Human telomerase expression regulation

Biochemistry and Cell Biology

Volumn 89, Issue 4, 2011, Pages 359-376

  Gladych, Marta ^a   Wojtyla, Aneta ^a   Rubis, Blazej ^a  

^a POZNAN UNIVERSITY OF MEDICAL SCIENCES   (Poland)

Author keywords

Cancer; Gene expression regulation; Gene therapy; Telomerase; Telomeres

Indexed keywords

ANTI-CANCER THERAPIES; ANTI-ESTROGENS; ANTISENSE; CANCER; CANCER CELLS; CANCER THERAPY; HUMAN TELOMERASE; MULTISUBUNIT; REGULATION MECHANISMS; RETINOIC ACIDS; RIBOZYMES; TELOMERASE; TELOMERASE EXPRESSION; TELOMERE LENGTH; TELOMERES; TRANSCRIPTION LEVEL; VITAMIN-D;

CHROMOSOMES; COMPLEXATION; GENE THERAPY; GENETIC ENGINEERING; ONCOLOGY; STEM CELLS; TRANSCRIPTION;

DISEASES;

BINDING PROTEIN; BRCA1 PROTEIN; GAMMA INTERFERON; IMMUNOGLOBULIN ENHANCER BINDING PROTEIN; MESSENGER RNA; MYC PROTEIN; PEROXISOME PROLIFERATOR ACTIVATED RECEPTOR GAMMA; PROTEIN P53; PROTEIN P73; RAF PROTEIN; RAS PROTEIN; TELOMERASE; TRANSCRIPTION FACTOR ETS; WT1 PROTEIN;

BINDING SITE; CANCER CELL; CARCINOGENESIS; CATABOLITE REPRESSION; CATALYSIS; CHROMOSOME 5P; ENZYME ACTIVITY; ENZYME BINDING; ENZYME INHIBITION; ENZYME PHOSPHORYLATION; EPIGENETICS; GENE DELETION; GENE EXPRESSION; GENETIC TRANSCRIPTION; HUMAN; LYMPHOCYTE; MACROPHAGE; METHYLATION; PROMOTER REGION; PROTEIN EXPRESSION; PROTEIN FUNCTION; PROTEIN MOTIF; PROTEIN PROCESSING; REGULATORY MECHANISM; REVIEW; STEM CELL; TRANSCRIPTION INITIATION;

DNA-BINDING PROTEINS; EPIGENESIS, GENETIC; GENE EXPRESSION REGULATION; GENE THERAPY; GONADAL STEROID HORMONES; HUMANS; NEOPLASMS; TELOMERASE; TELOMERE; TRANSCRIPTION, GENETIC;

EID: 80051533444     PISSN: 08298211     EISSN: 12086002     Source Type: Journal    
DOI: 10.1139/o11-037     Document Type: Review

References (205)

