{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T13:18:42Z","timestamp":1785417522849,"version":"3.56.0"},"reference-count":76,"publisher":"Rockefeller University Press","issue":"3","license":[{"start":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T00:00:00Z","timestamp":1767916800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["F99 CA274517"],"award-info":[{"award-number":["F99 CA274517"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["F31 AR038249"],"award-info":[{"award-number":["F31 AR038249"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["T32 AR007197"],"award-info":[{"award-number":["T32 AR007197"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01 AR079418"],"award-info":[{"award-number":["R01 AR079418"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100003185","name":"National Psoriasis Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100003185","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["rupress.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2026,3,2]]},"abstract":"<jats:p>The type I intermediate filament proteins keratin 14 (K14) and keratin 15 (K15) are common to all complex epithelia. K14 is highly expressed by progenitor keratinocytes, in which it provides essential mechanical integrity and gates keratinocyte entry into differentiation by sequestering YAP1, a transcriptional effector of Hippo signaling, to the cytoplasm. K15 has long been used as a marker of hair bulge stem cells, though its specific role in skin epithelia is unknown. Here, we show that the lack of two biochemical determinants, a cysteine residue within the stutter motif of the central rod domain and a 14-3-3 binding site in the N-terminal head domain, renders K15 unable to effectively sequester YAP1 in the cytoplasm like K14 does. We combine insight obtained from cell culture and transgenic mouse models with computational analyses of transcriptomics data and propose a model in which a higher K15:K14 ratio promotes a progenitor state and antagonizes differentiation in keratinocytes of the epidermis.<\/jats:p>","DOI":"10.1083\/jcb.202503046","type":"journal-article","created":{"date-parts":[[2025,12,5]],"date-time":"2025-12-05T07:21:14Z","timestamp":1764919274000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":4,"title":["Keratin 15 promotes a progenitor cell state in basal keratinocytes of skin epidermis"],"prefix":"10.1083","volume":"225","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9474-2803","authenticated-orcid":true,"given":"Catherine J.","family":"Redmond","sequence":"first","affiliation":[{"name":"University of Michigan Medical School 1 Department of Cell & Developmental Biology, , Ann Arbor, MI,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1269-3900","authenticated-orcid":true,"given":"Sarah N.","family":"Steiner","sequence":"additional","affiliation":[{"name":"University of Michigan Medical School 1 Department of Cell & Developmental Biology, , Ann Arbor, MI,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2390-3707","authenticated-orcid":true,"given":"Erez","family":"Cohen","sequence":"additional","affiliation":[{"name":"University of Michigan Medical School 1 Department of Cell & Developmental Biology, , Ann Arbor, MI,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3955-7394","authenticated-orcid":true,"given":"Craig N.","family":"Johnson","sequence":"additional","affiliation":[{"name":"University of Michigan Medical School 1 Department of Cell & Developmental Biology, , Ann Arbor, MI,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5157-8780","authenticated-orcid":true,"given":"Nurhan","family":"\u00d6zl\u00fc","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/00jzwgz36","id-type":"ROR","asserted-by":"publisher"}],"name":"Ko\u00e7 University 2 Department of Molecular Biology and Genetics, Science Faculty, , Istanbul,","place":["Turkey"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0680-2373","authenticated-orcid":true,"given":"Pierre A.","family":"Coulombe","sequence":"additional","affiliation":[{"name":"University of Michigan Medical School 1 Department of Cell & Developmental Biology, , Ann