Affinage

PSMD11

26S proteasome non-ATPase regulatory subunit 11 · UniProt O00231

Round 2 corrected
Length
422 aa
Mass
47.5 kDa
Annotated
2026-04-28
130 papers in source corpus 10 papers cited in narrative 10 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PSMD11 (Rpn6) is a non-ATPase subunit of the 19S regulatory particle lid that serves as a rate-limiting determinant of 26S proteasome assembly and activity. Rpn6 is essential for incorporating a subset of lid subunits (Rpn3, Rpn7, Rpn12, Sem1/Rpn15) into the 19S complex, and its abundance directly controls the proportion of assembled, doubly-capped 26S proteasomes capable of degrading ubiquitinated substrates (PMID:15611133, PMID:22972301). PKA-mediated phosphorylation at Ser14, triggered by cAMP-raising hormones (glucagon, epinephrine, vasopressin) and physiological states such as fasting and exercise, acutely stimulates 26S proteasome activity and enhances clearance of aggregation-prone proteins in a tissue-specific manner (PMID:26669444, PMID:30782827). PSMD11 expression is transcriptionally regulated by FOXO4 in human embryonic stem cells and is additionally modulated by AMPK and MEK1/ERK1/2 signaling, positioning PSMD11 as a convergence point through which multiple signaling pathways tune cellular proteostasis (PMID:22972301, PMID:19616115, PMID:29665121).

Mechanistic history

Synthesis pass · year-by-year structured walk · 9 steps
  1. 2002 Medium

    Establishing that Rpn6 physically interfaces with the COP9 signalosome via its PCI domain addressed whether the 19S lid shares functional crosstalk with structurally related PCI-containing complexes.

    Evidence Yeast two-hybrid and PCI domain mapping in Drosophila; P-element insertion alleles showing developmental lethality

    PMID:12426099

    Open questions at the time
    • Interaction detected by yeast two-hybrid without reciprocal Co-IP in Drosophila cells
    • Functional consequence of Rpn6–CSN2 interaction on proteasome or signalosome activity unknown
    • Not tested in mammalian systems
  2. 2004 High

    Defining Rpn6 as essential for lid assembly resolved how the 19S regulatory particle is built and identified the specific subunit recruitment step that depends on Rpn6.

    Evidence Temperature-sensitive yeast rpn6 mutants accumulating sub-lid complexes; native gel and affinity-purified proteasome analysis

    PMID:15611133

    Open questions at the time
    • Structural basis for how Rpn6 bridges the sub-lid to late-incorporating subunits not resolved
    • Whether the assembly role is conserved quantitatively in mammalian cells was untested
  3. 2009 Medium

    Identifying AMPK as a PSMD11-binding kinase raised the possibility that metabolic energy sensing directly modulates proteasome function.

    Evidence Yeast two-hybrid screen with constitutively active AMPKα2 bait from human skeletal muscle library; Co-IP and in vitro phosphorylation

    PMID:19616115

    Open questions at the time
    • Specific AMPK phosphorylation site(s) on PSMD11 not mapped
    • Functional consequence on proteasome activity not determined
    • Single-lab study without independent replication
  4. 2012 High

    Demonstrating that PSMD11 abundance is rate-limiting for 26S proteasome assembly in human stem cells established a direct link between proteostasis capacity and stem cell identity, with FOXO4 as the upstream transcriptional regulator.

    Evidence Ectopic PSMD11 overexpression increasing 26S/30S assembly and activity; FOXO4 knockdown reducing PSMD11; proteasome inhibition compromising pluripotency markers in hESCs

    PMID:22972301

    Open questions at the time
    • Whether FOXO4-driven PSMD11 expression is the dominant mechanism in somatic tissues unknown
    • Contribution of other lid subunit levels to the rate-limiting step not fully dissected
  5. 2015 High

    Identifying Ser14 as the PKA phosphorylation site on Rpn6 that directly stimulates 26S proteasome activity provided the first acute, post-translational mechanism for hormonal regulation of proteasome function.

    Evidence In vitro PKA phosphorylation of purified 26S proteasomes; S14D phosphomimetic and S14A mutants; degradation of neurodegenerative-disease-associated substrates in multiple cell lines

    PMID:26669444

    Open questions at the time
    • Structural mechanism by which Ser14 phosphorylation enhances gate opening or substrate engagement unresolved
    • Relative contribution of Rpn6-Ser14 versus other PKA proteasome targets not fully delineated
  6. 2018 Medium

    Finding that MEK1/ERK1/2 signaling acutely induces PSMD11 protein synthesis in cancer cells revealed an additional kinase pathway converging on PSMD11 expression, with therapeutic implications for drug resistance.

