Transcriptional regulation of the telomerase hTERT gene as a target for cellular and viral oncogenic mechanisms

Carcinogenesis

Volumn 24, Issue 7, 2003, Pages 1167-1176

  Horikawa, Izumi ^a   Barrett, J Carl ^a  

^a NATIONAL CANCER INSTITUTE   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

HISTONE; MYC PROTEIN; PROTEIN P53; REPRESSOR PROTEIN; RNA DIRECTED DNA POLYMERASE; TELOMERASE; TRANSCRIPTION FACTOR; TUMOR SUPPRESSOR PROTEIN; VIRUS PROTEIN; WT1 PROTEIN;

CANCER CELL; CARCINOGENESIS; CELL IMMORTALIZATION; CHROMATIN STRUCTURE; CHROMOSOME 3P; CYTOLOGY; DISEASE ASSOCIATION; ENZYME ACTIVATION; ENZYME ACTIVE SITE; ENZYME ACTIVITY; ENZYME INACTIVATION; ENZYME REGULATION; ENZYME REPRESSION; GENE CONTROL; GENE FUNCTION; GENE OVEREXPRESSION; GENE TARGETING; GENE TRANSFER; GENETIC TRANSCRIPTION; HEPATITIS B VIRUS; HUMAN; HUMAN CELL; KERATINOCYTE; LIVER CELL CARCINOMA; MALIGNANT TRANSFORMATION; MOLECULAR MECHANICS; NUCLEOSOME; PRIORITY JOURNAL; PROTEIN ANALYSIS; PROTEIN MODIFICATION; REVIEW; TELOMERE; TRANSACTIVATION; TRANSCRIPTION INITIATION; TRANSCRIPTION REGULATION; TUMOR SUPPRESSOR GENE; VIRUS ONCOGENE; WART VIRUS;

CELL TRANSFORMATION, VIRAL; DNA METHYLATION; DNA-BINDING PROTEINS; GENE EXPRESSION REGULATION, ENZYMOLOGIC; HUMANS; PAPILLOMAVIRIDAE; TELOMERASE; TELOMERE; TRANS-ACTIVATION (GENETICS); TRANSCRIPTION, GENETIC;

EID: 0042169025     PISSN: 01433334     EISSN: None     Source Type: Journal    
DOI: 10.1093/carcin/bgg085     Document Type: Review

References (120)

1. Chiu,C.P. and Harley,C.B. (1997) Replicative senescence and cell immortality: the role of telomeres and telomerase. Proc. Soc. Exp. Biol. Med., 214, 99-106.
2. Mathon,N.F. and Lloyd,A.C. (2001) Cell senescence and cancer. Naure Rev. Cancer, 1, 203-213.
3. Campisi,J. (2001) Cellular senescence as a tumor-suppressor mechanism. Trends Cell Biol., 11, S27-31.
4. Hanahan,D. and Weinberg,R.A. (2000) The hallmarks of cancer. Cell, 100, 57-70.
5. Campisi,J., Kim,S.H., Lim,C.S. and Rubio,M. (2001) Cellular senescence, cancer and aging: the telomere connection. Exp. Gerontol., 36, 1619-1637.
6. Horikawa,I., Yawata,T. and Barrett,J.C. (2000) Cellular senescence mechanisms independent of telomere shortening and telomerase: other barriers to cell immortalization and carcinogenesis. J. Anti-Aging Med., 3, 373-382.
7. Bryan,T.M. and Cech,T.R. (1999) Telomerase and the maintenance of chromosome ends. Curr. Opin. Cell Biol., 11, 318-324.
8. Meyerson,M. (2000) Role of telomerase in normal and cancer cells. J. Clin. Oncol., 18, 2626-2634.
9. Granger,M.P., Wright,W.E. and Shay,J.W. (2002) Telomerase in cancer and aging. Crit. Rev. Oncol. Hematol., 41, 29-40.
10. 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.
11. Meyerson,M., Counter,C.M., Eaton,E.N. et al. (1997) hEST2, the putative human telomerase catalytic subunit gene, is up-regulated in tumor cells and during immortalization. Cell, 90, 785-795.
