{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:02:11Z","timestamp":1792036931908,"version":"4.4.0"},"reference-count":305,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2020,11,27]],"date-time":"2020-11-27T00:00:00Z","timestamp":1606435200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Joint CNRS \/ Ales Group grant","award":["171416"],"award-info":[{"award-number":["171416"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>One of the most important functions of skin is to act as a protective barrier. To fulfill this role, the structural integrity of the skin depends on the dermal-epidermal junction\u2014a complex network of extracellular matrix macromolecules that connect the outer epidermal layer to the underlying dermis. This junction provides both a structural support to keratinocytes and a specific niche that mediates signals influencing their behavior. It displays a distinctive microarchitecture characterized by an undulating pattern, strengthening dermal-epidermal connectivity and crosstalk. The optimal stiffness arising from the overall molecular organization, together with characteristic anchoring complexes, keeps the dermis and epidermis layers extremely well connected and capable of proper epidermal renewal and regeneration. Due to intrinsic and extrinsic factors, a large number of structural and biological changes accompany skin aging. These changes progressively weaken the dermal\u2013epidermal junction substructure and affect its functions, contributing to the gradual decline in overall skin physiology. Most changes involve reduced turnover or altered enzymatic or non-enzymatic post-translational modifications, compromising the mechanical properties of matrix components and cells. This review combines recent and older data on organization of the dermal-epidermal junction, its mechanical properties and role in mechanotransduction, its involvement in regeneration, and its fate during the aging process.<\/jats:p>","DOI":"10.3390\/biom10121607","type":"journal-article","created":{"date-parts":[[2020,11,27]],"date-time":"2020-11-27T09:16:49Z","timestamp":1606468609000},"page":"1607","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":163,"title":["The Human Epidermal Basement Membrane: A Shaped and Cell Instructive Platform That Aging Slowly Alters"],"prefix":"10.3390","volume":"10","author":[{"given":"Eva","family":"Roig-Rosello","sequence":"first","affiliation":[{"name":"Laboratoire de Biologie Tissulaire et Ing\u00e9nierie Th\u00e9rapeutique, UMR 5305, CNRS-Universit\u00e9 Lyon 1, SFR BioSciences Gerland-Lyon Sud, 7 Passage du Vercors, 69367 Lyon, France"},{"name":"Roger Gallet SAS, 4 rue Euler, 75008 Paris, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patricia","family":"Rousselle","sequence":"additional","affiliation":[{"name":"Laboratoire de Biologie Tissulaire et Ing\u00e9nierie Th\u00e9rapeutique, UMR 5305, CNRS-Universit\u00e9 Lyon 1, SFR BioSciences Gerland-Lyon Sud, 7 Passage du Vercors, 69367 Lyon, France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,11,27]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1146\/annurev.cellbio.23.090506.123357","article-title":"Mechanisms regulating epithelial stratification","volume":"23","author":"Koster","year":"2007","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1155\/2013\/179784","article-title":"Skin basement membrane: The foundation of epidermal integrity\u2014BM functions and diverse roles of bridging molecules nidogen and perlecan","volume":"2013","author":"Breitkreutz","year":"2013","journal-title":"Biomed Res. Int."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"a004911","DOI":"10.1101\/cshperspect.a004911","article-title":"Basement membranes: Cell scaffoldings and signaling platforms","volume":"3","author":"Yurchenco","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Pozzi, A., Yurchenco, P.D., and Iozzo, R.V. (2017). The nature and biology of basement membranes. Matrix Biol., 1\u201311.","DOI":"10.1016\/j.matbio.2016.12.009"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"jcs241794","DOI":"10.1242\/jcs.241794","article-title":"Atypical basement membranes and basement membrane diversity\u2013what is normal anyway?","volume":"133","year":"2020","journal-title":"J. Cell Sci."},{"key":"ref_6","first-page":"213","article-title":"Structure and molecular assembly of the dermal-epidermal attachment complex in skin","volume":"30","author":"Nishiyama","year":"1998","journal-title":"Connect. Tissue"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/S0923-1811(03)00045-8","article-title":"Epidermal basement membrane zone components: Ultrastructural distribution and molecular interactions","volume":"31","author":"McMillan","year":"2003","journal-title":"J. Dermatol. Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1242\/jcs.72.1.163","article-title":"Anchoring filaments of the amphibian epidermal-dermal junction transverse the basal lamina entirely from plasma membranes of hemidesmosomes to the dermis","volume":"72","author":"Ellison","year":"1984","journal-title":"J. Cell. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1083\/jcb.114.3.567","article-title":"Kalinin: An epithelium-specific basement membrane adhesion molecule that is a component of anchoring filaments","volume":"114","author":"Rousselle","year":"1991","journal-title":"J. Cell. Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1083\/jcb.104.3.611","article-title":"Type VII collagen forms an extended network of anchoring fibrils","volume":"104","author":"Keene","year":"1987","journal-title":"J. Cell Biol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"24506","DOI":"10.1074\/jbc.M802415200","article-title":"Supramolecular interactions in the dermo-epidermal junction zone: Anchoring fibril-collagen VII tightly binds to banded collagen fibrils","volume":"283","author":"Villone","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1007\/s00441-015-2216-6","article-title":"Molecular architecture and function of the hemidesmosome","volume":"360","author":"Walko","year":"2015","journal-title":"Cell Tissue Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1007\/s12079-017-0417-3","article-title":"Restoration of the basement membrane after wounding: A hallmark of young human skin altered with aging","volume":"12","author":"Fisher","year":"2018","journal-title":"J. Cell Commun. Signal."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/bs.ctdb.2018.02.005","article-title":"Basement membranes in development and disease","volume":"130","author":"Sekiguchi","year":"2018","journal-title":"Curr. Top. Dev. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Rousselle, P., Montmasson, M., and Garnier, C. (2019). Extracellular matrix contribution to skin wound re-epithelialization. Matrix Biol., 12\u201326.","DOI":"10.1016\/j.matbio.2018.01.002"},{"key":"ref_16","first-page":"30073-1","article-title":"A novel mechanism in wound healing: Laminin 332 drives MMP9\/14 activity by recruiting syndecan-1 and CD44","volume":"S0945-053X","author":"Michopoulou","year":"2020","journal-title":"Matrix Biol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Itoh, Y. (2015). Membrane-type matrix metalloproteinases: Their functions and regulations. Matrix Biol., 207\u2013223.","DOI":"10.1016\/j.matbio.2015.03.004"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"122","DOI":"10.4161\/cam.23132","article-title":"Laminin 332 processing impacts cellular behavior","volume":"7","author":"Rousselle","year":"2013","journal-title":"Cell Adhes. Migr."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"646","DOI":"10.1038\/nrm1702","article-title":"Basement membrane proteoglycans: From cellar to ceiling","volume":"6","author":"Iozzo","year":"2005","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/j.cub.2017.02.006","article-title":"Basement membranes","volume":"27","author":"Jayadev","year":"2017","journal-title":"Curr. Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"4466","DOI":"10.1111\/febs.13495","article-title":"New concepts in basement membrane biology","volume":"282","author":"Halfter","year":"2015","journal-title":"FEBS J."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"28098","DOI":"10.3390\/ijms161226078","article-title":"Dermal contributions to human interfollicular epidermal architecture and self-renewal","volume":"16","author":"Lawlor","year":"2015","journal-title":"Int. J. Mol. Sci."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2566","DOI":"10.1001\/jama.1988.03720170042031","article-title":"Burn wounds resurfaced by cultured epidermal autografts show abnormal reconstitution of anchoring fibrils","volume":"259","author":"Woodley","year":"1988","journal-title":"JAMA"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1111\/bjd.15353","article-title":"Organization of the dermal matrix impacts the biomechanical properties of skin","volume":"177","author":"Langton","year":"2017","journal-title":"Br. J. Dermatol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1046\/j.1087-0024.2000.00010.x","article-title":"The cutaneous microcirculation","volume":"5","author":"Braverman","year":"2000","journal-title":"J. Investig. Dermatol. Symp. Proc."},{"key":"ref_26","first-page":"399","article-title":"The morphology of the attachment between the dermis and the epidermis","volume":"108","author":"Odland","year":"1950","journal-title":"Anat. Rec. Adv. Integr. Anat. Evol. Biol."},{"key":"ref_27","unstructured":"Fawcett, D.W. (1994). Skin. Bloom and Fawcett: A Textbook of Histology, Chapman & Hall. [6th ed.]."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1111\/srt.12312","article-title":"Novel approaches to characterize age-related remodelling of the dermal-epidermal junction in 2D, 3D and in vivo","volume":"23","author":"Newton","year":"2017","journal-title":"Skin Res. Technol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/pde.12796","article-title":"Developmental changes in neonatal and infant skin structures during the first 6 months: In vivo observation","volume":"33","author":"Miyauchi","year":"2016","journal-title":"Pediatr. Dermatol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1117\/1.1645795","article-title":"Characterization of age-related effects in human skin: A comparative study that applies confocal laser scanning microscopy and optical coherence tomography","volume":"9","author":"Neerken","year":"2004","journal-title":"J. Biomed. Opt."