Affinage

TCOF1

Treacle protein · UniProt Q13428

Length
1488 aa
Mass
152.1 kDa
Annotated
2026-06-10
100 papers in source corpus 22 papers cited in narrative 21 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

Treacle (TCOF1) is a low-complexity, heavily phosphorylated nucleolar phosphoprotein that organizes ribosome biogenesis and the nucleolar genome-stability response (PMID:9096354, PMID:15249688). It nucleates liquid-like phase condensates—initiated by its C-terminal domain and stabilized by an alternating-charge central region—that build the nucleolar fibrillar center and are required for high-level rDNA transcription, accurate rRNA processing, and damage-response recruitment (PMID:40223701). Through its central repeated domain it binds RNA polymerase I while its C-terminus mediates rDNA promoter recognition and UBF recruitment, making Treacle the essential platform that loads the Pol I transcription machinery onto the nucleolus (PMID:15249688, PMID:19527688). Treacle also enters hNop56/NOP56-associated pre-ribosomal RNP complexes and is required for 2'-O-methylation of pre-rRNA, linking it directly to rRNA modification (PMID:12777385, PMID:15930015). Casein kinase 2 binds and phosphorylates Treacle, and this phosphorylation, together with ATM-dependent modification of conserved C-terminal Ser/Thr residues, gates its damage functions: Treacle recruits NBS1 and the phospho-peptide-binding BRCT domains of TOPBP1 into nucleoli upon rDNA double-strand breaks or replication stress, silencing rRNA synthesis, driving nucleolar segregation, and supporting ATR signaling at stalled forks (PMID:10982400, PMID:25064736, PMID:25512513, PMID:31913317, PMID:34100862). Beyond the nucleolus, Treacle redistributes during S phase to telomeres via TRF2 to inhibit RNA Pol II, limiting TERRA and preventing R-loop-driven replication-fork stalling, and in mitosis it localizes to centrosomes/kinetochores and binds Plk1 to control spindle orientation (PMID:22479190, PMID:33082515). Tcof1 haploinsufficiency causes a cell-autonomous deficiency of neural crest cells through reduced mature ribosome production, the basis of Treacher Collins syndrome craniofacial malformation (PMID:16938878, PMID:9736782).

Mechanistic history

Synthesis pass · year-by-year structured walk · 10 steps
  1. 1997 Medium

    Establishing TCOF1's primary structure defined it as a low-complexity nucleolar phosphoprotein, framing the hypothesis that phosphorylation and nucleolar trafficking underlie its function.

    Evidence Genomic sequencing and bioinformatic domain analysis of the 1,411-aa protein

    PMID:9096354

    Open questions at the time
    • Predicted phosphorylation and localization signals not yet experimentally tested
    • No functional assay of the repeated motifs
  2. 1998 High

    Mapping nucleolar/nuclear localization signals and showing disease-truncated protein is mislocalized connected Treacle's subcellular targeting to Treacher Collins pathology.

    Evidence GFP-fusion deletion mapping and mutagenesis of TCS mutations with live microscopy

    PMID:10982400 PMID:9736782 PMID:9811939

    Open questions at the time
    • Does not define what Treacle does once in the nucleolus
    • Functional consequence of mislocalization at the molecular level not resolved
  3. 2000 Medium

    Identifying CK2 as the kinase associating with and phosphorylating Treacle gave a mechanistic basis for its predicted heavy phosphorylation.

    Evidence Co-immunoprecipitation, in vitro kinase assays, immunofluorescence

    PMID:10545604 PMID:10982400

    Open questions at the time
    • Specific phosphosites and their functional impact not mapped
    • Co-IP does not prove direct kinase-substrate engagement in vivo
  4. 2005 High

    Linking Treacle to NOP56-associated RNP complexes and pre-rRNA 2'-O-methylation defined a direct role in rRNA modification beyond simple localization.

    Evidence Affinity purification/MS, RNase controls, antisense knockdown in Xenopus, Tcof1+/- mouse rRNA methylation analysis, Co-IP

    PMID:12777385 PMID:15930015

    Open questions at the time
    • Whether Treacle directly stimulates the methyltransferase or acts as a scaffold unresolved
    • Specificity for individual methylation sites not fully mapped
  5. 2009 High

    Domain dissection showed Treacle is the essential platform recruiting Pol I and UBF to rDNA, establishing a mechanistic hierarchy for nucleolar transcription initiation.