12. Nakamura,T.M., Morin,G.B., Chapman,K.B., Weinrich,S.L., Andrews,W.H., Lingner,J., Harley,C.B. and Cech,T.R. (1997) Telomerase catalytic subunit homologs from fission yeast and human. Science, 277, 955-959.
13. Kilian,A., Bowtell,D.D., Abud,H.E., Hime,G.R., Venter,D.J., Keese,P.K., Duncan,E.L., Reddel,R.R. and Jefferson,R.A. (1997) Isolation of a candidate human telomerase catalytic subunit gene, which reveals complex splicing patterns in different cell types. Hum. Mol. Genet., 6, 2011-2019.
14. Nakayama,J., Tahara,H., Tahara,E., Saito,M., Ito,K., Nakamura,H., Nakanishi,T., Tahara,E., Ide,T. and Ishikawa,F. (1998) Telomerase activation by hTRT in human normal fibroblasts and hepatocellular carcinomas. Nature Genet., 18, 65-68.
15. Aisner,D.L., Wright,W.E. and Shay,J.W. (2002) Telomerase regulation: not just flipping the switch. Curr. Opin. Genet. Dev., 12, 80-85.
16. Poole,J.C., Andrews,L.G. and Tollefsbol,T.O. (2001) Activity, function, and gene regulation of the catalytic subunit of telomerase (hTERT). Gene, 269, 1-12.
17. Horikawa,I., Cable,P.L., Afshari,C. and Barrett,J.C. (1999) Cloning and characterization of the promoter region of human telomerase reverse transcriptase gene. Cancer Res., 59, 826-830.
18. Takakura,M., Kyo,S., Kanaya,T., Hirano,H., Takeda,J., Yutsudo,M. and Inoue,M. (1999) Cloning of human telomerase catalytic subunit (hTERT) gene promoter and identification of proximal core promoter sequences essential for transcriptional activation in immortalized and cancer cells. Cancer Res., 59, 551-557.
19. Wu,K.J., Grandori,C., Amacker,M., Simon-Vermot,N., Polack,A., Lingner,J. and Dalla-Favera,R. (1999) Direct activation of TERT transcription by c-MYC. Nature Genet., 21, 220-224.
20. Greenberg,R.A., O'Hagan,R.C., Deng,H., Xiao,Q., Hann,S.R., Adams,R.R., Lichtsteiner,S., Chin,L., Morin,G.B. and DePinho,R.A. (1999) Telomerase reverse transcriptase gene is a direct target of c-Myc but is not functionally equivalent in cellular transformation. Oncogene, 18, 1219-1226.
21. Oh,S., Song,Y.H., Kim,U.J., Yim,J. and Kim,T.K. (1999) In vivo and in vitro analyses of Myc for differential promoter activities of the human telomerase (hTERT) gene in normal and tumor cells. Biochem. Biophys. Res. Commun., 263, 361-365.
22. Cong,Y.S., Wen,J. and Bacchetti,S. (1999) The human telomerase catalytic subunit hTERT: organization of the gene and characterization of the promoter. Hum. Mol. Genet., 8, 137-142.
23. Grandori,C., Cowley,S.M., James,L.P. and Eisenman,R.N. (2000) The Myc/Max/Mad network and the transcriptional control of cell behavior. Annu. Rev. Cell. Dev. Biol., 16, 653-699.
24. Kyo,S., Takakura,M., Taira,T., Kanaya,T., Itoh,H., Yutsudo,M., Ariga,H. and Inoue,M. (2000) Sp1 cooperates with c-Myc to activate transcription of the human telomerase reverse transcriptase gene (hTERT). Nucleic Acids Res., 28, 669-677.
25. Wang,J., Xie,L.Y., Allan,S., Beach,D. and Hannon,G.J. (1998) Myc activates telomerase. Genes Dev., 12, 1769-1774.
26. Günes,C., Lichtsteiner,S., Vasserot,A.P. and Englert,C. (2000) Expression of the hTERT gene is regulated at the level of transcriptional initiation and repressed by Mad1. Cancer Res., 60, 2116-2121.
