Affinage

RPS2

Small ribosomal subunit protein uS5 · UniProt P15880

Round 2 corrected
Length
293 aa
Mass
31.3 kDa
Annotated
2026-04-28
130 papers in source corpus 13 papers cited in narrative 13 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

RPS2 (uS5) is a universally conserved structural protein of the 40S small ribosomal subunit that plays essential roles in both ribosome biogenesis and extraribosomal signaling. Its incorporation into pre-40S particles requires dedicated co-translational chaperones—Tsr4 in budding yeast and PDCD2 in metazoans—that bind nascent uS5 in the cytoplasm and escort it to nuclear assembly sites, and its depletion blocks pre-rRNA processing and nuclear export of pre-40S subunits (PMID:18820293, PMID:31182640, PMID:33245768). In the cytoplasm, uS5 is arginine-methylated by PRMT3, a modification that cannot be compensated by other methyltransferases, and PRMT3 and ZNF277 compete for binding to extraribosomal uS5 through structurally related zinc-finger domains (PMID:17439947, PMID:30530495). Under ribosomal stress, free uS5 binds the MDM2 RING domain to suppress its E3 ligase activity and stabilize p53—a response counter-regulated by the deubiquitinase USP47, which removes MDM2-mediated ubiquitin from uS5 under homeostatic conditions—and uS5 also directly binds pre-let-7a-1 RNA to block miRNA maturation (PMID:31928715, PMID:32370049, PMID:21148031).

Mechanistic history

Synthesis pass · year-by-year structured walk · 8 steps
  1. 1992 Medium

    Molecular cloning of human RPS2 established it as a conserved ribosomal protein with elevated expression in transformed cells, providing the sequence identity needed for subsequent functional studies.

    Evidence Differential cDNA screening of ras-transformed human tumor cells with DNA sequencing

    PMID:1586449

    Open questions at the time
    • No mechanistic link between RPS2 expression and transformation was established
    • Expression correlation does not demonstrate causality
  2. 2007 High

    Identification of RPS2 as the dedicated in vivo substrate of PRMT3 resolved how cytoplasmic arginine methylation is targeted to a specific ribosomal protein, showing that no other PRMT compensates for PRMT3 loss.

    Evidence PRMT3-knockout mice with mass spectrometry-based methylation analysis and ribosome fractionation

    PMID:17439947

    Open questions at the time
    • Functional consequence of RPS2 methylation on translation remained undefined
    • Whether methylation affects RPS2 incorporation into 40S subunits was not tested
  3. 2008 High

    Depletion of the fission yeast ortholog revealed that uS5 is required for pre-rRNA processing at site A2 and for nuclear export competence of pre-40S particles, establishing its essential role in 40S biogenesis beyond mere structural occupancy.

    Evidence Conditional Rps2 depletion in S. pombe with pulse-chase rRNA labeling, Northern blotting, and fluorescence microscopy

    PMID:18820293

    Open questions at the time
    • Which assembly factors depend on Rps2 presence for recruitment was not determined
    • Whether the human ortholog shows identical processing defects upon depletion was not tested
  4. 2010 Medium

    Discovery that extraribosomal RPS2 directly binds pre-let-7a-1 RNA and blocks its maturation revealed an unexpected non-ribosomal function linking RPS2 to miRNA-mediated gene regulation and tumorigenesis.

    Evidence EMSA, antibody supershift, co-immunoprecipitation, RPS2 overexpression/knockdown in prostate cancer cells, and SCID mouse xenograft

    PMID:21148031

    Open questions at the time
    • Structural basis of the RPS2–pre-let-7a-1 interaction is unknown
    • Whether RPS2 regulates other pre-miRNAs beyond pre-let-7a-1 was not explored
    • Single-lab finding without independent replication
  5. 2013 High

    Cryo-EM structures of human and Drosophila 80S ribosomes placed uS5 at atomic resolution within the 40S subunit, defining its rRNA contacts and metazoan-specific structural features.

    Evidence High-resolution cryo-EM with atomic model building

    PMID:23636399 PMID:25901680

    Open questions at the time
    • Structures captured the mature ribosome; uS5 contacts in pre-40S intermediates remained unresolved
  6. 2018 High

    Identification of ZNF277 as a competing extraribosomal partner of uS5 that uses the same zinc-finger architecture as PRMT3 revealed a regulatory network governing the fate of free uS5 in the cytoplasm and nucleolus.

