Affinage

WRNIP1

ATPase WRNIP1 · UniProt Q96S55

Length
665 aa
Mass
72.1 kDa
Annotated
2026-06-11
63 papers in source corpus 27 papers cited in narrative 27 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

WRNIP1 is a conserved AAA+ ATPase that safeguards genome stability at stalled and damaged replication forks through a combination of ATPase-dependent and ATPase-independent activities (PMID:15670210, PMID:27242363). It localizes to replication factories and accumulates on chromatin and at sites of DNA damage upon fork stalling, with recruitment governed by its N-terminal ubiquitin-binding zinc finger (UBZ) domain and C-terminal leucine-zipper oligomerization region rather than its ATPase activity (PMID:18842586, PMID:22209848); a crystal structure of its UBZ domain bound to ubiquitin defines a binding surface distinct from that of pol η (PMID:27062441). At stalled forks WRNIP1 cooperates with RAD51 to stabilize the recombinase on ssDNA and block MRE11-mediated degradation of nascent DNA—an activity independent of its ATPase—while its ATPase activity is required for fork restart (PMID:27242363), and it additionally protects reversed-fork junctions from SLX4 endonucleolytic cleavage in a BRCA2-independent manner (PMID:31654852). WRNIP1 links replication stress to checkpoint signaling by bridging monoubiquitinated PCNA to the ATM cofactor ATMIN to drive ATM-dependent activation and 53BP1 focus formation (PMID:26549024). It biochemically stimulates DNA polymerase delta processivity and initiation and functions genetically in the pol δ pathway from yeast to vertebrates (PMID:15670210, PMID:12436259). Through its UBZ domain it promotes interstrand-crosslink repair by facilitating chromatin loading of the FANCD2/FANCI complex (PMID:32640220), and it restrains R-loop accumulation and transcription-replication conflicts to enable restart of transcription-stalled forks (PMID:38488661). Beyond replication, WRNIP1 (as WHIP) assembles a mitochondrial WHIP-TRIM14-PPP6C signalosome in which its UBZ domain bridges polyubiquitinated RIG-I to MAVS to promote antiviral innate immune signaling (PMID:29053956).

Mechanistic history

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

    Established that the WRNIP1 ortholog acts within the DNA polymerase delta replication pathway, defining its core genome-maintenance context before any biochemical activity was known.

    Evidence Synthetic dosage lethality and suppressor genetics in S. cerevisiae MGS1/WHIP with pol δ, RFC, PCNA, RPA, and Sgs1 mutants

    PMID:12436259 PMID:12509289

    Open questions at the time
    • Did not define a direct biochemical activity for the protein
    • Relationship to human WRNIP1 function not yet tested
  2. 2005 High

    Defined WRNIP1 as a DNA-end-stimulated octameric ATPase that physically stimulates pol δ synthesis, providing the first biochemical mechanism for its replication role.

    Evidence In vitro reconstitution with purified proteins, ATPase and DNA-synthesis assays, ATPase-dead mutant analysis

    PMID:15670210

    Open questions at the time
    • Did not establish the cellular consequence of pol δ stimulation
    • ATP-suppression of stimulation left mechanistically unexplained
  3. 2006 High

    Connected WRNIP1 to WRN and to camptothecin-induced lesion repair, while DT40 epistasis placed it in pathways parallel to RAD18 and other recombination factors.

    Evidence In vitro MBP pull-down with Walker A mutants and DT40 double-knockout epistasis (WRN, RAD18) with SCE and CPT-sensitivity readouts

    PMID:16769258 PMID:17077513

    Open questions at the time
    • Functional consequence of the WRN interaction not resolved
    • Mechanism of the parallel-to-RAD18 activity undefined
  4. 2008 Medium

    Showed WRNIP1 recruitment to replication factories and damage sites depends on its UBZ and oligomerization domains and is regulated by complex polyubiquitination, linking its targeting to the ubiquitin system.

    Evidence Immunofluorescence, chromatin fractionation, live-cell imaging with domain mutants; denaturing tandem affinity purification and MS of ubiquitin chains; parallel-pathway epistasis with BLM in DT40

    PMID:18379138 PMID:18613717 PMID:18842586

    Open questions at the time
    • Identity of the ubiquitinated chromatin ligand bound by the UBZ not yet defined
    • E3 ligase responsible for WRNIP1 polyubiquitination unknown
  5. 2009 Medium

    Demonstrated ATP-dependent binding to fork-mimicking DNA and a reciprocal regulatory interaction with RAD18, refining how WRNIP1 engages replication intermediates.