    Evidence Co-IP, ChIP, siRNA knockdown, domain mapping

    PMID:15249688 PMID:19527688

    Open questions at the time
    • Structural basis of Pol I and rDNA promoter recognition not solved
    • How phosphorylation modulates these interactions not addressed here
  6. 2006 High

    In vivo knockout established that Tcof1 haploinsufficiency depletes neural crest cells via reduced ribosome production, explaining Treacher Collins craniofacial defects.

    Evidence Germline Tcof1 mouse KO, neural crest lineage tracing, ribosome biogenesis assays

    PMID:16938878

    Open questions at the time
    • Cell-type specificity of the requirement not fully explained
    • Does not address Treacle's non-ribosomal roles
  7. 2014 High

    Discovery that Treacle recruits NBS1 to nucleoli upon DNA damage, in a CK2- and ATM-phosphorylation-dependent manner, defined a nucleolar DNA-damage response that silences rRNA synthesis.

    Evidence Reciprocal Co-IP, ATM-site mutagenesis, siRNA, relocalization assays, survival assays

    PMID:25064736 PMID:25512513

    Open questions at the time
    • Precise ATM phosphosites incompletely defined
    • How pan-nuclear silencing is propagated from the nucleolus unclear
  8. 2020 High

    Phospho-dependent BRCT-mediated TOPBP1 recruitment by Treacle, and a TRF2-mediated telomeric role suppressing TERRA, extended Treacle's scaffold function to rDNA break response and telomere replication integrity.

    Evidence Phospho-peptide binding, mutagenesis, Co-IP (TOPBP1, TRF2), ChIP, R-loop detection, RNase H1 rescue

    PMID:31913317 PMID:33082515

    Open questions at the time
    • Cell-cycle switch governing nucleolar-to-telomere redistribution not mechanistically defined
    • How Treacle inhibits Pol II at telomeres versus recruiting Pol I at rDNA unresolved
  9. 2021 High

    Treacle–TOPBP1 foci were shown to drive ATR signaling at stalled forks within the nucleolus, integrating Treacle into the replication stress response.

    Evidence Co-IP, immunofluorescence, siRNA, ATR signaling assays

    PMID:34100862

    Open questions at the time
    • Distinction between cap-forming and non-cap responses incompletely defined
    • Downstream effector recruitment partly uncharacterized
  10. 2025 High

    Demonstrating that Treacle drives liquid-like phase separation unified its diverse functions under a condensate-scaffolding mechanism required for fibrillar center assembly, transcription, processing, and TOPBP1 recruitment.

    Evidence Phase condensate reconstitution, domain-deletion and condensate-deficient mutants, transcription/processing and TOPBP1 recruitment assays

    PMID:40223701

    Open questions at the time
    • In vivo regulation of condensate formation by phosphorylation not fully defined
    • How condensate composition is tuned for damage versus transcription modes unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how a single phosphorylation- and condensate-dependent scaffold is dynamically partitioned across nucleolar transcription, the nucleolar DNA-damage response, telomere protection, and mitotic spindle control.
  • No integrated model linking cell-cycle state to Treacle's location switching
  • Structural basis of condensate-mediated partner selection unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 4 GO:0140110 transcription regulator activity 3 GO:0003723 RNA binding 2
Localization
GO:0005730 nucleolus 5 GO:0005634 nucleus 1 GO:0005815 microtubule organizing center 1
Pathway
R-HSA-73894 DNA Repair 4 R-HSA-74160 Gene expression (Transcription) 2 R-HSA-8953854 Metabolism of RNA 2 R-HSA-1266738 Developmental Biology 1 R-HSA-1640170 Cell Cycle 1
Partners
CK2FGF12NBS1NOP56PLK1TOPBP1TRF2UBF
Complex memberships
NOP56-associated pre-ribosomal RNP complexRNA Pol I transcription complex

