{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T05:49:34Z","timestamp":1790142574768,"version":"4.0.1"},"reference-count":156,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T00:00:00Z","timestamp":1788998400000},"content-version":"vor","delay-in-days":45,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Cell Commun Signal"],"DOI":"10.1186\/s12964-026-03086-6","type":"journal-article","created":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T09:04:46Z","timestamp":1785143086000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Aging of hair follicle stem cells and their niche: mechanisms and regenerative therapeutic strategies"],"prefix":"10.1186","volume":"24","author":[{"given":"Siqi","family":"Xu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiming","family":"Guo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ying","family":"Kang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinjing","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yue","family":"Ming","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weixing","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhijie","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yisheng","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7178-424X","authenticated-orcid":false,"given":"Bo","family":"Jia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,27]]},"reference":[{"issue":"1","key":"3086_CR1","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1046\/j.0022-202x.2001.01377.x","volume":"117","author":"S M\u00fcller-R\u00f6ver","year":"2001","unstructured":"M\u00fcller-R\u00f6ver S, Foitzik K, Paus R, et al. A Comprehensive Guide for the Accurate Classification of Murine Hair Follicles in Distinct Hair Cycle Stages. J Invest Dermatology. 2001;117(1):3\u201315.","journal-title":"J Invest Dermatology"},{"issue":"4","key":"3086_CR2","doi-asserted-by":"publisher","first-page":"532","DOI":"10.1016\/j.stem.2020.09.011","volume":"27","author":"E Fuchs","year":"2020","unstructured":"Fuchs E, Blau HM. Tissue Stem Cells: Architects of Their Niches. Cell Stem Cell. 2020;27(4):532\u201356.","journal-title":"Cell Stem Cell"},{"issue":"3","key":"3086_CR3","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1016\/j.stem.2009.06.004","volume":"5","author":"YV Zhang","year":"2009","unstructured":"Zhang YV, Cheong J, Ciapurin N, et al. Distinct self-renewal and differentiation phases in the niche of infrequently dividing hair follicle stem cells. Cell Stem Cell. 2009;5(3):267\u201378.","journal-title":"Cell Stem Cell"},{"key":"3086_CR4","doi-asserted-by":"publisher","first-page":"1192149","DOI":"10.3389\/fragi.2023.1192149","volume":"4","author":"C Zhang","year":"2023","unstructured":"Zhang C, Wang D, Dowell R, et al. Single cell analysis of transcriptome and open chromatin reveals the dynamics of hair follicle stem cell aging. Front Aging. 2023;4:1192149.","journal-title":"Front Aging"},{"key":"3086_CR5","doi-asserted-by":"publisher","first-page":"204173142412487","DOI":"10.1177\/20417314241248753","volume":"15","author":"MJ Kim","year":"2024","unstructured":"Kim MJ, Ahn HJ, Kong D, et al. Modeling of solar UV-induced photodamage on the hair follicles in human skin organoids. J Tissue Eng. 2024;15:20417314241248753.","journal-title":"J Tissue Eng"},{"key":"3086_CR6","doi-asserted-by":"publisher","first-page":"527","DOI":"10.2147\/CCID.S310487","volume":"14","author":"X Zhai","year":"2021","unstructured":"Zhai X, Gong M, Peng Y, et al. Effects of UV Induced-Photoaging on the Hair Follicle Cycle of C57BL6\/J Mice. Clin Cosmet Investig Dermatol. 2021;14:527\u201339.","journal-title":"Clin Cosmet Investig Dermatol"},{"issue":"4","key":"3086_CR7","doi-asserted-by":"publisher","first-page":"722","DOI":"10.1111\/j.1474-9726.2012.00837.x","volume":"11","author":"ME Waaijer","year":"2012","unstructured":"Waaijer ME, Parish WE, Strongitharm BH, et al. The number of p16INK4a positive cells in human skin reflects biological age. Aging Cell. 2012;11(4):722\u20135.","journal-title":"Aging Cell"},{"issue":"8","key":"3086_CR8","doi-asserted-by":"publisher","first-page":"1676","DOI":"10.1242\/dev.090662","volume":"140","author":"W Chi","year":"2013","unstructured":"Chi W, Wu E, Morgan BA. Dermal papilla cell number specifies hair size, shape and cycling and its reduction causes follicular decline. Development. 2013;140(8):1676\u201383.","journal-title":"Development"},{"key":"3086_CR9","doi-asserted-by":"publisher","unstructured":"Chen J, Fan ZX, Zhu DC, Guo YL, Ye K, Dai D, Guo Z, Hu ZQ, Miao Y, Qu Q. Emerging Role of Dermal White Adipose Tissue in Modulating Hair Follicle Development During Aging. Front Cell Dev Biol. 2021 Oct 15;9:728188. 10.3389\/fcell.2021.728188. PMID: 34722509; PMCID: PMC8554130. https:\/\/doi.org\/10.3389\/fcell.2021.728188.","DOI":"10.3389\/fcell.2021.728188"},{"issue":"1","key":"3086_CR10","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1016\/j.stem.2021.09.009","volume":"29","author":"Y Xie","year":"2022","unstructured":"Xie Y, Chen D, Jiang K, et al. Hair shaft miniaturization causes stem cell depletion through mechanosensory signals mediated by a Piezo1-calcium-TNF-\u03b1 axis. Cell Stem Cell. 2022;29(1):70\u2013e856.","journal-title":"Cell Stem Cell"},{"key":"3086_CR11","doi-asserted-by":"publisher","unstructured":"Xiong L, Zhevlakova I, West XZ, Gao D, Murtazina R, Horak A, Brown JM, Molokotina I, Podrez EA, Byzova TV. TLR2 regulates hair follicle cycle and regeneration via BMP signaling. Elife. 2024 Mar 14;12:RP89335. https:\/\/doi.org\/10.7554\/eLife.89335","DOI":"10.7554\/eLife.89335"},{"key":"3086_CR12","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1016\/j.jare.2024.05.006","volume":"70","author":"S Zhou","year":"2025","unstructured":"Zhou S, Li Z, Li X, et al. Crosstalk between endothelial cells and dermal papilla entails hair regeneration and angiogenesis during aging. J Adv Res. 2025;70:339\u201353.","journal-title":"J Adv Res"},{"issue":"1","key":"3086_CR13","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.cell.2014.02.013","volume":"157","author":"DT Scadden","year":"2014","unstructured":"Scadden DT. Nice neighborhood: emerging concepts of the stem cell niche. Cell. 2014;157(1):41\u201350.","journal-title":"Cell"},{"issue":"1","key":"3086_CR14","doi-asserted-by":"publisher","first-page":"46","DOI":"10.1046\/j.1523-1747.2003.12171.x","volume":"8","author":"VA Botchkarev","year":"2003","unstructured":"Botchkarev VA, Kishimoto J. Molecular control of epithelial-mesenchymal interactions during hair follicle cycling. J Investig Dermatol Symp Proc. 2003;8(1):46\u201355.","journal-title":"J Investig Dermatol Symp Proc"},{"key":"3086_CR15","doi-asserted-by":"publisher","unstructured":"Wang Q, Oh JW, Lee HL, Dhar A, Peng T, Ramos R, Guerrero-Juarez CF, Wang X, Zhao R, Cao X, Le J, Fuentes MA, Jocoy SC, Rossi AR, Vu B, Pham K, Wang X, Mali NM, Park JM, Choi JH, Lee H, Legrand JMD, Kandyba E, Kim JC, Kim M, Foley J, Yu Z, Kobielak K, Andersen B, Khosrotehrani K, Nie Q, Plikus MV. A multi-scale model for hair follicles reveals heterogeneous domains driving rapid spatiotemporal hair growth patterning. Elife. 2017 Jul 11;6:e22772. https:\/\/doi.org\/10.7554\/eLife.22772","DOI":"10.7554\/eLife.22772"},{"issue":"7176","key":"3086_CR16","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1038\/nature06457","volume":"451","author":"MV Plikus","year":"2008","unstructured":"Plikus MV, Mayer JA, De La Cruz D, et al. Cyclic dermal BMP signalling regulates stem cell activation during hair regeneration. Nature. 