1. Ahmed S. Passos J.F. Birket M.J. Beckmann T. Brings S. Peters H. et al. 2008. Telomerase does not counteract telomere shortening but protects mitochondrial function under oxidative stress. J. Cell Sci. 121 (7): 1046-1053. 10.1242/jcs.019372 18334557 (Pubitemid 351639423)
2. Akiyama M. Hideshima T. Hayashi T. Tai Y.T. Mitsiades C.S. Mitsiades N. et al. 2002. Cytokines modulate telomerase activity in a human multiple myeloma cell line. Cancer Res. 62 (13): 3876-3882. 12097303 (Pubitemid 34728875)
3. Akiyama M. Kawano T. Mikami-Terao Y. Agawa-Ohta M. Yamada O. Ida H. Yamada H. 2010. Erythropoietin activates telomerase through transcriptional and posttranscriptional regulation in human erythroleukemic JAS-REN-A cells. Leuk. Res. 35 (3): 416-418. 21163530
4. Akutagawa O. Nishi H. Kyo S. Terauchi F. Yamazawa K. Higuma C. et al. 2008. Early growth response-1 mediates downregulation of telomerase in cervical cancer. Cancer Sci. 99 (7): 1401-1406. 10.1111/j.1349-7006.2008.00835.x 18460021 (Pubitemid 351761730)
5. Alder J.K. Chen J.J. Lancaster L. Danoff S. Su S.C. Cogan J.D. et al. 2008. Short telomeres are a risk factor for idiopathic pulmonary fibrosis. Proc. Natl. Acad. Sci. U.S.A. 105 (35): 13051-13056. 10.1073/pnas.0804280105 18753630
6. Alfonso-De Matte M.Y. Yang H. Evans M.S. Cheng J.Q. Kruk P.A. 2002. Telomerase is regulated by c-Jun NH2-terminal kinase in ovarian surface epithelial cells. Cancer Res. 62 (16): 4575-4578. 12183409
7. Armanios M. Chen J.L. Chang Y.P. Brodsky R.A. Hawkins A. Griffin C.A. et al. 2005. Haploinsufficiency of telomerase reverse transcriptase leads to anticipation in autosomal dominant dyskeratosis congenita. Proc. Natl. Acad. Sci. U.S.A. 102 (44): 15960-15964. 10.1073/pnas.0508124102 16247010 (Pubitemid 41552850)
8. Armanios M.Y. Chen J.J. Cogan J.D. Alder J.K. Ingersoll R.G. Markin C. et al. 2007. Telomerase mutations in families with idiopathic pulmonary fibrosis. N. Engl. J. Med. 356 (13): 1317-1326. 10.1056/NEJMoa066157 17392301 (Pubitemid 46513297)
9. Artandi S.E. DePinho R.A. 2010. Telomeres and telomerase in cancer. Carcinogenesis, 31 (1): 9-18. 10.1093/carcin/bgp268 19887512
10. Azouz A. Wu Y.L. Hillion J. Tarkanyi I. Karniguian A. Aradi J. et al. 2010. Epigenetic plasticity of hTERT gene promoter determines retinoid capacity to repress telomerase in maturation-resistant acute promyelocytic leukemia cells. Leukemia, 24 (3): 613-622. 10.1038/leu.2009.283 20072159
11. Ballal R.D. Saha T. Fan S. Haddad B.R. Rosen E.M. 2009. BRCA1 localization to the telomere and its loss from the telomere in response to DNA damage. J. Biol. Chem. 284 (52): 36083-36098. 10.1074/jbc.M109.025825 19797051
12. Bayne S. Liu J.P. 2005. Hormones and growth factors regulate telomerase activity in ageing and cancer. Mol. Cell. Endocrinol. 240 (1-2): 11-22. 10.1016/j.mce.2005.05.009 16005142 (Pubitemid 41133367)
13. Beato M. Sánchez-Pacheco A. 1996. Interaction of steroid hormone receptors with the transcription initiation complex. Endocr. Rev. 17 (6): 587-609. 8969970 (Pubitemid 26425258)
14. Bechter O.E. Eisterer W. Dlaska M. Kühr T. Thaler J. 2002. CpG island methylation of the hTERT promoter is associated with lower telomerase activity in B-cell lymphocytic leukemia. Exp. Hematol. 30 (1): 26-33. 10.1016/S0301-472X(01)00760-3 11823034 (Pubitemid 34059214)
15. Beitzinger M. Oswald C. Beinoraviciute-Kellner R. Stiewe T. 2006. Regulation of telomerase activity by the p53 family member p73. Oncogene, 25 (6): 813-826. 10.1038/sj.onc.1209125 16205639 (Pubitemid 43237700)
16. Bellon M. Nicot C. 2008. Regulation of telomerase and telomeres: human tumor viruses take control. J. Natl. Cancer Inst. 100 (2): 98-108. 18182620 (Pubitemid 351480562)
17. Beyne-Rauzy O. Prade-Houdellier N. Demur C. Recher C. Ayel J. Laurent G. Mansat-De Mas V. 2005. Tumor necrosis factor-α inhibits hTERT gene expression in human myeloid normal and leukemic cells. Blood, 106 (9): 3200-3205. 10.1182/blood-2005-04-1386 16020509 (Pubitemid 41565919)
18. Bilsland A.E. Stevenson K. Atkinson S. Kolch W. Keith W.N. 2006. Transcriptional repression of telomerase RNA gene expression by c-Jun-NH2-kinase and Sp1/Sp3. Cancer Res. 66 (3): 1363-1370. 10.1158/0008-5472.CAN-05-1941 16452190 (Pubitemid 43259915)
19. Block T.M. Mehta A.S. Fimmel C.J. Jordan R. 2003. Molecular viral oncology of hepatocellular carcinoma. Oncogene, 22 (33): 5093-5107. 10.1038/sj.onc.1206557 12910247 (Pubitemid 37059591)
20. Boggess J.F. Zhou C. Bae-Jump V.L. Gehrig P.A. Whang Y.E. 2006. Estrogen-receptor-dependent regulation of telomerase activity in human endometrial cancer cell lines. Gynecol. Oncol. 103 (2): 417-424. 10.1016/j.ygyno.2006.03.032 16690106 (Pubitemid 44602145)
21. Bougel S. Renaud S. Braunschweig R. Loukinov D. Morse III.H.C. Bosman F.T. et al. 2010. PAX5 activates the transcription of the human telomerase reverse transcriptase gene in B cells. J. Pathol. 220 (1): 87-96. 10.1002/path.2620 19806612
22. Brandl A. Angele P. Roll C. Prantl L. Kujat R. Kinner B. 2010. Influence of the growth factors PDGF-BB, TGF-beta1 and bFGF on the replicative aging of human articular chondrocytes during in vitro expansion. J. Orthop. Res. 28 (3): 354-360. 19798738
23. Bu D.X. Johansson M.E. Ren J. Xu D.W. Johnson F.B. Edfeldt K. Yan Z.Q. 2010. Nuclear factor κB-mediated transactivation of telomerase prevents intimal smooth muscle cell from replicative senescence during vascular repair. Arterioscler. Thromb. Vasc. Biol. 30 (12): 2604-2610. 10.1161/ATVBAHA.110.213074 20864668
24. Buchkovich K.J. Greider C.W. 1996. Telomerase regulation during entry into the cell cycle in normal human T cells. Mol. Biol. Cell, 7 (9): 1443-1454. 8885238 (Pubitemid 26304205)
25. Calado R.T. Yewdell W.T. Wilkerson K.L. Regal J.A. Kajigaya S. Stratakis C.A. Young N.S. 2009. Sex hormones, acting on the TERT gene, increase telomerase activity in human primary hematopoietic cells. Blood, 114 (11): 2236-2243. 10.1182/blood-2008-09-178871 19561322
26. Cao Y. Li H. Mu F.T. Ebisui O. Funder J.W. Liu J.P. 2002. Telomerase activation causes vascular smooth muscle cell proliferation in genetic hypertension. FASEB J. 16 (1): 96-98. 11772940
27. Casillas M.A. Brotherton S.L. Andrews L.G. Ruppert J.M. Tollefsbol T.O. 2003. Induction of endogenous telomerase (hTERT) by c-Myc in WI-38 fibroblasts transformed with specific genetic elements. Gene, 316: 57-65. 10.1016/S0378-1119(03)00739-X 14563552 (Pubitemid 37249353)
28. Cerone M.A. Londoño-Vallejo J.A. Autexier C. 2006. Mutated telomeres sensitize tumor cells to anticancer drugs independently of telomere shortening and mechanisms of telomere maintenance. Oncogene, 25 (56): 7411-7420. 10.1038/sj.onc.1209727 16767163 (Pubitemid 44853432)
29. Cesare A.J. Reddel R.R. 2010. Alternative lengthening of telomeres: models, mechanisms and implications. Nat. Rev. Genet. 11 (5): 319-330. 10.1038/nrg2763 20351727
30. Cha Y. Kwon S.J. Seol W. Park K.S. 2008. Estrogen receptor-alpha mediates the effects of estradiol on telomerase activity in human mesenchymal stem cells. Mol. Cells, 26 (5): 454-458. 18719354