Evidence

Reading pass · 21 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1997 TCOF1 encodes a low-complexity nucleolar phosphoprotein (treacle) of 1,411 amino acids with repeated motifs shared with nucleolar trafficking proteins, predicted to be highly phosphorylated by casein kinase, and containing putative nuclear and nucleolar localization signals. Genomic sequencing, bioinformatic analysis of protein structure Proceedings of the National Academy of Sciences of the United States of America Medium 9096354
1998 Treacle localizes to the nucleolus, with the last 41 residues of the C-terminus (residues 1262-1302) sufficient for nucleolar localization; at least two functional nuclear localization signals exist in the C-terminal region. GFP fusion constructs with full-length and deleted domains of murine treacle expressed in cells; live fluorescence microscopy Human molecular genetics High 10982400 9736782 9811939
1998 TCS mutations that create premature termination codons produce truncated treacle proteins that are mislocalized within the cell (not nucleolar), supporting the hypothesis that nucleolar shuttling is integral to treacle function. Site-directed mutagenesis recreating TCS mutations; GFP-fusion protein localization assays Human molecular genetics High 9736782
1999 Treacle fusion peptides can be phosphorylated by casein kinase II (CKII) and protein kinase C (PKC) in vitro; branchial arch tissue extracts contain a kinase activity consistent with CKII site recognition that phosphorylates treacle peptides, coinciding with peak treacle expression. In vitro kinase assays using GST-treacle fusion peptides with recombinant kinases and tissue extracts Human molecular genetics Medium 10545604
2000 Treacle is a highly phosphorylated nucleolar protein; casein kinase 2 (CK2) is responsible for its high degree of phosphorylation, as evidenced by co-immunoprecipitation of CK2 with treacle. Immunofluorescence, co-immunoprecipitation, phosphorylation analysis Molecular biology of the cell Medium 10982400
2003 Treacle is a constituent of hNop56p-associated pre-ribosomal ribonucleoprotein complexes; the association of treacle with hNop56p is independent of rRNA integrity, indicating a direct protein-protein interaction. Affinity purification of Nop56p complexes, mass spectrometry, RNase treatment controls The Journal of biological chemistry Medium 12777385
2004 Treacle interacts physically with upstream binding factor (UBF) and is involved in ribosomal DNA (rDNA) gene transcription; siRNA knockdown of treacle inhibits rDNA transcription and cell growth. Co-immunoprecipitation, yeast two-hybrid, immunofluorescence co-localization, siRNA knockdown with rDNA transcription assay Proceedings of the National Academy of Sciences of the United States of America High 15249688
2005 Treacle participates in 2'-O-methylation of pre-rRNA; antisense knockdown of treacle in Xenopus oocytes reduces pre-rRNA methylation, and Tcof1+/- mouse embryos show significant reduction in 2'-O-methylation at specific rRNA residues. Treacle's role in methylation is mediated through direct physical interaction with NOP56, a component of the ribonucleoprotein methylation complex. Antisense knockdown in Xenopus oocytes, rRNA methylation analysis, co-immunoprecipitation of treacle with NOP56, immunofluorescence co-localization Human molecular genetics High 15930015
2006 Tcof1/Treacle is required cell-autonomously for neural crest cell formation and proliferation; haploinsufficiency of Tcof1 leads to deficiency in migrating neural crest cells causing craniofacial malformations. Tcof1/Treacle regulates proliferation by controlling production of mature ribosomes. Germ-line Tcof1 mouse knockout, neural crest cell lineage tracing, ribosome biogenesis assays Proceedings of the National Academy of Sciences of the United States of America High 16938878
2009 The central repeated domain of treacle binds RNA polymerase I (Pol I), while the C-terminus is involved in rDNA promoter recognition and UBF recruitment. Treacle, but not UBF, is essential for nucleolar recruitment of the Pol I transcription complex; knockdown of treacle disperses Pol I and UBF from the nucleolus, but treacle-Pol I interactions and treacle-rDNA promoter interactions are not disrupted by UBF depletion. Co-immunoprecipitation, siRNA knockdown, chromatin immunoprecipitation (ChIP), domain-mapping experiments Biochemical and biophysical research communications High 19527688
2012 Treacle is a novel centrosome- and kinetochore-associated protein that controls spindle orientation and mitotic progression via a direct interaction with Polo-like kinase 1 (Plk1); Tcof1 loss-of-function disrupts spindle orientation and cell cycle progression, perturbing neural progenitor maintenance during cortical neurogenesis. Co-immunoprecipitation (Treacle-Plk1 interaction), Tcof1 loss-of-function mouse model, immunofluorescence for centrosome/kinetochore localization, spindle orientation analysis PLoS genetics High 22479190