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"e201960063","DOI":"10.1002\/jbio.201960063","article-title":"Comparative analysis of intrinsic skin aging between Caucasian and Asian subjects by slide-free in vivo harmonic generation microscopy","volume":"13","author":"Lin","year":"2020","journal-title":"J. Biophotonics"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1038\/jid.2009.297","article-title":"Human skin aging is associated with reduced expression of the stem cell markers beta1 integrin and MCSP","volume":"130","author":"Giangreco","year":"2010","journal-title":"J. Investig. Dermatol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1111\/bjd.14280","article-title":"Morphological, structural and biophysical properties of French and Brazilian photoaged skin","volume":"174","author":"Mercurio","year":"2016","journal-title":"Br. J. Dermatol."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Cinotti, E., Bovi, C., Tonini, G., Labeille, B., Heus\u00e8le, C., Nizard, C., Schnebert, S., Aubailly, S., Barth\u00e9l\u00e9my, J., and Cambazard, F. (2020). Structural skin changes in elderly people investigated by reflectance confocal microscopy. J. Eur. Acad. Dermatol. Venereol.","DOI":"10.1111\/jdv.16466"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1111\/j.1600-0625.2012.01451.x","article-title":"In vivo quantification of epidermis pigmentation and dermis papilla density with reflectance confocal microscopy: Variations with age and skin phototype","volume":"21","author":"Lagarrigue","year":"2012","journal-title":"Exp. Dermatol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"101527","DOI":"10.1016\/j.bspc.2019.04.004","article-title":"Automatic evaluation of stratum basale and dermal papillae using ultrahigh resolution optical coherence tomography","volume":"53","author":"Xie","year":"2019","journal-title":"Biomed. Signal. Process. Control."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"024003","DOI":"10.1117\/1.JMI.6.2.024003","article-title":"Three-dimensional conditional random field for the dermal\u2013epidermal junction segmentation","volume":"6","author":"Robic","year":"2019","journal-title":"J. Med. Imaging"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1159\/000342926","article-title":"Morphogenesis of rete ridges in human oral mucosa: A pioneering morphological and immunohistochemical study","volume":"197","author":"Wu","year":"2013","journal-title":"Cells Tissues Organs"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"2121","DOI":"10.1002\/term.2952","article-title":"Mechanical stretching stimulates growth of the basal layer and rete ridges in the epidermis","volume":"13","author":"Topczewska","year":"2019","journal-title":"J. Tissue Eng. Regen. Med."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"eaay0244","DOI":"10.1126\/sciadv.aay0244","article-title":"Morphology and composition play distinct and complementary roles in the tolerance of plantar skin to mechanical load","volume":"5","author":"Boyle","year":"2019","journal-title":"Sci. Adv."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1136\/jmg.10.3.201","article-title":"The development of the epidermal ridges","volume":"10","author":"Penrose","year":"1973","journal-title":"J. Med. Genet."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1039\/C5IB00238A","article-title":"Mimicking the topography of the epidermal-dermal interface with elastomer substrates","volume":"8","author":"Viswanathan","year":"2016","journal-title":"Integr. Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.aanat.2018.05.001","article-title":"The development of human digital Meissner\u2019s and Pacinian corpuscles","volume":"219","author":"Feito","year":"2018","journal-title":"Ann. Anat. Anat. Anz."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1136\/jmg.12.3.243","article-title":"Development of dermal ridges in the fetus","volume":"12","author":"Okajima","year":"1975","journal-title":"J. Med. Genet."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1111\/1523-1747.ep12532761","article-title":"Structural changes in aging human skin","volume":"73","author":"Montagna","year":"1979","journal-title":"J. Investig. Dermatol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1196\/annals.1354.044","article-title":"Prevention and treatment of skin aging","volume":"1067","author":"McCullough","year":"2006","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1111\/srt.12180","article-title":"Changes in dermal papilla structures due to aging in the facial cheek region","volume":"21","author":"Mizukoshi","year":"2015","journal-title":"Skin Res. Technol."},{"key":"ref_48","first-page":"665","article-title":"Skin aging: Molecular pathology, dermal remodelling and the imaging revolution","volume":"150","author":"Newton","year":"2015","journal-title":"G. Ital. Dermatol. Venereol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"31s","DOI":"10.1038\/jid.1983.8","article-title":"Transplantation of psoriatic skin onto nude mice","volume":"80","author":"Fraki","year":"1983","journal-title":"J. Investig. Dermatol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/j.alit.2014.06.001","article-title":"Periostin contributes to epidermal hyperplasia in psoriasis common to atopic dermatitis","volume":"64","author":"Arima","year":"2015","journal-title":"Allergol. Int."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1018","DOI":"10.1111\/cup.13033","article-title":"Psoriasis or not? Review of 51 clinically confirmed cases reveals an expanded histopathologic spectrum of psoriasis","volume":"44","author":"Chau","year":"2017","journal-title":"J. Cutan. Pathol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1126\/science.7058342","article-title":"Heterogeneity in epidermal basal keratinocytes: Morphological and functional correlations","volume":"215","author":"Lavker","year":"1982","journal-title":"Science"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"13473","DOI":"10.1073\/pnas.250380097","article-title":"Epidermal stem cells: Properties, markers, and location","volume":"97","author":"Lavker","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_54","doi-asserted-by":"crossref","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":"ref_55","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/0092-8674(95)90453-0","article-title":"Stem cell patterning and fate in human epidermis","volume":"80","author":"Jones","year":"1995","journal-title":"Cell"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"2409","DOI":"10.1242\/dev.126.11.2409","article-title":"The spatial relationship between stem cells and their progeny in the basal layer of human epidermis: A new view based on whole-mount labelling and lineage analysis","volume":"126","author":"Jensen","year":"1999","journal-title":"Development"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.mad.2018.03.007","article-title":"Laminin-332 regulates differentiation of human interfollicular epidermal stem cells","volume":"171","author":"Yamada","year":"2018","journal-title":"Mech. Ageing Dev."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1111\/j.0022-202X.2004.23599.x","article-title":"Organization of stem cells and their progeny in human epidermis","volume":"124","author":"Ghazizadeh","year":"2005","journal-title":"J. Investig. Dermatol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1214","DOI":"10.1016\/S0140-6736(13)61897-8","article-title":"Allogeneic blood and bone marrow cells for the treatment of severe epidermolysis bullosa: Repair of the extracellular matrix","volume":"382","author":"Tolar","year":"2013","journal-title":"Lancet"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"206","DOI":"10.1016\/j.jmbbm.2019.07.025","article-title":"Multimodal investigation of a keloid scar by combining mechanical tests in vivo with diverse imaging techniques","volume":"99","author":"Chambert","year":"2019","journal-title":"J. Mech. Behav. Biomed. Mater."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1111\/j.1067-1927.2004.012311.x","article-title":"Molecular, histologic, and gross phenotype of skin wound healing in red Duroc pigs reveals an abnormal healing phenotype of hypercontracted, hyperpigmented scarring","volume":"12","author":"Olson","year":"2004","journal-title":"Wound Repair Regen."},{"key":"ref_62","first-page":"e29","article-title":"A translational animal model for scar compression therapy using an automated pressure delivery system","volume":"15","author":"Alkhalil","year":"2015","journal-title":"Eplasty"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1111\/exd.13875","article-title":"Regeneration of rete ridges in Lanyu pig (Sus scrofa): Insights for human skin wound healing","volume":"28","author":"Lin","year":"2019","journal-title":"Exp. Dermatol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1016\/j.addr.2018.06.019","article-title":"Re-epithelialization of adult skin wounds: Cellular mechanisms and therapeutic strategies","volume":"146","author":"Rousselle","year":"2019","journal-title":"Adv. Drug Deliv. Rev."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1097\/00000637-200044040-00019","article-title":"Reconstruction of aplasia cutis congenita of the trunk in a newborn infant using a cellular allogenic dermal graft and cultured epithelial autografts","volume":"44","author":"Simman","year":"2000","journal-title":"Ann. Plast. Surg."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1111\/j.1468-3083.2007.02326.x","article-title":"Decellularized dermis in combination with cultivated keratinocytes in a short- and long-term animal experimental investigation","volume":"22","author":"Bannasch","year":"2008","journal-title":"J. Eur. Acad. Dermatol. Venereol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/j.burns.2009.10.018","article-title":"The use of AlloDerm on major burn patients: AlloDerm prevents post-burn joint contracture","volume":"36","author":"Yim","year":"2010","journal-title":"Burns"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Dussoyer, M., Michopoulou, A., and Rousselle, P. (2020). Decellularized scaffolds for skin repair and regeneration. Appl. Sci., 10.","DOI":"10.3390\/app10103435"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"662","DOI":"10.1097\/00005373-199305000-00008","article-title":"Wound closure and outcome in extensively burned patients treated with cultured autologous keratinocytes","volume":"34","author":"Rue","year":"1993","journal-title":"J. Trauma"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1016\/S0305-4179(99)00143-6","article-title":"Cultured epithelial autografts in extensive burn coverage of severely traumatized patients: A five year single-center experience with 30 patients","volume":"26","author":"Carsin","year":"2000","journal-title":"Burns"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1016\/j.burns.2006.11.002","article-title":"Cultured epithelial autograft (CEA) in burn treatment: Three decades later","volume":"33","author":"Atiyeh","year":"2007","journal-title":"Burns"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1016\/j.burns.2012.10.004","article-title":"Chronological histological findings of cultured epidermal autograft over bilayer artificial dermis","volume":"39","author":"Matsumura","year":"2013","journal-title":"Burns"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"544","DOI":"10.1002\/jbm.a.31689","article-title":"In vitro engineering of a palatal mucosa equivalent with acellular porcine dermal matrix","volume":"86","author":"Xiong","year":"2008","journal-title":"J. Biomed. Mater. Res. A"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1111","DOI":"10.1089\/ten.tec.2009.0725","article-title":"Development of a three-dimensional human skin equivalent wound model for investigating novel wound healing therapies","volume":"16","author":"Xie","year":"2010","journal-title":"Tissue Eng. Part C Methods"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1089\/ten.tea.2016.0296","article-title":"In vivo evaluation and imaging of a bilayered self-assembled skin substitute using a decellularized dermal matrix grafted on mice","volume":"23","author":"Goyer","year":"2017","journal-title":"Tissue Eng. Part A"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1096\/fasebj.14.3.593","article-title":"Microfabrication of an analog of the basal lamina: Biocompatible membranes with complex topographies","volume":"14","author":"Pins","year":"2000","journal-title":"FASEB J."