Solar radiation induced skin damage: Review of protective and preventive options

International Journal of Radiation Biology

Volumn 86, Issue 12, 2010, Pages 999-1030

  Svobodová, Alena ^a   Vostálová, Jitka ^a  

^a PALACKÝ UNIVERSITY   (Czech Republic)

Author keywords

antioxidant system; DNA repair; exogenous protection; melanin synthesis; skin; solar radiation

Indexed keywords

CATALASE; GLUTATHIONE; GLUTATHIONE PEROXIDASE; GLUTATHIONE REDUCTASE; GLUTATHIONE TRANSFERASE; HEME OXYGENASE; MELANIN; POLYPHENOL; SUNSCREEN; THIOREDOXIN; THIOREDOXIN REDUCTASE;

ANATOMY; APOPTOSIS; AWARENESS; CELL DEATH; CELL PROLIFERATION; CELL SURVIVAL; DNA DAMAGE; DNA REPAIR; GENE EXPRESSION; HUMAN; MEDICAL RESEARCH; MELANOGENESIS; PHOTOAGING; PRIORITY JOURNAL; PROTECTIVE CLOTHING; REVIEW; SKIN DEFECT; SKIN PIGMENTATION; SKIN PROTECTION; SKINFOLD; SOLAR RADIATION; SUN EXPOSURE; SUNLIGHT; ULTRAVIOLET RADIATION;

EID: 78649751072     PISSN: 09553002     EISSN: 13623095     Source Type: Journal    
DOI: 10.3109/09553002.2010.501842     Document Type: Review

References (256)

1. Aburatani H, Hippo Y, Ishida T, Takashima R, Matsuba C, Kodama T, Takao M, Yasui A, Yamamoto K, Asano M. 1997. Cloning and characterization of mammalian 8-hydro-xy-guanine-specific DNA glycosylase/apurinic, apyrimidinic lyase, a functional mutM homologue. Cancer Research 57:2151-2156.
2. Adhami VM, Syed DN, Khan N, Afaq F. 2008. Phytochemicals for prevention of solar ultraviolet radiation-induced damages. Photochemistry and Photobiology 84:489-500.
3. Afaq F, Mukhtar H. 2001. Effects ofsolar radiation on cutaneous detoxification pathways. Journal ofPhotochemistry and Photo-biology B Biology 63:61-69.
4. Afaq F, Malik A, Syed D, Maes D, Matsui MS, Mukhtar H. 2005a. Pomegranate fruit extract modulates UV-B-mediated phosphorylation of mitogen-activated protein kinases and activation of nuclear factor kappa B in normal human epidermal keratinocytes paragraph sign. Photochemistry and Photobiology 81:38-45.
5. Afaq F, Zaid MA, Khan N, Dreher M, Mukhtar H. 2009. Protective effect of pomegranate-derived products on UVB-mediated damage in human reconstituted skin. Experimental Dermatology 18:553-561.
6. Afaq F, Adhami VM, Mukhtar H. 2005b. Photochemoprevention of ultraviolet B signaling and photocarcinogenesis. Mutation Research 571:153-173.
7. Akerboom TP, Bilzer M, Sies H. 1982. The relationship ofbiliary glutathione disulfide efflux and intracellular glutathione disulfide content in perfused rat liver. Journal of Biological Chemistry 257:4248-4252.
8. Allanson M, Reeve VE. 2005. Ultraviolet A (320-400 nm) modulation of ultraviolet B (290-320 nm)-induced immune suppression is mediated by carbon monoxide. Journal of Investigative Dermatology 124:644-650.
9. Alonso-Lebrero JL, Doḿinguez-Jiménez C, Tejedor R, Brieva A, Pivel JP. 2003. Photoprotective properties of a hydrophilic extract of the fern Polypodium leucotomos on human skin cells. Journal of Photochemistry and Photobiology B Biology 70:31-37.
10. Aricioglu A, Bozkurt M, Balabanli B, Kilinç M, Nazaroglu NK, Türközkan N. 2001. Changes in zinc levels and superoxide dismutase activities in the skin ofacute, ultraviolet-B-irradiated mice after treatment with Ginkgo biloba extract. Biological Trace Element Research 80:175-179.
11. Armeni T, Damiani E, Battino M, Greci L, Principato G. 2004. Lack of in vitro protection by a common sunscreen ingredient on UVA-induced cytotoxicity in keratinocytes. Toxicology 203:165-178.
12. Athar M, Back JH, Tang X, Kim KK, Kopelovich L, Bickers DR, Kim AL. 2007. Resveratrol: A review ofpreclinical studies for human cancer prevention. Toxicology and Applied Pharma-cology 224:274-283.
13. Autier P, Boniol M, Doré JF. 2007. Sunscreen use and increased duration of intentional sun exposure: Still a burning issue. International Journal ofCancer 121:1-5.
14. Aziz MH, Afaq F, Ahmad N. 2005. Prevention of ultraviolet-B radiation damage by resveratrol in mouse skin is mediated via modulation in survivin. Photochemistry and Photobiology 81:25-31.
15. Balajee AS, Bohr VA. 2000. Genomic heterogeneity ofnucleotide excision repair. Gene 250:15-30.
16. Baron ED, Kirkland BE, Domingo DS. 2008. Advances in photoprotection. Dermatology Nursing 20:265-272.
17. BartekJ, Lukas J. 2001. Mammalian G1-and S-phase checkpoints in response to DNA damage. Current Opinion in Cell Biology 13:738-747.
18. Basu-Modak S, Gordon MJ, Dobson LH, Spencer JP, Rice-Evans C, Tyrrell RM. 2003. Epicatechin and its methylated metabolite attenuate UVA-induced oxidative damage to hu-man skin fibroblasts. Free Radical Biology and Medicine 35:910-921.
19. Basu-Modak S, Luscher P, Tyrrell RM. 1996. Lipid metabolite involvement in the activation of the human heme oxygenase-1 gene. Free Radical Biology and Medicine 20:887-897.
20. Baumann L. 2007. Skin ageing and its treatment. Journal of Pathology 211:241-251.
21. Beyer TA, Auf dem Keller U, Braun S, Schäfer M, Werner S. 2007. Roles and mechanisms of action of the Nrf2 transcrip-tion factor in skin morphogenesis, wound repair and skin cancer. Cell Death and Differentiation 14:1250-1254.
22. Bito T, Ueda M, Nagano T, Fujii S, Ichihashi M. 1995. Reduction of ultraviolet-induced skin cancer in mice by topical applica-tion of DNA excision repair enzymes. Photodermatology, Photoimmunology and Photomedicine 11:9-13.
23. Böhm F, Edge R, Lange L, Truscott TG. 1998. Enhanced protection of human cells against ultraviolet light by antiox-idant combinations involving dietary carotenoids. Journal of Photochemistry and Photobiology B Biology 44:211-215.
24. Boiteux S, RadicellaJP. 2000. Thehuman OGG1 gene: Structure, functions, and its implication in the process ofcarcinogenesis. Archives of Biochemistry and Biophysics 377:1-8.
25. Brigelius-Flohé R. 2006. Glutathione peroxidases and redox-regulated transcription factors. Journal of Biological Chemistry 387:1329-1335.
26. Brown DA. 2001. Skin pigmentation enhancers. Journal of Photochemistry and Photobiology B Biology 63:148-161.
27. Burke KE, Clive J, Combs GF Jr, Nakamura RM. 2003. Effects of topical L-selenomethionine with topical and oral vitamin E on pigmentation and skin cancer induced by ultraviolet irradiation in Skh:2 hairless mice. Journal of the American Academy of Dermatology 49:458-472.
