Common chromosomal fragile sites and cancer: Focus on FRA16D

Cancer Letters

Volumn 232, Issue 1, 2006, Pages 37-47

  O'Keefe, Louise V ^a   Richards, Robert I ^a  

^a UNIVERSITY OF ADELAIDE   (Australia)

Author keywords

Common chromosomal fragile site; In vitro fragility in vivo instability; Protective gene function

Indexed keywords

APHIDICOLIN; FRAGILE HISTIDINE TRIAD PROTEIN; TRANS ACTING FACTOR;

CANCER; CELL PROTECTION; CHROMOSOME BREAKAGE; CHROMOSOME FRAGILE SITE; CONFERENCE PAPER; ENVIRONMENT; GENE FUNCTION; HUMAN; MUTATION; NONHUMAN; PRIORITY JOURNAL; STRESS; TUMOR SUPPRESSOR GENE; WWOX GENE;

ACID ANHYDRIDE HYDROLASES; ANIMALS; APOPTOSIS; CHROMOSOME FRAGILE SITES; CHROMOSOME FRAGILITY; CHROMOSOMES, HUMAN, PAIR 16; CHROMOSOMES, HUMAN, PAIR 3; GENES, TUMOR SUPPRESSOR; HUMANS; LOSS OF HETEROZYGOSITY; NEOPLASM PROTEINS; NEOPLASMS; OXIDOREDUCTASES;

EID: 30144436775     PISSN: 03043835     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.canlet.2005.07.041     Document Type: Conference Paper

References (96)

