Affinage

PSMC1

26S proteasome regulatory subunit 4 · UniProt P62191

Round 2 corrected
Length
440 aa
Mass
49.2 kDa
Annotated
2026-04-28
130 papers in source corpus 5 papers cited in narrative 5 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PSMC1 (also called S4; yeast ortholog Rpt2) is an AAA-ATPase subunit of the 19S regulatory particle of the 26S proteasome that couples ATP hydrolysis to the unfolding and translocation of ubiquitinated substrates into the 20S core particle for degradation (PMID:8811196). Within the 19S base, PSMC1 forms a specific heterodimer with S7/PSMC2 and assembles into higher-order subcomplexes with S2 and S5b through distinct N-terminal and C-terminal binding surfaces, and its C-terminal hydrophobic motif is required for docking the ATPase ring onto the core particle (PMID:10625621, PMID:22158466). N-myristoylation of the PSMC1 N-terminus governs nuclear localization of the 26S proteasome; loss of this modification causes proteasome redistribution to the cytoplasm, accumulation of polyubiquitinated nuclear proteins, altered chaperone levels, and impaired protein quality control under stress (PMID:23102099, PMID:26344132).

Mechanistic history

Synthesis pass · year-by-year structured walk · 5 steps
  1. 1996 High

    Establishing that PSMC1 is one of several ATPase subunits in the 19S regulatory particle answered the question of how the 26S proteasome recognizes, unfolds, and translocates ubiquitinated substrates into the 20S core for proteolysis.

    Evidence Biochemical reconstitution, peptidase activity assays, and structural analysis across multiple laboratories

    PMID:8811196

    Open questions at the time
    • Individual contributions of each ATPase subunit to substrate processing were not resolved
    • No structural model of the intact 19S base at atomic resolution at this time
  2. 2000 High

    Mapping the subunit interaction surfaces of PSMC1 revealed that it forms a specific heterodimer with S7 and that S2 and S5b bind distinct regions (N-terminal and C-terminal, respectively), defining the modular assembly pathway of the 19S base.

    Evidence In vitro co-translation/sedimentation, immunoprecipitation of reticulocyte lysate-translated subunits, chimeric ATPase domain-swap experiments

    PMID:10625621

    Open questions at the time
    • Whether these subcomplexes represent on-pathway assembly intermediates in vivo was not determined
    • Structural basis for selectivity of S2 vs S5b binding to different PSMC1 surfaces was not resolved
  3. 2011 Medium

    Genetic analysis of Arabidopsis RPT2 (PSMC1 ortholog) demonstrated that both the C-terminal CP-docking motif and ATPase catalytic activity are essential for 26S proteasome stability and ubiquitin-dependent proteolysis in vivo.

    Evidence Genetic knockout/complementation with wild-type and mutant RPT2 in Arabidopsis, 26S complex stability assays, polyubiquitin accumulation assays

    PMID:22158466

    Open questions at the time
    • Findings are from a plant ortholog; direct validation in mammalian PSMC1 was not performed
    • Contribution of the C-terminal motif relative to other RPT subunit tails was not individually resolved
  4. 2012 High

    Discovery that N-myristoylation of Rpt2/PSMC1 controls nuclear localization of the 26S proteasome revealed a post-translational mechanism for compartment-specific proteasome targeting, independent of assembly or catalytic activity.

    Evidence Site-directed mutagenesis of N-myristoylation site in yeast, fluorescence microscopy, peptidase activity assays, polyubiquitin accumulation assays, growth assays under stress

    PMID:23102099

    Open questions at the time
    • Mechanism by which myristoylation anchors proteasomes in the nucleus is unknown
    • Whether mammalian PSMC1 N-myristoylation has the same localization function was not tested
  5. 2015 Medium

    Quantitative ubiquitinome profiling of myristoylation-deficient Rpt2 mutants demonstrated that loss of nuclear proteasome enrichment specifically upregulates ubiquitination of nucleo-cytoplasmic proteins and alters chaperone networks, establishing PSMC1 myristoylation as a regulator of compartment-specific protein quality control.