2008;451(7176):340\u20134.","journal-title":"Nature"},{"issue":"1","key":"3086_CR17","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/j.cell.2010.11.049","volume":"144","author":"YC Hsu","year":"2011","unstructured":"Hsu YC, Pasolli HA, Fuchs E. Dynamics between stem cells, niche, and progeny in the hair follicle. Cell. 2011;144(1):92\u2013105.","journal-title":"Cell"},{"issue":"5","key":"3086_CR18","doi-asserted-by":"publisher","first-page":"635","DOI":"10.1016\/j.cell.2004.08.012","volume":"118","author":"C Blanpain","year":"2004","unstructured":"Blanpain C, Lowry WE, Geoghegan A, et al. Self-renewal, multipotency, and the existence of two cell populations within an epithelial stem cell niche. Cell. 2004;118(5):635\u201348.","journal-title":"Cell"},{"issue":"2","key":"3086_CR19","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1038\/s41580-023-00662-3","volume":"25","author":"B Zhang","year":"2024","unstructured":"Zhang B, Chen T. Local and systemic mechanisms that control the hair follicle stem cell niche. Nat Rev Mol Cell Biol. 2024;25(2):87\u2013100.","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"12","key":"3086_CR20","doi-asserted-by":"publisher","first-page":"3001","DOI":"10.1016\/j.celrep.2016.02.078","volume":"14","author":"A Rezza","year":"2016","unstructured":"Rezza A, Wang Z, Sennett R, et al. Signaling Networks among Stem Cell Precursors, Transit-Amplifying Progenitors, and their Niche in Developing Hair Follicles. Cell Rep. 2016;14(12):3001\u201318.","journal-title":"Cell Rep"},{"issue":"2","key":"3086_CR21","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1016\/j.stem.2008.12.009","volume":"4","author":"V Greco","year":"2009","unstructured":"Greco V, Chen T, Rendl M, et al. A two-step mechanism for stem cell activation during hair regeneration. Cell Stem Cell. 2009;4(2):155\u201369.","journal-title":"Cell Stem Cell"},{"issue":"4","key":"3086_CR22","doi-asserted-by":"publisher","first-page":"935","DOI":"10.1016\/j.cell.2014.02.057","volume":"157","author":"YC Hsu","year":"2014","unstructured":"Hsu YC, Li L, Fuchs E. Transit-amplifying cells orchestrate stem cell activity and tissue regeneration. Cell. 2014;157(4):935\u201349.","journal-title":"Cell"},{"issue":"4","key":"3086_CR23","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1016\/S0092-8674(00)00050-7","volume":"102","author":"G Taylor","year":"2000","unstructured":"Taylor G, Lehrer MS, Jensen PJ, et al. Involvement of follicular stem cells in forming not only the follicle but also the epidermis. Cell. 2000;102(4):451\u201361.","journal-title":"Cell"},{"issue":"2","key":"3086_CR24","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1016\/S0092-8674(01)00208-2","volume":"104","author":"H Oshima","year":"2001","unstructured":"Oshima H, Rochat A, Kedzia C, et al. Morphogenesis and renewal of hair follicles from adult multipotent stem cells. Cell. 2001;104(2):233\u201345.","journal-title":"Cell"},{"issue":"6883","key":"3086_CR25","doi-asserted-by":"publisher","first-page":"854","DOI":"10.1038\/416854a","volume":"416","author":"EK Nishimura","year":"2002","unstructured":"Nishimura EK, Jordan SA, Oshima H, et al. Dominant role of the niche in melanocyte stem-cell fate determination. Nature. 2002;416(6883):854\u201360.","journal-title":"Nature"},{"issue":"6","key":"3086_CR26","first-page":"1601","volume":"151","author":"G Lindner","year":"1997","unstructured":"Lindner G, Botchkarev VA, Botchkareva NV, et al. Analysis of apoptosis during hair follicle regression (catagen). Am J Pathol. 1997;151(6):1601\u201317.","journal-title":"Am J Pathol"},{"issue":"1","key":"3086_CR27","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/S1534-5807(01)00022-3","volume":"1","author":"E Fuchs","year":"2001","unstructured":"Fuchs E, Merrill BJ, Jamora C, et al. At the Roots of a Never-Ending Cycle. Dev Cell. 2001;1(1):13\u201325.","journal-title":"Dev Cell"},{"issue":"10","key":"3086_CR28","doi-asserted-by":"publisher","first-page":"5339","DOI":"10.1073\/pnas.1901720117","volume":"117","author":"Y Ge","year":"2020","unstructured":"Ge Y, Miao Y, Gur-Cohen S, et al. The aging skin microenvironment dictates stem cell behavior. Proc Natl Acad Sci U S A. 2020;117(10):5339\u201350.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"8","key":"3086_CR29","doi-asserted-by":"publisher","first-page":"2086","DOI":"10.1038\/jid.2014.139","volume":"134","author":"CC Chen","year":"2014","unstructured":"Chen CC, Murray PJ, Jiang TX, et al. Regenerative hair waves in aging mice and extra-follicular modulators follistatin, dkk1, and sfrp4. J Invest Dermatol. 2014;134(8):2086\u201396.","journal-title":"J Invest Dermatol"},{"issue":"9","key":"3086_CR30","doi-asserted-by":"publisher","first-page":"795","DOI":"10.1038\/s43587-021-00094-8","volume":"1","author":"Y Yu","year":"2021","unstructured":"Yu Y, Zhang X, Liu F, et al. A stress-induced miR-31-CLOCK-ERK pathway is a key driver and therapeutic target for skin aging. Nat Aging. 2021;1(9):795\u2013809.","journal-title":"Nat Aging"},{"issue":"51","key":"3086_CR31","doi-asserted-by":"publisher","first-page":"E4950","DOI":"10.1073\/pnas.1320301110","volume":"110","author":"BE Keyes","year":"2013","unstructured":"Keyes BE, Segal JP, Heller E, et al. Nfatc1 orchestrates aging in hair follicle stem cells. Proc Natl Acad Sci U S A. 2013;110(51):E4950\u20139.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"3086_CR32","doi-asserted-by":"publisher","first-page":"3105","DOI":"10.1007\/s11357-023-01042-7","volume":"46","author":"A Ungvari","year":"2024","unstructured":"Ungvari A, Kiss T, Gulej R, et al. Irradiation-induced hair graying in mice: an experimental model to evaluate the effectiveness of interventions targeting oxidative stress, DNA damage prevention, and cellular senescence. Geroscience. 2024;46(3):3105\u201322.","journal-title":"Geroscience"},{"issue":"2","key":"3086_CR33","doi-asserted-by":"publisher","first-page":"1215","DOI":"10.1007\/s11357-022-00592-6","volume":"45","author":"DM Dai","year":"2023","unstructured":"Dai DM, He Y, Guan Q, et al. Modeling human gray hair by irradiation as a valuable tool to study aspects of tissue aging. Geroscience. 2023;45(2):1215\u201330.","journal-title":"Geroscience"},{"issue":"8","key":"3086_CR34","doi-asserted-by":"publisher","first-page":"3571","DOI":"10.7150\/thno.103221","volume":"15","author":"M Wang","year":"2025","unstructured":"Wang M, Wang M, Jiang J, et al. THSD4 promotes hair growth by facilitating dermal papilla and hair matrix interactions. Theranostics. 