{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,21]],"date-time":"2026-04-21T06:54:07Z","timestamp":1776754447974,"version":"3.51.2"},"reference-count":81,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2013,1,18]],"date-time":"2013-01-18T00:00:00Z","timestamp":1358467200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/3.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Exposure to sunlight is the major cause of skin cancer. Ultraviolet radiation (UV) from the sun causes damage to DNA by direct absorption and can cause skin cell death. UV also causes production of reactive oxygen species that may interact with DNA to indirectly cause oxidative DNA damage. UV increases accumulation of p53 in skin cells, which upregulates repair genes but promotes death of irreparably damaged cells.  A benefit of sunlight is vitamin D, which is formed following exposure of  7-dehydrocholesterol in skin cells to UV. The relatively inert vitamin D is metabolized to various biologically active compounds, including 1,25-dihydroxyvitamin D3. Therapeutic use of vitamin D compounds has proven beneficial in several cancer types, but more recently these compounds have been shown to prevent UV-induced cell death and DNA damage in human skin cells. Here, we discuss the effects of vitamin D compounds in skin cells that have been exposed to UV. Specifically, we examine the various signaling pathways involved in the vitamin D-induced protection of skin cells from UV.<\/jats:p>","DOI":"10.3390\/ijms14011964","type":"journal-article","created":{"date-parts":[[2013,1,18]],"date-time":"2013-01-18T11:04:23Z","timestamp":1358507063000},"page":"1964-1977","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":38,"title":["Vitamin D and Death by Sunshine"],"prefix":"10.3390","volume":"14","author":[{"given":"Katie","family":"Dixon","sequence":"first","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"},{"name":"Discipline of Anatomy and Histology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wannit","family":"Tongkao-On","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Vanessa","family":"Sequeira","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sally","family":"Carter","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eric","family":"Song","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mark","family":"Rybchyn","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Clare","family":"Gordon-Thomson","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Rebecca","family":"Mason","sequence":"additional","affiliation":[{"name":"Discipline of Physiology, Bosch Institute, School of Medical Sciences, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2013,1,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1016\/j.taap.2003.08.019","article-title":"Toxic effects of ultraviolet radiation on the skin","volume":"195","author":"Matsumura","year":"2004","journal-title":"Toxicol. Appl. Pharmacol"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"564","DOI":"10.1039\/b9pp00184k","article-title":"Vitamin D-fence","volume":"9","author":"Dixon","year":"2010","journal-title":"Photochem. Photobiol. Sci"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1111\/1523-1747.ep12479237","article-title":"The cutaneous photosynthesis of previtamin D3: A unique photoendocrine system","volume":"77","author":"Holick","year":"1981","journal-title":"J. Invest. Dermatol"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1545","DOI":"10.1021\/bi00355a013","article-title":"Neonatal human foreskin keratinocytes produce 1,25-dihydroxyvitamin D3","volume":"25","author":"Bikle","year":"1986","journal-title":"Biochemistry"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1034\/j.1600-0625.2000.009002097.x","article-title":"Conversion of vitamin D3 to 1alpha,25-dihydroxyvitamin D3 in human skin equivalents","volume":"9","author":"Lehmann","year":"2000","journal-title":"Exp. Dermatol"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Pattison, D.I., and Davies, M.J. (2006). Actions of ultraviolet light on cellular structures. Cancer, 131\u2013157.","DOI":"10.1007\/3-7643-7378-4_6"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1111\/j.1476-5381.1995.tb13341.x","article-title":"Release by ultraviolet b (U.V.B) radiation of nitric oxide (no) from human keratinocytes: A potential role for nitric oxide in erythema production","volume":"114","author":"Deliconstantinos","year":"1995","journal-title":"Br. J. Pharmacol"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1046\/j.1523-1747.1998.00084.x","article-title":"Nitric oxide in human skin: Current status and future prospects","volume":"110","author":"Ruzicka","year":"1998","journal-title":"J. Invest. Dermatol"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1016\/j.niox.2004.04.005","article-title":"Nitric oxide function in the skin","volume":"10","author":"Ormerod","year":"2004","journal-title":"Nitric Oxide"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.freeradbiomed.2004.11.018","article-title":"Enzyme-independent nitric oxide formation during UVA challenge of human skin: Characterization, molecular sources, and mechanisms","volume":"38","author":"Paunel","year":"2005","journal-title":"Free Radic. Biol. Med"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"834","DOI":"10.1038\/jid.2008.296","article-title":"Enzyme-independent no stores in human skin: Quantification and influence of UV radiation","volume":"129","author":"Mowbray","year":"2009","journal-title":"J. Invest. Dermatol"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"E46","DOI":"10.1038\/35055152","article-title":"S-nitrosylation: Spectrum and specificity","volume":"3","author":"Hess","year":"2001","journal-title":"Nat. Cell Biol"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1016\/j.mrfmmm.2004.09.013","article-title":"Inflammation, gene mutation and photoimmunosuppression in response to UVR-induced oxidative damage contributes to photocarcinogenesis","volume":"571","author":"Halliday","year":"2005","journal-title":"Mutat. Res"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1089\/ars.2006.8.1033","article-title":"8-nitroguanine, a product of nitrative DNA damage caused by reactive nitrogen species: Formation, occurrence, and implications in inflammation and carcinogenesis","volume":"8","author":"Ohshima","year":"2006","journal-title":"Antioxid. Redox Signal"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1152\/physrev.00029.2006","article-title":"Nitric oxide and peroxynitrite in health and disease","volume":"87","author":"Pacher","year":"2007","journal-title":"Physiol. Rev"},{"key":"ref_16","first-page":"375","article-title":"The therapeutic potential of poly(adp-ribose) polymerase inhibitors","volume":"54","year":"2002","journal-title":"Physiol. Rev"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1179\/135100000101535447","article-title":"UVB-induced experimental carcinogenesis: Dysregulation of apoptosis and p53 signalling pathway","volume":"5","author":"Ouhtit","year":"2000","journal-title":"Redox Report"},{"key":"ref_18","first-page":"184","article-title":"Inflammatory cytokines induce DNA damage and inhibit DNA repair in cholangiocarcinoma cells by a nitric oxide-dependent mechanism","volume":"60","author":"Jaiswal","year":"2000","journal-title":"Cancer Res"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1093\/carcin\/22.5.709","article-title":"Nitric oxide is involved in arsenite inhibition of pyrimidine dimer excision","volume":"22","author":"Bau","year":"2001","journal-title":"Carcinogenesis"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"2861","DOI":"10.1038\/sj.onc.1207523","article-title":"Toxic proteins released from mitochondria in cell death","volume":"23","author":"Saelens","year":"2004","journal-title":"Oncogene"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1016\/0960-0760(96)00055-6","article-title":"1,25-dihydroxyvitamin D3 induces morphological and biochemical markers of apoptosis in mcf-7 breast cancer cells","volume":"58","author":"Narvaez","year":"1996","journal-title":"J. Steroid. Biochem. Mol. Biol"},{"key":"ref_22","first-page":"667","article-title":"Apoptosis induction by 1\u03b1,25-dihydroxyvitamin D3 in prostate cancer","volume":"1","author":"Guzey","year":"2002","journal-title":"Mol. Cancer Ther"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/0304-3835(95)03958-Y","article-title":"Effects of vitamin D3 derivatives on growth, differentiation and apoptosis in tumoral colonic ht 29 cells: Possible implication of intracellular calcium","volume":"97","author":"Vandewalle","year":"1995","journal-title":"Cancer Lett"},{"key":"ref_24","unstructured":"Norman, A., Bouillon, R., and Thomasset, M. (2000). Vitamin D Endocrine System: Structural, Biological, Genetic and Clinical Aspects, University of California."