{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T11:01:09Z","timestamp":1785409269497,"version":"3.56.0"},"reference-count":46,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2025,6,10]],"date-time":"2025-06-10T00:00:00Z","timestamp":1749513600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001348","name":"Agency for Science","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001348","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000268","name":"Biotechnology and Biological Sciences Research Council","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000268","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000770","name":"a Wellcome Trust Institutional Strategic Support Fund to the University of Manchester","doi-asserted-by":"publisher","award":["204796\/Z\/16\/Z"],"award-info":[{"award-number":["204796\/Z\/16\/Z"]}],"id":[{"id":"10.13039\/501100000770","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000272","name":"National Institute for Health and Care Research","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000272","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100014653","name":"Manchester Biomedical Research Centre","doi-asserted-by":"publisher","award":["NIHR203308"],"award-info":[{"award-number":["NIHR203308"]}],"id":[{"id":"10.13039\/100014653","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001349","name":"National Medical Research Council","doi-asserted-by":"publisher","award":["(OF-IRG) MOH-001217"],"award-info":[{"award-number":["(OF-IRG) MOH-001217"]}],"id":[{"id":"10.13039\/501100001349","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,10,17]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:sec>\n                  <jats:title>Background<\/jats:title>\n                  <jats:p>Chronic skin wounds exhibit impaired re-epithelialization due to defective keratinocyte migration and excessive inflammatory signals. Arginase 1 (ARG1) is an enzyme expressed by keratinocytes that is important for skin wound healing. However, the mechanisms underpinning keratinocyte ARG1 function in wound closure are not fully understood.<\/jats:p>\n               <\/jats:sec>\n               <jats:sec>\n                  <jats:title>Objectives<\/jats:title>\n                  <jats:p>To investigate the role of ARG1 in keratinocyte wound closure and inflammatory responses.<\/jats:p>\n               <\/jats:sec>\n               <jats:sec>\n                  <jats:title>Methods<\/jats:title>\n                  <jats:p>In vitro two-dimensional wounding assays using ARG1-inhibited keratinocytes were used to explore the function of ARG1 in keratinocyte scratch closure. ARG1 was also assessed in human skin wounds.<\/jats:p>\n               <\/jats:sec>\n               <jats:sec>\n                  <jats:title>Results<\/jats:title>\n                  <jats:p>ARG1 was strongly expressed in the perilesional regions of acute wounds, but its expression was dysregulated in chronic ulcers, with high expression in the upper layers of the hyperplastic epidermis. ARG1 inhibition in keratinocytes led to a significant decrease in two-dimensional scratch closure, augmented interleukin-1 family- and tumour necrosis factor-\u03b1-driven proinflammatory signalling, and significant downregulation of lipocalin 2 (LCN2). LCN2 expression was partially dependent on Janus kinase\/signal transducer and activator of transcription signalling. Keratinocyte LCN2 neutralization led to delayed scratch wound closure, while LCN2 expression in response to bacterial challenge was impaired upon ARG1 inhibition. ARG1 metabolic products, putrescine and urea, could rescue keratinocyte migration and LCN2 expression in ARG1-inhibited cells.<\/jats:p>\n               <\/jats:sec>\n               <jats:sec>\n                  <jats:title>Conclusions<\/jats:title>\n                  <jats:p>ARG1 plays a major role in keratinocyte re-epithelialization, regulating inflammation and LCN2 production in sterile and infected wound conditions. Dysregulated expression in chronic ulcers may reflect altered enzymatic capacity to drive re-epithelialization or attempts to reduce inflammation and promote antimicrobial function. In culture, impaired wound responses of ARG1 inhibited cells can be rescued by its downstream products putrescine and urea, highlighting the complexity of ARG1 control of wound healing. Manipulation of the ARG1 pathway may have the potential to be used for the management of skin conditions such as infected chronic lesions; however, further study of the pathway is needed to fully understand the role of ARG1 in chronic wounds.