    Evidence DNA-binding assays on forked/template-primer substrates, co-immunoprecipitation, competition binding

    PMID:19556710

    Open questions at the time
    • In vivo significance of mutual RAD18 competition not established
    • Single-lab biochemistry without cellular validation
  6. 2010 Medium

    Identified an association of WRNIP1/WHIP with the Nup107-160 nuclear pore subcomplex, raising a nuclear-rim localization not tied to its WRN interaction.

    Evidence MS of isolated Nup107-160 subcomplex, reciprocal Co-IP, immunofluorescence, nuclear envelope fractionation, cell synchronization

    PMID:20676042

    Open questions at the time
    • Functional role of the NPC association unknown
    • Not integrated with the replication-fork functions
  7. 2014 High

    Placed WRNIP1 upstream of Pol η in translesion synthesis, defining its position in the UV damage response pathway.

    Evidence WRNIP1/POLH double-knockout DT40 epistasis with UV-sensitivity, CPD-repair, mutation-frequency, SCE, and fiber-progression readouts

    PMID:25139235

    Open questions at the time
    • Molecular step by which WRNIP1 acts before Pol η not defined
    • Direct biochemical link to TLS polymerase regulation not shown
  8. 2015 Medium

    Defined WRNIP1 as a molecular bridge coupling monoubiquitinated PCNA at stalled forks to ATMIN-dependent ATM checkpoint signaling.

    Evidence Co-immunoprecipitation, siRNA knockdown, 53BP1 focus and ATM-signaling assays distinguishing replication stress from ionizing radiation

    PMID:26549024

    Open questions at the time
    • Structural basis of the PCNA-WRNIP1-ATMIN bridge not resolved
    • Single-lab Co-IP for the bridging interaction
  9. 2016 High

    Separated WRNIP1's two fork-protective modes—ATPase-independent stabilization of RAD51 to block MRE11 degradation versus ATPase-dependent fork restart—and resolved the atomic basis of UBZ-ubiquitin recognition.

    Evidence DNA fiber assay, iPOND, ATPase-dead mutant, FBH1 epistasis; X-ray crystallography of the UBZ-ubiquitin complex

    PMID:27062441 PMID:27242363

    Open questions at the time
    • How RAD51 stabilization is mechanistically achieved at the fork unresolved
    • ATPase-driven restart step not biochemically reconstituted
  10. 2017 High

    Revealed a non-genome-maintenance role: as WHIP, WRNIP1 nucleates a mitochondrial signalosome that bridges polyubiquitinated RIG-I to MAVS for antiviral signaling, repurposing its UBZ and ATPase domains.

    Evidence Pooled RNAi screen, yeast two-hybrid, reciprocal Co-IP, domain mutants, antiviral signaling assays

    PMID:29053956

    Open questions at the time
    • Whether nuclear and mitochondrial pools are functionally distinct unclear
    • Role of PPP6C phosphatase within the signalosome undefined
  11. 2019 High

    Extended fork protection to reversed-fork junctions, showing an isoform-specific, BRCA2-independent shielding of these structures from SLX4 cleavage, and revealed a regulatory link to PrimPol stability.

    Evidence DNA fiber assay, SLX4/BRCA2 epistasis, EM of replication intermediates; Co-IP and proteasome-dependent degradation assays for PrimPol

    PMID:31061318 PMID:31654852

    Open questions at the time
    • Mechanism by which the short isoform recognizes reversed junctions unknown
    • Whether PrimPol degradation is direct or pathway-mediated not established
  12. 2020 High

    Broadened WRNIP1's protective repertoire to interstrand-crosslink repair via UBZ-dependent FANCD2/FANCI loading, to suppression of R-loop-driven instability, and to a yeast recombination-salvage restraint, unifying its activities as conflict resolution at forks.

    Evidence Live-cell imaging, MS of FANCD2 complex, ICL-sensitivity and UBZ-mutant assays; chromatin fractionation, S9.6 R-loop detection and RAD51 ChIP; multi-mutant genetic dissection in S. cerevisiae

    PMID:32046194 PMID:32285001 PMID:32640220

    Open questions at the time
    • How UBZ-mediated recruitment selects between ICL, R-loop, and recombination contexts unresolved
    • Direct substrate of the ATPase in these pathways not identified
  13. 2024 High

    Solidified the role of WRNIP1 in resolving transcription-replication conflicts and uncovered a context-specific partnership with nuclear AXL for fork protection in cancer cells.