2025;15(8):3571\u201388.","journal-title":"Theranostics"},{"issue":"2","key":"3086_CR35","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1016\/j.devcel.2020.03.019","volume":"53","author":"W Shin","year":"2020","unstructured":"Shin W, Rosin NL, Sparks H, et al. Dysfunction of Hair Follicle Mesenchymal Progenitors Contributes to Age-Associated Hair Loss. Dev Cell. 2020;53(2):185\u2013e1987.","journal-title":"Dev Cell"},{"issue":"7","key":"3086_CR36","doi-asserted-by":"publisher","first-page":"771","DOI":"10.1038\/s41556-021-00705-x","volume":"23","author":"J Koester","year":"2021","unstructured":"Koester J, Miroshnikova YA, Ghatak S, et al. Niche stiffening compromises hair follicle stem cell potential during ageing by reducing bivalent promoter accessibility. Nat Cell Biol. 2021;23(7):771\u201381.","journal-title":"Nat Cell Biol"},{"issue":"1","key":"3086_CR37","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1186\/s12929-022-00800-7","volume":"29","author":"YH Jung","year":"2022","unstructured":"Jung YH, Chae CW, Choi GE, et al. Cyanidin 3-O-arabinoside suppresses DHT-induced dermal papilla cell senescence by modulating p38-dependent ER-mitochondria contacts. J Biomed Sci. 2022;29(1):17.","journal-title":"J Biomed Sci"},{"issue":"1","key":"3086_CR38","doi-asserted-by":"publisher","first-page":"e14353","DOI":"10.1111\/acel.14353","volume":"24","author":"A Pappalardo","year":"2025","unstructured":"Pappalardo A, Kim JY, Abaci HE, et al. Restoration of hair follicle inductive properties by depletion of senescent cells. Aging Cell. 2025;24(1):e14353.","journal-title":"Aging Cell"},{"issue":"5611","key":"3086_CR39","doi-asserted-by":"publisher","first-page":"1355","DOI":"10.1126\/science.1079161","volume":"299","author":"P Hasty","year":"2003","unstructured":"Hasty P, Campisi J, Hoeijmakers J, van Steeg H, Vijg J. Aging and genome maintenance: lessons from the mouse? Science. 2003 Feb 28;299(5611):1355-9. doi: 10.1126\/science.1079161. PMID: 12610296.","journal-title":"Science"},{"issue":"2","key":"3086_CR40","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1016\/j.molcel.2010.10.002","volume":"40","author":"MC Haigis","year":"2010","unstructured":"Haigis MC, Yankner BA. The aging stress response. Mol Cell. 2010;40(2):333\u201344.","journal-title":"Mol Cell"},{"issue":"7372","key":"3086_CR41","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1038\/nature10600","volume":"479","author":"DJ Baker","year":"2011","unstructured":"Baker DJ, Wijshake T, Tchkonia T, et al. Clearance of p16Ink4a-positive senescent cells delays ageing-associated disorders. Nature. 2011;479(7372):232\u20136.","journal-title":"Nature"},{"issue":"10","key":"3086_CR42","doi-asserted-by":"publisher","first-page":"889","DOI":"10.1038\/s43587-021-00103-w","volume":"1","author":"C Zhang","year":"2021","unstructured":"Zhang C, Wang D, Wang J, et al. Escape of hair follicle stem cells causes stem cell exhaustion during aging. Nat Aging. 2021;1(10):889\u2013903.","journal-title":"Nat Aging"},{"issue":"6273","key":"3086_CR43","doi-asserted-by":"publisher","first-page":"aad4395","DOI":"10.1126\/science.aad4395","volume":"351","author":"H Matsumura","year":"2016","unstructured":"Matsumura H, Mohri Y, Binh NT, et al. Hair follicle aging is driven by transepidermal elimination of stem cells via COL17A1 proteolysis. Science. 2016;351(6273):aad4395.","journal-title":"Science"},{"issue":"2","key":"3086_CR44","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1016\/j.stem.2010.11.029","volume":"8","author":"S Tanimura","year":"2011","unstructured":"Tanimura S, Tadokoro Y, Inomata K, et al. Hair follicle stem cells provide a functional niche for melanocyte stem cells. Cell Stem Cell. 2011;8(2):177\u201387.","journal-title":"Cell Stem Cell"},{"issue":"7","key":"3086_CR45","doi-asserted-by":"publisher","first-page":"1358","DOI":"10.1096\/fj.06-6926com","volume":"21","author":"V Levy","year":"2007","unstructured":"Levy V, Lindon C, Zheng Y, et al. Epidermal stem cells arise from the hair follicle after wounding. Faseb j. 2007;21(7):1358\u201366.","journal-title":"Faseb j"},{"issue":"4","key":"3086_CR46","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1038\/nbt950","volume":"22","author":"RJ Morris","year":"2004","unstructured":"Morris RJ, Liu Y, Marles L, et al. Capturing and profiling adult hair follicle stem cells. Nat Biotechnol. 2004;22(4):411\u20137.","journal-title":"Nat Biotechnol"},{"key":"3086_CR47","doi-asserted-by":"crossref","unstructured":"Ito M, Kizawa K, Hamada K et al. Hair follicle stem cells in the lower bulge form the secondary germ, a biochemically distinct but functionally equivalent progenitor cell population, at the termination of catagen. Differentiation. 2004;72(9\u201310): 548\u2009\u2013\u200957.","DOI":"10.1111\/j.1432-0436.2004.07209008.x"},{"issue":"2","key":"3086_CR48","doi-asserted-by":"publisher","first-page":"190","DOI":"10.1038\/s43587-021-00033-7","volume":"1","author":"H Matsumura","year":"2021","unstructured":"Matsumura H, Liu N, Nanba D, et al. Distinct types of stem cell divisions determine organ regeneration and aging in hair follicles. Nat Aging. 2021;1(2):190\u2013204.","journal-title":"Nat Aging"},{"issue":"7958","key":"3086_CR49","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1038\/s41586-023-05960-6","volume":"616","author":"Q Sun","year":"2023","unstructured":"Sun Q, Lee W, Hu H, et al. Dedifferentiation maintains melanocyte stem cells in a dynamic niche. Nature. 2023;616(7958):774\u201382.","journal-title":"Nature"},{"issue":"7792","key":"3086_CR50","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/s41586-020-1935-3","volume":"577","author":"B Zhang","year":"2020","unstructured":"Zhang B, Ma S, Rachmin I, et al. Hyperactivation of sympathetic nerves drives depletion of melanocyte stem cells. Nature. 2020;577(7792):676\u201381.","journal-title":"Nature"},{"issue":"7866","key":"3086_CR51","doi-asserted-by":"publisher","first-page":"266","DOI":"10.1038\/s41586-021-03624-x","volume":"595","author":"H Morinaga","year":"2021","unstructured":"Morinaga H, Mohri Y, Grachtchouk M, et al. Obesity accelerates hair thinning by stem cell-centric converging mechanisms. Nature. 2021;595(7866):266\u201371.","journal-title":"Nature"},{"issue":"5","key":"3086_CR52","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1016\/j.devcel.2014.10.022","volume":"31","author":"W Rahmani","year":"2014","unstructured":"Rahmani W, Abbasi S, Hagner A, et al. Hair Follicle Dermal Stem Cells Regenerate the Dermal Sheath, Repopulate the Dermal Papilla, and Modulate Hair Type. Dev Cell. 2014;31(5):543\u201358.","journal-title":"Dev Cell"},{"issue":"14","key":"3086_CR53","doi-asserted-by":"publisher","first-page":"1758","DOI":"10.1016\/j.devcel.2022.06.005","volume":"57","author":"Y Liu","year":"2022","unstructured":"Liu Y, Guerrero-Juarez CF, Xiao F, et al. Hedgehog signaling reprograms hair follicle niche fibroblasts to a hyper-activated state. Dev Cell. 2022;57(14):1758\u2013e17757.","journal-title":"Dev Cell"},{"issue":"6474","key":"3086_CR54","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1126\/science.aax9131","volume":"367","author":"N Heitman","year":"2020","unstructured":"Heitman N, Sennett R, Mok KW, et al. Dermal sheath contraction powers stem cell niche relocation during hair cycle regression. Science. 