{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T06:47:18Z","timestamp":1785826038517,"version":"3.56.0"},"reference-count":63,"publisher":"MDPI AG","issue":"4","license":[{"start":{"date-parts":[[2026,2,21]],"date-time":"2026-02-21T00:00:00Z","timestamp":1771632000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Background: The interaction between keratinocytes and proinflammatory cytokines is essential in the development of psoriatic lesions. The synergism among these cytokines and their involvement in ferroptosis are not yet elucidated. This study aimed at evaluating the early impact of a complete proinflammatory microenvironment on keratinocyte differentiation, intercellular adhesion, proliferation, and induction of ferroptosis. Methods: HaCaT cells were differentiated with 1.8 mM CaCl2 and treated with a cytokine combination (MIX) containing IL-17A, IL-22, IL-23, and TNF-alpha for 24 and 48 h. Claudin 1 (CLDN-1), Zonula Occludens 1 (ZO-1), and keratins (K)10\/K14 expression was analyzed by immunofluorescence and immunoblot analysis, paralleled by proliferation and ultrastructural analysis. Ferroptosis was induced with erastin and RSL3 and evaluated by testing glutathione (GSH)\/glutathione peroxidase 4 (GPX4) protein expression, GSH levels, cell availability\/toxicity, intracellular iron and ATP levels. Results: After MIX incubation at T48, CLDN-1 and ZO-1 immunofluorescences were reduced in HaCaT cells, while K10 and K14 were unaffected. The proliferative activity was reduced. Psoriatic-like MIX triggered the ferroptotic pathway, as shown by the increase in intracellular iron levels as well as by the reduction in GPX4 protein expression, the decrease in GSH levels, cell availability, and ATP levels. Conclusions: This experimental model mimics the early pathogenetic processes underlying psoriatic plaque formation\/progression paving the way for new therapeutic strategies.<\/jats:p>","DOI":"10.3390\/biology15040362","type":"journal-article","created":{"date-parts":[[2026,2,23]],"date-time":"2026-02-23T10:00:52Z","timestamp":1771840852000},"page":"362","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["A Proinflammatory Psoriatic Microenvironment Has Early Effects on Keratinocyte Proliferation\/Differentiation and Induces Ferroptosis in HaCaT Cells"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9000-1696","authenticated-orcid":false,"given":"Federica","family":"Riva","sequence":"first","affiliation":[{"name":"Department of Public Health, Experimental and Forensic Medicine, Histology and Embryology Unit, University of Pavia, 27100 Pavia, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elena","family":"Gammella","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, University of Milan, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1187-3988","authenticated-orcid":false,"given":"Margherita","family":"Correnti","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, University of Milan, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-2505-4122","authenticated-orcid":false,"given":"Davide","family":"Daluiso","sequence":"additional","affiliation":[{"name":"Department of Public Health, Experimental and Forensic Medicine, Histology and Embryology Unit, University of Pavia, 27100 Pavia, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5997-2045","authenticated-orcid":false,"given":"Francesca","family":"Prignano","sequence":"additional","affiliation":[{"name":"Department of Health Sciences, Section of Dermatology, University of Florence, 50125 Florence, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefania","family":"Recalcati","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, University of Milan, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5634-5863","authenticated-orcid":false,"given":"Elena","family":"Donetti","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, University of Milan, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,2,21]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1111\/exd.12832","article-title":"The dynamic anatomy and patterning of skin","volume":"25","author":"Wong","year":"2016","journal-title":"Exp. Dermatol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"12222","DOI":"10.3390\/ijms140612222","article-title":"UV radiation and the skin","volume":"14","author":"Jarrett","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1824624","DOI":"10.1155\/2019\/1824624","article-title":"The Role of Toll-Like Receptors in Skin Host Defense, Psoriasis, and Atopic Dermatitis","volume":"2019","author":"Sun","year":"2019","journal-title":"J. Immunol. Res."