    Evidence R-loop IF, DNA fiber assay, RNase H1 and transcription-inhibitor rescue, UBZ mutants; nuclear-fraction Co-IP with AXL and pharmacological inhibition

    PMID:38190717 PMID:38488661

    Open questions at the time
    • Mechanism by which AXL kinase modulates WRNIP1 at forks undefined
    • Whether R-loop resolution is direct or via RAD51 stabilization unresolved
  14. 2025 Medium

    Assigned WRNIP1 ATPase activity a fidelity function in restraining Y-family polymerase misincorporation and uncovered a new G1-phase DSB repair role independent of NHEJ.

    Evidence In vivo TLS fidelity assays with ATPase-dead WRN/WRNIP1 alleles and mutation-spectrum analysis; WRNIP1/Ku70 double-knockout DT40 epistasis with cell-cycle and γH2AX readouts

    PMID:40484680 PMID:40900148

    Open questions at the time
    • How the ATPase tightens the TLS polymerase active site not structurally shown
    • Molecular nature of the NHEJ-independent G1 DSB repair pathway undefined

Open questions

Synthesis pass · forward-looking unresolved questions
  • How a single AAA+ ATPase coordinates its many UBZ-dependent recruitment events—fork protection, ICL repair, checkpoint bridging, R-loop suppression, and mitochondrial antiviral signaling—and what determines pathway selection remains unresolved.
  • No unifying model linking the nuclear and mitochondrial functions
  • Determinants of UBZ ligand selectivity across pathways unknown
  • In vivo regulation by the ATPase cycle across contexts not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140657 ATP-dependent activity 3 GO:0003677 DNA binding 2 GO:0016787 hydrolase activity 2 GO:0060090 molecular adaptor activity 2 GO:0098772 molecular function regulator activity 2
Localization
GO:0000228 nuclear chromosome 3 GO:0005634 nucleus 2 GO:0005739 mitochondrion 2 GO:0005635 nuclear envelope 1
Pathway
R-HSA-69306 DNA Replication 3 R-HSA-73894 DNA Repair 3 R-HSA-8953897 Cellular responses to stimuli 2 R-HSA-168256 Immune System 1
Partners
ATMINAXLPOLD1PRIMPOLRAD18RAD51TRIM14WRN
Complex memberships
FANCD2/FANCI-associated complexNup107-160 nuclear pore subcomplexWHIP-TRIM14-PPP6C mitochondrial signalosome

