Affinage

TERT

Telomerase reverse transcriptase · UniProt O14746

Round 2 corrected
Length
1132 aa
Mass
127.0 kDa
Annotated
2026-04-28
130 papers in source corpus 42 papers cited in narrative 41 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

TERT is the catalytic reverse transcriptase subunit of the telomerase holoenzyme, which extends chromosome ends by synthesizing TTAGGG telomeric repeats using its integral RNA component (TERC/hTR) as a template; the minimal active human enzyme comprises two molecules each of TERT, TERC, and dyskerin (PMID:9398860, PMID:17395830). TERT expression is the rate-limiting determinant of telomerase activity in normal and cancer cells: its transcription is activated by c-MYC via E-box elements and SP1 via GC-rich sites in the core promoter, while cancer-specific reactivation frequently occurs through hotspot promoter mutations (-124C>T, -146C>T) that generate de novo ETS/GABP binding sites (PMID:9988278, PMID:23348506, PMID:33758097). Beyond telomere synthesis, TERT functions as a transcriptional co-regulator in the Wnt/β-catenin pathway through interaction with BRG1 and occupancy of Wnt target gene promoters, regulates pentose phosphate pathway flux and oxidative defense through an Nrf2-TERT regulatory loop, localizes to mitochondria under oxidative stress to preserve mitochondrial function, and is post-transcriptionally regulated by alternative splicing (RBM10-promoted exon skipping generates non-functional isoforms) and miR-128-mediated mRNA degradation (PMID:19571879, PMID:27148686, PMID:36901881, PMID:33520366). Heterozygous loss-of-function TERT mutations cause telomere shortening by haploinsufficiency and are causal for aplastic anemia and familial idiopathic pulmonary fibrosis (PMID:15814878, PMID:17392301).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 1989 High

    The question of whether human cells possess a telomere-synthesizing enzyme was answered by demonstrating that HeLa cell extracts contain an RNase-sensitive ribonucleoprotein that adds TTAGGG repeats to telomeric primers, establishing that telomerase-mediated telomere maintenance is conserved in humans.

    Evidence In vitro enzymatic assay with crude HeLa cell extracts plus RNase sensitivity testing

    PMID:2805070

    Open questions at the time
    • Catalytic protein subunit not yet identified
    • Stoichiometry and composition unknown
  2. 1997 High

    Identification and functional validation of TERT as the catalytic subunit resolved the molecular identity of the telomerase enzyme: cloning of hTERT/hEST2 revealed conserved reverse transcriptase motifs, mutagenesis of these motifs abolished catalytic activity, in vitro reconstitution of hTERT with hTR was sufficient for telomerase activity, and hTERT expression correlated with telomerase-positive status across cell types.

    Evidence Mass spectrometry identification (Euplotes), cDNA cloning by homology, site-directed mutagenesis of RT motifs, in vitro reconstitution, expression surveys across cell lines, and gene disruption in S. pombe and S. cerevisiae

    PMID:9110970 PMID:9252327 PMID:9288757 PMID:9328464 PMID:9398860

    Open questions at the time
    • Holoenzyme stoichiometry unresolved
    • Role of accessory subunits not defined
    • Alternative splicing regulation not characterized
  3. 1998 High

    Ectopic hTERT expression was shown to be sufficient to reconstitute telomerase activity and bypass replicative senescence in normal human cells without inducing malignant transformation, establishing TERT as the rate-limiting component and causally linking telomere shortening to cellular senescence.

    Evidence Retroviral expression of hTERT in normal fibroblasts and RPE cells; TRAP assay, telomere length measurement, cell counting, senescence staining, karyotyping

    PMID:9454332 PMID:9528864

    Open questions at the time
    • Whether TERT has functions beyond telomere elongation unknown
    • In vivo relevance not tested
  4. 1999 High

    Dissection of the TERT promoter revealed how TERT transcription is controlled: c-MYC directly activates TERT through E-box elements, and SP1 binds GC-rich sites in the minimal 181-bp core promoter, explaining cancer-specific versus normal-cell differential expression.

    Evidence Promoter cloning, deletion analysis, EMSA for c-MYC/E-box and SP1 binding, luciferase reporter assays in cancer vs. normal cells, c-MYC overexpression with RT-PCR

    PMID:9973199 PMID:9988278

    Open questions at the time
    • Additional trans-acting regulators not yet surveyed
    • Epigenetic regulation of the locus unexplored
  5. 2003 High

    The assumption that normal somatic cells completely lack TERT was overturned: hTERT is expressed in cycling presenescent fibroblasts and its disruption alters telomeric overhang structure and restricts lifespan, revealing dynamic telomerase regulation even in normal cells.

