Affinage

RPL5

Large ribosomal subunit protein uL18 · UniProt P46777

Length
297 aa
Mass
34.4 kDa
Annotated
2026-04-28
100 papers in source corpus 25 papers cited in narrative 25 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

RPL5 (uL18) is a core component of the 60S ribosomal subunit that serves dual roles in ribosome biogenesis and p53-dependent tumor suppression. As a ribosomal protein, RPL5 binds 5S rRNA through its N-terminal domain to form a ~7S extraribosomal RNP that is chaperoned by nucleophosmin (NPM/B23) via CRM1-dependent export for incorporation into nascent 60S subunits, and it anchors peptidyl-tRNA at the ribosomal P-site to maintain translational reading frame fidelity (PMID:3279045, PMID:16648475, PMID:11497428). Under ribosomal biogenesis stress, RPL5 and RPL11 are mutually stabilized against proteasomal degradation, accumulate in the ribosome-free fraction, and cooperatively inhibit MDM2 E3 ubiquitin ligase activity toward p53, thereby activating p53-dependent cell cycle arrest; RPL5 additionally activates TAp73 independently of MDM2 and recruits the RISC complex to c-Myc mRNA for its degradation (PMID:23169665, PMID:15308643, PMID:25301064, PMID:24141778). Cancer-associated somatic RPL5 mutations disrupt HDM2 binding and impair this checkpoint, and HEATR3 variants that block RPL5 nuclear import cause Diamond-Blackfan anemia-like phenotypes (PMID:32108164, PMID:35213692).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 1988 High

    Identifying RPL5 as the protein component of the extraribosomal 5S rRNA–protein complex established it as a key assembly intermediate in ribosome biogenesis, resolving the identity of the ~7S pre-ribosomal RNP.

    Evidence Autoantibody immunoprecipitation, pulse-chase, immunofluorescence, and sucrose gradient sedimentation in human cells

    PMID:3279045

    Open questions at the time
    • Chaperone pathway for 5S RNP assembly not yet identified
    • Stoichiometry and order of incorporation into pre-60S particles unknown
  2. 1994 High

    Discovery that RPL5 co-purifies with MDM2 and p53 revealed the first link between a ribosomal protein and the p53 regulatory pathway, opening the field of ribosomal stress signaling.

    Evidence Co-purification, reciprocal co-IP with multiple antibodies, N-terminal sequencing from human cells

    PMID:7935455

    Open questions at the time
    • Functional consequence of RPL5–MDM2 interaction on p53 stability unknown
    • Whether the interaction is direct or bridged by 5S rRNA unclear
  3. 1996 High

    Mapping RPL5's modular domain architecture—N-terminal 5S rRNA-binding domain and C-terminal nucleolar localization domain—established how a single protein coordinates RNA binding with subnuclear targeting.

    Evidence In vitro RNA-binding assay with deletion mutants, transfection and immunofluorescence in human cells

    PMID:10766838 PMID:8626719

    Open questions at the time
    • Structural basis of 5S rRNA recognition unresolved at atomic level
    • Signals governing cytoplasmic re-export after nucleolar assembly not mapped
  4. 2001 High

    Yeast rpl5 mutant analysis demonstrated that RPL5 anchors peptidyl-tRNA at the P-site, providing the first ribosomal function beyond structural scaffolding—maintaining translational reading frame fidelity.

    Evidence Frameshifting reporter assays, peptidyl-tRNA binding biochemistry, sparsomycin rescue in S. cerevisiae rpl5 mutants

    PMID:11497428

    Open questions at the time
    • Whether human RPL5 P-site function is identical to yeast ortholog not directly tested
    • Structural contacts between RPL5 and peptidyl-tRNA not resolved
  5. 2004 High

    Demonstrating that RPL5 overexpression inhibits MDM2-mediated p53 ubiquitination and induces G1 arrest converted the earlier physical interaction into a functional mechanism: RPL5 is a bona fide p53 activator.

    Evidence Overexpression and siRNA knockdown in H1299/U2OS cells, p53 ubiquitination assay, cell cycle analysis

    PMID:15308643

    Open questions at the time
    • Whether RPL5 alone suffices or requires RPL11 cooperation unclear
    • Endogenous stoichiometry of MDM2–RPL5 complex not determined
  6. 2006 High

    Identification of nucleophosmin (NPM/B23) as a CRM1-dependent chaperone for RPL5/5S rRNA nuclear export resolved how the extraribosomal RNP reaches maturing 60S subunits in the cytoplasm.