Evidence

Reading pass · 27 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2005 Human WRNIP1 purified as a homo-octameric complex with ATPase activity stimulated by double-stranded DNA ends; it directly interacts with DNA polymerase delta (pol δ) and stimulates pol δ DNA synthesis activity more than fivefold, increasing both processivity and initiation frequency. ATP addition partially suppressed this stimulation, and an ATPase-dead WRNIP1 mutant could still stimulate pol δ but was insensitive to ATP suppression. In vitro biochemical reconstitution (purified proteins), ATPase assay, DNA synthesis assay, gel filtration, mutant analysis Genes to cells High 15670210
2002 S. cerevisiae WHIP/MGS1 (ortholog of WRNIP1) functionally interacts with DNA polymerase delta; its RFC-like motifs are essential for function. Overexpression causes synthetic dosage lethality with pol δ, RFC, PCNA, and RPA mutants. Loss of MGS1 suppresses hydroxyurea sensitivity of pol31 and pol32 (pol δ subunit) mutants, placing Mgs1 in the same functional pathway as pol δ. Genetic epistasis (synthetic dosage lethality screen, suppressor analysis, double-mutant growth assays in S. cerevisiae) Molecular genetics and genomics High 12436259
2002 In S. cerevisiae, simultaneous deletion of WHIP/MGS1 and SGS1 (RecQ helicase) causes elevated G2/M arrest, increased spontaneous sister chromatid recombination, and shortened lifespan. Complementation of the synthetic growth defect requires both the helicase/Top3-binding activity of Sgs1 and the ATPase activity of Mgs1. Genetic epistasis (double-deletion analysis, complementation with domain mutants), flow cytometry, SCR measurement in S. cerevisiae DNA repair High 12509289
2006 MBP-tagged WRNIP1 directly interacts with WRN in vitro; interaction is enhanced by ATP addition. Walker A motif mutations showed that WRNIP1, but not WRN, must bind ATP for efficient interaction. In DT40 cells, WRNIP1/WRN double knockout showed synergistic increase in camptothecin sensitivity and elevated sister-chromatid exchange compared to single knockouts. In vitro pull-down (MBP-tagged proteins), Walker A mutant analysis, DT40 knockout cell lines, camptothecin sensitivity assay, SCE measurement DNA repair High 16769258
2008 Human WRNIP1 localizes to DNA replication factories in unperturbed cells and its chromatin association increases upon replication fork stalling (UVC). Localization to subnuclear structures (replication factories and PML bodies) depends on its N-terminal ubiquitin-binding zinc finger (UBZ) domain and C-terminal oligomerization domain. Immunofluorescence, chromatin fractionation, live-cell imaging, domain-deletion mutant analysis, co-localization with replication markers The Journal of biological chemistry High 18842586
2008 WRNIP1 is polyubiquitinated in vivo with complex chains linked through K11, K48, and K63 of ubiquitin. This polyubiquitination depends on an intact UBZ domain and is enhanced by UV radiation. Sumoylation and phosphorylation are also detected as post-translational modifications. Denaturing tandem affinity purification, mass spectrometry, UV treatment, UBZ domain mutant analysis Journal of proteome research Medium 18613717
2009 Human WRNIP1 binds forked DNA and template/primer DNA in an ATP-dependent manner. WRNIP1 physically interacts with RAD18 and interferes with RAD18 binding to forked and template/primer DNA. Reciprocally, RAD18 enhances WRNIP1 binding to these DNA substrates. DNA-binding assay (forked/template-primer substrates), co-immunoprecipitation, competition binding assay Genes & genetic systems Medium 19556710
2011 WRNIP1 accumulates rapidly at laser-irradiated DNA damage sites via its UBZ domain and C-terminal leucine zipper region. The ATPase domain and WRNIP1 ubiquitination (lysine acceptor sites) are dispensable for this recruitment. Laser microirradiation, live-cell imaging, domain-deletion and lysine mutant analysis Biochemical and biophysical research communications Medium 22209848
2015 WRNIP1 bridges monoubiquitinated PCNA (a stalled replication fork marker) with the ATM cofactor ATMIN, connecting PCNA monoubiquitination to ATM signaling. WRNIP1, ATMIN, and RAD18 are specifically required for ATM activation and 53BP1 focus formation in response to replication stress (but not ionising radiation). Co-immunoprecipitation, siRNA knockdown, 53BP1 focus formation assay, ATM signaling assays Oncogene Medium 26549024
