Affinage

NONO

Non-POU domain-containing octamer-binding protein · UniProt Q15233

Length
471 aa
Mass
54.2 kDa
Annotated
2026-04-29
100 papers in source corpus 40 papers cited in narrative 39 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

NONO (p54nrb) is a multifunctional nuclear RNA- and DNA-binding protein of the DBHS family that couples transcription by RNA polymerase II to pre-mRNA splicing, 3′-end cleavage, and transcription termination, while also participating in DNA double-strand break repair and transcriptional co-regulation. As an obligate heterodimer with PSF (SFPQ), NONO binds the RNA Pol II CTD and U5 snRNA/spliceosomal snRNPs to coordinate cotranscriptional RNA processing, recruits XRN2 for torpedo-model transcription termination, stimulates polyadenylation as a component of the snRNP-free U1A complex, and promotes NHEJ by cooperating with Ku to align DNA ends (PMID:12358429, PMID:17639083, PMID:16373496, PMID:15590677). NONO is an essential structural component of NEAT1-dependent paraspeckles, where the NONO–PSF–matrin 3 complex selectively retains hyperedited inosine-containing mRNAs in the nucleus; CARM1-mediated methylation of the NONO coiled-coil domain attenuates this RNA binding and releases retained transcripts (PMID:11525732, PMID:19716791, PMID:25792598). NONO also functions as a context-dependent transcriptional coactivator or corepressor for nuclear receptors (AR, PR, TR/RXR, SF-1) and other factors (Sox9, CREB/TORC2, SREBP-1a), recruiting Sin3A/HDAC complexes for repression or stabilizing activator complexes, with physiological consequences including regulation of chondrocyte differentiation, rod photoreceptor survival, lipogenesis, and early mouse embryonic cell-fate specification (PMID:11259580, PMID:18677406, PMID:24301678, PMID:26148231, PMID:30550788).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 1993 High

    Identification of p54nrb as a dual-RRM nuclear protein sharing the DBHS domain with PSF established it as a candidate RNA-processing factor.

    Evidence Protein purification from HeLa nuclear extract and cDNA cloning with sequence analysis

    PMID:8371983

    Open questions at the time
    • No functional assay demonstrated; role in splicing was inferred from domain homology
  2. 1996 Medium

    Discovery that Spi-1/PU.1 competes with RNA for p54nrb binding and alters splicing in vitro provided the first evidence that p54nrb participates directly in pre-mRNA splicing.

    Evidence In vitro RNA-binding competition and splicing assays with purified proteins

    PMID:8626664

    Open questions at the time
    • In vitro system only; no in vivo splicing targets identified
    • Single study without independent replication
  3. 1998 High

    Demonstration that the PSF–p54nrb heterodimer stimulates topoisomerase I catalytic activity and DNA strand transfer revealed that the complex has direct biochemical activity on DNA topology, beyond RNA processing.

    Evidence In vitro topoisomerase activity assays with reconstituted purified complex

    PMID:10858305 PMID:9756848

    Open questions at the time
    • Physiological relevance of topoisomerase I stimulation not established in vivo
  4. 2001 High

    Two contemporaneous discoveries established that the NONO–PSF complex both retains hyperedited mRNAs in the nucleus and functions as a transcriptional corepressor via Sin3A/HDAC recruitment to nuclear receptors, revealing dual post-transcriptional and transcriptional roles.

    Evidence Nuclear extract fractionation with inosine-RNA binding and Xenopus oocyte retention assay; co-IP with Sin3A and HDAC inhibitor treatment with reporter assays

    PMID:11259580 PMID:11525732

    Open questions at the time
    • Structural basis for selective recognition of inosine-containing RNA unknown
    • Whether corepressor function requires the PSF partner or is p54nrb-autonomous was unclear
  5. 2002 High

    Identification of NONO and PSF as paraspeckle components and as Pol II CTD-binding proteins that simultaneously engage RNA established the physical framework for cotranscriptional RNA processing.

    Evidence Live-cell imaging of paraspeckles; CTD affinity chromatography and co-IP with Pol II

    PMID:11790299 PMID:12358429 PMID:12403470

    Open questions at the time
    • Paraspeckle function was unknown at this point
    • Whether CTD binding and snRNP association are simultaneous or sequential was not resolved
  6. 2004 High

    Reconstitution of NHEJ stimulation by the PSF–p54nrb–Ku complex and demonstration that NONO associates with 5′ splice sites on elongating Pol II complexes expanded the functional repertoire to DNA repair and cotranscriptional splicing.

    Evidence In vitro NHEJ reconstitution with purified proteins; immobilized DNA template assay with HeLa nuclear extracts

    PMID:15057275 PMID:15590677

    Open questions at the time
    • In vivo DSB repair role not yet demonstrated
    • Whether NONO itself contacts DNA ends or acts as scaffold was unclear
  7. 2005 High

    Establishing that NONO is required for activator/CTD-stimulated splicing and 3′-end cleavage, and that its heterodimerization with PSP1 is needed for paraspeckle targeting, linked its molecular interactions to functional RNA-processing outcomes.