    Evidence Anti-K-ε-GG antibody enrichment and quantitative mass spectrometry in yeast myristoylation-site mutants

    PMID:26344132

    Open questions at the time
    • Single-lab proteomics study; independent confirmation in additional systems is lacking
    • Which specific nuclear substrates are direct clients of myristoylation-dependent proteasome targeting is unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • The mechanism by which N-myristoylation physically anchors the proteasome in the nucleus, and whether this modification regulates PSMC1 function in human cells, remains unresolved.
  • No structural or biophysical model for how myristoylation mediates nuclear retention
  • Mammalian validation of myristoylation-dependent localization is absent
  • Individual catalytic contribution of PSMC1 ATPase activity versus other RPT subunits in human proteasomes is not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140657 ATP-dependent activity 2 GO:0016787 hydrolase activity 1 GO:0044183 protein folding chaperone 1
Localization
GO:0005634 nucleus 2
Pathway
R-HSA-392499 Metabolism of proteins 4
Partners
PSMC2PSMD1PSMD11
Complex memberships
19S regulatory particle26S proteasome

Evidence

Reading pass · 5 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2000 The 26S proteasome ATPase subunit S4 (PSMC1) forms specific heterodimeric and higher-order complexes within the 19S regulatory complex. S4 pairs with S7 as an ATPase dimer, and both the non-ATPase subunit S2 (binding the NH2-terminal region of S4) and S5b (binding the COOH-terminal two-thirds of S4) assemble with the S4-S7 dimer to form a tetramer. Domain-mapping with chimeric ATPases identified distinct binding surfaces on S4 for S2 and S5b. In vitro co-translation/sedimentation, immunoprecipitation of reticulocyte lysate-translated subunits, chimeric ATPase domain-swap experiments The Journal of biological chemistry High 10625621
2012 N-myristoylation of the Rpt2 subunit (yeast ortholog of PSMC1) at its N-terminus controls the intracellular localization of the 26S proteasome. Mutation of the N-myristoylation site causes diffusion of nuclear proteasomes into the cytoplasm where they form aggregates, without altering molecular assembly or peptidase activity. Loss of N-myristoylation leads to accumulation of polyubiquitinated proteins and a growth defect under protein-misfolding stress. Site-directed mutagenesis of N-myristoylation site, fluorescence microscopy of proteasome localization, peptidase activity assays, polyubiquitin accumulation assays, growth assays Biochemistry High 23102099
2015 N-myristoylation of yeast Rpt2 (PSMC1 ortholog) specifically regulates the nuclear versus cytoplasmic distribution of the proteasome and thereby controls compartment-specific protein quality control. In N-myristoylation-deficient mutants, nuclear ubiquitination of proteins with nucleo-cytoplasmic localization is upregulated, and levels of nuclear-import-associated Hsp70 chaperones (Ssa, Sse1) are elevated while the nuclear-export chaperone Ssb is reduced, indicating disruption of nuclear protein quality control. Quantitative ubiquitinome profiling (anti-K-ε-GG antibody enrichment, MS), protein-level quantification in myristoylation-site mutant yeast strains Journal of proteomics Medium 26344132
2011 The Arabidopsis RPT2 subunit (plant ortholog of PSMC1) is essential for 26S proteasome assembly. The C-terminal hydrophobic sequence of RPT2, which docks the RPT ATPase ring onto the core particle (CP), is required for function, as is ATP binding/hydrolysis activity. Loss of RPT2 destabilizes the 26S complex and causes accumulation of ubiquitinated proteins, impaired histone dynamics, and developmental defects. Functional redundancy exists between RPT2a and RPT2b paralogs. Genetic analysis of double null mutants, complementation with wild-type and ATP-binding/C-terminal mutants, 26S complex stability assays, polyubiquitin accumulation assays The Plant cell Medium 22158466
1996 The 26S proteasome contains a 19S regulatory complex composed of multiple ATPases (including the S4/PSMC1 subunit) that form a ring and are necessary for binding and degrading ubiquitinated protein substrates. The ATPase subunits are essential for unfolding substrates and translocating them into the 20S core particle. The regulatory complex ATPases bind ubiquitinated substrates and couple ATP hydrolysis to substrate processing. Biochemical reconstitution, peptidase activity assays, review of structural and functional data across multiple studies Annual review of biochemistry High 8811196

