Affinage

USP12

Ubiquitin carboxyl-terminal hydrolase 12 · UniProt O75317

Length
370 aa
Mass
42.9 kDa
Annotated
2026-04-28
33 papers in source corpus 25 papers cited in narrative 25 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

USP12 is a deubiquitinating enzyme that functions as the catalytic subunit of a ternary complex with the WD40-repeat proteins UAF1 (WDR48) and WDR20, which are required for full enzymatic activation and proper subcellular localization (PMID:20147737, PMID:27650958, PMID:30466959). The USP12–UAF1–WDR20 complex deubiquitinates a remarkably broad substrate repertoire—including histones H2A/H2B, the androgen receptor, PHLPP phosphatases, Notch, MDM2, NF-κB pathway components (BCL10, p65, PPM1B), innate immune sensors (IFI16), and endosomal cargo (β1 integrin)—thereby stabilizing these proteins and controlling transcription, PI3K–Akt signaling, TCR and innate immune signaling, Notch trafficking, and integrin recycling (PMID:21183687, PMID:24056413, PMID:24145035, PMID:22778262, PMID:26811477, PMID:33941870, PMID:37410794). In addition to its catalytic deubiquitinase function, USP12 inhibits CBP acetyltransferase activity through a non-catalytic mechanism, sustaining nuclear phospho-STAT1 during interferon signaling (PMID:31899788). USP12 dynamically shuttles between the plasma membrane, cytoplasm, and nucleus via CRM1-dependent export directed by WDR20, with TCR or IFN stimulation triggering redistribution that dictates substrate access (PMID:30466959, PMID:26811477).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2010 High

    Identification of USP12 as a histone-directed DUB and discovery of its obligate activator UAF1/WDR48 established that USP12 requires a WD40-repeat cofactor for catalytic competence, setting it apart from most USP-family members.

    Evidence In vitro deubiquitination of nucleosomal H2A/H2B, co-IP with WDR48, Xenopus knockdown/overexpression with ChIP

    PMID:21183687

    Open questions at the time
    • Whether UAF1 binding induces conformational change or allosteric activation was unresolved
    • Relative contributions of H2A vs H2B deubiquitination to transcriptional output not determined
  2. 2010 High

    Discovery that WDR20 forms a ternary complex with USP12–UAF1 and specifically stimulates USP12 (but not USP1) activity revealed a second cofactor layer that distinguishes USP12/USP46 from other UAF1-dependent DUBs.

    Evidence Tandem affinity purification, in vitro DUB activity assay with recombinant ternary complex

    PMID:20147737

    Open questions at the time
    • Mechanism by which WDR20 stimulates activity unknown at this point
    • Physiological substrates of the full ternary complex not yet identified
  3. 2012 High

    Demonstrating that USP12 deubiquitinates non-activated Notch to promote its lysosomal trafficking revealed the first non-histone substrate and placed USP12 in endosomal sorting, where deubiquitination paradoxically promotes degradation rather than stabilization.

    Evidence shRNA screen, in vitro DUB assay on Notch, trafficking assay by flow cytometry and confocal imaging, Drosophila homolog validation

    PMID:22778262

    Open questions at the time
    • Whether USP12 acts on Notch at the plasma membrane vs endosome not resolved
    • Identity of the E3 ligase counteracting USP12 on Notch unknown
  4. 2013 High

    Identification of the androgen receptor and PHLPP phosphatases as USP12 substrates connected the enzyme to two converging oncogenic pathways—AR transcription and PI3K–Akt signaling—in prostate cancer.

    Evidence Co-IP, ubiquitination/deubiquitination assays, Akt phosphorylation and AR reporter assays in prostate cancer cells

    PMID:24056413 PMID:24145035

    Open questions at the time
    • Whether AR and PHLPP are deubiquitinated by the same pool of USP12 complex unknown
    • No structural basis for substrate recognition
  5. 2016 High

    The crystal structure of USP12–Ub bound to UAF1 resolved the activation mechanism, revealing a 1:2 stoichiometry with high- and low-affinity UAF1 binding sites mediated by the USP12 fingers subdomain.