    Evidence Immunodepletion/reconstitution polyadenylation assays; domain mutagenesis with live-cell paraspeckle imaging

    PMID:15701524 PMID:16024807 PMID:16148043

    Open questions at the time
    • NEAT1 dependence of paraspeckles not yet discovered
    • Cdk1 phosphorylation sites fully mapped but functional consequences during mitosis not characterized
  8. 2007 High

    Discovery that p54nrb recruits XRN2 for transcription termination and regulates alternative splicing of caspase-9 and ARE-containing mRNAs demonstrated its role in termination and post-transcriptional gene regulation.

    Evidence ChIP along transcribed genes, siRNA knockdown with termination readthrough; co-IP and alternative splicing assays

    PMID:17639083 PMID:17965020 PMID:32332923

    Open questions at the time
    • Whether XRN2 recruitment is direct or via an RNA intermediate was not resolved
    • Specificity of ARE binding relative to PSF contribution unclear
  9. 2008 High

    In vivo evidence that NONO loss causes dwarfism in transgenic mice (via Sox9 target gene disruption) and interacts with SWI/SNF to regulate TERT splicing demonstrated physiological consequences of NONO dysfunction.

    Evidence Transgenic mouse model with RRM-deletion; shRNA knockdown with RT-PCR splicing analysis and ChIP

    PMID:18042045 PMID:18677406

    Open questions at the time
    • Whether chondrocyte phenotype reflects splicing defects, transcriptional coactivation, or both was not separated
  10. 2009 High

    Identification of NEAT1 as the essential architectural RNA of paraspeckles, whose disruption releases NONO-retained hyperedited mRNAs, and demonstration that NONO loss impairs DSB repair in vivo, completed the picture of NONO's dual nuclear retention and DNA repair functions.

    Evidence NEAT1 knockdown with mRNA export assays; γ-H2AX focus assay, chromosomal aberration analysis, and clonogenic survival after IR

    PMID:19188602 PMID:19716791 PMID:19759212

    Open questions at the time
    • Whether NONO contributes to HR in addition to NHEJ was unknown
    • Mechanism of NONO recruitment to DSBs was unclear — PSF drives relocalization, not NONO itself
  11. 2015 High

    Discovery that CARM1 methylates the NONO coiled-coil domain to attenuate RNA binding and reduce paraspeckle-mediated mRNA retention provided the first post-translational regulatory mechanism controlling NONO's RNA-retention function.

    Evidence CARM1 KO, methylation assays, RNA-IP, and nuclear/cytoplasmic fractionation

    PMID:25792598

    Open questions at the time
    • Whether methylation affects other NONO functions (NHEJ, transcriptional coregulation) was not tested
    • Demethylase that reverses this mark unknown
  12. 2018 High

    Demonstration that NONO and NEAT1 are required for CARM1-dependent H3R26 methylation and inner-cell-mass specification in mouse embryos established a developmental role upstream of epigenetic programming.

    Evidence siRNA microinjection into mouse embryos with epistasis analysis and developmental assays

    PMID:30550788

    Open questions at the time
    • Mechanism by which paraspeckle-localized NONO influences CARM1 histone methylation not fully defined
    • Whether NONO's RNA-retention versus transcriptional function is critical for this phenotype is unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the structural basis by which the NONO–PSF heterodimer selectively recognizes inosine-containing versus unedited RNA, the full spectrum of NONO-dependent alternative splicing events in normal tissues, and how Cdk1-dependent mitotic phosphorylation functionally alters NONO activity during cell division.
  • No high-resolution structure of NONO–RNA complex with inosine substrate
  • Genome-wide map of NONO-dependent splicing events in primary tissues lacking
  • Functional consequences of mitotic phosphorylation on NONO's multiple activities untested

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 9 GO:0003723 RNA binding 6 GO:0003677 DNA binding 4 GO:0060090 molecular adaptor activity 3 GO:0098772 molecular function regulator activity 3
Localization
GO:0005634 nucleus 4 GO:0005654 nucleoplasm 4 GO:0005730 nucleolus 1
Pathway
R-HSA-74160 Gene expression (Transcription) 11 R-HSA-8953854 Metabolism of RNA 8 R-HSA-1266738 Developmental Biology 2 R-HSA-73894 DNA Repair 2
Partners
CARM1MATR3NEAT1PSPC1SFPQSREBF1TOP1XRN2
Complex memberships
DBHS heterodimer (NONO-SFPQ/PSF)Paraspeckle (NONO-SFPQ-NEAT1)snRNP-free U1A (SF-A) complex

