Affinage

N4BP1

NEDD4-binding protein 1 · UniProt O75113

Length
896 aa
Mass
100.4 kDa
Annotated
2026-06-10
23 papers in source corpus 15 papers cited in narrative 15 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

N4BP1 is a multifunctional immune-restriction protein that couples post-transcriptional silencing of inflammatory transcripts to direct suppression of NF-κB signaling, integrating its activity into checkpoints that limit antiviral and inflammatory output (PMID:32971525, PMID:38697117). As an endoribonuclease, it uses both its KH domain and a PilT N-terminus (NYN/YacP-like) RNase domain to bind and degrade target mRNAs through their coding sequences—recognizing a polyC-containing element near the 5' CDS—in a manner independent of nonsense-mediated decay (PMID:31133753, PMID:39491646); through this activity it restricts HIV-1 by degrading viral mRNA (PMID:31133753) and limits inflammation by destabilizing JunB, FosB, and chemokine transcripts in keratinocytes and neutrophils (PMID:33990547), while a parallel mode blocks translation of the IL-17 adaptor Act1 (TRAF3IP2) to restrain p38 MAPK–driven cytokine production (PMID:41504891). In NF-κB control, N4BP1 is a constitutive suppressor of MyD88-dependent TLR (TLR1/2, TLR7, TLR9) and TNF responses (PMID:32971525); it binds the NEMO COZI domain via UBA-like and CUE-like regions to block IKKγ oligomerization (PMID:33654074), reads linear (M1) polyubiquitin chains at the nascent TNFR1 complex through a dimerization-dependent LUBIN module to limit signaling duration and restrain complex II–driven cell death (PMID:38643192), and operates together with the non-canonical kinases TBK1/IKKε and the adaptor TANK in a polyubiquitin-binding-dependent manner (PMID:38697117). N4BP1 is switched off by proteolysis: caspase-8 cleaves it downstream of TRIF, death receptors, and MALT1 to license TLR3/4 and death-receptor cytokine output and HIV reactivation (PMID:31133753, PMID:32971525, PMID:33654074), and enteroviral 3C protease cleaves it at Q816 to relieve NF-κB restriction (PMID:39655957). The protein additionally binds the E3 ligase Itch to inhibit substrate ubiquitylation (PMID:17592138) and antagonizes Notch signaling by promoting Trim21-dependent degradation of NICD via its CoCUN domain (PMID:37807845). Its abundance and localization are regulated by Nedd4-mediated polyubiquitylation opposed by SUMO1 conjugation (PMID:20233849), an NLS (residues 279–299) governing CRM1-dependent nucleocytoplasmic shuttling, and CoCUN-dependent phase separation with neddylated cullins under stress (PMID:40701250).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2007 High

    Established the first molecular activity for N4BP1—as a competitive inhibitor of the E3 ligase Itch—answering whether it participates in ubiquitin pathway regulation.

    Evidence Co-IP, in vitro and in vivo ubiquitylation assays, RNAi and knockout cells

    PMID:17592138

    Open questions at the time
    • Did not address RNase or NF-κB functions later attributed to N4BP1
    • Physiological context of Itch antagonism not defined
  2. 2010 High

    Defined how N4BP1 protein levels and subnuclear localization are controlled, showing a Nedd4-ubiquitylation/SUMO1 balance and nucleolar/PML body residence.

    Evidence Ubiquitylation and SUMOylation assays, immunofluorescence, SENP1-mutant fibroblasts, proteasome inhibition

    PMID:20233849

    Open questions at the time
    • Functional consequence of nucleolar vs PML localization unresolved
    • Link between stability control and downstream signaling roles not established
  3. 2019 High

    Identified N4BP1 as an NYN-domain endoribonuclease that restricts HIV-1 and is inactivated by MALT1 cleavage, defining both an enzymatic activity and a switch-off mechanism.