},{"key":"ref_77","first-page":"47","article-title":"The influence of microtextured basal lamina analog topography on keratinocyte function and epidermal organization","volume":"72","author":"Downing","year":"2004","journal-title":"J. Biomed. Mater. Res. A"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1002\/mabi.201100443","article-title":"Construction of a microstructured collagen membrane mimicking the papillary dermis architecture and guiding keratinocyte morphology and gene expression","volume":"12","author":"Lammers","year":"2012","journal-title":"Macromol. Biosci."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"2343","DOI":"10.1089\/ten.tea.2011.0479","article-title":"Development of microfabricated dermal epidermal regenerative matrices to evaluate the role of cellular microenvironments on epidermal morphogenesis","volume":"18","author":"Bush","year":"2012","journal-title":"Tissue Eng. Part A"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"9474","DOI":"10.1016\/j.actbio.2013.08.017","article-title":"Micropatterned dermal-epidermal regeneration matrices create functional niches that enhance epidermal morphogenesis","volume":"9","author":"Clement","year":"2013","journal-title":"Acta Biomater."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1452","DOI":"10.1016\/j.biomaterials.2005.08.004","article-title":"Electrospinning of collagen nanofibers: Effects on the behavior of normal human keratinocytes and early-stage wound healing","volume":"27","author":"Rho","year":"2006","journal-title":"Biomaterials"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1496","DOI":"10.1016\/j.biomaterials.2013.11.013","article-title":"Regulation of migratory activity of human keratinocytes by topography of multiscale collagen-containing nanofibrous matrices","volume":"35","author":"Fu","year":"2014","journal-title":"Biomaterials"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1016\/j.actbio.2019.01.063","article-title":"Patterning of human epidermal stem cells on undulating elastomer substrates reflects differences in cell stiffness","volume":"87","author":"Mobasseri","year":"2019","journal-title":"Acta Biomater."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1089\/ten.tea.2018.0125","article-title":"Dynamic culture substrates that mimic the topography of the epidermal-dermal junction","volume":"25","author":"Helling","year":"2019","journal-title":"Tissue Eng. Part A"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.actbio.2019.11.051","article-title":"Fractional CO2 laser micropatterning of cell-seeded electrospun collagen scaffolds enables rete ridge formation in 3D engineered skin","volume":"102","author":"Blackstone","year":"2020","journal-title":"Acta Biomater."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1138","DOI":"10.1089\/ten.tea.2020.0090","article-title":"Cultured epithelial autograft combined with micropatterned dermal template forms rete ridges in vivo","volume":"26","author":"Malara","year":"2020","journal-title":"Tissue Eng. Part A"},{"key":"ref_87","first-page":"1661","article-title":"An active role for basement membrane assembly and modification in tissue sculpting","volume":"128","author":"Morrissey","year":"2015","journal-title":"J. Cell Sci."},{"key":"ref_88","first-page":"72","article-title":"Basement membrane mechanics shape development: Lessons from the fly","volume":"75\u201376","year":"2019","journal-title":"Matrix Biol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1950","DOI":"10.1038\/jid.2011.146","article-title":"Migration of immunocytes across the basement membrane in skin: The role of basement membrane pores","volume":"131","author":"Oakford","year":"2011","journal-title":"J. Investig. Dermatol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1016\/j.matbio.2016.07.001","article-title":"Mechanical properties of basement membrane in health and disease","volume":"57\u201358","author":"Miller","year":"2017","journal-title":"Matrix Biol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"2897","DOI":"10.1111\/j.1742-4658.2007.05823.x","article-title":"Biomechanical properties of native basement membranes","volume":"274","author":"Candiello","year":"2007","journal-title":"FEBS J."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.jsb.2009.03.012","article-title":"Determining the mechanical properties of human corneal basement membranes with atomic force microscopy","volume":"167","author":"Last","year":"2009","journal-title":"J. Struct. Biol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"e24958","DOI":"10.7554\/eLife.24958","article-title":"Organ sculpting by patterned extracellular matrix stiffness","volume":"6","author":"Crest","year":"2017","journal-title":"Elife"},{"key":"ref_94","first-page":"4350","article-title":"Variations in basement membrane mechanics are linked to epithelial morphogenesis","volume":"144","author":"Chlasta","year":"2017","journal-title":"Development"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"2456","DOI":"10.1083\/jcb.201903066","article-title":"Beyond proteases: Basement membrane mechanics and cancer invasion","volume":"218","author":"Chang","year":"2019","journal-title":"J. Cell Biol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1111\/j.1600-0625.2012.01482.x","article-title":"A direct method to determine the strength of the dermal-epidermal junction in a mouse model for epidermolysis bullosa","volume":"21","author":"Sproule","year":"2012","journal-title":"Exp. Dermatol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"3231","DOI":"10.1002\/jbm.a.36519","article-title":"A 3D in vitro model of the dermoepidermal junction amenable to mechanical testing","volume":"106","author":"Jung","year":"2018","journal-title":"J. Biomed. Mater. Res. Part A"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1089\/ten.tec.2019.0268","article-title":"Peel test to assess the adhesion strength of the dermal-epidermal junction in tissue-engineered skin","volume":"26","author":"LaRose","year":"2020","journal-title":"Tissue Eng. Part C Methods"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1016\/j.mechrescom.2011.12.001","article-title":"Papillary network in the dermal-epidemal junction of skin: A biochemical model","volume":"42","author":"Ciarletta","year":"2012","journal-title":"Mech. Res. Commun."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"806","DOI":"10.1039\/C5SM02082D","article-title":"Understanding geometric instabilities in thin films via a multi-layer model","volume":"12","author":"Lejeune","year":"2016","journal-title":"Soft Matter"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"291","DOI":"10.3233\/NHA-170037","article-title":"Skin aging as a mechanical phenomenon: The main weak links","volume":"4","author":"Kruglikov","year":"2018","journal-title":"Nutr. Healthy Aging"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1038\/nrm2594","article-title":"Mechanotransduction at a distance: Mechanically coupling the extracellular matrix with the nucleus","volume":"10","author":"Wang","year":"2009","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1146\/annurev.cellbio.19.111301.153011","article-title":"Adhesion-dependent cell mechanosensitivity","volume":"19","author":"Bershadsky","year":"2003","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s10237-005-0012-z","article-title":"An Introductory Review of Cell Mechanobiology","volume":"5","author":"Wang","year":"2006","journal-title":"Biomech. Model. Mechanobiol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1139","DOI":"10.1126\/science.1116995","article-title":"Tissue cells feel and respond to the stiffness of their substrate","volume":"310","author":"Discher","year":"2005","journal-title":"Science"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"1103","DOI":"10.1038\/nature07765","article-title":"A mechanosensitive transcriptional mechanism that controls angiogenesis","volume":"457","author":"Mammoto","year":"2009","journal-title":"Nature"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"1425","DOI":"10.1126\/science.276.5317.1425","article-title":"Geometric control of cell life and death","volume":"276","author":"Chen","year":"1997","journal-title":"Science"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"1972","DOI":"10.1073\/pnas.032668799","article-title":"Engineering gene expression and protein synthesis by modulation of nuclear shape","volume":"99","author":"Thomas","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"1573","DOI":"10.1016\/S0142-9612(97)00144-0","article-title":"Topographical control of cells","volume":"18","author":"Curtis","year":"1997","journal-title":"Biomaterials"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1016\/j.jmbbm.2013.04.023","article-title":"Epithelial mechanobiology, skin wound healing, and the stem cell niche","volume":"28","author":"Evans","year":"2013","journal-title":"J. Mech. Behav. Biomed. Mater."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1097\/00006534-199302000-00001","article-title":"Mitotic activity in expanded human skin","volume":"91","author":"Olenius","year":"1993","journal-title":"Plast. Reconstr. Surg."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1002\/(SICI)1097-4644(19971201)67:3<327::AID-JCB5>3.0.CO;2-Y","article-title":"Cyclic strain stimulates isoform-specific PKC activation and translocation in cultured human keratinocytes","volume":"67","author":"Takei","year":"1997","journal-title":"J. Cell. Biochem."