Affinage

Showing TERF1 — TRF1 is a alias.

TERF1

Telomeric repeat-binding factor 1 · UniProt P54274

Length
439 aa
Mass
50.2 kDa
Annotated
2026-06-10
100 papers in source corpus 59 papers cited in narrative 59 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 9/9 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

TERF1/TRF1 is a sequence-specific duplex telomere-binding protein and core shelterin component that defines telomere structure, controls telomere length, and ensures faithful replication of telomeric repeats (PMID:9130722, PMID:10669743, PMID:19596237). It recognizes the duplex TTAGGG repeat through a single C-terminal Myb/homeodomain that engages the TAGGGTT site, and dimerizes via an N-terminal TRFH domain so that the homodimer contacts a bipartite recognition element with extreme spatial flexibility, looping intervening DNA and bending the duplex (PMID:9130722, PMID:10523316, PMID:15608617, PMID:11738049). TRF1 dimers pair and synapse telomeric tracts and can engage telomeric nucleosomes by binding unwrapped DNA ends and the histone octamer, remodeling nucleosomal arrays (PMID:9571035, PMID:25999344, PMID:37624885). Within shelterin, TRF1 bridges to TRF2 through TIN2 and, via the TIN2-PIP1-POT1 module, governs POT1 loading onto single-stranded telomeric DNA to transmit telomere-length information and restrict telomerase access; consistent with this, TRF1 overexpression progressively shortens telomeres without altering telomerase activity, and TRF1 loss destabilizes TIN2/TRF2 at telomeres and produces end-to-end fusions and chromosomal instability (PMID:10669743, PMID:12768206, PMID:14559908, PMID:15316005). TRF1 is essential for efficient replication of telomeric repeats: its loss causes a fragile-telomere phenotype, ATR activation, and fork stalling, and it acts upstream of the BLM and RTEL1 helicases and recruits TFIIH to telomeres to resolve replication stress (PMID:19596237, PMID:36229075, PMID:36864251). TRF1 abundance and DNA-binding are tightly tuned by a layered post-translational program—tankyrase-mediated ADP-ribosylation releases it from telomeres and licenses FBX4/SCF-dependent ubiquitination and proteasomal degradation, while phosphorylation by Cdk1-primed Plk1, Nek7, and Aurora kinases controls its mitotic DNA-binding activity, stability, and telomere replication (PMID:12782650, PMID:16275645, PMID:18625707, PMID:28216227, PMID:29040668). Its N-terminal acidic domain restrains the TRFH condensation activity and counteracts TRF2-driven invasion of TERRA into telomeric duplex DNA, suppressing harmful telomeric R-loops and recombination (PMID:22139926, PMID:29358759, PMID:31934863). Beyond telomeres, TRF1 contributes to mitotic chromosome segregation through centromeric Aurora-B recruitment, forms a meiosis-specific heterocomplex with TERB1 that recruits the SUN-KASH and cohesin machinery for chromosome movement, and is an Oct3/4 transcriptional target required for stem cell pluripotency and reprogramming (PMID:23735977, PMID:24413433, PMID:24752893, PMID:29083416). Targeted deletion of mouse Trf1 causes early embryonic lethality independent of telomere length, underscoring an essential cellular role (PMID:12944479).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 1997 High

    Established the fundamental biochemical identity of TRF1 as a sequence-specific duplex telomere-binding protein, defining how it recognizes and architecturally distorts telomeric DNA.

    Evidence Yeast two-hybrid, gel-shift/footprinting, and EM showing homodimeric Myb-domain binding and ~120° bending of TTAGGG DNA; reciprocal assays distinguishing it from TRF2

    PMID:9130722 PMID:9326950

    Open questions at the time
    • Did not resolve atomic-level recognition determinants
    • In vivo functional consequence of DNA bending unaddressed
  2. 1999 High

    Defined the bipartite, spatially flexible recognition mode of the TRF1 dimer, explaining how it accommodates variably spaced telomeric half-sites.

    Evidence SELEX, affinity assays, footprinting, and EM showing two Myb domains contacting two YTAGGGTTR half-sites with looping of intervening DNA

    PMID:10523316

    Open questions at the time
    • Functional role of intervening-DNA looping in vivo not established
  3. 2000 High

    Showed TRF1 is a cis-acting negative regulator of telomere length that limits telomerase access without inhibiting the enzyme itself.

    Evidence Inducible overexpression over >300 doublings with Southern blot length tracking and telomerase activity assays

    PMID:10669743

    Open questions at the time
    • Molecular mechanism transmitting length to the terminus not yet defined at this stage
  4. 2003 High

    Placed TRF1 within shelterin as a length-sensing platform and demonstrated it is genetically essential, linking molecular partners to organismal viability.

    Evidence TRF1-TIN2-PIP1-POT1 co-IP and dominant-negative POT1 epistasis; conditional/germline Trf1 deletion in mouse ES cells and embryos with telomere-independent lethality

    PMID:12768206 PMID:12944479 PMID:14559908

    Open questions at the time
    • Precise signal relayed from duplex TRF1 to ssDNA POT1 not mechanistically resolved
    • Cause of embryonic lethality not attributable to a single molecular defect
  5. 2004 High

    Connected TRF1 to the shelterin bridge and to replication/helicase control, showing it both stabilizes TRF2 via TIN2 and modulates fork progression and BLM activity at telomeres.

    Evidence Multi-method (MS, co-IP, Far-Western, Y2H) TIN2 bridging; in vitro helicase and SV40 replication assays with telomeric substrate specificity

    PMID:15007108 PMID:15229185 PMID:15316005

    Open questions at the time
    • In vivo consequence of TRF1-mediated fork stalling not yet demonstrated
    • How TRF1 toggles between inhibiting and enabling replication unresolved
  6. 2005 High

    Provided high-resolution structural and damage-sensitivity insight into TRF1-DNA recognition, including the chemical features that disrupt binding.

    Evidence X-ray crystallography of the DNA-binding domain on telomeric DNA and quantitative binding assays with oxidatively damaged substrates

    PMID:11738049 PMID:15608617 PMID:15731343

    Open questions at the time
    • Cellular impact of damage-induced TRF1 loss not addressed in these structural/biochemical studies
  7. 2005 High

    Defined the tankyrase-ubiquitin axis controlling TRF1 abundance and telomere occupancy, establishing ordered post-translational regulation.

    Evidence ADP-ribosylation-then-ubiquitination sequence shown by ubiquitination assays and proteasome inhibition; FBX4/SCF-dependent TRF1 ubiquitination with bidirectional length effects

    PMID:12782650 PMID:16275645

    Open questions at the time
    • Signals triggering TRF1 release versus retention in homeostasis not defined
  8. 2008 High

    Revealed mitotic kinase control of TRF1 DNA-binding activity, linking cell-cycle phosphorylation to peak telomere engagement.