28. Byrne SN, Spinks N, Halliday GM. 2002. Ultraviolet a irradiation of C57BL/6 mice suppresses systemic contact hypersensitivity or enhances secondary immunity depending on dose. Journal of Investigative Dermatology 119:858-864.
29. Cao C, Wan Y. 2009. Parameters of protection against ultraviolet radiation-induced skin cell damage. Journal of Cellular Physiology 220:277-284.
30. Carell T, Burgdorf LT, Kundu LM, Cichon M. 2001. The mechanism of action of DNA photolyases. Current Opinion in Chemical Biology 5:491-498.
31. Carless MA, Lea RA, Curran JE, Appleyard B, Gaffney P, Green A, Griffiths LR. 2002. The GSTM1 null genotype confers an increased risk for solar keratosis development in an Australian Caucasian population. Journal of Investigative Dermatology 119:1373-1378.
32. Casso D, Beach D. 1996. A mutation in a thioredoxin reductase homolog suppresses p53-induced growth inhibition in the fission yeast Schizosaccharomyces pombe. Archives of Dermatology 123:615-619.
33. Chen ML, Li J, Xiao WR, Sun L, Tang H, Wang L, Wu LY, Chen X, Xie HF. 2006. Protective effect of resveratrol against oxidative damage of UVA irradiated HaCaT cells. Zhong Nan Da Xue Xue Bao Yi Xue Ban 31:635-639.
34. Chiu A, Kimball AB. 2003. Topical vitamins, minerals and botanical ingredients as modulators of environmental and chronological skin damage. British Journal of Dermatology 149:681-691.
35. Cook N, Freeman S. 2001. Report of 19 cases ofphotoallergic contact dermatitis to sunscreens seen at the Skin and Cancer Foundation. Australasian Journal of Dermatology 42:257-259.
36. Coroneo MT. 1993. Pterygium as an early indicator ofultraviolet insolation: A hypothesis. British Journal of Ophthalmology 77:734-739.
37. Costa MR, Chiganeas V, Galhardo RS, Calvarho H, Menck CF. 2003. The euraryotic nucleotide excision repair pathway. Biochimie 85:1083-1099.
38. Cotter MA, Thomas J, Cassidy P, Robinette K, Jenkins N, Florell SR, Leachman S, Samlowski WE, Grossman D. 2007. N-acetylcysteine protects melanocytes against oxidative stress/damage and delays onset of ultraviolet-induced melanoma in mice. Clinical Cancer Research 13:5952-5958.
39. D'Agostiny F, Balansky RM, Camoirano A, De Flora S. 2005. Modulation of light-induced skin tumors by N-acetylcysteine and/or ascorbic acid in hairless mice. Carcinogenesis 26:657-664.
40. Dahle J, Brunborg G, Svendsrud DH, Stokke T, Kvam E. 2008. Overexpression of human OGG1 in mammalian cells decreases ultraviolet A induced mutagenesis. Cancer Letters 267:18-25.
41. Dany AL, Tissier A. 2001. A functional OGG1 homologue from Arabidopsis thaliana. Molecular Genetics and Genomics 265: 293-301.
42. Darr D, Combs S, Dunston S, Manning T, Pinnell S. 1992. Topical vitamin C protects porcine skin from ultraviolet radiation-induced damage. British Journal of Dermatology 127:247-253.
43. Darr D, Dunston S, Faust H, Pinnell S. 1996. Effectiveness of antioxidants (vitamin C and E) with and without sunscreens as topical photoprotectants. Acta Dermato-Venereologica 76: 264-268.
44. Darvin M, Zastrow L, Sterry W, Lademann J. 2006. Effect of supplemented and topically applied antioxidant substances on human tissue. Skin Pharmacology and Physiology 19:238-247.
45. Davies KJ. 2000. Oxidative stress, antioxidant defenses, and damage removal, repair, and replacement systems. IUBMB Life 50:279-289.
46. de Boer J, Hoeijmakers JH. 2000. Nucleotide excision repair and human syndromes. Carcinogenesis 21:453-460.
47. de Winter S, Vink AA, Roza L, Pavel S. 2001. Solar-simulated skin adaptation and its effect on subsequent UV-induced epidermal DNA damage. Journal of Investigative Dermatology 117:678-682.
48. Declercq L, Sente I, Hellemans L, Corst1ens H, Maes D. 2004. Use of the synthetic superoxide dismutase/catalase mimetic EUK-134 to compensate for seasonal antioxidant deficiency by reducing pre-existing lipid peroxides at the human skin surface. International Journal ofCosmetic Science 26:255-263.
49. Deep G, Agarwal R. 2007. Chemopreventive efficacy of silymarin in skin and prostate cancer. Integrative Cancer Therapies 6:130-145.
50. Deliconstantinos G, Villitou V, Stavrides JC. 1997. Inhibition of ultraviolet B-induced skin erythema by N-nitro-L-arginine and N-monomethyl-L-arginine. Journal of Dermatological Science 15:23-35.
51. D'Errico M, Teson M, Calcagnile A, Proietti De Santis L, Nikaido O, Botta E, Zambruno G, Stefanini M, Dogliotti E. 2003. Apoptosis and efficient repair of DNA damage protect human keratinocytes against UVB. Cell Death and Differentiation 10:754-756.
52. Didier C, Kerblat I, Drouet C, Favier A, Béani JC, Richard MJ. 2001a. Induction of thioredoxin by ultraviolet-A radiation prevents oxidative-mediated cell death in human skin fibroblasts. Free Radical Biology and Medicine 31:585-598. (Pubitemid 32777859)
53. Didier C, Pouget JP, Cadet J, Favier A, Béani JC, Richard MJ. 2001b. Modulation of exogenous and endogenous levels of thioredoxin in human skin fibroblasts prevents DNA damaging effect of ultraviolet A radiation. Free Radical Biology and Medicine 30:537-546.
54. Diffey B. 2001b. Sunscreenisn't enough. Journal of Photochemistry and Photobiology B Biology 64:105-108.
55. Dinkova-Kostova AT. 2008. Phytochemicals as protectors against ultraviolet radiation: Versatility of effects and mechanisms. Planta Medica 74:1548-1559.
56. El Sayed F, Ammoury A, Nakhle F, Dhaybi R, Marguery MC. 2006. Photoprotection in teenagers. Photodermatology, Photoimmunology and Photomedicine 22:18-21.
57. Eller MS, Ostrom K, Gilchrest BA. 1996. DNA damage enhances melanogenesis. Proceedings of the National Academy of Sciences of the USA 93:1087-1092.
58. Eller MS, Yaar M, Gilchrest BA. 1994. DNA damage and melanogenesis. Nature 372:413-414.
59. Eller MS, Maeda T, Magnoni C, Arwal D, Gilchrest BA. 1997. Enhancement of DNA repair in human skin cells by thymidine dinucleotides: Evidence for p53-mediated mammalian SOS response. Proceedings of the National Academy of Sciences of the USA 94:12627-12632.
60. Erden IM, Kahraman A, Köken T. 2001. Beneficial effects of quercetin on oxidative stress induced by ultraviolet A. Clinical and Experimental Dermatology 26:536-539.
61. Essen LO, Klar T. 2006. Light-driven DNA repairbyphotolyases. Cellular and Molecular Life Sciences 63:1266-1277.
62. Ettler K. 2004. Fotoprotekce ku°ze. Ochrana ku°zepredú cinky ultrafialového záreńi. Praque: Triton.
63. F'guyer S, Afaq F, Mukhtar H. 2003. Photochemoprevention of skin cancer by botanical agents. Photodermatology, Photoimmunology and Photomedicine 19:56-72.
