Affinage

UBE2A

Ubiquitin-conjugating enzyme E2 A · UniProt P49459

Length
152 aa
Mass
17.3 kDa
Annotated
2026-04-28
55 papers in source corpus 18 papers cited in narrative 18 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

UBE2A (RAD6A/HHR6A) is an E2 ubiquitin-conjugating enzyme that partners with multiple E3 ligases to regulate the N-end rule protein degradation pathway, DNA damage tolerance, histone modification, and mitophagy. It forms functional complexes with UBR1 for N-end rule substrate ubiquitination (PMID:1651502), RAD18 for PCNA monoubiquitination in post-replication repair (PMID:10908344, PMID:21967848), RNF20/RNF40 for H2B K120 monoubiquitylation—allosterically stimulated by H2B S112 GlcNAcylation contacting UBE2A directly (PMID:41495224)—Parkin for mitochondrial protein ubiquitination during mitophagy (PMID:23685073), and UBR4 via an atypical hemiRING module that exploits UBE2A's intrinsically high lysine reactivity (PMID:38182926). Its catalytic activity is regulated by CDK1/2-mediated Ser120 phosphorylation, which enhances histone H2B ubiquitylation at G2/M while suppressing N-end rule degradation (PMID:11953320, PMID:21041297), and relies on a catalytic microenvironment centered on Q93 that deprotonates the incoming substrate lysine (PMID:30531907). Loss-of-function mutations in UBE2A cause X-linked intellectual disability with mitochondrial dysfunction, impaired mGluR-dependent long-term depression, and defective myeloid differentiation (PMID:23685073, PMID:26476408, PMID:30531907).

Mechanistic history

Synthesis pass · year-by-year structured walk · 16 steps
  1. 1991 High

    Establishing that RAD6/UBC2 is the obligate E2 for N-end rule degradation resolved how substrate recognition (UBR1) couples to ubiquitin conjugation in this pathway.

    Evidence Genetic epistasis and biochemical association of UBC2 with UBR1 in yeast

    PMID:1651502

    Open questions at the time
    • Molecular interface between E2 and UBR1 unknown
    • Whether mammalian UBE2A behaves identically not yet tested
  2. 1993 High

    Defining that the acidic C-terminal tail of RAD6 mediates E3 binding while the active-site cysteine mediates catalysis separated the recognition and catalytic functions of this E2.

    Evidence C88A mutagenesis and yeast two-hybrid mapping of UBR1 interaction domain

    PMID:8505328

    Open questions at the time
    • Role of the acidic tail with non-UBR1 E3 partners not addressed
    • No structural model of the E2–E3 interface
  3. 1996 Medium

    Demonstrating constitutive nuclear localization enriched in euchromatin linked UBE2A to chromatin biology beyond its known cytoplasmic degradation roles.

    Evidence Immunogold EM and subcellular fractionation in mammalian tissues

    PMID:8575614

    Open questions at the time
    • Mechanism of nuclear import/export not defined
    • Chromatin substrates not identified
  4. 1997 High

    Showing that RAD6 ubiquitin-conjugating activity is required for telomeric silencing independently of UBR1 and RAD18 revealed a distinct chromatin-regulatory function for this E2.

    Evidence Genetic epistasis with C88 mutants and silencing reporter assays in yeast

    PMID:9343433

    Open questions at the time
    • E3 partner for silencing function unknown
    • Direct chromatin substrate not identified
  5. 2000 Medium

    Purification of stable human RAD18–UBE2A complexes established that the yeast post-replication repair E2–E3 partnership is conserved in mammals.

    Evidence Co-expression and co-purification of recombinant human RAD18–HHR6A/B complexes

    PMID:10908344

    Open questions at the time
    • PCNA ubiquitination by this complex not yet demonstrated
    • Stoichiometry unresolved
  6. 2002 High

    Discovery that CDK1/2-mediated Ser120 phosphorylation activates UBE2A's conjugating activity and peaks at G2/M with increased H2B ubiquitylation revealed cell-cycle-dependent regulation of histone modification through E2 phosphorylation.

