Affinage

RIF1

Insulin-like peptide INSL6 · UniProt Q9Y581

Length
213 aa
Mass
24.9 kDa
Annotated
2026-06-10
100 papers in source corpus 48 papers cited in narrative 48 extracted findings
Cross-family judge faithfulness: 8/8 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

RIF1 is a large multifunctional scaffold that governs DNA double-strand break (DSB) repair pathway choice and the spatiotemporal control of DNA replication, in both cases largely by acting as a targeting subunit for Protein Phosphatase 1 (PP1) (PMID:23333306, PMID:24532715, PMID:28522851). In DSB repair, ATM-phosphorylated 53BP1 recruits RIF1 to break sites, where RIF1 acts as the critical 53BP1 effector that inhibits 5'-end resection and channels repair toward NHEJ in G1, a function antagonized by BRCA1/CtIP in S/G2; loss of RIF1 restores resection and RAD51 loading in BRCA1-deficient cells and underlies class switch recombination and dysfunctional-telomere fusion (PMID:23333306, PMID:23333305, PMID:23306437, PMID:23306439). RIF1 is a phosphopeptide-binding protein that directly engages 53BP1 LxL-phospho motifs, and its recruitment is further reinforced by methylated histone marks (H3K4 via SETD1A-BOD1L; H4K20me2-dependent assembly of the 53BP1-RIF1-MAD2L2 complex) (PMID:35216668, PMID:35439434, PMID:29160738). Downstream of recruitment, RIF1 enforces NHEJ by directly binding SHLD3 through its HEAT-repeat domain to nucleate the shieldin complex, which together with CST and Pol-alpha fills in resected ends, and by recruiting the histone chaperone ASF1 to compact flanking chromatin (PMID:30022158, PMID:37306046, PMID:35177609). In replication, RIF1-PP1 reverses DDK (Cdc7-Dbf4)-mediated phosphorylation of the MCM helicase (notably MCM4) and protects ORC1 to control origin licensing and firing timing, a mechanism conserved from budding and fission yeast through Xenopus and human cells (PMID:24532715, PMID:24685139, PMID:24656819, PMID:28077461, PMID:28273463). RIF1 organizes nuclear architecture and replication-timing domains by coating late-replicating, Lamin B1-associated chromatin and binding G-quadruplex DNA through its HEAT-repeat and C-terminal segments and oligomerization (PMID:22850674, PMID:26725008, PMID:26436827, PMID:29348174). RIF1-PP1 additionally protects stalled/reversed replication forks from DNA2-WRN-mediated degradation independently of its NHEJ role, and recruits PP1 to the midbody to time cytokinetic abscission by counteracting Aurora B-driven CHMP4C phosphorylation (PMID:31141682, PMID:31337767, PMID:30905608). In yeast, RIF1 builds higher-order telomeric architecture via Rap1 binding and oligomerization and restrains Tel1/ATM, and is anchored to the inner nuclear membrane by Pfa4-mediated S-acylation that supports DSB repair (PMID:23746845, PMID:19217405, PMID:31182712).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2004 High

    Established that human Rif1 is a damage-responsive factor acting downstream of ATM and 53BP1 rather than a constitutive telomere protein, defining its entry point into the DNA damage response.

    Evidence siRNA, immunofluorescence co-localization at dysfunctional telomeres, ATM/53BP1 epistasis, intra-S checkpoint and clonogenic survival assays

    PMID:15342490 PMID:15583028

    Open questions at the time
    • Did not define the molecular function of RIF1 at breaks
    • Anaphase midzone localization left mechanistically unexplained
  2. 2009 High

    Showed RIF1 has essential roles in replication-stress survival and fork-associated repair in mammals, beyond telomere biology.

    Evidence Conditional Rif1 deletion in MEFs, HDR reporter, clonogenic survival, immunofluorescence at stalled forks

    PMID:19948482

    Open questions at the time
    • Did not separate replication-timing from fork-protection functions
    • Molecular partners at forks undefined
  3. 2010 High

    Identified RIF1 as a BLM-complex component with intrinsic branched-DNA-binding activity, providing a biochemical basis for its fork association.

    Evidence Co-IP, domain mapping, in vitro fork/Holliday-junction DNA binding, DT40 replication-stress epistasis

    PMID:20711169

    Open questions at the time
    • Relationship between BLM-bound and 53BP1-bound RIF1 pools unresolved
  4. 2012 High

    Defined RIF1 as a global regulator of DNA replication timing and nuclear chromatin-domain architecture, answering how it shapes the replication program.

    Evidence Genome-wide replication timing, chromatin-loop and nuclear-fractionation analysis in human cells and conditional mouse MEFs

    PMID:22850673 PMID:22850674

    Open questions at the time
    • Did not identify the enzymatic effector linking RIF1 to timing control
    • Mechanism of chromatin-loop organization unspecified
  5. 2013 High

    Resolved RIF1 as the principal 53BP1 effector inhibiting end resection to promote NHEJ and antagonize BRCA1, establishing its central role in DSB pathway choice.

    Evidence Reciprocal Co-IP with phospho-53BP1, Rif1 knockout mice, CSR and telomere-fusion assays, resection/RAD51 readouts, epistasis with CtIP/BLM/Exo1; four concurrent studies

    PMID:23306437 PMID:23306439 PMID:23333305 PMID:23333306 PMID:23486525

    Open questions at the time
    • Did not define the structural basis of 53BP1 phosphopeptide recognition
    • Downstream resection-blocking effectors not yet identified
  6. 2014 High

    Identified RIF1 as a PP1 phosphatase-targeting subunit that counteracts DDK phosphorylation of MCM via RVxF/SILK motifs, providing the molecular mechanism of replication-timing control.

    Evidence In vitro phosphorylation and PP1 Co-IP, RVxF/SILK mutagenesis, genome-wide timing analysis, genetic suppressors in budding and fission yeast

    PMID:24532715 PMID:24656819 PMID:24685139

    Open questions at the time
    • Conservation of the PP1-MCM mechanism to humans not yet shown at this stage
  7. 2014 Medium

    Characterized intrinsic DNA-structure-binding activities of RIF1's C-terminal region and described H3K9me3-dependent telomere/chromatin regulation in mESCs.