    Evidence Biochemical purification of NPM-bound complexes from HeLa cells, co-IP, colocalization with polysomes, NPM shuttling inhibition

    PMID:16648475

    Open questions at the time
    • Whether NPM chaperones RPL5 independently of 5S rRNA not addressed
    • Regulation of NPM–RPL5 handoff to pre-60S particles unknown
  7. 2008 High

    Cooperative inhibition of MDM2 by RPL5 and RPL11 (requiring L11's 5S rRNA-binding capacity) established the 5S RNP as the functional unit of the ribosomal stress checkpoint, not individual ribosomal proteins.

    Evidence MDM2 E3 activity assay with L11 5S rRNA-binding mutant, overexpression; nucleolar stress induction with MPA plus siRNA epistasis

    PMID:18305114 PMID:18560357

    Open questions at the time
    • Whether the intact 5S RNP or a remodeled form engages MDM2 in vivo unresolved
    • Quantitative threshold of free 5S RNP required for p53 activation unknown
  8. 2012 High

    Showing that RPL5 and RPL11 are mutually protected from proteasomal degradation and selectively accumulate in the ribosome-free fraction upon Pol I inhibition explained how the cell detects ribosomal biogenesis stress: excess free 5S RNP is the signal.

    Evidence Pulse-chase with proteasome inhibition, subcellular fractionation, co-IP, immunofluorescence colocalization with PML bodies

    PMID:23169665

    Open questions at the time
    • Ubiquitin ligase(s) responsible for RPL5/RPL11 turnover when incorporated into ribosomes not identified
    • Role of PML bodies as activation platforms versus bystander not resolved
  9. 2013 High

    Three concurrent studies expanded RPL5 function beyond MDM2–p53: RPL5/RPL11 recruit RISC to c-Myc mRNA for degradation, participate in oncogene-induced senescence via SRSF1, and RPL5 loss impedes cell cycle independently of p53 by reducing translational capacity.

    Evidence Co-IP of RPL5 with Ago2/TRBP and c-Myc mRNA; SRSF1–MDM2–RPL5 co-IP in primary fibroblast OIS model; polysome profiling and cyclin expression in RPL5-depleted primary cells

    PMID:23478443 PMID:24061479 PMID:24141778

    Open questions at the time
    • Specificity of RPL5-RISC targeting to c-Myc versus other mRNAs unclear
    • Whether SRSF1-containing RPL5-MDM2 complex is distinct from the canonical 5S RNP-MDM2 complex not resolved
  10. 2018 High

    SPIN1 was identified as a nucleolar sequestrator of RPL5, revealing an upstream negative regulator that prevents RPL5 from engaging MDM2 under non-stress conditions.

    Evidence Co-IP, nucleolar colocalization imaging, SPIN1 siRNA epistasis with p53 activation in cancer cells

    PMID:29547122

    Open questions at the time
    • Mechanism by which stress releases RPL5 from SPIN1 not determined
    • Whether SPIN1 also sequesters the 5S RNP as a unit or RPL5 alone unclear
  11. 2020 High

    Cancer-associated somatic RPL5 mutations were shown to impair HDM2 binding and p53 checkpoint activation, directly demonstrating that RPL5 functions as a tumor suppressor disrupted in human cancer.

    Evidence Biochemical and functional analyses of cancer-derived RPL5 variants including co-IP, ubiquitination assay, p53 stabilization in cancer cell models

    PMID:32108164

    Open questions at the time
    • Structural basis of how specific mutations disrupt the RPL5–MDM2 interface not resolved
    • Contribution of RPL5 haploinsufficiency versus dominant-negative effects in tumors unclear
  12. 2022 High

    HEATR3 was identified as the nuclear import factor for RPL5; loss-of-function HEATR3 variants cause Diamond-Blackfan anemia-like disease by impairing RPL5 nuclear accumulation, pre-rRNA processing, and erythroid differentiation.

    Evidence Patient-derived fibroblasts, HEATR3 siRNA, immunofluorescence for RPL5 nuclear import, pre-rRNA processing assay, yeast complementation

    PMID:35213692

    Open questions at the time
    • Whether HEATR3 directly binds RPL5 or acts via an adaptor not established
    • How impaired RPL5 import specifically affects erythroid lineage over other lineages unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the atomic-resolution structure of the RPL5/RPL11/5S rRNA–MDM2 inhibitory complex, the mechanism by which ribosomal stress triggers release of RPL5 from nucleolar sequestrators like SPIN1, and whether RPL5's extraribosomal functions in RISC recruitment and c-Myc regulation are broadly relevant across tissue types.
  • No high-resolution structure of RPL5–MDM2 complex available
  • Tissue-specific regulation of RPL5 extraribosomal functions unexplored
  • In vivo genetic models of RPL5 checkpoint function in mammals limited

