de Lange, T. Shelterin: the protein complex that shapes and safeguards human telomeres. Genes Dev. 19, 2100–2110 (2005).
Riha, K., Heacock, M.L. & Shippen, D.E. The role of the nonhomologous end-joining DNA double-strand break repair pathway in telomere biology. Annu. Rev. Genet. 40, 237–277 (2006).
O'Connor, M.S., Safari, A., Xin, H., Liu, D. & Songyang, Z. A critical role for TPP1 and TIN2 interaction in high-order telomeric complex assembly. Proc. Natl. Acad. Sci. USA 103, 11874–11879 (2006).
Wang, F. et al. The POT–TTP1 telomere complex is a telomerase processivity factor. Nature 445, 506–510 (2007).
d'Adda di Fagagna, F. et al. A DNA damage checkpoint response in telomere-initiated senescence. Nature 426, 194–198 (2003).
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).
Tseng, S.F., Lin, J.J. & Teng, S.C. The telomerase-recruitment domain of the telomere binding protein Cdc13 is regulated by Mec1p/Tel1p-dependent phosphorylation. Nucleic Acids Res. 34, 6327–6336 (2006).
Wu, Y., Xiao, S. & Zhu, X-D. MRE11/RAD50/NBS1 and ATM function as co-mediators of TRF1 in telomere length control. Nat. Struct. Mol. Biol. 14, 832–840 (2007).
Verdun, R.E. & Karlseder, J. Replication and protection of telomeres. Nature 447, 924–931 (2007).
Hector, R.E. et al. Tel1p preferentially associates with short telomeres to stimulate their elongation. Mol. Cell 27, 851–858 (2007).
Verdun, R.E., Crabbe, L., Haggblom, C. & Karlseder, J. Functional human telomeres are recognized as DNA damage in G2 of the cell cycle. Mol. Cell 20, 551–561 (2005).
Verdun, R.E. & Karlseder, J. The DNA damage machinery and homologous recombination pathway act consecutively to protect human telomeres. Cell 127, 709–720 (2006).
Herbig, U., Jobling, W.A., Chen, B.P., Chen, D.J. & Sedivy, J.M. Telomere shortening triggers senescence of human cells through a pathway involving ATM, p53, and p21(CIP1), but not p16(INK4a). Mol. Cell 14, 501–513 (2004).
Celli, G.B. & de Lange, T. DNA processing is not required for ATM-mediated telomere damage response after TRF2 deletion. Nat. Cell Biol. 7, 712–718 (2005).
Baumann, P. & Cech, T.R. Pot1, the putative telomere end-binding protein in fission yeast and humans. Science 292, 1171–1175 (2001).
Hockemeyer, D., Daniels, J.P., Takai, H. & de Lange, T. Recent expansion of the telomeric complex in rodents: two distinct Pot1 proteins protect mouse telomeres. Cell 126, 63–77 (2006).
Churikov, D., Wei, C. & Price, C.M. Vertebrate Pot1 restricts G-overhang length and prevents activation of a telomeric DNA damage checkpoint but is dispensable for overhang protection. Mol. Cell. Biol. 26, 6971–6982 (2006).
Wu, L. et al. Pot1 deficiency initiates DNA damage checkpoint activation and aberrant homologous recombination at telomeres. Cell 126, 49–62 (2006).
Zou, L. Single- and double-stranded DNA: building a trigger of ATR-mediated DNA damage response. Genes Dev. 21, 879–885 (2007).
Sarkaria, J.N. et al. Inhibition of phosphoinositide 3-kinase related kinases by the radiosensitizing agent wortmannin. Cancer Res. 58, 4375–4382 (1998).
Bartek, J. & Lukas, J. Chk1 and Chk2 kinases in checkpoint control and cancer. Cancer Cell 3, 421–429 (2003).
Kohn, E.A., Yoo, C.J. & Eastman, A. The protein kinase C inhibitor Go6976 is a potent inhibitor of DNA damage-induced S and G2 cell cycle checkpoints. Cancer Res. 63, 31–35 (2003).
Zachos, G., Rainey, M.D. & Gillespie, D.A. Chk1-dependent S-M checkpoint delay in vertebrate cells is linked to maintenance of viable replication structures. Mol. Cell. Biol. 25, 563–574 (2005).
Gaestel, M. MAPKAP kinases—MKs—two's company, three's a crowd. Nat. Rev. Mol. Cell Biol. 7, 120–130 (2006).