    Evidence In vitro kinase assays, S120A/S120D mutagenesis, cell cycle synchronization and histone ubiquitylation analysis

    PMID:11953320

    Open questions at the time
    • Whether phosphorylation differentially affects different E3 partnerships not tested
    • In vivo kinase identity not confirmed by knockdown
  7. 2004 High

    The finding that UBE2A knockout female mice are sterile due to two-cell-stage embryonic arrest established UBE2A as a maternal-effect gene essential for early development.

    Evidence Conditional knockout mouse with embryo culture and histology

    PMID:15169909

    Open questions at the time
    • Substrate responsible for developmental arrest not identified
    • Whether UBE2B compensates partially not determined
  8. 2005 High

    Demonstrating that RAD6/UBR1-mediated ubiquitination of Cut8 drives nuclear proteasome enrichment linked this E2 to a previously unrecognized role in spatial regulation of protein degradation.

    Evidence In vivo ubiquitination of Cut8, nuclear fractionation of proteasome, and DNA damage sensitivity in fission yeast

    PMID:16096059

    Open questions at the time
    • Whether mammalian UBE2A performs an analogous proteasome-targeting function unknown
    • Whether ubiquitination is degradative or serves as a localization signal unclear
  9. 2010 High

    Showing that Ser120 phosphorylation suppresses N-end rule degradation 20-fold while enhancing histone ubiquitylation resolved how a single phosphorylation switch redirects UBE2A between its E3 partners.

    Evidence Quantitative in vitro ubiquitination kinetics with S120A/S120D mutants and UBR1/E3α

    PMID:21041297

    Open questions at the time
    • Structural basis for differential E3 responsiveness to phosphorylation not determined
    • Physiological consequence of pathway switch not demonstrated in vivo
  10. 2011 High

    Resolving the asymmetric RAD6A–(RAD18)₂ ternary complex architecture clarified how a single E2 is activated by a dimeric RING E3 for PCNA ubiquitination.

    Evidence Differential tagging, co-IP, domain deletion mutagenesis, and in vitro ubiquitin ligase assays

    PMID:21967848

    Open questions at the time
    • High-resolution structure of the ternary complex not obtained
    • How complex assembles on chromatin-bound PCNA unknown
  11. 2013 High

    Identification of UBE2A as the E2 for Parkin-mediated mitochondrial ubiquitination connected UBE2A loss to mitochondrial dysfunction and synaptic impairment, explaining neurological features of UBE2A deficiency.

    Evidence In vitro ubiquitination reconstitution with Parkin, Drosophila and mouse knockouts, patient-derived cell analysis

    PMID:23685073

    Open questions at the time
    • Specific mitochondrial substrates not fully catalogued
    • Whether UBE2A is the sole or preferred E2 for Parkin in neurons not resolved
  12. 2015 High

    Demonstrating that Ube2a knockout mice have impaired spatial learning and defective mGluR-LTD but normal LTP identified a specific synaptic plasticity mechanism downstream of UBE2A.

    Evidence Knockout mouse behavioral testing and hippocampal electrophysiology

    PMID:26476408

    Open questions at the time
    • Substrate responsible for mGluR-LTD regulation unknown
    • Whether mitophagy defect underlies the plasticity phenotype not tested
  13. 2018 High

    Characterization of the pathogenic Q93E mutation established that UBE2A uses a catalytic microenvironment to deprotonate substrate lysines, defining its aminolysis mechanism at the chemical level.

    Evidence Patient mutation, in vitro ubiquitination with pH and nucleophile rescue, RAD18-PCNA assay

    PMID:30531907

    Open questions at the time
    • Whether Q93 plays the same role with all E3 partners not tested
    • No structural visualization of lysine deprotonation step
  14. 2023 Medium

    Discovery of force-dependent nucleocytoplasmic shuttling of UBE2A/B that regulates transcription through histone ubiquitination revealed a YAP-independent mechanotransduction pathway.

    Evidence DHS-proteomics, nucleocytoplasmic fractionation, myosin/actin inhibitor treatments, NGS

    PMID:37818922

    Open questions at the time
    • Nuclear import/export signals mediating shuttling not mapped
    • Whether this pathway operates in all cell types unknown
    • Independent replication in a second lab needed
  15. 2024 High

    Crystal structure of UBR4's atypical hemiRING–UBE2A complex explained how this N-degron E3 specifically recruits UBE2A/B and exploits their intrinsically high lysine reactivity for substrate modification.