Evidence

Reading pass · 39 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1993 p54nrb was purified from HeLa cells and found to contain two RNA recognition motifs (RRMs); it shares 71% identity with splicing factor PSF within a 320 aa region including both RRMs, defining the conserved DBHS (Drosophila behavior, human splicing) domain. The protein was identified by cross-reactivity with antibodies against yeast splicing factor PRP18. Protein purification, cDNA cloning, sequence analysis Nucleic acids research High 8371983
2001 p54nrb forms a multiprotein complex with PSF and matrin 3 that binds specifically and cooperatively to inosine-containing (hyperedited) RNAs and anchors them to the nuclear matrix, thereby retaining promiscuously A-to-I edited RNAs in the nucleus and preventing their export. HeLa cell nuclear extract fractionation, affinity purification, RNA-binding assays, Xenopus oocyte retention assay Cell High 11525732
2002 p54nrb/NonO and PSF are components of paraspeckles, a novel subnuclear compartment in the interchromatin nucleoplasmic space; both proteins interact dynamically with nucleoli in a transcription-dependent manner and relocalize to nucleolar cap structures upon transcription inhibition. Immunofluorescence, live-cell imaging with YFP-PSP1 stable cell line, transcription inhibition experiments Current biology : CB High 11790299
2002 PSF and p54nrb/NonO bind specifically to the CTD of RNA polymerase II (both hypo- and hyperphosphorylated forms), co-purify with a pol II holoenzyme, and can interact with the CTD and RNA simultaneously, providing a physical link between pol II and pre-mRNA processing. Affinity chromatography on immobilized CTD matrices, co-immunoprecipitation with pol II antibodies, RNA-binding assays RNA (New York, N.Y.) High 12358429
2002 PSF and p54nrb bind U5 snRNA at stem 1b in a sequence- and structure-dependent manner; both proteins associate with spliceosomes and with U4/U6.U5 tri-snRNP. SELEX/iterative selection, filter-binding assays, RNA affinity selection, sedimentation analysis RNA (New York, N.Y.) High 12403470
2004 p54nrb associates with the 5' splice site within large complexes containing hyperphosphorylated RNA pol II (RNAPIIO) and U1/U2 snRNPs; p54nrb and PSF can interact directly with the 5' splice site, indicating they mediate contacts between RNAPII and snRNPs during coupled transcription/splicing. HeLa nuclear extract pull-down, immobilized DNA template assay, co-immunoprecipitation The EMBO journal High 15057275
2004 The PSF·p54nrb heterodimeric complex stimulates DNA double-strand break rejoining in vitro by cooperating with Ku protein to form a preligation complex with substrate DNA; the four RRM domains in the heterodimer are proposed to align separate DNA molecules. In vitro NHEJ reconstitution assay with purified proteins, DNA-binding studies The Journal of biological chemistry High 15590677
2005 p54nrb forms a heterodimer with PSP1 in vivo; the interaction domain in PSP1 is required for its correct localization to paraspeckles, and an RNA-binding-competent RRM is also required for paraspeckle targeting. Paraspeckle formation depends on ongoing RNA Pol II transcription. Co-immunoprecipitation, domain mapping, DRB transcription inhibition, live-cell imaging Molecular biology of the cell High 16148043
2005 PSF and p54nrb preferentially bind strong transcriptional activation domains and are required for activator- and CTD-mediated stimulation of pre-mRNA splicing and 3'-end cleavage; PSF overexpression bypasses the requirement for a strong activator. Affinity chromatography, overexpression in 293 cells, CTD truncation experiments, splicing assays Molecular and cellular biology High 16024807
2007 p54nrb/NonO (p54) physically associates with XRN2 and is required for degradation of 3'-cleaved RNA in vitro; p54 is present along the length of transcribed genes and its siRNA-mediated knockdown leads to defects in XRN2 recruitment and transcription termination. Co-immunoprecipitation, in vitro 3'-processing assays, ChIP, siRNA knockdown Genes & development High 17639083
2007 NONO/p54nrb was identified as a TORC2-interacting protein in the cAMP signaling pathway; NONO is necessary for cAMP-dependent activation of CREB target genes and acts as a bridge between the CREB/TORC complex and RNA polymerase II. High-throughput coactivator trap screen, RNAi knockdown, ChIP Proceedings of the National Academy of Sciences of the United States of America Medium 18077367
2007 PSF and p54nrb bind AU-rich element (ARE)-containing RNAs including TNFα mRNA; Mnk kinases phosphorylate PSF at two sites, and Mnk-mediated phosphorylation increases PSF binding to TNFα mRNA in living cells. Proteomic substrate screen, in vitro kinase assays, in vivo phosphorylation, RNA immunoprecipitation The Journal of biological chemistry Medium 17965020
2008 p54nrb physically interacts with Sox9 transcription factor, enhances Sox9-dependent transcriptional activation of the Col2a1 promoter, and is required for maturation of Sox9-target gene mRNA. Loss of p54nrb RNA recognition motifs disrupts paraspeckle bodies, inhibits chondrocyte differentiation, and causes dwarfism in transgenic mice. Co-immunoprecipitation, reporter assays, siRNA knockdown, transgenic mouse model The Journal of clinical investigation High 18677406