    Evidence NMR of CRII, in vitro cruciform-DNA binding and mutagenesis; telomere FISH, H3K9me3 ChIP and methyltransferase Co-IP in ESCs

    PMID:24634216 PMID:24735877

    Open questions at the time
    • Functional contribution of cruciform binding in vivo not established
    • ESC chromatin role rests on single-lab Co-IP/ChIP
  8. 2015 High

    Linked RIF1 to nuclear genome organization and to anaphase chromosome-segregation fidelity, broadening its cell-cycle functions.

    Evidence ChIP-seq, Hi-C topology and super-resolution imaging in mESCs; G4-binding ChIP-seq and origin-firing assays in fission yeast; UFB analysis with PICH/BLM/53BP1 epistasis

    PMID:26256213 PMID:26436827 PMID:26725008

    Open questions at the time
    • UFB role characterized in a single lab
    • Causal link between G4 binding and domain architecture in mammals not direct
  9. 2017 High

    Demonstrated that the RIF1-PP1-MCM mechanism is conserved in human and Xenopus systems and that RIF1 also stabilizes ORC1, unifying licensing and firing control; a crystal structure defined the DNA-encasing Rif1-NTD.

    Evidence Phosphoproteomics and ORC1 stability assays in human cells, Xenopus extract reconstitution, DNA fiber analysis; X-ray crystallography of yeast Rif1-NTD; PP1 biophysics (NMR/ITC/SPR)

    PMID:28077461 PMID:28273463 PMID:28522851 PMID:28604726

    Open questions at the time
    • Structure of full-length human RIF1 not determined
    • Selectivity of RIF1-PP1 toward distinct substrates not fully mapped
  10. 2017 Medium

    Identified chromatin remodelers and additional adaptors (CSB, SCAI) that modulate RIF1 retention and turnover at breaks, refining the dynamics of pathway choice.

    Evidence Co-IP, domain mapping, histone-eviction and foci dynamics assays, HDR reporters in human cells

    PMID:28700933 PMID:29203878

    Open questions at the time
    • Single-lab Co-IP evidence
    • Quantitative ordering of CSB/SCAI/BRCA1 displacement of RIF1 incomplete
  11. 2018 High

    Placed RIF1 upstream of the shieldin-CST-Pol-alpha fill-in machinery and dissected histone-mark and oligomerization determinants of its recruitment and DNA binding.

    Evidence Co-IP/resection/PARPi assays for CST-shieldin; full-length murine Rif1 hydrodynamics and G4-binding; H4K20me2 ChIP and 53BP1-RIF1-MAD2L2 Co-IP

    PMID:29160738 PMID:29348174 PMID:30022158

    Open questions at the time
    • Direct RIF1-shieldin contact not yet defined at this stage
    • MAD2L2-recruitment complex shown by single-lab Co-IP
  12. 2019 High

    Separated RIF1's PP1-dependent fork-protection function from its NHEJ function and established a PP1-dependent role in cytokinetic abscission timing.

    Evidence DNA fiber and PLA assays with DNA2/WRN epistasis and NHEJ-separation-of-function mutants; live-cell abscission imaging with Aurora B/PP1 epistasis and CHMP4C phospho-readout

    PMID:30905608 PMID:31141682 PMID:31337767

    Open questions at the time
    • How RIF1 selects fork substrates of PP1 versus break substrates unresolved
    • Abscission role established in a single study
  13. 2022 High

    Defined the structural logic of RIF1 recruitment to 53BP1 and chromatin: direct LxL-phosphopeptide recognition and direct binding to methylated H3K4 via SETD1A-BOD1L.

    Evidence In vitro phosphopeptide pulldowns and 53BP1-motif mutagenesis with IR-foci readout; in vitro RIF1 binding to H3K4me peptides, depletion epistasis, CSR and PARPi assays including patient cells

    PMID:35216668 PMID:35439434

    Open questions at the time
    • Structural model of the RIF1-53BP1 interface not solved
    • Integration of phosphopeptide and histone-mark cues for recruitment unresolved
  14. 2023 High

    Established a direct RIF1 HEAT-repeat-SHLD3 binding interface as the molecular link nucleating shieldin recruitment for NHEJ.

    Evidence AlphaFold2-Multimer prediction validated by in vitro pulldown, co-localization, CSR and PARPi sensitivity assays

    PMID:37306046

    Open questions at the time
    • Predicted interface not confirmed by experimental structure
    • Stoichiometry of RIF1-shieldin assembly unknown

Open questions

Synthesis pass · forward-looking unresolved questions
  • How RIF1 partitions its single PP1-targeting and DNA/chromatin-binding scaffold across replication timing, fork protection, DSB pathway choice, abscission, and Wnt signaling within one cell remains unresolved.
  • No structure of full-length human RIF1 or its multivalent assemblies
  • Determinants selecting context-specific PP1 substrates unknown
  • Whether distinct RIF1 functional pools are spatially/temporally segregated is unclear

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 6 GO:0003677 DNA binding 5 GO:0060090 molecular adaptor activity 4 GO:0042393 histone binding 1
Localization
GO:0005634 nucleus 3 GO:0000228 nuclear chromosome 2 GO:0005635 nuclear envelope 1
Pathway
R-HSA-73894 DNA Repair 6 R-HSA-69306 DNA Replication 5 R-HSA-1640170 Cell Cycle 3 R-HSA-4839726 Chromatin organization 2
Partners
ASF1BLMCTC1MAD2L2PPP1CARAP1SHLD3TP53BP1
Complex memberships
53BP1-RIF1-MAD2L253BP1-RIF1-shieldinBLM complex