    Evidence Homoharringtonine treatment of pancreatic cancer cells; MEK1/ERK1/2 inhibition by sorafenib; in vivo validation in genetically engineered mouse model

    PMID:29665121

    Open questions at the time
    • Mechanism of selective PSMD11 translational upregulation under global translation inhibition unexplained
    • Single-lab study in pancreatic cancer context
  7. 2019 High

    Demonstrating that diverse cAMP-raising hormones and physiological states activate proteasomes through Rpn6-Ser14 phosphorylation in vivo extended the PKA–Rpn6 axis from biochemistry to whole-organism physiology, encompassing liver, heart, skeletal muscle, and kidney.

    Evidence In vivo proteasome activity and Rpn6 phosphorylation in multiple tissues; PKA-knockout cells; fasting and exercise models; human muscle biopsies

    PMID:30782827

    Open questions at the time
    • Tissue-specific quantitative contribution of Rpn6 phosphorylation versus transcriptional upregulation not resolved
    • Long-term physiological consequences of chronic Rpn6-Ser14 phosphorylation unexplored
  8. 2019 Medium

    Establishing that PSMD11 is a direct miR-451 target linked to NF-κB signaling in glomerular mesangial cells connected proteasome subunit expression to inflammatory pathway activation in diabetic nephropathy.

    Evidence Dual-luciferase reporter assay for miR-451/PSMD11; PSMD11 overexpression rescue; NF-κB pathway Western blots in high-glucose-treated mesangial cells

    PMID:31652441

    Open questions at the time
    • Whether PSMD11 activates NF-κB via proteasome-mediated IκBα degradation or a proteasome-independent mechanism not distinguished
    • Single cell type and single lab
  9. 2022 Medium

    Identification of Rpn6/PSMD11 as the direct molecular target of the small molecule DDO-7263 that blocks 26S proteasome assembly validated PSMD11 as a druggable node for modulating Nrf2 stability.

    Evidence Affinity chromatography with biotin/fluorescein probes; mass spectrometry target identification; proteasome assembly and Nrf2 degradation assays

    PMID:35253427

    Open questions at the time
    • Binding site and selectivity of DDO-7263 on Rpn6 not structurally resolved
    • In vivo pharmacokinetics and off-target effects not fully characterized

Open questions

Synthesis pass · forward-looking unresolved questions
  • The structural basis for how Ser14 phosphorylation and small-molecule binding to Rpn6 alter 26S proteasome conformation, gate opening, or substrate translocation remains unresolved, as does the functional significance of the AMPK–PSMD11 interaction.
  • No cryo-EM or crystal structure of phospho-Ser14 Rpn6 within the 26S proteasome
  • AMPK phosphorylation site(s) and functional outcome on proteasome activity not mapped
  • Whether PSMD11 has proteasome-independent scaffolding roles in NF-κB or Wnt/β-catenin signaling is unresolved

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 3 GO:0005198 structural molecule activity 2
Localization
GO:0005829 cytosol 2
Pathway
R-HSA-392499 Metabolism of proteins 4 R-HSA-162582 Signal Transduction 3 R-HSA-8953897 Cellular responses to stimuli 3
Partners
CSN2FOXO4PRKAA2PRKACAPSMD12PSMD3PSMD6
Complex memberships
19S regulatory particle lid26S proteasome

