• 1
    Gates, F. L. (1928) On nuclear derivatives and lethal action of ultraviolet light. Science 68, 479480.
  • 2
    Hollaender, A. and C. W. Emmons (1941) Wavelength dependence of mutation production in the ultraviolet with special emphasis on fungi. Cold Spring Harb. Symp. Quant. Biol. 9, 179186.
  • 3
    Goldblatt, C., T. M. Lenton and A. J. Watson (2006) Bistability of atmospheric oxygen and the Great Oxidation. Nature 443, 683686.
  • 4
    Kojo, K., C. T. Jansen, P. Nybom, L. Huurto, J. Laihia, T. Ilus and A. Auvinen (2006) Population exposure to ultraviolet radiation in Finland 1920–1995: Exposure trends and a time-series analysis of exposure and cutaneous melanoma incidence. Environ. Res. 101, 123131.
  • 5
    Wassberg, C., M. Thorn, A. M. Johansson, R. Bergstrom, B. Berne and U. Ringborg (2001) Increasing incidence rates of squamous cell carcinoma of the skin in Sweden. Acta Derm. Venereol. 81, 268272.
  • 6
    De Vries, E., J. W. Coebergh and H. Van Der Rhee (2006) Trends, causes, approach and consequences related to the skin-cancer epidemic in the Netherlands and Europe. Ned. Tijdschr. Geneeskd. 150, 11081115.
  • 7
    Cokkinides, V., M. Weinstock, K. Glanz, J. Albano, E. Ward and M. Thun (2006) Trends in sunburns, sun protection practices, and attitudes toward sun exposure protection and tanning among US adolescents, 1998–2004. Pediatrics 118, 853864.
  • 8
    Housman, T. S., S. R. Feldman, P. M. Williford, A. B. Fleischer Jr, N. D. Goldman, J. M. Acostamadiedo and G. J. Chen (2003) Skin cancer is among the most costly of all cancers to treat for the Medicare population. J. Am. Acad. Dermatol. 48, 425429.
  • 9
    Ramsay, H. M., A. A. Fryer, C. M. Hawley, A. G. Smith and P. N. Harden (2002) Non-melanoma skin cancer risk in the Queensland renal transplant population. Br. J. Dermatol. 147, 950956.
  • 10
    Hartevelt, M. M., J. N. Bavinck, A. M. Kootte, B. J. Vermeer and J. P. Vandenbroucke (1990) Incidence of skin cancer after renal transplantation in The Netherlands. Transplantation 49, 506509.
  • 11
    Gupta, S. and H. Mukhtar (2001) Chemoprevention of skin cancer through natural agents. Skin Pharmacol. Appl. Skin Physiol. 14, 373385.
  • 12
    Saleem, M., F. Afaq, V. M. Adhami and H. Mukhtar (2004) Lupeol modulates NF-kappaB and PI3K/Akt pathways and inhibits skin cancer in CD-1 mice. Oncogene 23, 52035214.
  • 13
    Cooke, A. and B. E. Johnson (1978) Dose response, wavelength dependence and rate of excision of ultraviolet radiation-induced pyrimidine dimers in mouse skin DNA. Biochim. Biophys. Acta 517, 2430.
  • 14
    Johnson, B. E. (1978) Formation of thymine containing dimers in skin exposed to ultraviolet radiation. Bull. Cancer 65, 283297.
  • 15
    De Gruijl, F. R., H. J. Sterenborg, P. D. Forbes, R. E. Davies, C. Cole, G. Kelfkens, H. Van Weelden, H. Slaper and J. C. Van Der Leun (1993) Wavelength dependence of skin cancer induction by ultraviolet irradiation of albino hairless mice. Cancer Res. 53, 5360.
  • 16
    De Gruijl, F. R. and J. C. Van Der Leun (1994) Estimate of the wavelength dependency of ultraviolet carcinogenesis in humans and its relevance to the risk assessment of a stratospheric ozone depletion. Health Phys. 67, 319325.
