic analysis of nuclear morphology Experimental cell research Medium 12413887
2010 Inducible knockout of Ubc9 in adult mouse intestinal epithelium causes rapid depletion of intestinal stem cells, loss of the proliferative compartment, disruption of enterocyte polarity (nucleus positioning and actin organization), and detachment from basal lamina. Keratin 8 was identified as a SUMO substrate in intestinal epithelial cells. Inducible conditional knockout mouse (4-OHT), histology, immunofluorescence, in vitro SUMO conjugation assay for keratin 8 Gastroenterology High 20951138
2014 Ubc9 knockdown stimulates apoptosis in embryonic stem cells (ESCs) but not MEFs; Ubc9 is required for reprogramming MEFs to iPS cells (knockdown dramatically inhibits reprogramming); Ubc9 knockdown decreases expression of pluripotency markers Nanog, Klf4, Oct4, and Sox2 in ESCs. siRNA knockdown, apoptosis assays, reprogramming efficiency measurement, qRT-PCR for pluripotency markers Stem cells (Dayton, Ohio) Medium 24706591
2005 p14ARF interacts with Ubc9 and enhances SUMO-1 modification of its binding partners (hdm2, E2F-1, HIF-1α, TBP-1, p120E4F); melanoma-associated p14ARF mutations abrogate this sumoylation-enhancing activity, suggesting p14ARF acts as a SUMO E3-like factor through Ubc9. Co-immunoprecipitation, in vivo sumoylation assays, mutagenesis of p14ARF cancer-associated mutations Cell cycle (Georgetown, Tex.) Medium 15876874
2018 Ubc9 directly binds CRMP2 with low micromolar affinity (measured by microscale thermophoresis and AlphaLISA); disrupting this interaction with a tat-conjugated CRMP2 SUMOylation motif peptide (t-CSM) decreases CRMP2 SUMOylation, reduces NaV1.7 surface trafficking and sodium currents in sensory neurons, and reverses mechanical and thermal hypersensitivity in a spinal nerve injury model. Microscale thermophoresis, AlphaLISA binding assay, peptide inhibitor, patch clamp electrophysiology, in vivo neuropathic pain model Pain Medium 29847471
2008 MEL-18 (polycomb protein) interacts with both HSF2 and UBC9; MEL-18 overexpression decreases HSF2 sumoylation and broad cellular protein sumoylation, while MEL-18 knockdown increases them. MEL-18 inhibits UBC9's ability to transfer SUMO to target proteins, functioning as an anti-E3 inhibitor of UBC9 activity. MEL-18 binding to HSF2 decreases during mitosis, correlating with increased mitotic HSF2 sumoylation. Co-immunoprecipitation, RNAi knockdown, overexpression, in vitro SUMO transfer assay The Journal of biological chemistry Medium 18211895
2022 A chimeric E1 enzyme (containing the Ub fold domain of SUMO E1 and remaining domains of Ub E1) activates and loads native ubiquitin onto Ubc9, enabling Ubc9-mediated site-specific ubiquitylation of target proteins bearing a short peptide tag (LACE system) in vitro and in E. coli. Protein engineering, directed evolution of chimeric E1, in vitro ubiquitylation assay, E. coli co-expression ACS central science High 35237717
2019 Beta cell-specific knockout of Ubc9 in mice causes loss of beta cell mass, increased reactive oxygen species, and diabetes; SUMOylation of NRF2 by Ubc9 promotes NRF2 nuclear expression and transcriptional activity, thereby preventing ROS accumulation. Conversely, Ubc9 overexpression preserves normal blood glucose but impairs insulin secretion. Conditional knockout and transgenic mouse models (beta cell-specific), ROS measurement, NRF2 subcellular fractionation, immunoprecipitation for SUMOylation Diabetologia Medium 29299635
2019 Loss of oocyte-specific Ubc9 in mice causes female infertility with defects in primordial follicle pool stability, folliculogenesis, ovulation and meiosis, and impairs expression of NOBOX and its target genes, demonstrating that SUMOylation in oocytes regulates both oocyte development and communication with granulosa cells. Oocyte-specific conditional knockout mouse, histology, transcriptomic profiling of ovaries Development (Cambridge, England) Medium 31704792
2006 siRNA-mediated Ubc9 knockdown severely compromises C2C12 myoblast terminal differentiation and myotube formation without affecting expression, localization, or activation of MyoD or myogenin, placing SUMO-dependent targets downstream of these regulators as required for myogenesis. siRNA knockdown, Oil Red O staining, immunofluorescence, Western blotting for myogenic factors Experimental cell research Medium 16631162
