Large ribosomal subunit protein uL4 · UniProt P36578
RPL4 (uL4) is a universally conserved large-ribosomal-subunit protein that makes direct, extended contacts with the large-subunit rRNA and is among the earliest proteins to engage the pre-rRNA during subunit biogenesis (PMID:814400, PMID:9838082). In the assembled ribosome it contacts the 23S/large-subunit rRNA at defined positions (Tyr35–U615 and nucleotides 320–325 in E. coli) and lines the peptide exit tunnel, uniquely contacting nascent chains along the entire tunnel length (PMID:6998491, PMID:3278299, PMID:24072706). A conserved internal loop is critical for late assembly steps, while full-length depletion blocks early assembly, defining two temporally separable biogenesis roles (PMID:25246649); in eukaryotes the nascent protein is co-translationally captured by the dedicated chaperone Acl4, whose TPR domain sequesters the exposed internal loop to prevent premature rRNA insertion and whose binding site overlaps that of the transport factor Kap104 to provide continuous protection during nuclear escort, with a eukaryote-specific extension contacting Rpl18 to orchestrate chaperone release and pre-60S incorporation (PMID:25936803, PMID:26447800, PMID:28148929). Beyond its structural role, bacterial L4 is the autogenous regulator of its own S10 operon, acting both translationally and by stabilizing a NusA-dependent transcriptional pause at an attenuator hairpin through direct binding to the mRNA leader using an RNA-binding surface structurally distinct from its rRNA-binding interface (PMID:6996835, PMID:6157482, PMID:2157208, PMID:12738792, PMID:8846294); L4 also binds and inhibits RNase E to broadly stabilize stress-response transcripts (PMID:19144914). Mutations in the conserved loop tip confer macrolide (and in some organisms chloramphenicol/linezolid) resistance by perturbing higher-order 23S rRNA conformation in domains II, III and V and slowing peptide-bond formation and elongation (PMID:10369764, PMID:17956547, PMID:10898684, PMID:16048983). In humans, RPL4 acts extraribosomally by binding the MDM2 acidic domain to suppress p53 ubiquitination and promote an RPL4–RPL5–RPL11–MDM2 complex (PMID:26908445), and supports rRNA processing through interaction with the RNA helicase Guα (PMID:16045751).
Establishing that L4 directly touches rRNA in the assembled ribosome anchored its structural role within the large subunit.
Evidence UV cross-linking of E. coli 50S subunits with immunological identification
Mapping the cross-link to Tyr35 of L4 and U615 of 23S rRNA pinpointed the precise rRNA contact and demonstrated the molecular intimacy of the L4–rRNA interface.
Evidence Protease/nuclease digestion of isolated cross-linked complexes with oligonucleotide analysis
Showing that L4 alone among S10 operon proteins represses translation of the operon, both in vitro and on overexpression in vivo, identified L4 as a self-limiting feedback regulator of ribosomal protein synthesis.
Evidence DNA-directed in vitro synthesis with purified L4 plus in vivo oversynthesis assays
Discovery that L4 also stimulates premature transcription termination in the S10 leader, independent of translational control, revealed a second, transcriptional layer of autogenous regulation.
Evidence In vivo RNA end-mapping plus reconstituted in vitro transcription with purified L4 and NusA
Dissecting the attenuator showed NusA establishes the polymerase pause while distinct leader elements allow L4 to stabilize it, separating the pause and stabilization functions.
Evidence Genetic and antisense-oligonucleotide dissection of a purified in vitro transcription system
PMID:1285127 PMID:7531246 PMID:7844821
Isolation of mutants that separate ribosome assembly from autogenous control proved L4 uses non-identical RNA-binding surfaces for rRNA versus mRNA recognition.
Evidence Genetic isolation of L4 mutants with orthogonal assembly and regulation readouts
Demonstrating direct, specific L4 binding to a hairpin in the S10 mRNA leader, with three-dimensional mimicry of the rRNA site, explained how one protein recognizes both RNAs.
Evidence In vitro binding, competition with the rRNA site, and phosphorothioate footprinting
Showing L4 erythromycin-resistance mutations perturb 23S rRNA domains II/III/V and lower peptidyl-transferase activity established that L4 mutations act allosterically through rRNA conformation rather than direct drug contact.
Evidence Chemical probing of 23S rRNA in defined resistance-mutant ribosomes
A 1.7 Å crystal structure of L4 provided an atomic framework, identifying separate N-terminal rRNA and C-terminal mRNA/protein binding sites and a large disordered loop.
Evidence X-ray crystallography of Thermotoga maritima L4
Identifying physical and functional partnership with RNA helicase Guα extended RPL4's role into eukaryotic rRNA processing.
Evidence Co-IP, siRNA knockdown, and rescue with interaction-deficient RPL4 in mammalian cells
Discovery that L4 binds and inhibits RNase E and broadly stabilizes stress-response mRNAs revealed an extraribosomal moonlighting role in RNA turnover.
Evidence Affinity purification, two-hybrid, in vitro cleavage and in vivo mRNA half-life and microarray assays
Separating the consequences of deleting the internal loop versus depleting full L4 defined two distinct biogenesis roles and showed the eukaryote-specific extension is dispensable for viability.
