Affinage

DHX9

ATP-dependent RNA helicase A · UniProt Q08211

Length
1270 aa
Mass
141.0 kDa
Annotated
2026-06-09
100 papers in source corpus 46 papers cited in narrative 46 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

DHX9 is an abundant nuclear DExH-box helicase that maintains genome stability and shapes gene expression by recognizing and unwinding diverse non-canonical nucleic acid structures, including R-loops, DNA/RNA G-quadruplexes, intramolecular triplex H-DNA, and inverted-repeat Alu dsRNA (PMID:21561811, PMID:20669935, PMID:28355180). Its catalytic core is a RecA-like helicase domain structurally closer to viral NS3 helicases than to DEAD-box proteins, lacking base-selective contacts so that it hydrolyzes ATP, GTP, CTP, and UTP, and it unwinds substrates with 3'→5' polarity in an ATP-dependent manner (PMID:20510246, PMID:20669935). At the genome level, DHX9 resolves transcription-associated R-loops and processes triplex DNA to suppress mutagenesis and replication stress, and its loss drives premature senescence, aberrant R-loop accumulation, DNA double-strand breaks, and cGAS-STING- or dsRNA-triggered innate immune activation (PMID:30341290, PMID:24049074, PMID:24990949, PMID:38594251, PMID:38189443). It promotes homologous recombination by resolving R-loops at double-strand breaks and recruiting BRCA1, RPA, and RAD51, an activity coordinated by partners such as TDRD3, USP42, and Polη, and gated by ATR-mediated S321 phosphorylation and SUMO2 modification at K120 that license its association with R-loop-bound factors including γH2AX, RPA, PARP1, and DDX21 (PMID:34226554, PMID:34329467, PMID:32541651, PMID:35905379, PMID:37930853, PMID:39019926). As a transcriptional co-activator, DHX9 associates with RNA Polymerase II and bridges it to sequence-specific factors—NF-κB p65, STAT1, and the androgen receptor—and to Nup98, with its ATPase/helicase activity required for RNAPII recruitment to target promoters (PMID:30137501, PMID:36735791, PMID:35590370, PMID:28221134). Through its dsRNA-binding domains it controls post-transcriptional outcomes: it is required for RISC assembly, regulates A-to-I editing bidirectionally with ADAR, suppresses circular RNA biogenesis from inverted-repeat Alu elements, and unwinds rG4 structures in 5'-UTRs to govern translation (PMID:23361462, PMID:29796672, PMID:28355180, PMID:30591072). DHX9 additionally functions in innate antiviral immunity as a dsRNA sensor signaling through IPS-1/MAVS (PMID:21957149), and is exploited by numerous viruses as a host factor. Pathogenic DHX9 variants cause neurodevelopmental disorders and Charcot-Marie-Tooth disease, with distinct variants causing mislocalization, altered ATPase activity, or elevated R-loops and DNA damage (PMID:37467750).

Mechanistic history

Synthesis pass · year-by-year structured walk · 24 steps
  1. 2010 High

    Establishing the structural and biochemical basis of DHX9 catalysis answered how this helicase engages nucleotides and nucleic acid substrates, distinguishing it mechanistically from DEAD-box enzymes.

    Evidence X-ray crystallography of helicase domain 1 plus NTPase assays, and in vitro triplex helicase assays with kinetic analysis

    PMID:20510246 PMID:20669935

    Open questions at the time
    • No full-length structure with substrate engaged
    • Mechanism of substrate-structure selectivity not resolved at atomic level
  2. 2010 Medium

    Demonstrating cooperative unwinding with WRN at RNA-containing replication intermediates placed DHX9 at the replication fork, though without in vivo validation.

    Evidence In vitro helicase assays with Okazaki-fragment-like and chicken-foot substrates plus protein-protein interaction assays

    PMID:20385589

    Open questions at the time
    • No in vivo confirmation of cooperative mechanism
    • Direct interaction interface not mapped
  3. 2011 High

    Defining R-loops and G-quadruplexes as preferred substrates over simple duplexes identified the physiologically relevant structures DHX9 acts on.

    Evidence In vitro helicase assays on defined partial duplex, D-loop, R-loop, and G-quadruplex substrates

    PMID:21561811

    Open questions at the time
    • Cellular substrate preference not directly tested in this study
    • No genome-wide substrate map
  4. 2011 High

    Identifying DHX9 as a dsRNA sensor signaling through IPS-1/MAVS established a role in innate immune detection of viral RNA, defining the domain interfaces involved.

    Evidence siRNA knockdown, dsRNA binding assays, co-IP with HelicC-HA2-DUF/CARD domain mapping, cytokine readouts in dendritic cells

    PMID:21957149

    Open questions at the time
    • Relative contribution versus canonical RNA sensors unclear
    • In vivo requirement not established here
  5. 2013 High

    Structural and functional dissection of the dsRNA-binding domains showed how DHX9 is incorporated into RISC, linking its dsRNA recognition to gene silencing.

    Evidence Crystal structures of dsRBD1/dsRBD2 with dsRNA, ITC, mutagenesis-validated co-IP

    PMID:23361462

    Open questions at the time
    • Coupling between dsRBD binding and helicase core not resolved
    • Step in RISC assembly not pinpointed
  6. 2013 High

    Showing that DHX9 depletion increases H-DNA-induced mutagenesis demonstrated an in vivo genome-protective role at triplex-forming sequences.

    Evidence siRNA mutation-reporter assay, in vitro binding, ChIP at H-DNA regions in U2OS cells

    PMID:24049074

    Open questions at the time
    • Mechanism converting unresolved triplex to deletions unclear
    • Downstream repair factors not identified
  7. 2014 High

    Linking DHX9 to p53-dependent senescence and origin firing established its essentiality for DNA replication and proliferation.

    Evidence siRNA/shRNA knockdown, SA-β-gal assay, ChIP at replication origins, nascent DNA labeling in human fibroblasts

    PMID:24990949

    Open questions at the time
    • Molecular role at origins not defined
    • Connection to R-loop resolution at origins untested here
  8. 2017 High

    Revealing DHX9 control of inverted-repeat Alu dsRNA and circular RNA production, with an RNA-independent ADAR1-p150 interaction, connected dsRNA structure resolution to circRNA biogenesis and editing.

    Evidence Biochemical purification, RIP, reporter assays, circRNA quantification, ADAR/DHX9 co-depletion and co-IP

    PMID:28355180

    Open questions at the time
    • Catalytic mechanism on Alu dsRNA in cells not directly shown
    • Functional consequence of ADAR1-p150 interaction incompletely defined
  9. 2017 High

    Defining DHX9 processing of IGS-rRNA into pRNA to guide TIP5 placed DHX9 in heterochromatin establishment and stem cell differentiation.