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 6 GO:0003723 RNA binding 4 GO:0005198 structural molecule activity 2
Localization
GO:0005730 nucleolus 4 GO:0005634 nucleus 3 GO:0005840 ribosome 3 GO:0005829 cytosol 2
Pathway
R-HSA-1640170 Cell Cycle 4 R-HSA-392499 Metabolism of proteins 4 R-HSA-5357801 Programmed Cell Death 3 R-HSA-74160 Gene expression (Transcription) 3 R-HSA-1643685 Disease 2
Partners
AGO2HEATR3MDM2NPM1RPL11SPIN1SRSF1TP53
Complex memberships
5S RNP (RPL5/RPL11/5S rRNA)60S ribosomal subunitMDM2-RPL5-RPL11 checkpoint complex

Evidence

Reading pass · 25 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1994 RPL5 (ribosomal protein L5) co-purifies with MDM2 and MDM2-p53 complexes and binds 5S RNA, forming a ribonucleoprotein complex containing 5S RNA, L5, MDM2, and p53. Protein co-purification, monoclonal antibody co-immunoprecipitation, N-terminal amino acid sequencing, partial peptide mapping, RNA binding assay Molecular and cellular biology High 7935455
1988 RPL5 is the protein component of a ~7S extraribosomal 5S rRNA-protein complex that is a precursor to ribosome assembly, concentrated in the nucleolus, and distinct from ribosome-associated 5S rRNA. Patient autoantibody immunoprecipitation, pulse-chase experiments, indirect immunofluorescence microscopy, sucrose gradient sedimentation The Journal of cell biology High 3279045
1996 RPL5 has a modular domain structure: the N-terminal 93 amino acids are necessary and sufficient for 5S rRNA binding in vitro, while the C-terminal residues (amino acids 151–296) mediate nucleolar localization. In vitro RNA-binding assay, transfection experiments with deletion mutants, immunofluorescence microscopy The Journal of biological chemistry High 8626719
2000 Human RPL5 contains defined nuclear import/nucleolar localization signals at amino acids 21–37 and 255–297, a nuclear export signal (NES) at amino acids 101–111 with leucine-rich motif, and RNA-binding domains at both N- and C-termini. Deletion mutant transfection, microinjection studies in somatic cells, nuclear export assay The Journal of biological chemistry High 10766838
1999 RPL5 in Xenopus oocytes contains three separate nuclear localization sequences (NLS-1, -2, -3); NLS-1 and NLS-3 promote nuclear transfer of attached RNP complexes; NLS-1 strongly binds import receptors; nucleolar accumulation requires the full-length protein. Xenopus oocyte microinjection, NLS mapping with deletion constructs, import receptor binding assay The Journal of biological chemistry High 10567357
2006 Nucleophosmin (NPM/B23) directly interacts with RPL5 via a CRM1-dependent nuclear export sequence, chaperoning the nuclear export of RPL5 and 5S rRNA; inhibition of NPM shuttling or NPM loss blocks RPL5/5S rRNA nuclear export, resulting in cell cycle arrest. Biochemical purification of NPM-bound complexes from HeLa cells, co-IP, colocalization with maturing 60S subunits and polysomes, NPM shuttling inhibition experiments Molecular and cellular biology High 16648475
2004 RPL5 activates p53 by inhibiting MDM2-mediated p53 ubiquitination; L5 overexpression stabilizes p53, enhances p53 transcriptional activity, and induces G1 cell cycle arrest; L5 forms at least two MDM2 complexes: MDM2-L5-L11-L23 and p53-MDM2-L5-L11-L23; L5 interaction with MDM2 is enhanced by low-dose actinomycin D treatment. Overexpression in H1299 and U2OS cells, p53 ubiquitination assay in cells, reciprocal co-IP, siRNA knockdown of L5, cell cycle analysis The Journal of biological chemistry High 15308643