2016 WRNIP1 localizes to stalled replication forks and cooperates with RAD51 to prevent MRE11-mediated nucleolytic degradation of nascent DNA at stalled forks by stabilizing RAD51 on ssDNA. Fork protection does not require WRNIP1 ATPase activity, but ATPase activity is required for recovery/restart of perturbed replication forks. Loss of WRNIP1 causes DNA damage and chromosomal aberrations; downregulation of anti-recombinase FBH1 rescues fork degradation and aberrations in WRNIP1-deficient cells. DNA fiber assay, iPOND, siRNA/shRNA knockdown, ATPase-dead mutant, chromosomal aberration analysis, epistasis with FBH1 knockdown The EMBO journal High 27242363
2016 Crystal structure of the WRNIP1 UBZ domain in complex with ubiquitin was determined, revealing a novel ubiquitin-binding surface composed of the first β-strand and the C-terminal α-helix, distinct from the pol η UBZ binding mode. X-ray crystallography (GFP-fusion crystallization), structural comparison The FEBS journal High 27062441
2017 WRNIP1 (as WHIP) assembles into a mitochondrial signaling complex with TRIM14 and PPP6C. The UBZ (ubiquitin-binding) domain of WRNIP1 bridges RIG-I with MAVS by binding polyubiquitin chains on RIG-I at K164. The ATPase domain of WRNIP1 stabilizes the RIG-I–dsRNA interaction. This WHIP-TRIM14-PPP6C signalosome promotes RIG-I-mediated innate antiviral signaling. Pooled RNAi screen, yeast two-hybrid, co-immunoprecipitation, domain mutant analysis, antiviral signaling assays Molecular cell High 29053956
2019 WRNIP1 protects the junction point of reversed replication forks from SLX4-mediated endonucleolytic cleavage, functioning downstream of fork reversal. This protective activity is specific to the shorter, less abundant isoform of WRNIP1 and is independent of the BRCA2-dependent fork protection branch. DNA fiber assay, siRNA knockdown, epistasis with SLX4/BRCA2 depletion, electron microscopy of replication intermediates iScience High 31654852
2019 WRNIP1 and PrimPol form a complex in cells. Overexpression of WRNIP1 reduces PrimPol protein levels via proteasome-dependent degradation, while WRNIP1 depletion increases PrimPol levels. The ATPase domain of WRNIP1 is involved in regulating PrimPol amount. Co-immunoprecipitation, overexpression/knockdown, proteasome inhibitor treatment, domain mutant analysis Biological & pharmaceutical bulletin Medium 31061318
2020 WRNIP1 is recruited to DNA interstrand crosslinks (ICLs) rapidly after their formation, promoting ICL repair and facilitating subsequent recruitment of the FANCD2/FANCI complex. Ubiquitination of WRNIP1 and integrity of its UBZ domain are required for FANCD2/FANCI chromatin loading and efficient ICL repair. WRNIP1 was identified in a FANCD2-containing protein complex by MS. Live-cell imaging, mass spectrometry of FANCD2 complex, siRNA depletion, ICL-drug sensitivity assay, UBZ mutant analysis Cell reports High 32640220
2020 In S. cerevisiae, Mgs1/WRNIP1 ATPase activity prevents a recombination salvage pathway at stalled replication forks. In the absence of Rad5 (DDT factor), loss of Mgs1 activates a RAD52/RAD59-dependent recombination bypass pathway that requires pol δ and PCNA-K164 modification and is enabled by Esc2 and Elg1. Mgs1 normally inhibits this pathway to favor Rad5-dependent template switching. Genetic epistasis (double/triple mutants in S. cerevisiae), DNA replication assay, viability under replication stress, mutant analysis Science advances High 32285001
2020 WRNIP1 is retained on chromatin and required to maintain genome integrity in cells with compromised ATR checkpoint. WRNIP1 mediates ATM-dependent CHK1 phosphorylation. WRNIP1 chromatin retention stabilizes RAD51 association with ssDNA near R-loops, and loss of WRNIP1 increases R-loop-dependent genomic instability. Chromatin fractionation, immunofluorescence, siRNA knockdown, ATM/ATR inhibition, R-loop detection (S9.6 antibody), RAD51 ChIP Cancers Medium 32046194
2010 WRNIP1/WHIP was found to associate in vivo with the Nup107-160 subcomplex of the nuclear pore complex (NPC), identified by mass spectrometry. Reciprocal immunoprecipitation confirmed WHIP-Nup107 interaction. WHIP localizes to the nuclear rim and nuclear matrix by immunofluorescence. This NPC association is dynamic through the cell cycle and occurs without interaction with WRN. Mass spectrometry of isolated Nup107-160 subcomplex, reciprocal co-immunoprecipitation, immunofluorescence, nuclear envelope fractionation, cell synchronization Cell cycle Medium 20676042