27. Horikawa,I., Cable,P.L., Mazur,S.J., Appella,E., Afshari,C.A. and Barrett,J.C. (2002) Downstream E-box-mediated regulation of the human telomerase reverse transcriptase (hTERT) gene transcription: evidence for an endogenous mechanism of transcriptional repression. Mol. Biol. Cell., 13, 2585-2597.
28. Veldman,T., Horikawa,I., Barrett,J.C. and Schlegel,R. (2001) Transcriptional activation of the telomerase hTERT gene by human papillomavirus type 16 E6 oncoprotein. J. Virol., 75, 4467-4472.
29. Gewin,L. and Galloway,D.A. (2001) E box-dependent activation of telomerase by human papillomavirus type 16 E6 does not require induction of c-myc. J. Virol., 75, 7198-7201.
30. Xiao,X., Phogat,S.K., Sidorov,I.A., Yang,J., Horikawa,I., Prieto,D., Adelesberger,J., Lempicki,R., Barrett,J.C. and Dimitrov,D.S. (2002) Identification and characterization of rapidly dividing U937 clones with differential telomerase activity and gene expression profiles: role of c-Myc/Mad1 and Id/Ets proteins. Leukemia, 16, 1877-1880.
31. Xu,D., Popov,N., Hou,M., Wang,Q., Bjorkholm,M., Gruber,A., Menkel,A.R. and Henriksson,M. (2001) Switch from Myc/Max to Mad1/Max binding and decrease in histone acetylation at the telomerase reverse transcriptase promoter during differentiation of HL60 cells. Proc. Natl Acad. Sci. USA, 98, 3826-3831.
32. Hannon,G.J. (2002) RNA interference. Nature, 418, 244-251.
33. Yago,M., Ohki,R., Hatakeyama,S., Fujita,T. and Ishikawa,F. (2002) Variant forms of upstream stimulatory factors (USFs) control the promoter activity of hTERT, the human gene encoding the catalytic subunit of telomerase. FEBS Lett., 520, 40-46.
34. Maida,Y., Kyo,S., Kanaya,T., Wang,Z., Yatabe,N, Tanaka,M., Nakamura,M., Ohmichi,M., Gotoh,N., Murakami,S. and Inoue,M. (2002) Direct activation of telomerase by EGF through Ets-mediated transactivation of TERT via MAP kinase signaling pathway. Oncogene, 21, 4071-4079.
35. Oh,S., Song,Y., Yim,J. and Kim,T.K. (1999) The Wilms' tumor 1 tumor suppressor gene represses transcription of the human telomerase reverse transcriptase gene. J. Biol. Chem., 274, 37473-37478.
36. Oh,S., Song,Y.H., Yim,J. and Kim,T.K. (2000) Identification of Mad as a repressor of the human telomerase (hTERT) gene. Oncogene, 19, 1485-1490.
37. Haber,D.A. and Buckler,A.J. (1992) WT1: a novel tumor suppressor gene inactivated in Wilms' tumor. N. Biol., 4, 97-106.
38. Foley,K.P. and Eisenman,R.N. (1999) Two MAD tails: what the recent knockouts of Mad1 and Mxi1 tell us about the MYC/MAX/MAD network. Biochim. Biophys. Acta, 1423, M37-47.
39. Fujimoto,K., Kyo,S., Takakura,M., Kanaya,T., Kitagawa,Y., Itoh,H., Takahashi,M. and Inoue,M. (2000) Identification and characterization of negative regulatory elements of the human telomerase catalytic subunit (hTERT) gene promoter: possible role of MZF-2 in transcriptional repression of hTERT. Nucleic Acids Res., 28, 2557-2562.
40. Crowe,D.L., Nguyen,D.C., Tsang,K.J. and Kyo,S. (2001) E2F-1 represses transcription of the human telomerase reverse transcriptase gene. Nucleic Acids Res., 29, 2789-2794.
41. Yamasaki,L., Jacks,T., Bronson,R., Goillot,E., Harlow,E. and Dyson,N.J. (1996) Tumor induction and tissue atrophy in mice lacking E2F-1. Cell, 85, 537-548.