Evidence Yeast conditional depletion/deletion mutants with ribosome profiling and pulse-chase
Identification of the dedicated chaperone Acl4, which sequesters the internal loop and is escorted to the nucleus, explained how the aggregation-prone nascent L4 is protected and hierarchically incorporated via an Rpl18 co-evolved site.
Evidence Yeast genetics, co-IP, co-translational capture assays and mutagenesis of contact surfaces
Demonstrating RPL4 binds the MDM2 acidic domain to suppress p53 ubiquitination connected RPL4 to the ribosomal stress–p53 surveillance axis.
Evidence Reciprocal Co-IP, ubiquitination and p53 stability assays in human cells
Finding that RPL4 cooperates with Nucleolin to support EBNA1-mediated EBV episome maintenance revealed a host-factor role hijacked by virus.
Evidence Co-IP, shRNA knockdown, oriP reporter, ChIP and genome copy-number assays
Showing that bacterial uL4 loop deletion causes cold-sensitive assembly defects and altered response to a translation-pause peptide tied the loop to both assembly and nascent-chain sensing.
Evidence E. coli strains expressing only loop-deleted uL4 with sedimentation and in vivo pausing assays
| Year | Finding | Method | Journal | Conf | PMIDs |
|---|---|---|---|---|---|
| 1980 | E. coli ribosomal protein L4 autogenously regulates the S10 operon by inhibiting translation of the promoter-proximal proteins (S10, L3, L4, L23, L2) in vitro; no other S10 operon protein caused selective inhibition, and L4 did not inhibit other operons. | Lambda fus3 DNA-directed in vitro protein synthesis system with purified L4 protein addition | Cell | High | 6996835 |
| 1980 | E. coli ribosomal protein L4, when overproduced in vivo, reduces mRNA synthesis from at least four genes of the S10 operon, identifying L4 as the sole autogenous regulator among eleven S10 operon products. | In vivo oversynthesis of individual S10 operon ribosomal proteins with measurement of individual protein synthesis rates | Cell | High | 6157482 |
| 1975 | Ribosomal protein L4 is the primary RNA-crosslinked protein in the E. coli 50S subunit upon UV irradiation, demonstrating direct RNA-protein contact in the ribosome. | UV cross-linking of 50S subunits followed by Sarkosyl gel analysis, 2D electrophoresis, and immunological identification with protein-specific antisera | Molecular & general genetics : MGG | High | 814400 |
| 1980 | The UV cross-link between L4 and 23S rRNA is located at tyrosine-35 of L4 and uridine-615 of 23S rRNA, establishing the precise RNA-protein contact site. | Isolation of cross-linked L4-oligonucleotide complexes, successive protease and nuclease digestions, 2D gel electrophoresis, oligonucleotide analysis | Biochemistry | High | 6998491 |
| 1988 | L4 is cross-linked to nucleotides 320–325 of E. coli 23S rRNA when 50S subunits are treated with 2-iminothiolane followed by UV irradiation, identifying an additional RNA contact site. | Chemical cross-linking (2-iminothiolane + UV), affinity chromatography with antibodies, partial RNA digestion, complex isolation | Nucleic acids research | Medium | 3278299 |
| 1990 | L4 stimulates premature transcription termination approximately 140 bases from the transcription start site within the S10 operon leader, upstream of the first structural gene, by a mechanism independent of its inhibition of translation. | In vivo mapping of 5′ and 3′ ends of newly synthesized RNA; deletion of sequences downstream of termination site | Journal of molecular biology | High | 1692593 |
| 1990 | L4 stimulates termination of transcription at the S10 attenuator in vitro, but RNA polymerase requires NusA to terminate at this site; L4 increases termination efficiency at the NusA-dependent pause site. | Purified in vitro transcription system with purified L4 and NusA proteins | Proceedings of the National Academy of Sciences of the United States of America | High | 2157208 |
| 1992 | NusA is required for RNA polymerase pausing at the S10 attenuation site; L4 further stabilizes the NusA-modified paused complex. These two activities require genetically separable regions of the S10 leader RNA. | In vitro transcription with purified components; isolation of paused complexes; genetic deletion analysis of leader RNA | Genes & development | High | 1285127 |
| 1995 | The upper stem-loop of the attenuator hairpin is the major determinant for NusA-dependent pausing; upstream hairpin and the ascending side of the attenuator are required for L4 stabilization of the paused complex; NusA is not absolutely required for initial RNA polymerase pausing at low UTP concentration, but is required for L4 to stabilize the pause. | Genetic and antisense oligonucleotide competition with purified in vitro transcription system; isolation of paused complexes | Journal of molecular biology | High | 7531246 7844821 |
