Telomere length, telomeric proteins and genomic instability during the multistep carcinogenic process

Allsopp, 1992, Telomere length predicts replicative capacity of human fibroblasts, Proc Natl Acad Sci USA, 89, 10114, 10.1073/pnas.89.21.10114 Levy, 1992, Telomere end-replication problem and cell aging, J Mol Biol, 225, 951, 10.1016/0022-2836(92)90096-3 Ancelin, 2002, Targeting assay to study the cis functions of human telomeric proteins: evidence for inhibition of telomerase by TRF1 and for activation of telomere degradation by TRF2, Mol Cell Biol, 22, 3474, 10.1128/MCB.22.10.3474-3487.2002 Callen, 2004, Telomere dysfunction in genome instability syndromes, Mutat Res, 567, 85, 10.1016/j.mrrev.2004.06.003 Hara, 1991, Cooperative effect of antisense-Rb and antisense-p53 oligomers on the extension of life span in human diploid fibroblasts, TIG-1, Biochem Biophys Res Commun, 179, 528, 10.1016/0006-291X(91)91403-Y Desmaze, 2003, Telomere-driven genomic instability in cancer cells, Cancer Lett, 194, 173, 10.1016/S0304-3835(02)00704-8 de Lange, 2005, Shelterin: the protein complex that shapes and safeguards human telomeres, Genes Dev, 19, 2100, 10.1101/gad.1346005 Soria, 2003, Chemoprevention of lung cancer, Lancet Oncol, 4, 659, 10.1016/S1470-2045(03)01244-0 Hong, 1997, Recent advances in chemoprevention of cancer, Science, 278, 1073, 10.1126/science.278.5340.1073 Mao, 1998, Phenotype and genotype of advanced premalignant head and neck lesions after chemopreventive therapy, J Natl Cancer Inst, 90, 1545, 10.1093/jnci/90.20.1545 Aaltonen, 1996, The multistep process of colon carcinogenesis, Cytokines Mol Ther, 2, 111 Schedin, 2004, Multistep tumorigenesis and the microenvironment, Breast Cancer Res, 6, 93, 10.1186/bcr772 Hanahan, 2000, The hallmarks of cancer, Cell, 100, 57, 10.1016/S0092-8674(00)81683-9 Henderson, 1989, An overhanging 3′ terminus is a conserved feature of telomeres, Mol Cell Biol, 9, 345, 10.1128/MCB.9.1.345 Griffith, 1999, Mammalian telomeres end in a large duplex loop, Cell, 97, 503, 10.1016/S0092-8674(00)80760-6 Makarov, 1993, Nucleosomal organization of telomere-specific chromatin in rat, Cell, 73, 775, 10.1016/0092-8674(93)90256-P Lejnine, 1995, Conserved nucleoprotein structure at the ends of vertebrate and invertebrate chromosomes, Proc Natl Acad Sci USA, 92, 2393, 10.1073/pnas.92.6.2393 Campisi, 2001, Cellular senescence, cancer and aging: the telomere connection, Exp Gerontol, 36, 1619, 10.1016/S0531-5565(01)00160-7 Zhong, 1992, A mammalian factor that binds telomeric TTAGGG repeats in vitro, Mol Cell Biol, 12, 4834, 10.1128/MCB.12.11.4834 Bilaud, 1997, Telomeric localization of TRF2, a novel human telobox protein, Nat Genet, 17, 236, 10.1038/ng1097-236 Court, 2005, How the human telomeric proteins TRF1 and TRF2 recognize telomeric DNA: a view from high-resolution crystal structures, EMBO Rep, 6, 39, 10.1038/sj.embor.7400314 Karlseder, 2003, Telomere repeat binding factors: keeping the ends in check, Cancer Lett, 194, 189, 10.1016/S0304-3835(02)00706-1 Bunch, 2005, Distinct requirements for Pot1 in limiting telomere length and maintaining chromosome stability, Mol Cell Biol, 25, 5567, 10.1128/MCB.25.13.5567-5578.2005 Wellinger, 1997, The DNA structures at the ends of eukaryotic chromosomes, Eur J Cancer, 33, 735, 10.1016/S0959-8049(97)00067-1 Verdun, 2005, Functional human telomeres are recognized as DNA damage in G2 of the cell cycle, Mol Cell, 20, 551, 10.1016/j.molcel.2005.09.024 Slijepcevic, 2006, The role of DNA damage response proteins at telomeres–an “integrative” model, DNA Repair (Amst), 5, 1299, 10.1016/j.dnarep.2006.05.038 Shay, 1997, A survey of telomerase activity in human cancer, Eur J Cancer, 33, 787, 