},{"key":"ref_25","unstructured":"Tsambos, D., and Merk, H. (2002). Modern Trends in Skin Pharmacology, Parissianos Medical Publications S.A. Athens. [1st ed]."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1002\/jcb.10540","article-title":"1,25-dihydroxyvitamin D3 inhibits ultraviolet B-induced apoptosis, jun kinase activation, and interleukin-6 production in primary human keratinocytes","volume":"89","author":"Garmyn","year":"2003","journal-title":"J. Cell Biochem"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1007\/s00403-004-0451-x","article-title":"Two 14-epi analogues of 1,25-dihydroxyvitamin D3 protect human keratinocytes against the effects of UVB","volume":"295","author":"Garmyn","year":"2004","journal-title":"Arch. Dermatol. Res"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1016\/j.jsbmb.2004.03.072","article-title":"1,25-dihydroxyvitamin D and three low-calcemic analogs decrease UV-induced DNA damage via the rapid response pathway","volume":"89\u201390","author":"Wong","year":"2004","journal-title":"J. Steroid. Biochem. Mol. Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1038\/sj.jid.5700597","article-title":"Photoprotection by 1,25 dihydroxyvitamin D3 is associated with an increase in p53 and a decrease in nitric oxide products","volume":"127","author":"Gupta","year":"2007","journal-title":"J. Invest. Dermatol"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1241","DOI":"10.1046\/j.0022-202x.2001.01496.x","article-title":"1\u03b1,25-dihydroxyvitamin D-3 protects human keratinocytes from apoptosis by the formation of sphingosine-1-phosphate","volume":"117","author":"Manggau","year":"2001","journal-title":"J. Invest. Dermatol"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/S0923-1811(98)00015-2","article-title":"The photoprotective effect of 1,25-dihydroxyvitamin D3 on ultraviolet light B-induced damage in keratinocyte and its mechanism of action","volume":"18","author":"Lee","year":"1998","journal-title":"J. Dermatol. Sci"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"276","DOI":"10.1111\/1523-1747.ep12335756","article-title":"1,25-dihydroxyvitamin D3, transforming growth factor beta1, calcium, and ultraviolet B radiation induce apoptosis in cultured human keratinocytes","volume":"109","author":"Benassi","year":"1997","journal-title":"J. Invest. Dermatol"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1111\/j.1600-0781.1997.tb00126.x","article-title":"Photoprotective effect of calcipotriol upon skin photoreaction to UVA and UVB","volume":"13","author":"Youn","year":"1997","journal-title":"Photodermatol. Photoimmunol. Photomed"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1825","DOI":"10.1039\/c2pp25208b","article-title":"Differential photoprotective effects of 1,25-dihydroxyvitamin D3 and a low calcaemic deltanoid","volume":"11","author":"Dixon","year":"2012","journal-title":"Photochem. Photobiol. Sci"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1039\/b108291d","article-title":"The sunburn cell revisited: An update on mechanistic aspects","volume":"1","author":"Sheehan","year":"2002","journal-title":"Photochem. Photobiol. Sci"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1039\/b504970a","article-title":"Pathways involved in sunburn cell formation: Deregulation in skin cancer","volume":"5","author":"Claerhout","year":"2006","journal-title":"Photochem. Photobiol. Sci"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/0923-1811(94)00378-R","article-title":"Possible role of 1,25-dihydroxyvitamin D3-induced metallothionein in photoprotection against UVB injury in mouse skin and cultured rat keratinocytes","volume":"9","author":"Hanada","year":"1995","journal-title":"J. Dermatol. Sci"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1016\/j.jsbmb.2006.11.016","article-title":"In vivo relevance for photoprotection by the vitamin D rapid response