    Evidence X-ray crystallography at 2.8 Å, mutagenesis, binding stoichiometry assays

    PMID:27650958

    Open questions at the time
    • Structure of the full ternary complex including WDR20 not determined
    • Structural basis for substrate selectivity remains unknown
  6. 2016 High

    Showing that USP12 is phosphorylated and translocates from nucleus to cytosol upon TCR stimulation to deubiquitinate LAT and Trat1 established USP12 as a signal-dependent effector in adaptive immunity that maintains proximal TCR signaling complexes.

    Evidence Activity-based DUB probes, USP12-KO Jurkat cells with rescue, BioID proximity labeling, co-IP, flow cytometry

    PMID:26811477

    Open questions at the time
    • Kinase responsible for USP12 phosphorylation not identified
    • Whether USP12 acts similarly in primary human T cells not confirmed
  7. 2018 Medium

    Characterization of USP12's dynamic subcellular shuttling—WDR20 recruits USP12 to the plasma membrane while CRM1-dependent export controls nuclear exit—explained how a single DUB accesses substrates in distinct compartments.

    Evidence Confocal and live-cell microscopy, mutagenesis of N-terminal MEIL motif, leptomycin B treatment

    PMID:30466959

    Open questions at the time
    • Functional consequence of membrane localization for specific substrates not defined
    • Whether phosphorylation regulates this shuttling is untested
  8. 2020 Medium

    Discovery that USP12 inhibits CBP acetyltransferase activity independently of its DUB catalytic function revealed a non-enzymatic role in sustaining nuclear phospho-STAT1 during IFN signaling, broadening USP12's functional repertoire beyond deubiquitination.

    Evidence CBP HAT activity assay with catalytic-dead USP12 mutant, subcellular fractionation, p-STAT1 dephosphorylation assay

    PMID:31899788

    Open questions at the time
    • Structural basis for CBP HAT domain inhibition not resolved
    • Whether this non-catalytic function extends to other acetyltransferases untested
    • Single-study finding
  9. 2021 Medium

    A burst of substrate identifications—HMGB1, PPM1B, BCL10, p300, midkine, Bax, and DMWD as an alternative WDR20-like cofactor—collectively demonstrated that USP12 is a broadly acting stabilizer of signaling regulators across NF-κB, autophagy, apoptosis, and epigenetic pathways.

    Evidence Multiple independent Co-IP and ubiquitination assays, KO mouse models (PPM1B, BCL10), autophagy flux (HMGB1), cardiomyocyte models (p300), Bax mutagenesis

    PMID:33844468 PMID:33941870 PMID:34339675 PMID:34381028 PMID:34759262 PMID:34997217 PMID:36361894

    Open questions at the time
    • Most substrates validated by single labs; independent replication needed
    • Hierarchy or competition among substrates for USP12 binding is unexplored
    • Whether DMWD and WDR20 direct USP12 to distinct substrate pools is unresolved
  10. 2023 Medium

    Identification of IFI16 and RRM2 as USP12 substrates extended its roles to innate antiviral immunity (IFI16–STING–IRF3 axis) and DNA replication (dNTP supply via RRM2 stability), linking USP12 to genome maintenance.

    Evidence USP12 KO/KD with Co-IP and ubiquitination assays, IFN-β/ISG induction and viral replication assays (IFI16), DNA replication stress markers and in vivo tumor growth (RRM2)

    PMID:37341611 PMID:37410794

    Open questions at the time
    • Whether USP12 regulates RRM2 during normal S-phase or only under oncogenic stress unclear
    • Redundancy with USP46 for these substrates not assessed
  11. 2024 Medium

    Studies on YAP, c-Myc, β1 integrin, and FOXO3 further expanded the USP12 substrate repertoire to Hippo signaling, oncogene stabilization, endosomal integrin recycling, and sepsis-induced apoptosis, underscoring USP12 as a context-dependent stabilizer of diverse signaling nodes.