10.1016/S0959-8049(97)00062-2 Kolquist, 1998, Expression of TERT in early premalignant lesions and a subset of cells in normal tissues, Nat Genet, 19, 182, 10.1038/554 Herbert, 2001, Telomerase and breast cancer, Breast Cancer Res, 3, 146, 10.1186/bcr288 Yashima, 1998, Telomerase enzyme activity and RNA expression during the multistage pathogenesis of breast carcinoma, Clin Cancer Res, 4, 229 Luzar, 2005, Expression of human telomerase catalytic protein in gall bladder carcinogenesis, J Clin Pathol, 58, 820, 10.1136/jcp.2004.023143 Lantuejoul, 2007, Telomerase expression in lung preneoplasia and neoplasia., Int J Cancer, 120, 1835, 10.1002/ijc.22473 Oh, 2003, Telomere shortening and telomerase reactivation in dysplastic nodules of human hepatocarcinogenesis, J Hepatol, 39, 786, 10.1016/S0168-8278(03)00395-7 Yan, 1999, Telomerase activation in colorectal carcinogenesis, J Pathol, 189, 207, 10.1002/(SICI)1096-9896(199910)189:2<207::AID-PATH424>3.0.CO;2-H Koering, 2002, Human telomeric position effect is determined by chromosomal context and telomeric chromatin integrity, EMBO Rep, 3, 1055, 10.1093/embo-reports/kvf215 Baur, 2001, Telomere position effect in human cells, Science, 292, 2075, 10.1126/science.1062329 Benetti, 2007, Telomere length regulates the epigenetic status of mammalian telomeres and subtelomeres, Nat Genet, 39, 243, 10.1038/ng1952 Fraga, 2005, Loss of acetylation at Lys16 and trimethylation at Lys20 of histone H4 is a common hallmark of human cancer, Nat Genet, 37, 391, 10.1038/ng1531 Blasco, 2007, The epigenetic regulation of mammalian telomeres, Nat Rev Genet, 8, 299, 10.1038/nrg2047 Gupta, 2005, Involvement of human MOF in ATM function, Mol Cell Biol, 25, 5292, 10.1128/MCB.25.12.5292-5305.2005 Meeker, 2004, Telomere length abnormalities occur early in the initiation of epithelial carcinogenesis, Clin Cancer Res, 10, 3317, 10.1158/1078-0432.CCR-0984-03 Meeker, 2002, Telomere shortening is an early somatic DNA alteration in human prostate tumorigenesis, Cancer Res, 62, 6405 Hansel, 2006, Telomere length variation in biliary tract metaplasia, dysplasia, and carcinoma, Mod Pathol, 10.1038/modpathol.3800591 Meeker, 2004, Telomere shortening occurs in subsets of normal breast epithelium as well as in situ and invasive carcinoma, Am J Pathol, 164, 925, 10.1016/S0002-9440(10)63180-X Lantuejoul, 2005, Telomere shortening and telomerase reverse transcriptase expression in preinvasive bronchial lesions, Clin Cancer Res, 11, 2074, 10.1158/1078-0432.CCR-04-1376 Chin, 2004, In situ analyses of genome instability in breast cancer, Nat Genet, 36, 984, 10.1038/ng1409 Vukovic, 2003, Evidence of multifocality of telomere erosion in high-grade prostatic intraepithelial neoplasia (HPIN) and concurrent carcinoma, Oncogene, 22, 1978, 10.1038/sj.onc.1206227 Kondo, 2004, Expression of POT1 is associated with tumor stage and telomere length in gastric carcinoma, Cancer Res, 64, 523, 10.1158/0008-5472.CAN-03-1196 Loayza, 2003, POT1 as a terminal transducer of TRF1 telomere length control, Nature, 423, 1013, 10.1038/nature01688 Yang, 2005, POT1 and TRF2 cooperate to maintain telomeric integrity, Mol Cell Biol, 25, 1070, 10.1128/MCB.25.3.1070-1080.2005 Nakanishi, 2003, Expression of mRNAs for telomeric repeat binding factor (TRF)-1 and TRF2 in atypical adenomatous hyperplasia and adenocarcinoma of the lung, Clin Cancer Res, 9, 1105 Oh, 2005, Up-regulation of telomere-binding proteins, TRF1, TRF2, and TIN2 is related to telomere shortening during human multistep hepatocarcinogenesis, Am J Pathol, 166, 73, 10.1016/S0002-9440(10)62233-X Garcia-Aranda, 2006, Correlations of telomere length, telomerase activity, and telomeric-repeat binding factor 1 expression in colorectal