2009 Paraspeckle assembly and function depend on the long noncoding RNA NEAT1; NEAT1 is absent in hESCs and induced upon differentiation, coinciding with paraspeckle formation. Knockdown of NEAT1 in HeLa cells abolishes paraspeckles and enhances nucleocytoplasmic export of mRNAs containing inverted Alu repeats that are normally retained in paraspeckle-associated complexes containing p54nrb. NEAT1 knockdown, siRNA, mRNA export assays, immunofluorescence Molecular cell High 19716791
2009 MENβ (NEAT1_2) is stabilized by selective association with p54nrb and PSF, which is required for paraspeckle structural integrity; knockdown of MENβ leads to paraspeckle disintegration and prevents reassembly after transcriptional arrest. RNAi knockdown, immunoprecipitation, transcriptional arrest/recovery assays Proceedings of the National Academy of Sciences of the United States of America High 19188602
2009 p54nrb is phosphorylated during mitosis at its carboxy-terminal extremity by Cdk1 and at amino-terminal sites by a distinct kinase; p54nrb interacts with the peptidylprolyl isomerase Pin1 through three phosphorylated threonine residues in its proline-rich C-terminus, suggesting regulation by phosphorylation-dependent conformational changes upon mitotic entry. Mitosis-specific phospho-antibodies (CC-3, MPM-2), CDK inhibitor (roscovitine), immunodepletion of cyclin B1, site-directed mutagenesis, Pin1 interaction assays Journal of molecular biology High 15701524
2009 p54nrb contributes to DNA double-strand break repair in vivo; siRNA-mediated p54nrb knockdown delays DSB repair (γ-H2AX focus assay), increases ionizing radiation-induced chromosomal aberrations, and increases radiosensitivity in clonogenic survival assays. siRNA knockdown, γ-H2AX focus assay, chromosomal aberration analysis, clonogenic survival assay Nucleic acids research High 19759212
2001 PSF acts as a transcriptional corepressor through interaction with Sin3A; both PSF and NonO/p54nrb interact with the DNA-binding domains of nuclear hormone receptors (TR and RXR) and mediate silencing through HDAC recruitment to the receptor DBD. Biochemical purification from nuclear extracts, co-immunoprecipitation, reporter assays, HDAC inhibitor treatment Molecular and cellular biology Medium 11259580
2002 p54nrb/NonO (identified as IPEB) functions as a transcriptional activator that binds DNA at the IAP proximal enhancer element; its N-terminal half contains the DNA-binding domain and C-terminal half contains the activation domain. Protein purification, cDNA cloning, reporter gene assays, domain deletion analysis Molecular and cellular biology Medium 9001221
2002 p54nrb/NonO and PSF form a complex with SF-1 that binds the hCYP17 promoter; PSF represses basal transcription by interacting with mSin3A, while the SF-1/PSF/p54nrb/NonO complex mediates cAMP-induced transcriptional activation; PSF-p54nrb interaction with mSin3A recruits HDAC activity. EMSA, co-immunoprecipitation, reporter gene assays, HDAC inhibitor (trichostatin A) treatment Endocrinology Medium 11897684
1998 The PSF/p54nrb complex directly interacts with DNA topoisomerase I; the complex stimulates topoisomerase I catalytic activity 16-fold, and PSF/p54nrb stimulates the ability of topoisomerase I to transfer ('jump') between separate DNA helices without affecting cleavage or religation half-reactions. Co-purification, co-immunoprecipitation, far Western blotting, in vitro topoisomerase activity assays, chemical crosslinking The Journal of biological chemistry High 10858305 9756848
2007 p54nrb interacts with PSF in a complex that modulates the activity of SR protein SRSF2 on caspase-9 RNA splicing, decreasing SRSF2 binding to caspase-9 RNA and favoring expression of the antiapoptotic alternatively spliced form. Co-immunoprecipitation, RNA-binding assays, alternative splicing assays, siRNA knockdown Oncogene Medium 32332923
2015 CARM1 methylates the coiled-coil domain of p54nrb, resulting in reduced binding of p54nrb to mRNAs containing inverted repeated Alu elements (IRAlus), thus attenuating nuclear retention of these transcripts. CARM1 also suppresses NEAT1 transcription, reducing paraspeckle formation. CARM1 knockdown/knockout, RNA immunoprecipitation, methylation assays, mRNA nuclear/cytoplasmic fractionation Genes & development High 25792598
2018 In mouse embryos, p54nrb and NEAT1 are required for CARM1 association with paraspeckles and for H3R26 methylation; depletion of p54nrb causes arrest at the 16- to 32-cell stage with elevated Cdx2 expression, indicating paraspeckles/p54nrb act upstream of CARM1 in early embryonic cell fate specification. Microinjection of siRNA into mouse embryos, immunofluorescence, embryonic developmental assays Cell High 30550788
2006 N-WASP forms a large nuclear complex with PSF-NonO (p54nrb), nuclear actin, and RNA polymerase II; interaction of N-WASP with PSF-NonO couples N-WASP to RNA pol II and regulates transcription; nuclear actin polymerization promoted by N-WASP is important in this process. Co-immunoprecipitation, nuclear fractionation, knockdown/overexpression with transcription readout Nature cell biology Medium 16767080
2006 p54nrb is a component of the snRNP-free U1A (SF-A) complex; immunodepletion and reconstitution experiments show that p54nrb is critical for pre-mRNA cleavage during polyadenylation. TAP-tag purification, mass spectrometry, immunodepletion, reconstitution assays RNA (New York, N.Y.) High 16373496
