Hanahan, D. & Weinberg, R. A. The hallmarks of cancer. Cell 100, 57–70 (2000). This paper identifies the key hallmarks of cancer, one of which is infinite replicative potential that can be enabled by activation of a TMM.

CAS  PubMed  Article  Google Scholar 

Guterres, A. N. & Villanueva, J. Targeting telomerase for cancer therapy. Oncogene 39, 5811–5824 (2020).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Rousseau, P. & Autexier, C. Telomere biology: rationale for diagnostics and therapeutics in cancer. RNA Biol. 12, 1078–1082 (2015).

PubMed  PubMed Central  Article  Google Scholar 

Meyne, J., Ratliff, R. L. & Moyzis, R. K. Conservation of the human telomere sequence (TTAGGG)n among vertebrates. Proc. Natl Acad. Sci. USA 86, 7049–7053 (1989).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Harley, C. B., Futcher, A. B. & Greider, C. W. Telomeres shorten during ageing of human fibroblasts. Nature 345, 458–460 (1990).

CAS  PubMed  Article  Google Scholar 

Griffith, J. D. et al. Mammalian telomeres end in a large duplex loop. Cell 97, 503–514 (1999).

CAS  PubMed  Article  Google Scholar 

Larrivee, M., LeBel, C. & Wellinger, R. J. The generation of proper constitutive G-tails on yeast telomeres is dependent on the MRX complex. Genes Dev. 18, 1391–1396 (2004).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Makarov, V. L., Hirose, Y. & Langmore, J. P. Long G tails at both ends of human chromosomes suggest a C strand degradation mechanism for telomere shortening. Cell 88, 657–666 (1997).

CAS  PubMed  Article  Google Scholar 

Wright, W. E., Tesmer, V. M., Huffman, K. E., Levene, S. D. & Shay, J. W. Normal human chromosomes have long G-rich telomeric overhangs at one end. Genes Dev. 11, 2801–2809 (1997).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Erdel, F. et al. Telomere recognition and assembly mechanism of mammalian shelterin. Cell Rep. 18, 41–53 (2017).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Chong, L. et al. A human telomeric protein. Science 270, 1663–1667 (1995).

CAS  PubMed  Article  Google Scholar 

de Lange, T. Shelterin: the protein complex that shapes and safeguards human telomeres. Genes Dev. 19, 2100–2110 (2005).

PubMed  Article  CAS  Google Scholar 

Van Ly, D. et al. Telomere loop dynamics in chromosome end protection. Mol. Cell 71, 510–525.e6 (2018).

PubMed  Article  CAS  Google Scholar 

Tomaska, L., Nosek, J., Kar, A., Willcox, S. & Griffith, J. D. A new view of the T-loop junction: implications for self-primed telomere extension, expansion of disease-related nucleotide repeat blocks, and telomere evolution. Front. Genet. https://doi.org/10.3389/fgene.2019.00792 (2019).

Article  PubMed  PubMed Central  Google Scholar 

Ghanim, G. E. et al. Structure of human telomerase holoenzyme with bound telomeric DNA. Nature 593, 449–453 (2021). This paper shows sub-4 Å resolution of the human telomerase holoenzyme bound to telomeric DNA.

CAS  PubMed  PubMed Central  Article  Google Scholar 

Zhang, Q., Kim, N.-K. & Feigon, J. Architecture of human telomerase RNA. Proc. Natl Acad. Sci. USA 108, 20325–20332 (2011).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Zhang, J.-M. & Zou, L. Alternative lengthening of telomeres: from molecular mechanisms to therapeutic outlooks. Cell Biosci. 10, 30 (2020).

PubMed  PubMed Central  Article  Google Scholar 

Sobinoff, A. P. & Pickett, H. A. Mechanisms that drive telomere maintenance and recombination in human cancers. Curr. Opin. Genet. Dev. 60, 25–30 (2020). This paper provides an in-depth review of the current understanding of the ALT mechanism.