Arbor, MI,","place":["USA"]},{"name":"University of Michigan Medical School 3 Department of Dermatology, , Ann Arbor, MI,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/00jmfr291","id-type":"ROR","asserted-by":"publisher"}],"name":"Rogel Cancer Center, University of Michigan 4 , Ann Arbor, MI,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"291","published-online":{"date-parts":[[2026,1,9]]},"reference":[{"key":"2026010907260363100_bib1","doi-asserted-by":"publisher","first-page":"284","DOI":"10.1016\/j.jid.2019.06.137","article-title":"Mechanical forces in the skin: Roles in tissue architecture, stability, and function","volume":"140","author":"Biggs","year":"2020","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib2","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1038\/nrm2636","article-title":"Epidermal homeostasis: A balancing act of stem cells in the skin","volume":"10","author":"Blanpain","year":"2009","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026010907260363100_bib3","doi-asserted-by":"publisher","first-page":"1202","DOI":"10.1126\/science.1720261","article-title":"Epidermolysis bullosa simplex: Evidence in two families for keratin gene abnormalities","volume":"254","author":"Bonifas","year":"1991","journal-title":"Science"},{"key":"2026010907260363100_bib4","doi-asserted-by":"publisher","first-page":"19385","DOI":"10.3390\/ijms141019385","article-title":"Keratin 15 as a biomarker of epidermal stem cells","volume":"14","author":"Bose","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"2026010907260363100_bib5","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1016\/j.jid.2016.08.018","article-title":"The X-ray crystal structure of the keratin 1-keratin 10 helix 2B heterodimer reveals molecular surface properties and biochemical insights into human skin disease","volume":"137","author":"Bunick","year":"2017","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib6","doi-asserted-by":"publisher","first-page":"2574","DOI":"10.1101\/gad.8.21.2574","article-title":"A human keratin 14 \u201cknockout\u201d: The absence of K14 leads to severe epidermolysis bullosa simplex and a function for an intermediate filament protein","volume":"8","author":"Chan","year":"1994","journal-title":"Genes Dev."},{"key":"2026010907260363100_bib7","doi-asserted-by":"publisher","first-page":"871","DOI":"10.1016\/j.celrep.2018.09.006","article-title":"Transcriptional programming of normal and inflamed human epidermis at single-cell resolution","volume":"25","author":"Cheng","year":"2018","journal-title":"Cell Rep."},{"key":"2026010907260363100_bib8","doi-asserted-by":"publisher","first-page":"1692","DOI":"10.1038\/s41556-022-01021-8","article-title":"Gradual differentiation uncoupled from cell cycle exit generates heterogeneity in the epidermal stem cell layer","volume":"24","author":"Cockburn","year":"2022","journal-title":"Nat. Cell Biol."},{"key":"2026010907260363100_bib9","doi-asserted-by":"publisher","first-page":"jcs260594","DOI":"10.1242\/jcs.260594","article-title":"Revisiting the significance of keratin expression in complex epithelia","volume":"135","author":"Cohen","year":"2022","journal-title":"J. Cell Sci."},{"key":"2026010907260363100_bib10","doi-asserted-by":"publisher","first-page":"108805","DOI":"10.1016\/j.isci.2024.108805","article-title":"Significance of stress keratin expression in normal and diseased epithelia","volume":"27","author":"Cohen","year":"2024","journal-title":"iScience"},{"key":"2026010907260363100_bib11","doi-asserted-by":"publisher","first-page":"1301","DOI":"10.1016\/0092-8674(91)90051-y","article-title":"Point mutations in human keratin 14 genes of epidermolysis bullosa simplex patients: Genetic and functional analyses","volume":"66","author":"Coulombe","year":"1991","journal-title":"Cell"},{"key":"2026010907260363100_bib12","doi-asserted-by":"publisher","first-page":"1784","DOI":"10.1172\/JCI38177","article-title":"Epidermolysis bullosa simplex: A paradigm for disorders of tissue fragility","volume":"119","author":"Coulombe","year":"2009","journal-title":"J. Clin. Invest."