64. Faurschou A, Wulf HC. 2007. The relation between sun protection factor and amount of sunscreen applied in vivo. Photobiology 156:716-719.
65. Fitzpatrick TB, Breathnach AS. 1963. The epidermal melanin unit system. Dermatologische Wochenschrift 147:481-489.
66. Fonnum F, Lock EA. 2004. The contributions of excitotoxicity, glutathione depletion and DNA repair in chemically induced injury to neurones: Exemplified with toxic effects on cerebellar ganule cells. Journal of Neurochemistry 88:513-531.
67. Frank S, Kämpfer H, Wetzler C, PfeilschifterJ. 2002. Nitric oxide drives skin repair: Novel functions of an established mediator. Kidney International 61:882-888.
68. Fuller B, Smith D, Howerton A, Kern D. 2006. Antiinflammatory effects of CoQ10 and colorless carotenoids. Journal of Cosmetic Dermatology 5:30-38.
69. Gallardo A, Teixidó J, Miralles R, Raga M, Guglietta A, Marquillas F, Sallarès J, Nonell S. 2010. Dose-dependent progressive sunscreens. A new strategy for photoprotection? Photochemical and Photobiological Sciences 9:530-534.
70. Gambichler T, Laperre J, Hoffmann K. 2006. The European standard for sun-protective clothing: EN 13758. Journal of the European Academy of Dermatology and Venereology 20:125-130.
71. Garmyn M, Ribaya-Mercado JD, Russel RM, Bhawan J, Gilchrest BA. 1995. Effect of beta-carotene supplementation on the human sunburn reaction. Experimental Dermatology 4:104-111.
72. Gaspar LR, Campos PM. 2007. Photostability and efficacy studies of topical formulations containing UV-filters combination and vitamins A, C and E. International Journal of Pharmaceutics 343:181-189.
73. Gerber B, Mathys P, Moser M, Bressoud D, Braun-Fahrländer C. 2002. Ultraviolet emission spectra of sunbeds. Photochemistry and Photobiology 76:664-668.
74. Gies P. 2007. Photoprotection by clothing. Photodermatology, Photoimmunology and Photomedicine 23:264-274.
75. Gonzalez S, Alonso-Lebrero JL, Del Rio R, Jaen P. 2007. Polypodium leucotomos extract: A nutraceutical with photo-protective properties. Drugs of Today (Barcelona) 43:475-485.
76. Goukassian DA, Eller MS, Yaar M, Gilchrest BA. 1999. Thymidine dinucleotide mimics the effect of solar simulated irradiation on p53 and p53-regulated proteins. Journal of Investigative Dermatology 112:25-31.
77. Greul AK, Grudmann JU, Heinrich F, Pfitzner I, Bernhardt J, Ambach A, Biesalski HK, Gollnick H. 2002. Photoprotection ofUV-irradiated human skin: An antioxidative combination of vitamins E and C, carotenoids, selenium and proanthocyanidins. Skin Pharmacology and Applied Skin Physiology 15:307-315.
78. Hadshiew IM, Eller MS, Gasparro FP, Gilchrest BA. 2001. Stimulation of melanogenesis by DNA nucleotides. Effects of size, sequence and 5' phosphorilation. Journal of Dermatological Science 5:127-138.
79. Hall AG. 1999. The role of glutathione in the regulation of apoptosis. European Journal ofClinical Investigation 29:238-245.
80. Halliday GM. 2005. Inflammation, gene mutation and photoimmunosuppression in response to UVR-induced oxidative damage contributes to photocarcinogenesis. Mutation Research 571:107-120.
81. Halliday GM, Rana S. 2008. Waveband and dose dependency of sunlight-induced immunomodulation and cellular changes. Photochemistry and Photobiology 84:35-46.
82. Hanada K, Sawamura D, Tamai K, Hashimoto I, Kobayashi S. 1997. Photoprotective effect of esterified glutathione against ultraviolet B-induced sunburn cell formation in the hairless mice. Journal of Investigative Dermatology 108:727-730.
83. Hayes JD, Strange RC. 2000. Glutathione-S-transferase polymorphisms and their biological consequence. Pharmacology 61:154-166.
84. Haywood R, Wardman P, Sanders R, Linge C. 2003. Sunscreens inadequately protect against ultraviolet-A-induced free radicals in skin: Implications for skin aging and melanoma? Journal of Investigative Dermatology 121:862-868.
85. Hellemans L, Corstjens H, Neven A, Declercq L, Maes D. 2003. Antioxidant enzyme activity in human stratum corneum shows seasonal variations with an age-dependent recovery. Journal of Investigative Dermatology 120:434-439.
86. Herzog B, Mongiat S, Quass K, Deshayes C. 2004. Prediction of sun protection factors and UVA parameters of sunscreens by using a calibrated step film model. Journal of Pharmaceutical Sciences 93:1780-1795.
87. Hirota A, Kawachi Y, Itoh K, Nakamura Y, Xu X, Banno T, Takahashi T, Yamamoto M, Otsuka F. 2005. Ultraviolet A irradiation induces NF-E2-related factor 2 activation in dermal fibroblasts: Protective role in UVA-induced apoptosis. Journal of Investigative Dermatology 124:825-832.
88. Hirota K, Murata M, Sachi Y, Nakamura H, Takeuchi J, Mori K, Yodoi J. 1999. Distinct roles of thioredoxin in the cytoplasm and in the nucleus. A two-step mechanism of redox regulation of transcription factor NF-kappaB. Journal of Biological Chemistry 274:27891-27897.
89. Hoffmann K, Laperre J, Avermaete A, Altmeyer P, Gambichler T. 2001. Defined UV protection by apparel textiles. Archives of Dermatology 137:1089-1094.
90. Ibrahim SF, Brown MD. 2008. Tanning and cutaneous malignancy. Dermatologic Surgery 34:460-474.
91. Inui M, Ooe M, Fujii K, Matsunaka H, Yoshida M, Ichihashi M. 2008. Mechanisms of inhibitory effects of CoQ10 on UVB-induced wrinkle formation in vitro and in vivo. Biofactors 32:237-243.
92. Jaakola L, Määttä-Riihinen K, Kärenlampi S, Hohtola A. 2004. Activation of flavonoid biosynthesis by solar radiation in bilberry (Vaccinium myrtillus L.) leaves. Planta 218:721-728.
93. Javeri A, Huang XX, Bernerd F, Mason RS, Halliday GM. 2008. Human 8-oxoguanine-DNA glycosylase 1 protein and gene are expressed more abundantly in the superficial than basal layer of human epidermis. DNA Repair (Amst) 7:1542-1550.
94. Jung K, Seifert M, Herrling T. 2008a. The fatal effects of self-tanning agents during UV radiation. SOFW-Journal 3:12-17.
95. Jung K, Seifert M, Herrling T, Fuchs J. 2008b. UV generated free radicals (FR) in the skin: Their prevention by sunscreens and their induction by self-tanning agents. Spectrochimica acta. Part A, Molecular and biomolecular spectroscopy 69:1423-1428.
96. Junqueira LC, Carneiro J, Kelley RO. 1992. Basic histology. Norwalk: Prentice Hall International Inc. pp 339-351.
97. Kaidbey KH, Agin PP, Sayre RM, Kligman AM. 1979. Photoprotection by melanin. A comparison ofblack and Caucasian skin. Journal of the American Academy of Dermatology 1:249-260.