31. Chan S.S. Chang S. 2010. Defending the end zone: studying the players involved in protecting chromosome ends. FEBS Lett. 584 (17): 3773-3778. 10.1016/j.febslet.2010.06.016 20579983
32. Chang W. Dynek J.N. Smith S. 2003. TRF1 is degraded by ubiquitin mediated proteolysis after release from telomeres. Genes Dev. 17 (11): 1328-1333. 10.1101/gad.1077103 12782650 (Pubitemid 36666001)
33. Chang J.T. Yang H.T. Wang T.C. Cheng A.J. 2005. Upstream stimulatory factor (USF) as a transcriptional suppressor of human telomerase reverse transcriptase (hTERT) in oral cancer cells. Mol. Carcinog. 44 (3): 183-192. 10.1002/mc.20129 16010690 (Pubitemid 41552751)
34. Chebel A. Rouault J.P. Urbanowicz I. Baseggio L. Chien W.W. Salles G. Ffrench M. 2009. Transcriptional activation of hTERT, the human telomerase reverse transcriptase, by nuclear factor of activated T cells. J. Biol. Chem. 284 (51): 35725-35734. 10.1074/jbc.M109.009183 19843528
35. Choi J.H. Min N.Y. Park J. Kim J.H. Park S.H. Ko Y.J. et al. 2010. TSA-induced DNMT1 down-regulation represses hTERT expression via recruiting CTCF into demethylated core promoter region of hTERT in HCT116. Biochem. Biophys. Res. Commun. 391 (1): 449-454. 10.1016/j.bbrc.2009.11.078 19914212
36. Choubey D. Deka R. Ho S.M. 2008. Interferon-inducible IFI16 protein in human cancers and autoimmune diseases. Front. Biosci. 13 (13): 598-608. 10.2741/2705 17981573 (Pubitemid 351599770)
37. Clément G. Braunschweig R. Pasquier N. Bosman F.T. Benhattar J. 2006. Methylation of APC, TIMP3, and TERT: a new predictive marker to distinguish Barrett's oesophagus patients at risk for malignant transformation. J. Pathol. 208 (1): 100-107. 10.1002/path.1884 16278815 (Pubitemid 43025708)
38. Comolli L.R. Smirnov I. Xu L. Blackburn E.H. James T.L. 2002. A molecular switch underlies a human telomerase disease. Proc. Natl. Acad. Sci. U.S.A. 99 (26): 16998-17003. 10.1073/pnas.262663599 12482936 (Pubitemid 36034087)
39. Cong Y.S. Wen J. Bacchetti S. 1999. The human telomerase catalytic subunit hTERT: organization of the gene and characterization of the promoter. Hum. Mol. Genet. 8 (1): 137-142. 10.1093/hmg/8.1.137 9887342 (Pubitemid 29039059)
40. Cong Y.S. Wright W.E. Shay J.W. 2002. Human telomerase and its regulation. Microbiol. Mol. Biol. Rev. 66 (3): 407-425. 10.1128/MMBR.66.3.407- 425.2002 12208997 (Pubitemid 35001137)
41. Cook B.D. Dynek J.N. Chang W. Shostak G. Smith S. 2002. Role for the related poly(ADP-Ribose) polymerases tankyrase 1 and 2 at human telomeres. Mol. Cell. Biol. 22 (1): 332-342. 10.1128/MCB.22.1.332-342.2002 11739745 (Pubitemid 33144279)
42. de Wilde J. Kooter J.M. Overmeer R.M. Claassen-Kramer D. Meijer C.J. Snijders P.J. Steenbergen R.D. 2010. hTERT promoter activity and CpG methylation in HPV-induced carcinogenesis. BMC Cancer, 10 (1): 271. 10.1186/1471-2407-10- 271 20534141
43. Dessain S.K. Yu H. Reddel R.R. Beijersbergen R.L. Weinberg R.A. 2000. Methylation of the human telomerase gene CpG island. Cancer Res. 60 (3): 537-541. 10676632 (Pubitemid 30094546)
44. Devereux T.R. Horikawa I. Anna C.H. Annab L.A. Afshari C.A. Barrett J.C. 1999. DNA methylation analysis of the promoter region of the human telomerase reverse transcriptase (hTERT) gene. Cancer Res. 59 (24): 6087-6090. 10626795 (Pubitemid 30035625)
45. Dez C. Henras A. Faucon B. Lafontaine D. Caizergues-Ferrer M. Henry Y. 2001. Stable expression in yeast of the mature form of human telomerase RNA depends on its association with the box H/ACA small nucleolar RNP proteins Cbf5p, Nhp2p and Nop10p. Nucleic Acids Res. 29 (3): 598-603. 10.1093/nar/29.3.598 11160879 (Pubitemid 32124368)
46. Dong X. Liu A. Zer C. Feng J. Zhen Z. Yang M. Zhong L. 2009. siRNA inhibition of telomerase enhances the anti-cancer effect of doxorubicin in breast cancer cells. BMC Cancer, 9 (1): 133. 10.1186/1471-2407-9-133 19416503
47. Doshida M. Ohmichi M. Tsutsumi S. Kawagoe J. Takahashi T. Du B. et al. 2006. Raloxifene increases proliferation and up-regulates telomerase activity in human umbilical vein endothelial cells. J. Biol. Chem. 281 (34): 24270-24278. 10.1074/jbc.M513251200 16798746 (Pubitemid 44274200)
48. Ducrest A.L. Szutorisz H. Lingner J. Nabholz M. 2002. Regulation of the human telomerase reverse transcriptase gene. Oncogene, 21 (4): 541-552. 10.1038/sj.onc.1205081 11850779 (Pubitemid 34146254)
49. Dwyer J.M. Liu J.P. 2010. Ets2 transcription factor, telomerase activity and breast cancer. Clin. Exp. Pharmacol. Physiol. 37 (1): 83-87. 10.1111/j.1440-1681.2009.05236.x 19566835
50. Dwyer J. Li H. Xu D. Liu J.P. 2007. Transcriptional regulation of telomerase activity: roles of the the Ets transcription factor family. Ann. N.Y. Acad. Sci. 1114 (1): 36-47. 10.1196/annals.1396.022 17986575 (Pubitemid 350070389)
51. Flores I. Blasco M.A. 2010. The role of telomeres and telomerase in stem cell aging. FEBS Lett. 584 (17): 3826-3830. 10.1016/j.febslet.2010.07.042 20674573
52. Fogarty P.F. Yamaguchi H. Wiestner A. Baerlocher G.M. Sloand E. Zeng W.S. et al. 2003. Late presentation of dyskeratosis congenita as apparently acquired aplastic anaemia due to mutations in telomerase RNA. Lancet, 362 (9396): 1628-1630. 10.1016/S0140-6736(03)14797-6 14630445 (Pubitemid 37443532)
53. Ford L.P. Suh J.M. Wright W.E. Shay J.W. 2000. Heterogeneous nuclear ribonucleoproteins C1 and C2 associate with the RNA component of human telomerase. Mol. Cell. Biol. 20 (23): 9084-9091. 10.1128/MCB.20.23.9084-9091. 2000 11074006 (Pubitemid 32245937)
54. Ford L.P. Shay J.W. Wright W.E. 2001. The La antigen associates with the human telomerase ribonucleoprotein and influences telomere length in vivo. RNA, 7 (8): 1068-1075. 10.1017/S1355838201010159 11497426 (Pubitemid 32726906)
55. Fujiki T. Udono M. Kadooka K. Yamashita S. Miura T. Shirahata S. Katakura Y. 2010. Regulatory mechanisms of human and mouse telomerase reverse transcriptase gene transcription: distinct dependency on c-Myc. Cytotechnology, 62 (4): 333-339. 10.1007/s10616-010-9276-y 20454928
56. Fujimoto K. Kyo S. Takakura M. Kanaya T. Kitagawa Y. Itoh H. et al. 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 (13): 2557-2562. 10.1093/nar/28.13.2557 10871406 (Pubitemid 30420854)
57. Furuya M. Tsuji N. Kobayashi D. Watanabe N. 2009. Interaction between survivin and aurora-B kinase plays an important role in survivin-mediated up-regulation of human telomerase reverse transcriptase expression. Int. J. Oncol. 34 (4): 1061-1068. 19287963
58. García-Cao M. O'Sullivan R. Peters A.H. Jenuwein T. Blasco M.A. 2004. Epigenetic regulation of telomere length in mammalian cells by the Suv39h1 and Suv39h2 histone methyltransferases. Nat. Genet. 36 (1): 94-99. 10.1038/ng1278 14702045 (Pubitemid 38056214)
59. Gaudray G. Gachon F. Basbous J. Biard-Piechaczyk M. Devaux C. Mesnard J.M. 2002. The complementary strand of the human T-cell leukemia virus type 1 RNA genome encodes a bZIP transcription factor that down-regulates viral transcription. J. Virol. 76 (24): 12813-12822. 10.1128/JVI.76.24.12813-12822. 2002 12438606 (Pubitemid 35386965)
