The roles of tolemeres and telomerase in cell life span

Mutation Research - Reviews in Genetic Toxicology

Volumn 366, Issue 1, 1996, Pages 45-63

  Counter, Christopher M ^a  

^a WHITEHEAD INSTITUTE FOR BIOMEDICAL RESEARCH   (United States)

Author keywords

Cell life span; Telomer; Telomerase

Indexed keywords

TELOMERASE; TRANSFERASE;

CELL CYCLE; CELL DIVISION; CELL SURVIVAL; DNA REPLICATION; DNA SEQUENCE; ENZYME ACTIVITY; HUMAN; NONHUMAN; PRIORITY JOURNAL; REVIEW; SENESCENCE; TELOMERE; ANIMAL; CELL AGING; CILIATE; CYTOLOGY; ENZYMOLOGY; PHYSIOLOGY; SACCHAROMYCES CEREVISIAE;

CILIOPHORA; EUKARYOTA;

ANIMALS; CELL AGING; CELL SURVIVAL; CILIOPHORA; HUMANS; SACCHAROMYCES CEREVISIAE; TELOMERASE; TELOMERE;

EID: 0030271719     PISSN: 01651110     EISSN: None     Source Type: Journal    
DOI: 10.1016/S0165-1110(96)90006-8     Document Type: Article

References (178)

1. Blackburn E.H., Greider C.W. Telomeres. 1995;Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
2. Henderson E. Telomere DNA structure. Blackburn E.H., Greider C.W. Telomeres. 1995;11-34 Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
3. Henderson E.R., Blackburn E.H. An overhanging 3′ terminus is a conserved feature of telomeres. Mol. Cell Biol. 9:1989;345-348.
4. Klobutcher L.A., Swanton M.T., Donini P., Prescott D.M. All gene-sized DNA molecules in four species of hypotrichs have the same terminal sequence and an unusual 3′ terminus. Proc. Natl. Acad. Sci. USA. 78:1981;3015-3019.
5. Pluta A.F., Kaine B.P., Spear B.B. The terminal organization of macronuclear DNA in Oxytricha fallax. Nucleic Acids Res. 10:1982;8145-8154.
6. Wellinger R.J., Wolf A.J., Zakian V.A. Saccharomyces telomeres acquire single-strand TG1-3 tails late in S phase. Cell. 72:1993;51-60.
7. Liu Z., Lee A., Gilbert W. Gene disruption of a G4-DNA-dependent nuclease in yeast leads to cellular senescence and telomere shortening. Proc. Natl. Acad. Sci. USA. 92:1995;6002-6006.
8. Blackburn E.H., Chiou S.S. Non-nucleosomal packaging of a tandemly repeated DNA sequence at termini of extrachromosomal DNA coding for rRNA in Tetrahymena. Proc. Natl. Acad. Sci. USA. 78:1981;2263-2267.
9. Edwards C.A., Firtel R.A. Site-specific phasing in the chromatin of the rDNA in Dictyostelium discoideum. J. Mol. Biol. 180:1984;73-90.
10. Gottschling D.E., Cech T.R. Chromatin structure of the molecular ends of Oxytricha macronuclear DNA: phased nucleosomes and a telomeric complex. Cell. 38:1984;501-510.
11. Lucchini R., Pauli U., Braun R., Koller T., Sogo J.M. Structure of the extrachromosomal ribosomal RNA chromatin of Physarum polycephalum. J. Mol. Biol. 196:1987;829-843.
12. Wright J.H., Gottschling D.E., Zakian V.A. Saccharomyces telomeres assume a non-nucleosomal chromatin structure. Genes Dev. 6:1992;197-210.
13. Lejnine S., Makarov V.L., Langmore J.P. Conserved nucleoprotein structure at the ends of vertebrate and invertebrate chromosomes. Proc. Natl. Acad. Sci. USA. 92:1995;2393-2397.
14. Tommerup H., Dousmanis A., de Lange T. Unusual chromatin in human telomeres. Mol. Cell Biol. 14:1994;5777-5785.
15. Biessmann H., Mason J.M. Genetics and molecular biology of telomeres. Adv. Genet. 30:1992;185-249.
16. Blackburn E.H., Szostak J.W. The molecular structure of centromeres and telomeres. Annu. Rev. Biochem. 53:1984;163-194.