  • 17
    Longstreth, J., F. R. De Gruijl, M. L. Kripke, S. Abseck, F. Arnold, H. Slaper, G. Velders, Y. Takizawa and J. C. Van Der Leun (1998) Health risks. J. Photochem. Photobiol. B, Biol. 46, 2039.
  • 18
    Anders, A., H. J. Altheide, M. Knalmann and H. Tronnier (1995) Action spectrum for erythema in humans investigated with dye lasers. Photochem. Photobiol. 61, 200205.
  • 19
    De Gruijl, F. R. (1999) Skin cancer and solar UV radiation. Eur. J. Cancer 35, 20032009.
  • 20
    Freeman, S. E., H. Hacham, R. W. Gange, D. J. Maytum, J. C. Sutherland and B. M. Sutherland (1989) Wavelength dependence of pyrimidine dimer formation in DNA of human skin irradiated in situ with ultraviolet light. Proc. Natl Acad. Sci. USA 86, 56055609.
  • 21
    Jans, J., W. Schul, Y. G. Sert, Y. Rijksen, H. Rebel, A. P. Eker, S. Nakajima, H. Van Steeg, F. R. De Gruijl, A. Yasui, J. H. Hoeijmakers and G. T. Van Der Horst (2005) Powerful skin cancer protection by a CPD-photolyase transgene. Curr. Biol. 15, 105115.
  • 22
    Brash, D. E. and W. A. Haseltine (1982) UV-induced “mutation hotspots” occur at damage “hotspots.” Nature 298, 189192.
  • 23
    Brash, D. E., J. A. Rudolph, J. A. Simon, A. Lin, G. J. McKenna, H. P. Baden, A. J. Halperin and J. Ponten (1991) A role for sunlight in skin cancer: UV-induced p53 mutations in squamous cell carcinoma. Proc. Natl Acad. Sci. USA 88, 1012410128.
  • 24
    Dumaz, N., H. J. Van Kranen, A. De Vries, R. J. Berg, P. W. Wester, C. F. Van Kreijl, A. Sarasin, L. Daya-Grosjean and F. R. De Gruijl (1997) The role of UV-B light in skin carcinogenesis through the analysis of p53 mutations in squamous cell carcinomas of hairless mice. Carcinogenesis 18, 897904.
  • 25
    Ruven, H. J., R. J. Berg, C. M. Seelen, J. A. Dekkers, P. H. Lohman, L. H. Mullenders and A. A. Van Zeeland (1993) Ultraviolet-induced cyclobutane pyrimidine dimers are selectively removed from transcriptionally active genes in the epidermis of the hairless mouse. Cancer Res. 53, 16421645.
  • 26
    You, Y. H., P. E. Szabo and G. P. Pfeifer (2000) Cyclobutane pyrimidine dimers form preferentially at the major p53 mutational hotspot in UVB-induced mouse skin tumors. Carcinogenesis 21, 21132117.
  • 27
    Kielbassa, C., L. Roza and B. Epe (1997) Wavelength dependence of oxidative DNA damage induced by UV and visible light. Carcinogenesis 18, 811816.
  • 28
    Van Kranen, H. J., A. De Laat, J. Van De Ven, P. W. Wester, A. De Vries, R. J. Berg, C. F. Van Kreijl and F. R. De Gruijl (1997) Low incidence of p53 mutations in UVA (365-nm)-induced skin tumors in hairless mice. Cancer Res. 57, 12381240.
  • 29
    Li, G., V. C. Ho, K. Berean and V. A. Tron (1995) Ultraviolet radiation induction of squamous cell carcinomas in p53 transgenic mice. Cancer Res. 55, 20702074.
  • 30
    Jiang, W., H. N. Ananthaswamy, H. K. Muller and M. L. Kripke (1999) p53 protects against skin cancer induction by UV-B radiation. Oncogene 18, 42474253.