    Evidence RIP, ChIP, knockdown with rescue by defined RNA species, ESC differentiation assays

    PMID:28588071

    Open questions at the time
    • Whether helicase activity per se generates pRNA not isolated
    • Generality across rRNA loci unclear
  10. 2017 High

    Identifying Nup98 as a stimulator of DHX9 ATPase and co-occupant of gene loci connected nucleoporin function to DHX9-dependent transcription.

    Evidence In vitro binding and ATPase assays, ChIP-seq, transcriptional reporter, co-IP

    PMID:28221134

    Open questions at the time
    • Mechanism of ATPase stimulation not structurally defined
    • Subnuclear site of action unclear
  11. 2018 High

    Showing DHX9-driven R-loop formation through prolonged RNAPII association upon splicing impairment defined a context where DHX9 generates rather than resolves hybrids that trap Pol II and block replication.

    Evidence S9.6 IF, ChIP, DRIP-seq, proximity ligation, splicing-factor knockdown

    PMID:30341290

    Open questions at the time
    • Reconciling R-loop-forming versus R-loop-resolving roles unresolved
    • Determinants of switch between activities unknown
  12. 2018 High

    Transcriptome-wide profiling showed DHX9 binds 5'-UTR rG4 substrates to regulate uORF and downstream translation, extending its role to translational control.

    Evidence Ribo-seq, CLIP-seq, knockdown, luciferase reporters

    PMID:30591072

    Open questions at the time
    • Division of labor with DHX36 not fully separated
    • In vivo translational impact untested
  13. 2018 High

    Establishing the DHX9-p65-RNAPII axis showed DHX9 is required for RNAPII recruitment to NF-κB promoters via its helicase activity and contributes to antiviral immunity against DNA viruses.

    Evidence Macrophage-specific KO, fibroblast KD, multiprotein co-IP, ChIP for p65/RNAPII, helicase-dead mutant, viral assays

    PMID:30137501

    Open questions at the time
    • How helicase activity enables Pol II loading mechanistically unclear
    • Promoter structures unwound not identified
  14. 2018 Medium

    Bidirectional regulation of A-to-I editing demonstrated DHX9 differentially modulates ADAR1 versus ADAR2 substrate editing, partly via helicase activity.

    Evidence siRNA knockdown, transcriptome-wide editing analysis, helicase-dead mutant in cancer cells

    PMID:29796672

    Open questions at the time
    • Mechanistic basis of ADAR isoform specificity unresolved
    • Single lab without structural support
  15. 2021 High

    Defining DHX9-mediated BRCA1 recruitment to RNA at damage sites placed DHX9 mechanistically within homologous recombination and end resection.

    Evidence Knockdown, RPA/RAD51 foci, co-IP, HR reporter, camptothecin/Olaparib sensitivity

    PMID:34226554

    Open questions at the time
    • Direct biochemical role in resection not reconstituted
    • RNA species recruiting BRCA1 not defined
  16. 2021 High

    Identifying TDRD3 as a recruiter and stimulator of DHX9 helicase activity at promoters established a mechanism for targeted R-loop resolution.

    Evidence Reciprocal co-IP with domain mapping, ChIP, DRIP, in vitro helicase stimulation

    PMID:34329467

    Open questions at the time
    • Promoter selectivity determinants unclear
    • Interplay with other R-loop resolvers not addressed
  17. 2020 Medium

    USP42 was shown to partner DHX9 for R-loop resolution at DSBs and HR, reinforcing the helicase-dependent R-loop/BRCA1 axis.

    Evidence Co-IP, knockdown, HR reporter, DRIP, BRCA1 foci

    PMID:32541651

    Open questions at the time
    • Whether USP42 deubiquitinates DHX9 not established
    • Single lab
  18. 2022 Medium

    Polη was identified as recruiting DHX9 to G4 loci, defining cooperative action enabling replication through G-quadruplex regions.

    Evidence BioID, affinity pulldown-MS, ChIP-seq, co-IP, replication assay at G4 loci

    PMID:35905379

    Open questions at the time
    • Order of recruitment versus unwinding unclear
    • Direct interaction interface unmapped
  19. 2023 High

    Pathogenic DHX9 variants causally linking it to neurodevelopmental disorders and CMT defined genotype-specific mechanisms (mislocalization, altered ATPase, elevated R-loops/DSBs).

    Evidence Exome sequencing with family analysis, variant expression and localization, ATPase assays, R-loop and γH2AX measurement, Dhx9 knockout mice

    PMID:37467750

    Open questions at the time
    • Cell-type basis of neuronal versus sensory phenotypes unclear
    • How distinct molecular defects map to distinct diseases not fully resolved
  20. 2023 High

    Defining DHX9-STAT1-RNAPII recruitment to ISG promoters established a MAVS-independent, interferon-downstream antiviral transcription role distinct from its dsRNA-sensor function.

    Evidence Cell-type-specific KO mice, double-KO epistasis with MAVS and STAT1, co-IP, ChIP at ISG promoters, viral infection

    PMID:36735791

    Open questions at the time
    • Reconciling MAVS-dependent sensing with MAVS-independent transcription unclear
    • Promoter-level helicase substrate not defined
  21. 2024 High

    ATR-mediated S321 phosphorylation was shown to gate DHX9 association with R-loops and damage factors, providing a damage-responsive regulatory switch.

    Evidence Phospho-mapping MS, ATR inhibition, phosphomimetic/non-phosphorylatable mutants, co-IP, DRIP, in vitro DHX9-RPA binding, drug sensitivity

    PMID:37930853

    Open questions at the time
    • Structural effect of S321 phosphorylation unknown
    • Kinetics relative to damage signaling unresolved
  22. 2024 High

    SUMO2 modification at K120 was shown to license SIM-dependent interactions with PARP1 and DDX21 required for genome stability, adding a second post-translational control of R-loop function.

    Evidence K120R mutagenesis, SUMO2 fusion rescue, co-IP with SIM-dependent partners, DRIP, γH2AX IF, viability assays

    PMID:39019926

    Open questions at the time
    • SUMO E3 ligase not identified
    • Interplay between K120 SUMO and S321 phosphorylation untested
  23. 2024 High

    Genetic studies established DHX9 as a therapeutic target by showing its loss triggers dual cytoplasmic dsRNA innate immunity and R-loop-driven DNA damage, enhancing checkpoint-blockade response in SCLC and driving cGAS-STING inflammation in intestinal stem cells.

    Evidence CRISPR screen and KO with in vivo tumor models and immune profiling; conditional Dhx9 KO mice with DRIP and cGAS-STING readouts and colitis models

    PMID:38189443 PMID:38594251

    Open questions at the time
    • Relative contribution of dsRNA versus R-loop arms to phenotypes unclear
    • Tissue-specific thresholds for DHX9 loss unknown
  24. 2024 High

    Discovery of DHX9-specific stress granules enriched in damaged intron RNA defined a distinct cytoplasmic dsRNA-handling compartment promoting survival after UV damage.