    Evidence Quantitative AP-MS, co-IP, bimolecular fluorescence complementation, competition assays with PRMT3 overexpression/depletion

    PMID:30530495

    Open questions at the time
    • Functional consequence of ZNF277–uS5 complex formation on ribosome biogenesis or translation is unclear
    • Whether ZNF277 modulates PRMT3-dependent methylation of uS5 physiologically remains open
  7. 2019 High

    Discovery that Tsr4 co-translationally captures Rps2 via its eukaryote-specific N-terminal extension and escorts it to nuclear assembly established the first dedicated chaperone pathway for uS5 biogenesis.

    Evidence Co-IP, ribosome fractionation, genetic depletion, polysome profiling, and fluorescence microscopy in S. cerevisiae

    PMID:31182640

    Open questions at the time
    • How the Tsr4–Rps2 complex is disassembled at the nuclear import step was not defined
  8. 2020 High

    Human PDCD2 was shown to be the functional ortholog of yeast Tsr4, confirming a conserved co-translational chaperone mechanism for uS5 from yeast to mammals, while RPS2 was simultaneously linked to the ribosomal stress–p53 pathway through MDM2 binding and to its counter-regulation by USP47.

    Evidence AP-MS, siRNA knockdown, pulse-chase labeling for PDCD2 chaperone function; co-IP, ubiquitination assays, siRNA, and xenograft models for MDM2/USP47 axis

    PMID:31928715 PMID:32370049 PMID:33245768

    Open questions at the time
    • Structural basis of the PDCD2–uS5 or MDM2–uS5 interface is lacking
    • How the pool of free uS5 is partitioned between the PDCD2 chaperone pathway and the MDM2-p53 signaling axis is unknown
    • Whether USP47 regulation of RPS2 ubiquitination has been independently replicated is unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the structural basis of uS5 recognition by its chaperones and signaling partners, the functional role of PRMT3-mediated arginine methylation on translation fidelity, and how cells partition extraribosomal uS5 among its multiple non-ribosomal functions.
  • No reconstitution of the PDCD2–uS5 handoff to pre-40S particles
  • Functional significance of uS5 methylation for translational accuracy untested in mammalian cells
  • Physiological conditions under which ZNF277 versus PRMT3 versus MDM2 pathways dominate are undefined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005198 structural molecule activity 3 GO:0098772 molecular function regulator activity 2 GO:0003723 RNA binding 1
Localization
GO:0005840 ribosome 4 GO:0005829 cytosol 3 GO:0005634 nucleus 2 GO:0005730 nucleolus 2
Pathway
R-HSA-392499 Metabolism of proteins 6 GO:0005840 ribosome 3 R-HSA-5357801 Programmed Cell Death 2 R-HSA-8953854 Metabolism of RNA 2
Partners
MDM2PDCD2PRMT3USP47ZNF277
Complex memberships
40S small ribosomal subunit80S ribosome