    Evidence Immunofluorescence and RT-PCR detection of hTERT in primary fibroblasts; dominant-negative hTERT expression; G-tail assay for overhang integrity

    PMID:12887925

    Open questions at the time
    • Mechanism by which TERT regulates overhang independent of bulk telomere length unclear
    • Cell-cycle-dependent regulation not resolved
  6. 2005 High

    The genetic basis of telomere biology disorders was established when heterozygous TERT mutations were shown to cause aplastic anemia through haploinsufficiency rather than dominant-negative effects, demonstrating that even partial reduction in telomerase activity is clinically consequential.

    Evidence Sequencing of TERT in aplastic anemia patients; coexpression of WT and mutant TERT in telomerase-deficient cells with TRAP assay

    PMID:15814878

    Open questions at the time
    • Genotype–phenotype correlations for specific mutations incomplete
    • Tissue specificity of haploinsufficiency not explained
  7. 2007 High

    Two advances defined the enzyme's composition and disease spectrum: purification of catalytically active human telomerase revealed a dimeric complex of two TERT, two TERC, and two dyskerin molecules, while germline TERT mutations were linked to familial idiopathic pulmonary fibrosis.

    Evidence ~10^8-fold purification with nucleotide addition-dependent elution plus mass spectrometry (stoichiometry); family-based cosegregation analysis with telomere length measurement (IPF)

    PMID:17392301 PMID:17395830 PMID:17460043

    Open questions at the time
    • Structural basis for holoenzyme assembly unknown at atomic resolution
    • Why lung is specifically vulnerable in TERT haploinsufficiency unclear
  8. 2009 High

    A non-canonical function of TERT was uncovered when it was shown to act as a transcriptional co-regulator in the Wnt/β-catenin pathway: TERT interacts with BRG1, occupies Wnt target gene promoters, and Tert-null mice exhibit Wnt-related developmental defects independent of telomere length changes.

    Evidence Reciprocal Co-IP (TERT–BRG1–β-catenin); ChIP at Wnt target promoters; Wnt reporter assays in cells and Xenopus; Tert−/− mouse phenotyping

    PMID:19571879

    Open questions at the time
    • Whether Wnt co-activation is relevant in all TERT-expressing tissues unknown
    • Structural basis of BRG1–TERT interaction not determined
  9. 2013 High

    The most frequent non-coding driver mutations in cancer were identified: recurrent TERT promoter mutations at -124C>T and -146C>T create de novo ETS factor binding sites, increase TERT transcription 2–4-fold, and occur in ~71% of melanomas and many other cancers, explaining how cancer cells reactivate telomerase.

    Evidence Systematic cancer genome sequencing (melanoma, multiple cancer types); luciferase reporter assays comparing WT and mutant promoters

    PMID:23348503 PMID:23348506

    Open questions at the time
    • Specific ETS family member mediating activation not yet identified
    • Chromatin context effects not characterized
  10. 2015 High

    The specific ETS factor was identified: GABP (GA-binding protein), particularly complexes containing the GABPB1L isoform, selectively binds the mutant TERT promoter and mediates TERT reactivation; TERT promoter mutations also correlate with increased TERT mRNA, protein, activity, and telomere length across cancer cell lines.

    Evidence ChIP for GABP at mutant vs. WT TERT promoter; GABPB1L knockdown with TERT expression readout; TERT mRNA/protein/activity/telomere length across 23 urothelial cancer lines

    PMID:25722414 PMID:33758097

    Open questions at the time
    • Why GABPB1L isoform is specifically required unclear
    • Contribution of additional cofactors to mutant promoter activation not resolved
  11. 2016 Medium

    A metabolic function of TERT was revealed: TERT regulates pentose phosphate pathway enzyme expression (G6PD, TKT) and oxidative defense through a feed-forward loop with Nrf2, positioning TERT as a metabolic regulator beyond its telomere elongation role.

