Affinage

IPO9

Importin-9 · UniProt Q96P70

Length
1041 aa
Mass
116.0 kDa
Annotated
2026-06-10
36 papers in source corpus 18 papers cited in narrative 18 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

IPO9 is an importin-β family nuclear transport receptor that imports a structurally diverse cargo repertoire using non-classical, combinatorial binding surfaces rather than linear NLS motifs (PMID:11823430, PMID:42182196). It recognizes cargos by wrapping around their globular folds: crystallographic and cryo-EM analyses show IPO9 engaging the H2A-H2B core and the winged-helix ETS domain of EHF through distinct interaction hotspots, with both the HEAT-repeat inner cavity and unstructured loops (H8, H18-19) contributing to selective recognition (PMID:30855230, PMID:42066049, PMID:42182196). Its validated cargos span ribosomal proteins rpS7/rpL18a (PMID:11823430), HMG-box (Sox2/SRY) and homeodomain (Arx) transcription factors (PMID:19349578, PMID:19494118), the stress-activated MAPKs JNK and p38 (imported in heterotrimeric Imp3/Imp9/MAPK complexes) (PMID:24216760), the kinase NUAK1 (PMID:31090959), monomeric actin (PMID:35278073, PMID:41478570), and the 20S proteasome via the AKIRIN2 scaffold (PMID:41639071). Beyond transport, IPO9 functions as a histone chaperone that sequesters H2A-H2B from premature contacts with DNA and H3-H4; rather than dissociating the complex, nuclear RanGTP assembles a stable RanGTP·IPO9·H2A-H2B ternary intermediate that selectively releases histone-binding surfaces at HEAT repeats 4-5 to license nucleosome assembly near chromatin (PMID:30855230, PMID:37379840). For actin, IPO9 binds the monomer barbed face with mid-nanomolar affinity in competition with cofilin and profilin, and this import underlies nuclear actin pools driving β-catenin entry, RelA/p65 regulation, and nuclear F-actin formation during ferroptosis (PMID:35278073, PMID:35563720, PMID:41478570, PMID:41450740). IPO9 additionally acts as a posttranscriptional repressor by binding the 5'UTR stem-loop of IFN-ε mRNA (PMID:23851686), and is a host dependency factor for flavivirus replication (PMID:30650657).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2002 High

    Established IPO9's founding identity as a nuclear import receptor that doubles as a cytoplasmic chaperone, answering how highly basic ribosomal proteins reach the nucleus without aggregating.

    Evidence In vitro nuclear import and aggregation-prevention assays with mouse Imp9 orthologs and rpS7/rpL18a

    PMID:11823430

    Open questions at the time
    • Structural basis of ribosomal-protein recognition not defined
    • Did not address cargo diversity beyond ribosomal proteins
  2. 2009 Medium

    Extended the cargo range to DNA-binding transcription factors, showing IPO9 recognizes the HMG box and homeodomain folds where import determinants overlap DNA-binding residues.

    Evidence In vitro import, co-IP, RNAi and domain deletion for Sox2/SRY and Arx

    PMID:19349578 PMID:19494118

    Open questions at the time
    • No structure of the importin-cargo complex
    • Redundancy with parallel receptors (Exp4, Imp-β/7) not fully resolved
  3. 2013 Medium

    Revealed a signaling role: IPO9 escorts stimulated JNK/p38 into the nucleus as part of a heterotrimeric Imp3/Imp9/MAPK complex, coupling MAPK activation to nuclear transcription factor phosphorylation.

    Evidence Co-IP, PLA, gel filtration, immunostaining and RNAi in stimulated cells

    PMID:24216760

    Open questions at the time
    • The stimulus-induced post-translational modification of Imp9 not molecularly defined
    • Division of labor between Imp3 and Imp9 at the envelope unresolved
  4. 2013 Medium

    Identified a transport-independent function: IPO9 represses IFN-ε mRNA by binding a 5'UTR stem-loop, defining a posttranscriptional RNA-regulatory activity.

    Evidence RNA affinity pulldown, overexpression/silencing and luciferase reporter assays

    PMID:23851686

    Open questions at the time
    • Mechanism linking RNA binding to translational/stability repression unknown
    • Generality across loop-forming mRNAs only partly tested
  5. 2016 High

    Began mapping the histone-recognition logic, showing IPO9 binds H3/H4 tail elements and that H3K14 acetylation reduces binding, linking histone PTMs to import competence.

