Regulation of the human catalytic subunit of telomerase (hTERT)

Gene

Volumn 498, Issue 2, 2012, Pages 135-146

  Daniel, Michael ^a   Peek, Gregory W ^a   Tollefsbol, Trygve O ^a,b  

^a UNIVERSITY OF ALABAMA AT BIRMINGHAM   (United States)

^b UNIVERSITY OF ALABAMA AT BIRMINGHAM   (United States)

Author keywords

Cancer; Gene regulation; Telomerase; Telomeres

Indexed keywords

CYCLIN D; CYCLIN DEPENDENT KINASE 2; CYCLIN DEPENDENT KINASE 4; CYCLIN E; EPIDERMAL GROWTH FACTOR; HISTONE; I KAPPA B KINASE; MAMMALIAN TARGET OF RAPAMYCIN; MESSENGER RNA; MITOGEN ACTIVATED PROTEIN KINASE; PHOSPHATIDYLINOSITOL 3 KINASE; PHOSPHATIDYLINOSITOL 3,4,5 TRISPHOSPHATE 3 PHOSPHATASE; PROTEIN KINASE B; PROTEIN MDM2; PROTEIN P14; PROTEIN P16; PROTEIN P21; PROTEIN P27; PROTEIN P53; RETINOBLASTOMA PROTEIN; SMAD3 PROTEIN; SMAD4 PROTEIN; TELOMERASE; TRANSCRIPTION FACTOR E2F1; TRANSCRIPTION FACTOR E2F4; TRANSCRIPTION FACTOR E2F5; TRANSFORMING GROWTH FACTOR BETA; TRANSFORMING GROWTH FACTOR BETA RECEPTOR 1; TRANSFORMING GROWTH FACTOR BETA RECEPTOR 2; UNINDEXED DRUG;

CARCINOGENESIS; CELL DIFFERENTIATION; CELL IMMORTALIZATION; DNA METHYLATION; ENZYME ACTIVATION; ENZYME REGULATION; EPIGENETICS; HISTONE MODIFICATION; HUMAN; ONCOGENE C MYC; PRIORITY JOURNAL; PROMOTER REGION; PROTEIN EXPRESSION; REVIEW; TRANSCRIPTION REGULATION; TUMOR SUPPRESSOR GENE;

CATALYTIC DOMAIN; EPIGENESIS, GENETIC; HUMANS; ONCOGENES; PROMOTER REGIONS, GENETIC; REPRESSOR PROTEINS; TELOMERASE; TRANSCRIPTION FACTORS; TUMOR SUPPRESSOR PROTEINS;

EID: 84858750992     PISSN: 03781119     EISSN: 18790038     Source Type: Journal    
DOI: 10.1016/j.gene.2012.01.095     Document Type: Review

References (199)

1. Abe S., et al. Localization of an hTERT repressor region on human chromosome 3p21.3 using chromosome engineering. Genome Integr. 2010, 1(1):6.
2. Artandi S.E., et al. Constitutive telomerase expression promotes mammary carcinomas in aging mice. Proc. Natl. Acad. Sci. U. S. A. 2002, 99(12):8191-8196.
3. Asker C., Wiman K.G., Selivanova G. p-53-induced apoptosis as a safeguard against cancer. Biochem. Biophys. Res. Commun. 1999, 265(1):1-6.
4. Atkinson S.P., Hoare S.F., Glasspool R.M., Keith W.N. Lack of telomerase gene expression in alternative lengthening of telomere cells is associated with chromatin remodeling of the hTR and hTERT gene promoters. Cancer Res. 2005, 65(17):7585-7590.
5. Bai D., Ueno L., Vogt P.K. Akt-mediated regulation of NFkappaB and the essentialness of NFkappaB for the oncogenicity of P13K and Akt. Int. J. Cancer 2009, 125(12):2863-2870.
6. Beattie T.L., Zhou W., Robinson M.O., Harrington L. Reconstitution of human telomerase activity in vitro. Curr. Biol. 1998, 8(3):177-180.
7. Beijersbergen R.L., Hijmans E.M., Zhu L., Bernards R. Interaction of c-Myc with the pRb-related protein p107 results in inhibition of c-Myc-mediated transactivation. EMBO J. 1994, 13(17):4080-4086.
8. Berletch J.B., et al. Epigenetic and genetic mechanisms contribute to telomerase inhibition by EGCG. J. Cell. Biochem. 2008, 103(2):509-519.
9. Bilaud T., et al. Telomeric localization of TRF2, a novel human telobox protein. Nat. Genet. 1997, 17(2):236-239.
10. Blackburn E.H. Switching and signaling at the telomere. Cell 2001, 106(6):661-673.
11. Blackwood E.M., Eisenman R.N. Max: a helix-loop-helix zipper protein that forms a sequence-specific DNA-binding complex with Myc. Science 1991, 251(4998):1211-1217.
12. Broccoli D., Godley L.A., Donehower L.A., Varmus H.E., de Lange T. Telomerase activation in mouse mammary tumors: lack of detectable telomere shortening and evidence for regulation of telomerase RNA with cell proliferation. Mol. Cell. Biol. 1996, 16(7):3765-3772.
13. Bryce L.A., Morrison N., Hoare S.F., Muir S., Keith W.N. Mapping of the gene for the human telomerase reverse transcriptase, hTERT, to chromosome 5p15.33 by fluorescence in situ hybridization. Neoplasia 2000, 2(3):197-201.