| 1999 | E. coli L4 binds directly and specifically to the S10 mRNA leader in vitro; the binding site is a small hairpin structure within the leader but a 64-nucleotide sequence is required for full interaction. Phosphorothioate footprinting shows structural mimicry between the mRNA and rRNA binding sites in three dimensions. | In vitro binding assays, competition with 23S rRNA L4-binding site, phosphorothioate footprinting, mutational analysis | The Journal of biological chemistry | High | 12738792 |
| 1996 | L4 uses non-identical determinants for ribosome assembly and autogenous S10 operon regulation: mutations were isolated that eliminate autogenous control but allow ribosome assembly, and vice versa, showing the two RNA-binding interfaces are distinct. | Genetic isolation of L4 mutants; functional assays for ribosome assembly and autogenous control | RNA | High | 8846294 |
| 2000 | Crystal structure of Thermotoga maritima L4 at 1.7 Å resolution reveals an alternating α/β fold with a large disordered loop; two separate RNA-binding sites are identified: an N-terminal site (disordered loop with flanking helices) for rRNA binding and a C-terminal site (two non-consecutive helices) candidate for S10 mRNA leader interaction; a C-terminal protein-binding interface is also identified. | X-ray crystallography at 1.7 Å resolution | The EMBO journal | High | 10698923 |
| 1999 | Erythromycin resistance mutations in ribosomal protein L4 (but not L22) of E. coli result in decreased peptide-bond-forming activity and perturb the conformation of 23S rRNA nucleotides in domains II, III, and V, without affecting the A2058 region—indicating L4 mutations act by altering rRNA higher-order structure. | Chemical modification/probing of 23S rRNA structure in ribosomes from resistance mutants; multiple chemical probes at multiple nucleotide positions | Journal of molecular biology | High | 10369764 |
| 2004 | L4 mutant ribosomes with altered L4 protein show increased frameshifting, missense decoding, stop codon readthrough, non-AUG initiation, and mutant initiator tRNA utilization, demonstrating that L4 mutations in the 50S subunit also alter 30S subunit structure and function upon association. | In vivo translational fidelity assays (frameshifting, readthrough, missense) in E. coli L4 and L22 mutant strains; antibiotic interaction assays | Nucleic acids research | High | 15509870 |
| 2007 | Eight new L4 mutations in the tentacle region of E. coli L4 confer erythromycin resistance; all reduce in vivo peptide-chain elongation rates and increase precursor 23S rRNA levels; large insertions in L4 cause accumulation of abnormal ribosomal subunits, highlighting L4's role in ribosome assembly and function. | In vivo isolation of erythromycin-resistant mutants; sequencing; growth rate measurements; ribosome profiling; rRNA precursor analysis | Molecular microbiology | High | 17956547 |
| 2009 | E. coli ribosomal protein L4 interacts with RNase E outside its catalytic domain and inhibits RNase E endoribonucleolytic activity in vitro; in vivo, ectopic L4 stabilizes RNase E-targeted mRNAs and stabilizes an antisense regulatory RNA controlling plasmid replication; L4 overexpression broadens mRNA stability for stress-response transcripts. | Affinity purification, immunoprecipitation, E. coli two-hybrid screening, in vitro cleavage assays, mRNA half-life measurements, DNA microarray | Proceedings of the National Academy of Sciences of the United States of America | High | 19144914 |
| 2015 | Eukaryotic Rpl4 (yeast) requires a dedicated assembly chaperone Acl4, which binds the universally conserved internal loop of newly synthesized Rpl4 via a superhelical TPR domain, preventing premature rRNA insertion. Rpl4's eukaryote-specific extension makes distinct interactions with the 60S surface (including a co-evolved site on Rpl18) that orchestrate Acl4 release and Rpl4 pre-ribosome incorporation. | Yeast genetics, biochemical pulldowns, co-IP, mutational analysis of RpL4 and RpL18 contact sites, ribosome assembly assays | Molecular cell | High | 25936803 |
| 2015 | Acl4 acts as dedicated chaperone that binds the long internal loop of Rpl4 (yeast), accompanies it from the cytoplasm to the nucleus, and is required for 60S subunit production; both the internal loop and C-terminal eukaryote-specific extension are essential for Rpl4 function; Rpl4 contains at least five nuclear localization signals. | Yeast genetics (slow-growth and 60S deficiency phenotypes), biochemical co-fractionation, co-translational capture assays, localization studies | PLoS genetics | High | 26447800 |
| 2017 | Crystal structure of yeast Rpl4 bound to its chaperone Acl4 reveals extensive interactions sequestering 70 exposed residues of the extended Rpl4 loop; the eukaryote-specific Rpl4 extension harbors overlapping binding sites for Acl4 and nuclear transport factor Kap104, enabling continuous protection from cellular degradation machinery. | X-ray crystallography of Rpl4–Acl4 complex; biochemical binding assays for Kap104 competition; degradation assays | Nature communications | High | 28148929 |