},{"key":"2026010907260363100_bib13","doi-asserted-by":"publisher","first-page":"2295","DOI":"10.1083\/jcb.109.5.2295","article-title":"Expression of keratin K14 in the epidermis and hair follicle: Insights into complex programs of differentiation","volume":"109","author":"Coulombe","year":"1989","journal-title":"J. Cell Biol."},{"key":"2026010907260363100_bib14","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1038\/jid.2011.450","article-title":"Defining keratin protein function in skin epithelia: Epidermolysis bullosa simplex and its aftermath","volume":"132","author":"Coulombe","year":"2012","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib15","doi-asserted-by":"publisher","first-page":"2079","DOI":"10.1016\/j.jid.2022.01.007","article-title":"KLHL24-mediated hair follicle stem cells structural disruption causes alopecia","volume":"142","author":"Cui","year":"2022","journal-title":"J. Invest Dermatol."},{"key":"2026010907260363100_bib16","doi-asserted-by":"publisher","first-page":"22507","DOI":"10.1074\/jbc.M115.654749","article-title":"Complementary roles of specific cysteines in keratin 14 toward the assembly, organization, and dynamics of intermediate filaments in skin keratinocytes","volume":"290","author":"Feng","year":"2015","journal-title":"J. Biol. Chem."},{"key":"2026010907260363100_bib17","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1083\/jcb.201408079","article-title":"A role for disulfide bonding in keratin intermediate filament organization and dynamics in skin keratinocytes","volume":"209","author":"Feng","year":"2015","journal-title":"J. Cell Biol."},{"key":"2026010907260363100_bib18","doi-asserted-by":"publisher","first-page":"850","DOI":"10.1038\/jid.2012.397","article-title":"Keratin intracellular concentration revisited: Implications for keratin function in surface epithelia","volume":"133","author":"Feng","year":"2013","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib19","doi-asserted-by":"publisher","first-page":"baz046","DOI":"10.1093\/database\/baz046","article-title":"PanglaoDB: A web server for exploration of mouse and human single-cell RNA sequencing data","volume":"2019","author":"Franzen","year":"2019","journal-title":"Database"},{"key":"2026010907260363100_bib20","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1146\/annurev.cb.11.110195.001011","article-title":"Keratins and the skin","volume":"11","author":"Fuchs","year":"1995","journal-title":"Ann. Rev. Cell Dev. Biol."},{"key":"2026010907260363100_bib21","doi-asserted-by":"publisher","first-page":"1033","DOI":"10.1016\/0092-8674(80)90094-x","article-title":"Changes in keratin gene expression during terminal differentiation of the keratinocyte","volume":"19","author":"Fuchs","year":"1980","journal-title":"Cell"},{"key":"2026010907260363100_bib22","doi-asserted-by":"publisher","first-page":"979","DOI":"10.12688\/f1000research.73600.2","article-title":"Doublet identification in single-cell sequencing data using scDblFinder","volume":"10","author":"Germain","year":"2021","journal-title":"F1000Res."},{"key":"2026010907260363100_bib23","doi-asserted-by":"publisher","DOI":"10.15252\/embj.2021110488","article-title":"Binary organization of epidermal basal domains highlights robustness to environmental exposure","volume":"41","author":"Ghuwalewala","year":"2022","journal-title":"EMBO J."},{"key":"2026010907260363100_bib24","doi-asserted-by":"publisher","first-page":"2378","DOI":"10.1172\/JCI88941","article-title":"Long-lived keratin 15+ esophageal progenitor cells contribute to homeostasis and regeneration","volume":"127","author":"Giroux","year":"2017","journal-title":"J. Clin. Invest."},{"key":"2026010907260363100_bib25","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.53165","article-title":"Keratin 14-dependent disulfides regulate epidermal homeostasis and barrier function via 14-3-3\u03c3 and YAP1","volume":"9","author":"Guo","year":"2020","journal-title":"Elife"},{"key":"2026010907260363100_bib26","doi-asserted-by":"publisher","DOI":"10.1172\/jci.insight.162041","article-title":"Krt14 and Krt15 differentially regulate regenerative properties and differentiation potential of airway basal cells","volume":"8","author":"Ievlev","year":"2023","journal-title":"JCI Insight"},{"key":"2026010907260363100_bib27","doi-asserted-by":"publisher","first-page":"844","DOI":"10.1038\/labinvest.2009.48","article-title":"Differential expression of stem-cell-associated markers in human hair follicle epithelial cells","volume":"89","author":"Inoue","year":"2009","journal-title":"Lab. Invest."