UV exposure, genetic targets in melanocytic tumors and transgenic mouse models

Photochemistry and Photobiology

Volumn 81, Issue 1, 2005, Pages 52-64

  De Gruijl, Frank R ^a   Van Kranen, Henk J ^b   Van Schanke, Arne ^a  

^a LEIDEN UNIVERSITY MEDICAL CENTER   (Netherlands)

^b NATIONAL INSTITUTE FOR PUBLIC HEALTH AND THE ENVIRONMENT   (Netherlands)

Author keywords

[No Author keywords available]

Indexed keywords

PROTEIN TYROSINE KINASE; RETINOBLASTOMA PROTEIN;

ANIMAL CELL; ANIMAL EXPERIMENT; ANIMAL MODEL; ANIMAL TISSUE; CANCER INHIBITION; CARCINOGENESIS; CONFERENCE PAPER; CONTROLLED STUDY; ENHANCER REGION; MELANOCYTE; MELANOCYTOSIS; MELANOMA; MOUSE; NONHUMAN; PIGMENTATION; PROMOTER REGION; RADIATION EXPOSURE; TRANSCRIPTION REGULATION; TRANSGENIC MOUSE; ULTRAVIOLET A RADIATION; ULTRAVIOLET B RADIATION; ULTRAVIOLET RADIATION;

ANIMALS; DISEASE MODELS, ANIMAL; GENETIC PREDISPOSITION TO DISEASE; MELANOMA; MICE; MICE, TRANSGENIC; SIGNAL TRANSDUCTION; SKIN NEOPLASMS; ULTRAVIOLET RAYS;

ANIMALIA; MUS MUSCULUS;

EID: 14744279538     PISSN: 00318655     EISSN: None     Source Type: Journal    
DOI: 10.1562/2004-09-26-IR-328.1     Document Type: Conference Paper

References (165)