    Evidence DUB siRNA screens, BioID and genetic screens (integrin), Co-IP with ubiquitination site mutagenesis (YAP K315), patient-derived organoids and transgenic mice (c-Myc), LPS-SIMD models (FOXO3)

    PMID:38605077 PMID:39432777 PMID:39662205

    Open questions at the time
    • Integrin deubiquitination data from preprint awaiting peer review
    • How USP12 distinguishes >20 substrates through a single catalytic domain lacks structural explanation
    • Relative physiological importance of individual substrates in vivo not ranked

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include: (1) the structural basis for WDR20/DMWD-dependent activation and substrate selectivity in the full ternary complex, (2) whether distinct cofactor combinations (WDR20 vs DMWD) channel USP12 to specific substrate pools in vivo, and (3) how signal-dependent post-translational modifications orchestrate USP12 localization and substrate access across its many reported pathways.
  • No structure of USP12–UAF1–WDR20 ternary complex
  • Kinase(s) and phosphorylation sites controlling USP12 relocalization unidentified
  • Systematic comparison of USP12 vs USP46 substrate specificity lacking

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140096 catalytic activity, acting on a protein 17 GO:0098772 molecular function regulator activity 1
Localization
GO:0005634 nucleus 4 GO:0005768 endosome 2 GO:0005829 cytosol 2 GO:0005886 plasma membrane 1
Pathway
R-HSA-162582 Signal Transduction 8 R-HSA-1643685 Disease 5 R-HSA-168256 Immune System 4 R-HSA-74160 Gene expression (Transcription) 3 R-HSA-5357801 Programmed Cell Death 2 R-HSA-9609507 Protein localization 2 R-HSA-4839726 Chromatin organization 1 R-HSA-9612973 Autophagy 1
Complex memberships
USP12–UAF1(WDR48)–DMWD ternary complexUSP12–UAF1(WDR48)–WDR20 ternary complex