42. Nakamura,T., Monden,Y., Kawashima,K., Naruke,T. and Nishimura,S. (1996) Failure to detect mutations in the retinoblastoma protein-binding domain of the transcription factor E2F-1 in human cancers. Jpn. J. Cancer Res., 87, 1204-1209.
43. Ho,G.H., Calvano,J.E., Bisogna,M. and Van Zee,K.J. (2001) Expression of E2F-1 and E2F-4 is reduced in primary and metastatic breast carcinomas. Breast Cancer Res. Treat., 69, 115-122.
44. Yagi,K., Furuhashi,M., Aoki,H., Goto,D., Kuwano,H., Sugamura,K., Miyazono,K. and Kato,M. (2002) c-myc is a downstream target of the Smad pathway. J. Biol. Chem., 277, 854-861.
45. Chen,C.R., Kang,Y. and Massague,J. (2001) Defective repression of c-myc in breast cancer cells: a loss at the core of the transforming growth factor beta growth arrest program. Proc. Natl Acad. Sci. USA, 98, 992-999.
46. Yang,H., Kyo,S., Takatura,M. and Sun,L. (2001) Autocrine transforming growth factor beta suppresses telomerase activity and transcription of human telomerase reverse transcriptase in human cancer cells. Cell Growth. Differ., 12, 119-127.
47. Cerezo,A., Kalthoff,H., Schuermann,M., Schafer,B. and Boukamp,P. (2002) Dual regulation of telomerase activity through c-Myc-dependent inhibition and alternative splicing of hTERT. J. Cell Sci., 115, 1305-1312.
48. Venkitaraman,A.R. (2002) Cancer susceptibility and the functions of BRCA1 and BRCA2. Cell, 108, 171-182.
49. Li,H., Lee,T.H. and Avraham,H. (2002) A novel tricomplex of BRCA1, Nmi, and c-Myc inhibits c-Myc-induced human telomerase reverse transcriptase gene (hTERT) promoter activity in breast cancer. J. Biol. Chem., 277, 20965-20973.
50. Zhou,C. and Liu,J. (2003) Inhibition of human telomerase reverse transcriptase gene expression by BRCA1 in human ovarian cancer cells. Biochem. Biophys. Res. Commun., 303, 130-136.
51. Xu,D., Wang,Q., Gruber,A., Bjorkholm,M., Chen,Z.G., Zaid,A., Selivanova,G., Peterson,C., Wiman,K.G. and Pisa,P. (2000) Downregulation of telomerase reverse transcriptase mRNA expression by wild type p53 in human tumor cells. Oncogene, 19, 5123-5133.
52. Kanaya,T., Kyo,S., Hamada,K., Takakura,M., Kitagawa,Y., Harada,H. and Inoue,M. (2000) Adenoviral expression of p53 represses telomerase activity through down-regulation of human telomerase reverse transcriptase transcription. Clin. Cancer Res., 6, 1239-1247.
53. Oshimura,M. and Barrett,J.C. (1997) Multiple pathways to cellular senescence: role of telomerase repressors. Eur. J. Cancer, 33, 710-715.
54. Ran,Q. and Pereira-Smith,O.M. (2000) Genetic approaches to the study of replicative senescence. Exp. Gerontol., 35, 7-13.
55. Horikawa,I., Oshimura,M. and Barrett,J.C. (1998) Repression of the telomerase catalytic subunit by a gene on human chromosome 3 that induces cellular senescence. Mol. Carcinogen., 22, 65-72.
56. Tanaka,H., Shimizu,M., Horikawa,I., Kugoh,H., Yokota,J., Barret,J.C. and Oshimura,M. (1998) Evidence for a putative telomerase repressor gene in the 3p14.2-p21.1 region. Genes Chromosomes Cancer, 23, 123-133.
57. Cuthbert,A.P., Bond,J., Trott,D.A., Gill,S., Broni,J., Marriott,A., Khoudoli,G., Parkinson,E.K., Cooper,C.S. and Newbold,R.F. (1999) Telomerase repressor sequences on chromosome 3 and induction of permanent growth arrest in human breast cancer cells. J. Natl Cancer Inst., 91, 37-45.