1. De Vries, E., F. Bray, J. W. W. Coeberg and D. M. Parkin, ENCR (2003) Changing epidemiology of malignant cutaneous melanoma in Europe 1969-1997; rising trends in incidence and mortality, but recent stabilizations in Western Europe and decreases in Scandinavia. Int. J. Cancer 107, 119-126.
2. Holman, C. D. J. and B. K. Armstrong (1984) Cutaneous malignant melanoma and indicators of total accumulated exposure to the sun: an analysis separating histogenic types. J. Natl. Cancer Inst. 73, 75-82.
3. Whiteman, D. C., C. A. Whiteman and A. C. Green (2001) Childhood sun exposure a as risk factor for melanoma: a systematic review of epidemiological studies. Cancer Causes Control 12, 69-82.
4. Gallagher, R. P., D. I. McLean, G. P. Yang, A. J. Coldman, H. K. Silver and J. J. Spinelli (1990) Suntan, sunburn and pigmentation factors and the frequency of acquired nevi in children. Similarities to melanoma: the Vancouver Mole Study. Arch. Dermatol. 126, 770-776.
5. Darlington, S., V. Siskind, L. Green and A. Green (2002) Longitudinal study of melanocytic nevi in adolescents. J. Am. Acad. Dermatol. 46, 715-722.
6. Wiecker, T. S., H. Luther, P. Buettner, J. Bauer and C. Garbe (2003) Moderate sun exposure and nevus counts in parents are associated with development of melanocytic nevi in childhood: a risk factor study in 1,812 kindergarten children. Cancer 97, 628-638.
7. Kraemer, K. H., M. M. Lee and J. Scotto (1987) Xeroderma pigmentosum; cutaneous ocular, and neurologic abnormalities in 830 published cases. Arch. Dermatol. 123, 241-250.
8. Valverde, P., E. Healy, I. Jackson, J. L. Rees and A. J. Thody (1995) Variants of the melanocyte-stimulating hormone receptor gene are associated with red hair and fair skin in humans. Nature Genet. 11, 328-330.
9. Palmer, J. S., D. L. Duffy, N. F. Box, J. F. Aitken, L. E. O'Gorman, A. C. Green, N. K. Hayward, N. G. Martin and R. A. Strum (2000) Melanocortin-1 receptor polymorphisms and risk of melanoma: is the association explained solely by pigmentation phenotype? Am. J. Hum. Genet. 66, 176-186.
10. Goldstein, A. M., J. P. Struewing, A. Chidambaran, M. C. Fraser and M. A. Tucker (2000) Genotype-phenotype relationships in U.S. melanoma prone families with CDKN2A and CDK4 mutations. J. Natl. Cancer Inst. 92, 1006-1010.
11. Bishop, D. T., F. Demenais, A. M. Goldstein, W. Bergman, J. N. Bishop, B. Bressac-de Paillerets, A. Chompret, P. Ghiorzo, N. Gruis, J. Hansson, M. Harland, N. Hayward, E. A. Holland, G. J. Mann, M. Mantelli, D. Nancarrow, A. Platz and M. A. Tucker (2002) Geographical variation in the penetrance of CDKN2A mutations for melanoma. J. Natl. Cancer Inst. 94, 894-903.
12. Box, N. F., D. L. Duffy, W. Chen, M. Stark, N. G. Martin, R. A. Sturm and N. K. Hayward (2001) MC1R genotype modifies risk of melanoma in families segregating CDKN2A mutations. Am. J. Hum. Genet. 69, 765-773.
13. Van der Velden, P. A., L. A. Sandkuijl, W. Bergman, S. Pavel, L. van Mourik, R. R. Frants and N. A. Gruis (2001) Melanocortin-1 receptor variant R151C modifies melanoma risk in Dutch families with melanoma. Am. J. Hum. Genet. 69, 774-779.
14. Chaudru, V., A. Chompret, B. Bressac-de Paillerets, A. Spatz, M. F. Avril and F. Demenais (2004) Influence of genes, nevi, and sun sensitivity on melanoma risk in a family sample unselected by history and in melanoma-prone families. J. Natl. Cancer Inst. 96, 785-795.
15. Eskandarpour, M., J. Hashemi, L. Kanter, U. Ringborg and J. Hansson (2003) Frequency of UV-inducible NRAS mutations in melanomas of patients with germline CDKN2A mutations. J. Natl. Cancer Inst. 95, 790-798.
16. Van Elsas, A., S. F. Zerp, S. van der Flier, K. M. Kruse, C. Aarnoudse, N. K. Hayward, D. J. Ruiter and P. I. Schrier (1996) Relevance of ultraviolet-induced N-ras oncogene point mutations in development of primary human cutaneous melanoma. Am. J. Pathol. 143, 883-893.
17. Van't Veer, L. J., B. M. T. Burgering, R. Versteeg, A. J. M. Boot, D. J. Ruiter, S. Osanto, P. I. Schrier and J. L. Bos (1989) N-ras mutations in human cutaneous melanoma from sun-exposed body sites. Mol. Cell. Biol. 9, 3114-3116.
18. Brash, D. E., J. A. Rudolph, J. A. Simon, A. Lin, G. J. McKenna, H. P. Baden, A. J. Halperin and J. A. Ponten (1991) Role for sunlight in skin cancer: UV-induced p53 mutations in squamous cell carcinomas. Proc. Natl. Acad. Sci. USA 88, 10124-10128.
19. Davies, H., G. R. Bignell, C. Cox, P. Stephens, S. Edkins, S. S. Clegg, J. Teague, H. Woffendin, M. J. Garnett, W. Bottomley, N. Davis. E. Dicks, R. Ewing, Y. Floyd, K. Gray, S. Hall, R. Hawes, J. Hughes, V. Kosmidou, A. Menzies, C. Mould, A. Parker, C. Stevens, S. Watt, S. Hooper, R. Wilson, H. Jayatilake, B. A. Gusterson, C. Cooper, J. Shipley, D. Hargrave, K. Pritchard-Jones, N. Maitland, G. Chenevix-Trench, G. J. Riggins, D. D. Bigner, G. Palmieri, A. Cossu, A. Flanagan, A. Nicholson, J. W. Ho, S. Y. Leung, S. T. Yuen, B. L. Weber, H. F. Seigler, T. L. Darrow, H. Paterson, R. Marais, C. J. Marshall, R. Wooster, M. R. Stratton and P. A. Futreal (2002) Mutations of the B-RAF gene in human cancer. Nature 417, 949-954.
20. Pollock, F. M., U. L. Harper, K. S. Hansen, L. M. Yudt, M. Stark, C. M. Robbins, T. Y. Moses, G. Hostetter, U. Wagner, J. Kakareka, G. Salem, T. Pohida, P. Heenan, P. Duray, O. Kallioniemi, N. K. Hayward, J. M. Trent and P. S. Meltzer (2003) High frequency of BRAF mutations in nevi. Nat. Genet. 33, 19-20.