14. Buchholz M., et al. Overexpression of c-myc in pancreatic cancer caused by ectopic activation of NFATc1 and the Ca2+/calcineurin signaling pathway. EMBO J. 2006, 25(15):3714-3724.
15. Buchkovich K.J., Greider C.W. Telomerase regulation during entry into the cell cycle in normal human T cells. Mol. Biol. Cell 1996, 7(9):1443-1454.
16. Burge S., Parkinson G.N., Hazel P., Todd A.K., Neidle S. Quadruplex DNA: sequence, topology and structure. Nucleic Acids Res. 2006, 34(19):5402-5415.
17. Casillas M.A., Brotherton S.L., Andrews L.G., Ruppert J.M., Tollefsbol T.O. Induction of endogenous telomerase (hTERT) by c-Myc in WI-38 fibroblasts transformed with specific genetic elements. Gene 2003, 316:57-65.
18. Casillas M.A., Lopatina N., Andrews L.G., Tollefsbol T.O. Transcriptional control of the DNA methyltransferases is altered in aging and neoplastically-transformed human fibroblasts. Mol. Cell. Biochem. 2003, 252(1-2):33-43.
19. Chebel A., Ffrench M. Transcriptional regulation of the human telomerase reverse transcriptase: New insights. Transcription 2010, 1(1):27-31.
20. Chebel A., et al. Transcriptional activation of hTERT, the human telomerase reverse transcriptase, by nuclear factor of activated T cells. J. Biol. Chem. 2009, 284(51):35725-35734.
21. Chen C.R., Kang Y., Siegel P.M., Massagué J. E2F4/5 and p107 as Smad cofactors linking the TGFbeta receptor to c-myc repression. Cell 2002, 110(1):19-32.
22. Chen Y., et al. Decreased PITX1 homeobox gene expression in human lung cancer. Lung Cancer 2007, 55(3):287-294.
23. Chen Y.N., Chen H., Xu Y., Zhang X., Luo Y. Expression of pituitary homeobox 1 gene in human gastric carcinogenesis and its clinicopathological significance. World J. Gastroenterol. 2008, 14(2):292-297.
24. Cherlet T., Murphy L.C. Estrogen receptors inhibit Smad3 transcriptional activity through Ap-1 transcription factors. Mol. Cell. Biochem. 2007, 306(1-2):33-42.
25. Chikhirzhina G.I., Al'-Shekhadat R.I., Chikhirzhina E.V. Transcription factors of the nuclear factor 1 (NF1) family. Role in chromatin remodelation. Mol. Biol. (Mosk) 2008, 42(3):388-404.
26. Chiu C.P., Harley C.B. Replicative senescence and cell immortality: the role of telomeres and telomerase. Proc. Soc. Exp. Biol. Med. 1997, 214(2):99-106.
27. Choi K.C., et al. Epigallocatechin-3-gallate, a histone acetyltransferase inhibitor, inhibits EBV-induced B lymphocyte transformation via suppression of RelA acetylation. Cancer Res. 2009, 69(2):583-592.
28. Choi J.H., et al. TSA-induced DNMT1 down-regulation represses hTERT expression via recruiting CTCF into demethylated core promoter region of hTERT in HCT116. Biochem. Biophys. Res. Commun. 2010, 391(1):449-454.
29. Chong L., et al. A human telomeric protein. Science 1995, 270(5242):1663-1667.
30. Chou C.K., et al. The suppression of MAD1 by AKT-mediated phosphorylation activates MAD1 target genes transcription. Mol. Carcinog. 2009, 48(11):1048-1058.
31. Chun A.C., Jin D.Y. Transcriptional regulation of mitotic checkpoint gene MAD1 by p53. J. Biol. Chem. 2003, 278(39):37439-37450.
32. Cline M.S., et al. Integration of biological networks and gene expression data using Cytoscape. Nat. Protoc. 2007, 2(10):2366-2382.
33. Cohen S.B., et al. Protein composition of catalytically active human telomerase from immortal cells. Science 2007, 315(5820):1850-1853.
34. Collins K., Mitchell J.R. Telomerase in the human organism. Oncogene 2002, 21(4):564-579.
35. Cong Y.S., Bacchetti S. Histone deacetylation is involved in the transcriptional repression of hTERT in normal human cells. J. Biol. Chem. 2000, 275(46):35665-35668.
36. Cong Y.S., Wen J., Bacchetti S. The human telomerase catalytic subunit hTERT: organization of the gene and characterization of the promoter. Hum. Mol. Genet. 1999, 8(1):137-142.
37. Cong Y.S., Wright W.E., Shay J.W. Human telomerase and its regulation. Microbiol. Mol. Biol. Rev. 2002, 66(3):407-425. table of contents.
38. Crowe D.L., Nguyen D.C., Tsang K.J., Kyo S. E2F-1 represses transcription of the human telomerase reverse transcriptase gene. Nucleic Acids Res. 2001, 29(13):2789-2794.
39. Cukusić A., Skrobot Vidacek N., Sopta M., Rubelj I. Telomerase regulation at the crossroads of cell fate. Cytogenet. Genome Res. 2008, 122(3-4):263-272.
40. Davis J.T. G-quartets 40 years later: from 5'-GMP to molecular biology and supramolecular chemistry. Angew. Chem. Int. Ed. Engl. 2004, 43(6):668-698.
