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UBL4A

Ubiquitin-like protein 4A · UniProt P11441

Length
157 aa
Mass
17.8 kDa
Annotated
2026-06-10
37 papers in source corpus 24 papers cited in narrative 23 extracted findings
Cross-family judge vs UniProt: tie faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

UBL4A (GdX/Get5/Mdy2) is a ubiquitin-like domain adaptor that operates at the core of the cytosolic machinery delivering tail-anchored (TA) membrane proteins to the ER, while also serving as a versatile scaffold in actin dynamics, immune signaling, and nuclear stress responses (PMID:23142665, PMID:26195787). Within the BAG6-UBL4A-TRC35/GET4 holdase pathway, the unconventional UBL domain of UBL4A engages the N-terminal dimerization domain of SGTA/Sgt2 through a high-affinity electrostatic and hydrophobic interface (one UBL per SGTA dimer), recruiting SGTA to the BAG6 complex to enhance substrate loading and prevent nondegradable aggregates during ERAD (PMID:23246001, PMID:23142665, PMID:23297211, PMID:24100326). In the upstream targeting step, the Get4-Get5/UBL4A subcomplex positions Get3/TRC40 and uses Get4 to prime Get3 ATPase activity for optimal TA-substrate capture (PMID:20554915, PMID:20106980, PMID:24727835). UBL4A binds the BAG6 C-terminus through a distinct BAGS-domain interface that controls UBL4A stability, and proteotoxic stresses such as polyglutamine inclusions, proteasome inhibition, and mitochondrial depolarization dissociate the BAG6-UBL4A complex (PMID:25713138, PMID:37747814). Beyond protein biogenesis, UBL4A directly binds the Arp2/3 complex via its C-terminus—a contact requiring residue D122—to drive actin branching, supporting insulin-induced Akt plasma-membrane translocation and glycogen synthesis, cell migration, and stress-induced mitochondrial fusion (PMID:26195787, PMID:30146258, PMID:33211772). UBL4A additionally tunes innate immunity by trapping TC45/PTPN2 to sustain NF-κB p65 phosphorylation and by promoting K63-linked ubiquitination of TRAF6 to enhance type I IFN signaling (PMID:30867837, PMID:31451677), and translocates to the nucleus with BAG6 upon DNA damage to support BRCA1 recruitment and damage-induced cell death (PMID:23723067). In yeast, the ortholog mediates NLS-dependent nuclear import, heat-stress stress-granule formation via Pab1, and microtubule-dependent nuclear migration through association with α-tubulin and dynactin (PMID:16390866, PMID:20722039, PMID:23285234).

Mechanistic history

Synthesis pass · year-by-year structured walk · 21 steps
  1. 2006 Medium

    Established that the UBL4A ortholog is a UBL-domain protein lacking canonical ubiquitin processing and is needed for cytoskeletal organization, answering whether this UBL behaves like ubiquitin or has a distinct cellular role.

    Evidence Yeast mdy2Δ deletion phenotyping, His-tag processing assay, and microtubule/Kar9 immunofluorescence

    PMID:16390866

    Open questions at the time
    • Molecular basis linking Mdy2 to microtubule/Kar9 localization not defined
    • Relevance to mammalian UBL4A function untested
  2. 2007 Medium

    Identified the Sgt2 N-terminus and the chaperone Ydj1 as physical partners of the ortholog, placing it within a chaperone-adaptor network rather than acting alone.

    Evidence Co-immunoprecipitation and genetic interaction assays in yeast

    PMID:17441508

    Open questions at the time
    • Direct vs. bridged nature of the Ydj1 association unresolved
    • Structural interface undefined
  3. 2010 High

    Positioned Get4/Get5 upstream of Get3 in the TA-targeting pathway by showing nucleotide-dependent Get3 binding to a conserved Get4 surface, defining the pathway's directional architecture.

    Evidence Crystal structure of the Get4/Get5 N-terminal fragment with ITC and co-IP

    PMID:20106980 PMID:20554915

    Open questions at the time
    • How substrate is handed from Get5/Sgt2 to Get3 not yet resolved at this stage
  4. 2010 Medium

    Linked the ortholog to cytoskeletal and stress-granule machinery, showing it associates with α-tubulin and dynactin and relocalizes to stress granules under heat, expanding its role beyond TA targeting.

