Affinage

IPO9

Importin-9 · UniProt Q96P70

Round 2 corrected
Length
1041 aa
Mass
116.0 kDa
Annotated
2026-04-28
64 papers in source corpus 19 papers cited in narrative 19 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

IPO9 (Importin-9) is a versatile importin-β family nuclear transport receptor that imports structurally diverse cargoes—including ribosomal proteins, histones H2A-H2B and H3/H4, homeodomain and HMG-box transcription factors (Sox2, SRY, Arx), ETS-family transcription factors, MAPKs (JNK, p38), NUAK1 kinase, monomeric actin, and the 20S proteasome (via the AKIRIN2 scaffold)—by recognizing globular folds and basic domains rather than classical linear NLS sequences (PMID:11823430, PMID:19349578, PMID:30855230, PMID:41542470, PMID:41639071). Structural studies reveal that IPO9 wraps its HEAT-repeat solenoid around cargo globular cores using distinct combinatorial binding surfaces for each substrate, and for H2A-H2B it employs an unusual mechanism in which RanGTP forms a ternary complex that partially releases histone contacts at HEAT repeats 4–5, enabling DNA-driven nucleosome assembly rather than simple cargo ejection (PMID:30855230, PMID:37379840). IPO9 also functions as a cytoplasmic chaperone that shields the basic domains of ribosomal proteins and histones from aggregation with polyanions, directly binds monomeric actin at its barbed face independently of cofilin to mediate nuclear actin import required for mechanosensing and ferroptotic nuclear F-actin assembly, and suppresses expression of specific mRNAs (IFN-ε, HIF-1α) by binding 5′UTR stem-loop structures (PMID:11823430, PMID:41478570, PMID:35278073, PMID:23851686). Nuclear import of JNK/p38 MAPKs proceeds through stimulus-induced IPO9–importin-3 heterotrimeric complexes requiring post-translational modification of IPO9 (PMID:24216760, PMID:30946556).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2002 High

    Establishing IPO9 as a bona fide importin-β family member resolved how ribosomal proteins reach the nucleus and revealed an unexpected cytoplasmic chaperone function that prevents aggregation of basic cargo domains with polyanions.

    Evidence Reconstituted import assays and solubility/aggregation assays with recombinant ribosomal proteins rpS7 and rpL18a

    PMID:11823430

    Open questions at the time
    • Full range of ribosomal protein cargoes not mapped
    • Structural basis of chaperone function unresolved at this stage
    • Relationship between chaperone and import functions not dissected
  2. 2009 High

    Identifying homeodomain (Arx) and HMG-box (Sox2, SRY) transcription factors as IPO9 cargoes demonstrated that IPO9 recognizes DNA-binding folds rather than canonical linear NLS sequences, establishing a pattern of fold-based recognition.

    Evidence In vitro import assays, co-immunoprecipitation, siRNA knockdown, and domain deletion analysis in mammalian cells

    PMID:19349578 PMID:19494118

    Open questions at the time
    • Structural basis for HMG-box or homeodomain recognition not determined
    • Redundancy with other importins (Imp-β/7, Exp4) for these cargoes not quantified in vivo
  3. 2013 High

    Discovery that IPO9 forms stimulus-induced heterotrimeric complexes with importin-3 and JNK/p38 MAPKs revealed a non-canonical, NLS-independent import mechanism regulated by post-translational modification of the importin itself.

    Evidence Co-immunoprecipitation, proximity ligation assay, gel filtration, siRNA knockdown with transcription factor phosphorylation readout; confirmed in independent follow-up study

    PMID:24216760 PMID:30946556

    Open questions at the time
    • Identity of the post-translational modification on IPO9 not determined
    • Structural basis of the IPO9–Imp3–MAPK heterotrimer unknown
    • Whether other MAPKs use this pathway not tested
  4. 2013 Medium

    Finding that IPO9 binds 5′UTR stem-loops of IFN-ε and HIF-1α mRNAs and suppresses their expression introduced an unexpected post-transcriptional regulatory role distinct from nuclear transport.