41. De Boeck G., Forsyth R.G., Praet M., Hogendoorn P.C. Telomere-associated proteins: cross-talk between telomere maintenance and telomere-lengthening mechanisms. J. Pathol. 2009, 217(3):327-344.
42. Deng W.G., et al. Tumor-specific activation of human telomerase reverses transcriptase promoter activity by activating enhancer-binding protein-2beta in human lung cancer cells. J. Biol. Chem. 2007, 282(36):26460-26470.
43. Dessain S.K., Yu H., Reddel R.R., Beijersbergen R.L., Weinberg R.A. Methylation of the human telomerase gene CpG island. Cancer Res. 2000, 60(3):537-541.
44. Devereux T.R., et al. DNA methylation analysis of the promoter region of the human telomerase reverse transcriptase (hTERT) gene. Cancer Res. 1999, 59(24):6087-6090.
45. Eickelberg O., Centrella M., Reiss M., Kashgarian M., Wells R.G. Betaglycan inhibits TGF-beta signaling by preventing type I-type II receptor complex formation. Glycosaminoglycan modifications alter betaglycan function. J. Biol. Chem. 2002, 277(1):823-829.
46. Farwell D.G., et al. Genetic and epigenetic changes in human epithelial cells immortalized by telomerase. Am. J. Pathol. 2000, 156(5):1537-1547.
47. Feng Z., et al. The regulation of AMPK beta1, TSC2, and PTEN expression by p53: stress, cell and tissue specificity, and the role of these gene products in modulating the IGF-1-AKT-mTOR pathways. Cancer Res. 2007, 67(7):3043-3053.
48. Filippova G.N., et al. An exceptionally conserved transcriptional repressor, CTCF, employs different combinations of zinc fingers to bind diverged promoter sequences of avian and mammalian c-myc oncogenes. Mol. Cell. Biol. 1996, 16(6):2802-2813.
49. Fujimoto K., et al. 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. 2000, 28(13):2557-2562.
50. Gao J., Chen D., Tian Y., Zhang J., Cai K. Effect of estrogen on telomerase activity in human breast cancer cells. J. Huazhong Univ. Sci. Technolog. Med. Sci. 2003, 23(3):286-287. 293.
51. Gellert M., Lipsett M.N., Davies D.R. Helix formation by guanylic acid. Proc. Natl. Acad. Sci. U. S. A. 1962, 48:2013-2018.
52. Goerttler P.S., et al. Gene expression profiling in polycythaemia vera: overexpression of transcription factor NF-E2. Br. J. Haematol. 2005, 129(1):138-150.
53. Gombert W.M., Krumm A. Targeted deletion of multiple CTCF-binding elements in the human C-MYC gene reveals a requirement for CTCF in C-MYC expression. PLoS One 2009, 4(7):e6109.
54. Gomez D., et al. Telomerase downregulation induced by the G-quadruplex ligand 12459 in A549 cells is mediated by hTERT RNA alternative splicing. Nucleic Acids Res. 2004, 32(1):371-379.
55. González-Suárez E., et al. Increased epidermal tumors and increased skin wound healing in transgenic mice overexpressing the catalytic subunit of telomerase, mTERT, in basal keratinocytes. EMBO J. 2001, 20(11):2619-2630.
56. González-Suárez E., Flores J.M., Blasco M.A. Cooperation between p53 mutation and high telomerase transgenic expression in spontaneous cancer development. Mol. Cell. Biol. 2002, 22(20):7291-7301.
57. Goueli B.S., Janknecht R. Regulation of telomerase reverse transcriptase gene activity by upstream stimulatory factor. Oncogene 2003, 22(39):8042-8047.
58. Grand C.L., et al. The cationic porphyrin TMPyP4 down-regulates c-MYC and human telomerase reverse transcriptase expression and inhibits tumor growth in vivo. Mol. Cancer Ther. 2002, 1(8):565-573.
59. Griffith J.D., et al. Mammalian telomeres end in a large duplex loop. Cell 1999, 97(4):503-514.
60. Gros J., Guédin A., Mergny J.L., Lacroix L. G-Quadruplex formation interferes with P1 helix formation in the RNA component of telomerase hTERC. ChemBioChem 2008, 9(13):2075-2079.
61. Günes C., Lichtsteiner S., Vasserot A.P., Englert C. Expression of the hTERT gene is regulated at the level of transcriptional initiation and repressed by Mad1. Cancer Res. 2000, 60(8):2116-2121.
62. Guo C., Armbruster B.N., Price D.T., Counter C.M. In vivo regulation of hTERT expression and telomerase activity by androgen. J. Urol. 2003, 170(2 Pt 1):615-618.
63. Guo Y., Pajovic S., Gallie B.L. Expression of p14ARF, MDM2, and MDM4 in human retinoblastoma. Biochem. Biophys. Res. Commun. 2008, 375(1):1-5.
64. Hahn W.C., et al. Creation of human tumour cells with defined genetic elements. Nature 1999, 400(6743):464-468.
65. Han D.F., et al. Polymorphisms of estrogen-metabolizing genes and breast cancer risk: a multigenic study. Chin. Med. J. (Engl) 2005, 118(18):1507-1516.
66. Harley C.B., Futcher A.B., Greider C.W. Telomeres shorten during ageing of human fibroblasts. Nature 1990, 345(6274):458-460.