    Evidence Fluorescence anisotropy/ITC binding assays with histone tail mutants

    PMID:27528606

    Open questions at the time
    • Did not yet show that the globular core, not tails, dominates H2A-H2B binding
    • Functional consequence of acetylation-modulated import not tested in cells
  6. 2019 High

    Solved the H2A-H2B complex structure and redefined IPO9 as a histone chaperone whose RanGTP response builds a remodeling intermediate rather than simple cargo release.

    Evidence X-ray crystallography, binding assays and in vitro nucleosome assembly

    PMID:30855230

    Open questions at the time
    • Dynamics of the RanGTP ternary complex not resolved at residue level
    • How chaperone hand-off to assembly factors occurs in cells unknown
  7. 2019 Medium

    Broadened cargo scope to a stress-regulated kinase and showed import is redox-sensitive, linking IPO9 transport to oxidative stress.

    Evidence Interactome MS, co-IP, RNAi, fractionation and importazole inhibition for NUAK1

    PMID:31090959

    Open questions at the time
    • The bipartite NLS recognition mode not structurally defined
    • Redundancy with IPO7 not dissected
  8. 2021 Medium

    Genetic in vivo evidence established essential developmental roles in chromosome segregation, protamine exchange and proteasome nuclear localization, connecting the receptor to fertility.

    Evidence Drosophila Ipo9 knockout with IF, FISH and co-IP

    PMID:33632744

    Open questions at the time
    • Which cargo defects drive each phenotype not separable
    • Mammalian in vivo confirmation absent in corpus
  9. 2022 Medium

    Placed IPO9-mediated nuclear actin import upstream of mechanotransduction and inflammatory signaling, showing β-catenin and NF-κB/RelA outputs depend on actin transport.

    Evidence RNAi, nuclear fractionation, imaging, NF-κB reporter and proteasome-inhibitor experiments in MSCs

    PMID:35278073 PMID:35563720

    Open questions at the time
    • Direct versus indirect coupling of actin import to β-catenin/RelA not biochemically isolated
    • Stoichiometry with cofilin in cells unknown
  10. 2023 High

    Resolved the RanGTP-driven release mechanism for histones, showing selective contact release at HEAT repeats 4-5 and affinity tuning that confines release to high-RanGTP chromatin regions.

    Evidence HDX-MS of the RanGTP·IPO9·H2A-H2B complex with binding and nucleosome assembly assays

    PMID:37379840

    Open questions at the time
    • Whether the same partial-release logic applies to other cargos untested at the time
    • In-cell visualization of the intermediate lacking
  11. 2025 High

    Quantitative reconstitution overturned the prior actin-import model, showing IPO9 binds the actin barbed face in competition with cofilin and profilin and that no IPO9-actin-RanGTP ternary complex forms.

    Evidence Competitive in vitro binding, polymerization kinetics and affinity measurements

    PMID:41478570

    Open questions at the time
    • How actin is released in the nucleus if RanGTP does not form a ternary complex is unresolved
    • Reconciliation with cell-based cofilin-cooperative models needed
  12. 2025 Medium

    Linked IPO9-dependent nuclear actin import to a specific cell-death program, showing it is required for nuclear F-actin assembly during ferroptosis.

    Evidence RNAi with phalloidin staining and nuclear actin chromobody imaging in HT-1080 cells

    PMID:41450740

    Open questions at the time
    • Functional consequence of nuclear F-actin for ferroptotic death not established
    • Single cell line, no in vivo validation
  13. 2026 High

    Structures of ETS-domain and proteasome cargo recognition demonstrated combinatorial use of distinct binding surfaces and a scaffold-based import route, generalizing IPO9's non-classical recognition principle.

    Evidence Cryo-EM, binding assays and mutagenesis for EHF; saturation mutagenesis and reconstitution of the AKIRIN2-proteasome import module

    PMID:41639071 PMID:42066049

    Open questions at the time
    • Predictive rules mapping cargo fold to IPO9 surface not yet derived
    • How RanGTP releases each structurally distinct cargo not uniformly defined
  14. 2026 Medium

    Systematic interactome and footprinting defined the full cargo cohort (~79 cargos) and showed recognition uses the inner cavity plus specific loops with no linear motif, plus cargo-specific RanGTP sensitivity.

