{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T03:55:58Z","timestamp":1791518158235,"version":"4.3.4"},"reference-count":44,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T00:00:00Z","timestamp":1780272000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T00:00:00Z","timestamp":1780272000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,9,21]],"date-time":"2025-09-21T00:00:00Z","timestamp":1758412800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012226","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","award":["2022CDJYGRH-003"],"award-info":[{"award-number":["2022CDJYGRH-003"]}],"id":[{"id":"10.13039\/501100012226","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012226","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","award":["2023CDJKYJH066"],"award-info":[{"award-number":["2023CDJKYJH066"]}],"id":[{"id":"10.13039\/501100012226","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82373509"],"award-info":[{"award-number":["82373509"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000009","name":"Foundation for the National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000009","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012165","name":"Key Technologies Research and Development Program","doi-asserted-by":"publisher","award":["2023YFC2508200"],"award-info":[{"award-number":["2023YFC2508200"]}],"id":[{"id":"10.13039\/501100012165","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002858","name":"China Postdoctoral Science Foundation","doi-asserted-by":"publisher","award":["2024M763171"],"award-info":[{"award-number":["2024M763171"]}],"id":[{"id":"10.13039\/501100002858","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Journal of Advanced Research"],"published-print":{"date-parts":[[2026,6]]},"DOI":"10.1016\/j.jare.2025.09.040","type":"journal-article","created":{"date-parts":[[2025,9,24]],"date-time":"2025-09-24T09:20:18Z","timestamp":1758705618000},"page":"345-359","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":2,"special_numbering":"C","title":["Roles of Msx2 in exogen control: modulating the stem cell niche during the transition from hair shedding to regeneration"],"prefix":"10.1016","volume":"84","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-9255-3207","authenticated-orcid":false,"given":"Qi","family":"Chen","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5460-9653","authenticated-orcid":false,"given":"Wen-Chien","family":"Jea","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting-Xin","family":"Jiang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ping","family":"Wu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0743-2079","authenticated-orcid":false,"given":"Chi","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ji","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mingxing","family":"Lei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cheng-Ming","family":"Chuong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7813-8921","authenticated-orcid":false,"given":"Ya-Chen","family":"Liang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"3","key":"10.1016\/j.jare.2025.09.040_b0005","doi-asserted-by":"crossref","DOI":"10.3390\/jcm12030893","article-title":"Integrative and mechanistic approach to the hair growth cycle and hair loss","volume":"12","author":"Natarelli","year":"2023","journal-title":"J Clin Med"},{"issue":"3","key":"10.1016\/j.jare.2025.09.040_b0010","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1046\/j.1523-1747.2002.01842.x","article-title":"Exogen, shedding phase of the hair growth cycle: characterization of a mouse model","volume":"119","author":"Milner","year":"2002","journal-title":"J Invest Dermatol"},{"issue":"2","key":"10.1016\/j.jare.2025.09.040_b0015","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1016\/j.jaad.2004.07.040","article-title":"Exogen is an active, separately controlled phase of the hair growth cycle","volume":"52","author":"Stenn","year":"2005","journal-title":"J Am Acad Dermatol"},{"issue":"9","key":"10.1016\/j.jare.2025.09.040_b0020","doi-asserted-by":"crossref","first-page":"793","DOI":"10.1111\/j.1600-0625.2008.00833.x","article-title":"Exogen involves gradual release of the hair club fibre in the vibrissa follicle model","volume":"18","author":"Higgins","year":"2009","journal-title":"Exp Dermatol"},{"issue":"4","key":"10.1016\/j.jare.2025.09.040_b0025","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1111\/j.1600-0846.2007.00248.x","article-title":"Exogen hair characterization in human scalp.","volume":"13","author":"Van Neste","year":"2007","journal-title":"Skin Res Technol"},{"issue":"4","key":"10.1016\/j.jare.2025.09.040_b0030","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1159\/000460300","article-title":"Exogen hairs in women with and without hair loss","volume":"3","author":"Guarrera","year":"2017","journal-title":"Skin Append Disord."