Evidence

Reading pass · 48 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2013 ATM-dependent phosphorylation of 53BP1 physically recruits RIF1 to DSB sites; RIF1 acts as the critical effector of 53BP1 to inhibit end resection and promote NHEJ in G1 phase. BRCA1 and CtIP antagonize RIF1 accumulation at DSBs in S phase. Depletion of RIF1 restores end resection and RAD51 loading in BRCA1-depleted cells. Co-immunoprecipitation, siRNA knockdown, cell-cycle-specific analysis of DSB repair foci, RAD51 loading assays Molecular cell High 23333306
2013 RIF1 is recruited to DSBs via the N-terminal phospho-SQ/TQ domain of 53BP1 in an ATM-dependent manner. RIF1 deficiency in mice severely compromises 53BP1-dependent class switch recombination (CSR) and fusion of dysfunctional telomeres. DSBs are hyperresected in the absence of RIF1. Deletion of Rif1 suppresses toxic NHEJ induced by PARP inhibition in Brca1-deficient cells. Rif1 knockout mice, CSR assay, telomere fusion assay, end-resection analysis, Co-immunoprecipitation, domain mapping Molecular cell High 23333305
2013 Rif1 is the main factor used by 53BP1 to impair 5' end resection at DSBs. Rif1 inhibits resection involving CtIP, BLM, and Exo1; limits BRCA1/BARD1 accumulation at damage sites; and underlies 53BP1-caused chromosomal abnormalities in Brca1-deficient cells. Genetic deletion of Rif1 in mouse cells, dysfunctional telomere assay, genome-wide DSB analysis, epistasis with CtIP/BLM/Exo1 Science High 23306437
2013 Rif1 is an ATM phosphorylation-dependent interactor of 53BP1 and its absence results in 5'-3' DNA-end resection. Rif1 deficiency impairs DSB repair in G1 and S phases, interferes with class switch recombination in B lymphocytes, and leads to accumulation of chromosomal DSBs. Co-immunoprecipitation with phospho-53BP1, Rif1 knockout mouse, CSR assay, cell cycle analysis of resection Science High 23306439
2013 RIF1 counteracts BRCA1-mediated end resection downstream of 53BP1. RIF1 also regulates foci formation and chromatin loading of BLM helicase, a function distinct from 53BP1. siRNA knockdown, resection assays, BLM chromatin loading assay, Co-immunoprecipitation Journal of Biological Chemistry Medium 23486525
2018 The 53BP1-RIF1-shieldin complex counteracts DSB resection through CST (CTC1-STN1-TEN1) and Polymerase-α-mediated fill-in of resected DNA ends. CST interacts with shieldin and localizes with Polα to sites of DNA damage in a 53BP1- and shieldin-dependent manner. Co-immunoprecipitation, siRNA depletion, DNA-end resection assays, RAD51 loading, PARP inhibitor sensitivity assays Nature High 30022158
2022 RIF1 is a phosphopeptide-binding protein that directly interacts with three phosphorylated 53BP1 epitopes sharing an essential LxL motif followed by two closely apposed phosphorylated residues. Simultaneous mutation of these sites abrogates RIF1 accumulation at IR-induced foci. RIF1 also modifies shieldin action independently of 53BP1 binding. In vitro pulldown, mutagenesis of 53BP1 phosphopeptide motifs, IR-induced foci analysis, biochemical binding assays Molecular cell High 35216668
2023 AlphaFold2 modeling and in vitro pulldown identified a direct binding interface between the HEAT-repeat domain of RIF1 and the eIF4E-like domain of SHLD3. This RIF1-SHLD3 direct interaction is essential for shieldin recruitment to DNA damage sites, antibody class switch recombination, and PARP inhibitor sensitivity. AlphaFold2-Multimer structural prediction, in vitro pulldown, cellular co-localization assays, CSR assay, PARP inhibitor sensitivity EMBO reports High 37306046
2017 Crystal structure of the ~125-kDa N-terminal domain of yeast Rif1 (Rif1-NTD) reveals an α-helical fold shaped like a shepherd's crook. The Rif1-NTD contains a high-affinity DNA-binding site that fully encases DNA as a head-to-tail dimer. This DNA-binding activity is essential for checkpoint control and telomere length regulation in yeast, and Rif1-NTD also promotes NHEJ at DNA breaks. X-ray crystallography, in vitro DNA binding assays, yeast genetics, checkpoint assay Nature structural & molecular biology High 28604726
2014 Budding yeast Rif1 controls DNA replication genome-wide by directing Protein Phosphatase 1 (PP1) to dephosphorylate the MCM complex, counteracting DDK (Cdc7-Dbf4)-mediated phosphorylation of Mcm4. PP1 interaction motifs (RVxF/SILK) in the Rif1 N-terminal domain are critical for its replication-repressive effect. DDK phosphorylates near these motifs to downregulate Rif1-PP1 interaction. In vitro phosphorylation assay, Co-immunoprecipitation, genetic suppressor analysis, Mcm4 phosphorylation assay, domain mutagenesis Genes & development High 24532715
2014 Budding yeast Rif1 inhibits activation of pre-replication complexes via two N-terminal PP1-docking motifs (RVxF and SILK) that recruit PP1 phosphatase (Glc7). Rif1 interaction with Glc7 is required to suppress DDK-mediated phosphorylation of Mcm4 and Sld3. Rif1 also interacts with Dbf4 in yeast two-hybrid assays. RVxF/SILK motif mutagenesis, PP1 Co-immunoprecipitation, genome-wide replication timing analysis, yeast two-hybrid Cell reports High 24685139
2014 Fission yeast Rif1 inhibits origin firing through PP1 phosphatase. Mutations of two PP1 docking motifs in Rif1 lead to early replication of telomeres and misregulation of origin firing. Rif1/PP1 counteract DDK activity on the replicative MCM helicase. PP1 docking motif mutagenesis, replication timing assay, origin firing analysis Cell reports High 24656819
2017 Human RIF1 forms a complex with PP1 that limits phosphorylation-mediated activation of the MCM replicative helicase. Specific residues on four MCM helicase subunits (especially the N-terminal domain of MCM4) show hyperphosphorylation upon RIF1 depletion. In addition, RIF1-PP1 protects ORC1 from untimely phosphorylation and proteasomal degradation during G1, promoting efficient origin licensing. RIF1 depletion, phosphoproteomics (MS), ORC1 stability assays, origin spacing analysis by DNA combing EMBO reports High 28077461