pathway","volume":"103","author":"Dixon","year":"2007","journal-title":"J. Steroid. Biochem. Mol. Biol"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1485","DOI":"10.1158\/1940-6207.CAPR-11-0165","article-title":"1\u03b1,25(OH)2-vitamin D and a nongenomic vitamin D analogue inhibit ultraviolet radiation-induced skin carcinogenesis","volume":"4","author":"Dixon","year":"2011","journal-title":"Cancer Prev. Res"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"E23","DOI":"10.1111\/j.1600-0625.2009.00955.x","article-title":"Topical calcitriol protects from UV-induced genetic damage but suppresses cutaneous immunity in humans","volume":"19","author":"Damian","year":"2010","journal-title":"Exp. Dermatol"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1016\/j.jsbmb.2010.03.082","article-title":"Photoprotection by 1\u03b1,25-dihydroxyvitamin D and analogs: Further studies on mechanisms and implications for UV-damage","volume":"121","author":"Mason","year":"2010","journal-title":"J. Steroid. Biochem. Mol. Biol"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1038\/ejcn.2011.105","article-title":"Vitamin D: The light side of sunshine","volume":"65","author":"Mason","year":"2011","journal-title":"Eur. J. Clin. Nut"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Song, E.J., Gordon-Thomson, C., Coleb, L., Sternc, H., Halliday, G.M., Damian, D.L., Reeve, V.E., and Mason, R.S. (2012). 1\u03b1,25-dihydroxyvitamin D3 reduces several types of UV-induced DNA damage and contributes to photoprotection. J. Steroid. Biochem. Mol. Biol., submitted for publication.","DOI":"10.1016\/j.jsbmb.2012.11.003"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Gordon-Thomson, C., Gupta, R., Tongkao-on, W., Ryan, A., Halliday, G.M., and Mason, R.S. (2012). 1\u03b1,25 dihydroxyvitamin D3 enhances cellular defences against UV-induced oxidative and other forms of DNA damage in skin. Photochem. Photobiol. Sci.","DOI":"10.1039\/c2pp25202c"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1210\/me.2011-1161","article-title":"The role of the vitamin D receptor and ERp57 in photoprotection by 1\u03b1,25-dihydroxyvitamin D-3","volume":"26","author":"Sequeira","year":"2012","journal-title":"Mol. Endocrinol"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"11678","DOI":"10.1074\/jbc.275.16.11678","article-title":"Formation of the main UV-induced thymine dimeric lesions within isolated and cellular DNA as measured by high performance liquid chromatography-tandem mass spectrometry","volume":"275","author":"Douki","year":"2000","journal-title":"J. Biol. Chem"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/S0022-1759(03)00269-2","article-title":"Immunochemical detection of UV-induced DNA damage and repair","volume":"280","author":"Cooke","year":"2003","journal-title":"J. Immunol. Methods"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"13765","DOI":"10.1073\/pnas.0604213103","article-title":"Cyclobutane pyrimidine dimers are predominant DNA lesions in whole human skin exposed to UVA radiation","volume":"103","author":"Mouret","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2379","DOI":"10.1093\/carcin\/18.12.2379","article-title":"Induction of oxidative DNA base damage in human skin cells by UV and near visible radiation","volume":"18","author":"Kvam","year":"1997","journal-title":"Carcinogenesis"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"4954","DOI":"10.1073\/pnas.0401141101","article-title":"The basal layer in human squamous tumors harbors more UVA than UVB fingerprint mutations: A role for UVA in human skin carcinogenesis","volume":"101","author":"Agar","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/j.jphotobiol.2004.09.010","article-title":"1,25-dihydroxyvitamin D3 and analogues protect primary human keratinocytes against UVB-induced DNA damage","volume":"78","author":"Garmyn","year":"2005","journal-title":"J. Photochem. Photobiol. B"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/j.jsbmb.2005.06.006","article-title":"Skin cancer prevention: A possible role of 1,25dihydroxyvitamin D3 and its analogs","volume":"97","author":"Dixon","year":"2005","journal-title":"J. Steroid. Biochem. Mol. Biol"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Sequeira, V.B., Rybchyn, M.S., Gordon-Thomson, C., Tongkao-On, W., Mizwicki, M.T., Norman, A.W., Reeve, V.E., Halliday, G.M., and