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1046\/j.1523-1747.2000.00915.x","article-title":"Signaling of mechanical stretch in human keratinocytes via MAP kinases","volume":"114","author":"Kippenberger","year":"2000","journal-title":"J. Investig. Dermatol."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1111\/j.0022-202X.2004.22328.x","article-title":"Mechanical stretching in vitro regulates signal transduction pathways and cellular proliferation in human epidermal keratinocytes","volume":"122","author":"Yano","year":"2004","journal-title":"J. Investig. Dermatol."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"5576","DOI":"10.1073\/pnas.85.15.5576","article-title":"Cell shape controls terminal differentiation of human epidermal keratinocytes","volume":"85","author":"Watt","year":"1988","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1038\/ncb2074","article-title":"Actin and serum response factor transduce physical cues from the microenvironment to regulate epidermal stem cell fate decisions","volume":"12","author":"Connelly","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/j.actbio.2018.12.003","article-title":"Micro-scaled topographies direct differentiation of human epidermal stem cells","volume":"84","author":"Zijl","year":"2019","journal-title":"Acta Biomater."},{"key":"ref_118","doi-asserted-by":"crossref","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. Investig. Dermatol."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1038\/nrc2544","article-title":"A tense situation: Forcing tumour progression","volume":"9","author":"Butcher","year":"2009","journal-title":"Nat. Rev. Cancer"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/j.ccr.2005.08.010","article-title":"Tensional homeostasis and the malignant phenotype","volume":"8","author":"Paszek","year":"2005","journal-title":"Cancer Cell"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1083\/jcb.200305010","article-title":"ROCK-generated contractility regulates breast epithelial cell differentiation in response to the physical properties of a three-dimensional collagen matrix","volume":"163","author":"Wozniak","year":"2003","journal-title":"J. Cell Biol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"776","DOI":"10.1016\/j.ccr.2011.05.008","article-title":"Actomyosin-mediated cellular tension drives increased tissue stiffness and \u03b2-catenin activation to induce epidermal hyperplasia and tumor growth","volume":"19","author":"Samuel","year":"2011","journal-title":"Cancer Cell"},{"key":"ref_123","doi-asserted-by":"crossref","unstructured":"Tokuyama, E., Nagai, Y., Takahashi, K., Kimata, Y., and Naruse, K. (2015). Mechanical stretch on human skin equivalents increases the epidermal thickness and develops the basement membrane. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0141989"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"642","DOI":"10.1038\/nmat3339","article-title":"Extracellular-matrix tethering regulates stem-cell fate","volume":"11","author":"Trappmann","year":"2012","journal-title":"Nat. Mater."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"15206","DOI":"10.1038\/ncomms15206","article-title":"YAP\/TAZ link cell mechanics to Notch signalling to control epidermal stem cell fate","volume":"8","author":"Totaro","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1038\/nphys1269","article-title":"Physical forces during collective cell migration","volume":"5","author":"Trepat","year":"2009","journal-title":"Nat. Phys."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/j.burns.2011.09.002","article-title":"Substrate stiffness regulates the proliferation, migration, and differentiation of epidermal cells","volume":"38","author":"Wang","year":"2012","journal-title":"Burns"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/srep20139","article-title":"Hierarchy of cellular decisions in collective behavior: Implications for wound healing","volume":"6","author":"Wickert","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"3539","DOI":"10.1039\/C5RA19947F","article-title":"Substrate stiffness does affect the fate of human keratinocytes","volume":"6","author":"Gupta","year":"2016","journal-title":"RSC Adv."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"jcs215780","DOI":"10.1242\/jcs.215780","article-title":"Tissue stiffening promotes keratinocyte proliferation through activation of epidermal growth factor signaling","volume":"131","author":"Kenny","year":"2018","journal-title":"J. Cell Sci."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1007\/s00403-019-01962-5","article-title":"Substrate softness promotes terminal differentiation of human keratinocytes without altering their ability to proliferate back into a rigid environment","volume":"311","author":"Ya","year":"2019","journal-title":"Arch. Dermatol. Res."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1007\/s12195-015-0377-8","article-title":"Substrate stiffness affects human keratinocyte colony formation","volume":"8","author":"Zarkoob","year":"2015","journal-title":"Cell. Mol. Bioeng."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1096\/fasebj.4.6.2180767","article-title":"Molecular architecture of basement membranes","volume":"4","author":"Yurchenco","year":"1990","journal-title":"FASEB J."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1038\/nrc1094","article-title":"Basement membranes: Structure, assembly and role in tumour angiogenesis","volume":"3","author":"Kalluri","year":"2003","journal-title":"Nat. Rev. Cancer"},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"18700","DOI":"10.1074\/jbc.M111.336073","article-title":"The epidermal basement membrane is a composite of separate laminin- or collagen IV-containing networks connected by aggregated perlecan, but not by nidogens","volume":"287","author":"Behrens","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1016\/j.matbio.2018.07.004","article-title":"A scar-like lesion is apparent in basement membrane after wound repair in vivo","volume":"74","author":"LaFever","year":"2018","journal-title":"Matrix Biol."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.semcdb.2018.07.025","article-title":"Matrix molecules and skin biology","volume":"89","author":"Nystroem","year":"2019","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"21437","DOI":"10.1074\/jbc.M702963200","article-title":"Role of laminin terminal globular domains in basement membrane assembly","volume":"282","author":"McKee","year":"2007","journal-title":"J. Biol. Chem."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1083\/jcb.200401058","article-title":"Laminin: The crux of basement membrane assembly","volume":"164","author":"Sasaki","year":"2004","journal-title":"J. Cell Biol."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1046\/j.1469-7580.1998.19310001.x","article-title":"The role of laminins in basement membrane function","volume":"193","author":"Aumailley","year":"1998","journal-title":"J. Anat."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1042\/EBC20180075","article-title":"Structural biology of laminins","volume":"63","author":"Hohenester","year":"2019","journal-title":"Essays Biochem."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"987","DOI":"10.1016\/j.tcb.2019.10.001","article-title":"Laminins in cellular differentiation","volume":"29","author":"Yap","year":"2019","journal-title":"Trends Cell Biol."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1083\/jcb.117.5.1119","article-title":"Laminin forms an independent network in basement membranes","volume":"117","author":"Yurchenco","year":"1992","journal-title":"J. Cell. Biol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1016\/S1534-5807(03)00128-X","article-title":"The role of laminin in embryonic cell polarization and tissue organization","volume":"4","author":"Li","year":"2003","journal-title":"Dev. Cell"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"31525","DOI":"10.1074\/jbc.272.50.31525","article-title":"Self-assembly of laminin isoforms","volume":"272","author":"Cheng","year":"1997","journal-title":"J. Biol. Chem."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"56","DOI":"10.4161\/cam.21831","article-title":"Laminins in basement membrane assembly","volume":"7","author":"Hohenester","year":"2013","journal-title":"Cell Adhes. Migr."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/S0945-053X(98)00004-3","article-title":"Laminins of the dermo-epidermal junction","volume":"18","author":"Aumailley","year":"1999","journal-title":"Matrix Biol."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1369\/jhc.5A6701.2005","article-title":"Colocalization of multiple laminin isoforms predominantly beneath hemidesmosomes in the upper lamina densa of the epidermal basement membrane","volume":"54","author":"McMillan","year":"2006","journal-title":"J. Histochem. Cytochem."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.matbio.2016.05.001","article-title":"Laminin \u03b15 in the keratinocyte basement membrane is required for epidermal-dermal intercommunication","volume":"56","author":"Wegner","year":"2016","journal-title":"Matrix Biol."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1136\/jmedgenet-2017-104555","article-title":"Identification of the first dominant mutation of LAMA5 gene causing a complex multisystem syndrome due to dysfunction of the extracellular matrix","volume":"54","author":"Sampaolo","year":"2017","journal-title":"J. Med. Genet."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"2400","DOI":"10.1093\/emboj\/cdg239","article-title":"Laminin-10 is crucial for hair morphogenesis","volume":"22","author":"Julia","year":"2003","journal-title":"EMBO J."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"2111","DOI":"10.1101\/gad.1689908","article-title":"Laminin-511 is an epithelial message promoting dermal papilla development and function during early hair morphogenesis","volume":"22","author":"Gao","year":"2008","journal-title":"Genes Dev."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"dev189183","DOI":"10.1242\/dev.189183","article-title":"A mutation affecting laminin alpha 5 polymerisation gives rise to a syndromic developmental disorder","volume":"147","author":"Jones","year":"2020","journal-title":"Development"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"390","DOI":"10.1172\/JCI200419140","article-title":"Extensive tissue-regenerative capacity of neonatal human keratinocyte stem cells and their progeny","volume":"113","author":"Li","year":"2004","journal-title":"J. Clin. Investig."