67. Harper J.W., Adami G.R., Wei N., Keyomarsi K., Elledge S.J. The p21 Cdk-interacting protein Cip1 is a potent inhibitor of G1 cyclin-dependent kinases. Cell 1993, 75(4):805-816.
68. Hein N., Jiang K., Cornelissen C., Lüscher B. TGFβ1 enhances MAD1 expression and stimulates promoter-bound Pol II phosphorylation: basic functions of C/EBP, SP and SMAD3 transcription factors. BMC Mol. Biol. 2011, 12:9.
69. Henderson Y.C., Breau R.L., Liu T.J., Clayman G.L. Telomerase activity in head and neck tumors after introduction of wild-type p53, p21, p16, and E2F-1 genes by means of recombinant adenovirus. Head Neck 2000, 22(4):347-354.
70. Hooker C.W., Hurlin P.J. Of Myc and Mnt. J. Cell Sci. 2006, 119(Pt 2):208-216.
71. Horikawa I., Cable P.L., Afshari C., Barrett J.C. Cloning and characterization of the promoter region of human telomerase reverse transcriptase gene. Cancer Res. 1999, 59(4):826-830.
72. Horikawa I., et al. 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 2002, 13(8):2585-2597.
73. Hsu M.C., Chang H.C., Hung W.C. HER-2/neu transcriptionally activates Jab1 expression via the AKT/beta-catenin pathway in breast cancer cells. Endocr. Relat. Cancer 2007, 14(3):655-667.
74. Hsu M.C., et al. Jab1 is overexpressed in human breast cancer and is a downstream target for HER-2/neu. Mod. Pathol. 2008, 21(5):609-616.
75. Iliopoulos D., Satra M., Drakaki A., Poultsides G.A., Tsezou A. Epigenetic regulation of hTERT promoter in hepatocellular carcinomas. Int. J. Oncol. 2009, 34(2):391-399.
76. Imachi H., et al. Menin, a product of the MENI gene, binds to estrogen receptor to enhance its activity in breast cancer cells: possibility of a novel predictive factor for tamoxifen resistance. Breast Cancer Res. Treat. 2010, 122(2):395-407.
77. Ishikawa F. Regulation mechanisms of mammalian telomerase. A review. Biochemistry (Mosc) 1997, 62(11):1332-1337.
78. James L., Eisenman R.N. Myc and Mad bHLHZ domains possess identical DNA-binding specificities but only partially overlapping functions in vivo. Proc. Natl. Acad. Sci. U. S. A. 2002, 99(16):10429-10434.
79. Jiang K., Hein N., Eckert K., Lüscher-Firzlaff J., Lüscher B. Regulation of the MAD1 promoter by G-CSF. Nucleic Acids Res. 2008, 36(5):1517-1531.
80. Jin W., et al. Involvement of MyoD and c-myb in regulation of basal and estrogen-induced transcription activity of the BRCA1 gene. Breast Cancer Res. Treat. 2011, 125(3):699-713.
81. Johnson T.L., Lai M.B., Lai J.C., Bhushan A. Inhibition of Cell Proliferation and MAP Kinase and Akt Pathways in Oral Squamous cell Carcinoma by Genistein and Biochanin A. Evid. Based Complement. Altern. Med. 2010, 7(3):351-358.
82. Jung P., Hermeking H. The c-MYC-AP4-p21 cascade. Cell Cycle 2009, 8(7):982-989.
83. Jung P., Menssen A., Mayr D., Hermeking H. AP4 encodes a c-MYC-inducible repressor of p21. Proc. Natl. Acad. Sci. U. S. A. 2008, 105(39):15046-15051.
84. Kamata H., Manabe T., Oka S., Kamata K., Hirata H. Hydrogen peroxide activates IkappaB kinases through phosphorylation of serine residues in the activation loops. FEBS Lett. 2002, 519(1-3):231-237.
85. Kanaya T., et al. Adenoviral expression of p53 represses telomerase activity through down-regulation of human telomerase reverse transcriptase transcription. Clin. Cancer Res. 2000, 6(4):1239-1247.
86. Kanduri C., et al. Functional association of CTCF with the insulator upstream of the H19 gene is parent of origin-specific and methylation-sensitive. Curr. Biol. 2000, 10(14):853-856.
87. Karlseder J., Broccoli D., Dai Y., Hardy S., de Lange T. p53- and ATM-dependent apoptosis induced by telomeres lacking TRF2. Science 1999, 283(5406):1321-1325.
88. Karlseder J., Smogorzewska A., de Lange T. Senescence induced by altered telomere state, not telomere loss. Science 2002, 295(5564):2446-2449.
89. Khan S.J., et al. Interactions of TRF2 with model telomeric ends. Biochem. Biophys. Res. Commun. 2007, 363(1):44-50.
90. Kikuno N., et al. Genistein mediated histone acetylation and demethylation activates tumor suppressor genes in prostate cancer cells. Int. J. Cancer 2008, 123(3):552-560.
91. Kim J.H., et al. Ubiquitin ligase MKRN1 modulates telomere length homeostasis through a proteolysis of hTERT. Genes Dev. 2005, 19(7):776-781.
92. Kimura A., et al. Induction of hTERT expression and phosphorylation by estrogen via Akt cascade in human ovarian cancer cell lines. Oncogene 2004, 23(26):4505-4515.