60. Ge Z. Li W. Wang N. Liu C. Zhu Q. Björkholm M. et al. 2010. Chromatin remodeling: recruitment of histone demethylase RBP2 by Mad1 for transcriptional repression of a Myc target gene, telomerase reverse transcriptase. FASEB J. 24 (2): 579-586. 10.1096/fj.09-140087 19762557
61. George J. Banik N.L. Ray S.K. 2009. Combination of hTERT knockdown and IFN-gamma treatment inhibited angiogenesis and tumor progression in glioblastoma. Clin. Cancer Res. 15 (23): 7186-7195. 10.1158/1078-0432.CCR-09- 1425 19934306
62. Gewin L. Myers H. Kiyono T. Galloway D.A. 2004. Identification of a novel telomerase repressor that interacts with the human papillomavirus type-16 E6/E6-AP complex. Genes Dev. 18 (18): 2269-2282. 10.1101/gad.1214704 15371341 (Pubitemid 39209575)
63. Gigek C.O. Leal M.F. Silva P.N. Lisboa L.C. Lima E.M. Calcagno D.Q. et al. 2009. hTERT methylation and expression in gastric cancer. Biomarkers, 14 (8): 630-636. 10.3109/13547500903225912 20001710
64. INK4a pathway inhibits vascular smooth muscle cell proliferation by repressing cell cycle-dependent telomerase activation. Circ. Res. 103 (10): 1155-1163. 10.1161/CIRCRESAHA.108.186205 18818403
65. Gizard F. Heywood E.B. Findeisen H.M. Zhao Y. Jones K.L. Cudejko C. et al. 2011. Telomerase activation in atherosclerosis and induction of telomerase reverse transcriptase expression by inflammatory stimuli in macrophages. Arterioscler. Thromb. Vasc. Biol. 31 (2): 245-252. 10.1161/ATVBAHA.110.219808 21106948
66. Goldman F. Bouarich R. Kulkarni S. Freeman S. Du H.Y. Harrington L. et al. 2005. The effect of TERC haploinsufficiency on the inheritance of telomere length. Proc. Natl. Acad. Sci. U.S.A. 102 (47): 17119-17124. 10.1073/pnas.0505318102 16284252 (Pubitemid 41692693)
67. Goueli B.S. Janknecht R. 2003. Regulation of telomerase reverse transcriptase gene activity by upstream stimulatory factor. Oncogene, 22 (39): 8042-8047. 10.1038/sj.onc.1206847 12970752 (Pubitemid 37175868)
68. Goueli B.S. Janknecht R. 2004. Upregulation of the catalytic telomerase subunit by the transcription factor ER81 and oncogenic HER2/Neu, Ras, or Raf. Mol. Cell. Biol. 24 (1): 25-35. 10.1128/MCB.24.1.25-35.2004 14673140 (Pubitemid 38010032)
69. Grasselli A. Nanni S. Colussi C. Aiello A. Benvenuti V. Ragone G. et al. 2008. Estrogen receptor-α and endothelial nitric oxide synthase nuclear complex regulates transcription of human telomerase. Circ. Res. 103 (1): 34-42. 10.1161/CIRCRESAHA.107.169037 18519947
70. Guilleret I. Benhattar J. 2004. Unusual distribution of DNA methylation within the hTERT CpG island in tissues and cell lines. Biochem. Biophys. Res. Commun. 325 (3): 1037-1043. 10.1016/j.bbrc.2004.10.137 15541393 (Pubitemid 39487975)
71. Günes C. Lichtsteiner S. Vasserot A.P. Englert C. 2000. Expression of the hTERT gene is regulated at the level of transcriptional initiation and repressed by Mad1. Cancer Res. 60 (8): 2116-2121. 10786671 (Pubitemid 30225170)
72. Hao H. Nancai Y. Lei F. Xiong W. Wen S. Guofu H. et al. 2008. siRNA directed against c-Myc inhibits proliferation and downregulates human telomerase reverse transcriptase in human colon cancer Colo 320 cells. J. Exp. Clin. Cancer Res. 27 (1): 27. 10.1186/1756-9966-27-27 18694522
73. Hashimoto M. Kyo S. Hua X. Tahara H. Nakajima M. Takakura M. et al. 2008. Role of menin in the regulation of telomerase activity in normal and cancer cells. Int. J. Oncol. 33 (2): 333-340. 18636154
74. Hayflick L. Moorhead P.S. 1961. The serial cultivation of human diploid cell strains. Exp. Cell Res. 25 (3): 585-621. 10.1016/0014-4827(61)90192-6
75. Henson J.D. Reddel R.R. 2010. Assaying and investigating Alternative Lengthening of Telomeres activity in human cells and cancers. FEBS Lett. 584 (17): 3800-3811. 10.1016/j.febslet.2010.06.009 20542034
76. Hiyama E. Hiyama K. 2002. Clinical utility of telomerase in cancer. Oncogene, 21 (4): 643-649. 10.1038/sj.onc.1205070 11850791 (Pubitemid 34146266)
77. Horikawa I. Barrett C.J. 2003. Transcriptional regulation of the telomerase hTERT gene as a target for cellular and viral oncogenic mechanisms. Carcinogenesis, 24 (7): 1167-1176. 10.1093/carcin/bgg085 12807729 (Pubitemid 36927056)
78. Horikawa I. Cable P.L. Afshari C. Barrett J.C. 1999. Cloning and characterization of the promoter region of human telomerase reverse transcriptase gene. Cancer Res. 59 (4): 826-830. 10029071 (Pubitemid 29086735)
79. Horikawa I. Cable P.L. Mazur S.J. Appella E. Afshari C.A. 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 (8): 2585-2597. 10.1091/mbc.E01-11-0107 12181331
80. Jagadeesh S. Banerjee P.P. 2006. Inositol hexaphosphate represses telomerase activity and translocates TERT from the nucleus in mouse and human prostate cancer cells via the deactivation of Akt and PKCα. Biochem. Biophys. Res. Commun. 349 (4): 1361-1367. 10.1016/j.bbrc.2006.09.002 16979586 (Pubitemid 44416407)
81. Ji X.M. Xie C.H. Fang M.H. Zhou F.X. Zhang W.J. Zhang M.S. Zhou Y.F. 2006. Efficient inhibition of human telomerase activity by antisense oligonucleotides sensitizes cancer cells to radiotherapy. Acta Pharmacol. Sin. 27 (9): 1185-1191. 10.1111/j.1745-7254.2006.00417.x 16923339 (Pubitemid 44410999)
82. Jiang S. Galindo M.R. Jarrett H.W. 2010. Purification and identification of a transcription factor, USF-2, binding to E-box element in the promoter of human telomerase reverse transcriptase (hTERT). Proteomics, 10 (2): 203-211. 10.1002/pmic.200800693 19899074
83. Jin X. Beck S. Sohn Y.W. Kim J.K. Kim S.H. Yin J. et al. 2010. Human telomerase catalytic subunit (hTERT) suppresses p53-mediated anti-apoptotic response via induction of basic fibroblast growth factor. Exp. Mol. Med. 42 (8): 574-582. 10.3858/emm.2010.42.8.058 20628269
84. Jones P.A. Laird P.W. 1999. Cancer epigenetics comes of age. Nat. Genet. 21 (2): 163-167. 10.1038/5947 9988266 (Pubitemid 29070360)
85. Jost C.A. Marin M.C. Kaelin W.G., Jr. 1997. p73 is a human p53-related protein that can induce apoptosis. Nature, 389 (6647): 191-194. 10.1038/38298 9296498 (Pubitemid 27400602)
86. Kaghad M. Bonnet H. Yang A. Creancier L. Biscan J.C. Valent A. et al. 1997. Monoallelically expressed gene related to p53 at 1p36, a region frequently deleted in neuroblastoma and other human cancers. Cell, 90 (4): 809-819. 10.1016/S0092-8674(00)80540-1 9288759 (Pubitemid 27357971)
87. Kanaya T. Kyo S. Hamada K. Takakura M. Kitagawa Y. Harada H. Inoue M. 2000. Adenoviral expression of p53 represses telomerase activity through down-regulation of human telomerase reverse transcriptase transcription. Clin. Cancer Res. 6 (4): 1239-1247. 10778946 (Pubitemid 30226203)
88. Kawagoe J. Ohmichi M. Takahashi T. Ohshima C. Mabuchi S. Takahashi K. et al. 2003. Raloxifene inhibits estrogen-induced up-regulation of telomerase activity in a human breast cancer cell line. J. Biol. Chem. 278 (44): 43363-43372. 10.1074/jbc.M304363200 12917431 (Pubitemid 37345958)