    Evidence FANCI damaged-RNA mapping, IF, SG-fraction RNA-seq, p62 knockout/knockdown, viability assays

    PMID:38503283

    Open questions at the time
    • Trigger of DHX9 SG nucleation not fully defined
    • Relationship to nuclear DHX9 functions unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • How DHX9 switches between R-loop formation and resolution, and how its many post-translational modifications and partner proteins are integrated to direct it among genome-protective, transcriptional, translational, and antiviral roles, remains unresolved.
  • No unified model coordinating modification states with substrate choice
  • Structural basis for partner-directed targeting lacking
  • Determinants of nuclear versus cytoplasmic functional partitioning unclear

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003723 RNA binding 4 GO:0140097 catalytic activity, acting on DNA 3 GO:0140098 catalytic activity, acting on RNA 3 GO:0140110 transcription regulator activity 3 GO:0140657 ATP-dependent activity 3 GO:0003677 DNA binding 2 GO:0016787 hydrolase activity 2 GO:0140299 molecular sensor activity 1
Localization
GO:0005829 cytosol 4 GO:0005634 nucleus 3 GO:0000228 nuclear chromosome 2 GO:0005730 nucleolus 2
Pathway
R-HSA-168256 Immune System 4 R-HSA-74160 Gene expression (Transcription) 4 R-HSA-8953854 Metabolism of RNA 4 R-HSA-1643685 Disease 3 R-HSA-69306 DNA Replication 3 R-HSA-73894 DNA Repair 3 R-HSA-8953897 Cellular responses to stimuli 2
Partners
ADARARBRCA1NUP98RELARPASTAT1TDRD3
Complex memberships
RISCRNA Polymerase II transcription complex