93. Kirkpatrick K.L., et al. hTERT expression in human breast cancer and non-cancerous breast tissue: correlation with tumour stage and c-Myc expression. Breast Cancer Res. Treat. 2003, 77(3):277-284.
94. Klenova E.M., et al. CTCF, a conserved nuclear factor required for optimal transcriptional activity of the chicken c-myc gene, is an 11-Zn-finger protein differentially expressed in multiple forms. Mol. Cell. Biol. 1993, 13(12):7612-7624.
95. Ko L.J., Prives C., et al. p53: puzzle and paradigm. Genes Dev. 1996, 10(9):1054-1072.
96. Kyo S., et al. Estrogen activates telomerase. Cancer Res. 1999, 59(23):5917-5921.
97. Kyo S., et al. Sp1 cooperates with c-Myc to activate transcription of the human telomerase reverse transcriptase gene (hTERT). Nucleic Acids Res. 2000, 28(3):669-677.
98. Kyo S., Takakura M., Fujiwara T., Inoue M. Understanding and exploiting hTERT promoter regulation for diagnosis and treatment of human cancers. Cancer Sci. 2008, 99(8):1528-1538.
99. Lacerte A., et al. Transforming growth factor-beta inhibits telomerase through SMAD3 and E2F transcription factors. Cell. Signal. 2008, 20(1):50-59.
100. Lai S.R., Cunningham A.P., Huynh V.Q., Andrews L.G., Tollefsbol T.O. Evidence of extra-telomeric effects of hTERT and its regulation involving a feedback loop. Exp. Cell Res. 2007, 313(2):322-330.
101. Lee S.H., et al. Interferon regulatory factor-1 (IRF-1) is a mediator for interferon-gamma induced attenuation of telomerase activity and human telomerase reverse transcriptase (hTERT) expression. Oncogene 2003, 22(3):381-391.
102. Lee J.H., Lee S.Y., Lee S.H. p21 WAF1 is involved in interferon-β-induced attenuation of telomerase activity and human telomerase reverse transcriptase (hTERT) expression in ovarian cancer. Mol. Cells 2010, 30(4):327-333.
103. Leem S., et al. The human telomerase gene: complete genomic sequence and analysis of tandem repeat polymorphisms in intronic regions. Oncogene 2002, 21(5):769-777.
104. Leng X., Noble M., Adams P.D., Qin J., Harper J.W. Reversal of growth suppression by p107 via direct phosphorylation by cyclin D1/cyclin-dependent kinase 4. Mol. Cell. Biol. 2002, 22(7):2242-2254.
105. Li Y., Sarkar F.H. Inhibition of nuclear factor kappaB activation in PC3 cells by genistein is mediated via Akt signaling pathway. Clin. Cancer Res. 2002, 8(7):2369-2377.
106. Li H., Lee T.H., Avraham H. 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. 2002, 277(23):20965-20973.
107. Li Y., Liu L., Andrews L.G., Tollefsbol T.O. Genistein depletes telomerase activity through cross-talk between genetic and epigenetic mechanisms. Int. J. Cancer 2009, 125(2):286-296.
108. Li A.Y., et al. High-Mobility Group A2 Protein Modulates hTERT Transcription To Promote Tumorigenesis. Mol. Cell. Biol. 2011, 31(13):2605-2617.
109. Li J., Huang X., Xie X., Wang J., Duan M. Human telomerase reverse transcriptase regulates cyclin D1 and G1/S phase transition in laryngeal squamous carcinoma. Acta Otolaryngol. 2011, 131(5):546-551.
110. Lim K.W., et al. Coexistence of two distinct G-quadruplex conformations in the hTERT promoter. J. Am. Chem. Soc. 2010, 132(35):12331-12342.
111. Liu L., Lai S., Andrews L.G., Tollefsbol T.O. Genetic and epigenetic modulation of telomerase activity in development and disease. Gene 2004, 340(1):1-10.
112. Liu L., Saldanha S.N., Pate M.S., Andrews L.G., Tollefsbol T.O. Epigenetic regulation of human telomerase reverse transcriptase promoter activity during cellular differentiation. Genes Chromosomes Cancer 2004, 41(1):26-37.
113. Liu C., et al. The telomerase reverse transcriptase (hTERT) gene is a direct target of the histone methyltransferase SMYD3. Cancer Res. 2007, 67(6):2626-2631.
114. Liu L., et al. MCAF1/AM is involved in Sp1-mediated maintenance of cancer-associated telomerase activity. J. Biol. Chem. 2009, 284(8):5165-5174.
115. Lopatina N.G., et al. Control mechanisms in the regulation of telomerase reverse transcriptase expression in differentiating human teratocarcinoma cells. Biochem. Biophys. Res. Commun. 2003, 306(3):650-659.
116. Lutz M., et al. Transcriptional repression by the insulator protein CTCF involves histone deacetylases. Nucleic Acids Res. 2000, 28(8):1707-1713.
117. Majid S., et al. Genistein induces the p21WAF1/CIP1 and p16INK4a tumor suppressor genes in prostate cancer cells by epigenetic mechanisms involving active chromatin modification. Cancer Res. 2008, 68(8):2736-2744.
118. Makarov V.L., Hirose Y., Langmore J.P. Long G tails at both ends of human chromosomes suggest a C strand degradation mechanism for telomere shortening. Cell 1997, 88(5):657-666.