<\/jats:p>\n               <\/jats:sec>","DOI":"10.1093\/bjd\/ljaf223","type":"journal-article","created":{"date-parts":[[2025,6,10]],"date-time":"2025-06-10T14:16:06Z","timestamp":1749564966000},"page":"924-935","source":"Crossref","is-referenced-by-count":2,"title":["Keratinocyte arginase 1 regulates proinflammatory responses and drives re-epithelialization via lipocalin 2"],"prefix":"10.1093","volume":"193","author":[{"given":"Denis C","family":"Szondi","sequence":"first","affiliation":[{"name":"Lydia Becker Institute of Immunology and Inflammation, Manchester Academic Health Science Centre, Faculty of Biology, Medicine and Health, University of Manchester , Manchester ,","place":["UK"]},{"name":"A*STAR Skin Research Labs, Agency for Science, Technology and Research (A*STAR) , Singapore ,","place":["Singapore"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel A","family":"Crompton","sequence":"additional","affiliation":[{"name":"Lydia Becker Institute of Immunology and Inflammation, Manchester Academic Health Science Centre, Faculty of Biology, Medicine and Health, University of Manchester , Manchester ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Linus","family":"Oon","sequence":"additional","affiliation":[{"name":"A*STAR Skin Research Labs, Agency for Science, Technology and Research (A*STAR) , Singapore ,","place":["Singapore"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nagavidya","family":"Subramaniam","sequence":"additional","affiliation":[{"name":"A*STAR Skin Research Labs, Agency for Science, Technology and Research (A*STAR) , Singapore ,","place":["Singapore"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sze Han","family":"Lee","sequence":"additional","affiliation":[{"name":"A*STAR Skin Research Labs, Agency for Science, Technology and Research (A*STAR) , Singapore ,","place":["Singapore"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gloria","family":"Lopez-Castejon","sequence":"additional","affiliation":[{"name":"Lydia Becker Institute of Immunology and Inflammation, Manchester Academic Health Science Centre, Faculty of Biology, Medicine and Health, University of Manchester , Manchester ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jason","family":"Wong","sequence":"additional","affiliation":[{"name":"Blond McIndoe Laboratories, Division of Cell Matrix Biology and Regenerative Medicine, Manchester Academic Health Science Centre, School of Biological Sciences, Faculty of Biology, Medicine and Health, The University of Manchester , Manchester ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3047-5475","authenticated-orcid":false,"given":"Sheena M","family":"Cruickshank","sequence":"additional","affiliation":[{"name":"Lydia Becker Institute of Immunology and Inflammation, Manchester Academic Health Science Centre, Faculty of Biology, Medicine and Health, University of Manchester , Manchester ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leah A","family":"Vardy","sequence":"additional","affiliation":[{"name":"A*STAR Skin Research Labs, Agency for Science, Technology and Research (A*STAR) , Singapore ,","place":["Singapore"]},{"name":"Skin Research Institute of Singapore (SRIS) , Immunos ,","place":["Singapore"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,6,10]]},"reference":[{"key":"2025101708280927400_ljaf223-B1","doi-asserted-by":"crossref","first-page":"4047","DOI":"10.1242\/dev.152587","article-title":"Fibroblast growth factors: key players in regeneration and tissue repair","volume":"144","author":"Maddaluno","year":"2017","journal-title":"Development"},{"key":"2025101708280927400_ljaf223-B2","doi-asserted-by":"crossref","first-page":"265sr6","DOI":"10.1126\/scitranslmed.3009337","article-title":"Wound repair and regeneration: mechanisms, signaling, and translation","volume":"6","author":"Eming","year":"2014","journal-title":"Sci Transl Med"},{"key":"2025101708280927400_ljaf223-B3","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1111\/iwj.13292","article-title":"Non-healing wounds: can it take different diagnosis?","volume":"17","author":"Calis","year":"2020","journal-title":"Int Wound J"},{"key":"2025101708280927400_ljaf223-B4","doi-asserted-by":"crossref","first-page":"1206","DOI":"10.1016\/j.jid.2021.09.009","article-title":"An epidermal-specific role for arginase 1 during cutaneous wound repair","volume":"142","author":"Crompton","year":"2021","journal-title":"J Invest Dermatol"},{"key":"2025101708280927400_ljaf223-B5","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1016\/S0039-6109(02)00207-4","article-title":"Nitric oxide and wound