    Evidence Co-precipitation and GFP live imaging with stress-granule markers in yeast

    PMID:20722039

    Open questions at the time
    • Functional consequence of tubulin/dynactin binding unclear
    • Whether mammalian UBL4A shares these contacts untested
  5. 2011 Medium

    Distinguished cooperative from independent roles of Sgt2 and Get5, showing they act together in TA sorting yet Sgt2 retains a Get5-independent delivery function to Get3.

    Evidence Genetic epistasis of sgt2Δ get5Δ double mutants and overexpression toxicity assays

    PMID:21619481

    Open questions at the time
    • Biochemical basis of the Get5-independent route not defined
  6. 2012 High

    Defined the molecular handshake of the pathway: the noncanonical SGTA UBD engages the unconventional UBL4A UBL domain via electrostatics, recruiting SGTA to BAG6 and preventing aggregate formation during ERAD.

    Evidence NMR, crystal structures (yeast and human), ITC, chemical shift perturbation, and co-IP across two studies

    PMID:23142665 PMID:23246001

    Open questions at the time
    • Substrate transfer dynamics across the interface not directly visualized
  7. 2012 Medium

    Established NLS-dependent nuclear import as a prerequisite for the ortholog's heat-stress response and Pab1 interaction, revealing a nuclear-history requirement for its stress function.

    Evidence NLS/NES deletion mutants, GFP localization, co-precipitation with Pab1, and heat-stress viability in yeast

    PMID:23285234

    Open questions at the time
    • Why nuclear transit is required for cytoplasmic stress-granule function is unexplained
  8. 2013 High

    Resolved the binding stoichiometry and atomic interface (1 Sgt2 dimer : 1 Get5 UBL) and validated the contact residues, providing structural ground truth for the SGTA-UBL4A complex.

    Evidence NMR solution structure, crystal structure, ITC, and site-directed mutagenesis

    PMID:23297211 PMID:24100326

    Open questions at the time
    • Conformational changes during substrate loading not captured
  9. 2013 Medium

    Extended UBL4A into the DNA-damage response, showing nuclear translocation with BAG6/GET4 and a requirement for BRCA1 recruitment and damage-induced cell death.

    Evidence siRNA knockdown, viability assays, immunofluorescence, and BRCA1 recruitment assays

    PMID:23723067

    Open questions at the time
    • Direct mechanistic link between the complex and BRCA1 recruitment undefined
    • Single-lab finding
  10. 2014 High

    Showed how the pathway is kinetically regulated: Get4 primes Get3 in an ATP-bound ternary complex to optimize TA-substrate capture, defining the catalytic logic of targeting.

    Evidence Crystal structure of the Get3-Get4-Get5 ternary complex with ATPase and structure-guided assays

    PMID:24727835

    Open questions at the time
    • Full hand-off to the ER membrane downstream of Get3 not captured in this complex
  11. 2014 High

    Demonstrated that UBL4A and BAG6 UBL domains compete for the same SGTA site, establishing a regulatory switch at the SGTA dimerization domain.

    Evidence NMR structure, ITC, microscale thermophoresis, and HADDOCK docking

    PMID:25415308

    Open questions at the time
    • Physiological trigger that selects UBL4A vs. BAG6 occupancy unknown
  12. 2015 High

    Defined the BAG6-UBL4A heterodimerization through a novel C-terminal BAGS interface and showed BAG6 binding stabilizes UBL4A, explaining co-dependence of complex members.

    Evidence Crystal structure with biochemical and cell-based stability assays

    PMID:25713138

    Open questions at the time
    • How stability control feeds back on TA-targeting flux not addressed
  13. 2015 High

    Opened a TA-independent role by showing UBL4A directly binds Arp2/3 to accelerate actin branching, enabling insulin-induced Akt membrane translocation and glycogen synthesis.

    Evidence In vitro actin branching, interaction assays, Ubl4A KO mouse, and live-cell Akt imaging

    PMID:26195787

    Open questions at the time
    • Whether the UBL and Arp2/3-binding functions are spatially partitioned in cells unclear
  14. 2016 Medium

    Generalized the actin role to motility, showing UBL4A loss impairs membrane protrusion, fibroblast and macrophage migration, and wound healing.