    Evidence X-ray crystallography, mutagenesis, in vitro ubiquitination, allosteric activation analysis

    PMID:38182926

    Open questions at the time
    • In vivo substrates of UBR4–UBE2A not identified
    • Whether the hemiRING mechanism is shared by other E3s unknown
  16. 2026 High

    Cryo-EM of the RNF20/RNF40–RAD6A–nucleosome complex with H2B S112 GlcNAcylation showed that the sugar contacts the E2 directly to allosterically enhance K120 nucleophilicity, revealing crosstalk between O-GlcNAcylation and ubiquitylation at the structural level.

    Evidence Cryo-EM structure, chemically synthesized GlcNAc-nucleosomes, mutagenesis, kinetic analysis

    PMID:41495224

    Open questions at the time
    • In vivo relevance of H2BS112GlcNAc stimulation not confirmed in cells
    • Whether other histone PTMs modulate UBE2A activity through similar direct E2 contacts unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How UBE2A is allocated among its multiple E3 partners in vivo—and whether pathway switching is regulated beyond Ser120 phosphorylation—remains unresolved.
  • No systems-level quantification of E2–E3 complex partitioning in cells
  • Structural basis of phosphorylation-dependent E3 selectivity unknown
  • Full catalogue of in vivo substrates across E3 partnerships lacking

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140096 catalytic activity, acting on a protein 8 GO:0016740 transferase activity 6
Localization
GO:0005694 chromosome 4 GO:0005634 nucleus 3 GO:0005829 cytosol 1
Pathway
R-HSA-392499 Metabolism of proteins 7 R-HSA-4839726 Chromatin organization 4 R-HSA-73894 DNA Repair 3 R-HSA-1640170 Cell Cycle 1 R-HSA-9612973 Autophagy 1
Partners
CDK1PRKNRAD18RNF20RNF40UBR1UBR4ZMYM2
Complex memberships
Parkin–UBE2ARAD18–UBE2ARNF20/RNF40–UBE2AUBR1–UBE2A