    Evidence RNA affinity pulldown, luciferase reporter assays with UTR constructs, overexpression and siRNA knockdown

    PMID:23851686

    Open questions at the time
    • Direct RNA-binding site on IPO9 not mapped
    • Scope of mRNA targets genome-wide unknown
    • Mechanism of translational or mRNA stability regulation not defined
  5. 2016 High

    Quantitative binding studies with histone tail peptides revealed that IPO9 recognizes two separate basic segments in the H3 tail and analogous elements in H4, with acetylation of H3K14 reducing binding, linking histone modification state to import efficiency.

    Evidence Fluorescence polarization and ITC with systematic histone tail peptide mutants across seven importins

    PMID:27528606

    Open questions at the time
    • Whether tail recognition contributes to import in vivo vs. chaperone shielding not separated
    • Interplay with H2A-H2B import not addressed
  6. 2016 Medium

    Demonstrating that IPO9 knockdown reduces nuclear/perinuclear F-actin and that cofilin alone cannot transport actin into the nucleus established IPO9 as the essential receptor for nuclear actin import.

    Evidence siRNA knockdown, F-actin fluorescence quantification, western blotting in MCF-7 cells

    PMID:26934847

    Open questions at the time
    • Direct IPO9–actin binding not demonstrated at this stage
    • Mechanism of actin recognition by IPO9 undefined
  7. 2019 High

    The crystal structure of IPO9 bound to H2A-H2B revealed an extensive interface wrapping around the histone globular core and uncovered a unique cargo-release mechanism: RanGTP forms a stable ternary complex rather than ejecting cargo, allowing DNA-driven histone transfer for nucleosome assembly.

    Evidence X-ray crystallography, ITC, fluorescence binding assays, deletion mutagenesis, DNA competition, in vitro nucleosome assembly

    PMID:30855230

    Open questions at the time
    • In vivo contribution of IPO9 vs. other H2A-H2B chaperones/importins not quantified
    • Whether ternary complex mechanism applies to other cargoes unknown
  8. 2023 High

    HDX-MS mapping of the RanGTP•Imp9•H2A-H2B ternary complex resolved how RanGTP selectively releases H2A-H2B contacts at HEAT repeats 4–5 while maintaining contacts at 18–19, explaining the partial release that licenses DNA-mediated nucleosome assembly at chromatin-proximal high-RanGTP concentrations.

    Evidence Hydrogen-deuterium exchange mass spectrometry comparing binary and ternary complexes, quantitative binding assays, nucleosome assembly assay

    PMID:37379840

    Open questions at the time
    • Full kinetic pathway from ternary complex to assembled nucleosome not captured
    • Whether chromatin remodelers cooperate with the ternary complex in vivo untested
  9. 2025 High

    Reconstituted binding studies overturned the classical model by showing that IPO9 directly binds monomeric actin at the barbed face with mid-nanomolar affinity independently of cofilin, which instead competitively inhibits IPO9–actin binding, redefining the actin import complex.

    Evidence In vitro competitive binding assays, fluorescence polarization, actin polymerization kinetics

    PMID:41478570

    Open questions at the time
    • In vivo stoichiometry and regulation of IPO9–actin vs. cofilin–actin pools not determined
    • How RanGTP releases actin in the nucleus (no tripartite complex forms) remains mechanistically unclear
  10. 2025 Medium

    IPO9-mediated nuclear actin import was shown to be functionally required for nuclear F-actin assembly during ferroptosis and for cAMP-induced proteasomal degradation of RelA/p65, establishing physiological contexts where nuclear actin import has defined downstream consequences.