    Evidence Ubl4A KO mouse, wound-healing, ex vivo migration, and motility assays

    PMID:27998771

    Open questions at the time
    • Direct dependence on Arp2/3 binding not tested with domain mutants here
  15. 2018 Medium

    Mapped the Arp2/3 contact to the UBL4A C-terminus and identified D122 as the critical residue whose mutation abolishes binding, destabilizes the protein, and removes pro-death activity in colon cancer.

    Evidence In vitro pull-down, D122A mutagenesis, and cell death assays

    PMID:30146258

    Open questions at the time
    • Structural basis of D122-mediated Arp2/3 contact not solved
    • Single-study finding
  16. 2019 Medium

    Assigned UBL4A an immune-signaling role by trapping TC45/PTPN2 to disrupt the phosphatase complex that dephosphorylates p65, sustaining NF-κB activation.

    Evidence Reciprocal co-IP, luciferase reporters, and tissue-specific KO mice in LPS and DSS colitis models

    PMID:30867837

    Open questions at the time
    • Whether this competes with TA-targeting functions of UBL4A unknown
  17. 2019 Medium

    Extended immune function to antiviral signaling, showing UBL4A binds TRAF6 and promotes its K63-linked ubiquitination to amplify type I IFN responses.

    Evidence Co-IP, K63-specific ubiquitination assays, knockdown/overexpression, and phosphorylation blots

    PMID:31451677

    Open questions at the time
    • Whether UBL4A acts as or recruits the E3 ligase for TRAF6 not determined
  18. 2019 Medium

    Linked UBL4A to lysosomal control by showing direct LAMP1 interaction that impairs lysosomal function and autophagic degradation in pancreatic cancer.

    Evidence Co-IP, autophagy marker blots, and in vitro/orthotopic functional rescue

    PMID:31288830

    Open questions at the time
    • Single Co-IP method for direct interaction without reciprocal structural validation
    • Mechanism of LAMP1-mediated lysosomal impairment unclear
  19. 2020 Medium

    Tied the Arp2/3 axis to mitochondrial homeostasis, showing UBL4A promotes nutrient-stress mitochondrial fusion and that an Arp2/3-binding-deficient mutant fails to rescue fragmentation and apoptosis.

    Evidence Ubl4A KO cells with WT vs. Arp2/3-binding mutant rescue, mitochondrial imaging, ROS, and caspase-9 assays

    PMID:33211772

    Open questions at the time
    • How perimitochondrial primed Arp2/3 drives fusion mechanically not defined
  20. 2023 Medium

    Showed that the BAG6-UBL4A complex is disrupted by proteotoxic stress, connecting complex integrity to impaired TA biogenesis under disease-relevant conditions.

    Evidence Co-IP under polyglutamine, proteasome inhibition, and mitochondrial depolarization

    PMID:37747814

    Open questions at the time
    • Functional consequence on TA-protein flux measured indirectly
    • Single-study finding
  21. 2024 Low

    Proposed a UBL4A-STAT3 axis in which UBL4A reduces STAT3 phosphorylation and downstream oncogenic target expression in breast cancer.

    Evidence Dual-luciferase reporter, immunofluorescence colocalization, phosphorylation blots, and KO mouse tumor model

    PMID:39487760

    Open questions at the time
    • No direct binding assay (co-IP/pull-down) reported for the UBL4A-STAT3 interaction
    • Single-lab study

Open questions

Synthesis pass · forward-looking unresolved questions
  • How UBL4A partitions among its multiple roles—TA-protein holdase, Arp2/3-driven actin scaffold, immune signaling regulator, and nuclear damage-response factor—and what governs the switch between them remains unresolved.
  • No unifying model coordinating cytosolic, immune, and nuclear functions
  • Structural basis of the C-terminal Arp2/3 contact unsolved
  • Triggers selecting BAG6 vs. SGTA vs. non-canonical partners unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008092 cytoskeletal protein binding 3 GO:0060090 molecular adaptor activity 3 GO:0098772 molecular function regulator activity 2
Localization
GO:0005829 cytosol 3 GO:0005634 nucleus 2 GO:0005783 endoplasmic reticulum 2
Pathway
R-HSA-392499 Metabolism of proteins 3 R-HSA-9609507 Protein localization 3 R-HSA-168256 Immune System 2 R-HSA-9612973 Autophagy 1
Partners
ARPC (ARP2/3 COMPLEX)BAG6GET3GET4LAMP1PTPN2SGTATRAF6
Complex memberships
BAG6-UBL4A-TRC35/GET4 holdase complexGet3-Get4-Get5 targeting complex

