Affinage

HNRNPU

Heterogeneous nuclear ribonucleoprotein U · UniProt Q00839

Length
825 aa
Mass
90.6 kDa
Annotated
2026-04-28
100 papers in source corpus 49 papers cited in narrative 48 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

HNRNPU (SAF-A) is a multifunctional nuclear scaffold protein that organizes large-scale chromatin architecture, regulates RNA processing and gene expression, participates in DNA damage responses, and contributes to chromosome segregation during mitosis. It binds scaffold/matrix attachment region (SAR/MAR) DNA via its SAP domain and chromatin-associated RNAs via its RGG domain, cycling between oligomeric and monomeric states through an intrinsic AAA+ ATPase domain; this ATP-dependent oligomerization with chromatin-associated RNAs decompacts chromatin and maintains topologically associating domain boundaries and compartment organization in concert with CTCF-cohesin (PMID:28622508, PMID:29273625). HNRNPU broadly regulates U2 snRNP maturation and global alternative splicing through direct binding to snRNAs and pre-mRNAs, is required for Polycomb-mediated silencing directed by imprinting lncRNAs and XIST on the inactive X chromosome, and participates in DNA double-strand break repair by undergoing DNA-PK-dependent Ser59 phosphorylation that temporally coordinates base excision repair with NHEJ (PMID:22325991, PMID:25030905, PMID:27303920, PMID:40493679, PMID:40791421). Loss of HNRNPU in the developing cortex causes rapid cell death and dysregulated splicing of neurodevelopmental genes, and tissue-specific knockouts demonstrate essential roles in spermatogenesis, hepatocyte chromatin organization, and skeletal muscle homeostasis (PMID:35864088, PMID:34815802, PMID:31469911, PMID:38797891).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 1992 High

    Identification of SAF-A as a nuclear DNA-binding protein with specificity for SAR/MAR elements established the founding concept that a nuclear matrix-associated factor directly organizes looped chromatin domains.

    Evidence Protein purification with DNA binding assays and electron microscopy of loop structures

    PMID:1324173

    Open questions at the time
    • Binding specificity beyond A/T-rich sequences undefined
    • In vivo relevance of looped structures not demonstrated
    • No RNA-binding activity yet recognized
  2. 1994 High

    Demonstrating that SAF-A and hnRNP-U are identical and that the protein binds both DNA and RNA in vivo unified two research lines and raised the question of how dual nucleic acid binding contributes to nuclear architecture.

    Evidence UV cross-linking in vivo, filter-binding assays, biochemical fractionation, and isoform purification with electron microscopy

    PMID:8068679 PMID:8174554

    Open questions at the time
    • DNA- and RNA-binding domains not yet mapped to distinct regions
    • Functional consequence of dual binding unknown
  3. 1997 High

    Mapping a bipartite SAR-binding domain distinct from the RGG RNA-binding domain, and showing caspase cleavage separates DNA-binding from hnRNP functions during apoptosis, established modular domain architecture and functional separability.

    Evidence Domain mapping, in vitro caspase cleavage, nuclear fractionation, and in vivo cross-linking

    PMID:10671544 PMID:9204873 PMID:9405365

    Open questions at the time
    • ATPase domain not yet recognized
    • Structural basis of SAR recognition unknown
    • How apoptotic cleavage contributes to nuclear disassembly unclear
  4. 2003 High

    SAF-A enrichment on the inactive X chromosome via its RGG domain provided the first evidence that RNA-dependent chromatin association underlies a specific epigenetic process—X chromosome inactivation.

    Evidence Immunofluorescence, nuclear matrix extraction, and RGG domain deletion analysis in female cells

    PMID:14608463

    Open questions at the time
    • Direct XIST RNA interaction not yet biochemically confirmed
    • Whether SAF-A is required for Xi silencing or merely recruited unknown
  5. 2006 High

    Discovery that hnRNPU acts as a pseudosubstrate retaining SCF(β-TrCP) in the nucleus, and that it stabilizes specific mRNAs via 3′-UTR binding, expanded the functional repertoire beyond structural scaffolding to include signaling regulation and post-transcriptional gene control.

    Evidence Affinity purification-MS, competition assays, mRNA stability assays, and RNA immunoprecipitation

    PMID:11850407 PMID:17174306

    Open questions at the time
    • mRNA stabilization targets largely uncharacterized genome-wide
    • β-TrCP pseudosubstrate mechanism not confirmed in vivo
  6. 2009 High

    Identification of Ser59 phosphorylation by DNA-PK specifically in response to DSBs, with prolonged phosphorylation in NHEJ-deficient cells, placed SAF-A in the DNA damage signaling network and linked it to DSB repair pathway choice.

    Evidence Cell-free kinase assay, phospho-specific antibody, DNA-PK inhibitors, and NHEJ-deficient cell lines

    PMID:19351595 PMID:19844162

    Open questions at the time
    • Downstream effector mechanism of pS59 not identified
    • Whether SAF-A directly participates in NHEJ unknown
  7. 2012 High

    Genome-wide CLIP-seq revealed that hnRNPU binds virtually all snRNA classes and regulates U2 snRNP maturation, establishing it as a global regulator of splicing machinery rather than a gene-specific splicing factor; concurrently, reconstituted assays showed direct stimulation of NEIL1 DNA glycosylase, linking hnRNPU to base excision repair.

    Evidence CLIP-seq, RNA-seq splicing analysis, Cajal body imaging, in vitro BER reconstitution with binding affinity determination

    PMID:22325991 PMID:22902625

    Open questions at the time
    • How snRNP maturation defects propagate to specific splicing outcomes unresolved
    • Whether NEIL1 stimulation occurs at endogenous damage sites in vivo unclear
  8. 2014 High

    Biphasic dynamics at DNA damage sites—PAR-dependent recruitment followed by ATM/ATR/DNA-PK-dependent exclusion coupled to R-loop resolution—revealed SAF-A as an active participant in the transcription-coupled damage response rather than a passive scaffold.

    Evidence Laser micro-irradiation, live-cell imaging, kinase and PARP inhibitors, R-loop reporter, domain mutants

    PMID:25030905

    Open questions at the time
    • Mechanism of R-loop removal by SAF-A exclusion unknown
    • Whether PAR binding is direct or via intermediary not confirmed
  9. 2015 High

    PLK1 was identified as the mitotic Ser59 kinase (with PP2A as phosphatase), and S59A mutation caused chromosome misalignment and delayed mitotic exit, establishing that the same regulatory site serves distinct kinases in interphase (DNA-PK) vs. mitosis (PLK1).