17. Zakian V.A. Structure and function of telomeres. Annu. Rev. Genet. 23:1989;579-604.
18. Longtine M.S., Enomoto S., Finstad S.L., Berman J. Yeast telomere repeat sequence (TRS) improves circular plasmid segregation, and TRS plasmid segregation involves the RAP1 gene product. Mol. Cell Biol. 12:1992;1997-2009.
19. Chikashige Y., Ding D.-Q., Funabiki H., Haraguchi T., Mashiko S., Yanagida M., Hiraoka Y. Telomere-led premiotic chromosome movement in fission yeast. Science. 264:1994;270-273.
20. Dernburg A.F., Sedat J.W., Cande W.Z., Bass H.W. Cytology of telomeres. Blackburn E.H., Greider C.W. Telomeres. 1995;295-338 Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
21. Bayne R.A., Broccoli D., Taggart M.H., Thomson E.J., Farr C.J., Cooke H.J. Sandwiching of a gene within 12 kb of a functional telomere and alpha satellite does not result in silencing. Hum. Mol. Genet. 3:1994;539-546.
22. Huang Y.-J., Stoffel R., Tobler H., Mueller F. A newly formed telomere in Ascaris suum does not exhert a telomere position effect on a nearby gene. Mol. Cell. Biol. 16:1996;130-134.
23. Horn D., Cross G.A.M. A developmentally regulated position effect at the telomeric locus in Trypanosoma brucei. Cell. 83:1995;555-561.
24. Karpen G.H. Position-effect variegation and the new biology of heterochromatin. Curr. Opin. Genet. Dev. 4:1994;281-291.
25. Nimmo E.R., Cranston G., Allshire R.C. Telomere-associated chromosome breakage in fission yeast results in variegated expression of adjacent genes. EMBO J. 13:1994;3801-3811.
26. Rudenko G., Blundell P.A., Dirks-Mulder A., Kieft R., Borst P. A ribosomal DNA promoter replacing the promoter of a telomeric VSG gene expression site can be efficiently switched on and off in T. brucei. Cell. 83:1995;547-553.
27. Shore D. Telomere position effects and transcriptional silencing in the yeast Saccharomyces cerevisiae. Blackburn E.H., Greider C.W. Telomeres. 1995;139-192 Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
28. Olovnikov A.M. [Principle of marginotomy in template synthesis of polynucleotides]. Dokl Akad Nauk SSSR. 201:1971;1496-1499.
29. Olovnikov A.M. A theory of marginotomy. The incomplete copying of template margin in enzymic synthesis of polynucleotides and biological significance of the phenomenon. J. Theor. Biol. 41:1973;181-190.
30. Watson J.D. Origin of concatemeric T7 DNA. Nature New Biol. 239:1972;197-201.
31. Lingner J., Cooper J.P., Cech T.R. Telomerase and DNA end replication: no longer a lagging strand problem? Science. 269:1995;1533-1534.
32. Biessmann H., Mason J.M., Ferry K., d'Hulst M., Valgeirsdottir K., Traverse K.L., Pardue M.L. Addition of telomere-associated HeT DNA sequences 'heals' broken chromosome ends in Drosophila. Cell. 61:1990;663-673.
33. Biessmann H., Champion L.E., O'Hair M., Ikenaga K., Kasravi B., Mason J.M. Frequent transpositions of Drosophila melanogaster HeT-A transposable elements to receding chromosome ends. EMBO J. 11:1992;4459-4469.
34. Traverse K.L., Pardue M.L. A spontaneously opened ring chromosome of Drosophila melanogaster has acquired He-T DNA sequences at both new telomeres. Proc. Natl. Acad. Sci. USA. 85:1988;8116-8120.
35. - senescence. Cell. 73:1993;347-360.
36. Bryan T.M., Englezou A., Gupta J., Bacchetti S., Reddel R.R. Telomere elongation in immortal human cells without detectable telomerase activity. EMBO J. 14:1995;4240-4248.
37. Murnane J.P., Sabatier L., Marder B.A., Morgan W.F. Telomere dynamics in an immortal human cell line. EMBO J. 13:1994;4953-4962.
38. Rogan E.M., Bryan T.M., Hukku B., Maclean K., Chang A.C., Moy E.L., Englezou A., Warneford S.G., Dalla-Pozza L., Reddel R.R. Alterations in p53 and p16INK4 expression and telomere length during spontaneous immortalization of Li-Fraumeni syndrome fibroblasts. Mol. Cell. Biol. 15:1995;4745-4753.
39. Greider C.W., Blackburn E.H. Identification of a specific telomere terminal transferase activity in Tetrahymena extracts. Cell. 43:1985;405-413.