| 2014 | In yeast, deletion of Rpl4's conserved internal loop impairs growth and reduces large ribosomal subunit levels, affecting later steps in assembly; depletion of full Rpl4 blocks early assembly steps, revealing two distinct roles for L4 in ribosome biogenesis. Deletion of the entire eukaryote-specific C-terminal extension has no effect on viability or 60S production. | Yeast genetics (conditional depletion, deletion mutants), ribosome profiling, polysome analysis, pulse-chase labeling | RNA | High | 25246649 |
| 2016 | In E. coli, deletion of the uL4 loop causes cold-sensitive ribosome assembly defects and accumulation of immature particles; the uL4 loop deletion also impairs response to the cmlA(crb) translation pause peptide but not to the secM pause peptide, while uL22 loop deletion affects different assembly intermediates. | Genetic analysis using strains expressing only loop-deleted uL4; ribosome sedimentation, rRNA processing analysis, in vivo pausing assays | Nucleic acids research | High | 27257065 |
| 2013 | In yeast, ribosomal tunnel proteins Rpl4, Rpl17, and Rpl39 all contact the signal anchor of nascent chains within the exit tunnel; Rpl4 uniquely contacts nascent chain residues throughout the entire ribosomal tunnel length, whereas Rpl17 contacts the middle region and Rpl39 the exit region. | UV cross-linking of ribosome-bound nascent chains, FLAG exposure assay for nascent chain secondary structure, antibody-based identification | The Journal of biological chemistry | High | 24072706 |
| 1998 | Yeast Rpl4 binds specifically to the 35S precursor rRNA in vitro (Ka = 4.4×10⁶/M) but not to mature 25S rRNA, indicating L4 is one of the earliest ribosomal proteins to engage the pre-rRNA during 60S biogenesis. | Modified membrane filtration assay and agarose gel mobility shift assay with in vitro synthesized 35S pre-rRNA and purified ribosomal proteins | Biochimica et biophysica acta | Medium | 9838082 |
| 2016 | Human RPL4 directly interacts with the central acidic domain of MDM2, suppresses MDM2-mediated p53 ubiquitination and degradation, leading to p53 stabilization and activation; RPL4 overexpression promotes MDM2 binding to RPL5 and RPL11 and formation of an RPL4–RPL5–RPL11–MDM2 complex; RPL4 knockdown also induces p53 in an RPL5/RPL11-dependent manner, indicating ribosomal stress. | Co-immunoprecipitation, ubiquitination assays, Western blot for p53 stability, cell cycle analysis | Oncotarget | Medium | 26908445 |
| 2016 | Human RPL4 is essential for EBV Nuclear Antigen 1 (EBNA1) function: EBNA1 binds RPL4; RPL4 knockdown reduces EBNA1 activation of oriP, EBNA1 DNA binding, and EBV genome maintenance. EBV infection redistributes RPL4 to cell nuclei. RPL4 and Nucleolin form a scaffold for an EBNA1-oriP complex. | Co-immunoprecipitation, shRNA knockdown, oriP luciferase reporter assay, chromatin immunoprecipitation, EBV genome copy number measurement | Proceedings of the National Academy of Sciences of the United States of America | Medium | 26858444 |
| 2005 | Human RPL4 physically interacts with RNA helicase II/Guα; the ATPase activity of Guα is required for this interaction; RPL4 knockdown inhibits 47/45S, 32S, 28S, and 18S rRNA production, and this inhibition is rescued by wild-type but not Guα-interaction-deficient RPL4, placing RPL4 as a functional partner of Guα in rRNA processing. | Co-immunoprecipitation in mammalian cells, siRNA knockdown, rRNA processing assays, rescue with wild-type vs. mutant RPL4 | The FEBS journal | Medium | 16045751 |
| 2000 | Translational control of RPL4 mRNA is required for rapid transcription-independent neurite regeneration in PC12 cells and conditioned sensory neurons; antisense oligonucleotides to RPL4 mRNA inhibit neurite regeneration from differentiated PC12 cells and axonal elongation from conditioned sensory neurons. | Subtractive hybridization to isolate RPL4 mRNA; antisense oligonucleotide inhibition; neurite regeneration assays in PC12 and sensory neurons | Neurobiology of disease | Medium | 10964612 |
| 1998 | La autoantigen and CNBP bind mutually exclusively to the 5′UTR of Xenopus RPL4 mRNA, assisted by Ro60 autoantigen as an ancillary factor; CNBP binds as a dimer; mutations in the 5′UTR that disrupt translational control in vivo also disrupt protein binding, implicating these interactions in translational regulation of RPL4. | In vitro RNA–protein binding assays, mutational analysis of 5′UTR, competition assays, antibody perturbation, in vivo translational control assays | Journal of molecular biology | Medium | 9710533 |