2014 Treacle (TCOF1) interacts with NBS1 (Nijmegen breakage syndrome protein 1) and mediates NBS1 translocation into nucleoli in response to DNA damage, thereby triggering pan-nuclear silencing of rRNA transcription. NBS1 nucleolar accumulation is Treacle-dependent. Co-immunoprecipitation, siRNA knockdown, immunofluorescence relocalization assays after DNA damage induction Nature cell biology High 25064736 25512513
2014 NBS1 relocalization to nucleoli after DNA damage requires TCOF1 and is dependent on casein kinase II and ATM-mediated phosphorylation of TCOF1; a putative ATM phosphorylation site in TCOF1 is required for NBS1 nucleolar relocalization. TCOF1 promotes cellular resistance to DNA-damaging agents. Co-immunoprecipitation, site-directed mutagenesis of ATM phosphorylation site, siRNA knockdown, cell survival assays after DNA damage Proceedings of the National Academy of Sciences of the United States of America High 25512513
2020 Treacle recruits TOPBP1 into nucleoli in an ATM- and NBS1-dependent manner in response to rDNA double-strand breaks; this is mediated by phosphorylation-dependent interactions between three BRCT domains of TOPBP1 and conserved phosphorylated Ser/Thr residues at the C-terminus of Treacle. TOPBP1 recruitment is required for inhibition of rRNA synthesis and nucleolar segregation. Co-immunoprecipitation, phospho-peptide binding assays, siRNA knockdown, immunofluorescence, site-directed mutagenesis of phosphorylation sites Nature communications High 31913317
2020 During S phase, TCOF1 leaves the nucleolus and is recruited to telomeres by interacting with TRF2; at telomeres, TCOF1 suppresses telomere transcription by binding and inhibiting RNA Pol II, thereby limiting TERRA levels. Depletion of TCOF1 causes elevated TERRA, DNA/RNA R-loops, replication fork stalling, and fragile telomeres. Co-immunoprecipitation (TCOF1-TRF2), chromatin immunoprecipitation, siRNA knockdown, R-loop detection, RNase H1 rescue experiment, TERRA quantification Cell death and differentiation High 33082515
2021 TOPBP1 interacts with Treacle (TCOF1) and forms large foci inside the nucleolus in response to replication stress; Treacle and TOPBP1 together facilitate ATR signaling at stalled replication forks inside the nucleolus and promote recruitment of downstream replication stress response proteins without forming nucleolar caps. Co-immunoprecipitation, immunofluorescence, siRNA knockdown, ATR signaling assays The Journal of cell biology High 34100862
2022 FGF12 is localized to the nucleolus and interacts with both NOLC1 and TCOF1 in a phosphorylation-dependent manner requiring the C-terminal region of FGF12; NOLC1 and TCOF1 are unable to interact with each other in the absence of FGF12, indicating FGF12 bridges this complex. Co-immunoprecipitation, immunofluorescence localization, domain-deletion experiments, phosphorylation-dependence assays Cell communication and signaling : CCS Medium 36411431
2022 The matrix (M) proteins of Henipaviruses (including Hendra and Nipah viruses) and rabies virus P3 protein target nucleolar Treacle protein and suppress ribosomal RNA synthesis, mimicking the cellular nucleolar DNA-damage response mechanism; Treacle depletion impacts mumps virus particle production and MuV matrix protein interacts with and colocalizes with Treacle in the nucleolus. Co-immunoprecipitation (viral M protein with Treacle), siRNA knockdown, rRNA synthesis assays, viral growth kinetics Traffic (Copenhagen, Denmark) / Journal of virology Medium 36135364 36479968
2025 Treacle forms liquid-like phase condensates through electrostatic interactions; the C-terminal domain initiates phase separation while the central domain (with alternating charged residues) maintains condensate stability. These condensates are essential for nucleolar fibrillar center assembly, high-level rRNA gene transcription, accurate rRNA processing, and TOPBP1 recruitment for nucleolar DNA damage response. Mutant Treacle forms unable to phase-separate fail to support these functions. Phase condensate assays, domain-deletion mutants, overexpression of condensate-deficient mutants, rRNA transcription and processing assays, TOPBP1 recruitment assays eLife High 40223701
2008 Transcription of the Tcof1 promoter is regulated by differential repression; binding of Cebpb, Zfp161, and Sp1 transcription factors to the regulatory region was detected, with Zfp161 conferring cell-type-specific repression and Sp1/Sp3 conferring cell-type-specific activation. The minimal promoter (-253 to +43 bp) directs constitutive expression. Deletion and mutation reporter assays, chromatin binding (ChIP) studies, in neural and non-neural cell lines DNA and cell biology Medium 18771418
2010 Treacle is downregulated by oxidative stress (H2O2) in lung cells through proteasome-dependent degradation independently of mRNA changes, suggesting a role in oxidant defense. Treacle suppression by siRNA increases cellular sensitivity to H2O2, and treacle reduction is independent of p53 responses. SILAC proteomics, siRNA knockdown, proteasome inhibitor experiments, H2O2 sensitivity assays Proteomics Medium 20340163