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Yin, J., Zhang, S., Yang, C., Wang, Y., Shi, B., Zheng, Q., Zeng, N., and Huang, H. (2022). Mechanotransduction in skin wound healing and scar formation: Potential therapeutic targets for controlling hypertrophic scarring. Front. Immunol., 13.","DOI":"10.3389\/fimmu.2022.1028410"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"NP19","DOI":"10.1177\/12034754241227623","article-title":"Interleukins and Psoriasis","volume":"28","author":"Zhu","year":"2024","journal-title":"J. Cutan. Med. Surg."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1038\/s41577-018-0084-5","article-title":"The immunological anatomy of the skin","volume":"19","author":"Kabashima","year":"2019","journal-title":"Nat. Rev. Immunol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1056\/NEJMra0804595","article-title":"Psoriasis","volume":"361","author":"Nestle","year":"2009","journal-title":"N. Engl. J. Med."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.cytogfr.2014.10.009","article-title":"An inflammatory triangle in psoriasis: TNF, type I IFNs and IL-17","volume":"26","author":"Grine","year":"2015","journal-title":"Cytokine Growth Factor. Rev."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1159\/000524119","article-title":"Looking at Interleukin-22 from a New Dermatological Perspective: From Epidermal Homeostasis to Its Role in Chronic Skin Diseases","volume":"238","author":"Prignano","year":"2022","journal-title":"Dermatology"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1242\/dmm.009092","article-title":"Putting together the psoriasis puzzle: An update on developing targeted therapies","volume":"5","author":"Lowes","year":"2012","journal-title":"Dis. Model. Mech."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"3218","DOI":"10.4081\/ejh.2021.3218","article-title":"Evaluation of expression of Toll-Like Receptors 7 and 9, proliferation, and cytoskeletal biomarkers in plaque and guttate psoriasis: A pilot morphological study","volume":"65","author":"Prignano","year":"2021","journal-title":"Eur. J. Histochem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1038\/onc.2010.456","article-title":"Keratins in health and cancer: More than mere epithelial cell markers","volume":"30","author":"Karantza","year":"2011","journal-title":"Oncogene"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Bouchard, C., Grenier, A., Cardinal, S., B\u00e9langer, S., Voyer, N., and Pouliot, R. (2022). Antipsoriatic Potential of Quebecol and Its Derivatives. Pharmaceutics, 14.","DOI":"10.3390\/pharmaceutics14061129"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1515\/bmc.2011.060","article-title":"Keratin function and regulation in tissue homeostasis and pathogenesis","volume":"3","author":"Roth","year":"2012","journal-title":"Biomol. Concepts"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1016\/j.semcdb.2014.08.011","article-title":"Architecture of tight junctions and principles of molecular composition","volume":"36","author":"Anderson","year":"2014","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1301","DOI":"10.1016\/j.ajpath.2017.02.006","article-title":"Tight Junction Proteins Claudin-1 and Occludin Are Important for Cutaneous Wound Healing","volume":"187","author":"Volksdorf","year":"2017","journal-title":"Am. J. Pathol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1095","DOI":"10.2353\/ajpath.2009.080973","article-title":"Alteration of tight junction proteins is an early event in psoriasis: Putative involvement of proinflammatory cytokines","volume":"175","author":"Kirschner","year":"2009","journal-title":"Am. J. Pathol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"673\u2013681; quiz 682-673","DOI":"10.1590\/S0365-05962012000500001","article-title":"Tumor necrosis factor-alpha and the cytokine network in psoriasis","volume":"87","author":"Brotas","year":"2012","journal-title":"An. Bras. Dermatol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1177\/1753425919852156","article-title":"The contribution of IL-17 to the development of autoimmunity in psoriasis","volume":"25","author":"Furue","year":"2019","journal-title":"Innate Immun."