2017 In Xenopus egg extracts and human cells, RIF1-PP1 reverses DDK-mediated MCM phosphorylation (specifically hyperphosphorylation of Mcm4). Loss of Rif1 increases MCM phosphorylation and the rate of replication initiation. RIF1 can also mediate MCM dephosphorylation at replication forks, and loss of Rif1 compromises the ability of cells to block initiation under replication stress. Xenopus egg extract reconstitution, DDK inhibitors, phospho-specific antibodies, DNA fiber analysis Cell reports High 28273463
2017 Mouse Rif1 is a high-affinity PP1 adaptor protein. Using NMR, isothermal calorimetry, and surface plasmon resonance, Rif1 was shown to bind PP1 with higher affinity than the established PP1-inhibitor I2 in vitro. Novel Rif1 interactors involved in chromatin metabolism and phosphorylation were also identified. NMR, isothermal calorimetry, surface plasmon resonance, mutagenesis, mass spectrometry interactome Scientific reports High 28522851
2012 Human Rif1 depletion results in specific loss of mid-S replication foci profiles, stimulation of early-S-phase initiation events, and changes in long-range replication timing domain structures. Rif1 binds to nuclear-insoluble structures at late-M-to-early-G1 and regulates chromatin-loop sizes. Rif1 colocalizes specifically with mid-S replication foci. siRNA depletion, BrdU incorporation genome-wide, chromatin loop size analysis, nuclear fractionation, fluorescence microscopy EMBO journal High 22850674
2012 Mouse Rif1 deficiency causes an unprecedented global alteration in the temporal order of replication in primary cells, already in the first S-phase after deletion. Rif1 deficiency also leads to defective G1/S transition and chromatin re-organization after DNA replication, without altering the transcriptional landscape or major heterochromatin identity. Conditional Rif1 deletion in primary MEFs, genome-wide replication timing analysis, chromatin analysis EMBO journal High 22850673
2012 Fission yeast Rif1 is a global regulator of replication origin firing timing. Rif1 binds not only to telomeres but also to specific arm segment locations near late/dormant origins (in a Taz1-independent manner during M-to-G1). Deletion of rif1 leads to deregulation of dormant origins and suppression/delay of many early-firing origins. ChIP-seq, rif1 deletion, BrdU-IP-seq replication timing, genetic suppressor analysis of hsk1 Genes & development High 22279046
2015 In mouse embryonic stem cells, Rif1 coats late-replicating domains and, with Lamin B1, identifies most of the late-replicating genome. Rif1 defines and restricts interactions between replication-timing domains during G1 phase, organizing nuclear architecture. Loss of Rif1 affects number and replication-timing specificity of domain interactions. During S phase, Rif1 ensures temporally coordinated replication of interacting domains. ChIP-seq, Hi-C/genome topology analysis, super-resolution microscopy, conditional knockout in mESCs Molecular cell High 26725008
2015 Fission yeast Rif1 binds to G quadruplex (G4)-like structures at 35 high-affinity chromosomal binding sites containing conserved CNWWGTGGGGG motifs. Base substitution within these motifs abolishes Rif1 binding and activates late-firing or dormant origins up to 50 kb away, demonstrating long-range suppressive effects. ChIP-seq, in vitro G4 binding assay with purified Rif1, base-substitution mutagenesis, origin firing analysis Nature structural & molecular biology High 26436827
2018 Purified murine Rif1 forms elongated homo-oligomers in solution and binds G-quadruplex (G4) DNA with high specificity and affinity. Both the N-terminal (HEAT-repeat) and C-terminal segments participate in oligomer formation and G4 binding. The central intrinsically disordered segment increases G4 affinity. Rif1 can simultaneously bind multiple G4 molecules. Hydrodynamic analysis, in vitro G4 binding assay, pulldown with multiple G4 molecules, full-length murine Rif1 purification Journal of Biological Chemistry High 29348174
2014 The C-terminal region of murine Rif1 contains conserved region II (CRII) that is partially folded and binds cruciform DNA with high selectivity and micromolar affinity. Mutational analysis identified a critical α-helical region of CRII required for cruciform DNA binding. NMR structural analysis, ESPRIT truncation library, in vitro cruciform DNA binding assay, mutagenesis Journal of Biological Chemistry High 24634216
2004 Human Rif1 localizes to dysfunctional telomeres and to telomeric DNA clusters in ALT cells after DSB induction. This localization depends strictly on ATM and 53BP1, but not ATR, BRCA1, Chk2, Nbs1, or Mre11. Rif1 inhibition causes radiosensitivity and a defect in the intra-S-phase checkpoint, acting in a pathway distinct from Nbs1. siRNA inhibition, immunofluorescence co-localization with damage markers, clonogenic survival after IR, S-phase checkpoint assay, ATM/53BP1 epistasis Genes & development High 15342490
2004 Human Rif1 does not accumulate at functional telomeres or interact with TRF1, TRF2, or hRap1. During early anaphase specifically, hRif1 aligns along a subset of midzone microtubules between separating chromosomes, a novel cell-cycle-regulated subcellular localization. Immunofluorescence, siRNA RNAi, co-immunoprecipitation (negative for TRF1/TRF2/hRap1), cell cycle fractionation and imaging Journal of Cell Biology Medium 15583028