    Evidence In vitro kinase assays, mutagenesis (S435), and DNA-binding readouts for Cdk1-primed Plk1 phosphorylation; Xenopus egg extract Plx1-dependent mitotic chromatin association

    PMID:16424898 PMID:18625707

    Open questions at the time
    • Functional purpose of heightened mitotic DNA binding not fully resolved
  9. 2009 High

    Demonstrated that TRF1 is required for replication of telomeric repeats and that its loss triggers replication stress and senescence-associated genome instability in vivo.

    Evidence Conditional KO with single-molecule replication analysis, ATR/ATM-CHK signaling, FISH fragility, and epistasis with BLM/RTEL1; mouse tissue KO with tumorigenesis in p53-null background

    PMID:19596237 PMID:19679647

    Open questions at the time
    • How TRF1 recruits/coordinates helicases mechanistically not yet pinpointed
  10. 2014 High

    Uncovered telomere-independent mitotic and meiotic chromosome-segregation roles for TRF1, broadening it beyond a telomere-capping factor.

    Evidence Telomere-unbound TRF1 rescue of centromeric Aurora-B/segregation defects; TERB1 knockout and co-IP placing TRF1 in meiotic telomere-LINC recruitment

    PMID:24413433 PMID:24752893

    Open questions at the time
    • Mechanism of TRF1 targeting to centromeres unclear
    • Whether mitotic and meiotic roles share a common molecular basis unknown
  11. 2018 High

    Defined the acidic N-terminal domain as the element that suppresses TRF2-driven TERRA R-loop formation and chromatin condensation, assigning a protective genome-integrity function to a previously enigmatic domain.

    Evidence In vitro RNA-invasion and condensation assays with domain deletions plus in vivo acidic-domain-deleted variant inducing telR loops and telomere loss

    PMID:22139926 PMID:29358759 PMID:36142374

    Open questions at the time
    • Quantitative contribution of TRF1 R-loop suppression to overall telomere maintenance not fully delineated
  12. 2020 High

    Showed TRF1 actively suppresses recombination-based telomere maintenance, identifying the proteins and pathway it excludes from telomeres.

    Evidence Quantitative telomere proteomics with siRNA epistasis (SMC5/6, PML, BRCA1) and POLD3-dependent break-induced replication assays

    PMID:31934863

    Open questions at the time
    • How TRF1 mechanistically excludes recombination factors not resolved
  13. 2022 High

    Identified the direct effector through which TRF1 promotes telomere replication, separating this activity from DNA binding via a structure-guided mutant.

    Evidence Separation-of-function TRF1 mutant, co-IP, and TFIIH-subunit knockdown phenocopying TRF1 deletion fragility; PARP1-dependent S-phase modulation of TRF1 and helicase recruitment

    PMID:36229075 PMID:36864251

    Open questions at the time
    • How TFIIH resolves telomeric replication impediments mechanistically unclear
    • Interplay between PARP1 and TFIIH-helicase recruitment not integrated
  14. 2023 High

    Resolved how TRF1 engages telomeric chromatin at the nucleosome level, showing it reads unwrapped nucleosomal DNA and the octamer in a phosphorylation-dependent manner.

    Evidence Cryo-EM of telomeric nucleosome with and without TRF1, with MS phospho-site mapping and binding-surface mutagenesis

    PMID:37624885

    Open questions at the time
    • In vivo importance of nucleosomal-register shift not established

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the many post-translational modifications, partner exchanges, and chromatin states are integrated to switch TRF1 between length control, replication support, recombination suppression, and segregation roles remains unresolved.
  • No unified model coordinating phosphorylation, PARylation, and ubiquitination across the cell cycle
  • Relative in vivo contributions of telomeric versus non-telomeric TRF1 functions not quantified

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 5 GO:0060090 molecular adaptor activity 3 GO:0098772 molecular function regulator activity 3 GO:0003723 RNA binding 2 GO:0042393 histone binding 2
Localization
GO:0000228 nuclear chromosome 4 GO:0005634 nucleus 2 GO:0005730 nucleolus 2 GO:0005856 cytoskeleton 2 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-4839726 Chromatin organization 3 R-HSA-69306 DNA Replication 3 R-HSA-73894 DNA Repair 3 R-HSA-1474165 Reproduction 2 R-HSA-1640170 Cell Cycle 2
Partners
FBX4GNL3LPINX1POT1TERB1TERF2TINF2TNKS
Complex memberships
shelterin