Evidence

Reading pass · 23 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2012 SGTA contains a noncanonical ubiquitin-like-binding domain (UBD) that interacts specifically with the unconventional ubiquitin-like domain of Ubl4A, at least in part via electrostatics. This interaction recruits SGTA to the BAG6 complex, enhances substrate loading to BAG6, and prevents formation of nondegradable protein aggregates during ERAD. NMR spectroscopy, biochemical binding assays, co-immunoprecipitation Cell Reports High 23246001
2010 Get4 (TRC35) and Get5 (UBL4A/Mdy2) form a tight complex in yeast, and Get3 specifically binds to a conserved surface on Get4 in a nucleotide-dependent manner, placing Get4/5 upstream of Get3 in the tail-anchored protein targeting pathway. Crystal structure of Get4/Get5 N-terminal fragment, co-immunoprecipitation, isothermal titration calorimetry Proceedings of the National Academy of Sciences High 20106980 20554915
2012 The N-terminal dimerization domain of Sgt2/SGTA directly binds the UBL domain of Get5/UBL4A via electrostatics, forming a high-affinity complex with rapid kinetics, structurally characterized for both yeast and human protein pairs. Crystal structures (yeast and human UBD/UBL complex), biophysical studies (ITC, NMR chemical shift perturbation) Cell Reports High 23142665
2013 Solution structure of Sgt2_NT (unique helical fold) and crystal structure of Get5_UBL determined; NMR chemical shift perturbation and ITC establish a 1:2 stoichiometry (one Get5 UBL to one Sgt2 dimer); site-directed mutagenesis validated the binding interface. NMR solution structure, crystal structure, isothermal titration calorimetry, site-directed mutagenesis, relaxation experiments Proceedings of the National Academy of Sciences High 23297211
2014 Crystal structure of yeast Get3-Get4-Get5 ternary complex in an ATP-bound state reveals that Get4 primes Get3 by promoting an optimal configuration for TA substrate capture; Get4-mediated regulation of Get3 ATP hydrolysis is essential for efficient TA-protein targeting. Crystal structure, structure-guided biochemical assays, ATPase activity measurements Nature Structural & Molecular Biology High 24727835
2015 Crystal structure of the C-terminal heterodimerization domains of BAG6 and Ubl4A reveals a novel binding interface; the BAG6 C-terminus (designated BAGS domain) is structurally distinct from canonical BAG domains. Tight association of BAG6 and Ubl4A modulates Ubl4A protein stability in cells. Crystal structure, biochemical binding assays, cell-based stability assays Journal of Biological Chemistry High 25713138
2014 NMR solution structure of the SGTA dimerization domain determined; UBL domains from UBL4A and BAG6 compete for binding to SGTA at the same site, characterized by NMR chemical shift perturbation, ITC, and microscale thermophoresis; HADDOCK docking models of SGTA-UBL complexes generated. NMR structure determination, ITC, microscale thermophoresis, NMR chemical shift perturbation PLoS ONE High 25415308
2013 Nuclear BAG6-UBL4A-GET4 complex mediates DNA damage signaling and cell death. Upon DNA damage, UBL4A and GET4 translocate to the nucleus. Depletion of BAG6 causes loss of both UBL4A and GET4 proteins. Co-depletion of UBL4A and GET4 (but not single depletion) confers resistance to DNA damage-induced cell death. All three components regulate BRCA1 recruitment to DNA damage sites. siRNA knockdown, cell viability assays, immunofluorescence for nuclear translocation, BRCA1 recruitment assays Journal of Biological Chemistry Medium 23723067
2015 Ubl4A directly interacts with the Arp2/3 complex to accelerate actin branching and networking, enabling insulin-induced Akt translocation to the plasma membrane. Ubl4A knockout mice have defective Akt-dependent glycogen synthesis. Akt binds actin filaments and colocalizes with Arp2/3 in membrane ruffles. Protein-protein interaction assays, actin branching in vitro assays, Ubl4A knockout mouse model, live-cell imaging of Akt translocation, glycogen synthesis assays Proceedings of the National Academy of Sciences High 26195787
2016 Ubl4A, F-actin, and Arp2/3 co-localize at cell leading edges during wound closure. Knockout of Ubl4A reduces actin-mediated membrane protrusion, delays wound healing in primary mouse embryonic fibroblasts, impairs fibroblast migration from corneal tissue ex vivo, and decreases macrophage motility without affecting phagocytosis. Ubl4A KO mouse model, wound-healing assay, immunofluorescence co-localization, ex vivo migration assay, macrophage motility and phagocytosis assays Biochemical and Biophysical Research Communications Medium 27998771
2018 The C-terminal region of Ubl4A directly interacts with the Arp2/3 complex in pull-down assays. A point mutation D122A in the C-terminus abolishes Arp2/3 binding and destabilizes Ubl4A. Wild-type Ubl4A induces cell death in colon cancer cells via its C-terminal Arp2/3-binding activity, while D122A mutant loses this pro-death activity. In vitro protein pull-down, site-directed mutagenesis (D122A), cell death assays Biochemical and Biophysical Research Communications Medium 30146258