    Evidence Cytoplasmic IP-MS, oxidative footprinting and loop-perturbation IP-MS (preprint)

    PMID:42182196

    Open questions at the time
    • Preprint not yet peer reviewed
    • Functional import of most newly identified cargos not validated

Open questions

Synthesis pass · forward-looking unresolved questions
  • A unifying code that predicts which globular folds IPO9 recognizes and how RanGTP release affinity is tuned across structurally unrelated cargos remains undefined.
  • No general structural rule linking cargo fold to IPO9 binding surface
  • Mechanism of nuclear release for non-histone cargos lacking a stable ternary intermediate unresolved

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140104 molecular carrier activity 7 GO:0044183 protein folding chaperone 3 GO:0008092 cytoskeletal protein binding 2 GO:0042393 histone binding 2 GO:0003723 RNA binding 1 GO:0060090 molecular adaptor activity 1
Localization
GO:0005634 nucleus 3 GO:0005829 cytosol 2 GO:0005635 nuclear envelope 1
Pathway
R-HSA-9609507 Protein localization 4 R-HSA-162582 Signal Transduction 3 R-HSA-4839726 Chromatin organization 3 R-HSA-74160 Gene expression (Transcription) 3 R-HSA-1643685 Disease 1 R-HSA-5357801 Programmed Cell Death 1
Partners
ACTBAKIRIN2CFL1EHFH2A-H2BIPO7NUAK1RAN
Complex memberships
AKIRIN2-proteasome import moduleImp3/Imp9/MAPK heterotrimerRanGTP·IPO9·H2A-H2B ternary complex

