Telomeric DNA in ALT Cells Is Characterized by Free Telomeric Circles and Heterogeneous t-Loops

REFERENCES

• Bechter, O. E., Shay J. W., and Wright W. E.. 2004. The frequency of homologous recombination in human ALT cells. Cell Cycle 3:547–549.
   PubMedGoogle Scholar
• Bechter, O. E., Zou Y., Shay J. W., and Wright W. E.. 2003. Homologous recombination in human telomerase-positive and ALT cells occurs with the same frequency. EMBO Rep. 4:1138–1143.
   PubMed Web of Science ®Google Scholar
• Bechter, O. E., Zou Y., Walker W., Wright W. E., and Shay J. W.. 2004. Telomeric recombination in mismatch repair deficient human colon cancer cells after telomerase inhibition. Cancer Res. 64:3444–3451.
   PubMed Web of Science ®Google Scholar
• Bodnar, A. G., Ouellette M., Frolkis M., Holt S. E., Chiu C. P., Morin G. B., Harley C. B., Shay J. W., Lichtsteiner S., and Wright W. E.. 1998. Extension of life-span by introduction of telomerase into normal human cells. Science 279:349–352.
   PubMed Web of Science ®Google Scholar
• Bryan, T. M., Englezou A., Dalla-Pozza L., Dunham M. A., and Reddel R. R.. 1997. Evidence for an alternative mechanism for maintaining telomere length in human tumors and tumor-derived cell lines. Nat. Med. 3:1271–1274.
   PubMed Web of Science ®Google Scholar
• Bryan, T. M., Englezou A., Gupta J., Bacchetti S., and Reddel R. R.. 1995. Telomere elongation in immortal human cells without detectable telomerase activity. EMBO J. 14:4240–4248.
   PubMed Web of Science ®Google Scholar
• Cesare, A. J., Quinney N., Willcox S., Subramanian D., and Griffith J. D.. 2003. Telomere looping in P. sativum (common garden pea). Plant J. 36:271–279.
   PubMedGoogle Scholar
• Cohen, S., and Mechali M.. 2002. Formation of extrachromosomal circles from telomeric DNA in Xenopus laevis. EMBO Rep. 3:1168–1174.
   PubMedGoogle Scholar
• Counter, C. M., Avilion A. A., LeFeuvre C. E., Stewart N. G., Greider C. W., Harley C. B., and Bacchetti S.. 1992. Telomere shortening associated with chromosome instability is arrested in immortal cells which express telomerase activity. EMBO J. 11:1921–1929.
   PubMed Web of Science ®Google Scholar
• de Lange, T., Shiue L., Myers R. M., Cox D. R., Naylor S. L., Killery A. M., and Varmus H. E.. 1990. Structure and variability of human chromosome ends. Mol. Cell. Biol. 10:518–527.
   PubMed Web of Science ®Google Scholar
• Dunham, M. A., Neumann A. A., Fasching C. L., and Reddel R. R.. 2000. Telomere maintenance by recombination in human cells. Nat. Genet. 26:447–450.
   PubMed Web of Science ®Google Scholar
• Girardi, A. J., Jensen F. C., and Koprowski H.. 1965. SV40-induced transformation of human diploid cells: crisis and recovery. J. Cell Physiol. 65:69–83.
   Google Scholar
• Griffith, J. D., Comeau L., Rosenfield S., Stansel R. M., Bianchi A., Moss H., and de Lange T.. 1999. Mammalian telomeres end in a large duplex loop. Cell 97:503–514.
   PubMed Web of Science ®Google Scholar
• Harley, C. B., Futcher A. B., and Greider C. W.. 1990. Telomeres shorten during ageing of human fibroblasts. Nature 345:458–460.
   PubMed Web of Science ®Google Scholar
• Harley, C. B., Kim N. W., Prowse K. R., Weinrich S. L., Hirsch K. S., West M. D., Bacchetti S., Hirte H. W., Counter C. M., Greider C. W., et al. 1994. Telomerase, cell immortality, and cancer. Cold Spring Harbor Symp. Quant. Biol. 59:307–315.
   PubMedGoogle Scholar
• Henson, J. D., Neumann A. A., Yeager T. R., and Reddel R. R.. 2002. Alternative lengthening of telomeres in mammalian cells. Oncogene 21:598–610.