carcinoma, Cancer, 106, 541, 10.1002/cncr.21625 Matsutani, 2001, Expression of telomeric repeat binding factor 1 and 2 and TRF1-interacting nuclear protein 2 in human gastric carcinomas, Int J Oncol, 19, 507 Smogorzewska, 2000, Control of human telomere length by TRF1 and TRF2, Mol Cell Biol, 20, 1659, 10.1128/MCB.20.5.1659-1668.2000 Saito, 2002, Gene expression for suppressors of telomerase activity (telomeric-repeat binding factors) in breast cancer, Jpn J Cancer Res, 93, 253, 10.1111/j.1349-7006.2002.tb02166.x Kishi, 2001, Telomeric protein Pin2/TRF1 induces mitotic entry and apoptosis in cells with short telomeres and is down-regulated in human breast tumors, Oncogene, 20, 1497, 10.1038/sj.onc.1204229 Yamada, 2002, Down-regulation of TRF1, TRF2 and TIN2 genes is important to maintain telomeric DNA for gastric cancers, Anticancer Res, 22, 3303 Miyachi, 2002, Correlation between telomerase activity and telomeric-repeat binding factors in gastric cancer, J Exp Clin Cancer Res, 21, 269 Blackburn, 1999, The telomere and telomerase: how do they interact?, Mt Sinai J Med, 66, 292 van Steensel, 1998, TRF2 protects human telomeres from end-to-end fusions, Cell, 92, 401, 10.1016/S0092-8674(00)80932-0 Ancelin, 1998, Role of the telomeric DNA-binding protein TRF2 in the stability of human chromosome ends, Bioessays, 20, 879, 10.1002/(SICI)1521-1878(199811)20:11<879::AID-BIES2>3.0.CO;2-I Smith, 2000, Tankyrase promotes telomere elongation in human cells, Curr Biol, 10, 1299, 10.1016/S0960-9822(00)00752-1 Hsiao SJ, Smith S, Tankyrase function at telomeres, spindle poles, and beyond. Biochimie; 2007. Marcand, 1997, Rap1p and telomere length regulation in yeast, Ciba Found Symp, 211, 76 Hardy, 1992, A RAP1-interacting protein involved in transcriptional silencing and telomere length regulation, Genes Dev, 6, 801, 10.1101/gad.6.5.801 Wotton, 1997, A novel Rap1p-interacting factor, Rif2p, cooperates with Rif1p to regulate telomere length in Saccharomyces cerevisiae, Genes Dev, 11, 748, 10.1101/gad.11.6.748 Teixeira, 2004, Telomere length homeostasis is achieved via a switch between telomerase-extendible and -nonextendible states, Cell, 117, 323, 10.1016/S0092-8674(04)00334-4 Loayza, 2004, Telomerase regulation at the telomere: a binary switch, Cell, 117, 279, 10.1016/S0092-8674(04)00409-X Wang, 2007, The POT1-TPP1 telomere complex is a telomerase processivity factor, Nature, 445, 506, 10.1038/nature05454 Kim, 1999, TIN2, a new regulator of telomere length in human cells, Nat Genet, 23, 405, 10.1038/70508 Kim, 2004, TIN2 mediates functions of TRF2 at human telomeres, J Biol Chem, 279, 43799, 10.1074/jbc.M408650200 Zhou, 2001, The Pin2/TRF1-interacting protein PinX1 is a potent telomerase inhibitor, Cell, 107, 347, 10.1016/S0092-8674(01)00538-4 Banik, 2004, Characterization of interactions between PinX1 and human telomerase subunits hTERT and hTR, J Biol Chem, 279, 51745, 10.1074/jbc.M408131200 Zhou, 2000, The DNA damage response: putting checkpoints in perspective, Nature, 408, 433, 10.1038/35044005 Khanna, 2001, DNA double-strand breaks: signaling, repair and the cancer connection, Nat Genet, 27, 247, 10.1038/85798 Bartek, 2003, Chk1 and Chk2 kinases in checkpoint control and cancer, Cancer Cell, 3, 421, 10.1016/S1535-6108(03)00110-7 Gorgoulis, 2005, Activation of the DNA damage checkpoint and genomic instability in human precancerous lesions, Nature, 434, 907, 10.1038/nature03485 Bartkova, 2005, DNA damage response as a candidate anti-cancer barrier in early human tumorigenesis, Nature, 434, 864, 10.1038/nature03482 D’Amours, 2002, The Mre11 complex: at the crossroads of DNA repair and checkpoint signalling, Nat Rev Mol Cell Biol, 3, 317, 10.1038/nrm805 Uziel, 