2019 GdX/UBL4A positively regulates NF-κB signaling in dendritic cells and macrophages by trapping TC45 (PTPN2), thereby disrupting the TC45/PP2A/p65 complex that mediates p65 dephosphorylation and preventing p65 dephosphorylation. Immunoprecipitation, luciferase reporter assay, LPS challenge of GdX-deficient mice, cytokine ELISA, DSS colitis model with tissue-specific KO mice Theranostics Medium 30867837
2019 UBL4A interacts with TRAF6 via co-immunoprecipitation, and this interaction is enhanced upon viral infection. UBL4A promotes K63-linked ubiquitination of TRAF6, enhancing TBK1, IRF3, and IKKα/β phosphorylation and type I IFN transcription. Co-immunoprecipitation, ubiquitination assay (K63-specific), siRNA knockdown and overexpression, phosphorylation western blots Journal of Immunology Medium 31451677
2019 UBL4A directly interacts with LAMP1 by co-immunoprecipitation and causes lysosomal dysfunction leading to impaired autophagic degradation and autophagosome accumulation in pancreatic cancer cells. LAMP1 overexpression reverses the antitumor effects of UBL4A. Co-immunoprecipitation, western blot autophagy markers, in vitro and in vivo (orthotopic) functional assays Journal of Experimental & Clinical Cancer Research Medium 31288830
2020 Ubl4A promotes mitochondrial fusion under nutrient deprivation via its interaction with the Arp2/3 complex. Ubl4A deficiency reduces the primed Arp2/3 complex pool around mitochondria, impairs fusion, causes mitochondrial fragmentation, ROS accumulation, and caspase 9-dependent apoptosis. An Arp2/3-binding-deficient Ubl4A mutant fails to rescue this phenotype. Ubl4A KO cells and rescue with wild-type vs. Arp2/3-binding mutant, mitochondrial morphology imaging, ROS measurement, caspase 9 activity assay PLoS ONE Medium 33211772
2023 Disease-associated polyglutamine inclusions disrupt formation of the BAG6-UBL4A complex. UBL4A also dissociates from BAG6 in response to proteasomal inhibition and mitochondrial depolarization (proteotoxic stresses), implying that disruption of the BAG6-UBL4A complex contributes to impaired TA protein biogenesis under stress. Co-immunoprecipitation under proteotoxic stress conditions (polyglutamine, proteasome inhibitor, mitochondrial depolarizing agent) Biochemical Journal Medium 37747814
2007 Yeast Mdy2 (UBL4A ortholog) interacts with the N-terminal region of Sgt2, and Mdy2 also physically interacts with the chaperone Ydj1, potentially mediating association between Ydj1 and Sgt2. MDY2 interacts genetically with YDJ1. Co-immunoprecipitation, genetic interaction (synthetic lethality/mating efficiency assay) Cell Stress & Chaperones Medium 17441508
2006 Yeast Mdy2 (UBL4A ortholog) contains a UBL domain but shows no evidence of C-terminal processing typical of ubiquitin. In mdy2Δ cells, microtubule bundles and the MT end-binding protein Kar9 fail to localize properly to the shmoo tip, causing defects in nuclear migration and karyogamy during mating. Deletion mutant phenotypic analysis, GFP localization, His-tag processing assay, immunofluorescence of microtubules and Kar9 Journal of Cell Science Medium 16390866
2010 Yeast Mdy2 (UBL4A ortholog) associates with α-tubulin and the dynactin subunit p150(Glued)/Nip100. Under heat stress, nuclear Mdy2 relocalizes to cytoplasmic stress granules co-marked by Pab1 (poly(A)-binding protein). Co-precipitation, GFP live imaging, colocalization with stress granule markers Cytoskeleton Medium 20722039
2012 Nuclear import of yeast Mdy2 (UBL4A ortholog) is mediated by an N-terminal nuclear localization signal (NLS) and is required for heat stress response. Mdy2 physically interacts with Pab1 and this interaction (and accumulation in stress granules) depends on nuclear history (NLS intact), not nuclear retention (NES deletion has no effect). NLS/NES deletion mutants, GFP localization, co-precipitation with Pab1, heat stress viability assays PLoS ONE Medium 23285234
2013 Crystal structure of the Sgt2 dimerization domain complexed with the Get5 UBL domain (yeast) reveals one Sgt2 dimer binding one Get5 monomer via hydrophobic residues from both proteins; these hydrophobic interface residues are important for cell survival under heat stress. Crystal structure, mutagenesis, heat stress cell survival assay Acta Crystallographica Section D High 24100326
2024 GdX/UBL4A directly interacts with STAT3 (confirmed by dual-luciferase reporter and immunofluorescence) and overexpression of GdX reduces phosphorylation of STAT3, inhibiting downstream STAT3 target genes BCL-XL, Cyclin D1, and c-Myc in breast cancer cells. Dual-luciferase reporter assay, immunofluorescence co-localization, western blot phosphorylation assay, KO mouse tumor model Cancer Biology & Therapy Low 39487760
2011 In yeast, sgt2Δ get5Δ double mutants show more severe TA protein sorting defects than either single knockout, indicating cooperative functions. Overproduction of Sgt2 is toxic in get3Δ but not get5Δ cells, indicating a Get5-independent role for Sgt2 in TA protein delivery to Get3. Genetic epistasis (double-mutant analysis), TA protein sorting assays, overexpression toxicity assay Biological Chemistry Medium 21619481