98. Kanitakis J. 2002. Anatomy, histology and immunohistochemistry of normal human skin. European Journal of Dermatology 12:390-399.
99. Kappes UP, Luo D, Potter M, Schulmeister K, Runger TM. 2006. Short-and long-wave UV light (UVB and UVA) induce similar mutations in human skin cells. Journal of Investigative Dermatology 126:667-675.
100. Katiyar S, Elmets CA, Katiyar SK. 2007. Green tea and skin cancer: Photoimmunology, angiogenesis and DNA repair. Journal ofNutritional Biochemistry 18:287-296.
101. Katiyar SK. 2005. Silymarin and skin cancer prevention: Antiinflammatory, antioxidant and immunomodulatory effects (Review). International Journal of Oncology 26:169-176.
102. Katiyar SK. 2007. UV-induced immune suppression and photo-carcinogenesis: Chemoprevention by dietary botanical agents. Cancer Letters 255:1-11.
103. Katiyar SK. 2008. Grape seed proanthocyanidines and skin cancer prevention: Inhibition of oxidative stress and protection of immune system. Molecular Nutrition and Food Research 52:S71-76.
104. Katiyar SK, Afaq F, Perez A, Mukhtar H. 2001. Green tea polyphenol (-)-epigallocatechin-3-gallate treatment of human skin inhibits ultraviolet radiation-induced oxidative stress. Carcinogenesis 22:287-294.
105. Kawachi Y, Xu X, Taguchi S, Sakurai H, Nakamura Y, Ishii Y, Fujisawa Y, Furuta J, Takahashi T, Itoh K, Yamamoto M, Yamazaki F, Otsuka F. 2008. Attenuation of UVB-induced sunburn reaction and oxidative DNA damage with no alterations in UVB-induced skin carcinogenesis in Nrf2 gene-deficient mice. Journal of Investigative Dermatology 128:1773-1779.
106. Kerb R, Brockmöller J, Reum T, Roots I. 1997. Deficiency of glutathione S-transferases T1 and M1 as heritable factors of increased cutaneous UV sensitivity. Journal of Investigative Dermatology 108:229-232.
107. Kim DW, Hwang IK, Kim DW, Yoo KY, Won CK, Moon WK, Won MH. 2007. Coenzyme Q effects on manganese super-oxide dismutase and glutathione peroxidase in the hairless mouse skin induced by ultraviolet B irradiation. Biofactors 30:139-147.
108. Kobayashi S, Takehana M, Tohyama C. 1996. Glutathione isopropyl ester reduces UVB-induced skin damage in hairless mice. Photochemistry and Photobiology 63:106-110.
109. Koháryová M, Kolárová M. 2008. Oxidative stress and thioredox-in system. General Physiology and Biophysics 27:71-84.
110. Kohen R, Gati I. 2000. Skin low molecular weight antioxidants and their role in aging and in oxidative stress. Toxikology 148:149-157.
111. Korkina LG, Pastore S, De Luca C, Kostyuk VA. 2008. Metabolism of plant polyphenols in the skin: Beneficial versus deleterious effects. Current Drug Metabolism 9:710-729.
112. Krol ES, Kramrp-Stickland KA, Lieber DC 2000. Photoprotec-tive actions of topically applied vitamin E. Drug Metabolism Reviews 32:413-420.
113. Kulms D, Schwarz T. 2002. Independent contribution of three different pathways to ultraviolet-B-induced apoptosis. Bio-chemical Pharmacology 64:837-841.
114. Kvam E, Noel A, Basu-Modak S, Tyrrell RM. 1999. Cycloox-ygenase dependent release of heme from microsomal heme-proteins correlates with induction of heme oxygenase 1 transcription in human fibroblasts. Free Radical Biology and Medicine 26:511-517.
115. Lautenschlanger S, WulfHC, Pittelkow MR. 2007. Photoprotection. The Lancet 370:528-537.
116. Lautier D, Luscher P, Tyrrell RM. 1992. Endogenous glutathione levels modulate both constitutive and UVA radiation/hydrogen peroxide inducible expression of the human heme oxygenase gene. Carcinogenesis 13:227-232.
117. Lavker RM, Gerberick GF, Veres D, Irwin CJ, Kaidbey KH. 1995. Cumulative effects from repeated exposures to suber-ythemal doses of UVB and UVA in human skin. Journal of the American Academy of Dermatology 32:53-62.
118. Leccia MT, Yaar M, Allen N, Gleason M, Gilchrest BA. 2001. Solar simulated irradiation modulates gene expression and activity of antioxidants enzymes in cultured human dermal fibroblasts. Experimental Dermatology 10:272-279.
119. Leccia MT, Richard MJ, Beani JC, Faure H, Monjo AM, Cadet J, Amblard P, Favier A. 1993. Protective effect ofselenium and zinc on UV-A damage in human skin fibroblasts. Photo Chemistry and Photobiology 58:548-553.
120. Lee JH, An HT, Chung JH, Kim KH, Eun HC, Cho KH. 2002. Acute effects of UVB radiation on the proliferation and differentiation of keratinocytes. Photodermatology, Photoim-munology and Photomedicine 18:253-261.
121. Lloyd RS. 2005. Investigations ofpyrimidine dimer glycosylases-a paradigm for DNA base excision repair enzymology. Mutation Research 577:77-91.
122. Lopez-Torres M, Thiele JJ, Shindo Y, Han D, Packer L. 1998. Topical application of tocopherol modulates the antioxidant network and diminishes ultraviolet-induced oxidative damage in murine skin. British Journal of Dermatology 138:207-215.
123. Lucas CR, Ke MS, Matsui MS, Maes D, Cooper KD, Stevens SR, Baron ED. 2008. Immune protective effect of a moisturizer with DNA repair ingredients. Journal of Cosmetic Dermatol-ogy 7:132-135.
124. Mahmoud BH, Hexsel CL, Hamzavi IH. 2008. Effects of visible light on the skin. Photo Chemistry and Photobiology 84:450-462.
125. Mahroos MA, Yaar M, Phillips TJ, Bhawan J, Gilchrest BA. 2002. Effect of sunscreen application on UV-induced thymine dimers. Archives of Dermatology 138:1480-1485.
126. Maier T, Korting HC. 2005. Sunscreens-which and what for? Skin Pharmacology and Physiology 18:253-262.
127. Manzanas Garćia A, López Carrioza MC, Vallejo Ocana C, Samper Ots P, Delgado Pérez JM, Carretero Accame E, Gómez- Serranillos P, de la Morena del Valle L. 2008. Superoxide dismutase (SOD) topical use in oncologic patients: Treatment of acute cutaneous toxicity secondary to radio-therapy. Clinical and Translational Oncology 10:163-167.
128. Marrot L, Jones C, Perez P, MeunierJR. 2007. The significance of Nrf2 pathway in (photo)-oxidative stress response in melano-cytes and keratinocytes of the human epidermis. Pigment Cell and Melanoma Research 21:79-88.
129. Marrot L, Belaidi J, MeunierJ, Perez P, Agapakis-Causse C. 1999. The human melanocyte as a particular target for UVA radiation and an endpoint for photoprotection assessment. Photo Chemistry and Photobiology 69:686-693.
130. Masella R, Di Benedetto R, Vari R, Filesi C, Giovannini C. 2005. Novel mechanisms of natural antioxidant compounds in biological systems: Involvement of glutathione and glu-tathione-related enzymes. Journal of Nutritional Biochemistry 16:577-586.