2009 Mammalian Rif1 deficiency leads to failure in embryonic development. Conditional deletion causes S-phase progression defects and hypersensitivity to replication poisons. Rif1 knockdown decreases the efficiency of homology-directed repair (HDR), and Rif1 deficiency results in aberrant Rad51 aggregates. Rif1 accumulates at stalled replication forks, preferentially around pericentromeric heterochromatin. Conditional gene deletion in MEFs, clonogenic survival, HDR reporter assay, siRNA, immunofluorescence at stalled forks Journal of Cell Biology High 19948482
2010 Rif1 is a novel component of the BLM complex, interacting through a conserved C-terminal domain. Rif1 stability depends on BLM complex presence. Rif1 is recruited to stalled replication forks with similar kinetics as BLM, and recruitment is delayed in BLM-deficient cells. Vertebrate Rif1 contains a DNA-binding domain resembling the αCTD of bacterial RNA polymerase α that preferentially binds fork and Holliday junction DNA in vitro. Co-immunoprecipitation, domain mapping, replication stress survival assay in DT40 cells, in vitro DNA binding assay, immunofluorescence at stalled forks EMBO journal High 20711169
2019 RIF1 protects nascent DNA at stalled replication forks from over-degradation by DNA2 nuclease (operating with WRN as accessory helicase). This protection requires RIF1's interaction with PP1. RIF1 limits phosphorylation of WRN at resection-control sites. Loss of RIF1-mediated fork protection leads to DNA breakage accumulation. DNA fiber assay, RIF1 depletion, DNA2/WRN knockdown epistasis, WRN phosphorylation analysis, PP1 interaction mutants Cell reports High 31141682
2019 RIF1 is enriched at stalled replication forks and protects reversed forks from DNA2 nuclease-mediated degradation independently of its NHEJ function but dependently on its PP1 interaction. RIF1 deficiency delays fork restart and results in exposure of under-replicated DNA. Proximity ligation assay at forks, DNA fiber assay, RIF1 NHEJ-separation-of-function mutants, DNA2 inhibitor/knockdown Nature communications High 31337767
2019 RIF1 and its binding partner PP1 are critical for regulation of abscission timing in human cells. RIF1 promotes cytokinesis by recruiting PP1 to the midbody, which counteracts Aurora B kinase activity, leading to dephosphorylation of the abscission regulator CHMP4C. siRNA depletion, live-cell imaging of abscission, immunofluorescence of midbody localization, CHMP4C phosphorylation assay, Aurora B inhibition epistasis Current Biology High 30905608
2015 Rif1 is required for resolution of ultrafine DNA bridges (UFBs) in anaphase. Rif1 is recruited to UFBs in a PICH-dependent but 53BP1- and BLM-independent manner. Loss of Rif1 increases the frequency of nucleoplasmic bridges and RPA70-positive UFBs, and increases nuclear bodies with damaged DNA in G1. siRNA depletion, immunofluorescence of UFBs in anaphase, epistasis with PICH/BLM/53BP1 Developmental cell Medium 26256213
2013 Yeast Rif1 and Rif2 have separable and independent Rap1-binding epitopes (crystal structures determined). Rif1 tetramerization and Rif2 polymerization modules, combined with long-range Rap1 binding, generate a higher-order architecture (molecular Velcro) that interlinks Rap1 units on telomeric arrays. This architecture is required for telomere homeostasis in vivo. X-ray crystallography of Rif1-Rap1C and Rif2-Rap1C complexes; biochemical domain dissection; genetic analysis Cell High 23746845
2009 Budding yeast Rif1 and Rif2 inhibit Tel1 (ATM) recruitment to DNA ends through distinct mechanisms. Both inhibit Tel1 localization but not MRX complex localization to adjacent DNA ends. Rif2 competes with Tel1 for binding to the C terminus of Xrs2. ChIP analysis, genetic epistasis, Tel1/MRX localization assays at defined DNA ends Molecular cell High 19217405
2011 Palmitoylation of budding yeast Rif1 by the palmitoyltransferase Pfa4 anchors Rif1 to the inner nuclear membrane, influencing its role in heterochromatin dynamics at HMR/HML mating-type loci. Loss of palmitoylation causes Rif1-GFP to disperse from nuclear peripheral foci into the nucleoplasm. Palmitoylation detection in cell extracts, fluorescence microscopy of Rif1-GFP localization, genetic epistasis (pfa4Δ upstream of rif1) PNAS Medium 21844336
2019 Yeast Rif1 is S-acylated by Pfa4 palmitoyl acyltransferase at cysteine residues C466 and C473 within its conserved N-terminal domain. Rif1 S-acylation facilitates accumulation at DSBs, attenuation of DNA end-resection, and DSB repair by NHEJ. S-acylated Rif1 mounts a localized DNA-damage response at the inner nuclear membrane. S-acylation biochemical detection, Rif1 cysteine mutagenesis (C466A, C473A), DSB end-resection assay, NHEJ assay, inner nuclear membrane localization by imaging Nature communications High 31182712
2022 H3K4 methylation by the SETD1A-BOD1L complex facilitates RIF1 recruitment to DSBs. RIF1 directly binds methylated H3K4. Compromising SETD1A or BOD1L leads to uncontrolled DNA end resection, impairs telomere end-joining, and abrogates class switch recombination. In BRCA1-deficient cells, loss of SETD1A-dependent RIF1 recruitment restores HR and causes PARP inhibitor resistance. RIF1 foci analysis in SETD1A/BOD1L-depleted cells, in vitro direct binding of RIF1 to methylated H3K4 peptides, genetic epistasis, CSR assay, PARP inhibitor sensitivity Molecular cell High 35439434
2018 MAD2L2 is recruited to DSBs by forming a protein complex with 53BP1 and RIF1 in H4K20 dimethylated (pre-replicative) chromatin. Saturating H4K20me2 in non-replicated DNA promotes robust 53BP1-RIF1-MAD2L2 recruitment and exclusion of BRCA1. Replication-associated dilution of H4K20me2 releases the 53BP1-RIF1-MAD2L2 complex and favors BRCA1 access. Co-immunoprecipitation of 53BP1-RIF1-MAD2L2 complex, H4K20me2 ChIP, cell cycle analysis, DSB foci analysis Cell cycle Medium 29160738