| 1993 | Human RPL4 encodes a 425-amino acid protein (47,821 Da) with a 5′UTR initiating with 12 pyrimidines characteristic of vertebrate ribosomal protein mRNAs; comparison across species reveals a conserved N-terminus and a divergent C-terminus. | cDNA cloning and sequencing; deduced amino acid sequence and comparison | Biochimica et biophysica acta | Low | 8268230 |
| 2015 | RPL4 interacts with viral protein VP3 of infectious bursal disease virus (IBDV); RPL4 knockdown reduces viral protein pVP2 expression and virus titers in DF1 cells, indicating RPL4 supports IBDV replication. | Immunoprecipitation–mass spectrometry screening, co-immunoprecipitation, confocal colocalization, siRNA knockdown | Virus research | Medium | 26415754 |
| 2022 | TTC22 directly interacts with RPL4 and promotes binding of WTAP mRNA to RPL4, enhancing WTAP mRNA stability and translation; RPL4 knockdown diminishes TTC22-induced increases in m6A modification levels and downstream SNAI1 upregulation. | Co-immunoprecipitation, mRNA stability assays, siRNA knockdown, m6A quantification | Oncogene | Low | 35798874 |
| 1999 | L4 mutations in S. pneumoniae (G69C substitution; SQ insertion between Q67 and K68 in the conserved KPWRQKGTGRAR motif) confer macrolide resistance in strains without known resistance determinants, establishing that L4 loop mutations can cause macrolide resistance in a pathogen. | Serial passage selection, sequencing of 23S rRNA and L4 genes, MIC determination | Antimicrobial agents and chemotherapy | Medium | 10898684 |
| 2005 | 6-bp deletions in the L4 ribosomal protein gene of S. pneumoniae confer resistance to macrolides and chloramphenicol and nonsusceptibility to linezolid; gene transformation of susceptible strain R6 confirmed causality, with a fitness cost observed. | Sequencing, gene transformation/complementation, MIC determination | Antimicrobial agents and chemotherapy | High | 16048983 |
| Year | Title | Journal | Citations | PMID |
|---|---|---|---|---|
| 2000 | Mutations in 23S rRNA and ribosomal protein L4 account for resistance in pneumococcal strains selected in vitro by macrolide passage. | Antimicrobial agents and chemotherapy | 236 | 10898684 |
| 1991 | The open reading frames UL3, UL4, UL10, and UL16 are dispensable for the replication of herpes simplex virus 1 in cell culture. | Journal of virology | 161 | 1846207 |
| 1980 | Protein L4 of the E. coli ribosome regulates an eleven gene r protein operon. | Cell | 124 | 6157482 |
| 2014 | Modality-specific thalamocortical inputs instruct the identity of postsynaptic L4 neurons. | Nature | 117 | 24828045 |
| 1999 | Erythromycin resistance mutations in ribosomal proteins L22 and L4 perturb the higher order structure of 23 S ribosomal RNA. | Journal of molecular biology | 112 | 10369764 |
| 2005 | Novel mechanism of resistance to oxazolidinones, macrolides, and chloramphenicol in ribosomal protein L4 of the pneumococcus. | Antimicrobial agents and chemotherapy | 107 | 16048983 |
| 1975 | Specific cross-linking of proteins S7 and L4 to ribosomal RNA, by UV irradiation of Escherichia coli ribosomal subunits. | Molecular & general genetics : MGG | 102 | 814400 |
| 1980 | E. coli ribosomal protein L4 is a feedback regulatory protein. | Cell | 100 | 6996835 |
| 1999 | CD39-L4 is a secreted human apyrase, specific for the hydrolysis of nucleoside diphosphates. | The Journal of biological chemistry | 96 | 10400613 |
| 1990 | The adenovirus L4 100-kilodalton protein is necessary for efficient translation of viral late mRNA species. | Journal of virology | 95 | 2335816 |
| 2000 | Identification and characterization of KLK-L4, a new kallikrein-like gene that appears to be down-regulated in breast cancer tissues. | The Journal of biological chemistry | 81 | 10766816 |
| 2000 | Translational control of ribosomal protein L4 mRNA is required for rapid neurite regeneration. | Neurobiology of disease | 79 | 10964612 |
| 2007 | Novel mutations in ribosomal proteins L4 and L22 that confer erythromycin resistance in Escherichia coli. | Molecular microbiology | 77 | 17956547 |
| 2006 | ATF3 expression in L4 dorsal root ganglion neurons after L5 spinal nerve transection. | The European journal of neuroscience | 77 | 16420444 |
| 2002 | Susceptibilities to telithromycin and six other agents and prevalence of macrolide resistance due to L4 ribosomal protein mutation among 992 Pneumococci from 10 central and Eastern European countries. | Antimicrobial agents and chemotherapy | 70 | 11796344 |
| 2019 | LncRNA SNHG7 accelerates the proliferation, migration and invasion of hepatocellular carcinoma cells via regulating miR-122-5p and RPL4. | Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie | 66 | 31545291 |
| 1988 | RNA-protein cross-linking in Escherichia coli 50S ribosomal subunits; determination of sites on 23S RNA that are cross-linked to proteins L2, L4, L24 and L27 by treatment with 2-iminothiolane. | Nucleic acids research | 66 | 3278299 |
| 2001 | Adenovirus L4-100K assembly protein is a granzyme B substrate that potently inhibits granzyme B-mediated cell death. | Immunity | 65 | 11420045 |