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"5085","DOI":"10.1038\/s41598-018-23510-3","article-title":"Regulation of IL-22BP in psoriasis","volume":"8","author":"Voglis","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1007\/s00011-020-01339-9","article-title":"Interleukin 23 and autoimmune diseases: Current and possible future therapies","volume":"69","author":"Abdo","year":"2020","journal-title":"Inflamm. Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"5263","DOI":"10.4049\/jimmunol.0902464","article-title":"Skin Inflammation Induced by the Synergistic Action of IL-17A, IL-22, Oncostatin M, IL-1{alpha}, and TNF-{alpha} Recapitulates Some Features of Psoriasis","volume":"184","author":"Guilloteau","year":"2010","journal-title":"J. Immunol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Rabeony, H., Petit-Paris, I., Garnier, J., Barrault, C., Pedretti, N., Guilloteau, K., Jegou, J.F., Guillet, G., Huguier, V., and Lecron, J.C. (2014). Inhibition of keratinocyte differentiation by the synergistic effect of IL-17A, IL-22, IL-1\u03b1, TNF\u03b1 and oncostatin M. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0101937"},{"key":"ref_24","first-page":"e61537","article-title":"Establishing Transcription Profile of Psoriasiform Cutaneous In Vitro using HaCaT Cells Stimulated with Combination of Cytokines","volume":"169","author":"Zheng","year":"2021","journal-title":"J. Vis. Exp."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Zima, K., Purzycka-Bohdan, D., Szczerkowska-Dobosz, A., and Gabig-Cimi\u0144ska, M. (2024). Keratinocyte-Mediated Antigen Presentation in Psoriasis: Preliminary Insights from In Vitro Studies. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms252413387"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"115021","DOI":"10.1016\/j.intimp.2025.115021","article-title":"Psoriasis unveiled: The cellular ballet, molecular symphony, and genetic puzzle","volume":"161","author":"Sharma","year":"2025","journal-title":"Int. Immunopharmacol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1517\/14728222.2014.889686","article-title":"Role of IL-23 in the pathogenesis of psoriasis: A novel potential therapeutic target?","volume":"18","author":"Chiricozzi","year":"2014","journal-title":"Expert. Opin. Ther. Targets"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Li, S., Luo, X., Zhang, S., Su, Y., Deng, M., Zhu, Y., Zhang, P., Wu, R., and Zhao, M. (2023). Ferroptosis Activation Contributes to the Formation of Skin Lesions in Psoriasis Vulgaris. Antioxidants, 12.","DOI":"10.3390\/antiox12020310"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"e183219","DOI":"10.1172\/JCI183219","article-title":"Ferroptosis of select skin epithelial cells initiates and maintains chronic systemic immune-mediated psoriatic disease","volume":"135","author":"Vats","year":"2024","journal-title":"J. Clin. Investig."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1009","DOI":"10.1038\/s41419-021-04284-5","article-title":"Inhibition of keratinocyte ferroptosis suppresses psoriatic inflammation","volume":"12","author":"Shou","year":"2021","journal-title":"Cell Death Dis."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1060","DOI":"10.1016\/j.cell.2012.03.042","article-title":"Ferroptosis: An iron-dependent form of nonapoptotic cell death","volume":"149","author":"Dixon","year":"2012","journal-title":"Cell"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/j.freeradbiomed.2020.02.027","article-title":"Lipid peroxidation and ferroptosis: The role of GSH and GPx4","volume":"152","author":"Ursini","year":"2020","journal-title":"Free Radic. Biol. Med."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1007\/s00403-024-03345-x","article-title":"Pilot study of the role of ferroptosis in abnormal biological behaviour of keratinocytes in psoriasis vulgaris","volume":"316","author":"Wu","year":"2024","journal-title":"Arch. Dermatol. Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"101675","DOI":"10.1016\/j.tice.2021.101675","article-title":"Inside-out and outside-in organotypic normal human skin culture: JAK-STAT pathway is activated after pro-inflammatory psoriatic cytokine exposure","volume":"74","author":"Cornaghi","year":"2022","journal-title":"Tissue Cell"},{"key":"ref_35","first-page":"4","article-title":"3D skin model to investigate the early epidermal morphological psoriatic features","volume":"6","author":"Donetti","year":"2019","journal-title":"J. Transl. Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1038\/s41420-022-01297-7","article-title":"Recent progress in ferroptosis: Inducers and inhibitors","volume":"8","author":"Du","year":"2022","journal-title":"Cell Death Discov."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1016\/j.cell.2013.12.010","article-title":"Regulation of ferroptotic cancer cell death by GPX4","volume":"156","author":"Yang","year":"2014","journal-title":"Cell"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.reth.2018.04.001","article-title":"Respective optimal calcium concentrations for proliferation on type I collagen fibrils in two keratinocyte line cells, HaCaT and FEPE1L-8","volume":"8","author":"Fujisaki","year":"2018","journal-title":"Regen. Ther."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"3695","DOI":"10.4049\/jimmunol.174.6.3695","article-title":"IL-22 inhibits epidermal differentiation and induces proinflammatory gene expression and migration of human keratinocytes","volume":"174","author":"Boniface","year":"2005","journal-title":"J. Immunol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.cyto.2014.03.003","article-title":"An innovative three-dimensional model of normal human skin to study the proinflammatory psoriatic effects of tumor necrosis factor-alpha and interleukin-17","volume":"68","author":"Donetti","year":"2014","journal-title":"Cytokine"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/978-1-60761-380-0_3","article-title":"Growth and stratification of epithelial cells in minimal culture conditions","volume":"Volume 585","author":"Riva","year":"2010","journal-title":"Epidermal Cells: Methods in Molecular Biology"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/0014-4827(83)90115-5","article-title":"Calcium regulation of cell-cell contact and differentiation of epidermal cells in culture. An ultrastructural study","volume":"143","author":"Hennings","year":"1983","journal-title":"Exp. Cell Res."},{"key":"ref_43","first-page":"607","article-title":"Up-modulation of the expression of functional keratinocyte growth factor receptors induced by high cell density in the human keratinocyte HaCaT cell line","volume":"11","author":"Capone","year":"2000","journal-title":"Cell Growth Differ."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1111\/exd.13674","article-title":"Molecular basis of the skin barrier structures revealed by electron microscopy","volume":"27","author":"Igawa","year":"2018","journal-title":"Exp. Dermatol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1111\/j.1600-0625.2011.01412.x","article-title":"IL-17 downregulates filaggrin and affects keratinocyte expression of genes associated with cellular adhesion","volume":"21","author":"Schaupp","year":"2012","journal-title":"Exp. Dermatol."},{"key":"ref_46","first-page":"2715","article-title":"Evaluation of the effects of IL-22 on the proliferation and differentiation of keratinocytes in vitro","volume":"22","author":"Zhuang","year":"2020","journal-title":"Mol. Med. Rep."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"398","DOI":"10.5021\/ad.2012.24.4.398","article-title":"Interleukin-17 and Interleukin-22 Induced Proinflammatory Cytokine Production in Keratinocytes via Inhibitor of Nuclear Factor \u03baB Kinase-\u03b1 Expression","volume":"24","author":"Cho","year":"2012","journal-title":"Ann. Dermatol."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Ghaffarinia, A., Poliska, S., Ayaydin, F., Goblos, A., Parvaneh, S., Manczinger, M., Balogh, F., Erdei, L., Vereb, Z., and Szabo, K. (2023). Unraveling Transcriptome Profile, Epigenetic Dynamics, and Morphological Changes in Psoriasis-like Keratinocytes: \u201cInsights into Similarity with Psoriatic Lesional Epidermis\u201d. Cells, 12.","DOI":"10.3390\/cells12242825"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Kobayashi, M., Shu, S., Marunaka, K., Matsunaga, T., and Ikari, A. (2020). Weak Ultraviolet B Enhances the Mislocalization of Claudin-1 Mediated by Nitric Oxide and Peroxynitrite Production in Human Keratinocyte-Derived HaCaT Cells. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21197138"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Marunaka, K., Kobayashi, M., Shu, S., Matsunaga, T., and Ikari, A. (2019). Brazilian Green Propolis Rescues Oxidative Stress-Induced Mislocalization of Claudin-1 in Human Keratinocyte-Derived HaCaT Cells. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20163869"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"906","DOI":"10.1038\/sj.jid.5701070","article-title":"Regulation of epidermal tight-junctions (TJ) during infection with exfoliative toxin-negative Staphylococcus strains","volume":"128","author":"Ohnemus","year":"2008","journal-title":"J. Investig. Dermatol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1038\/nrm1055","article-title":"Signalling to and from tight junctions","volume":"4","author":"Matter","year":"2003","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"e13441","DOI":"10.1111\/cpr.13441","article-title":"Zonula occludens-1 distribution and barrier functions are affected by epithelial proliferation and turnover rates","volume":"56","author":"Imafuku","year":"2023","journal-title":"Cell Prolif."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1007\/BF00371784","article-title":"Normal psoriatic epidermis expression of hyperproliferation-associated keratins","volume":"283","author":"Thewes","year":"1991","journal-title":"Arch. Dermatol. Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1083\/jcb.200304017","article-title":"S100C\/A11 is a key mediator of Ca(2+)-induced growth inhibition of human epidermal keratinocytes","volume":"163","author":"Sakaguchi","year":"2003","journal-title":"J. Cell Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"7435621","DOI":"10.1155\/2017\/7435621","article-title":"HaCaT Cells as a Reliable In Vitro Differentiation Model to Dissect the Inflammatory\/Repair Response of Human Keratinocytes","volume":"2017","author":"Colombo","year":"2017","journal-title":"Mediators Inflamm."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1007\/7651_2013_42","article-title":"Growth and differentiation of HaCaT keratinocytes","volume":"Volume 1195","author":"Wilson","year":"2014","journal-title":"Epidermal Cells: Methods in Molecular Biology"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1111\/j.1600-0625.2008.00775.x","article-title":"Effects of extracellular calcium on the growth-differentiation switch in immortalized keratinocyte HaCaT cells compared with normal human keratinocytes","volume":"18","author":"Micallef","year":"2009","journal-title":"Exp. Dermatol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"171","DOI":"10.4062\/biomolther.2012.20.2.171","article-title":"HaCaT Keratinocytes and Primary Epidermal Keratinocytes Have Different Transcriptional Profiles of Cornified Envelope-Associated Genes to T Helper Cell Cytokines","volume":"20","author":"Seo","year":"2012","journal-title":"Biomol. Ther."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"2558","DOI":"10.1016\/j.jid.2021.04.005","article-title":"Down-Regulated Calcium-Sensing Receptor in Keratinocytes and Skin from Aged Mice and Humans Impairs Function","volume":"141","author":"Bollag","year":"2021","journal-title":"J. Investig. Dermatol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"149553","DOI":"10.1016\/j.bbrc.2024.149553","article-title":"Induction of ferroptosis in human keratinocyte HaCaT cells by squalene hydroperoxide: Possible prevention of skin ferroptosis by botanical extracts","volume":"698","author":"Kato","year":"2024","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"2518","DOI":"10.1016\/j.bbagen.2018.05.018","article-title":"Selenium unmasks protective iron armor: A possible defense against cutaneous inflammation and cancer","volume":"1862","author":"Arbiser","year":"2018","journal-title":"Biochim. Biophys. Acta Gen. Subj."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1159\/000495291","article-title":"Psoriasis: Keratinocytes or Immune Cells\u2014Which Is the Trigger?","volume":"235","author":"Benhadou","year":"2019","journal-title":"Dermatology"}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/4\/362\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,23]],"date-time":"2026-02-23T10:22:43Z","timestamp":1771842163000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/4\/362"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,21]]},"references-count":63,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2026,2]]}},"alternative-id":["biology15040362"],"URL":"https:\/\/doi.org\/10.3390\/biology15040362","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,21]]}}}