Affinage

PSMD4

26S proteasome non-ATPase regulatory subunit 4 · UniProt P55036

Round 2 corrected
Length
377 aa
Mass
40.7 kDa
Annotated
2026-04-28
130 papers in source corpus 38 papers cited in narrative 38 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PSMD4 (S5a/Rpn10) is the principal polyubiquitin-chain receptor of the 26S proteasome 19S regulatory particle, recognizing K48- and K63-linked polyubiquitin chains through tandem ubiquitin-interacting motifs (UIMs) and delivering ubiquitinated substrates—including p53 and cell-cycle regulators—for proteasomal degradation (PMID:8530351, PMID:15826667, PMID:24121268). Its VWA domain serves as a secondary, ubiquitin-independent receptor for UbL-domain proteins (hHR23A/B, ubiquilin/hPLIC, FAT10, NUB1L), and Rpn10 cooperates redundantly with Rpn13 and UBL-UBA shuttle factors (Rad23, Dsk2) to maintain ubiquitin-dependent proteostasis (PMID:10488153, PMID:22434192, PMID:26222436). Monoubiquitination of Rpn10 at a conserved lysine (written by Rsp5/NEDD4, erased by Ubp2) promotes its dissociation from the proteasome and reduces UIM-mediated substrate and shuttle-factor binding, dynamically switching proteasome receptor composition (PMID:20542005, PMID:27698474, PMID:26450923). Beyond canonical proteolysis, PSMD4 localizes to neuronal centrosomes to support dendrite elaboration, participates in sperm–zona pellucida penetration, and in plants functions as a dual ATG8/ubiquitin receptor directing selective autophagy of inactive proteasomes (proteaphagy) (PMID:23831032, PMID:20526895, PMID:26004230).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 1995 High

    Identified PSMD4 (S5a) as the polyubiquitin-conjugate recognition subunit of the 26S proteasome, answering the long-standing question of how the proteasome selects ubiquitinated substrates.

    Evidence Cell-free ubiquitin-dependent proteolysis inhibition in reticulocyte lysate and Xenopus egg extracts with recombinant S5a/MBP1

    PMID:8530351

    Open questions at the time
    • Number and identity of ubiquitin-binding sites within S5a unknown
    • Whether S5a is the sole ubiquitin receptor of the proteasome unresolved
  2. 1999 High

    Established that Rpn10/S5a and UBL-UBA shuttle factors (Rad23/hHR23) function as parallel, partially redundant pathways for delivering ubiquitinated substrates to the proteasome, resolving why Rpn10 deletion alone is non-lethal in yeast.

    Evidence Yeast double-deletion epistasis (rad23Δ rpn10Δ) causing synthetic growth defects and substrate stabilization; biochemical hHR23B–S5a interaction via UbL domain

    PMID:10471701 PMID:10488153

    Open questions at the time
    • Relative contribution of each pathway to specific substrate classes unclear
    • Whether additional shuttle factors cooperate with Rpn10 unknown
  3. 2002 High

    Defined the structural basis for UbL–UIM recognition at S5a, showing that UbL domains of hPLIC-2 and hHR23a mimic ubiquitin's S5a-contact surface, unifying substrate and shuttle-factor recruitment mechanisms.

    Evidence NMR structure of UbL domains and chemical shift perturbation mapping of S5a-binding surfaces

    PMID:11827521 PMID:12832454

    Open questions at the time
    • How S5a discriminates between ubiquitin and UbL in vivo not resolved
    • Full-length S5a structure within the proteasome lacking
  4. 2003 High

    Demonstrated that Rpn10 loss is lethal in Drosophila and required for 26S proteasome integrity, revealing that in higher eukaryotes Rpn10 has essential, non-redundant functions beyond yeast.

    Evidence Drosophila p54/Rpn10 null showing larval-pupal lethality, accumulation of ubiquitin conjugates, and defective proteasome particles

    PMID:12584246

    Open questions at the time
    • Whether lethality reflects substrate-recognition or assembly defects not distinguished
    • Mammalian in vivo null phenotype not yet reported
  5. 2005 High

    Solved the crystal structure of S5a UIMs bound to ubiquitin, revealing that the two UIMs employ distinct binding modes and prefer longer polyubiquitin chains, providing a structural framework for polyubiquitin selectivity at the proteasome.

    Evidence X-ray crystallography and ITC of S5a(196–306) in complex with monoubiquitin and polyubiquitin

    PMID:15826667

    Open questions at the time
    • How UIM-chain length preference translates to substrate selectivity in cells unclear
    • Structure of full-length S5a in the intact proteasome not determined
  6. 2010 High

    Discovered that monoubiquitination of Rpn10 at the UIM region (written by Rsp5, erased by Ubp2) inhibits ubiquitin-receptor function and is stress-regulated, establishing a reversible post-translational switch controlling proteasome substrate recognition capacity.

    Evidence In vivo ubiquitination assays, mass spectrometry, genetic deletion of Rsp5/Ubp2, substrate-binding assays under stress in yeast

    PMID:20542005

    Open questions at the time
    • Whether the Rsp5/Ubp2 writer-eraser pair operates on mammalian PSMD4 unknown
    • Downstream physiological consequences of monoubiquitination toggle not fully explored
  7. 2012 High

    Identified the VWA domain of PSMD4 as a ubiquitin-independent receptor for FAT10 and NUB1L, demonstrating that S5a has two functionally distinct receptor domains—UIMs for ubiquitin chains and VWA for ubiquitin-like modifiers.

    Evidence Co-IP, yeast complementation with VWA domain, RNAi knockdown in human cells showing FAT10-conjugate accumulation

    PMID:22434192

    Open questions at the time
    • Structural basis of VWA–FAT10 recognition not determined
    • Physiological importance of FAT10 disposal via PSMD4 in vivo unclear
  8. 2013 High

    Showed that PSMD4 localizes to neuronal centrosomes and is required for centrosomal proteasome activity and dendrite elaboration, extending its function beyond generic proteolysis to a spatially regulated developmental role.