Evidence

Reading pass · 13 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1992 Human ribosomal protein S2 (RPS2) mRNA was identified as elevated in ras-transformed human tumor cells. DNA sequence analysis confirmed this clone encodes the human ribosomal S2 protein, closely related to the yeast omnipotent suppressor SUP44 (yeast ribosomal protein S4) and mouse LLRep3, establishing RPS2 as a conserved ribosomal protein with potential roles in oncogenesis. Differential cDNA library screening, DNA sequencing, in situ hybridization Molecular carcinogenesis Medium 1586449
2007 PRMT3, a cytoplasmic type I arginine methyltransferase, methylates ribosomal protein rpS2 (RPS2) in vivo. In PRMT3-knockout mice, rpS2 is hypomethylated, demonstrating that rpS2 is a bona fide, dedicated in vivo substrate of PRMT3 that cannot be compensated by other PRMTs. PRMT3 is tethered to ribosomes through its interaction with rpS2. PRMT3-deficient mice display Minute-like characteristics (small embryo size) but survive to adulthood, and polyribosome profiles are unaffected. Targeted gene disruption (knockout mouse), mass spectrometry, ribosome fractionation, co-immunoprecipitation The Journal of biological chemistry High 17439947
2008 In fission yeast (Schizosaccharomyces pombe), Rps2 (ortholog of human RPS2) is essential for cell viability and production of 40S ribosomal subunits. Depletion of Rps2 blocks pre-rRNA processing at site A2 within 32S pre-rRNA, causing accumulation of 21S rRNA precursors. Pre-40S particles formed in the absence of Rps2 are retained in the nucleolus, revealing that Rps2 is required for nuclear export competence of pre-40S subunits. Conditional gene depletion, pulse-chase rRNA labeling, Northern blotting, fluorescence microscopy Nucleic acids research High 18820293
2009 In budding yeast Saccharomyces cerevisiae, Rps2 carries asymmetric dimethylarginine and monomethylarginine modifications. Despite lacking a zinc-finger-containing Rmt3/PRMT3 homolog, arginine methylation of Rps2 is dependent on the major arginine methyltransferase Rmt1, revealing that different organisms use distinct methyltransferases to modify Rps2. Genetic deletion of Rmt1, mass spectrometry detection of methylarginine, biochemical fractionation Biochemical and biophysical research communications Medium 20035717
2010 Human RPS2 specifically binds pre-let-7a-1 RNA, blocking its processing to mature let-7a/let-7f miRNA. Electrophoretic mobility shift assays, antibody supershift assays, and co-immunoprecipitation demonstrated direct RPS2–pre-let-7a-1 interaction. RPS2 overexpression suppresses let-7a levels and enhances colony-forming and invasive activity of prostate cancer cells; RPS2 knockdown reduces tumorigenesis in SCID mice. EMSA, antibody supershift assay, co-immunoprecipitation, stable transfection, shRNA knockdown, xenograft mouse model Molecular cancer research : MCR Medium 21148031
2018 Human uS5 (RPS2) forms an extraribosomal complex with ZNF277, a conserved zinc finger protein, in the cytoplasm and nucleolus. ZNF277 uses a C2H2-type zinc finger domain to recognize uS5, the same domain architecture used by PRMT3. ZNF277 and PRMT3 compete for uS5 binding: overexpression of PRMT3 inhibits ZNF277-uS5 complex formation and vice versa. ZNF277 recognizes nascent uS5 co-translationally, suggesting co-translational assembly of the ZNF277-uS5 complex. Quantitative proteomics (AP-MS), co-immunoprecipitation, bimolecular fluorescence complementation, ribosome fractionation, overexpression and depletion experiments The Journal of biological chemistry High 30530495