| 1980 | Precise localization of the site of cross-linking between protein L4 and 23S ribonucleic acid induced by mild ultraviolet irradiation of Escherichia coli 50S ribosomal subunits. | Biochemistry | 65 | 6998491 |
| 1998 | Involvement of the Xenopus laevis Ro60 autoantigen in the alternative interaction of La and CNBP proteins with the 5'UTR of L4 ribosomal protein mRNA. | Journal of molecular biology | 64 | 9710533 |
| 2015 | Coordinated Ribosomal L4 Protein Assembly into the Pre-Ribosome Is Regulated by Its Eukaryote-Specific Extension. | Molecular cell | 63 | 25936803 |
| 2009 | Regulation of ribonuclease E activity by the L4 ribosomal protein of Escherichia coli. | Proceedings of the National Academy of Sciences of the United States of America | 63 | 19144914 |
| 2015 | The Dedicated Chaperone Acl4 Escorts Ribosomal Protein Rpl4 to Its Nuclear Pre-60S Assembly Site. | PLoS genetics | 61 | 26447800 |
| 2006 | The L4 22-kilodalton protein plays a role in packaging of the adenovirus genome. | Journal of virology | 60 | 16809303 |
| 2004 | Activation of the early-late switch in adenovirus type 5 major late transcription unit expression by L4 gene products. | Journal of virology | 59 | 14747543 |
| 2015 | Small GTPase Rab2B and Its Specific Binding Protein Golgi-associated Rab2B Interactor-like 4 (GARI-L4) Regulate Golgi Morphology. | The Journal of biological chemistry | 56 | 26209634 |
| 1992 | Identification and transcriptional analyses of the UL3 and UL4 genes of equine herpesvirus 1, homologs of the ICP27 and glycoprotein K genes of herpes simplex virus. | Journal of virology | 54 | 1323700 |
| 1999 | Translational effects of mutations and polymorphisms in a repressive upstream open reading frame of the human cytomegalovirus UL4 gene. | Journal of virology | 49 | 10482583 |
| 1995 | Nucleotide sequence analysis of a 30-kb region of the bovine herpesvirus 1 genome which exhibits a colinear gene arrangement with the UL21 to UL4 genes of herpes simplex virus. | Virology | 44 | 7793062 |
| 2016 | Ribosomal protein L4 is a novel regulator of the MDM2-p53 loop. | Oncotarget | 43 | 26908445 |
| 2021 | The Zinc Finger Antiviral Protein ZAP Restricts Human Cytomegalovirus and Selectively Binds and Destabilizes Viral UL4/UL5 Transcripts. | mBio | 42 | 33947766 |
| 1990 | Escherichia coli ribosomal protein L4 stimulates transcription termination at a specific site in the leader of the S10 operon independent of L4-mediated inhibition of translation. | Journal of molecular biology | 41 | 1692593 |
| 2019 | Proteome profiling of L3 and L4 Anisakis simplex development stages by TMT-based quantitative proteomics. | Journal of proteomics | 40 | 30978463 |
| 2006 | L4-33K, an adenovirus-encoded alternative RNA splicing factor. | The Journal of biological chemistry | 40 | 17028184 |
| 2015 | Increased expression of HCN2 channel protein in L4 dorsal root ganglion neurons following axotomy of L5- and inflammation of L4-spinal nerves in rats. | Neuroscience | 39 | 25813712 |
| 2010 | Adenovirus late-phase infection is controlled by a novel L4 promoter. | Journal of virology | 39 | 20444889 |
| 2000 | Crystal structure of ribosomal protein L4 shows RNA-binding sites for ribosome incorporation and feedback control of the S10 operon. | The EMBO journal | 39 | 10698923 |
| 1990 | Ribosomal protein L4 stimulates in vitro termination of transcription at a NusA-dependent terminator in the S10 operon leader. | Proceedings of the National Academy of Sciences of the United States of America | 39 | 2157208 |
| 2001 | Identification and molecular characterization of five novel kallikrein gene 13 (KLK13; KLK-L4) splice variants: differential expression in the human testis and testicular cancer. | Anticancer research | 38 | 11848466 |
| 2012 | Failure of translation of human adenovirus mRNA in murine cancer cells can be partially overcome by L4-100K expression in vitro and in vivo. | Molecular therapy : the journal of the American Society of Gene Therapy | 37 | 22735379 |
| 2004 | The coat protein of tobamovirus acts as elicitor of both L2 and L4 gene-mediated resistance in Capsicum. | The Journal of general virology | 37 | 15218193 |
| 2006 | The adenovirus L4 33-kilodalton protein binds to intragenic sequences of the major late promoter required for late phase-specific stimulation of transcription. | Journal of virology | 36 | 17093188 |
| 2004 | Multiple defects in translation associated with altered ribosomal protein L4. | Nucleic acids research | 35 | 15509870 |
| 2007 | Molecular cloning and characterization of the gene encoding cold-active beta-galactosidase from a psychrotrophic and halotolerant Planococcus sp. L4. | Journal of agricultural and food chemistry | 34 | 17326654 |
| 2007 | Ternary complex formation on the adenovirus packaging sequence by the IVa2 and L4 22-kilodalton proteins. | Journal of virology | 34 | 17804492 |