    Evidence Immunofluorescence and subcellular fractionation in neurons; in vivo RNAi in rodent brain via in utero electroporation

    PMID:23831032

    Open questions at the time
    • Identity of critical centrosomal substrates degraded via PSMD4 unknown
    • Whether centrosomal proteasome function generalizes to non-neuronal contexts untested
  9. 2015 High

    Conditional mammalian double knockout of Rpn10 and Rpn13 proved that these two subunits are the principal docking receptors for UBL-UBA shuttle proteins in vivo and act redundantly to maintain ubiquitin conjugate homeostasis.

    Evidence Liver-specific Cre-lox double knockout in mice with massive ubiquitin-conjugate accumulation and loss of shuttle-factor docking

    PMID:26222436

    Open questions at the time
    • Tissue-specific differences in Rpn10/Rpn13 dependence unexplored
    • Whether other tissues tolerate single-subunit loss differently unknown
  10. 2015 High

    Revealed that Arabidopsis RPN10 acts as a dual ATG8/ubiquitin receptor for selective autophagy of inhibited proteasomes (proteaphagy), providing the first evidence of a proteasome subunit serving as its own autophagy receptor.

    Evidence GFP-proteasome imaging, genetic epistasis with ATG mutants, Co-IP of RPN10 with ATG8 and ubiquitin in Arabidopsis

    PMID:26004230

    Open questions at the time
    • Whether mammalian PSMD4 performs a proteaphagy receptor function not demonstrated
    • Structural basis of simultaneous ATG8 and ubiquitin binding by RPN10 not solved
  11. 2016 High

    Crystal structure of ubiquitylated Rpn10 showed that the conjugated ubiquitin clashes sterically with Rpn9 within the proteasome, providing a structural mechanism for how monoubiquitination drives cyclic Rpn10 dissociation and receptor exchange.

    Evidence X-ray crystallography of ubiquitylated Rpn10, cryo-EM superimposition, in vitro dissociation assay, in vivo K84R mutant Co-IP

    PMID:27698474

    Open questions at the time
    • Kinetics of Rpn10 cycling on and off the proteasome in living cells not measured
    • Whether other ubiquitylation sites drive additional regulatory outcomes unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • Open questions remain: whether monoubiquitination-driven Rpn10 cycling operates in mammalian cells with the same writer/eraser pair, whether PSMD4 mediates proteaphagy in animals, the full structural view of PSMD4 within the intact human 26S proteasome engaging substrate, and the physiological importance of tissue-specific isoforms generated by alternative splicing.
  • Mammalian writer/eraser for PSMD4 monoubiquitination not confirmed
  • Proteaphagy receptor function in mammals untested
  • Full cryo-EM structure of substrate-engaged PSMD4 within human proteasome not published in this timeline

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 6 GO:0005198 structural molecule activity 2
Localization
GO:0005829 cytosol 3 GO:0005634 nucleus 1 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-392499 Metabolism of proteins 8 R-HSA-1640170 Cell Cycle 2 R-HSA-9612973 Autophagy 1
Partners
FAT10ID1NEDD4NUB1LRAD23ARAD23BRPN13UBQLN2
Complex memberships
26S proteasome 19S regulatory particle