1. T.W. Glover, C. Berger, J. Coyle, and B. Echo DNA polymerase alpha inhibition by aphidicolin induces gaps and breaks at common fragile sites in human chromosomes Hum. Genet. 67 1984 136 142
2. J.J. Yunis, and A.L. Soreng Constitutive fragile sites and cancer Science 226 1984 1199 1204
3. G.R. Sutherland, and R.N. Simmers No statistical association between common fragile sites and nonrandom chromosome breakpoints in cancer cells Cancer Genet. Cytogenet. 31 1988 9 15
4. M. Le Beau, H. Drabkin, T. Glover, R. Gemmill, F. Rassool, T. McKeithan, and D. Smith An FHIT tumor suppressor gene? Genes Chromosomes Cancer 21 1998 281 289
5. M. Finnis, S. Dayan, L. Hobson, G. Chenevix-Trench, K. Friend, and K. Ried Common chromosomal fragile site FRA16D mutation in cancer cells Hum. Mol. Genet. 14 2005 1341 1349
6. S. Corbin, M.E. Neilly, R. Espinosa 3rd, E.M. Davis, T.W. McKeithan, and M.M. Le Beau Identification of unstable sequences within the common fragile site at 3p14.2: implications for the mechanism of deletions within fragile histidine triad gene/common fragile site at 3p14.2 in tumors Cancer Res. 62 2002 3477 3484
7. K. Huebner, and C.M. Croce Cancer and the FRA3B/FHIT fragile locus: it's a HIT Br. J. Cancer 88 2003 1501 1506
8. M.F. Arlt, D.E. Miller, D.G. Beer, and T.W. Glover Molecular characterization of FRAXB and comparative common fragile site instability in cancer cells Genes Chromosomes Cancer 33 2002 82 92
9. K. Huebner, and C.M. Croce FRA3B and other common fragile sites: the weakest links Nat. Rev. Cancer 1 2001 214 221
10. L.V. O'Keefe, Y. Liu, A. Perkins, S. Dayan, R. Saint, and R.I. Richards FRA16D common chromosomal fragile site oxido-reductase (FOR/WWOX) protects against the effects of ionizing radiation in Drosophila Oncogene 2005 Advance on-line publication 20th Jun, 2005
11. A.M. Casper, P. Nghiem, M.F. Arlt, and T.W. Glover ATR regulates fragile site stability Cell 111 2002 779 789
12. R.I. Richards Dynamic mutations: a decade of unstable expanded repeats in human genetic disease Hum. Mol. Genet. 10 2001 2187 2194
13. D. Mishmar, A. Rahat, S.W. Scherer, G. Nyakatura, B. Hinzmann, and Y. Kohwi Molecular characterization of a common fragile site (FRA7H) on human chromosome 7 by the cloning of a simian virus 40 integration site Proc. Natl Acad. Sci. USA 95 1998 8141 8146
14. E. Zlotorynski, A. Rahat, J. Skaug, N. Ben-Porat, E. Ozeri, and R. Hershberg Molecular basis for expression of common and rare fragile sites Mol. Cell Biol. 23 2003 7143 7151
15. M.F. Arlt, B. Xu, S.G. Durkin, A.M. Casper, M.B. Kastan, and T.W. Glover BRCA1 is required for common-fragile-site stability via its G2/M checkpoint function Mol. Cell Biol. 24 2004 6701 6709
16. N.G. Howlett, T. Taniguchi, S.G. Durkin, A.D. D'Andrea, and T.W. Glover The Fanconi anemia pathway is required for the DNA replication stress response and for the regulation of common fragile site stability Hum. Mol. Genet. 14 2005 693 701
17. A. Musio, C. Montagna, T. Mariani, M. Tilenni, M.L. Focarelli, and L. Brait SMC1 involvement in fragile site expression Hum. Mol. Genet. 14 2005 525 533
18. M.M. Le Beau, F.V. Rassool, M.E. Neilly, R. Espinosa 3rd, T.W. Glover, D.I. Smith, and T.W. McKeithan Replication of a common fragile site, FRA3B, occurs late in S phase and is delayed further upon induction: implications for the mechanism of fragile site induction Hum. Mol. Genet. 7 1998 755 761
19. L. Wang, J. Darling, J.S. Zhang, H. Huang, W. Liu, and D.I. Smith Allele-specific late replication and fragility of the most active common fragile site, FRA3B Hum. Mol. Genet. 8 1999 431 437
20. A. Palakodeti, Y. Han, Y. Jiang, and M.M. Le Beau The role of late/slow replication of the FRA16D in common fragile site induction Genes Chromosomes Cancer 39 2004 71 76
21. A. Hellman, A. Rahat, S.W. Scherer, A. Darvasi, L.C. Tsui, and B. Kerem Replication delay along FRA7H, a common fragile site on human chromosome 7, leads to chromosomal instability Mol. Cell Biol. 20 2000 4420 4427