Evidence

Reading pass · 18 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2002 Imp9a and Imp9b (mouse orthologs of human IPO9) mediate nuclear import of ribosomal proteins rpS7 and rpL18a, and act as cytoplasmic chaperones by covering their exposed basic domains to prevent aggregation with cytoplasmic polyanions such as RNA. In vitro nuclear import assay, chaperone/aggregation prevention assays The EMBO journal High 11823430
2009 IPO9 mediates nuclear import of Sox2 and SRY transcription factors via their HMG box domain, acting in parallel with Exp4 and the Imp-beta/7 heterodimer; import signals overlap with conserved residues critical for DNA binding. In vitro nuclear import assay, co-immunoprecipitation, RNAi knockdown The Journal of cell biology Medium 19349578
2009 IPO9 mediates nuclear import of the homeodomain protein Arx via its NLS2 (within the DNA-binding homeodomain); binding to IPO9 is RanGTP-sensitive and NLS2 can be co-precipitated with IPO9. In vitro nuclear import assay, co-immunoprecipitation, RNAi knockdown, domain deletion analysis The Journal of biological chemistry Medium 19494118
2013 IPO9 participates in stimulation-induced nuclear translocation of JNK and p38 MAPKs by forming heterotrimeric complexes (Imp3/Imp9/MAPK); JNK1/2 and p38α/β bind Imp9 upon stimulated post-translational modification of Imp9; IPO9 escorts MAPKs into the nucleus while Imp3 remains at the nuclear envelope. Knockdown of IPO9 inhibits MAPK nuclear translocation and downstream transcription factor phosphorylation. Co-immunoprecipitation, proximity ligation assay, gel filtration, immunostaining, RNAi knockdown Molecular and cellular biology Medium 24216760
2013 IPO9 binds the 5'UTR stem-loop structure 1 of IFN-ε mRNA; IPO9 overexpression decreases and IPO9 silencing increases basal IFN-ε mRNA expression, defining a negative posttranscriptional regulatory role for IPO9. This effect extends to other mRNAs capable of forming similar loop structures (e.g., HIF-1α). RNA affinity pulldown (agarose-bound RNA with HeLa cell extracts), overexpression, RNAi knockdown, luciferase reporter assay Journal of immunology Medium 23851686
2016 IPO9 binds histone H3 and H4 tails at two separate elements: the segment at residues 11-27 and an isoleucine-lysine NLS (IK-NLS) motif at residues 35-40 of the H3 tail; acetylation of H3 Lys14 substantially decreases binding to IPO9 and several other importins. Quantitative binding assays (fluorescence anisotropy/ITC), mutagenic analysis of histone tail deletion mutants The Journal of biological chemistry High 27528606
2019 Crystal structure of IPO9 bound to the H2A-H2B dimer reveals that IPO9 wraps around the globular core region of H2A-H2B forming an extensive interface; the NLS-like sequences in H2A-H2B tails play a minor role. IPO9 precludes H2A-H2B interactions with DNA and H3-H4 (acting as a chaperone/sequestrant). RanGTP does not dissociate IPO9•H2A-H2B but assembles a stable RanGTP•IPO9•H2A-H2B ternary complex that can facilitate H2A-H2B dissociation by DNA and nucleosome assembly. X-ray crystallography, quantitative binding assays, nucleosome assembly assay, deletion mutagenesis eLife High 30855230
2019 IPO9 mediates nuclear import of NUAK1 (a serine/threonine AMPK-family kinase) via a bipartite NLS at the N-terminal domain; knockdown of IPO9 (or IPO7) inhibits NUAK1 nuclear import. Oxidative stress induces NUAK1 cytoplasmic accumulation, indicating that oxidative stress affects IPO9-mediated nuclear transport. Mass spectrometry (interactome), co-immunoprecipitation, RNAi knockdown, subcellular fractionation, importazole inhibition Journal of cellular biochemistry Medium 31090959
2019 IPO9 (validated by silencing) is required for optimal replication of yellow fever virus (YFV) and West Nile virus (WNV) in human cells, identifying it as a host dependency factor for flavivirus replication. Genome-wide gain-of-function cDNA screen, RNAi knockdown validation, virological assays Viruses Medium 30650657
2021 Drosophila Importin-9 (Ipo9/Ranbp9, ortholog) is required for chromosome condensation and segregation during meiosis, protamine exchange during spermatogenesis, and nuclear localization of proteasome components; Ipo9 physically interacts with proteasome proteins. Loss of Ipo9 causes female and male sterility. Genetic knockout (Ipo9KO), immunofluorescence, FISH, co-immunoprecipitation Journal of cell science Medium 33632744
2022 IPO9 (together with cofilin-1/CFL1) co-mediates nuclear transfer of G-actin; knockdown of IPO9 prevents dynamic strain-mediated nuclear transfer of both actin and β-catenin in mesenchymal stem cells, indicating that β-catenin nuclear entry depends on actin transport via IPO9. RNAi knockdown of IPO9, nuclear fractionation, fluorescence imaging, mechanical strain application Stem cells Medium 35278073
2022 Silencing IPO9 or CFL1 (components of the nuclear actin import complex) prevents cAMP-induced nuclear actin monomer increase and rescues RelA/p65 levels and NF-κB reporter activity, placing IPO9-mediated nuclear actin import upstream of proteasomal degradation of RelA/p65 in the cAMP anti-inflammatory pathway. RNAi knockdown, NF-κB reporter assay, western blotting, proteasome inhibitor experiment Cells Medium 35563720
2023 HDX-MS analysis of the RanGTP•IPO9•H2A-H2B ternary complex shows that RanGTP releases H2A-H2B contacts at IPO9 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 IPO9 when H2A-H2B is bound, ensuring release only at high nuclear RanGTP concentrations near chromatin. Hydrogen-deuterium exchange mass spectrometry (HDX-MS), quantitative binding assays, in vitro nucleosome assembly Structure High 37379840
2025 IPO9 directly binds monomeric actin with mid-nanomolar affinity; contrary to the established model, cofilin competitively inhibits (rather than promotes) IPO9-actin complex formation. Profilin similarly competes with IPO9 for actin binding at the barbed face. RanGTP binds monomeric actin but a tripartite IPO9-actin-RanGTP complex does not form. IPO9 modestly decreases the rate of actin filament assembly and exhibits minimal binding to actin filaments. In vitro binding assays (competitive), actin polymerization kinetics assay, quantitative affinity measurements The Journal of biological chemistry High 41478570
2025 IPO9 knockdown markedly reduces nuclear F-actin assembly during ferroptosis in HT-1080 cells, establishing that IPO9-dependent nuclear import of G-actin is required for nuclear F-actin formation during ferroptotic cell death. RNAi knockdown, phalloidin staining, live imaging with nuclear actin chromobody (nAC-TagGFP2) Frontiers in cell and developmental biology Medium 41450740
2026 Cryo-EM structure of IPO9 bound to the ETS domain transcription factor EHF reveals that IPO9 wraps around the winged-helix fold (ETS domain) and engages structural features throughout; the DNA-binding helix of the ETS domain is critical for importin recognition and NLS activity. IPO9 uses distinct interaction hotspots compared to its H2A-H2B binding surfaces, demonstrating combinatorial use of binding surfaces for structurally diverse cargos. ETS domains constitute a structure-encoded (globular) NLS class recognized by IPO9 with nanomolar affinity. Cryo-electron microscopy, biochemical binding assays, mutagenesis, cellular NLS activity assays Proceedings of the National Academy of Sciences of the United States of America High 42066049
2026 AKIRIN2 acts as a multivalent scaffold that simultaneously binds the 20S proteasome and IPO9 (as well as KPNA2/KPNB1), recruiting an importin cluster to mediate nuclear import of the proteasome. In the nucleus, RanGTP triggers IPO9 dissociation to release the proteasome. Identified by saturation mutagenesis screens, cryo-EM, and biochemical reconstitution. Protein-wide saturation mutagenesis, cryo-EM, biochemical reconstitution, co-immunoprecipitation Nature communications High 41639071
2026 Systematic cytoplasmic IP-MS identifies 79 bona fide IPO9-bound cargos (including H2A-H2B, TFIIB, actin, proteasome subunits, and 20 previously validated cargos); IPO9 does not use classical NLS motifs nor any linear peptide motif for cargo recognition. Oxidative footprinting shows both the inner cavity and unstructured loops (H8, H18-19) of IPO9 are protected by bound cargo. Loop perturbation IP-MS shows H8 and H18-19 loops mediate selective cargo recognition; H7, H8, H18-19 loops restrict binding of a secondary set of potential cargos. RanGTP sensitivity for cargo release varies by orders of magnitude across the cargo cohort. Cytoplasmic immunoprecipitation/mass spectrometry, oxidative protein footprinting, systematic loop-perturbation IP-MS bioRxivpreprint Medium 42182196

