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This alignment established the prevailing paradigm in which erythema serves as a surrogate for mutagenic risk and ultraviolet B radiation (UVB) is responsible for biologically meaningful DNA damage in skin. However, the discovery of delayed or \u201cdark\u201d CPDs, generated through post\u2010exposure biochemical processes such as melanin\u2010dependent chemiexcitation, extends and refines this classical view. It is increasingly well established that CPD formation is not limited to direct photon absorption but can additionally arise through longer\u2010wavelength, redox\u2010driven chemiexcitation pathways, particularly within the UVA range. Emerging evidence further suggests that non\u2010melanin oxidative pathways, including lipid peroxidation, can generate chemiexcited states potentially relevant to DNA damage. In light of this evidence, we propose that the biological action spectrum for CPD formation is determined not only by direct DNA absorption, but also by the capacity of cells to generate DNA damaging excited states through redox chemistry. This novel framework reshapes how light exposure is linked to mutagenesis and underscores the need for next\u2010generation photoprotective strategies targeting both direct and indirect DNA damage pathways.<\/jats:p>","DOI":"10.1111\/php.70154","type":"journal-article","created":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T06:42:38Z","timestamp":1790232158000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Revisiting the action spectrum of\n                    <scp>DNA<\/scp>\n                    damage: Chemiexcitation and the expanded biological spectrum of cyclobutane pyrimidine dimer formation"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3146-9317","authenticated-orcid":false,"given":"Jyoti","family":"Srivastava","sequence":"first","affiliation":[{"name":"Department of Tumor Microenvironment and Metastasis H. Lee Moffitt Cancer Center and Research Institute  Tampa Florida USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0369-862X","authenticated-orcid":false,"given":"Marcus S.","family":"Cooke","sequence":"additional","affiliation":[{"name":"Oxidative Stress Group, Department of Molecular Biosciences University of South Florida  Tampa Florida USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5492-5469","authenticated-orcid":false,"given":"Sanjay","family":"Premi","sequence":"additional","affiliation":[{"name":"Department of Tumor Microenvironment and Metastasis H. Lee Moffitt Cancer Center and Research Institute  Tampa Florida USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"e_1_2_13_2_1","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1126\/science.1135428","article-title":"Thymine dimerization in DNA is an ultrafast photoreaction","volume":"315","author":"Schreier WJ","year":"2007","journal-title":"Science"},{"key":"e_1_2_13_3_1","doi-asserted-by":"crossref","first-page":"3363","DOI":"10.1073\/pnas.71.9.3363","article-title":"The wavelengths in sunlight effective in producing skin cancer: a theoretical analysis","volume":"71","author":"Setlow RB","year":"1974","journal-title":"Proc Natl Acad Sci USA"},{"key":"e_1_2_13_4_1","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1111\/j.1751-1097.2010.00871.x","article-title":"Know your standard: clarifying the CIE erythema action spectrum","volume":"87","author":"Webb AR","year":"2011","journal-title":"Photochem Photobiol"},{"key":"e_1_2_13_5_1","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.pbiomolbio.2006.02.005","article-title":"Acute effects of UVR on human eyes and skin","volume":"92","author":"Young AR","year":"2006","journal-title":"Prog Biophys Mol Biol"},{"key":"e_1_2_13_6_1","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/S1046-2023(02)00205-0","article-title":"Ultraviolet radiation\u2010induced erythema in human skin","volume":"28","author":"Harrison GI","year":"2002","journal-title":"Methods"},{"key":"e_1_2_13_7_1","doi-asserted-by":"crossref","first-page":"982","DOI":"10.1046\/j.1523-1747.1998.00436.x","article-title":"The similarity of action spectra for thymine dimers in human epidermis and erythema suggests that DNA is the chromophore for erythema","volume":"111","author":"Young AR","year":"1998","journal-title":"J Invest Dermatol"},{"key":"e_1_2_13_8_1","doi-asserted-by":"crossref","first-page":"1816","DOI":"10.1039\/c7pp00395a","article-title":"Formation of UV\u2010induced DNA damage contributing to skin cancer development","volume":"17","author":"Cadet J","year":"2018","journal-title":"Photochem Photobiol Sci"},{"key":"e_1_2_13_9_1","first-page":"53","article-title":"Wavelength dependence of skin cancer induction by ultraviolet irradiation of albino hairless mice","volume":"53","author":"Gruijl FR","year":"1993","journal-title":"Cancer Res"},{"key":"e_1_2_13_10_1","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1111\/j.1751-1097.1995.tb03961.x","article-title":"Action spectrum for erythema in humans investigated with dye lasers","volume":"61","author":"Anders A","year":"1995","journal-title":"Photochem Photobiol"},{"key":"e_1_2_13_11_1","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1038\/jid.2011.283","article-title":"UVA1 induces cyclobutane pyrimidine dimers but not 6\u20134 photoproducts in human skin in\u00a0vivo","volume":"132","author":"Tewari A","year":"2012","journal-title":"J Invest Dermatol"},{"key":"e_1_2_13_12_1","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 S","year":"2006","journal-title":"Proc Natl Acad Sci USA"},{"key":"e_1_2_13_13_1","doi-asserted-by":"crossref","first-page":"842","DOI":"10.1126\/science.1256022","article-title":"Photochemistry. 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