},{"issue":"12","key":"10.1016\/j.jare.2025.09.040_b0035","doi-asserted-by":"crossref","first-page":"2349","DOI":"10.1038\/jid.2011.227","article-title":"Modulation in proteolytic activity is identified as a hallmark of exogen by transcriptional profiling of hair follicles","volume":"131","author":"Higgins","year":"2011","journal-title":"J Invest Dermatol"},{"issue":"9","key":"10.1016\/j.jare.2025.09.040_b0040","doi-asserted-by":"crossref","first-page":"2100","DOI":"10.1038\/jid.2009.66","article-title":"From telogen to exogen: mechanisms underlying formation and subsequent loss of the hair club fiber","volume":"129","author":"Higgins","year":"2009","journal-title":"J Invest Dermatol"},{"issue":"2","key":"10.1016\/j.jare.2025.09.040_b0045","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1242\/dev.00201","article-title":"'Cyclic alopecia' in Msx2 mutants: defects in hair cycling and hair shaft differentiation","volume":"130","author":"Ma","year":"2003","journal-title":"Development"},{"issue":"3","key":"10.1016\/j.jare.2025.09.040_b0050","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/0092-8674(93)90379-5","article-title":"A mutation in the homeodomain of the human MSX2 gene in a family affected with autosomal dominant craniosynostosis","volume":"75","author":"Jabs","year":"1993","journal-title":"Cell"},{"issue":"41","key":"10.1016\/j.jare.2025.09.040_b0055","doi-asserted-by":"crossref","first-page":"20528","DOI":"10.1073\/pnas.1905973116","article-title":"The FBXW2-MSX2-SOX2 axis regulates stem cell property and drug resistance of cancer cells","volume":"116","author":"Yin","year":"2019","journal-title":"PNAS"},{"issue":"37","key":"10.1016\/j.jare.2025.09.040_b0060","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2105130118","article-title":"MSX2 safeguards syncytiotrophoblast fate of human trophoblast stem cells","volume":"118","author":"Hornbachner","year":"2021","journal-title":"Proc Natl Acad Sci USA"},{"issue":"9","key":"10.1016\/j.jare.2025.09.040_b0065","doi-asserted-by":"crossref","first-page":"2041","DOI":"10.1016\/j.jid.2018.02.043","article-title":"Msx2 supports epidermal competency during wound-induced hair follicle neogenesis","volume":"138","author":"Hughes","year":"2018","journal-title":"J Invest Dermatol"},{"issue":"2","key":"10.1016\/j.jare.2025.09.040_b0070","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.jdermsci.2013.06.017","article-title":"Microarray analysis of androgenetic and senescent alopecia: comparison of gene expression shows two distinct profiles","volume":"72","author":"Karnik","year":"2013","journal-title":"J Dermatol Sci"},{"issue":"1","key":"10.1016\/j.jare.2025.09.040_b0075","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1186\/s13287-020-01653-3","article-title":"MSX2 suppression through inhibition of TGF\u03b2 signaling enhances hematopoietic differentiation of human embryonic stem cells","volume":"11","author":"Wang","year":"2020","journal-title":"Stem Cell Res Ther"},{"issue":"19","key":"10.1016\/j.jare.2025.09.040_b0080","doi-asserted-by":"crossref","first-page":"4007","DOI":"10.1016\/j.cell.2023.07.036","article-title":"TGF-\u03b2 signaling in health and disease","volume":"186","author":"Massagu\u00e9","year":"2023","journal-title":"Cell."},{"issue":"4","key":"10.1016\/j.jare.2025.09.040_b0085","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1038\/74231","article-title":"Msx2 deficiency in mice causes pleiotropic defects in bone growth and ectodermal organ formation","volume":"24","author":"Satokata","year":"2000","journal-title":"Nat Genet"},{"key":"10.1016\/j.jare.2025.09.040_b0090","article-title":"Isolating hair follicle stem cells and epidermal keratinocytes from dorsal mouse skin","volume":"110","author":"Soteriou","year":"2016","journal-title":"J Visual Exp: JoVE"},{"issue":"10","key":"10.1016\/j.jare.2025.09.040_b0095","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1038\/nmeth.4396","article-title":"An improved ATAC-seq protocol reduces background and enables interrogation of frozen tissues","volume":"14","author":"Corces","year":"2017","journal-title":"Nat