    Evidence siRNA and dominant-negative TERT in GBM cells; G6PD and TKT activity assays; Nrf2 overexpression rescue; xenograft model

    PMID:27148686

    Open questions at the time
    • Whether metabolic regulation requires catalytic activity or is structure-dependent unknown
    • Generalizability beyond glioblastoma not tested
  12. 2020 High

    Precision editing validated the causal role of TERT promoter mutations: adenine base editing to revert the -124C>T mutation blocked ETS factor binding, reduced TERT expression, and induced senescence in glioma cells in vitro and in vivo, while GABPB1L knockdown selectively suppressed TERT-promoter-mutant tumors and synergized with temozolomide.

    Evidence CjABE base editing with ChIP and expression readout; inducible GABPB1L shRNA in intracranial GBM xenografts; CRISPR TERT disruption with telomere and xenograft assays

    PMID:31963842 PMID:32066906 PMID:33758097

    Open questions at the time
    • Delivery of base editors for clinical application not solved
    • Long-term effects of TERT promoter correction on normal tissues unknown
  13. 2021 Medium

    Post-transcriptional regulation of TERT was detailed: intron retention controls nuclear compartmentalization of TERT mRNA with mitosis-specific splicing, RBM10 promotes generation of non-functional splice isoforms, and Nrf2 operates upstream of TERT to regulate ferroptosis via SLC7A11.

    Evidence Single-molecule RNA FISH; thiomorpholino antisense perturbation; RBM10 RNA-IP and TERT isoform switching by RT-PCR; Nrf2 knockout mouse and TERT overexpression with ferroptosis markers

    PMID:33520366 PMID:34083519 PMID:34716298

    Open questions at the time
    • Cell-cycle machinery linking mitotic splicing to TERT not identified
    • Relative contribution of splicing vs. transcription to TERT protein levels in vivo unclear
    • SLC7A11 regulation by TERT mechanism not defined
  14. 2023 Medium

    TERT's mitochondrial protective role was directly demonstrated: TERT localizes to mitochondria and this localization increases under oxidative stress, where TERT overexpression reduces ROS, preserves membrane potential, and upregulates antioxidant proteins.

    Evidence Mitochondrial fractionation and immunofluorescence in hTERT-overexpressing fibroblasts; ROS and membrane potential assays after H2O2 and radiation

    PMID:36901881

    Open questions at the time
    • Mitochondrial import signal and mechanism not defined
    • Whether mitochondrial TERT requires catalytic activity unknown
    • Single laboratory, awaits independent replication
  15. 2024 High

    Pharmacologic TERT activation was shown to reverse aging phenotypes: a TERT-activating compound acts via MEK/ERK/AP-1 signaling and promotes DNMT3B-mediated p16INK4a silencing, reducing senescence, stimulating neurogenesis, and alleviating neuroinflammation in aged mice.

    Evidence Small-molecule screen; MEK/ERK/AP-1 pathway inhibition; DNMT3B ChIP at p16 promoter; telomere length and senescence assays; BrdU labeling for neurogenesis in aged mice

    PMID:38908367

    Open questions at the time
    • Long-term oncogenic risk of pharmacologic TERT activation not assessed
    • Whether p16 silencing is required for all rejuvenation phenotypes unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include: the atomic-resolution structure of the complete human telomerase holoenzyme on a telomere substrate, the precise mechanism by which TERT exerts telomere-independent functions (Wnt co-activation, metabolic regulation, mitochondrial protection) and whether these require catalytic activity, and how tissue-specific pathology arises from systemic TERT haploinsufficiency.
  • No high-resolution cryo-EM structure of human telomerase engaged with telomeric DNA
  • Catalytic-activity dependence of non-canonical functions not systematically tested
  • Tissue-specific vulnerability in telomere biology disorders mechanistically unexplained

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140097 catalytic activity, acting on DNA 5 GO:0003723 RNA binding 3 GO:0140110 transcription regulator activity 2
Localization
GO:0005694 chromosome 3 GO:0005634 nucleus 2 GO:0005739 mitochondrion 1
Pathway
R-HSA-74160 Gene expression (Transcription) 5 R-HSA-1643685 Disease 4 R-HSA-162582 Signal Transduction 3 R-HSA-1640170 Cell Cycle 3 R-HSA-73894 DNA Repair 2 R-HSA-4839726 Chromatin organization 1
Partners
AGO2BRG1DKC1GABPB1LRBM10SP1TERCTPP1
Complex memberships
Telomerase holoenzyme (TERT–TERC–dyskerin)β-catenin–BRG1 transcriptional complex