Evidence

Reading pass · 59 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1997 TRF1 binds duplex TTAGGG telomeric DNA as a homodimer, using a single Myb-type helix-turn-helix motif; dimerization is mediated by a conserved N-terminal TRFH domain and both Myb repeats of the dimer are required for stable DNA binding. TRF1 also bends telomeric DNA by approximately -120 degrees. Yeast two-hybrid, gel-shift/DNase I footprinting, electron microscopy, biochemical dimerization assays The EMBO journal High 9130722
1997 TRF1 and TRF2 are two distinct Myb-related proteins that both bind duplex TTAGGG repeats at human telomeres; their dimerization domains do not interact with each other, indicating they exist predominantly as homodimers. TRF1 has an acidic N-terminus while TRF2 has a basic N-terminus. Cloning, in vitro DNA binding assays, indirect immunofluorescence on metaphase chromosomes Nature genetics High 9326950
1998 TRF1 dimers promote parallel pairing and synapsis of telomeric DNA tracts in vitro, forming filamentous structures on long telomeric arrays; mass analysis revealed TRF1 dimers and tetramers; DNA pairing confirmed by ligation assay. Electron microscopy, mass analysis, ligation assay Journal of molecular biology High 9571035
1999 TRF1 homodimers bind a bipartite recognition site consisting of two 5'-YTAGGGTTR-3' half-sites with extreme spatial flexibility (no constraint on distance or orientation between half-sites); the two Myb domains contact both half-sites simultaneously with looping of intervening DNA. SELEX, binding affinity assays, DNase I footprinting, electron microscopy The EMBO journal High 10523316
2000 TRF1 overexpression progressively shortens telomere length without affecting telomerase expression or activity in vitro, consistent with TRF1 acting as a negative regulator of telomerase access in cis (t-loop model). Inducible overexpression, Southern blotting for telomere length, telomerase activity assay Molecular and cellular biology High 10669743
2001 PinX1 interacts with TRF1 (Pin2/TRF1) and binds the telomerase catalytic subunit hTERT to potently inhibit its activity; overexpression shortens telomeres and induces crisis, while PinX1 depletion increases telomerase activity and elongates telomeres. Co-immunoprecipitation, in vitro telomerase inhibition assay, siRNA/overexpression with telomere length measurement Cell High 11701125
2001 ATM kinase co-immunoprecipitates with Pin2/TRF1 and directly phosphorylates it on Ser219 in vitro and in vivo in response to ionizing radiation. Phosphorylation-refractory mutants potentiate apoptosis and mitotic entry, while phosphomimetic mutants suppress these effects, demonstrating ATM phosphorylation negatively regulates Pin2/TRF1 pro-apoptotic activity. Co-immunoprecipitation, in vitro kinase assay, site-directed mutagenesis, functional rescue experiments The Journal of biological chemistry High 11375976
2001 Pin2/TRF1 protein levels are cell-cycle regulated, peaking in G2/M; overexpression induces G2/M accumulation and then apoptosis via caspase-3 activation. Pin2 (shorter isoform) is the major cellular product and forms homo- and heterodimers with TRF1. Cell cycle analysis, co-immunoprecipitation, immunofluorescence, caspase-3 assay Proceedings of the National Academy of Sciences of the United States of America Medium 9391075
2001 Pin2/TRF1 physically interacts with microtubules in a cell cycle-specific (mitosis-specific) manner via its C-terminal domain, localizes to the mitotic spindle, and promotes microtubule polymerization in vitro. Co-immunoprecipitation, indirect immunofluorescence, in vitro microtubule polymerization assay, domain deletion mapping Current biology : CB Medium 11591318
2001 SALL1 (mutated in Townes-Brocks syndrome) interacts with PIN2/TRF1; the interaction was identified by yeast two-hybrid and confirmed by GST pulldown; SALL1 colocalizes with heterochromatin and pericentromeric regions. Yeast two-hybrid, GST pulldown, co-localization by fluorescence microscopy Human molecular genetics Medium 11751684
2002 Human TRF1 contains an RXXPDG motif that mediates direct binding to the ankyrin-repeat domain of tankyrase-1 and tankyrase-2; mouse TRF1 lacks this motif and does not bind tankyrase. This motif is shared with IRAP and TAB182. Yeast two-hybrid, mutagenesis, binding assays The Journal of biological chemistry High 12080061
2002 Pin2/TRF1 interacts with EB1 (a microtubule plus end-binding protein) in vitro and in vivo, co-localizes with EB1 at the mitotic spindle; EB1 inhibits Pin2/TRF1-mediated microtubule polymerization in vitro. Inhibition of Pin2/TRF1 in ATM-deficient cells restores mitotic spindle defects. Yeast two-hybrid, co-immunoprecipitation, in vitro microtubule polymerization assay, dominant-negative rescue in A-T cells FEBS letters Medium 11943150
2003 TRF1 is degraded by ubiquitin-mediated proteasomal proteolysis after ADP-ribosylation by tankyrase 1 releases it from telomeres; only the telomere-unbound form of TRF1 is ubiquitinated, demonstrating sequential post-translational modification (ADP-ribosylation then ubiquitination) regulates telomere access. Co-immunoprecipitation, ubiquitination assay, proteasome inhibitor experiments, tankyrase 1 overexpression Genes & development High 12782650
2003 Targeted deletion of mouse Trf1 exon 1 causes embryonic lethality at day 5-6 post coitus with severe growth defect of inner cell mass and apoptosis; this lethality is independent of telomere elongation by telomerase and p53 deficiency only briefly delays it. Gene targeting (Trf1 knockout mice), histological analysis, blastocyst culture, telomere length measurement Molecular and cellular biology High 12944479
2003 The TRF1 complex (containing TRF1, TIN2, PIP1, POT1) interacts with POT1 to regulate POT1 loading onto single-stranded telomeric DNA in a telomere-length-dependent manner, transmitting length information to the telomere terminus where telomerase is regulated. A POT1 mutant lacking the DNA-binding domain abrogates TRF1-mediated length control. Co-immunoprecipitation, dominant-negative POT1 overexpression, telomere length measurement Nature High 12768206
2003 TRF1 deletion in mouse embryonic stem cells leads to loss of telomeric association of TIN2 and decreased TRF2 telomere association, chromosomal instability, and end-to-end fusions with detectable telomere signals, demonstrating TRF1 is required for functional telomere structure. Conditional gene deletion, immunofluorescence, metaphase chromosome analysis The Journal of biological chemistry High 14559908
