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Showing PSMD6RPN7 is a alias.

PSMD6

26S proteasome non-ATPase regulatory subunit 6 · UniProt Q15008

Length
389 aa
Mass
45.5 kDa
Annotated
2026-06-10
48 papers in source corpus 14 papers cited in narrative 14 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PSMD6 (Rpn7) is a PCI-domain subunit of the lid of the 19S regulatory particle of the 26S proteasome, where it is required for correct lid assembly and thereby for ubiquitin-dependent protein degradation (PMID:15102831, PMID:17135287). Studies in yeast established that Rpn7 is specifically necessary for lid formation—not base or core particle assembly—and that its loss leaves an incomplete lid and stabilizes ubiquitin-proteasome substrates including cell-cycle regulators (PMID:15102831, PMID:17135287). Within an ordered, multi-step lid assembly pathway, Rpn7 incorporates as part of lid module 2 together with Rpn3 and Sem1, after a core module forms and before Rpn12 addition (PMID:20471955, PMID:26182356). Sem1/DSS1 acts as a molecular tether that grasps both Rpn3 and Rpn7 through acidic segments to enforce ordered formation and stabilization of the Rpn3-Sem1-Rpn7 ternary complex; its C-terminal helix engages the PCI domain of Rpn7, a domain whose conserved leucine also mediates high-affinity binding to Rpn9 (PMID:17761670, PMID:23643786, PMID:24412063). Beyond assembly, PSMD6 undergoes DNA-damage-induced phosphorylation and nuclear translocation, colocalizes with and stabilizes DNA damage foci, and supports DNA repair and senescence following genotoxic insult (PMID:22473755, PMID:24987841). Loss of psmd6 in zebrafish causes accumulation of polyubiquitylated proteins and APC/C substrate-dependent defects in lens fiber differentiation as well as defects in cardiac morphogenesis, linking proteasome function to organ development (PMID:20724448, PMID:40857331).

Mechanistic history

Synthesis pass · year-by-year structured walk · 10 steps
  1. 2004 High

    Established that Rpn7 is required for structural integrity of the 26S proteasome, answering whether this lid subunit is essential for degradative function.

    Evidence Temperature-sensitive rpn7-3 yeast mutant with 26S affinity purification and ubiquitin-substrate stabilization assays

    PMID:15102831

    Open questions at the time
    • Did not localize Rpn7 within the lid architecture
    • Did not distinguish lid-specific versus general assembly defects
  2. 2006 High

    Resolved that Rpn7 is specifically required for lid assembly rather than base or core particle formation, placing it precisely in proteasome biogenesis.

    Evidence Biochemical fractionation and fluorescence localization of proteasome modules in yeast lid and base mutants

    PMID:17135287

    Open questions at the time
    • Order of subunit addition within the lid not yet defined
    • Molecular partners of Rpn7 in the lid not identified
  3. 2007 Medium

    Identified the PCI domain and a conserved leucine as the structural determinant of Rpn7 function and its high-affinity binding to Rpn9, and showed DSS1/Sem1 connects Rpn7 to Rpn3 and BRCA2.

    Evidence PCI-domain site-directed mutagenesis with binding assays in S. pombe; reciprocal Co-IP and domain mapping in human tumor cells

    PMID:17563742 PMID:17761670

    Open questions at the time
    • BRCA2/RPN7 interaction functional significance unresolved
    • Single-lab Co-IP without structural validation
  4. 2010 High

    Defined the ordered, multi-step lid assembly pathway and placed Rpn3-Rpn7-Sem1 as a discrete module added after a core module, and linked Psmd6 loss in vivo to APC/C substrate degradation during lens development.

    Evidence Mass spectrometry of subassemblies in yeast lid mutants; zebrafish volvox/psmd6 forward-genetic mutant with polyubiquitylation and APC/C epistasis

    PMID:20471955 PMID:20724448

    Open questions at the time
    • Mechanism tethering Rpn3-Rpn7 within the module not yet defined
    • Tissue-specificity of developmental requirement unexplained
  5. 2012 Medium

    Revealed a non-assembly role: DNA-damage-induced nuclear translocation of Rpn7 stabilizes long-lived DNA damage foci to enable senescence.