    Evidence siRNA knockdown with live 3D imaging of nuclear actin chromobody during ferroptosis; siRNA knockdown with NF-κB reporter assay and ubiquitin pulldown for cAMP signaling

    PMID:35563720 PMID:41450740

    Open questions at the time
    • Molecular mechanism linking nuclear actin to RelA degradation not fully elucidated
    • Whether IPO9-dependent nuclear actin import is rate-limiting in these processes in vivo uncertain
  11. 2026 High

    Cryo-EM of IPO9 bound to the ETS domain of EHF and structural comparison with the H2A-H2B complex revealed that IPO9 uses distinct combinatorial HEAT-repeat surfaces to recognize structurally unrelated globular folds, explaining its cargo versatility; separately, cryo-EM and saturation mutagenesis showed that AKIRIN2 scaffolds an importin cluster (including IPO9) around the 20S proteasome for nuclear import.

    Evidence Cryo-EM structures, saturation mutagenesis screens, biochemical reconstitution, mammalian cell NLS activity assays

    PMID:41542470 PMID:41639071

    Open questions at the time
    • ETS-domain cryo-EM is from a preprint and awaits peer review
    • Whether IPO9 contacts the proteasome directly or solely via AKIRIN2 not fully resolved
    • Quantitative contribution of IPO9 vs. KPNA2/KPNB1 in proteasome import not determined

Open questions

Synthesis pass · forward-looking unresolved questions
  • How IPO9 selects among its many structurally diverse cargoes in the crowded cytoplasmic milieu, and whether post-translational modifications or adaptor proteins regulate cargo priority, remains unresolved.
  • No systematic in vivo cargo prioritization study exists
  • Post-translational modification of IPO9 identified for MAPK import but identity unknown
  • Full spectrum of IPO9 cargoes likely incomplete

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005215 transporter activity 9 GO:0008092 cytoskeletal protein binding 3 GO:0042393 histone binding 3 GO:0044183 protein folding chaperone 2 GO:0003723 RNA binding 1
Localization
GO:0005634 nucleus 5 GO:0005829 cytosol 3
Pathway
R-HSA-9609507 Protein localization 9 R-HSA-162582 Signal Transduction 3 R-HSA-4839726 Chromatin organization 3
Partners
ACTBAKIRIN2CFL1H2AFZH2BC1IPO3NUAK1RAN
Complex memberships
AKIRIN2-IPO9-proteasome import complexIPO9-Imp3-MAPK heterotrimerRanGTP-Imp9-H2A-H2B ternary complex