},{"key":"ref_155","first-page":"2795","article-title":"A role for pericytes as microenvironmental regulators of human skin tissue regeneration","volume":"119","author":"Li","year":"2009","journal-title":"J. Clin. Investig."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/bs.asn.2020.06.002","article-title":"The aging epidermal skin niche","volume":"Volume 4","author":"Nilsson","year":"2020","journal-title":"Advances in Stem Cells and Their Niches"},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1002\/jemt.20564","article-title":"Mammalian collagen IV","volume":"71","author":"Khoshnoodi","year":"2008","journal-title":"Microsc. Res. Tech."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1159\/000237090","article-title":"The collagen superfamily","volume":"107","author":"Brown","year":"1995","journal-title":"Int. Arch. Allergy Immunol."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1111\/1523-1747.ep12275360","article-title":"Secretion of type IV collagen by keratinocytes of human adult","volume":"85","author":"Oguchi","year":"1985","journal-title":"J. Investig. Dermatol."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1111\/1523-1747.ep12277381","article-title":"Differential expression of type IV procollagen and laminin genes by fetal vs adult skin fibroblasts in culture: Determination of subunit mRNA steady-state levels","volume":"93","author":"Olsen","year":"1989","journal-title":"J. Investig. Dermatol."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"1","DOI":"10.4103\/1947-2714.92892","article-title":"Collagen IV in normal and in disease process","volume":"4","author":"Howard","year":"2012","journal-title":"N. Am. J. Med. Sci."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"38117","DOI":"10.1074\/jbc.R600025200","article-title":"Molecular recognition in the assembly of collagens: Terminal noncollagenous domains are key recognition modules in the formation of triple helical protomers","volume":"281","author":"Khoshnoodi","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1016\/0014-5793(79)81022-4","article-title":"Identification of a new basement membrane collagen by the aid of a large fragment resistant to bacterial collagenase","volume":"101","author":"Timpl","year":"1979","journal-title":"FEBS Lett."},{"key":"ref_164","first-page":"239","article-title":"7-S Collagen: Characterization of an unusual basement membrane structure","volume":"108","author":"Risteli","year":"1980","journal-title":"JBIC J. Biol. Inorg. Chem."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"2543","DOI":"10.1056\/NEJMra022296","article-title":"Alport\u2019s syndrome, Goodpasture\u2019s syndrome, and type IV collagen","volume":"348","author":"Hudson","year":"2003","journal-title":"N. Engl. J. Med."},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1016\/j.devcel.2011.06.026","article-title":"Shaping cells and organs in drosophila by opposing roles of fat body-secreted collagen IV and perlecan","volume":"21","author":"Xu","year":"2011","journal-title":"Dev. Cell"},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"5154","DOI":"10.1002\/j.1460-2075.1996.tb00899.x","article-title":"Site-directed mutagenesis and structural interpretation of the nidogen binding site of the laminin gamma1 chain","volume":"15","author":"Mayer","year":"1996","journal-title":"EMBO J."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"3137","DOI":"10.1002\/j.1460-2075.1991.tb04875.x","article-title":"Recombinant nidogen consists of three globular domains and mediates binding of laminin to collagen type IV","volume":"10","author":"Fox","year":"1991","journal-title":"EMBO J."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"6846","DOI":"10.1128\/MCB.25.15.6846-6856.2005","article-title":"Compound genetic ablation of nidogen 1 and 2 causes basement membrane defects and perinatal lethality in mice","volume":"25","author":"Bader","year":"2005","journal-title":"Mol. Cell. Biol."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"2259","DOI":"10.1038\/jid.2008.65","article-title":"Basement membranes in skin are differently affected by lack of nidogen 1 and 2","volume":"128","author":"Mokkapati","year":"2008","journal-title":"J. Investig. Dermatol."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"10881","DOI":"10.1016\/S0021-9258(18)82067-1","article-title":"Mapping of nidogen binding sites for collagen type IV, heparan sulfate proteoglycan, and zinc","volume":"268","author":"Reinhardt","year":"1993","journal-title":"J. Biol. Chem."},{"key":"ref_172","first-page":"1","article-title":"Integrating Activities of laminins that drive basement membrane assembly and function","volume":"76","author":"Yurchenco","year":"2015","journal-title":"Transp. Syst."},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1016\/j.matbio.2016.09.003","article-title":"A current view of perlecan in physiology and pathology: A mosaic of functions","volume":"57\u201358","author":"Gubbiotti","year":"2017","journal-title":"Matrix Biol."},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"5178","DOI":"10.1074\/jbc.M509500200","article-title":"Targeting perlecan in human keratinocytes reveals novel roles for perlecan in epidermal formation","volume":"281","author":"Sher","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"751","DOI":"10.18632\/aging.100928","article-title":"Perlecan expression influences the keratin 15-positive cell population fate in the epidermis of aging skin","volume":"8","author":"Michopoulou","year":"2016","journal-title":"Aging"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1016\/j.matbio.2010.03.004","article-title":"Age-dependent changes in the structure, composition and biophysical properties of a human basement membrane","volume":"29","author":"Candiello","year":"2010","journal-title":"Matrix Biol."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1016\/j.matbio.2010.04.002","article-title":"Molecular interactions in the retinal basement membrane system: A proteomic approach","volume":"29","author":"Balasubramani","year":"2010","journal-title":"Matrix Biol."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1038\/nsb0196-67","article-title":"Structure of a novel extracellular Ca(2+)-binding module in BM-40","volume":"3","author":"Hohenester","year":"1996","journal-title":"Nat. Struct. Biol."},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"18273","DOI":"10.1073\/pnas.0808452105","article-title":"Structural basis of sequence-specific collagen recognition by SPARC","volume":"105","author":"Hohenester","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/s12079-009-0062-6","article-title":"The role of SPARC in extracellular matrix assembly","volume":"3","author":"Bradshaw","year":"2009","journal-title":"J. Cell Commun. Signal."},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"721","DOI":"10.1002\/jcb.21688","article-title":"A prototypic matricellular protein in the tumor microenvironment-where there\u2019s SPARC, there\u2019s fire","volume":"104","author":"Clark","year":"2008","journal-title":"J. Cell. Biochem."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"967","DOI":"10.1093\/carcin\/bgu072","article-title":"Molecular mechanisms underlying the divergent roles of SPARC in human carcinogenesis","volume":"35","author":"Nagaraju","year":"2014","journal-title":"Carcinogenesis"},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1002\/jcb.240570218","article-title":"SPARC mediates focal adhesion disassembly in endothelial cells through a follistatin-like region and the Ca(2+)-binding EF-hand","volume":"57","author":"Lane","year":"1995","journal-title":"J. Cell. Biochem."},{"key":"ref_184","doi-asserted-by":"crossref","unstructured":"Chlenski, A., Guerrero, L.J., Salwen, H.R., Yang, Q., Tian, Y., La Madrid, A.M., Mirzoeva, S., Bouyer, P.G., Xu, D., and Walker, M. (2011). Secreted protein acidic and rich in cysteine is a matrix scavenger chaperone. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0023880"},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"H841","DOI":"10.1152\/ajpheart.01247.2010","article-title":"SPARC regulates collagen interaction with cardiac fibroblast cell surfaces","volume":"301","author":"Harris","year":"2011","journal-title":"Am. J. Physiol. Heart Circ. Physiol."},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1083\/jcb.109.1.341","article-title":"SPARC, a secreted protein associated with cellular proliferation, inhibits cell spreading in vitro and exhibits Ca+2-dependent binding to the extracellular matrix","volume":"109","author":"Sage","year":"1989","journal-title":"J. Cell Biol."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"1671","DOI":"10.1242\/jcs.021931","article-title":"Haemocyte-derived SPARC is required for collagen-IV-dependent stability of basal laminae in drosophila embryos","volume":"121","author":"Martinek","year":"2008","journal-title":"J. Cell Sci."},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"212","DOI":"10.1016\/j.ydbio.2015.08.018","article-title":"Dynamic regulation of basement membrane protein levels promotes egg chamber elongation in Drosophila","volume":"406","author":"Isabella","year":"2015","journal-title":"Dev. Biol."},{"key":"ref_189","doi-asserted-by":"crossref","unstructured":"Morrissey, M.A., Jayadev, R., Miley, G.R., Blebea, C.A., Chi, Q., Ihara, S., and Sherwood, D.R. (2016). SPARC promotes cell invasion in vivo by decreasing type IV collagen levels in the basement membrane. PLoS Genet., 12.","DOI":"10.1101\/037218"},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.ydbio.2017.09.037","article-title":"Collagen IV trafficking: The inside-out and beyond story","volume":"431","author":"Chioran","year":"2017","journal-title":"Dev. Biol."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1046\/j.1523-1747.1999.00483.x","article-title":"Confocal laser scanning analysis of the association of fibulin-2 with fibrillin-1 and fibronectin define different stages of skin regeneration","volume":"112","author":"Raghunath","year":"1999","journal-title":"J. Investig. Dermatol."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"906","DOI":"10.1046\/j.1523-1747.1999.00802.x","article-title":"Spatiotemporal changes of fibronectin, tenascin-C, fibulin-1, and fibulin-2 in the skin during the development of chronic contact dermatitis","volume":"113","author":"Kusubata","year":"1999","journal-title":"J. Investig. Dermatol."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1038\/nrm1130","article-title":"Fibulins: A versatile family of extracellular matrix proteins","volume":"4","author":"Timpl","year":"2003","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"1609","DOI":"10.1038\/jid.2014.10","article-title":"Reduced fibulin-2 contributes to loss of basement membrane integrity and skin blistering in mice lacking integrin \u03b13\u03b21 in the epidermis","volume":"134","author":"Longmate","year":"2014","journal-title":"J. Investig. Dermatol."