2007 p54nrb/NonO and three interacting proteins (hnRNP M, Ywhaz, Ppp1ca) bind to the rhodopsin distal enhancer region and augment rhodopsin expression; shRNA knockdown of NonO in mouse retina leads to loss of rhodopsin expression and rod cell death, with splicing defects in phototransduction genes. Mass spectrometry, ChIP-seq, shRNA knockdown, RNA-seq, rescue experiments Human molecular genetics High 24301678
2009 p54nrb functions as a transcriptional corepressor of the progesterone receptor (PR) independent of PSF; p54nrb directly interacts with PR, recruits mSin3A through its N-terminus to the PR-DNA complex, and inhibits PR-mediated transactivation. Knockdown of p54nrb alleviates PR-mediated repression of connexin 43 (Cx43/Gja1) transcription. Co-immunoprecipitation, reporter gene assays, siRNA knockdown, ChIP, overexpression Molecular endocrinology (Baltimore, Md.) Medium 19423654
2003 p54nrb interacts with the AF-1 (N-terminal A/B) domain of the androgen receptor (AR) in a ligand-dependent manner and potentiates AR AF-1 transcriptional activity; p54nrb was purified from HEK293 cells along with PSF, PSP1, and PSP2 as AR AF-1 region-interacting proteins. Biochemical purification, MALDI-TOF MS, co-immunoprecipitation, reporter gene assays Biochemical and biophysical research communications Medium 12810069
2007 p54nrb and PSF synergistically form protein complexes with androgen receptor (AR) in a ligand-independent manner, interact directly with mSin3A, and inhibit AR transcriptional activity through HDAC recruitment; suppression is reversed by HDAC inhibition. Co-immunoprecipitation, reporter gene assays, HDAC inhibitor treatment, siRNA knockdown Molecular and cellular biology Medium 17452459
2008 p54nrb binds to the SWI/SNF complex components BRG1, Brm, and BAF60a through its N-terminal region; together with Brm, p54nrb regulates TERT gene transcription and alternative splicing of TERT exons. Brm and p54nrb co-localize at TERT exon 7 alternative splicing acceptor site. Pull-down assays, shRNA knockdown, RT-PCR for alternative splicing, ChIP, co-localization The Biochemical journal Medium 18042045
2012 YM155 induces disruption of the ILF3/p54nrb complex; p54nrb binds to the survivin promoter and regulates survivin expression; disruption of the ILF3/p54nrb complex by YM155 results in altered subcellular localization of ILF3 and p54nrb. Co-immunoprecipitation, promoter binding assays, small-molecule perturbation Biochemical and biophysical research communications Medium 22842455
2015 p54nrb/NONO interacts with the nuclear form of SREBP-1a through its conserved Y267 residue, increasing nuclear SREBP-1a protein stability and stimulating SREBP-1-mediated transcription of lipogenic genes and lipid production in breast cancer cells. Biochemical screen, co-immunoprecipitation, mutagenesis (Y267), reporter assays, siRNA knockdown, in vivo tumor model Oncogene High 26148231
2013 p54/NONO and PSF heterodimer bind the occludin enhancer element (OEE) in retinal endothelial cells; siRNA knockdown of p54 demonstrates it is necessary for glucocorticoid induction of occludin and claudin-5 expression and for pro-barrier effects. DNA affinity purification, MALDI-TOF MS/MS, ChIP, siRNA knockdown, permeability assays, TER measurement Investigative ophthalmology & visual science Medium 23640037
2020 NONO (p54nrb) promotes oncogenic splicing switch of BIN1 from a short tumor-suppressor isoform to a long isoform (containing exon 12a) in hepatocellular carcinoma; NONO interacts with DHX9 and SFPQ to form a complex that promotes BIN1 exon 12a inclusion. RNA-seq, siRNA knockdown, co-immunoprecipitation, splicing assays Hepatology (Baltimore, Md.) Medium 31815296
2019 Toxic phosphorothioate ASOs form a complex including RNase H1, p54nrb, and PSF; p54nrb binds PS-ASOs via RRM1 and RRM2 domains, while RNase H1 binds via its hybrid-binding domain; RNase H1/p54nrb complexes form only in cells treated with toxic (not safe) PS-ASOs, and the p54nrb core domains are required for association with RNase H1. NanoLuciferase structural complementation reporter, live-cell imaging, in vitro binding assays, domain mutagenesis Nucleic acids research Medium 31495875
2010 PSF (not its dimerization partner p54nrb/NONO) drives relocalization of PSF-containing complexes to sites of dense laser-induced DNA damage in living cells; sequences in PSF's N-terminal domain mediate both radioresistance and DNA damage site relocalization. miRNA-mediated knockdown, genetic rescue with mutant PSF constructs, real-time relocalization assay in live cells, laser-induced DNA damage DNA repair Medium 21144806
2016 p54nrb transcriptionally regulates the lncRNA PCGEM1 by binding its promoter; p54nrb-induced PCGEM1 upregulates AR splice variant AR3, contributing to castration resistance. DIM (3,3'-diindolylmethane) suppresses PCGEM1 expression by preventing p54nrb from binding the PCGEM1 promoter. siRNA/CRISPR knockout, rescue experiments, ChIP, reporter assays, xenograft mouse model Scientific reports Medium 27682980
1996 p54nrb was identified as a binding partner of the transcription factor Spi-1/PU.1; Spi-1/PU.1 impedes the binding of p54nrb to RNA and alters splicing in vitro, suggesting a mechanism for functional interference in erythropoiesis. Immobilized protein binding assay, microsequencing, in vitro RNA-binding competition, in vitro splicing assay The Journal of biological chemistry Medium 8626664