58. Ducrest,A.L., Amacker,M., Mathieu,Y.D., Cuthbert,A.P., Trott,D.A., Newbold,R.F., Nabholz,M. and Lingner,J. (2001) Regulation of human telomerase activity: repression by normal chromosome 3 abolishes nuclear telomerase reverse transcriptase transcripts but does not affect c-Myc activity. Cancer Res., 61, 7594-7602.
59. Backsch,C., Wagenbach,N., Nonn,M., Leistritz,S., Stanbridge,E., Schneider,A. and Durst,M. (2001) Microcell-mediated transfer of chromosome 4 into HeLa cells suppresses telomerase activity. Genes Chromosomes Cancer, 31, 196-198.
60. Steenbergen,R.D.M., Kramer,D., Meijer,C.J., Walboomers,J.M., Trott,D.A., Cuthbert,A.P., Newbold,R.F., Overkamp,W.J., Zdzienicka,M.Z. and Snijders,P.J. (2001) Telomerase suppression by chromosome 6 in a human papillomavirus type 16-immortalized keratinocyte cell line and in a cervical cancer cell line. J. Natl Cancer Inst., 93, 865-872.
61. Nakabayashi,K., Ogino,H., Michishita,E., Satoh,N. and Ayusawa,D. (1999) Introduction of chromosome 7 suppresses telomerase with shortening of telomeres in a human mesothelial cell line. Exp. Cell Res., 252, 376-382.
62. Nishimoto,A., Miura,N., Horikawa,I. et al. (2001) Functional evidence for a telomerase repressor gene on human chromosome 10p15.1. Oncogene, 20, 828-835.
63. Yang,X., Tahin,Q., Hu,Y.F., Russo,I.H., Balsara,B.R., Mihaila,D., Slater,C., Barrett,J.C. and Russo,J. (1999) Functional roles of chromosomes 11 and 17 in the transformation of human breast epithelial cells in vitro. Int. J. Oncol., 15, 629-638.
64. Greider,C.W. (1998) Telomerase activity, cell proliferation, and cancer. Proc. Natl Acad. Sci. USA, 95, 90-92.
65. Horikawa,I., Parker,E.S., Solomon,G.G. and Barrett,J.C. (2001) Upregulation of the gene encoding a cytoplasmic dynein intermediate chain in senescent human cells. J. Cell. Biochem., 82, 415-421.
66. Oh,S.T., Kyo,S. and Laimins,L.A. (2001) Telomerase activation by human papillomavirus type 16 E6 protein: induction of human telomerase reverse transcriptase expression through Myc and GC-rich Sp1 binding sites. J. Virol., 75, 5559-5566.
67. Klingelhutz,A.J., Foster,S.A. and McDougall,J.K. (1996) Telomerase activation by the E6 gene product of human papillomavirus type 16. Nature, 380, 79-82.
68. Gao,Q., Singh,L., Kumar,A., Srinivasan,S., Wazer,D.E. and Band,V. (2001) Human papillomavirus type 16 E6-induced degradation of E6TP1 correlates with its ability to immortalize human mammary epithelial cells. J. Virol., 75, 4459-4466.
69. Lee,D., Kim,H.Z., Jeong,K.W., Shim,Y.S., Horikawa,I., Barrett,J.C. and Choe,J. (2002) Human papillomavirus E2 down-regulates the human telomerase reverse transcriptase promoter. J. Biol. Chem., 277, 27748-27756.
70. Ham,J., Dostatni,N., Gauthier,J.M. and Yaniv,M. (1991) The papillomavirus E2 protein: a factor with many talents. Trends Biochem. Sci., 16, 440-444.
71. Desaintes,C. and Demeret,C. (1996) Control of papillomavirus DNA replication and transcription. Semin. Cancer Biol., 7, 339-347.
72. Lee,C.J., Suh,E.J., Kang,H.T., Im,J.S., Um,S.J., Park,J.S. and Hwang,E.S. (2002) Induction of senescence-like state and suppression of telomerase activity through inhibition of HPV E6/E7 gene expression in cells immortalized by HPV16 DNA. Exp. Cell Res., 277, 173-182.