| 1998 | Overexpression of ribosomal proteins L4 and L5 and the putative alternative elongation factor PTI-1 in the doxorubicin resistant human colon cancer cell line LoVoDxR. | European journal of cancer (Oxford, England : 1990) | 34 | 9713282 |
| 2012 | The adenovirus L4-22K protein is multifunctional and is an integral component of crucial aspects of infection. | Journal of virology | 32 | 22811519 |
| 1999 | Phylogenetic analysis of L4-mediated autogenous control of the S10 ribosomal protein operon. | Journal of bacteriology | 32 | 10498727 |
| 2015 | Regulation of human adenovirus alternative RNA splicing by the adenoviral L4-33K and L4-22K proteins. | International journal of molecular sciences | 31 | 25636034 |
| 2003 | RNA-structural mimicry in Escherichia coli ribosomal protein L4-dependent regulation of the S10 operon. | The Journal of biological chemistry | 31 | 12738792 |
| 2013 | Interaction of nascent chains with the ribosomal tunnel proteins Rpl4, Rpl17, and Rpl39 of Saccharomyces cerevisiae. | The Journal of biological chemistry | 30 | 24072706 |
| 2007 | Invariant Ser211 is involved in the catalysis of PD-L4, type I RIP from Phytolacca dioica leaves. | Proteins | 30 | 17243169 |
| 1993 | RNA-binding properties of a translational activator, the adenovirus L4 100-kilodalton protein. | Journal of virology | 30 | 8497066 |
| 2018 | Combined exposure to methylmercury and manganese during L1 larval stage causes motor dysfunction, cholinergic and monoaminergic up-regulation and oxidative stress in L4 Caenorhabditis elegans. | Toxicology | 29 | 30336192 |
| 1999 | The role of the L4 33K gene in adenovirus infection. | Virology | 29 | 10544122 |
| 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 | 28 | 32971090 |
| 1999 | Colocalization of the herpes simplex virus 1 UL4 protein with infected cell protein 22 in small, dense nuclear structures formed prior to onset of DNA synthesis. | Journal of virology | 28 | 10233976 |
| 2019 | Unilateral L4-dorsal root ganglion stimulation evokes pain relief in chronic neuropathic postsurgical knee pain and changes of inflammatory markers: part II whole transcriptome profiling. | Journal of translational medicine | 27 | 31217010 |
| 2017 | Molecular basis for protection of ribosomal protein L4 from cellular degradation. | Nature communications | 27 | 28148929 |
| 2016 | The extended loops of ribosomal proteins uL4 and uL22 of Escherichia coli contribute to ribosome assembly and protein translation. | Nucleic acids research | 27 | 27257065 |
| 1992 | Ribosomal protein L4 and transcription factor NusA have separable roles in mediating terminating of transcription within the leader of the S10 operon of Escherichia coli. | Genes & development | 27 | 1285127 |
| 1991 | The organization and expression of the Saccharomyces cerevisiae L4 ribosomal protein genes and their identification as the homologues of the mammalian ribosomal protein gene L7a. | Molecular & general genetics : MGG | 27 | 2046660 |
| 2020 | Impact of L4 lymph node dissection on long-term survival in left-side operable non-small-cell lung cancer: a propensity score matching study. | European journal of cardio-thoracic surgery : official journal of the European Association for Cardio-thoracic Surgery | 26 | 32016340 |
| 2015 | Molecular characterisation of new organisation of plnEF and plw loci of bacteriocin genes harbour concomitantly in Lactobacillus plantarum I-UL4. | Microbial cell factories | 26 | 26077560 |
| 2000 | Differentiation of human cytomegalovirus genotypes in immunocompromised patients on the basis of UL4 gene polymorphisms. | The Journal of infectious diseases | 25 | 11110652 |
| 2019 | Propriospinal Neurons of L3-L4 Segments Involved in Control of the Rat External Urethral Sphincter. | Neuroscience | 24 | 31785359 |
| 2016 | Ribosome Protein L4 is essential for Epstein-Barr Virus Nuclear Antigen 1 function. | Proceedings of the National Academy of Sciences of the United States of America | 24 | 26858444 |
| 2014 | Deletion of L4 domains reveals insights into the importance of ribosomal protein extensions in eukaryotic ribosome assembly. | RNA (New York, N.Y.) | 24 | 25246649 |
| 1995 | Role of NusA in L4-mediated attenuation control of the S10 r-protein operon of Escherichia coli. | Journal of molecular biology | 24 | 7844821 |
| 1994 | Identification of the pseudorabies virus UL4 and UL5 (helicase) genes. | Virology | 24 | 8030256 |
| 2022 | TTC22 promotes m6A-mediated WTAP expression and colon cancer metastasis in an RPL4 binding-dependent pattern. | Oncogene | 22 | 35798874 |
| 2005 | Mutation of L4 ribosomal protein conferring unusual macrolide resistance in two independent clinical isolates of Staphylococcus aureus. | Microbial drug resistance (Larchmont, N.Y.) | 21 | 15770089 |