  • 31
    Van Kranen, H. J., A. Westerman, R. J. Berg, N. Kram, C. F. Van Kreijl, P. W. Wester and F. R. De Gruijl (2005) Dose-dependent effects of UVB-induced skin carcinogenesis in hairless p53 knockout mice. Mutat. Res. 571, 8190.
  • 32
    Wijnhoven, S. W., E. N. Speksnijder, X. Liu, E. Zwart, C. T. Van Oostrom, R. B. Beems, E. M. Hoogervorst, M. M. Schaap, L. D. Attardi, T. Jacks, H. Van Steeg, J. Jonkers and A. De Vries (2007) Dominant-negative but not gain-of-function effects of a p53.R270H mutation in mouse epithelium tissue after DNA damage. Cancer Res. 67, 46484656.
  • 33
    Berg, R. J., H. Rebel, G. T. Van Der Horst, H. J. Van Kranen, L. H. Mullenders, W. A. Van Vloten and F. R. De Gruijl (2000) Impact of global genome repair versus transcription-coupled repair on ultraviolet carcinogenesis in hairless mice. Cancer Res. 60, 28582863.
  • 34
    Berg, R. J., A. De Vries, H. Van Steeg and F. R. De Gruijl (1997) Relative susceptibilities of XPA knockout mice and their heterozygous and wild-type littermates to UVB-induced skin cancer. Cancer Res. 57, 581584.
  • 35
    Van Zeeland, A. A., M. P. Vreeswijk, F. R. De Gruijl, H. J. Van Kranen, H. Vrieling and L. H. Mullenders (2005) Transcription-coupled repair: Impact on UV-induced mutagenesis in cultured rodent cells and mouse skin tumors. Mutat. Res. 577, 170178.
  • 36
    De Vries, A., R. J. Berg, S. Wijnhoven, A. Westerman, P. W. Wester, C. F. Van Kreijl, P. J. Capel, F. R. De Gruijl, H. J. Van Kranen and H. Van Steeg (1998) XPA-deficiency in hairless mice causes a shift in skin tumor types and mutational target genes after exposure to low doses of U.V.B. Oncogene 16, 22052212.
  • 37
    Pazzaglia, S., M. Mancuso, B. Primerano, S. Rebessi, G. Biozzi, V. Covelli and A. Saran (2001) Analysis of c-Ha-ras gene mutations in skin tumors induced in carcinogenesis-susceptible and carcinogenesis-resistant mice by different two-stage protocols or tumor promoter alone. Mol. Carcinog. 30, 111118.
  • 38
    Abrahams, P. J., A. A. Van Der Kleij, R. Schouten and A. J. Van Der Eb (1988) Absence of induction of enhanced reactivation of herpes simplex virus in cells from xeroderma pigmentosum patients without skin cancer. Cancer Res. 48, 60546057.
  • 39
    Terleth, C., T. Van Laar, R. Schouten, H. Van Steeg, H. Hodemaekers, T. Wormhoudt, P. D. Cornelissen-Steijger, P. J. Abrahams and A. J. Van Der Eb (1997) A lack of radiation-induced ornithine decarboxylase activity prevents enhanced reactivation of herpes simplex virus and is linked to non-cancer proneness in xeroderma pigmentosum patients. Cancer Res. 57, 43844392.
  • 40
    Rebel, H., H. Van Steeg, R. B. Beems, R. Schouten, F. R. De Gruijl and C. Terleth (2002) Suppression of UV carcinogenesis by difluoromethylornithine in nucleotide excision repair-deficient Xpa knockout mice. Cancer Res. 62, 13381342.
  • 41
    Fischer, S. M., H. H. Lo, G. B. Gordon, K. Seibert, G. Kelloff, R. A. Lubet and C. J. Conti (1999) Chemopreventive activity of celecoxib, a specific cyclooxygenase-2 inhibitor, and indomethacin against ultraviolet light-induced skin carcinogenesis. Mol. Carcinog. 25, 231240.