2004 TIN2 directly interacts with both TRF1 and TRF2 simultaneously (confirmed by co-IP, Far-Western, two-hybrid), linking the two telomeric complexes; TIN2 depletion reduces TRF2 and hRap1 at telomeres, demonstrating TIN2 stabilizes TRF2 on telomeres through cooperative binding. Mass spectrometry, co-immunoprecipitation, Far-Western assay, yeast two-hybrid, siRNA knockdown The Journal of biological chemistry High 15316005
2004 TRF1 and TRF2 inhibit BLM helicase unwinding activity in vitro: TRF1 specifically inhibits BLM unwinding of telomeric (but not non-telomeric) substrates, while TRF2 stimulates BLM unwinding of both; TRF1 and TRF2 co-complex with BLM. Co-immunoprecipitation, in vitro helicase unwinding assay with telomeric and non-telomeric substrates Human molecular genetics High 15229185
2004 TRF1 (and TRF2) stall replication fork progression at telomeric repeats in vitro using an SV40-based replication system; TRF1 overexpression in HeLa cells increases 4N DNA content and overlap of replication foci with telomere signals. In vitro SV40 replication system, flow cytometry, immunofluorescence of replication foci and telomeres Nucleic acids research High 15007108
2005 A single 8-oxo-guanine lesion in telomeric DNA reduces TRF1 (and TRF2) binding by at least 50%; multiple lesions or base-excision repair intermediates (abasic sites, single-nucleotide gaps) further disrupt TRF1 binding, showing oxidative DNA damage impairs TRF1-telomere association. In vitro DNA binding assays with defined oxidatively damaged telomeric substrates Nucleic acids research High 15731343
2005 X-ray crystal structures of TRF1 and TRF2 DNA-binding domains (Dbds) in complex with telomeric DNA at 2.0 and 1.8 Å resolution show recognition of the TAGGGTT site via a homeodomain; both Myb motifs adopt similar folds with water-mediated contacts; TRF1 uses an arginine to recognize the TT sequence in the minor groove (vs. lysine in TRF2). X-ray crystallography at 2.0 Å (TRF1) and 1.8 Å (TRF2) EMBO reports High 15608617
2005 FBX4 (PinX3), an F-box protein and SCF ubiquitin ligase subunit, interacts with Pin2/TRF1 isoforms and promotes their ubiquitination in vitro and in vivo; FBX4 overexpression reduces endogenous TRF1 and causes telomere elongation, while FBX4 RNAi stabilizes TRF1 and promotes telomere shortening. Yeast two-hybrid, co-immunoprecipitation, in vitro ubiquitination assay, RNAi knockdown, telomere length measurement The Journal of biological chemistry High 16275645
2001 NMR structure of the TRF1 DNA-binding domain in complex with telomeric DNA shows three helices; the third helix recognizes TAGGG in the major groove and the N-terminal arm interacts with TT in the minor groove; a single Myb domain is sufficient for sequence-specific recognition of AGGGTT. NMR structure determination Structure (London, England : 1993) High 11738049
2006 TRF1 binds to telomeric binding sites within nucleosomes, forming a ternary TRF1-DNA-nucleosome complex; binding is orientation-dependent (dependent on position of binding site on nucleosome surface) and causes nucleosome structural alterations without histone dissociation. In vitro reconstitution of TRF1 binding to telomeric nucleosomes, DNase I footprinting, electrophoretic mobility shift Journal of molecular biology Medium 16756990
2007 ATM-mediated phosphorylation of TRF1 impairs its telomeric DNA binding in vitro; MRE11-RAD50-NBS1 (MRN) complex is required for TRF1 phosphorylation by ATM, and this phosphorylation releases TRF1 from telomeres to promote telomerase access. RAD50 overexpression at telomeres downregulates TRF1 telomeric association. In vitro kinase/DNA binding assay, siRNA knockdown of MRN components, ATM inhibition, chromatin immunoprecipitation Nature structural & molecular biology High 17070170
2008 Polo-like kinase 1 (Plk1) associates with TRF1 and phosphorylates it at Ser-435 in vivo; Cdk1 serves as a priming kinase creating the Plk1 docking site. Plk1 phosphorylation dramatically increases TRF1 telomeric DNA binding activity, which peaks during mitosis. Co-immunoprecipitation, in vitro kinase assay, site-directed mutagenesis, cell cycle analysis, telomeric DNA binding assay The Journal of biological chemistry High 18625707
2006 Xenopus TRF1 (xTRF1) associates with telomere chromatin specifically in mitotic Xenopus egg extracts and dissociates upon mitotic exit; this cell-cycle-dependent chromatin association requires both the TRFH domain and linker region, and is mediated by Polo-like kinase (Plx1) phosphorylation of xTRF1. Xenopus egg extract biochemistry, immunodepletion of Plx1, in vitro Plx1 kinase assay, telomere chromatin fractionation The EMBO journal High 16424898
2009 TRF1 is required for efficient replication of TTAGGG repeat arrays; TRF1 deletion (via conditional gene deletion) causes telomere fragility (fragile-site phenotype), ATR activation in S phase, and fork stalling at telomeres as shown by single-molecule analysis. BLM and RTEL1 helicases are epistatic to TRF1 in suppressing fragile telomeres. Conditional gene deletion, single-molecule replication analysis, metaphase FISH, ATR kinase assays, epistasis with BLM/RTEL1 Cell High 19596237
2009 Conditional deletion of TRF1 in mouse embryonic fibroblasts causes rapid senescence with abundant telomeric γ-H2AX foci and activation of ATM/ATR-CHK1/CHK2; abrogation of p53/RB pathways bypasses senescence but causes chromosomal instability including sister chromatid fusions and multitelomeric signals (fragile sites); TRF1 deletion in epithelial tissues causes perinatal lethality and cancer in p53-deficient mice. Conditional mouse knockout, flow cytometry, immunofluorescence, metaphase FISH, ATM/ATR inhibition, tumor formation assay Genes & development High 19679647
2009 GNL3L (a nucleolar GTP-binding protein) interacts with TRF1 in the nucleoplasm, promotes TRF1 homodimerization, enhances TRF1 telomeric association, prevents PML body recruitment of TRF1, and stabilizes TRF1 by inhibiting its ubiquitylation and FBX4 binding. GNL3L-mediated TRF1 stabilization drives the mitotic increase of TRF1 and promotes metaphase-to-anaphase transition. Co-immunoprecipitation, FRAP, ubiquitination assay, siRNA knockdown, cell cycle analysis The Journal of cell biology High 19487455