2019 Tsr4 is a dedicated cytoplasmic chaperone for Rps2 (uS5) in Saccharomyces cerevisiae. Tsr4 associates co-translationally with Rps2, requiring the eukaryote-specific N-terminal extension of Rps2 for this interaction. Perturbation of Tsr4 results in decreased Rps2 protein levels and phenocopies Rps2 depletion (40S biogenesis defects). Tsr4 is restricted to the cytoplasm despite Rps2 ultimately joining nuclear pre-40S particles, establishing Tsr4 as a cytoplasmic chaperone that hands off Rps2 for nuclear assembly. Co-immunoprecipitation, ribosome fractionation, genetic depletion, polysome profiling, fluorescence microscopy Molecular and cellular biology High 31182640
2020 Human PDCD2 (ortholog of yeast Tsr4 and Drosophila zfrp8) functions as a dedicated ribosomal protein chaperone for uS5 (RPS2). PDCD2 specifically interacts with uS5 and the PDCD2-uS5 complex is assembled co-translationally. Loss of PDCD2 causes defects in 40S ribosomal subunit synthesis that phenocopy uS5 deficiency. PDCD2 is important for the accumulation of soluble uS5 protein and its incorporation into 40S subunits, and accompanies uS5 from cytoplasmic translation sites to ribosome assembly sites in the nucleus. Quantitative proteomics (AP-MS), ribosome fractionation, siRNA knockdown, pulse-chase labeling, fluorescence microscopy Nucleic acids research High 33245768
2020 RPS2 interacts with MDM2 through the RING finger domain of MDM2. MDM2 ubiquitinates RPS2, and the ubiquitination status of RPS2 regulates p53 protein stability. Under ribosomal stress, RPS2 binds MDM2 to suppress MDM2 E3 ligase activity, leading to p53 stabilization. RPS2 knockdown prevents p53 induction even under ribosomal stress conditions, demonstrating that RPS2 is essential for the ribosomal stress–p53 response. Co-immunoprecipitation, ubiquitination assay, siRNA knockdown, Western blotting, stress induction experiments Biochemical and biophysical research communications Medium 31928715
2020 USP47 (Ubiquitin Specific Protease 47) deubiquitinates RPS2 that has been ubiquitinated by MDM2. USP47 inhibits the RPS2-MDM2 interaction under normal conditions, alleviating RPS2-mediated MDM2 suppression and keeping p53 levels low. Under ribosomal stress, dissociation of USP47 allows RPS2 to bind and suppress MDM2, inducing p53. Depletion of USP47 inhibits cell proliferation, colony formation, and tumor progression in xenograft models. Co-immunoprecipitation, ubiquitination assay, siRNA knockdown, xenograft mouse model, colony formation assay Cancers Medium 32370049
2016 In Drosophila, Zfrp8/PDCD2 (ortholog of human PDCD2) interacts directly with RpS2 (uS5) of the 40S small ribosomal subunit. Zfrp8/PDCD2 knockdown in ovaries leads to nuclear accumulation of specific mRNAs and TE transcripts. Zfrp8/PDCD2 regulates cytoplasmic levels of small (40S) ribosomal subunit components but does not control nucleolar localization of ribosomal proteins, suggesting a role at late stages of ribosome assembly and in selective mRNA translation. Co-immunoprecipitation, RNAi knockdown, fluorescence imaging of ribosomal protein distribution, RNA analysis PloS one Medium 26807849
2013 Cryo-EM structures of the human and Drosophila 80S ribosomes revealed the position of uS5 (RPS2) as a component of the small (40S) ribosomal subunit, showing its co-evolution with metazoan-specific ribosomal RNA and the presence of metazoan-specific structural layers in the ribosome. High-resolution cryo-electron microscopy, atomic model building Nature High 23636399
2015 The near-atomic structure of the human 80S ribosome (3.6 Å average, 2.9 Å in stable regions) by cryo-EM defined the precise position and interactions of RPS2 (uS5) within the 40S subunit, providing atomic-level detail of ribosomal RNA contacts and amino acid side chains. Single-particle cryo-electron microscopy, atomic model building Nature High 25901680