73. Baege,A.C., Berger,A., Schlegel,R., Veldman,T. and Schlegel R. (2002) Cervical epithelial cells transduced with the papillomavirus E6/E7 oncogenes maintain stable levels of oncoprotein expression but exhibit progressive, major increases in hTERT gene expression and telomerase activity. Am. J. Pathol., 160, 1251-1257.
74. Knight,J.S., Cotter,M.A. and Robertson,E.S. (2001) The latency-associated nuclear antigen of Kaposi's sarcoma-associated herpesvirus transactivates the telomerase reverse transcriptase promoter. J. Biol. Chem., 276, 22971-22978.
75. Kirch,H.C., Ruschen,S., Brockmann,D., Esche,H., Horikawa,I., Barrett,J.C., Opalka,B. and Hengge,U.R. (2002) Tumor-specific activation of hTERT-derived promoters by tumor suppressive E1A-mutants involves recruitment of p300/CBP/HAT and suppression of HDAC-1 and defines a combined tumor targeting and suppression system. Oncogene, 21, 7991-8000.
76. Makowski,D.R., Rothberg,P.G. and Astrin,S.M. (1984) The role of promoter insertion in the induction of neoplasia. Surv. Synth. Path. Res., 3, 342-349.
77. Horikawa,I. and Barrett,J.C. (2001) cis-Activation of the human telomerase gene (hTERT) by the hepatitis B virus genome. J. Natl Cancer Inst., 93, 1171-1173.
78. Gozuacik,D., Murakami,Y., Saigo,K., Chami,M., Mugnier,C., Lagorce,D., Okanoue,T., Urashima,T., Brechot,C. and Paterlini-Brechot,P. (2001) Identification of human cancer-related genes by naturally occurring Hepatitis B Virus DNA tagging. Oncogene, 20, 6233-6240.
79. Hahn,W.C., Counter,C.M., Lundberg,A.S., Beijersbergen,R.L., Brooks,M.W. and Weinberg,R.A. (1999) Creation of human tumour cells with defined genetic elements. Nature, 400, 464-468.
80. Elenbaas,B., Spirio,L., Koerner,F., Fleming,M.D., Zimonjic,D.B., Donaher,J.L., Popescu,N.C., Hahn,W.C. and Weinberg,R.A. (2001) Human breast cancer cells generated by oncogenic transformation of primary mammary epithelial cells. Genes Dev., 15, 50-65.
81. Harley,C.B. (2002) Telomerase is not an oncogene. Oncogene, 21, 494-502.
82. Jiang,X.R., Jimenez,G., Chang,E. et al. (1999) Telomerase expression in human somatic cells does not induce changes associated with a transformed phenotype. Nature Genet., 21, 111-114.
83. Morales,C.P., Holt,S.E., Ouellette,M., Kaur,K.J., Yan,Y., Wilson.K.S., White,M.A., Wright,W.E. and Shay,J.W. (1999) Absence of cancer-associated changes in human fibroblasts immortalized with telomerase. Nature Genet., 21, 115-118.
84. Serrano,M., Lin,A.W., McCurrach,M.E., Beach,D. and Lowe,S.W. (1997) Oncogenic ras provokes premature cell senescence associated with accumulation of p53 and p16INK4a. Cell, 88, 593-602.
85. Zhang,A., Zheng, C., Lindvall,C. et al. (2000) Frequent amplification of the telomerase reverse transcriptase gene in human tumors. Cancer Res., 60, 6230-6235.
86. Zhang,A., Zheng,C., Hou,M. et al. (2002) Amplification of the telomerase reverse transcriptase (hTERT) gene in cervical carcinomas. Genes Chromosomes Cancer, 34, 269-275.
87. Esteller,M. (2002) CpG island hypermethylation and tumor suppressor genes: a booming present, a brighter future. Oncogene, 21, 5427-5440.
88. Devereux,T.R., Horikawa,I., Anna,C.H., Annab,L.A., Afshari,C.A. and Barrett,J.C. (1999) DNA methylation analysis of the promoter region of the human telomerase reverse transcriptase (hTERT) gene. Cancer Res., 59, 6087-6090.