Evidence

Reading pass · 18 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1991 UBC2/RAD6 (yeast ortholog of UBE2A) is essential for multiubiquitination and degradation of N-end rule substrates, and physically associates with UBR1 (N-recognin), the recognition component of the N-end rule pathway. Genetic epistasis, in vivo ubiquitination assays, physical association demonstrated biochemically Proceedings of the National Academy of Sciences of the United States of America High 1651502
1993 The physical stability and functional activity of the N-recognin/UBR1–Ubc2/RAD6 complex requires the highly acidic 23-residue C-terminal region of Ubc2. The active-site Cys-88 is required for catalysis but not for N-recognin binding. A ~170-residue C-terminal fragment of UBR1 was identified as the Ubc2-interacting domain by two-hybrid analysis. Yeast two-hybrid, active-site mutagenesis (C88A), in vivo N-end rule pathway assays, mRNA stability analysis The Journal of biological chemistry High 8505328
1996 Human HHR6A (UBE2A) and HHR6B proteins are constitutively expressed in all mammalian tissues and localize to the nucleus, preferentially to euchromatin, with elevated expression in testis coinciding with histone replacement during spermatogenesis, supporting a role in ubiquitin-mediated histone degradation and chromatin remodeling. 2D immunoblot, immunohistochemistry, electron microscopy with immunogold labeling, subcellular fractionation Developmental biology Medium 8575614
1997 RAD6/UBC2 ubiquitin-conjugating activity (dependent on active-site Cys-88 and the N-terminus of Rad6) is required for telomeric silencing in yeast, through a mechanism independent of N-end rule protein degradation (Ubr1) and DNA repair (Rad18). Genetic epistasis with rad6 null and point mutants (C88A, C88S), silencing reporter assays Molecular and cellular biology High 9343433
2000 Human RAD18 physically interacts with HHR6A (UBE2A) and HHR6B, forming stable ternary complexes that can be purified, implicating RAD18-UBE2A complex formation in DNA lesion bypass/post-replication repair in humans. Co-expression in yeast, co-purification, protein complex isolation Nucleic acids research Medium 10908344
2002 HHR6A (UBE2A) is phosphorylated by CDK1 and CDK2 on Ser120, resulting in a ~4-fold increase in ubiquitin-conjugating activity in vitro. In vivo, HHR6A phosphorylation peaks at G2/M with a concomitant increase in histone H2B ubiquitylation. Mutation of Ser120 to alanine abolishes activity, while Ser120Asp (phosphomimetic) increases activity ~3-fold. Solid-phase phosphorylation screen, in vitro kinase assay, site-directed mutagenesis, cell cycle analysis, histone ubiquitylation assay, yeast complementation The EMBO journal High 11953320
2003 ZNF198 forms a protein complex with HHR6A/6B (UBE2A/UBE2B) and RAD18, as demonstrated by yeast two-hybrid and co-immunoprecipitation. The ZNF198/FGFR1 fusion oncoprotein also binds HHR6 but disrupts the RAD18 interaction, leading to dominant-negative impairment of DNA repair. Yeast two-hybrid, co-immunoprecipitation, UV sensitivity assay Oncogene Medium 12776193
2004 mHR6A (UBE2A) is a maternal factor essential for early embryonic development: mHR6A knockout females fail to produce offspring because oocytes lacking mHR6A cannot develop beyond the two-cell stage, establishing a specific role for UBE2A in maternal-effect developmental control. Conditional knockout mouse, histology, embryo culture, histone H3 methylation analysis Molecular and cellular biology High 15169909
2005 Fission yeast Rhp6/Ubc2/Rad6 (ortholog of UBE2A) and E3 ligase Ubr1 ubiquitinate Cut8, a nuclear envelope protein, to promote nuclear enrichment of the proteasome. Non-ubiquitinatable Cut8 and rhp6/ubr1 null mutants both fail to enrich nuclear proteasome, causing hypersensitivity to DNA damage. In vivo ubiquitination assays, nuclear fractionation, mutant analysis (K-all-R Cut8), proteasome localization studies, DNA damage sensitivity assays Cell High 16096059
2010 CDK1/2-mediated phosphorylation of Ubc2/Rad6 at Ser120 negatively regulates N-end rule-dependent degradation: Ser120Asp (phosphomimetic) is 20-fold less active than wild-type for E3α/Ubr1-catalyzed conjugation in vitro, while Ser120Ala is 8-fold less active, demonstrating differential regulation of distinct E2 activities by phosphorylation. In vitro ubiquitination kinetics, ectopic expression in human cells, N-end rule substrate degradation assays, transthiolation assays The Journal of biological chemistry High 21041297