40. Greider C.W., Blackburn E.H. A telomeric sequence in the RNA of Tetrahymena telomerase required for telomere repeat synthesis. Nature. 337:1989;331-337.
41. Collins K., Kobayashi R., Greider C.W. Purification of Tetrahymena telomerase and cloning of genes encoding the two protein components of the enzyme. Cell. 81:1995;677-686.
42. Autexier C., Greider C.W. Functional reconstitution of wild-type and mutant Tetrahymena telomerase. Genes Dev. 8:1994;563-575.
43. Yu G.L., Bradley J.D., Attardi L.D., Blackburn E.H. In vivo alteration of telomere sequences and senescence caused by mutated Tetrahymena telomerase RNAs. Nature. 344:1990;126-132.
44. Yu G.L., Blackburn E.H. Developmentally programmed healing of chromosomes by telomerase in Tetrahymena. Cell. 67:1991;823-832.
45. Greider C.W. Telomerase biochemistry and regulation. Blackburn E.H., Greider C.W. Telomeres. 1995;35-68 Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
46. Collins K., Greider C.W. Tetrahymena telomerase catalyzes nucleolytic cleavage and nonprocessive elongation. Genes Dev. 7:1993;1364-1376.
47. Morin G.B. The human telomere terminal transferase enzyme is a ribonucleoprotein that synthesizes TTAGGG repeats. Cell. 59:1989;521-529.
48. Prowse K.R., Avilion A.A., Greider C.W. Identification of a nonprocessive telomerase activity from mouse cells. Proc. Natl. Acad. Sci. USA. 90:1993;1493-1497.
49. Mantell L.L., Greider C.W. Telomerase activity in germline and embryonic cells of Xenopus. EMBO J. 13:1994;3211-3217.
50. Cohn M., Blackburn E.H. Telomerase in yeast. Science. 269:1995;396-400.
51. Lin J.J., Zakian V.A. An in vitro assay for Saccharomyces telomerase requires EST1. Cell. 81:1995;1127-1135.
52. Zahler A.M., Prescott D.M. Telomere terminal transferase activity in the hypotrichous ciliate Oxytricha nova and a model for replication of the ends of linear DNA molecules. Nucleic Acids Res. 16:1988;6953-6972.
53. Shippen-Lentz D., Blackburn E.H. Functional evidence for an RNA template in telomerase. Science. 247:1990;546-552.
54. Lingner J., Hendrick L.L., Cech T.R. Telomerase RNAs of different ciliates have a common secondary structure and a permuted template. Genes Dev. 8:1994;1984-1998.
55. Melek M., Davis B.T., Shippen D.E. Oligonucleotides complementary to the Oxytricha nova telomerase RNA delineate the template domain and uncover a novel mode of primer utilization. Mol. Cell. Biol. 14:1994;7827-7838.
56. Singer M.S., Gottschling D.E. TLC1: template RNA component of Saccharomyces cerevisiae telomerase. Science. 266:1994;404-409.
57. McEachern M.J., Blackburn E.H. Runaway telomere elongation caused by telomerase RNA gene mutations. Nature. 376:1995;403-409.
58. Blasco M.A., Funk W., Villeponteau B., Greider C.W. Functional characterization and developmental regulation of mouse telomerase RNA. Science. 269:1995;1267-1270.
59. Feng J., Funk W.D., Wang S.S., Weinrich S.L., Avilion A.A., Chiu C.P., Adams R.R., Chang E., Allsopp R.C., Yu J., et al. The RNA component of human telomerase. Science. 269:1995;1236-1241.
60. Hayflick L., Moorhead P.S. The serial cultivation of human diploid cell strains. Exp. Cell Res. 25:1961;585-621.
61. Goldstein S. Replicative senescence: the human fibroblast comes of age. Science. 249:1990;1129-1133.
62. Sager R. Senescence as a mode of tumor suppression. Environ. Health Perspect. 93:1991;59-62.
63. Stanulis-Praeger B.M. Cellular senescence revisited: a review. Mech. Ageing. Dev. 38:1987;1-48.
64. Dell'Orco R.T., Mertens J.G., Kruse P.F. Jr. Doubling potential, calendar time, and senescence of human diploid cells in culture. Exp. Cell. Res. 77:1973;356-360.
65. Harley C.B., Goldstein S. Cultured human fibroblasts: distribution of cell generations and a critical limit. J. Cell Physiol. 97:1978;509-516.