2008 L5 and L11 cooperate to robustly inhibit MDM2 E3 ubiquitin ligase activity toward p53; the ability of L11 to bind 5S rRNA is required for this cooperation, as an L11 mutant unable to bind 5S rRNA cannot cooperate with L5. Overexpression, MDM2 E3 activity assay, L11 5S rRNA-binding mutant analysis Oncogene High 18560357
2008 Mycophenolic acid (MPA) treatment disrupts the nucleolus and inhibits pre-rRNA synthesis, enhancing MDM2-L5 and MDM2-L11 interaction; knockdown of L5 or L11 markedly impairs MPA-induced p53 stabilization and G1 arrest, demonstrating that L5 and L11 are required for MPA-induced nucleolar stress p53 activation. siRNA knockdown, co-IP, p53 induction assay, cell cycle analysis The Journal of biological chemistry High 18305114
2010 Perturbation of 60S ribosomal biogenesis (by knockdown of L29 or L30) activates p53 specifically through L5 and L11; L29/L30 depletion increases L11 NEDDylation and nuclear retention, enhancing L11 and L5 binding to MDM2 and inhibiting MDM2-mediated p53 ubiquitination; L29 and L30 themselves do not bind MDM2. siRNA knockdown, co-IP, MDM2-p53 ubiquitination assay, NEDD8 knockdown epistasis The Journal of biological chemistry High 20554519
2012 Upon inhibition of Pol I activity, L5 and L11 selectively accumulate in the ribosome-free fraction due to their mutual protection from proteasomal degradation, then bind MDM2 and co-localize with MDM2, p53, and PML in disrupted nucleoli, which serve as a platform for p53 activation under ribosomal biogenesis stress. Pulse-chase/proteasome inhibition, subcellular fractionation, co-IP, immunofluorescence colocalization Proceedings of the National Academy of Sciences of the United States of America High 23169665
2013 RPL5 (co-operatively with RPL11) guides the RNA-induced silencing complex (RISC) to c-Myc mRNA and mediates its degradation; RPL5 binds the 3'UTR of c-Myc mRNA and RISC subunits TRBP and Ago2, suppressing c-Myc expression at both mRNA and protein levels. siRNA knockdown, overexpression, immunoprecipitation of RPL5 with RISC subunits and c-Myc mRNA, c-Myc mRNA/protein level assays Oncogene High 24141778
2013 SRSF1 (oncogenic splicing factor) is a component of an extraribosomal MDM2/RPL5 complex; RPL5-MDM2 complex is implicated in oncogene-induced senescence (OIS) triggered by SRSF1 overexpression, linking spliceosomal and ribosomal components in a p53 checkpoint. Co-IP of SRSF1 with MDM2/RPL5, p53 stabilization assay, primary fibroblast OIS assay Molecular cell High 23478443
2013 Loss of RPL5 or RPL11 does not trigger a p53-dependent cell cycle arrest checkpoint but impedes cell cycle progression by reducing ribosome content and translational capacity, which suppresses cyclin accumulation at the translational level. siRNA knockdown in primary human lung fibroblasts, ribosome profiling, polysome analysis, cell cycle analysis, cyclin expression by western blot Molecular and cellular biology High 24061479
2014 RPL5 and RPL11 directly associate with the transactivation domain of TAp73 independently of MDM2, preventing MDM2-TAp73 interaction and consequently enhancing TAp73 transcriptional activity and TAp73-mediated apoptosis in response to ribosomal stress. Co-IP, overexpression/knockdown, reporter assays for TAp73 transcriptional activity, apoptosis assay Cell death and differentiation High 25301064
1998 RPL5 interacts with eukaryotic initiation factor 5A (eIF-5A), which connects HIV-1 Rev with cellular RNA transport; overexpression of L5 enhances Rev activity and antibodies against L5 or eIF-5A inhibit Rev nuclear export, implicating L5 in 5S rRNA/mRNA export pathways shared with Rev. Co-immunoprecipitation of eIF-5A and L5, overexpression functional assay, antibody microinjection nuclear export inhibition Proceedings of the National Academy of Sciences of the United States of America Medium 9465063