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2009 Altered nuclear retention of mRNAs containing inverted repeats in human embryonic stem cells: functional role of a nuclear noncoding RNA. Molecular cell 552 19716791
2009 MENepsilon/beta noncoding RNAs are essential for structural integrity of nuclear paraspeckles. Proceedings of the National Academy of Sciences of the United States of America 522 19188602
2002 Paraspeckles: a novel nuclear domain. Current biology : CB 448 11790299
2006 Translation repression in human cells by microRNA-induced gene silencing requires RCK/p54. PLoS biology 413 16756390
2001 The fate of dsRNA in the nucleus: a p54(nrb)-containing complex mediates the nuclear retention of promiscuously A-to-I edited RNAs. Cell 404 11525732
2002 PSF and p54(nrb)/NonO--multi-functional nuclear proteins. FEBS letters 297 12417296
2008 Alu element-mediated gene silencing. The EMBO journal 279 18497743
1997 Fusion of splicing factor genes PSF and NonO (p54nrb) to the TFE3 gene in papillary renal cell carcinoma. Oncogene 279 9393982
1998 The African swine fever virus proteins p54 and p30 are involved in two distinct steps of virus attachment and both contribute to the antibody-mediated protective immune response. Virology 201 9568043
2005 P54nrb forms a heterodimer with PSP1 that localizes to paraspeckles in an RNA-dependent manner. Molecular biology of the cell 196 16148043
2008 Enhanced discrimination of African swine fever virus isolates through nucleotide sequencing of the p54, p72, and pB602L (CVR) genes. Virus genes 173 19009341
2001 African swine fever virus protein p54 interacts with the microtubular motor complex through direct binding to light-chain dynein. Journal of virology 170 11559815
2002 Splicing and transcription-associated proteins PSF and p54nrb/nonO bind to the RNA polymerase II CTD. RNA (New York, N.Y.) 159 12358429
2004 p54(nrb) associates with the 5' splice site within large transcription/splicing complexes. The EMBO journal 156 15057275
2001 PSF is a novel corepressor that mediates its effect through Sin3A and the DNA binding domain of nuclear hormone receptors. Molecular and cellular biology 153 11259580
2007 The multifunctional protein p54nrb/PSF recruits the exonuclease XRN2 to facilitate pre-mRNA 3' processing and transcription termination. Genes & development 151 17639083
1993 Purification and cDNA cloning of HeLa cell p54nrb, a nuclear protein with two RNA recognition motifs and extensive homology to human splicing factor PSF and Drosophila NONA/BJ6. Nucleic acids research 151 8371983
2006 Regulation of RNA-polymerase-II-dependent transcription by N-WASP and its nuclear-binding partners. Nature cell biology 137 16767080
1997 Xp54, the Xenopus homologue of human RNA helicase p54, is an integral component of stored mRNP particles in oocytes. Nucleic acids research 128 9023105
1992 Activating transcription factor-2 DNA-binding activity is stimulated by phosphorylation catalyzed by p42 and p54 microtubule-associated protein kinases. Molecular endocrinology (Baltimore, Md.) 119 1337144
2018 CARM1 and Paraspeckles Regulate Pre-implantation Mouse Embryo Development. Cell 113 30550788
2001 Predominant identification of RNA-binding proteins in Fas-induced apoptosis by proteome analysis. The Journal of biological chemistry 109 11352910
2002 Transcriptional activation of human CYP17 in H295R adrenocortical cells depends on complex formation among p54(nrb)/NonO, protein-associated splicing factor, and SF-1, a complex that also participates in repression of transcription. Endocrinology 106 11897684
2009 Role of p54 RNA helicase activity and its C-terminal domain in translational repression, P-body localization and assembly. Molecular biology of the cell 105 19297524
2005 Role for PSF in mediating transcriptional activator-dependent stimulation of pre-mRNA processing in vivo. Molecular and cellular biology 103 16024807
2002 PSF and p54nrb bind a conserved stem in U5 snRNA. RNA (New York, N.Y.) 103 12403470
2004 Identification of the polypyrimidine tract binding protein-associated splicing factor.p54(nrb) complex as a candidate DNA double-strand break rejoining factor. The Journal of biological chemistry 101 15590677
1994 Interleukin-1 activates p54 mitogen-activated protein (MAP) kinase/stress-activated protein kinase by a pathway that is independent of p21ras, Raf-1, and MAP kinase kinase. The Journal of biological chemistry 99 7527398
2004 African swine fever virus structural protein p54 is essential for the recruitment of envelope precursors to assembly sites. Journal of virology 96 15047843
2007 A coactivator trap identifies NONO (p54nrb) as a component of the cAMP-signaling pathway. Proceedings of the National Academy of Sciences of the United States of America 92 18077367
2010 DDX6 (Rck/p54) is required for efficient hepatitis C virus replication but not for internal ribosome entry site-directed translation. Journal of virology 91 20392846
1999 PKC-dependent activation of p46/p54 JNKs during ischemic preconditioning in conscious rabbits. The American journal of physiology 90 10564130
2015 Protein arginine methyltransferase CARM1 attenuates the paraspeckle-mediated nuclear retention of mRNAs containing IRAlus. Genes & development 89 25792598
2007 Transcriptional activity of androgen receptor is modulated by two RNA splicing factors, PSF and p54nrb. Molecular and cellular biology 89 17452459
1996 Functional properties of p54, a novel SR protein active in constitutive and alternative splicing. Molecular and cellular biology 89 8816452