Evidence

Reading pass · 10 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2012 Human embryonic stem cells (hESCs) exhibit elevated proteasome activity correlated with increased PSMD11 (Rpn6) protein levels and enhanced 26S/30S proteasome assembly. Ectopic expression of PSMD11 alone is sufficient to increase proteasome assembly and activity, establishing PSMD11 as a rate-limiting regulator of proteasome assembly. FOXO4 transcription factor (downstream of insulin/IGF-I signaling) regulates PSMD11 expression in hESCs, linking proteostasis to longevity pathways. Ectopic overexpression of PSMD11 with proteasome activity assays; immunoprecipitation of 26S/30S complexes; FOXO4 knockdown; proteasome inhibition with pluripotency marker readout Nature High 22972301
2015 PKA (cAMP-dependent protein kinase) directly phosphorylates the 19S regulatory subunit Rpn6/PSMD11 at Ser14. This phosphorylation enhances 26S proteasome activity and the selective degradation of ubiquitinated proteins, short-lived proteins, and aggregation-prone neurodegenerative disease proteins (mutant FUS, SOD1, TDP43, tau). A phosphomimetic Rpn6 S14D mutant activates proteasomes, whereas a non-phosphorylatable S14A mutant decreases activity. cAMP-raising agents also increase formation of doubly-capped 26S proteasomes. In vitro PKA phosphorylation of purified 26S proteasomes; site-directed mutagenesis (S14D and S14A); proteasome activity assays with ubiquitinated substrates and peptides; protein phosphatase treatment; degradation assays in multiple mammalian cell lines Proceedings of the National Academy of Sciences of the United States of America High 26669444
2019 In vivo, diverse hormones and physiological states that raise cAMP (glucagon, epinephrine, vasopressin, fasting, exercise) activate 26S proteasomes through PKA-mediated phosphorylation of Rpn6/PSMD11 in a tissue-specific manner. Rpn6 phosphorylation and proteasome activation were demonstrated in hepatocytes, working hearts, skeletal muscles of exercising humans, electrically stimulated rat muscles, and kidney cells. PKA-knockout cells failed to respond to vasopressin, confirming the PKA→Rpn6 axis. In vivo and ex vivo proteasome activity assays; Rpn6 phosphorylation measurements by immunoblot; PKA knockout cells; hormone treatment of primary hepatocytes and intact tissues; exercise physiology samples Proceedings of the National Academy of Sciences of the United States of America High 30782827
2004 Yeast Rpn6p (PSMD11 ortholog) is essential for the structural integrity and assembly of the 19S lid complex. Temperature-sensitive rpn6 mutants accumulate a sub-lid complex containing only Rpn5p, Rpn8p, Rpn9p, and Rpn11p, demonstrating that Rpn6p is specifically required for the incorporation of Rpn3p, Rpn7p, Rpn12p, and Sem1p/Rpn15p into the lid. The defect is at the assembly level rather than in proteolytic function of pre-assembled proteasomes. Temperature-sensitive yeast mutants; affinity-purified 26S proteasome activity assays; polyubiquitinated substrate (Sic1p) degradation; native gel analysis of lid sub-complexes; in vivo protein degradation assays The Journal of biological chemistry High 15611133
2002 Drosophila Rpn6 (PSMD11 ortholog) physically interacts with the COP9 signalosome subunit Alien/CSN2 via its PCI domain. Rpn6 is essential for Drosophila development, as P-element insertion and newly generated alleles are lethal. The temporal and spatial expression patterns of Rpn6 and alien/CSN2 overlap during development, consistent with an in vivo interaction between the 19S lid and COP9 signalosome. Yeast two-hybrid screen; PCI domain interaction mapping; P-element insertion mutant analysis; developmental expression pattern analysis Gene Medium 12426099
2009 PSMD11 was identified as a novel physical interaction partner of AMP-activated protein kinase (AMPK) by yeast two-hybrid screening of a human skeletal muscle cDNA library using constitutively active AMPKα2 as bait. Co-IP confirmed the physical interaction, and AMPK was shown to modify the phosphorylation status of PSMD11, suggesting AMPK regulates proteasome function through PSMD11. Yeast two-hybrid screening; co-immunoprecipitation; in vitro phosphorylation assays The international journal of biochemistry & cell biology Medium 19616115
2022 The 1,2,4-oxadiazole derivative DDO-7263 (an Nrf2-ARE activator) directly binds Rpn6/PSMD11, as identified by affinity chromatography and mass spectrometry using biotin-labeled and fluorescein-based probes. DDO-7263 binding to Rpn6 blocks 26S proteasome assembly, preventing degradation of ubiquitinated Nrf2 and thereby activating the Nrf2 signaling pathway for cytoprotection. Affinity chromatography with biotin-labeled probe; mass spectrometry target identification; in vitro 26S proteasome assembly assays; Nrf2 ubiquitination and degradation assays Journal of medicinal chemistry Medium 35253427
2018 Homoharringtonine (HHT), a protein synthesis inhibitor, paradoxically induces rapid PSMD11 protein synthesis in pancreatic cancer cells (MiaPaCa-2, PANC-1) through activation of the MEK1/ERK1/2 signaling pathway. This acute PSMD11 upregulation protects cancer cells from HHT-induced apoptosis. Co-inhibition of MEK1/ERK1/2 with sorafenib improves HHT cytotoxicity in vitro and in a genetically engineered mouse model of pancreatic cancer. Western blot analysis of PSMD11 protein levels; MEK1/ERK1/2 pathway inhibition with sorafenib; in vitro proliferation/apoptosis assays; genetically engineered mouse model experiments Journal of cellular biochemistry Medium 29665121
2020 In a mouse epilepsy model, Psmd11 expression is downregulated in hippocampal tissues. Overexpression of Psmd11 improved spatial learning and memory, protected hippocampal neurons from injury, inhibited apoptosis, suppressed microglial/astrocyte activation, and reduced inflammatory responses. Mechanistically, Psmd11 inactivates the Wnt/β-catenin pathway. The lncRNA Peg13 upregulates Psmd11 by sponging miR-490-3p (which targets Psmd11 mRNA), establishing a Peg13/miR-490-3p/Psmd11/Wnt-β-catenin axis in epilepsy. Epileptic mouse model; stereotaxic overexpression; HE and Nissl staining; ELISA for cytokines; luciferase reporter assays for miR-490-3p/Psmd11 interaction; Western blot for Wnt/β-catenin pathway components American journal of translational research Medium 33437373
2019 MicroRNA-451 directly targets PSMD11 (validated by dual-luciferase reporter assay). In high-glucose-treated glomerular mesangial cells, PSMD11 promotes inflammation and proliferation through NF-κB p65 signaling (as evidenced by increased p-IκBα, NF-κB p65, COX-2, and cyclinD1). miR-451 mimics suppress these effects, which are reversed by PSMD11 overexpression, establishing a miR-451/PSMD11/NF-κB axis. Dual-luciferase reporter assay; miR-451 mimic transfection; PSMD11 overexpression; ELISA for inflammatory cytokines; Western blot for NF-κB pathway; flow cytometry for cell cycle Bioscience reports Medium 31652441