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
2010 Network organization of the human autophagy system. Nature 1286 20562859
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
2014 A proteome-scale map of the human interactome network. Cell 977 25416956
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
2004 A physical and functional map of the human TNF-alpha/NF-kappa B signal transduction pathway. Nature cell biology 841 14743216
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
2008 An empirical framework for binary interactome mapping. Nature methods 652 19060904
2003 HIV-1 Vif blocks the antiviral activity of APOBEC3G by impairing both its translation and intracellular stability. Molecular cell 607 14527406
1995 Reversal of the interferon-sensitive phenotype of a vaccinia virus lacking E3L by expression of the reovirus S4 gene. Journal of virology 176 7527085
2001 CD8beta endows CD8 with efficient coreceptor function by coupling T cell receptor/CD3 to raft-associated CD8/p56(lck) complexes. The Journal of experimental medicine 173 11714755
2004 RPT2 is a signal transducer involved in phototropic response and stomatal opening by association with phototropin 1 in Arabidopsis thaliana. The Plant cell 163 15031408
1998 Uncharged S4 residues and cooperativity in voltage-dependent potassium channel activation. The Journal of general physiology 148 9482709
2005 Association of systemic chemokine concentrations with impaired glucose tolerance and type 2 diabetes: results from the Cooperative Health Research in the Region of Augsburg Survey S4 (KORA S4). Diabetes 135 16306328
2001 The S4-S5 linker couples voltage sensing and activation of pacemaker channels. Proceedings of the National Academy of Sciences of the United States of America 131 11553787
2003 Viral stress-inducible protein p56 inhibits translation by blocking the interaction of eIF3 with the ternary complex eIF2.GTP.Met-tRNAi. The Journal of biological chemistry 129 12885778
1998 Role of the S4 in cooperativity of voltage-dependent potassium channel activation. The Journal of general physiology 126 9482708
1989 Mutations in ribosomal proteins S4 and S12 influence the higher order structure of 16 S ribosomal RNA. Journal of molecular biology 112 2477554
2001 Tyrosine phosphorylation-dependent activation of NF-kappa B. Requirement for p56 LCK and ZAP-70 protein tyrosine kinases. European journal of biochemistry 102 11231305
1996 N-type inactivation and the S4-S5 region of the Shaker K+ channel. The Journal of general physiology 99 8882863
1994 Activation of Src-like p56/p53lyn tyrosine kinase by ionizing radiation. The Journal of biological chemistry 90 8051175
2000 Hsp90 is essential for the synthesis and subsequent membrane association, but not the maintenance, of the Src-kinase p56(lck). Molecular biology of the cell 78 10793137
1997 Intrinsic signals in the unique domain target p56(lck) to the plasma membrane independently of CD4. The Journal of cell biology 78 9166404
2005 On the mechanisms of the internalization of S4(13)-PV cell-penetrating peptide. The Biochemical journal 77 15907190
1998 Zinc is essential for binding of p56(lck) to CD4 and CD8alpha. The Journal of biological chemistry 77 9830036
2010 State-dependent electrostatic interactions of S4 arginines with E1 in S2 during Kv7.1 activation. The Journal of general physiology 76 20479111
1993 Proteins homologous to the Xenopus germ cell-specific RNA-binding proteins p54/p56 are temporally expressed in mouse male germ cells. Developmental biology 71 8330676
2016 Potato NPH3/RPT2-Like Protein StNRL1, Targeted by a Phytophthora infestans RXLR Effector, Is a Susceptibility Factor. Plant physiology 70 26966171
1997 Characterization of Borna disease virus p56 protein, a surface glycoprotein involved in virus entry. Journal of virology 70 9060684
2008 Direct and indirect impairment of human dendritic cell function by virulent Francisella tularensis Schu S4. Infection and immunity 69 18981246