22. A. Coquelle, L. Rozier, B. Dutrillaux, and M. Debatisse Induction of multiple double-strand breaks within an hsr by meganucleaseI-SceI expression or fragile site activation leads to formation of double minutes and other chromosomal rearrangements Oncogene 21 2002 7671 7679
23. J.J. Yunis, A.L. Soreng, and A.E. Bowe Fragile sites are targets of diverse mutagens and carcinogens Oncogene 1 1987 59 69
24. C.K. Stein, T.W. Glover, J.L. Palmer, and B.S. Glisson Direct correlation between FRA3B expression and cigarette smoking Genes Chromosomes Cancer 34 2002 333 340
25. T.W. Glover, and C.K. Stein Induction of sister chromatid exchanges at common fragile sites Am J Hum Genet 41 1987 882 890
26. T.W. Glover, and C.K. Stein Chromosome breakage and recombination at fragile sites Am. J. Hum. Genet. 43 1988 265 273
27. F.V. Rassool, T.W. McKeithan, M.E. Neilly, E. van Melle, R. Espinosa, and M.M. Le Beau Preferential integration of marker DNA into the chromosomal fragile site at 3p14: an approach to cloning fragile sites Proc. Natl Acad. Sci. USA 88 1991 6657 6661
28. E.C. Thorland, S.L. Myers, D.H. Persing, G. Sarkar, R.M. McGovern, B.S. Gostout, and D.I. Smith Human papillomavirus type 16 integrations in cervical tumors frequently occur in common fragile sites Cancer Res. 60 2000 5916 5921
29. M. Ciullo, M.A. Debily, L. Rozier, M. Autiero, A. Billault, and V. Mayau Initiation of the breakage-fusion-bridge mechanism through common fragile site activation in human breast cancer cells: the model of PIP gene duplication from a break at FRA7I Hum. Mol. Genet. 11 2002 2887 2894
30. A. Hellman, E. Zlotorynski, S.W. Scherer, J. Cheung, J.B. Vincent, and D.I. Smith A role for common fragile site induction in amplification of human oncogenes Cancer Cell. 1 2002 89 97
31. A. Latil, O. Cussenot, G. Fournier, K. Driouch, and R. Lidereau Loss of heterozygosity at chromosome 16q in prostate adenocarcinoma: identification of three independent regions Cancer Res. 57 1997 1058 1062
32. T. Chen, A. Sahin, and C.M. Aldaz Deletion map of chromosome 16q in ductal carcinoma in situ of the breast: refining a putative tumor suppressor gene region Cancer Res. 56 1996 5605 5609
33. H. Ishii, K. Ozawa, and Y. Furukawa Alteration of the fragile histidine triad gene early in carcinogenesis: an update J. Exp. Ther. Oncol. 3 2003 291 296
34. G. Guler, A. Uner, N. Guler, S.Y. Han, D. Iliopoulos, P. McCue, and K. Huebner Concordant loss of fragile gene expression early in breast cancer development Pathol. Int. 55 2005 471 478
35. D. Iliopoulos, G. Guler, S.Y. Han, D. Johnston, T. Druck, and K.A. McCorkell Fragile genes as biomarkers: epigenetic control of WWOX and FHIT in lung, breast and bladder cancer Oncogene 24 2005 1625 1633
36. H. Ishii, A. Vecchione, Y. Furukawa, K. Sutheesophon, S.Y. Han, and T. Druck Expression of FRA16D/WWOX and FRA3B/FHIT genes in hematopoietic malignancies Mol. Cancer Res. 1 2003 940 947
37. R.V. Jamieson, R. Perveen, B. Kerr, M. Carette, J. Yardley, and E. Heon Domain disruption and mutation of the bZIP transcription factor, MAF, associated with cataract, ocular anterior segment dysgenesis and coloboma Hum. Mol. Genet. 11 2002 33 42
38. M. Chesi, P.L. Bergsagel, O.O. Shonukan, M.L. Martelli, L.A. Brents, and T. Chen Frequent dysregulation of the c-maf proto-oncogene at 16q23 by translocation to an Ig locus in multiple myeloma Blood 91 1998 4457 4463
39. N.A. Becker, E.C. Thorland, S.R. Denison, L.A. Phillips, and D.I. Smith Evidence that instability within the FRA3B region extends four megabases Oncogene 21 2002 8713 8722
40. L.Y. Fong, V. Fidanza, N. Zanesi, L.F. Lock, L.D. Siracusa, and R. Mancini Muir-Torre-like syndrome in Fhit-deficient mice Proc. Natl Acad. Sci. USA 97 2000 4742 4747
41. N. Zanesi, V. Fidanza, L.Y. Fong, R. Mancini, T. Druck, and M. Valtieri The tumor spectrum in FHIT-deficient mice Proc. Natl Acad. Sci. USA 98 2001 10250 10255