    Evidence mRNA degradation assays, Arg509 mutagenesis, MALT1 KO, primary T cell/macrophage infection

    PMID:31133753

    Open questions at the time
    • Substrate sequence determinants not yet defined
    • Mechanism of RNA target selection unknown
  4. 2019 High

    Structurally characterized the C-terminal CoCUN domain as a ubiquitin-binding module distinct from CUBAN, defining how N4BP1 reads ubiquitin.

    Evidence NMR chemical shift perturbation, mutagenesis, circular dichroism, sequence analysis

    PMID:31319543

    Open questions at the time
    • In-cell function of CoCUN ubiquitin reading not yet linked to a pathway
    • Selectivity for chain linkage types not addressed
  5. 2020 High

    Placed N4BP1 as a constitutive suppressor of MyD88-dependent TLRs and TNF that is licensed off by caspase-8 cleavage, establishing it as an inflammatory checkpoint.

    Evidence N4BP1 KO mice, caspase-8-deficient macrophages, N4BP1/caspase-8 double-KO rescue, cytokine and cleavage assays

    PMID:32971525

    Open questions at the time
    • Molecular mechanism by which N4BP1 suppresses signaling not defined here
    • Whether RNase activity mediates the cytokine suppression unaddressed
  6. 2021 High

    Provided a molecular mechanism for NF-κB suppression—direct binding to the NEMO COZI domain to block IKK oligomerization—and mapped the caspase-8 cleavage that abolishes it.

    Evidence Co-IP, in vitro binding, domain deletion mapping, N4bp1-deficient mice, cleavage site mutagenesis

    PMID:33654074

    Open questions at the time
    • Relationship between NEMO binding and ubiquitin reading not resolved
    • Stoichiometry of NEMO inhibition unknown
  7. 2021 Medium

    Connected N4BP1 RNase activity to specific inflammatory mRNA targets and disease, showing destabilization of JunB/FosB and chemokine transcripts and a psoriasis-like phenotype in KO mice.

    Evidence RNA-IP, transcriptome profiling, N4BP1 KO mice, keratinocyte proliferation assays, IMQ psoriasis model

    PMID:33990547

    Open questions at the time
    • Single lab
    • Direct demonstration that catalytic RNase activity drives each phenotype not shown
  8. 2023 Medium

    Extended N4BP1 function to Notch signaling, showing CoCUN-dependent, Trim21-mediated NICD degradation and a role in neural progenitor differentiation.

    Evidence Protein binding assays, Phe-Pro motif mutagenesis, Trim21 KO, progenitor overexpression/KO, cortical development assays

    PMID:37807845

    Open questions at the time
    • Single lab
    • How CoCUN ubiquitin binding directs NICD to Trim21 mechanistically unclear
  9. 2024 Medium

    Defined the LUBIN module as a dimerization-dependent linear-ubiquitin reader that recruits N4BP1 to the TNFR1 complex to limit signaling duration and cell death.

    Evidence Structural modeling, mutagenesis, TNFR1 complex IP, KO functional assays, ubiquitin-chain binding, caspase-8 cleavage

    PMID:38643192

    Open questions at the time
    • Single lab
    • High-resolution structure of LUBIN–M1 chain interaction not determined
  10. 2024 High

    Demonstrated genetic cooperation of N4BP1 with TBK1/IKKε and TANK in a ubiquitin-binding-dependent manner to terminate TLR-driven IKK signaling.

    Evidence N4BP1 KO and ubiquitin-binding mutant knock-in mice, TBK1/IKKε/TANK KO macrophages, cytokine assays, epistasis

    PMID:38697117

    Open questions at the time
    • Biochemical nature of the N4BP1–TBK1/TANK complex not resolved
    • Whether the same module mediates both TLR and TNFR1 effects unaddressed
  11. 2024 High

    Resolved the RNA-degradation mechanism, showing CDS-targeted, polyC-element-dependent cleavage requiring KH and NYN domains and independent of NMD.