119. Mao B., et al. Sirt1 deacetylates c-Myc and promotes c-Myc/Max association. Int. J. Biochem. Cell Biol. 2011, 43(11):1573-1581.
120. Masutomi K., et al. Telomerase maintains telomere structure in normal human cells. Cell 2003, 114(2):241-253.
121. McArthur G.A., et al. MAD1 and p27(KIP1) cooperate to promote terminal differentiation of granulocytes and to inhibit Myc expression and cyclin E-CDK2 activity. Mol. Cell. Biol. 2002, 22(9):3014-3023.
122. Meeran S.M., Katiyar S., Katiyar S.K. Berberine-induced apoptosis in human prostate cancer cells is initiated by reactive oxygen species generation. Toxicol. Appl. Pharmacol. 2008, 229(1):33-43.
123. Meeran S.M., Ahmed A., Tollefsbol T.O. Epigenetic targets of bioactive dietary components for cancer prevention and therapy. Clin. Epigenetics 2010, 1(3-4):101-116.
124. Meeran S.M., Patel S.N., Tollefsbol T.O. Sulforaphane causes epigenetic repression of hTERT expression in human breast cancer cell lines. PLoS One 2010, 5(7):e11457.
125. Meeran S.M., Patel S.N., Chan T.H., Tollefsbol T.O. A Novel Prodrug of Epigallocatechin-3-gallate: Differential Epigenetic hTERT Repression in Human Breast Cancer Cells. Cancer Prev. Res. (Phila) 2011, 4(8):1243-1254.
126. Meyerson M., et al. hEST2, the putative human telomerase catalytic subunit gene, is up-regulated in tumor cells and during immortalization. Cell 1997, 90(4):785-795.
127. Misiti S., et al. Induction of hTERT expression and telomerase activity by estrogens in human ovary epithelium cells. Mol. Cell. Biol. 2000, 20(11):3764-3771.
128. Moore L., et al. Cooperativity of p19ARF, Mdm2, and p53 in murine tumorigenesis. Oncogene 2003, 22(49):7831-7837.
129. Moyzis R.K., et al. A highly conserved repetitive DNA sequence, (TTAGGG)n, present at the telomeres of human chromosomes. Proc. Natl. Acad. Sci. U. S. A. 1988, 85(18):6622-6626.
130. Murphree A.L., Benedict W.F. Retinoblastoma: clues to human oncogenesis. Science 1984, 223(4640):1028-1033.
131. Nakamura T.M., et al. Telomerase catalytic subunit homologs from fission yeast and human. Science 1997, 277(5328):955-959.
132. Nanni S., et al. Signaling through estrogen receptors modulates telomerase activity in human prostate cancer. J. Clin. Invest. 2002, 110(2):219-227.
133. Nguyen D.C., Crowe D.L. Intact functional domains of the retinoblastoma gene product (pRb) are required for downregulation of telomerase activity. Biochim. Biophys. Acta 1999, 1445(2):207-215.
134. Ogawa H., Murayama A., Nagata S., Fukunaga R. Regulation of myeloid zinc finger protein 2A transactivation activity through phosphorylation by mitogen-activated protein kinases. J. Biol. Chem. 2003, 278(5):2921-2927.
135. Ogawara Y., et al. Akt enhances Mdm2-mediated ubiquitination and degradation of p53. J. Biol. Chem. 2002, 277(24):21843-21850.
136. Oh S., Song Y.H., Yim J., Kim T.K. Identification of Mad as a repressor of the human telomerase (hTERT) gene. Oncogene 2000, 19(11):1485-1490.
137. Oh W., et al. Jab1 induces the cytoplasmic localization and degradation of p53 in coordination with Hdm2. J. Biol. Chem. 2006, 281(25):17457-17465.
138. Papanikolaou V., et al. hTERT regulation by NF-κB and c-myc in irradiated HER2-positive breast cancer cells. Int. J. Radiat. Biol. 2011, 87(6):609-621.
139. Parisi F., Wirapati P., Naef F. Identifying synergistic regulation involving c-Myc and sp1 in human tissues. Nucleic Acids Res. 2007, 35(4):1098-1107.
140. Patel D.J., Phan A.T., Kuryavyi V. Human telomere, oncogenic promoter and 5'-UTR G-quadruplexes: diverse higher order DNA and RNA targets for cancer therapeutics. Nucleic Acids Res. 2007, 35(22):7429-7455.
141. Perrault S.D., Hornsby P.J., Betts D.H. Global gene expression response to telomerase in bovine adrenocortical cells. Biochem. Biophys. Res. Commun. 2005, 335(3):925-936.
142. Phuong N.T., et al. Role of PTEN promoter methylation in tamoxifen-resistant breast cancer cells. Breast Cancer Res. Treat. 2011, 130(1):73-83.
143. Potischman N., Swanson C.A., Siiteri P., Hoover R.N. Reversal of relation between body mass and endogenous estrogen concentrations with menopausal status. J. Natl. Cancer Inst. 1996, 88(11):756-758.
144. Qi D.L., et al. Identification of PITX1 as a TERT suppressor gene located on human chromosome 5. Mol. Cell. Biol. 2011, 31(8):1624-1636.
145. Quintana A.M., Liu F., O'Rourke J.P., Ness S.A. Identification and regulation of c-Myb target genes in MCF-7 cells. BMC Cancer 2011, 11:30.