Methods"},{"issue":"3","key":"10.1016\/j.jare.2025.09.040_b0100","doi-asserted-by":"crossref","DOI":"10.1016\/j.cell.2017.03.038","article-title":"Epithelial-mesenchymal micro-niches govern stem cell lineage choices","volume":"169","author":"Yang","year":"2017","journal-title":"Cell"},{"issue":"11","key":"10.1016\/j.jare.2025.09.040_b0105","doi-asserted-by":"crossref","first-page":"E1506","DOI":"10.1073\/pnas.1601569113","article-title":"FOXC1 maintains the hair follicle stem cell niche and governs stem cell quiescence to preserve long-term tissue-regenerating potential","volume":"113","author":"Lay","year":"2016","journal-title":"PNAS"},{"issue":"1","key":"10.1016\/j.jare.2025.09.040_b0110","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1016\/j.cell.2010.11.049","article-title":"Dynamics between stem cells, niche, and progeny in the hair follicle","volume":"144","author":"Hsu","year":"2011","journal-title":"Cell"},{"issue":"2","key":"10.1016\/j.jare.2025.09.040_b0115","doi-asserted-by":"crossref","DOI":"10.1016\/j.cell.2011.07.001","article-title":"SnapShot: hair follicle stem cells","volume":"146","author":"Woo","year":"2011","journal-title":"Cell"},{"issue":"4","key":"10.1016\/j.jare.2025.09.040_b0120","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/j.cell.2011.01.014","article-title":"The basement membrane of hair follicle stem cells is a muscle cell niche","volume":"144","author":"Fujiwara","year":"2011","journal-title":"Cell"},{"key":"10.1016\/j.jare.2025.09.040_b0125","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1038\/s43587-021-00103-w","article-title":"Escape of hair follicle stem cells causes stem cell exhaustion during aging","volume":"1","author":"Zhang","year":"2021","journal-title":"Nat Aging"},{"issue":"1","key":"10.1016\/j.jare.2025.09.040_b0130","doi-asserted-by":"crossref","first-page":"2577","DOI":"10.1038\/s41467-021-22881-y","article-title":"Mapping the molecular and structural specialization of the skin basement membrane for inter-tissue interactions","volume":"12","author":"Tsutsui","year":"2021","journal-title":"Nat Commun"},{"issue":"6","key":"10.1016\/j.jare.2025.09.040_b0135","doi-asserted-by":"crossref","DOI":"10.1016\/j.cell.2018.10.012","article-title":"Identity noise and adipogenic traits characterize dermal fibroblast aging","volume":"175","author":"Salzer","year":"2018","journal-title":"Cell."},{"issue":"7866","key":"10.1016\/j.jare.2025.09.040_b0140","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/s41586-021-03624-x","article-title":"Obesity accelerates hair thinning by stem cell-centric converging mechanisms","volume":"595","author":"Morinaga","year":"2021","journal-title":"Nature"},{"issue":"12","key":"10.1016\/j.jare.2025.09.040_b0145","doi-asserted-by":"crossref","DOI":"10.15252\/embr.202255478","article-title":"The extracellular matrix fibulin 7 maintains epidermal stem cell heterogeneity during skin aging","volume":"23","author":"Raja","year":"2022","journal-title":"EMBO Rep"},{"issue":"6687","key":"10.1016\/j.jare.2025.09.040_b0150","doi-asserted-by":"crossref","DOI":"10.1126\/science.adi7342","article-title":"Vitamin a resolves lineage plasticity to orchestrate stem cell lineage choices","volume":"383","author":"Tierney","year":"2024","journal-title":"Science (New York NY)"},{"issue":"6273","key":"10.1016\/j.jare.2025.09.040_b0155","doi-asserted-by":"crossref","DOI":"10.1126\/science.aad4395","article-title":"Hair follicle aging is driven by transepidermal elimination of stem cells via COL17A1 proteolysis","volume":"351","author":"Matsumura","year":"2016","journal-title":"Science (New York, NY)"},{"issue":"1","key":"10.1016\/j.jare.2025.09.040_b0160","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.stem.2011.11.005","article-title":"Paracrine TGF-\u03b2 signaling counterbalances BMP-mediated repression in hair follicle stem cell activation","volume":"10","author":"Oshimori","year":"2012","journal-title":"Cell Stem Cell"},{"issue":"D1","key":"10.1016\/j.jare.2025.09.040_b0165","doi-asserted-by":"crossref","first-page":"D39","DOI":"10.1093\/nar\/gkac907","article-title":"AnimalTFDB 4.0: a comprehensive animal transcription factor database updated with variation and expression annotations","volume":"51","author":"Shen","year":"2023","journal-title":"Nucleic Acids Res"},{"key":"10.1016\/j.jare.2025.09.040_b0170","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.60066","article-title":"Lef1 expression in fibroblasts maintains developmental potential