Evidence

Reading pass · 38 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1995 S5a (PSMD4) and its plant homolog MBP1 bind multiubiquitin chains with high affinity and act as the ubiquitin-conjugate recognition component of the 26S proteasome; free MBP1/S5a potently inhibits ubiquitin-dependent proteolysis in reticulocyte lysates and Xenopus egg extracts by competing for ubiquitinated substrate recognition, without affecting peptidase activity or ubiquitin removal. Cell-free ubiquitin-dependent proteolysis assay (reticulocyte lysate, Xenopus egg extracts), multiubiquitin chain binding assay The Journal of biological chemistry High 8530351
1997 S5a (PSMD4) interacts specifically with Id1 (and less strongly with MyoD and E12) through the N-terminal half of S5a; this interaction is ubiquitin-independent and results in restoration of DNA binding by MyoD-Id1 and E12-Id1 heterodimers, reversal of Id1-mediated repression of the muscle creatine kinase promoter, identifying a non-proteolytic function for S5a in myogenic gene regulation. Yeast two-hybrid, co-immunoprecipitation, DNA-binding assays (EMSA), luciferase/CAT reporter assays, deletion mutagenesis The Journal of biological chemistry Medium 9235903
1997 The yeast S5a ortholog Sun1p (encoded by SUN1, a multicopy suppressor of nin1-1) binds ubiquitin-lysozyme conjugates in vitro; both Sun1p and Sun2p co-migrate with known proteasome components in glycerol density gradients, establishing them as components of the 19S regulatory module of the yeast 26S proteasome. Multicopy suppressor screen, glycerol density gradient sedimentation, ubiquitin-conjugate binding assay Molecular biology of the cell Medium 9017604
1999 hHR23B and hHR23A interact specifically with S5a (PSMD4), a subunit of the human 26S proteasome, via the ubiquitin-like (UbL) domain of hHR23; co-sedimentation in glycerol gradients of HeLa extracts confirms in vivo association; hHR23B inhibits degradation of ubiquitinated lysozyme in reticulocyte lysate, linking hHR23 to the proteasomal proteolytic pathway. Yeast two-hybrid, glycerol gradient centrifugation, co-precipitation with deletion mutants, in vitro degradation assay The Journal of biological chemistry High 10488153
1999 Double deletion of RAD23 and RPN10 (S5a ortholog) in S. cerevisiae causes pleiotropic defects (slow growth, cold sensitivity, G2/M delay, canavanine sensitivity) not seen in single mutants; a Rad23 mutant unable to bind the proteasome (ΔUbL) fails to suppress these defects, and the double mutant accumulates multiubiquitinated proteins and stabilizes a specific proteolytic substrate, demonstrating overlapping roles of Rad23 and Rpn10 in proteasomal substrate delivery. Yeast genetics (double deletion, epistasis), pulse-chase, western blot for ubiquitin conjugates, flow cytometry (cell cycle) Genetics High 10471701
2000 The fission yeast Rpn10 ortholog Pus1 is not essential for viability but is synthetically lethal with mts3 (Rpn12), pad1 (Rpn11), and mts4 (Rpn1) mutations; overexpression of wild-type Pus1 rescues mts3-1, whereas a polyubiquitin-binding-deficient mutant cannot, demonstrating that the polyubiquitin-binding activity of Rpn10 is essential when other proteasomal functions are compromised. Pus1 and Mts3 interact in vitro. Genetic epistasis (synthetic lethality), multicopy suppression, in vitro polyubiquitin binding assay, in vitro protein interaction The Journal of biological chemistry High 10809753
2000 Mouse Rpn10 mRNA is alternatively spliced to generate at least five isoforms (Rpn10a–e); Rpn10a is ubiquitously expressed while Rpn10e is embryo/brain-specific. Both isoforms incorporate into 26S proteasome with similar affinity for multiubiquitylated lysozyme in vitro, but exert markedly divergent effects on B-type cyclin destruction in Xenopus egg extracts, demonstrating that alternative splicing generates functionally distinct proteasome forms. RT-PCR/cDNA cloning, Xenopus egg extract cyclin destruction assay, in vitro binding assay with [125I]-ubiquitylated lysozyme The EMBO journal High 10921894
2001 Immobilized GST-S5a affinity chromatography purifies polyubiquitinated proteins (requiring chains of ≥4 ubiquitins) from mammalian tissue extracts; the major non-ubiquitinated S5a-binding protein purified from human placental extracts is hHR23B, interacting via its N-terminal UbL domain. GST pulldown affinity chromatography, 2D gel electrophoresis, western blot, Edman degradation protein sequencing Proteomics Medium 11677784
2002 NMR structures of the UbL domains of hPLIC-2 and hHR23a reveal that S5a binds these UbL modules using surfaces overlapping with ubiquitin's S5a-contact surface; the S5a-binding surface on hPLIC-2's UbL is required for its proteasome interaction, identifying the molecular basis for how polyubiquitinated proteins are targeted to the proteasome via UbL-UIM interactions. NMR spectroscopy (solution structure determination), homology modeling, NMR chemical shift perturbation mapping, GST pulldown Biochemistry High 11827521
2002 Triple deletion of RAD23, DSK2, and RPN10 in S. cerevisiae causes massive accumulation of polyubiquitinated proteins; Dsk2, Rad23, and Rpn10 have different capacities to bind multiubiquitin chains in vitro; Ddi1 has similar chain-binding activity to Rad23 and Dsk2, establishing cooperative redundancy among these ubiquitin-like proteins and Rpn10 in ubiquitin-dependent proteolysis. Yeast genetics (triple deletion), western blot for ubiquitin conjugates, in vitro multiubiquitin chain binding assay Biochemical and biophysical research communications Medium 12051757
2003 NMR spectroscopy of full-length hHR23a (40 kDa) reveals four structured domains connected by flexible linkers; the N-terminal UbL domain interacts intramolecularly with the two UBA domains, giving hHR23a a closed conformation. Binding of the proteasomal subunit S5a disrupts these intramolecular interactions, causing hHR23a to adopt an open conformation, demonstrating that S5a regulates hHR23a protein structure. NMR spectroscopy (structure determination, residual dipolar coupling, chemical shift perturbation), SAXS-like analysis Proceedings of the National Academy of Sciences of the United States of America High 14557549
2003 NMR structure of the UbL domain of hHR23B and NMR chemical shift perturbation mapping reveal that UbL and ubiquitin share similar surfaces for binding the UBA1, UBA2, and PUbS2 (S5a polyubiquitin-binding site 2) domains; UBA domains bind ubiquitin at Lys-48 (required for multiubiquitin assembly), explaining inhibition of multiubiquitination by hHR23B; intramolecular UbL-UBA interaction is also demonstrated. NMR spectroscopy (structure determination, chemical shift perturbation) The Journal of biological chemistry High 12832454
2003 Deletion of S5a/Rpn10/p54 in Drosophila causes larval-pupal lethality, multiple mitotic defects, accumulation of high-molecular-weight ubiquitinated proteins, and massive accumulation of defective 26S proteasome particles; deletion does not destabilize the 19S regulatory complex assembly; lethality reflects depletion of maternally provided proteasome and sudden increased proteolytic demand at pupation, establishing an essential role for S5a/Rpn10 in higher eukaryotes. Drosophila genetics (null deletion), immunofluorescence, western blot, electron microscopy of proteasome particles, cell cycle analysis Journal of cell science High 12584246