Evidence

Reading pass · 25 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2010 USP12 and USP46 deubiquitinate histone H2A and H2B (preferring nucleosomal substrates) in vitro and in vivo; WDR48 (a WD40 repeat protein) interacts with USP12 and USP46 and is required for this histone deubiquitination activity. In vitro deubiquitination assay with nucleosomal substrates, co-immunoprecipitation, Xenopus knockdown/overexpression with ChIP The Journal of biological chemistry High 21183687
2010 WDR20, a WD40-repeat protein, forms a ternary complex with USP12 and UAF1 (WDR48), and WDR20 stimulates the enzymatic activity of the USP12·UAF1 complex but not of USP1·UAF1, distinguishing the USP12/USP46 complexes from the USP1 complex. Tandem affinity purification, co-immunoprecipitation, in vitro deubiquitinase activity assay, siRNA depletion The Journal of biological chemistry High 20147737
2013 USP12, in complex with UAF1 and WDR20, deubiquitinates the androgen receptor (AR), enhancing AR protein stability and transcriptional activity in prostate cancer cells. siRNA screen, co-immunoprecipitation, ubiquitination assay, AR stability and transcriptional reporter assays The Journal of biological chemistry High 24056413
2013 The WDR48·USP12 complex deubiquitinates PHLPP1, stabilizing it and thereby suppressing Akt activation and promoting apoptosis; identified via tandem affinity purification. Tandem affinity purification, co-immunoprecipitation, ubiquitination/deubiquitination assay, Akt phosphorylation assay, proliferation assay The Journal of biological chemistry High 24145035
2012 USP12 (with activator UAF1) deubiquitinates the non-activated form of Notch receptor, promoting its trafficking to lysosomes for degradation; USP12 silencing interrupts Notch lysosomal trafficking, increasing surface Notch and Notch signaling activity. shRNA library screen, in vitro deubiquitination assay, Notch trafficking assay (flow cytometry, confocal imaging), Drosophila homolog validation The Journal of biological chemistry High 22778262
2016 Crystal structure of USP12-Ub/UAF1 complex (2.8 Å) revealed two UAF1 binding sites; USP12/UAF1 complex has 1:2 stoichiometry with high-affinity (4 nM) and low-affinity (325 nM) binding steps; the fingers subdomain of USP12 mediates the high-affinity UAF1 interface required for activation. X-ray crystallography, mutagenesis, binding assays (stoichiometry determination), enzymatic activity assays Journal of structural biology High 27650958
2016 Upon TCR stimulation, USP12 undergoes phosphorylation and translocates from the nucleus to the cytosol; USP12 deubiquitylates LAT and Trat1 (TCR adaptor proteins), preventing their lysosomal degradation and maintaining the proximal TCR complex for sustained NFκB, NFAT, and MAPK signaling. Activity-based DUB probes (HA-Ub-VME), USP12-KO Jurkat cells, proximity-based BirA labeling, co-immunoprecipitation, rescue experiments, flow cytometry Proceedings of the National Academy of Sciences of the United States of America High 26811477
2014 USP12, in complex with UAF1 and WDR20, directly deubiquitinates and stabilizes PHLPP and PHLPPL phosphatases, reducing active pAkt levels and consequently decreasing AR phosphorylation at Ser213, thereby enhancing AR stability and transcriptional activity. Co-immunoprecipitation, ubiquitination/deubiquitination assay, Akt phosphorylation assay, AR transcriptional reporter Oncotarget High 25216524
2018 USP12 deubiquitinates MDM2 and AR, controlling TP53 tumor suppressor levels and AR oncogene levels in prostate cancer; identified via transcriptome analysis and denaturing immunoprecipitations. Transcriptome analysis, denaturing co-immunoprecipitation, immunohistochemistry Oncogene Medium 29755129
2018 WDR20 promotes recruitment of USP12 (but not USP46) to the plasma membrane, and the USP12/UAF1/WDR20 complex dynamically shuttles between plasma membrane, cytoplasm, and nucleus via CRM1-dependent nuclear export; this requires a short N-terminal motif (1MEIL4) in USP12 and a NES in WDR20. Confocal and live microscopy, site-directed mutagenesis, CRM1 inhibitor (leptomycin B) treatment, co-immunoprecipitation European journal of cell biology Medium 30466959
2020 USP12 interacts with the HAT domain of CBP and inhibits CBP acetyltransferase activity independently of its deubiquitinase activity; during IFN signaling, USP12 translocates from cytoplasm to nucleus, where nuclear USP12 accumulation blocks CBP-mediated acetylation of phospho-STAT1, inhibiting TCPTP-dependent dephosphorylation and maintaining nuclear p-STAT1 levels. Co-immunoprecipitation, CBP acetyltransferase activity assay, subcellular fractionation, catalytic mutant USP12, p-STAT1 dephosphorylation assay PLoS pathogens Medium 31899788
2021 USP12 interacts with, deubiquitylates, and stabilizes HMGB1, sustaining HMGB1-mediated pro-survival autophagy and contributing to bortezomib resistance in multiple myeloma. Co-immunoprecipitation, ubiquitination assay, USP12 knockdown, autophagy flux assay Oncogene Medium 34997217