    Evidence Co-immunoprecipitation with PLK1, kinase inhibitors, phosphomimetic/phospho-null mutants, live-cell imaging

    PMID:25986610

    Open questions at the time
    • Mitotic substrates or partners of phospho-S59 SAF-A not identified
    • Whether PLK1-SAF-A axis feeds into spindle assembly checkpoint unclear
  10. 2016 High

    Demonstrating that DNA-PK-phosphorylated SAF-A releases chromatin-bound NEIL1 and that dephosphorylated SAF-A relieves Ku inhibition of glycosylases resolved the temporal coordination problem—how cells delay BER at DSBs to prioritize NHEJ.

    Evidence Reconstituted in vitro glycosylase/Ku assay with phosphomimetic mutants, chromatin fractionation after irradiation

    PMID:27303920

    Open questions at the time
    • In vivo validation of temporal repair ordering limited
    • Whether other glycosylases are similarly regulated unknown
  11. 2017 High

    Discovery of the AAA+ ATPase domain that drives ATP-dependent oligomerization cycles with chromatin-associated RNAs to decompact chromatin, combined with Hi-C showing loss of TAD boundaries and compartment structure upon depletion, provided the first unified mechanistic model for SAF-A's genome-organizing function.

    Evidence ATPase and oligomerization assays, Hi-C, FRAP, DamID, ChIP-seq in conditional KO hepatocytes

    PMID:28622508 PMID:29273625

    Open questions at the time
    • Structural basis of ATPase-driven oligomerization unknown
    • Which specific caRNAs are critical at individual loci undefined
    • Whether chromatin decompaction is cause or consequence of transcription unclear
  12. 2020 High

    Tissue-specific conditional knockouts revealed that hnRNPU maintains chromatin accessibility and transcriptional programs in hepatocytes (preventing NASH-like signatures) and cortical neurons, establishing its essential in vivo roles beyond cell lines.

    Evidence Hepatocyte-specific KO with RNA-seq, ChIP-seq, NASH dietary model; cortical conditional truncation with RNA-seq and pharmacological rescue

    PMID:31469911 PMID:35864088

    Open questions at the time
    • Whether chromatin disorganization is the primary driver of tissue pathology or secondary to splicing defects unresolved
    • Human disease mutations not yet functionally characterized in these systems
  13. 2022 High

    hnRNPU was shown to promote class-switch recombination via C-NHEJ condensates, binding S-region G-quadruplexes and regulating R-loops, revealing a new role in adaptive immunity that integrates its RNA/DNA-binding, phase separation, and NHEJ functions; concurrently, its role in DNA replication origin licensing was established.

    Evidence S–S joining assay, G-quadruplex binding, DRIP, phase separation inhibitors, DNA fiber assays, single-cell replication timing

    PMID:34888666 PMID:36943867

    Open questions at the time
    • Whether condensate formation is required or coincidental for CSR not formally tested
    • Mechanism of origin licensing promotion unknown
    • Replication timing boundary maintenance could be indirect via chromatin organization
  14. 2025 High

    Systematic allelic reconstitution demonstrated that the SAP domain (with serines S14/S26), ATPase domain, and RGG repeats each contribute non-redundantly to XIST localization, facultative heterochromatin maintenance, splicing, and cell proliferation; α-satellite RNAs were identified as SAF-A ligands essential for post-mitotic chromatin re-association and nuclear lamina reassembly.

    Evidence Allelic reconstitution of SAP/ATPase/RGG mutants, FRAP, RNA FISH, ChIP, α-satellite RNA depletion with chromosome segregation and lamina assembly assays

    PMID:40219970 PMID:40493679

    Open questions at the time
    • How SAP domain phosphorylation is regulated in vivo unknown
    • Whether α-satellite RNA function generalizes beyond the tested cell types unclear
    • Structural model integrating all three functional domains lacking
  15. 2025 Medium

    hnRNPU was identified as required for long-range Polycomb recruitment by multiple imprinting lncRNAs, connecting its RNA-binding scaffold function to PRC-mediated gene silencing beyond X inactivation.

    Evidence Systematic RIP (27 antibodies), HNRNPU KO, ChIP for PRC marks, RNA FISH (preprint)

    PMID:40791421

    Open questions at the time
    • Preprint awaiting peer review
    • Mechanism by which hnRNPU bridges lncRNA to PRC undefined
    • Whether this is direct or via chromatin architecture changes not resolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include: the structural basis of ATP-dependent oligomerization, how SAF-A distinguishes its chromatin-organizing versus splicing-regulatory RNA targets, the identity and hierarchy of caRNAs at specific genomic loci, and whether human HNRNPU mutations cause a defined neurodevelopmental disorder through chromatin, splicing, or combined mechanisms.
  • No high-resolution structure of full-length SAF-A or its oligomeric state
  • Genome-wide caRNA target hierarchy unresolved
  • Human genetic disease mechanism not formally delineated

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003723 RNA binding 6 GO:0003677 DNA binding 5 GO:0140110 transcription regulator activity 4 GO:0098772 molecular function regulator activity 2 GO:0008092 cytoskeletal protein binding 1 GO:0140657 ATP-dependent activity 1
Localization
GO:0005634 nucleus 6 GO:0005694 chromosome 4 GO:0005654 nucleoplasm 2 GO:0005856 cytoskeleton 1
Pathway
R-HSA-73894 DNA Repair 5 R-HSA-4839726 Chromatin organization 4 R-HSA-74160 Gene expression (Transcription) 4 R-HSA-1640170 Cell Cycle 3 R-HSA-8953854 Metabolism of RNA 3 R-HSA-1266738 Developmental Biology 2 R-HSA-5357801 Programmed Cell Death 2 R-HSA-168256 Immune System 1 R-HSA-69306 DNA Replication 1
Partners
BTRCCTCFDDX5NEIL1PLK1PRKDCSMARCA4WT1
Complex memberships
CTCF-cohesinSWI/SNF (BAF)hnRNP complex