131. Masnec IS, Voda K, Situm M. 2007. UV radiation: What we know and do we protect ourselves adequates? Collegium Antropo-logicum 1:97-100.
132. Matsumura Y, Ananthaswamy HN. 2004. Toxic effects of ultraviolet radiation on the skin. Toxicology and Applied Pharmacology 195:298-308.
133. McArdle F, Rhodes LE, Parslew RA, Close GL, Jack CI, Friedmann PS, Jackson MJ. 2004. Effects of oral vitamin E and beta-carotene supplementation on ultraviolet radiation-induced oxidative stress in human skin. American Journal of Clinical Nutrition 80:1270-1275.
134. Meewes C, Brenneisen P, Wenk J, Kuhr L, Ma W, Alikoski J, Poswig A, Krieg T, Scharffetter-Kochanek K. 2001. Adaptive antioxidant response protects dermal fibroblasts from UVA-induced phototoxicity. Free Radical Biology and Medicine 3:238-247.
135. Meira LB, Cheo DL, Reis AM, Claij N, Burns DK, te Riele H, Friedberg EC 2002. Mice defective in the mismatch repair gene Msh2 show increased predisposition to UVB radiation-induced skin cancer. DNA Repair (Amst) 1:929-934.
136. Melnikova VO, Ananthaswamy NH. 2005. Cellular and molecular events leading to the development of skin cancer. Mutation Research 571:91-106.
137. Mistry P, Herbert KE. 2003. Modulation of hOGG1 DNA repair enzyme in human cultured cells in response to prooxidant and antioxidant challenge. Free Radical Biology and Medicine 35:397-405.
138. Monteiro-Riviere NA. 2004. Anatomical factors affecting barrier function. In: Zhai H, Maibach HI, editors. Dermatotoxicology, Danvers, MA: CRC Press LLC. pp 43-70.
139. Morley N, Curnow A, Salter L, Campbell S, Gould D. 2003. N-acetyl-L-cysteine prevents DNA damage induced by UVA, UVB and visible radiation in human fibroblasts. Journal of Photo Chemistry and Photobiology B Biology 72:55-60.
140. Mouret S, Baudouin C, Charveron M, Favier A, Cadet J, Douki T. 2006. Cyclobutane pyrimidine dimers are predominant DNA lesions in whole human skin exposed to UVA radiation. Proceedings of the National Academy of Sciences of the USA 103:13765-13770.
141. Mouret S, Philippe C, Gracia-Chantegrel J, Banyasz A, Karpati S, Markovitsi D, Douki T. 2010. UVA-induced cyclobutane pyrimidine dimers in DNA: A direct photoche-mical mechanism? Organic and Biomolecular Chemistry 8:1706-1711.
142. Mullenders LH, Berneburg M. 2001. Photoimmunology and nucleotide excision repair: Impact of transcription coupled and global genome excision repair. Journal of Photo Chemistry and Photobiology B Biology 2/3:97-100.
143. Murray JC, Burch JA, Streilein RD, Iannacchione MA, Hall RP, Pinnell RS. 2008. A topical antioxidant solution containing vitamins C and E stabilized by ferulic acid provides protection for human skin against damage caused by ultraviolet irradia-tion. Journal of the American Academy of Dermatology 59:418-425.
144. Mustacich D, Powis G. 2000. Thioredoxin reductase. Biochemical Journal 346:1-8.
145. Naderi-Hachtroudi L, Peters T, Brenneisen P, Meewes C, Hommel C, Razi-Wolf Z, Schneider LA, Schuller J, Wlaschek M, Scharffetter-Kochanek K. 2002. Induction of manganese superoxide dismutase in human dermal fibroblasts: A UV-B-mediated paracrine mechanism with the release of epidermal interleukin 1 alpha, interleukin 1 beta, and tumor necrosis factor alpha. Archives of Dermatology 138:1473-1479.
146. Nair S, Kekatpure VD, Judson BL, Rifkind AB, Granstein RD, Boyle JO, Subbaramaiah K, Guttenplan JB, Dannenberg AJ. 2009. UVR exposure sensitizes keratinocytes to DNA adduct formation. Cancer Prevention Research (Phila Pa) 2:895-902.
147. Nakamura Y, Yoshida C, Murakami A, Ohigashi H, Osawa T, Uchida K. 2004. Zerumbone, a tropical ginger sesquiterpene, activates phase II drug metabolizing enzymes. FEBS Letters 572:245-250.
148. Nandakumar V, Singh T, Katiyar SK. 2008 Multi-targeted prevention and therapy of cancer by proanthocyanidins. Cancer Letters 269:378-387.
149. Nash JF. 2006. Human safety and efficacy ofultraviolet filters and sunscreen products. Dermatologic Clinics 24:35-51.
150. Nohynek GJ, Schaefer H. 2001. Benefit and risk of organic ultraviolet filters. Regulatory Toxicology and Pharmacology 33:285-291.
151. Nohynek GJ, Dufour EK, Roberts MS. 2008. Nanotechnology, cosmetics and the skin: Is there a health risk? Skin Pharmacol-ogy and Physiology 21:136-149.
152. Nolan BV, Taylor SL, Liguori A, Feldman SR.2009.Tanning as an addictive behavior: A literature review. Photodermatology, Photoimmunology and Photomedicine 25:12-19.
153. Obermuller-Jevic UC, Schlegel B, Flaccus A, Biesalski HK. 2001. The effect of beta-carotene on the expression of interleukin-6 and heme oxygenase-1 in UV-irradiated human skin fibroblasts in vitro. FEBS Letters 509:186-190.
154. Offord EA, GautierJC, Avanti O, Scaletta C, Runge F, Krämer K, Applegate LA. 2002. Photoprotective potential of lycopene, beta-carotene, vitamin E, vitamin C and carnosic acid in UVA-irradiated human skin fibroblasts. Free Radical Biology and Medicine 32:1293-1303.
155. Okada K, Takahashi Y, Ohnishi K, Ishikawa O, Miyachi Y. 1994. Time-dependent effect of chronic UV irradiation on super-oxide dismutase and catalase activity in hairless mice skin. Journal of Dermatological Science 8:183-186.
156. Park HY, Mosmadaki M, Yaar M, Gilchrest BA. 2009. Cellular mechanisms regulating human melanogenesis. Cellular and Molecular Life Sciences 66:1493-1506.
157. Pearse AD, Gaskell SA, Marks R. 1987. Epidermal changes in human skin following irradiation with either UVB or UVA. Journal of Investigative Dermatology 88:83-87.
158. Pence BC, Naylor MF. 1990. Effects of single-dose ultraviolet radiation on skin superoxide dismutase, catalase, and xanthine oxidase in hairless mice. Journal of Investigative Dermatology 95:213-216.
159. Persson AE, Edström DW, Bäckvall H, Lundeberg J, Potén F, Ros AM, Williams C. 2002. The mutagenic effect ofultraviolet-A1 on human skin demonstrated by sequencing the p53 gene in single keratinocytes. Photodermatology, Photoimmunology and Photomedicine 18:287-293.
160. Petropoulos I, Friguet B. 2006. Maintenance of proteins and aging: The role of oxidized protein repair. Free Radical Research 40:1269-1276.
161. Phan TA, Halliday GM, Barnetson RC, Damian DL. 2006. Melanin differentially protects from the initiation and progres-sion of threshold UV-induced erythema depending on UV waveband. Photodermatology, Photoimmunology and Photo-medicine 22:174-180.