2003, Requirement of the MRN complex for ATM activation by DNA damage, Embo J, 22, 5612, 10.1093/emboj/cdg541 Weizman, 2003, Contribution of the Atm protein to maintaining cellular homeostasis evidenced by continuous activation of the AP-1 pathway in Atm-deficient brains, J Biol Chem, 278, 6741, 10.1074/jbc.M211168200 d’Adda di Fagagna, 2003, A DNA damage checkpoint response in telomere-initiated senescence, Nature, 426, 194, 10.1038/nature02118 Lustig, 1986, Identification of yeast mutants with altered telomere structure, Proc Natl Acad Sci USA, 83, 1398, 10.1073/pnas.83.5.1398 Hande, 2001, Extra-chromosomal telomeric DNA in cells from Atm (−/−) mice and patients with ataxia-telangiectasia, Hum Mol Genet, 10, 519, 10.1093/hmg/10.5.519 Hollstein, 1991, p53 mutations in human cancers, Science, 253, 49, 10.1126/science.1905840 Kastan, 2004, Cell-cycle checkpoints and cancer, Nature, 432, 316, 10.1038/nature03097 Halazonetis, 2004, Constitutively active DNA damage checkpoint pathways as the driving force for the high frequency of p53 mutations in human cancer, DNA Repair (Amst), 3, 1057, 10.1016/j.dnarep.2004.03.036 Graeber, 1996, Hypoxia-mediated selection of cells with diminished apoptotic potential in solid tumours, Nature, 379, 88, 10.1038/379088a0 Takai, 2003, DNA damage foci at dysfunctional telomeres, Curr Biol, 13, 1549, 10.1016/S0960-9822(03)00542-6 van Steensel, 1997, Control of telomere length by the human telomeric protein TRF1, Nature, 385, 740, 10.1038/385740a0 de Lange, 2002, Protection of mammalian telomeres, Oncogene, 21, 532, 10.1038/sj.onc.1205080 Karlseder, 1999, p53- and ATM-dependent apoptosis induced by telomeres lacking TRF2, Science, 283, 1321, 10.1126/science.283.5406.1321 Celli, 2005, DNA processing is not required for ATM-mediated telomere damage response after TRF2 deletion, Nat Cell Biol, 7, 712, 10.1038/ncb1275 Karlseder, 2004, The telomeric protein TRF2 binds the ATM kinase and can inhibit the ATM-dependent DNA damage response, PLoS Biol, 2, E240, 10.1371/journal.pbio.0020240 d’Adda di Fagagna, 2001, Effects of DNA nonhomologous end-joining factors on telomere length and chromosomal stability in mammalian cells, Curr Biol, 11, 1192, 10.1016/S0960-9822(01)00328-1 Song, 2001, Human Ku70 interacts with heterochromatin protein 1alpha, J Biol Chem, 276, 8321, 10.1074/jbc.M008779200 Jaco, 2004, Role of human Ku86 in telomere length maintenance and telomere capping, Cancer Res, 64, 7271, 10.1158/0008-5472.CAN-04-1381 Espejel, 2002, Mammalian Ku86 mediates chromosomal fusions and apoptosis caused by critically short telomeres, Embo J, 21, 2207, 10.1093/emboj/21.9.2207 Goytisolo, 2001, The absence of the dna-dependent protein kinase catalytic subunit in mice results in anaphase bridges and in increased telomeric fusions with normal telomere length and G-strand overhang, Mol Cell Biol, 21, 3642, 10.1128/MCB.21.11.3642-3651.2001 Espejel, 2002, Functional interaction between DNA-PKcs and telomerase in telomere length maintenance, Embo J, 21, 6275, 10.1093/emboj/cdf593 Zhu, 2000, Cell-cycle-regulated association of RAD50/MRE11/NBS1 with TRF2 and human telomeres, Nat Genet, 25, 347, 10.1038/77139 Ranganathan, 2001, Rescue of a telomere length defect of Nijmegen breakage syndrome cells requires NBS and telomerase catalytic subunit, Curr Biol, 11, 962, 10.1016/S0960-9822(01)00267-6 Meeker, 2002, Telomere length assessment in human archival tissues: combined telomere fluorescence in situ hybridization and immunostaining, Am J Pathol, 160, 1259, 10.1016/S0002-9440(10)62553-9 Wiltshire, 1995, Direct visualization of the clonal progression of primary cutaneous melanoma: application of tissue microdissection and comparative genomic