21. Kumar, R., S. Angelini, E. Snellman and K. Hemminki (2004) BRAF mutations are common somatic events in melanocytic nevi. J. Investig. Dermatol. 122, 342-348.
22. Saldana, G., D. Purnell, A. Fletcher, L. Potter, A. Gillies and J. H. Pringle (2004) High BRAF mutation frequency does not characterize all melanocytic tumor types. Int. J. Cancer 111, 705-710.
23. Kamiya, H. and H. Kasia (1997) Mutations induced by 2-hydroxyadenine on a shuttle vector during leading and lagging strand synthesis in mammalian cells. Biochemistry 36, 11125-11130.
24. Omholt, K., A. Platz, L. Kanter, U. Ringborg and J. Hansson (2003) NRAS and BRAF mutations arise early during melanoma pathogenesis and are preserved throughout tumor progression. Clin. Cancer Res. 9, 6483-6488.
25. Pavey, S., P. Johansson, L. Packer, J. Taylor, M. Stark, P. M. Pollock, G. J. Walker, G. M. Boyle, U. Harper, S. J. Corri, K. Hansen, L. Yudt, C. Schmidt, P. Hersey, K. A. Ellem, M. G. O'Rourke, P. G. Parsons, P. Meltzer, M. Ringner and N. K. Hayward (2004) Microarray expression profiling in melanoma reveals BRAF mutation signature. Oncogene 23, 4060-4067.
26. Maldonado, J. L., J. Fridlyand, H. Patel, A. N. Jain, K. Busan, T. Kageshita, T. Ono, D. G. Albertson, D. Finkel and B. C. Bastian (2003) Determinants of BRAF mutations in primary melanomas. J. Natl. Cancer Inst. 95, 1878-1890.
27. Cohen, Y., E. Rosenbaum, S. Begun, D. Goldenberg, C. Esche, O. Lavis, D. Sidransky and W. H. Westra (2004) Exon 15 BRAF mutations are uncommon in melanoma arising in nonsun-exposed sites. Clin. Cancer Res. 10, 3444-3447.
28. INK4a/CDKN2A tumor suppressor and its relatives. Biochim. Biophys. Acta 1378, F115-F177.
29. Funk, J. O., P. I. Schiller, M. T. Barrett, D. J. Wong, P. Kind and C. A. Sander (1998) p16INK4a expression is frequently decreased and associated with 9p21 loss of heterozygosity in sporadic melanoma. J. Cutan. Pathol. 25, 291-296.
30. Daniotti, M., M. Oggionini, T. Ranzani, V. Vallacchi, V. Campi, D. Di Statsi, G. D. Torre, F. Perrone, C. Luonni, S. Suardi, M. Frattini, S. Pilotti, A. Anichi, G. Tragni, G. Parmiani, M. A. Pierotti and M. Rodolfo (2004) BRAF alterations are associated with complex mutational profiles in malignant melanoma. Oncogene 23, 5968-5977.
31. Rahdi, J. M. (1999) Malignant melanoma arising from nevi, p53, p16, and Bcl-2: expression in benign versus malignant components. J. Cutan. Med. Surg. 3, 293-297.
32. Bogdan, I., J. Smolle, H. Kerl, G. Burg and R. Boni (2003) Melanoma ex naevo: a study of associated naevus. Melanoma Res. 13, 213-217.
33. Tran, T. P., L. Titus-Ernstoff, A. E. Perry, M. S. Emstoff, and I. F. Newsham (2002) Alteration of chromosome 9p21 and/or p16 in benign and dysplastic nevi suggests a role in early melanoma progression (United States). Cancer Causes Control 13, 675-682.
34. Park, W. S., A. O. Vortmeyer, S. Pack, P. H. Duray, R. Boni, A. A. Guerami, M. R. Emmert-Buck, L. A. Liotta and Z. Zhuang (1998) Allelic deletion at chromosome 9p21 (p16) and 17p13 (p53) in microdissected sporadic dysplastic naevus. Hum. Pathol. 29, 127-130.
35. Lee, J. Y., S. M. Doog, M. S. Shin, S. Y. Kim, S. H. Lee, S. J. Kang, J. D. Lee, C. S. Kim, S. H. Kim and N. J. Yoo (1997) Genetic alterations of p16INK4a and p53 genes in sporadic dysplastic nevus. Biochem. Biophys. Res. Commun. 237, 667-672.
36. Sparrow, L. E., M. J. Eldon, D. R. English and P. J. Heenan (1998) P16 and p21WAF1 protein expression in melanocytic tumors by immunohistochemistry. Am. J. Dermapathol. 20, 255-261.
37. Keller-Melchior, R., R. Schmidt and M. Piepkorn (1998) Expression of the tumor suppressor gene product p16INK4a in benign and malignant melanocytic lesions. J. Investig. Dermatol. 110, 932-938.
38. Talve, L., I. Sauroja, Y. Collan, K. Punnonen and T. Ekfors (1997) Loss of expression of the p16INK4/CDKN2 gene in cutaneous malignant melanoma correlates with tumor cell proliferation and invasive stage. Int. J. Cancer 74, 255-259.
39. Wang, Y. L., H. Uhara, Y. Yamazaki, T. Nikaido and T. Saida (1996) Immunohistochemical detection of CDK4 and p16INK4 proteins in cutaneous malignant melanoma. Br. J. Dermatol. 134, 269-275.
40. Bennett, D. C. (2003) Human melanocytes senescence and melanoma susceptibility genes. Oncogene 22, 3063-3069.
41. Tsao, H., M. C. Mihm and C. Sheenan (2003) PTEN expression in normal skin, acquired melanocytic nevi, and cutaneous melanoma. J. Am. Acad. Dermatol. 49, 865-872.
42. Whiteman, D. C., X. P. Zhiu, M. C. Cummings, S. Pavey, N. K. Hayward and C. Eng (2002) Nuclear PTEN expression and clinicopathological features in a population-based series of primary cutaneous melanoma. Int. J. Cancer 99, 63-67.
43. Zhou, X. P., O. Gimm, H. Hampel, T. Niemann, M. J. Walker and C. Eng (2000) Epigenetic PTEN silencing in malignant melanomas without PTEN mutation. Am. J. Pathol. 157, 1123-1128.
44. Birck, A., V. Ahrenkiel, J. Zeuthen, K. Hou-Jensen and P. Guldberg (2000) Mutation and allelic loss of the PTEN/MMAC1 gene in primary and metastatic melanoma biopsies. J. Investig. Dermatol. 114, 277-280.