2020;367(6474):161\u20136.","journal-title":"Science"},{"issue":"4","key":"3086_CR55","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1016\/j.cell.2011.01.014","volume":"144","author":"H Fujiwara","year":"2011","unstructured":"Fujiwara H, Ferreira M, Donati G, et al. The basement membrane of hair follicle stem cells is a muscle cell niche. Cell. 2011;144(4):577\u201389.","journal-title":"Cell"},{"issue":"24","key":"3086_CR56","doi-asserted-by":"publisher","first-page":"7025","DOI":"10.1016\/j.cell.2024.10.019","volume":"187","author":"S Ma","year":"2024","unstructured":"Ma S, Ji Z, Zhang B, et al. Spatial transcriptomic landscape unveils immunoglobin-associated senescence as a hallmark of aging. Cell. 2024;187(24):7025\u2013e704434.","journal-title":"Cell"},{"issue":"5","key":"3086_CR57","doi-asserted-by":"publisher","first-page":"334","DOI":"10.1038\/s41580-022-00568-6","volume":"24","author":"A De Morree","year":"2023","unstructured":"De Morree A, Rando TA. Regulation of adult stem cell quiescence and its functions in the maintenance of tissue integrity. Nat Rev Mol Cell Biol. 2023;24(5):334\u201354.","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"1","key":"3086_CR58","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1038\/s41580-022-00510-w","volume":"24","author":"A Brunet","year":"2023","unstructured":"Brunet A, Goodell MA, Rando TA. Ageing and rejuvenation of tissue stem cells and their niches. Nat Rev Mol Cell Biol. 2023;24(1):45\u201362.","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"4","key":"3086_CR59","doi-asserted-by":"publisher","first-page":"1351","DOI":"10.1073\/pnas.1121312110","volume":"110","author":"E Kandyba","year":"2013","unstructured":"Kandyba E, Leung Y, Chen YB, et al. Competitive balance of intrabulge BMP\/Wnt signaling reveals a robust gene network ruling stem cell homeostasis and cyclic activation. Proc Natl Acad Sci U S A. 2013;110(4):1351\u20136.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"3086_CR60","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1038\/11078","volume":"1","author":"VA Botchkarev","year":"1999","unstructured":"Botchkarev VA, Botchkareva NV, Roth W, et al. Noggin is a mesenchymally derived stimulator of hair-follicle induction. Nat Cell Biol. 1999;1(3):158\u201364.","journal-title":"Nat Cell Biol"},{"issue":"6029","key":"3086_CR61","doi-asserted-by":"publisher","first-page":"586","DOI":"10.1126\/science.1201647","volume":"332","author":"MV Plikus","year":"2011","unstructured":"Plikus MV, Baker RE, Chen CC, et al. Self-organizing and stochastic behaviors during the regeneration of hair stem cells. Science. 2011;332(6029):586\u20139.","journal-title":"Science"},{"issue":"1","key":"3086_CR62","doi-asserted-by":"publisher","first-page":"62","DOI":"10.1093\/jmcb\/mjv005","volume":"7","author":"Z Deng","year":"2015","unstructured":"Deng Z, Lei X, Zhang X, et al. mTOR signaling promotes stem cell activation via counterbalancing BMP-mediated suppression during hair regeneration. J Mol Cell Biol. 2015;7(1):62\u201372.","journal-title":"J Mol Cell Biol"},{"issue":"4","key":"3086_CR63","doi-asserted-by":"publisher","first-page":"4778","DOI":"10.18632\/aging.202694","volume":"13","author":"RL Tiwari","year":"2021","unstructured":"Tiwari RL, Mishra P, Martin N, et al. A Wnt5a-Cdc42 axis controls aging and rejuvenation of hair-follicle stem cells. Aging. 2021;13(4):4778\u201393.","journal-title":"Aging"},{"issue":"4","key":"3086_CR64","doi-asserted-by":"publisher","first-page":"979","DOI":"10.1038\/jid.2010.397","volume":"131","author":"S Kauser","year":"2011","unstructured":"Kauser S, Westgate GE, Green MR, et al. Human hair follicle and epidermal melanocytes exhibit striking differences in their aging profile which involves catalase. J Invest Dermatol. 2011;131(4):979\u201382.","journal-title":"J Invest Dermatol"},{"issue":"1","key":"3086_CR65","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1111\/brv.12648","volume":"96","author":"JDB O\u2019sullivan","year":"2021","unstructured":"O\u2019sullivan JDB, Nicu C, Picard M, et al. The biology of human hair greying. Biol Rev Camb Philos Soc. 2021;96(1):107\u201328.","journal-title":"Biol Rev Camb Philos Soc"},{"issue":"5710","key":"3086_CR66","doi-asserted-by":"publisher","first-page":"720","DOI":"10.1126\/science.1099593","volume":"307","author":"EK Nishimura","year":"2005","unstructured":"Nishimura EK, Granter SR, Fisher DE. Mechanisms of hair graying: incomplete melanocyte stem cell maintenance in the niche. Science. 2005;307(5710):720\u20134.","journal-title":"Science"},{"issue":"6","key":"3086_CR67","doi-asserted-by":"publisher","first-page":"1088","DOI":"10.1016\/j.cell.2009.03.037","volume":"137","author":"K Inomata","year":"2009","unstructured":"Inomata K, Aoto T, Binh NT, et al. Genotoxic stress abrogates renewal of melanocyte stem cells by triggering their differentiation. Cell. 2009;137(6):1088\u201399.","journal-title":"Cell"},{"issue":"3","key":"3086_CR68","doi-asserted-by":"publisher","first-page":"435","DOI":"10.1046\/j.1365-2133.2004.05787.x","volume":"150","author":"S Commo","year":"2004","unstructured":"Commo S, Gaillard O, Bernard BA. Human hair greying is linked to a specific depletion of hair follicle melanocytes affecting both the bulb and the outer root sheath. Br J Dermatol. 2004;150(3):435\u201343.","journal-title":"Br J Dermatol"},{"issue":"2","key":"3086_CR69","doi-asserted-by":"publisher","first-page":"e13083","DOI":"10.1111\/acel.13083","volume":"19","author":"CSL Tuttle","year":"2020","unstructured":"Tuttle CSL, Waaijer MEC, Slee-Valentijn MS, et al. Cellular senescence and chronological age in various human tissues: A systematic review and meta-analysis. Aging Cell. 2020;19(2):e13083.","journal-title":"Aging Cell"},{"issue":"1","key":"3086_CR70","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1186\/s13024-021-00507-7","volume":"17","author":"P Xu","year":"2022","unstructured":"Xu P, Wang M, Song WM, et al. The landscape of human tissue and cell type specific expression and co-regulation of senescence genes. Mol Neurodegener. 2022;17(1):5.","journal-title":"Mol Neurodegener"},{"key":"3086_CR71","doi-asserted-by":"publisher","first-page":"102686","DOI":"10.1016\/j.arr.2025.102686","volume":"105","author":"H Thau","year":"2025","unstructured":"Thau H, Gerjol BP, Hahn K, et al. Senescence as a molecular target in skin aging and disease. Ageing Res Rev. 2025;105:102686.","journal-title":"Ageing Res Rev"},{"issue":"3","key":"3086_CR72","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1016\/j.cell.2017.03.038","volume":"169","author":"H Yang","year":"2017","unstructured":"Yang H, Adam RC, Ge Y, et al. Epithelial-Mesenchymal Micro-niches Govern Stem Cell Lineage Choices. Cell. 2017;169(3):483\u2013e49613.","journal-title":"Cell"},{"issue":"1","key":"3086_CR73","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/j.stem.2011.11.005","volume":"10","author":"N Oshimori","year":"2012","unstructured":"Oshimori N, Fuchs E. Paracrine TGF-\u03b2 signaling counterbalances BMP-mediated repression in hair follicle stem cell activation. Cell Stem Cell. 2012;10(1):63\u201375.","journal-title":"Cell Stem Cell"},{"issue":"6859","key":"3086_CR74","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1038\/35102160","volume":"414","author":"A Spradling","year":"2001","unstructured":"Spradling A, Drummond-Barbosa D, Kai T. Stem cells find their niche. Nature. 2001;414(6859):98\u2013104.","journal-title":"Nature"},{"issue":"6","key":"3086_CR75","doi-asserted-by":"publisher","first-page":"640","DOI":"10.1002\/wsbm.86","volume":"2","author":"J Seita","year":"2010","unstructured":"Seita J, Weissman IL. Hematopoietic stem cell: self-renewal versus differentiation. Wiley Interdiscip Rev Syst Biol Med. 2010;2(6):640\u201353.","journal-title":"Wiley Interdiscip Rev Syst Biol Med"},{"issue":"4","key":"3086_CR76","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1016\/j.cell.2008.01.025","volume":"132","author":"SH Orkin","year":"2008","unstructured":"Orkin SH, Zon LI. Hematopoiesis: an evolving paradigm for stem cell biology. Cell. 2008;132(4):631\u201344.","journal-title":"Cell"},{"issue":"2","key":"3086_CR77","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1016\/j.cell.2022.11.001","volume":"186","author":"C L\u00f3pez-Ot\u00edn","year":"2023","unstructured":"L\u00f3pez-Ot\u00edn C, Blasco MA, Partridge L, et al. Hallmarks of aging: An expanding universe. Cell. 2023;186(2):243\u201378.","journal-title":"Cell"},{"issue":"2","key":"3086_CR78","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.molmed.2021.12.003","volume":"28","author":"AC Franco","year":"2022","unstructured":"Franco AC, Aveleira C, Cavadas C. Skin senescence: mechanisms and impact on whole-body aging. Trends Mol Med. 2022;28(2):97\u2013109.","journal-title":"Trends Mol Med"},{"key":"3086_CR79","doi-asserted-by":"publisher","unstructured":"Harn HI, Chen CC, Wang SP, Lei M, Chuong CM. Tissue Mechanics in Haired Murine Skin: Potential Implications for Skin Aging. Front Cell Dev Biol. 2021 Feb 19;9:635340. https:\/\/doi.org\/10.3389\/fcell.2021.635340","DOI":"10.3389\/fcell.2021.635340"},{"key":"3086_CR80","doi-asserted-by":"publisher","unstructured":"Li KN, Jain P, He CH, Eun FC, Kang S, Tumbar T. Skin vasculature and hair follicle cross-talking associated with stem cell activation and tissue homeostasis. Elife. 