2011 RAD6A (UBE2A) forms an asymmetric ternary complex with two RAD18 subunits (RAD6A–(RAD18)₂). Only one of the two RAD18 R6BD (RAD6-binding domain) is required for complex formation and ligase activity. Both subunits of RING and SAP domains (but not UBZ) are required for full ligase activity; mutations in only one RING or SAP subunit are without effect. Differential tagging of RAD18 subunits, co-immunoprecipitation, in vitro ubiquitin ligase assay, domain deletion mutagenesis Nucleic acids research High 21967848
2013 RAD6A (UBE2A) functions as an E2 ubiquitin-conjugating enzyme that, together with E3 ligase Parkin, ubiquitinates mitochondrial proteins to facilitate clearance of dysfunctional mitochondria (mitophagy). Loss of Ube2a in Drosophila and mouse causes mitochondrial dysfunction and impairs synaptic function; patient-derived mutant cells show defective mitochondria. In vitro and in vivo ubiquitination assays, Drosophila dRad6 knockout, mouse Ube2a knockout, patient-derived cell lines, synaptic function assays Molecular cell High 23685073
2015 Ube2a knockout mice show a major deficit in spatial learning tasks and a deficit in mGLUR-dependent long-term depression in the hippocampus, with normal synaptic transmission and LTP, establishing a causal role for UBE2A in learning and mGLUR-LTD. Knockout mouse, behavioral assays (Morris water maze, contextual fear conditioning), hippocampal electrophysiology (LTP, mGLUR-LTD) Human molecular genetics High 26476408
2018 A pathogenic Q93E mutation in UBE2A impairs aminolysis activity (ubiquitin transfer to target lysine) without affecting ubiquitin conjugation to the catalytic cysteine. The defect is not rescued by the cognate E3 RAD18 for PCNA ubiquitination but is reversed at high pH or with a low-pKa amine nucleophile, demonstrating that Q93 is required for deprotonation of the incoming lysine in the UBE2A catalytic microenvironment. Patient mutation identification, in vitro ubiquitination assays, active-site mutagenesis, high-pH rescue experiments, PCNA ubiquitination assay with RAD18 Nature chemical biology High 30531907
2019 UBE2A mutations acquired during CML blast crisis decrease UBE2A enzymatic activity in vitro, leading to impairment of myeloid differentiation in CML cells. Parallel sequencing of blast crisis vs. chronic phase samples, in vitro UBE2A activity assays, myeloid differentiation assays Haematologica Medium 30819912
2023 UBE2A/B undergoes force- and contact-inhibition-dependent nucleocytoplasmic shuttling and, once nuclear, regulates transcription of downstream genes (including YAP) through histone ubiquitination, defining a YAP-independent mechanotransduction and contact inhibition pathway. DHS-proteomics, nucleocytoplasmic fractionation, myosin/actin inhibitor treatments, next-generation sequencing of downstream gene targets The Biochemical journal Medium 37818922
2024 UBR4 contains an atypical E3 module (hemiRING zinc finger + UZI subdomain + N-terminal affinity region) that specifically recruits UBE2A and UBE2B. Crystal structure of the E2–E3 complex provides atomic-level insight into hemiRING specificity for UBE2A/UBE2B. The UZI subdomain allosterically and modestly activates Ub-loaded UBE2A, with the intrinsically high lysine reactivity of UBE2A complementing attenuated activation to impart substrate specificity. Crystal structure determination, co-IP, in vitro ubiquitination assays, mutagenesis, allosteric activation analysis Nature structural & molecular biology High 38182926
2026 H2B S112 GlcNAcylation (H2BS112GlcNAc) allosterically stimulates ubiquitin transfer by the RNF20/RNF40–RAD6A (UBE2A) E3–E2 complex to H2B K120. Cryo-EM of a chemically trapped complex shows that H2BS112GlcNAc interacts directly with the E2 enzyme RAD6A (not the E3), enhancing the nucleophilicity of H2B K120. Mutagenesis and kinetics confirmed the mechanism; the C2 N-acetyl group and β-configuration of GlcNAc C1 are essential. Chemical synthesis of modified nucleosomes, cryo-EM structure determination, mutagenesis, in vitro ubiquitination kinetics, structure-activity relationship analysis Nature chemical biology High 41495224