146. Rahmati-Yamchi M., Zarghami N., Rahbani M., Montazeri A. Plasma Leptin, hTERT Gene Expression, and Anthropometric Measures in Obese and Non-Obese Women with Breast Cancer. Breast Cancer (Auckl) 2011, 5:27-35.
147. Ray A., James M.K., Larochelle S., Fisher R.P., Blain S.W. p27Kip1 inhibits cyclin D-cyclin-dependent kinase 4 by two independent modes. Mol. Cell. Biol. 2009, 29(4):986-999.
148. Ren Y., et al. Dual effects of TGF-beta on ERalpha-mediated estrogenic transcriptional activity in breast cancer. Mol. Cancer 2009, 8:111.
149. Ren H., et al. Leptin upregulates telomerase activity and transcription of human telomerase reverse transcriptase in MCF-7 breast cancer cells. Biochem. Biophys. Res. Commun. 2010, 394(1):59-63.
150. Renaud S., Loukinov D., Bosman F.T., Lobanenkov V., Benhattar J. CTCF binds the proximal exonic region of hTERT and inhibits its transcription. Nucleic Acids Res. 2005, 33(21):6850-6860.
151. Renaud S., et al. Dual role of DNA methylation inside and outside of CTCF-binding regions in the transcriptional regulation of the telomerase hTERT gene. Nucleic Acids Res. 2007, 35(4):1245-1256.
152. Renaud S., et al. BORIS/CTCFL-mediated transcriptional regulation of the hTERT telomerase gene in testicular and ovarian tumor cells. Nucleic Acids Res. 2011, 39(3):862-873.
153. Rose D.P., Haffner S.M., Baillargeon J. Adiposity, the metabolic syndrome, and breast cancer in African-American and white American women. Endocr. Rev. 2007, 28(7):763-777.
154. Saito T., et al. Comparative gene mapping of the human and mouse TEP1 genes, which encode one protein component of telomerases. Genomics 1997, 46(1):46-50.
155. Salvatico J., Kim J.H., Chung I.K., Muller M.T. Differentiation linked regulation of telomerase activity by Makorin-1. Mol. Cell. Biochem. 2010, 342(1-2):241-250.
156. Sasaki R., et al. Oncogenic transformation of human ovarian surface epithelial cells with defined cellular oncogenes. Carcinogenesis 2009, 30(3):423-431.
157. Shats I., et al. p53-dependent down-regulation of telomerase is mediated by p21waf1. J. Biol. Chem. 2004, 279(49):50976-50985.
158. Shin K.H., Kang M.K., Dicterow E., Park N.H. Hypermethylation of the hTERT promoter inhibits the expression of telomerase activity in normal oral fibroblasts and senescent normal oral keratinocytes. Br. J. Cancer 2003, 89(8):1473-1478.
159. Simonsson T. G-quadruplex DNA structures-variations on a theme. Biol. Chem. 2001, 382(4):621-628.
160. Sitaram R.T., et al. Wilms' tumour 1 can suppress hTERT gene expression and telomerase activity in clear cell renal cell carcinoma via multiple pathways. Br. J. Cancer 2010, 103(8):1255-1262.
161. Smith L.L., Coller H.A., Roberts J.M. Telomerase modulates expression of growth-controlling genes and enhances cell proliferation. Nat. Cell Biol. 2003, 5(5):474-479.
162. Song J., et al. Genomic organization and expression of a human gene for Myc-associated zinc finger protein (MAZ). J. Biol. Chem. 1998, 273(32):20603-20614.
163. Song J., et al. Transcriptional regulation by zinc-finger proteins Sp1 and MAZ involves interactions with the same cis-elements. Int. J. Mol. Med. 2003, 11(5):547-553.
164. Stanelle J., Stiewe T., Theseling C.C., Peter M., Pützer B.M. Gene expression changes in response to E2F1 activation. Nucleic Acids Res. 2002, 30(8):1859-1867.
165. Stansel R.M., de Lange T., Griffith J.D. T-loop assembly in vitro involves binding of TRF2 near the 3' telomeric overhang. EMBO J. 2001, 20(19):5532-5540.
166. Stefanou N., Papanikolaou V., Furukawa Y., Nakamura Y., Tsezou A. Leptin as a critical regulator of hepatocellular carcinoma development through modulation of human telomerase reverse transcriptase. BMC Cancer 2010, 10:442.
167. Stein G.S., Montecino M., van Wijnen A.J., Stein J.L., Lian J.B. Nuclear structure-gene expression interrelationships: implications for aberrant gene expression in cancer. Cancer Res. 2000, 60(8):2067-2076.
168. Stewart S.A., et al. Telomerase contributes to tumorigenesis by a telomere length-independent mechanism. Proc. Natl. Acad. Sci. U. S. A. 2002, 99(20):12606-12611.
169. Sun D., et al. Inhibition of human telomerase by a G-quadruplex-interactive compound. J. Med. Chem. 1997, 40(14):2113-2116.
170. Suphapeetiporn K., Greally J.M., Walpita D., Ashley T., Bale A.E. MEN1 tumor-suppressor protein localizes to telomeres during meiosis. Genes Chromosomes Cancer 2002, 35(1):81-85.
171. Suske G. The Sp-family of transcription factors. Gene 1999, 238(2):291-300.
172. Takakura M., et al. 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. 1999, 59(3):551-557.
173. Takakura M., Kyo S., Inoue M., Wright W.E., Shay J.W. Function of AP-1 in transcription of the telomerase reverse transcriptase gene (TERT) in human and mouse cells. Mol. Cell. Biol. 2005, 25(18):8037-8043.