66. Hayflick L. The cell biology of human aging. N. Engl. J. Med. 295:1976;1302-1308.
67. Barrett J.C., Annab L.A., Alcorta D., Preston G., Vojta P., Yin Y. Cellular senescence and cancer. Cold Spring Harbour Symp. Quant. Biol. 59:1994;411-418.
68. Dimri G.P., Campisi J. Molecular and cell biology of replicative senescence. Cold Spring Harbour Symp. Quant. Biol. 59:1994;67-73.
69. Shay J.W., Wright W.E., Werbin H. Defining the molecular mechanisms of human cell immortalization. Biochim. Biophys. Acta. 1072:1991;1-7.
70. Yoshida M. HTLV-1 Tax: regulation of gene expression and disease. Trends Microbiol. 1:1993;131-135.
71. Harris C.C. Human tissues and cells in carcinogenesis research. Cancer Res. 47:1987;1-10.
72. Bischoff F.Z., Yim S.O., Pathak S., Grant G., Siciliano M.J., Giovanella B.C., Strong L.C., Tainsky M.A. Spontaneous abnormalities in normal fibroblasts from patients with Li-Fraumeni cancer sydrome: aneuploidy and immortalization. Cancer Res. 50:1990;7979-7984.
73. his). Oncogene. 1996;. in press.
74. Shay J.W., Tomlinson G., Piatyszek M.A., Gollahon L.S. Spontaneous in vitro immortalization of breast epithelial cells from a patient with Li-Fraumeni syndrome. Mol. Cell. Biol. 15:1995;425-432.
75. 40-transformed human cells. J. Cell Comp. Physiol. 62:1963;57-84.
76. Moorhead P.S., Saksela E. The sequence of chromosome aberrations during SV40 transformation of a human diploid cell strain. Hereditas. 52:1965;271-284.
77. Sack G.H. Jr. Human cell transformation by simian virus 40 - a review. In Vitro. 17:1981;1-19.
78. Stewart N., Bacchetti S. Expression of SV40 large T antigen, but not small t antigen, is required for the induction of chromosomal aberrations in transformed human cells. Virology. 180:1991;49-57.
79. 40-infected human fibroblast cultures. Cytogenetics. 3:1964;45-61.
80. Shay J.W., Wright W.E. Quantitation of the frequency of immortalization of normal human diploid fibroblasts by SV40 large T-antigen. Exp. Cell Res. 184:1989;109-118.
81. Strauss M., Griffin B.E. Cellular immortalization - an essential step or merely a risk factor in DNA virus-induced transformation? Cancer Cells. 2:1990;360-365.
82. Rubelj I., Pereira-Smith O.M. SV40-transformed human cells in crisis exhibit changes that occur in normal cellular senescence. Exp. Cell Res. 211:1994;82-89.
83. Band V., Zajchowski D., Kulesa V., Sager R. Human papilloma virus DNAs immortalize normal human mammary epithelial cells and reduce their growth factor requirements. Proc. Natl. Acad. Sci., USA. 87:1990;463-467.
84. Huschtscha L.I., Holliday R. Limited and unlimited growth of SV40-transformed cells from human diploid MRC-5 fibroblasts. J. Cell Sci. 63:1983;77-99.
85. Shay J.W., Van Der Haegen B.A., Ying Y., Wright W.E. The frequency of immortalization of human fibroblasts and mammary epithelial cells transfected with SV40 large T-antigen. Exp. Cell Res. 209:1993;45-52.
86. Chang S.E. In vitro transformation of human epithelial cells. Biochim. Biophys. Acta. 823:1986;161-194.
87. Wright W.E., Pereira-Smith O.M., Shay J.W. Reversible cellular senescence: implications for immortalization of normal human diploid fibroblasts. Mol. Cell. Biol. 9:1989;3088-3092.
88. Cairns J. Mutation selection and the natural history of cancer. Nature. 255:1975;197-200.
89. Fearon E.R., Vogelstein B. A genetic model for colorectal tumorigenesis. Cell. 61:1990;759-767.
90. Nowell P.C. The clonal evolution of tumor cell populations. Science. 194:1976;23-28.
91. Weinberg R.A. Oncogenes, antioncogenes, and the molecular bases of multistep carcinogenesis. Cancer Res. 49:1989;3713-3721.
92. Brown W.R.A. Molecular cloning of human telomeres in yeast. Nature. 338:1989;774-776.
93. Cross S.H., Allshire R.C., McKay S.J., McGill N.I., Cooke H.J. Cloning of human telomeres by complementation in yeast. Nature. 338:1989;771-774.