Source papers

Stage 0 corpus · 55 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 Deficiency in the Ubiquitin Conjugating Enzyme UBE2A in Alzheimer's Disease (AD) is Linked to Deficits in a Natural Circular miRNA-7 Sponge (circRNA; ciRS-7). Genes 266 27929395
1991 The N-end rule is mediated by the UBC2(RAD6) ubiquitin-conjugating enzyme. Proceedings of the National Academy of Sciences of the United States of America 230 1651502
1997 The ubiquitin-conjugating enzyme Rad6 (Ubc2) is required for silencing in Saccharomyces cerevisiae. Molecular and cellular biology 109 9343433
2004 The ubiquitin-conjugating DNA repair enzyme HR6A is a maternal factor essential for early embryonic development in mice. Molecular and cellular biology 99 15169909
1992 A chimeric ubiquitin conjugating enzyme that combines the cell cycle properties of CDC34 (UBC3) and the DNA repair properties of RAD6 (UBC2): implications for the structure, function and evolution of the E2s. The EMBO journal 87 1639076
2006 UBE2A, which encodes a ubiquitin-conjugating enzyme, is mutated in a novel X-linked mental retardation syndrome. American journal of human genetics 85 16909393
1996 Expression of the ubiquitin-conjugating DNA repair enzymes HHR6A and B suggests a role in spermatogenesis and chromatin modification. Developmental biology 81 8575614
2020 E2 conjugases UBC1 and UBC2 regulate MYB42-mediated SOS pathway in response to salt stress in Arabidopsis. The New phytologist 76 32167578
1993 N-recognin/Ubc2 interactions in the N-end rule pathway. The Journal of biological chemistry 76 8505328
2013 Mutations in the intellectual disability gene Ube2a cause neuronal dysfunction and impair parkin-dependent mitophagy. Molecular cell 75 23685073
2005 Regulation of nuclear proteasome by Rhp6/Ubc2 through ubiquitination and destruction of the sensor and anchor Cut8. Cell 71 16096059
2002 Regulation of the ubiquitin-conjugating enzyme hHR6A by CDK-mediated phosphorylation. The EMBO journal 65 11953320
2000 The human RAD18 gene product interacts with HHR6A and HHR6B. Nucleic acids research 61 10908344
2001 The ubc2 gene of Ustilago maydis encodes a putative novel adaptor protein required for filamentous growth, pheromone response and virulence. Molecular microbiology 54 11580841
1992 Localization of two human homologs, HHR6A and HHR6B, of the yeast DNA repair gene RAD6 to chromosomes Xq24-q25 and 5q23-q31. Genomics 49 1559696
2010 Novel missense mutations in the ubiquitination-related gene UBE2A cause a recognizable X-linked mental retardation syndrome. Clinical genetics 47 20412111
1993 The ubc-2 gene of Caenorhabditis elegans encodes a ubiquitin-conjugating enzyme involved in selective protein degradation. Molecular and cellular biology 41 8441382
2010 UBE2A deficiency syndrome: Mild to severe intellectual disability accompanied by seizures, absent speech, urogenital, and skin anomalies in male patients. American journal of medical genetics. Part A 32 21108393
2011 Asymmetric nature of two subunits of RAD18, a RING-type ubiquitin ligase E3, in the human RAD6A-RAD18 ternary complex. Nucleic acids research 29 21967848
2017 A novel UBE2A mutation causes X-linked intellectual disability type Nascimento. Human genome variation 27 28611923
2015 An essential role for UBE2A/HR6A in learning and memory and mGLUR-dependent long-term depression. Human molecular genetics 27 26476408
2020 Leishmania differentiation requires ubiquitin conjugation mediated by a UBC2-UEV1 E2 complex. PLoS pathogens 26 33108402
2018 Mechanistic insights revealed by a UBE2A mutation linked to intellectual disability. Nature chemical biology 24 30531907
2010 Novel deletion at Xq24 including the UBE2A gene in a patient with X-linked mental retardation. Journal of human genetics 24 20339384
2019 De novo UBE2A mutations are recurrently acquired during chronic myeloid leukemia progression and interfere with myeloid differentiation pathways. Haematologica 23 30819912
2003 ZNF198 protein, involved in rearrangement in myeloproliferative disease, forms complexes with the DNA repair-associated HHR6A/6B and RAD18 proteins. Oncogene 22 12776193
2014 UBE2A deficiency syndrome: a report of two unrelated cases with large Xq24 deletions encompassing UBE2A gene. American journal of medical genetics. Part A 20 25287747
2015 Cellular Ubc2/Rad6 E2 ubiquitin-conjugating enzyme facilitates tombusvirus replication in yeast and plants. Virology 19 26135843