    Evidence siRNA knockdown, immunofluorescence colocalization, and foci lifespan quantification

    PMID:22473755

    Open questions at the time
    • Molecular basis of foci stabilization unknown
    • Single-lab observation
  6. 2013 High

    Structurally localized the Sem1-Rpn7 interaction, showing Sem1 acts as molecular glue stabilizing the Rpn3/Rpn7 heterodimer via the PCI domain of Rpn7.

    Evidence Cryo-EM single-particle reconstruction of sem1-deletion yeast proteasomes with site-specific cross-linking

    PMID:23643786

    Open questions at the time
    • Dynamics of the interaction during assembly not captured
    • Human-complex structure not determined
  7. 2014 High

    Demonstrated mechanistically that Sem1 enforces ordered Rpn3/Rpn7 incorporation by grasping both subunits, with tethering essential for ternary complex biogenesis but dispensable once incorporated, and that PSMD6 phosphorylation/nuclear import in DNA repair is uPAR-dependent.

    Evidence In vitro reconstitution with engineered TEV-cleavable Sem1 and domain mutagenesis; uPAR knockdown with phosphorylation, localization, and survival assays in vascular smooth muscle cells

    PMID:24412063 PMID:24987841

    Open questions at the time
    • Kinase responsible for PSMD6 phosphorylation not identified
    • Mechanism of uPAR-to-proteasome signaling not defined
  8. 2015 Medium

    Placed Rpn7 in lid module 2 and showed Rpn11 bridges modules 1 and 2 to recruit Rpn7-containing subunits, refining the assembly hierarchy.

    Evidence RPN11 suppression, mutant proteasome isolation, and stepwise in vitro module reconstitution

    PMID:26182356

    Open questions at the time
    • Precise contacts between Rpn11 and module 2 not resolved
    • Single-lab reconstitution
  9. 2023 Medium

    Defined the distinct subcellular distribution of PSMD6-containing 19S RP versus uncapped 20S core particles, including stress-induced nuclear proteasomal granules.

    Evidence CRISPR endogenous mScarlet/YFP tagging of PSMD6 and PSMB6, live-cell colocalization, osmotic stress, and PSMD1 knockdown

    PMID:37371572

    Open questions at the time
    • Functional role of nuclear granules unknown
    • Cause of PSMD6 cytoplasmic redistribution upon PSMD1 loss not mechanistically explained
  10. 2025 Medium

    Extended the in vivo developmental requirement of psmd6 to cardiac morphogenesis, linking proteasome function to a congenital heart disease subnetwork.

    Evidence CRISPR loss-of-function zebrafish with cardiac morphology/function phenotyping and PPI network analysis

    PMID:40857331

    Open questions at the time
    • Cardiac-specific substrates of the proteasome not identified
    • Direct disease causation in humans not established

Open questions

Synthesis pass · forward-looking unresolved questions
  • How PSMD6's assembly role is coupled to its DNA-damage signaling and tissue-specific developmental functions remains unresolved.
  • Kinase and signaling pathway driving PSMD6 phosphorylation unidentified
  • Substrates underlying lens and cardiac developmental defects unknown
  • Human structural model of the Rpn3-Sem1-Rpn7 module not determined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 3
Localization
GO:0005634 nucleus 4 GO:0005829 cytosol 1
Pathway
R-HSA-392499 Metabolism of proteins 3 R-HSA-1266738 Developmental Biology 2 R-HSA-1640170 Cell Cycle 2 R-HSA-73894 DNA Repair 2
Complex memberships
26S proteasome 19S regulatory particle lidRpn3-Sem1-Rpn7 ternary complex