94. Kipling D., Cooke H.J. Hypervariable ultra-long telomeres in mice. Nature. 347:1990;400-402.
95. Moyzis R.K., Buckingham J.M., Cram L.S., Dani M., Deaven L.L., Jones M.D., Meyne J., Thailandliff R.L., Wu J.R. A highly conserved repetitive DNA sequence, (TTAGGG)n, present doubling the telomeres of human chromosomes. Proc. Natl. Acad. Immunoelectrophoresis USA. 85:1988;6622-6626.
96. Starling J.A., Maule J., Hastie N.D., Allshire R.C. Extensive telomere repeat arrays in mouse are hypervariable. Nucleic Acids Res. 18:1990;6881-6888.
97. Allshire R.C., Dempster M., Hastie N.D. Human telomeres contain at least three types of G-rich repeat distributed non-randomly. Nucleic Acids Res. 17:1989;4611-4627.
98. Brown W.R., MacKinnon P.J., Villasante A., Spurr N., Buckle V.J., Dobson M.J. Structure and polymorphism of human telomere-associated DNA. Cell. 63:1990;119-132.
99. de Lange T., Shiue L., Myers R.M., Cox D.R., Naylor S.L., Killery A.M., Varmus H.E. Structure and variability of human chromosome ends. Mol. Cell. Biol. 10:1990;518-527.
100. Harley C.B., Futcher A.B., Greider C.W. Telomeres shorten during ageing of human fibroblasts. Nature. 345:1990;458-460.
101. de Lange T. Telomere dynamics and genome instability in human cancer. Blackburn E.H., Greider C.W. Telomeres. 1995;265-294 Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
102. Allshire R.C., Gosden J.R., Cross S.H., Cranston G., Rout D., Sugawara N., Szostak J.W., Fantes P.A., Hastie N.D. Telomeric repeat from T. thermophila cross hybridizes with human telomeres. Nature. 332:1988;656-659.
103. Allsopp R.C., Vaziri H., Patterson C., Goldstein S., Younglai E.V., Futcher A.B., Greider C.W., Harley C.B. Telomere length predicts replicative capacity of human fibroblasts. Proc. Natl. Acad. Sci. USA. 89:1992;10114-10118.
104. Cooke H.J., Smith B.A. Variability at the telomeres of the human X/Y pseudoautosomal region. Cold Spring Harbour Symp. Quant. Biol. 1:1986;213-219.
105. Cross S., Lindsey J., Fantes J., McKay S., McGill N., Cooke H. The structure of a subterminal repeated sequence present on many human chromosomes. Nucleic Acids Res. 18:1990;6649-6657.
106. Hastie N.D., Dempster M., Dunlop M.G., Thompson A.M., Green D.K., Allshire R.C. Telomere reduction in human colorectal carcinoma and with ageing. Nature. 346:1990;866-868.
107. Allsopp R.C., Chang E., Kashefi-Aazam M., Rogaev E.I., Piatyszek M.A., Shay J.W., Harley C.B. Telomere shortening is associated with cell division in vitro and in vivo. Exp. Cell Res. 220:1995;194-200.
108. Allsopp R.C., Harley C.B. Evidence for a critical telomere length in senescent human fibroblasts. Exp. Cell Res. 219:1995;130-136.
109. Chang E., Harley C.B. Telomere length and replicative aging in human vascular tissues. Proc. Natl. Acad. Sci. USA. 92:1995;11190-11194.
110. Counter C.M., Avilion A.A., Le Feuvre C.E., Stewart N.G., Greider C.W., Harley C.B., Bacchetti S. Telomere shortening associated with chromosome instability is arrested in immortal cells which express telomerase activity. EMBO J. 11:1992;1921-1929.
111. Kitada T., Seki S., Kawakita N., Kuroki T., Monna T. Telomere shortening in chronic liver diseases. Biochem. Biophys. Res. Commun. 211:1995;33-39.
112. Lindsey J., McGill N.I., Lindsey L.A., Green D.K., Cooke H.J. In vivo loss of telomeric repeats with age in humans. Mutation Res. 256:1991;45-48.
113. Shay J.W., Wright W.E., Brasiskyte D., Van der Haegen B.A. E6 of human papillomavirus type 16 can overcome the M1 stage of immortalization in human mammary epithelial cells but not in human fibroblasts. Oncogene. 8:1993;1407-1413.
114. Vaziri H., Schachter F., Uchida I., Wei L., Zhu X., Effros R., Cohen D., Harley C.B. Loss of telomeric DNA during aging of normal and trisomy 21 human lymphocytes. Am. J. Hum. Genet. 52:1993;661-667.