2008 Ubc2, an ortholog of the yeast Ste50p adaptor, possesses a basidiomycete-specific carboxy terminal extension essential for pathogenicity independent of pheromone response. Molecular plant-microbe interactions : MPMI 18 18052888
2024 UBE2A and UBE2B are recruited by an atypical E3 ligase module in UBR4. Nature structural & molecular biology 17 38182926
2010 Ser(120) of Ubc2/Rad6 regulates ubiquitin-dependent N-end rule targeting by E3{alpha}/Ubr1. The Journal of biological chemistry 17 21041297
2000 Characterization of novel rad6/ubc2 ubiquitin-conjugating enzyme mutants in yeast. Current genetics 17 10803884
1999 Heat-induced cell cycle arrest of Saccharomyces cerevisiae: involvement of the RAD6/UBC2 and WSC2 genes in its reversal. Molecular microbiology 14 10361277
2011 A mutation in the Cc.ubc2 gene affects clamp cell morphogenesis as well as nuclear migration for dikaryosis in Coprinopsis cinerea. Fungal genetics and biology : FG & B 13 21281729
1995 Characterization of novel yeast RAD6 (UBC2) ubiquitin-conjugating enzyme mutants constructed by charge-to-alanine scanning mutagenesis. Journal of bacteriology 12 7836290
2017 UBE2A deficiency in two siblings: A novel splicing variant inherited from a maternal germline mosaicism. American journal of medical genetics. Part A 11 29283210
1995 Caenorhabditis elegans UBC-1, a ubiquitin-conjugating enzyme homologous to yeast RAD6/UBC2, contains a novel carboxy-terminal extension that is conserved in nematodes. DNA and cell biology 11 7546294
2022 The circular RNA hsa_circ_0001394 promotes hepatocellular carcinoma progression by targeting the miR-527/UBE2A axis. Cell death discovery 10 35210429
2020 Delineation of Clinical Manifestations of the Inherited Xq24 Microdeletion Segregating with sXCI in Mothers: Two Novel Cases with Distinct Phenotypes Ranging from UBE2A Deficiency Syndrome to Recurrent Pregnancy Loss. Cytogenetic and genome research 10 32485717
2019 A novel splice site mutation in the UBE2A gene leads to aberrant mRNA splicing in a Chinese patient with X-linked intellectual disability type Nascimento. Molecular genetics & genomic medicine 9 31566921
2004 Caenorhabditis elegans UBC-2 functions with the anaphase-promoting complex but also has other activities. Journal of cell science 9 15466891
2013 Deletion of the ubiquitin-conjugating enzyme Ubc2 confers resistance to methylmercury in budding yeast by promoting Whi2 degradation. The Journal of toxicological sciences 8 23535409
2019 Alternative Splicing of RAD6B and Not RAD6A is Selectively Increased in Melanoma: Identification and Functional Characterization. Cells 7 31683936
2020 Refinement of the clinical and mutational spectrum of UBE2A deficiency syndrome. Clinical genetics 6 32415735
2023 UBE2A/B is the trans-acting factor mediating mechanotransduction and contact inhibition. The Biochemical journal 5 37818922
2020 A novel UBE2A mutation in a Chinese family with X-linked intellectual disability. The journal of gene medicine 5 32222108
2024 Formation of functional E3 ligase complexes with UBC2 and UEV1 of Leishmania mexicana. Molecular and biochemical parasitology 4 38556171
2021 The Role of the Reanalysis of Genetic Test Results in the Diagnosis of Dysmorphic Syndrome Caused by Inherited xq24 Deletion including the UBE2A and CXorf56 Genes. Genes 4 33673493
2012 One stone, two birds: CDK9-directed activation of UBE2A regulates monoubiquitination of both H2B and PCNA. Cell cycle (Georgetown, Tex.) 4 22722497
2022 A novel UBE2A splice site variant causing intellectual disability type Nascimento. Clinical case reports 3 35846913
2021 A novel missense mutation in the UBE2A gene causes intellectual disability in the large X-linked family. The journal of gene medicine 2 33368912
2026 Allosteric activation of RNF20/RNF40-RAD6A-mediated H2BK120 monoubiquitylation by H2BS112 GlcNAcylation. Nature chemical biology 1 41495224
2025 Sudden infant death in a neonate with X-linked intellectual disability type Nascimento because of UBE2A deletion. Clinical dysmorphology 0 40156274
2025 [A large family of Nascimento form of syndromic X-linked intellectual developmental disorder caused by large segment deletion of the UBE2A gene: a case report and literature review]. Zhongguo dang dai er ke za zhi = Chinese journal of contemporary pediatrics 0 40695520
2016 Identification and characterization of a ubiquitinconjugating enzyme UBE2A gene from lamprey. Fish physiology and biochemistry 0 26463350