Evidence

Reading pass · 19 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 IPO9 (Imp9a and Imp9b) was identified as a novel importin beta-family member that mediates nuclear import of ribosomal proteins rpS7 and rpL18a, and functions as a cytoplasmic chaperone by covering the basic domains of these substrates to prevent their aggregation with cytoplasmic polyanions such as RNA. Import assays, co-precipitation, solubility/aggregation assays with recombinant proteins The EMBO journal High 11823430
2009 IPO9 (Imp9) mediates nuclear import of Sox2 and SRY via their HMG box domain, acting in parallel with Exp4 and the Imp-beta/7 heterodimer; import signals overlap with conserved HMG box residues critical for DNA binding. Co-immunoprecipitation, in vitro nuclear import assay, RanGTP sensitivity assay The Journal of cell biology High 19349578
2009 IPO9 mediates nuclear import of the homeodomain protein Arx via its NLS2 (within the DNA-binding homeodomain), with binding sensitive to RanGTP; Arx co-precipitates with importin 9 in an NLS2-dependent manner. In vitro nuclear import assay, co-immunoprecipitation, siRNA knockdown, domain deletion analysis The Journal of biological chemistry High 19494118
2013 IPO9 forms stimulation-induced heterotrimers with Imp3 and JNK1/2 or p38α/β to mediate Ran-dependent, NLS-independent nuclear translocation of JNK and p38 MAPKs; Imp9 undergoes stimulated post-translational modification that enables MAPK binding, and escorts MAPKs into the nucleus while Imp3 remains at the nuclear envelope. Co-immunoprecipitation, proximity ligation assay, gel filtration, immunostaining, siRNA knockdown with transcription factor phosphorylation readout Molecular and cellular biology High 24216760
2013 IPO9 associates with stem-loop structures in the 5'UTR of IFN-ε mRNA (specifically loop 1) and acts as a negative posttranscriptional regulator of IFN-ε expression; IPO9 overexpression decreased and IPO9 silencing increased basal IFN-ε mRNA levels. This regulatory role extends to additional mRNAs containing specific loop structures, including HIF-1α. RNA affinity pulldown from cell extracts, luciferase reporter assays with UTR constructs, overexpression and siRNA knockdown Journal of immunology Medium 23851686
2016 IPO9 binds two separate elements in the H3 tail (residues 11-27 and an IK-NLS motif at residues 35-40) and similarly binds two basic segments of the H4 tail; acetylation of H3 Lys14 substantially decreases binding to IPO9 and several other importins. Quantitative binding assays (fluorescence polarization/ITC) with histone tail peptides and deletion/point mutants The Journal of biological chemistry High 27528606
2019 Crystal structure of Importin-9 bound to histones H2A-H2B reveals that IPO9 wraps around the globular core of H2A-H2B through an extensive interface, sequesters H2A-H2B from DNA and H3-H4 interactions (acting as a storage chaperone), and that RanGTP does not dissociate the complex but instead forms a stable RanGTP•Imp9•H2A-H2B ternary complex in which H2A-H2B can be released by DNA to assemble into a nucleosome. X-ray crystallography, quantitative binding assays (ITC/fluorescence), deletion mutagenesis, DNA competition assay, in vitro nucleosome assembly eLife High 30855230
2019 IPO9 mediates nuclear import of NUAK1 kinase; NUAK1 interacts with IPO9 as identified by mass spectrometry, and knockdown of IPO9 inhibits NUAK1 nuclear import. Oxidative stress induces NUAK1 cytoplasmic accumulation, indicating that this import pathway is stress-regulated. Mass spectrometry interactome, co-immunoprecipitation confirmation, siRNA knockdown with subcellular localization readout, importazole inhibition Journal of cellular biochemistry Medium 31090959
2019 IPO9 is required for optimal flavivirus (YFV, WNV) replication in human cells, as validated by siRNA silencing approaches in a genome-wide gain-of-function screen. Genome-wide cDNA gain-of-function screen, siRNA silencing validation Viruses Low 30650657
2019 IPO9 forms stimulation-induced heterotrimers with Imp3 and JNK/p38 MAPKs; binding of JNK1/2 and p38α/β to Imp7 or Imp9 requires stimulated post-translational modification of the importins, confirmed by coimmunoprecipitation and proximity ligation assay. Coimmunoprecipitation, proximity ligation assay, gel filtration, immunostaining, knockdown Cellular physiology and biochemistry High 30946556