2003 The crystal/NMR structure of S5a's two ubiquitin-interacting motifs (UIMs) bound to the UbL domain of HR23B reveals that UbL presents one hydrophobic and two polar contact sites for UIM interaction; a histidine residue in ubiquitin (absent in UbL) interferes with UIM/UBA binding in a pH-dependent manner, identifying a regulatory mechanism for ubiquitin vs. UbL discrimination. NMR spectroscopy (structure determination, chemical shift perturbation), site-directed mutagenesis, binding affinity measurements The Journal of biological chemistry High 14585839
2004 S5a/Rpn10 directly recognizes ubiquitin chains via its UIM; Rad23 recognizes chains indirectly as a reversibly bound proteasome cofactor. A block substitution in the UIM of RPN10 combined with RAD23 null mutation causes synthetic protein degradation defects in vivo. Rad23 and Ubp6 both bind Rpn1 via N-terminal UbL domains but do not compete with each other, suggesting Rpn1 acts as a scaffold for multiple ubiquitin-handling proteins. Purified biochemical system (in vitro ubiquitin chain binding), yeast genetics (double mutant), co-immunoprecipitation, in vitro binding assay The Journal of biological chemistry High 15117949
2005 Crystal structure of S5a(196–306) alone and complexed with two monoubiquitin molecules reveals that the two UIMs of S5a have different ubiquitin-binding mechanisms and distinct specificities for ubiquitin-like domains; S5a(196–306) binds K63-linked and K48-linked polyubiquitin chains, preferring longer chains in both cases, providing a structural model for polyubiquitin recognition by the proteasome. X-ray crystallography, NMR spectroscopy, ITC (isothermal titration calorimetry) Journal of molecular biology High 15826667
2005 Proteomic analysis of polyubiquitin conjugates accumulating in rpn10Δ yeast identifies 54 proteins uniquely recovered from rpn10Δ cells (including Sic1 and Gcn4), establishing that the Rpn10 substrate receptor contributes to the turnover of multiple proteasome targets beyond cell-cycle regulators. Quantitative mass spectrometry of polyubiquitin conjugates, IMAC enrichment, H6-ubiquitin pulldown, pulse-chase Molecular & cellular proteomics : MCP High 15699485
2005 NUB1 interacts directly with S5a (PSMD4) through its C-terminal region (residues 536–584), not its UbL domain, as shown by GST pulldown and yeast two-hybrid; the UbL domain of NUB1 is required for its function but not for S5a binding, establishing a distinct docking mechanism for NUB1 at the proteasome. GST pulldown, yeast two-hybrid, deletion mutagenesis Biochemical and biophysical research communications Medium 16171779
2006 Knockdown of the C. elegans rpn-10 gene (but not other proteasome subunit genes) sexually transforms hermaphrodites to females by eliminating hermaphrodite spermatogenesis; feminization is rescued by knockdown of tra-2, indicating RPN-10 controls sex determination via the TRA-2-mediated pathway; TRA-2 protein accumulates in rpn-10-defective worms; co-knockdown of rpn-10 and ufd-2 (ubiquitin ligase) overcomes germline-masculinizing effects, placing RPN-10 in a specific ubiquitin-dependent sex determination pathway. RNAi knockdown (C. elegans), genetic epistasis, western blot for TRA-2 accumulation Molecular biology of the cell High 17050737
2007 NMR mapping of the interaction between the UIM of S5a and K48-linked or K63-linked di-ubiquitin shows that UIM binding involves a conformational transition in K48-linked di-ubiquitin that opens the hydrophobic interdomain interface, allowing UIM to enter and bind the same hydrophobic patch as in monoubiquitin:UIM complexes; up to two UIM molecules can bind di-ubiquitin with essentially the same interface for both chain types. NMR spectroscopy (chemical shift perturbation, titration), stoichiometry analysis Journal of molecular biology High 17368669
2009 S5a/PSMD4 can be ubiquitinated by all types of E3 ligases tested (RING, U-box, HECT) using UbcH5-related E2s but not by UbcH1 or UbcH13/Uev1a; ubiquitination requires S5a's UIM domains (UIM-deleted mutant is not ubiquitinated) and occurs through S5a binding to ubiquitin chains on E3s or substrates, rather than through conventional degron recognition; S5a is rapidly degraded in vivo, linking UIM-mediated ubiquitin-chain binding to S5a's own turnover. In vitro ubiquitination assays with multiple E3/E2 combinations, mutagenesis (UIM deletion, UIM-GST fusion), pulse-chase in vivo The Journal of biological chemistry High 19240029
2009 NMR spectroscopy reveals that the parkin UbL domain preferentially binds UIM I of S5a, with K48A substitution in parkin UbL strongly diminishing this interaction; in contrast, parkin recruits Eps15 using both UIM sequences and a larger interaction surface including β1 and β2 strands, demonstrating that the parkin UbL domain uses differential surfaces to recognize the S5a proteasomal subunit versus a ubiquitination substrate. NMR spectroscopy (structure determination, chemical shift perturbation), mutagenesis The Journal of biological chemistry High 19875440
2010 Rpn10/S5a is monoubiquitinated in vivo at its UIM; monoubiquitination inhibits the UIM's ability to interact with ubiquitinated substrates and shuttling factors. Rsp5 (NEDD4 family E3 ligase) catalyzes Rpn10 monoubiquitination while Ubp2 (deubiquitinase) reverses it; monoubiquitination decreases under stress conditions, identifying a regulated mechanism of proteasome ubiquitin-receptor availability. In vivo ubiquitination assays, mass spectrometry, genetic deletion of Rsp5/Ubp2, substrate-binding assays, stress experiments Molecular cell High 20542005
2010 PSMD4 (S5a/Rpn10) is localized to the acrosomal surface of porcine sperm; a monoclonal anti-PSMD4 antibody blocks sperm-zona pellucida (ZP) penetration but not binding during in vitro fertilization; mutant ubiquitins (Ub+1 and Ub5+1) refractory to 19S processing also inhibit fertilization; PSMD4 co-immunoprecipitates with acrosin inhibitor (AI), and ubiquitinated AI species are isolated from sperm, establishing that PSMD4-mediated substrate recognition is required for sperm-ZP penetration. Immunofluorescence localization, antibody blocking in IVF, co-immunoprecipitation, proteomic analysis, affinity purification of ubiquitinated proteins, fractionation Cell and tissue research Medium 20526895
2010 The structure of full-length S. pombe Rpn10 is determined by X-ray crystallography (VWA domain) and NMR (full-length characterization); the single UIM of SpRpn10 forms a 1:1 complex with K48-linked di-ubiquitin and binds it selectively over monoubiquitin and K63-linked di-ubiquitin; SpRpn10's UIM also binds SpRpn12 (lid subunit) with comparable affinity, the first observation of a UIM binding a non-ubiquitin fold, suggesting Rpn12 modulates Rpn10's ubiquitin receptor activity. X-ray crystallography, NMR spectroscopy, isothermal titration calorimetry (ITC), surface plasmon resonance The Journal of biological chemistry High 20739285
2011 In Arabidopsis, the null rpn10-2 mutant shows decreased double-capped proteasomes and increased 20S core complexes, indicating RPN10 is required for proper 26S proteasome assembly; a RPN10 variant defective in ubiquitylated substrate recognition rescues all rpn10-2 phenotypes, demonstrating that the proteasome assembly/structural function (not substrate recognition) is primarily responsible for null phenotypes, while multiple redundant recognition pathways exist in plants. T-DNA knockout, RNAi knockdown, proteasome sedimentation analysis, plant phenotype rescue with domain mutants The Plant cell High 21764993