2022 RIF1 interacts with the histone chaperone ASF1 (in a manner similar to CAF-1 and HIRA interactions with ASF1). ASF1 is recruited to distal chromatin flanking DSBs by 53BP1-RIF1. ASF1 uses its histone chaperone activity to promote NHEJ. Epistasis shows ASF1 acts in the same NHEJ pathway as RIF1 but via a parallel pathway with shieldin. ASF1 compacts adjacent chromatin by heterochromatinization to protect broken ends from BRCA1-mediated resection. Co-immunoprecipitation (RIF1-ASF1 interaction), chromatin fractionation, epistasis analysis, resection assay, NHEJ assay in BRCA1-deficient cells Nature communications Medium 35177609
2017 CSB interacts via its winged helix domain with RIF1, mediating CSB recruitment to DSBs in S phase. At DSBs, CSB remodels chromatin by evicting histones, which limits RIF1 and MAD2L2 accumulation but promotes BRCA1 accumulation. This chromatin remodeling requires ATM-dependent phosphorylation of CSB at S10 and CDK2-dependent phosphorylation at S158, both modulating the CSB N-terminal/ATPase domain interaction. Co-immunoprecipitation (CSB-RIF1), domain mapping, histone eviction assay, RIF1/MAD2L2/BRCA1 foci analysis, ATM/CDK2 phosphorylation analysis Nature communications Medium 29203878
2017 SCAI binds to 53BP1 phosphorylated at S/TP sites and facilitates HDR. RIF1 immediately accumulates at damage sites after DSB induction and then gradually dissociates from 53BP1 and is replaced by SCAI. SCAI inhibits RIF1 function to allow BRCA1-mediated repair. Co-immunoprecipitation, time-lapse imaging of foci, HDR reporter assay, depletion of SCAI Cell reports Medium 28700933
2014 Rif1 maintains telomere length homeostasis in mouse ESCs by negatively regulating Zscan4 expression through maintaining H3K9me3 levels at subtelomeric regions. Rif1 interacts with and stabilizes H3K9 methylation complexes. Depletion of Rif1 results in telomere hyperrecombination, length heterogeneity, and chromosomal fusions. Rif1 RNAi/gene deletion in ESCs, telomere FISH, ChIP for H3K9me3, Co-immunoprecipitation with H3K9 methyltransferase complex, Zscan4 expression analysis Developmental cell Medium 24735877
2017 Rif1 promotes a repressive chromatin state at endogenous retroviruses (ERVs) in mouse ESCs. Rif1 directly occupies ERV regions, is required for H3K9me3 and H3K27me3 assembly and DNA methylation at ERVs, and interacts with histone methyltransferases to facilitate their recruitment. Loss of Rif1 increases ERV transcription and chromatin accessibility. The HEAT-like domain is essential for this function. RNAi screen, gene deletion, ChIP for repressive marks, ATAC-seq, Co-immunoprecipitation with histone methyltransferases, Rif1 ChIP-seq Nucleic acids research Medium 29040764
2018 In Drosophila, Rif1 is a repressor of DNA replication whose chromatin association is inhibited by Cdk1 activity. At the mid-blastula transition (MBT), as Cdk1 activity drops, Rif1 binds selectively to satellite sequences. Rif1 dissociates from different satellites in an orderly schedule anticipating their replication. Rif1 lacking potential phosphorylation sites fails to dissociate and dominantly prevents completion of replication. Loss of Rif1 shortens post-MBT S phase. Live imaging of Rif1 chromatin association, genetic Rif1 loss-of-function, phosphorylation-site mutant Rif1 expression, Cdc7 depletion epistasis PLoS biology High 29746464
2018 In Drosophila polyploid cells, Rif1 interacts with the SNF2-domain-containing SUUR protein and acts downstream of SUUR to inhibit replication fork progression and control DNA copy number. Rif1 localizes to active replication forks in a partially SUUR-dependent manner. Co-immunoprecipitation (SUUR-Rif1), DNA copy-number analysis, Rif1/SUUR genetic epistasis, Rif1 localization to forks by imaging eLife Medium 30277458
2018 Rif1 telomere-length control in budding yeast requires its PP1 interaction via RVxF/SILK motifs. Rif1-PP1 suppresses telomere lengthening and prevents inappropriate Tel1 kinase recruitment at de novo telomeres, independent of replication timing regulation. Rif1 PP1-interaction mutants (RVxF/SILK), telomere length assay, Tel1 ChIP at telomeres, origin deletion for replication-timing bypass Nucleic acids research Medium 29529242
2018 In budding yeast, Rif1 primarily controls late-replicating origins within 100 kb of a telomere. ChEC-seq detects Rif1 binding directly at late-replicating origins that are targets of its inhibitory action. Abrogation of Rif1 telomere association by mutation of its Rap1-binding module increases Rif1 binding and origin inhibition at chromosome-internal sites. ChEC-seq genome-wide Rif1 binding, replication timing analysis, Rap1-binding module mutants Cell reports Medium 29694906
2018 Budding yeast Rif1 binds to replication origins (in addition to telomeres) in a Rap1-independent manner, associating with both early and late-initiating origins. Rif1 also associates with blocked replication forks under hydroxyurea, and stabilizes recently synthesized DNA at blocked forks. ChIP-seq, DNA combing/fiber analysis, nascent DNA labeling, Rap1-interaction domain mutants EMBO reports Medium 30104203
2018 RIF1 promotes cancer cell survival after multifractionated radiotherapy as the critical effector of 53BP1 in this context. 53BP1-mediated NHEJ and RIF1 activity account for the enhanced survival of cells after multifractionated vs. single-dose radiation. Genetic inhibition of 53BP1/RIF1, clonogenic survival assay after multifractionated vs. single-dose irradiation Nucleic acids research Medium 31822909
2018 RIF1 promotes PP1-AXIN interaction and thereby activates Wnt/β-catenin signaling. RIF1 overexpression in NSCLC cells promotes cell growth, cell cycle progression, and cancer stem cell-like properties through PP1-mediated dephosphorylation of AXIN. PP1 inhibition counteracts RIF1 effects on these phenotypes. Co-immunoprecipitation (RIF1-PP1, PP1-AXIN), siRNA knockdown/overexpression, Wnt reporter assay, xenograft model Cell death & disease Medium 30237512