2017 UBC9 is physiologically targeted for degradation by autophagy in human cells; HPV E6/E7 oncoproteins inhibit autophagosome-lysosome fusion, blocking autophagic degradation of UBC9 and leading to p53-dependent UBC9 accumulation during viral transformation. Ultrastructural analysis (EM), pharmacological and genetic autophagy inhibition/activation, HPV E6/E7 expression, p53 manipulation PLoS pathogens Medium 28253371
2015 UBC9 interacts with the cytoplasmic domain of calnexin (an ER membrane chaperone) and SUMOylates it; this interaction modulates calnexin's association with PTP1B, forming UBC9-dependent calnexin–PTP1B complexes at the ER membrane, linking SUMOylation to ER protein quality control and insulin/leptin signaling regulation. Co-immunoprecipitation, GST pulldown, in vitro and in vivo SUMO modification assay, immunofluorescence colocalization The Journal of biological chemistry Medium 25586181
2005 Zebrafish Ubc9 interacts with and SUMOylates DeltaNp63alpha; Ubc9-mediated sumoylation (together with Nedd4-mediated ubiquitination) destabilizes DeltaNp63alpha protein on the dorsal side of the embryo. Mutant DeltaNp63alpha unable to bind Ubc9 is stabilized, leading to more widespread neural repression. Yeast two-hybrid, in vivo sumoylation assay in HEK293 and zebrafish embryos, mutational analysis, rescue experiments in zebrafish Cell cycle (Georgetown, Tex.) Medium 15908775
2011 Ubc9 mediates nuclear localization of BRCA1 and BRCA1a; siRNA knockdown of Ubc9 in MCF-7 cells causes enhanced cytoplasmic localization of BRCA1 and exclusive cytoplasmic retention of BRCA1a/BRCA1b, which is associated with loss of growth suppression and ER-α repression activities of BRCA1. Live cell imaging (YFP/GFP/RFP-tagged BRCA1), siRNA knockdown, colony formation assay Journal of cellular physiology Medium 21344391
2022 FOSL1 promotes UBC9-dependent SUMOylation of CYLD, inducing K63-linked polyubiquitination of NF-κB intermediaries and NF-κB activation, which drives proneural-to-mesenchymal transition in glioblastoma stem cells. Co-immunoprecipitation, SUMO modification assay, NF-κB reporter, FOSL1 knockdown/overexpression, in vivo tumor model Molecular therapy : the journal of the American Society of Gene Therapy Medium 35351656

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2001 Polymeric chains of SUMO-2 and SUMO-3 are conjugated to protein substrates by SAE1/SAE2 and Ubc9. The Journal of biological chemistry 697 11451954
2005 Insights into E3 ligase activity revealed by a SUMO-RanGAP1-Ubc9-Nup358 complex. Nature 418 15931224
2006 Ubc9- and mms21-mediated sumoylation counteracts recombinogenic events at damaged replication forks. Cell 247 17081974
1998 The ubiquitin-like proteins SMT3 and SUMO-1 are conjugated by the UBC9 E2 enzyme. Proceedings of the National Academy of Sciences of the United States of America 195 9435231
2008 Ubc9 sumoylation regulates SUMO target discrimination. Molecular cell 186 18691969
2007 Noncovalent interaction between Ubc9 and SUMO promotes SUMO chain formation. The EMBO journal 174 17491593
2005 A role for Ubc9 in tumorigenesis. Oncogene 159 15735760
2003 SUMO-1/Ubc9 promotes nuclear accumulation and metabolic stability of tumor suppressor Smad4. The Journal of biological chemistry 155 12813045
1999 Ubc9 interacts with the androgen receptor and activates receptor-dependent transcription. The Journal of biological chemistry 153 10383460
1996 Mammalian ubiquitin-conjugating enzyme Ubc9 interacts with Rad51 recombination protein and localizes in synaptonemal complexes. Proceedings of the National Academy of Sciences of the United States of America 144 8610150
2004 Unique binding interactions among Ubc9, SUMO and RanBP2 reveal a mechanism for SUMO paralog selection. Nature structural & molecular biology 128 15608651
1996 Associations of UBE2I with RAD52, UBL1, p53, and RAD51 proteins in a yeast two-hybrid system. Genomics 127 8921390
2007 Structure and analysis of a complex between SUMO and Ubc9 illustrates features of a conserved E2-Ubl interaction. Journal of molecular biology 125 17466333
2009 MicroRNA-mediated regulation of Ubc9 expression in cancer cells. Clinical cancer research : an official journal of the American Association for Cancer Research 104 19223510