162. Pinnell SR. 2003. Cutaneous photodamage, oxidative stress, and topical antioxidant protection. Journal of the American Academy of Dermatology 48:1-19.
163. Pitsikas P, Lee D, Rainbow AJ. 2007. Reduced host cell reactivation of oxidative DNA damage in human cells deficient in the mismatch repair gene hMSH2. Mutagenesis 22:235-243.
164. Podda M, Grundmann-Kollmann M. 2001. Low molecular weight antioxidants and their role in skin ageing. Clinical and Experimental Dermatology 26:578-582. (Pubitemid 32989582)
165. Poon TS, Barnetson RS, Halliday GM. 2005. Sunlight-induced immunosuppression in humans is initially because of UVB, then UVA, followed by interactive effects. Journal of Investi-gative Dermatology 125:840-846.
166. Poswig A, Wenk J, Brenneisen P, Wlaschek M, Hommel C, Quel G, Faisst K, Dissemond J, Briviba K, Krieg T, Scharffetter-Kochanek K. 1999. Adaptive antioxidant response of manganese-superoxide dismutase following repe-titive UVA irradiaton. Journal of Investigative Dermatology 112:13-18.
167. Psotová J, Svobodová A, Kolárová H, Walterová D. 2006. Photoprotective properties of Prunella vulgaris and rosmarinic acid on human keratinocytes. Journal of Photo Chemistry and Photobiology B Biology 84:167-174.
168. Rafferty TS, Walker C, Hunter JA, Beckett GJ, McKenzie RC. 2002. Inhibition of ultraviolet B radiation-induced interleukin 10 expression in murine keratinocytes by selenium com-pounds. British Journal of Dermatology 146:485-489.
169. Rahman I, MacNee W. 2000. Regulation of redox glutathione levels and gene transcription in lung inflammation: Therapeu-tic approaches. Free Radical Biology and Medicine 28:1405-1420.
170. Reagan-Shaw S, Mukhtar H, Ahmad N. 2008. Resveratrol imparts photoprotection of normal cells and enhances the efficacy of radiation therapy in cancer cells. Photo Chemistry and Photo-biology 84:415-421.
171. Reedy MV, Faraco CD, Erickson CA. 1998. Specification and migration of melanoblasts at the vagal level and in hyperpig-mented Silkie chickens. Developmental Dynamics 213:476-485.
172. Reeve VE, Tyrrell RM. 1999. Heme oxygenase induction mediates the photoimmunoprotective activity of UVA radiation in the mouse. Proceedings of the National Academy of Sciences of the USA 96:9317-9321.
173. Rezvani HR, Mazurier F, Cario-André M, Pain C, Ged C, Taïeb A, de Verneuil H. 2006a. Protective effect of catalase overexpression on UVB-induced apoptosis in normal human keratinocytes. Journal of Biological Chemistry 281:17999-18007.
174. Rezvani HR, Cario-Andre M, Pain C, Ged C, de Verneuil H, Täieb A. 2006b. Protection of normal human reconstructed epidermis from UV by catalase overexpression. Cancer Gene Therapy 14:174-186.
175. Ridley AJ, Whiteside JR, McMillan TJ, Allinson SL. 2009. Cellular and sub-cellular responses to UVA in relation to carcinogenesis. International Journal of Radiation Biology 85:177-195.
176. Routaboul C, Denis A, Vinche A. 1999. Immediate pigment darkening: Description, kinetic and biological function. European Journal of Dermatology 9:95-99.
177. Saitoh T, Shinmura K, Yamaguchi S, Tani M, Seki S, Murakami H, Nojima Y, Yokota J. 2001. Enhancement ofOGG1 protein AP lyase activity by increase of APEX protein. Mutation Research 486:31-40.
178. Sánchez-Campillo M, Gabaldon JA, Castillo J, Benavente- García O, Del Bano MJ, Alcaraz M, Vicente V, Alvarez N, Lozano JA. 2009. Rosmarinic acid, a photo-protective agent against UV and other ionizing radiations. Food and Chemical Toxicology 47:386-392.
179. Sasaki H, Akamatsu H, Horio T. 1997. Effects of a single exposure to UVB radiation on the activities and protein levels of copper-zinc and manganese superoxide dismutase in cultured human keratinocytes. Photo Chemistry and Photobiol-ogy 65:707-713.
180. Schäfer M, Werner S. 2008. Oxidative stress in normal and impaired wound repair. Pharmacological Research 58:165-171.
181. Schallreuter KU, Wood JM. 1986. The role of thioredoxin reductase in the reduction of free radicals at the surface of the epidermis. Biochemical and Biophysical Research Com-munications 136:630-637.
182. Schallreuter KU, Wood JM. 2001. Thioredoxin reductase-its role in epidermal redox status. Journal of Photo Chemistry and Photobiology B Biology 64:179-184.
183. Schallreuter KU, Hordinsky MK, Wood JM. 1987. Thioredoxin reductase. Role in free radical reduction in different hypopig-mentation disorders. Archives of Dermatology 123:615-619.
184. Schieke SM, Ruwiedel K, Gers-Barlag H, Grether-Beck S, Krutmann J. 2005. Molecular crosstalk of the ultraviolet A and ultraviolet B signaling responses at the level of mitogen-activated protein kinases. Journal of Investigative Dermatology 124:857-859.
185. Schieke SM, Schroeder P, KrutmannJ. 2003. Cutaneous effects of infrared radiation: From clinical observations to molecular response mechanisms. Photodermatology, Photoimmunology and Photomedicine 19:228-234.
186. Schilling K, Bradford B, Castelli D, Dufour E, Nash JF, Pape W, Schulte S, Tooley I, van den Bosch J, Schellauf F. 2010. Human safety review of'nano' titanium dioxide and zinc oxide. Photochemical and Photobiological Sciences 9:495-509.
187. Schroeder P, Haendeler J, Krutmann J. 2008. The role of near infrared radiation in photoaging of the skin. Experimental Gerontology 43:629-632.
188. Seifert M, Scherer SJ, Edelmann W, Böhm M, Meineke V, Löbrich M, Tilgen W, Reichrath J. 2008. The DNA-mismatch repair enzyme hMSH2 modulates UV-B-induced cell cycle arrest and apoptosis in melanoma cells. Journal of Investigative Dermatology 128:203-213.
189. Shaath NA. 2010. Ultraviolet filters. Photochemical and Photo-biological Sciences 9:464-469.
190. Sharma SD, Meeran SM, Katiyar SK. 2007. Dietary grape seed proanthocyanidins inhibit UVB-induced oxidative stress and activation of mitogen-activated protein kinases and nuclear factor-B signaling in vivo SKH-1 hairless mice. Molecular Cancer Therapeutics 6:995-1005.
191. Sheedy JE, Edlich RF. 2004. Ultraviolet eye radiation: The problem and solutions. Journal of Long-Term Effects of Medical Implants 14:67-71.
192. Shen J, Bao S, Reeve VE. 1999. Modulation ofIL-10, IL-12, and IFN-gamma in the epidermis of hairless mice by UVA (320-400 nm) and UVB (280-320 nm) radiation. Journal of Investigative Dermatology 113:1059-1064.
193. Shin-Darlak CY, Skinner AM, Turker MS. 2005. A role for Pms2 in the prevention of tandem CC→ TT substitutions induced by ultraviolet radiation and oxidative stress. DNA Repair (Amst) 4:51-57.
194. Shindo Y, Hashimoto T. 1997. Time course of changes in antioxidant enzymes in human skin fibroblasts after UVA irradiation. Journal of Dermatological Science 14:225-232.
