Mass Cultured Human Fibroblasts Overexpressing hTERT Encounter a Growth Crisis Following an Extended Period of Proliferation

Hayflick, 1961, The serial cultivation of human diploid cell strains, Exp. Cell. Res., 25, 585, 10.1016/0014-4827(61)90192-6

Olovnikov, 1973, A theory of marginotomy: The incomplete copying of template margin in enzymic synthesis of polynucleotides and the biological significance of the phenomenon, J. Theor. Biol., 41, 181, 10.1016/0022-5193(73)90198-7

Moyzis, 1988, A highly conserved repetitive DNA sequence, (TTAGGG)n, present at the telomeres of human chromosomes, Proc. Natl. Acad. Sci. USA., 85, 6622, 10.1073/pnas.85.18.6622

Bilaud, 1997, Telomeric localization of TRF2, a novel human telobox protein, Nature Genet., 17, 236, 10.1038/ng1097-236

Broccoli, 1997, Human telomeres contain two disting Myb-related proteins, TRF1 and TRF2, Nature Genet., 17, 231, 10.1038/ng1097-231

Griffith, 1999, Mammalian telomeres end in a large duplex loop, Cell, 97, 503, 10.1016/S0092-8674(00)80760-6

Kim, 1999, TIN2, a new regulator of telomere length in human cells, Nature Genet., 23, 312, 10.1038/13854

Harley, 1990, Telomeres shorten with ageing of human fibroblasts, Nature, 345, 458, 10.1038/345458a0

Chin, 1999, p53 deficiency rescues the adverse effects of telomere loss and cooperates with telomere dysfunction to accelerate carcinogenesis, Cell, 97, 527, 10.1016/S0092-8674(00)80762-X

Giradi, 1965, SV40-induced transformation of human diploid cells: crisis and recovery, J. Cell. Comp. Physiol., 65, 69, 10.1002/jcp.1030650110

Smith, 1996, Replicative senescence: Implications for in vivo aging and tumor suppression, Science, 273, 63, 10.1126/science.273.5271.63

Counter, 1992, Telomere shortening associated with chromosome instability is arrested in immortal cells which express telomerase activity, EMBO J., 11, 1921, 10.1002/j.1460-2075.1992.tb05245.x

Klingelhutz, 1994, Restoration of telomeres in human papillomavirus-immortalized human angogenital epithelial cells, Mol. Cell. Biol., 14, 961, 10.1128/MCB.14.2.961

Shay, 1989, Quantitation of the frequency of immortalization of normal human diploid fibroblasts by SV40 large T-antigen, Exp. Cell Res., 184, 109, 10.1016/0014-4827(89)90369-8

Dimri, 1994, Molecular and cell biology of replicative senescence, Cold Spring Harbor Symp. Quant. Biol., 59, 67, 10.1101/SQB.1994.059.01.010

Greider, 1989, A telomeric sequence in the RNA of tetrahymena telomerase required for telomere repeat synthesis, Nature, 337, 331, 10.1038/337331a0

Yu, 1990, In vivo alteration of telomere sequences and senescence caused by mutated tetrahymena telomerase RNAs, Nature, 344, 126, 10.1038/344126a0

Bryan, 1995, Telomere elongation in immortal human cells without detectable telomerase activity, EMBO J., 33

Kim, 1994, Specific association of human telomerase activity with immortal cells and cancer, Science, 266, 2011, 10.1126/science.7605428

Bryan, 1997, Evidence for an alternative mechanism for maintaining telomere length in human tumors and tumor-derived cell lines, Nature Med., 3, 1271, 10.1038/nm1197-1271

Feng, 1995, The RNA component of human telomerase, Science, 269, 1236, 10.1126/science.7544491

Nakamura, 1997, Telomerase catalytic subunit homologues from fission yeast and human, Science, 277, 955, 10.1126/science.277.5328.955

Nakayama, 1998, Telomerase activation by hTRT in human normal fibroblasts and hepatocellular carcinomas, Nature Genet., 18, 65, 10.1038/ng0198-65

Weinrich, 1997, Reconstitution of human telomerase with the template RNA component hTR and the catalytic protein subunit hTRT, Nature Genet., 17, 498, 10.1038/ng1297-498

Bodnar, 1998, Extension of life-span by introduction of telomerase into normal human cells, Science, 279, 349, 10.1126/science.279.5349.349

Vaziri, 1998, Reconstitution of telomerase activity in normal human cells leads to elongation of telomeres and extended replicative lifespan, Curr. Biol., 8, 279, 10.1016/S0960-9822(98)70109-5

Oulette, 1999, Telomerase activity does not always imply telomere maintenance, Biochem. Biophys. Res. Commun., 254, 795, 10.1006/bbrc.1998.0114

Yang, 1999, Human endothelial cell life extension by telomerase expression, J. Biol. Chem., 274, 26141, 10.1074/jbc.274.37.26141

Dickinson, 2000, Human keratinocytes that express hTERT and also bypass a p16INK4a-enforced mechanism that limits lifespan become immortal yet retain normal growth and differentiation characteristics, Mol. Cell Biol., 20, 1436, 10.1128/MCB.20.4.1436-1447.2000

Kiyono, 1998, Both Rb/p16INK4a inactivation and telomerase activity are required to immortalize human epithelial cells, Nature, 396, 84, 10.1038/23962

Counter, 1998, Dissociation among in vitro telomerase activity, telomere maintenance and cellular immortalization, Proc. Natl. Acad. Sci. USA, 95, 14723, 10.1073/pnas.95.25.14723

Halvorsen, 1999, Telomerase activity is sufficient to allow transformed cells to escape from crisis, Mol. Cell. Biol., 19, 1864, 10.1128/MCB.19.3.1864