89. Kim N.W. Piatyszek M.A. Prowse K.R. Harley C.B. West M.D. Ho P.L. et al. 1994. Specific association of human telomerase activity with immortal cells and cancer. Science, 266 (5193): 2011-2015. 10.1126/science.7605428 7605428
90. Kim M.O. Moon D.O. Kang S.H. Heo M.S. Choi Y.H. Jung J.H. et al. 2008. Pectenotoxin-2 represses telomerase activity in human leukemia cells through suppression of hTERT gene expression and Akt-dependent hTERT phosphorylation. FEBS Lett. 582 (23-24): 3263-3269. 10.1016/j.febslet.2008.08.030 18775701
91. Klenova E.M. Nicolas R.H. Paterson H.F. Carne A.F. Heath C.M. Goodwin G.H. et al. 1993. 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. 13 (12): 7612-7624. 8246978 (Pubitemid 23354286)
92. Knight J.S. Cotter M.A. II 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 (25): 22971-22978. 10.1074/jbc.M101890200 11313352
93. Konnikova L. Simeone M.C. Kruger M.M. Kotecki M. Cochran B.H. 2005. Signal transducer and activator of transcription 3 (STAT3) regulates human telomerase reverse transcriptase (hTERT) expression in human cancer and primary cells. Cancer Res. 65 (15): 6516-6520. 10.1158/0008-5472.CAN-05-0924 16061629 (Pubitemid 41060685)
94. Koshiji M. Kageyama Y. Pete E.A. Horikawa I. Barrett J.C. Huang L.E. 2004. HIF-1α induces cell cycle arrest by functionally counteracting Myc. EMBO J. 23 (9): 1949-1956. 10.1038/sj.emboj.7600196 15071503 (Pubitemid 38902073)
95. Kumari A. Srinivasan R. Vasishta R.K. Wig J.D. 2009. Positive regulation of human telomerase reverse transcriptase gene expression and telomerase activity by DNA methylation in pancreatic cancer. Ann. Surg. Oncol. 16 (4): 1051-1059. 10.1245/s10434-009-0333-8 19194757
96. Kunimura C. Kikuchi K. Ahmed N. Shimizu A. Yasumoto S. 1998. Telomerase activity in a specific cell subset co-expressing integrinbeta1/EGFR but not p75NGFR/bcl2/integrin beta4 in normal human epithelial cells. Oncogene, 17 (2): 187-197. 10.1038/sj.onc.1201916 9674703 (Pubitemid 28371892)
97. Kuranaga N. Shinomiya N. Mochizuki H. 2001. Long-term cultivation of colorectal carcinoma cells with anti-cancer drugs induces drug resistance and telomere elongation: an in vitro study. BMC Cancer, 1 (1): 10. 10.1186/1471-2407-1-10 11518543 (Pubitemid 38846509)
98. Kyo S. Takakura M. Kanaya T. Zhuo W. Fujimoto K. Nishio Y. et al. 1999. Estrogen activates telomerase. Cancer Res. 59 (23): 5917-5921. 10606235 (Pubitemid 30004951)
99. Kyo S. Takakura M. Fujiwara T. Inoue M. 2008. Understanding and exploiting hTERT promoter regulation for diagnosis and treatment of human cancers. Cancer Sci. 99 (8): 1528-1538. 10.1111/j.1349-7006.2008.00878.x 18754863
100. Lafferty-Whyte K. Bilsland A. Hoare S.F. Burns S. Zaffaroni N. Cairney C.J. Keith W.N. 2010. TCEAL7 inhibition of c-Myc activity in alternative lengthening of telomeres regulates hTERT expression. Neoplasia, 12 (5): 405-414. 20454512
101. Le S. Sternglanz R. Greider C.W. 2000. Identification of two RNA binding proteins associated with human telomerase RNA. Mol. Biol. Cell, 11 (3): 999-1010. 10712515 (Pubitemid 30159317)
102. Lee J.S. Igarashi T. Fraga S. Dahl E. Hovgaard P. Yasumoto T. 1989. Determination of diarrhetic shellfish toxins in various dinoflagellate species. J. Appl. Phycol. 1 (2): 147-152. 10.1007/BF00003877
103. Lee S.H. Kim J.W. Lee H.W. Cho Y.S. Oh S.H. Kim Y.J. et al. 2003. Interferon regulatory factor-1 (IRF-1) is a mediator for interferon-γ induced attenuation of telomerase activity and human telomerase reverse transcriptase (hTERT) expression. Oncogene, 22 (3): 381-391. 10.1038/sj.onc.1206133 12545159 (Pubitemid 36194334)
104. Lee S.H. Kim J.W. Oh S.H. Kim Y.J. Rho S.B. Park K. et al. 2005. IFN-γ/IRF-1-induced p27kip1 down-regulates telomerase activity and human telomerase reverse transcriptase expression in human cervical cancer. FEBS Lett. 579 (5): 1027-1033. 10.1016/j.febslet.2005.01.005 15710386 (Pubitemid 40248664)
105. Li H. Lee T.H. 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 (23): 20965-20973. 10.1074/jbc.M112231200 11916966 (Pubitemid 34967407)
106. Li Y. Liu L. Andrews L.G. Tollefsbol T.O. 2009. Genistein depletes telomerase activity through cross-talk between genetic and epigenetic mechanisms. Int. J. Cancer, 125 (2): 286-296. 10.1002/ijc.24398 19358274
107. Li W. Zeng J. Li Q. Zhao L. Liu T. Björkholm M. et al. 2010. Reptin is required for the transcription of telomerase reverse transcriptase and over-expressed in gastric cancer. Mol. Cancer, 9 (1): 132. 10.1186/1476-4598-9- 132 20509972
108. Lin S.Y. Elledge S.J. 2003. Multiple tumor suppressor pathways negatively regulate telomerase. Cell, 113 (7): 881-889. 10.1016/S0092-8674(03)00430-6 12837246 (Pubitemid 36798050)
109. Lindkvist A. Franzén A. Ren Z.P. Heldin N.E. Paulsson-Karlsson Y. 2005. Differential effects of TGF-β1 on telomerase activity in thyroid carcinoma cell lines. Biochem. Biophys. Res. Commun. 338 (3): 1625-1633. 10.1016/j.bbrc.2005.10.131 16288728 (Pubitemid 41608436)
110. Liu H. Shi W. Luan F. Xu S. Yang F. Sun W. et al. 2010. Hepatitis B virus X protein upregulates transcriptional activation of human telomerase reverse transcriptase. Virus Genes, 40 (2): 174-182. 10.1007/s11262-009-0441-3 20107884
111. Lou F. Chen X. Jalink M. Zhu Q. Ge N. Zhao S. et al. 2007. The opposing effect of hypoxia-inducible factor-2α on expression of telomerase reverse transcriptase. Mol. Cancer Res. 5 (8): 793-800. 10.1158/1541-7786.MCR-07-0065 17699105 (Pubitemid 47294807)
112. Maida Y. Kyo S. Kanaya T. Wang Z. Yatabe N. Tanaka M. et al. 2002. Direct activation of telomerase by EGF through Ets-mediated transactivation of TERT via MAP kinase signaling pathway. Oncogene, 21 (26): 4071-4079. 10.1038/sj.onc.1205509 12037663 (Pubitemid 34712874)
113. Martínez-Campa C.M. Alonso-González C. Mediavilla M.D. Cos S. González A. Sanchez-Barcelo E.J. 2008. Melatonin down-regulates hTERT expression induced by either natural estrogens (17β-estradiol) or metalloestrogens (cadmium) in MCF-7 human breast cancer cells. Cancer Lett. 268 (2): 272-277. 10.1016/j.canlet.2008.04.001 18479810
114. Massard C. Zermati Y. Pauleau A.L. Larochette N. Métivier D. Sabatier L. et al. 2006. hTERT: a novel endogenous inhibitor of the mitochondrial cell death pathway. Oncogene, 25 (33): 4505-4514. 10.1038/sj.onc.1209487 16619047 (Pubitemid 44175110)
115. Matsubara Y. Murata M. Watanabe K. Saito I. Miyaki K. Omae K. et al. 2006. Coronary artery disease and a functional polymorphism of hTERT. Biochem. Biophys. Res. Commun. 348 (2): 669-672. 10.1016/j.bbrc.2006.07.103 16890917 (Pubitemid 44188616)
116. McMurray H.R. McCance D.J. 2003. Human papillomavirus type 16 E6 activates TERT gene transcription through induction of c-Myc and release of USF-mediated repression. J. Virol. 77 (18): 9852-9861. 10.1128/JVI.77.18.9852- 9861.2003 12941894 (Pubitemid 37122066)