Mason, R.S. (2012). Opening of chloride channels by 1\u03b1,25-dihydroxyvitamin D(3) contributes to photoprotection against UVR-induced thymine dimers in keratinocytes. J. Invest. Dermatol.","DOI":"10.1038\/jid.2012.343"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2508","DOI":"10.1038\/jid.2008.131","article-title":"Inactivation of the vitamin D receptor enhances susceptibility of murine skin to UV-induced tumorigenesis","volume":"128","author":"Ellison","year":"2008","journal-title":"J. Invest. Dermatol"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1016\/j.steroids.2005.09.012","article-title":"1\u03b1,25(OH)2 vitamin D3 actions on ion channels in osteoblasts","volume":"71","author":"Zanello","year":"2006","journal-title":"Steroids"},{"key":"ref_56","first-page":"203","article-title":"High levels of p53 protein in UV-irradiated normal human skin","volume":"8","author":"Hall","year":"1993","journal-title":"Oncogene"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"916","DOI":"10.1111\/j.1600-0625.2008.00722.x","article-title":"Cyclobutane pyrimidine dimer formation and p53 production in human skin after repeated UV irradiation","volume":"17","author":"Yamaguchi","year":"2008","journal-title":"Exp. Dermatol"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1002\/jcb.21061","article-title":"Expression profiling of vitamin D treated primary human keratinocytes","volume":"100","author":"Moll","year":"2007","journal-title":"J. Cell Biochem"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"843","DOI":"10.1093\/carcin\/bgg031","article-title":"P53 responsive nucleotide excision repair gene products p48 and xpc, but not p53, localize to sites of UV-irradiation-induced DNA damage, in vivo","volume":"24","author":"Fitch","year":"2003","journal-title":"Carcinogenesis"},{"key":"ref_60","first-page":"37","article-title":"UVA induces immediate and enzyme-independent nitric oxide formation in healthy human skin leading to no-specific signalling","volume":"84","author":"Dejam","year":"2005","journal-title":"Eur. J. Cell Biol"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/978-1-60327-530-9_1","article-title":"Immunohistochemical Analysis of 8-Nitroguanine, a Nitrative DNA Lesion, in Relation to Inflammation-Associated Carcinogenesis","volume":"512","author":"Kozlov","year":"2009","journal-title":"Inflammation and Cancer"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1034\/j.1600-0625.2001.100407.x","article-title":"Solar simulated irradiation modulates gene expression and activity of antioxidant enzymes in cultured human dermal fibroblasts","volume":"10","author":"Leccia","year":"2001","journal-title":"Exp. Dermatol"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"525","DOI":"10.1677\/joe.0.1730525","article-title":"Vitamin D inhibits the activation of stress-activated protein kinases by physiological and environmental stresses in keratinocytes","volume":"173","author":"Ravid","year":"2002","journal-title":"J. Endocrinol"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1196\/annals.1299.064","article-title":"Vitamin D protects keratinocytes from apoptosis induced by osmotic shock, oxidative stress, and tumor necrosis factor","volume":"1010","author":"Koren","year":"2003","journal-title":"Ann. N. Y. Acad. Sci"},{"key":"ref_65","first-page":"111","article-title":"Protective effects of cadmium chloride against UVB injury in mouse skin and in cultured human cells: A possible role of cadmium-induced metallothionein","volume":"8","author":"Hanada","year":"1991","journal-title":"Photodermatol. Photoimmunol. Photomed"},{"key":"ref_66","first-page":"209","article-title":"Possible role of cutaneous metallothionein in protection against photo-oxidative stress-epidermal localization and scavenging activity for superoxide and hydroxyl radicals","volume":"9","author":"Hanada","year":"1992","journal-title":"Photodermatol. Photoimmunol. Photomed"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1046\/j.1365-2567.2000.00026.x","article-title":"Lack of metallothionein-i and -ii exacerbates the immunosuppressive effect of ultraviolet B radiation and cis-urocanic acid in