2009 Ubc9 promotes breast cell invasion and metastasis in a sumoylation-independent manner. Oncogene 97 20023705
2002 Ubc9 is essential for viability of higher eukaryotic cells. Experimental cell research 97 12413887
2001 Regulation of STRA13 by the von Hippel-Lindau tumor suppressor protein, hypoxia, and the UBC9/ubiquitin proteasome degradation pathway. The Journal of biological chemistry 93 11278694
2003 Role of an N-terminal site of Ubc9 in SUMO-1, -2, and -3 binding and conjugation. Biochemistry 84 12924945
1996 Identification of the structural and functional human homolog of the yeast ubiquitin conjugating enzyme UBC9. Nucleic acids research 82 8668529
2010 Expression analysis of Ubc9, the single small ubiquitin-like modifier (SUMO) E2 conjugating enzyme, in normal and malignant tissues. Human pathology 81 20561671
2002 Ubc9 is a novel modulator of the induction properties of glucocorticoid receptors. The Journal of biological chemistry 78 11812797
2005 Targeting Ubc9 for cancer therapy. Expert opinion on therapeutic targets 73 16300471
2002 Transcription factor AP-2 interacts with the SUMO-conjugating enzyme UBC9 and is sumolated in vivo. The Journal of biological chemistry 72 12072434
2006 Coactivation of the N-terminal transactivation of mineralocorticoid receptor by Ubc9. The Journal of biological chemistry 69 17105732
2022 FOSL1 promotes proneural-to-mesenchymal transition of glioblastoma stem cells via UBC9/CYLD/NF-κB axis. Molecular therapy : the journal of the American Society of Gene Therapy 67 35351656
2015 SUMO modification of Akt regulates global SUMOylation and substrate SUMOylation specificity through Akt phosphorylation of Ubc9 and SUMO1. Oncogene 67 25867063
2013 Ubc9 sumoylation controls SUMO chain formation and meiotic synapsis in Saccharomyces cerevisiae. Molecular cell 67 23644018
1996 Interaction of the Ubc9 human homologue with c-Jun and with the glucocorticoid receptor. Steroids 67 8733011
2016 UBC9-Mediated Sumoylation Favorably Impacts Cardiac Function in Compromised Hearts. Circulation research 65 27142163
2005 p14ARF interacts with the SUMO-conjugating enzyme Ubc9 and promotes the sumoylation of its binding partners. Cell cycle (Georgetown, Tex.) 65 15876874
2018 Both conditional ablation and overexpression of E2 SUMO-conjugating enzyme (UBC9) in mouse pancreatic beta cells result in impaired beta cell function. Diabetologia 64 29299635
2019 Nuclear accumulation of UBC9 contributes to SUMOylation of lamin A/C and nucleophagy in response to DNA damage. Journal of experimental & clinical cancer research : CR 63 30744690
2007 The SUMO conjugating enzyme Ubc9 is a regulator of GLUT4 turnover and targeting to the insulin-responsive storage compartment in 3T3-L1 adipocytes. Diabetes 63 17536066
2006 Ubc9 regulates mitosis and cell survival during zebrafish development. Molecular biology of the cell 63 17035631
2010 Sumoylation by Ubc9 regulates the stem cell compartment and structure and function of the intestinal epithelium in mice. Gastroenterology 59 20951138
2010 Over-accumulation of nuclear IGF-1 receptor in tumor cells requires elevated expression of the receptor and the SUMO-conjugating enzyme Ubc9. Biochemical and biophysical research communications 59 21147068
2008 UBE2I (UBC9), a SUMO-conjugating enzyme, localizes to nuclear speckles and stimulates transcription in mouse oocytes. Biology of reproduction 57 18703419
2017 Manipulating PML SUMOylation via Silencing UBC9 and RNF4 Regulates Cardiac Fibrosis. Molecular therapy : the journal of the American Society of Gene Therapy 56 28143738
2014 Sumo E2 enzyme UBC9 is required for efficient protein quality control in cardiomyocytes. Circulation research 56 25097219
2005 Destabilization of DeltaNp63alpha by Nedd4-mediated ubiquitination and Ubc9-mediated sumoylation, and its implications on dorsoventral patterning of the zebrafish embryo. Cell cycle (Georgetown, Tex.) 56 15908775
2004 Smt3/SUMO and Ubc9 are required for efficient APC/C-mediated proteolysis in budding yeast. Molecular microbiology 56 14982631
2023 UBC9 deficiency enhances immunostimulatory macrophage activation and subsequent antitumor T cell response in prostate cancer. The Journal of clinical investigation 54 36626227