Evidence

Reading pass · 14 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2004 Rpn7 (PSMD6 ortholog in yeast) is required for the structural integrity of the 26S proteasome; temperature-sensitive rpn7-3 mutant cells accumulate poly-ubiquitinated proteins and stabilize ubiquitin-proteasome substrates (N-end rule, UFD pathway substrates, cell cycle regulators Pds1 and Clb2). Analysis showed the lid subcomplex in rpn7-3 cells contained only 5 of 8 lid components, indicating Rpn7 is required for correct lid structure and 26S holoenzyme assembly. Temperature-sensitive yeast mutant analysis, affinity purification of 26S proteasome, substrate stabilization assays (beta-galactosidase reporters), immunoblotting The Journal of biological chemistry High 15102831
2006 The 26S proteasome modules (core particle, base, and lid) can be formed and imported into the yeast nucleus independently of each other. In rpn7-3 mutant cells at restrictive temperature, lid formation was specifically disrupted while an intact base was produced and localized to the nucleus, establishing that Rpn7 is specifically required for lid assembly and not for base or core particle formation. Biochemical characterization and microscopic localization of proteasome in yeast lid mutants (rpn5-1, rpn7-3) and base mutant (ΔN rpn2); subcellular fractionation and fluorescence microscopy Molecular biology of the cell High 17135287
2007 Human DSS1 (Sem1) interacts with the proteasome subunits RPN3 and RPN7 (PSMD6) in human tumor cells; BRCA2 also interacts with RPN3 and RPN7; the BRCA2/RPN7 interaction is independent of DSS1. Defined regions of DSS1 are important for interactions with RPN3, RPN7, and BRCA2. Co-immunoprecipitation, domain mapping/deletion analysis in human tumor cell lines Oncogene Medium 17563742
2007 A conserved leucine residue in the PCI domain of Rpn7 (PSMD6 ortholog in fission yeast) is critical for protein function; replacing it with aspartate inactivated Rpn7 and abolished high-affinity binding to Rpn9. Rpn7 and Rpn5 bind Rpn9 with high affinity via their PCI domains, and this interaction is disrupted by the leucine-to-aspartate mutation. High-copy suppressor screen, site-directed mutagenesis of PCI domain leucine residue, protein-binding assays in S. pombe The Journal of biological chemistry Medium 17761670
2010 Dissection of lid assembly using yeast rpn7-3, Δrpn9, and rpn12-1 mutants identified lid subassemblies including an Rpn3-Rpn7 pair and a lid-like complex lacking Rpn12 by mass spectrometry, demonstrating that the lid assembles in an ordered, multi-step process: first a core module (Rpn5, 6, 8, 9, 11) forms, then a second module consisting of Rpn3, Rpn7 (PSMD6 ortholog), and Sem1 is attached, followed by Rpn12 incorporation. Yeast lid mutant analysis, affinity purification, mass spectrometry identification of subassemblies Biochemical and biophysical research communications High 20471955
2010 Psmd6 (zebrafish ortholog) is required for cell proliferation of lens epithelium and for proper elongation and differentiation of lens fiber cells. Loss of psmd6 in the volvox mutant causes accumulation of polyubiquitylated proteins, and the anaphase-promoting complex/cyclosome (APC/C) E3 ubiquitin ligase is involved in the lens defects, placing Psmd6 in the ubiquitin-proteasome pathway upstream of APC/C substrate degradation during lens development. Zebrafish volvox mutant characterization, genetic mapping to psmd6, polyubiquitylation assays, epistasis with APC/C Development (Cambridge, England) High 20724448
2012 The 19S subunit Rpn7 (PSMD6) undergoes increased protein levels and nuclear translocation in response to DNA damage. Rpn7 colocalizes with DNA damage foci throughout their lifespan. Silencing of Rpn7 promotes faster resolution of DNA damage foci and specifically decreases long-lived foci frequencies without affecting repair of short-lived foci, indicating Rpn7 stabilizes DNA damage foci and enables senescence following genotoxic insult. siRNA knockdown, immunofluorescence colocalization with DNA damage foci markers, foci lifespan analysis, subcellular fractionation (nuclear translocation) IUBMB life Medium 22473755