2019 Jul 25;8:e45977. https:\/\/doi.org\/10.7554\/eLife.45977","DOI":"10.7554\/eLife.45977"},{"issue":"5","key":"3086_CR81","doi-asserted-by":"publisher","first-page":"909","DOI":"10.1046\/j.1523-1747.2000.00954.x","volume":"114","author":"L Mecklenburg","year":"2000","unstructured":"Mecklenburg L, Tobin DJ, M\u00fcller-R\u00f6ver S, et al. Active hair growth (anagen) is associated with angiogenesis. J Invest Dermatol. 2000;114(5):909\u201316.","journal-title":"J Invest Dermatol"},{"issue":"4","key":"3086_CR82","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1172\/JCI11317","volume":"107","author":"K Yano","year":"2001","unstructured":"Yano K, Brown LF, Detmar M. Control of hair growth and follicle size by VEGF-mediated angiogenesis. J Clin Invest. 2001;107(4):409\u201317.","journal-title":"J Clin Invest"},{"issue":"6","key":"3086_CR83","first-page":"398","volume":"14","author":"Q Zhao","year":"2023","unstructured":"Zhao Q, Zheng Y, Zhao D, et al. Single-cell profiling reveals a potent role of quercetin in promoting hair regeneration. Protein Cell. 2023;14(6):398\u2013415.","journal-title":"Protein Cell"},{"issue":"2","key":"3086_CR84","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1016\/j.cell.2015.02.016","volume":"161","author":"CC Chen","year":"2015","unstructured":"Chen CC, Wang L, Plikus MV, et al. Organ-level quorum sensing directs regeneration in hair stem cell populations. Cell. 2015;161(2):277\u201390.","journal-title":"Cell"},{"issue":"6","key":"3086_CR85","doi-asserted-by":"publisher","first-page":"461","DOI":"10.1038\/nri3464","volume":"13","author":"S Sakaguchi","year":"2013","unstructured":"Sakaguchi S, Vignali DA, Rudensky AY, et al. The plasticity and stability of regulatory T cells. Nat Rev Immunol. 2013;13(6):461\u20137.","journal-title":"Nat Rev Immunol"},{"issue":"7350","key":"3086_CR86","doi-asserted-by":"publisher","first-page":"216","DOI":"10.1038\/nature10160","volume":"474","author":"J Fujisaki","year":"2011","unstructured":"Fujisaki J, Wu J, Carlson AL, et al. In vivo imaging of Treg cells providing immune privilege to the haematopoietic stem-cell niche. Nature. 2011;474(7350):216\u20139.","journal-title":"Nature"},{"issue":"3","key":"3086_CR87","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1016\/j.stem.2018.01.017","volume":"22","author":"Y Hirata","year":"2018","unstructured":"Hirata Y, Furuhashi K, Ishii H, et al. CD150(high) Bone Marrow Tregs Maintain Hematopoietic Stem Cell Quiescence and Immune Privilege via Adenosine. Cell Stem Cell. 2018;22(3):445\u2013e4535.","journal-title":"Cell Stem Cell"},{"issue":"6","key":"3086_CR88","doi-asserted-by":"publisher","first-page":"1119","DOI":"10.1016\/j.cell.2017.05.002","volume":"169","author":"N Ali","year":"2017","unstructured":"Ali N, Zirak B, Rodriguez RS, et al. Regulatory T Cells in Skin Facilitate Epithelial Stem Cell Differentiation. Cell. 2017;169(6):1119\u2013e112911.","journal-title":"Cell"},{"issue":"7","key":"3086_CR89","doi-asserted-by":"publisher","first-page":"1086","DOI":"10.1038\/s41590-022-01244-9","volume":"23","author":"Z Liu","year":"2022","unstructured":"Liu Z, Hu X, Liang Y, et al. Glucocorticoid signaling and regulatory T cells cooperate to maintain the hair-follicle stem-cell niche. Nat Immunol. 2022;23(7):1086\u201397.","journal-title":"Nat Immunol"},{"issue":"1","key":"3086_CR90","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1634\/stemcells.2007-0563","volume":"26","author":"SL Tomchuck","year":"2008","unstructured":"Tomchuck SL, Zwezdaryk KJ, Coffelt SB, et al. Toll-like receptors on human mesenchymal stem cells drive their migration and immunomodulating responses. Stem Cells. 2008;26(1):99\u2013107.","journal-title":"Stem Cells"},{"issue":"6","key":"3086_CR91","doi-asserted-by":"publisher","first-page":"801","DOI":"10.1016\/j.immuni.2006.04.008","volume":"24","author":"Y Nagai","year":"2006","unstructured":"Nagai Y, Garrett KP, Ohta S, et al. Toll-like receptors on hematopoietic progenitor cells stimulate innate immune system replenishment. Immunity. 2006;24(6):801\u201312.","journal-title":"Immunity"},{"issue":"12","key":"3086_CR92","doi-asserted-by":"publisher","first-page":"875","DOI":"10.1038\/nri3547","volume":"13","author":"AC Shaw","year":"2013","unstructured":"Shaw AC, Goldstein DR, Montgomery RR. Age-dependent dysregulation of innate immunity. Nat Rev Immunol. 2013;13(12):875\u201387.","journal-title":"Nat Rev Immunol"},{"key":"3086_CR93","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1038\/s41536-020-0095-2","volume":"5","author":"W Rahmani","year":"2020","unstructured":"Rahmani W, Sinha S, Biernaskie J. Immune modulation of hair follicle regeneration. NPJ Regen Med. 2020;5:9.","journal-title":"NPJ Regen Med"},{"issue":"5","key":"3086_CR94","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1016\/j.cell.2011.07.019","volume":"146","author":"E Festa","year":"2011","unstructured":"Festa E, Fretz J, Berry R, et al. Adipocyte lineage cells contribute to the skin stem cell niche to drive hair cycling. Cell. 2011;146(5):761\u201371.","journal-title":"Cell"},{"issue":"6","key":"3086_CR95","doi-asserted-by":"publisher","first-page":"1575","DOI":"10.1016\/j.cell.2018.10.012","volume":"175","author":"MC Salzer","year":"2018","unstructured":"Salzer MC, Lafzi A, Berenguer-Llergo A, et al. Identity Noise and Adipogenic Traits Characterize Dermal Fibroblast Aging. Cell. 2018;175(6):1575\u2013e159022.","journal-title":"Cell"},{"key":"3086_CR96","doi-asserted-by":"publisher","first-page":"101150","DOI":"10.1016\/j.arr.2020.101150","volume":"63","author":"A Angelini","year":"2020","unstructured":"Angelini A, Trial J, Ortiz-Urbina J, et al. Mechanosensing dysregulation in the fibroblast: A hallmark of the aging heart. Ageing Res Rev. 2020;63:101150.","journal-title":"Ageing Res Rev"},{"issue":"2","key":"3086_CR97","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/j.yexcr.2018.11.016","volume":"374","author":"L He","year":"2019","unstructured":"He L, Ahmad M, Perrimon N. Mechanosensitive channels and their functions in stem cell differentiation. Exp Cell Res. 2019;374(2):259\u201365.","journal-title":"Exp Cell Res"},{"key":"3086_CR98","doi-asserted-by":"publisher","unstructured":"Chermnykh E, Kalabusheva E, Vorotelyak E. Extracellular Matrix as a Regulator of Epidermal Stem Cell Fate. Int J Mol Sci. 2018 Mar 27;19(4):1003. https:\/\/doi.org\/10.3390\/ijms19041003","DOI":"10.3390\/ijms19041003"},{"issue":"6","key":"3086_CR99","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1016\/j.stem.2009.05.001","volume":"4","author":"E Fuchs","year":"2009","unstructured":"Fuchs E. Finding one\u2019s niche in the skin. Cell Stem Cell. 2009;4(6):499\u2013502.","journal-title":"Cell Stem Cell"},{"key":"3086_CR100","doi-asserted-by":"publisher","unstructured":"Watt FM, Fujiwara H. Cell-extracellular matrix interactions in normal and diseased skin. Cold Spring Harb Perspect Biol. 2011 Apr 1;3(4):a005124. https:\/\/doi.org\/10.1101\/cshperspect.a005124","DOI":"10.1101\/cshperspect.a005124"},{"key":"3086_CR101","doi-asserted-by":"publisher","first-page":"8198","DOI":"10.1038\/ncomms9198","volume":"6","author":"J Morgner","year":"2015","unstructured":"Morgner J, Ghatak S, Jakobi T, et al. Integrin-linked kinase regulates the niche of quiescent epidermal stem cells. Nat Commun. 