Affinage

NUP54

Nucleoporin p54 · UniProt Q7Z3B4

Length
507 aa
Mass
55.4 kDa
Annotated
2026-06-10
32 papers in source corpus 16 papers cited in narrative 16 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

NUP54 is a central FG-nucleoporin of the nuclear pore complex (NPC) that builds the diffusion barrier and transport conduit of the central channel and contributes to nuclear envelope dynamics during mitosis (PMID:26046439, PMID:29065307). It is the central subunit of the Nup62 channel complex, assembling in 1:1:1 stoichiometry by directly binding both Nup62 and Nup58 through conserved coiled-coil segments, an architecture conserved in the yeast Nsp1 complex (PMID:24574455). Within this assembly, the ordered Nup54·Nup58 and Nup54·Nup62 interactomes form dynamic, interconvertible hetero- and homo-oligomeric rings and triple-helix "fingers," and multivalent engagement of the transport factor Kapβ1 shifts these equilibria to provide a mechanism for central-channel constriction and dilation (PMID:26025361, PMID:26046439); consistent with this, these inner-ring nucleoporins undergo transport-state-dependent conformational changes in live cells (PMID:33346731). The disordered FG-repeat region of Nup54 forms labile cross-β polymers required for the import barrier, and Nup54 is specifically required for importin α/β-dependent nuclear import (PMID:28069952, PMID:17682050). These FG polymers are pathologically locked by C9orf72-derived PRn poly-dipeptides, inhibiting nucleocytoplasmic transport, while restoring Nup54 rescues import defects caused by C9orf72 poly-GA aggregates (PMID:28069952, PMID:28040728). Beyond constitutive transport, Nup54 recruits PLK-1 to the NPC via Polo-docking sites to drive efficient nuclear envelope breakdown, and the complex is remodeled at mitotic entry by TIP60-mediated Nup62 acetylation to enable correct spindle orientation and chromosome segregation (PMID:29065307, PMID:36190325). Nup54 also functions in homologous-recombination DNA repair epistatically with Rad51 (PMID:29986057), supports locus-specific piRNA biogenesis (PMID:33856346), and promotes nuclear import of the methyltransferase CARM1 in gastric cancer (PMID:34725461). Biallelic variants clustered in the NUP62-interaction region cause early-onset dystonia with striatal lesions, establishing that the NUP54–NUP62 interaction is essential for NPC function in the basal ganglia (PMID:36333996).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 2007 Medium

    Established that Nup54 is not a generic channel filler but is selectively required for one directionality of transport, distinguishing its role in import from export.

    Evidence RNAi knockdown in Drosophila S2 cells with import/export reporter assays

    PMID:17682050

    Open questions at the time
    • Does not define which FG segment mediates importin α/β selectivity
    • Drosophila ortholog; human pathway specificity not directly tested
  2. 2011 Low

    Placed NUP54 among host factors supporting influenza polymerase activity, hinting that pathogens co-opt its transport function.

    Evidence Yeast two-hybrid screen for polymerase interactors plus siRNA replication validation

    PMID:21994455

    Open questions at the time
    • Single Y2H screen plus knockdown; no direct binding or mechanism for NUP54 specifically
    • Does not distinguish a specific interaction from a general transport requirement
  3. 2014 High

    Defined the stoichiometric architecture of the channel complex, showing Nup54 is the central organizing subunit bridging Nup62 and Nup58.

    Evidence In vitro reconstitution with gel filtration and analytical ultracentrifugation; conservation to yeast Nsp1

    PMID:24574455

    Open questions at the time
    • Stoichiometry shown in solution, not within the assembled NPC
    • Does not address FG-region behavior
  4. 2015 High

    Resolved how the ordered and disordered segments couple, providing a molecular mechanism for channel dilation/constriction via Kapβ1-driven oligomer equilibria.

    Evidence Crystallography, AUC, SAXS and equilibrium binding analysis of reconstituted sub-complexes

    PMID:26025361 PMID:26046439

    Open questions at the time
    • Allosteric model derived from in vitro equilibria, not direct NPC measurement
    • Does not capture FG-polymer contribution to gating
  5. 2017 High

    Showed that the Nup54 FG region forms labile cross-β polymers that constitute the diffusion barrier and become a target locked by C9orf72 PRn poly-dipeptides.