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Tcof1/Treacle is required for neural crest cell formation and proliferation deficiencies that cause craniofacial abnormalities. Proceedings of the National Academy of Sciences of the United States of America 297 16938878
2004 The Treacher Collins syndrome (TCOF1) gene product is involved in ribosomal DNA gene transcription by interacting with upstream binding factor. Proceedings of the National Academy of Sciences of the United States of America 238 15249688
2014 The NBS1-Treacle complex controls ribosomal RNA transcription in response to DNA damage. Nature cell biology 139 25064736
1997 TCOF1 gene encodes a putative nucleolar phosphoprotein that exhibits mutations in Treacher Collins Syndrome throughout its coding region. Proceedings of the National Academy of Sciences of the United States of America 127 9096354
2003 Proteomic analysis of human Nop56p-associated pre-ribosomal ribonucleoprotein complexes. Possible link between Nop56p and the nucleolar protein treacle responsible for Treacher Collins syndrome. The Journal of biological chemistry 109 12777385
2005 The Treacher Collins syndrome (TCOF1) gene product is involved in pre-rRNA methylation. Human molecular genetics 108 15930015
2014 Treacher Collins syndrome TCOF1 protein cooperates with NBS1 in the DNA damage response. Proceedings of the National Academy of Sciences of the United States of America 100 25512513
2000 High mutation detection rate in TCOF1 among Treacher Collins syndrome patients reveals clustering of mutations and 16 novel pathogenic changes. Human mutation 87 11013442
2000 Characterization of the nucleolar gene product, treacle, in Treacher Collins syndrome. Molecular biology of the cell 82 10982400
2008 Treacher Collins syndrome: unmasking the role of Tcof1/treacle. The international journal of biochemistry & cell biology 78 19027870
2016 Triclosan (TCS) exposure impairs lipid metabolism in zebrafish embryos. Aquatic toxicology (Amsterdam, Netherlands) 77 26828895
2004 Genetic background has a major effect on the penetrance and severity of craniofacial defects in mice heterozygous for the gene encoding the nucleolar protein Treacle. Developmental dynamics : an official publication of the American Association of Anatomists 74 15042714
1997 Sequence analysis, identification of evolutionary conserved motifs and expression analysis of murine tcof1 provide further evidence for a potential function for the gene and its human homologue, TCOF1. Human molecular genetics 69 9158147
2020 Treacle controls the nucleolar response to rDNA breaks via TOPBP1 recruitment and ATR activation. Nature communications 64 31913317
1998 The Treacher Collins syndrome (TCOF1) gene product, treacle, is targeted to the nucleolus by signals in its C-terminus. Human molecular genetics 63 9811939
2016 Triclosan (TCS) and Triclocarban (TCC) cause lifespan reduction and reproductive impairment through oxidative stress-mediated expression of the defensome in the monogonont rotifer (Brachionus koreanus). Comparative biochemistry and physiology. Toxicology & pharmacology : CBP 61 27067728
2012 Mammalian neurogenesis requires Treacle-Plk1 for precise control of spindle orientation, mitotic progression, and maintenance of neural progenitor cells. PLoS genetics 57 22479190
1998 Mutations in the Treacher Collins syndrome gene lead to mislocalization of the nucleolar protein treacle. Human molecular genetics 57 9736782
2014 Autosomal recessive POLR1D mutation with decrease of TCOF1 mRNA is responsible for Treacher Collins syndrome. Genetics in medicine : official journal of the American College of Medical Genetics 54 24603435
2016 Face off against ROS: Tcof1/Treacle safeguards neuroepithelial cells and progenitor neural crest cells from oxidative stress during craniofacial development. Development, growth & differentiation 53 27481486
2009 Treacle recruits RNA polymerase I complex to the nucleolus that is independent of UBF. Biochemical and biophysical research communications 46 19527688
2015 TCS: a web server for multiple sequence alignment evaluation and phylogenetic reconstruction. Nucleic acids research 44 25855806