42. L. Sard, P. Accornero, S. Tornielli, D. Delia, G. Bunone, and M. Campiglio The tumor-suppressor gene FHIT is involved in the regulation of apoptosis and in cell cycle control Proc. Natl Acad. Sci. USA 96 1999 8489 8492
43. B. Hu, S.Y. Han, X. Wang, M. Ottey, M.B. Potoczek, and A. Dicker Involvement of the Fhit gene in the ionizing radiation-activated ATR/CHK1 pathway J. Cell Physiol. 202 2005 518 523
44. M. Ottey, S.Y. Han, T. Druck, B.L. Barnoski, K.A. McCorkell, and C.M. Croce Fhit-deficient normal and cancer cells are mitomycin C and UVC resistant Br. J. Cancer 91 2004 1669 1677
45. K. Ried, M. Finnis, L. Hobson, M. Mangelsdorf, S. Dayan, and J.K. Nancarrow Common chromosomal fragile site FRA16D sequence: identification of the FOR gene spanning FRA16D and homozygous deletions and translocation breakpoints in cancer cells Hum. Mol. Genet. 9 2000 1651 1663
46. M. Mangelsdorf, K. Ried, E. Woollatt, S. Dayan, H. Eyre, and M. Finnis Chromosomal fragile site FRA16D and DNA instability in cancer Cancer Res. 60 2000 1683 1689
47. K.A. Krummel, L.R. Roberts, M. Kawakami, T.W. Glover, and D.I. Smith The characterization of the common fragile site FRA16D and its involvement in multiple myeloma translocations Genomics 69 2000 37 46
48. H. Ishii, and Y. Furukawa Alterations of common chromosome fragile sites in hematopoietic malignancies Int. J. Hematol. 79 2004 238 242
49. R.I. Aqeilan, T. Kuroki, Y. Pekarsky, O. Albagha, F. Trapasso, and R. Baffa Loss of WWOX expression in gastric carcinoma Clin. Cancer Res. 10 2004 3053 3058
50. K. Driouch, H. Prydz, R. Monese, H. Johansen, R. Lidereau, and E. Frengen Alternative transcripts of the candidate tumor suppressor gene, WWOX, are expressed at high levels in human breast tumors Oncogene 21 2002 1832 1840
51. C. Gourley, A.J. Paige, K.J. Taylor, D. Scott, N.J. Francis, and R. Rush WWOX mRNA expression profile in epithelial ovarian cancer supports the role of WWOX variant 1 as a tumour suppressor, although the role of variant 4 remains unclear Int. J. Oncol. 26 2005 1681 1689
52. G. Guler, A. Uner, N. Guler, S.Y. Han, D. Iliopoulos, and W.W. Hauck The fragile genes FHIT and WWOX are inactivated coordinately in invasive breast carcinoma Cancer 100 2004 1605 1614
53. T. Kuroki, F. Trapasso, T. Shiraishi, H. Alder, K. Mimori, M. Mori, and C.M. Croce Genetic alterations of the tumor suppressor gene WWOX in esophageal squamous cell carcinoma Cancer Res. 62 2002 2258 2260
54. T. Kuroki, S. Yendamuri, F. Trapasso, A. Matsuyama, R.I. Aqeilan, and H. Alder The tumor suppressor gene WWOX at FRA16D is involved in pancreatic carcinogenesis Clin. Cancer Res. 10 2004 2459 2465
55. M.I. Nunez, J. Ludes-Meyers, M.C. Abba, H. Kil, N.W. Abbey, and R.E. Page Frequent loss of WWOX expression in breast cancer: correlation with estrogen receptor status Breast Cancer Res. Treat. 89 2005 99 105
56. M.I. Nunez, D.G. Rosen, J.H. Ludes-Meyers, M.C. Abba, H. Kil, and R. Page WWOX protein expression varies among ovarian carcinoma histotypes and correlates with less favorable outcome BMC Cancer 5 2005 64
57. A.J. Paige, K.J. Taylor, C. Taylor, S.G. Hillier, S. Farrington, and D. Scott WWOX: a candidate tumor suppressor gene involved in multiple tumor types Proc. Natl Acad. Sci. USA 98 2001 11417 11422
58. S.W. Park, J. Ludes-Meyers, D.B. Zimonjic, M.E. Durkin, N.C. Popescu, and C.M. Aldaz Frequent downregulation and loss of WWOX gene expression in human hepatocellular carcinoma Br. J. Cancer 91 2004 753 759
59. M.C. Yakicier, P. Legoix, C. Vaury, L. Gressin, E. Tubacher, and F. Capron Identification of homozygous deletions at chromosome 16q23 in aflatoxin B1 exposed hepatocellular carcinoma Oncogene 20 2001 5232 5238
60. S. Yendamuri, T. Kuroki, F. Trapasso, A.C. Henry, K.R. Dumon, and K. Huebner containing oxidoreductase gene expression is altered in non-small cell lung cancer Cancer Res. 63 2003 878 881