Evidence

Reading pass · 48 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1992 SAF-A (hnRNPU) was identified as a novel nuclear DNA-binding protein with high affinity for scaffold/matrix attachment region (SAR/MAR) DNA elements; the purified protein binds at multiple sites to SAR elements, particularly A/T-rich stretches, forms large protein aggregates, and mediates formation of looped DNA structures. Protein purification, DNA binding assays, electron microscopy, competition studies with synthetic polynucleotides The EMBO journal High 1324173
1994 SAF-A and hnRNP-U are the same protein; it binds chromosomal DNA in vivo (UV cross-linking), binds both double-stranded and single-stranded DNA and RNA in vitro, likely at different binding sites, forming higher-order nucleic acid-protein complexes. UV cross-linking in vivo, filter-binding assays, biochemical fractionation European journal of biochemistry High 8174554
1994 Two isoforms of hnRNP-U (form 1 and form 2) were purified; they differ in primary structure but both bind single- and double-stranded DNA and RNA and specifically bind/aggregate the human SAR element MII; the isoforms form morphologically distinct nucleic acid-protein complexes (unbranched filaments vs. spheres ~35 nm diameter). Chromatographic purification, filter binding, electron microscopy Biochemistry High 8068679
1997 SAF-A contains a novel bipartite SAR-specific DNA-binding domain independent of the RGG RNA-binding domain; during apoptosis, caspase-dependent cleavage within this DNA-binding domain abolishes DNA binding and detaches SAF-A from nuclear structural sites, while hnRNP complex association (RNA metabolism function) is preserved. Domain mapping, in vitro cleavage assays, nuclear fractionation, immunofluorescence The EMBO journal High 9405365
1997 hnRNP-U/SAF-A is directly bound to chromosomal DNA in vivo, demonstrated by formaldehyde cross-linking of living cells followed by CsCl density gradient purification; dimethylsulfate cross-linking and limited protease digestion confirmed direct DNA binding (not via other proteins). Chemical cross-linking (formaldehyde, dimethylsulfate), CsCl density gradient centrifugation, western blotting, protease digestion Biochemistry High 9204873
2000 Caspase-3 cleaves SAF-A at a noncanonical site (SALD after Asp-100); point mutation D100A abrogates cleavage by recombinant caspase-3 in vitro and during apoptosis in vivo. In vitro cleavage with recombinant caspase-3, MALDI-TOF mass spectrometry, Edman sequencing, site-directed mutagenesis The Journal of biological chemistry High 10671544
2002 hnRNP-U is the major nuclear binding partner of the SCF(β-TrCP) ubiquitin ligase subunit β-TrCP/E3RS; it acts as a pseudosubstrate that occupies E3RS stoichiometrically, stabilizes the E3 component, and is responsible for its nuclear localization; hnRNP-U binding is abolished by phospho-IκBα peptide competition or a point mutation in the E3RS WD region, and hnRNP-U is stable (not degraded) unlike canonical substrates. Affinity purification, mass spectrometry, co-immunoprecipitation, competition assays, point mutagenesis Genes & development High 11850407
2002 hnRNP-U/SAF-A binds to an episomally replicating vector (pEPI-1) containing an S/MAR element in vivo, demonstrated by cis-diamminedichloroplatinum II cross-linking followed by nuclear matrix co-purification and immunoprecipitation of the crosslinked DNA-protein complex. Cis-DDP cross-linking, nuclear matrix co-purification, southwestern analysis, immunoprecipitation EMBO reports High 11897664
2003 SAF-A is enriched on the inactive X chromosome (Xi) through its RGG RNA-binding domain; after near-complete removal of DNA and chromatin proteins, SAF-A remains with the nuclear matrix at the Xi position, suggesting RNA-dependent (possibly XIST RNA) association contributing to Xi nuclear architecture. Immunofluorescence, nuclear matrix extraction, domain deletion analysis (RGG box mutants) Chromosoma High 14608463
2006 hnRNP-U directly interacts with the Wilms' tumor suppressor WT1; the interaction involves WT1 zinc-fingers and the middle domain of hnRNP-U, does not require other proteins or nucleic acids, and hnRNP-U modulates WT1 transcriptional activation of a bona fide WT1 target gene. Co-immunoprecipitation of endogenous proteins, domain mapping pulldowns, transcriptional reporter assay Oncogene High 16924231
2006 hnRNP-U enhances expression of specific genes including TNF-α, GADD45A, HEXIM1, HOXA2, IER3, NHLH2, and ZFY by binding to and stabilizing their mRNAs (particularly via 3' UTR binding), increasing mRNA half-life. mRNA stability assays, RNA immunoprecipitation, overexpression/knockdown experiments FEBS letters Medium 17174306
2008 SAF-A binds the 3'-flanking region of the Bmal1 promoter with circadian timing (in vivo footprinting), correlating with Bmal1 expression; this binding is required for circadian transcriptional regulation of Bmal1. In vivo footprinting, chromatin immunoprecipitation, reporter gene assay Molecular and cellular biology Medium 18332112
2009 hnRNP-U/SAF-A is phosphorylated at Ser59 specifically by DNA-PK in vitro and in cells in response to DNA double-strand breaks; this was identified using a cell-free system and confirmed in cells with DSB-inducing agents. Cell-free kinase assay, mass spectrometry mapping, western blotting with phospho-specific antibody, DNA damage induction Biochemical and biophysical research communications High 19351595
2009 SAF-A Ser59 phosphorylation by DNA-PK in response to DSB inducers is exclusively dependent on DNA-PK and inversely correlates with the cell's capacity to repair DSBs by nonhomologous end joining (NHEJ); cells deficient in NHEJ show prolonged phosphorylation. Phospho-specific antibody, DNA-PK inhibitor, NHEJ-deficient cell lines, kinase mapping Cell cycle High 19844162
2010 B23/NPM1, upon transcription repression-induced nucleolar segregation, translocates to the cytoplasm and forms a sequential complex with hnRNPU and hnRNPA1; this complex is regulated by RNA Polymerase I inhibition (not Pol II) and by hnRNPU-bound mRNAs (3'-UTR of Bcl-xL); disruption of the complex by ectopic Bcl-xL 3'-UTR enhances apoptosis, suggesting the complex promotes cell survival. Co-immunoprecipitation, subcellular fractionation, RNA immunoprecipitation, ectopic RNA overexpression, apoptosis assays Oncogene Medium 20101230
2010 SAF-A/Saf-A is recruited to the inactive X chromosome (Xi) by Xist RNA at the developmental transition to Xi maintenance, concomitantly with Ash2l and macroH2A; a mutant Xist that does not repress genes still recruits SAF-A, indicating recruitment is separable from gene silencing. Immunofluorescence, RNA FISH, genetic analysis with mutant Xist in ES cell differentiation Development High 20150277
2011 SAF-A regulates kinetochore-microtubule attachment and spindle organization during mitosis; it localizes to spindles, spindle midzone, and cytoplasmic bridge; co-immunoprecipitates with nucleolin, Aurora-A, and TPX2; SAF-A can bind microtubules directly and contributes to targeting Aurora-A to mitotic spindle MTs; depletion causes mitotic delay, chromosome misalignment, and spindle assembly defects. RNAi depletion, immunofluorescence, co-immunoprecipitation, microtubule binding assay, live-cell imaging Journal of cell science High 21242313