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Global, in vivo, and site-specific phosphorylation dynamics in signaling networks. Cell 2861 17081983
1996 Structure and functions of the 20S and 26S proteasomes. Annual review of biochemistry 2108 8811196
2002 Isolation of a human gene that inhibits HIV-1 infection and is suppressed by the viral Vif protein. Nature 1924 12167863
2005 A human protein-protein interaction network: a resource for annotating the proteome. Cell 1704 16169070
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2009 Recognition and processing of ubiquitin-protein conjugates by the proteasome. Annual review of biochemistry 1398 19489727
2009 Defining the human deubiquitinating enzyme interaction landscape. Cell 1282 19615732
2003 Broad antiretroviral defence by human APOBEC3G through lethal editing of nascent reverse transcripts. Nature 1236 12808466
2003 DNA deamination mediates innate immunity to retroviral infection. Cell 1150 12809610
2015 The BioPlex Network: A Systematic Exploration of the Human Interactome. Cell 1118 26186194
2017 Architecture of the human interactome defines protein communities and disease networks. Nature 1085 28514442
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2003 Induction of APOBEC3G ubiquitination and degradation by an HIV-1 Vif-Cul5-SCF complex. Science (New York, N.Y.) 1006 14564014
2003 The cytidine deaminase CEM15 induces hypermutation in newly synthesized HIV-1 DNA. Nature 912 12808465
2013 Landscape of the PARKIN-dependent ubiquitylome in response to mitochondrial depolarization. Nature 870 23503661
2020 A reference map of the human binary protein interactome. Nature 849 32296183
2018 VIRMA mediates preferential m6A mRNA methylation in 3'UTR and near stop codon and associates with alternative polyadenylation. Cell discovery 829 29507755
2003 The antiretroviral enzyme APOBEC3G is degraded by the proteasome in response to HIV-1 Vif. Nature medicine 798 14528300
2003 Species-specific exclusion of APOBEC3G from HIV-1 virions by Vif. Cell 763 12859895
2003 Complete sequencing and characterization of 21,243 full-length human cDNAs. Nature genetics 754 14702039
2007 Large-scale mapping of human protein-protein interactions by mass spectrometry. Molecular systems biology 733 17353931
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2012 A census of human soluble protein complexes. Cell 689 22939629
2003 HIV-1 Vif protein binds the editing enzyme APOBEC3G and induces its degradation. Nature medicine 679 14528301
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2003 HIV-1 Vif blocks the antiviral activity of APOBEC3G by impairing both its translation and intracellular stability. Molecular cell 607 14527406
2011 Global landscape of HIV-human protein complexes. Nature 593 22190034
2003 Hypermutation of HIV-1 DNA in the absence of the Vif protein. Science (New York, N.Y.) 570 12750511
2021 Multilevel proteomics reveals host perturbations by SARS-CoV-2 and SARS-CoV. Nature 532 33845483
2006 Hsp90 cochaperone Aha1 downregulation rescues misfolding of CFTR in cystic fibrosis. Cell 517 17110338
2012 Increased proteasome activity in human embryonic stem cells is regulated by PSMD11. Nature 331 22972301
1997 Large conductance voltage- and calcium-dependent K+ channel, a distinct member of voltage-dependent ion channels with seven N-terminal transmembrane segments (S0-S6), an extracellular N terminus, and an intracellular (S9-S10) C terminus. Proceedings of the National Academy of Sciences of the United States of America 248 9391153
2015 cAMP-induced phosphorylation of 26S proteasomes on Rpn6/PSMD11 enhances their activity and the degradation of misfolded proteins. Proceedings of the National Academy of Sciences of the United States of America 202 26669444
2019 High-resolution, strand-specific R-loop mapping via S9.6-based DNA-RNA immunoprecipitation and high-throughput sequencing. Nature protocols 195 31053798
1999 Genomic organization of the S locus: Identification and characterization of genes in SLG/SRK region of S(9) haplotype of Brassica campestris (syn. rapa). Genetics 195 10471721
2003 Potent estrogenic metabolites of bisphenol A and bisphenol B formed by rat liver S9 fraction: their structures and estrogenic potency. Toxicological sciences : an official journal of the Society of Toxicology 133 14691209