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Global, in vivo, and site-specific phosphorylation dynamics in signaling networks. Cell 2861 17081983
2012 Insights into RNA biology from an atlas of mammalian mRNA-binding proteins. Cell 1718 22658674
2005 A human protein-protein interaction network: a resource for annotating the proteome. Cell 1704 16169070
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2006 A probability-based approach for high-throughput protein phosphorylation analysis and site localization. Nature biotechnology 1336 16964243
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
2012 The mRNA-bound proteome and its global occupancy profile on protein-coding transcripts. Molecular cell 973 22681889
2005 Nucleolar proteome dynamics. Nature 934 15635413
2020 A reference map of the human binary protein interactome. Nature 849 32296183
2018 VIRMA mediates preferential m6A mRNA methylation in 3'UTR and near stop codon and associates with alternative polyadenylation. Cell discovery 829 29507755
2003 Arabidopsis RIN4 is a target of the type III virulence effector AvrRpt2 and modulates RPS2-mediated resistance. Cell 721 12581527
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
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2020 Structural basis for translational shutdown and immune evasion by the Nsp1 protein of SARS-CoV-2. Science (New York, N.Y.) 653 32680882
2003 Initiation of RPS2-specified disease resistance in Arabidopsis is coupled to the AvrRpt2-directed elimination of RIN4. Cell 616 12581526
2018 High-Density Proximity Mapping Reveals the Subcellular Organization of mRNA-Associated Granules and Bodies. Molecular cell 580 29395067
2017 Anticancer sulfonamides target splicing by inducing RBM39 degradation via recruitment to DCAF15. Science (New York, N.Y.) 533 28302793
2011 Analysis of the myosin-II-responsive focal adhesion proteome reveals a role for β-Pix in negative regulation of focal adhesion maturation. Nature cell biology 490 21423176
2013 Structures of the human and Drosophila 80S ribosome. Nature 481 23636399
2020 SARS-CoV-2 Nsp1 binds the ribosomal mRNA channel to inhibit translation. Nature structural & molecular biology 466 32908316
2020 SARS-CoV-2 Disrupts Splicing, Translation, and Protein Trafficking to Suppress Host Defenses. Cell 449 33080218
2004 The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). Genome research 438 15489334
2022 OpenCell: Endogenous tagging for the cartography of human cellular organization. Science (New York, N.Y.) 432 35271311
2010 Global analysis of TDP-43 interacting proteins reveals strong association with RNA splicing and translation machinery. Journal of proteome research 422 20020773
2013 The intracellular interactome of tetraspanin-enriched microdomains reveals their function as sorting machineries toward exosomes. The Journal of biological chemistry 413 23463506
2015 Panorama of ancient metazoan macromolecular complexes. Nature 407 26344197
2004 The molecular mechanics of eukaryotic translation. Annual review of biochemistry 396 15189156
2004 Proteomic, functional, and domain-based analysis of in vivo 14-3-3 binding proteins involved in cytoskeletal regulation and cellular organization. Current biology : CB 386 15324660
2015 Structure of the human 80S ribosome. Nature 380 25901680
2003 HSP90 interacts with RAR1 and SGT1 and is essential for RPS2-mediated disease resistance in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America 358 14504384
1996 Isolation of Arabidopsis genes that differentiate between resistance responses mediated by the RPS2 and RPM1 disease resistance genes. The Plant cell 228 8742710
2006 Double-stranded RNA is internalized by scavenger receptor-mediated endocytosis in Drosophila S2 cells. The Journal of biological chemistry 205 16531407
1999 Herpes simplex virus inhibits apoptosis through the action of two genes, Us5 and Us3. Journal of virology 189 10516000
1986 Typical and atypical antipsychotic occupancy of D2 and S2 receptors: an autoradiographic analysis in rat brain. Brain research bulletin 170 2872945
2000 Mutational analysis of the Arabidopsis nucleotide binding site-leucine-rich repeat resistance gene RPS2. The Plant cell 141 11148296
2023 Broadly neutralizing anti-S2 antibodies protect against all three human betacoronaviruses that cause deadly disease. Immunity 140 36889306
2010 Expression in aneuploid Drosophila S2 cells. PLoS biology 139 20186269