61. J.E. Watson, N.A. Doggett, D.G. Albertson, A. Andaya, A. Chinnaiyan, and H. van Dekken Integration of high-resolution array comparative genomic hybridization analysis of chromosome 16q with expression array data refines common regions of loss at 16q23-qter and identifies underlying candidate tumor suppressor genes in prostate cancer Oncogene 23 2004 3487 3494
62. A.K. Bednarek, C.L. Keck-Waggoner, R.L. Daniel, K.J. Laflin, P.L. Bergsagel, and K. Kiguchi WWOX, the FRA16D gene, behaves as a suppressor of tumor growth Cancer Res. 61 2001 8068 8073
63. A. Watanabe, Y. Hippo, H. Taniguchi, H. Iwanari, M. Yashiro, and K. Hirakawa An opposing view on WWOX protein function as a tumor suppressor Cancer Res. 63 2003 8629 8633
64. A.K. Bednarek, K.J. Laflin, R.L. Daniel, Q. Liao, K.A. Hawkins, and C.M. Aldaz WWOX, a novel WW domain-containing protein mapping to human chromosome 16q23.3-24.1, a region frequently affected in breast cancer Cancer Res. 60 2000 2140 2145
65. N.S. Chang, N. Pratt, J. Heath, L. Schultz, D. Sleve, G.B. Carey, and N. Zevotek Hyaluronidase induction of a WW domain-containing oxidoreductase that enhances tumor necrosis factor cytotoxicity J. Biol. Chem. 276 2001 3361 3370
66. N.S. Chang, J. Doherty, and A. Ensign JNK1 physically interacts with WW domain-containing oxidoreductase (WOX1) and inhibits WOX1-mediated apoptosis J. Biol. Chem. 278 2003 9195 9202
67. R.I. Aqeilan, Y. Pekarsky, J.J. Herrero, A. Palamarchuk, J. Letofsky, and T. Druck Functional association between Wwox tumor suppressor protein and p73, a p53 homolog Proc. Natl Acad. Sci. USA 101 2004 4401 4406
68. C.I. Sze, M. Su, S. Pugazhenthi, P. Jambal, L.J. Hsu, and J. Heath Down-regulation of WW domain-containing oxidoreductase induces Tau phosphorylation in vitro. A potential role in Alzheimer's disease J. Biol. Chem. 279 2004 30498 30506
69. J.H. Ludes-Meyers, H. Kil, A.K. Bednarek, J. Drake, M.T. Bedford, and C.M. Aldaz WWOX binds the specific proline-rich ligand PPXY: identification of candidate interacting proteins Oncogene 23 2004 5049 5055
70. R.I. Aqeilan, A. Palamarchuk, R.J. Weigel, J.J. Herrero, Y. Pekarsky, and C.M. Croce Physical and functional interactions between the Wwox tumor suppressor protein and the AP-2gamma transcription factor Cancer Res. 64 2004 8256 8261
71. A.M. Brumby, and H.E. Richardson Using Drosophila melanogaster to map human cancer pathways Nat. Rev. Cancer 5 2005 626 639
72. N. Sogame, M. Kim, and J.M. Abrams Drosophila p53 preserves genomic stability by regulating cell death Proc. Natl Acad. Sci. USA 100 2003 4696 4701
73. J.H. Lee, E. Lee, J. Park, E. Kim, J. Kim, and J. Chung In vivo p53 function is indispensable for DNA damage-induced apoptotic signaling in Drosophila FEBS Lett. 550 2003 5 10
74. T.W. Glover, A.W. Hoge, D.E. Miller, J.E. Ascara-Wilke, A.N. Adam, and S.L. Dagenais The murine Fhit gene is highly similar to its human orthologue and maps to a common fragile site region Cancer Res. 58 1998 3409 3414
75. K.A. Krummel, S.R. Denison, E. Calhoun, L.A. Phillips, and D.I. Smith The common fragile site FRA16D and its associated gene WWOX are highly conserved in the mouse at Fra8E1 Genes Chromosomes Cancer 34 2002 154 167
76. L. Rozier, E. El-Achkar, F. Apiou, and M. Debatisse Characterization of a conserved aphidicolin-sensitive common fragile site at human 4q22 and mouse 6C1: possible association with an inherited disease and cancer Oncogene 23 2004 6872 6880
77. M. Iida, C.H. Anna, W.M. Holliday, J.B. Collins, M.L. Cunningham, R.C. Sills, and T.R. Devereux Unique patterns of gene expression changes in liver after treatment of mice for 2 weeks with different known carcinogens and non-carcinogens Carcinogenesis 26 2005 689 699
78. H. Ishii, K. Mimori, T. Inageta, Y. Murakumo, A. Vecchione, M. Mori, and Y. Furukawa Components of DNA damage checkpoint pathway regulate UV exposure-dependent alterations of gene expression of FHIT and WWOX at chromosome fragile sites Mol. Cancer Res. 3 2005 130 138