    Evidence Domain/point mutagenesis, reporter and CDS/3'UTR swap assays, UPF1/UPF3A/UPF3B KO cells, RNA-IP

    PMID:39491646

    Open questions at the time
    • Full consensus of the targeted RNA element not defined
    • How CDS engagement is coordinated with translation unknown
  12. 2024 Medium

    Showed that enteroviral 3C protease cleaves N4BP1 at Q816 to relieve NF-κB restriction, identifying a viral evasion strategy and species-specific susceptibility.

    Evidence Motif search, cleavage assays, 3Cpro expression in cells, Q816 mutagenesis, NF-κB readouts

    PMID:39655957

    Open questions at the time
    • Single lab
    • Whether cleavage affects RNase function in addition to NF-κB not tested
  13. 2025 Medium

    Characterized N4BP1 trafficking and biophysics, defining an NLS, CRM1-dependent export, and CoCUN-dependent phase separation with neddylated cullins that confers stress protection.

    Evidence Leptomycin B, NLS deletion/GFP fusion, live imaging, 1,6-hexanediol assay, co-localization with neddylated cullins, heat shock

    PMID:40701250

    Open questions at the time
    • Single lab
    • Functional link between phase separation and RNase/NF-κB roles not established
  14. 2026 Medium

    Identified a translational-control mechanism, showing N4BP1 blocks Act1 (TRAF3IP2) mRNA translation to restrain IL-17/p38 signaling, distinct from mRNA destabilization.

    Evidence Polysome profiling, mRNA stability assays, N4BP1 KO cells/mice, Act1 shRNA rescue, p38 inhibition, IMQ skin model

    PMID:41504891

    Open questions at the time
    • Single lab
    • How N4BP1 selects translational vs degradative fate for different mRNAs unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how N4BP1 integrates its distinct activities—CDS-directed RNA cleavage, translational repression, NEMO/linear-ubiquitin-based NF-κB suppression, and E3-ligase modulation—into a unified regulatory logic, and which activity dominates in a given cell type or stimulus.
  • No single-molecule or structural framework unifying the RNase and ubiquitin-reader modules
  • Determinants selecting between degradation, translational block, and signaling roles undefined
  • Crosstalk between localization/phase separation and effector functions unmapped

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 4 GO:0003723 RNA binding 3 GO:0031386 protein tag activity 3 GO:0140098 catalytic activity, acting on RNA 2 GO:0045182 translation regulator activity 1
Localization
GO:0005634 nucleus 2 GO:0005730 nucleolus 1 GO:0005829 cytosol 1
Pathway
R-HSA-162582 Signal Transduction 3 R-HSA-168256 Immune System 3 R-HSA-1643685 Disease 2 R-HSA-8953854 Metabolism of RNA 2 R-HSA-5357801 Programmed Cell Death 1
Complex memberships
TNFR1 signaling complex