    Evidence Chemical footprinting, polymerization-blocking mutagenesis, and 1,6-hexanediol melting with functional transport readout

    PMID:28069952

    Open questions at the time
    • Polymer state characterized in vitro; in-NPC polymer geometry unknown
    • Stoichiometry of PRn binding to FG repeats not quantified
  6. 2017 High

    Extended Nup54 function beyond steady-state transport by showing it recruits PLK-1 to the NPC to drive nuclear envelope breakdown at mitosis.

    Evidence PLK-1 PBD pulldowns, phospho-site mutagenesis, RNAi loss-of-function with NEBD live imaging in C. elegans

    PMID:29065307

    Open questions at the time
    • Direct Nup54-PLK1 contact vs complex-level recruitment not fully separated
    • Human NEBD relevance shown but mechanism mapped chiefly in C. elegans
  7. 2017 Medium

    Linked Nup54 abundance to rescue of import defects in C9orf72 disease models, placing it as a downstream effector in the importin α/β pathway.

    Evidence Overexpression rescue of a TDP-43 NLS reporter against poly-GA aggregates in cell culture

    PMID:28040728

    Open questions at the time
    • Rescue may reflect general transport buffering rather than specific repair of NUP54 function
    • Limited mechanistic depth
  8. 2018 High

    Revealed a non-transport role for Nup54 in homologous recombination, with loss causing radiosensitivity and mitotic catastrophe in replicated cells.

    Evidence siRNA knockdown, HR reporters, SCE and chromosome aberration analysis, Rad51 epistasis, IR survival

    PMID:29986057

    Open questions at the time
    • Whether Nup54 acts at the pore or at repair sites is unresolved
    • Direct repair-factor contacts not identified
  9. 2020 Medium

    Demonstrated in living cells that inner-ring Nup54 physically changes conformation in response to transport state, validating the gating model.

    Evidence FRET conformational sensors with transport perturbation in live cells

    PMID:33346731

    Open questions at the time
    • Conformational change not mapped to specific structural transitions
    • Causal link to dilation/constriction inferred
  10. 2021 Medium

    Uncovered specialized, non-canonical roles: locus-specific piRNA biogenesis, neuronal circuit wiring, and oncogenic CARM1 nuclear import.

    Evidence Drosophila RNAi/small-RNA-seq, forward-genetic mutant and behavioral analysis, and Co-IP/fractionation with cancer functional assays

    PMID:33856346 PMID:34666772 PMID:34725461

    Open questions at the time
    • How a transport channel achieves locus-specific piRNA selectivity is unexplained
    • CARM1 interaction rests on single-lab Co-IP and fractionation
  11. 2022 Medium

    Showed the channel complex is actively remodeled at mitosis by TIP60 acetylation of Nup62, coupling Nup54 complex disassembly to spindle orientation and chromosome segregation.

    Evidence Acetyltransferase assay, Lys432 mutagenesis, Co-IP of complex dissociation, live imaging of spindle/segregation

    PMID:36190325

    Open questions at the time
    • Direct functional consequence for Nup54 vs Nup62 redistribution not separated
    • Whether Nup54 itself is post-translationally regulated unknown
  12. 2022 Medium

    Established human disease causation, tying biallelic NUP54 variants in the NUP62-interaction domain to early-onset dystonia with striatal lesions.

    Evidence Patient sequencing with in silico and protein-biochemical validation of variant effects on the NUP62 interface

    PMID:36333996

    Open questions at the time
    • Tissue-specific basis for basal ganglia vulnerability unknown
    • Single study; functional disease model not yet established

Open questions

Synthesis pass · forward-looking unresolved questions
  • How Nup54's structured channel architecture, its FG-polymer barrier, and its diverse moonlighting roles (HR repair, piRNA biogenesis, mitotic signaling) are mechanistically integrated within the assembled NPC remains unresolved.
  • No structure of the intact channel including FG polymers in situ
  • No unifying model connecting transport-independent roles to NPC architecture

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005198 structural molecule activity 3 GO:0008092 cytoskeletal protein binding 3 GO:0060089 molecular transducer activity 3
Localization
GO:0005635 nuclear envelope 3
Pathway
R-HSA-9609507 Protein localization 3 R-HSA-1640170 Cell Cycle 2 R-HSA-73894 DNA Repair 1 R-HSA-8953854 Metabolism of RNA 1
Complex memberships
Nup62-Nup54-Nup58 channel complex