2013 The sub-nanomolar binding of DNA-RNA hybrids by the single-chain Fv fragment of antibody S9.6. Journal of molecular recognition : JMR 126 23784994
2017 The Affinity of the S9.6 Antibody for Double-Stranded RNAs Impacts the Accurate Mapping of R-Loops in Fission Yeast. Journal of molecular biology 111 29289567
2001 Metabolic activation of bisphenol A by rat liver S9 fraction. Toxicological sciences : an official journal of the Society of Toxicology 109 11452134
2021 Recognition of RNA by the S9.6 antibody creates pervasive artifacts when imaging RNA:DNA hybrids. The Journal of cell biology 108 33830170
2002 The conformationally flexible S9-S10 linker region in the core domain of p53 contains a novel MDM2 binding site whose mutation increases ubiquitination of p53 in vivo. The Journal of biological chemistry 103 11925449
1998 Detection and assignment of proteins encoded by rice black streaked dwarf fijivirus S7, S8, S9 and S10. The Journal of general virology 102 9634092
2019 26S Proteasomes are rapidly activated by diverse hormones and physiological states that raise cAMP and cause Rpn6 phosphorylation. Proceedings of the National Academy of Sciences of the United States of America 98 30782827
2006 Dermaseptin S9, an alpha-helical antimicrobial peptide with a hydrophobic core and cationic termini. Biochemistry 88 16401077
2022 Structural basis of R-loop recognition by the S9.6 monoclonal antibody. Nature communications 74 35347133
1976 The isolation of eukaryotic ribosomal proteins. The purification and characterization of the 40 S ribosomal subunit proteins S2, S3, S4, S5, S6, S7, S8, S9, S13, S23/S24, S27, and S28. The Journal of biological chemistry 72 947902
2010 Silencing of ribosomal protein S9 elicits a multitude of cellular responses inhibiting the growth of cancer cells subsequent to p53 activation. PloS one 71 20221446
2007 Cidofovir and (S)-9-[3-hydroxy-(2-phosphonomethoxy)propyl]adenine are highly effective inhibitors of vaccinia virus DNA polymerase when incorporated into the template strand. Antimicrobial agents and chemotherapy 68 18056278
1988 Probing the assembly of the 3' major domain of 16 S ribosomal RNA. Quaternary interactions involving ribosomal proteins S7, S9 and S19. Journal of molecular biology 67 3373531
1985 The action of anticlastogens in human lymphocyte cultures and their modification by rat-liver S9 mix. II. Studies with vitamins C and E. Mutation research 67 2579333
1975 Nucleotide sequences of Escherichia coli 16-S RNA associated with ribosomal proteins S7, S9, S10, S14 and S19. European journal of biochemistry 64 1100389
2004 Functional analysis of Rpn6p, a lid component of the 26 S proteasome, using temperature-sensitive rpn6 mutants of the yeast Saccharomyces cerevisiae. The Journal of biological chemistry 61 15611133
2004 A mutation in the nuclear-encoded plastid ribosomal protein S9 leads to early embryo lethality in maize. The Plant journal : for cell and molecular biology 60 14675435
1977 Isolation of eukaryotic ribosomal proteins. Purification and characterization of the 40 S ribosomal subunit proteins Sa, Sc, S3a, S3b, S5', S9, S10, S11, S12, S14, S15, S15', S16, S17, S18, S19, S20, S21, S26, S27', and S29. The Journal of biological chemistry 60 925037
1973 Requirement of proteins S5 and S9 from 30S subunits for the ribosome-dependent GTPase activity of elongation factor G. Proceedings of the National Academy of Sciences of the United States of America 57 4346030
2019 Comparative Proteomics Analysis of Human Liver Microsomes and S9 Fractions. Drug metabolism and disposition: the biological fate of chemicals 55 31699809
1981 Greater effectiveness of hepatocyte and liver S9 preparations from hamsters than rat preparations in activating N-nitroso compounds to metabolites mutagenic to Salmonella. Journal of the National Cancer Institute 55 6946249
2013 Utility of rat liver S9 fractions to study skin-sensitizing prohaptens in a modified KeratinoSens assay. Toxicological sciences : an official journal of the Society of Toxicology 52 23872582
2012 Assessment of metabolic stability using the rainbow trout (Oncorhynchus mykiss) liver S9 fraction. Current protocols in toxicology 52 22896006