115. Vaziri H., Dragowska W., Allsopp R.C., Thomas T.E., Harley C.B., Lansdorp P.M. Evidence for a mitotic clock in human hematopoietic stem cells: loss of telomeric DNA with age. Proc. Natl. Acad. Sci. USA. 91:1994;9857-9860.
116. Weng N.P., Levine B.L., June C.H., Hodes R.J. Human naive and memory T lymphocytes differ in telomeric length and replicative potential. Proc. Natl. Acad. Sci. USA. 92:1995;11091-11094.
117. 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., Shay J.W. Specific association of human telomerase activity with immortal cells and cancer. Science. 266:1994;2011-2015.
118. Bacchetti S., Counter C.M. Telomeres and telomerase in human cancer - Review. Int. J. Oncol. 7:1995;423-432.
119. Tahara H., Nakanishi T., Kitamoto M., Nakashio R., Shay J.W., Tahara E., Kajiyama G., Ide T. Telomerase activity in human liver tissues: comparison between chronic liver disease and hepatocellular carcinomas. Cancer Res. 55:1995;2734-2736.
120. Broccoli D., Young J.W., de Lange T. Telomerase activity in normal and malignant hematopoietic cells. Proc. Natl. Acad. Sci. USA. 92:1995;9082-9086.
121. Counter C.M., Gupta J., Harley C.B., Leber B., Bacchetti S. Telomerase activity in normal leukocytes and in hematologic malignancies. Blood. 85:1995;2315-2320.
122. Hiyama K., Hirai Y., Kyoizumi S., Akiyama M., Hiyama E., Piatyszek M.A., Shay J.W., Ishioka S., Yamakido M. Activation of telomerase in human lymphocytes and hematopoietic progenitor cells. J. Immunol. 155:1995;3711-3715.
123. Kruk P.A., Rampino N.J., Bohr V.A. DNA damage and repair in telomeres: relation to aging. Proc. Natl. Acad. Sci. USA. 92:1995;258-262.
124. Wright W.E., Shay J.W. Telomere positional effects and the regulation of cellular senescence. Trends Genet. 8:1992;193-197.
125. Harley C.B. Telomere loss: mitotic clock or genetic time bomb? Mutation Res. 256:1991;271-282.
126. Counter C.M., Botelho F.M., Wang P., Harley C.B., Bacchetti S. Stabilization of short telomeres and telomerase activity accompany immortalization of Epstein-Barr virus-transformed human B lymphocytes. J. Virol. 68:1994a;3410-3414.
127. Levy M.Z., Allsopp R.C., Futcher A.B., Greider C.W., Harley C.B. Telomere end-replication problem and cell aging. J. Mol. Biol. 225:1992;951-960.
128. Benn P.A. Specific chromosome aberrations in senescent fibroblast cell lines derived from human embryos. Am. J. Hum. Genet. 28:1976;465-473.
129. Sherwood S.W., Rush D., Ellsworth J.L., Schimke R.T. Defining cellular senescence in IMR-90 cells: a flow cytometric analysis. Proc. Natl. Acad. Sci. USA. 85:1988;9086-9090.
130. Di Leonardo A., Linke S.P., Clarkin K., Wahl G.M. DNA damage triggers a prolonged p53-dependent Gl arrest and long-term induction of Cipl in normal human fibroblasts. Genes Dev. 8:1994;2540-2551.
131. Weirich-Schwaiger H., Weirich H.G., Gruber B., Schweiger M., Hirsch-Kauffmann M. Correlation between senescence and DNA repair in cells from young and old individuals and in premature aging syndromes. Mutation Res. 316:1994;37-48.
132. Bernstein L., Bernstein H. Aging, Sex and DNA Repair. 1991;Academic Press, San Diego, California.
133. Wright W.E., Brasiskyte D., Piatyszek M.A., Shay J.W. Experimental elongation of telomeres extends the life span of immortal×normal cell hybrids. EMBO J. 1996;. in press.
134. Avilion A.A., Piatyszek M.A., Gupta J., Shay J.W., Bacchetti S., Greider C.W. Human telomerase RNA and telomerase activity in immortal cell lines and tumor tissues. Cancer Res. 56:1996;645-650.
135. Holt S.E., Gollahon L.S., Willingham T., Barbosa M.S., Shay J.W. p53 levels in human mammary epithelial cells expressing wild-type and mutant human papillomavirus type 16 (HPV-16) E6 proteins: relationship to reactivation of telomerase and immortalization. Int. J. Oncol. 8:1996;263-270.