Evidence

Reading pass · 16 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2017 The FG repeat domain of Nup54 forms labile cross-β polymers (characterized by chemical footprinting), and mutations within the footprinted region block both polymerization and binding by the toxic PRn poly-dipeptide encoded by C9orf72 repeat expansion. PRn binding locks FG repeats in the polymerized state, inhibiting nuclear import/export. Chemical footprinting of FG domain polymers, site-directed mutagenesis blocking polymerization, in vitro 1,6-hexanediol treatment to melt polymers Proceedings of the National Academy of Sciences of the United States of America High 28069952
2017 In C. elegans, channel nucleoporins NPP-4/Nup54 (along with NPP-1/Nup58 and NPP-11/Nup62) recruit PLK-1 to nuclear pore complexes prior to nuclear envelope breakdown (NEBD) via physical interaction between multiple Cdk1/PLK-1-primed Polo-docking sites and the PLK-1 Polo-box domain (PBD). This localization is required for efficient NEBD. Co-IP/pulldown of PLK-1 PBD with nucleoporins, genetic loss-of-function (RNAi) with NEBD phenotype, phosphorylation-site mutagenesis, live imaging Developmental cell High 29065307
2015 Nup54 is the central subunit of the Nup62 channel complex, directly binding both Nup62 and Nup58 via coiled-coil segments. The structured domain of Nup58 allosterically couples with its neighboring disordered domain in interaction with Nup54's disordered domain and transport factor Kapβ1; multivalent Kapβ1 interactions with disordered domains of Nup58 stabilize the structured domain associated with Nup54, shifting equilibria from homo-oligomers to hetero-oligomers, providing a mechanism for NPC central channel constriction/dilation. Crystallography, analytical ultracentrifugation, equilibrium binding analysis, in vitro reconstitution of complexes Cell High 26046439
2014 The Nup62 complex (Nup62, Nup54, Nup58) has a 1:1:1 stoichiometry in solution, with Nup54 central and directly binding both Nup62 and Nup58 via conserved coiled-coil segments. This stoichiometry is conserved in the homologous yeast Nsp1 complex. Gel filtration chromatography, analytical ultracentrifugation, in vitro reconstitution Molecular biology of the cell High 24574455
2015 The ordered regions of Nup62, Nup54, and Nup58 form a dynamic 'triple complex' in solution built from characterized Nup54·Nup58 and Nup54·Nup62 interactomes. Nup54·Nup58 segments form inter-convertible hetero- and homo-oligomers proposed as 'mid-plane rings' for NPC channel dilation/constriction, while Nup54·Nup62 forms 1:2 triple-helix 'fingers' projecting from the ring. Solution analysis (SEC, AUC, SAXS), crystal structure analysis, in vitro reconstitution of sub-complexes The Journal of biological chemistry High 26025361
2007 In Drosophila, RNAi-mediated inactivation of Nup54 (fly homolog) selectively impairs importin α/β-mediated nuclear import but not CRM1-dependent export, identifying Nup54 FG nucleoporin as specifically required for the import pathway. RNAi knockdown in Drosophila S2 cells, nuclear import/export reporter assays The Journal of cell biology Medium 17682050
2018 Nup54 (along with its complex partners Nup62 and Nup58) is required for homologous recombination (HR) repair. Nup54 depletion decreased HR repair activity (measured by specific reporters), reduced HR-linked DNA synthesis foci and sister chromatid exchanges after ionizing radiation, and increased chromosome aberrations. Nup54 is epistatic with HR factor Rad51, and its loss caused mitotic catastrophe and enhanced radiosensitivity specifically in cells with replicated DNA. siRNA knockdown, HR reporter assays, FACS cell cycle analysis, chromosome aberration analysis, epistasis analysis with Rad51, ionizing radiation survival assays Nucleic acids research High 29986057