1996 The transcription factor Spi-1/PU.1 binds RNA and interferes with the RNA-binding protein p54nrb. The Journal of biological chemistry 88 8626664
2015 2'-Fluoro-modified phosphorothioate oligonucleotide can cause rapid degradation of P54nrb and PSF. Nucleic acids research 87 25855809
1995 The rck/p54 candidate proto-oncogene product is a 54-kilodalton D-E-A-D box protein differentially expressed in human and mouse tissues. Cancer research 81 7614484
2009 Involvement of p54(nrb), a PSF partner protein, in DNA double-strand break repair and radioresistance. Nucleic acids research 79 19759212
1998 The RNA-splicing factor PSF/p54 controls DNA-topoisomerase I activity by a direct interaction. The Journal of biological chemistry 79 9756848
2012 Multiple binding of repressed mRNAs by the P-body protein Rck/p54. RNA (New York, N.Y.) 78 22836354
1997 The intracisternal A-particle proximal enhancer-binding protein activates transcription and is identical to the RNA- and DNA-binding protein p54nrb/NonO. Molecular and cellular biology 78 9001221
2007 Specific trans-acting proteins interact with auxiliary RNA polyadenylation elements in the COX-2 3'-UTR. RNA (New York, N.Y.) 76 17507659
2007 DjCBC-1, a conserved DEAD box RNA helicase of the RCK/p54/Me31B family, is a component of RNA-protein complexes in planarian stem cells and neurons. Developmental dynamics : an official publication of the American Association of Anatomists 74 17994545
2008 Paraspeckle protein p54nrb links Sox9-mediated transcription with RNA processing during chondrogenesis in mice. The Journal of clinical investigation 73 18677406
2004 The African swine fever virus dynein-binding protein p54 induces infected cell apoptosis. FEBS letters 71 15225638
1993 Proteins homologous to the Xenopus germ cell-specific RNA-binding proteins p54/p56 are temporally expressed in mouse male germ cells. Developmental biology 71 8330676
2015 p54(nrb)/NONO regulates lipid metabolism and breast cancer growth through SREBP-1A. Oncogene 70 26148231
2009 p54nrb is a transcriptional corepressor of the progesterone receptor that modulates transcription of the labor-associated gene, connexin 43 (Gja1). Molecular endocrinology (Baltimore, Md.) 70 19423654
1999 Overexpression of rck/p54, a DEAD box protein, in human colorectal tumours. British journal of cancer 70 10360675
2007 The PSF.p54nrb complex is a novel Mnk substrate that binds the mRNA for tumor necrosis factor alpha. The Journal of biological chemistry 69 17965020
1994 Characterization and molecular basis of heterogeneity of the African swine fever virus envelope protein p54. Journal of virology 69 7933107
2020 NONO and tumorigenesis: More than splicing. Journal of cellular and molecular medicine 66 32168434
2011 p54nrb is a new regulator of progression of malignant melanoma. Carcinogenesis 66 21642354
2002 A single c-Jun N-terminal kinase isoform (JNK3-p54) is an effector in both neuronal differentiation and cell death. The Journal of biological chemistry 66 12401814
2012 Sepantronium bromide (YM155) induces disruption of the ILF3/p54(nrb) complex, which is required for survivin expression. Biochemical and biophysical research communications 65 22842455
2005 Induction and mode of action of the viral stress-inducible murine proteins, P56 and P54. Virology 61 16023166
2002 RNA helicase p54 (DDX6) is a shuttling protein involved in nuclear assembly of stored mRNP particles. Journal of cell science 61 11839790
2002 Adrenocorticotropin/cyclic adenosine 3',5'-monophosphate-mediated transcription of the human CYP17 gene in the adrenal cortex is dependent on phosphatase activity. Endocrinology 61 11956159
2010 Sequences in PSF/SFPQ mediate radioresistance and recruitment of PSF/SFPQ-containing complexes to DNA damage sites in human cells. DNA repair 60 21144806
2003 p54nrb acts as a transcriptional coactivator for activation function 1 of the human androgen receptor. Biochemical and biophysical research communications 60 12810069
1991 Structural basis for the inactivation of the P54 mutant of beta-lactamase from Staphylococcus aureus PC1. Biochemistry 60 1892849
2020 Splicing Regulator p54nrb /Non-POU Domain-Containing Octamer-Binding Protein Enhances Carcinogenesis Through Oncogenic Isoform Switch of MYC Box-Dependent Interacting Protein 1 in Hepatocellular Carcinoma. Hepatology (Baltimore, Md.) 58 31815296
2020 Splicing factor proline- and glutamine-rich (SFPQ) protein regulates platinum response in ovarian cancer-modulating SRSF2 activity. Oncogene 58 32332923
1994 Interleukin 1 alpha activates two forms of p54 alpha mitogen-activated protein kinase in rabbit liver. The Journal of experimental medicine 58 7964479
2003 Deletion of proteasomal subunit S5a/Rpn10/p54 causes lethality, multiple mitotic defects and overexpression of proteasomal genes in Drosophila melanogaster. Journal of cell science 57 12584246
1995 Macromolecular interactions in the nucleoporin p62 complex of rat nuclear pores: binding of nucleoporin p54 to the rod domain of p62. The Journal of cell biology 54 7531196
2006 Visualization of the African swine fever virus infection in living cells by incorporation into the virus particle of green fluorescent protein-p54 membrane protein chimera. Virology 52 16490226
2001 Co-overexpression of DEAD box protein rck/p54 and c-myc protein in human colorectal adenomas and the relevance of their expression in cultured cell lines. Carcinogenesis 51 11751426