117. Meeran S.M. Ahmed A. Tollefsbol T.O. 2010 a. Epigenetic targets of bioactive dietary components for cancer prevention and therapy. Clin. Epigenetics, 1 (3-4): 101-116. 10.1007/s13148-010-0011-5 21258631
118. Meeran S.M. Patel S.N. Tollefsbol T.O. 2010 b. Sulforaphane causes epigenetic repression of hTERT expression in human breast cancer cell lines. PLoS ONE, 5 (7): e11457. 10.1371/journal.pone.0011457 20625516
119. Meyerson M. Counter C.M. Eaton E.N. Ellisen L.W. Steiner P. Caddle S.D. et al. 1997. hEST2, the putative human telomerase catalytic subunit gene, is up-regulated in tumor cells and during immortalization. Cell, 90 (4): 785-795. 10.1016/S0092-8674(00)80538-3 9288757 (Pubitemid 27357969)
120. Minamino T. Komuro I. 2003. The role of telomerase activation in the regulation of vascular smooth muscle cell proliferation. Drug News Perspect. 16 (4): 211-216. 10.1358/dnp.2003.16.4.829332 12942150 (Pubitemid 37039402)
121. Minamino T. Mitsialis S.A. Kourembanas S. 2001. Hypoxia extends the life span of vascular smooth muscle cells through telomerase activation. Mol. Cell. Biol. 21 (10): 3336-3342. 10.1128/MCB.21.10.3336-3342.2001 11313459 (Pubitemid 32381769)
122. Misiti S. Nanni S. Fontemaggi G. Cong Y.S. Wen J. Hirte H.W. et al. 2000. Induction of hTERT expression and telomerase activity by estrogens in human ovary epithelium cells. Mol. Cell. Biol. 20 (11): 3764-3771. 10.1128/MCB.20.11.3764-3771.2000 10805720 (Pubitemid 30314476)
123. Mitchell J.R. Cheng J. Collins K. 1999 a. A box H/ACA small nucleolar RNA-like domain at the human telomerase RNA 3-2 end. Mol. Cell. Biol. 19 (1): 567-576. 9858580 (Pubitemid 29018458)
124. Mitchell J.R. Wood E. Collins K. 1999 b. A telomerase component is defective in the human disease dyskeratosis congenita. Nature, 402 (6761): 551-555. 10.1038/990141 10591218
125. Moon D.O. Kim M.O. Heo M.S. Lee J.D. Choi Y.H. Kim G.Y. 2009 a. Gefitinib induces apoptosis and decreases telomerase activity in MDA-MB-231 human breast cancer cells. Arch. Pharm. Res. 32 (10): 1351-1360. 10.1007/s12272-009-2002-7 19898796
126. Moon D.O. Kim M.O. Lee J.D. Choi Y.H. Kim G.Y. 2009 b. Butein suppresses c-Myc-dependent transcription and Akt-dependent phosphorylation of hTERT in human leukemia cells. Cancer Lett. 286 (2): 172-179. 10.1016/j.canlet.2009.05. 028 19560862
127. Murillo-Ortiz B. Astudillo-De la Vega H. Castillo-Medina S. Malacara J.M. Benitez-Bribiesca L. 2006. Telomerase activity, estrogen receptors (α, β), Bcl-2 expression in human breast cancer and treatment response. BMC Cancer, 6 (1): 206. 10.1186/1471-2407-6-206 16911782 (Pubitemid 44382760)
128. Murnane J.P. 2010. Telomere loss as a mechanism for chromosome instability in human cancer. Cancer Res. 70 (11): 4255-4259. 10.1158/0008-5472.CAN-09-4357 20484032
129. Nakamura T.M. Morin G.B. Chapman K.B. Weinrich S.L. Andrews W.H. Lingner J. et al. 1997. Telomerase catalytic subunit homologs from fission yeast and human. Science, 277 (5328): 955-959. 10.1126/science.277.5328.955 9281073 (Pubitemid 27366681)
130. Nanni S. Narducci M. Della Pietra L. Moretti F. Grasselli A. De Carli P. et al. 2002. Signaling through estrogen receptors modulates telomerase activity in human prostate cancer. J. Clin. Invest. 110 (2): 219-227. 12122114 (Pubitemid 34787359)
131. Natarajan S. Chen Z. Wancewicz E.V. Monia B.P. Corey D.R. 2004. Telomerase reverse transcriptase (hTERT) mRNA and telomerase RNA (hTR) as targets for downregulation of telomerase activity. Oligonucleotides, 14 (4): 263-273. 10.1089/oli.2004.14.263 15665594 (Pubitemid 40144364)
132. Nishi H. Nakada T. Kyo S. Inoue M. Shay J.W. Isaka K. 2004. Hypoxia-inducible factor 1 mediates upregulation of telomerase (hTERT). Mol. Cell. Biol. 24 (13): 6076-6083. 10.1128/MCB.24.13.6076-6083.2004 15199161 (Pubitemid 38787990)
133. Nittis T. Guittat L. Stewart S.A. 2008. Alternative lengthening of telomeres (ALT) and chromatin: is there a connection? Biochimie, 90 (1): 5-12. 10.1016/j.biochi.2007.08.009 17935854 (Pubitemid 350216214)
134. Nomoto K. Maekawa M. Sugano K. Ushiama M. Fukayama N. Fujita S. Kakizoe T. 2002. Methylation status and expression of human telomerase reverse transcriptase mRNA in relation to hypermethylation of the p16 gene in colorectal cancers as analyzed by bisulfite PCR-SSCP. Jpn. J. Clin. Oncol. 32 (1): 3-8. 10.1093/jjco/hyf001 11932355 (Pubitemid 34201907)
135. Noreen F. Heinrich J. Moelling K. 2009. Antitumor activity of small double-stranded oligodeoxynucleotides targeting telomerase RNA in malignant melanoma cells. Oligonucleotides, 19 (2): 169-178. 10.1089/oli.2008.0170 19441892
136. Ogawa D. Nomiyama T. Nakamachi T. Heywood E.B. Stone J.F. Berger J.P. et al. 2006. Activation of peroxisome proliferator-activated receptor γ suppresses telomerase activity in vascular smooth muscle cells. Circ. Res. 98 (7): e50-e59. 10.1161/01.RES.0000218271.93076.c3 16556873
137. Oh S. Song Y. Yim J. Kim T.K. 1999. The Wilms tumor 1 tumor suppressor gene represses transcription of the human telomerase reverse transcriptase gene. J. Biol. Chem. 274 (52): 37473-37478. 10.1074/jbc.274.52.37473 10601322 (Pubitemid 30016836)
138. Oh S.T. Kyo S. 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 (12): 5559-5566. 10.1128/JVI.75.12.5559-5566.2001 11356963 (Pubitemid 32488149)
139. Patel R. Shervington A. 2009. Telomerase and DNA repair in glioma. Biochim. Biophys. Acta, 1792 (4): 275-279. 19232390
140. Phipps S.M. Love W.K. White T. Andrews L.G. Tollefsbol T.O. 2009. Retinoid-induced histone deacetylation inhibits telomerase activity in estrogen receptor-negative breast cancer cells. Anticancer Res. 29 (12): 4959-4964. 20044602
141. Pogacić V. Dragon F. Filipowicz W. 2000. Human H/ACA small nucleolar RNPs and telomerase share evolutionarily conserved proteins NHP2 and NOP10. Mol. Cell. Biol. 20 (23): 9028-9040. 10.1128/MCB.20.23.9028-9040.2000 11074001 (Pubitemid 32245932)
142. Poole J.C. Andrews L.G. Tollefsbol T.O. 2001. Activity, function, and gene regulation of the catalytic subunit of telomerase (hTERT). Gene, 269 (1-2): 1-12. 10.1016/S0378-1119(01)00440-1 11376932 (Pubitemid 32488901)
143. Poynter K.R. Sachs P.C. Bright A.T. Breed M.S. Nguyen B.N. Elmore L.W. Holt S.E. 2009. Genetic inhibition of telomerase results in sensitization and recovery of breast tumor cells. Mol. Cancer Ther. 8 (5): 1319-1327. 10.1158/1535-7163.MCT-08-0849 19417141
144. Qi D.L. Ohhira T. Oshimura M. Kugoh H. 2010. Human chromosome 5 carries a transcriptional regulator of human telomerase reverse transcriptase (hTERT). Biochem. Biophys. Res. Commun. 398 (4): 695-701. 10.1016/j.bbrc.2010.07.003 20621064
145. Racek T. Mise N. Li Z. Stoll A. Pützer B.M. 2005. C-terminal p73 isoforms repress transcriptional activity of the human telomerase reverse transcriptase (hTERT) promoter. J. Biol. Chem. 280 (49): 40402-40405. 10.1074/jbc.C500193200 16234237 (Pubitemid 41780526)
146. Rama S. Suresh Y. Rao A.J. 2003. TGF β1 induces multiple independent signals to regulate human trophoblastic differentiation: mechanistic insights. Mol. Cell. Endocrinol. 206 (1-2): 123-136. 10.1016/S0303-7207(03)00202-8 12943995 (Pubitemid 37011257)