Source papers

Stage 0 corpus · 36 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 Importins fulfil a dual function as nuclear import receptors and cytoplasmic chaperones for exposed basic domains. The EMBO journal 260 11823430
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 49 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 44 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 25 19494118
2019 Uncovering Flavivirus Host Dependency Factors through a Genome-Wide Gain-of-Function Screen. Viruses 22 30650657
2022 Mechanically Induced Nuclear Shuttling of β-Catenin Requires Co-transfer of Actin. Stem cells (Dayton, Ohio) 20 35278073
2019 Identification of a nuclear localization signal and importin beta members mediating NUAK1 nuclear import inhibited by oxidative stress. Journal of cellular biochemistry 19 31090959
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
2022 Enhancer promoter interactome and Mendelian randomization identify network of druggable vascular genes in coronary artery disease. Human genomics 9 35246263
2023 Molecular basis of RanGTP-activated release of Histones H2A-H2B from Importin-9. Structure (London, England : 1993) 8 37379840
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 5 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 3 40149595
2024 Core biomarkers analysis benefit for diagnosis on human intrahepatic cholestasis of pregnancy. BMC pregnancy and childbirth 2 39127651
2026 RHOJ-induced chemotherapy resistance through epithelial-mesenchymal transition in drug-tolerant persister cells of head and neck cancer. Translational oncology 1 41548474
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
2025 A revised model of nuclear actin import: Importin 9 competes with cofilin, profilin, and RanGTP for actin binding. The Journal of biological chemistry 1 41478570
2026 Importins recognize the winged-helix fold of ETS transcription factors to mediate nuclear import. bioRxiv : the preprint server for biology 0 41542470
2026 A multivalent adaptor mechanism drives the nuclear import of proteasomes. Nature communications 0 41639071
2026 Importin-9 recognizes the winged-helix fold of ETS transcription factors to mediate nuclear import. Proceedings of the National Academy of Sciences of the United States of America 0 42066049
2026 Bound for the nucleus: defining the molecular principles of cargo selection by importin 9. bioRxiv : the preprint server for biology 0 42182196
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

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