136. Klingelhutz A.J., Foster S.A., McDougall J.K. Telomerase activation by the E6 gene product of human papillomavirus type 16. Nature. 380:1996;79-82.
137. Klingelhutz A.J., Barber S.A., Smith P.P., Dyer K., McDougall J.K. Restoration of telomeres in human papillomavirus-immortalized human anogenital epithelial cells. Mol. Cell. Biol. 14:1994;961-969.
138. Strahl C., Blackburn E.H. Effects of reverse transcriptase inhibitors on telomere length and telomerase activity in two immortalized human cell lines. Mol. Cell. Biol. 16:199;53-65.
139. Counter C.M., Hirte H.W., Bacchetti S., Harley C.B. Telomerase activity in human ovarian carcinoma. Proc. Natl. Acad. Sci. USA. 91:1994;2900-2904.
140. Ohmura H., Tahara H., Suzuki M., Ide T., Shimizu M., Yoshida M.A., Tahara E., Shay J.W., Barrett J.C., Oshimura M. Restoration of the cellular senescence program and repression of telomerase by human chromosome 3. Jpn. J. Cancer Res. 86:1995;899-904.
141. Metz T., Harris A.W., Adams J.M. Absence of p53 allows direct immortalization of hematopoietic cells by the myc and raf oncogenes. Cell. 82:1995;29-36.
142. Todaro G.J., Green H. Quantitative studies on the growth of mouse embryo cells in culture and their development into established lines. J. Cell. Biol. 17:1963;299-313.
143. Garagna S., Broccoli D., Redi C.A., Searle J.B., Cooke H.J., Capanna E. Robertsonian metacentrics of the house mouse lose telomeric sequences but retain some minor satellite DNA in the pericentromeric area. Chromosoma. 103:1995;685-692.
144. Hayashi T., Mafune Y., Takada T., Hatakeyama K., Takagi N., Kominami R. Telomere change and loss of heterozygosity of mouse primary tumors and cell lines. Jpn. J. Cancer Res. 84:1993;1292-1299.
145. Prowse K.R., Greider C.W. Developmental and tissue-specific regulation of mouse telomerase and telomere length. Proc. Natl. Acad. Sci. USA. 92:1995;4818-4822.
146. Meyne J., Ratliff R.L., Moyzis R.K. Conservation of the human telomere sequence (TTAGGG), among vertebrates. Proc. Natl. Acad. Sci. USA. 86:1989;7049-7053.
147. Meyne J., Baker R.J., Hobart H.H., Hsu T.C., Ryder O.A., Ward O.G., Wiley J.E., Wurster-Hill D.H., Yates T.L., Moyzis R.K. Distribution of non-telomeric sites of the (TTAGGG), telomeric sequence in vertebrate chromosomes. Chromosoma. 99:1990;3-10.
148. Bednarek A., Budunova I., Slaga T.J., Aldaz C.M. Increased telomerase activity in mouse skin premalignant progression. Cancer Res. 55:1995;4566-4569.
149. Chadeneau C., Siegel P., harley C.B., Muller W.J., Bacchetti S. Telomerase activity in normal and malignant murine tissues. Oncogene. 11:1995;893-898.
150. Sharma H.W., Sokoloski J.A., Perez J.R., Maltese J.Y., Sartorelli A.C., Stein C.A., Nichols G., Khaled Z., Telang N.T., Narayananan R. Differentiation of immortal cells inhibits telomerase activity. Proc. Natl. Acad. Sci. USA. 92:1995;12343-12346.
151. Blasco M.A., Rizen M., Greider C.W., Hanahan D. Differential regulation of telomerase activity and telomerase RNA during multi-stage tumorigenesis. Nature Genet. 12:1996;200-204.
152. Mortimer R.K., Johnston J.R. Life span of individual yeast cells. Nature. 183:1959;1751-1752.
153. Kennedy B.K., Austriaco N.R. Jr., Guarente L. Daughter cells of Saccharomyces cerevisiae from old mothers display a reduced life span. J Cell Biol. 127:1994;1985-1993.
154. Müller I., Zimmermann M., Becker D., Flomer M. Calendar life span versus budding life span of Saccharomyces cerevisiae. Mech. Ageing Dev. 12:1980;47-52.
155. Egilmez N.K., Chen J.B., Jazwinski S.M. Preparation and partial characterization of old yeast cells. J. Gerontol. 45:1990;B9-B17.