2015;6:8198.","journal-title":"Nat Commun"},{"issue":"22","key":"3086_CR102","doi-asserted-by":"publisher","first-page":"eadt2771","DOI":"10.1126\/sciadv.adt2771","volume":"11","author":"J Wang","year":"2025","unstructured":"Wang J, Fu C, Chang S, et al. PIEZO1-mediated calcium signaling reinforces mechanical properties of hair follicle stem cells to promote quiescence. Sci Adv. 2025;11(22):eadt2771.","journal-title":"Sci Adv"},{"issue":"1","key":"3086_CR103","doi-asserted-by":"publisher","first-page":"1524","DOI":"10.1038\/s41467-019-09402-8","volume":"10","author":"SY Chu","year":"2019","unstructured":"Chu SY, Chou CH, Huang HD, et al. Mechanical stretch induces hair regeneration through the alternative activation of macrophages. Nat Commun. 2019;10(1):1524.","journal-title":"Nat Commun"},{"key":"3086_CR104","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1016\/j.actbio.2023.11.032","volume":"175","author":"C-Y Lin","year":"2024","unstructured":"Lin C-Y, Sugerman GP, Kakaletsis S, et al. Sex- and age-dependent skin mechanics\u2014A detailed look in mice. Acta Biomater. 2024;175:106\u201313.","journal-title":"Acta Biomater"},{"key":"3086_CR105","doi-asserted-by":"publisher","first-page":"317","DOI":"10.2147\/CCID.S51958","volume":"6","author":"Q Zheng","year":"2013","unstructured":"Zheng Q, Chen S, Chen Y, et al. Investigation of age-related decline of microfibril-associated glycoprotein-1 in human skin through immunohistochemistry study. Clin Cosmet Investig Dermatol. 2013;6:317\u201323.","journal-title":"Clin Cosmet Investig Dermatol"},{"key":"3086_CR106","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1016\/j.actbio.2019.10.020","volume":"101","author":"WD Meador","year":"2020","unstructured":"Meador WD, Sugerman GP, Story HM, et al. The regional-dependent biaxial behavior of young and aged mouse skin: A detailed histomechanical characterization, residual strain analysis, and constitutive model. Acta Biomater. 2020;101:403\u201313.","journal-title":"Acta Biomater"},{"issue":"3","key":"3086_CR107","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1016\/j.stem.2020.11.002","volume":"28","author":"W Ning","year":"2021","unstructured":"Ning W, Muroyama A, Li H, et al. Differentiated Daughter Cells Regulate Stem Cell Proliferation and Fate through Intra-tissue Tension. Cell Stem Cell. 2021;28(3):436\u2013e4525.","journal-title":"Cell Stem Cell"},{"issue":"8","key":"3086_CR108","first-page":"701","volume":"83","author":"S Devjani","year":"2023","unstructured":"Devjani S, Ezemma O, Kelley KJ, et al. Androgenetic Alopecia: Therapy Update Drugs. 2023;83(8):701\u201315.","journal-title":"Androgenetic Alopecia: Therapy Update Drugs"},{"issue":"1","key":"3086_CR109","doi-asserted-by":"publisher","first-page":"128","DOI":"10.1177\/1090820X12468314","volume":"33","author":"JE Vogel","year":"2013","unstructured":"Vogel JE, Jimenez F, Cole J, et al. Hair restoration surgery: the state of the art. Aesthet Surg J. 2013;33(1):128\u201351.","journal-title":"Aesthet Surg J"},{"key":"3086_CR110","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1016\/j.mad.2017.07.001","volume":"170","author":"JP De Magalh\u00e3es","year":"2018","unstructured":"De Magalh\u00e3es JP, Passos JF. Stress, cell senescence and organismal ageing. Mech Ageing Dev. 2018;170:2\u20139.","journal-title":"Mech Ageing Dev"},{"issue":"2","key":"3086_CR111","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1158\/2159-8290.CD-16-0241","volume":"7","author":"M Demaria","year":"2017","unstructured":"Demaria M, O\u2019leary MN, Chang J, et al. Cellular Senescence Promotes Adverse Effects of Chemotherapy and Cancer Relapse. Cancer Discov. 2017;7(2):165\u201376.","journal-title":"Cancer Discov"},{"key":"3086_CR112","doi-asserted-by":"publisher","first-page":"685","DOI":"10.1146\/annurev-physiol-030212-183653","volume":"75","author":"J Campisi","year":"2013","unstructured":"Campisi J. Aging, cellular senescence, and cancer. Annu Rev Physiol. 2013;75:685\u2013705.","journal-title":"Annu Rev Physiol"},{"issue":"2","key":"3086_CR113","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1038\/s41580-020-00314-w","volume":"22","author":"R Di Micco","year":"2021","unstructured":"Di Micco R, Krizhanovsky V, Baker D, et al. Cellular senescence in ageing: from mechanisms to therapeutic opportunities. Nat Rev Mol Cell Biol. 2021;22(2):75\u201395.","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"7589","key":"3086_CR114","doi-asserted-by":"publisher","first-page":"184","DOI":"10.1038\/nature16932","volume":"530","author":"DJ Baker","year":"2016","unstructured":"Baker DJ, Childs BG, Durik M, et al. Naturally occurring p16(Ink4a)-positive cells shorten healthy lifespan. Nature. 2016;530(7589):184\u20139.","journal-title":"Nature"},{"key":"3086_CR115","doi-asserted-by":"publisher","unstructured":"Wang Y, Yang J, Luo Y, Zhao Z, Yuan Y, Li J, Liu Y, Yi Y, Xu X, Lan Y, Zou J, Li Q, Wang L, Pan Y, Yang Y, Xiong M, Wu M, Li J, Li W, Zhang Y, Cao Y, Zhu Y, Xiao ZJ. Targeting IGF1-Induced Cellular Senescence to Rejuvenate Hair Follicle Aging. Aging Cell. 2025 Jul;24(7):e70053. https:\/\/doi.org\/10.1111\/acel.70053","DOI":"10.1111\/acel.70053"},{"key":"3086_CR116","doi-asserted-by":"publisher","first-page":"11190","DOI":"10.1038\/ncomms11190","volume":"7","author":"R Yosef","year":"2016","unstructured":"Yosef R, Pilpel N, Tokarsky-Amiel R, et al. Directed elimination of senescent cells by inhibition of BCL-W and BCL-XL. Nat Commun. 2016;7:11190.","journal-title":"Nat Commun"},{"issue":"6616","key":"3086_CR117","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1126\/science.abf3326","volume":"378","author":"NS Reyes","year":"2022","unstructured":"Reyes NS, Krasilnikov M, Allen NC, et al. Sentinel p16(INK4a+) cells in the basement membrane form a reparative niche in the lung. Science. 2022;378(6616):192\u2013201.","journal-title":"Science"},{"issue":"4","key":"3086_CR118","doi-asserted-by":"publisher","first-page":"412","DOI":"10.1038\/nm.4054","volume":"22","author":"A Helman","year":"2016","unstructured":"Helman A, Klochendler A, Azazmeh N, et al. p16(Ink4a)-induced senescence of pancreatic beta cells enhances insulin secretion. Nat Med. 2016;22(4):412\u201320.","journal-title":"Nat Med"},{"issue":"6","key":"3086_CR119","doi-asserted-by":"publisher","first-page":"722","DOI":"10.1016\/j.devcel.2014.11.012","volume":"31","author":"M Demaria","year":"2014","unstructured":"Demaria M, Ohtani N, Youssef SA, et al. An essential role for senescent cells in optimal wound healing through secretion of PDGF-AA. Dev Cell. 2014;31(6):722\u201333.","journal-title":"Dev Cell"},{"issue":"7966","key":"3086_CR120","doi-asserted-by":"publisher","first-page":"808","DOI":"10.1038\/s41586-023-06172-8","volume":"618","author":"X Wang","year":"2023","unstructured":"Wang X, Ramos R, Phan AQ, et al. Signalling by senescent melanocytes hyperactivates hair growth. Nature. 