Source papers

Stage 0 corpus · 37 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2012 SGTA recognizes a noncanonical ubiquitin-like domain in the Bag6-Ubl4A-Trc35 complex to promote endoplasmic reticulum-associated degradation. Cell reports 80 23246001
2010 Crystal structure of Get4-Get5 complex and its interactions with Sgt2, Get3, and Ydj1. The Journal of biological chemistry 77 20106980
2010 Structural characterization of the Get4/Get5 complex and its interaction with Get3. Proceedings of the National Academy of Sciences of the United States of America 71 20554915
2011 A structural model of the Sgt2 protein and its interactions with chaperones and the Get4/Get5 complex. The Journal of biological chemistry 57 21832041
2012 Structures of the Sgt2/SGTA dimerization domain with the Get5/UBL4A UBL domain reveal an interaction that forms a conserved dynamic interface. Cell reports 56 23142665
2019 UBL4A inhibits autophagy-mediated proliferation and metastasis of pancreatic ductal adenocarcinoma via targeting LAMP1. Journal of experimental & clinical cancer research : CR 54 31288830
2014 Crystal structure of ATP-bound Get3-Get4-Get5 complex reveals regulation of Get3 by Get4. Nature structural & molecular biology 51 24727835
2007 SGT2 and MDY2 interact with molecular chaperone YDJ1 in Saccharomyces cerevisiae. Cell stress & chaperones 41 17441508
2015 Ubl4A is required for insulin-induced Akt plasma membrane translocation through promotion of Arp2/3-dependent actin branching. Proceedings of the National Academy of Sciences of the United States of America 39 26195787
2013 Nuclear BAG6-UBL4A-GET4 complex mediates DNA damage signaling and cell death. The Journal of biological chemistry 35 23723067
2010 Structures of Get3, Get4, and Get5 provide new models for TA membrane protein targeting. Structure (London, England : 1993) 35 20696390
2011 Cooperative and independent activities of Sgt2 and Get5 in the targeting of tail-anchored proteins. Biological chemistry 29 21619481
2019 Absence of GdX/UBL4A Protects against Inflammatory Diseases by Regulating NF-кB Signaling in Macrophages and Dendritic Cells. Theranostics 28 30867837
2015 Structure of a BAG6 (Bcl-2-associated athanogene 6)-Ubl4a (ubiquitin-like protein 4a) complex reveals a novel binding interface that functions in tail-anchored protein biogenesis. The Journal of biological chemistry 28 25713138
2013 Structure of the Sgt2/Get5 complex provides insights into GET-mediated targeting of tail-anchored membrane proteins. Proceedings of the National Academy of Sciences of the United States of America 28 23297211
1990 Linkage and sequence conservation of the X-linked genes DXS253E (P3) and DXS254E (GdX) in mouse and man. Genomics 21 1973144
2014 Solution structure of the SGTA dimerisation domain and investigation of its interactions with the ubiquitin-like domains of BAG6 and UBL4A. PloS one 17 25415308
2018 GdX/UBL4A null mice exhibit mild kyphosis and scoliosis accompanied by dysregulation of osteoblastogenesis and chondrogenesis. Cell biochemistry and function 14 29464742