2009 PML3 isoform directly interacts with TRF1 (identified as a new TRF1-binding protein), recruits TRF1 to PML bodies in U2OS cells, and is required for assembly of ALT-associated PML bodies (APBs); depletion of PML3 by siRNA inhibits recruitment of both TRF1 and TRF2 to APBs. Co-immunoprecipitation, siRNA knockdown, immunofluorescence co-localization Cancer letters Medium 19900757
2010 Aurora-A overexpression-induced mitotic defects (chromosome misalignment, cytokinetic failure, tetraploidization) depend on TRF1; Aurora-A binds and phosphorylates TRF1, and a non-phosphorylatable TRF1 mutant fails to restore Aurora-A-induced mitotic abnormalities. siRNA knockdown, site-directed mutagenesis, co-immunoprecipitation, in vitro kinase assay, live cell imaging Cancer research High 20160025
2011 The N-terminal acidic domain of TRF1 represses the ability of the TRFH domain to condense telomeric DNA (an activity intrinsic to the isolated TRFH domain); TERRA-like RNA binding to the N-terminus of TRF2 similarly represses TRF2-mediated condensation. In vitro DNA condensation assays, atomic force microscopy, domain deletion/truncation biochemistry Nucleic acids research High 22139926
2011 Nucleostemin (NS) binds TRF1 directly but competitively with GNL3L through two separate TRF1 domains; NS prevents TRF1 dimerization and shortens TRF1 dynamic association with telomeres (measured by FRAP) without reducing total telomere-bound TRF1, and protects against telomere dysfunction foci. Co-immunoprecipitation, FRAP, domain deletion mapping, immunofluorescence Journal of cell science Medium 22045740
2013 TRF1 is a direct transcriptional target of Oct3/4; TRF1 is essential for both induction and maintenance of induced pluripotent stem cell (iPSC) identity; eGFP-TRF1 expression is highest in adult stem cell compartments. Knock-in reporter mouse (eGFP-TRF1), TRF1 deletion in reprogramming assay, ChIP for Oct3/4 at TRF1 promoter Nature communications Medium 23735977
2013 Single-molecule fluorescence imaging shows TRF1 uses limited 1D diffusion to search for telomeric DNA (in contrast to TRF2 which uses extensive 1D sliding via its basic domain); both TRF proteins show transient binding (~9-17 s) and slow 1D diffusion at telomeric sites, proposed to facilitate shelterin complex assembly. Single-molecule fluorescence imaging of quantum dot-labeled proteins on DNA tightropes Nucleic acids research High 24271387
2013 Cdk1-phosphorylated TRF1 (pT371-TRF1) is recruited to sites of non-telomeric DNA double-strand breaks in an ATM- and MRN-dependent manner; depletion of TRF1 or the T371A mutation impairs DNA end resection, homologous recombination repair, and G2/M checkpoint activation. Immunofluorescence of damage-induced foci, siRNA knockdown, site-directed mutagenesis, HR reporter assay Nucleic acids research Medium 23997120
2014 TERB1, a meiosis-specific Myb-domain protein, forms a heterocomplex with TRF1 at meiotic telomeres, binds telomeric DNA, recruits SUN-KASH complex and cohesin to telomeres, and is required for meiotic chromosomal movement, homologue pairing, and synapsis in mice. Co-immunoprecipitation, Terb1 knockout mice, meiotic chromosome spreads, telomere FISH, SUN-KASH localization Nature cell biology High 24413433
2014 TRF1 is required for centromeric recruitment of Aurora-B; TRF1 depletion abolishes centromeric Aurora-B, loosens sister centromere cohesion, and causes merotelic attachments, lagging chromosomes, micronuclei, and aneuploidy. A telomere-unbound TRF1 mutant rescues these defects, indicating this role is telomere-independent. siRNA knockdown, overexpression of telomere-unbound mutant, immunofluorescence, chromosome segregation analysis Molecular and cellular biology High 24752893
2015 TRF1 binds efficiently to telomeric DNA in nucleosomal arrays and remodels telomeric nucleosomal arrays, while TRF2 binding is strongly hampered by nucleosomes; the differential behavior partly depends on the divergent N-terminal domains interacting with histone tails. In vitro reconstitution of telomeric nucleosomal arrays, binding assays, domain deletion/histone tail interaction assays Nucleic acids research High 25999344
2016 SA1 (cohesin subunit) uses 1D diffusion on DNA and subdiffusive sliding at telomeric regions; TRF1 tethers SA1 to telomeric regions; together SA1 and TRF1 form longer DNA-DNA pairing tracts than TRF1 alone, supporting a model where TRF1-SA1 interaction mediates sister telomere cohesion. Single-molecule fluorescence imaging, atomic force microscopy, co-immunoprecipitation Nucleic acids research Medium 27298259
2016 Cdk-dependent phosphorylation of TRF1 at T371 is required for recruitment of TRF1 to ALT-associated PML bodies (APBs) independently of telomeric DNA binding; this (pT371)TRF1 pool is required for C-circle production (ALT hallmark) and APB formation, dependent on ATM, Mre11, and BRCA1. Phospho-specific antibodies, siRNA knockdown, site-directed mutagenesis, immunofluorescence, C-circle assay Journal of cell science Medium 27185864
2017 Nek7 kinase is recruited to telomeres after oxidative damage in an ATM-dependent manner and phosphorylates TRF1 on Ser114, preventing TRF1 binding to FBX4 E3 ligase and thereby preventing proteasomal degradation of TRF1; Nek7 deficiency leads to telomere aberrations and augmented cell death upon oxidative telomeric DNA damage. In vitro kinase assay, co-immunoprecipitation, site-directed mutagenesis (S114A), siRNA knockdown, mass spectrometry phosphosite identification Molecular cell High 28216227
2017 Aurora Kinase B (AURKB) localizes to telomeres in mouse ES cells, interacts with TERF1, and phosphorylates TERF1 on Serine 404 in vitro; S404 phosphomimetic and phospho-dead mutants cause fragile telomere formation in vivo. Co-localization, co-immunoprecipitation, in vitro kinase assay, site-directed mutagenesis, telomere FISH Nucleic acids research Medium 29040668
2017 Tankyrase1 (TNKS1) is recruited to damaged telomeres via TRF1 interaction (through TRF1's tankyrase-binding motif, TBM) and PARylates TRF1 after telomeric DNA damage; TNKS1-mediated TRF1 PARylation facilitates recruitment of XRCC1 and polymerase β for single-strand break repair at telomeres. Mutation of the TBM on TRF1 disrupts TNKS1 recruitment and sensitizes cells to telomeric oxidative damage. Co-immunoprecipitation, site-directed mutagenesis of TBM, TNKS inhibition, immunofluorescence of repair factor recruitment Nucleic acids research Medium 28160604