2002 The S4 genome segment of baboon reovirus is bicistronic and encodes a novel fusion-associated small transmembrane protein. Journal of virology 69 11836390
1991 Studies on the role of the S4 substrate binding site of HIV proteinases. FEBS letters 63 2001747
2000 p56(dok-2) as a cytokine-inducible inhibitor of cell proliferation and signal transduction. The EMBO journal 62 11013214
2005 Induction and mode of action of the viral stress-inducible murine proteins, P56 and P54. Virology 61 16023166
1999 Expression of the p56(Lck) Y505F mutation in CD45-deficient mice rescues thymocyte development. Molecular and cellular biology 57 10330160
1998 Requirement for p56(lck) tyrosine kinase activation in Th subset differentiation. International immunology 55 9645606
2000 Mapping subunit contacts in the regulatory complex of the 26 S proteasome. S2 and S5b form a tetramer with ATPase subunits S4 and S7. The Journal of biological chemistry 53 10625621
1982 Role of the S4 gene in the establishment of persistent reovirus infection in L cells. Cell 53 7074685
2010 The inhibitory action of P56 on select functions of E1 mediates interferon's effect on human papillomavirus DNA replication. Journal of virology 52 20926571
2000 Calcium-induced ERK activation in human T lymphocytes occurs via p56(Lck) and CaM-kinase. Molecular immunology 52 11164895
2021 Regulation of plant phototropic growth by NPH3/RPT2-like substrate phosphorylation and 14-3-3 binding. Nature communications 49 34675214
1997 Reovirus variants selected during persistent infections of L cells contain mutations in the viral S1 and S4 genes and are altered in viral disassembly. Journal of virology 49 8995660
1999 Trafficking of an acylated cytosolic protein: newly synthesized p56(lck) travels to the plasma membrane via the exocytic pathway. The Journal of cell biology 48 10225948
1991 The rpsD gene, encoding ribosomal protein S4, is autogenously regulated in Bacillus subtilis. Journal of bacteriology 48 1906866
2008 Novel characteristics of the function and induction of murine p56 family proteins. Journal of virology 47 18768971
1996 Mutations in yeast ribosomal proteins S28 and S4 affect the accuracy of translation and alter the sensitivity of the ribosomes to paromomycin. Biochimica et biophysica acta 46 8950190
2012 N-myristoylation of the Rpt2 subunit regulates intracellular localization of the yeast 26S proteasome. Biochemistry 45 23102099
2012 The soluble pyocins S2 and S4 from Pseudomonas aeruginosa bind to the same FpvAI receptor. MicrobiologyOpen 45 23170226
1997 Defective TCR stimulation in anergized type 2 T helper cells correlates with abrogated p56(lck) and ZAP-70 tyrosine kinase activities. Journal of immunology (Baltimore, Md. : 1950) 45 9200438
1997 Interleukin-18 induces activation and association of p56(lck) and MAPK in a murine TH1 clone. Biochemical and biophysical research communications 45 9266843
1996 Molecular cloning of the epidermal growth factor-stimulated protein kinase p56 KKIAMRE. Oncogene 43 9000130
1992 Translational effects and sequence comparisons of the three serotypes of the reovirus S4 gene. Virology 43 1736524
2009 Global stabilization of rRNA structure by ribosomal proteins S4, S17, and S20. Journal of molecular biology 42 19616559
2020 Evidence that the TRPV1 S1-S4 membrane domain contributes to thermosensing. Nature communications 41 32820172
2003 Molecular design, synthesis, and structure-Activity relationships leading to the potent and selective p56(lck) inhibitor BMS-243117. Bioorganic & medicinal chemistry letters 41 12798323
1999 Voltage-shift of the current activation in HERG S4 mutation (R534C) in LQT2. Cardiovascular research 41 10690305
2021 How the Discovery of the CD4/CD8-p56lck Complexes Changed Immunology and Immunotherapy. Frontiers in cell and developmental biology 40 33791292
2011 The RPT2 subunit of the 26S proteasome directs complex assembly, histone dynamics, and gametophyte and sporophyte development in Arabidopsis. The Plant cell 40 22158466