repair","volume":"83","author":"Schwentker","year":"2003","journal-title":"Surg Clin North Am"},{"key":"2025101708280927400_ljaf223-B6","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1046\/j.1524-475X.2003.11605.x","article-title":"Arginine physiology and its implication for wound healing","volume":"11","author":"Witte","year":"2003","journal-title":"Wound Repair Regen"},{"key":"2025101708280927400_ljaf223-B7","doi-asserted-by":"crossref","first-page":"1561","DOI":"10.1038\/jid.2012.42","article-title":"Urea uptake enhances barrier function and antimicrobial defense in humans by regulating epidermal gene expression","volume":"132","author":"Grether-Beck","year":"2012","journal-title":"J Invest Dermatol"},{"key":"2025101708280927400_ljaf223-B8","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1016\/j.tips.2015.03.006","article-title":"Arginase: an old enzyme with new tricks","volume":"36","author":"Caldwell","year":"2015","journal-title":"Trends Pharmacol Sci"},{"key":"2025101708280927400_ljaf223-B9","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1042\/BJ20090598","article-title":"Polyamine catabolism and disease","volume":"421","author":"Casero","year":"2009","journal-title":"Biochem J"},{"key":"2025101708280927400_ljaf223-B10","first-page":"681","article-title":"Roles of cellular polyamines in mucosal healing in the gastrointestinal tract","volume":"64","author":"Gao","year":"2013","journal-title":"J Physiol Pharmacol"},{"key":"2025101708280927400_ljaf223-B11","doi-asserted-by":"crossref","first-page":"2653","DOI":"10.1016\/j.jid.2018.05.029","article-title":"Polyamine regulator AMD1 promotes cell migration in epidermal wound healing","volume":"138","author":"Lim","year":"2018","journal-title":"J Invest Dermatol"},{"key":"2025101708280927400_ljaf223-B12","doi-asserted-by":"crossref","first-page":"2011","DOI":"10.3945\/jn.117.256404","article-title":"Proline precursors and collagen synthesis: biochemical challenges of nutrient supplementation and wound healing","volume":"147","author":"Albaugh","year":"2017","journal-title":"J Nutr"},{"key":"2025101708280927400_ljaf223-B13","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1177\/0022034509359125","article-title":"Factors affecting wound healing","volume":"89","author":"Guo","year":"2010","journal-title":"J Dent Res"},{"key":"2025101708280927400_ljaf223-B14","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1152\/physrev.2003.83.3.835","article-title":"Regulation of wound healing by growth factors and cytokines","volume":"83","author":"Werner","year":"2003","journal-title":"Physiol Rev"},{"key":"2025101708280927400_ljaf223-B15","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1038\/s41392-021-00791-1","article-title":"The JAK\/STAT signaling pathway: from bench to clinic","volume":"6","author":"Hu","year":"2021","journal-title":"Signal Transduct Target Ther"},{"key":"2025101708280927400_ljaf223-B16","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.cytogfr.2017.10.001","article-title":"The JAK\/STAT signaling pathway and photobiomodulation in chronic wound healing","volume":"38","author":"Jere","year":"2007","journal-title":"Cytokine Growth Factor Rev"},{"key":"2025101708280927400_ljaf223-B17","doi-asserted-by":"crossref","first-page":"155214","DOI":"10.1016\/j.cyto.2020.155214","article-title":"TNF-\u03b1 controls lipocalin-2 expression in PC-3 prostate cancer cells","volume":"135","author":"Schr\u00f6der","year":"2020","journal-title":"Cytokine"},{"key":"2025101708280927400_ljaf223-B18","doi-asserted-by":"crossref","first-page":"151328","DOI":"10.1016\/j.ejcb.2023.151328","article-title":"Lipocalin-2 regulates the expression of interferon-stimulated genes and the susceptibility of prostate cancer cells to oncolytic virus infection","volume":"102","author":"Barer","year":"2023","journal-title":"Eur J Cell Biol"},{"key":"2025101708280927400_ljaf223-B19","doi-asserted-by":"crossref","first-page":"1436","DOI":"10.1128\/MCB.20.4.1436-1447.2000","article-title":"Human keratinocytes that express hTERT and also bypass a p16(INK4a)-enforced mechanism that limits life span become immortal yet retain normal growth and differentiation characteristics","volume":"20","author":"Dickson","year":"2000","journal-title":"Mol Cell Biol"},{"key":"2025101708280927400_ljaf223-B20","doi-asserted-by":"crossref","first-page":"5157","DOI":"10.1128\/MCB.22.14.5157-5172.2002","article-title":"A two-stage, p16(INK4A)- and p53-dependent keratinocyte senescence mechanism that limits replicative potential independent of telomere status","volume":"22","author":"Rheinwald","year":"2002","journal-title":"Mol