Evidence

Reading pass · 41 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1989 Human telomerase (identified in HeLa cell extracts) is a ribonucleoprotein that synthesizes TTAGGG repeats onto telomeric primers, establishing that telomerase-mediated telomere maintenance is conserved in human cells. In vitro enzymatic assay with crude HeLa cell extracts; RNase sensitivity assay establishing ribonucleoprotein nature Cell High 2805070
1997 The catalytic subunit of telomerase contains reverse transcriptase motifs; single amino acid substitutions within these RT motifs in the yeast homolog Est2 cause telomere shortening and senescence, demonstrating these motifs are essential for catalysis of telomere elongation. Mass spectrometry identification of Euplotes p123; yeast Est2 in vitro telomeric DNA extension assay; site-directed mutagenesis of RT motifs Science High 9110970
1997 hEST2 (TERT), cloned based on homology to lower eukaryote telomerase catalytic subunit genes, is expressed in telomerase-positive tumor cells and tissues but not in telomerase-negative differentiated cells, and is upregulated concomitant with telomerase activation during immortalization, identifying it as the putative human telomerase catalytic subunit. cDNA cloning by homology; Northern blot expression analysis across telomerase-positive and -negative cell lines; mRNA quantification during cellular immortalization and differentiation Cell High 9288757
1997 hTRT (TERT) together with hTR (telomerase RNA) reconstitutes telomerase activity in vitro; single amino acid changes in conserved telomerase-specific and RT motifs abolish activity, directly demonstrating hTRT is the catalytic protein component; hTRT is the limiting component for telomerase activity in normal human diploid cells. In vitro transcription/translation reconstitution of hTRT + hTR; site-directed mutagenesis of RT and telomerase-specific motifs; transient transfection of hTRT into normal human cells with telomerase activity assay (TRAP) Nature genetics High 9398860
1997 The fission yeast TERT homolog (SpTrt1) is required for telomere maintenance; disruption leads to telomere shortening and senescence, establishing functional conservation of TERT across eukaryotes. Gene disruption in S. pombe; telomere length analysis; senescence phenotype Science High 9252327
1998 Ectopic expression of hTERT in telomerase-negative cells induces telomerase activity to levels comparable to immortal cells, and the expressed hTERT protein is physically associated with cellular telomerase activity, demonstrating hTERT is the rate-limiting determinant of telomerase activity. Ectopic expression of epitope-tagged hTERT; TRAP assay; co-immunoprecipitation of hTERT with cellular telomerase activity Oncogene High 9528864
1998 Introduction of the human telomerase catalytic subunit (hTERT) into normal human cells (retinal pigment epithelial cells and foreskin fibroblasts) results in telomere elongation, continued vigorous division beyond normal lifespan without senescence, and normal karyotype, establishing a causal relationship between telomere shortening and replicative senescence. Retroviral transfection of hTERT into normal human cells; telomere length measurement; cell division counting; beta-galactosidase senescence staining; karyotyping Science High 9454332
1999 c-MYC directly activates TERT transcription by binding E-box elements in the TERT core promoter; c-MYC-induced TERT expression is rapid and independent of cell proliferation and de novo protein synthesis, identifying TERT as a direct transcriptional target of c-MYC. Promoter cloning and reporter assays; gel shift assays (EMSA) for c-MYC binding to E-box in TERT promoter; c-MYC overexpression experiments; RT-PCR for TERT mRNA Nature genetics High 9988278
1999 The hTERT core promoter (181 bp upstream of transcription start) is GC-rich, lacks TATA and CAAT boxes, contains E-box and Sp1 binding sites, and is selectively activated in cancer and immortalized cells but repressed in normal primary cells; c-Myc overexpression significantly increases core promoter transcriptional activity. Promoter cloning (~3.5 kb 5'-flanking sequence); transient transfection reporter assays in cancer vs. normal cells; deletion analysis; gel shift assay (E-box and Sp1 binding); CapSite Hunting for transcription start site Cancer research High 9973199