2013 The C-terminal helix of Sem1 (DSS1, the mammalian ortholog partner) binds to the PCI domain of Rpn7 (PSMD6 ortholog) within the 26S proteasome. Cryo-EM single-particle reconstruction of proteasomes from sem1-deletion yeast localized this interaction, and site-specific cross-linking data indicated the N-terminal region of Sem1 bridges the cleft between Rpn7 and Rpn3, with Sem1 acting as molecular glue stabilizing the Rpn3/Rpn7 heterodimer. Cryo-EM single particle reconstruction, sem1 deletion yeast strains, site-specific cross-linking Biochemical and biophysical research communications High 23643786
2014 Sem1 (DSS1 in mammals) enforces ordered incorporation of subunits Rpn3 and Rpn7 (PSMD6 ortholog) into the assembling proteasome lid by grasping both using conserved acidic segments separated by a flexible linker. TEV protease cleavage of Sem1 showed that its tethering function is important for biogenesis and integrity of the Rpn3-Sem1-Rpn7 ternary complex but becomes dispensable once the ternary complex is incorporated into larger lid precursors. In vitro reconstitution, TEV protease-cleavage site engineering into Sem1, biochemical assembly assays, domain mutagenesis Molecular cell High 24412063
2014 PSMD6 undergoes DNA damage-induced phosphorylation, nuclear import, and recruitment to proteasome at DNA damage sites; these events are mediated by urokinase-type plasminogen activator receptor (uPAR) signaling in vascular smooth muscle cells. uPAR is essential for functional assembly of the 26S proteasome, and deficiency of uPAR or PSMD6 delays DNA repair and reduces cell survival. uPAR knockdown, phosphorylation assays, subcellular fractionation (nuclear import), colocalization/immunofluorescence, cell survival assays in vascular smooth muscle cells PloS one Medium 24987841
2015 Rpn11 plays a key role in bridging lid module 1 and module 2 subunits; module 2 includes Rpn3, Rpn7 (PSMD6 ortholog), Rpn12, and Rpn15. Suppression of RPN11 halted lid assembly but allowed base and 20S CP to pre-assemble. Re-introducing the C-terminal portion of Rpn11 enabled recruitment of missing module 2 subunits including Rpn7, and module 1 was reconstituted in vitro stepwise. RPN11 expression suppression, mutant proteasome isolation, in vitro module reconstitution, protein interaction assays Bioscience reports Medium 26182356
2021 AT1R colocalizes with PSMD6 (used as a proteasomal marker) in aortic cells, and this colocalization is reduced after SNX1 knockdown. Proteasomal inhibition (not lysosomal) increases AT1R protein content and this is accompanied by decreased ubiquitin/AT1R binding after SNX1 knockdown, suggesting PSMD6-containing 26S proteasome mediates SNX1-directed degradation of AT1R. Confocal colocalization microscopy (PSMD6 and AT1R), proteasomal vs. lysosomal inhibitor treatment, cycloheximide chase assay, SNX1 siRNA knockdown in A10 cells Hypertension research Low 33972750
2023 PSMD6 (Rpn7) tagged with mScarlet at the endogenous locus colocalizes with YFP-tagged PSMB6 (20S CP subunit) in cells, predominantly with nuclear accumulation. Under osmotic stress, nuclear proteasomal granules are positive for both tags. Upon PSMD1 knockdown (which increases free 20S CPs), 20S-YFP remained nuclear while PSMD6-mScarlet redistributed to cytoplasm, demonstrating distinct subcellular distribution of 19S RP (containing PSMD6) versus uncapped 20S CPs. CRISPR endogenous YFP/mScarlet tagging of PSMB6 and PSMD6, live-cell fluorescence colocalization microscopy, osmotic stress treatment, PSMD1 siRNA knockdown Biomolecules Medium 37371572
2025 CRISPR-based loss of psmd6 in zebrafish leads to defects in heart development including dysmorphic hearts, myocardial cell blebbing, and reduced outflow tracts, as well as deficits in cardiac function. PPI network analysis placed PSMD6 in a subnetwork with known CHD genes and other proteasome factors (POMP, PSMA6, PSMA7, PSMD3), indicating a role for proteasome function in cardiac morphogenesis. CRISPR targeting and mutant zebrafish strain generation for psmd6, cardiac morphology and function assessment, PPI network analysis PLoS genetics Medium 40857331