2011 Endogenous SAF-A interacts with the C-terminal domain (CTD) of RNA Polymerase II independently of CTD phosphorylation and mRNA; SAF-A also exists in complexes with Sox2, Oct4, and STAT3 in ES cells; SAF-A binds the Oct4 proximal promoter and its depletion reduces Oct4 expression. Co-immunoprecipitation of endogenous proteins, ChIP, RNA interference, reporter assays Cellular reprogramming Medium 21235343
2011 SAF-A interacts with BRG1 (ATPase motor of the SWI/SNF chromatin remodeling complex) in mouse ES cells; interaction is RNA-independent and persists upon differentiation; dual depletion of SAF-A and BRG1 abolishes global RNA Pol II transcription while leaving RNA Pol I transcription unaffected. Co-immunoprecipitation, co-localization, proximity ligation assay, RNA interference, transcription assay PloS one Medium 22162999
2012 hnRNP U regulates U2 snRNP maturation and Cajal body morphology; genome-wide CLIP-seq shows hnRNP U binds virtually all classes of regulatory noncoding RNAs including all snRNAs required for splicing; hnRNP U loss globally alters alternative splicing through modulation of core splicing machinery. RNAi screen, CLIP-seq, RNA-seq, nuclear fractionation, Cajal body imaging Molecular cell High 22325991
2012 hnRNPUL proteins 1 and 2 (hnRNPU-like, not hnRNPU itself) bind the DSB sensor complex MRN (MRE11-RAD50-NBS1) and are recruited to DNA damage in an MRN-dependent, interdependent manner to stimulate DNA-end resection and promote BLM helicase recruitment; this is noted as a related but distinct protein family. Co-immunoprecipitation, laser micro-irradiation, RNAi depletion, DNA repair assays Molecular cell Medium 22365830
2012 hnRNP-U directly interacts with NEIL1 DNA glycosylase via NEIL1's C-terminal domain; hnRNP-U stimulates NEIL1 base excision activity for 5-hydroxyuracil primarily by enhancing product release; the interacting regions in hnRNP-U map to both N and C termini (Kd ~54 nM); depletion of hnRNP-U epistatically sensitizes cells to low-level oxidative damage together with NEIL1 depletion. Co-immunoprecipitation, in vitro BER assay, domain mapping pulldown, circular dichroism, PONDR prediction, genetic epistasis (siRNA) The Journal of biological chemistry High 22902625
2014 SAF-A exhibits biphasic dynamics at DNA damage sites: rapid recruitment dependent on poly(ADP-ribose) binding, followed by prolonged exclusion dependent on ATM, ATR, and DNA-PK activity; the RNA-binding domain recapitulates this behavior; SAF-A exclusion is coupled to R-loop removal at transcribed damage sites, forming part of an anti-R-loop mechanism. Laser micro-irradiation, live-cell imaging, PARP inhibitors, ATM/ATR/DNA-PK inhibitors, R-loop reporter, domain deletion mutants Nucleic acids research High 25030905
2015 SAF-A Ser59 is phosphorylated in mitosis by PLK1 (not DNA-PKcs); SAF-A interacts with PLK1 in nocodazole-treated cells; Ser59 is dephosphorylated by PP2A in mitosis; cells expressing S59A SAF-A show misaligned chromosomes, lagging chromosomes, polylobed nuclei, and delayed mitotic exit. Phospho-specific antibody, kinase inhibitors, PLK1 knockdown, co-immunoprecipitation, site-directed mutagenesis (S59A), live-cell imaging, PP2A inhibitor Molecular and cellular biology High 25986610
2016 SAF-A, phosphorylated at Ser59 by DNA-PK early after ionizing radiation, promotes transient release of chromatin-bound NEIL1, preventing premature BER at DSB sites; dephosphorylated (but not phosphomimetic S59D) SAF-A relieves Ku inhibition of DNA glycosylases in vitro, coordinating temporal repair order. In vitro DNA glycosylase assay, Ku inhibition assay, phosphomimetic mutants, cell irradiation, chromatin fractionation Oncotarget High 27303920
2017 SAF-A/hnRNPU interacts with chromatin-associated RNAs (caRNAs) via its RGG domain to regulate interphase chromatin structure; SAF-A's AAA+ ATPase domain mediates cycles of protein oligomerization with caRNAs in response to ATP binding and hydrolysis; SAF-A oligomerization decompacts large-scale chromatin structure, while monomerization or loss causes aberrant chromosome folding and genome damage accumulation. RGG domain mutants, ATPase mutants, Hi-C, FRAP, ATPase assay, DNA damage assays, oligomerization assays Cell High 28622508
2017 HNRNPU depletion in mouse hepatocytes increases lamina-associated domain (LAD) coverage, causes global chromatin condensation, compartment switching on 7.5% of genome, decreased TAD boundary strengths at A/B compartment borders, and reduced chromatin loop intensities; HNRNPU mainly associates with active chromatin and 80% of peaks coincide with CTCF or RAD21 binding. In situ Hi-C, DamID, ChIP-seq, RNA-seq, conditional knockout in hepatocytes Genome research High 29273625
2018 HPSE eRNA binds hnRNPU to facilitate its interaction with p300 acetyltransferase and their co-enrichment on a super enhancer, driving chromatin looping between super enhancer and HPSE promoter and p300-mediated transactivation of EGR1. RNA immunoprecipitation, co-immunoprecipitation, ChIP-seq, chromatin conformation capture, gain/loss-of-function Oncogene Medium 29511351
2020 hnRNPU retains miR-30c-5p and other microRNAs in the nucleus, preventing their export into large extracellular vesicles; nuclear binding of miR-30c-5p to hnRNPU stabilizes miRNAs and reduces cytoplasmic availability; binding confirmed by RNA-immunoprecipitation, EMSA, and miR-pulldown; a sequence motif AAMRUGCU was identified as a potential sorting signal. RNA immunoprecipitation, EMSA, miRNA pulldown, gain/loss-of-function, Taqman array, nanoparticle tracking analysis Journal of extracellular vesicles Medium 32944175
2020 HNF4A-AS1 lncRNA binds hnRNPU and facilitates its interaction with CTCF, resulting in CTCF transactivation and transcriptional alteration of HNF4A and glycolysis genes; a small peptide blocking HNF4A-AS1-hnRNPU interaction suppresses this axis. RNA immunoprecipitation, co-immunoprecipitation, luciferase reporter, peptide competition, lentiviral shRNA Journal of hematology & oncology Medium 32216806
2020 Cytoplasmic hnRNPU directly binds the 3'-UTR of IL-6 mRNA to regulate mRNA stability; this cytoplasmic interaction was identified by optimized HITS-CLIP and validated, revealing a nucleocytoplasmic shuttling function of hnRNPU in mRNA stability control. HITS-CLIP (optimized BrdU-CLIP), subcellular fractionation, RNA immunoprecipitation PloS one Medium 32302342
2020 Hepatocyte-specific inactivation of hnRNPU disrupts liver chromatin accessibility, activates molecular signature of NASH, and sensitizes mice to diet-induced NASH; mechanistically, hnRNPU deficiency stimulates expression of a truncated TrkB isoform (TRKB-T1) promoting inflammatory signaling and stress-induced cell death. Hepatocyte-specific conditional KO, RNA-seq, ChIP-seq, BDNF treatment rescue, NASH dietary model Hepatology High 31469911
2021 hnRNPU interacts with WT1 and SOX9 in Sertoli cells and enhances transcription of Sox8 and Sox9 by directly binding their promoter regions (ChIP-qPCR); conditional KO of hnRNPU in Sertoli cells causes testicular atrophy, depletion of Sertoli and germ cells, and male sterility. Conditional KO (Cre/loxP), ChIP-qPCR, co-immunoprecipitation, luciferase assay, RNA-seq Theranostics High 34815802