| 2015 | Ribosomal protein L4 interacts with viral protein VP3 and regulates the replication of infectious bursal disease virus. | Virus research | 20 | 26415754 |
| 2009 | Arginine methylation of human adenovirus type 5 L4 100-kilodalton protein is required for efficient virus production. | Journal of virology | 20 | 19264777 |
| 1992 | Primary structures of ribosomal proteins L3 and L4 from Bacillus stearothermophilus. | European journal of biochemistry | 20 | 1499563 |
| 2013 | Adenoviral E2 IVa2 protein interacts with L4 33K protein and E2 DNA-binding protein. | The Journal of general virology | 19 | 23388198 |
| 1995 | RNA determinants required for L4-mediated attenuation control of the S10 r-protein operon of Escherichia coli. | Journal of molecular biology | 19 | 7531246 |
| 2010 | Mutations in 23S rRNA and ribosomal protein L4 account for resistance in Chlamydia trachomatis strains selected in vitro by macrolide passage. | Andrologia | 18 | 20629652 |
| 2005 | Functional interaction between RNA helicase II/Gu(alpha) and ribosomal protein L4. | The FEBS journal | 18 | 16045751 |
| 1990 | Characteristics of PHA-L4, the mitogenic isolectin of phytohemagglutinin, as an ideal biologic response modifier. | Molecular biotherapy | 18 | 2185793 |
| 2019 | Effects of microencapsulated olfactory ensheathing cell transplantation on neuropathic pain and P2X7 receptor expression in the L4-5 spinal cord segment. | Neuroscience letters | 17 | 30753909 |
| 2010 | Adenovirus L4-22K stimulates major late transcription by a mechanism requiring the intragenic late-specific transcription factor-binding site. | Virus research | 17 | 20621673 |
| 1979 | Cat L4-S1 dermatomes determined using signal averaging. | Neuroscience letters | 17 | 460741 |
| 2006 | Linker chains of the gigantic hemoglobin of the earthworm Lumbricus terrestris: primary structures of linkers L2, L3, and L4 and analysis of the connectivity of the disulfide bonds in linker L1. | Proteins | 16 | 16425180 |
| 2023 | Silencing of the immune gene BmPGRP-L4 in the midgut affects the growth of silkworm (Bombyx mori) larvae. | Insect molecular biology | 15 | 36705338 |
| 2021 | LIFU Alleviates Neuropathic Pain by Improving the KCC2 Expression and Inhibiting the CaMKIV-KCC2 Pathway in the L4-L5 Section of the Spinal Cord. | Neural plasticity | 15 | 33953740 |
| 2015 | Laminin L4 domain structure resembles adhesion modules in ephrin receptor and other transmembrane glycoproteins. | The FEBS journal | 15 | 25962468 |
| 2003 | Protein disulphide isomerase of Ostertagia ostertagi: an excretory-secretory product of L4 and adult worms? | International journal for parasitology | 15 | 12633650 |
| 2000 | Activation of transcription of the human cytomegalovirus early UL4 promoter by the Ets transcription factor binding element. | Journal of virology | 15 | 11024111 |
| 1996 | Ribosomal protein L4 from Escherichia coli utilizes nonidentical determinants for its structural and regulatory functions. | RNA (New York, N.Y.) | 15 | 8846294 |
| 2022 | Low-Intensity Focused Ultrasound Alleviates Spasticity and Increases Expression of the Neuronal K-Cl Cotransporter in the L4-L5 Sections of Rats Following Spinal Cord Injury. | Frontiers in cellular neuroscience | 14 | 35634464 |
| 2022 | Macrolide therapy in Pseudomonas aeruginosa infections causes uL4 ribosomal protein mutations leading to high-level resistance. | Clinical microbiology and infection : the official publication of the European Society of Clinical Microbiology and Infectious Diseases | 13 | 35988850 |
| 2021 | A Water Soluble Pd2 L4 Cage for Selective Binding of Neu5Ac. | Chemistry (Weinheim an der Bergstrasse, Germany) | 13 | 34486179 |
| 2015 | Adenoviral L4 33K forms ring-like oligomers and stimulates ATPase activity of IVa2: implications in viral genome packaging. | Frontiers in microbiology | 13 | 25954255 |
| 2015 | Expression of Genes Encoding the Enzymes for Glycogen and Trehalose Metabolism in L3 and L4 Larvae of Anisakis simplex. | Journal of parasitology research | 13 | 26783451 |
| 1998 | Yeast ribosomal proteins L4, L17, L20, and L25 exhibit different binding characteristics for the yeast 35S precursor rRNA. | Biochimica et biophysica acta | 13 | 9838082 |
| 1993 | Human ribosomal protein L4: cloning and sequencing of the cDNA and primary structure of the protein. | Biochimica et biophysica acta | 13 | 8268230 |
| 1991 | Ribosomal protein L4 of Escherichia coli: in vitro analysis of L4-mediated attenuation control. | Biochimie | 13 | 1764518 |
| 1978 | Inhibitory effects of phytohemagglutinin isolectins L4 and E4 on L1210 cells. | Cancer research | 13 | 565679 |
| 2022 | Tandem Mass Tagging (TMT) Reveals Tissue-Specific Proteome of L4 Larvae of Anisakis simplex s. s.: Enzymes of Energy and/or Carbohydrate Metabolism as Potential Drug Targets in Anisakiasis. | International journal of molecular sciences | 12 | 35457153 |
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