  • 42
    Pentland, A. P., J. W. Schoggins, G. A. Scott, K. N. Khan and R. Han (1999) Reduction of UV-induced skin tumors in hairless mice by selective COX-2 inhibition. Carcinogenesis 20, 19391944.
  • 43
    Berg, R. J., H. J. Van Kranen, H. G. Rebel, A. De Vries, W. A. Van Vloten, C. F. Van Kreijl, J. C. Van Der Leun and F. R. De Gruijl (1996) Early p53 alterations in mouse skin carcinogenesis by UVB radiation: Immunohistochemical detection of mutant p53 protein in clusters of preneoplastic epidermal cells. Proc. Natl Acad. Sci. USA 93, 274278.
  • 44
    Jonason, A. S., S. Kunala, G. J. Price, R. J. Restifo, H. M. Spinelli, J. A. Persing, D. J. Leffell, R. E. Tarone and D. E. Brash (1996) Frequent clones of p53-mutated keratinocytes in normal human skin. Proc. Natl Acad. Sci. USA 93, 1402514029.
  • 45
    Rebel, H., L. O. Mosnier, R. J. Berg, A. Westerman-de Vries, H. Van Steeg, H. J. Van Kranen and F. R. De Gruijl (2001) Early p53-positive foci as indicators of tumor risk in ultraviolet-exposed hairless mice: Kinetics of induction, effects of DNA repair deficiency, and p53 heterozygosity. Cancer Res. 61, 977983.
  • 46
    Ling, G., A. Persson, B. Berne, M. Uhlen, J. Lundeberg and F. Ponten (2001) Persistent p53 mutations in single cells from normal human skin. Am. J. Pathol. 159, 12471253.
  • 47
    Zhang, W., E. Remenyik, D. Zelterman, D. E. Brash and N. M. Wikonkal (2001) Escaping the stem cell compartment: Sustained UVB exposure allows p53-mutant keratinocytes to colonize adjacent epidermal proliferating units without incurring additional mutations. Proc. Natl Acad. Sci. USA 98, 1394813953.
  • 48
    Backvall, H., S. Stromberg, A. Gustafsson, A. Asplund, A. Sivertsson, J. Lundeberg and F. Ponten (2004) Mutation spectra of epidermal p53 clones adjacent to basal cell carcinoma and squamous cell carcinoma. Exp. Dermatol. 13, 643650.
  • 49
    Einspahr, J. G., D. S. Alberts, J. A. Warneke, P. Bozzo, J. Basye, T. M. Grogan, M. A. Nelson and G. T. Bowden (1999) Relationship of p53 mutations to epidermal cell proliferation and apoptosis in human UV-induced skin carcinogenesis. Neoplasia 1, 468475.
  • 50
    Ahmadian, A., Z. Ren, C. Williams, F. Ponten, J. Odeberg, J. Ponten, M. Uhlen and J. Lundeberg (1998) Genetic instability in the 9q22.3 region is a late event in the development of squamous cell carcinomas. Oncogene 17, 18371843.
  • 51
    Ananstaswamy, H. N., A. Fourtanier, R. L. Evans, S. Tison, S. Medaisko, S. E. Ullrich and M. L. Kripke (1998) P53 mutations in hairless SKH-hr1 mouse skin tumors induced by a solar simulator. Photochem. Photobiol. 67, 227232.
  • 52
    Rebel, H., N. Kram, A. Westerman, S. Banus, H. J. Van Kranen and F. R. De Gruijl (2005) Relationship between UV-induced mutant p53 patches and skin tumours, analysed by mutation spectra and by induction kinetics in various DNA-repair-deficient mice. Carcinogenesis 26, 21232130.
  • 53
    Kramata, P., Y. P. Lu, Y. R. Lou, R. N. Singh, S. M. Kwon and A. H. Conney (2005) Patches of mutant p53-immunoreactive epidermal cells induced by chronic UVB irradiation harbor the same p53 mutations as squamous cell carcinomas in the skin of hairless SKH-1 mice. Cancer Res. 65, 35773585.