2014 WRNIP1 functions upstream of DNA polymerase η (Polη) in the UV-induced DNA damage response. Disruption of WRNIP1 in Polη-deficient DT40 cells suppresses Polη-loss phenotypes: UV sensitivity, delayed CPD repair, elevated mutation frequency, elevated UV-induced SCE, and reduced fork progression after UV. This epistasis places WRNIP1 upstream of Polη in translesion synthesis. Genetic epistasis (WRNIP1/POLH double-knockout DT40 cells), UV sensitivity assay, CPD repair assay, mutation frequency, SCE assay, DNA fiber assay Biochemical and biophysical research communications High 25139235
2022 The UBZ domain of WRNIP1 is responsible for reducing UV-induced PCNA monoubiquitylation in Polη-deficient cells. The ATPase domain regulates PrimPol protein levels. The leucine zipper domain is required for interaction with RAD18 and with DNA pol δ catalytic subunit POLD1. Domain-deletion and point mutants of WRNIP1 expressed in DT40 double-knockout cells, PCNA monoubiquitylation assay, co-immunoprecipitation with RAD18 and POLD1, UV sensitivity Biological & pharmaceutical bulletin Medium 35110507
2024 WRNIP1 co-localizes with transcription/replication complexes and R-loops upon replication perturbation. Loss of WRNIP1 leads to R-loop accumulation and replication-transcription conflicts. WRNIP1 is required for replication restart from transcription-induced fork stalling. The UBZ domain is critical for preventing pathological R-loop persistence. Transcription inhibition or RNase H1 overexpression rescues defects caused by WRNIP1 loss. Immunofluorescence, S9.6 R-loop detection, DNA fiber assay, RNase H1 overexpression rescue, transcription inhibitor rescue, siRNA/shRNA depletion, UBZ mutant analysis eLife High 38488661
2024 Nuclear AXL (TAM receptor tyrosine kinase) interacts with WRNIP1 and this interaction promotes protection of stalled replication forks in HER2+ breast cancer cells. Knockdown or inhibition of AXL or WRNIP1 attenuates fork protection. Co-immunoprecipitation (nuclear fraction), siRNA knockdown, pharmacological inhibition, DNA fiber assay Cancer research Medium 38190717
2023 WRNIP1 is essential for the cellular response to mitochondrial DNA double-strand breaks (mtDSBs). CRISPR screen identified WRNIP1 as specifically essential under mitochondrial (not nuclear) DNA damage. WRNIP1 is implicated in mitochondrial turnover in response to mtDSBs and in innate immune signaling downstream of mtDNA damage. CRISPR/Cas9 genome-wide screen with mitochondria-targeted doxorubicin, functional validation by WRNIP1 knockdown/knockout, mitochondrial damage assays ACS chemical biology Medium 38054633
2025 WRN and WRNIP1 ATPase activities are required for high-fidelity translesion synthesis (TLS) by Y-family DNA polymerases in human cells. Defects in WRN or WRNIP1 ATPase activity cause diverse nucleotide misincorporations opposite DNA lesions by Y-family Pols, indicating these ATPase activities restrain misincorporations (possibly by tightening the TLS Pol active site), while WRN exonuclease removes misinserted nucleotides. In vivo TLS fidelity assays in human cells with ATPase-dead mutants of WRN and WRNIP1, mutation spectrum analysis eLife Medium 40900148
2025 WRNIP1 has a novel function in G1-phase DNA double-strand break (DSB) repair, independent of non-homologous end joining (NHEJ). Deletion of WRNIP1 in NHEJ-deficient DT40 cells causes slow growth, G1 accumulation, increased dead cells, and accumulation of γH2AX-marked DSBs. Genetic epistasis (WRNIP1/Ku70 double-knockout DT40 cells), flow cytometry (cell cycle), γH2AX immunofluorescence, growth assay Biological & pharmaceutical bulletin Medium 40484680
2006 WRNIP1-deficient DT40 cells and RAD18-deficient DT40 cells both show moderate camptothecin sensitivity; double knockout cells show additive/synergistic SCE elevation and slower growth. Unexpectedly, the severe CPT sensitivity of RAD18-deficient cells is slightly suppressed by loss of WRNIP1, suggesting WRNIP1 acts in a pathway parallel to RAD18 in vertebrate cells. Genetic epistasis (WRNIP1/RAD18 double-knockout DT40 cells), CPT sensitivity, SCE measurement, growth assay Biological & pharmaceutical bulletin Medium 17077513
2008 WRNIP1 and BLM (RecQ helicase) independently suppress sister chromatid exchange in vertebrate cells; WRNIP1/BLM double-knockout DT40 cells show additive SCE elevation and enhanced sensitivity to camptothecin but not MMS, indicating they function in parallel pathways for CPT-induced lesion repair. Genetic epistasis (wrnip1/blm double-knockout DT40 cells), SCE measurement, CPT/MMS sensitivity assay Genes & genetic systems Medium 18379138