2018 TRF2 stimulates invasion of TERRA-like RNA into telomeric dsDNA via its N-terminal basic domain, forming telomeric R loops (telR loops); TRF1 counteracts this via its N-terminal acidic domain but does not block ssDNA invasion. In vivo, TRF1 depletion or replacement with an acidic-domain-deleted variant induces TRF2-dependent telR loop formation and telomere loss. In vitro RNA invasion assay, domain deletion mutagenesis, siRNA knockdown, telomere FISH, R-loop detection Nature structural & molecular biology High 29358759
2017 Crystal structure of the TERB1 TRF1-binding motif in complex with the TRF1 TRFH domain reveals the structural basis of TERB1-TRF1 interaction; a point mutation specifically disrupting this interaction in mice causes male-specific infertility by arresting spermatocytes at zygotene-early pachytene and failing X-Y pairing at PAR. X-ray crystallography, knock-in point mutant mice, meiotic chromosome spreads, FISH Nature structural & molecular biology High 29083416
2019 TRF1 regulates genome-wide polycomb (PRC2) binding and H3K27me3 marks at pluripotency genes through TERRA; TRF1 abrogation increases TERRA levels and TERRA binding to pluripotency and differentiation genes, coinciding with loss of naïve pluripotency. TRF1 deletion in mouse ES cells, RNA immunoprecipitation for TERRA-chromatin binding, ChIP-seq for H3K27me3 and PRC2, TERRA level measurement eLife Medium 31426913
2020 TRF1 suppresses accumulation of PML protein, BRCA1, and the SMC5/6 complex at telomeres; TRF1 loss leads to increased homologous recombination, TERRA transcription, and POLD3-dependent break-induced replication (BIR) at telomeres. SMC5 is epistatic to TRF1 in suppressing telomere recombination. Telomere proteomics (quantitative MS), siRNA knockdown epistasis, TERRA measurement, BIR assay with POLD3 depletion eLife High 31934863
2022 TRF1 recruits TFIIH (transcription/NER factor) to telomeres to promote efficient telomere replication; Helix2 of TRFH and Helix1 of the Myb domain are required for this interaction. A separation-of-function mutant (TRF1-E83K/LW-TI) fails to co-immunoprecipitate TFIIH and causes severe telomere replication defects; loss of TFIIH subunits (but no other NER factors) phenocopies TRF1 deletion telomere fragility. Site-directed mutagenesis, co-immunoprecipitation, siRNA knockdown of TFIIH subunits, telomere fragility assay, separation-of-function genetic analysis Genes & development High 36229075
2022 TERRA binding to both TRF1 and TRF2 requires TERRA to form G-quadruplex structures; the TRF2 B domain cluster of arginines mediates TERRA binding; TRF1's N-terminal glutamate patch (A domain) inhibits TERRA-TRF2-B domain complex formation. Mouse TRF1 A domain lacks this inhibitory activity. In vitro RNA-protein binding assays, G-quadruplex disruption experiments, mutagenesis of TERRA-binding domains International journal of molecular sciences Medium 36142374
2023 Cryo-EM structures of a human telomeric nucleosome unbound and bound to TRF1 reveal that TRF1 binds unwrapped nucleosomal DNA ends by engaging both nucleosomal DNA and the histone octamer; TRF1 binding shifts nucleosomal DNA register by 1 bp; phosphorylation of the TRF1 C terminus and a noncanonical DNA binding surface are critical for nucleosomal recognition. Cryo-electron microscopy, mass spectrometry phosphorylation site identification, mutagenesis of binding surfaces Science advances High 37624885
2023 PARP1 interacts with and PARylates TRF1 in S phase, modifying TRF1 DNA affinity; PARP1 inhibition impairs dynamic TRF1 association with replicating telomeres and reduces BrdU incorporation at telomeres; PARP1 inhibition also reduces WRN and BLM helicase recruitment to TRF1-containing complexes during S phase, causing telomere fragility. Co-immunoprecipitation, in vitro PARylation assay, PARP1 inhibitor and siRNA, BrdU incorporation at telomeres, telomere fragility analysis Communications biology Medium 36864251
2021 Naked mole rat TRF1, when substituted for mouse TRF1 in cellular models, supports maximal glycolytic capacity under hypoxia and shows increased nuclear localization and telomere association under low oxygen; this metabolic gain of function maps to specific amino acid changes in the homodimerization domain. Species substitution cellular model, glycolytic capacity assays, immunofluorescence of nuclear/telomere localization, homodimerization domain mutagenesis Science advances Medium 33608273
2006 Nucleostemin (NS) interacts with TRF1 and enhances TRF1 protein degradation (but not ubiquitination) in mouse embryonic fibroblasts; NS loss leads to premature senescence associated with altered TRF1 levels. Co-immunoprecipitation, NS knockout and transgenic mice, Western blot for TRF1 protein levels and ubiquitination Molecular and cellular biology Medium 17000763
2009 PinX1 overexpression forces endogenous TRF1 accumulation in the nucleolus and upregulates TRF1 binding to telomeres; nucleolar PinX1 interacts with TRF1 but does not itself bind telomeres; in ALT cells (WI-38 VA-13), this nucleolar TRF1 accumulation by PinX1 is absent. Overexpression/co-localization by fluorescence microscopy, ChIP, domain deletion analysis Journal of molecular biology Medium 19265708
2012 PinX1 localizes to telomeres primarily at mitosis; PinX1 knockdown causes delayed mitotic entry and reduced TRF1 accumulation on telomeres during mitosis, suggesting PinX1 stabilizes TRF1 at telomeres to promote M-phase entry. siRNA knockdown, cell cycle synchronization, co-immunoprecipitation, immunofluorescence Molecular and cellular biology Medium 22331467
2019 NCAPH2 (condensin II subunit) co-immunoprecipitates with TRF1 and co-localizes at telomeres; NCAPH2 depletion causes ATR-dependent DNA damage foci at telomeres, fragile telomere phenotype, and apparent sister-telomere fusions. Co-immunoprecipitation, immunofluorescence co-localization, siRNA knockdown, telomere FISH Journal of cellular physiology Medium 31026066
2014 TRF1 promotes Rad51-mediated telomeric displacement loop (D-loop) formation in vitro, which may facilitate HR-mediated replication fork restart at telomeres. In vitro fluorescent D-loop assay with Cy3-labeled telomeric substrates and plasmid templates Biochemistry Medium 25115914