Zhu, 1999, Telomerase extends the lifespan of virus-transformed human cells without net telomere lengthening, Proc. Natl. Acad. Sci. USA, 96, 3723, 10.1073/pnas.96.7.3723

Riviere, 1995, Effects of retroviral vector design on expression of human adenosine deaminase in murine bone marrow transplant recipients engrafted with genetically modified cells, Proc. Natl. Acad. Sci. USA, 92, 6733, 10.1073/pnas.92.15.6733

Ory, 1996, A stable human-derived packaging cell line for production of high titer retrovirus/vesicular stomatitis G pseudotypes, Proc. Natl. Acad. Sci. USA, 93, 11400, 10.1073/pnas.93.21.11400

Miller, 1991, Construction and properties of retrovirus packaging cells based on gibbon ape leukemia virus, J. Virol., 65, 2220, 10.1128/JVI.65.5.2220-2224.1991

Albanell, 1996, Telomerase activity is repressed during differentiation of maturation sensitive but not resistant human tumor cell lines, Cancer Res., 56, 1503

Engelhardt, 1997, Telomerase regulation, cell cycle, and telomere stability in primitive hematopoietic cells, Blood, 90, 182, 10.1182/blood.V90.1.182

Dimri, 1995, A biomarker that identifies senescent human cells in culture and in aging skin in vivo, Proc. Natl. Acad. Sci. USA, 92, 9363, 10.1073/pnas.92.20.9363

Dolbeare, 1983, Flow cytometric measurement of total DNA content and incorporated bromodeoxyuridine, Proc. Natl. Acad. Sci. USA, 80, 5573, 10.1073/pnas.80.18.5573

Franco, S, MacKenzie, K. L, Dias, S, Rafii, S, and, Moore, M. A. S. 2000, Clonal variations in phenotype of human fibroblasts expressing hTERT. Submitted for publication.

Jiang, 1999, Telomerase expression in human somatic cells does not induce changes associated with a transformed phenotype, Nature Genet., 21, 111, 10.1038/5056

Morales, 1999, Absence of cancer associated changes in human fibroblasts immortalized with telomerase, Nature Genet., 21, 115, 10.1038/5063

Rubelj, 1994, SV40-Transformed human cells in crisis exhibit changes that occur in normal cellular senescence, Exp. Cell Res., 211, 82, 10.1006/excr.1994.1062

Bond, 1999, Control of replicative life span in human cells: Barriers to clonal expansion intermediate between M1 senescence and M2 crisis, Mol. Cell Biol., 19, 3103, 10.1128/MCB.19.4.3103

Shay, 1993, The frequency of immortalization of human fibroblasts and mammary epithelial cells transfected with SV40 large T-antigen, Exp. Cell. Res., 209, 45, 10.1006/excr.1993.1283

Mondello, 1997, Chromosomal instability and telomere length variations during the life span of human fibroblast clones, Exp. Cell Res., 236, 385, 10.1006/excr.1997.3756

Filatov, 1998, Chromosomal instability is correlated with telomere erosion and inactivation of G2 checkpoint function in human fibroblasts expressing human papilloma virus type 16 oncoprotein, Oncogene, 16, 1825, 10.1038/sj.onc.1201711

Ducray, 1999, Telomere dynamics, end-to end fusions and telomerase activation during the human fibroblast immortalization process, Oncogene, 18, 4211, 10.1038/sj.onc.1202797

Minn, 1996, Expression of Bcl-XL and loss of p53 can co-operate to overcome cell cycle checkpoint induced mitotic spindle damage, Genes Dev., 10, 2621, 10.1101/gad.10.20.2621

Hastie, 1990, Telomere reduction in human colorectal carcinoma and with aging, Nature, 346, 866, 10.1038/346866a0

Allsop, 1992, Telomere length predicts replicative capacity of human fibroblasts, Proc. Natl. Acad. Sci. USA, 89, 10114, 10.1073/pnas.89.21.10114

Karlseder, 1999, p53- and ATM-dependent apoptosis induced by telomeres lacking TRF-2, Science, 283, 1321, 10.1126/science.283.5406.1321

Sohal, 1996, Oxidative stress, caloric restriction, and aging, Science, 273, 59, 10.1126/science.273.5271.59

Johnson, 1998, Telomeres, the nucleolus and aging, Curr. Opin. Cell Biol., 10, 332, 10.1016/S0955-0674(98)80008-2

Hahn, 1999, Creation of human tumor cells with defined genetic elements, Nature, 400, 464, 10.1038/22780

Seshadri, 1990, Repression of c-fos transcription and an altered genetic program in senescent human fibroblasts, Science, 247, 205, 10.1126/science.2104680

Shay, 1991, A role for both RB and p53 in the regulation of human cellular senescence, Exp. Cell. Res., 196, 33, 10.1016/0014-4827(91)90453-2

Rogan, 1995, Alterations in p53 and p16INK4 expression and telomere length during spontaneous immortalization of Li-Fraumeni syndrome fibroblasts, Mol. Cell. Biol., 15, 4745, 10.1128/MCB.15.9.4745

Bryan, 1994, SV40-induced immortalization of human cells, Crit. Rev. Oncol., 5, 331, 10.1615/CritRevOncog.v5.i4.10

Porcu, 1992, The growth-stimulating effect of simian virys 40 T antigen requires the interaction of insulinlike growth factor 1, Mol. Cell Biol., 12, 5069, 10.1128/MCB.12.11.5069

Candal, 1996, Human bone marrow microvascular endothelial cell line with primary cell characteristics, Microvasc. Res., 52, 221, 10.1006/mvre.1996.0060