2001 Xenopus TERT (xTERT) co-synthesized with Xenopus telomerase RNA (xTR) in a rabbit reticulocyte lysate reconstitutes active telomerase, confirming functional conservation; xTERT expression and telomerase activity are both high at early blastula and decline at gastrulation, with TERT expression level being the primary regulatory mechanism for telomerase activity during Xenopus development. cDNA cloning of xTERT; in vitro reconstitution in rabbit reticulocyte lysate with xTR; TRAP assay at different developmental stages; RT-PCR expression analysis Gene High 11602347
2003 hTERT is expressed in cycling primary presenescent human fibroblasts previously believed to lack telomerase; disruption of telomerase activity in normal human cells slows proliferation, restricts lifespan, and alters maintenance of the 3' single-stranded telomeric overhang without changing the rate of overall telomere shortening, revealing that telomerase and telomere structure are dynamically regulated in normal cells. Immunofluorescence and RT-PCR for hTERT in primary fibroblasts; dominant-negative hTERT expression to disrupt telomerase; telomere overhang measurement (G-tail assay); cell proliferation and lifespan assays Cell High 12887925
2004 Telomerase is regulated in cis at the telomere terminus by single-stranded DNA-binding proteins (POT1/Cdc13) that recruit telomerase, and along the telomere length by TRF1/TIN2/tankyrase (humans) and Rap1/Rif1/Rif2 (yeast) as part of a negative feedback loop regulating telomere length; the Est2 (TERT) reverse transcriptase associates with telomerase RNA Tlc1 and accessory factors Est1, Est3, and Sm proteins. Review synthesizing genetic and biochemical epistasis data, co-IP, reconstitution, and structural studies from multiple labs Annual Review of Biochemistry High 15189140
2004 In Saccharomyces cerevisiae, a specific TLC1 stem-loop structure (not sequence) mediates the interaction with the Est2 (TERT) reverse transcriptase protein; replacement of a 95-nt region required for Est2 interaction with a 39-nt pseudoknot from a distantly related telomerase RNA reconstitutes functional telomerase, demonstrating that Est2-RNA association depends on a highly structured region rather than specific sequences. Site-directed mutagenesis of TLC1 stem-loop; co-immunoprecipitation of Est2 with TLC1 variants; in vivo telomere length assay; functional complementation Molecular and Cellular Biology High 15314178
2005 Heterozygous missense mutations in TERT impair telomerase activity by haploinsufficiency in aplastic anemia patients; coexpression of wild-type TERT and mutant TERT in a telomerase-deficient cell line demonstrated that mutations do not exert dominant-negative effects but reduce enzyme activity by ~50%. Sequencing of TERT in aplastic anemia patients; telomere length measurement; telomerase activity assay (TRAP); cell-based coexpression of wild-type and mutant TERT in telomerase-deficient cells New England Journal of Medicine High 15814878
2007 Purification of catalytically active human telomerase from immortal cells revealed its protein composition: two molecules each of TERT, telomerase RNA (hTR), and dyskerin, establishing the minimal stoichiometry of the functional human telomerase holoenzyme. ~10^8-fold purification by affinity chromatography with nucleotide addition-dependent elution; mass spectrometric sequencing of protein components; molecular size determination Science High 17395830
2007 Germline heterozygous mutations in TERT cause short telomeres and cosegregate with familial idiopathic pulmonary fibrosis; mutant telomerase resulted in short telomeres in asymptomatic carriers, establishing TERT loss-of-function as a cause of adult-onset pulmonary fibrosis. Sequencing of TERT and TERC in familial IPF probands; telomere length measurement by flow-FISH and Southern blot; cosegregation analysis in families New England Journal of Medicine High 17392301 17460043
2009 TERT directly modulates Wnt/β-catenin signaling by serving as a cofactor in a β-catenin transcriptional complex; TERT interacts with BRG1 (SMARCA4), a SWI/SNF chromatin remodeling protein, and activates Wnt-dependent reporters; TERT physically occupies promoters of Wnt-dependent genes in mouse gastrointestinal tract; Tert-/- mice show homeotic vertebral transformations consistent with Wnt signaling defects. Co-immunoprecipitation of TERT with BRG1 and β-catenin; Wnt reporter assays in cultured cells and in vivo (Xenopus axis formation); chromatin immunoprecipitation (ChIP) of endogenous TERT at Wnt target gene promoters; analysis of Tert-/- mouse phenotype Nature High 19571879