45. Guldberg, P., P. thor Straten, A. Birck, V. Ahrenkiel, A. F. Kirkin and J. Zeuthen (1997) Disruption of the MMAC1/PTEN gene by deletion or mutation is a frequent event in malignant melanoma. Cancer Res. 57, 3660-3663.
46. Boni, R., A. O. Vortmeyer, G. Burg, G. Hofbauer and Z. Zhuang (1998) The PTEN tumor suppressor gene and malignant melanoma. Melanoma Res. 8, 300-302.
47. Deichmann, H., M. Thome, A. Benner, U. Egner, W. Hartschuh and H. Naher (2002) PTEN/MMAC1 expression in melanoma resection specimens. Br. J. Cancer 87, 1431-1436.
48. Tsao, H., X. Zhang, K. Fowlkes and F. G. Haluska (2000) Relative reciprocity of NRAS and PTEN/MACC1 alterations in cutaneous melanoma cell lines. Cancer Res. 60, 1800-1804.
49. Tsao, H., V. Goel, H. Wu, G. Yang and F. G. Haluska (2004) Genetic interaction between NRAS and BRAF mutations and PTEN/MMAC1 inactivation in melanoma. J. Investig. Dermatol. 122, 337-341.
50. Cohen, C., A. Zavala-Pompa, J. H. Sequeira, M. Shoji, D. G. Sexton, G. Cotsonis, F. Cerimele, B. Govindarajan, N. Macaron and J. L. Arbiser (2002) Mitogen-activated protein kinase activation is an early event in melanoma progression. Clin. Cancer Res. 8, 3728-3733.
51. Satyamoorthy, K., G. Li, M. R. Gerrerro, M. S. Brose, P. Volpe, B. L. Weber, P. Van Belle, D. E. Elder and M. Herlyn (2003) Constitutive mitogen-activated protein kinase activation in melanoma is mediated by both BRAF mutations and autocrine growth factor stimulation. Cancer Res. 63, 756-759.
52. Dawan, P., A. B. Singh, D. L. Ellis and A. Richmond (2002) Constitutive activation of Akt/protein kinase B in melanoma leads to up-regulation of nuclear factor-kappaB and tumor progression. Cancer Res. 62, 7335-7342.
53. Bastian, B., J. Curtin, J. Fridlyand, H. Patel, D. Pinkel, K. Busam, H. Kutzner and T. Kageshita (2004) Braf and melanocytic diseases. Pigment Cell Res. 17, 428-428.
54. De Gruijl, F. R., H. J. van Kranen and L. H. Mullenders (2001) UV-induced DNA damage, repair, mutations and oncogenic pathways in skin cancer. J. Photochem. Photobiol. B. Biol. 63, 19-27.
55. Rodeck, U. (1993) Growth factor independence and growth regulation pathways in human melanoma development. Cancer Metastasis Rev. 12, 219-226.
56. Halaban, R., B. Fan, J. Ahn, Y. Funasaka, H. Gitay-Goren and G. Neufeld (1992) Growth factors, receptor kinases, and protein tyrosine phosphatases in normal and malignant melanocytes. J. Immonother. 12, 154-161.
57. Easty, D. J., M. Herlyn and D. C. Bennett (1995) Abnormal protein tyrosine kinase gene expression during melanoma progression and metastasis. Int. J. Cancer 60, 129-136.
58. Easty, D. J. and D. C. Bennett (2000) Protein tyrosine kinases in malignant melanoma. Melanoma Res. 10, 401-411.
59. Kakurai, M., N. Fujita, T. Kiyosawa, T. Inoue, S. Ishibashi, Y. Furukawa, T. Demitsu and H. Nakagawa (2002) Vasoactive intestinal peptide and cytokines enhance stem cell factor production from epidermal keratinocytes DJM-1. J. Investig. Dermatol. 119, 1183-1188.
60. Hachiya, A., A. Kobayashi, A. Chuchi, Y. Takema and G. Imokawa (2001) The paracrine role of stem cell factor/c-kit signaling in the activation of human melanocyte in ultraviolet-B-induced pigmentation. J. Investig. Dermatol. 116, 578-586.
61. McCarthy, M. F., D. R. Bielenberg, M. B. Nilsson, J. E. Gershenwald, R. L. Barnhill, P. Ahearne, C. D. Bucana and I. J. Fidler (2003) Epidermal hyperplasia overlying human melanoma correlates with tumour depth and angiogenesis. Melanoma Res. 13, 379-387.
62. Bielenberg, D. R., C. D. Bucana, R. Sanchez, C. K. Donawho, M. L. Kripke, and I. J. Fidler (1998) Molecular regulation of UVB-induced cutaneous angiogenesis. J. Investig. Dermatol. 111, 864-872.
63. Nesbit, M., H. K. Nesbit, J. Bennett, T. Ani, M. Y. Hsu, E. Dejesus, M. McBrian, A. R. Gupta, S. L. Eck and M. Herlyn (1999) Basic fibroblast growth factor induces a transformed phenotype in normal human melanocytes. Oncogene 18, 6469-6476.
64. Castagnino, P., M. V. Lorenzi, J. Yeh, D. Breckenridge, H. Sakatp, B. Munz, S. Wrener and D. P. Bottaro (2000) Neu differentiation factor/heregulin induction by hepatocyte and keratinocyte growth factors. Oncogene 19, 640-648.
65. Rudman, S. M., M. P. Philpott, G. A. Thomas and T. Kesley (1997) The role of IGF-1 in human skin and its appendages: morphogen as well as mitogen? J. Investig. Dermatol. 109, 770-777.
66. Blaudschun, R., C. Sunderkotter, P. Brenneisen, R. Hinrichs, T. Peters, L. Schneider, Z. Razi-Wolf, N. Hunzelman and K. Scharffetter-Kochanek (2002) Vascular endothelial growth factor causally contributes to the angiogenic response upon ultraviolet B irradiation in vivo. Br. J. Dermatol. 146, 581-587.
67. Trompezinski, S., I. Pernet, D. Schmitt and J. Viac (2001) UV radiation and prostaglandin E2 up-regulate vascular endothelial growth factor (VEGF) in cultured human fibroblasts. Inflamm. Res. 50, 422-427.
68. Gron, B., K. Stoltze, A. Anderson, and E. Dabelsteen (2002) Oral fibroblasts produce more HGF and KGF than skin fibroblasts in response to co-culture with keratinocytes. APMIS 110, 892-898.
69. Larcher, F., M. Franco, M. Bolontrade, M. Rodriguez-Puebla, L. Casanova, M. Navarro, G. Yancopoulos, J. L. Jorcano and C. J. Conti (2003) Modulation of the angiogenesis response through Ha-ras control, placenta growth factor, and angiopoietin expression in mouse skin carcinogenesis. Mol. Carcinog. 37, 83-90.