},{"key":"2026010907260363100_bib28","doi-asserted-by":"publisher","first-page":"1291","DOI":"10.1038\/ng.239","article-title":"Lgr5 marks cycling, yet long-lived, hair follicle stem cells","volume":"40","author":"Jaks","year":"2008","journal-title":"Nat. Genet."},{"key":"2026010907260363100_bib29","doi-asserted-by":"publisher","first-page":"764","DOI":"10.1111\/1523-1747.ep12365805","article-title":"Effects of keratin 14 ablation on the clinical and cellular phenotype in a kindred with recessive epidermolysis bullosa simplex","volume":"107","author":"Jonkman","year":"1996","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib30","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1038\/s41586-020-2308-7","article-title":"The mutational constraint spectrum quantified from variation in 141,456 humans","volume":"581","author":"Karczewski","year":"2020","journal-title":"Nature"},{"key":"2026010907260363100_bib31","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1038\/nature04659","article-title":"A keratin cytoskeletal protein regulates protein synthesis and epithelial cell growth","volume":"441","author":"Kim","year":"2006","journal-title":"Nature"},{"key":"2026010907260363100_bib32","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1038\/nmeth.4256","article-title":"MSFragger: Ultrafast and comprehensive peptide identification in mass spectrometry-based proteomics","volume":"14","author":"Kong","year":"2017","journal-title":"Nat. Methods"},{"key":"2026010907260363100_bib33","doi-asserted-by":"publisher","first-page":"1892","DOI":"10.1093\/emboj\/17.7.1892","article-title":"Phosphorylation of human keratin 18 serine 33 regulates binding to 14-3- 3 proteins","volume":"17","author":"Ku","year":"1998","journal-title":"EMBO J."},{"key":"2026010907260363100_bib34","doi-asserted-by":"publisher","first-page":"W90","DOI":"10.1093\/nar\/gkw377","article-title":"Enrichr: A comprehensive gene set enrichment analysis web server 2016 update","volume":"44","author":"Kuleshov","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"2026010907260363100_bib35","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1083\/jcb.200810196","article-title":"Self-organization of keratin intermediate filaments into cross-linked networks","volume":"186","author":"Lee","year":"2009","journal-title":"J. Cell Biol."},{"key":"2026010907260363100_bib36","doi-asserted-by":"publisher","first-page":"707","DOI":"10.1038\/nsmb.2330","article-title":"Structural basis for heteromeric assembly and perinuclear organization of keratin filaments","volume":"19","author":"Lee","year":"2012","journal-title":"Nat. Struct. Mol. Biol."},{"key":"2026010907260363100_bib37","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1016\/j.jaci.2022.03.002","article-title":"Single-cell transcriptome profile of mouse skin undergoing antigen-driven allergic inflammation recapitulates findings in atopic dermatitis skin lesions","volume":"150","author":"Leyva-Castillo","year":"2022","journal-title":"J. Allergy Clin. Immunol."},{"key":"2026010907260363100_bib38","doi-asserted-by":"publisher","first-page":"102264","DOI":"10.1016\/j.ceb.2023.102264","article-title":"Posttranslational modifications of keratins and their associated proteins as therapeutic targets in keratin diseases","volume":"85","author":"Li","year":"2023","journal-title":"Curr. Opin. Cell Biol."},{"key":"2026010907260363100_bib39","doi-asserted-by":"publisher","first-page":"5434","DOI":"10.1038\/s41467-020-19234-6","article-title":"Murine interfollicular epidermal differentiation is gradualistic with GRHL3 controlling progression from stem to transition cell states","volume":"11","author":"Lin","year":"2020","journal-title":"Nat. Commun."