Source papers

Stage 0 corpus · 48 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2011 Meta-analysis of genome-wide association studies identifies eight new loci for type 2 diabetes in east Asians. Nature genetics 495 22158537
2006 The assembly pathway of the 19S regulatory particle of the yeast 26S proteasome. Molecular biology of the cell 94 17135287
2006 Array-comparative genomic hybridization to detect genomewide changes in microdissected primary and metastatic oral squamous cell carcinomas. Molecular carcinogenesis 73 16676365
2011 Gene networks associated with conditional fear in mice identified using a systems genetics approach. BMC systems biology 68 21410935
2014 The intrinsically disordered Sem1 protein functions as a molecular tether during proteasome lid biogenesis. Molecular cell 63 24412063
2010 Dissection of the assembly pathway of the proteasome lid in Saccharomyces cerevisiae. Biochemical and biophysical research communications 50 20471955
2007 The proteasome is involved in determining differential utilization of double-strand break repair pathways. Oncogene 47 17563742
2010 The ubiquitin proteasome system is required for cell proliferation of the lens epithelium and for differentiation of lens fiber cells in zebrafish. Development (Cambridge, England) 45 20724448
2004 Rpn7 Is required for the structural integrity of the 26 S proteasome of Saccharomyces cerevisiae. The Journal of biological chemistry 45 15102831
2015 Interplay between promoter methylation and chromosomal loss in gene silencing at 3p11-p14 in cervical cancer. Epigenetics 43 26291246
2013 Identification of eight candidate target genes of the recurrent 3p12-p14 loss in cervical cancer by integrative genomic profiling. The Journal of pathology 40 23335387
2013 Replication study for the association of 9 East Asian GWAS-derived loci with susceptibility to type 2 diabetes in a Japanese population. PloS one 39 24086726
2015 Increased New lncRNA-mRNA Gene Pair Levels in Human Cumulus Cells Correlate With Oocyte Maturation and Embryo Development. Reproductive sciences (Thousand Oaks, Calif.) 35 25670720
2006 Functional study of hot pepper 26S proteasome subunit RPN7 induced by Tobacco mosaic virus from nuclear proteome analysis. Biochemical and biophysical research communications 34 17070775
2005 A combination of molecular cytogenetic analyses reveals complex genetic alterations in conventional renal cell carcinoma. Cancer genetics and cytogenetics 34 15860350
2018 Protein interactions of FAM134B with EB1 and APC/beta-catenin in vitro in colon carcinoma. Molecular carcinogenesis 30 29964340
2011 Effects of sex steroids on indices of protein turnover in rainbow trout (Oncorhynchusmykiss) white muscle. General and comparative endocrinology 29 21878334
2013 Localization of the regulatory particle subunit Sem1 in the 26S proteasome. Biochemical and biophysical research communications 27 23643786
2021 The mustard leaf beetle, Phaedon cochleariae, as a screening model for exogenous RNAi-based control of coleopteran pests. Pesticide biochemistry and physiology 26 34119215
2000 The p44S10 locus, encoding a subunit of the proteasome regulatory particle, is amplified during progression of cutaneous malignant melanoma. Oncogene 26 10723133
2021 The Silence of PSMC6 Inhibits Cell Growth and Metastasis in Lung Adenocarcinoma. BioMed research international 24 34239933
2019 Molecular mechanisms linking peri-implantitis and type 2 diabetes mellitus revealed by transcriptomic analysis. PeerJ 24 31275749
2020 Construction of relapse-related lncRNA-mediated ceRNA networks in Hodgkin lymphoma. Archives of medical science : AMS 23 33224341
2014 Loss of urokinase receptor sensitizes cells to DNA damage and delays DNA repair. PloS one 23 24987841