195. Shindo Y, Witt E, Han D, Parker L. 1994a. Dose-response effects of acute ultraviolet irradiation on antioxidants and molecular markers of oxidation in murine epidermis and dermis. Journal of Investigative Dermatology 102:470-475.
196. Shindo Y, Witt E, Han D, Epstein W, Packer L. 1994b. Enzymatic and non-enzymatic antioxidants in epidermis and dermis of human skin. Journal of Investigative Dermatology 102:122-124.
197. Shindo Y, Witt E, Hand D, Tzeng B, Aziz T, Nguyen L, Packer L. 1994c. Recovery of antioxidants and reduction in lipid hydroperoxides in murine epidermis and dermis after acute ultraviolet radiation exposure. Photodermatology, Photoim-munology and Photomedicine 10:183-191.
198. Sime S, Reeve VE. 2004. Protection from inflammation, immunosuppression and carcinogenesis induced by UV radia-tion in mice by topical Pycnogenol. Photo Chemistry and Photobiology 79:193-198.
199. Sindhu RK, Wagner FE, Kikkawa Y. 2003. Induction of cytochrome p450 1A1 and 1B1 by photooxidized tryptophan in transformed human keratinocytes. Advances in Experimental Medicine and Biology 527:297-306.
200. Singh RP, Agarwal R. 2009. Cosmeceuticals and silibinin. Clinical Dermatology 27:479-484.
201. Singh RP, Agarwal R. 2005. Mechanisms and preclinical efficacy of silibinin in preventing skin cancer. European Journal of Cancer 41:1969-1979.
202. Song XZ, Xia JP, Bi ZG. 2004. Effects of (-)-epigallocatechin-3-gallate on expression of matrix metalloproteinase-1 and tissue inhibitor of metalloproteinase-1 in fibroblasts irra-diated with ultraviolet A. Chinese Medical Journal (Engl) 117:1838-1841.
203. Stahl W, Heinrich U, Aust O, Tronnier H, Sies H. 2006. Lycopene-rich products and dietary photoprotection. Photo-chemical and Photobiological Sciences 5:238-242.
204. Steenvoorden DP, Beijersbergen van Henegouwen GM. 1998a. Glutathione ethylester protects against local and systemic suppression of contact hypersensitivity induced by ultraviolet B radiation in mice. Radiation Research 150:292-297.
205. Steenvoorden DP, Beijersbergen van Henegouwen GM. 1998. Glutathione ethylester protects against local and systemic suppression of contact hypersensitivity induced by ultraviolet B radiation in mice. Radiation Research 150:292-297.
206. Steenvoorden DP, Hasselbaink DM, Beijersbergen van Hene-gouwen GM. 1998b. Protection against UV-induced reactive intermediates in human cells and mouse skin by glutathione precursors: A comparison of N-acetylcysteine and glutathione ethylester. Photo Chemistry and Photobiology 67:651-656.
207. Stege H, Roza L, Vink AA, Grewe M, Ruzicka T, Grether-Beck S, Krutmann J. 2000. Enzyme plus light therapy to repair DNA damage in ultraviolet-B- irradiated human skin. PNAS 97:1790-1795.
208. Steinberg ML, Hubbard K, Utti C, Clas B, Hwang BJ, Hill HZ, Orlow I. 2009. Patterns ofpersistent DNA damage associated with sun exposure and the glutathione S-transferase M1 genotype in melanoma patients. Photo Chemistry and Photo-biology 85:379-386.
209. Sturm RA. 1998. Human pigmentation genes and their response to solar UV radiation. Mutation Research 422:69-76.
210. Svobodová A, Zdarilová A, Malisková J, Mikulková H, Walterová D, Vostálová J. 2007. Attenuation of UVA-induced oxidative damage to human keratinocytes by silymarin. Journal of Dermatological Science 46:21-30. (Pubitemid 46366634)
211. Svobodová A. 2006. Effects of natural polyphenols on UV-induced damage to keratinocytes. PhD Thesis. Olomouc: Palacký University.
212. Svobodová A, Walterová D, Vostálová J. 2006. Ultraviolet light induced alteration to the skin. Biomedical papers of the Medical Faculty of the University Palacký, Olomouc, Czecho-slovakia 150:25-38.
213. Syed DN, MalikA, Hadi N, Sarfaraz S, Afaq F, Mukhtar H. 2006. Photochemopreventive effect of pomegranate fruit extract on UVA-mediated activation of cellular pathways in normal human epidermal keratinocytes. Photo Chemistry and Photo-biology 82:398-405.
214. Takahashi H, Hashimoto Y, Aoki N, Kinouchy M, Ishida-Yamamoto A, Izuka H. 2000. Copper, zinc-superoxiddismu-tase protects from ultraviolet B-induced apoptosis of SV-40 transformed human kerationcytes: The protection is associated with the increased levels of antioxidant enzymes. Journal of Dermatological Science 23:12-21.
215. Takeuchi S, Zhang W, Wakamatsu K, Ito S, Hearing VJ, Kraemer KH, Brash DE. 2004. Melanin acts as a potent UVB photosensitizer to cause an atypical mode of cell death in murine skin. Proceedings of the National Academy ofSciences of the USA 101:15076-15081.
216. Tanaka K, Sekiguchi M, Okada Y. 1975. Restoration of ultraviolet-induced unscheduled DNA synthesis of xeroderma pigmentosum cells by the concomitant treatment with bacter-iophage T4 endonuclease V and HVJ (Sendai virus). Proceedings of the National Academy ofSciences of the USA 72:4071-4075.
217. Thiele JJ, Schroeter C, Hsieh SN, Podda M, Parker L. 2001. The antioxidant network of the stratum corneum. In: Thiele JJ, Elsner P, editors. Current problems in Dermatology, Vol. 29. Basel: Karger.
218. Thiele JJ, Traber MG, Packer L. 1998. Depletion of stratum corneum vitamin E: An early and sensitive in vivo marker of UV-induced photooxidation. Journal of Investigative Derma-tology 110:756-761.
219. Thody AJ, Higgins EM, Wakamatsu K, Ito S, Burchill SA, Marks JM. 1991. Pheomelanin as well as eumelanin is present in human epidermis. Journal of Investigative Dermatology 97:340-344.
220. Trevithick JR, Shum DT, Redae S, Mitton KP, Norley C, Karlik SJ, Groom AC, Schmidt EE. 1993. Reduction of sunburn damage to skin by topical application of vitamin E acetate following exposure to ultraviolet B radiation: Effect ofdelaying application or of reducing concentration of vitamin E acetate applied. Scanning Microscopy 7:1269-1281.
221. Tsatmali M, Ancans J, Thody AJ. 2002. Melanocyte function and its control by melanocortin peptides. Journal of Histochemistry and Cytochemistry 50:125-133.
222. Tyrrell RM. 2004. Solar ultraviolet A radiation: An oxidizing skin carcinogen that activates heme oxygenase-1. Antioxidants and Redox Signaling 6:835-840.
223. Van der Broeke LT, Beijersbergen van Henegouwen GM. 1995. Topically applied N-acetylcysteine as a protector against UVB-induced systemic immunosuppression. Journal of Photo Chemistry and Photobiology B Biology 27:61-65.
224. Van der Kemp PA, Blais JC, Bazin M, Boiteux S, Santus R. 2002. Ultraviolet-B-induced inactivation of human OGG1, the repair enzyme for removal of 8-oxoguanine in DNA. Photo Chemistry and Photobiology 76:640-648.