2011 Yeast Est3p (structural homolog of human shelterin protein TPP1) directly binds the TEN domain of the Est2p (TERT) catalytic subunit; surface mutations in Est3p that disrupt this interaction reduce Est3p assembly with telomerase in vivo and cause telomere shortening and senescence; recombinant Est3p stimulates telomerase activity in vitro in a TEN domain-dependent manner. In vitro binding assay of recombinant Est3p with purified TEN domain of Est2p; site-directed mutagenesis; co-immunoprecipitation from yeast; in vitro telomerase activity assay; telomere Southern blot Journal of Biological Chemistry High 21659533
2013 Two recurrent hotspot mutations at -124C>T and -146C>T in the TERT core promoter occur in 71% of melanomas and 16% of diverse cancer cell lines; these mutations create de novo consensus binding motifs for ETS transcription factors and increase TERT transcriptional activity 2–4-fold in reporter assays. Systematic cancer genome sequencing; reporter assays with wild-type vs. mutant TERT promoter constructs; ETS binding site bioinformatics prediction Science High 23348503 23348506
2015 TERT promoter mutations (C228T and C250T) in urothelial cancer correlate with higher TERT mRNA, TERT protein, telomerase enzymatic activity, and telomere length across 23 human UC cell lines, establishing that these promoter mutations functionally increase telomerase activity in cancer. TERT promoter mutation sequencing; quantitative RT-PCR for TERT mRNA; TERT protein immunoblot; TRAP assay for telomerase activity; telomere length measurement Science High 25722414
2015 TERT promoter mutations create de novo ETS factor binding sites preferentially recruited by GABP (GA-binding protein) transcription factor complex; increased binding of GABPB1L-containing complex to mutant TERT promoter mediates TERT reactivation in glioblastoma. Chromatin immunoprecipitation (ChIP) of GABP factors at mutant vs. wild-type TERT promoter; GABPB1L knockdown with measurement of TERT expression; reporter assays Proceedings of the National Academy of Sciences High 33758097
2016 TERT regulates the pentose phosphate pathway (PPP) in glioblastoma: TERT knockdown abrogates expression and activity of G6PD and Transketolase (TKT), and increases glycogen accumulation; this is mediated through an Nrf2-TERT regulatory loop in which TERT maintains Nrf2 levels and Nrf2 in turn regulates TERT expression. siRNA knockdown and dominant-negative TERT; pharmacological inhibition with Costunolide; G6PD and TKT activity assays; glycogen measurement; xenograft mouse model; Nrf2 overexpression rescue experiments Cell Death & Disease Medium 27148686
2018 Argonaute 2 (AGO2) promotes telomerase activity and stimulates the physical association between TERT and TERC; AGO2 depletion results in shorter telomeres and reduced proliferation in vitro and in vivo; AGO2 interacts with TERC and a small RNA (terc-sRNA) derived from the H/ACA box of TERC, and terc-sRNA overexpression is sufficient to enhance telomerase activity. AGO2 gain- and loss-of-function (siRNA, overexpression); TRAP assay; TERT-TERC co-immunoprecipitation; telomere length measurement; in vivo xenograft proliferation assay; RNA immunoprecipitation of AGO2 with TERC EMBO Reports Medium 30591524
2018 miR-128 directly interacts with the coding sequence of TERT mRNA (not 3'UTR) to reduce TERT mRNA and protein levels and inhibit telomerase activity; the miR-128 binding site in TERT mRNA is conserved with the reverse transcriptase of LINE-1. Anti-miR library screen; luciferase reporter assays with TERT coding sequence; RNA immunoprecipitation; quantitative RT-PCR and western blot in multiple cancer cell lines Oncotarget Medium 29568354
2020 TERT promoter mutations create a binding site for a specific GABPB1L-isoform-containing complex; GABPB1L knockdown selectively suppresses TERT expression and proliferation in TERT-promoter-mutant GBM cells but not wild-type cells; combined GABPB1L knockdown with temozolomide impairs DNA damage response and profoundly reduces intracranial GBM tumor growth. ChIP for GABPB1L at mutant TERT promoter; inducible shRNA knockdown of GABPB1L in GBM cells and in vivo intracranial xenografts; TERT expression measurement; cell proliferation and DNA damage response assays; temozolomide combination treatment PNAS High 33758097
2020 CRISPR/Cas9-mediated TERT disruption causes telomere attrition and growth retardation in vitro; TERT haploinsufficiency in tumor cells is sufficient to prevent xenograft formation in vivo, demonstrating TERT is required for cancer cell survival. CRISPR/Cas9 gene editing of TERT in cancer cells; telomere length measurement; in vitro proliferation assays; nude mouse xenograft transplantation International Journal of Molecular Sciences Medium 31963842