Evidence

Reading pass · 15 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2007 N4BP1 binds the WW2 domain of the E3 ubiquitin ligase Itch, competing with Itch substrates (p73α, c-Jun, p63) for the same binding site, thereby inhibiting Itch-mediated ubiquitylation of these substrates and Itch autoubiquitylation activity both in vitro and in vivo. Co-immunoprecipitation, in vitro and in vivo ubiquitylation assays, RNAi knockdown, N4BP1 knockout cells Proceedings of the National Academy of Sciences of the United States of America High 17592138
2010 N4BP1 is a substrate for Nedd4-mediated polyubiquitylation and proteasomal degradation. SUMOylation of N4BP1 by SUMO1 abrogates its ubiquitylation, stabilizing it. Endogenous N4BP1 localizes predominantly to the nucleolus in primary cells, with a fraction at PML nuclear bodies; loss of the desumoylase SENP1 or proteasome inhibition causes accumulation of N4BP1 at PML nuclear bodies. Ubiquitylation assays, SUMOylation assays, immunofluorescence localization, subcellular fractionation, SENP1-mutant primary embryonic fibroblasts, proteasome inhibitor (MG132) treatment Journal of cell science High 20233849
2019 N4BP1 harbors a PilT N-terminus (NYN)-like RNase domain and inhibits HIV-1 replication by binding to and degrading viral mRNA species. N4BP1 is cleaved at Arg509 by the paracaspase MALT1 following T cell activation, and this cleavage inactivates its HIV-1 restriction activity and facilitates reactivation of latent HIV-1 proviruses. RNA-binding and mRNA degradation assays, mutational analysis (Arg509), MALT1 knockout studies, primary T cell and macrophage infection models Nature microbiology High 31133753
2019 N4BP1 contains a novel ubiquitin-binding domain (CoCUN) in its C-terminal ~50 residues that recognizes the canonical hydrophobic patch of ubiquitin (Ile44) through a Phe-Pro motif, resembling CUE domain recognition but evolutionarily distinct. CoCUN does not bind NEDD8 (unlike the related CUBAN domain). Both CoCUN and CUBAN are polyubiquitinated in cells. NMR spectroscopy (15N chemical shift perturbation), mutagenesis, circular dichroism, protein sequence analysis Biomolecules High 31319543
2020 N4BP1 is a suppressor of TLR1/2-, TLR7-, and TLR9-driven cytokine production (MyD88-dependent TLRs), but not TLR3 or TLR4 responses in wild-type macrophages. N4BP1 is cleaved and inactivated by caspase-8 downstream of TRIF signaling (TLR3, TLR4) or TNF/death receptor engagement, thereby licensing those pathways to produce higher cytokine levels. Caspase-8-deficient macrophages retain intact N4BP1, explaining their impaired cytokine responses to TLR3/4; co-deletion of N4BP1 in caspase-8-deficient macrophages largely rescues this defect. N4BP1 knockout mice, caspase-8-deficient macrophages, N4BP1/caspase-8 double-knockout genetic rescue, cytokine production assays, cleavage assays Nature High 32971525
2021 N4BP1 inhibits TLR-dependent NF-κB activation by directly interacting with the NEMO (IKKγ) COZI domain through its UBA-like and CUE-like domains, attenuating NEMO dimerization/oligomerization. Caspase-8, activated downstream of TRIF (TLR3/TLR4), cleaves N4BP1 distal to residues D424 and D490, abolishing its inhibitory effect on NF-κB. Co-immunoprecipitation, in vitro binding assays, domain deletion mapping, N4bp1-deficient mice, caspase-8 cleavage site mutagenesis Nature communications High 33654074
2021 In keratinocytes, N4BP1 binds JunB and FosB mRNAs and reduces their stability, thereby suppressing keratinocyte proliferation. In neutrophils, N4BP1 limits survival and tissue infiltration by targeting CXCL1, CCL20, and S100A8 mRNAs. N4BP1-deficient mice develop severe IMQ-induced psoriasis-like disease. RNA immunoprecipitation, transcriptome profiling, N4BP1 KO mice, primary keratinocyte proliferation assays, IMQ psoriasis model Cell death & disease Medium 33990547
2023 N4BP1 negatively regulates Notch signaling by binding NICD (specifically via the NICD RAM domain) and promoting its ubiquitin-mediated proteasomal degradation after Notch1 S3 cleavage. The CoCUN domain Phe-Pro motif (residues 862/863) in N4BP1 is essential for NICD degradation. The E3 ligase Trim21 is required for N4BP1-regulated NICD degradation. In cortical neural progenitors, N4BP1 overexpression promotes differentiation while N4BP1 loss maintains stem-like properties. Protein binding assays, domain mutagenesis (Phe-Pro motif), Trim21 KO, overexpression and KO in neural progenitor cells, cortical development assays The EMBO journal Medium 37807845