2021 In Drosophila, loss of Importin-9 (Ipo9/Ranbp9) causes female and male sterility with chromosome condensation and segregation defects during meiosis, abnormal sperm structure, failure to exchange histones for protamines in males, and disruption of nuclear localization of proteasome components; Ipo9 physically interacts with proteasome proteins. Genetic knockout, immunofluorescence, FISH, co-immunoprecipitation Journal of cell science High 33632744
2022 IPO9, together with cofilin-1 (CFL1), co-mediates nuclear transfer of G-actin in response to dynamic mechanical strain in mesenchymal stem cells; knockdown of IPO9 prevented dynamic strain-mediated nuclear transfer of both actin and β-catenin, indicating that β-catenin nuclear access depends on the actin transport pathway mediated by IPO9. siRNA knockdown, live-cell imaging, subcellular fractionation, dynamic vs. static strain comparison Stem cells Medium 35278073
2022 Silencing components of the nuclear actin import complex IPO9 and CFL1 prevented the cAMP-induced increase in nuclear actin monomer and rescued RelA/p65 levels and NF-κB reporter gene activity, demonstrating that IPO9-mediated nuclear actin import is required for cAMP-induced proteasomal degradation of RelA/p65. siRNA knockdown, NF-κB reporter assay, western blotting, ubiquitin affinity pulldown Cells Medium 35563720
2023 HDX-MS analysis of the RanGTP•Imp9•H2A-H2B ternary complex shows that RanGTP binding releases H2A-H2B contacts at Imp9 HEAT repeats 4-5 but not 18-19, exposing DNA- and histone-binding surfaces of H2A-H2B to facilitate nucleosome assembly; RanGTP has weaker affinity for Imp9 when H2A-H2B is bound, ensuring release only at high RanGTP concentrations near chromatin. Hydrogen-deuterium exchange mass spectrometry (HDX-MS), quantitative binding assays, in vitro nucleosome assembly Structure High 37379840
2016 Downregulation of IPO9 in MCF-7 breast cancer cells reduces F-actin content in the nuclear/perinuclear area and is correlated with increased post-translational expression of cofilin-1 (CFL1), and that CFL1 alone does not transport actin into the nucleus but requires functional IPO9 expression for this transport. siRNA knockdown, western blotting, F-actin fluorescence quantification, flow cytometry for apoptosis Oncology reports Medium 26934847
2025 IPO9 directly binds monomeric actin with mid-nanomolar affinity independently of cofilin; cofilin and profilin competitively inhibit IPO9-actin binding (likely via overlapping barbed-face interaction), IPO9 modestly decreases actin filament assembly rate, and RanGTP binds monomeric actin but a tripartite IPO9-actin-RanGTP complex does not form—revising the classical model in which cofilin anchors IPO9 to actin monomers. In vitro binding assays (competitive binding, fluorescence polarization), actin polymerization kinetics assay, filamentous actin binding assay The Journal of biological chemistry High 41478570
2025 IPO9 knockdown markedly reduces nuclear F-actin assembly during ferroptosis, indicating that nuclear G-actin import via IPO9 is required for the nuclear F-actin that assembles under conditions of intracellular acidification during ferroptotic cell death. siRNA knockdown, phalloidin staining, live 3D/time-lapse imaging with nuclear actin chromobody, pH-sensitive reporter Frontiers in cell and developmental biology Medium 41450740
2026 IPO9 directly recognizes the ETS domain (a winged-helix fold) of ETS family transcription factors to mediate their nuclear import; cryo-EM of the EHF:IPO9 complex shows IPO9 wrapping around the ETS domain engaging structural features throughout the fold, with the DNA-binding helix critical for importin recognition. Comparison with the IPO9•H2A-H2B structure reveals distinct interaction hotspots, demonstrating that IPO9 employs unique combinatorial binding surfaces for structurally diverse cargos. Cryo-electron microscopy, biochemical binding assays, mutagenesis, mammalian cell NLS activity assay bioRxivpreprint High 41542470
2026 AKIRIN2 acts as a multivalent scaffold that simultaneously binds the 20S proteasome and importin IPO9 (as well as KPNA2/KPNB1), coordinating assembly of an importin cluster around the proteasome for nuclear import; in the nucleus, RanGTP triggers importin dissociation releasing the proteasome, while AKIRIN2 undergoes ubiquitin-independent degradation. Protein-wide saturation mutagenesis screens, cryo-EM, biochemical reconstitution, co-immunoprecipitation Nature communications High 41639071