2019 Loss of ubiquitin-conjugating enzyme E2 (Ubc9) in macrophages exacerbates multiple low-dose streptozotocin-induced diabetes by attenuating M2 macrophage polarization. Cell death & disease 54 31767832
2013 Ubc9 acetylation modulates distinct SUMO target modification and hypoxia response. The EMBO journal 53 23395904
2000 Bovine papillomavirus E1 protein is sumoylated by the host cell Ubc9 protein. The Journal of biological chemistry 53 10871618
2018 Inhibition of the Ubc9 E2 SUMO-conjugating enzyme-CRMP2 interaction decreases NaV1.7 currents and reverses experimental neuropathic pain. Pain 52 29847471
2017 Autophagy regulates UBC9 levels during viral-mediated tumorigenesis. PLoS pathogens 51 28253371
2012 Phosphorylation of Ubc9 by Cdk1 enhances SUMOylation activity. PloS one 50 22509284
2011 DNA repair and global sumoylation are regulated by distinct Ubc9 noncovalent complexes. Molecular and cellular biology 48 21444718
2011 Ubc9 expression predicts chemoresistance in breast cancer. Chinese journal of cancer 46 21880185
2001 Perturbation of SUMOlation enzyme Ubc9 by distinct domain within nucleoporin RanBP2/Nup358. The Journal of biological chemistry 46 11709548
1996 Cloning, expression, and mapping of UBE2I, a novel gene encoding a human homologue of yeast ubiquitin-conjugating enzymes which are critical for regulating the cell cycle. Cytogenetics and cell genetics 46 8565643
2012 microRNA-214-mediated UBC9 expression in glioma. BMB reports 45 23187003
2004 Ubc9 and Protein Inhibitor of Activated STAT 1 Activate Chicken Ovalbumin Upstream Promoter-Transcription Factor I-mediated Human CYP11B2 Gene Transcription. The Journal of biological chemistry 45 15611122
2011 Correlations among ERCC1, XPB, UBE2I, EGF, TAL2 and ILF3 revealed by gene signatures of histological subtypes of patients with epithelial ovarian cancer. Oncology reports 42 21971700
2000 Interaction of Daxx, a Fas binding protein, with sentrin and Ubc9. Biochemical and biophysical research communications 42 11112409
2004 FHL2, UBC9, and PIAS1 are novel estrogen receptor alpha-interacting proteins. Endocrine research 41 15666801
2013 Ubc9 promotes invasion and metastasis of lung cancer cells. Oncology reports 39 23381475
2024 Stabilization of Pin1 by USP34 promotes Ubc9 isomerization and protein sumoylation in glioma stem cells. Nature communications 37 38167292
2006 Role of SUMO/Ubc9 in DNA damage repair and tumorigenesis. Journal of molecular histology 37 16758298
2009 Common variants in the UBC9 gene encoding the SUMO-conjugating enzyme are associated with breast tumor grade. International journal of cancer 33 19358266
2009 cDNA cloning and expression of Ubc9 in the developing embryo and ovary of Oriental river prawn, Macrobrachium nipponense. Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology 33 19944179
2019 Loss of the E2 SUMO-conjugating enzyme Ube2i in oocytes during ovarian folliculogenesis causes infertility in mice. Development (Cambridge, England) 32 31704792
2020 SUMOylation of the transcription factor ZFHX3 at Lys-2806 requires SAE1, UBC9, and PIAS2 and enhances its stability and function in cell proliferation. The Journal of biological chemistry 31 32249212
2009 Targeting the SUMO E2 conjugating enzyme Ubc9 interaction for anti-cancer drug design. Anti-cancer agents in medicinal chemistry 31 19149481
2004 Ubc9 interacts with chicken ovalbumin upstream promoter-transcription factor I and represses receptor-dependent transcription. Journal of molecular endocrinology 31 14765993
2014 The SUMO conjugating enzyme Ubc9 is required for inducing and maintaining stem cell pluripotency. Stem cells (Dayton, Ohio) 30 24706591
2011 Ubc9 mediates nuclear localization and growth suppression of BRCA1 and BRCA1a proteins. Journal of cellular physiology 30 21344391
2009 Human Ubc9 contributes to production of fully infectious human immunodeficiency virus type 1 virions. Journal of virology 30 19640976
2007 Ubc9 fusion-directed SUMOylation identifies constitutive and inducible SUMOylation. Nucleic acids research 28 17709345
2018 Ubc9 overexpression and SUMO1 deficiency blunt inflammation after intestinal ischemia/reperfusion. Laboratory investigation; a journal of technical methods and pathology 26 29472640