Evidence

Reading pass · 46 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2017 DHX9 binds specifically to inverted-repeat Alu elements transcribed as parts of genes; loss of DHX9 increases circular RNA production, causes translational repression of reporters containing inverted-repeat Alu elements, and produces transcriptional rewiring. DHX9 interacts with the interferon-inducible ADAR isoform p150 (but not p110) in an RNA-independent manner. Co-depletion of ADAR and DHX9 augments dsRNA accumulation and circular RNA production, revealing a functional link between the two enzymes. Biochemical purification, RNA immunoprecipitation, reporter assays, siRNA knockdown, circular RNA quantification, co-IP Nature High 28355180
2011 DHX9 preferentially unwinds RNA-containing displacement loops (R-loops) and DNA/RNA-based G-quadruplexes over simple DNA-DNA duplexes in vitro, suggesting these structures are biologically relevant substrates. In vitro helicase assays using defined nucleic acid substrates (partial duplexes, D-loops, R-loops, G-quadruplexes) DNA repair High 21561811
2011 DHX9 functions as a dsRNA sensor in myeloid dendritic cells, binding polyinosine-polycytidylic acid via its dsRNA-binding motifs and interacting with the adaptor IPS-1 (MAVS) via DHX9's HelicC-HA2-DUF domain and IPS-1's CARD domain to activate NF-κB and IRF3 signaling in response to dsRNA. siRNA knockdown, dsRNA binding assays, co-immunoprecipitation with domain mapping, cytokine production assays Journal of immunology High 21957149
2018 DHX9 (RNA helicase A) promotes the formation of R-loops in cells with impaired splicing through prolonged association with RNA Polymerase II, producing DNA-RNA hybrids that trap RNA Pol II on chromatin and can block DNA replication. R-loop immunofluorescence (S9.6 antibody), chromatin immunoprecipitation, DRIP-seq, proximity ligation assay, siRNA knockdown of splicing factors Nature communications High 30341290
2018 DHX9 and DHX36 helicase depletion promotes translation of rG4-associated upstream open reading frames (uORFs) while reducing translation of downstream coding regions; transcriptome-wide binding analysis shows DHX9 directly interacts with rG4 substrates in 5'-UTRs to regulate translation. Ribosome profiling (Ribo-seq), CLIP-seq for DHX9 binding site identification, siRNA knockdown, luciferase reporter assays Genome biology High 30591072
2010 DHX9 unwinds triple-helical DNA structures (H-DNA/intramolecular triplexes) in vitro with 3'→5' polarity with respect to the displaced third strand, requiring a 3'-single-stranded overhang and ATP hydrolysis; DHX9 co-immunoprecipitates with triplex DNA structures in mammalian cells. In vitro helicase assay with triplex substrates, co-immunoprecipitation with triplex DNA, ATP hydrolysis assays, kinetic analysis Biochemistry High 20669935
2010 DHX9 stimulates WRN helicase unwinding of RNA-containing Okazaki fragment-like substrates and RNA-containing 'chicken-foot' structures in vitro, suggesting cooperative action at replication forks; DHX9 and WRN engage in direct protein-protein interaction. In vitro helicase assays with synthetic substrates, protein-protein interaction assays Nucleic acids research Medium 20385589
2013 DHX9 is required for RISC assembly via its dsRNA-binding domains (dsRBD1 and dsRBD2); crystal structures of dsRBD1 and dsRBD2 in complex with dsRNA reveal that dsRBD1 has higher siRNA-binding affinity and a crystallographic dsRBD1 pair cooperatively recognizes dsRNA. Both dsRBDs are required for RISC association mediated by dsRNA. Crystal structure determination, isothermal titration calorimetry (ITC), co-immunoprecipitation with mutagenesis Nucleic acids research High 23361462
2013 DHX9 depletion increases mutagenesis at H-DNA structures in human cells, predominantly causing deletions; DHX9 associates with H-DNA in supercoiled plasmids and is enriched at H-DNA regions in chromatin by ChIP, implicating DHX9 in processing intramolecular triplex DNA in vivo. siRNA knockdown mutation-reporter assay, in vitro binding assay, chromatin immunoprecipitation (ChIP) in U2OS cells Nucleic acids research High 24049074
2010 Crystal structure of the conserved helicase domain 1 (DEIH motif) of human DHX9 reveals a RecA-like core more similar to viral NS3 helicases than DEAD-box proteins; DHX9 lacks base-selective contacts and forms an unspecific stacking interaction with the nucleotide base, allowing hydrolysis of ATP, GTP, CTP, and UTP. X-ray crystallography, NTPase biochemical assays Journal of molecular biology High 20510246
2017 DHX9 binds to IGS-rRNA at rRNA genes upon embryonic stem cell (ESC) differentiation, processes IGS-rRNA into pRNA, and thereby guides the repressor TIP5 to establish heterochromatin at rRNA genes; DHX9 depletion prevents ESC differentiation, a phenotype rescued by addition of pRNA but not IGS-rRNA or TIP5-binding-deficient pRNA mutants. RNA immunoprecipitation, ChIP, siRNA knockdown, differentiation assays, rescue experiments with RNA addition EMBO reports High 28588071
2021 DHX9 promotes BRCA1 recruitment to RNA at sites of DNA damage as part of the RNA Polymerase II transcription complex, facilitating DNA end resection and homologous recombination (HR). DHX9-deficient cells fail to recruit RPA and RAD51 to DSBs, are defective in HR, hypersensitive to camptothecin and Olaparib, and show impaired ATR-mediated damage signaling and replication restart. siRNA knockdown, immunofluorescence for RPA/RAD51 foci, co-immunoprecipitation, HR reporter assay, drug sensitivity assays Nature communications High 34226554
2021 TDRD3 directly interacts with DHX9 via its Tudor domain, recruits DHX9 to target gene promoters to resolve R-loops in a helicase-activity-dependent manner, and stimulates DHX9 helicase activity via its OB-fold domain that likely binds the ssDNA in the R-loop. Co-immunoprecipitation, ChIP, DRIP (DNA-RNA immunoprecipitation), helicase activity assays, domain-mapping mutagenesis Nucleic acids research High 34329467
2014 Suppression of DHX9 in primary human diploid fibroblasts induces premature senescence in a p53-dependent manner; DHX9 associates with origins of replication by ChIP, and its suppression reduces nascent DNA synthesis, demonstrating an essential role in DNA replication and cell cycle progression. siRNA/shRNA knockdown, senescence-associated β-galactosidase assay, ChIP at replication origins, nascent DNA labeling, gene expression profiling Journal of biological chemistry High 24990949
2018 DHX9 functions as a bidirectional regulator of A-to-I RNA editing in cancer cells, at least partially dependent on its helicase activity: DHX9 silencing preferentially represses ADAR1-specific editing while augmenting ADAR2-specific substrate editing. siRNA knockdown, transcriptome-wide RNA editing analysis, helicase-dead mutant expression Nucleic acids research Medium 29796672
2018 DHX9 forms a complex with RNA Polymerase II and EWS-FLI1 to enhance transcription in Ewing sarcoma cells; UV irradiation causes enhanced RNAPII phosphorylation and decreased processivity, leading to alternative splicing inclusion of DHX9 exon 6A that targets DHX9 mRNA to NMD-mediated decay, reducing DHX9 protein levels. Co-immunoprecipitation, ChIP, siRNA knockdown, minigene splicing assays, RT-PCR for alternative isoforms, UV irradiation Oncotarget Medium 26450900