1984 Serotonin-S2 receptor binding sites and functional correlates. Neuropharmacology 136 6396526
2003 Natural selection for polymorphism in the disease resistance gene Rps2 of Arabidopsis thaliana. Genetics 133 12618410
2005 Molecular basis for the RIN4 negative regulation of RPS2 disease resistance. The Plant cell 126 15749765
1996 Molecular recognition of pathogen attack occurs inside of plant cells in plant disease resistance specified by the Arabidopsis genes RPS2 and RPM1. Proceedings of the National Academy of Sciences of the United States of America 125 8986840
2005 Cell cycle-dependent dynamics and regulation of mitotic kinesins in Drosophila S2 cells. Molecular biology of the cell 119 15958489
1985 Pharmacology of potent and selective S2-serotonergic antagonists. Journal of cardiovascular pharmacology 118 2412048
2007 Ribosomal protein rpS2 is hypomethylated in PRMT3-deficient mice. The Journal of biological chemistry 113 17439947
2015 Enhanced immune response to hepatitis B vaccination through immunization with a Pre-S1/Pre-S2/S vaccine. Medical microbiology and immunology 110 25557605
1999 Diversity and molecular evolution of the RPS2 resistance gene in Arabidopsis thaliana. Proceedings of the National Academy of Sciences of the United States of America 110 9874813
2021 The Case for S2: The Potential Benefits of the S2 Subunit of the SARS-CoV-2 Spike Protein as an Immunogen in Fighting the COVID-19 Pandemic. Frontiers in immunology 106 33767706
2004 Expression of SCC-S2, an antiapoptotic molecule, correlates with enhanced proliferation and tumorigenicity of MDA-MB 435 cells. Oncogene 104 14724590
2022 SARS-CoV-2 spike engagement of ACE2 primes S2' site cleavage and fusion initiation. Proceedings of the National Academy of Sciences of the United States of America 100 34930824
2022 From delta to Omicron: S1-RBD/S2 mutation/deletion equilibrium in SARS-CoV-2 defined variants. Gene 100 34990799
2005 A duplicated pair of Arabidopsis RING-finger E3 ligases contribute to the RPM1- and RPS2-mediated hypersensitive response. The Plant journal : for cell and molecular biology 98 16212605
1993 Arabidopsis mutations at the RPS2 locus result in loss of resistance to Pseudomonas syringae strains expressing the avirulence gene avrRpt2. Molecular plant-microbe interactions : MPMI 98 8400373
1982 Cloning the double-stranded RNA genes of reovirus: sequence of the cloned S2 gene. Proceedings of the National Academy of Sciences of the United States of America 98 6961439
2014 Pathway for Mn-cluster oxidation by tyrosine-Z in the S2 state of photosystem II. Proceedings of the National Academy of Sciences of the United States of America 94 24889635
2004 SNS: Adhesive properties, localization requirements and ectodomain dependence in S2 cells and embryonic myoblasts. Mechanisms of development 89 15511638
1988 The state of cluster SH and S2- of aconitase during cluster interconversions and removal. A convenient preparation of apoenzyme. The Journal of biological chemistry 88 2836417
2011 Physical association of Arabidopsis hypersensitive induced reaction proteins (HIRs) with the immune receptor RPS2. The Journal of biological chemistry 81 21757708
2008 Strong correlation between SHAPE chemistry and the generalized NMR order parameter (S2) in RNA. Journal of the American Chemical Society 81 18710236
1993 Molecular structures and functions of pyocins S1 and S2 in Pseudomonas aeruginosa. Journal of bacteriology 74 8491711
1984 Loxapine and clozapine decrease serotonin (S2) but do not elevate dopamine (D2) receptor numbers in the rat brain. Psychiatry research 74 6239298
2015 Reciprocal interareal connections to corticospinal neurons in mouse M1 and S2. The Journal of neuroscience : the official journal of the Society for Neuroscience 68 25698734
2011 Gallic acid inhibits the migration and invasion of A375.S2 human melanoma cells through the inhibition of matrix metalloproteinase-2 and Ras. Melanoma research 65 21734530
2001 Physicochemical and immunological characterization of hepatitis B virus envelope particles exclusively consisting of the entire L (pre-S1 + pre-S2 + S) protein. Vaccine 63 11312011
1992 Elevated expression of the ribosomal protein S2 gene in human tumors. Molecular carcinogenesis 60 1586449
2021 BNT162b2 vaccine induces divergent B cell responses to SARS-CoV-2 S1 and S2. Nature immunology 58 34848871
2006 The Erwinia amylovora avrRpt2EA gene contributes to virulence on pear and AvrRpt2EA is recognized by Arabidopsis RPS2 when expressed in pseudomonas syringae. Molecular plant-microbe interactions : MPMI 57 16776298