CAS  PubMed  Article  Google Scholar 

Jafri, M. A., Ansari, S. A., Alqahtani, M. H. & Shay, J. W. Roles of telomeres and telomerase in cancer, and advances in telomerase-targeted therapies. Genome Med. 8, 69 (2016).

PubMed  PubMed Central  Article  CAS  Google Scholar 

Vulliamy, T. et al. Mutations in the telomerase component NHP2 cause the premature ageing syndrome dyskeratosis congenita. Proc. Natl Acad. Sci. USA 105, 8073–8078 (2008).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Pogacic, V., Dragon, F. & Filipowicz, W. Human H/ACA small nucleolar RNPs and telomerase share evolutionarily conserved proteins NHP2 and NOP10. Mol. Cell Biol. 20, 9028–9040 (2000).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Cerone, M. A., Ward, R. J., Londono-Vallejo, J. A. & Autexier, C. Telomerase RNA mutated in autosomal dyskeratosis congenita reconstitutes a weakly active telomerase enzyme defective in telomere elongation. Cell Cycle 4, 585–589 (2005).

CAS  PubMed  Article  Google Scholar 

Zhong, F. L. et al. TPP1 OB-fold domain controls telomere maintenance by recruiting telomerase to chromosome ends. Cell 150, 481–494 (2012).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Nandakumar, J. et al. The TEL patch of telomere protein TPP1 mediates telomerase recruitment and processivity. Nature 492, 285–289 (2012).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Chen, L. Y., Redon, S. & Lingner, J. The human CST complex is a terminator of telomerase activity. Nature 488, 540–544 (2012).

CAS  PubMed  Article  Google Scholar 

Latrick, C. M. & Cech, T. R. POT1-TPP1 enhances telomerase processivity by slowing primer dissociation and aiding translocation. EMBO J. 29, 924–933 (2010).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Kelleher, C., Kurth, I. & Lingner, J. Human protection of telomeres 1 (POT1) is a negative regulator of telomerase activity in vitro. Mol. Cell Biol. 25, 808–818 (2005).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Smogorzewska, A. & de Lange, T. Regulation of telomerase by telomeric proteins. Annu. Rev. Biochem. 73, 177–208 (2004).

CAS  PubMed  Article  Google Scholar 

Ye, J. Z. & de Lange, T. TIN2 is a tankyrase 1 PARP modulator in the TRF1 telomere length control complex. Nat. Genet. 36, 618–623 (2004).

CAS  PubMed  Article  Google Scholar 

Lee, S. S., Bohrson, C., Pike, A. M., Wheelan, S. J. & Greider, C. W. ATM kinase is required for telomere elongation in mouse and human cells. Cell Rep. 13, 1623–1632 (2015).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Leteurtre, F., Li, X., Gluckman, E. & Carosella, E. D. Telomerase activity during the cell cycle and in gamma-irradiated hematopoietic cells. Leukemia 11, 1681–1689 (1997).

CAS  PubMed  Article  Google Scholar 

Schmidt, J. C., Zaug, A. J. & Cech, T. R. Live cell imaging reveals the dynamics of telomerase recruitment to telomeres. Cell 166, 1188–1197.e9 (2016).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Sarek, G., Vannier, J. B., Panier, S., Petrini, J. H. J. & Boulton, S. J. TRF2 recruits RTEL1 to telomeres in S phase to promote t-loop unwinding. Mol. Cell 57, 622–635 (2015).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Zhao, Y. et al. Telomere extension occurs at most chromosome ends and is uncoupled from fill-in in human cancer cells. Cell 138, 463–475 (2009).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Barthel, F. P. et al. Systematic analysis of telomere length and somatic alterations in 31 cancer types. Nat. Genet. 49, 349–357 (2017).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Hiyama, E. & Hiyama, K. Telomere and telomerase in stem cells. Br. J. Cancer 96, 1020–1024 (2007).

CAS  PubMed  PubMed Central  Article  Google Scholar 

Blasco, M. A., Funk, W., Villeponteau, B. & Greider, C. W. Functional characterisation and developmental regulation of mouse telomerase RNA. Science 269, 1267–1270 (1995).

CAS  PubMed  Article