2012 FAT10 and NUB1L dock with the 26S proteasome through hRpn10/S5a (PSMD4) via its VWA domain (not the UIM); NUB1L can additionally bind Rpn1/S2. Human Rpn10's VWA domain is sufficient to enable FAT10 degradation in Rpn10-deficient yeast. Depletion of hRpn10 in human cells causes accumulation of FAT10 conjugates, establishing the VWA domain of hRpn10 as a ubiquitin-independent receptor for ubiquitin-like proteins within the proteasome. Co-immunoprecipitation, yeast complementation, RNAi knockdown in human cells, VWA domain deletion/mutation analysis Nature communications High 22434192
2012 In vivo ubiquitylation of Drosophila p54/Rpn10 occurs at the conserved C-terminal lysine cluster; extraproteasomal p54 is extensively multiubiquitylated while proteasome-assembled p54 is only modestly modified. Ubiquitylation of p54 impairs its UIM-mediated interaction with the UbL domains of Dsk2 and Rad23; deletion of the conserved lysine cluster shifts lethality of Δp54 from early pupa to pharate adult, indicating ubiquitylated extraproteasomal p54 has an essential role in the pupa-adult transition. In vivo ubiquitylation mass spectrometry, transgenic rescue in Δp54 Drosophila, in vitro UIM-UbL binding assay, proteasome fractionation Biochemistry High 22364263
2013 Knockdown of S5a/PSMD4 in human cells inhibits p53 protein degradation and causes accumulation of ubiquitinated p53; overexpression of a dominant-negative S5a lacking UIMs (but still incorporating into proteasome) stabilizes p53; siRNA rescue confirms UIMs are required for maintaining low p53 levels. In contrast, S5a knockdown has no effect on Mdm2 degradation rate, establishing that proteasomal recognition of p53 is selectively S5a-dependent while Mdm2 uses an S5a-independent pathway. S5a depletion causes p53-dependent decrease in cell proliferation. siRNA knockdown, dominant-negative overexpression, pulse-chase, ubiquitination assay, proliferation assay Oncogene High 24121268
2013 S5a/Rpn10 (PSMD4) localizes to the centrosome in mammalian neurons and is essential for proteasomal activity at centrosomes; loss of S5a reduces centrosomal proteasome activity and impairs dendrite arbor elaboration in rodent brain in vivo; the helix-loop-helix protein Id1 disrupts the interaction of S5a with the proteasomal lid, thereby inhibiting centrosomal proteasome activity and dendrite elaboration. Immunofluorescence/subcellular fractionation, in vivo RNAi in rodent brain (in utero electroporation), co-immunoprecipitation, proteasome activity assay at isolated centrosomes Cell reports High 23831032
2013 Ube3a (Drosophila UBE3A ortholog) directly ubiquitinates Rpn10 in neuronal cells; among Ube3a substrates identified, only Rpn10 is targeted for degradation upon ubiquitination; overexpression of Ube3a and the C-terminal part of Rpn10 leads to enhanced accumulation of ubiquitinated proteins; genetic interaction between Ube3a and Rpn10 C-terminus is demonstrated in vivo. In vitro ubiquitination assay, mass spectrometry substrate screen, western blot for protein levels, Drosophila genetics (in vivo overexpression) Cellular and molecular life sciences : CMLS Medium 24292889
2015 Liver-specific deletion of both Rpn10 and Rpn13 in mice causes severe liver injury with massive accumulation of ubiquitin conjugates, whereas single deletions show only modest impairment; mHR23B and ubiquilin/Plic-1/4 fail to bind the proteasome in the absence of both Rpn10 and Rpn13, establishing that these two subunits are the main docking receptors for UBL-UBA shuttle proteins in mammals and act redundantly in ubiquitinated protein recognition and homeostasis. Liver-specific conditional knockout (Cre-lox), western blot for ubiquitin conjugates, co-immunoprecipitation, liver histology, re-expression rescue PLoS genetics High 26222436
2015 Rpn10 monoubiquitination decreases both Rpn10-proteasome and Rpn10-Dsk2 associations; this facilitates formation of Dsk2-proteasomes in vivo, switching the proteasome between an 'Rpn10-high/Dsk2-low' and 'Rpn10-low/Dsk2-high' state; functional yeast proteasomes can associate and dissociate with Rpn10 dynamically. In vivo ubiquitination assays, quantitative co-immunoprecipitation, yeast genetics with UIM/UBL mutants The Biochemical journal High 26450923
2015 In Arabidopsis, autophagic turnover of inactive 26S proteasomes (proteaphagy) requires the proteasome subunit RPN10, which simultaneously binds ATG8 (via an AIM motif) and ubiquitin (via UIM), acting as a selective autophagy receptor targeting ubiquitylated, inhibited proteasomes for vacuolar delivery; proteasome inhibition stimulates comprehensive ubiquitylation of the complex prior to proteaphagy. GFP-proteasome imaging in Arabidopsis, autophagy pathway genetic analysis (ATG mutants), co-immunoprecipitation of RPN10 with ATG8 and ubiquitin, nitrogen starvation and proteasome inhibitor experiments Molecular cell High 26004230
2016 Crystal structure of ubiquitylated Rpn10 (K84 ubiquitylation site) reveals a novel ubiquitin-binding patch directing K84 modification; superimposition onto proteasome EM models shows that the conjugated ubiquitin clashes with Rpn9, suggesting ubiquitylation releases Rpn10 from the proteasome; in vitro, ubiquitylation of immobilized-proteasome Rpn10 dissociates it from the complex while unmodified Rpn10 remains associated; in vivo, Rpn10-K84R mutant is stably associated with Rpn9, confirming that ubiquitylation promotes cyclic Rpn10 dissociation from the proteasome. X-ray crystallography (bacterial ubiquitylation system), cryo-EM model superimposition, in vitro proteasome dissociation assay, in vivo co-immunoprecipitation with K84R mutant Nature communications High 27698474
2016 Cytoplasmic Nrf2 (cNrf2) persistently increases PSMD4 expression via the HIF1α/β-catenin axis; in a feed-forward loop, elevated PSMD4 reciprocally enhances Nrf2 nuclear export by increasing CRM1 expression through p53 degradation; this cascade promotes tumor invasion and growth in colorectal cancer cells and xenograft models, which is suppressed by the proteasomal inhibitor carfilzomib or the β-catenin inhibitor XAV939. Cell line overexpression/knockdown, reporter assays, western blot for pathway components, xenograft mouse model with inhibitor treatment, immunohistochemistry Free radical biology & medicine Medium 27033953
2017 PSMD4 is amplified and overexpressed in breast cancer; knockdown of PSMD4 in amplified breast cancer cells results in knockdown of PARP1 protein; breast cancer cells with PSMD4 copy-number gain are significantly more sensitive to talazoparib (PARP inhibitor); loss of PSMD4 amplicon during acquired PARPi resistance correlates with PSMD4 downregulation and decreased PARP1, establishing PSMD4 as a regulator of PARP1 protein levels. Array-CGH, siRNA knockdown, western blot, cell viability assay, comparison of sensitive vs. resistant cell lines Genes, chromosomes & cancer Medium 28316110
2019 Rpn10 promotes proliferation in hepatocellular carcinoma (HCC) cells; Rpn10 knockdown induces G1 phase cell cycle arrest; Rpn10 directly promotes PTEN degradation through the ubiquitin-proteasome system; HIF1α directly binds the Rpn10 promoter and increases its expression in HCC; the PTEN/Akt signaling pathway mediates Rpn10's effect on cell proliferation, as active Akt overexpression rescues Rpn10-knockdown growth suppression. siRNA knockdown, overexpression of active Akt (rescue), cell cycle analysis, ChIP (HIF1α on Rpn10 promoter), pulse-chase/cycloheximide chase for PTEN stability, western blot Cancer letters Medium 30673593