2024 N4BP1 acts as a dimerization-dependent linear ubiquitin reader through a novel ubiquitin-binding module named LUBIN, which positions two non-selective ubiquitin-binding domains to preferentially recognize linear (M1) ubiquitin chains. N4BP1 is recruited to the nascent TNFR1 signaling complex and limits NF-κB signaling duration in a LUBIN-dependent manner. N4BP1 deficiency accelerates TNFα-induced cell death by increasing complex II assembly. Under apoptotic conditions, caspase-8 cleaves N4BP1, leading to its rapid 26S proteasomal degradation. Structural modeling, mutagenesis, TNFR1 signaling complex immunoprecipitation, N4BP1 KO functional assays, ubiquitin-chain binding assays, caspase-8 cleavage assays Cell death discovery Medium 38643192
2024 N4BP1 acts in concert with non-canonical IκB kinases (TBK1 and IKKε) and their adaptor TANK to limit the duration of canonical IKKα/β signaling after MyD88-dependent TLR activation. Deletion of TBK1/IKKε or TANK phenocopies N4BP1 deficiency. Optimal suppression requires N4BP1's ability to bind polyubiquitin chains, as inactivating mutations in the ubiquitin-binding motif of N4BP1 increase TLR-induced cytokine production. N4BP1 KO and ubiquitin-binding mutant knock-in mice, TBK1/IKKε/TANK KO macrophages, cytokine production assays, genetic epistasis Immunity High 38697117
2024 N4BP1 degrades its mRNA targets via their coding sequences (CDS) rather than the 3'-UTR. A critical 33-nt sequence (nt 289–322) near the 5' end of FosC CDS, containing a polyC motif, is required for N4BP1-mediated degradation. Both the KH domain and the NYN (YacP-like nuclease) domain are essential for N4BP1's mRNA-degrading activity. N4BP1-mediated mRNA turnover is independent of nonsense-mediated decay (NMD), as it is preserved in UPF1/UPF3A/UPF3B-deficient cells. Domain deletion mutagenesis, point mutations, reporter assays, CDS/3'UTR swap experiments, NMD-deficient cell lines (UPF1/UPF3A/UPF3B KO), RNA immunoprecipitation The Journal of biological chemistry High 39491646
2024 Enteroviral 3C protease (3Cpro) from multiple human enteroviruses cleaves N4BP1 at residue Q816, destroying its ability to regulate TNFα-induced NF-κB signaling. Mouse N4BP1, which has a threonine at the P1' site, is resistant to human enteroviral 3Cpro, but rodent EMCV 3Cpro cleaves both human and mouse N4BP1 at species-specific sites. FIMO motif search, biochemical cleavage assays, cell biology experiments with 3Cpro expression, site mutagenesis (Q816), NF-κB signaling readouts Journal of virology Medium 39655957
2025 N4BP1 is a nucleocytoplasmic shuttling protein. Nuclear accumulation is induced by leptomycin B (blocking CRM1-mediated export). A nuclear localization signal (NLS) spanning residues 279–299 is required for nuclear import; its deletion abolishes nuclear import and fusion to GFP drives GFP into the nucleus. N4BP1 forms liquid-liquid phase separation aggregates in both cytoplasm and nucleus that colocalize with NEDD8-modified proteins (including cullin-1 and cullin-2), dependent on the CoCUN domain. Heat shock promotes N4BP1 aggregation, which confers cellular protection under stress. Leptomycin B treatment, NLS deletion and GFP fusion constructs, live imaging, 1,6-hexanediol phase separation assay, co-localization with neddylated proteins, heat shock experiments The Journal of biological chemistry Medium 40701250
2026 N4BP1 suppresses IL-17 signaling by blocking translation of Act1 (TRAF3IP2) mRNA rather than affecting its stability. In N4BP1-deficient cells, Act1 mRNA is enriched in the translationally active polysome fraction, leading to elevated Act1 protein, hyperactivation of p38 MAPK, and increased downstream cytokines (CXCL1, CCL20, MMP9). Polysome profiling, mRNA stability assays, N4BP1 KO cells and mice, Act1 shRNA rescue experiments, p38 pharmacological inhibition, IMQ-induced skin model Inflammation research Medium 41504891
2025 N4BP1 is identified as a negative regulator of NF-κB that is targeted by poxvirus effector myxoma virus M3.1; N4BP1 inactivation by M3.1 unleashes an NF-κB ETI response, placing N4BP1 functionally alongside ZC3H12A and TBK1 as anti-viral NF-κB suppressors that can serve as pathogen sensors. Virulence factor expression screen in human monocytes, RNA-seq transcriptional response assays bioRxiv (preprint)preprint Low bio_10.1101_2025.03.04.641538