147. Ren H. Zhao T. Wang X. Gao C. Wang J. Yu M. Hao J. 2010. Leptin upregulates telomerase activity and transcription of human telomerase reverse transcriptase in MCF-7 breast cancer cells. Biochem. Biophys. Res. Commun. 394 (1): 59-63. 10.1016/j.bbrc.2010.02.093 20171193
148. Renaud S. Loukinov D. Bosman F.T. Lobanenkov V. Benhattar J. 2005. CTCF binds the proximal exonic region of hTERT and inhibits its transcription. Nucleic Acids Res. 33 (21): 6850-6860. 10.1093/nar/gki989 16326864 (Pubitemid 41830489)
149. Renaud S. Loukinov D. Abdullaev Z. Guilleret I. Bosman F.T. Lobanenkov V. Benhattar J. 2007. Dual role of DNA methylation inside and outside of CTCF-binding regions in the transcriptional regulation of the telomerase hTERT gene. Nucleic Acids Res. 35 (4): 1245-1256. 10.1093/nar/gkl1125 17267411 (Pubitemid 46522976)
150. Royle N.J. Méndez-Bermúdez A. Gravani A. Novo C. Foxon J. Williams J. et al. 2009. The role of recombination in telomere length maintenance. Biochem. Soc. Trans. 37 (3): 589-595. 10.1042/BST0370589 19442255
151. Rubis B. Kaczmarek M. Szymanowska N. Galezowska E. Czyrski A. Juskowiak B. et al. 2009. The biological activity of G-quadruplex DNA binding papaverine-derived ligand in breast cancer cells. Invest. New Drugs, 27 (4): 289-296. 10.1007/s10637-008-9173-9 18773143
152. Sahin E. Depinho R.A. 2010. Linking functional decline of telomeres, mitochondria and stem cells during ageing. Nature, 464 (7288): 520-528. 10.1038/nature08982 20336134
153. Seimiya H. 2006. The telomeric PARP, tankyrases, as targets for cancer therapy. Br. J. Cancer, 94 (3): 341-345. 10.1038/sj.bjc.6602951 16421589 (Pubitemid 43237557)
154. Seimiya H. Tanji M. Oh-hara T. Tomida A. Naasani I. Tsuruo T. 1999. Hypoxia up-regulates telomerase activity via mitogen-activated protein kinase signaling in human solid tumor cells. Biochem. Biophys. Res. Commun. 260 (2): 365-370. 10.1006/bbrc.1999.0910 10403776 (Pubitemid 29352071)
155. waf1. J. Biol. Chem. 279 (49): 50976-50985. 10.1074/jbc.M402502200 15371422 (Pubitemid 40017838)
156. Shay J.W. Wright W.E. 2010. Telomeres and telomerase in normal and cancer stem cells. FEBS Lett. 584 (17): 3819-3825. 10.1016/j.febslet.2010.05.026 20493857
157. Shin K.H. Kang M.K. Dicterow E. Park N.H. 2003. Hypermethylation of the hTERT promoter inhibits the expression of telomerase activity in normal oral fibroblasts and senescent normal oral keratinocytes. Br. J. Cancer, 89 (8): 1473-1478. 10.1038/sj.bjc.6601291 14562019 (Pubitemid 37386602)
158. Sikand K. Kaul D. Varma N. 2006. Receptor Ck-dependent signaling regulates hTERT gene transcription. BMC Cell Biol. 7 (1): 2. 10.1186/1471-2121-7-2 16405739 (Pubitemid 43180955)
159. Siva A.C. Raval-Fernandes S. Stephen A.G. LaFemina M.J. Scheper R.J. Kickhoefer V.A. Rome L.H. 2001. Up-regulation of vaults may be necessary but not sufficient for multidrug resistance. Int. J. Cancer, 92 (2): 195-202. 10.1002/1097-0215(200102)9999:9999<::AID-IJC1168>3.0.CO;2-7 11291045 (Pubitemid 32225369)
160. Smith S. de Lange T. 1999. Cell cycle dependent localization of the telomeric PARP, tankyrase, to nuclear pore complexes and centrosomes. J. Cell Sci. 112 (Pt 21): 3649-3656. 10523501 (Pubitemid 129783932)
161. Smith S. Giriat I. Schmitt A. de Lange T. 1998. Tankyrase, a poly(ADP-ribose) polymerase at human telomeres. Science, 282 (5393): 1484-1487. 10.1126/science.282.5393.1484 9822378 (Pubitemid 28538609)
162. Smith V. Dai F. Spitz M. Peters G.J. Fiebig H.H. Hussain A. Burger A.M. 2009. Telomerase activity and telomere length in human tumor cells with acquired resistance to anticancer agents. J. Chemother. 21 (5): 542-549. 19933046
163. Song Z. Krishna S. Thanos D. Strominger J.L. Ono S.J. 1994. A novel cysteine-rich sequence-specific DNA-binding protein interacts with the conserved X-box motif of the human major histocompatibility complex class II genes via a repeated Cys-His domain and functions as a transcriptional repressor. J. Exp. Med. 180 (5): 1763-1774. 10.1084/jem.180.5.1763 7964459 (Pubitemid 24328567)
164. Song L.L. Ponomareva L. Shen H. Duan X. Alimirah F. Choubey D. 2010. Interferon-inducible IFI16, a negative regulator of cell growth, down-regulates expression of human telomerase reverse transcriptase (hTERT) gene. PLoS ONE, 5 (1): e8569. 10.1371/journal.pone.0008569 20052289
165. Stein G.S. Montecino M. van Wijnen A.J. Stein J.L. Lian J.B. 2000. Nuclear structure-gene expression interrelationships: implications for aberrant gene expression in cancer. Cancer Res. 60 (8): 2067-2076. 10786661 (Pubitemid 30225160)
166. Stiewe T. Zimmermann S. Frilling A. Esche H. Pützer B.M. 2002. Transactivation-deficient ΔTA-p73 acts as an oncogene. Cancer Res. 62 (13): 3598-3602. 12097259
167. Strååt K. Liu C. Rahbar A. Zhu Q. Liu L. Wolmer-Solberg N. et al. 2009. Activation of telomerase by human cytomegalovirus. J. Natl. Cancer Inst. 101 (7): 488-497. 10.1093/jnci/djp031 19318640
168. Suphapeetiporn K. Greally J.M. Walpita D. Ashley T. Bale A.E. 2002. MEN1 tumor-suppressor protein localizes to telomeres during meiosis. Genes Chromosomes Cancer, 35 (1): 81-85. 10.1002/gcc.10113 12203793 (Pubitemid 34847872)
169. Takakura M. Kyo S. Kanaya T. Hirano H. Takeda J. Yutsudo M. 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 (3): 551-557. 9973199 (Pubitemid 29076049)
170. Takakura M. Kyo S. Sowa Y. Wang Z. Yatabe N. Maida Y. et al. 2001. Telomerase activation by histone deacetylase inhibitor in normal cells. Nucleic Acids Res. 29 (14): 3006-3011. 10.1093/nar/29.14.3006 11452025 (Pubitemid 32685226)
171. Theimer C.A. Finger L.D. Feigon J. 2003. YNMG tetraloop formation by a dyskeratosis congenita mutation in human telomerase RNA. RNA, 9 (12): 1446-1455. 10.1261/rna.5152303 14624001 (Pubitemid 37463471)
172. Toaldo C. Pizzimenti S. Cerbone A. Pettazzoni P. Menegatti E. Daniela B. et al. 2010. PPARγ ligands inhibit telomerase activity and hTERT expression through modulation of the Myc/Mad/Max network, in colon cancer cells. J. Cell. Mol. Med. 14 (6a): 1347-1357. 10.1111/j.1582-4934.2009.00966.x 19912441
173. Toh W.H. Kyo S. Sabapathy K. 2005. Relief of p53-mediated telomerase suppression by p73. J. Biol. Chem. 280 (17): 17329-17338. 10.1074/jbc.M500044200 15734740 (Pubitemid 41389202)
174. Tsakiri K.D. Cronkhite J.T. Kuan P.J. Xing C. Raghu G. Weissler J.C. et al. 2007. Adult-onset pulmonary fibrosis caused by mutations in telomerase. Proc. Natl. Acad. Sci. U.S.A. 104 (18): 7552-7557. 10.1073/pnas.0701009104 17460043 (Pubitemid 47185944)
175. Uziel O. Fenig E. Nordenberg J. Beery E. Reshef H. Sandbank J. et al. 2005. Imatinib mesylate (Gleevec) downregulates telomerase activity and inhibits proliferation in telomerase-expressing cell lines. Br. J. Cancer, 92 (10): 1881-1891. 10.1038/sj.bjc.6602592 15870711 (Pubitemid 40826137)