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2013 A cell cycle-dependent regulatory circuit composed of 53BP1-RIF1 and BRCA1-CtIP controls DNA repair pathway choice. Molecular cell 742 23333306
2013 RIF1 is essential for 53BP1-dependent nonhomologous end joining and suppression of DNA double-strand break resection. Molecular cell 520 23333305
2013 53BP1 regulates DSB repair using Rif1 to control 5' end resection. Science (New York, N.Y.) 503 23306437
1980 A new mouse tumor model system (RIF-1) for comparison of end-point studies. Journal of the National Cancer Institute 430 6928244
2018 53BP1-RIF1-shieldin counteracts DSB resection through CST- and Polα-dependent fill-in. Nature 356 30022158
2013 Rif1 prevents resection of DNA breaks and promotes immunoglobulin class switching. Science (New York, N.Y.) 340 23306439
2013 RIF1 counteracts BRCA1-mediated end resection during DNA repair. The Journal of biological chemistry 230 23486525
2012 Rif1 regulates the replication timing domains on the human genome. The EMBO journal 203 22850674
2012 Mouse Rif1 is a key regulator of the replication-timing programme in mammalian cells. The EMBO journal 198 22850673
2012 Rif1 is a global regulator of timing of replication origin firing in fission yeast. Genes & development 197 22279046
2014 Rif1 controls DNA replication by directing Protein Phosphatase 1 to reverse Cdc7-mediated phosphorylation of the MCM complex. Genes & development 183 24532715
2004 Human Rif1, ortholog of a yeast telomeric protein, is regulated by ATM and 53BP1 and functions in the S-phase checkpoint. Genes & development 169 15342490
2015 Nuclear Architecture Organized by Rif1 Underpins the Replication-Timing Program. Molecular cell 141 26725008
1993 Apoptosis during photodynamic therapy-induced ablation of RIF-1 tumors in C3H mice: electron microscopic, histopathologic and biochemical evidence. Photochemistry and photobiology 141 8309997
2014 Protein phosphatase 1 recruitment by Rif1 regulates DNA replication origin firing by counteracting DDK activity. Cell reports 137 24656819
2014 Rif1 controls DNA replication timing in yeast through the PP1 phosphatase Glc7. Cell reports 133 24685139
2015 Rif1 binds to G quadruplexes and suppresses replication over long distances. Nature structural & molecular biology 128 26436827
2017 Human RIF1 and protein phosphatase 1 stimulate DNA replication origin licensing but suppress origin activation. EMBO reports 112 28077461
2009 Mammalian Rif1 contributes to replication stress survival and homology-directed repair. The Journal of cell biology 112 19948482
2009 Rif1 and rif2 inhibit localization of tel1 to DNA ends. Molecular cell 104 19217405
2014 Rif1 maintains telomere length homeostasis of ESCs by mediating heterochromatin silencing. Developmental cell 101 24735877
2005 Taz1, Rap1 and Rif1 act both interdependently and independently to maintain telomeres. The EMBO journal 101 16096639
2004 Human Rif1 protein binds aberrant telomeres and aligns along anaphase midzone microtubules. The Journal of cell biology 97 15583028
1986 Radiosensitization by cisplatin of RIF1 tumour cells in vitro. International journal of radiation biology and related studies in physics, chemistry, and medicine 97 3490450
2017 Reversal of DDK-Mediated MCM Phosphorylation by Rif1-PP1 Regulates Replication Initiation and Replisome Stability Independently of ATR/Chk1. Cell reports 95 28273463
2019 RIF1 promotes replication fork protection and efficient restart to maintain genome stability. Nature communications 88 31337767
2010 Rif1 provides a new DNA-binding interface for the Bloom syndrome complex to maintain normal replication. The EMBO journal 85 20711169
2013 Rif1 and Rif2 shape telomere function and architecture through multivalent Rap1 interactions. Cell 84 23746845
2015 Rif1 Is Required for Resolution of Ultrafine DNA Bridges in Anaphase to Ensure Genomic Stability. Developmental cell 82 26256213
1980 Response to the RIF-1 tumor in vitro and in C3H/Km mice to X-radiation (cell survival, regrowth delay, and tumor control), chemotherapeutic agents, and activated macrophages. Journal of the National Cancer Institute 79 6928245
1988 Antitumor effects of recombinant human interleukin 1 alpha in RIF-1 and Panc02 solid tumors. Cancer research 76 3262418
2014 Rif1 regulates initiation timing of late replication origins throughout the S. cerevisiae genome. PloS one 72 24879017
2002 Photodynamic therapy with hypericin induces vascular damage and apoptosis in the RIF-1 mouse tumor model. International journal of cancer 72 11857421
2017 ATM and CDK2 control chromatin remodeler CSB to inhibit RIF1 in DSB repair pathway choice. Nature communications 63 29203878
2012 Functional diversification of yeast telomere associated protein, Rif1, in higher eukaryotes. BMC genomics 58 22712556
1981 Enhancing effect of misonidazole on the response of the RIF-1 tumour to cyclophosphamide. British journal of cancer 58 7272187
2013 Replication timing regulation of eukaryotic replicons: Rif1 as a global regulator of replication timing. Trends in genetics : TIG 57 23809990
2018 Rif1 prolongs the embryonic S phase at the Drosophila mid-blastula transition. PLoS biology 56 29746464
2024 Longitudinal profiling identifies co-occurring BRCA1/2 reversions, TP53BP1, RIF1 and PAXIP1 mutations in PARP inhibitor-resistant advanced breast cancer. Annals of oncology : official journal of the European Society for Medical Oncology 55 38244928
2019 Human RIF1-Protein Phosphatase 1 Prevents Degradation and Breakage of Nascent DNA on Replication Stalling. Cell reports 55 31141682
2011 Palmitoylation controls the dynamics of budding-yeast heterochromatin via the telomere-binding protein Rif1. Proceedings of the National Academy of Sciences of the United States of America 55 21844336
2011 Rif1 supports the function of the CST complex in yeast telomere capping. PLoS genetics 54 21437267
1989 Characterization of a cisplatin-resistant subline of murine RIF-1 cells and reversal of drug resistance by hyperthermia. Cancer research 54 2713851
1994 Isolation and identification of fresh tumor-derived endothelial cells from a murine RIF-1 fibrosarcoma. Cancer research 49 8275463
2017 Rif1 promotes a repressive chromatin state to safeguard against endogenous retrovirus activation. Nucleic acids research 48 29040764
2009 Human RIF1 encodes an anti-apoptotic factor required for DNA repair. Carcinogenesis 45 19483192
2019 The RIF1-PP1 Axis Controls Abscission Timing in Human Cells. Current biology : CB 44 30905608
2018 H4K20me2 distinguishes pre-replicative from post-replicative chromatin to appropriately direct DNA repair pathway choice by 53BP1-RIF1-MAD2L2. Cell cycle (Georgetown, Tex.) 44 29160738
2017 Rif1 maintains telomeres and mediates DNA repair by encasing DNA ends. Nature structural & molecular biology 44 28604726
2011 A novel checkpoint and RPA inhibitory pathway regulated by Rif1. PLoS genetics 43 22194703
2017 Mouse Rif1 is a regulatory subunit of protein phosphatase 1 (PP1). Scientific reports 41 28522851