},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.matbio.2016.10.002","article-title":"Collagen XVIII in tissue homeostasis and dysregulation\u2014Lessons learned from model organisms and human patients","volume":"57\u201358","author":"Heljasvaara","year":"2017","journal-title":"Matrix Biol."},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"25404","DOI":"10.1074\/jbc.273.39.25404","article-title":"Collagen XVIII is a basement membrane heparan sulfate proteoglycan","volume":"273","author":"Halfter","year":"1998","journal-title":"J. Biol. Chem."},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1016\/S0002-9440(10)65603-9","article-title":"The short and long forms of type XVIII collagen show clear tissue specificities in their expression and location in basement membrane zones in humans","volume":"153","author":"Saarela","year":"1998","journal-title":"Am. J. Pathol."},{"key":"ref_198","doi-asserted-by":"crossref","first-page":"494","DOI":"10.1006\/geno.1994.1098","article-title":"Cloning of cDNA and genomic DNA encoding human type XVIII collagen and localization of the alpha1(XVIII) collagen gene to mouse chromosome 10 and human chromosome 21","volume":"19","author":"Oh","year":"1994","journal-title":"Genomics"},{"key":"ref_199","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1038\/sj.emboj.7600014","article-title":"Collagen XVIII\/endostatin is essential for vision and retinal pigment epithelial function","volume":"23","author":"Marneros","year":"2004","journal-title":"EMBO J."},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1016\/S0002-9440(10)62246-8","article-title":"Endostatin overexpression specifically in the lens and skin leads to cataract and ultrastructural alterations in basement membranes","volume":"166","author":"Elamaa","year":"2006","journal-title":"Am. J. Pathol."},{"key":"ref_201","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1177\/002215540305100303","article-title":"The collagen type XVIII endostatin domain is co-localized with perlecan in basement membranes in vivo","volume":"51","author":"Miosge","year":"2003","journal-title":"J. Histochem. Cytochem."},{"key":"ref_202","doi-asserted-by":"crossref","first-page":"716","DOI":"10.1096\/fj.04-2134rev","article-title":"Physiological role of collagen XVIII and endostatin","volume":"19","author":"Marneros","year":"2005","journal-title":"FASEB J."},{"key":"ref_203","doi-asserted-by":"crossref","first-page":"1535","DOI":"10.1093\/emboj\/21.7.1535","article-title":"Lack of collagen XVIII\/endostatin results in eye abnormalities","volume":"21","author":"Fukai","year":"2002","journal-title":"EMBO J."},{"key":"ref_204","first-page":"35","article-title":"Collagen XVIII: A key interfacial component of the skin architecture","volume":"68","author":"Bonnet","year":"2017","journal-title":"J. Cosmet. Sci."},{"key":"ref_205","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1016\/0092-8674(91)90092-D","article-title":"Epiligrin, a new cell adhesion ligand for integrin alpha 3 beta 1 in epithelial basement membranes","volume":"65","author":"Carter","year":"1991","journal-title":"Cell"},{"key":"ref_206","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/S0738-081X(00)00175-9","article-title":"Molecular organization of the cutaneous basement membrane zone","volume":"19","author":"Ghohestani","year":"2001","journal-title":"Clin. Dermatol."},{"key":"ref_207","doi-asserted-by":"crossref","first-page":"376","DOI":"10.1016\/j.tcb.2006.05.004","article-title":"Current insights into the formation and breakdown of hemidesmosomes","volume":"16","author":"Litjens","year":"2006","journal-title":"Trends Cell Biol."},{"key":"ref_208","doi-asserted-by":"crossref","first-page":"969","DOI":"10.1083\/jcb.149.4.969","article-title":"The tetraspan molecule CD151, a novel constituent of hemidesmosomes, associates with the integrin alpha6beta4 and may regulate the spatial organization of hemidesmosomes","volume":"149","author":"Sterk","year":"2000","journal-title":"J. Cell Biol."},{"key":"ref_209","first-page":"3714","article-title":"The molecular architecture of hemidesmosomes, as revealed with super-resolution microscopy","volume":"128","author":"Nahidiazar","year":"2015","journal-title":"J. Cell Sci."},{"key":"ref_210","doi-asserted-by":"crossref","first-page":"e201904137","DOI":"10.1083\/jcb.201904137","article-title":"Hemidesmosomes modulate force generation via focal adhesions","volume":"219","author":"Wang","year":"2020","journal-title":"J. Cell Biol."},{"key":"ref_211","doi-asserted-by":"crossref","first-page":"1189","DOI":"10.1083\/jcb.132.6.1189","article-title":"Human amnion contains a novel laminin variant, laminin 7, which like laminin 6, covalently associates with laminin 5 to promote stable epithelial-stromal attachment","volume":"132","author":"Champliaud","year":"1996","journal-title":"J. Cell Biol."},{"key":"ref_212","doi-asserted-by":"crossref","first-page":"12204","DOI":"10.1074\/jbc.M111.300061","article-title":"Cell surface proteoglycans syndecan-1 and -4 bind overlapping but distinct sites in laminin \u03b13 LG45 protein domain","volume":"287","author":"Carulli","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_213","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1159\/000357032","article-title":"Processing of laminin \u03b1 chains generates peptides involved in wound healing and host defense","volume":"6","author":"Kempf","year":"2014","journal-title":"J. Innate Immun."},{"key":"ref_214","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1083\/jcb.103.4.1577","article-title":"Type VII collagen is a major structural component of anchoring fibrils","volume":"103","author":"Sakai","year":"1986","journal-title":"J. Cell Biol."},{"key":"ref_215","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1111\/1346-8138.14222","article-title":"Epidermal aspects of type VII collagen: Implications for dystrophic epidermolysis bullosa and epidermolysis bullosa acquisita","volume":"45","author":"Watanabe","year":"2018","journal-title":"J. Dermatol."},{"key":"ref_216","doi-asserted-by":"crossref","first-page":"252","DOI":"10.1111\/1523-1747.ep12365129","article-title":"Type VII collagen, anchoring fibrils and epidermolysis bullosa","volume":"101","author":"Burgeson","year":"1993","journal-title":"J. Investig. Dermatol."},{"key":"ref_217","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1083\/jcb.138.3.719","article-title":"Laminin 5 binds the NC-1 domain of type VII collagen","volume":"138","author":"Rousselle","year":"1997","journal-title":"J. Cell Biol."},{"key":"ref_218","doi-asserted-by":"crossref","first-page":"14516","DOI":"10.1074\/jbc.272.23.14516","article-title":"Interactions of the amino-terminal noncollagenous (NC1) domain of type VII collagen with extracellular matrix components","volume":"272","author":"Chen","year":"1997","journal-title":"J. Biol. Chem."},{"key":"ref_219","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1046\/j.1523-1747.1999.00491.x","article-title":"NC1 domain of type VII collagen binds to the beta 3 chain of laminin 5 via a unique subdomain within the fibronectin-like repeats","volume":"112","author":"Chen","year":"1999","journal-title":"J. Investig. Dermatol."},{"key":"ref_220","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1093\/jb\/mvi153","article-title":"Regulation of cell adhesion and type VII collagen binding by the beta3 chain short arm of laminin-5: Effect of its proteolytic cleavage","volume":"138","author":"Nakashima","year":"2005","journal-title":"J. Biochem."},{"key":"ref_221","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.matbio.2010.10.005","article-title":"Type XVII collagen (BP180) can function as a cell\u2212matrix adhesion molecule via binding to laminin 332","volume":"30","author":"Eliason","year":"2011","journal-title":"Matrix Biol."},{"key":"ref_222","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1016\/j.ajpath.2011.04.019","article-title":"Dynamic interactions of epidermal collagen XVII with the extracellular matrix: Laminin 332 as a major binding partner","volume":"179","author":"Nishie","year":"2011","journal-title":"Am. J. Pathol."},{"key":"ref_223","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1016\/S0945-053X(03)00051-9","article-title":"Collagenous transmembrane proteins: Collagen XVII as a prototype","volume":"22","author":"Franzke","year":"2003","journal-title":"Matrix Biol."},{"key":"ref_224","doi-asserted-by":"crossref","first-page":"352","DOI":"10.3389\/fonc.2020.00352","article-title":"The role of collagen XVII in cancer: Squamous cell carcinoma and beyond","volume":"10","author":"Jones","year":"2020","journal-title":"Front. Oncol."},{"key":"ref_225","doi-asserted-by":"crossref","first-page":"1135","DOI":"10.1111\/exd.13550","article-title":"Life before and beyond blistering: The role of collagen XVII in epidermal physiology","volume":"28","author":"Natsuga","year":"2019","journal-title":"Exp. Dermatol."},{"key":"ref_226","doi-asserted-by":"crossref","first-page":"1103","DOI":"10.1016\/j.jaad.2014.01.903","article-title":"Inherited epidermolysis bullosa: Updated recommendations on diagnosis and classification","volume":"70","author":"Fine","year":"2014","journal-title":"J. Am. Acad. Dermatol."},{"key":"ref_227","doi-asserted-by":"crossref","first-page":"658","DOI":"10.5021\/ad.2015.27.6.658","article-title":"Recently identified forms of epidermolysis bullosa","volume":"27","author":"McGrath","year":"2015","journal-title":"Ann. Dermatol."},{"key":"ref_228","doi-asserted-by":"crossref","unstructured":"Murrell, D.F. (2015). Laminin 332 in junctional epidermolysis and as an autoantigen in mucous membrane pemphigoid. Blistering Diseases, Springer.","DOI":"10.1007\/978-3-662-45698-9"},{"key":"ref_229","doi-asserted-by":"crossref","first-page":"1","DOI":"10.12688\/f1000research.14974.1","article-title":"Recent advances in understanding and managing epidermolysis bullosa","volume":"7","author":"Kiritsi","year":"2018","journal-title":"F1000Research"},{"key":"ref_230","first-page":"adv00053-101","article-title":"Skin fragility: Perspectives on evidence-based therapies","volume":"100","year":"2020","journal-title":"Acta Derm. Venereol."},{"key":"ref_231","doi-asserted-by":"crossref","first-page":"1176","DOI":"10.1111\/exd.13979","article-title":"Epidermolysis bullosa: Advances in research and treatment","volume":"28","author":"Prodinger","year":"2019","journal-title":"Exp. Dermatol."},{"key":"ref_232","doi-asserted-by":"crossref","first-page":"1154","DOI":"10.1111\/exd.13446","article-title":"Structural proteins of the dermal-epidermal junction targeted by autoantibodies in pemphigoid diseases","volume":"26","author":"Goletz","year":"2017","journal-title":"Exp. Dermatol."