2023;618(7966):808\u201317.","journal-title":"Nature"},{"issue":"8","key":"3086_CR121","doi-asserted-by":"publisher","first-page":"S183","DOI":"10.1016\/j.jid.2024.06.1199","volume":"144","author":"X Wang","year":"2024","unstructured":"Wang X, Ramos R, Almet A, et al. LB1031 Immune Cell-Mediated Amplification of Stem Cell Activation in Hairy Melanocytic Nevus via Osteopontin-CD44 Axis. J Invest Dermatology. 2024;144(8):S183.","journal-title":"J Invest Dermatology"},{"key":"3086_CR122","doi-asserted-by":"publisher","unstructured":"Li K, Liu F, He Y, Qu Q, Sun P, Du L, Wang J, Chen R, Gan Y, Fu D, Fan Z, Liu B, Hu Z, Miao Y. The homing of exogenous hair follicle mesenchymal stem cells into hair follicle niches. JCI Insight. 2023 Dec 22;8(24):e173549. https:\/\/doi.org\/10.1172\/jci.insight.173549","DOI":"10.1172\/jci.insight.173549"},{"issue":"1","key":"3086_CR123","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1038\/s41392-020-00441-y","volume":"6","author":"S Ji","year":"2021","unstructured":"Ji S, Zhu Z, Sun X, et al. Functional hair follicle regeneration: an updated review. Signal Transduct Target Therapy. 2021;6(1):66.","journal-title":"Signal Transduct Target Therapy"},{"key":"3086_CR124","doi-asserted-by":"publisher","first-page":"101303","DOI":"10.1016\/j.mtbio.2024.101303","volume":"29","author":"Y Lv","year":"2024","unstructured":"Lv Y, Yang W, Kannan PR, et al. Materials-based hair follicle engineering: Basic components and recent advances. Mater Today Bio. 2024;29:101303.","journal-title":"Mater Today Bio"},{"issue":"49","key":"3086_CR125","doi-asserted-by":"publisher","first-page":"19679","DOI":"10.1073\/pnas.1309970110","volume":"110","author":"CA Higgins","year":"2013","unstructured":"Higgins CA, Chen JC, Cerise JE, et al. Microenvironmental reprogramming by three-dimensional culture enables dermal papilla cells to induce de novo human hair-follicle growth. Proc Natl Acad Sci U S A. 2013;110(49):19679\u201388.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"30","key":"3086_CR126","doi-asserted-by":"publisher","first-page":"eaba1685","DOI":"10.1126\/sciadv.aba1685","volume":"6","author":"S Hu","year":"2020","unstructured":"Hu S, Li Z, Lutz H, et al. Dermal exosomes containing miR-218-5p promote hair regeneration by regulating \u03b2-catenin signaling. Sci Adv. 2020;6(30):eaba1685.","journal-title":"Sci Adv"},{"issue":"2","key":"3086_CR127","doi-asserted-by":"publisher","first-page":"442","DOI":"10.1016\/j.biomaterials.2012.09.083","volume":"34","author":"Y-C Huang","year":"2013","unstructured":"Huang Y-C, Chan C-C, Lin W-T, et al. Scalable production of controllable dermal papilla spheroids on PVA surfaces and the effects of spheroid size on hair follicle regeneration. Biomaterials. 2013;34(2):442\u201351.","journal-title":"Biomaterials"},{"issue":"7","key":"3086_CR128","doi-asserted-by":"publisher","first-page":"2006","DOI":"10.1021\/acsptsci.4c00114","volume":"7","author":"M Bejaoui","year":"2024","unstructured":"Bejaoui M, Oliva Mizushima AK, Ngoc Linh T, et al. Triethylene Glycol Squalene Improves Hair Regeneration by Maintaining the Inductive Capacity of Human Dermal Papilla Cells and Preventing Premature Aging. ACS Pharmacol Transl Sci. 2024;7(7):2006\u201322.","journal-title":"ACS Pharmacol Transl Sci"},{"key":"3086_CR129","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/j.jare.2020.10.006","volume":"30","author":"CM Abreu","year":"2021","unstructured":"Abreu CM, Cerqueira MT, Pirraco RP, et al. Rescuing key native traits in cultured dermal papilla cells for human hair regeneration. J Adv Res. 2021;30:103\u201312.","journal-title":"J Adv Res"},{"key":"3086_CR130","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.actbio.2022.11.004","volume":"165","author":"Z Liu","year":"2023","unstructured":"Liu Z, Huang J, Kang D, et al. Microenvironmental reprogramming of human dermal papilla cells for hair follicle tissue engineering. Acta Biomater. 2023;165:31\u201349.","journal-title":"Acta Biomater"},{"issue":"1","key":"3086_CR131","doi-asserted-by":"publisher","first-page":"624","DOI":"10.1038\/s41598-020-79722-z","volume":"11","author":"T Kageyama","year":"2021","unstructured":"Kageyama T, Chun Y-S, Fukuda J. Hair follicle germs containing vascular endothelial cells for hair regenerative medicine. Sci Rep. 2021;11(1):624.","journal-title":"Sci Rep"},{"issue":"25","key":"3086_CR132","doi-asserted-by":"publisher","first-page":"11673","DOI":"10.7150\/thno.48723","volume":"10","author":"P Chen","year":"2020","unstructured":"Chen P, Miao Y, Zhang F, et al. Nanoscale microenvironment engineering based on layer-by-layer self-assembly to regulate hair follicle stem cell fate for regenerative medicine. Theranostics. 2020;10(25):11673\u201389.","journal-title":"Theranostics"},{"issue":"14","key":"3086_CR133","doi-asserted-by":"publisher","first-page":"6938","DOI":"10.7150\/thno.107659","volume":"15","author":"M Shrestha","year":"2025","unstructured":"Shrestha M, Nguyen TT, Kausar R, et al. Enhancing hair regrowth using rapamycin-primed mesenchymal stem cell-derived exosomes. Theranostics. 2025;15(14):6938\u201356.","journal-title":"Theranostics"},{"issue":"4","key":"3086_CR134","doi-asserted-by":"publisher","first-page":"4354","DOI":"10.1021\/acsnano.8b09573","volume":"13","author":"G Yang","year":"2019","unstructured":"Yang G, Chen Q, Wen D, et al. A Therapeutic Microneedle Patch Made from Hair-Derived Keratin for Promoting Hair Regrowth. ACS Nano. 2019;13(4):4354\u201360.","journal-title":"ACS Nano"},{"issue":"3","key":"3086_CR135","doi-asserted-by":"publisher","first-page":"1454","DOI":"10.7150\/thno.39566","volume":"10","author":"Y Chen","year":"2020","unstructured":"Chen Y, Huang J, Chen R, et al. Sustained release of dermal papilla-derived extracellular vesicles from injectable microgel promotes hair growth. Theranostics. 2020;10(3):1454\u201378.","journal-title":"Theranostics"},{"key":"3086_CR136","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.actbio.2022.03.011","volume":"165","author":"D Kang","year":"2023","unstructured":"Kang D, Liu Z, Qian C, et al. 3D bioprinting of a gelatin-alginate hydrogel for tissue-engineered hair follicle regeneration. Acta Biomater. 2023;165:19\u201330.","journal-title":"Acta Biomater"},{"key":"3086_CR137","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1016\/j.actbio.2022.06.021","volume":"165","author":"A Nanmo","year":"2023","unstructured":"Nanmo A, Yan L, Asaba T, et al. Bioprinting of hair follicle germs for hair regenerative medicine. Acta Biomater. 2023;165:50\u20139.","journal-title":"Acta Biomater"},{"key":"3086_CR138","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1016\/j.mad.2017.02.009","volume":"163","author":"A Chatterjee","year":"2017","unstructured":"Chatterjee A, Georgiev G, Iannacchione G. Aging and efficiency in living systems: Complexity, adaptation and self-organization. Mech Ageing Dev. 2017;163:2\u20137.","journal-title":"Mech Ageing Dev"},{"key":"3086_CR139","doi-asserted-by":"publisher","unstructured":"Bera S, Loeffler D. Cell polarity: cell type-specific regulators, common pathways, and polarized vesicle transport. Leukemia. 