Cell Biol"},{"key":"2025101708280927400_ljaf223-B21","author":"Wang"},{"key":"2025101708280927400_ljaf223-B22","doi-asserted-by":"crossref","first-page":"1447","DOI":"10.1021\/acs.jproteome.9b00640","article-title":"Skyline for small molecules: a unifying software package for quantitative metabolomics","volume":"19","author":"Adams","year":"2020","journal-title":"J\u00a0Proteome Res"},{"key":"2025101708280927400_ljaf223-B23","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1007\/s001250051224","article-title":"The role of nitric oxide synthase isoforms and arginase in the pathogenesis of diabetic foot ulcers: possible modulatory effects by transforming growth factor beta 1","volume":"42","author":"Jude","year":"1999","journal-title":"Diabetologia"},{"key":"2025101708280927400_ljaf223-B24","doi-asserted-by":"crossref","first-page":"434","DOI":"10.1002\/1096-9896(2000)9999:9999<::AID-PATH654>3.0.CO;2-S","article-title":"Expression of nitric oxide synthase isoforms and arginase in normal human skin and chronic venous leg ulcers","volume":"191","author":"Abd-El-Aleem","year":"2000","journal-title":"J Pathol"},{"key":"2025101708280927400_ljaf223-B25","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1016\/j.addr.2018.06.019","article-title":"Re-epithelialization of adult skin wounds: cellular mechanisms and therapeutic strategies","volume":"146","author":"Rousselle","year":"2019","journal-title":"Adv Drug Deliv Rev"},{"key":"2025101708280927400_ljaf223-B26","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1097\/00005537-200302000-00009","article-title":"Interleukin-1 facilitates airway epithelial migration in response to injury","volume":"113","author":"Hicks","year":"2003","journal-title":"Laryngoscope"},{"key":"2025101708280927400_ljaf223-B27","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1111\/1523-1747.ep12606970","article-title":"Interleukin-1 alpha stimulates keratinocyte migration through an epidermal growth factor\/transforming growth factor-alpha-independent pathway","volume":"104","author":"Chen","year":"1995","journal-title":"J Invest Dermatol"},{"key":"2025101708280927400_ljaf223-B28","first-page":"1812","article-title":"IL (interleukin)-1\u03b1 promotes nuclear factor-kappa B and AP-1-induced IL-8 expression, cell survival, and proliferation in head and neck squamous cell carcinomas","volume":"7","author":"Wolf","year":"2001","journal-title":"Clin Cancer Res"},{"key":"2025101708280927400_ljaf223-B29","doi-asserted-by":"crossref","first-page":"16514","DOI":"10.1074\/jbc.270.28.16514","article-title":"NF-kappa-B activation by interleukin-1 (IL-1) requires an IL-1 receptor-associated protein-kinase activity","volume":"270","author":"Croston","year":"1995","journal-title":"J Biol Chem"},{"key":"2025101708280927400_ljaf223-B30","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1186\/s12974-015-0448-8","article-title":"NF-\u03baB transcriptional activation by TNF\u03b1 requires phospholipase C, extracellular signal-regulated kinase 2 and poly(ADP-ribose) polymerase-1","volume":"12","author":"Vuong","year":"2015","journal-title":"J\u00a0Neuroinflammation"},{"key":"2025101708280927400_ljaf223-B31","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1016\/j.cyto.2011.07.021","article-title":"Lipocalin 2 regulation and its complex role in inflammation and cancer","volume":"56","author":"Li","year":"2011","journal-title":"Cytokine"},{"key":"2025101708280927400_ljaf223-B32","doi-asserted-by":"crossref","first-page":"1905","DOI":"10.1007\/s13555-021-00611-y","article-title":"Urea in dermatology: a review of its emollient, moisturizing, keratolytic, skin barrier enhancing and antimicrobial properties","volume":"11","author":"Piquero-Casals","year":"2021","journal-title":"Dermatol Ther"},{"key":"2025101708280927400_ljaf223-B33","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1111\/j.1468-3083.2011.04415.x","article-title":"Cytokines, chemokines and growth factors in wound healing","volume":"26","author":"Behm","year":"2012","journal-title":"J Eur Acad Dermatol Venereol"},{"key":"2025101708280927400_ljaf223-B34","doi-asserted-by":"crossref","first-page":"222969","DOI":"10.1371\/journal.pone.0222969","article-title":"Interleukin (IL)-17\/IL-36 axis participates to the crosstalk between endothelial cells and keratinocytes during inflammatory skin responses","volume":"15","author":"Mercurio","year":"2020","journal-title":"PLOS ONE"},{"key":"2025101708280927400_ljaf223-B35","doi-asserted-by":"crossref","first-page":"919973","DOI":"10.3389\/fimmu.2022.919973","article-title":"Interleukin-18 cytokine in