1995 RPL5 interacts with the catalytic subunit of type 1 protein phosphatase (PP1) via a yeast two-hybrid screen; L5-5S RNA complex activates PP1 phosphatase activity in vitro; PP1 co-sediments with ribosomal subunits containing L5 but not with L5-deficient subunits. Yeast two-hybrid, in vitro phosphatase activity assay, ribosome subunit co-sedimentation The Journal of biological chemistry Medium 7649987
1996 The beta subunit of casein kinase II (CKII) interacts with RPL5 (but not the alpha/alpha' catalytic subunits) in a yeast two-hybrid assay, suggesting the beta subunit targets CKII to L5 and a potential role for CKII in RPL5 phosphorylation, ribosomal assembly, or transport. Yeast two-hybrid Biochemical and biophysical research communications Low 8806611
2001 In yeast (S. cerevisiae), rpl5 mutants increase both -1 and +1 programmed ribosomal frameshifting efficiencies and decrease affinity of ribosomes for peptidyl-tRNA at the P-site; sparsomycin (which increases peptidyl-tRNA binding) antagonizes the frameshifting defects, demonstrating that L5 functions to anchor peptidyl-tRNA to the ribosomal P-site. rpl5 mutant analysis, frameshifting reporter assay, biochemical peptidyl-tRNA binding assay, sparsomycin pharmacology RNA High 11497428
2018 SPIN1 (Spindlin 1) binds RPL5/uL18 in the nucleolus, sequestering it from interacting with MDM2; SPIN1 ablation increases ribosome-free uL18 and uL5 (RPL11), which are required for SPIN1 depletion-induced p53 activation; SPIN1 negatively regulates the uL18-MDM2 axis to suppress p53. Co-IP, nucleolar localization by imaging, SPIN1 siRNA ablation with p53 activation assay, cancer cell growth and apoptosis assays eLife High 29547122
2017 HEATR1 (a nucleolar protein that positively regulates rRNA synthesis) depletion disrupts nucleolar structure and activates the RPL5/RPL11-MDM2-p53 ribosome biogenesis stress checkpoint, causing p53-dependent cell cycle arrest. HEATR1 siRNA knockdown, nucleolar disruption imaging, p53 stabilization assay, cell cycle analysis, epistasis with RPL5/RPL11 knockdown Cell cycle Medium 29143558
2020 Cancer-associated somatic RPL5 mutations impair the ability of RPL5 to bind HDM2 and inhibit its ubiquitin ligase activity toward p53, disrupting the RPL5/RPL11/5S rRNA-HDM2-p53 ribosome biogenesis checkpoint in human cancer cells. Computational analysis of mutation databases, biochemical/functional analyses in cancer cell models, Co-IP, ubiquitination assay, p53 stabilization assay Oncogene High 32108164
2022 HEATR3 variants impair nuclear import of uL18/RPL5 (and uL5/RPL11), reducing their nuclear accumulation, and cause Diamond-Blackfan anemia-like phenotypes including impaired pre-rRNA processing, ribosomal subunit formation, and erythroid differentiation. Patient-derived fibroblasts, HEATR3 siRNA knockdown, immunofluorescence for nuclear accumulation of uL18, pre-rRNA processing assay, yeast complementation model Blood High 35213692
2020 MeCP2 represses RPL5 (and RPL11) transcription by binding to their promoter regions, reducing RPL5 levels and consequently increasing MDM2-mediated p53 ubiquitination and degradation, promoting breast cancer cell proliferation. ChIP showing MeCP2 binding to RPL5 promoter, RPL5 overexpression with p53/MDM2 ubiquitination assay, siRNA knockdown, cell proliferation and apoptosis assays Oncogenesis Medium 32483207
2020 WDR74 modulates RPL5 protein levels and consequently regulates MDM2 activity and p53 ubiquitination; WDR74 overexpression reduces RPL5, while WDR74 loss increases RPL5 and p53 stability. iTRAQ proteomics, gain/loss-of-function assays, co-IP, p53 ubiquitination assay, in vivo xenograft Oncogene Medium 32005977