2017 Human Nup98 binds to DHX9 via its FG/GLFG repeat region interacting with N- and C-terminal regions of DHX9 in an RNA-facilitated manner; Nup98 binding stimulates DHX9 ATPase activity and supports DHX9-stimulated transcription; Nup98 and DHX9 bind interdependently to similar gene loci. In vitro binding assays, ATPase activity assays, transcriptional reporter assay, ChIP-seq, co-immunoprecipitation eLife High 28221134
2013 Myxoma virus M029 protein interacts with DHX9/RHA in a dsRNA-independent manner in infected cells; M029 not only inhibits PKR but also conscripts DHX9 as a pro-viral effector to promote viral replication specifically in myeloid THP1 cells, as PKR depletion rescue of M029-minus viruses is reversed by DHX9 knockdown. Proteomic co-immunoprecipitation, siRNA knockdown of DHX9, virus replication assays, epistasis (PKR KD + DHX9 KD double knockdown) PLoS pathogens Medium 23853588
2016 PRRSV nucleocapsid (N) protein interacts with cellular DHX9 and redistributes it from the nucleus to the cytoplasm; knockdown of DHX9 increases production of short subgenomic mRNAs while DHX9 overexpression promotes synthesis of longer subgenomic mRNAs and genomic RNA, suggesting DHX9 acts as an antiattenuation factor for continuous elongation during viral RNA synthesis. Co-immunoprecipitation, siRNA knockdown, overexpression, viral RNA quantification by RT-qPCR, immunofluorescence for localization Journal of virology Medium 27009951
2016 DHX9/RHA binds to the double-stranded PBS-segment of the HIV-1 5'-UTR dimer; ITC shows one RHA molecule binds per RNA strand independently of nucleocapsid protein; NMR with deuterium labeling defines the structural basis of this interaction; disruption of this binding (single A-to-C substitution) attenuates virion infectivity. Isothermal titration calorimetry (ITC), NMR spectroscopy, cell-based viral infectivity assays, mutagenesis Journal of molecular biology High 27107641
2019 Virion-associated DHX9/RHA enhances the processivity of HIV-1 reverse transcriptase (RT) during elongation phase of (-)cDNA synthesis; pre-steady-state and steady-state kinetics show RHA has little impact on single-nucleotide incorporation rate but increases RT processivity, as demonstrated by primer extension in the presence of trap dsDNA. In vitro reverse transcription reconstitution, pre-steady-state and steady-state kinetic assays, primer extension with trap assay, quantification of reverse-transcription intermediates in infected T cells Journal of biological chemistry High 31175158
2019 DHX9 is recruited to CHIKV replication complexes at the plasma membrane via interaction with nsP3's C-terminal hypervariable domain; during early infection DHX9 negatively controls viral RNA synthesis but is required for optimal translation of mature nonstructural proteins; at later stages DHX9 is degraded via the proteasome. Co-immunoprecipitation, siRNA knockdown, immunofluorescence for colocalization, viral RNA quantification, proteasome inhibitor experiments Journal of virology Medium 30463980
2014 During FMDV infection, JMJD6 arginine demethylase physically interacts with DHX9 (preferring JMJD6 monomers) and demethylates RHA arginine residues at its C-terminus; this demethylation promotes DHX9 redistribution from nucleus to cytoplasm and facilitates viral protein synthesis and virus replication. Reciprocal co-immunoprecipitation, NOG inhibitor of JmjC demethylases, modified biochemical demethylation assay, immunofluorescence for subcellular localization Virology Medium 24606677
2004 DHX9/RHA is SUMOylated by Ubc9 (E2 SUMO ligase); the N-terminal domain (residues 1–137) of RHA is sufficient for Ubc9 interaction; Ubc9 interaction activates RHA transcriptional activity in a CREB-mediated reporter assay, and Ubc9 catalytic activity is dispensable for the transcription activation function. Co-immunoprecipitation, domain-mapping truncation mutants, in vitro and in vivo SUMO conjugation assay, transcriptional reporter assay Journal of molecular biology Medium 15312759
2024 ATR kinase phosphorylates DHX9 at serine 321 (S321) upon DNA damage; this phosphorylation promotes DHX9 interaction with γH2AX, BRCA1, and RPA, and is required for DHX9 association with R-loops under genotoxic stress. Non-phosphorylatable DHX9-S321A causes R-loop accumulation and hypersensitivity to genotoxic agents; phosphomimetic DHX9-S321D prevents R-loop accumulation and confers resistance. In vitro binding confirms direct DHX9–RPA interaction. Phospho-mapping mass spectrometry, ATR inhibitor treatment, phosphomimetic/non-phosphorylatable mutants, co-immunoprecipitation, DRIP assay, in vitro binding assay, drug sensitivity assays Nucleic acids research High 37930853
2024 DHX9 SUMOylation at lysine 120 (K120) is required for genome stability; SUMO2 modification at K120 enables DHX9 to interact with R-loop-associated proteins PARP1 and DDX21 via SUMO-interacting motifs. The K120R mutant (non-SUMOylatable) shows R-loop accumulation, increased DNA damage, and cell death; fusing SUMO2 to K120R rescues these defects. Site-directed mutagenesis (K120R), SUMO2 fusion rescue construct, co-immunoprecipitation, DRIP assay, γH2AX immunofluorescence, cell viability assays Nature communications High 39019926
2024 DHX9 depletion in SCLC leads to cytoplasmic dsRNA accumulation triggering tumor-intrinsic innate immunity, and simultaneously causes aberrant R-loop accumulation leading to DNA damage-derived cytoplasmic DNA and replication stress; in vivo, DHX9 deletion reduces tumor growth and enhances immunogenic tumor microenvironment, improving response to immune-checkpoint blockade. CRISPR-based screen, CRISPR knockout, immunofluorescence for dsRNA and R-loops, DNA damage markers, in vivo tumor models, immune profiling Cancer discovery High 38189443
2024 DHX9 forms stress granules (SGs) distinct from classical SGs upon UV irradiation; DHX9 SGs are enriched in damaged intron RNA rather than mature mRNA. UV-induced RNA crosslinking impedes intron splicing and decay, triggering DHX9 SGs in daughter cells. DHX9 modulates dsRNA abundance within SGs and promotes cell survival; autophagy receptor p62 is required for DHX9 SG disassembly. FANCI technology for damaged RNA identification, immunofluorescence, RNA sequencing of SG fractions, p62 knockout/knockdown, cell viability assays Cell High 38503283
2024 DHX9 deficiency in intestinal stem cells (ISCs) causes abnormal R-loop accumulation leading to genomic instability and cGAS-STING-mediated inflammatory response, impairing ISC function; conditional Dhx9 knockout in intestinal epithelium increases susceptibility to experimental colitis with reduction in ISC and Paneth cell numbers. Conditional ISC-specific and Paneth cell-specific Dhx9 knockout mice, DRIP for R-loops, cGAS-STING pathway marker analysis, histology, colitis models Nature communications High 38594251
2023 Nuclear DHX9 directly binds STAT1 and recruits RNA Pol II to interferon-stimulated gene (ISG) promoters to participate in STAT1-mediated ISG transcription; myeloid- or hepatocyte-specific DHX9 knockout mice are more susceptible to RNA virus infections, and this antiviral role is downstream of type I interferon, dependent on STAT1 (not MAVS). Myeloid- and hepatocyte-specific DHX9 knockout mice, double-KO with MAVS or STAT1, co-immunoprecipitation, ChIP for Pol II at ISG promoters, viral infection models Science advances High 36735791
2018 DHX9 interacts with p65 (NF-κB subunit) and RNA Polymerase II in chromatin at NF-κB binding sites; nuclear DHX9 is essential for recruitment of RNAPII (but not p65) to NF-κB target gene promoters, requiring its ATPase/helicase activity; macrophage- and fibroblast-specific DHX9 knockout/knockdown impairs antiviral innate immunity against DNA viruses. Macrophage-specific KO, fibroblast-specific KD, co-immunoprecipitation of multiprotein complex, ChIP for p65 and RNAPII at NF-κB promoters, helicase-dead mutant, viral replication assays Nucleic acids research High 30137501