Source papers

Stage 0 corpus · 64 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2009 Defining the human deubiquitinating enzyme interaction landscape. Cell 1282 19615732
2015 The BioPlex Network: A Systematic Exploration of the Human Interactome. Cell 1118 26186194
2017 Architecture of the human interactome defines protein communities and disease networks. Nature 1085 28514442
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2018 VIRMA mediates preferential m6A mRNA methylation in 3'UTR and near stop codon and associates with alternative polyadenylation. Cell discovery 829 29507755
2003 Complete sequencing and characterization of 21,243 full-length human cDNAs. Nature genetics 754 14702039
2007 Large-scale mapping of human protein-protein interactions by mass spectrometry. Molecular systems biology 733 17353931
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2012 A census of human soluble protein complexes. Cell 689 22939629
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2011 Global landscape of HIV-human protein complexes. Nature 593 22190034
2006 Hsp90 cochaperone Aha1 downregulation rescues misfolding of CFTR in cystic fibrosis. Cell 517 17110338
2004 The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). Genome research 438 15489334
2015 A Dynamic Protein Interaction Landscape of the Human Centrosome-Cilium Interface. Cell 433 26638075
2022 OpenCell: Endogenous tagging for the cartography of human cellular organization. Science (New York, N.Y.) 432 35271311
2011 Defining human ERAD networks through an integrative mapping strategy. Nature cell biology 427 22119785
2005 Diversification of transcriptional modulation: large-scale identification and characterization of putative alternative promoters of human genes. Genome research 409 16344560
2015 Panorama of ancient metazoan macromolecular complexes. Nature 407 26344197
2007 Systematic analysis of the protein interaction network for the human transcription machinery reveals the identity of the 7SK capping enzyme. Molecular cell 367 17643375
2021 A proximity-dependent biotinylation map of a human cell. Nature 339 34079125
2012 A high-throughput approach for measuring temporal changes in the interactome. Nature methods 273 22863883
2002 Importins fulfil a dual function as nuclear import receptors and cytoplasmic chaperones for exposed basic domains. The EMBO journal 257 11823430
2017 Optimized fragmentation schemes and data analysis strategies for proteome-wide cross-link identification. Nature communications 221 28524877
2014 Proximity biotinylation and affinity purification are complementary approaches for the interactome mapping of chromatin-associated protein complexes. Journal of proteomics 215 25281560
2016 An organelle-specific protein landscape identifies novel diseases and molecular mechanisms. Nature communications 211 27173435
2015 ∆F508 CFTR interactome remodelling promotes rescue of cystic fibrosis. Nature 209 26618866
2021 N6-Methyladenosine on mRNA facilitates a phase-separated nuclear body that suppresses myeloid leukemic differentiation. Cancer cell 207 34048709
2020 UFMylation maintains tumour suppressor p53 stability by antagonizing its ubiquitination. Nature cell biology 168 32807901
2000 Cloning and functional analysis of cDNAs with open reading frames for 300 previously undefined genes expressed in CD34+ hematopoietic stem/progenitor cells. Genome research 161 11042152
2019 A protein-interaction network of interferon-stimulated genes extends the innate immune system landscape. Nature immunology 159 30833792
2009 Exportin 4 mediates a novel nuclear import pathway for Sox family transcription factors. The Journal of cell biology 71 19349578
2019 Importin-9 wraps around the H2A-H2B core to act as nuclear importer and histone chaperone. eLife 45 30855230
2013 OprD mutations and inactivation in imipenem-resistant Pseudomonas aeruginosa isolates from China. Infection, genetics and evolution : journal of molecular epidemiology and evolutionary genetics in infectious diseases 43 24211415
2006 bla(IMP-9) and its association with large plasmids carried by Pseudomonas aeruginosa isolates from the People's Republic of China. Antimicrobial agents and chemotherapy 42 16377710
2007 Characterization of the 12q amplicons by high-resolution, oligonucleotide array CGH and expression analyses of a novel liposarcoma cell line. Cancer letters 41 18160213