79. H. Ishii, K. Mimori, A. Vecchione, K. Sutheesophon, T. Fujiwara, M. Mori, and Y. Furukawa Effect of exogenous E2F-1 on the expression of common chromosome fragile site genes, FHIT and WWOX Biochem. Biophys. Res. Commun. 316 2004 1088 1093
80. J. Bartkova, Z. Horejsi, K. Koed, A. Kramer, F. Tort, and K. Zieger DNA damage response as a candidate anti-cancer barrier in early human tumorigenesis Nature 434 2005 864 870
81. V.G. Gorgoulis, L.V. Vassiliou, P. Karakaidos, P. Zacharatos, A. Kotsinas, and T. Liloglou Activation of the DNA damage checkpoint and genomic instability in human precancerous lesions Nature 434 2005 907 913
82. J. Fang, M. Arlt, A. Burgess, S. Dagenais, D. Beer, and T. Glover Translocation breakpoints in FHIT and FRA3B in both homologs of chromosome 3 in an esophageal adenocarcinoma Genes Chromosomes Cancer 30 2001 292 298
83. T. Glover, J. Coyle-Morris, F. Li, R. Brown, C. Berger, R. Gemmill, and F. Hecht Translocation t(3;8)(p14.2;q24.1) in renal cell carcinoma affects expression of the common fragile site at 3p14(FRA3B) in lymphocytes Cancer Genet. Cytogenet. 31 1988 69 73
84. K. Mimori, T. Druck, H. Inoue, H. Alder, L. Berk, and M. Mori Cancer-specific chromosome alterations in the constitutive fragile region FRA3B Proc. Natl Acad. Sci. USA 96 1999 7456 7461
85. S.R. Denison, F. Wang, N.A. Becker, B. Schule, N. Kock, and L.A. Phillips Alterations in the common fragile site gene Parkin in ovarian and other cancers Oncogene 22 2003 8370 8378
86. S.R. Denison, G. Callahan, N.A. Becker, L.A. Phillips, and D.I. Smith Characterization of FRA6E and its potential role in autosomal recessive juvenile parkinsonism and ovarian cancer Genes Chromosomes Cancer 38 2003 40 52
87. F. Wang, S. Denison, J.P. Lai, L.A. Philips, D. Montoya, and N. Kock Parkin gene alterations in hepatocellular carcinoma Genes Chromosomes Cancer 40 2004 85 96
88. M.C. Picchio, E.S. Martin, R. Cesari, G.A. Calin, S. Yendamuri, and T. Kuroki Alterations of the tumor suppressor gene Parkin in non-small cell lung cancer Clin. Cancer Res. 10 2004 2720 2724
89. R. Cesari, E.S. Martin, G.A. Calin, F. Pentimalli, R. Bichi, and H. McAdams Parkin, a gene implicated in autosomal recessive juvenile parkinsonism, is a candidate tumor suppressor gene on chromosome 6q25-q27 Proc. Natl Acad. Sci. USA 100 2003 5956 5961
90. O. Bluteau, J.C. Beaudoin, P. Pasturaud, J. Belghiti, D. Franco, and P. Bioulac-Sage Specific association between alcohol intake, high grade of differentiation and 4q34-q35 deletions in hepatocellular carcinomas identified by high resolution allelotyping Oncogene 21 2002 1225 1232
91. K.O. Robinson, A.M. Petersen, S.N. Morrison, C.M. Elso, and L. Stubbs Two reciprocal translocations provide new clues to the high mutability of the Grid2 locus Mamm. Genome 16 2005 32 40
92. H. Huang, J. Qian, J. Proffit, K. Wilber, R. Jenkins, and D.I. Smith FRA7G extends over a broad region: coincidence of human endogenous retroviral sequences (HERV-H) and small polydispersed circular DNAs (spcDNA) and fragile sites Oncogene 16 1998 2311 2319
93. C. Tatarelli, A. Linnenbach, K. Mimori, and C.M. Croce Characterization of the human TESTIN gene localized in the FRA7G region at 7q31.2 Genomics 68 2000 1 12
94. M.Z. Limongi, F. Pelliccia, and A. Rocchi Characterization of the human common fragile site FRA2G Genomics 81 2003 93 97
95. G. Callahan, S.R. Denison, L.A. Phillips, V. Shridhar, and D.I. Smith Characterization of the common fragile site FRA9E and its potential role in ovarian cancer Oncogene 22 2003 590 601
96. C. Morelli, E. Karayianni, C. Magnanini, A.J. Mungall, E. Thorland, and M. Negrini Cloning and characterization of the common fragile site FRA6F harboring a replicative senescence gene and frequently deleted in human tumors Oncogene 21 2002 7266 7276