2017 Human Cytomegalovirus MicroRNAs miR-US5-1 and miR-UL112-3p Block Proinflammatory Cytokine Production in Response to NF-κB-Activating Factors through Direct Downregulation of IKKα and IKKβ. mBio 56 28270578
2001 The leucine-rich repeat domain can determine effective interaction between RPS2 and other host factors in arabidopsis RPS2-mediated disease resistance. Genetics 55 11333251
2004 Copper homoeostasis in Drosophila melanogaster S2 cells. The Biochemical journal 54 15239669
2023 SARS-CoV-2 spike S2-specific neutralizing antibodies. Emerging microbes & infections 53 37254830
2017 Robotic Guidance for S2-Alar-Iliac Screws in Spinal Deformity Correction. Clinical spine surgery 52 28107243
2008 Genomic RNAi screening in Drosophila S2 cells: what have we learned about host-pathogen interactions? Current opinion in microbiology 50 18539520
1995 The S2 subunit of the murine coronavirus spike protein is not involved in receptor binding. Journal of virology 49 7474149
2012 The soluble pyocins S2 and S4 from Pseudomonas aeruginosa bind to the same FpvAI receptor. MicrobiologyOpen 45 23170226
2003 Inhibition of apoptosis by Z-VAD-fmk in SMN-depleted S2 cells. The Journal of biological chemistry 45 12783893
2011 Drosophila melanogaster S2 cells for expression of heterologous genes: From gene cloning to bioprocess development. Biotechnology advances 44 22079894
1995 Expression analysis of the individual bovine beta-, alpha s2- and kappa-casein genes in transgenic mice. The Biochemical journal 43 7487953
2003 Cloning of a Heliothis virescens 110 kDa aminopeptidase N and expression in Drosophila S2 cells. Insect biochemistry and molecular biology 40 12706629
2003 Intracellular regulation of evodiamine-induced A375-S2 cell death. Biological & pharmaceutical bulletin 38 14600398
1994 Sequence diversity within the reovirus S2 gene: reovirus genes reassort in nature, and their termini are predicted to form a panhandle motif. Journal of virology 38 8289378
2019 The retroviral accessory proteins S2, Nef, and glycoMA use similar mechanisms for antagonizing the host restriction factor SERINC5. The Journal of biological chemistry 37 30862674
2015 Down-regulation of deacetylase HDAC6 inhibits the melanoma cell line A375.S2 growth through ROS-dependent mitochondrial pathway. PloS one 36 25774669
2012 Triptolide induced DNA damage in A375.S2 human malignant melanoma cells is mediated via reduction of DNA repair genes. Oncology reports 35 23233170
2011 Proteomics analyses of microvesicles released by Drosophila Kc167 and S2 cells. Proteomics 34 21901833
2011 The influence of ciprofloxacin on viability of A549, HepG2, A375.S2, B16 and C6 cell lines in vitro. Acta poloniae pharmaceutica 34 22125950
2020 Ptr1 evolved convergently with RPS2 and Mr5 to mediate recognition of AvrRpt2 in diverse solanaceous species. The Plant journal : for cell and molecular biology 32 32391580
1976 Identification of genes for elongation factor Ts and ribosomal protein S2 in E. coli. Cell 32 782723
2021 Human Cytomegalovirus UL7, miR-US5-1, and miR-UL112-3p Inactivation of FOXO3a Protects CD34+ Hematopoietic Progenitor Cells from Apoptosis. mSphere 31 33408225
2007 Characterization of Components of Partial Resistance, Rps2, and Root Resistance to Phytophthora sojae in Soybean. Phytopathology 31 18943586
1996 Acid xylanase from yeast Cryptococcus sp. S-2: purification, characterization, cloning, and sequencing. Bioscience, biotechnology, and biochemistry 31 8987553
2016 Timosaponin AIII induces apoptosis and autophagy in human melanoma A375-S2 cells. Archives of pharmacal research 29 27271334
2004 Norcantharidin induces human melanoma A375-S2 cell apoptosis through mitochondrial and caspase pathways. Journal of Korean medical science 29 15308848
2018 The 40S ribosomal protein uS5 (RPS2) assembles into an extraribosomal complex with human ZNF277 that competes with the PRMT3-uS5 interaction. The Journal of biological chemistry 28 30530495
2020 Human Cytomegalovirus miR-US5-2 Downregulation of GAB1 Regulates Cellular Proliferation and UL138 Expression through Modulation of Epidermal Growth Factor Receptor Signaling Pathways. mSphere 26 32759334
2006 Development of a virus-induced gene-silencing system for functional analysis of the RPS2-dependent resistance signalling pathways in Arabidopsis. Plant molecular biology 26 16845481
2006 Herpes simplex virus genes Us3, Us5, and Us12 differentially regulate cytotoxic T lymphocyte-induced cytotoxicity. Viral immunology 26 16987059