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2004 Inhibition of MDM2-mediated p53 ubiquitination and degradation by ribosomal protein L5. The Journal of biological chemistry 467 15308643
1994 The ribosomal L5 protein is associated with mdm-2 and mdm-2-p53 complexes. Molecular and cellular biology 289 7935455
1999 The inhibitory receptor LIR-1 uses a common binding interaction to recognize class I MHC molecules and the viral homolog UL18. Immunity 278 10591185
2012 Mutual protection of ribosomal proteins L5 and L11 from degradation is essential for p53 activation upon ribosomal biogenesis stress. Proceedings of the National Academy of Sciences of the United States of America 184 23169665
2006 Nucleophosmin is essential for ribosomal protein L5 nuclear export. Molecular and cellular biology 170 16648475
1988 A 5S rRNA/L5 complex is a precursor to ribosome assembly in mammalian cells. The Journal of cell biology 162 3279045
1997 Preventing the loss of competence for neural induction: HGF/SF, L5 and Sox-2. Development (Cambridge, England) 146 9102306
1993 Superinfection immunity of mycobacteriophage L5: applications for genetic transformation of mycobacteria. Molecular microbiology 125 8459767
2007 The human cytomegalovirus MHC class I homolog UL18 inhibits LIR-1+ but activates LIR-1- NK cells. Journal of immunology (Baltimore, Md. : 1950) 114 17372005
2008 Cooperation between the ribosomal proteins L5 and L11 in the p53 pathway. Oncogene 112 18560357
2015 Mechanism of UCH-L5 activation and inhibition by DEUBAD domains in RPN13 and INO80G. Molecular cell 106 25702870
2008 Replication of KIAA0350, IL2RA, RPL5 and CD58 as multiple sclerosis susceptibility genes in Australians. Genes and immunity 106 18650830
2017 The ribosomal protein gene RPL5 is a haploinsufficient tumor suppressor in multiple cancer types. Oncotarget 101 28147343
2008 Mycophenolic acid activation of p53 requires ribosomal proteins L5 and L11. The Journal of biological chemistry 94 18305114
2013 Splicing-factor oncoprotein SRSF1 stabilizes p53 via RPL5 and induces cellular senescence. Molecular cell 93 23478443
2013 Ribosomal proteins L5 and L11 co-operatively inactivate c-Myc via RNA-induced silencing complex. Oncogene 88 24141778
2013 Loss of tumor suppressor RPL5/RPL11 does not induce cell cycle arrest but impedes proliferation due to reduced ribosome content and translation capacity. Molecular and cellular biology 83 24061479
2008 Structure of UL18, a peptide-binding viral MHC mimic, bound to a host inhibitory receptor. Proceedings of the National Academy of Sciences of the United States of America 79 18632577
2015 Plasma L5 levels are elevated in ischemic stroke patients and enhance platelet aggregation. Blood 77 26679863
2009 Diamond-Blackfan anemia: genotype-phenotype correlations in Italian patients with RPL5 and RPL11 mutations. Haematologica 77 19773262
2014 A comparison of RNA-seq and exon arrays for whole genome transcription profiling of the L5 spinal nerve transection model of neuropathic pain in the rat. Molecular pain 73 24472155
1997 Transcriptional silencing by the mycobacteriophage L5 repressor. The EMBO journal 71 9312049
1980 The overall synthesis of L-5,6-dihydroorotate by multienzymatic protein pyr1-3 from hamster cells. Kinetic studies, substrate channeling, and the effects of inhibitors. The Journal of biological chemistry 70 6108323
2010 Perturbation of 60 S ribosomal biogenesis results in ribosomal protein L5- and L11-dependent p53 activation. The Journal of biological chemistry 68 20554519
1998 Interaction of the HIV-1 rev cofactor eukaryotic initiation factor 5A with ribosomal protein L5. Proceedings of the National Academy of Sciences of the United States of America 67 9465063
1992 Construction and characterization of a human cytomegalovirus mutant with the UL18 (class I homolog) gene deleted. Journal of virology 64 1328689
2000 Human ribosomal protein L5 contains defined nuclear localization and export signals. The Journal of biological chemistry 61 10766838
1996 Distinct domains in ribosomal protein L5 mediate 5 S rRNA binding and nucleolar localization. The Journal of biological chemistry 59 8626719
2004 brinker and optomotor-blind act coordinately to initiate development of the L5 wing vein primordium in Drosophila. Development (Cambridge, England) 57 15073155
2016 L5-LDL from ST-elevation myocardial infarction patients induces IL-1β production via LOX-1 and NLRP3 inflammasome activation in macrophages. American journal of physiology. Heart and circulatory physiology 55 27864235
2004 Genetic analysis of RpL38 and RpL5, two minute genes located in the centric heterochromatin of chromosome 2 of Drosophila melanogaster. Genetics 55 15520262
2001 Ribosomal protein L5 helps anchor peptidyl-tRNA to the P-site in Saccharomyces cerevisiae. RNA (New York, N.Y.) 53 11497428
2018 SPIN1 promotes tumorigenesis by blocking the uL18 (universal large ribosomal subunit protein 18)-MDM2-p53 pathway in human cancer. eLife 51 29547122
2016 RPL5 on 1p22.1 is recurrently deleted in multiple myeloma and its expression is linked to bortezomib response. Leukemia 51 27909306