2021 An IncP-2 plasmid sublineage associated with dissemination of blaIMP-45 among carbapenem-resistant Pseudomonas aeruginosa. Emerging microbes & infections 36 33620296
2018 Differential Proteomic Analysis between Small Cell Lung Carcinoma (SCLC) and Pulmonary Carcinoid Tumors Reveals Molecular Signatures for Malignancy in Lung Cancer. Proteomics. Clinical applications 35 29888431
2017 Rapid and simple identification of carbapenemase genes, bla NDM, bla OXA-48, bla VIM, bla IMP-14 and bla KPC groups, in Gram-negative bacilli by in-house loop-mediated isothermal amplification with hydroxynaphthol blue dye. World journal of microbiology & biotechnology 35 28585170
2016 Recognition Elements in the Histone H3 and H4 Tails for Seven Different Importins. The Journal of biological chemistry 34 27528606
2013 Beta-like importins mediate the nuclear translocation of mitogen-activated protein kinases. Molecular and cellular biology 32 24216760
2009 The roles of multiple importins for nuclear import of murine aristaless-related homeobox protein. The Journal of biological chemistry 24 19494118
2019 Uncovering Flavivirus Host Dependency Factors through a Genome-Wide Gain-of-Function Screen. Viruses 21 30650657
2019 Identification of a nuclear localization signal and importin beta members mediating NUAK1 nuclear import inhibited by oxidative stress. Journal of cellular biochemistry 18 31090959
2022 Mechanically Induced Nuclear Shuttling of β-Catenin Requires Co-transfer of Actin. Stem cells (Dayton, Ohio) 17 35278073
2021 Importin-9 regulates chromosome segregation and packaging in Drosophila germ cells. Journal of cell science 16 33632744
2019 Beta-Like Importins Mediate the Nuclear Translocation of MAPKs. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 14 30946556
2013 Novel role for molecular transporter importin 9 in posttranscriptional regulation of IFN-ε expression. Journal of immunology (Baltimore, Md. : 1950) 12 23851686
2023 Molecular basis of RanGTP-activated release of Histones H2A-H2B from Importin-9. Structure (London, England : 1993) 8 37379840
2022 Enhancer promoter interactome and Mendelian randomization identify network of druggable vascular genes in coronary artery disease. Human genomics 8 35246263
2020 Integrative Genomic Analysis Predicts Regulatory Role of N 6-Methyladenosine-Associated SNPs for Adiposity. Frontiers in cell and developmental biology 8 32733881
2016 Downregulation of importin-9 protects MCF-7 cells against apoptosis induced by the combination of garlic-derived alliin and paclitaxel. Oncology reports 8 26934847
2024 PGRMC2 influences the onset of postmenopausal osteoporosis through disulfidptosis in monocytes: Evidence from experimental validation and Mendelian randomization. Heliyon 4 39263088
2023 Molecular basis of RanGTP-activated nucleosome assembly with Histones H2A-H2B bound to Importin-9. bioRxiv : the preprint server for biology 4 36747879
2022 Cyclic-AMP Increases Nuclear Actin Monomer Which Promotes Proteasomal Degradation of RelA/p65 Leading to Anti-Inflammatory Effects. Cells 4 35563720
2025 Genomic Characterisation of the Relationship and Causal Links Between Vascular Calcification, Alzheimer's Disease, and Cognitive Traits. Biomedicines 2 40149595
2024 Core biomarkers analysis benefit for diagnosis on human intrahepatic cholestasis of pregnancy. BMC pregnancy and childbirth 2 39127651
2026 IPO9 Promotes Ovarian Cancer Progression by Suppressing HMOX1-Dependent Ferroptosis. Human mutation 1 41584724
2025 pH-regulated nuclear F-actin assembly during ferroptosis. Frontiers in cell and developmental biology 1 41450740
2026 Importins recognize the winged-helix fold of ETS transcription factors to mediate nuclear import. bioRxiv : the preprint server for biology 0 41542470
2026 RHOJ-induced chemotherapy resistance through epithelial-mesenchymal transition in drug-tolerant persister cells of head and neck cancer. Translational oncology 0 41548474
2026 A multivalent adaptor mechanism drives the nuclear import of proteasomes. Nature communications 0 41639071
2025 A revised model of nuclear actin import: Importin 9 competes with cofilin, profilin, and RanGTP for actin binding. bioRxiv : the preprint server for biology 0 41040170
2025 A revised model of nuclear actin import: Importin 9 competes with cofilin, profilin, and RanGTP for actin binding. The Journal of biological chemistry 0 41478570