2014 A functional genomic screen for evolutionarily conserved genes required for lifespan and immunity in germline-deficient C. elegans. PloS one 21 25093668
2015 Base-CP proteasome can serve as a platform for stepwise lid formation. Bioscience reports 19 26182356
2021 Increased AT1 receptor expression mediates vasoconstriction leading to hypertension in Snx1-/- mice. Hypertension research : official journal of the Japanese Society of Hypertension 16 33972750
2012 The 19S proteasome subunit Rpn7 stabilizes DNA damage foci upon genotoxic insult. IUBMB life 16 22473755
2007 Isolation of the Schizosaccharomyces pombe proteasome subunit Rpn7 and a structure-function study of the proteasome-COP9-initiation factor domain. The Journal of biological chemistry 13 17761670
2009 Karyotype variability in KP1(+) and KP1(-) strains of Trypanosoma rangeli isolated in Brazil and Colombia. Acta tropica 11 19283897
2021 Housekeeping Genes for Parkinson's Disease in Humans and Mice. Cells 10 34571901
2024 Comparison of differentially expressed genes in longissimus dorsi muscle of Diannan small ears, Wujin and landrace pigs using RNA-seq. Frontiers in veterinary science 9 38249550
2022 Silencing suppressors of rice black-streaked dwarf virus and rice stripe virus hijack the 26S proteasome of Laodelphax striatellus to facilitate virus accumulation and transmission. Pest management science 8 35439336
2022 Proteasomal subunit depletions differentially affect germline integrity in C. elegans. Frontiers in cell and developmental biology 8 36060813
2023 Exploration of anti-stress mechanisms in high temperature exposed juvenile golden cuttlefish (Sepia esculenta) based on transcriptome profiling. Frontiers in physiology 7 37234426
2024 Bioinformatics to analyze the differentially expressed genes in different degrees of Alzheimer's disease and their roles in progress of the disease. Journal of applied genetics 6 38315405
2022 Risk Stratification and Validation of Eleven Autophagy-Related lncRNAs for Esophageal Squamous Cell Carcinoma. Frontiers in genetics 6 35832188
2022 Comparative Ubiquitome Analysis Reveals Deubiquitinating Effects Induced by Wolbachia Infection in Drosophila melanogaster. International journal of molecular sciences 6 36012723
2019 Umbilical cord blood-based gene signatures related to prenatal major depressive disorder. Medicine 5 31305436
2025 Overexpression pattern, function, and clinical value of proteasome 26S subunit non-ATPase 6 in hepatocellular carcinoma. World journal of clinical oncology 2 39995557
2024 Overexpression of proteasome 26S subunit non-ATPase 6 protein and its clinicopathological significance in intrahepatic cholangiocarcinoma. World journal of hepatology 2 39606161
2024 Identification of PSMD2 as a promising biomarker for pancreatic cancer patients based on comprehensive bioinformatics and in vitro studies. Heliyon 2 39634424
2023 Method of Monitoring 26S Proteasome in Cells Revealed the Crucial Role of PSMA3 C-Terminus in 26S Integrity. Biomolecules 2 37371572
2022 Genetic analysis of Caenorhabditis elegans pry-1/Axin suppressors identifies genes involved in reproductive structure development, stress responses, and aging. G3 (Bethesda, Md.) 2 35100345
2025 A systems genetics approach identifies roles for proteasome factors in heart development and congenital heart defects. PLoS genetics 1 40857331
2024 Is 26S proteasome non-ATPase regulatory subunit 6 a potential molecular target for intrahepatic cholangiocarcinoma? World journal of hepatology 1 39606166
2026 Targeting muscle wasting: Bioinformatics-derived ubiquitination genes as potential therapeutic targets for sarcopenia. Medicine 0 41560007
2026 Mapping Genetic Modifiers of Polyp Formation in Smad4-Deficient Juvenile Polyposis Using the Collaborative Cross Mouse Population. Cells 0 42193864

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