1995 Interaction of the ribosomal protein, L5, with protein phosphatase type 1. The Journal of biological chemistry 51 7649987
1997 Characterization of the mycobacteriophage L5 attachment site, attP. Journal of molecular biology 50 9054972
2014 Ribosomal proteins L11 and L5 activate TAp73 by overcoming MDM2 inhibition. Cell death and differentiation 46 25301064
2009 Selective expression of ligand-gated ion channels in L5 pyramidal cell axons. The Journal of neuroscience : the official journal of the Society for Neuroscience 46 19759293
2004 Specific recognition of the viral protein UL18 by CD85j/LIR-1/ILT2 on CD8+ T cells mediates the non-MHC-restricted lysis of human cytomegalovirus-infected cells. Journal of immunology (Baltimore, Md. : 1950) 45 15100307
1992 The virulence-determining genomic BamHI fragment 4 of pseudorabies virus contains genes corresponding to the UL15 (partial), UL18, UL19, UL20, and UL21 genes of herpes simplex virus and a putative origin of replication. Virology 45 1333128
2020 Cancer-associated mutations in the ribosomal protein L5 gene dysregulate the HDM2/p53-mediated ribosome biogenesis checkpoint. Oncogene 43 32108164
2016 Ubiquitin carboxyl-terminal hydrolase-L5 promotes TGFβ-1 signaling by de-ubiquitinating and stabilizing Smad2/Smad3 in pulmonary fibrosis. Scientific reports 42 27604640
1976 RNA sequences associated with proteins L1, L9, and L5, L18, L25, in ribonucleoprotein fragments isolated from the 50-S subunit of Escherichia coli ribosomes. European journal of biochemistry 42 827440
1999 Functional modules in ribosomal protein L5 for ribonucleoprotein complex formation and nucleocytoplasmic transport. The Journal of biological chemistry 41 10567357
2020 MeCP2 facilitates breast cancer growth via promoting ubiquitination-mediated P53 degradation by inhibiting RPL5/RPL11 transcription. Oncogenesis 40 32483207
2003 Control of directionality in L5 integrase-mediated site-specific recombination. Journal of molecular biology 40 12581642
2000 Identification and characterization of mycobacteriophage L5 excisionase. Molecular microbiology 38 10652095
2009 Crystal structure of the de-ubiquitinating enzyme UCH37 (human UCH-L5) catalytic domain. Biochemical and biophysical research communications 37 19836345
2020 Ubiquitin C-Terminal Hydrolase L5 (UCHL5) Accelerates the Growth of Endometrial Cancer via Activating the Wnt/β-Catenin Signaling Pathway. Frontiers in oncology 36 32596150
2007 Human cytomegalovirus-derived protein UL18 alters the phenotype and function of monocyte-derived dendritic cells. Journal of leukocyte biology 36 17898320
2023 Supplementation with high-GABA-producing Lactobacillus plantarum L5 ameliorates essential tremor triggered by decreased gut bacteria-derived GABA. Translational neurodegeneration 35 38093327
2017 Perturbation of RNA Polymerase I transcription machinery by ablation of HEATR1 triggers the RPL5/RPL11-MDM2-p53 ribosome biogenesis stress checkpoint pathway in human cells. Cell cycle (Georgetown, Tex.) 35 29143558
2014 p53-Independent cell cycle and erythroid differentiation defects in murine embryonic stem cells haploinsufficient for Diamond Blackfan anemia-proteins: RPS19 versus RPL5. PloS one 35 24558476
2018 Serotonin Differentially Regulates L5 Pyramidal Cell Classes of the Medial Prefrontal Cortex in Rats and Mice. eNeuro 34 29445767
2006 Spontaneous mutations in the human CMV HLA class I homologue UL18 affect its binding to the inhibitory receptor LIR-1/ILT2/CD85j. European journal of immunology 34 16479538
1996 Interaction of the beta subunit of casein kinase II with the ribosomal protein L5. Biochemical and biophysical research communications 34 8806611
2019 Risk factors of instrumentation failure and pseudarthrosis after stand-alone L5-S1 anterior lumbar interbody fusion: a retrospective cohort study. Journal of neurosurgery. Spine 33 31151106
2014 JNK1 controls dendritic field size in L2/3 and L5 of the motor cortex, constrains soma size, and influences fine motor coordination. Frontiers in cellular neuroscience 33 25309320
2011 Chemical composition-oriented receptor selectivity of L5, a naturally occurring atherogenic low-density lipoprotein. Pure and applied chemistry. Chimie pure et appliquee 33 24198440
2015 The Fate of L5-S1 With Low-Dose BMP-2 and Pelvic Fixation, With or Without Interbody Fusion, in Adult Deformity Surgery. Spine 32 25768688
2014 Cross-protective effect of a combined L5 plus L3 Leishmania major ribosomal protein based vaccine combined with a Th1 adjuvant in murine cutaneous and visceral leishmaniasis. Parasites & vectors 32 24382098
1996 Characterization of Mycobacterium smegmatis gene that confers resistance to phages L5 and D29 when overexpressed. Molecular microbiology 32 8843442
2008 The natural history of age-related disc degeneration: the influence of age and pathology on cell populations in the L4-L5 disc. Spine 31 19050583
2022 HEATR3 variants impair nuclear import of uL18 (RPL5) and drive Diamond-Blackfan anemia. Blood 30 35213692
2016 Transcriptome analysis reveals a ribosome constituents disorder involved in the RPL5 downregulated zebrafish model of Diamond-Blackfan anemia. BMC medical genomics 30 26961822