2012 Gross deletions in TCOF1 are a cause of Treacher-Collins-Franceschetti syndrome. European journal of human genetics : EJHG 40 22317976
2010 Defects in middle ear cavitation cause conductive hearing loss in the Tcof1 mutant mouse. Human molecular genetics 39 20106873
2004 Another face of the Treacher Collins syndrome (TCOF1) gene: identification of additional exons. Gene 37 15019983
2020 Morphological and immunohistochemical phenotype of TCs in the intestinal bulb of Grass carp and their potential role in intestinal immunity. Scientific reports 35 32820212
2012 Balancing neural crest cell intrinsic processes with those of the microenvironment in Tcof1 haploinsufficient mice enables complete enteric nervous system formation. Human molecular genetics 34 22228097
2009 A synonymous mutation in TCOF1 causes Treacher Collins syndrome due to mis-splicing of a constitutive exon. American journal of medical genetics. Part A 34 19572402
2020 PI3K/Akt/FoxO pathway mediates glycolytic metabolism in HepG2 cells exposed to triclosan (TCS). Environment international 33 31918333
2013 Zar1 represses translation in Xenopus oocytes and binds to the TCS in maternal mRNAs with different characteristics than Zar2. Biochimica et biophysica acta 33 23827238
2001 Canine TCOF1; cloning, chromosome assignment and genetic analysis in dogs with different head types. Mammalian genome : official journal of the International Mammalian Genome Society 33 11471057
2014 Possible causes of excess sludge reduction adding metabolic uncoupler, 3,3',4',5-tetrachlorosalicylanilide (TCS), in sequence batch reactors. Bioresource technology 32 25285765
2017 Effects of triclosan (TCS) on hormonal balance and genes of hypothalamus-pituitary- gonad axis of juvenile male Yellow River carp (Cyprinus carpio). Chemosphere 30 29175396
2005 TCOF1 mutation database: novel mutation in the alternatively spliced exon 6A and update in mutation nomenclature. Human mutation 30 15832313
2021 The Role of TCOF1 Gene in Health and Disease: Beyond Treacher Collins Syndrome. International journal of molecular sciences 29 33804586
2011 Novel mutations of TCOF1 gene in European patients with Treacher Collins syndrome. BMC medical genetics 29 21951868
1985 A novel cell type carrying both Th and Tc/s markers in the blood of cyclosporine-treated, allografted rats. Transplantation 28 3159136
2018 Comparison of hepatotoxicity and mechanisms induced by triclosan (TCS) and methyl-triclosan (MTCS) in human liver hepatocellular HepG2 cells. Toxicology research 27 30713659
2021 TCOF1 coordinates oncogenic activation and rRNA production and promotes tumorigenesis in HCC. Cancer science 26 34904330
2011 SMP production by activated sludge in the presence of a metabolic uncoupler, 3,3',4',5-tetrachlorosalicylanilide (TCS). Applied microbiology and biotechnology 25 22159739
2020 TRF2 recruits nucleolar protein TCOF1 to coordinate telomere transcription and replication. Cell death and differentiation 22 33082515
2019 New Insights into Multistep-Phosphorelay (MSP)/ Two-Component System (TCS) Regulation: Are Plants and Bacteria that Different? Plants (Basel, Switzerland) 22 31835810
2017 Different culture conditions affect the growth of human tendon stem/progenitor cells (TSPCs) within a mixed tendon cells (TCs) population. Journal of experimental orthopaedics 22 28244027
2010 SILAC analysis of oxidative stress-mediated proteins in human pneumocytes: new role for treacle. Proteomics 22 20340163
2022 FGF12 is a novel component of the nucleolar NOLC1/TCOF1 ribosome biogenesis complex. Cell communication and signaling : CCS 21 36411431
2022 Treacle Sticks the Nucleolar Responses to DNA Damage Together. Frontiers in cell and developmental biology 20 35646927
1999 Detection of an appropriate kinase activity in branchial arches I and II that coincides with peak expression of the Treacher Collins syndrome gene product, treacle. Human molecular genetics 20 10545604
2020 Genome-wide characterization of two-component system (TCS) genes in melon (Cucumis melo L.). Plant physiology and biochemistry : PPB 18 32229405
2019 TCOF1 pathogenic variants identified by Whole-exome sequencing in Chinese Treacher Collins syndrome families and hearing rehabilitation effect. Orphanet journal of rare diseases 18 31307516