2007 Multispectral quantification of tissue types in a RIF-1 tumor model with histological validation. Part I. Magnetic resonance in medicine 41 17326181
1986 Effect of dexamethasone on vascular function in RIF-1 tumors. Cancer research 40 3708563
2008 yKu70/yKu80 and Rif1 regulate silencing differentially at telomeres in Candida glabrata. Eukaryotic cell 38 18836091
2022 RIF1 acts in DNA repair through phosphopeptide recognition of 53BP1. Molecular cell 36 35216668
2019 Rif1 S-acylation mediates DNA double-strand break repair at the inner nuclear membrane. Nature communications 36 31182712
2014 RIF1: a novel regulatory factor for DNA replication and DNA damage response signaling. DNA repair 36 24462468
2016 Rif1: A Conserved Regulator of DNA Replication and Repair Hijacked by Telomeres in Yeasts. Frontiers in genetics 35 27066066
2018 RIF1 promotes tumor growth and cancer stem cell-like traits in NSCLC by protein phosphatase 1-mediated activation of Wnt/β-catenin signaling. Cell death & disease 34 30237512
1996 13C and 31P NMR investigation of effect of 6-aminonicotinamide on metabolism of RIF-1 tumor cells in vitro. The Journal of biological chemistry 34 8626749
1988 Effect of cyclophosphamide on the pathophysiology of RIF-1 solid tumors. Cancer research 34 3390814
2018 Oligomer formation and G-quadruplex binding by purified murine Rif1 protein, a key organizer of higher-order chromatin architecture. The Journal of biological chemistry 33 29348174
2018 Rif1 Binding and Control of Chromosome-Internal DNA Replication Origins Is Limited by Telomere Sequestration. Cell reports 33 29694906
2018 Rif1 inhibits replication fork progression and controls DNA copy number in Drosophila. eLife 32 30277458
2006 The response of RIF-1 fibrosarcomas to the vascular-disrupting agent ZD6126 assessed by in vivo and ex vivo 1H magnetic resonance spectroscopy. Neoplasia (New York, N.Y.) 31 16867218
2022 H3K4 methylation by SETD1A/BOD1L facilitates RIF1-dependent NHEJ. Molecular cell 29 35439434
2014 Structural and biophysical characterization of murine rif1 C terminus reveals high specificity for DNA cruciform structures. The Journal of biological chemistry 29 24634216
1990 Cytotoxicity of lonidamine alone and in combination with other drugs against murine RIF-1 and human HT1080 cells in vitro. Cancer research 29 2253227
1983 Discrepancies between patterns of potentially lethal damage repair in the RIF-1 tumor system in vitro and in vivo. Radiation research 29 6337381
2016 Rif1 Regulates the Fate of DNA Entanglements during Mitosis. Cell reports 28 27320927
1997 Quantitation of metabolic and radiobiological effects of 6-aminonicotinamide in RIF-1 tumor cells in vitro. Cancer research 28 9307279
2019 Rif1 promotes association of G-quadruplex (G4) by its specific G4 binding and oligomerization activities. Scientific reports 27 31197198
2015 Drosophila Rif1 is an essential gene and controls late developmental events by direct interaction with PP1-87B. Scientific reports 27 26022086
2013 RIF1 in DNA break repair pathway choice. Molecular cell 27 23473603
2005 Application of 23Na MRI to monitor chemotherapeutic response in RIF-1 tumors. Neoplasia (New York, N.Y.) 26 16026645
1994 Potentiation by interleukin 1 alpha of cisplatin and carboplatin antitumor activity: schedule-dependent and pharmacokinetic effects in the RIF-1 tumor model. Cancer research 26 7923169
1993 Vascular morphometry of KHT and RIF-1 murine sarcomas. Radiotherapy and oncology : journal of the European Society for Therapeutic Radiology and Oncology 26 7694322
1982 Sensitivity of normal mouse marrow and RIF-1 tumour to hyperthermia combined with cyclophosphamide or BCNU: a lack of therapeutic gain. British journal of cancer 26 6758828
2018 Budding yeast Rif1 binds to replication origins and protects DNA at blocked replication forks. EMBO reports 25 30104203
2017 Inhibition of RIF1 by SCAI Allows BRCA1-Mediated Repair. Cell reports 25 28700933
2016 Impaired 53BP1/RIF1 DSB mediated end-protection stimulates CtIP-dependent end resection and switches the repair to PARP1-dependent end joining in G1. Oncotarget 25 27494840
2012 Enforcement of a lifespan-sustaining distribution of Sir2 between telomeres, mating-type loci, and rDNA repeats by Rif1. Aging cell 25 23082874
1989 Comparison between tumor pH and cell sensitivity to heat in RIF-1 tumors. Cancer research 25 2736514
2016 Budding Yeast Rif1 Controls Genome Integrity by Inhibiting rDNA Replication. PLoS genetics 24 27820830
2022 RIF1-ASF1-mediated high-order chromatin structure safeguards genome integrity. Nature communications 23 35177609
2007 Multispectral tissue characterization in a RIF-1 tumor model: monitoring the ADC and T2 responses to single-dose radiotherapy. Part II. Magnetic resonance in medicine 23 17326182
1999 The high photoactivity of m-THPC in photodynamic therapy. Unusually strong retention of m-THPC by RIF-1 cells in culture. Photochemistry and photobiology 23 10089829
2022 Rif1-Dependent Control of Replication Timing. Genes 22 35328102
2018 Rif1 acts through Protein Phosphatase 1 but independent of replication timing to suppress telomere extension in budding yeast. Nucleic acids research 22 29529242
2018 Shepherding DNA ends: Rif1 protects telomeres and chromosome breaks. Microbial cell (Graz, Austria) 22 29992129
2007 Monitoring chemotherapeutic response in RIF-1 tumors by single-quantum and triple-quantum-filtered (23)Na MRI, (1)H diffusion-weighted MRI and PET imaging. Magnetic resonance imaging 22 17707164
1993 Synergistic enhancement by interleukin-1 alpha of cisplatin-mediated antitumor activity in RIF-1 tumor-bearing C3H/HeJ mice. Cancer chemotherapy and pharmacology 22 8339383
1986 Cell kill and tumor control after heat treatment with and without vascular occlusion in RIF-1 tumors. Radiation research 22 3704113
2023 An AlphaFold2 map of the 53BP1 pathway identifies a direct SHLD3-RIF1 interaction critical for shieldin activity. EMBO reports 21 37306046
2018 Downregulation of RIF1 Enhances Sensitivity to Platinum-Based Chemotherapy in Epithelial Ovarian Cancer (EOC) by Regulating Nucleotide Excision Repair (NER) Pathway. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 21 29719287
2018 RIF1 promotes human epithelial ovarian cancer growth and progression via activating human telomerase reverse transcriptase expression. Journal of experimental & clinical cancer research : CR 21 30075819
2017 Establishment of expression-state boundaries by Rif1 and Taz1 in fission yeast. Proceedings of the National Academy of Sciences of the United States of America 21 28096402
2014 Saccharomyces cerevisiae Rif1 cooperates with MRX-Sae2 in promoting DNA-end resection. EMBO reports 21 24692507
1990 Effects of hydralazine on the blood flow in RIF-1 tumors and normal tissues of mice. Radiation research 21 2247597
2020 53BP1/RIF1 signaling promotes cell survival after multifractionated radiotherapy. Nucleic acids research 20 31822909

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