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1996 Structure and functions of the 20S and 26S proteasomes. Annual review of biochemistry 2108 8811196
2005 Towards a proteome-scale map of the human protein-protein interaction network. Nature 2090 16189514
2002 Isolation of a human gene that inhibits HIV-1 infection and is suppressed by the viral Vif protein. Nature 1924 12167863
2012 Insights into RNA biology from an atlas of mammalian mRNA-binding proteins. Cell 1718 22658674
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
2001 The co-chaperone CHIP regulates protein triage decisions mediated by heat-shock proteins. Nature cell biology 829 11146632
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
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
2008 Genome-scale RNAi screen for host factors required for HIV replication. Cell host & microbe 627 18976975
2003 HIV-1 Vif blocks the antiviral activity of APOBEC3G by impairing both its translation and intracellular stability. Molecular cell 607 14527406
2004 Sequestosome 1/p62 is a polyubiquitin chain binding protein involved in ubiquitin proteasome degradation. Molecular and cellular biology 587 15340068
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
2023 Pulsed Field Ablation for the Treatment of Atrial Fibrillation: PULSED AF Pivotal Trial. Circulation 453 36877118
2017 Screening for Atrial Fibrillation: A Report of the AF-SCREEN International Collaboration. Circulation 440 28483832
2015 Autophagic Degradation of the 26S Proteasome Is Mediated by the Dual ATG8/Ubiquitin Receptor RPN10 in Arabidopsis. Molecular cell 336 26004230
2004 Rad23 and Rpn10 serve as alternative ubiquitin receptors for the proteasome. The Journal of biological chemistry 274 15117949
1999 Differential regulation of glucocorticoid receptor transcriptional activation via AF-1-associated proteins. The EMBO journal 229 10508170
1999 Interaction of hHR23 with S5a. The ubiquitin-like domain of hHR23 mediates interaction with S5a subunit of 26 S proteasome. The Journal of biological chemistry 224 10488153
2003 The pleiotropic role of the 26S proteasome subunit RPN10 in Arabidopsis growth and development supports a substrate-specific function in abscisic acid signaling. The Plant cell 201 12671091
2002 Structural studies of the interaction between ubiquitin family proteins and proteasome subunit S5a. Biochemistry 199 11827521
1996 Identification of AF-6 and canoe as putative targets for Ras. The Journal of biological chemistry 184 8557659
2010 Bisphenol AF is a full agonist for the estrogen receptor ERalpha but a highly specific antagonist for ERbeta. Environmental health perspectives 168 20427257
2022 SPEACH_AF: Sampling protein ensembles and conformational heterogeneity with Alphafold2. PLoS computational biology 159 35994486
2005 Structure of S5a bound to monoubiquitin provides a model for polyubiquitin recognition. Journal of molecular biology 159 15826667
2002 The AF-1 activation-function of ERalpha may be dispensable to mediate the effect of estradiol on endothelial NO production in mice. Proceedings of the National Academy of Sciences of the United States of America 147 11854517
2011 Pharmacogenomics of bortezomib test-dosing identifies hyperexpression of proteasome genes, especially PSMD4, as novel high-risk feature in myeloma treated with Total Therapy 3. Blood 146 21628408
2003 DNA-repair protein hHR23a alters its protein structure upon binding proteasomal subunit S5a. Proceedings of the National Academy of Sciences of the United States of America 129 14557549
1999 Pleiotropic defects caused by loss of the proteasome-interacting factors Rad23 and Rpn10 of Saccharomyces cerevisiae. Genetics 128 10471701
2000 Structural basis for autorepression of retinoid X receptor by tetramer formation and the AF-2 helix. Genes & development 126 10970886
2010 Monoubiquitination of RPN10 regulates substrate recruitment to the proteasome. Molecular cell 120 20542005
2002 Ubiquitin-like proteins and Rpn10 play cooperative roles in ubiquitin-dependent proteolysis. Biochemical and biophysical research communications 111 12051757
2010 AF-353, a novel, potent and orally bioavailable P2X3/P2X2/3 receptor antagonist. British journal of pharmacology 105 20590629
2003 Binding surface mapping of intra- and interdomain interactions among hHR23B, ubiquitin, and polyubiquitin binding site 2 of S5a. The Journal of biological chemistry 97 12832454
1997 Yeast counterparts of subunits S5a and p58 (S3) of the human 26S proteasome are encoded by two multicopy suppressors of nin1-1. Molecular biology of the cell 93 9017604
2005 Analysis of polyubiquitin conjugates reveals that the Rpn10 substrate receptor contributes to the turnover of multiple proteasome targets. Molecular & cellular proteomics : MCP 87 15699485
2008 Coordination of synaptic adhesion with dendritic spine remodeling by AF-6 and kalirin-7. The Journal of neuroscience : the official journal of the Society for Neuroscience 81 18550750
2008 Extraproteasomal Rpn10 restricts access of the polyubiquitin-binding protein Dsk2 to proteasome. Molecular cell 80 18995839
2017 Bisphenol AF negatively affects oocyte maturation of mouse in vitro through increasing oxidative stress and DNA damage. Chemico-biological interactions 77 29102535
2003 The Bcr kinase downregulates Ras signaling by phosphorylating AF-6 and binding to its PDZ domain. Molecular and cellular biology 76 12808105