1997 High level expression of the major antigenic African swine fever virus proteins p54 and p30 in baculovirus and their potential use as diagnostic reagents. Journal of virological methods 51 9029527
2013 The transcription-splicing protein NonO/p54nrb and three NonO-interacting proteins bind to distal enhancer region and augment rhodopsin expression. Human molecular genetics 49 24301678
2006 Coregulator exchange and sphingosine-sensitive cooperativity of steroidogenic factor-1, general control nonderepressed 5, p54, and p160 coactivators regulate cyclic adenosine 3',5'-monophosphate-dependent cytochrome P450c17 transcription rate. Molecular endocrinology (Baltimore, Md.) 47 17121866
2009 hnRNP M interacts with PSF and p54(nrb) and co-localizes within defined nuclear structures. Experimental cell research 46 19874820
2015 New Noncoding Lytic Transcripts Derived from the Epstein-Barr Virus Latency Origin of Replication, oriP, Are Hyperedited, Bind the Paraspeckle Protein, NONO/p54nrb, and Support Viral Lytic Transcription. Journal of virology 45 25926645
2006 p54nrb is a component of the snRNP-free U1A (SF-A) complex that promotes pre-mRNA cleavage during polyadenylation. RNA (New York, N.Y.) 43 16373496
1997 Mapping the ubiquitin-binding domains in the p54 regulatory complex subunit of the Drosophila 26S protease. FEBS letters 42 9256246
2016 Regulation of PCGEM1 by p54/nrb in prostate cancer. Scientific reports 41 27682980
2005 The multifunctional nuclear protein p54nrb is multiphosphorylated in mitosis and interacts with the mitotic regulator Pin1. Journal of molecular biology 41 15701524
2020 Epitope mapping of African swine fever virus (ASFV) structural protein, p54. Virus research 40 32004574
2009 The hepatitis delta virus RNA genome interacts with eEF1A1, p54(nrb), hnRNP-L, GAPDH and ASF/SF2. Virology 40 19464723
2003 55 kDa nuclear matrix protein (nmt55) mRNA is expressed in human prostate cancer tissue and is associated with the androgen receptor. International journal of cancer 40 12672026
2009 Transcriptional activity of the murine retinol-binding protein gene is regulated by a multiprotein complex containing HMGA1, p54 nrb/NonO, protein-associated splicing factor (PSF) and steroidogenic factor 1 (SF1)/liver receptor homologue 1 (LRH-1). The international journal of biochemistry & cell biology 39 19389484
2008 Brm transactivates the telomerase reverse transcriptase (TERT) gene and modulates the splicing patterns of its transcripts in concert with p54(nrb). The Biochemical journal 39 18042045
2001 Immunodetection of nmt55/p54nrb isoforms in human breast cancer. BMC cancer 39 11710964
1996 The structural protein p54 is essential for African swine fever virus viability. Virus research 39 8725112
2008 Knockdown of RCK/p54 expression by RNAi inhibits proliferation of human colorectal cancer cells in vitro and in vivo. Cancer biology & therapy 38 18769115
2004 Expression and functional significance of mouse paraspeckle protein 1 on spermatogenesis. Biology of reproduction 38 15140795
2000 PSF/p54(nrb) stimulates "jumping" of DNA topoisomerase I between separate DNA helices. Biochemistry 38 10858305
2019 Kinetic and subcellular analysis of PS-ASO/protein interactions with P54nrb and RNase H1. Nucleic acids research 36 31495875
2005 Functional analysis of the promoter of the mitochondrial phosphate carrier human gene: identification of activator and repressor elements and their transcription factors. The Biochemical journal 36 15984930
2007 Carbon monoxide blocks oxidative stress-induced hepatocyte apoptosis via inhibition of the p54 JNK isoform. Free radical biology & medicine 35 18206660
2003 Overexpression of a DEAD box/RNA helicase protein, rck/p54, in human hepatocytes from patients with hepatitis C virus-related chronic hepatitis and its implication in hepatocellular carcinogenesis. Journal of viral hepatitis 35 12823589
1996 Identification of the nuclear localization signal of mouse DNA primase: nuclear transport of p46 subunit is facilitated by interaction with p54 subunit. Journal of cell science 35 8937981
2002 The calcium-binding protein p54/NEFA is a novel luminal resident of medial Golgi cisternae that traffics independently of mannosidase II. European journal of cell biology 34 11893086
1992 Cell culture propagation modifies the African swine fever virus replication phenotype in macrophages and generates viral subpopulations differing in protein p54. Virus research 34 1604931
2013 Glucocorticoid induction of occludin expression and endothelial barrier requires transcription factor p54 NONO. Investigative ophthalmology & visual science 32 23640037
1999 Role of interleukin (IL)-2 receptor beta-chain subdomains and Shc in p38 mitogen-activated protein (MAP) kinase and p54 MAP kinase (stress-activated protein Kinase/c-Jun N-terminal kinase) activation. IL-2-driven proliferation is independent of p38 and p54 MAP kinase activation. The Journal of biological chemistry 32 10066828
1997 Chloroplast endoribonuclease p54 involved in RNA 3'-end processing is regulated by phosphorylation and redox state. Nucleic acids research 32 9171092
2021 Characterization of Anti-p54 Monoclonal Antibodies and Their Potential Use for African Swine Fever Virus Diagnosis. Pathogens (Basel, Switzerland) 31 33562314
2021 Identification of one novel epitope targeting p54 protein of African swine fever virus using monoclonal antibody and development of a capable ELISA. Research in veterinary science 31 34638027