immunity, inflammation, and autoimmunity: biological role in induction, regulation, and treatment","volume":"13","author":"Ihim","year":"2022","journal-title":"Front Immunol"},{"key":"2025101708280927400_ljaf223-B36","doi-asserted-by":"crossref","first-page":"101755","DOI":"10.1016\/j.molmet.2023.101755","article-title":"Chronic inflammation and the hallmarks of aging","volume":"74","author":"Baechle","year":"2023","journal-title":"Mol Metab"},{"key":"2025101708280927400_ljaf223-B37","doi-asserted-by":"crossref","first-page":"R80","DOI":"10.1186\/gb-2008-9-5-r80","article-title":"Estrogen, not intrinsic aging, is the major regulator of delayed human wound healing in the elderly","volume":"9","author":"Hardman","year":"2008","journal-title":"Genome Biol"},{"key":"2025101708280927400_ljaf223-B38","doi-asserted-by":"crossref","first-page":"2526","DOI":"10.1096\/fj.201801513R","article-title":"Critical role of IL-1\u03b1 in IL-1\u03b2-induced inflammatory responses: cooperation with NF-kB p65 in transcriptional regulation","volume":"33","author":"Singh","year":"2019","journal-title":"FASEB J"},{"key":"2025101708280927400_ljaf223-B39","doi-asserted-by":"crossref","first-page":"317","DOI":"10.3389\/fcell.2019.00317","article-title":"Function and regulation of IL-36 signaling in inflammatory diseases and cancer development","volume":"7","author":"Queen","year":"2019","journal-title":"Front Cell Dev Biol"},{"key":"2025101708280927400_ljaf223-B40","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1038\/nri910","article-title":"NF-kappaB regulation in the immune system","volume":"2","author":"Li","year":"2002","journal-title":"Nat Rev Immunol"},{"key":"2025101708280927400_ljaf223-B41","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1111\/exd.14310","article-title":"Induction of IL-1\u03b2 and antimicrobial peptides as a potential mechanism for topical dithranol","volume":"30","author":"Benezeder","year":"2021","journal-title":"Exp Dermatol"},{"key":"2025101708280927400_ljaf223-B42","doi-asserted-by":"crossref","first-page":"4088","DOI":"10.1038\/ncomms5088","article-title":"Tcf3 promotes cell migration and wound repair through regulation of lipocalin 2","volume":"5","author":"Miao","year":"2014","journal-title":"Nat Commun"},{"key":"2025101708280927400_ljaf223-B43","doi-asserted-by":"crossref","first-page":"1544","DOI":"10.1046\/j.1523-1747.2003.12610.x","article-title":"Expression and activity of arginase isoenzymes during normal and diabetes-impaired skin repair","volume":"121","author":"Kampfer","year":"2003","journal-title":"J Invest Dermatol"},{"key":"2025101708280927400_ljaf223-B44","doi-asserted-by":"crossref","first-page":"23618","DOI":"10.1002\/jcp.28930","article-title":"Cellular and physiological upregulation of inducible nitric oxide synthase, arginase, and inducible cyclooxygenase in wound healing","volume":"234","author":"Abd El-Aleem","year":"2019","journal-title":"J Cell Physiol"},{"key":"2025101708280927400_ljaf223-B45","doi-asserted-by":"crossref","first-page":"2498","DOI":"10.3390\/nu13082498","article-title":"The effect of amino acids on wound healing: a systematic review and meta-analysis on arginine and glutamine","volume":"13","author":"Arribas-L\u00f3pez","year":"2021","journal-title":"Nutrients"},{"key":"2025101708280927400_ljaf223-B46","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1016\/S0002-9610(36)90704-3","article-title":"Use of urea to stimulate healing in chronic purulent wounds","volume":"33","author":"Robinson","year":"1936","journal-title":"Am J Surg"}],"container-title":["British Journal of Dermatology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/bjd\/advance-article-pdf\/doi\/10.1093\/bjd\/ljaf223\/63464380\/ljaf223.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/bjd\/article-pdf\/193\/5\/924\/63464380\/ljaf223.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/bjd\/article-pdf\/193\/5\/924\/63464380\/ljaf223.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,17]],"date-time":"2025-10-17T12:28:19Z","timestamp":1760704099000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/bjd\/article\/193\/5\/924\/8159930"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,6,10]]},"references-count":46,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2025,6,10]]},"published-print":{"date-parts":[[2025,10,17]]}},"URL":"https:\/\/doi.org\/10.1093\/bjd\/ljaf223","relation":{},"ISSN":["0007-0963","1365-2133"],"issn-type":[{"value":"0007-0963","type":"print"},{"value":"1365-2133","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2025,11]]},"published":{"date-parts":[[2025,6,10]]}}}