2012 SCC-S2 is overexpressed in colon cancers and regulates cell proliferation. Tumour biology : the journal of the International Society for Oncodevelopmental Biology and Medicine 25 22886548
1985 Complete sequence of ovine alpha s2-casein messenger RNA. Biochimie 25 3002499
1984 Purification and characterization of Serratia marcescens US5 pili. Infection and immunity 24 6209220
2019 Tsr4 Is a Cytoplasmic Chaperone for the Ribosomal Protein Rps2 in Saccharomyces cerevisiae. Molecular and cellular biology 23 31182640
2011 Cloning, purification, and functional characterization of Carocin S2, a ribonuclease bacteriocin produced by Pectobacterium carotovorum. BMC microbiology 23 21569432
2009 Rmt1 catalyzes zinc-finger independent arginine methylation of ribosomal protein Rps2 in Saccharomyces cerevisiae. Biochemical and biophysical research communications 23 20035717
2005 Dracorhodin perchlorate induces A375-S2 cell apoptosis via accumulation of p53 and activation of caspases. Biological & pharmaceutical bulletin 23 15684474
2004 Silymarin prevents UV irradiation-induced A375-S2 cell apoptosis. Biological & pharmaceutical bulletin 23 15256735
2020 USP47 Promotes Tumorigenesis by Negative Regulation of p53 through Deubiquitinating Ribosomal Protein S2. Cancers 22 32370049
1989 Mapping of linear B-cell epitopes of the S2 subunit of pertussis toxin. Infection and immunity 22 2463969
2022 Elucidating the biodegradation pathway and catabolic genes of benzophenone-3 in Rhodococcus sp. S2-17. Environmental pollution (Barking, Essex : 1987) 21 35085657
2020 PDCD2 functions as an evolutionarily conserved chaperone dedicated for the 40S ribosomal protein uS5 (RPS2). Nucleic acids research 21 33245768
2013 Organelle transport in cultured Drosophila cells: S2 cell line and primary neurons. Journal of visualized experiments : JoVE 21 24300413
2012 AvrRpm1 missense mutations weakly activate RPS2-mediated immune response in Arabidopsis thaliana. PloS one 21 22880057
2010 Preparation of Drosophila S2 cells for light microscopy. Journal of visualized experiments : JoVE 21 20543772
2007 Critical roles for the M3-S2 transduction linker domain in kainate receptor assembly and postassembly trafficking. The Journal of neuroscience : the official journal of the Society for Neuroscience 21 17898214
2002 A Drosophila melanogaster cell line (S2) facilitates post-genome functional analysis of receptors and ion channels. BioEssays : news and reviews in molecular, cellular and developmental biology 21 12386938
2008 Nuclear export competence of pre-40S subunits in fission yeast requires the ribosomal protein Rps2. Nucleic acids research 20 18820293
2002 Plant mitochondrial rps2 genes code for proteins with a C-terminal extension that is processed. Plant molecular biology 20 12369627
2024 Discovery and characterization of a pan-betacoronavirus S2-binding antibody. Structure (London, England : 1993) 18 39326419
2004 Induction and biochemical purification of RNA-induced silencing complex from Drosophila S2 cells. Methods in molecular biology (Clifton, N.J.) 18 15103068
2020 Ribosomal protein S2 interplays with MDM2 to induce p53. Biochemical and biophysical research communications 17 31928715
2018 A Glycine soja group S2 bZIP transcription factor GsbZIP67 conferred bicarbonate alkaline tolerance in Medicago sativa. BMC plant biology 17 30316294
2015 Immune responses and protection induced by Brucella suis S2 bacterial ghosts in mice. Veterinary immunology and immunopathology 17 26022514
1993 Co-transcription pattern of an introgressed operon in the maize chloroplast genome comprising four ATP synthase subunit genes and the ribosomal rps2. Plant molecular biology 17 8490127
1990 Rat brain monoamine and Serotonin S2 receptor changes during pregnancy. Neurochemical research 17 2077430
2005 Assaying the Drosophila negative feedback loop with RNA interference in S2 cells. Methods in enzymology 16 15817315
2023 Ribosomal Protein uS5 and Friends: Protein-Protein Interactions Involved in Ribosome Assembly and Beyond. Biomolecules 15 37238722
2019 Molecular characterization and antiapoptotic function analysis of the duck plague virus Us5 gene. Scientific reports 15 30890748
2016 Zfrp8/PDCD2 Interacts with RpS2 Connecting Ribosome Maturation and Gene-Specific Translation. PloS one 15 26807849
2010 Role of ribosomal protein RPS2 in controlling let-7a expression in human prostate cancer. Molecular cancer research : MCR 15 21148031