2007 NADPH-dependent covalent binding of [3H]paroxetine to human liver microsomes and S-9 fractions: identification of an electrophilic quinone metabolite of paroxetine. Chemical research in toxicology 50 17907785
1984 Mutagenicity of derivatives and metabolites of 1-nitropyrene: activation by rat liver S9 and bacterial enzymes. Mutation research 50 6392870
2009 S9, a novel anticancer agent, exerts its anti-proliferative activity by interfering with both PI3K-Akt-mTOR signaling and microtubule cytoskeleton. PloS one 49 19293927
2017 The monoclonal S9.6 antibody exhibits highly variable binding affinities towards different R-loop sequences. PloS one 47 28594954
2008 Cloning, expression, and characterization of a new xylanase with broad temperature adaptability from Streptomyces sp. S9. Applied microbiology and biotechnology 45 18521591
2015 The utility of metabolic activation mixtures containing human hepatic post-mitochondrial supernatant (S9) for in vitro genetic toxicity assessment. Mutagenesis 44 26712374
1999 Advantage of the use of human liver S9 in the Ames test. Mutation research 42 9858674
2019 Tools for studying the metabolism of new psychoactive substances for toxicological screening purposes - A comparative study using pooled human liver S9, HepaRG cells, and zebrafish larvae. Toxicology letters 41 30682400
2022 Circular RNA circ-MTHFD1L induces HR repair to promote gemcitabine resistance via the miR-615-3p/RPN6 axis in pancreatic ductal adenocarcinoma. Journal of experimental & clinical cancer research : CR 39 35459186
2020 How to Study the Metabolism of New Psychoactive Substances for the Purpose of Toxicological Screenings-A Follow-Up Study Comparing Pooled Human Liver S9, HepaRG Cells, and Zebrafish Larvae. Frontiers in chemistry 38 32766204
1991 Effects of capsaicin on rat liver S9-mediated metabolism and DNA binding of aflatoxin. Nutrition and cancer 38 1901992
2016 Application of the TGx-28.65 transcriptomic biomarker to classify genotoxic and non-genotoxic chemicals in human TK6 cells in the presence of rat liver S9. Environmental and molecular mutagenesis 34 26946220
2022 Lactobacillus plantarum S9 alleviates lipid profile, insulin resistance, and inflammation in high-fat diet-induced metabolic syndrome rats. Scientific reports 33 36109620
2013 The flavonoid apigenin downregulates CDK1 by directly targeting ribosomal protein S9. PloS one 33 24009741
1979 Effects of the hepatic S9 fraction from aroclor-1254-treated rats on the mutagenicity of benzo[alpha]pyrene and 2-aminoanthracene in the Salmonella/microsome assay. Mutation research 33 118381
2002 The proteasome regulatory particle subunit Rpn6 is required for Drosophila development and interacts physically with signalosome subunit Alien/CSN2. Gene 30 12426099
1983 Spindle disturbances in mammalian cells. II. Induction of viable aneuploid/polyploid cells and multiple chromatid exchanges after treatment of V79 Chinese hamster cells with carbaryl. Modifying effects of glutathione and S9. Mutation research 30 6402697
2020 Long-noncoding RNA Peg13 alleviates epilepsy progression in mice via the miR-490-3p/Psmd11 axis to inactivate the Wnt/β-catenin pathway. American journal of translational research 28 33437373
2003 Use of human liver S9 in the Ames test: assay of three procarcinogens using human S9 derived from multiple donors. Regulatory toxicology and pharmacology : RTP 28 12662906
2012 In vitro studies on the oxidative metabolism of 20(s)-ginsenoside Rh2 in human, monkey, dog, rat, and mouse liver microsomes, and human liver s9. Drug metabolism and disposition: the biological fate of chemicals 27 22829543
2012 Comparison of metabolism-mediated effects of pyrrolizidine alkaloids in a HepG2/C3A cell-S9 co-incubation system and quantification of their glutathione conjugates. Xenobiotica; the fate of foreign compounds in biological systems 26 22519982
1994 Evaluation of pyrazole and ethanol induced S9 fraction in bacterial mutagenicity testing. Mutagenesis 26 8208127
2016 γH2AX and p53 responses in TK6 cells discriminate promutagens and nongenotoxicants in the presence of rat liver S9. Environmental and molecular mutagenesis 25 27364561