},{"key":"ref_233","first-page":"1","article-title":"Prevalence and clinical significance of anti-Laminin 332 autoantibodies detected by a novel ELISA in mucous membrane pemphigoid","volume":"20","author":"Bernard","year":"2013","journal-title":"JAMA Dermatol."},{"key":"ref_234","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1196\/annals.1404.027","article-title":"Molecular mechanisms of skin aging: State of the art","volume":"1119","author":"Makrantonaki","year":"2007","journal-title":"Ann. N.Y. Acad. Sci."},{"key":"ref_235","doi-asserted-by":"crossref","first-page":"44s","DOI":"10.1111\/1523-1747.ep12468934","article-title":"Aged skin: A study by light, transmission electron, and scanning electron microscopy","volume":"88","author":"Lavker","year":"1987","journal-title":"J. Investig. Dermatol."},{"key":"ref_236","doi-asserted-by":"crossref","first-page":"908","DOI":"10.1111\/j.1524-4725.1990.tb01554.x","article-title":"Histologic changes in skin associated with aging","volume":"16","author":"Kurban","year":"1990","journal-title":"J. Dermatol. Surg. Oncol."},{"key":"ref_237","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1159\/000213172","article-title":"Skin thickness changes in normal aging skin","volume":"36","author":"Branchet","year":"1990","journal-title":"Gerontology"},{"key":"ref_238","doi-asserted-by":"crossref","first-page":"846","DOI":"10.1046\/j.0022-202x.2001.01337.x","article-title":"Topographic variations in normal skin, as viewed by in vivo reflectance confocal microscopy","volume":"116","author":"Huzaira","year":"2001","journal-title":"J. Investig. Dermatol."},{"key":"ref_239","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1046\/j.0909-752x.2001.10297.x","article-title":"Age related changes of human skin investigated with histometric measurements by confocal laser scanning microscopy in vivo","volume":"8","author":"Sauermann","year":"2002","journal-title":"Skin Res. Technol."},{"key":"ref_240","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1034\/j.1600-0846.2000.006001017.x","article-title":"Interdigitation index\u2013a parameter for differentiating between young and older skin specimens","volume":"6","author":"Szende","year":"2000","journal-title":"Skin Res. Technol."},{"key":"ref_241","doi-asserted-by":"crossref","first-page":"3266","DOI":"10.1364\/BOE.5.003266","article-title":"Quantitative analysis of intrinsic skin aging in dermal papillae by in vivo harmonic generation microscopy","volume":"5","author":"Liao","year":"2014","journal-title":"Biomed. Opt. Express"},{"key":"ref_242","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1111\/srt.12127","article-title":"Assessment of skin photoaging with reflectance confocal microscopy","volume":"20","author":"Haytoglu","year":"2014","journal-title":"Skin Res. Technol."},{"key":"ref_243","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1111\/1523-1747.ep12532763","article-title":"Structural alterations in exposed and unexposed aged skin","volume":"73","author":"Lavker","year":"1979","journal-title":"J. Investig. Dermatol."},{"key":"ref_244","doi-asserted-by":"crossref","first-page":"73","DOI":"10.2165\/00128071-200910020-00001","article-title":"Clinical implications of aging skin: Cutaneous disorders in the elderly","volume":"10","author":"Farage","year":"2009","journal-title":"Am. J. Clin. Dermatol."},{"key":"ref_245","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.clindermatol.2010.07.001","article-title":"Clinical aspects and molecular diagnostics of skin aging","volume":"29","author":"Zoubouliss","year":"2011","journal-title":"Clin. Dermatol."},{"key":"ref_246","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1111\/j.1532-5415.1982.tb01289.x","article-title":"Age-associated changes in the skin","volume":"30","author":"Gilchrest","year":"1982","journal-title":"J. Am. Geriatr. Soc."},{"key":"ref_247","doi-asserted-by":"crossref","first-page":"1861","DOI":"10.2353\/ajpath.2006.051302","article-title":"Decreased collagen production in chronologically aged skin: Roles of age-dependent alteration in fibroblast function and defective mechanical stimulation","volume":"168","author":"Varani","year":"2006","journal-title":"Am. J. Pathol."},{"key":"ref_248","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1111\/joa.12186","article-title":"Age-related dermal collagen changes during development, maturation and ageing\u2013a morphometric and comparative study","volume":"225","year":"2014","journal-title":"J. Anat."},{"key":"ref_249","doi-asserted-by":"crossref","unstructured":"Mine, S., Fortunel, N.O., Pageon, H., and Asselineau, D. (2008). Aging alters functionally human dermal papillary fibroblasts but not reticular fibroblasts: A new view of skin morphogenesis and aging. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0004066"},{"key":"ref_250","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1684\/ejd.2012.1693","article-title":"Distinct and complementary roles of papillary and reticular fibroblasts in skin morphogenesis and homeostasis","volume":"22","author":"Pageon","year":"2012","journal-title":"Eur. J. Dermatol. EJD"},{"key":"ref_251","doi-asserted-by":"crossref","first-page":"2565","DOI":"10.1038\/jid.2012.192","article-title":"Different gene expression patterns in human papillary and reticular fibroblasts","volume":"132","author":"Janson","year":"2012","journal-title":"J. Investig. Dermatol."},{"key":"ref_252","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1016\/j.mad.2018.03.006","article-title":"Age-related evolutions of the dermis: Clinical signs, fibroblast and extracellular matrix dynamics","volume":"177","author":"Haydont","year":"2019","journal-title":"Mech. Ageing Dev."},{"key":"ref_253","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1016\/j.matbio.2010.01.008","article-title":"Tissue section AFM: In situ ultrastructural imaging of native biomolecules","volume":"29","author":"Graham","year":"2010","journal-title":"Matrix Biol."},{"key":"ref_254","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1111\/j.1365-2133.2012.10943.x","article-title":"In vivo assessment of chronological ageing and photoageing in forearm skin using reflectance confocal microscopy","volume":"167","author":"Wurm","year":"2012","journal-title":"Br. J. Dermatol."},{"key":"ref_255","doi-asserted-by":"crossref","first-page":"e73","DOI":"10.1016\/j.jaad.2011.08.021","article-title":"Skin aging: In vivo microscopic assessment of epidermal and dermal changes by means of confocal microscopy","volume":"68","author":"Longo","year":"2013","journal-title":"J. Am. Acad. Dermatol."},{"key":"ref_256","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1007\/s00403-015-1575-x","article-title":"High-definition optical coherence tomography intrinsic skin ageing assessment in women: A pilot study","volume":"307","author":"Boone","year":"2015","journal-title":"Arch. Dermatol. Res."},{"key":"ref_257","doi-asserted-by":"crossref","unstructured":"Runel, G., Cario, M., Lopez-Ramirez, N., Malbouyres, M., Ruggiero, F., Bernard, L., Puisieux, A., Caramel, J., Chlasta, J., and Masse, I. (2020). Stiffness measurement is a biomarker of skin ageing in vivo. Exp. Dermatol.","DOI":"10.1111\/exd.14195"},{"key":"ref_258","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1111\/j.1751-1097.1996.tb01851.x","article-title":"Ultraviolet B wavelength dependence for the regulation of two major matrix-metalloproteinases and their inhibitor TIMP-1 in human dermal fibroblasts","volume":"64","author":"Brenneisen","year":"1996","journal-title":"Photochem. Photobiol."},{"key":"ref_259","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1002\/path.2730","article-title":"Low-dose ultraviolet radiation selectively degrades chromophore-rich extracellular matrix components","volume":"222","author":"Sherratt","year":"2010","journal-title":"J. Pathol."},{"key":"ref_260","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1016\/j.cellsig.2018.11.024","article-title":"Selective proteolysis by matrix metalloproteinases of photo-oxidised dermal extracellular matrix proteins","volume":"54","author":"Hibbert","year":"2019","journal-title":"Cell. Signal."},{"key":"ref_261","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/S0378-5122(96)01066-3","article-title":"Changes of the basement membrane and type IV collagen in human skin during aging","volume":"25","author":"Palacios","year":"1996","journal-title":"Maturitas"},{"key":"ref_262","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1038\/jidsymp.1998.34","article-title":"Age-related functional and structural changes in human dermo-epidermal junction components","volume":"3","author":"Chaudagne","year":"1998","journal-title":"J. Investig. Dermatol. Symp. Proc."},{"key":"ref_263","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.mad.2016.03.006","article-title":"The impact of intrinsic ageing on the protein composition of the dermal-epidermal junction","volume":"156","author":"Langton","year":"2016","journal-title":"Mech. Ageing Dev."},{"key":"ref_264","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1684\/ejd.2016.2782","article-title":"Aging decreases collagen IV expression in vivo in the dermo-epidermal junction and in vitro in dermal fibroblasts: Possible involvement of TGF-\u03b21","volume":"26","author":"Feru","year":"2016","journal-title":"Eur. J. Dermatol. EJD"},{"key":"ref_265","doi-asserted-by":"crossref","first-page":"454","DOI":"10.1111\/j.1600-0625.2011.01258.x","article-title":"Changes in glycosaminoglycans and related proteoglycans in intrinsically aged human skin in vivo","volume":"20","author":"Oh","year":"2011","journal-title":"Exp. Dermatol."},{"key":"ref_266","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1038\/s41586-019-1085-7","article-title":"Stem cell competition orchestrates skin homeostasis and ageing","volume":"568","author":"Liu","year":"2019","journal-title":"Nature"},{"key":"ref_267","doi-asserted-by":"crossref","first-page":"e26635","DOI":"10.7554\/eLife.26635","article-title":"Type XVII collagen coordinates proliferation in the interfollicular epidermis","volume":"6","author":"Watanabe","year":"2017","journal-title":"eLife"},{"key":"ref_268","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.tice.2019.06.001","article-title":"Expression of anti-aging type-XVII collagen (COL17A1\/BP180) in hair follicle-associated pluripotent (HAP) stem cells during differentiation","volume":"59","author":"Shirai","year":"2019","journal-title":"Tissue Cell"},{"key":"ref_269","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1126\/science.aad4395","article-title":"Hair follicle aging is driven by transepidermal elimination of stem cells via COL17A1 proteolysis","volume":"351","author":"Matsumura","year":"2016","journal-title":"Science"},{"key":"ref_270","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.mam.2011.10.015","article-title":"Human matrix metalloproteinases: An ubiquitarian class of enzymes involved in several pathological processes","volume":"33","author":"Sbardella","year":"2012","journal-title":"Mol. Asp. Med."