Source papers

Stage 0 corpus · 63 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 WRNIP1 protects stalled forks from degradation and promotes fork restart after replication stress. The EMBO journal 76 27242363
2017 Assembly of the WHIP-TRIM14-PPP6C Mitochondrial Complex Promotes RIG-I-Mediated Antiviral Signaling. Molecular cell 68 29053956
2008 Human Wrnip1 is localized in replication factories in a ubiquitin-binding zinc finger-dependent manner. The Journal of biological chemistry 62 18842586
2005 Human Werner helicase interacting protein 1 (WRNIP1) functions as a novel modulator for DNA polymerase delta. Genes to cells : devoted to molecular & cellular mechanisms 59 15670210
2002 The product of Saccharomyces cerevisiae WHIP/MGS1, a gene related to replication factor C genes, interacts functionally with DNA polymerase delta. Molecular genetics and genomics : MGG 54 12436259
1987 On non-cyclooxygenase prostaglandin synthesis in the sea whip coral, Plexaura homomalla: an 8(R)-lipoxygenase pathway leads to formation of an alpha-ketol and a Racemic prostanoid. The Journal of biological chemistry 54 2824470
2015 RAD18, WRNIP1 and ATMIN promote ATM signalling in response to replication stress. Oncogene 38 26549024
2019 miR-22 enhances the radiosensitivity of small-cell lung cancer by targeting the WRNIP1. Journal of cellular biochemistry 36 31190355
2006 Analyses of the interaction of WRNIP1 with Werner syndrome protein (WRN) in vitro and in the cell. DNA repair 32 16769258
2002 Characterization of the slow-growth phenotype of S. cerevisiae Whip/Mgs1 Sgs1 double deletion mutants. DNA repair 32 12509289
2020 Genomic resources and toolkits for developmental study of whip spiders (Amblypygi) provide insights into arachnid genome evolution and antenniform leg patterning. EvoDevo 28 32874529
2009 Physical and functional interaction between WRNIP1 and RAD18. Genes & genetic systems 28 19556710
1987 Morphology, sense organs, and regeneration of the forelegs (whips) of the whip spider Heterophrynus elaphus (Arachnida, Amblypygi). Journal of morphology 27 29921113
2017 The role of WRNIP1 in genome maintenance. Cell cycle (Georgetown, Tex.) 25 28118071
2022 R-Loop-Associated Genomic Instability and Implication of WRN and WRNIP1. International journal of molecular sciences 24 35163467
2017 Metabolome Dynamics of Smutted Sugarcane Reveals Mechanisms Involved in Disease Progression and Whip Emission. Frontiers in plant science 24 28620397
2019 WRNIP1 Protects Reversed DNA Replication Forks from SLX4-Dependent Nucleolytic Cleavage. iScience 22 31654852
2008 Conjugation of complex polyubiquitin chains to WRNIP1. Journal of proteome research 19 18613717
2017 Sugarcane smut: shedding light on the development of the whip-shaped sorus. Annals of botany 18 27568298
2016 A novel mode of ubiquitin recognition by the ubiquitin-binding zinc finger domain of WRNIP1. The FEBS journal 17 27062441
2020 Checkpoint Defects Elicit a WRNIP1-Mediated Response to Counteract R-Loop-Associated Genomic Instability. Cancers 16 32046194
2018 Also looking like Limulus? - retinula axons and visual neuropils of Amblypygi (whip spiders). Frontiers in zoology 16 30574172
2020 WRNIP1 Is Recruited to DNA Interstrand Crosslinks and Promotes Repair. Cell reports 15 32640220
2008 Vertebrate WRNIP1 and BLM are required for efficient maintenance of genome stability. Genes & genetic systems 15 18379138
2024 AXL/WRNIP1 Mediates Replication Stress Response and Promotes Therapy Resistance and Metachronous Metastasis in HER2+ Breast Cancer. Cancer research 14 38190717
1980 Giant neurons and associated synapses in the peripheral nervous system of whip spiders. Journal of neurocytology 13 7441302
2011 WRNIP1 accumulates at laser light irradiated sites rapidly via its ubiquitin-binding zinc finger domain and independently from its ATPase domain. Biochemical and biophysical research communications 12 22209848
2006 Functional relationships between Rad18 and WRNIP1 in vertebrate cells. Biological & pharmaceutical bulletin 12 17077513
2020 The Mgs1/WRNIP1 ATPase is required to prevent a recombination salvage pathway at damaged replication forks. Science advances 11 32285001
2019 WRNIP1 Controls the Amount of PrimPol. Biological & pharmaceutical bulletin 10 31061318
2014 WRNIP1 functions upstream of DNA polymerase η in the UV-induced DNA damage response. Biochemical and biophysical research communications 10 25139235
2020 A histone H2A-derived antimicrobial peptide, Hipposin from mangrove whip ray, Himantura walga: Molecular and functional characterisation. 3 Biotech 9 33088663