2021 In Drosophila ovarian follicle cells, Nup54 (along with Nup58) is specifically required for piRNA biogenesis from the flamenco locus, with loss of Nup54/Nup58 causing compromised piRNA production exclusively from this locus while knockdown of other NPC subunits has more widespread consequences, indicating a specialized role beyond general nuclear transport. RNAi knockdown in Drosophila ovary, small RNA sequencing, piRNA pathway reporter assays eLife Medium 33856346
2021 In Drosophila, Nup54 loss-of-function mutations cause defective wiring of eight adult brain pickpocket-expressing neurons required for egg-laying, and reduce sensitivity to sex-peptide, indicating a role for Nup54 in neuronal circuit differentiation underlying the female post-mating response. Nup54 promoter variants alter nucleo-cytoplasmic shuttling. Forward genetic screen, viable mutant allele characterization, neuroanatomical analysis (brain neuron wiring), behavioral assays (egg-laying, sex-peptide sensitivity) BMC biology Medium 34666772
2021 Nup54 promotes nuclear importation of the arginine methyltransferase CARM1; Nup54 was identified as a CARM1-interacting protein, and overexpression of Nup54 facilitates CARM1 nuclear import, which in turn activates Notch2 transcription and methylation in gastric cancer cells. Co-immunoprecipitation identifying Nup54-CARM1 interaction, nuclear fractionation showing CARM1 localization changes, functional proliferation and tumor formation assays Oncogene Medium 34725461
2017 Expression of Nup54 (or Nup62) rescues nuclear import of a TDP-43 NLS reporter that is impaired by cytoplasmic poly-GA aggregates in C9orf72 ALS/FTLD, placing Nup54 in the importin-α/β-dependent nuclear import pathway as a downstream effector. Rescue/overexpression experiments in cell culture with fluorescent NLS reporter, nuclear import quantification Human molecular genetics Medium 28040728
2020 In live cells, conformational changes of Nup54 (along with Nup58 and Nup62) within the NPC inner ring are detected when transport through the NPC is perturbed, while Nups elsewhere in the NPC show no such changes, indicating that these inner ring nucleoporins are flexible and undergo transport-state-dependent conformational changes. FRET-based conformational sensors with rigidly conjugated mEGFP on NPC proteins in live cells, transport perturbation assays eLife Medium 33346731
2022 TIP60-mediated acetylation of Nup62 at Lys432 dissolves the Nup62-Nup58-Nup54 complex during mitotic entry, promoting Nup62 redistribution to the mitotic spindle. This remodeling of the nucleoporin complex is required for correct spindle orientation and accurate chromosome segregation. Identification of TIP60 as acetyltransferase by biochemical assay, site-specific mutagenesis of Lys432, co-immunoprecipitation of Nup62-Nup58-Nup54 complex disruption, live imaging, spindle orientation and chromosome segregation assays Journal of molecular cell biology Medium 36190325
2022 Biallelic (recessive) variants in NUP54 clustered in the C-terminal region that interacts with NUP62 cause early-onset dystonia with striatal lesions in humans, a phenotype similar to NUP62-related infantile striatonigral degeneration, establishing that the NUP54–NUP62 interaction is required for NPC function in the basal ganglia. Patient sequencing, in silico variant analysis, protein-biochemical studies confirming pathogenicity of variants affecting NUP62-interaction domain Annals of neurology Medium 36333996
2011 NUP54 was identified as a host factor required for influenza virus polymerase replication/transcriptional activity in human cells, placing it in the nuclear transport function supporting viral RNA polymerase activity. Yeast two-hybrid screen for influenza polymerase interactors, siRNA knockdown validation of virus replication Journal of virology Low 21994455
2023 In C. elegans, nuclear pore component NPP-1/Nup54 promotes perinuclear localization of the anti-silencing CSR-1 Argonaute and is required for piRNA-mediated gene silencing initiation, linking NPC function to piRNA pathway regulation. Sensitized piRNA reporter screen, RNAi knockdown, fluorescence microscopy of Argonaute localization Genetics Low 37210214