},{"key":"2026010907260363100_bib40","doi-asserted-by":"publisher","first-page":"963","DOI":"10.1046\/j.1523-1747.2003.12600.x","article-title":"Keratin 15 promoter targets putative epithelial stem cells in the hair follicle bulge","volume":"121","author":"Liu","year":"2003","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib41","doi-asserted-by":"publisher","first-page":"1329","DOI":"10.1083\/jcb.129.5.1329","article-title":"The basal keratin network of stratified squamous epithelia: Defining K15 function in the absence of K14","volume":"129","author":"Lloyd","year":"1995","journal-title":"J. Cell Biol."},{"key":"2026010907260363100_bib42","doi-asserted-by":"publisher","first-page":"3179","DOI":"10.1242\/jcs.111.21.3179","article-title":"The C8\/144B monoclonal antibody recognizes cytokeratin 15 and defines the location of human hair follicle stem cells","volume":"111","author":"Lyle","year":"1998","journal-title":"J. Cell Sci."},{"key":"2026010907260363100_bib43","doi-asserted-by":"publisher","first-page":"17046","DOI":"10.1074\/jbc.M117.805655","article-title":"Hypoxia-inducible factor 2alpha (HIF-2alpha) promotes colon cancer growth by potentiating yes-associated protein 1 (YAP1) activity","volume":"292","author":"Ma","year":"2017","journal-title":"J. Biol. Chem."},{"key":"2026010907260363100_bib44","doi-asserted-by":"publisher","first-page":"2276","DOI":"10.1093\/bioinformatics\/btv133","article-title":"14-3-3-Pred: Improved methods to predict 14-3-3-binding phosphopeptides","volume":"31","author":"Madeira","year":"2015","journal-title":"Bioinformatics"},{"key":"2026010907260363100_bib45","doi-asserted-by":"publisher","first-page":"930","DOI":"10.1091\/mbc.E18-06-0373","article-title":"14-3-3 targets keratin intermediate filaments to mechanically sensitive cell-cell contacts","volume":"31","author":"Mariani","year":"2020","journal-title":"Mol. Biol. Cell"},{"key":"2026010907260363100_bib46","doi-asserted-by":"publisher","first-page":"1017","DOI":"10.1242\/jcs.109.5.1017","article-title":"Keratin 19 as a biochemical marker of skin stem cells in vivo and in vitro: Keratin 19 expressing cells are differentially localized in function of anatomic sites, and their number varies with donor age and culture stage","volume":"109","author":"Michel","year":"1996","journal-title":"J. Cell Sci."},{"key":"2026010907260363100_bib47","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1038\/s41556-017-0005-z","article-title":"Adhesion forces and cortical tension couple cell proliferation and differentiation to drive epidermal stratification","volume":"20","author":"Miroshnikova","year":"2018","journal-title":"Nat. Cell Biol."},{"key":"2026010907260363100_bib48","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1038\/nbt950","article-title":"Capturing and profiling adult hair follicle stem cells","volume":"22","author":"Morris","year":"2004","journal-title":"Nat. Biotechnol."},{"key":"2026010907260363100_bib49","doi-asserted-by":"publisher","first-page":"1053","DOI":"10.1038\/s41467-018-03414-6","article-title":"Desmosomal cadherin association with Tctex-1 and cortactin-Arp2\/3 drives perijunctional actin polymerization to promote keratinocyte delamination","volume":"9","author":"Nekrasova","year":"2018","journal-title":"Nat. Commun."},{"key":"2026010907260363100_bib50","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1016\/j.stem.2020.11.002","article-title":"Differentiated daughter cells regulate stem cell proliferation and fate through intra-tissue tension","volume":"28","author":"Ning","year":"2021","journal-title":"Cell Stem Cell"},{"key":"2026010907260363100_bib51","doi-asserted-by":"publisher","first-page":"5587","DOI":"10.1038\/s41388-018-0348-3","article-title":"The dynamic and stress-adaptive signaling hub of 14-3-3: Emerging mechanisms of regulation and context-dependent protein-protein interactions","volume":"37","author":"Pennington","year":"2018","journal-title":"Oncogene"},{"key":"2026010907260363100_bib52","doi-asserted-by":"publisher","first-page":"1775","DOI":"10.1091\/mbc.12.6.1775","article-title":"Complete cytolysis and neonatal lethality in keratin 5 knockout mice reveal its fundamental role in skin integrity and in epidermolysis bullosa simplex","volume":"12","author":"Peters","year":"2001","journal-title":"Mol. Biol. Cell"},{"key":"2026010907260363100_bib53","doi-asserted-by":"publisher","first-page":"jcs230425","DOI":"10.1242\/jcs.230425","article-title":"YAP\/TAZ functions and their regulation at a glance","volume":"133","author":"Pocaterra","year":"2020","journal-title":"J. Cell Sci."