2013 Distinctive contributions of the ribosomal P-site elements m2G966, m5C967 and the C-terminal tail of the S9 protein in the fidelity of initiation of translation in Escherichia coli. Nucleic acids research 25 23530111
1990 Nucleotide sequence of segment S9 of the genome of rice gall dwarf virus. The Journal of general virology 24 2391501
1985 Cloning and nucleotide sequencing of the genes for ribosomal proteins S9 (rpsI) and L13 (rplM) of Escherichia coli. Molecular & general genetics : MGG 24 3884974
2020 PSMD11, PTPRM and PTPRB as novel biomarkers of pancreatic cancer progression. Biochimica et biophysica acta. General subjects 23 32663515
2018 Homoharringtonine could induce quick protein synthesis of PSMD11 through activating MEK1/ERK1/2 signaling pathway in pancreatic cancer cells. Journal of cellular biochemistry 23 29665121
2009 Two-hybrid analysis identifies PSMD11, a non-ATPase subunit of the proteasome, as a novel interaction partner of AMP-activated protein kinase. The international journal of biochemistry & cell biology 23 19616115
1993 Influence of S9 mix composition on the SOS response in Escherichia coli PQ37 by polycyclic aromatic hydrocarbons. Mutation research 22 7678915
1987 The effectiveness of S9 and microsomal mix on activation of cyclophosphamide to induce genotoxicity in human lymphocytes. Mutation research 22 3821768
1981 Characterization of the binding of rat liver ribosomal proteins L6, L8, L19, S9, and S13 to 5.8 S ribosomal ribonucleic acid. The Journal of biological chemistry 22 7251593
2015 PUB22 and PUB23 U-BOX E3 ligases directly ubiquitinate RPN6, a 26S proteasome lid subunit, for subsequent degradation in Arabidopsis thaliana. Biochemical and biophysical research communications 21 26188517
2013 Inhibition of pathogenic bacterial biofilm by biosurfactant produced by Lysinibacillus fusiformis S9. Bioprocess and biosystems engineering 21 23719930
1981 Mutagenicity and rat-liver S9 demethylation kinetics of N-nitrosomethylaniline and its ring-substituted derivatives. Mutation research 21 7027031
2016 Identification of allocryptopine and protopine metabolites in rat liver S9 by high-performance liquid chromatography/quadrupole-time-of-flight mass spectrometry. Rapid communications in mass spectrometry : RCM 20 27321842
2012 Elucidation of motifs in ribosomal protein S9 that mediate its nucleolar localization and binding to NPM1/nucleophosmin. PloS one 20 23285058
1983 Activation of nitrosamines and other carcinogens by mouse-liver S9, mouse hepatocytes and in the host-mediated assay produces different mutagenic responses in Salmonella TA1535. Mutation research 20 6865998
2019 MicroRNA-451 inhibits inflammation and proliferation of glomerular mesangial cells through down-regulating PSMD11 and NF-κB p65. Bioscience reports 19 31652441
1999 Cytotoxicity of dental resin monomers in the presence of S9 mix enzymes. Dental materials journal 19 10786150
1988 Studies of an S9-based metabolic activation system used in the mouse lymphoma L5178Y cell mutation assay. Mutagenesis 19 3070287
1987 Mutagenicity of nitracrine analogues in Salmonella typhimurium: mutational specificity and activation by bacterial enzymes and rat-liver S9. Mutation research 19 3540655
2021 Nitric oxide donor sodium nitroprusside-induced transcriptional changes and hypocrellin biosynthesis of Shiraia sp. S9. Microbial cell factories 18 33910564
2021 Mapping R-Loops and RNA:DNA Hybrids with S9.6-Based Immunoprecipitation Methods. Journal of visualized experiments : JoVE 18 34515688
2015 Formation of GSH-trapped reactive metabolites in human liver microsomes, S9 fraction, HepaRG-cells, and human hepatocytes. Journal of pharmaceutical and biomedical analysis 18 26263063
1986 Multiple crosslinks of proteins S7, S9, S13 to domains 3 and 4 of 16S RNA in the 30S particle. Nucleic acids research 18 2423965
2007 Genome-wide polysomal analysis of a yeast strain with mutated ribosomal protein S9. BMC genomics 17 17711575
1984 The action of anticlastogens in human lymphocyte cultures and its modification by rat liver S9 mix. I. Studies with AET and sodium fluoride. Mutation research 17 6209573
1977 Sequence determination of protein S9 from the Escherichia coli ribosome. Hoppe-Seyler's Zeitschrift fur physiologische Chemie 17 336507