hybridization, Cancer Res, 55, 3954 du Manoir, 1993, Detection of complete and partial chromosome gains and losses by comparative genomic in situ hybridization, Hum Genet, 90, 590, 10.1007/BF00202476 Ried, 1995, Comparative genomic hybridization of formalin-fixed, paraffin-embedded breast tumors reveals different patterns of chromosomal gains and losses in fibroadenomas and diploid and aneuploid carcinomas, Cancer Res, 55, 5415 Speicher, 1993, Molecular cytogenetic analysis of formalin-fixed, paraffin-embedded solid tumors by comparative genomic hybridization after universal DNA-amplification, Hum Mol Genet, 2, 1907, 10.1093/hmg/2.11.1907 Speicher, 1995, Correlation of microscopic phenotype with genotype in a formalin-fixed, paraffin-embedded testicular germ cell tumor with universal DNA amplification, comparative genomic hybridization, and interphase cytogenetics, Am J Pathol, 146, 1332 Klein, 1999, Comparative genomic hybridization, loss of heterozygosity, and DNA sequence analysis of single cells, Proc Natl Acad Sci USA, 96, 4494, 10.1073/pnas.96.8.4494 Barrett, 2004, Comparative genomic hybridization using oligonucleotide microarrays and total genomic DNA, Proc Natl Acad Sci USA, 101, 17765, 10.1073/pnas.0407979101 Carvalho, 2004, High resolution microarray comparative genomic hybridisation analysis using spotted oligonucleotides, J Clin Pathol, 57, 644, 10.1136/jcp.2003.013029 Lucito, 2003, Representational oligonucleotide microarray analysis: a high-resolution method to detect genome copy number variation, Genome Res, 13, 2291, 10.1101/gr.1349003 Zhao, 2004, An integrated view of copy number and allelic alterations in the cancer genome using single nucleotide polymorphism arrays, Cancer Res, 64, 3060, 10.1158/0008-5472.CAN-03-3308 Pinkel, 1998, High-resolution analysis of DNA copy number variation using comparative genomic hybridization to microarrays, Nat Genet, 20, 207, 10.1038/2524 Davies, 2005, Array CGH technologies and their applications to cancer genomes, Chromosome Res, 13, 237, 10.1007/s10577-005-2168-x O’Hagan, 2002, Telomere dysfunction provokes regional amplification and deletion in cancer genomes, Cancer Cell, 2, 149, 10.1016/S1535-6108(02)00094-6 Rudolph, 1999, Longevity, stress response, and cancer in aging telomerase-deficient mice, Cell, 96, 701, 10.1016/S0092-8674(00)80580-2 Gisselsson, 2002, Centrosomal abnormalities, multipolar mitoses, and chromosomal instability in head and neck tumours with dysfunctional telomeres, Br J Cancer, 87, 202, 10.1038/sj.bjc.6600438 der-Sarkissian, 2004, The shortest telomeres drive karyotype evolution in transformed cells, Oncogene, 23, 1221, 10.1038/sj.onc.1207152 Soler, 2005, Telomere dysfunction drives chromosomal instability in human mammary epithelial cells, Genes Chromosomes Cancer, 44, 339, 10.1002/gcc.20244 Graakjaer, 2004, The relative lengths of individual telomeres are defined in the zygote and strictly maintained during life, Aging Cell, 3, 97, 10.1111/j.1474-9728.2004.00093.x Sabatier, 2005, The loss of a single telomere can result in instability of multiple chromosomes in a human tumor cell line, Mol Cancer Res, 3, 139, 10.1158/1541-7786.MCR-04-0194 Martens, 2000, Accumulation of short telomeres in human fibroblasts prior to replicative senescence, Exp Cell Res, 256, 291, 10.1006/excr.2000.4823 Lansdorp, 1996, Heterogeneity in telomere length of human chromosomes, Hum Mol Genet, 5, 685, 10.1093/hmg/5.5.685 Londono-Vallejo, 2004, Telomere length heterogeneity and chromosome instability, Cancer Lett, 212, 135, 10.1016/j.canlet.2004.05.008 Olaussen, 2006, Telomeres and telomerase as targets for anticancer drug development, Crit Rev Oncol Hematol, 57, 191, 10.1016/j.critrevonc.2005.08.007