2021 SAF-A mutants (but not RNAi depletion) cause dramatic chromatin condensation through dominant negative effects on C0T-1 RNA (repeat-rich heterogeneous nuclear RNA) association with euchromatin; results support a model where SAF-A, chromatin-associated RNAs, and other RNA-binding proteins form a complex dynamic meshwork integral to large-scale chromatin architecture. SAF-A domain mutant expression, RNA FISH (C0T-1 RNA), RNAi, chromatin condensation imaging (DAPI) Mammalian genome Medium 34859278
2022 SAF-A promotes origin licensing in G1 phase, maintains replication fork progression, and maintains boundaries between early- and late-replicating domains; SAF-A depletion reduces origin activation frequency, increases γ-H2AX, and causes cells to enter quiescence. SAF-A depletion (siRNA), DNA fiber assay, single-cell replication timing analysis, γ-H2AX immunofluorescence, flow cytometry Journal of cell science High 34888666
2022 CDC20-mediated ubiquitination of hnRNPU promotes its interaction with the CTCF-cohesin complex, modulating chromatin condensation; CDC20 binds hnRNPU through amino acid residues 461-653; loss of this axis contributes to tumor progression and drug resistance. Affinity purification-mass spectrometry, co-immunoprecipitation, domain mapping, ubiquitination assay, DAPI/H2B-mCherry chromatin condensation imaging Cancers Medium 35954396
2022 HNRNPU loss of function in the developing cortex leads to rapid cell death of both postmitotic neurons and neural progenitors (higher sensitivity in progenitors); alternative splicing and expression of genes involved in cell survival, motility, and synapse formation are dysregulated; pharmacological and genetic agents can partially reverse cortical structural defects. Conditional truncation mouse model, immunofluorescence, RNA-seq, alternative splicing analysis, pharmacological rescue Nature communications High 35864088
2022 HNRNPU promotes C-NHEJ-mediated switch region joining through the 53BP1-shieldin DNA repair complex, binds S-region RNA/DNA G-quadruplexes to regulate R-loop and ssDNA accumulation; HNRNPU is an intrinsically disordered protein that interacts with both C-NHEJ and R-loop complexes in an RNA-dependent manner; recruitment of HNRNPU and C-NHEJ factors is sensitive to liquid-liquid phase separation inhibitors suggesting condensate formation. Co-immunoprecipitation, S-S joining assay, G-quadruplex binding assay, R-loop detection (DRIP), phase separation inhibitors, HNRNPU KO Cell reports High 36943867
2022 HNRNPU promotes triple-negative breast cancer progression via association with DDX5; the HNRNPU-DDX5 complex prevents intron retention of MCM10 pre-mRNA (reducing nonsense-mediated decay and activating Wnt/β-catenin) and promotes LMO4 transcription at transcription start sites (activating PI3K-Akt-mTOR signaling). Co-immunoprecipitation, RNA-seq, splicing analysis, ChIP, KD/OE experiments Cell death & disease Medium 36347834
2022 Muscle-specific Hnrnpu KO mice develop adult-onset myopathy with selective atrophy of glycolytic muscle, constitutive Akt activation, and altered expression and splicing of metabolic and signal transduction genes; loss of hnRNPU also causes NF-κB activation and Ly6C+ monocyte infiltration preceding muscle atrophy. Muscle-specific conditional KO, RNA-seq, splicing analysis, histology, signaling assays, immune cell flow cytometry iScience High 32659719 38797891
2023 SAF-A/hnRNPU binds polyphosphoinositides (PPIn) directly via a lysine-rich polybasic motif (aa 9-24) in the N-terminal SAP domain; deletion of this polybasic region abolishes PPIn interaction while retaining protein structure. Quantitative interactomics, PPIn-protein binding assay, deletion mutagenesis microPublication biology Medium 37038481
2022 lncRHL binds directly to hnRNPU, enhancing its protein stability; hnRNPU then transcriptionally activates Bmal1, leading to inhibition of VLDL secretion in hepatocytes; lncRHL deficiency accelerates hnRNPU protein degradation and suppresses Bmal1 transcription. RNA immunoprecipitation, co-immunoprecipitation, protein stability assay, ChIP, gene manipulation in mice and hepatocytes Diabetes Medium 35771993
2025 The SAF-A SAP domain (and its serines S14 and S26) is required for XIST RNA localization on the inactive X chromosome, XIST-dependent histone modifications on Xi, normal protein dynamics (FRAP), and cell proliferation; the SAP domain provides a distinct Xi localization signal; SAF-A is a highly dynamic protein interacting with nascent transcripts rather than being part of a static nuclear scaffold. Allelic reconstitution, FRAP, immunofluorescence, RNA FISH, phosphomimetic/phospho-null SAP domain mutants, cell proliferation assays PLoS genetics High 40493679
2025 The SAF-A ATPase domain and RGG repeats control SAF-A nuclear dynamics; RGG repeats mediate interaction with nascent transcripts; both domains are required for maintaining XIST RNA and facultative heterochromatin marks on Xi, for proper mRNA splicing, and for cell proliferation; ATP binding and hydrolysis have distinct effects on Xi maintenance. Allelic reconstitution, FRAP, transcriptome analysis, RNA FISH, chromatin immunoprecipitation, cell proliferation assays, ATP binding/hydrolysis mutants bioRxivpreprint High 41473319
2025 α-Satellite RNAs bind SAF-A and are required for recruitment of SAF-A back to chromatin upon mitotic exit; depletion of either α-satellite RNA or SAF-A causes chromosome missegregation; SAF-A is excluded from chromatin genome-wide during mitosis and both α-satellite RNAs and SAF-A are essential for nuclear lamina reassembly. RNA depletion (siRNA), immunofluorescence, RNA FISH, chromatin fractionation, RNA immunoprecipitation, chromosome segregation assay Nucleic acids research High 40219970
2025 HNRNPU associates with the mammalian SWI/SNF chromatin-remodeling complex (including SMARCA4 and SMARCC2), has a previously unrecognized role in translation, associates with mRNAs encoding neurodevelopmental proteins, and HNRNPU silencing reprograms methylation at active and bivalent promoters of neurodevelopmental transcription factors. Protein-protein interaction mapping, RNA target identification (RIP/CLIP), genome-wide DNA methylation profiling (EPIC arrays), protein co-immunoprecipitation in neural stem cells Nucleic acids research Medium 41674383
2025 HNRNPU is required for long-range Polycomb recruitment by lncRNAs Airn, Kcnq1ot1, and Xist; HNRNPU exhibited enriched and correlated association with all three lncRNAs, and was required for PRC-directed histone modifications induced by all three, without being necessary for proper localization of Airn or Kcnq1ot1. RNA immunoprecipitation from formaldehyde-crosslinked cells (27 antibodies), HNRNPU KO, ChIP for PRC modifications, RNA FISH bioRxivpreprint Medium 40791421
2024 hnRNPU is essential for establishing the spermatogonial stem cell (SSC) pool; conditional loss in prospermatogonia arrests differentiation and migration to the basement membrane; hnRNPU binds Vrk1, Slx4, and Dazl transcripts that undergo aberrant alternative splicing upon hnRNPU depletion. Conditional KO (Cre/loxP), single-cell RNA-seq, RNA immunoprecipitation, splicing analysis Cell reports High 38625792