174. Takano H., Murasawa S., Asahara T. Functional and gene expression analysis of hTERT overexpressed endothelial cells. Biologics 2008, 2(3):547-554.
175. Tee M.K., et al. Estradiol and selective estrogen receptor modulators differentially regulate target genes with estrogen receptors alpha and beta. Mol. Biol. Cell 2004, 15(3):1262-1272.
176. Tomoda K., et al. The cytoplasmic shuttling and subsequent degradation of p27Kip1 mediated by Jab1/CSN5 and the COP9 signalosome complex. J. Biol. Chem. 2002, 277(3):2302-2310.
177. van Steensel B., Smogorzewska A., de Lange T. TRF2 protects human telomeres from end-to-end fusions. Cell 1998, 92(3):401-413.
178. Varfolomeev E., et al. IAP antagonists induce autoubiquitination of c-IAPs, NF-kappaB activation, and TNFalpha-dependent apoptosis. Cell 2007, 131(4):669-681.
179. Vostrov A.A., Quitschke W.W. The zinc finger protein CTCF binds to the APBbeta domain of the amyloid beta-protein precursor promoter. Evidence for a role in transcriptional activation. J. Biol. Chem. 1997, 272(52):33353-33359.
180. Wan M., et al. Jab1 antagonizes TGF-beta signaling by inducing Smad4 degradation. EMBO Rep. 2002, 3(2):171-176.
181. Wang Z., et al. Tamoxifen regulates human telomerase reverse transcriptase (hTERT) gene expression differently in breast and endometrial cancer cells. Oncogene 2002, 21(22):3517-3524.
182. Wang C., et al. Activation of p27Kip1 Expression by E2F1. A negative feedback mechanism. J. Biol. Chem. 2005, 280(13):12339-12343.
183. Wang P., Greiner T.C., Lushnikova T., Eischen C.M. Decreased Mdm2 expression inhibits tumor development induced by loss of ARF. Oncogene 2006, 25(26):3708-3718.
184. Wang J., et al. Jab1 is a target of EGFR signaling in ERalpha-negative breast cancer. Breast Cancer Res. 2008, 10(3):R51.
185. Weinrich S.L., et al. Reconstitution of human telomerase with the template RNA component hTR and the catalytic protein subunit hTRT. Nat. Genet. 1997, 17(4):498-502.
186. Wellinger R.J., Sen D. The DNA structures at the ends of eukaryotic chromosomes. Eur. J. Cancer 1997, 33(5):735-749.
187. Wick M., Zubov D., Hagen G. Genomic organization and promoter characterization of the gene encoding the human telomerase reverse transcriptase (hTERT). Gene 1999, 232(1):97-106.
188. Wisman G.B., et al. Telomerase in relation to expression of p53, c-Myc and estrogen receptor in ovarian tumours. Int. J. Oncol. 2003, 23(5):1451-1459.
189. Won J., Yim J., Kim T.K. Sp1 and Sp3 recruit histone deacetylase to repress transcription of human telomerase reverse transcriptase (hTERT) promoter in normal human somatic cells. J. Biol. Chem. 2002, 277(41):38230-38238.
190. Wu L., et al. Smad4 as a transcription corepressor for estrogen receptor alpha. J. Biol. Chem. 2003, 278(17):15192-15200.
191. Wu S., et al. TGF-beta enforces senescence in Myc-transformed hematopoietic tumor cells through induction of Mad1 and repression of Myc activity. Exp. Cell Res. 2009, 315(18):3099-3111.
192. Xu H.J., et al. Reexpression of the retinoblastoma protein in tumor cells induces senescence and telomerase inhibition. Oncogene 1997, 15(21):2589-2596.
193. Xu D., et al. 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. U. S. A. 2001, 98(7):3826-3831.
194. Zahler A.M., Williamson J.R., Cech T.R., Prescott D.M. Inhibition of telomerase by G-quartet DNA structures. Nature 1991, 350(6320):718-720.
195. Zheng L., Lee W.H. Retinoblastoma tumor suppressor and genome stability. Adv. Cancer Res. 2002, 85:13-50.
196. Zhou C., Bae-Jump V.L., Whang Y.E., Gehrig P.A., Boggess J.F. The PTEN tumor suppressor inhibits telomerase activity in endometrial cancer cells by decreasing hTERT mRNA levels. Gynecol. Oncol. 2006, 101(2):305-310.
197. Zhu J., Blenis J., Yuan J. Activation of PI3K/Akt and MAPK pathways regulates Myc-mediated transcription by phosphorylating and promoting the degradation of Mad1. Proc. Natl. Acad. Sci. U. S. A. 2008, 105(18):6584-6589.
198. Zinn R.L., Pruitt K., Eguchi S., Baylin S.B., Herman J.G. hTERT is expressed in cancer cell lines despite promoter DNA methylation by preservation of unmethylated DNA and active chromatin around the transcription start site. Cancer Res. 2007, 67(1):194-201.
199. Zou L., Zhang P.H., Luo C.L., Tu Z.G. Transcript regulation of human telomerase reverse transcriptase by c-myc and mad1. Acta Biochim. Biophys. Sin. (Shanghai) 2005, 37(1):32-38.