},{"key":"2026010907260363100_bib54","doi-asserted-by":"publisher","first-page":"1701","DOI":"10.1038\/labinvest.3780180","article-title":"K15 expression implies lateral differentiation within stratified epithelial basal cells","volume":"80","author":"Porter","year":"2000","journal-title":"Lab. Invest."},{"key":"2026010907260363100_bib55","doi-asserted-by":"publisher","first-page":"18513","DOI":"10.1073\/pnas.1313491110","article-title":"Keratins as the main component for the mechanical integrity of keratinocytes","volume":"110","author":"Ramms","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026010907260363100_bib56","doi-asserted-by":"publisher","first-page":"2563","DOI":"10.1101\/gad.8.21.2563","article-title":"A functional \u201cknockout\u201d of human keratin 14","volume":"8","author":"Rugg","year":"1994","journal-title":"Genes Dev."},{"key":"2026010907260363100_bib57","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1038\/ncb3359","article-title":"Defining the cellular lineage hierarchy in the interfollicular epidermis of adult skin","volume":"18","author":"Sada","year":"2016","journal-title":"Nat. Cell Biol."},{"key":"2026010907260363100_bib58","doi-asserted-by":"publisher","first-page":"1621","DOI":"10.1038\/jid.2015.42","article-title":"14-3-3sigma regulates keratinocyte proliferation and differentiation by modulating Yap1 cellular localization","volume":"135","author":"Sambandam","year":"2015","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib59","doi-asserted-by":"publisher","first-page":"782","DOI":"10.1016\/j.cell.2011.02.031","article-title":"Yap1 acts downstream of alpha-catenin to control epidermal proliferation","volume":"144","author":"Schlegelmilch","year":"2011","journal-title":"Cell"},{"key":"2026010907260363100_bib60","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1083\/jcb.200603161","article-title":"New consensus nomenclature for mammalian keratins","volume":"174","author":"Schweizer","year":"2006","journal-title":"J. Cell Biol."},{"key":"2026010907260363100_bib61","doi-asserted-by":"publisher","DOI":"10.1101\/2025.09.15.676359","article-title":"Under pressure: A unique mechanoresponsive mechanism of body site-specific keratin regulation in palmoplantar epidermis","author":"Steiner","year":"2025","journal-title":"bioRxiv"},{"key":"2026010907260363100_bib62","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1083\/jcb.107.2.427","article-title":"Use of monospecific antisera and cRNA probes to localize the major changes in keratin expression during normal and abnormal epidermal differentiation","volume":"107","author":"Stoler","year":"1988","journal-title":"J. Cell Biol."},{"key":"2026010907260363100_bib63","doi-asserted-by":"publisher","first-page":"2225","DOI":"10.1101\/gad.267708.115","article-title":"CALML5 is a ZNF750- and TINCR-induced protein that binds stratifin to regulate epidermal differentiation","volume":"29","author":"Sun","year":"2015","journal-title":"Genes Dev."},{"key":"2026010907260363100_bib64","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1002\/humu.20652","article-title":"The human intermediate filament database: Comprehensive information on a gene family involved in many human diseases","volume":"29","author":"Szeverenyi","year":"2008","journal-title":"Hum. Mutat."},{"key":"2026010907260363100_bib65","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1038\/s41556-018-0142-z","article-title":"YAP\/TAZ upstream signals and downstream responses","volume":"20","author":"Totaro","year":"2018","journal-title":"Nat. Cell Biol."