156. Hensler P.J., Pereira-Smith O.M. Human replicative senescence. A molecular study. Am. J. Pathol. 147:1995;1-8.
157. Müller I. Parental age and the life span of zygotes of Saccharomyces cerevisiae. J. Microbiol. Serol. 51:1985;1-10.
158. Norwood T.H., Pendergrass W.R., Sprague C.A., Martin G.M. Dominance of the senescent phenotype in heterokaryons between replicative and post-replicative human fibroblast-like cells. Proc. Natl. Acad. Sci. USA. 71:1974;2231-2235.
159. D'Mello N.P., Jazwinski S.M. Telomere length constancy during aging of Saccharomyces cerevisiae. J. Bacteriol. 173:1991;6709-6713.
160. Kennedy B.K., Austriaco N.R. Jr., Zhang J., Guarente L. Mutation in the silencing gene SIR4 can delay aging in S. cerevisiae. Cell. 80:1995;485-496.
161. Lundblad V., Szostak J.W. A mutant with a defect in telomere elongation leads to senescence in yeast. Cell. 57:1989;633-643.
162. Liu Z., Frantz J.D., Gilbert W., Tye B.K. Identification and characterization of a nuclease activity specific for G4 tetrastranded DNA. Proc. Natl. Acad. Sci. USA. 90:1993;3157-3161.
163. Liu Z., Gilbert W. The yeast KEM1 gene encodes a nuclease specific for G4 tetraplex DNA: implication of in vivo functions for this novel DNA structure. Cell. 77:1994;1083-1092.
164. Zahler A.M., Williamson J.R., Cech T.R., Prescott D.M. Inhibition of telomerase by G-quartet DNA structures. Nature. 350:1991;718-720.
165. Lundblad V., Blackburn E.H. RNA-dependent polymerase motifs in EST1: tentative identification of a protein component of an essential yeast telomerase [letter]. Cell. 60:1990;529-530.
166. Sandell L.L., Zakian V.A. Loss of a yeast telomere: arrest, recovery, and chromosome loss. Cell. 75:1993;729-739.
167. Garvik B., Carson M., Hartwell L. Single-stranded DNA arising at telomeres in cdc13 mutants may constitute a specific signal for the RAD9 checkpoint. Mol. Cell. Biol. 15:1995;6128-6138.
168. Greenwell P.W., Kronmal S.L., Porter S.E., Gassenhuber J., Obermaier B., Petes T.D. TEL1, a gene involved in controlling telomere length in S. cerevisiae, is homologous to the human ataxia telangiectasia gene. Cell. 82:1995;823-829.
169. Morrow D.M., Tagle D.A., Shiloh Y., Collins F.S., Hieter P. TEL1, an S. cerevisiae homolog of the human gene mutated in ataxia telangiectasia, is functionally related to the yeast checkpoint gene MEC1. Cell. 82:1995;831-840.
170. Kastan M.B., Zhan Q., el-Deiry W.S., Carrier F., Jacks T., Walsh W.V., Plunkett B.S., Vogelstein B., Fornace A.J. Jr. A mammalian cell cycle checkpoint pathway utilizing p53 and GADD45 is defective in ataxia-telangiectasia. Cell. 71:1992;587-597.
171. Kato R., Ogawa H. An essential gene, ESR1, is required for mitotic cell growth, DNA repair and meiotic recombination in Saccharomyces cerevisiae. Nucleic Acids Res. 22:1994;3104-3112.
172. Shiloh Y. Ataxia-telangiectasia: closer to unraveling the mystery. Eur. J. Hum. Genet. 3:1995;116-138.
173. Weinert T.A., Kiser G.L., Hartwell L.H. Mitotic checkpoint genes in budding yeast and the dependence of mitosis on DNA replication and repair. Genes Dev. 8:1994;652-665.
174. Lustig A.J., Petes T.D. Identification of yeast mutants with altered telomere structure. Proc. Natl. Acad. Sci. USA. 83:1986;1398-1402.
175. Smith-Sonneborn J. DNA repair and longevity assurance in Paramecium tetraurelia. Science. 203:1979;1115-1117.
176. Gilley D., Blackburn E.H. Lack of telomere shortening during senescence in Paramecium. Proc. Natl. Acad. Sci. USA. 91:1994;1955-1958.
177. Holmes G.E., Holmes N.R. Accumulation of DNA damages in aging Paramecium tetraurelia. Mol. Gen. Genet. 204:1986;108-114.
178. Larson D.D., Spangler E.A., Blackburn E.H. Dynamics of telomere length variation in Tetrahymena thermophila. Cell. 50:1987;477-483.