89. Dessain,S.K., Yu,H., Reddel,R.R., Beijersbergen,R.L. and Weinberg,R.A. (2000) Methylation of the human telomerase gene CpG island. Cancer Res., 60, 537-541.
90. 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.
91. Kitagawa,Y., Kyo,S., Takakura,M., Kanaya,T., Koshida,K., Namiki,M. and Inoue,M. (2000) Demethylating reagent 5-azacytidine inhibits telomerase activity in human prostate cancer cells through transcriptional repression of hTERT. Clin. Cancer Res., 6, 2868-2875.
92. McGregor,F., Muntoni,A., Fleming,J., Brown,J., Felix,D.H., MacDonald,D.G., Parkinson,E.K. and Harrison,P.R. (2002) Molecular changes associated with oral dysplasia progression and acquisition of immortality: potential for its reversal by 5-azacytidine. Cancer Res., 62, 4757-4766.
93. Robertson,K,D. (2002) DNA methylation and chromatin-unraveling the tangled web. Oncogene, 21, 5361-5379.
94. Dobosy,J.R. and Selker,E.U. (2001) Emerging connections between DNA methylation and histone acetylation. Cell. Mol. Life Sci., 58, 721-727.
95. Cong,Y.S. and Bacchetti,S. (2000) Histone deacetylation is involved in the transcriptional repression of hTERT in normal human cells. J. Biol. Chem., 275, 35665-35668.
96. Takakura,M., Kyo,S., Sowa,Y., Wang,Z., Yatabe,N., Maida,Y., Tanaka,M. and Inoue,M. (2001) Telomerase activation by histone deacetylase inhibitor in normal cells. Nucleic Acids Res., 29, 3006-3011.
97. Hou,M., Wang,X., Popov,N. et al. (2002) The histone deacetylase inhibitor trichostatin A derepresses the telomerase reverse transcriptase (hTERT) gene in human cells. Exp. Cell Res., 274, 25-34.
98. Won,J., Yim,J. and Kim,T.K. (2002) Sp1 and Sp3 recruit histone deacetylase to repress transcription of human telomerase reverse transcriptase (hTERT) promoter in normal human somatic cells. J. Biol. Chem., 277, 38230-38238.
99. Billon,N., Carlisi,D., Datto,M.B., van Grunsven,L.A., Watt,A., Wang,X.F. and Rudkin,B.B. (1999) Cooperation of Sp1 and p300 in the induction of the CDK inhibitor p21WAF1/CIP1 during NGF-mediated neuronal differentiation. Oncogene, 18, 2872-2882.
100. Suzuki,T., Kimura,A., Nagai,R. and Horikoshi,M. (2000) Regulation of interaction of the acetyltransferase region of p300 and the DNA-binding domain of Sp1 on and through DNA binding. Genes Cells, 5, 29-41.
101. Laherty,C.D., Yang,W.M., Sun,J.M., Davie,J.R., Seto,E. and Eisenman,R.N. (1997) Histone deacetylases associated with the mSin3 corepressor mediate mad transcriptional repression. Cell, 89, 349-356.
102. McMahon,S.B., Wood,M.A. and Cole,M.D. (2000) The essential cofactor TRRAP recruits the histone acetyltransferase hGCN5 to c-Myc. Mol. Cell. Biol., 20, 556-562.
103. Nikiforov,M.A., Chandriani,S., Park,J., Kotenko,I., Matheos,D., Johnsson,A., McMahon,S.B. and Cole,M.D. (2002) TRRAP-dependent and TRRAP-independent transcriptional activation by Myc family oncoproteins. Mol. Cell. Biol., 22, 5054-5063.
104. Wang,Z., Kyo,S., Takakura,M. et al. (2000) Progesterone regulates human telomerase reverse transcriptase gene expression via activation of mitogen-activated protein kinase signaling pathway. Cancer Res., 60, 5376-5381.
105. Alfonso-De Matte,M.Y., Yang,H., Evans,M.S, Cheng,J.Q. and Kruk,P.A. (2002) Telomerase is regulated by c-Jun NH2-terminal kinase in ovarian surface epithelial cells. Cancer Res., 62, 4575-4578.