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1992 Characterization of SAF-A, a novel nuclear DNA binding protein from HeLa cells with high affinity for nuclear matrix/scaffold attachment DNA elements. The EMBO journal 230 1324173
2017 SAF-A Regulates Interphase Chromosome Structure through Oligomerization with Chromatin-Associated RNAs. Cell 168 28622508
1994 Nucleic-acid-binding properties of hnRNP-U/SAF-A, a nuclear-matrix protein which binds DNA and RNA in vivo and in vitro. European journal of biochemistry 159 8174554
2012 Regulation of DNA-end resection by hnRNPU-like proteins promotes DNA double-strand break signaling and repair. Molecular cell 154 22365830
2014 DNA damage triggers SAF-A and RNA biogenesis factors exclusion from chromatin coupled to R-loops removal. Nucleic acids research 148 25030905
2012 Nuclear matrix factor hnRNP U/SAF-A exerts a global control of alternative splicing by regulating U2 snRNP maturation. Molecular cell 144 22325991
1997 The novel SAR-binding domain of scaffold attachment factor A (SAF-A) is a target in apoptotic nuclear breakdown. The EMBO journal 125 9405365
2003 Scaffold attachment factor A (SAF-A) is concentrated in inactive X chromosome territories through its RGG domain. Chromosoma 114 14608463
2017 The nuclear matrix protein HNRNPU maintains 3D genome architecture globally in mouse hepatocytes. Genome research 101 29273625
2002 Pseudosubstrate regulation of the SCF(beta-TrCP) ubiquitin ligase by hnRNP-U. Genes & development 101 11850407
2010 The Trithorax group protein Ash2l and Saf-A are recruited to the inactive X chromosome at the onset of stable X inactivation. Development (Cambridge, England) 99 20150277
2022 Targeting HNRNPU to overcome cisplatin resistance in bladder cancer. Molecular cancer 98 35130920
2018 HPSE enhancer RNA promotes cancer progression through driving chromatin looping and regulating hnRNPU/p300/EGR1/HPSE axis. Oncogene 93 29511351
2002 An episomally replicating vector binds to the nuclear matrix protein SAF-A in vivo. EMBO reports 90 11897664
1997 The scaffold/matrix attachment region binding protein hnRNP-U (SAF-A) is directly bound to chromosomal DNA in vivo: a chemical cross-linking study. Biochemistry 82 9204873
2020 The RNA-binding protein hnRNPU regulates the sorting of microRNA-30c-5p into large extracellular vesicles. Journal of extracellular vesicles 77 32944175
2006 hnRNP-U enhances the expression of specific genes by stabilizing mRNA. FEBS letters 73 17174306
2010 Four patients with speech delay, seizures and variable corpus callosum thickness sharing a 0.440 Mb deletion in region 1q44 containing the HNRPU gene. European journal of medical genetics 72 20382278
2017 Heterozygous HNRNPU variants cause early onset epilepsy and severe intellectual disability. Human genetics 68 28393272
2017 Genetic and phenotypic dissection of 1q43q44 microdeletion syndrome and neurodevelopmental phenotypes associated with mutations in ZBTB18 and HNRNPU. Human genetics 67 28283832
1994 Purification of two isoforms of hnRNP-U and characterization of their nucleic acid binding activity. Biochemistry 58 8068679
2020 HNF4A-AS1/hnRNPU/CTCF axis as a therapeutic target for aerobic glycolysis and neuroblastoma progression. Journal of hematology & oncology 54 32216806
2010 B23 acts as a nucleolar stress sensor and promotes cell survival through its dynamic interaction with hnRNPU and hnRNPA1. Oncogene 50 20101230
2012 Enhancement of NEIL1 protein-initiated oxidized DNA base excision repair by heterogeneous nuclear ribonucleoprotein U (hnRNP-U) via direct interaction. The Journal of biological chemistry 49 22902625
2008 Identifying functional neighborhoods within the cell nucleus: proximity analysis of early S-phase replicating chromatin domains to sites of transcription, RNA polymerase II, HP1gamma, matrin 3 and SAF-A. Journal of cellular biochemistry 47 18618731
2000 Apoptotic cleavage of scaffold attachment factor A (SAF-A) by caspase-3 occurs at a noncanonical cleavage site. The Journal of biological chemistry 45 10671544
2020 hnRNPU/TrkB Defines a Chromatin Accessibility Checkpoint for Liver Injury and Nonalcoholic Steatohepatitis Pathogenesis. Hepatology (Baltimore, Md.) 41 31469911
2017 De novo mutations in HNRNPU result in a neurodevelopmental syndrome. American journal of medical genetics. Part A 40 28944577
2017 Clinical and molecular characterization of de novo loss of function variants in HNRNPU. American journal of medical genetics. Part A 39 28815871
2023 piRNA-1742 promotes renal cell carcinoma malignancy by regulating USP8 stability through binding to hnRNPU and thereby inhibiting MUC12 ubiquitination. Experimental & molecular medicine 35 37332045
2021 hnRNPU in Sertoli cells cooperates with WT1 and is essential for testicular development by modulating transcriptional factors Sox8/9. Theranostics 35 34815802
2018 The role of nuclear matrix protein HNRNPU in maintaining the architecture of 3D genome. Seminars in cell & developmental biology 35 29981443
2022 Heterogeneous nuclear ribonucleoprotein U (HNRNPU) safeguards the developing mouse cortex. Nature communications 34 35864088
2009 hnRNP-U is a specific DNA-dependent protein kinase substrate phosphorylated in response to DNA double-strand breaks. Biochemical and biophysical research communications 34 19351595