2013 Large deletions encompassing the TCOF1 and CAMK2A genes are responsible for Treacher Collins syndrome with intellectual disability. European journal of human genetics : EJHG 18 23695276
2017 Mutation screening of Chinese Treacher Collins syndrome patients identified novel TCOF1 mutations. Molecular genetics and genomics : MGG 17 29230583
2009 Reduced transcription of TCOF1 in adult cells of Treacher Collins syndrome patients. BMC medical genetics 17 20003452
2005 Mutational analysis of the TCOF1 gene in 11 Japanese patients with Treacher Collins Syndrome and mechanism of mutagenesis. American journal of medical genetics. Part A 17 15759264
2012 First Report of a Single Exon Deletion in TCOF1 Causing Treacher Collins Syndrome. Molecular syndromology 16 22712005
2003 Novel mutation in the 5' splice site of exon 4 of the TCOF1 gene in the patient with Treacher Collins syndrome. American journal of medical genetics. Part A 16 14598341
2024 Current state of knowledge of triclosan (TCS)-dependent reactive oxygen species (ROS) production. Environmental research 15 38401681
2022 Triclosan (TCS) affects the level of DNA methylation in the human oral squamous cell carcinoma (SCC-15) cell line in a nontoxic concentration. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 15 35286965
2021 Treacle and TOPBP1 control replication stress response in the nucleolus. The Journal of cell biology 15 34100862
2013 A novel mutation in the TCOF1 gene found in two Chinese cases of Treacher Collins syndrome. International journal of pediatric otorhinolaryngology 15 23838542
2006 Treacher Collins syndrome with a de Novo 5-bp deletion in the TCOF1 gene. Journal of the Formosan Medical Association = Taiwan yi zhi 15 16801042
2004 Mosaicism of a TCOF1 mutation in an individual clinically unaffected with Treacher Collins syndrome. American journal of medical genetics. Part A 15 15039977
2024 Apoptotic metabolites ameliorate bone aging phenotypes via TCOF1/FLVCR1-mediated mitochondrial homeostasis. Journal of nanobiotechnology 14 39237990
2021 Resveratrol and Angiogenin-2 Combined With PEGDA/TCS Hydrogel for the Targeted Therapy of Hypoxic Bone Defects via Activation of the Autophagy Pathway. Frontiers in pharmacology 14 33927615
2021 Effect of triclosan (TCS) on the protein content and associated histological changes on tilapia, Oreochromis mossambicus (Peters, 1852). Environmental science and pollution research international 14 34148199
2022 Henipaviruses and lyssaviruses target nucleolar treacle protein and regulate ribosomal RNA synthesis. Traffic (Copenhagen, Denmark) 13 36479968
2016 Tcof1-Related Molecular Networks in Treacher Collins Syndrome. The Journal of craniofacial surgery 13 27300466
2023 OmpR (TCS response regulator) of Aeromonas veronii plays a major role in drug resistance, stress resistance and virulence by regulating biofilm formation. Microbial pathogenesis 12 37244492
2020 Mutation analysis of TCOF1 gene in Chinese Treacher Collins syndrome patients. Journal of clinical laboratory analysis 12 32909271
1997 Mouse TCOF1 is expressed widely, has motifs conserved in nucleolar phosphoproteins, and maps to chromosome 18. Biochemical and biophysical research communications 12 9299440
1990 Pneumopathogenicity of a Sendai virus protease-activation mutant, TCs, which is sensitive to trypsin and chymotrypsin. Journal of virology 12 2170692
2024 Knockdown of OsPHP1 Leads to Improved Yield Under Salinity and Drought in Rice via Regulating the Complex Set of TCS Members and Cytokinin Signalling. Plant, cell & environment 11 39696826
2022 Involvement of sirtuins (Sirt1 and Sirt3) and aryl hydrocarbon receptor (AhR) in the effects of triclosan (TCS) on production of neurosteroids in primary mouse cortical neurons cultures. Pesticide biochemistry and physiology 11 35715069
2022 A novel TcS allele conferring the high-theacrine and low-caffeine traits and having potential use in tea plant breeding. Horticulture research 11 36338849
2019 The DNA-binding mechanism of the TCS response regulator ArlR from Staphylococcus aureus. Journal of structural biology 11 31520693
2018 Identification of a novel TCOF1 mutation in a Chinese family with Treacher Collins syndrome. Experimental and therapeutic medicine 11 30186496