2019 The visual system of Thelyphonida (whip scorpions): Support for Arachnopulmonata. Arthropod structure & development 9 31176004
2010 The discovery of a Werner Helicase Interacting Protein (WHIP) association with the nuclear pore complex. Cell cycle (Georgetown, Tex.) 8 20676042
2002 Peripheral synapses and giant neurons in whip spiders. Microscopy research and technique 8 12214295
2024 WRNIP1 prevents transcription-associated genomic instability. eLife 6 38488661
2016 The water-repellent cerotegument of whip-spiders (Arachnida: Amblypygi). Arthropod structure & development 6 27751783
2024 Circular RNA WRNIP1 activates the PI3K-AKT and ERK1/2 signaling pathways by binding to miR-129-5p/IGF2 axis and facilitates ovarian follicle development in chickens. Poultry science 5 39756109
2020 Cryptic diversity within three South American whip spider species (Arachnida, Amblypygi). Zoological research 5 32738110
2013 Chromosomal characteristics of a Brazilian whip spider (Amblypygi) and evolutionary relationships with other arachnid orders. Genetics and molecular research : GMR 5 24085433
2023 CRISPR Screening in Tandem with Targeted mtDNA Damage Reveals WRNIP1 Essentiality. ACS chemical biology 4 38054633
2021 Insights into the Karyotype Evolution of Charinidae, the Early-Diverging Clade of Whip Spiders (Arachnida: Amblypygi). Animals : an open access journal from MDPI 4 34827965
2020 Whip-LAMP: a novel LAMP assay for the detection of Trichuris muris-derived DNA in stool and urine samples in a murine experimental infection model. Parasites & vectors 4 33160406
2006 Low-energy states of a semiflexible polymer chain with attraction and the whip-toroid transitions. The Journal of chemical physics 4 16942378
2024 Assembly of FAP93 at the proximal axoneme in Chlamydomonas cilia. Cytoskeleton (Hoboken, N.J.) 3 38224153
2016 WRNIP1: A new guardian of genome integrity at stalled replication forks. Molecular & cellular oncology 3 27857978
2015 Flipping a Lipid-Linked Oligosaccharide? You Must Whip It! Trends in biochemical sciences 3 26476576
2022 The first complete mitochondrial genome of the micro-whip-scorpion Schizomus zhensis (Arachnida: Schizomida) and phylogenetic analysis. Mitochondrial DNA. Part B, Resources 2 35493713
2017 Unexpected Alliance of WHIP-TRIM14-PPP6C to Combat Viruses. Molecular cell 2 29053952
2022 Functional Domain Mapping of Werner Interacting Protein 1 (WRNIP1). Biological & pharmaceutical bulletin 1 35110507
2021 A new species of whip spider, Weygoldtia hainanensis sp. nov., from Hainan, China (Arachnida: Amblypygi: Charinidae). Zootaxa 1 35390983
2016 The phylogenomic position of the smalleye whip ray Himantura microphthalma (Myliobatiformes: Dasyatidae) inferred from the mitochondrial genome. Mitochondrial DNA. Part B, Resources 1 33473557
1919 PHOTOREACTIONS OF PARTIALLY BLINDED WHIP-TAIL SCORPIONS. The Journal of general physiology 1 19871760
2025 Defective Repair of G1-Phase Double-Strand Breaks in WRNIP1/Ku70 Double Knockout Cells. Biological & pharmaceutical bulletin 0 40484680
2025 WRN and WRNIP1 ATPases impose high fidelity on translesion synthesis by Y-family DNA polymerases. eLife 0 40900148
2025 Comparative analysis of the mitochondrial genome of whip scorpion, Typopeltis sinensis (Butler, 1872) (Arachnida: Thelyphonidae) with phylogenetic implication. Journal of genetics 0 40919758
2025 Functions of a subunit of DNA polymerase δ, POLD3, revealed by depletion of WRNIP1. Biochemical and biophysical research communications 0 41319413
2024 Whole-Genome Sequencing Analyses Reveal the Whip-like Tail Formation, Innate Immune Evolution, and DNA Repair Mechanisms of Eupleurogrammus muticus. Animals : an open access journal from MDPI 0 38338077
2024 New mitochondrial genomes of three whip spider species from the Amazon (Arachnida, Amblypygi) with phylogenetic relationships and comparative analysis. Scientific reports 0 39487275
2023 CRISPR Screening in Tandem with Targeted mtDNA Damage Reveals WRNIP1 Essentiality. bioRxiv : the preprint server for biology 0 37873237
2022 A new species of whip spider, Sarax sinensis sp. nov., from Fujian, China (Arachnida: Amblypygi: Charinidae). Zootaxa 0 36095500
2022 Appearance of a "Whip-Like" Rash in a Young Male Undergoing Therapy for Testicular Embryonal Carcinoma. Cureus 0 36721590
2015 Phylogeographic Study of Whip Scorpions (Chelicerata: Arachnida: Thelyphonida) in Japan and Taiwan. Zoological science 0 26245222

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