176. Vageli D. Ioannou M.G. Koukoulis G.K. 2009. Transcriptional activation of hTERT in breast carcinomas by the Her2-ER81-related pathway. Oncol. Res. 17 (9): 413-423. 10.3727/096504009788912507 19718948
177. Venteicher A.S. Meng Z. Mason P.J. Veenstra T.D. Artandi S.E. 2008. Identification of ATPases pontin and reptin as telomerase components essential for holoenzyme assembly. Cell, 132 (6): 945-957. 10.1016/j.cell.2008.01.019 18358808 (Pubitemid 351381739)
178. Vulliamy T. Marrone A. Goldman F. Dearlove A. Bessler M. Mason P.J. Dokal I. 2001. The RNA component of telomerase is mutated in autosomal dominant dyskeratosis congenita. Nature, 413 (6854): 432-435. 10.1038/35096585 11574891 (Pubitemid 32928599)
179. Vulliamy T. Marrone A. Szydlo R. Walne A. Mason P.J. Dokal I. 2004. Disease anticipation is associated with progressive telomere shortening in families with dyskeratosis congenita due to mutations in TERC. Nat. Genet. 36 (5): 447-449. 10.1038/ng1346 15098033 (Pubitemid 38620026)
180. Wang J. Xie L.Y. Allan S. Beach D. Hannon G.J. 1998. a. Myc activates telomerase. Genes Dev. 12 (12): 1769-1774. 10.1101/gad.12.12.1769 9637678 (Pubitemid 28304329)
181. Wang Q. Zhang H. Kajino K. Greene M.I. 1998. b. BRCA1 binds c-Myc and inhibits its transcriptional and transforming activity in cells. Oncogene, 17 (15): 1939-1948. 10.1038/sj.onc.1202403 9788437 (Pubitemid 28488462)
182. Wang J. Liu X. Fang J. 1999. Expression and clinical significance of telomerase catalytic subunit gene in lung cancer and its correlations with genes related to drug resistance and apoptosis. Zhonghua Zhong Liu Za Zhi, 21 (5): 350-353. 11776572
183. Wang Z. Kyo S. Takakura M. Tanaka M. Yatabe N. Maida Y. et al. 2000. Progesterone regulates human telomerase reverse transcriptase gene expression via activation of mitogen-activated protein kinase signaling pathway. Cancer Res. 60 (19): 5376-5381. 11034074
184. Wick M. Zubov D. Hagen G. 1999. Genomic organisation and promoter characterization of the gene encoding the human telomerase reverse transcriptase (hTERT). Gene, 232 (1): 97-106. 10.1016/S0378-1119(99)00108-0 10333526 (Pubitemid 29305322)
185. Widschwendter A. Müller H.M. Hubalek M.M. Wiedemair A. Fiegl H. Goebel G. et al. 2004. Methylation status and expression of human telomerase reverse transcriptase in ovarian and cervical cancer. Gynecol. Oncol. 93 (2): 407-416. 10.1016/j.ygyno.2004.01.036 15099954 (Pubitemid 38507610)
186. Wojtyla A. Gladych M. Rubis B. 2011. Human telomerase activity regulation. Mol. Biol. Rep. 38 (5): 3339-3349. 10.1007/s11033-010-0439-x 21086176
187. Won J. Chang S. Oh S. Kim T.K. 2004. Small-molecule-based identification of dynamic assembly of E2F-pocket protein-histone deacetylase complex for telomerase regulation in human cells. Proc. Natl. Acad. Sci. U.S.A. 101 (31): 11328-11333. 10.1073/pnas.0401801101 15263087 (Pubitemid 39038346)
188. Wu K.J. Grandori C. Amacker M. Simon-Vermot N. Polack A. Lingner J. Dalla-Favera R. 1999. Direct activation of TERT transcription by c-MYC. Nat. Genet. 21 (2): 220-224. 10.1038/6010 9988278 (Pubitemid 29070372)
189. Xiao X. Athanasiou M. Sidorov I.A. Horikawa I. Cremona G. Blair D. et al. 2003. Role of Ets/Id proteins for telomerase regulation in human cancer cells. Exp. Mol. Pathol. 75 (3): 238-247. 10.1016/S0014-4800(03)00092-3 14611815 (Pubitemid 37373710)
190. Xin H. Curry J. Johnstone R.W. Nickoloff B.J. Choubey D. 2003. Role of IFI 16, a member of the interferon-inducible p200-protein family, in prostate epithelial cellular senescence. Oncogene, 22 (31): 4831-4840. 10.1038/sj.onc.1206754 12894224 (Pubitemid 36981271)
191. Xin H. Pereira-Smith O.M. Choubey D. 2004. Role of IFI 16 in cellular senescence of human fibroblasts. Oncogene, 23 (37): 6209-6217. 10.1038/sj.onc.1207836 15208661 (Pubitemid 39215291)
192. Xiong J. Fan S. Meng Q. Schramm L. Wang C. Bouzahza B. et al. 2003. BRCA1 inhibition of telomerase activity in cultured cells. Mol. Cell. Biol. 23 (23): 8668-8690. 10.1128/MCB.23.23.8668-8690.2003 14612409 (Pubitemid 37433370)
193. Xu D. Wang Q. Gruber A. Björkholm M. Chen Z. Zaid A. et al. 2000. Downregulation of telomerase reverse transcriptase mRNA expression by wild type p53 in human tumor cells. Oncogene, 19 (45): 5123-5133. 10.1038/sj.onc.1203890 11064449
194. Xu D. Popov N. Hou M. Wang Q. Björkholm M. Gruber A. et al. 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. U.S.A. 98 (7): 3826-3831. 10.1073/pnas.071043198 11274400 (Pubitemid 32249856)
195. Xu D. Dwyer J. Li H. Duan W. Liu J.P. 2008. a. Ets2 maintains hTERT gene expression and breast cancer cell proliferation by interacting with c-Myc. J. Biol. Chem. 283 (35): 23567-23580. 10.1074/jbc.M800790200 18586674
196. Xu M. Luo W. Elzi D.J. Grandori C. Galloway D.A. 2008. b. NFX1 interacts with mSin3A/histone deacetylase to repress hTERT transcription in keratinocytes. Mol. Cell. Biol. 28 (15): 4819-4828. 10.1128/MCB.01969-07 18505829
197. Yago M. Ohki R. Hatakeyama S. Fujita T. 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 (1-3): 40-46. 10.1016/S0014-5793(02)02757-6 12044867 (Pubitemid 34607270)
198. Yamaguchi H. Calado R.T. Ly H. Kajigaya S. Baerlocher G.M. Chanock S.J. et al. 2005. Mutations in TERT, the gene for telomerase reverse transcriptase, in aplastic anemia. N. Engl. J. Med. 352 (14): 1413-1424. 10.1056/NEJMoa042980 15814878 (Pubitemid 40471449)
199. Yang Y. Cai S. Yang L. Yu S. Jiang J. Yan X. et al. 2010. Targeting eradication of malignant cells derived from human bone marrow mesenchymal stromal cells. Exp. Cell Res. 316 (20): 3329-3341. 10.1016/j.yexcr.2010.02.014 20184882
200. Yatabe N. Kyo S. Maida Y. Nishi H. Nakamura M. Kanaya T. et al. 2004. HIF-1-mediated activation of telomerase in cervical cancer cells. Oncogene, 23 (20): 3708-3715. 10.1038/sj.onc.1207460 15048086 (Pubitemid 38658438)
201. Ye J.Z. de Lange T. 2004. TIN2 is a tankyrase 1 PARP modulator in the TRF1 telomere length control complex. Nat. Genet. 36 (6): 618-623. 10.1038/ng1360 15133513 (Pubitemid 38715990)
202. Zhang A. Zheng C. Hou M. Lindvall C. Li K.J. Erlandsson F. et al. 2003. Deletion of the telomerase reverse transcriptase gene and haploinsufficiency of telomere maintenance in Cri du chat syndrome. Am. J. Hum. Genet. 72 (4): 940-948. 10.1086/374565 12629597 (Pubitemid 36403312)
203. Zhou C. Liu J. 2003. Inhibition of human telomerase reverse transcriptase gene expression by BRCA1 in human ovarian cancer cells. Biochem. Biophys. Res. Commun. 303 (1): 130-136. 10.1016/S0006-291X(03)00318-8 12646176 (Pubitemid 36338217)
204. Zhou C. Bae-Jump V.L. Whang Y.E. Gehrig P.A. Boggess J.F. 2006. The PTEN tumor suppressor inhibits telomerase activity in endometrial cancer cells by decreasing hTERT mRNA levels. Gynecol. Oncol. 101 (2): 305-310. 10.1016/j.ygyno.2005.10.038 16380156
205. Zou L. Zhang P. Luo C. Tu Z. 2006. Mad1 suppresses bladder cancer cell proliferation by inhibiting human telomerase reverse transcriptase transcription and telomerase activity. Urology, 67 (6): 1335-1340. 10.1016/j.urology.2005.12. 029 16765199 (Pubitemid 43833462)