70. Pomyje, J., J. H. Zivny, T. Stopka, J. Simak, M. Vankova and E. Necas (2001) Angiopoietin-1, angiopoietin-2, and Tie-2 in tumour and non-tumour tissues during growth of experimental melanoma. Melanoma Res. 11, 639-643.
71. Easty, D. J., S. P. Hill, M. Y. Hsu, M. E. Fallowfield, V. A. Florenes, M. Herlyn and D, C. Bennett (1999) Up-regulation of ephrin-A1 during melanoma progression. Int. J. Cancer 84, 494-501.
72. Grinnich, J. M., J. A. Burch, J. Burchette and C. R. Shea (1998) The SCF/KIT pathway plays a critical role in the control of normal human melanocyte homeostasis. J. Investig. Dermatol. 111, 233-238.
73. Wehrle-Haller, B. and J. A. Weston (1995) Soluble and cell-bound forms of steel factor activity play distinct roles in melanocyte precursor dispersal and survival on the lateral neural crest migration pathway. Development 121, 731-742.
74. Huang, S., D. Jean, M. Luca, M. A. Tainsky and M. Bar-Eli (1998) Loss of AP-2 results in downregulation of c-KIT and enhancement of melanoma tumorigenicity and metastasis. EMBO J. 17, 4358-4369.
75. Zukat, R., R. Perills, S. Eliyahu, Y. Yarden, D. Givol, S. D. Lyman and R. Halaban (1993) KIT ligand (mast cell growth factor) inhibits the growth of KIT-expressing melanoma cells. Oncogene 8, 2221-2229.
76. Huang, S., M. Luca, M. Gutman, D. J. McConkey, K. E. Langley, S. D. Lyman and M. Bar-Eli (1996) Enforced c-KIT expression renders highly metastatic human melanoma cells susceptible to stem cell factor-induced apoptosis and inhibits their tumorigenic and metastatic potential. Oncogene 13, 2339-2347.
77. Natali, P. G., M. R. Nicotra, M. F. Di Rento, M. Prat, A. Bigotti, R. Cavaliere and P. M. Comoglio (1993) Expression of the c-Met/ HGF receptor in human melanocytic neoplasms: demonstration of the relationship to malignant melanoma tumour progression. Br. J. Cancer 68, 746-750.
78. Kos, L., A. Aronzon, H. Takayama, F. Maina, C. Ponzetto, G. Merlino and W. Pavan (1999) Hepatocyte growth factor/scatter factor-MET signaling in neural crest-derived melanocyte development. Pigment Cell Res. 12, 13-21.
79. Nicolau, A., S. E. Estdale, M. Tsatmali, D. P. Herrero and A. J. Thody (2004) Protaglandin production by melanocytic cells and the effect of alpha-melanocyte stimulating hormone. FEBS Lett. 570, 223-226.
80. Matsumoto, K., H. Okazaki and T. Nakamura (1995) Novel function of prostaglandins as inducers of gene expression of HGF and putative mediators of tissue regulation. J. Biochem. (Tokyo) 117, 458-464.
81. Matsumoto-Taniura, N., K. Matsumoto and T. Nakamura (1999) Prostaglandin production in mouse mammary tumour cells confers invasive growth potential by inducing hepatocyte growth factor in stromal fibroblasts. Br. J. Cancer 81, 194-202.
82. Zhao, J., R. Pestell, and J.-L. Guan (2001) Transcriptional activation of cyclin D1 promoter by FAK contributes to cell cycle progression. Mol. Biol. Cell 12, 4066-4077.
83. Page, K., J. Li, K. C. Cosbit, K. M. Rumilla, J. W. Soh, I. B. Weinstein, C. Albanese, R. G. Pestell, M. R. Rosner and M. B. Hershenson (2002) Regulation of airway smooth muscle cyclin D1 transcription by protein kinase C-delta. Am. J. Respir. Cell Mol. Biol. 27, 204-213.
84. Wandzioch, E., C. E. Edling, R. H. Palmer, L. Carlson, and B. Hallberg (2004) Activation of MAP kinase pathway by c-Kit is PI-3 kinase dependent in hematopoietic progenitor/stem cell lines. Blood 104, 51-57.
85. Recio, J. A. and G. Merlino (2002) Hepatocyte growth factor/scatter factor activates proliferation in melanoma cells through p38 MAPK, ATF-2 and cyclin D1. Oncogene 21, 1000-1008.
86. Hernandez, I., M. Malumbres, P. Leonardi and A. Pellicer (2000) The Rgr oncogene (homologous to RalGDS) induced transformation and gene expression by activating Ras, Ral and Rho mediated pathways. Oncology 19, 2745-2757.
87. Pruitt, K., W. M. Pruitt, G. K. Bilter, J. K. Westwick and C. J, Der (2002) Raf-independent deregulation of p38 and JNK mitogen-activated protein kinases are critical for Ras transformation. J. Biol. Chem. 277, 31808-31817.
88. Machta, F., D. Lallemand, C. M. Pfarr and M. Yaniv (1997) Transformation by ras modifies AP1 composition and activity. Oncogene 14, 837-847.
89. Liu, G., N. Chen, A. Kaji, A. M. Bode, C. A. Ryan and Z. Dong (2001) Proteinase inhibitors I and II from potatoes block UVB-induced AP-1 activity by regulating the AP-1 protein compositional patterns in JB6 cells. Proc. Natl. Acad. Sci. USA 98, 5786-5791.
90. Agarwal, S. K., E. A. Novotny, J. S. Crabtree, J. B. Weitzmann, M. Yaniv, A. L. Burns, S. C. Chandrasekharappa, F. S. Collins, A, M. Spiegel and S. J. Marx (2003) Transcription factor JunD, deprived of menin, switches from growth suppressor to growth promoter. Proc. Natl. Acad. Sci. USA 100, 10770-10775.
91. Vasudevan, K. M., S. Gurumurthy and V. M. Rangnekar (2004) Suppression of PTEN expression by NF-kappa B prevents apoptosis. Mol. Cell Biol. 24, 1007-1021.
92. Gu, J., M. Tamura and K. Yamada (1998) Tumor suppressor PTEN inhibits integrin- and growth factor-mediated mitogen-activated protein (MAP) kinase signaling pathways. J. Cell Biol. 143, 1375-1383.