in adult skin to regenerate wounds","volume":"9","author":"Phan","year":"2020","journal-title":"eLife"},{"key":"10.1016\/j.jare.2025.09.040_b0175","doi-asserted-by":"crossref","DOI":"10.1016\/j.cell.2024.08.014","article-title":"TGF-\u03b2 and RAS jointly unmask primed enhancers to drive metastasis","author":"Lee","year":"2024","journal-title":"Cell"},{"issue":"2","key":"10.1016\/j.jare.2025.09.040_b0180","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1016\/j.jdermsci.2019.04.008","article-title":"Keratinocyte-derived TGF\u03b2 is not required to maintain skin immune homeostasis","volume":"94","author":"Yang","year":"2019","journal-title":"J Dermatol Sci"},{"issue":"3","key":"10.1016\/j.jare.2025.09.040_b0185","doi-asserted-by":"crossref","first-page":"314","DOI":"10.1016\/j.stem.2013.06.018","article-title":"Architectural niche organization by LHX2 is linked to hair follicle stem cell function","volume":"13","author":"Folgueras","year":"2013","journal-title":"Cell Stem Cell"},{"issue":"4","key":"10.1016\/j.jare.2025.09.040_b0190","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1101\/gad.233247.113","article-title":"SOX9: a stem cell transcriptional regulator of secreted niche signaling factors","volume":"28","author":"Kadaja","year":"2014","journal-title":"Genes Dev"},{"issue":"10","key":"10.1016\/j.jare.2025.09.040_b0195","doi-asserted-by":"crossref","first-page":"2358","DOI":"10.1038\/jid.2015.187","article-title":"Lack of collagen VI promotes wound-induced hair growth","volume":"135","author":"Chen","year":"2015","journal-title":"J Invest Dermatol"},{"issue":"30","key":"10.1016\/j.jare.2025.09.040_b0200","doi-asserted-by":"crossref","DOI":"10.1097\/MD.0000000000004297","article-title":"An investigation of crosstalk between Wnt\/\u03b2-catenin and transforming growth factor-\u03b2 signaling in androgenetic alopecia","volume":"95","author":"Lu","year":"2016","journal-title":"Medicine"},{"issue":"16","key":"10.1016\/j.jare.2025.09.040_b0205","doi-asserted-by":"crossref","first-page":"9139","DOI":"10.1073\/pnas.161016098","article-title":"Conditional epidermal expression of TGFbeta 1 blocks neonatal lethality but causes a reversible hyperplasia and alopecia","volume":"98","author":"Liu","year":"2001","journal-title":"PNAS"},{"issue":"9","key":"10.1016\/j.jare.2025.09.040_b0210","doi-asserted-by":"crossref","first-page":"460","DOI":"10.5483\/BMBRep.2013.46.9.228","article-title":"Induction of transforming growth factor-beta 1 by androgen is mediated by reactive oxygen species in hair follicle dermal papilla cells","volume":"46","author":"Shin","year":"2013","journal-title":"BMB Rep"},{"issue":"8","key":"10.1016\/j.jare.2025.09.040_b0215","doi-asserted-by":"crossref","DOI":"10.1016\/j.jid.2022.01.003","article-title":"Androgen receptor-mediated paracrine signaling induces regression of blood vessels in the dermal papilla in androgenetic alopecia","volume":"142","author":"Deng","year":"2022","journal-title":"J Invest Dermatol"},{"issue":"1","key":"10.1016\/j.jare.2025.09.040_b0225","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1111\/j.1365-2230.2005.02026.x","article-title":"Dandruff-associated smouldering alopecia: a chronobiological assessment over 5 years","volume":"31","author":"Pi\u00e9rard-Franchimont","year":"2006","journal-title":"Clin Exp Dermatol"}],"container-title":["Journal of Advanced Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2090123225007477?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2090123225007477?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,6,3]],"date-time":"2026-06-03T19:49:45Z","timestamp":1780516185000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2090123225007477"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6]]},"references-count":44,"alternative-id":["S2090123225007477"],"URL":"https:\/\/doi.org\/10.1016\/j.jare.2025.09.040","relation":{},"ISSN":["2090-1232"],"issn-type":[{"value":"2090-1232","type":"print"}],"subject":[],"published":{"date-parts":[[2026,6]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Roles of Msx2 in exogen control: modulating the stem cell niche during the transition from hair shedding to regeneration","name":"articletitle","label":"Article Title"},{"value":"Journal of Advanced Research","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jare.2025.09.040","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2025 The Author(s). Published by Elsevier B.V. on behalf of Cairo University.","name":"copyright","label":"Copyright"}]}}