106. Dimri,G.P., Martinez,J.L., Jacobs,J.J., Keblusek,P., Itahana,K., Van Lohuizen,M., Campisi,J., Wazer,D.E. and Band,V. (2002) The Bmi-1 oncogene induces telomerase activity and immortalizes human mammary epithelial cells. Cancer Res., 62, 4736-4745.
107. Misiti,S., Nanni,S., Fontemaggi,G. et al. (2000) Induction of hTERT expression and telomerase activity by estrogens in human ovary epithelium cells. Mol. Cell. Biol., 20, 3764-3771.
108. Kyo,S., Takakura,M., Kanaya,T., Zhuo,W., Fujimoto,K., Nishio,Y., Orimo,A. and Inoue,M. (1999) Estrogen activates telomerase. Cancer Res., 59, 5917-5921.
109. Nanni,S., Narducci,M., Delia Pietra,L., Moretti,F., Grasselli,A., De Carli,P., Sacchi,A., Pontecorvi,A. and Farsetti,A. (2002) Signaling through estrogen receptors modulates telomerase activity in human prostate cancer. J. Clin. Invest., 110, 219-227.
110. Wang,Z., Kyo,S., Maida,Y. et al. (2002) Tamoxifen regulates human telomerase reverse transcriptase (hTERT) gene expression differently in breast and endometrial cancer cells. Oncogene, 21, 3517-3524.
111. Soria,J.C., Moon,C., Wang,L. et al. (2001) Effects of N-(4-hydroxyphenyl)retinamide on hTERT expression in the bronchial epithelium of cigarette smokers. J. Natl Cancer. Inst., 93, 1257-1263.
112. Lian,Z., Fujimoto,J., Yokoyama,Y., Niwa,K. and Tamaya,T. (2001) Herbal complex suppresses telomerase activity in chemo-endocrine resistant cancer cell lines. Eur. J. Gynaecol. Oncol., 22, 347-349.
113. Ogretmen,B., Kraveka,J.M., Schady,D., Usta,J., Hannun,Y.A. and Obeid,L.M. (2001) Molecular mechanisms of ceramide-mediated telomerase inhibition in the A549 human lung adenocarcinoma cell line. J. Biol. Chem., 276, 32506-32514.
114. Leem,S.H., Londoño-Vallejo,J.A., Kim,J.H. et al. (2002) The human telomerase gene: complete genomic sequence and analysis of tandem repeat polymorphisms, in intronic regions. Oncogene, 21, 769-777.
115. Shay,J.W. and Wright,W.E. (2000) Implications of mapping the human telomerase gene (hTERT) as the most distal gene on chromosome 5p. Neoplasia, 2, 195-196.
116. Bryce,L.A., Morrison,N., Hoare,S.F., Muir,S. and Keith,W.N. (2000) Mapping of the gene for the human telomerase reverse transcriptase, hTERT, to chromosome 5p15.33 by fluorescence in situ hybridization. Neoplasia, 2, 197-201.
117. Baur,J.A., Zou,Y., Shay,J.W. and Wright,W.E. (2001) Telomere position effect in human cells. Science, 292, 2075-2077.
118. Guo,Q.M., Malek,R.L., Kim,S., Chiao,C., He,M., Ruffy,M,, Sanka,K., Lee,N.H., Dang,C.V. and Liu,E.T. (2000) Identification of c-Myc responsive genes using rat cDNA microarray. Cancer Res., 60, 5922-5928.
119. Tzukerman,M., Shachaf,C., Ravel,Y., Braunstein,I., Cohen-Barak,O., Yalon-Hacohen,M. and Skorecki,K.L. (2000) Identification of a novel transcription factor binding element involved in the regulation by differentiation of the human telomerase (hTERT) promoter. Mol. Biol. Cell, 11, 4381-4391.
120. Braunstein,I., Cohen-Barak,O., Shachaf,C., Ravel,Y., Yalon-Hacohen,M., Mills,G.B., Tzukerman,M. and Skorecki,K.L. (2001) Human telomerase reverse transcriptase promoter regulation in normal and malignant human ovarian epithelial cells. Cancer Res., 61, 5529-5536.