2015 Sesamol reduces the atherogenicity of electronegative L5 LDL in vivo and in vitro. Journal of natural products 30 25692815
2007 Increased expression of leukocyte Ig-like receptor-1 and activating role of UL18 in the response to cytomegalovirus infection. Journal of immunology (Baltimore, Md. : 1950) 29 17339449
2004 Transfer of RPS14 and RPL5 from the mitochondrion to the nucleus in grasses. Gene 29 14693379
2017 Electronegative L5-LDL induces the production of G-CSF and GM-CSF in human macrophages through LOX-1 involving NF-κB and ERK2 activation. Atherosclerosis 28 29078142
2020 WDR74 modulates melanoma tumorigenesis and metastasis through the RPL5-MDM2-p53 pathway. Oncogene 27 32005977
2014 Lytic peptidase L5 of Lysobacter sp. XL1 with broad antimicrobial spectrum. Journal of molecular microbiology and biotechnology 27 24434599
2008 Human cytomegalovirus UL18 utilizes US6 for evading the NK and T-cell responses. PLoS pathogens 27 18688275
1996 The rpl5-rps14-cob gene arrangement in Solanum tuberosum: rps14 is a transcribed and unedited pseudogene. Plant molecular biology 27 8806426
2020 Cav3.2 overexpression in L4 dorsal root ganglion neurons after L5 spinal nerve cutting involves Egr-1, USP5 and HMGB1 in rats: An emerging signaling pathway for neuropathic pain. European journal of pharmacology 26 32971090
2017 Mitochondrial Retroprocessing Promoted Functional Transfers of rpl5 to the Nucleus in Grasses. Molecular biology and evolution 26 28541477
2022 Ribosomal protein L5 (RPL5)/ E2F transcription factor 1 (E2F1) signaling suppresses breast cancer progression via regulating endoplasmic reticulum stress and autophagy. Bioengineered 25 35293275
2011 Interaction between the G3 and L5 proteins of the vaccinia virus entry-fusion complex. Virology 25 21295816
2010 Tag-SNP analysis of the GFI1-EVI5-RPL5-FAM69 risk locus for multiple sclerosis. European journal of human genetics : EJHG 24 20087403
2006 L5, the most electronegative subfraction of plasma LDL, induces endothelial vascular cell adhesion molecule 1 and CXC chemokines, which mediate mononuclear leukocyte adhesion. Atherosclerosis 24 17022986
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2022 Ribosomal protein RPL5 regulates colon cancer cell proliferation and migration through MAPK/ERK signaling pathway. BMC molecular and cell biology 22 36384455
2007 Variability of UL18, UL40, UL111a and US3 immunomodulatory genes among human cytomegalovirus clinical isolates from renal transplant recipients. Journal of clinical virology : the official publication of the Pan American Society for Clinical Virology 22 17827058
2020 Increased APOE glycosylation plays a key role in the atherogenicity of L5 low-density lipoprotein. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 21 32501643
2016 Leukocyte Immunoglobulin-Like Receptor 1-Expressing Human Natural Killer Cell Subsets Differentially Recognize Isolates of Human Cytomegalovirus through the Viral Major Histocompatibility Complex Class I Homolog UL18. Journal of virology 21 26739048
2012 The most negatively charged low-density lipoprotein L5 induces stress pathways in vascular endothelial cells. Journal of vascular research 21 22627396
2008 Immune modulation by the human cytomegalovirus-encoded molecule UL18, a mystery yet to be solved. Journal of immunology (Baltimore, Md. : 1950) 21 18096997
1999 Protein-DNA complexes in mycobacteriophage L5 integrative recombination. Journal of bacteriology 21 9882658
2003 Existence of brain-derived neurotrophic factor and vanilloid receptor subtype 1 immunoreactive sensory DRG neurons innervating L5/6 intervertebral discs in rats. Journal of orthopaedic science : official journal of the Japanese Orthopaedic Association 20 12560892
2020 Heavy ion radiation-induced DNA damage mediates apoptosis via the Rpl27a-Rpl5-MDM2-p53/E2F1 signaling pathway in mouse spermatogonia. Ecotoxicology and environmental safety 19 32535367
2016 Development of Soft Tissue Sarcomas in Ribosomal Proteins L5 and S24 Heterozygous Mice. Journal of Cancer 19 26722357
2013 Cis association of leukocyte Ig-like receptor 1 with MHC class I modulates accessibility to antibodies and HCMV UL18. European journal of immunology 19 23348966
1999 Cotranscription of the rpl5-rps14-cob gene cluster in pea mitochondria. Molecular & general genetics : MGG 19 10323235
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2022 Translocator Protein (TSPO) Alleviates Neuropathic Pain by Activating Spinal Autophagy and Nuclear SIRT1/PGC-1α Signaling in a Rat L5 SNL Model. Journal of pain research 17 35356265
2022 Quantitative Fluorescence Analysis Reveals Dendrite-Specific Thalamocortical Plasticity in L5 Pyramidal Neurons during Learning. The Journal of neuroscience : the official journal of the Society for Neuroscience 17 36639912
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2002 Predominant neuronal B-cell loss in L5 DRG of p75 receptor-deficient mice. Journal of anatomy 17 11833656
2020 Ribosomal protein L5 mediated inhibition of c-Myc is critically involved in sanggenon G induced apoptosis in non-small lung cancer cells. Phytotherapy research : PTR 16 32935429
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