},{"key":"2026010907260363100_bib66","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1046\/j.1523-1747.2003.12088.x","article-title":"Enrichment for living murine keratinocytes from the hair follicle bulge with the cell surface marker CD34","volume":"120","author":"Trempus","year":"2003","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib67","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1126\/science.1092436","article-title":"Defining the epithelial stem cell niche in skin","volume":"303","author":"Tumbar","year":"2004","journal-title":"Science"},{"key":"2026010907260363100_bib68","doi-asserted-by":"publisher","first-page":"29772","DOI":"10.1074\/jbc.M001207200","article-title":"Calyculin A-induced vimentin phosphorylation sequesters 14-3-3 and displaces other 14-3-3 partners in vivo","volume":"275","author":"Tzivion","year":"2000","journal-title":"J. Biol. Chem."},{"key":"2026010907260363100_bib69","doi-asserted-by":"publisher","first-page":"3165","DOI":"10.3390\/ijms26073165","article-title":"Generation of the Krt24-Cre(ERT2) mouse line targeting outer bulge hair follicle cells","volume":"26","author":"Wang","year":"2025","journal-title":"Int. J. Mol. Sci."},{"key":"2026010907260363100_bib70","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1046\/j.1523-1747.1999.00535.x","article-title":"Keratin 15 expression in stratified epithelia: Downregulation in activated keratinocytes","volume":"112","author":"Waseem","year":"1999","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib71","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1111\/j.1432-0436.2004.07208005.x","article-title":"Location and phenotype of human adult keratinocyte stem cells of the skin","volume":"72","author":"Webb","year":"2004","journal-title":"Differentiation"},{"key":"2026010907260363100_bib72","doi-asserted-by":"publisher","first-page":"322","DOI":"10.1038\/s41586-018-0073-7","article-title":"Diverse mechanisms for endogenous regeneration and repair in mammalian organs","volume":"557","author":"Wells","year":"2018","journal-title":"Nature"},{"key":"2026010907260363100_bib73","doi-asserted-by":"publisher","first-page":"448","DOI":"10.1006\/excr.1998.4217","article-title":"Expression of the intermediate filament keratin gene, K15, in the basal cell layers of epithelia and the hair follicle","volume":"244","author":"Whitbread","year":"1998","journal-title":"Exp. Cell Res."},{"key":"2026010907260363100_bib74","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/j.jid.2021.06.020","article-title":"Oncogenic hedgehog-smoothened signaling depends on YAP1\u2012TAZ\/TEAD transcription to restrain differentiation in basal cell carcinoma","volume":"142","author":"Yuan","year":"2022","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib75","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1038\/sj.jid.5700583","article-title":"Cytokeratin15-positive basal epithelial cells targeted in graft-versus-host disease express a constitutive antiapoptotic phenotype","volume":"127","author":"Zhan","year":"2007","journal-title":"J. Invest. Dermatol."},{"key":"2026010907260363100_bib76","doi-asserted-by":"publisher","first-page":"2270","DOI":"10.1073\/pnas.1019603108","article-title":"Yes-associated protein (YAP) transcriptional coactivator functions in balancing growth and differentiation in skin","volume":"108","author":"Zhang","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"}],"container-title":["Journal of Cell Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/rupress.org\/jcb\/article-pdf\/doi\/10.1083\/jcb.202503046\/2023910\/jcb_202503046.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/rupress.org\/jcb\/article-pdf\/doi\/10.1083\/jcb.202503046\/2023910\/jcb_202503046.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T12:26:22Z","timestamp":1767961582000},"score":1,"resource":{"primary":{"URL":"https:\/\/rupress.org\/jcb\/article\/225\/3\/e202503046\/281370\/Keratin-15-promotes-a-progenitor-cell-state-in"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,9]]},"references-count":76,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2026,3,2]]}},"URL":"https:\/\/doi.org\/10.1083\/jcb.202503046","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.02.27.640633","asserted-by":"object"}]},"ISSN":["0021-9525","1540-8140"],"issn-type":[{"value":"0021-9525","type":"print"},{"value":"1540-8140","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,9]]},"article-number":"e202503046"}}