2014 Exposure to Bisphenol AF disrupts sex hormone levels and vitellogenin expression in zebrafish. Environmental toxicology 75 25213402
1995 Inhibition of ubiquitin-mediated proteolysis by the Arabidopsis 26 S protease subunit S5a. The Journal of biological chemistry 75 8530351
2019 Bisphenol AF compromises blood-testis barrier integrity and sperm quality in mice. Chemosphere 74 31362132
2012 FAT10 and NUB1L bind to the VWA domain of Rpn10 and Rpn1 to enable proteasome-mediated proteolysis. Nature communications 73 22434192
2015 Redundant Roles of Rpn10 and Rpn13 in Recognition of Ubiquitinated Proteins and Cellular Homeostasis. PLoS genetics 71 26222436
2003 Structure of the ubiquitin-interacting motif of S5a bound to the ubiquitin-like domain of HR23B. The Journal of biological chemistry 71 14585839
2000 Analysis of a gene encoding Rpn10 of the fission yeast proteasome reveals that the polyubiquitin-binding site of this subunit is essential when Rpn12/Mts3 activity is compromised. The Journal of biological chemistry 71 10809753
2013 Ube3a, the E3 ubiquitin ligase causing Angelman syndrome and linked to autism, regulates protein homeostasis through the proteasomal shuttle Rpn10. Cellular and molecular life sciences : CMLS 70 24292889
2013 Afadin/AF-6 and canoe: roles in cell adhesion and beyond. Progress in molecular biology and translational science 68 23481206
2004 Rad23 and Rpn10: perennial wallflowers join the melee. Trends in biochemical sciences 65 15544949
2014 Disrupted interaction between CFTR and AF-6/afadin aggravates malignant phenotypes of colon cancer. Biochimica et biophysica acta 60 24373847
2012 Neurotoxic effects of bisphenol AF on calcium-induced ROS and MAPKs. Neurotoxicity research 60 22996013
2008 Protein polarization is critical to stabilizing AF-2 and helix-2' domains in ligand binding to PPAR-gamma. Journal of the American Chemical Society 60 19007119
2010 Structure of Rpn10 and its interactions with polyubiquitin chains and the proteasome subunit Rpn12. The Journal of biological chemistry 58 20739285
2003 Deletion of proteasomal subunit S5a/Rpn10/p54 causes lethality, multiple mitotic defects and overexpression of proteasomal genes in Drosophila melanogaster. Journal of cell science 57 12584246
2019 Exosomal miR-320d derived from adipose tissue-derived MSCs inhibits apoptosis in cardiomyocytes with atrial fibrillation (AF). Artificial cells, nanomedicine, and biotechnology 54 31591913
2011 The defective proteasome but not substrate recognition function is responsible for the null phenotypes of the Arabidopsis proteasome subunit RPN10. The Plant cell 54 21764993
2016 The AF-1-deficient estrogen receptor ERα46 isoform is frequently expressed in human breast tumors. Breast cancer research : BCR 51 27927249
1999 In vivo interaction of AF-6 with activated Ras and ZO-1. Biochemical and biophysical research communications 51 10334923
2006 In vivo role of the HNF4alpha AF-1 activation domain revealed by exon swapping. The EMBO journal 48 16498401
2018 Endocrine activities and adipogenic effects of bisphenol AF and its main metabolite. Chemosphere 47 30408883
2021 Extracellular matrix remodeling precedes atrial fibrillation: Results of the PREDICT-AF trial. Heart rhythm 46 34332113
2018 Bisphenol AF exerts estrogenic activity in MCF-7 cells through activation of Erk and PI3K/Akt signals via GPER signaling pathway. Chemosphere 46 30590302
2006 Proteasomal ubiquitin receptor RPN-10 controls sex determination in Caenorhabditis elegans. Molecular biology of the cell 45 17050737
2001 Purification of poly-ubiquitinated proteins by S5a-affinity chromatography. Proteomics 45 11677784
2000 Developmentally regulated, alternative splicing of the Rpn10 gene generates multiple forms of 26S proteasomes. The EMBO journal 45 10921894
2009 PPAR-gamma AF-2 domain functions as a component of a ubiquitin-dependent degradation signal. Obesity (Silver Spring, Md.) 44 19148122
2023 Integrated Care for Atrial Fibrillation Using the ABC Pathway in the Prospective APHRS-AF Registry. JACC. Asia 43 37614548
2009 The ubiquitin-interacting motif protein, S5a, is ubiquitinated by all types of ubiquitin ligases by a mechanism different from typical substrate recognition. The Journal of biological chemistry 42 19240029
2019 Nature's Combinatorial Biosynthesis Produces Vatiamides A-F. Angewandte Chemie (International ed. in English) 41 31071229
2019 LncRNA-LINC00472 contributes to the pathogenesis of atrial fibrillation (Af) by reducing expression of JP2 and RyR2 via miR-24. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 40 31562981
2008 Active nuclear receptors exhibit highly correlated AF-2 domain motions. PLoS computational biology 40 18617990
2009 An improved manufacturing process for Xyntha/ReFacto AF. Haemophilia : the official journal of the World Federation of Hemophilia 39 20041957
2022 Bisphenol AF blocks Leydig cell regeneration from stem cells in male rats. Environmental pollution (Barking, Essex : 1987) 38 35026324
2017 Isolation, Biosynthesis and Chemical Modifications of Rubterolones A-F: Rare Tropolone Alkaloids from Actinomadura sp. 5-2. Chemistry (Weinheim an der Bergstrasse, Germany) 38 28463423
2002 Formation of an hER alpha-COUP-TFI complex enhances hER alpha AF-1 through Ser118 phosphorylation by MAPK. The EMBO journal 38 12093745
2002 Neuropeptide AF and FF modulation of adipocyte metabolism. Primary insights from functional genomics and effects on beta-adrenergic responsiveness. The Journal of biological chemistry 38 12149260