93. Persad, S., A. A. Troussard, T. R. McPhee, D. J. Mulholland and S. Dedhar (2001) Tumorsuppressor PTEN inhibits nuclear accumulation of beta-catenin and T cell/lymphoid enhancer factor 1-mediated transcriptional activation. J. Cell Biol. 153, 1161-1173.
94. Mayo, L. D., J. E. Dixon, D. L. Durden, N. K. Tonks and D. B. Donner (2002) PTEN protects p53 from Mdm2 and sensitizes cancer cells to chemotherapy. J. Biol. Chem. 277, 5484-5489.
95. Chang, C. J., D. J. Freeman and H. Wu (2004) PTEN regulates Mdm2 expression through the P1 promoter. J. Biol. Chem. 279, 29841-29848.
96. Freeman, D. J., A. G. Li, G. Wei, H.-H. Li, N. Kertesz, R. Lesche, A. D. Whale, H. Martinez-Diaz, N. Rozengurt, R. D. Cardiff, X. Liu and H. Wu (2003) PTEN tumor suppressor regulates p53 protein levels and activity through phosphatase-dependent and -independent mechanisms. Cancer Cell 3, 117-130.
97. Chung, T. W., Y. C. Lee, J. H. Ko and C. H. Kim (2003) Hepatitis B virus X protein modulates the expression of PTEN by inhibiting the function of p53, a transcriptional activator in liver cells. Cancer Res. 63, 3453-3458.
98. INK4a. Cell 88, 593-602.
99. Bandyopadhyay, D., N. Timenchenko, T. Suwa, P. J. Hornsby, J. Campisi and E. E. Medrano (2001) The human melanocyte: a model system to study the complexity of cellular aging and transformation in non-fibroblast cells. Exp. Gerontol. 36, 1265-1275.
100. Sviderskaya, E. V., S. P. Hill, T. J. Evans-Whipp, L. Chin, S. J. Orlow, D. J. Easty, S. C. Cheong, D. Beach, R. A. DePinho and D. C. Bennett (2002) P16(Ink4a) in melanocyte senescence and differentiation. J. Natl. Cancer Inst. 94, 446-454.
101. Kurihana, Y., E. Egawa, S. Kunimoto, T. Takmuchi and K. Nose (2002) Induction of p16/INK4a gene expression and expression and cellular senescence by toyocamycin. Biol. Pharm. Bull. 25, 1271-1276.
102. Ohtani, N., Z. Zebedee, T. J. Huot, J. A. Stinson, M. Sugimoto, Y. Ohashi, A. D. Sharrocks, G. Peters and E. Hara (2001) Opposing effects of Ets and Id proteins on p16INK4a expression during cellular senescence. Nature 409, 1067-1070.
103. Polsky, D., A. Z. Young, K. J. Busan and R. M. Alani (2001) The transcriptional repressor of p16/INK4a, Id1, is up-regulated in early melanomas. Cancer Res. 61, 6008-6011.
104. Bates, S., A. Philips, P. Clark, F. Scott, G. Peters, R. Ludwig and K. Voudsen (1999) p14ARF links tumour suppressors RB and p53. Nature 395, 124-125.
105. Bales, E. S., C. Dietrich, D. Bandyopadhyay, D. J. Schwahn, W. Xu, V. Didenko, P. Leiss, N. Conrad, O. Pereira-Smith, I. Orengo and E. E. Medrano (1999) High levels of expression of p27KIP and cyclin E in invasive primary malignant melanomas. J. Investig. Dermatol. 113, 1039-1046.
106. Bulavin, D. V., S. Saito, M. C. Hollander, K. Sakaguchi, C. W. Anderson, E. Appella and A. J. Fornace Jr. (1999) Phosphorylation of human p53 by p38 kinase coordinates N-terminal phosphorylation and apoptosis in response to UV radiation. EMBO J. 18, 6845-6854.
107. Heron-Milhavet, L., M. Karas, C. M. Goldsmith, B. J. Baum and D. LeRoith (2001) Insulin-like growth factor-I (IGF-1) receptor activation rescues UV-damaged cells through p38 signaling pathway. Potential role of the IGF-1 receptor in DNA repair. J. Biol. Chem. 276, 18185-18192.
108. Fan, C., J. Yang and J. F. Engelhardt (2002) Temporal pattern of NfkappaB activation influences apoptotic cell fate in a stimuli-dependent fashion. J. Cell Sci. 115, 4843-4853.
109. McNuity, S. E., N. B. Tohidian and F, L. Mayskens Jr. (2001) RelA, p50, and inhibitor of kappa B alpha are elevated in human metastatic melanoma cells and respond aberrantly to ultraviolet light B. Pigment Cell Res. 14, 456-465.
110. Huang, C., W. Ma and Z. Dong (1997) Signal transduction through atypical PKCs, but not EGF receptor, is necessary for UVC-induced AP-1 activation in immortal murine cells. Oncogene 24, 1945-1954.
111. Knebel, A., H. J. Rahmsdorf, A. Ullrich and P. Herrlich (1996) Dephosphorylation of receptor tyrosine kinases as target of regulation by radiation, oxidants or alkylating agents. EMBO J. 15, 5314-5325.
112. Coffer, P. J., B. M. Burgering, M. P. Peppelenbosch, J. L. Bos and W. Kruijer (1995) UV activation of receptor tyrosine kinase activity. Oncogene 11, 561-569.
113. Galibert, M. D., S. Carriera and C. R. Goding (2001) The Usf-1 transcription factor is a novel target for the stress-responsive p38 kinase and mediates UV-induced Tyrosinase expression. EMBO J. 20, 5022-5031.
114. Yanase, H., H. Ando, M. Horikawa, M. Wanatabe, T. Mori and N. Matsuda (2001) Possible involvement of ERK 1/2 in UVA-induced melanogenesis in cultured normal human melanocytes. Pigment Cell Res. 14, 103-109.
115. Kabuyama, Y., M. K. Homma, M. Sekimata and Y. Homma (2001) Wavelength-specific activation of MAP kinase family proteins monochromatic UV irradiation. Photochem. Photobiol. 73, 147-152.
116. Piepkorn, M. (2002) The expression of p16(INK4a), the product of the tumor suppressor genefor melanoma, is upregulated in human melanocytes by UVB irradiation. J. Am. Acad. Dermatol. 42, 741-745.
117. Medrano, E. E., S. Im, F. Yang and Z. A. Abdel-Malek (1995) Ultraviolet B light induces G1 arrest in human melanocytes by prolonged inhibition of retinoblastoma protein phosphorylation associated with long-term expression of the p21WAF-1/SDI-1/Cip-1 protein. Cancer Res. 55, 4047-4052.