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2009 Mammalian telomeres resemble fragile sites and require TRF1 for efficient replication. Cell 830 19596237
1997 Human telomeres contain two distinct Myb-related proteins, TRF1 and TRF2. Nature genetics 782 9326950
2000 Control of human telomere length by TRF1 and TRF2. Molecular and cellular biology 612 10669743
2003 POT1 as a terminal transducer of TRF1 telomere length control. Nature 550 12768206
2008 Differential degradation of PIN2 auxin efflux carrier by retromer-dependent vacuolar targeting. Proceedings of the National Academy of Sciences of the United States of America 322 19004783
2009 Increased telomere fragility and fusions resulting from TRF1 deficiency lead to degenerative pathologies and increased cancer in mice. Genes & development 320 19679647
1997 TRF1 is a dimer and bends telomeric DNA. The EMBO journal 272 9130722
2001 The Pin2/TRF1-interacting protein PinX1 is a potent telomerase inhibitor. Cell 266 11701125
2004 TIN2 binds TRF1 and TRF2 simultaneously and stabilizes the TRF2 complex on telomeres. The Journal of biological chemistry 260 15316005
2005 Oxidative damage in telomeric DNA disrupts recognition by TRF1 and TRF2. Nucleic acids research 226 15731343
2008 Endosidin1 defines a compartment involved in endocytosis of the brassinosteroid receptor BRI1 and the auxin transporters PIN2 and AUX1. Proceedings of the National Academy of Sciences of the United States of America 192 18550817
2003 TRF1 is degraded by ubiquitin-mediated proteolysis after release from telomeres. Genes & development 185 12782650
2005 How the human telomeric proteins TRF1 and TRF2 recognize telomeric DNA: a view from high-resolution crystal structures. EMBO reports 181 15608617
1999 TRF1 binds a bipartite telomeric site with extreme spatial flexibility. The EMBO journal 173 10523316
2014 The TRF1-binding protein TERB1 promotes chromosome movement and telomere rigidity in meiosis. Nature cell biology 151 24413433
2003 Targeted deletion reveals an essential function for the telomere length regulator Trf1. Molecular and cellular biology 139 12944479
2004 Association and regulation of the BLM helicase by the telomere proteins TRF1 and TRF2. Human molecular genetics 134 15229185
2002 Identification of a tankyrase-binding motif shared by IRAP, TAB182, and human TRF1 but not mouse TRF1. NuMA contains this RXXPDG motif and is a novel tankyrase partner. The Journal of biological chemistry 131 12080061
1998 TRF1 promotes parallel pairing of telomeric tracts in vitro. Journal of molecular biology 119 9571035
2005 Comparison between TRF2 and TRF1 of their telomeric DNA-bound structures and DNA-binding activities. Protein science : a publication of the Protein Society 108 15608118
2002 Tankyrase-2 oligomerizes with tankyrase-1 and binds to both TRF1 (telomere-repeat-binding factor 1) and IRAP (insulin-responsive aminopeptidase). The Biochemical journal 102 11802774
1997 TRF1, a mammalian telomeric protein. Trends in genetics : TIG 98 9009844
2007 MRE11-RAD50-NBS1 and ATM function as co-mediators of TRF1 in telomere length control. Nature structural & molecular biology 94 17694070
2003 Importance of TRF1 for functional telomere structure. The Journal of biological chemistry 93 14559908
2006 Nucleostemin delays cellular senescence and negatively regulates TRF1 protein stability. Molecular and cellular biology 92 17000763
2001 Telomeric protein Pin2/TRF1 as an important ATM target in response to double strand DNA breaks. The Journal of biological chemistry 92 11375976
2000 Promoter-selective properties of the TBP-related factor TRF1. Science (New York, N.Y.) 92 10797011
2004 Telomere-bound TRF1 and TRF2 stall the replication fork at telomeric repeats. Nucleic acids research 91 15007108
2005 The F-box protein FBX4 targets PIN2/TRF1 for ubiquitin-mediated degradation and regulates telomere maintenance. The Journal of biological chemistry 89 16275645
2004 Tomato TERF1 modulates ethylene response and enhances osmotic stress tolerance by activating expression of downstream genes. FEBS letters 79 15327984
1997 Characterization and cell cycle regulation of the related human telomeric proteins Pin2 and TRF1 suggest a role in mitosis. Proceedings of the National Academy of Sciences of the United States of America 79 9391075
2001 SALL1, the gene mutated in Townes-Brocks syndrome, encodes a transcriptional repressor which interacts with TRF1/PIN2 and localizes to pericentromeric heterochromatin. Human molecular genetics 77 11751684
2013 TRF1 is a stem cell marker and is essential for the generation of induced pluripotent stem cells. Nature communications 71 23735977
2001 Solution structure of a telomeric DNA complex of human TRF1. Structure (London, England : 1993) 71 11738049
2009 TRF1 controls telomere length and mitotic fidelity in epithelial homeostasis. Molecular and cellular biology 70 19124610
2001 Expression of telomeric repeat binding factor 1 and 2 and TRF1-interacting nuclear protein 2 in human gastric carcinomas. International journal of oncology 67 11494028
2018 TRF1 participates in chromosome end protection by averting TRF2-dependent telomeric R loops. Nature structural & molecular biology 65 29358759
2011 The N-terminal domains of TRF1 and TRF2 regulate their ability to condense telomeric DNA. Nucleic acids research 64 22139926
2017 Inhibition of TRF1 Telomere Protein Impairs Tumor Initiation and Progression in Glioblastoma Mouse Models and Patient-Derived Xenografts. Cancer cell 61 29136505
2010 Expression of TRF1, TRF2, TIN2, TERT, KU70, and BRCA1 proteins is associated with telomere shortening and may contribute to multistage carcinogenesis of gastric cancer. Journal of cancer research and clinical oncology 61 20127252
2003 Telomere repeat binding factors TRF1, TRF2, and hRAP1 modulate replication of Epstein-Barr virus OriP. Journal of virology 61 14581536
2013 TRF1 and TRF2 use different mechanisms to find telomeric DNA but share a novel mechanism to search for protein partners at telomeres. Nucleic acids research 59 24271387
2012 Post-translational modifications of TRF1 and TRF2 and their roles in telomere maintenance. Mechanisms of ageing and development 59 22634377
2001 A critical role for Pin2/TRF1 in ATM-dependent regulation. Inhibition of Pin2/TRF1 function complements telomere shortening, radiosensitivity, and the G(2)/M checkpoint defect of ataxia-telangiectasia cells. The Journal of biological chemistry 58 11744712
2001 A specific interaction between the telomeric protein Pin2/TRF1 and the mitotic spindle. Current biology : CB 55 11591318
2015 Age intrinsic loss of telomere protection via TRF1 reduction in endothelial cells. Biochimica et biophysica acta 53 26658719
2017 Nek7 Protects Telomeres from Oxidative DNA Damage by Phosphorylation and Stabilization of TRF1. Molecular cell 52 28216227
2001 Telomeric protein Pin2/TRF1 induces mitotic entry and apoptosis in cells with short telomeres and is down-regulated in human breast tumors. Oncogene 52 11313893
2018 Mechanistic framework for cell-intrinsic re-establishment of PIN2 polarity after cell division. Nature plants 48 30518833
2008 Expression of TERF1 in rice regulates expression of stress-responsive genes and enhances tolerance to drought and high-salinity. Plant cell reports 48 18777179
2021 Sphingolipids mediate polar sorting of PIN2 through phosphoinositide consumption at the trans-Golgi network. Nature communications 47 34257291