2021 DHX9 interacts with p65 and RNAPII to enhance p65 phosphorylation and nuclear translocation, facilitating NF-κB-mediated transcription of downstream targets (Survivin, Snail) to promote colorectal cancer malignant phenotypes. Co-immunoprecipitation, ChIP, siRNA knockdown, overexpression, xenograft and hepatic metastasis mouse models Cellular and molecular life sciences Medium 34773477
2019 DHX9 binds the 3'-UTR of CDK6 mRNA and post-transcriptionally suppresses CDK6 expression; lnc-UCID (lncRNA) interacts with DHX9 via its 850-1030 nt domain and sequesters DHX9 from CDK6-3'UTR, thereby enhancing CDK6 expression and promoting G1/S transition. RNA immunoprecipitation, RNA pulldown, co-immunoprecipitation, luciferase reporter of CDK6-3'UTR, siRNA knockdown, gain/loss-of-function studies, xenograft models Hepatology Medium 30865310
2018 DHX9 is associated with HBV viral RNAs; DHX9 knockdown increases HBV-derived circular RNA production while decreasing viral protein levels (without affecting HBV DNA levels), demonstrating DHX9 is a novel regulator of viral circular RNA biogenesis. Co-immunoprecipitation/RNA binding assay, siRNA knockdown, circular RNA quantification, viral protein and DNA assays in HBV-producing cell lines and primary hepatocytes Oncotarget Medium 29765512
2019 DHX9 interacts with the EBV SM protein (co-immunoprecipitation, nuclear co-localization) and restricts EBV lytic replication; DHX9 depletion increases EBV late gene expression and virion production; DHX9 enhances innate antiviral pathways (ISGs), and SM inhibits DHX9's transcription-activating function through cAMP response elements (CREs). Mass spectrometry, co-immunoprecipitation, immunofluorescence, siRNA knockdown, viral gene expression and titer assays, CRE reporter assay Journal of virology Medium 30541834
2019 DHX9 induces alternative splicing of KLF5 mRNA by bridging YB-1 protein in vascular smooth muscle cells; smooth muscle cell-specific DHX9 conditional knockout alters transcriptional and alternative splicing profiles of smooth muscle cell phenotype-related genes and exacerbates aortic dissection. Vascular smooth muscle cell-specific DHX9 conditional knockout mice, RNA-seq, alternative splicing analysis, co-immunoprecipitation (DHX9-YB-1 interaction), aortic dissection model American journal of physiology. Cell physiology Medium 31116584
2022 DHX9 interacts with Androgen Receptor (AR) in prostate cancer cells; AR binds the DHX9 promoter to induce DHX9 expression; DHX9 depletion reduces AR recruitment to target gene promoters and AR-driven transcription in response to DHT; enzalutamide-mediated AR inhibition represses DHX9 expression, establishing a DHX9/AR positive feedback axis. Co-immunoprecipitation, ChIP, RNA-seq, promoter reporter assay, siRNA knockdown, cell proliferation and migration assays Journal of experimental & clinical cancer research Medium 35590370
2022 Pol η (DNA polymerase eta) interacts with DHX9 (identified by proximity labeling and affinity pulldown-MS); Pol η promotes DHX9 recruitment to G4-structure loci in chromatin; deficiency in either Pol η or DHX9 impairs replication across G4 regions, indicating cooperative action at G4 sites during replication. Proximity labeling (BioID), affinity pulldown with LC-MS/MS, ChIP-seq, co-immunoprecipitation, replication assay at G4 loci, siRNA knockdown Journal of the American Chemical Society Medium 35905379
2020 USP42 interacts with DHX9 and promotes R-loop resolution at DNA double-strand breaks (DSBs); USP42 localization to nuclear speckles is required for efficient HR; DHX9's DNA-RNA helicase activity is required for DSB-induced R-loop resolution and BRCA1 recruitment. Co-immunoprecipitation, siRNA knockdown, HR reporter assay, DRIP for R-loops, BRCA1 foci immunofluorescence Oncogenesis Medium 32541651
2024 AKT1 kinase domain directly interacts with DHX9 and facilitates DHX9 recruitment to R-loops; AKT inhibition reduces DHX9 association with R-loops and increases ATRi-induced R-loop-mediated replication stress; the interaction is direct as shown by in vitro binding analysis. Quantitative high-throughput drug combination screen, co-immunoprecipitation, in vitro binding assay, DRIP for R-loops, replication stress markers Cancer research Medium 38241710
2020 Oxaliplatin induces PI3KK-mediated phosphorylation of DHX9 at two sites near substrate-binding domains; non-phosphorylatable DHX9 mutants or PI3KK inhibitor blockage decreases oxaliplatin-induced circCCDC66 circular RNA expression and impairs development of chemoresistance. Site-directed mutagenesis (non-phosphorylatable DHX9 mutants), PI3KK inhibitors, circular RNA quantification, drug resistance assays Cancers Medium 32187976
2023 DHX9 pathogenic variants cause neurodevelopmental disorders and Charcot-Marie-Tooth disease; NLS missense variants cause abnormal cytoplasmic DHX9 accumulation (loss of nuclear localization), CMT2-associated variants cause aberrant nucleolar DHX9 accumulation, and an NDD variant (p.Arg141Gln) increases R-loop levels and DNA double-strand breaks without altering localization. Two variants (p.Gly411Glu, p.Arg761Gln) alter DHX9 ATPase activity. Dhx9-/- mice show hypoactivity, tremor, and sensorineural hearing loss. Exome sequencing with family analysis, human cell line DHX9 variant expression (localization by immunofluorescence), patient fibroblast localization, ATPase activity assays, R-loop measurement (S9.6), γH2AX foci, Dhx9 knockout mice American journal of human genetics High 37467750
2018 MDM2 E3 ubiquitin ligase interacts with DHX9 and mediates its ubiquitin-proteasome-dependent degradation; lnc-CCDST lncRNA binds DHX9 and serves as a scaffold to facilitate the DHX9–MDM2 interaction, promoting DHX9 degradation and suppressing cervical cancer motility and angiogenesis. Co-immunoprecipitation, ubiquitination assay, siRNA knockdown, overexpression, in vitro and in vivo migration/invasion/angiogenesis assays Cell death and differentiation Medium 30518908
2021 SPOP E3 ubiquitin ligase binds DHX9 via a canonical SPOP-binding motif in DHX9 and induces its ubiquitination and proteasomal degradation; SPOP depletion increases DHX9 levels and promotes choriocarcinoma migration and invasion; DHX9 siRNA partially rescues the pro-migratory effect of SPOP knockdown. Co-immunoprecipitation, ubiquitination assay, siRNA knockdown, binding motif identification, migration/invasion assays American journal of cancer research Medium 32905556
2019 DHX9 interacts with APOBEC3B (A3B) in the presence of HBV; DHX9 does not affect A3B deaminase activity but inhibits A3B binding to HBV pregenomic RNA (pgRNA), thereby attenuating A3B's anti-HBV effect. Co-immunoprecipitation and mass spectrometry, in vitro deaminase assay, pgRNA binding assay, HBV DNA replication quantification Emerging microbes & infections Medium 32056513
2025 DHX9 unwinds inhibitory RNA G-quadruplex (rG4) structures near the translation initiation site of PCMT1 mRNA; circCLASP2 acts as a molecular scaffold bringing DHX9 to PCMT1 mRNA, increasing PCMT1 translation and promoting cytoskeletal changes in nasopharyngeal carcinoma. RNA immunoprecipitation, RNA pulldown, ribosomal gradient sedimentation, dual-luciferase reporter, immunofluorescence, in vitro and in vivo NPC assays Molecular cancer Medium 40050914