118. Petrocelli, T., R. Poon, D. J. Drucker, J. M. Slingerland and C. F. Rosen (1996) UVB radiation induces p21Cip1/WAF1 and mediates G1 and S phase checkpoints. Oncogene 12, 1387-1396.
119. Chazal, M., C. Marionnet, L. Michel, K. Mollier, J. E. Dazard, V. Della Valle, C. J. Larsen, M. P. Gras and N. Basset-Seguin (2002) P16(INK4a) is implicated in both the immediate and adaptive response of human keratinocytes to UVB radiation. Oncogene 21, 2652-2661.
120. Milligan, A., B. G. Gabrielli, J. M. Clark, N. K. Hayward and K. A. Ellem (1998) Involvement of p16INK4a in cell cycle delays after low dose UV irradiation. Mutat. Res. 422, 43-53.
121. Poon, R. Y., H. Toyoshima and T. Hunter (1995) Redistribution of the CDK inhibitor p27 between different cyclin. CDK complexes in the mouse fibroblast cell cycle and in cells arrested with lovastatin or ultraviolet irradiation. Mol. Biol. Cell 6, 1197-1213.
122. Matsumoto-Taniura, N., K. Matsumoto and T. Nakamura (1999) Prostaglandin production in mouse mammary tumour cells confers invasive growth potential by inducing hepatocyte growth factor in stromal fibroblasts. Br. J. Cancer 81, 194-202.
123. Maas-Szabowski, N. and N. E. Fusenig (1996) Interleukin-1-induced growth factor expression in postmitotic and resting fibroblasts. J. Investig. Dermatol. 107, 849-855.
124. Pandey, A., H. Shao, R. M. Marks, P. J. Polverini and V. M. Dixit (1995) Role of B61, the ligand for the Eck receptor tyrosine kinase, in TNF-alpha-induced angiogenesis. Science 268, 567-569.
125. Castagnino, P., M. W. Lorenzi, J. Yeh, D. Breckenridge, H. Sakata, B. Muntz, S. Werner and D. P. Bottaro (2000) Neu differentiation factor/heregulin induction by hepatocyte and keratinocyte growth factors. Oncogene 19, 640-648.
126. Tsatmali, M., J. Ancana, J. Yukitake and A. J. Thody (2000) Skin POMV peptides: their actions at the human MC-1 receptor and roles in the tanning response. Pigment Cell Res. 13 (Suppl. 8), 125-129.
127. Dissanayake, N. S. and R. S. Mason (1998) Modulation of skin cell functions by transforming growth factor-beta1 and ACTH after ultraviolet irradiation. J. Endocrinol. 159, 153-163.
128. Wu, M., T. J. Hemesath, C. M. Takemoto, M. A. Horstmann, A. G. Wells, E. R. Price, D. Z. Fisher and D. E. Fisher (2000) c-KIT triggers dual phosphorylation, which couple activation and degradation of the essential melanocyte Mi. Genes Dev. 14, 301-312.
129. Jiang, J., S. D. Sharma, S. Nakamura, J. Y. Lai, J. L. Fink, V. J. Hruby and M. E. Hadley (1995) The melanotropic peptide, (Nle4, D-Phe7) alpha-MSH, stimulates human melanoma tyrosinase activity and inhibits cell proliferation. Pigment Cell Res. 8, 314-323.
130. Sviderskaya, E. V., S. P. Hill, D. Balachander, G. S. Barsh and D. C. Bennett (2001) Agouti signaling protein and other factors modulating differentiation and proliferation of immortal melanoblasts. Dev. Dyn. 221, 373-379.
131. Smalley, K. S. and T. G. Eisen (2002) Differentiation of human melanoma cells through p38 MAP kinase is associated with decreased retinoblastoma protein phosphorylation and cell cycle arrest. Melanoma Res. 12, 187-192.
132. Hedley, S. J., C. Layton, M. Heaton, K. H. Chakrabarty, R. A. Dawson, D. J. Gawkrodger and S. MacNeil (2002) Fibroblasts play a regulatory role in the control of pigmentation in reconstructed human skin from skin types I and II. Pigment Cell Res. 15, 49-56.
133. Hedley, S. J., D. J. Gawkrodger, A. P. Weetman and S. MacNeil (1998) alpha-MSH and melanogenesis in normal human adult melanocytes. Pigment Cell Res. 11, 45-56.
134. De Luca, M., W. Siegrist, S. Bondanza, M. Mathor, M. Cancedda and A. N. Eberle (1993) Alpha melanocyte stimulating hormone (alpha MSH) stimulates normal human melanocytes growth by binding to high-affinity receptors. J. Cell Sci. 105, 1079-1084.
135. Kadekaro, A. L., H. Kanto, R. Kavanagh and Z. A. Abdel-Malek (2003) Significance of the melanocoitin 1 receptor in regulating human melanocyte pigmentation, proliferation, and survival. Ann. N.Y. Acad. Sci. 994, 359-365.
136. Imokawa, G., M. Miyagishi and T. Tada (1995) Endothelin-1 as a new melanogen: coordinated expression of its gene and the tyrosinase gene in UVB-exposed human epidermis. J. Investig. Dermatol. 105, 32-37.
137. Jamal, S. and P. J. Schneider (2002) UV-induction of keratinocyte endothelin-1 downregulates E-cadherin in melanocytes and melanoma cells. J. Clin. Investig. 110, 443-452.
138. Tronnier, M., P. Rudolph, T. Köser, B. Raasch and J. Brinckmann (1997) One single erythemagenic UV irradiation is more effective in increasing the proliferative activity of melanocytes in melanocytic naevi compared with fractionally applied high doses. Br. J. Dermatol. 137, 534-539.
139. An, H. T., J. Y. Yoo, M. K. Lee, M. H. Shin, G. E. Rhie, J. Y. Seo, H. Chung, C. Eun and H. Cho (2001) Single dose radiation is more effective for the UV-induced activation and proliferation of melanocytes than fractionated dose radiation. Photodermatol. Photoimmunol. Photomed. 17, 266-271.
140. WAFI/Cip1 reflecting proliferation and repair activity in UV-irradiated melanocytic nevi. Hum Pathol. 29, 1480-1486.
141. Otsuka, T., H. Takayama, R. Sharp, G. Celli, W. J. LaRochelle, D. P. Bottaro, N. Ellmore, W. Vieira, J. W. Owens, M. Anver and G. Merlino (1998) c-Met autocrine activation induces development of malignant melanoma and acquisition of the metastatic phenotype. Cancer Res. 58, 5157-5167.