   PubMed Web of Science ®Google Scholar
• Herbig, U., Jobling W. A., Chen B. P., Chen D. J., and Sedivy J. M.. 2004. 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.
   PubMed Web of Science ®Google Scholar
• Hirt, B. 1967. Selective extraction of polyoma DNA from infected mouse cell cultures. J. Mol. Biol. 26:365–369.
   PubMed Web of Science ®Google Scholar
• Kim, N. W., Piatyszek M. A., Prowse K. R., Harley C. B., West M. D., Ho P. L., Coviello G. M., Wright W. E., Weinrich S. L., and Shay J. W.. 1994. Specific association of human telomerase activity with immortal cells and cancer. Science 266:2011–2015.
   PubMed Web of Science ®Google Scholar
• Londono-Vallejo, J. A., Der-Sarkissian H., Cazes L., Bacchetti S., and Reddel R. R.. 2004. Alternative lengthening of telomeres is characterized by high rates of telomeric exchange. Cancer Res. 64:2324–2327.
   PubMed Web of Science ®Google Scholar
• Lundblad, V., and Blackburn E. H.. 1993. An alternative pathway for yeast telomere maintenance rescues est1-senescence. Cell 73:347–360.
   PubMed Web of Science ®Google Scholar
• Lustig, A. J. 2003. Clues to catastrophic telomere loss in mammals from yeast telomere rapid deletion. Nat. Rev. Genet. 4:916–923.
   PubMedGoogle Scholar
• Makarov, V. L., Hirose Y., and Langmore J. P.. 1997. Long G tails at both ends of human chromosomes suggest a C strand degradation mechanism for telomere shortening. Cell 88:657–666.
   PubMed Web of Science ®Google Scholar
• Masutomi, K., Yu E. Y., Khurts S., Ben-Porath I., Currier J. L., Metz G. B., Brooks M. W., Kaneko S., Murakami S., DeCaprio J. A., Weinberg R. A., Stewart S. A., and Hahn W. C.. 2003. Telomerase maintains telomere structure in normal human cells. Cell 114:241–253.
   PubMed Web of Science ®Google Scholar
• McEachern, M. J., and Blackburn E. H.. 1996. Cap-prevented recombination between terminal telomeric repeat arrays (telomere CPR) maintains telomeres in Kluyveromyces lactis lacking telomerase. Genes Dev. 10:1822–1834.
   PubMed Web of Science ®Google Scholar
• Moyzis, R. K., Buckingham J. M., Cram L. S., Dani M., Deaven L. L., Jones M. D., Meyne J., Ratliff R. L., and Wu J. R.. 1988. A highly conserved repetitive DNA sequence, (TTAGGG)n, present at the telomeres of human chromosomes. Proc. Natl. Acad. Sci. USA 85:6622–6626.
   PubMed Web of Science ®Google Scholar
• Munoz-Jordan, J. L., Cross G. A., de Lange T., and Griffith J. D.. 2001. T-loops at trypanosome telomeres. EMBO J. 20:579–588.
   PubMedGoogle Scholar
• Murti, K. G., and Prescott D. M.. 1999. Telomeres of polytene chromosomes in a ciliated protozoan terminate in duplex DNA loops. Proc. Natl. Acad. Sci. USA 96:14436–14439.
   PubMedGoogle Scholar
• Natarajan, S., Groff-Vindman C., and McEachern M. J.. 2003. Factors influencing the recombinational expansion and spread of telomeric tandem arrays in Kluyveromyces lactis. Eukaryot. Cell 2:1115–1127.
   PubMedGoogle Scholar
• Natarajan, S., and McEachern M. J.. 2002. Recombinational telomere elongation promoted by DNA circles. Mol. Cell. Biol. 22:4512–4521.
   PubMed Web of Science ®Google Scholar
• Nikitina, T., and Woodcock C. L.. 2004. Closed chromatin loops at the ends of chromosomes. J. Cell Biol. 166:161–165.
   PubMed Web of Science ®Google Scholar
• Ogino, H., Nakabayashi K., Suzuki M., Takahashi E., Fujii M., Suzuki T., and Ayusawa D.. 1998. Release of telomeric DNA from chromosomes in immortal human cells lacking telomerase activity. Biochem. Biophys. Res. Commun. 248:223–227.