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2017 DHX9 suppresses RNA processing defects originating from the Alu invasion of the human genome. Nature 456 28355180
2018 Structural basis of G-quadruplex unfolding by the DEAH/RHA helicase DHX36. Nature 267 29899445
2011 Human DHX9 helicase preferentially unwinds RNA-containing displacement loops (R-loops) and G-quadruplexes. DNA repair 209 21561811
2003 Divergent stress responses and coping styles in psychogenetically selected Roman high-(RHA) and low-(RLA) avoidance rats: behavioural, neuroendocrine and developmental aspects. Stress (Amsterdam, Netherlands) 184 12775328
2021 LINC00460/DHX9/IGF2BP2 complex promotes colorectal cancer proliferation and metastasis by mediating HMGA1 mRNA stability depending on m6A modification. Journal of experimental & clinical cancer research : CR 176 33526059
2011 DHX9 pairs with IPS-1 to sense double-stranded RNA in myeloid dendritic cells. Journal of immunology (Baltimore, Md. : 1950) 157 21957149
2018 DHX9 helicase promotes R-loop formation in cells with impaired RNA splicing. Nature communications 152 30341290
2016 The biology of DHX9 and its potential as a therapeutic target. Oncotarget 143 27034008
2018 A DHX9-lncRNA-MDM2 interaction regulates cell invasion and angiogenesis of cervical cancer. Cell death and differentiation 133 30518908
2018 RNA G-quadruplexes at upstream open reading frames cause DHX36- and DHX9-dependent translation of human mRNAs. Genome biology 128 30591072
2010 Human DHX9 helicase unwinds triple-helical DNA structures. Biochemistry 92 20669935
1997 Neuroendocrine correlates of emotional reactivity and coping in male rats from the Roman high (RHA/Verh)- and low (RLA/Verh)-avoidance lines. Behavior genetics 92 9476359
2019 Lnc-UCID Promotes G1/S Transition and Hepatoma Growth by Preventing DHX9-Mediated CDK6 Down-regulation. Hepatology (Baltimore, Md.) 90 30865310
2024 Targeting DHX9 Triggers Tumor-Intrinsic Interferon Response and Replication Stress in Small Cell Lung Cancer. Cancer discovery 77 38189443
2013 DHX9 helicase is involved in preventing genomic instability induced by alternatively structured DNA in human cells. Nucleic acids research 75 24049074
2024 RNA damage compartmentalization by DHX9 stress granules. Cell 66 38503283
2017 Transport Study of Egg-Derived Antihypertensive Peptides (LKP and IQW) Using Caco-2 and HT29 Coculture Monolayers. Journal of agricultural and food chemistry 66 28782363
2016 Porcine Reproductive and Respiratory Syndrome Virus Nucleocapsid Protein Interacts with Nsp9 and Cellular DHX9 To Regulate Viral RNA Synthesis. Journal of virology 62 27009951
2015 Regulation of DEAH/RHA helicases by G-patch proteins. BioMed research international 58 25692149
2014 5-HT(2A) and mGlu2 receptor binding levels are related to differences in impulsive behavior in the Roman Low- (RLA) and High- (RHA) avoidance rat strains. Neuroscience 56 24412375
1998 Long-term behavioural and neuroendocrine changes in Roman high-(RHA/Verh) and low-(RLA-Verh) avoidance rats following neonatal handling. International journal of developmental neuroscience : the official journal of the International Society for Developmental Neuroscience 56 9785113
2021 DHX9-dependent recruitment of BRCA1 to RNA promotes DNA end resection in homologous recombination. Nature communications 54 34226554
2013 Myxoma virus protein M029 is a dual function immunomodulator that inhibits PKR and also conscripts RHA/DHX9 to promote expanded host tropism and viral replication. PLoS pathogens 54 23853588
2020 Oxaliplatin-Induced DHX9 Phosphorylation Promotes Oncogenic Circular RNA CCDC66 Expression and Development of Chemoresistance. Cancers 53 32187976
1992 In vitro and in vivo biotransformation and covalent binding of benzo(a)pyrene in Gunn and RHA rats with a genetic deficiency in bilirubin uridine diphosphate-glucuronosyltransferase. The Journal of pharmacology and experimental therapeutics 53 1403794
2021 TDRD3 promotes DHX9 chromatin recruitment and R-loop resolution. Nucleic acids research 52 34329467
2016 Tumor cell survival dependence on the DHX9 DExH-box helicase. Oncogene 51 26973242
2018 Bidirectional regulation of adenosine-to-inosine (A-to-I) RNA editing by DEAH box helicase 9 (DHX9) in cancer. Nucleic acids research 49 29796672
2019 The Host DHX9 DExH-Box Helicase Is Recruited to Chikungunya Virus Replication Complexes for Optimal Genomic RNA Translation. Journal of virology 48 30463980
2016 In Vivo Selection Against Human Colorectal Cancer Xenografts Identifies an Aptamer That Targets RNA Helicase Protein DHX9. Molecular therapy. Nucleic acids 47 27115840
2010 WRN helicase unwinds Okazaki fragment-like hybrids in a reaction stimulated by the human DHX9 helicase. Nucleic acids research 46 20385589
2017 The RNA helicase DHX9 establishes nucleolar heterochromatin, and this activity is required for embryonic stem cell differentiation. EMBO reports 45 28588071
2013 Structural insights into RISC assembly facilitated by dsRNA-binding domains of human RNA helicase A (DHX9). Nucleic acids research 45 23361462
2020 The enigmatic helicase DHX9 and its association with the hallmarks of cancer. Future science OA 44 33437516
2021 DHX9 contributes to the malignant phenotypes of colorectal cancer via activating NF-κB signaling pathway. Cellular and molecular life sciences : CMLS 43 34773477
2018 A DNA-sensing-independent role of a nuclear RNA helicase, DHX9, in stimulation of NF-κB-mediated innate immunity against DNA virus infection. Nucleic acids research 42 30137501
2008 Diverse sensitivity of RHA/Verh and RLA/Verh rats to emotional and spatial aspects of a novel environment as a result of a distinct pattern of neuronal activation in the fear/anxiety circuit. Behavior genetics 42 18972198
2015 Genotoxic stress inhibits Ewing sarcoma cell growth by modulating alternative pre-mRNA processing of the RNA helicase DHX9. Oncotarget 41 26450900
2014 Suppression of the DHX9 helicase induces premature senescence in human diploid fibroblasts in a p53-dependent manner. The Journal of biological chemistry 40 24990949
2017 Structure of the DEAH/RHA ATPase Prp43p bound to RNA implicates a pair of hairpins and motif Va in translocation along RNA. RNA (New York, N.Y.) 39 28416566
2014 Redistribution of demethylated RNA helicase A during foot-and-mouth disease virus infection: role of Jumonji C-domain containing protein 6 in RHA demethylation. Virology 39 24606677
2009 Genetic sequence variations of BRCA1-interacting genes AURKA, BAP1, BARD1 and DHX9 in French Canadian families with high risk of breast cancer. Journal of human genetics 39 19197335
2016 FXIIa inhibitor rHA-Infestin-4: Safe thromboprotection in experimental venous, arterial and foreign surface-induced thrombosis. British journal of haematology 38 27018425
2024 DHX9 maintains epithelial homeostasis by restraining R-loop-mediated genomic instability in intestinal stem cells. Nature communications 37 38594251
2018 DHX9 regulates production of hepatitis B virus-derived circular RNA and viral protein levels. Oncotarget 37 29765512
2017 RNA helicase DHX9 may be a therapeutic target in lung cancer and inhibited by enoxacin. American journal of translational research 37 28337295
2010 Crystal structure of human RNA helicase A (DHX9): structural basis for unselective nucleotide base binding in a DEAD-box variant protein. Journal of molecular biology 34 20510246
2017 Human Nup98 regulates the localization and activity of DExH/D-box helicase DHX9. eLife 33 28221134
2024 ATR phosphorylates DHX9 at serine 321 to suppress R-loop accumulation upon genotoxic stress. Nucleic acids research 32 37930853
2020 USP42 enhances homologous recombination repair by promoting R-loop resolution with a DNA-RNA helicase DHX9. Oncogenesis 32 32541651
2024 Induction of Viral Mimicry Upon Loss of DHX9 and ADAR1 in Breast Cancer Cells. Cancer research communications 30 38530197