142. Kato, M., M. Takahashi, A. A. Akhand, W. Lui, Y. Dai, S. Shimizu, T. Iwamoto, H. Suzuki and I. Nakashima (1998) Transgenic mouse model for skin malignant melanoma. Oncogene 17, 1885-1888.
143. Kunisada, T., S. Z. Lu, H. Yoshida, S. Nishikawa, M. Mizoguchi, S. Hayashi, L. Ryrrell, D. A. Williams, X. Wang and B. J. Longley (1998) Murine cutaneous mastcytosis and epidermal melanocytosis induced by keratinocyte expression of transgenic stem cell factor. J. Exp. Med. 187, 1565-1573.
144. Chin, L., J. Pomerantz, D. Polsky, M. Jacobson, C. Cohen, C. Cordon-Cardo, J. W. Horner and R. A. DePinho (1997) Cooperative effects of Ink4a and ras in melanoma susceptibility in vivo. Genes Dev. 11, 2822-2843.
145. Bardeesy, N., B. C. Bastian, A. Hezel, D. Pinkel, R. A. DePinho and L. Chin (2001) Dual inactivation of RB and p53 pathways in RAS-induced melanomas. Mol. Cell Biol. 21, 2144-2153.
146. You, M. J., D. H. Castrillon, B. C. Bastian, R. C. O'Hagan, M. W. Bodenberg, R. Parsons, L. Chin and R. A. DePinho (2002) Genetic analysis of Pten and Ink4a/Arf interactions in the suppression of tumorigenesis in mice. Proc. Natl. Acad. Sci. USA 99, 1455-1460.
147. Bradl, M., A. Klein-Szanto, S. Porter and B. Mintz (1991) Malignant melanoma in transgenic mice. Proc. Natl. Acad. Sci. USA 88, 164-168.
148. Kato, M., W. Lui, A. A. Akland, K. Hossain, K. Takeda, M. Takashashi and I. Nakashima (2000) Ultraviolet radiation induces both full activation of Ret kinase and malignant melanocyte tumor promotion in RFP-RET-transgenic mice. J. Investig. Dermatol. 115, 1157-1158.
149. Noonan, F. P., T. Otsuka, S. Bang, M. R. Anver and G. Merlino (2000) Accelerated ultraviolet radiation-induced carcinogenesis in hepatocyte growth factor/scatter factor transgenic mice. Cancer Res. 60, 3738-3743.
150. Noonan, F. P., J. A. Recio, H. Takayama, P. Durray, M. R. Anver, W. L. Rush, E. C. DeFabo and G. Merlino (2001) Neonatal sunburn and melanoma in mice. Nature 413, 271-272.
151. Recio, J. A., F. P. Noonan, H. Takayama, M. R. Anver, P. Duray, W. L. Rush, G. Lindner, E. C. De Fabo, R. A. DePinho and G. Merlino (2002) Ink4a/Arf deficiency promotes ultraviolet radiation-induced melanomagenesis. Cancer Res. 62, 6724-6730.
152. Powell, M. B., P. R. Gause, P. Hyman, J. Gregus, M. Lluria-Prevat, R. Nagle and G. T. Bowden (1999) Induction of melanoma in TPras transgenic mice. Carcinogenesis 20, 1747-1753.
153. Kannan, K., N. E. Sharpless, J. Xu, R. O'Hagan, M. Bosenberg and L. Chin (2003) Components of the Rb pathway are critical targets of UV mutagenesis in a murine melanoma model. Proc. Natl. Acad. Sci. USA 100, 1221-1225.
154. Sotillo, R., J. F. Garcia, S. Onega, J. Martin, P. Dubus, M. Barbacid and M. Malumbres (2001) Invasive melanoma in Cdk4-targeted mice. Proc. Natl. Acad. Sci. USA 98, 13312-13317.
155. Krimpenfort, P., K. C. Quon, W. J. Mooi, A. Loonstra and A. Berns (2001) Loss of p16Ink4a confers susceptibility to metastatic melanoma in mice. Nature 413(6851), 83-86.
156. Yang, J., J. Luan, Y. Yu, C. Li, R. A. De Pinho, L. Chin and A. Richmond (2001) Induction of melanoma in murine macrophage inflammatory protein 2 transgenic mice heterozygous for inhibitor of kinase/alternate reading frame. Cancer Res. 61, 8150-8157.
157. Keisall, S. R. and B. Mintz (1998) Metastatic cutaneous melanoma promoted by ultraviolet radiation in mice with transgene-initiated low melanoma susceptibility. Cancer Res. 58, 4061-4065.
158. Nishimura, N., V. E. Reeve, H. Nishimura, M. Satoh and C. Tohyama (2000) Cutaneous metallothionein induction by ultraviolet B irradiation in interleukin-6 null mice. J. Investig. Dermatol. 114, 343-348.
159. Oguro, T. and T. Yoshida (2001) Effect of ultraviolet A on omithine decarboxylase and metallothionein gene expression in mouse skin. Photodermatol. Photoimmunol. Photomed. 17, 71-78.
160. Anstey, A., R. Marks, C. Long, H. Navabi, A. Pearse, D. Wynford-Thomas and B. Jasani (1996) In vivo photoinduction of metallothionein in human skin by ultraviolet irradiation. J. Pathol. 178, 84-88.
161. Halaban, R., J. S. Rubin, Y. Funasaka, M. Cobb, T. Boulton, D. Faletto, E. Rosen, A. Chan, K. Yoko and W. White (1992) Met and hepatocyte growth factor/scatter factor signal transduction in normal melanocytes and melanoma cells. Oncogene 7, 2195-2206.
162. Hirobe, T., M. Osawa and S. Nishikawa (2004) Hepatocyte growth factor controls the proliferation of cultured epidermal melanoblasts and melanocytes from newborn mice. Pigment Cell Res. 17, 51-61.
163. Recio, J. A., F. P. Noonan, H. Takayama, M. R. Anver, P. Duray, W. L. Rush, G. Lindner, E. C. De Fabo, R. A. DePinho and G. Merlino (2002) Ink4a/Arf deficiency promotes ultraviolet radiation-induced melanomagenesis. Cancer Res. 62, 6724-6730.
164. De Fabo, E. C., F. P. Noonan, T. Fears and G. Merlino (2004) Ultraviolet B but not ultraviolet A radiation initiates melanoma. Cancer Res. 64, 6372-6376.
165. Yamazaki, F., H. Okamoto, H. Miyauchi-Hashimoto, Y. Matsumura, T. Itoh, K. Tanaka, T. Kunisada and T. Horio (2004) XPA gene-deficient SCF-transgenic mice with epidermal melanin are resistant to UV-induced carcinogenesis. J. Investig. Dermatol. 123, 220-228.