mice","volume":"100","author":"Reeve","year":"2000","journal-title":"Immunology"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1038\/sj.jid.5700013","article-title":"Isoflavonoid photoprotection in mouse and human skin is dependent on metallothionein","volume":"126","author":"Widyarini","year":"2006","journal-title":"J. Invest. Dermatol"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"8810","DOI":"10.1073\/pnas.84.24.8810","article-title":"Regulation of metallothionein gene expression by 1\u03b1,25-dihydroxyvitamin D3 in cultured cells and in mice","volume":"84","author":"Karasawa","year":"1987","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_70","first-page":"90","article-title":"Ultraviolet radiation-induced apoptosis in keratinocytes: On the role of cytosolic factors","volume":"1755","author":"Assefa","year":"2005","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"RE2","DOI":"10.1126\/stke.2003.167.re2","article-title":"Mitogen-activated protein kinase activation in UV-induced signal transduction","volume":"2003","author":"Bode","year":"2003","journal-title":"Sci. STKE"},{"key":"ref_72","unstructured":"Gardner, J. (2005). Mechanisms of photoprotection by 1a,25-dihydroxyvitamin D3 in keratinocytes. [Honours Thesis, University of Sydney]."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"13681","DOI":"10.1073\/pnas.251194298","article-title":"Sp600125, an anthrapyrazolone inhibitor of jun N-terminal kinase","volume":"98","author":"Bennett","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1046\/j.1523-1747.1999.00584.x","article-title":"UVB activates erk1\/2 and p38 signaling pathways via reactive oxygen species in cultured keratinocytes","volume":"112","author":"Peus","year":"1999","journal-title":"J. Invest. Dermatol"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"45737","DOI":"10.1074\/jbc.M300574200","article-title":"Epidermal growth factor receptor-dependent, NF-\u03baB-independent activation of the phosphatidylinositol 3-kinase\/akt pathway inhibits ultraviolet irradiation-induced caspases-3, -8, and -9 in human keratinocytes","volume":"278","author":"Wang","year":"2003","journal-title":"J. Biol. Chem"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1007\/s10495-006-5115-1","article-title":"Programmed cell death of stressed keratinocytes and its inhibition by vitamin D: The role of death and survival signaling pathways","volume":"11","author":"Koren","year":"2006","journal-title":"Apoptosis"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1002\/jcb.20227","article-title":"Molecular pathways involved in the anti-apoptotic effect of 1,25-dihydroxyvitamin D3 in primary human keratinocytes","volume":"93","author":"Garmyn","year":"2004","journal-title":"J. Cell Biochem"},{"key":"ref_78","unstructured":"Sequeira, V.B. (2011). Studies on the mechanisms of vitamin D compounds. [Ph.D. Thesis, University of Sydney]."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"23771","DOI":"10.1074\/jbc.M111.251116","article-title":"An akt-dependent increase in canonical wnt signaling and a decrease in sclerostin protein levels are involved in strontium ranelate-induced osteogenic effects in human osteoblasts","volume":"286","author":"Rybchyn","year":"2011","journal-title":"J. Biol. Chem"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.mce.2011.05.017","article-title":"Vitamin D metabolism and function in the skin","volume":"347","author":"Bikle","year":"2011","journal-title":"Mol. Cell Endocrinol"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"7516","DOI":"10.1073\/pnas.89.16.7516","article-title":"Pyrimidine dimers in DNA initiate systemic immunosuppression in UV-irradiated mice","volume":"89","author":"Kripke","year":"1992","journal-title":"Proc. Natl. Acad. Sci. USA"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/14\/1\/1964\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T21:44:24Z","timestamp":1760219064000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/14\/1\/1964"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2013,1,18]]},"references-count":81,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2013,1]]}},"alternative-id":["ijms14011964"],"URL":"https:\/\/doi.org\/10.3390\/ijms14011964","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2013,1,18]]}}}