},{"key":"ref_271","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1111\/1523-1747.ep12371763","article-title":"Type VII collagen gene expression by human skin fibroblasts and keratinocytes in culture: Influence of donor age and cytokine responses","volume":"102","author":"Chen","year":"1994","journal-title":"J. Investig. Dermatol."},{"key":"ref_272","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1046\/j.1365-2133.1997.18471955.x","article-title":"Clinical features of photodamaged human skin are associated with a reduction in collagen VII","volume":"137","author":"Craven","year":"1997","journal-title":"Br. J. Dermatol."},{"key":"ref_273","doi-asserted-by":"crossref","first-page":"2015","DOI":"10.1016\/j.bbamcr.2017.05.007","article-title":"The role of matrix metalloproteinases in aging: Tissue remodeling and beyond","volume":"1864","author":"Folgueras","year":"2017","journal-title":"Biochim. Biophys. Acta Mol. Cell Res."},{"key":"ref_274","doi-asserted-by":"crossref","unstructured":"Pittayapruek, P., Meephansan, J., Prapapan, O., Komine, M., and Ohtsuki, M. (2016). Role of matrix metalloproteinases in photoaging and photocarcinogenesis. Int. J. Mol. Sci., 17.","DOI":"10.3390\/ijms17060868"},{"key":"ref_275","doi-asserted-by":"crossref","first-page":"9363","DOI":"10.1073\/pnas.92.20.9363","article-title":"A biomarker that identifies senescent human cells in culture and in aging skin in vivo","volume":"92","author":"Dimri","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_276","doi-asserted-by":"crossref","first-page":"2133","DOI":"10.1016\/j.jid.2016.06.621","article-title":"The senescence-associated secretory phenotype: Critical effector in skin cancer and aging","volume":"136","author":"Ghosh","year":"2016","journal-title":"J. Investig. Dermatol."},{"key":"ref_277","doi-asserted-by":"crossref","first-page":"247","DOI":"10.3389\/fgene.2018.00247","article-title":"Biomarkers of cellular senescence and skin aging","volume":"9","author":"Wang","year":"2018","journal-title":"Front. Genet."},{"key":"ref_278","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1038\/jidsymp.2009.5","article-title":"Possible involvement of basement membrane damage in skin photoaging","volume":"14","author":"Amano","year":"2009","journal-title":"J. Investig. Dermatol. Symp. Proc."},{"key":"ref_279","doi-asserted-by":"crossref","unstructured":"Farage, M., Miller, K., and Maibach, H. (2015). Pathology of Aging Skin. Textbook of Aging Skin, Springer.","DOI":"10.1007\/978-3-642-27814-3"},{"key":"ref_280","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1016\/j.jphotobiol.2011.10.013","article-title":"Ultraviolet-B irradiation induces epidermal up-regulation of heparanase expression and activity","volume":"106","author":"Kurdykowski","year":"2012","journal-title":"J. Photochem. Photobiol. B Biol."},{"key":"ref_281","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1146\/annurev-bioeng-071114-040829","article-title":"The mechanobiology of aging","volume":"17","author":"Phillip","year":"2015","journal-title":"Annu. Rev. Biomed. Eng."},{"key":"ref_282","doi-asserted-by":"crossref","first-page":"4201","DOI":"10.1242\/jcs.075150","article-title":"Micropatterning as a tool to decipher cell morphogenesis and functions","volume":"123","year":"2010","journal-title":"J. Cell Sci."},{"key":"ref_283","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1016\/j.ceb.2014.09.006","article-title":"Forms, forces, and stem cell fate","volume":"31","author":"Bellas","year":"2014","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_284","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.arr.2009.11.002","article-title":"Effect of aging on cellular mechanotransduction","volume":"10","author":"Wu","year":"2011","journal-title":"Ageing Res. Rev."},{"key":"ref_285","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1016\/j.arr.2009.10.005","article-title":"Mechanical properties of cells and ageing","volume":"10","author":"Starodubtseva","year":"2011","journal-title":"Ageing Res. Rev."},{"key":"ref_286","doi-asserted-by":"crossref","first-page":"1435","DOI":"10.1016\/j.ultramic.2010.07.009","article-title":"Single cell mechanics of keratinocyte cells","volume":"110","author":"Lulevich","year":"2010","journal-title":"Ultramicroscopy"},{"key":"ref_287","doi-asserted-by":"crossref","first-page":"e200","DOI":"10.1111\/j.1600-0846.2012.00628.x","article-title":"The influence of surfactants and hydrolyzed proteins on keratinocytes viability and elasticity","volume":"19","author":"Kobiela","year":"2013","journal-title":"Skin Res. Technol."},{"key":"ref_288","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1088\/0031-9155\/50\/1\/007","article-title":"Human epithelial cells increase their rigidity with ageing in vitro: Direct measurements","volume":"50","author":"Berdyyeva","year":"2005","journal-title":"Phys. Med. Biol."},{"key":"ref_289","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.nano.2005.12.002","article-title":"Recovery of elasticity of aged human epithelial cells in vitro","volume":"2","author":"Sokolov","year":"2006","journal-title":"Nanomedicine"},{"key":"ref_290","doi-asserted-by":"crossref","first-page":"259","DOI":"10.4161\/derm.22028","article-title":"Advanced glycation end products: Key players in skin aging?","volume":"4","author":"Gkogkolou","year":"2012","journal-title":"Dermatoendocrinology"},{"key":"ref_291","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1016\/j.exger.2008.04.004","article-title":"Reconstructed skin modified by glycation of the dermal equivalent as a model for skin aging and its potential use to evaluate anti-glycation molecules","volume":"43","author":"Pageon","year":"2008","journal-title":"Exp. Gerontol."},{"key":"ref_292","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1016\/S0047-6374(01)00225-1","article-title":"Molecular mechanisms of ageing in connective tissues","volume":"122","author":"Bailey","year":"2001","journal-title":"Mech. Ageing Dev."},{"key":"ref_293","doi-asserted-by":"crossref","first-page":"39027","DOI":"10.1074\/jbc.M006700200","article-title":"Effect of collagen turnover on the accumulation of advanced glycation end products","volume":"275","author":"Verzijl","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_294","doi-asserted-by":"crossref","first-page":"49","DOI":"10.2337\/diab.41.2.S49","article-title":"Structural and functional changes of laminin and type IV collagen after nonenzymatic glycation","volume":"41","author":"Charonis","year":"1992","journal-title":"Diabetes"},{"key":"ref_295","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1161\/CIRCULATIONAHA.106.621854","article-title":"Advanced glycation end products: Sparking the development of diabetic vascular injury","volume":"114","author":"Goldin","year":"2006","journal-title":"Circulation"},{"key":"ref_296","doi-asserted-by":"crossref","first-page":"1302","DOI":"10.1038\/ki.1997.455","article-title":"Nonenzymatic glycation of type IV collagen and matrix metalloproteinase susceptibility","volume":"52","author":"Mott","year":"1997","journal-title":"Kidney Int."},{"key":"ref_297","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/j.matbio.2013.07.002","article-title":"Differential expression of cathepsins K, S and V between young and aged Caucasian women skin epidermis","volume":"33","author":"Sage","year":"2014","journal-title":"Matrix Biol."},{"key":"ref_298","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1196\/annals.1333.060","article-title":"An in vitro approach to the chronological aging of skin by glycation of the collagen: The biological effect of glycation on the reconstructed skin model","volume":"1043","author":"Pageon","year":"2005","journal-title":"Ann. N.Y. Acad. Sci."},{"key":"ref_299","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1038\/jid.2009.269","article-title":"AGE-modified collagens I and III induce keratinocyte terminal differentiation through AGE receptor CD36: Epidermal-dermal interaction in acquired perforating dermatosis","volume":"130","author":"Fujimoto","year":"2010","journal-title":"J. Investig. Dermatol."},{"key":"ref_300","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1159\/000322087","article-title":"Reactive metabolites and AGE\/RAGE-mediated cellular dysfunction affect the aging process: A mini-review","volume":"57","author":"Fleming","year":"2011","journal-title":"Gerontologia"},{"key":"ref_301","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1007\/s00109-005-0688-7","article-title":"Understanding RAGE, the receptor for advanced glycation end products","volume":"83","author":"Bierhaus","year":"2005","journal-title":"J. Mol. Med."},{"key":"ref_302","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1038\/sj.jid.5700070","article-title":"The receptor for advanced glycation end products is highly expressed in the skin and upregulated by advanced glycation end products and tumor necrosis factor-alpha","volume":"126","author":"Lohwasser","year":"2006","journal-title":"J. Investig. Dermatol."},{"key":"ref_303","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1111\/j.1600-0625.2011.01408.x","article-title":"Involvement of RAGE, MAPK and NF-\u03baB pathways in AGEs-induced MMP-9 activation in HaCaT keratinocytes","volume":"21","author":"Zhu","year":"2012","journal-title":"Exp. Dermatol."},{"key":"ref_304","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1159\/000107621","article-title":"Dermatoporosis: A chronic cutaneous insufficiency\/fragility syndrome","volume":"215","author":"Kaya","year":"2007","journal-title":"Dermatology"},{"key":"ref_305","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1159\/000342344","article-title":"Age-related aspects of cutaneous wound healing: A mini-review","volume":"59","author":"Sgonc","year":"2013","journal-title":"Gerontology"}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/10\/12\/1607\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T10:38:16Z","timestamp":1760179096000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/10\/12\/1607"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,11,27]]},"references-count":305,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2020,12]]}},"alternative-id":["biom10121607"],"URL":"https:\/\/doi.org\/10.3390\/biom10121607","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,11,27]]}}}