2006 hnRNP-U directly interacts with WT1 and modulates WT1 transcriptional activation. Oncogene 32 16924231
2001 Specificity of SAF-A and lamin B binding in vitro correlates with the satellite DNA bending state. Journal of cellular biochemistry 32 11573239
2009 Cell nonhomologous end joining capacity controls SAF-A phosphorylation by DNA-PK in response to DNA double-strand breaks inducers. Cell cycle (Georgetown, Tex.) 31 19844162
2021 The role of SAF-A/hnRNP U in regulating chromatin structure. Current opinion in genetics & development 30 34823151
2023 circ-hnRNPU inhibits NONO-mediated c-Myc transactivation and mRNA stabilization essential for glycosylation and cancer progression. Journal of experimental & clinical cancer research : CR 28 37993881
2021 MicroRNA-132-3p alleviates neuron apoptosis and impairments of learning and memory abilities in Alzheimer's disease by downregulation of HNRNPU stabilized BACE1. Cell cycle (Georgetown, Tex.) 27 34585626
2008 Rhythmic SAF-A binding underlies circadian transcription of the Bmal1 gene. Molecular and cellular biology 27 18332112
2011 The nuclear scaffold protein SAF-A is required for kinetochore-microtubule attachment and contributes to the targeting of Aurora-A to mitotic spindles. Journal of cell science 25 21242313
2001 Effects of the heterogeneous nuclear ribonucleoprotein U (hnRNP U/SAF-A) on glucocorticoid-dependent transcription in vivo. The Journal of steroid biochemistry and molecular biology 25 11530285
2021 HNRNPU promotes the progression of hepatocellular carcinoma by enhancing CDK2 transcription. Experimental cell research 24 34737140
2011 SAF-A has a role in transcriptional regulation of Oct4 in ES cells through promoter binding. Cellular reprogramming 24 21235343
2022 HNRNPU promotes the progression of triple-negative breast cancer via RNA transcription and alternative splicing mechanisms. Cell death & disease 23 36347834
2020 The splicing regulatory factor hnRNPU is a novel transcriptional target of c-Myc in hepatocellular carcinoma. FEBS letters 22 33040326
2014 A genomic copy number variant analysis implicates the MBD5 and HNRNPU genes in Chinese children with infantile spasms and expands the clinical spectrum of 2q23.1 deletion. BMC medical genetics 22 24885232
2024 Microglial circ-UBE2K exacerbates depression by regulating parental gene UBE2K via targeting HNRNPU. Theranostics 21 38994030
2022 Nuclear organization by satellite DNA, SAF-A/hnRNPU and matrix attachment regions. Seminars in cell & developmental biology 21 35484025
2022 Long Noncoding RNA lncRHL Regulates Hepatic VLDL Secretion by Modulating hnRNPU/BMAL1/MTTP Axis. Diabetes 21 35771993
2022 LncRNA IL21-AS1 interacts with hnRNPU protein to promote IL21 overexpression and aberrant differentiation of Tfh cells in systemic lupus erythematosus. Clinical and translational medicine 20 36447054
2023 HNRNPU facilitates antibody class-switch recombination through C-NHEJ promotion and R-loop suppression. Cell reports 19 36943867
2017 Expression Profiling Identifies the Noncoding Processed Transcript of HNRNPU with Proliferative Properties in Pancreatic Ductal Adenocarcinoma. Non-coding RNA 19 29657295
2015 Phosphorylation of SAF-A/hnRNP-U Serine 59 by Polo-Like Kinase 1 Is Required for Mitosis. Molecular and cellular biology 19 25986610
2024 hnRNPU is required for spermatogonial stem cell pool establishment in mice. Cell reports 18 38625792
2018 An episode of acute encephalopathy with biphasic seizures and late reduced diffusion followed by hemiplegia and intractable epilepsy observed in a patient with a novel frameshift mutation in HNRNPU. Brain & development 17 29858110
2011 SAF-A forms a complex with BRG1 and both components are required for RNA polymerase II mediated transcription. PloS one 17 22162999
2023 DNA methylation episignature and comparative epigenomic profiling of HNRNPU-related neurodevelopmental disorder. Genetics in medicine : official journal of the American College of Medical Genetics 16 37120726
2023 Neurodevelopmental deficits and cell-type-specific transcriptomic perturbations in a mouse model of HNRNPU haploinsufficiency. PLoS genetics 16 37782669
2020 Analysis of the nucleocytoplasmic shuttling RNA-binding protein HNRNPU using optimized HITS-CLIP method. PloS one 16 32302342
2000 Specific interaction of mouse major satellite with MAR-binding protein SAF-A. European journal of cell biology 16 11139148
1994 hnRNP-U/SAF-A is encoded by two differentially polyadenylated mRNAs in human cells. Biochimica et biophysica acta 16 7509195
2021 HNRNPU-AS1 Regulates Cell Proliferation and Apoptosis via the MicroRNA 205-5p/AXIN2 Axis and Wnt/β-Catenin Signaling Pathway in Cervical Cancer. Molecular and cellular biology 15 34309414
2019 The Superoanterior Fasciculus (SAF): A Novel White Matter Pathway in the Human Brain? Frontiers in neuroanatomy 15 30890921
2024 MSC-derived exosomal circMYO9B accelerates diabetic wound healing by promoting angiogenesis through the hnRNPU/CBL/KDM1A/VEGFA axis. Communications biology 14 39725699
2023 Germline pathogenic variants in HNRNPU are associated with alterations in blood methylome. European journal of human genetics : EJHG 13 37407733
2023 RNA-binding protein hnRNPU regulates multiple myeloma resistance to selinexor. Cancer letters 13 37984724