2014 mGlu5 receptors regulate synaptic sumoylation via a transient PKC-dependent diffusional trapping of Ubc9 into spines. Nature communications 25 25311713
2012 SUMO-conjugating enzyme E2 UBC9 mediates viral immediate-early protein SUMOylation in crayfish to facilitate reproduction of white spot syndrome virus. Journal of virology 25 23097446
2008 Ubc9 promotes the stability of Smad4 and the nuclear accumulation of Smad1 in osteoblast-like Saos-2 cells. Bone 25 18321803
2007 RAP80 interacts with the SUMO-conjugating enzyme UBC9 and is a novel target for sumoylation. Biochemical and biophysical research communications 25 17698038
2006 Ubc9 expression is essential for myotube formation in C2C12. Experimental cell research 25 16631162
2013 UBC9 regulates the stability of XBP1, a key transcription factor controlling the ER stress response. Cell structure and function 24 23470653
2009 Polymorphisms in the UBC9 and PIAS3 genes of the SUMO-conjugating system and breast cancer risk. Breast cancer research and treatment 24 19760037
2020 Targeting UBC9-mediated protein hyper-SUMOylation in cystic cholangiocytes halts polycystic liver disease in experimental models. Journal of hepatology 23 32950589
2019 The SUMO-Conjugase Ubc9 Prevents the Degradation of the Dopamine Transporter, Enhancing Its Cell Surface Level and Dopamine Uptake. Frontiers in cellular neuroscience 23 30828290
2017 Down-regulation of UBC9 increases the sensitivity of hepatocellular carcinoma to doxorubicin. Oncotarget 23 28572537
2017 Site-specific inhibition of the small ubiquitin-like modifier (SUMO)-conjugating enzyme Ubc9 selectively impairs SUMO chain formation. The Journal of biological chemistry 23 28784659
2014 SUMO Ubc9 enzyme as a viral target. IUBMB life 23 24395713
2017 Ubc9 Is Required for Positive Selection and Late-Stage Maturation of Thymocytes. Journal of immunology (Baltimore, Md. : 1950) 22 28314856
2014 SUMO-conjugating enzyme UBC9 promotes proliferation and migration of fibroblast-like synoviocytes in rheumatoid arthritis. Inflammation 22 24531852
2006 Distinct functional domains of Ubc9 dictate cell survival and resistance to genotoxic stress. Molecular and cellular biology 22 16782883
2019 UBC9 regulates cardiac sodium channel Nav1.5 ubiquitination, degradation and sodium current density. Journal of molecular and cellular cardiology 21 30772377
2009 Ubc9 gene polymorphisms and late-onset Alzheimer's disease in the Korean population: a genetic association study. Neuroscience letters 21 19765634
2006 Regulation of bcl-2 expression by Ubc9. Experimental cell research 21 16566921
2015 UBC9-dependent association between calnexin and protein tyrosine phosphatase 1B (PTP1B) at the endoplasmic reticulum. The Journal of biological chemistry 20 25586181
2008 MEL-18 interacts with HSF2 and the SUMO E2 UBC9 to inhibit HSF2 sumoylation. The Journal of biological chemistry 20 18211895
2005 Purification and activity assays for Ubc9, the ubiquitin-conjugating enzyme for the small ubiquitin-like modifier SUMO. Methods in enzymology 20 16275321
2022 Site-Specific Protein Ubiquitylation Using an Engineered, Chimeric E1 Activating Enzyme and E2 SUMO Conjugating Enzyme Ubc9. ACS central science 19 35237717
2020 UBC9 coordinates inflammation affecting development of bladder cancer. Scientific reports 19 33244139
2014 The Role of SUMO-Conjugating Enzyme Ubc9 in the Neuroprotection of Isoflurane Preconditioning Against Ischemic Neuronal Injury. Molecular neurobiology 19 24961570
2013 Cloning, genomic structure and expression analysis of ubc9 in the course of development in the half-smooth tongue sole (Cynoglossus semilaevis). Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology 18 23507627
2011 Coactivation of SF-1-mediated transcription of steroidogenic enzymes by Ubc9 and PIAS1. Endocrinology 18 21467194
2010 Role of UBC9 in the regulation of the adipogenic program in 3T3-L1 adipocytes. Endocrinology 18 20881252
2008 Modulation of PLAGL2 transactivation activity by Ubc9 co-activation not SUMOylation. Biochemical and biophysical research communications 18 18655774
2002 Interaction of the developmental regulator SALL1 with UBE2I and SUMO-1. Biochemical and biophysical research communications 18 12200128

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