2020 Correction of the -124C>T TERT promoter mutation to -124C using adenine base editing (CjABE) blocks ETS transcription factor binding to the TERT promoter, reduces TERT transcription and protein expression, and induces cancer-cell senescence and proliferative arrest; local AAV injection of CjABE inhibits growth of TERT-promoter-mutant gliomas in vivo. CRISPR-based adenine base editing (CjABE) with sgRNA; ETS factor ChIP and binding assays; TERT mRNA and protein measurement; senescence assays (SA-β-gal); in vivo AAV injection in glioma mouse model Nature Cell Biology High 32066906
2020 NCOA3 (nuclear receptor coactivator-3) specifically binds the TERT promoter at the -234 to -144 region and transcriptionally activates TERT expression; NCOA3 recruits SP1 to the TERT promoter; NCOA3-mediated HCC cell growth and tumor progression in vitro and in vivo acts through upregulating TERT signaling. ChIP of NCOA3 at TERT promoter; Co-IP of NCOA3 with SP1; NCOA3 knockdown with TERT overexpression rescue; in vitro colony formation and viability; in vivo xenograft Cell Death & Disease Medium 33239622
2020 TERT promoter mutations give rise to monoallelic TERT expression in cancer; deep RNA sequencing of cancer cell lines shows that hotspot TERT promoter mutations without exception lead to re-expression of only one allele, accounting for ~50% of monoallelic TERT expression cases. Deep RNA sequencing with allele-specific analysis of TERT expression in cancer cell lines; genotyping of TERT promoter mutations Oncogenesis Medium 26657580
2020 KLF4 transcriptionally regulates TERT expression in alveolar epithelial cells by directly binding the TERT promoter; KLF4 overexpression preserves TERT expression and telomerase activity in senescent alveolar epithelial cells and suppresses bleomycin-induced pulmonary fibrosis in mice. ChIP of KLF4 at TERT promoter; siRNA knockdown and overexpression of KLF4; TRAP assay for telomerase activity; TERT expression by RT-PCR/western blot; AAV-6-mediated KLF4 overexpression in vivo; bleomycin mouse model Cell Death & Disease Medium 35508454
2021 Nuclear compartmentalization of TERT pre-mRNA is driven by intron retention; single-molecule RNA FISH reveals nuclear TERT transcripts uniformly retain specific introns; splicing of retained introns occurs during mitosis; blocking intron excision with thiomorpholino antisense oligonucleotides forces nuclear retention and has significant effects on cell viability. Single-molecule RNA FISH for subcellular localization; RNA-targeting thiomorpholino antisense oligonucleotides to block splicing; cell viability assays; RT-PCR for splicing status at different cell cycle stages Nature Communications Medium 34083519
2021 Most TERT rare missense variants found in MDS patients (90%) have severe or intermediate impairment of telomere elongation capacity in cell-based assays; homology modeling of human TERT bound to TPP1 suggests these variants disrupt domain-specific functions including catalysis, protein-RNA interactions, and telomere recruitment. Cell-based telomere elongation assay for each cloned TERT variant; TERT sequencing in 1514 MDS patients; telomere length measurement; homology structural modeling with TERT-TPP1 interface Blood Medium 34019641
2021 Nrf2 modulates TERT expression; Nrf2 knockout reduces TERT levels and exacerbates ferroptosis in lung epithelial cells; TERT overexpression elevates SLC7A11 levels and thereby inhibits ferroptosis, placing TERT downstream of Nrf2 in a pathway that regulates ferroptosis via SLC7A11. Nrf2 knockout mouse model; OGD/R cell model; Nrf2 siRNA knockdown; TERT overexpression; measurement of MDA, GSH, GPX4, Fe levels (STXM); SLC7A11 expression by western blot Cell Death & Disease Medium 34716298
2021 RBM10 promotes exclusion of exons 7 and 8 in TERT pre-mRNA splicing, generating a non-functional TERT-s isoform rather than full-length TERT, thereby suppressing telomerase activity and telomere shortening; gain or loss of RBM10 changes pancreatic cancer cell proliferation in vitro and in xenografts. RNA immunoprecipitation (RNA-IP) and RNA pull-down assays to show RBM10-TERT mRNA interaction; RT-PCR for TERT isoform switching; TRAP assay for telomerase activity; RBM10 gain/loss-of-function; xenograft experiments American Journal of Cancer Research Medium 33520366