2006 Mdy2, a ubiquitin-like (UBL)-domain protein, is required for efficient mating in Saccharomyces cerevisiae. Journal of cell science 14 16390866
2011 Interaction surface and topology of Get3-Get4-Get5 protein complex, involved in targeting tail-anchored proteins to endoplasmic reticulum. The Journal of biological chemistry 13 22190685
2014 Evaluation of retinal nerve fiber layer thickness measurements for glaucoma detection: GDx ECC versus spectral-domain OCT. Journal of glaucoma 10 23970337
2016 Deficiency in ubiquitin-like protein Ubl4A impairs migration of fibroblasts and macrophages. Biochemical and biophysical research communications 9 27998771
2013 Structure of the Sgt2 dimerization domain complexed with the Get5 UBL domain involved in the targeting of tail-anchored membrane proteins to the endoplasmic reticulum. Acta crystallographica. Section D, Biological crystallography 9 24100326
2023 Circ_0006944 aggravates LPS-induced HK2 cell injury via modulating miR-205-5p/UBL4A pathway. Autoimmunity 8 37994026
2019 UBL4A Augments Innate Immunity by Promoting the K63-Linked Ubiquitination of TRAF6. Journal of immunology (Baltimore, Md. : 1950) 8 31451677
2019 GdX/UBL4A-knockout mice resist collagen-induced arthritis by balancing the population of Th1/Th17 and regulatory T cells. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 7 31002527
2012 Generation of mice with conditional null allele for GdX/Ubl4A. Genesis (New York, N.Y. : 2000) 7 22139977
2012 1H, 13C and 15N assignments of Sgt2 N-terminal dimerisation domain and its binding partner, Get5 Ubiquitin-like domain. Biomolecular NMR assignments 6 23001946
2020 Ubl4A is critical for mitochondrial fusion process under nutrient deprivation stress. PloS one 5 33211772
2012 Nuclear import of UBL-domain protein Mdy2 is required for heat-induced stress response in Saccharomyces cerevisiae. PloS one 5 23285234
2010 The yeast ubiquitin-like domain protein Mdy2 is required for microtubule-directed nuclear migration and localizes to cytoplasmic granules in response to heat stress. Cytoskeleton (Hoboken, N.J.) 5 20722039
2024 GdX inhibits the occurrence and progression of breast cancer by negatively modulating the activity of STAT3. Cancer biology & therapy 3 39487760
2018 The C-terminus of Ubl4A is critical for pro-death activity and association with the Arp2/3 complex. Biochemical and biophysical research communications 2 30146258
2023 Proteotoxic stresses stimulate dissociation of UBL4A from the tail-anchored protein recognition complex. The Biochemical journal 1 37747814
2021 The Dispensable Roles of X-Linked Ubl4a and Its Autosomal Counterpart Ubl4b in Spermatogenesis Represent a New Evolutionary Type of X-Derived Retrogenes. Frontiers in genetics 1 34249105
2026 UBL4A as a suppressor of intracerebral hemorrhage by binding to MAPK14 and promoting MAPK14 ubiquitination. Genes & genomics 0 41575627
2025 Gdx mediates low-affinity Cs⁺/H⁺ antiport and confers cesium resistance in Escherichia coli. Engineering microbiology 0 41982384

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