2021 USP12 deubiquitinates and stabilizes PPM1B (a phosphatase), and USP12 downregulation causes insufficient PPM1B deubiquitination leading to NF-κB hyperactivation in tumor cells, creating an immunosuppressive microenvironment. Co-immunoprecipitation, ubiquitination assay, USP12 KO mouse lung tumor model, NF-κB reporter Nature communications Medium 34381028
2021 USP12 stabilizes BCL10 by deubiquitinating it, thereby activating NF-κB signaling specifically in CD4+ T cells to promote their activation, differentiation, and proliferation. USP12-KO mouse model, co-immunoprecipitation, ubiquitination assay, NF-κB signaling assay, CD4+ T cell functional assays Cell death and differentiation Medium 33941870
2021 USP12 binds and stabilizes p300 by deubiquitination, upregulating p300-dependent transcription of METTL3, which catalyzes m6A modification on mRNAs and promotes cardiac hypertrophy. Co-immunoprecipitation, ubiquitination assay, USP12 KO/OE in cardiomyocytes and in vivo, METTL3 rescue experiment Experimental cell research Medium 34339675
2021 USP12 deubiquitinates and stabilizes midkine (MDK) in breast cancer cells, preventing its polyubiquitination-mediated degradation and promoting tumor angiogenesis. Co-immunoprecipitation, ubiquitination assay, MDK overexpression rescue, HUVEC tube formation and migration assays, in vivo metastasis model Cell death & disease Medium 34759262
2022 USP12 deubiquitinates and stabilizes p65 (NF-κB subunit) in monocytic MDSCs, upregulating iNOS and PD-L1 expression to suppress CD8+ T cell responses and promote tumor immune evasion. Co-immunoprecipitation, ubiquitination assay, USP12-KO mouse model, T cell suppression assay Immunology Medium 35898171
2022 USP12 interacts with Bax and removes K63-linked ubiquitin chains from Bax (at K128 and K190 sites), affecting Bax protein half-life without targeting it for proteasomal degradation. Yeast two-hybrid, co-immunoprecipitation, site-directed mutagenesis of Bax ubiquitination sites, half-life assay International journal of molecular sciences Medium 36361894
2023 USP12 inhibits proteasome-dependent degradation of IFI16 through its deubiquitinase activity, maintaining IFI16 stability and promoting IFI16-STING-IRF3/p65-mediated antiviral signaling against HSV-1. USP12 KO/KD, co-immunoprecipitation, ubiquitination assay, IFN-β/ISG expression assay, viral replication assay PLoS pathogens Medium 37410794
2023 USP12 directly interacts with and deubiquitinates RRM2 (ribonucleotide reductase catalytic subunit), stabilizing it in non-small cell lung cancer cells; USP12 knockdown causes DNA replication stress. Co-immunoprecipitation, ubiquitination assay, USP12 KD, DNA replication stress markers, in vivo tumor growth assay Molecular carcinogenesis Medium 37341611
2024 USP12/46-WDR48-WDR20 ternary complex removes ubiquitin from the cytoplasmic tail of internalized β1 integrin (Itgb1) in early endosomes, preventing ESCRT-mediated sorting and lysosomal degradation, thereby stabilizing integrin surface levels. Genetic screen, proximity-dependent biotin identification (BioID), co-immunoprecipitation, integrin surface level assay, ESCRT pathway analysis bioRxivpreprint Medium bio_10.1101_2024.05.14.594138
2024 WDR20 and USP12/46 co-deubiquitinate c-Myc simultaneously to maintain its stability; WDR20 silencing disturbs c-Myc protein stability, promoting CDKN1A transcription and HCC cellular senescence. siRNA screen of WDR-domain proteins, co-immunoprecipitation, ubiquitination assay, ChIP, patient-derived organoids, xenograft and transgenic mouse models Proceedings of the National Academy of Sciences of the United States of America Medium 39432777
2024 USP12 interacts with YAP, inhibits K48-linked polyubiquitination of YAP at K315, and stabilizes YAP in the nucleus to promote Hippo/YAP signaling in gastric cancer. DUB siRNA library screen, co-immunoprecipitation, ubiquitination site mutagenesis (K315R), immunostaining for nuclear localization, in vitro and in vivo tumor growth assays Cell death discovery Medium 38605077
2021 DMWD (dystrophia myotonica WD repeat protein), analogous to WDR20, directly binds USP12 at the same interface as WDR20 (suggesting mutually exclusive binding), promotes USP12 enzymatic activity, but differentially modulates USP12 subcellular localization compared to WDR20. Co-immunoprecipitation of epitope-tagged proteins, enzymatic activity assay, subcellular localization imaging, phylogenetic analysis The FEBS journal Medium 33844468
2024 METTL3/YTHDF1-mediated m6A modification stabilizes USP12 mRNA; elevated USP12 then interacts with and deubiquitinates FOXO3, preventing its ubiquitin-mediated degradation and leading to PUMA upregulation and intrinsic apoptosis in sepsis-induced myocardial dysfunction. Proteomic profiling, m6A modification analysis, co-immunoprecipitation, ubiquitination assay, in vitro/in vivo LPS-SIMD model, pathway inhibitor experiments Molecular immunology Medium 39662205