2022 RNA-binding protein DHX9 promotes glioma growth and tumor-associated macrophages infiltration via TCF12. CNS neuroscience & therapeutics 30 36377508
2023 Monoallelic variation in DHX9, the gene encoding the DExH-box helicase DHX9, underlies neurodevelopment disorders and Charcot-Marie-Tooth disease. American journal of human genetics 29 37467750
2016 DHX9/RHA Binding to the PBS-Segment of the Genomic RNA during HIV-1 Assembly Bolsters Virion Infectivity. Journal of molecular biology 29 27107641
2013 RNAi screening uncovers Dhx9 as a modifier of ABT-737 resistance in an Eμ-myc/Bcl-2 mouse model. Blood 29 23440244
2023 Nucleic DHX9 cooperates with STAT1 to transcribe interferon-stimulated genes. Science advances 28 36735791
1990 Transcription from the rha operon psr promoter. Journal of molecular biology 28 1688950
2024 AKT1 interacts with DHX9 to Mitigate R Loop-Induced Replication Stress in Ovarian Cancer. Cancer research 26 38241710
2017 Functional link between DEAH/RHA helicase Prp43 activation and ATP base binding. Nucleic acids research 26 28180308
2020 Poison-Exon Inclusion in DHX9 Reduces Its Expression and Sensitizes Ewing Sarcoma Cells to Chemotherapeutic Treatment. Cells 25 32023846
2019 Virion-associated, host-derived DHX9/RNA helicase A enhances the processivity of HIV-1 reverse transcriptase on genomic RNA. The Journal of biological chemistry 25 31175158
2019 DHX9 inhibits epithelial-mesenchymal transition in human lung adenocarcinoma cells by regulating STAT3. American journal of translational research 24 31497206
2017 The crystal structure of human DEAH-box RNA helicase 15 reveals a domain organization of the mammalian DEAH/RHA family. Acta crystallographica. Section F, Structural biology communications 24 28580923
2006 Reduced ethanol response in the alcohol-preferring RHA rats and neuropeptide mRNAs in relevant structures. The European journal of neuroscience 24 16420460
2022 Polymerase η Recruits DHX9 Helicase to Promote Replication across Guanine Quadruplex Structures. Journal of the American Chemical Society 23 35905379
1999 Studies on beta-D-Gal(f)-(1-->4)-alpha-L-Rha(p) octyl analogues as substrates for mycobacterial galactosyl transferase activity. Bioorganic & medicinal chemistry 23 10632050
2021 MARCH6 promotes Papillary Thyroid Cancer development by destabilizing DHX9. International journal of biological sciences 21 34512155
2021 Neurobehavioral and neurodevelopmental profiles of a heuristic genetic model of differential schizophrenia- and addiction-relevant features: The RHA vs. RLA rats. Neuroscience and biobehavioral reviews 21 34571119
1995 Cerebral tryptophan hydroxylase activity, and 5-HT1A receptor, 5-HT2A receptor, and 5-HT transporter binding in grouped and isolated Roman RHA and RLA rats: relationships with behaviours in two models of anxiety. Psychopharmacology 21 8584622
1990 Cloning and expression in Escherichia coli of LKP pilus genes from a nontypeable Haemophilus influenzae strain. Infection and immunity 20 1969397
2020 DHX9 interacts with APOBEC3B and attenuates the anti-HBV effect of APOBEC3B. Emerging microbes & infections 19 32056513
2019 Involvement of DHX9/YB-1 complex induced alternative splicing of Krüppel-like factor 5 mRNA in phenotypic transformation of vascular smooth muscle cells. American journal of physiology. Cell physiology 19 31116584
2023 A pan-cancer analysis of the expression and molecular mechanism of DHX9 in human cancers. Frontiers in pharmacology 18 37214432
2021 High expression of DHX9 promotes the growth and metastasis of hepatocellular carcinoma. Journal of clinical laboratory analysis 18 34676915
2019 Cellular RNA Helicase DHX9 Interacts with the Essential Epstein-Barr Virus (EBV) Protein SM and Restricts EBV Lytic Replication. Journal of virology 18 30541834
2017 Differential effects of antipsychotic and propsychotic drugs on prepulse inhibition and locomotor activity in Roman high- (RHA) and low-avoidance (RLA) rats. Psychopharmacology 18 28154892
1999 The human RNA helicase A (DDX9) gene maps to the prostate cancer susceptibility locus at chromosome band 1q25 and its pseudogene (DDX9P) to 13q22, respectively. Somatic cell and molecular genetics 18 10925702
2025 Circular RNA circCLASP2 promotes nasopharyngeal carcinoma progression through binding to DHX9 to enhance PCMT1 translation. Molecular cancer 17 40050914
2024 DHX9 SUMOylation is required for the suppression of R-loop-associated genome instability. Nature communications 17 39019926
2019 Hepatitis B virus X protein modulates upregulation of DHX9 to promote viral DNA replication. Cellular microbiology 17 31829498
2013 Genetic mapping of rust resistance genes in confection sunflower line HA-R6 and oilseed line RHA 397. TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik 17 23719761
2025 A Potent, Selective, Small-Molecule Inhibitor of DHX9 Abrogates Proliferation of Microsatellite Instable Cancers with Deficient Mismatch Repair. Cancer research 16 39589774
2024 LncRNA DYNLRB2-AS1 promotes gemcitabine resistance of nasopharyngeal carcinoma by inhibiting the ubiquitination degradation of DHX9 protein. Drug resistance updates : reviews and commentaries in antimicrobial and anticancer chemotherapy 16 38908233
2020 Function of Auxiliary Domains of the DEAH/RHA Helicase DHX36 in RNA Remodeling. Journal of molecular biology 16 32087197
2022 Regulation of the DEAH/RHA helicase Prp43 by the G-patch factor Pfa1. Proceedings of the National Academy of Sciences of the United States of America 15 36409901
2019 Molecular characterization and expression of the teleost cytosolic DNA sensor genes cGAS, LSm14A, DHX9, and DHX36 in Japanese medaka, Oryzias latipes. Developmental and comparative immunology 15 31141705
2015 Evolution of the DEAD box helicase family in chicken: chickens have no DHX9 ortholog. Microbiology and immunology 15 26382053
2024 AURKB targets DHX9 to promote hepatocellular carcinoma progression via PI3K/AKT/mTOR pathway. Molecular carcinogenesis 14 38874176
2022 The DNA/RNA helicase DHX9 contributes to the transcriptional program of the androgen receptor in prostate cancer. Journal of experimental & clinical cancer research : CR 14 35590370
2020 C1orf109L binding DHX9 promotes DNA damage depended on the R-loop accumulation and enhances camptothecin chemosensitivity. Cell proliferation 14 32761833
2020 SPOP attenuates migration and invasion of choriocarcinoma cells by promoting DHX9 degradation. American journal of cancer research 14 32905556
2019 Two DExD/H-box helicases, DDX3 and DHX9, identified in rainbow trout are able to bind dsRNA. Fish & shellfish immunology 14 31340170
2024 Cooperation between SIX1 and DHX9 transcriptionally regulates integrin-focal adhesion signaling mediated metastasis and sunitinib resistance in KIRC. Oncogene 13 39174859
2023 Phosphorylation of UHRF2 affects malignant phenotypes of HCC and HBV replication by blocking DHX9 ubiquitylation. Cell death discovery 13 36690646
2022 LncRNA PCAT19 induced by SP1 and acted as oncogene in gastric cancer competitively binding to miR429 and upregulating DHX9. Journal of Cancer 13 34976174
2021 SNORA42 promotes oesophageal squamous cell carcinoma development through triggering the DHX9/p65 axis. Genomics 13 34182081
2004 A functional interaction between RHA and Ubc9, an E2-like enzyme specific for Sumo-1. Journal of molecular biology 13 15312759
1985 A phi 80 function inhibitory for growth of lambdoid phage in him mutants of Escherichia coli deficient in integration host factor. I. Genetic analysis of the Rha phenotype. Virology 13 3155885
2023 DHRS4-AS1 regulate gastric cancer apoptosis and cell proliferation by destabilizing DHX9 and inhibited the association between DHX9 and ILF3. Cancer cell international 12 38041141
1986 Behavioral characteristics of Roman high avoidance rats homozygous for diabetes insipidus (RHA: di/di). Behavioural brain research 12 3718662

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