2022 EGFR amplification drives TERT expression selectively from the mutant TERT promoter through an AMPK-GABP pathway; pharmacologic and genetic inhibition of EGFR decreases TERT expression in a GABP-dependent manner; EGFR-AMPK signaling promotes telomerase activity and telomere length maintenance in TERT-promoter-mutant glioblastoma. EGFR inhibition (pharmacologic and genetic) with TERT mRNA/protein measurement; AMPK activation/inhibition; GABP knockdown rescue experiments; ChIP for GABP at TERT promoter; telomere length and telomerase activity assays; analysis of TCGA copy number, mRNA, and protein data Cell Reports Medium 36130485
2022 TPP1 promoter variants (in the ACD gene, ~5% of cutaneous melanoma) co-occur with TERT promoter mutations, create or modify ETS transcription factor binding sites, increase TPP1 expression, and synergistically lengthen telomeres together with TERT activation, suggesting cooperative immortalization through dual telomere maintenance pathway upregulation. Whole-genome sequencing of melanomas; luciferase reporter assays for ACD promoter variants; CRISPR-mediated introduction of variants; telomere length measurement; expression analysis of TPP1 and TERT Science Medium 36356143
2022 TERT silencing (directly or via GABPB1 knockdown) in glioblastoma reduces lactate production, glutathione (GSH), and pentose phosphate pathway flux via reduced glucose-6-phosphate dehydrogenase activity and NADPH; these metabolic changes serve as MRS-detectable biomarkers of TERT modulation. 1H-MRS and hyperpolarized 13C-MRS in GBM cells and in vivo tumors; TERT shRNA and GABPB1 shRNA knockdown; metabolic enzyme activity assays (G6PD); NADPH and NADH measurement; lactate dehydrogenase and GLUT1 expression Neuro-oncology Medium 35460557
2022 Melatonin alleviates alcoholic liver disease via direct binding to EGFR on hepatocytes; downstream BRG1 regulates TERT expression; liver-specific knockdown of Tert or Egfr in ALD mice impairs melatonin-mediated liver protection, placing TERT as a downstream effector of the EGFR-BRG1 axis. Biophysical binding assay (MLT-EGFR); RNA-seq; liver-specific Tert and Egfr knockdown (AAV); ALD mouse model; BRG1 inhibition; cell-based loss-of-function studies Acta Pharmaceutica Sinica B Medium 36815038
2023 TERT localizes to mitochondria and this localization increases after oxidative stress (H2O2 treatment); TERT overexpression reduces ROS induction, increases antioxidant defense protein expression, preserves mitochondrial membrane potential and morphology, and reduces basal mitochondria quantity, demonstrating a non-canonical antioxidant and mitochondrial protective role for TERT. Mitochondrial fractionation; immunofluorescence for TERT mitochondrial localization; ROS measurement; antioxidant protein expression (western blot); mitochondrial membrane potential assay; H2O2 and X-ray treatment of hTERT-overexpressing fibroblasts International Journal of Molecular Sciences Medium 36901881
2024 A TERT activator compound (TAC) upregulates TERT transcription via the MEK/ERK/AP-1 cascade; in primary human cells and aged mice, TAC-induced TERT elevation promotes telomere synthesis, reduces cellular senescence and inflammatory cytokines, silences p16INK4a via DNMT3B-mediated promoter hypermethylation, stimulates adult neurogenesis, and alleviates neuroinflammation. Small-molecule screen for TERT activators; MEK/ERK/AP-1 pathway perturbation (inhibitors); DNMT3B ChIP at p16 promoter; telomere length measurement; senescence assays; neurogenesis (BrdU labeling); cognitive testing in aged mice; inflammatory cytokine measurement Cell High 38908367
1997 hTERT (hEST2) gene was cloned from human cells; the gene produces complex alternative splicing patterns in different cell types; it is expressed in telomerase-positive tumor lines and immortalized cells but absent from most normal pre-crisis and ALT cell lines, identifying alternative splicing as a potential regulatory mechanism for telomerase activity. cDNA library screening and RT-PCR cloning; RT-PCR with primers in reverse transcriptase domain revealing multiple splice products; Northern blot analysis across cell types Human Molecular Genetics Medium 9328464

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1998 Extension of life-span by introduction of telomerase into normal human cells. Science (New York, N.Y.) 3879 9454332
1997 Telomerase catalytic subunit homologs from fission yeast and human. Science (New York, N.Y.) 1975 9252327
1997 hEST2, the putative human telomerase catalytic subunit gene, is up-regulated in tumor cells and during immortalization. Cell 1596 9288757
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