2025 Jul;39(7):1558-1570. https:\/\/doi.org\/10.1038\/s41375-025-02601-x","DOI":"10.1038\/s41375-025-02601-x"},{"issue":"1","key":"3086_CR140","doi-asserted-by":"publisher","first-page":"10250","DOI":"10.1038\/s41467-024-54638-8","volume":"15","author":"T Schuster","year":"2024","unstructured":"Schuster T, Amoah A, Vollmer A, et al. Quantitative determination of the spatial distribution of components in single cells with CellDetail. Nat Commun. 2024;15(1):10250.","journal-title":"Nat Commun"},{"issue":"1","key":"3086_CR141","first-page":"33","volume":"184","author":"C L\u00f3pez-Ot\u00edn","year":"2021","unstructured":"L\u00f3pez-Ot\u00edn C, Kroemer G. Hallm Health Cell. 2021;184(1):33\u201363.","journal-title":"Hallm Health Cell"},{"key":"3086_CR142","doi-asserted-by":"publisher","unstructured":"Restrepo P, Wilder A, Houser A, Sandhu HS, Ramirez A, Grace Hren M, Gill R, Kazmi A, Chen L, Nigro A, Imanishi I, Demircioglu D, Hasson D, Soto A, McQuillan S, Gonzalez-Kozlova E, Brody R, Ungar B, Kasper M, Lu CP, Torina P, Lewin JM, Gnjatic S, Ma S, Ji AL. Single-cell spatial transcriptomic analysis of human skin anatomy. Nat Genet. 2026 Apr;58(4):903-915. https:\/\/doi.org\/10.1038\/s41588-026-02552-8","DOI":"10.1038\/s41588-026-02552-8"},{"key":"3086_CR143","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1016\/j.ydbio.2024.07.012","volume":"516","author":"H Fujiwara","year":"2024","unstructured":"Fujiwara H. Dynamic duo: Cell-extracellular matrix interactions in hair follicle development and regeneration. Dev Biol. 2024;516:20\u201334.","journal-title":"Dev Biol"},{"key":"3086_CR144","doi-asserted-by":"publisher","unstructured":"Wu S, Yu Y, Liu C, Zhang X, Zhu P, Peng Y, Yan X, Li Y, Hua P, Li Q, Wang S, Zhang L. Single-cell transcriptomics reveals lineage trajectory of human scalp hair follicle and informs mechanisms of hair graying. Cell Discov. 2022 May 24;8(1):49. https:\/\/doi.org\/10.1038\/s41421-022-00394-2","DOI":"10.1038\/s41421-022-00394-2"},{"key":"3086_CR145","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1007\/7651_2019_227","volume":"2150","author":"YF Wu","year":"2020","unstructured":"Wu YF, Tan HY, Lin SJ. Long-Term Intravital Imaging of the Cornea, Skin, and Hair Follicle by Multiphoton Microscope. Methods Mol Biol. 2020;2150:131\u201340.","journal-title":"Methods Mol Biol"},{"issue":"1","key":"3086_CR146","doi-asserted-by":"publisher","first-page":"418","DOI":"10.1186\/s13287-025-04538-5","volume":"16","author":"A Yu","year":"2025","unstructured":"Yu A, Zhang Y, Zhong S, et al. Human umbilical cord mesenchymal stem cell-derived exosomes enhance follicular regeneration in androgenetic alopecia via activation of Wnt\/\u03b2-catenin pathway. Stem Cell Res Ther. 2025;16(1):418.","journal-title":"Stem Cell Res Ther"},{"issue":"5","key":"3086_CR147","doi-asserted-by":"publisher","first-page":"e70080","DOI":"10.1002\/jev2.70080","volume":"14","author":"Z Cao","year":"2025","unstructured":"Cao Z, Li P, Zhang M, et al. Erythrocyte Extracellular Vesicles Amalgamate into the Hair and Skin to Maintain Homeostasis. J Extracell Vesicles. 2025;14(5):e70080.","journal-title":"J Extracell Vesicles"},{"issue":"1","key":"3086_CR148","doi-asserted-by":"publisher","first-page":"754","DOI":"10.1186\/s12951-025-03865-2","volume":"23","author":"Y Dong","year":"2025","unstructured":"Dong Y, Sun Y, Li A, et al. Rejuvenating senescent hair follicles: a novel conjugated linoleic acid based nanovesicle approach to treat androgenic alopecia. J Nanobiotechnol. 2025;23(1):754.","journal-title":"J Nanobiotechnol"},{"key":"3086_CR149","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1016\/j.actbio.2025.05.039","volume":"200","author":"Z Xing","year":"2025","unstructured":"Xing Z, Zhang X, Zhao C, et al. Microenvironment-responsive recombinant collagen XVII-based composite microneedles for the treatment of androgenetic alopecia. Acta Biomater. 2025;200:400\u201315.","journal-title":"Acta Biomater"},{"key":"3086_CR150","doi-asserted-by":"publisher","first-page":"636","DOI":"10.1016\/j.jconrel.2025.01.034","volume":"379","author":"Y Chen","year":"2025","unstructured":"Chen Y, Ren T, Wu W, et al. Gas-propelled anti-hair follicle aging microneedle patch for the treatment of androgenetic alopecia. J Control Release. 2025;379:636\u201351.","journal-title":"J Control Release"},{"issue":"3","key":"3086_CR151","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1038\/s43587-022-00357-y","volume":"3","author":"R Waziry","year":"2023","unstructured":"Waziry R, Ryan CP, Corcoran DL, et al. Effect of long-term caloric restriction on DNA methylation measures of biological aging in healthy adults from the CALERIE trial. Nat Aging. 2023;3(3):248\u201357.","journal-title":"Nat Aging"},{"issue":"11","key":"3086_CR152","doi-asserted-by":"publisher","first-page":"2678","DOI":"10.1016\/j.celrep.2017.08.052","volume":"20","author":"MF Forni","year":"2017","unstructured":"Forni MF, Peloggia J, Braga TT, et al. Caloric Restriction Promotes Structural and Metabolic Changes in the Skin. Cell Rep. 2017;20(11):2678\u201392.","journal-title":"Cell Rep"},{"key":"3086_CR153","doi-asserted-by":"publisher","first-page":"102158","DOI":"10.1016\/j.arr.2023.102158","volume":"93","author":"Y Chen","year":"2024","unstructured":"Chen Y, Yang S, Yu K, et al. Spatial omics: An innovative frontier in aging research. Ageing Res Rev. 2024;93:102158.","journal-title":"Ageing Res Rev"},{"issue":"7472","key":"3086_CR154","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1038\/nature12602","volume":"502","author":"P Rompolas","year":"2013","unstructured":"Rompolas P, Mesa KR, Greco V. Spatial organization within a niche as a determinant of stem-cell fate. Nature. 2013;502(7472):513\u20138.","journal-title":"Nature"},{"issue":"7408","key":"3086_CR155","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1038\/nature11218","volume":"487","author":"P Rompolas","year":"2012","unstructured":"Rompolas P, Deschene ER, Zito G, et al. Live imaging of stem cell and progeny behaviour in physiological hair-follicle regeneration. Nature. 2012;487(7408):496\u20139.","journal-title":"Nature"},{"issue":"3","key":"3086_CR156","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1016\/j.cels.2016.08.010","volume":"3","author":"S Joost","year":"2016","unstructured":"Joost S, Zeisel A, Jacob T, et al. Single-Cell Transcriptomics Reveals that Differentiation and Spatial Signatures Shape Epidermal and Hair Follicle Heterogeneity. Cell Syst. 2016;3(3):221\u2013e2379.","journal-title":"Cell Syst"}],"container-title":["Cell Communication and Signaling"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12964-026-03086-6","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12964-026-03086-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12964-026-03086-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T10:05:38Z","timestamp":1789034738000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12964-026-03086-6"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,27]]},"references-count":156,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["3086"],"URL":"https:\/\/doi.org\/10.1186\/s12964-026-03086-6","relation":{},"ISSN":["1478-811X"],"issn-type":[{"value":"1478-811X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,27]]},"assertion":[{"value":"23 January 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 July 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 July 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"486"}}