225. Vayalil PK, Elmets CA, Katiyar SK. 2003. Treatment of green tea polyphenols in hydrophilic cream prevents UVB-induced oxidation of lipids and proteins, depletion of antioxidant enzymes and phosphorylation of MAPK proteins in SKH-1 hairless mouse skin. Carcinogenesis 24:927-936.
226. Verschooten LS, Claerhout S, Van Laethem A, Agostinis P, Garmyn M. 2006. New strategies of photoprotection. Photo-chemistry and Photobiology 82:1016-1023.
227. Vignot S, Faivre S, Aguirre D, Raymond E. 2005. mTOR-targeted therapy of cancer with rapamycin derivatives. Annals of Oncology 16:525-537.
228. Wang SQ, Stanfiel JW, Osterwalder U. 2008. In vitro assessment of UVA protection by popular sunscreens available in the United States. Journal of the American Academy of Dermatol-ogy 59:934-942.
229. Weller R. 2003. Nitric oxide: A key mediator in cutaneous physiology. Clinical and Experimental Dermatology 28:511-514.
230. Wenczl E, Pool S, Timmerman A, van der Schans G, Roza L, Schothorst A. 1997. Physiological doses ofultraviolet irradia-tion induce DNA strand breaks in cultured human melano-cytes, as detected by means of an immunochemical assay. Photo Chemistry and Photobiology 66:826-830.
231. Werninghaus K, Meydani M, Bhawan J, Margolis R, Blumberg JB, Gilchrest BA. 1994. Evaluation of the photoprotective effect of oral vitamin E supplementation. Archives of Dermatology 130:1257-1261.
232. Whiteman DC, Parsons PG, Green AC 1999. Determinants of melanocyte density in adult human skin. Archives of Dermatological Research 291:511-516.
233. Wilhelm D, Bender K, Knebel A, Angel P. 1997. The level of intracellular glutathione is a key regulator for the induction of stress-activated signal transduction pathways including Jun N-terminal protein kinases and p38 kinase by alkylating agents. Molecular and Cellular Biology 17:4792-4800.
234. Winkel-Shirley B. 2002. Biosynthesis offlavonoids and effects of stress. Current Opinion in Plant Biology 5:218-223.
235. Witte MB, Barbul A. 2002. Role ofnitric oxide in wound repair. American Journal ofSurgery 183:406-412.
236. Wiswedel I, Keihoff G, Dörner L, Navaro A, Böckelmann R, Bonnekoh B, Gardemann A Gollnick H. 2007. UVB irradiation-induced impairment of keratinocytes and adaptive responses to oxidative stress. Free Radical Research 41:1017-1027.
237. Wolf P, Maier H, Müllegger RR, Chadwick CA, Hofmann-WellenhofR, Soyer HP, Hofer A, Smolle J, Cerroni L, Yarosh D, Klein J, Bucana C, Dunner K, Potten CS, Hönigsmann H, Kerl H, Kripke ML. 2000. Topical treatment with liposomes containing T4 endonuclease V protects human skin in vivo from ultraviolet-induced upregulation of interleukin-10 and tumor necrosis factor-alpha. Journal of Investigative Dermatology 114:149-156.
238. Wondrak GT. 2007. Let the sun shine in: Mechanisms and potential for therapeutics in skin photodamage. Current Opinion in Investigational Drugs 8:390-400.
239. Woollons A, Clingen PH, Price ML, Arlett CF, Green MH. 1997. Induction of mutagenic damage in human fibroblasts after exposure to artificial tanning lamps. British Journal of Dermatology 137:687-692.
240. Wulff BC, Schick JS, Thomas-Ahner JM, Kusewitt DF, Yarosh DB, Oberyszyn TM. 2008. Topical treatment with OGG1 enzyme affects UVB-induced skin carcinogenesis. Photo Chemistry and Photobiology 84:317-321.
241. Yaar M, Gilchrest BA. 2001. Ageing and photoageing of keratinocytes and melanocytes. Clinical and Experimental Dermatology 26:583-591.
242. Yaar M, Gilchrest BA. 2007. Photoaging: Mechanisms, preven-tion and therapy. British Journal of Dermatology 157:874-887.
243. Yamaguchi YK, Takahashi BZ, Zmudzka A, Kornhauser A, Miller SA, Tadokoro T, Berens W, Beer JZ, Hearing VJ. 2006. Human skin responses to UV radiation: Pigment in the upper epidermis protects against DNA damage in the lower epidermis and facilitates apoptosis. FASEB Journal 20:1486-1488.
244. Yarosh DB, O'Connorl A, Alas L, Potten C, Wolf P. 1999. Photoprotection by topical DNA repair enzymes: Molecular correlates ofclinical studies. Photo Chemistry and Photobiology 69:136-140.
245. Yarosh DB, Kibitel J, Green L, Spinowitz A. 1991. Enhanced unscheduled DNA synthesis in UV-irradiated human skin explants treated with T4N5 liposomes. Journal of Investigative Dermatology 97:147-150.
246. Yarosh DB, KleinJ, O'Connor A, HawkJ, Rafal E, Wolf P. 2001. Effect of topically applied T4 endonuclease V in liposomes on skin cancer in xeroderma pigmentosum: A randomised study. The Lancet 357:926-929.
247. Yarosh DB, Tsimis J, Yee V. 1989. Enhacement of DNA repair of UV damage in mouse and human skin by liposomes containing a DNA repair enzyme. Journal of the Society of Cosmetic Chemists 41:85-92.
248. Yarosh DB, Alas L, Yee V, Oheryszyn A, Kibitel J, Mitchell D, Rosenstein R, Spinowitz A, Citron M. 1992. Pyrimidine dimer removal enhanced by DNA repair liposomes reduces the incidence of UV skin cancer in mice. Cancer Research 52:4227-4231.
249. Young AR. 2006. Acute effects of UVR on human eyes and skin. Progress in Biophysics and Molecular Biology 92:80-85.
250. Zastrow L, Groth N, Klein F, Kockott D, Lademann J, Renneberg R, Ferrero L. 2009. The missing link-light-induced (280-1,600 nm) free radical formation in human skin. Skin Pharmacology and Physiology 22:31-44.
251. Zhao Q, Barakat BM, Qin S, Ray A, Elp-Mahdy MA, Wani G, Arafa ES, Mir SN, Wang QE, Wani AA. 2008. The p38 mitogen-activated protein kinase augments nucleotide excision repair by mediating DDB2 degradation and chromatin relaxation. Journal of Biological Chemistry 283:32553-32561.
252. Zhevago NA, Samoilova KA, Calderhead RG. 2006. Polychro-matic light similar to the terrestrial solar spectrum without its UV component stimulates DNA synthesis in human peripheral blood lymphocytes in vivo and in vitro. Photo Chemistry and Photobiology 82:1301-1308.
253. Zhevago NA, Samoilova KA, Obolenskaia KD, Sokolov DI. 2005. Changes in cytokine content in the peripheral blood of volunteers after their exposure to polychromatic visible and infrared light. Tsitologiia 47:450-463.
254. Zhou BB, Elledge JS. 2000. The DNA damage response: Putting checkpoints in perspective. Nature 408:433-439.
255. Zhu M, Bowden GT. 2004. Molecular mechanism(s) for UV-B irradiation-induced glutathione depletion in cultured human keratinocytes. Photo Chemistry and Photobiology 80:191-196.
256. Zvyagin AV, Zhao X, Gierden A, Sanchez W, Ross JA Roberts MS 2008. Imaging ofzinc oxide nanoparticle penetration in human skin in vitro and in vivo. Journal of Biomedical Optics 13:064031.