2021 Effect of chain length and branching on the in vitro metabolism of a series of parabens in human liver S9, human skin S9, and human plasma. Regulatory toxicology and pharmacology : RTP 16 33741472
2020 New incorporation of the S9 metabolizing system into methods for detecting acetylcholinesterase inhibition. Analytica chimica acta 15 32891393
2019 In vitro biotransformation of pharmaceuticals and pesticides by trout liver S9 in the presence and absence of carbamazepine. Ecotoxicology and environmental safety 15 31421535
2013 Role of the ribosomal P-site elements of m²G966, m⁵C967, and the S9 C-terminal tail in maintenance of the reading frame during translational elongation in Escherichia coli. Journal of bacteriology 15 23729652
1992 Mutagenicity of analgesics, their derivatives, and anti-inflammatory drugs with S-9 mix of several animal species. The Journal of Nihon University School of Dentistry 15 1287147
1987 Comparative activation of 3,3'-dichlorobenzidine and related benzidines to mutagens in the Salmonella typhimurium assay by hepatic S9 and microsomes from rats pretreated with different inducers of cytochrome P-450. Mutation research 15 3309638
2022 The SLC26A9 inhibitor S9-A13 provides no evidence for a role of SLC26A9 in airway chloride secretion but suggests a contribution to regulation of ASL pH and gastric proton secretion. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 14 36183361
2021 Hepatic in vitro metabolism of peptides; Comparison of human liver S9, hepatocytes and Upcyte hepatocytes with cyclosporine A, leuprorelin, desmopressin and cetrorelix as model compounds. Journal of pharmaceutical and biomedical analysis 14 33548873
2020 Using a high-throughput method in the micronucleus assay to compare animal-free with rat-derived S9. The Science of the total environment 14 33182016
2018 Process desired functional attributes of an endoxylanase of GH10 family from a new strain of Aspergillus terreus S9. International journal of biological macromolecules 14 29684454
1991 S9-mediated metabolism of 1-nitropyrene to a mutagen in Chinese hamster ovary cells by ring-oxidation under aerobic conditions and by nitroreduction under anaerobic conditions. Carcinogenesis 14 1747934
1987 An investigation of the metabolism of N-nitroso-N-methylaniline by phenobarbital- and pyrazole-induced Sprague-Dawley rat liver and esophagus derived S-9. Chemico-biological interactions 14 3568192
2022 Target Fishing Reveals a Novel Mechanism of 1,2,4-Oxadiazole Derivatives Targeting Rpn6, a Subunit of 26S Proteasome. Journal of medicinal chemistry 13 35253427
2019 Comparison of the metabolism of 10 cosmetics-relevant chemicals in EpiSkin™ S9 subcellular fractions and in vitro human skin explants. Journal of applied toxicology : JAT 13 31701564
1991 Arylamine activation following chronic ethanol ingestion by rats: studies on the liver S9, microsomal and cytosolic fractions and comparison with Aroclor 1254 pretreatment. Mutation research 13 2002800
1989 Effect of tannic acid on rat liver S9 mediated mutagenesis, metabolism and DNA binding of benzo[a]pyrene. Cancer letters 13 2517593
1987 DNA repair synthesis in cultured fish and human cells exposed to fish S9-activated aromatic hydrocarbons. Comparative biochemistry and physiology. C, Comparative pharmacology and toxicology 13 2882940
1977 Further evidence that the ribosomal 30S proteins S3, S5, S9, S11, S12, and S18 possess specific 16S RNA binding sites. Molecular & general genetics : MGG 13 331074
2024 Metabolic activation of short-chain alkyl N-nitrosamines using Aroclor 1254 or phenobarbital/beta-naphthoflavone-induced rat or hamster S9 - A comparative analysis. Toxicology reports 12 38322170
2016 Cloning, expression, and biochemical characterization of a cold-active GDSL-esterase of a Pseudomonas sp. S9 isolated from Spitsbergen island soil. Acta biochimica Polonica 12 26824293
2013 Isolation and characterization of multiple F-box genes linked to the S9- and S10-RNase in apple (Malus × domestica Borkh.). Plant reproduction 12 23686223
2001 Chromosome analysis of human spermatozoa following in vitro exposure to cyclophosphamide, benzo(a)pyrene and N-nitrosodimethylamine in the presence of rat liver S9. Mutation research 12 11287298