2009 GNL3L stabilizes the TRF1 complex and promotes mitotic transition. The Journal of cell biology 46 19487455
2019 TERRA regulate the transcriptional landscape of pluripotent cells through TRF1-dependent recruitment of PRC2. eLife 45 31426913
2016 Ethylene Response Factor TERF1, Regulated by ETHYLENE-INSENSITIVE3-like Factors, Functions in Reactive Oxygen Species (ROS) Scavenging in Tobacco (Nicotiana tabacum L.). Scientific reports 45 27435661
2016 Effects of Auxins on PIN-FORMED2 (PIN2) Dynamics Are Not Mediated by Inhibiting PIN2 Endocytosis. Plant physiology 43 27506239
2017 Telomeric TERB1-TRF1 interaction is crucial for male meiosis. Nature structural & molecular biology 41 29083416
2008 Plk1 phosphorylation of TRF1 is essential for its binding to telomeres. The Journal of biological chemistry 40 18625707
2006 Novel TRF1/BRF target genes revealed by genome-wide analysis of Drosophila Pol III transcription. The EMBO journal 39 17170711
2005 Expressing TERF1 in tobacco enhances drought tolerance and abscisic acid sensitivity during seedling development. Planta 39 15871029
2017 Gene therapy with the TRF1 telomere gene rescues decreased TRF1 levels with aging and prolongs mouse health span. Aging cell 33 28944611
2015 TRF1 and TRF2 binding to telomeres is modulated by nucleosomal organization. Nucleic acids research 33 25999344
2020 TRF1 averts chromatin remodelling, recombination and replication dependent-break induced replication at mouse telomeres. eLife 32 31934863
2003 Role of Pin2/TRF1 in telomere maintenance and cell cycle control. Journal of cellular biochemistry 31 12682905
2002 Involvement of the telomeric protein Pin2/TRF1 in the regulation of the mitotic spindle. FEBS letters 31 11943150
2014 TRF1 ensures the centromeric function of Aurora-B and proper chromosome segregation. Molecular and cellular biology 30 24752893
2010 TRF1 mediates mitotic abnormalities induced by Aurora-A overexpression. Cancer research 29 20160025
2006 The human telomeric protein TRF1 specifically recognizes nucleosomal binding sites and alters nucleosome structure. Journal of molecular biology 29 16756990
2001 Impaired telomere regulation mechanism by TRF1 (telomere-binding protein), but not TRF2 expression, in acute leukemia cells. International journal of oncology 29 11179492
2021 Nitrate triggered phosphoproteome changes and a PIN2 phosphosite modulating root system architecture. EMBO reports 28 34357701
2017 Tankyrase1-mediated poly(ADP-ribosyl)ation of TRF1 maintains cell survival after telomeric DNA damage. Nucleic acids research 27 28160604
2009 PML3 interacts with TRF1 and is essential for ALT-associated PML bodies assembly in U2OS cells. Cancer letters 26 19900757
2023 PARP1 allows proper telomere replication through TRF1 poly (ADP-ribosyl)ation and helicase recruitment. Communications biology 25 36864251
2016 Functional interplay between SA1 and TRF1 in telomeric DNA binding and DNA-DNA pairing. Nucleic acids research 25 27298259
2018 TRF1 as a major contributor for telomeres' shortening in the context of obesity. Free radical biology & medicine 23 30268887
2011 Nucleostemin inhibits TRF1 dimerization and shortens its dynamic association with the telomere. Journal of cell science 23 22045740
2019 Condensin II subunit NCAPH2 associates with shelterin protein TRF1 and is required for telomere stability. Journal of cellular physiology 21 31026066
2010 Altered mRNA expression of telomere binding proteins (TPP1, POT1, RAP1, TRF1 and TRF2) in ulcerative colitis and Crohn's disease. Digestive and liver disease : official journal of the Italian Society of Gastroenterology and the Italian Association for the Study of the Liver 21 20061197
2009 Human PinX1 mediates TRF1 accumulation in nucleolus and enhances TRF1 binding to telomeres. Journal of molecular biology 21 19265708
2006 Cell-cycle-dependent Xenopus TRF1 recruitment to telomere chromatin regulated by Polo-like kinase. The EMBO journal 21 16424898
2022 TRF1 uses a noncanonical function of TFIIH to promote telomere replication. Genes & development 20 36229075
2017 Excessive Cellular S-nitrosothiol Impairs Endocytosis of Auxin Efflux Transporter PIN2. Frontiers in plant science 20 29218054
2017 Aurora Kinase B, a novel regulator of TERF1 binding and telomeric integrity. Nucleic acids research 19 29040668
2015 Tricho- and atrichoblast cell files show distinct PIN2 auxin efflux carrier exploitations and are jointly required for defined auxin-dependent root organ growth. Journal of experimental botany 19 26041320
2013 Phosphorylated (pT371)TRF1 is recruited to sites of DNA damage to facilitate homologous recombination and checkpoint activation. Nucleic acids research 19 23997120
2012 PinX1 localizes to telomeres and stabilizes TRF1 at mitosis. Molecular and cellular biology 19 22331467
2016 Cdk-dependent phosphorylation regulates TRF1 recruitment to PML bodies and promotes C-circle production in ALT cells. Journal of cell science 17 27185864
2014 PinX1, a telomere repeat-binding factor 1 (TRF1)-interacting protein, maintains telomere integrity by modulating TRF1 homeostasis, the process in which human telomerase reverse Transcriptase (hTERT) plays dual roles. The Journal of biological chemistry 17 24415760
2014 TRF1 and TRF2 differentially modulate Rad51-mediated telomeric and nontelomeric displacement loop formation in vitro. Biochemistry 17 25115914
2020 Pin2 telomeric repeat factor 1-interacting telomerase inhibitor 1 (PinX1) inhibits nasopharyngeal cancer cell stemness: implication for cancer progression and therapeutic targeting. Journal of experimental & clinical cancer research : CR 16 32028978
2020 NRT1.1-Mediated Nitrate Suppression of Root Coiling Relies on PIN2- and AUX1-Mediated Auxin Transport. Frontiers in plant science 16 32582237
2019 Biophysical characterization of the insertion of two potent antimicrobial peptides-Pin2 and its variant Pin2[GVG] in biological model membranes. Biochimica et biophysica acta. Biomembranes 16 31682816
2025 ABP1/ABL3-TMK1 cell-surface auxin signaling targets PIN2-mediated auxin fluxes for root gravitropism. Cell 14 41043433
2023 Boron supply restores aluminum-blocked auxin transport by the modulation of PIN2 trafficking in the root apical transition zone. The Plant journal : for cell and molecular biology 14 36721978
2009 Nucleolar modulation of TRF1: a dynamic way to regulate telomere and cell cycle by nucleostemin and GNL3L. Cell cycle (Georgetown, Tex.) 14 19713769
2023 Structural basis of telomeric nucleosome recognition by shelterin factor TRF1. Science advances 13 37624885
2022 In Vitro Characterization of the Physical Interactions between the Long Noncoding RNA TERRA and the Telomeric Proteins TRF1 and TRF2. International journal of molecular sciences 13 36142374
2022 Plasma tRF-1:29-Pro-AGG-1-M6 and tRF-55:76-Tyr-GTA-1-M2 as novel diagnostic biomarkers for lung adenocarcinoma. Frontiers in oncology 13 36203461
2021 Naked mole rat TRF1 safeguards glycolytic capacity and telomere replication under low oxygen. Science advances 13 33608273
2015 Integrated pathway and epistasis analysis reveals interactive effect of genetic variants at TERF1 and AFAP1L2 loci on melanoma risk. International journal of cancer 13 25892537
2014 PTOP and TRF1 help enhance the radio resistance in breast cancer cell. Cancer cell international 13 24460895

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