2020 TreeSummarizedExperiment: a S4 class for data with hierarchical structure. F1000Research 39 33274053
2003 Discovery and initial SAR of 2-amino-5-carboxamidothiazoles as inhibitors of the Src-family kinase p56(Lck). Bioorganic & medicinal chemistry letters 38 14592495
2001 A potential role for protein tyrosine kinase p56(lck) in rheumatoid arthritis synovial fluid T lymphocyte hyporesponsiveness. International immunology 38 11222499
1999 Volume expansion stimulates p72(syk) and p56(lyn) in skate erythrocytes. The Journal of biological chemistry 38 10075687
1997 Sequestration of p56(lck) by gp120, a model for TCR desensitization. Journal of immunology (Baltimore, Md. : 1950) 38 9036944
1982 Ribosomal protein S4 acts in trans as a translational repressor to regulate expression of the alpha operon in Escherichia coli. Journal of bacteriology 37 6211432
2016 Enriching s4 U-RNA Using Methane Thiosulfonate (MTS) Chemistry. Current protocols in chemical biology 35 27925666
2011 The hERG K(+) channel S4 domain L532P mutation: characterization at 37°C. Biochimica et biophysica acta 35 21777565
2004 A theoretical study of spin states in Ni-S4 complexes and models of the [NiFe] hydrogenase active site. Journal of biological inorganic chemistry : JBIC : a publication of the Society of Biological Inorganic Chemistry 34 15365900
1994 p56/p53lyn tyrosine kinase activation in mammalian cells treated with mitomycin C. Oncogene 34 8084605
2001 Recognition of 16S rRNA by ribosomal protein S4 from Bacillus stearothermophilus. Biochemistry 33 11401563
1999 HIV-1 Nef plays an essential role in two independent processes in CD4 down-regulation: dissociation of the CD4-p56(lck) complex and targeting of CD4 to lysosomes. Virology 33 10208934
2008 Structure and function of colicin S4, a colicin with a duplicated receptor-binding domain. The Journal of biological chemistry 31 19056731
1978 Electron microscopic determination of the binding sites of ribosomal proteins S4 and S8 on 16S RNA. Proceedings of the National Academy of Sciences of the United States of America 31 343105
1996 Deficient expression of p56(lck) in Th2 cells leads to partial TCR signaling and a dysregulation in lymphokine mRNA levels. Journal of immunology (Baltimore, Md. : 1950) 30 8943376
1995 A mouse homologue of the Xenopus germ cell-specific ribonucleic acid/deoxyribonucleic acid-binding proteins p54/p56 interacts with the protamine 2 promoter. Biology of reproduction 30 7756448
1981 Tissue specificity and cellular distribution of Novikoff hepatoma antigenic proteins p39, p49, and p56. The Journal of biological chemistry 30 7263643
2020 A Salmochelin S4-Inspired Ciprofloxacin Trojan Horse Conjugate. ACS infectious diseases 29 32786274
2015 Characterization of equine urinary metabolites of selective androgen receptor modulators (SARMs) S1, S4 and S22 for doping control purposes. Drug testing and analysis 29 25560998
2012 Role of RPT2 in leaf positioning and flattening and a possible inhibition of phot2 signaling by phot1. Plant & cell physiology 28 22739508
1999 Inhibition of p56(lck) tyrosine kinase by isothiazolones. Archives of biochemistry and biophysics 28 10087161
1995 1-beta-D-arabinofuranosylcytosine activates tyrosine phosphorylation of p34cdc2 and its association with the Src-like p56/p53lyn kinase in human myeloid leukemia cells. Biochemistry 28 7530044
1981 Apparent association constants for E. coli ribosomal proteins S4, S7, S8, S15, S17 and S20 binding to 16S RNA. Nucleic acids research 28 7029472
1997 The cytoplasmic domain of rat NKR-P1 receptor interacts with the N-terminal domain of p56(lck) via cysteine residues. European journal of immunology 27 9022000
2015 Roles of reactive oxygen species-degrading enzymes of Francisella tularensis SCHU S4. Infection and immunity 26 25802058
1994 The conformational analysis of a synthetic S4 peptide corresponding to a voltage-gated potassium ion channel protein. FEBS letters 26 8050599