1996 Lysis and biological control of Aspergillus niger by Bacillus subtilis AF 1. Canadian journal of microbiology 38 8801004
2018 The adjuvant GLA-AF enhances human intradermal vaccine responses. Science advances 35 30221194
2016 Cytoplasmic localization of Nrf2 promotes colorectal cancer with more aggressive tumors via upregulation of PSMD4. Free radical biology & medicine 34 27033953
2016 Structure of ubiquitylated-Rpn10 provides insight into its autoregulation mechanism. Nature communications 34 27698474
2013 The degradation of p53 and its major E3 ligase Mdm2 is differentially dependent on the proteasomal ubiquitin receptor S5a. Oncogene 33 24121268
1997 Novel regulation of the helix-loop-helix protein Id1 by S5a, a subunit of the 26 S proteasome. The Journal of biological chemistry 32 9235903
2017 CLDN6 enhances chemoresistance to ADM via AF-6/ERKs pathway in TNBC cell line MDAMB231. Molecular and cellular biochemistry 31 29159771
2013 The ubiquitin receptor S5a/Rpn10 links centrosomal proteasomes with dendrite development in the mammalian brain. Cell reports 31 23831032
2022 Intergenerational toxic effects of parental exposure to bisphenol AF on offspring and epigenetic modulations in zebrafish. The Science of the total environment 28 35143790
2020 Spironolactone in Atrial Fibrillation With Preserved Cardiac Fraction: The IMPRESS-AF Trial. Journal of the American Heart Association 28 32909497
2013 Evaluation of Mucoadhesive Gels with Propolis (EPP-AF) in Preclinical Treatment of Candidiasis Vulvovaginal Infection. Evidence-based complementary and alternative medicine : eCAM 28 23997797
2007 Mapping the interactions between Lys48 and Lys63-linked di-ubiquitins and a ubiquitin-interacting motif of S5a. Journal of molecular biology 28 17368669
2003 Role of conserved hydrophobic amino acids in androgen receptor AF-1 function. Journal of molecular endocrinology 28 14664704
2000 Identification and functional analysis of sequence rearrangements in the long control region of human papillomavirus type 16 Af-1 variants isolated from Ugandan penile carcinomas. The Journal of general virology 28 11086128
2024 Effects and mechanisms of endocrine disruptor bisphenol AF on male reproductive health: A mini review. Ecotoxicology and environmental safety 27 38583312
2017 Proteasome ubiquitin receptor PSMD4 is an amplification target in breast cancer and may predict sensitivity to PARPi. Genes, chromosomes & cancer 27 28316110
2017 Brain Mitochondrial Subproteome of Rpn10-Binding Proteins and Its Changes Induced by the Neurotoxin MPTP and the Neuroprotector Isatin. Biochemistry. Biokhimiia 27 28320274
2014 Neuropeptide AF induces anxiety-like and antidepressant-like behavior in mice. Behavioural brain research 27 25116251
2013 Tamoxifen elicits atheroprotection through estrogen receptor α AF-1 but does not accelerate reendothelialization. The American journal of pathology 27 23669343
2010 Interference with the 19S proteasomal regulatory complex subunit PSMD4 on the sperm surface inhibits sperm-zona pellucida penetration during porcine fertilization. Cell and tissue research 27 20526895
2022 4S-AF scheme and ABC pathway guided management improves outcomes in atrial fibrillation patients. European journal of clinical investigation 26 35080269
2019 Bucindolol for the Maintenance of Sinus Rhythm in a Genotype-Defined HF Population: The GENETIC-AF Trial. JACC. Heart failure 26 31042551
2007 Regulation of c-Src by binding to the PDZ domain of AF-6. The EMBO journal 26 17491594
2005 Interaction of NUB1 with the proteasome subunit S5a. Biochemical and biophysical research communications 26 16171779
2000 Co-localization of FAM and AF-6, the mammalian homologues of Drosophila faf and canoe, in mouse eye development. Mechanisms of development 26 10704870
2023 Bisphenol B and bisphenol AF exposure enhances uterine diseases risks in mouse. Environment international 25 36881955
2022 Oxidative Stress and Apoptosis in Bisphenol AF-Induced Neurotoxicity in Zebrafish Embryos. Environmental toxicology and chemistry 25 35723417
2009 Endocrine, behavioral and autonomic effects of neuropeptide AF. Hormones and behavior 25 19269292
2023 Exposure to bisphenol A alternatives bisphenol AF and fluorene-9-bisphenol induces gonadal injuries in male zebrafish. Ecotoxicology and environmental safety 24 36801538
2018 Hyaluronic acid-endostatin2-alft1 (HA-ES2-AF) nanoparticle-like conjugate for the target treatment of diseases. Journal of controlled release : official journal of the Controlled Release Society 24 30170067
2001 Chitin-supplemented formulations improve biocontrol and plant growth promoting efficiency of Bacillus subtilis AF 1. Canadian journal of microbiology 24 11547881
2019 Rpn10 promotes tumor progression by regulating hypoxia-inducible factor 1 alpha through the PTEN/Akt signaling pathway in hepatocellular carcinoma. Cancer letters 23 30673593
2009 Differential interaction of the E3 ligase parkin with the proteasomal subunit S5a and the endocytic protein Eps15. The Journal of biological chemistry 21 19875440
2018 Bisphenol AF as an activator of human estrogen receptor β1 (ERβ1) in breast cancer cell lines. The Journal of toxicological sciences 20 29743443
2015 Rpn10 monoubiquitination orchestrates the association of the ubiquilin-type DSK2 receptor with the proteasome. The Biochemical journal 20 26450923
2012 Ubiquitylation of Drosophila p54/Rpn10/S5a regulates its interaction with the UBA-UBL polyubiquitin receptors. Biochemistry 20 22364263