   PubMedGoogle Scholar
• Olovnikov, A. M. 1973. A theory of marginotomy. The incomplete copying of template margin in enzymic synthesis of polynucleotides and biological significance of the phenomenon. J. Theor. Biol. 41:181–190.
   PubMed Web of Science ®Google Scholar
• Perrem, K., Bryan T. M., Englezou A., Hackl T., Moy E. L., and Reddel R. R.. 1999. Repression of an alternative mechanism for lengthening of telomeres in somatic cell hybrids. Oncogene 18:3383–3390.
   PubMedGoogle Scholar
• Perrem, K., Colgin L. M., Neumann A. A., Yeager T. R., and Reddel R. R.. 2001. Coexistence of alternative lengthening of telomeres and telomerase in hTERT-transfected GM847 cells. Mol. Cell. Biol. 21:3862–3875.
   PubMed Web of Science ®Google Scholar
• Regev, A., Cohen S., Cohen E., Bar-Am I., and Lavi S.. 1998. Telomeric repeats on small polydisperse circular DNA (spcDNA) and genomic instability. Oncogene 17:3455–3461.
   PubMedGoogle Scholar
• Shay, J. W., and Bacchetti S.. 1997. A survey of telomerase activity in human cancer. Eur. J. Cancer 33:787–791.
   PubMed Web of Science ®Google Scholar
• Smogorzewska, A., and de Lange T.. 2002. Different telomere damage signaling pathways in human and mouse cells. EMBO J. 21:4338–4348.
   PubMed Web of Science ®Google Scholar
• Stewart, S. A., Hahn W. C., O'Connor B. F., Banner E. N., Lundberg A. S., Modha P., Mizuno H., Brooks M. W., Fleming M., Zimonjic D. B., Popescu N. C., and Weinberg R. A.. 2002. Telomerase contributes to tumorigenesis by a telomere length-independent mechanism. Proc. Natl. Acad. Sci. USA 99:12606–12611.
   PubMed Web of Science ®Google Scholar
• Teng, S. C., and Zakian V. A.. 1999. Telomere-telomere recombination is an efficient bypass pathway for telomere maintenance in Saccharomyces cerevisiae. Mol. Cell. Biol. 19:8083–8093.
   PubMed Web of Science ®Google Scholar
• Tokutake, Y., Matsumoto T., Watanabe T., Maeda S., Tahara H., Sakamoto S., Niida H., Sugimoto M., Ide T., and Furuichi Y.. 1998. Extra-chromosomal telomere repeat DNA in telomerase-negative immortalized cell lines. Biochem. Biophys. Res. Commun. 247:765–772.
   PubMedGoogle Scholar
• Tomaska, L., McEachern M. J., and Nosek J.. 2004. Alternatives to telomerase: keeping linear chromosomes via telomeric circles. FEBS Lett. 567:142–146.
   PubMedGoogle Scholar
• Tomaska, L., Nosek J., Makhov A. M., Pastorakova A., and Griffith J. D.. 2000. Extragenomic double-stranded DNA circles in yeast with linear mitochondrial genomes: potential involvement in telomere maintenance. Nucleic Acids Res. 28:4479–4487.
   PubMedGoogle Scholar
• von Zglinicki, T., Saretzki G., Docke W., and Lotze C.. 1995. Mild hyperoxia shortens telomeres and inhibits proliferation of fibroblasts: a model for senescence? Exp. Cell Res. 220:186–193.
   PubMed Web of Science ®Google Scholar
• Watson, J. D. 1972. Origin of concatemeric T7 DNA. Nat. New Biol. 239:197–201.
   PubMedGoogle Scholar
• Wright, W. E., Tesmer V. M., Huffman K. E., Levene S. D., and Shay J. W.. 1997. Normal human chromosomes have long G-rich telomeric overhangs at one end. Genes Dev. 11:2801–2809.
   PubMed Web of Science ®Google Scholar
• Yeager, T. R., Neumann A. A., Englezou A., Huschtscha L. I., Noble J. R., and Reddel R. R.. 1999. Telomerase-negative immortalized human cells contain a novel type of promyelocytic leukemia (PML) body. Cancer Res. 59:4175–4179.
   PubMed Web of Science ®Google Scholar