Source papers

Stage 0 corpus · 33 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2010 Regulation of histone H2A and H2B deubiquitination and Xenopus development by USP12 and USP46. The Journal of biological chemistry 98 21183687
2010 WDR20 regulates activity of the USP12 x UAF1 deubiquitinating enzyme complex. The Journal of biological chemistry 80 20147737
2013 Deubiquitinating enzyme Usp12 is a novel co-activator of the androgen receptor. The Journal of biological chemistry 75 24056413
2013 WD repeat protein WDR48 in complex with deubiquitinase USP12 suppresses Akt-dependent cell survival signaling by stabilizing PH domain leucine-rich repeat protein phosphatase 1 (PHLPP1). The Journal of biological chemistry 62 24145035
2012 The ubiquitin-specific protease 12 (USP12) is a negative regulator of notch signaling acting on notch receptor trafficking toward degradation. The Journal of biological chemistry 60 22778262
2015 The EBNA3 family of Epstein-Barr virus nuclear proteins associates with the USP46/USP12 deubiquitination complexes to regulate lymphoblastoid cell line growth. PLoS pathogens 49 25855980
2014 Deubiquitinating enzyme Usp12 regulates the interaction between the androgen receptor and the Akt pathway. Oncotarget 47 25216524
2016 Usp12 stabilizes the T-cell receptor complex at the cell surface during signaling. Proceedings of the National Academy of Sciences of the United States of America 44 26811477
2021 USP12 downregulation orchestrates a protumourigenic microenvironment and enhances lung tumour resistance to PD-1 blockade. Nature communications 41 34381028
2018 Molecular mechanism of the TP53-MDM2-AR-AKT signalling network regulation by USP12. Oncogene 36 29755129
2022 Deubiquitylase USP12 induces pro-survival autophagy and bortezomib resistance in multiple myeloma by stabilizing HMGB1. Oncogene 35 34997217
2021 De-ubiquitination of p300 by USP12 Critically Enhances METTL3 Expression and Ang II-induced cardiac hypertrophy. Experimental cell research 31 34339675
2016 A conserved two-step binding for the UAF1 regulator to the USP12 deubiquitinating enzyme. Journal of structural biology 29 27650958
2018 The novel anti-androgen candidate galeterone targets deubiquitinating enzymes, USP12 and USP46, to control prostate cancer growth and survival. Oncotarget 24 29861848
2020 USP12 translocation maintains interferon antiviral efficacy by inhibiting CBP acetyltransferase activity. PLoS pathogens 22 31899788
2021 USP12 promotes breast cancer angiogenesis by maintaining midkine stability. Cell death & disease 20 34759262
2021 USP12 promotes CD4+ T cell responses through deubiquitinating and stabilizing BCL10. Cell death and differentiation 19 33941870
2015 USP12 regulates cell cycle progression by involving c-Myc, cyclin D2 and BMI-1. Gene 17 26680102
2023 USP12 promotes antiviral responses by deubiquitinating and stabilizing IFI16. PLoS pathogens 16 37410794
2022 USP12 positively regulates M-MDSC function to inhibit antitumour immunity through deubiquitinating and stabilizing p65. Immunology 15 35898171
2024 WDR20 prevents hepatocellular carcinoma senescence by orchestrating the simultaneous USP12/46-mediated deubiquitination of c-Myc. Proceedings of the National Academy of Sciences of the United States of America 13 39432777
2023 Spotlights on ubiquitin-specific protease 12 (USP12) in diseases: from multifaceted roles to pathophysiological mechanisms. Journal of translational medicine 11 37752518
2017 Deubiquitinase USP12 promotes LPS induced macrophage responses through inhibition of IκBα. Biochemical and biophysical research communications 11 28063927
2024 USP12 facilitates gastric cancer progression via stabilizing YAP. Cell death discovery 10 38605077
2023 USP12 promotes nonsmall cell lung cancer progression through deubiquitinating and stabilizing RRM2. Molecular carcinogenesis 10 37341611
2022 Deubiquitinating Enzyme USP12 Regulates the Pro-Apoptosis Protein Bax. International journal of molecular sciences 10 36361894
2023 USP12 regulates ER stress-associated osteogenesis in human periodontal ligament cells under tension stress. Cellular signalling 9 38113977
2018 WDR20 regulates shuttling of the USP12 deubiquitinase complex between the plasma membrane, cytoplasm and nucleus. European journal of cell biology 9 30466959
2020 Downregulation of USP12 inhibits tumor growth via the p38/MAPK pathway in hepatocellular carcinoma. Molecular medicine reports 8 33174033
2022 Grouper USP12 exerts antiviral activity against nodavirus infection. Fish & shellfish immunology 6 35032679
2024 METTL3/YTHDF1-mediated m6A modification stabilizes USP12 to deubiquitinate FOXO3 and promote apoptosis in sepsis-induced myocardial dysfunction. Molecular immunology 5 39662205
2024 Membrane palmitoylated protein MPP1 inhibits immune escape by regulating the USP12/ CCL5 axis in urothelial carcinoma. International immunopharmacology 3 39700963
2021 The dystrophia myotonica WD repeat-containing protein DMWD and WDR20 differentially regulate USP12 deubiquitinase. The FEBS journal 3 33844468