Source papers

Stage 0 corpus · 23 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 The Nedd4-binding partner 1 (N4BP1) protein is an inhibitor of the E3 ligase Itch. Proceedings of the National Academy of Sciences of the United States of America 98 17592138
2020 Integration of innate immune signalling by caspase-8 cleavage of N4BP1. Nature 93 32971525
2019 N4BP1 restricts HIV-1 and its inactivation by MALT1 promotes viral reactivation. Nature microbiology 63 31133753
2021 N4BP1 negatively regulates NF-κB by binding and inhibiting NEMO oligomerization. Nature communications 41 33654074
2010 N4BP1 is a newly identified nucleolar protein that undergoes SUMO-regulated polyubiquitylation and proteasomal turnover at promyelocytic leukemia nuclear bodies. Journal of cell science 36 20233849
2017 miR-28-5p promotes the development and progression of ovarian cancer through inhibition of N4BP1. International journal of oncology 26 28350078
2024 N4BP1 coordinates ubiquitin-dependent crosstalk within the IκB kinase family to limit Toll-like receptor signaling and inflammation. Immunity 23 38697117
2021 The endoribonuclease N4BP1 prevents psoriasis by controlling both keratinocytes proliferation and neutrophil infiltration. Cell death & disease 22 33990547
2019 CoCUN, a Novel Ubiquitin Binding Domain Identified in N4BP1. Biomolecules 16 31319543
2024 N4BP1 functions as a dimerization-dependent linear ubiquitin reader which regulates TNF signalling. Cell death discovery 10 38643192
2023 N4BP1 mediates RAM domain-dependent notch signaling turnover during neocortical development. The EMBO journal 8 37807845
2013 Identification of a novel N4BP1-like gene from grass carp (Ctenopharyngodon idella) in response to GCRV infection. Fish & shellfish immunology 8 24220004
2024 Enteroviral 3C protease cleaves N4BP1 to impair the host inflammatory response. Journal of virology 7 39655957
2024 The NEDD4-binding protein N4BP1 degrades mRNA substrates through the coding sequence independent of nonsense-mediated decay. The Journal of biological chemistry 6 39491646
2025 Exosomal miR-92b-5p regulates N4BP1 to enhance PTEN mono-ubiquitination in doxorubicin-resistant AML. Cancer drug resistance (Alhambra, Calif.) 4 40201312
2023 N4BP1 regulates keratinocytes development and plays protective role in burn- and adhesive-related skin injury via MMP9. Cellular signalling 4 37579928
2025 N4BP1 as a modulator of the NF-κB pathway. Cytokine & growth factor reviews 1 40312219
2026 N4BP1 acts as a potent negative regulator of IL-17 signaling by blocking the translation of Act1 mRNA. Inflammation research : official journal of the European Histamine Research Society ... [et al.] 0 41504891
2026 N4BP1 is essential for the development of oral cancer via controlling both cancer cells and immune microenvironment. Cell death & disease 0 41513619
2026 N4BP1 promotes the progression of acute glaucoma by promoting pyroptosis through the JAK2/STAT3 pathway. Experimental eye research 0 41611111
2026 Altered expression of ADAR1, N4BP1, and PSME1 in PBMCs correlated with therapeutic outcomes in HBeAg-negative chronic hepatitis B patients treated with Peg-IFN-α. Frontiers in cellular and infection microbiology 0 42052091
2025 N4BP1 is a nucleocytoplasmic shuttling protein and recognizes aggregates of the ubiquitin-like protein NEDD8 to protect cells under heat shock. The Journal of biological chemistry 0 40701250
2024 Non-canonical IKKs side with N4BP1 against the family. Immunity 0 38749393

Missed literature

Know a paper Affinage missed for N4BP1? Flag it for the maintainers and the community.

No submissions yet.