2021 SAF-A mutants disrupt chromatin structure through dominant negative effects on RNAs associated with chromatin. Mammalian genome : official journal of the International Mammalian Genome Society 13 34859278
2016 Scaffold attachment factor A (SAF-A) and Ku temporally regulate repair of radiation-induced clustered genome lesions. Oncotarget 13 27303920
1991 Analysis of the promoter region of saf, a Streptomyces griseus gene that increases production of extracellular enzymes. Gene 13 1761232
2024 LncRNA evf-2 Exacerbates Podocyte Injury in Diabetic Nephropathy by Inducing Cell Cycle Re-entry and Inflammation Through Distinct Mechanisms Triggered by hnRNPU. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 12 39470303
2022 CDC20-Mediated hnRNPU Ubiquitination Regulates Chromatin Condensation and Anti-Cancer Drug Response. Cancers 11 35954396
2022 Characterization and functional analysis of chicken dsRNA binding protein hnRNPU. Developmental and comparative immunology 9 36044969
2020 Adult-Onset Myopathy with Constitutive Activation of Akt following the Loss of hnRNP-U. iScience 9 32659719
2022 SAF-A promotes origin licensing and replication fork progression to ensure robust DNA replication. Journal of cell science 8 34888666
2009 SAF-A/hnRNP-U localization in interphase and metaphase. Cytogenetic and genome research 8 19556781
2022 HnRNPU-AS1 inhibits the proliferation, migration and invasion of HCC cells and induces autophagy through miR-556-3p/ miR-580-3p/SOCS6 axis. Cancer biomarkers : section A of Disease markers 7 35275521
2022 Evidence of shared transcriptomic dysregulation of HNRNPU-related disorder between human organoids and embryonic mice. iScience 7 36594023
2021 De novo frameshift variants of HNRNPU in patients with early infantile epileptic encephalopathy: Two case reports and literature review. International journal of developmental neuroscience : the official journal of the International Society for Developmental Neuroscience 7 33914968
2015 Genomic regions targeted by DNA topoisomerase IIβ frequently interact with a nuclear scaffold/matrix protein hnRNP U/SAF-A/SP120. Journal of cellular biochemistry 7 25418483
2025 hnRNPU-mediated pathogenic alternative splicing drives gastric cancer progression. Journal of experimental & clinical cancer research : CR 6 39773744
2023 HNRNPU's multi-tasking is essential for proper cortical development. BioEssays : news and reviews in molecular, cellular and developmental biology 5 37439444
2021 Pharmacological Treatment of Severe Breathing Abnormalities in a Case of HNRNPU Epileptic Encephalopathy. Molecular syndromology 5 34012379
1999 Identification of chURP, a nuclear calmodulin-binding protein related to hnRNP-U. European journal of biochemistry 5 10103044
2024 Circular RNA CircSLC22A23 Promotes Gastric Cancer Progression by Activating HNRNPU Expression. Digestive diseases and sciences 4 38400886
2025 Role of the SAF-A/HNRNPU SAP domain in X chromosome inactivation, nuclear dynamics, transcription, splicing, and cell proliferation. PLoS genetics 3 40493679
2024 Loss of HNRNPU in Skeletal Muscle Increases Intramuscular Infiltration of Ly6C Positive Cells, leading to Muscle Atrophy through Activation of NF-κB Signaling. Advanced biology 3 38797891
2024 The nuclear matrix protein HNRNPU restricts hepatitis B virus transcription by promoting OAS3-based activation of host innate immunity. Journal of medical virology 3 39011773
2025 Chromatin-associated α-satellite RNA maintains chromosome stability by reestablishing SAF-A in the mitotic cell cycle. Nucleic acids research 2 40219970
2023 ITGA9-AS1 up-regulates ITGA9 by targeting miR-4765 and recruiting HNRNPU to affect the proliferation and apoptosis of non-small cell lung cancer cells. Cellular and molecular biology (Noisy-le-Grand, France) 2 38279497
2025 The molecular axis hnRNPU/circKCNK2/EDC4/IL-11 aggravates osteolytic bone metastasis of RCC. Oncogene 1 40640337
2025 Correlated protein-RNA associations reveal a requirement for HNRNPU in long-range Polycomb recruitment by the lncRNAs Airn , Kcnq1ot1 , and Xist. bioRxiv : the preprint server for biology 1 40791421
2025 Long non-coding RNA STMN1P2 promotes breast cancer doxorubicin resistance by inhibiting pyroptosis through the hnRNPU-EZH2-TARF6-MALT1-caspase-1 pathway. Acta pharmacologica Sinica 1 40926024
2025 Role of the SAF-A/HNRNPU ATPase and RGG domains in X chromosome inactivation, nuclear dynamics, transcription, splicing, and cell proliferation. bioRxiv : the preprint server for biology 1 41473319
2024 Biological functions and clinic significance of SAF‑A (Review). Biomedical reports 1 38665420
2024 Case report: Early use of whole exome sequencing unveils HNRNPU-related neurodevelopmental disorder and answers additional clinical questions through reanalysis. Frontiers in genetics 1 38846959
2023 SAF-A/hnRNP U binds polyphosphoinositides via a lysine rich polybasic motif located in the SAP domain. microPublication biology 1 37038481
2026 Molecular interactome of HNRNPU reveals regulatory networks in neuronal differentiation and DNA methylation. Nucleic acids research 0 41674383
2025 MRO/HNRNPU/CCL5 feedback loop amplifies M2 macrophage and breast cancer cell crosstalk to drive progression. Journal of translational medicine 0 40817243