Reinhardt, H.C., Aslanian, A.S., Lees, J.A. & Yaffe, M.B. p53-deficient cells rely on ATM- and ATR-mediated checkpoint signaling through the p38MAPK/MK2 pathway for survival after DNA damage. Cancer Cell 11, 175–189 (2007).
Jazayeri, A. et al. ATM- and cell cycle-dependent regulation of ATR in response to DNA double-strand breaks. Nat. Cell Biol. 8, 37–45 (2006).
Takao, N. et al. Disruption of ATM in p53-null cells causes multiple functional abnormalities in cellular response to ionizing radiation. Oncogene 18, 7002–7009 (1999).
Dodson, H. et al. Centrosome amplification induced by DNA damage occurs during a prolonged G2 phase and involves ATM. EMBO J. 23, 3864–3873 (2004).
Chen, Y.S. & Chk, Y. 1 in the DNA damage response: conserved roles from yeasts to mammals. DNA Repair (Amst.) 3, 1025–1032 (2004).
Wei, C., Skopp, R., Takata, M., Takeda, S. & Price, C.M. Effects of double-strand break repair proteins on vertebrate telomere structure. Nucleic Acids Res. 30, 2862–2870 (2002).
Kim, S.H., Kaminker, P. & Campisi, J. TIN2, a new regulator of telomere length in human cells. Nat. Genet. 23, 405–412 (1999).
Loayza, D. & De Lange, T. POT1 as a terminal transducer of TRF1 telomere length control. Nature 423, 1013–1018 (2003).
Smith, S. & de Lange, T. Tankyrase promotes telomere elongation in human cells. Curr. Biol. 10, 1299–1302 (2000).
Li, B. & De Lange, T. Rap1 affects the length and heterogeneity of human telomeres. Mol. Biol. Cell 14, 5060–5068 (2003).
Ye, J.Z. et al. POT1-interacting protein PIP1: a telomere length regulator that recruits POT1 to the TIN2/TRF1 complex. Genes Dev. 18, 1649–1654 (2004).
Liu, D. et al. PTOP interacts with POT1 and regulates its localization to telomeres. Nat. Cell Biol. 6, 673–680 (2004).
van Steensel, B. & de Lange, T. Control of telomere length by the human telomeric protein TRF1. Nature 385, 740–743 (1997).
Jacob, N.K., Lescasse, R., Linger, B.R. & Price, C.M. Tetrahymena Pot1a regulates telomere length and prevents activation of a cell cycle checkpoint. Mol Cell Biol 27, 1592–1601 (2007).
Leung-Pineda, V., Ryan, C.E. & Piwnica-Worms, H. Phosphorylation of Chk1 by ATR is antagonized by a Chk1-regulated protein phosphatase 2A circuit. Mol. Cell. Biol. 26, 7529–7538 (2006).
Lei, M., Zaug, A.J., Podell, E.R. & Cech, T.R. Switching human telomerase on and off with hPOT1 protein in vitro. J. Biol. Chem. 280, 20449–20456 (2005).
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).
Hockemeyer, D., Sfeir, A.J., Shay, J.W., Wright, W.E. & de Lange, T. POT1 protects telomeres from a transient DNA damage response and determines how human chromosomes end. EMBO J. 24, 2667–2678 (2005).
Denchi, E.L. & de Lange, T. Protection of telomeres through independent control of ATM and ATR by TRF2 and POT1. Nature 448, 1068–1071 (2007).
Wei, C. & Price, C.M. Cell cycle localization, dimerization, and binding domain architecture of the telomere protein cPot1. Mol. Cell. Biol. 24, 2091–2102 (2004).
Majka, J.N. & Burgers, P.M. Clamping the Mec1/ATR checkpoint kinase into action. Cell Cycle 6, 1157–1160 (2007).
Karlseder, J. et al. The telomeric protein TRF2 binds the ATM kinase and can inhibit the ATM-dependent DNA damage response. PLoS Biol. 2, E240 (2004).
Smogorzewska, A. & de Lange, T. Regulation of telomerase by telomeric proteins. Annu. Rev. Biochem. 73, 177–208 (2004).
Xin, H. et al. TPP1 is a homolog of ciliate TEBP-beta and interacts with POT1 to recruit telomerase. Nature 445, 559–562 (2007).
Frank, C.J., Hyde, M. & Greider, C.W. Regulation of telomere elongation by the cyclin-dependent kinase CDK1. Mol. Cell 24, 423–432 (2006).
Vodenicharov, M.D. & Wellinger, R.J. DNA degradation at unprotected telomeres in yeast is regulated by the CDK1 (Cdc28/Clb) cell-cycle kinase. Mol. Cell 24, 127–137 (2006).