Source papers

Stage 0 corpus · 32 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2017 Toxic PRn poly-dipeptides encoded by the C9orf72 repeat expansion block nuclear import and export. Proceedings of the National Academy of Sciences of the United States of America 185 28069952
2017 Cytoplasmic poly-GA aggregates impair nuclear import of TDP-43 in C9orf72 ALS/FTLD. Human molecular genetics 101 28040728
2017 Channel Nucleoporins Recruit PLK-1 to Nuclear Pore Complexes to Direct Nuclear Envelope Breakdown in C. elegans. Developmental cell 72 29065307
2010 The gsdf gene locus harbors evolutionary conserved and clustered genes preferentially expressed in fish previtellogenic oocytes. Gene 66 21047546
2011 Generation and comprehensive analysis of an influenza virus polymerase cellular interaction network. Journal of virology 63 21994455
2007 Distinct functions of the Drosophila Nup153 and Nup214 FG domains in nuclear protein transport. The Journal of cell biology 51 17682050
2015 Allosteric Regulation in Gating the Central Channel of the Nuclear Pore Complex. Cell 37 26046439
2018 A Mechanically Weak Extracellular Membrane-Adjacent Domain Induces Dimerization of Protocadherin-15. Biophysical journal 28 30527337
2017 Expansion of divergent SEA domains in cell surface proteins and nucleoporin 54. Protein science : a publication of the Protein Society 26 27977898
2021 Nup54-induced CARM1 nuclear importation promotes gastric cancer cell proliferation and tumorigenesis through transcriptional activation and methylation of Notch2. Oncogene 23 34725461
2014 The stoichiometry of the nucleoporin 62 subcomplex of the nuclear pore in solution. Molecular biology of the cell 22 24574455
2021 Channel nuclear pore complex subunits are required for transposon silencing in Drosophila. eLife 20 33856346
2018 Nucleoporin 54 contributes to homologous recombination repair and post-replicative DNA integrity. Nucleic acids research 18 29986057
2018 Analysis of the transcriptome data in Litopenaeus vannamei reveals the immune basis and predicts the hub regulation-genes in response to high-pH stress. PloS one 17 30517152
2022 Recessive NUP54 Variants Underlie Early-Onset Dystonia with Striatal Lesions. Annals of neurology 15 36333996
2021 Prevalence and functionality of intrinsic disorder in human FG-nucleoporins. International journal of biological macromolecules 15 33548309
2020 Conformation of the nuclear pore in living cells is modulated by transport state. eLife 15 33346731
2015 Ordered Regions of Channel Nucleoporins Nup62, Nup54, and Nup58 Form Dynamic Complexes in Solution. The Journal of biological chemistry 15 26025361
2021 Channel nuclear pore protein 54 directs sexual differentiation and neuronal wiring of female reproductive behaviors in Drosophila. BMC biology 14 34666772
2020 Integrative Analysis of DNA Methylation Identified 12 Signature Genes Specific to Metastatic ccRCC. Frontiers in oncology 12 33134164
2018 Analysis of RNA-Seq datasets reveals enrichment of tissue-specific splice variants for nuclear envelope proteins. Nucleus (Austin, Tex.) 11 29912636
2017 The Nup62 Coiled-Coil Motif Provides Plasticity for Triple-Helix Bundle Formation. Biochemistry 11 28406021
2022 Acetylation of Nup62 by TIP60 ensures accurate chromosome segregation in mitosis. Journal of molecular cell biology 10 36190325
2023 Sensitized piRNA reporter identifies multiple RNA processing factors involved in piRNA-mediated gene silencing. Genetics 8 37210214
2023 Regulation of FLC nuclear import by coordinated action of the NUP62-subcomplex and importin β SAD2. Journal of integrative plant biology 8 37278318
2009 Proteomic analysis of the effect of iptakalim on human pulmonary arterial smooth muscle cell proliferation. Acta pharmacologica Sinica 6 19169269
2024 MicroRNAome profiling of breast cancer unveils hsa-miR-5683 as a tumor suppressor microRNA predicting favorable clinical outcome. Cancer cell international 4 39538254
2023 Indel driven rapid evolution of core nuclear pore protein gene promoters. Scientific reports 4 37198214
2009 FISH-mapping comparison between river buffalo chromosome 7 and sheep chromosome 6: assignment of new loci and comparison with HSA4. Cytogenetic and genome research 4 19372676
2025 The role of ABI2 in modulating nuclear proteins: Therapeutic implications for NUP54 and NUP153 in TNBC. Advances in protein chemistry and structural biology 2 39843146
2025 Transcriptomic insights into developmental arrest in fluorescent labeling transgenic Asian elephant (Elephas maximus) embryos via inter-order cloning. Frontiers in cell and developmental biology 1 40092629
2022 An introduction to dynamic nucleoporins in Leishmania species: Novel targets for tropical-therapeutics. Journal of parasitic diseases : official organ of the Indian Society for Parasitology 1 36457769

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