2014 A novel silent deletion, an insertion mutation and a nonsense mutation in the TCOF1 gene found in two Chinese cases of Treacher Collins syndrome. Molecular genetics and genomics : MGG 11 24994558
2008 Metabolic uncoupling of Shewanella oneidensis MR-1, under the influence of excess substrate and 3, 3', 4', 5-tetrachlorosalicylanilide (TCS). Biotechnology and bioengineering 11 17972334
2008 Dual regulation with Ser/Thr kinase cascade and a His/Asp TCS in Myxococcus xanthus. Advances in experimental medicine and biology 11 18792684
2007 Mutations and new polymorphic changes in the TCOF1 gene of patients with oculo-auriculo-vertebral spectrum and Treacher-Collins syndrome. Clinical dysmorphology 11 17786119
2023 Genome-wide identification and expression profiling of two-component system (TCS) genes in Brassica oleracea in response to shade stress. Frontiers in genetics 10 37323660
2022 The oncogenic role of treacle ribosome biogenesis factor 1 (TCOF1) in human tumors: a pan-cancer analysis. Aging 10 35093935
2008 Capturing the VirA/VirG TCS of Agrobacterium tumefaciens. Advances in experimental medicine and biology 10 18792688
2023 MiR-133b as a crucial regulator of TCS-induced cardiotoxicity via activating β-adrenergic receptor signaling pathway in zebrafish embryos. Environmental pollution (Barking, Essex : 1987) 9 37467918
2020 In-silico analysis of ligand-receptor binding patterns of α-MMC, TCS and MAP30 protein to LRP1 receptor. Journal of molecular graphics & modelling 9 32311663
2013 Treatability study of 3,3',4',5-tetrachlorosalicylanilide (TCS) combined with 2,4,6-trichlorophenol (TCP) to reduce excess sludge production in a sequence batch reactor. Bioresource technology 9 23856019
2008 Excess maternal transmission of markers in TCOF1 among cleft palate case-parent trios from three populations. American journal of medical genetics. Part A 9 18688869
2004 Treacher Collins syndrome with craniosynostosis, choanal atresia, and esophageal regurgitation caused by a novel nonsense mutation in TCOF1. American journal of medical genetics. Part A 9 15214011
2002 Novel autosomal dominant mandibulofacial dysostosis with ptosis: clinical description and exclusion of TCOF1. Journal of medical genetics 9 12114479
2025 Treacle's ability to form liquid-like phase condensates is essential for nucleolar fibrillar center assembly, efficient rRNA transcription and processing, and rRNA gene repair. eLife 8 40223701
2021 Identification of three novel TCOF1 mutations in patients with Treacher Collins Syndrome. Human genome variation 8 34580285
2020 Treacher Collins syndrome: A novel TCOF1 mutation and monopodial stapes. Clinical otolaryngology : official journal of ENT-UK ; official journal of Netherlands Society for Oto-Rhino-Laryngology & Cervico-Facial Surgery 8 32351010
2020 Identification of a novel gross deletion of TCOF1 in a Chinese prenatal case with Treacher Collins syndrome. Molecular genetics & genomic medicine 8 32543076
2012 Novel insertion in exon 5 of the TCOF1 gene in twin sisters with Treacher Collins syndrome. Journal of applied genetics 8 22415350
2010 Exclusion of MYF5, GSC, RUNX2, and TCOF1 mutation in a case of cerebro-costo-mandibular syndrome. Clinical dysmorphology 8 20177378
2020 Comparison of Transcriptomics Changes Induced by TCS and MTCS Exposure in Human Hepatoma HepG2 Cells. ACS omega 7 32455190
2020 Exploration of the effects of goose TCs on GCs at different follicular stages using a co-culture model. Bioscience reports 7 32706022
2008 Regulation of the mouse Treacher Collins syndrome homolog (Tcof1) promoter through differential repression of constitutive expression. DNA and cell biology 7 18771418
2024 FOSL1 is a key regulator of a super-enhancer driving TCOF1 expression in triple-negative breast cancer. Epigenetics & chromatin 6 39523372
2022 Nucleolar Protein Treacle Is Important for the Efficient Growth of Mumps Virus. Journal of virology 6 36135364
2008 Detection of a novel silent deletion, a missense mutation and a nonsense mutation in TCOF1. Pediatrics international : official journal of the Japan Pediatric Society 6 19067896

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