  • 54
    De Gruijl, F. R., J. B. Van Der Meer and J. C. Van Der Leun (1983) Dose-time dependency of tumor formation by chronic UV exposure. Photochem. Photobiol. 37, 5362.
  • 55
    De Gruijl, F. R. and J. C. Van Der Leun (1991) Development of skin tumors in hairless mice after discontinuation of ultraviolet irradiation. Cancer Res. 51, 979984.
  • 56
    Remenyik, E., N. M. Wikonkal, W. Zhang, V. Paliwal and D. E. Brash (2003) Antigen-specific immunity does not mediate acute regression of UVB-induced p53-mutant clones. Oncogene 22, 63696376.
  • 57
    Stout, G. J., D. Westdijk, D. M. Calkhoven, O. Pijper, C. M. Backendorf, R. Willemze, L. H. Mullenders and F. R. De Gruijl (2005) Epidermal transit of replication-arrested, undifferentiated keratinocytes in UV-exposed XPC mice: An alternative to in situ apoptosis. Proc. Natl Acad. Sci. USA 102, 1898018985.
  • 58
    Potten, C. S. (1975) Epidermal transit times. Br. J. Dermatol. 93, 649658.
  • 59
    Faurschou, A., M. Haedersdal, T. Poulsen and H. C. Wulf (2007) Squamous cell carcinoma induced by ultraviolet radiation originates from cells of the hair follicle in mice. Exp. Dermatol. 16, 485489.
  • 60
    Hedelund, L., M. Haedersdal, H. Egekvist, M. Heidenheim, H. C. Wulf and T. Poulsen (2004) CO2 laser resurfacing and photocarcinogenesis: An experimental study. Lasers Surg. Med. 35, 5861.
  • 61
    Morris, R. J., K. A. Tryson and K. Q. Wu (2000) Evidence that the epidermal targets of carcinogen action are found in the interfollicular epidermis of infundibulum as well as in the hair follicles. Cancer Res. 60, 226229.
  • 62
    Mitchell, D. L., R. Greinert, F. R. De Gruijl, K. L. Guikers, E. W. Breitbart, M. Byrom, M. M. Gallmeier, M. G. Lowery and B. Volkmer (1999) Effects of chronic low-dose ultraviolet B radiation on DNA damage and repair in mouse skin. Cancer Res. 59, 28752884.
  • 63
    Nijhof, J. G., C. Van Pelt, A. A. Mulder, D. L. Mitchell, L. H. Mullenders and F. R. De Gruijl (2007) Epidermal stem and progenitor cells in murine epidermis accumulate UV damage despite NER proficiency. Carcinogenesis 28, 792800.
  • 64
    Nijhof, J. G., A. A. Mulder, E. N. Speksnijder, E. M. Hoogervorst, L. H. Mullenders and F. R. De Gruijl (2007) Growth stimulation of UV-induced DNA damage retaining epidermal basal cells gives rise to clusters of p53 overexpressing cells. DNA Repair 6, 16421650.
  • 65
    El-Abaseri, T. B., S. Putta and L. A. Hansen (2006) Ultraviolet irradiation induces keratinocyte proliferation and epidermal hyperplasia through the activation of the epidermal growth factor receptor. Carcinogenesis 27, 225231.
  • 66
    Husain, Z., Q. M. Yang and D. K. Biswas (1990) cHa-ras proto-oncogene. Amplification and overexpression in UV-B-induced mouse skin papillomas and carcinomas. Arch. Dermatol. 126, 324330.
  • 67
    De Gruijl, F. R. and J. C. Van Der Leun (1982) Effect of chronic UV exposure on epidermal transmission in mice. Photochem. Photobiol. 36, 433438.
  • 68
    Berton, T. R., D. L. Mitchell, S. M. Fischer and M. F. Locniskar (1997) Epidermal proliferation but not quantity of DNA photodamage is correlated with UV-induced mouse skin carcinogenesis. J. Invest. Dermatol. 109, 340347.