1987 Biosynthesis of reovirus-specified polypeptides. Efficiency of expression of cDNAs of the reovirus S1 and S4 genes in transfected animal cells differs at the level of translation. Virology 26 3617502
1973 Alteration of ribosomal protein S4 by mutation linked to kasugamycin-resistance in Escherichia coli. Proceedings of the National Academy of Sciences of the United States of America 26 4567337
2012 TCR-induced T cell activation leads to simultaneous phosphorylation at Y505 and Y394 of p56(lck) residues. Cytometry. Part A : the journal of the International Society for Analytical Cytology 25 22674786
2002 Discovery and initial SAR of imidazoquinoxalines as inhibitors of the Src-family kinase p56(Lck). Bioorganic & medicinal chemistry letters 25 11992777
1997 Targets of p56(lck) activity in immature thymoblasts: stimulation of the Ras/Raf/MAPK pathway. International immunology 25 9040011
2004 Involvement of tyrosine kinase p56/Lck in apoptosis induction by anticancer drugs. Biochemical pharmacology 24 15130763
1995 Messenger RNA recognition by fragments of ribosomal protein S4. The Journal of biological chemistry 24 7559430
2015 N-Myristoylation of the Rpt2 subunit of the yeast 26S proteasome is implicated in the subcellular compartment-specific protein quality control system. Journal of proteomics 23 26344132
2014 Modulation of decoding fidelity by ribosomal proteins S4 and S5. Journal of bacteriology 23 25548247
2011 Slow formation of stable complexes during coincubation of minimal rRNA and ribosomal protein S4. Journal of molecular biology 23 21821049
2009 Discovery of okilactomycin and congeners from Streptomyces scabrisporus by antisense differential sensitivity assay targeting ribosomal protein S4. The Journal of antibiotics 23 19132063
1999 p56(lck), ZAP-70, SLP-76, and calcium-regulated effectors are involved in NF-kappaB activation by bisperoxovanadium phosphotyrosyl phosphatase inhibitors in human T cells. The Journal of biological chemistry 23 10574981
2000 A novel mutation in ribosomal protein S4 that affects the function of a mutated RF1. Biochimie 22 11018284
2000 Transforming growth factor-beta is an autocrine mitogen for a novel androgen-responsive murine prostatic smooth muscle cell line, PSMC1. Journal of cellular physiology 22 11056012
1999 Refining the overall structure and subdomain orientation of ribosomal protein S4 delta41 with dipolar couplings measured by NMR in uniaxial liquid crystalline phases. Journal of molecular biology 22 10493882
1999 Protein tyrosine kinase p53/p56(lyn) forms complexes with gamma-tubulin in rat basophilic leukemia cells. International immunology 22 10545487
2021 Conjugation to Enterobactin and Salmochelin S4 Enhances the Antimicrobial Activity and Selectivity of β-Lactam Antibiotics against Nontyphoidal Salmonella. ACS infectious diseases 20 33691061
2012 Structural interactions between lipids, water and S1-S4 voltage-sensing domains. Journal of molecular biology 20 22858867
2009 A minimized rRNA-binding site for ribosomal protein S4 and its implications for 30S assembly. Nucleic acids research 20 19190093
2007 Novel repressor of the human FMR1 gene - identification of p56 human (GCC)(n)-binding protein as a Krüppel-like transcription factor ZF5. The FEBS journal 20 17714511
2009 Mycoreovirus 1 S4-coded protein is dispensable for viral replication but necessary for efficient vertical transmission and normal symptom induction. Virology 18 20006368
1998 Biochemical analysis of the CD45-p56(lck) complex in Jurkat T cells lacking expression of lymphocyte phosphatase-associated phosphoprotein. International immunology 18 9533446
2018 Plasmodium yoelii S4/CelTOS is important for sporozoite gliding motility and cell traversal. Cellular microbiology 17 29253313
2018 Degradation and detoxification of azo dyes by a salt-tolerant yeast Cyberlindnera samutprakarnensis S4 under high-salt conditions. World journal of microbiology & biotechnology 17 30105649
2019 Hi-C detects novel structural variants in HL-60 and HL-60/S4 cell lines. Genomics 16 31095996