{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T22:06:14Z","timestamp":1791324374077,"version":"4.3.1"},"reference-count":67,"publisher":"MDPI AG","issue":"15","license":[{"start":{"date-parts":[[2022,8,5]],"date-time":"2022-08-05T00:00:00Z","timestamp":1659657600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"University of Milan","award":["PSR2020"],"award-info":[{"award-number":["PSR2020"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Duchenne muscular dystrophy (DMD) is a rare genetic disease leading to progressive muscle wasting, respiratory failure, and cardiomyopathy. Although muscle fibrosis represents a DMD hallmark, the organisation of the extracellular matrix and the molecular changes in its turnover are still not fully understood. To define the architectural changes over time in muscle fibrosis, we used an mdx mouse model of DMD and analysed collagen and glycosaminoglycans\/proteoglycans content in skeletal muscle sections at different time points during disease progression and in comparison with age-matched controls. Collagen significantly increased particularly in the diaphragm, quadriceps, and gastrocnemius in adult mdx, with fibrosis significantly correlating with muscle degeneration. We also analysed collagen turnover pathways underlying fibrosis development in cultured primary quadriceps-derived fibroblasts. Collagen secretion and matrix metalloproteinases (MMPs) remained unaffected in both young and adult mdx compared to wt fibroblasts, whereas collagen cross-linking and tissue inhibitors of MMP (TIMP) expression significantly increased. We conclude that, in the DMD model we used, fibrosis mostly affects diaphragm and quadriceps with a higher collagen cross-linking and inhibition of MMPs that contribute differently to progressive collagen accumulation during fibrotic remodelling. This study offers a comprehensive histological and molecular characterisation of DMD-associated muscle fibrosis; it may thus provide new targets for tailored therapeutic interventions.<\/jats:p>","DOI":"10.3390\/ijms23158735","type":"journal-article","created":{"date-parts":[[2022,8,7]],"date-time":"2022-08-07T22:51:46Z","timestamp":1659912706000},"page":"8735","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":33,"title":["Characterisation of Progressive Skeletal Muscle Fibrosis in the Mdx Mouse Model of Duchenne Muscular Dystrophy: An In Vivo and In Vitro Study"],"prefix":"10.3390","volume":"23","author":[{"given":"Matteo","family":"Giovarelli","sequence":"first","affiliation":[{"name":"Department of Biomedical and Clinical Sciences, Universit\u00e0 degli Studi di Milano, via G. B. Grassi 74, 20157 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2041-9928","authenticated-orcid":false,"given":"Francesca","family":"Arnaboldi","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, Universit\u00e0 degli Studi di Milano, via Mangiagalli 31, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Silvia","family":"Zecchini","sequence":"additional","affiliation":[{"name":"Department of Biomedical and Clinical Sciences, Universit\u00e0 degli Studi di Milano, via G. B. Grassi 74, 20157 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6871-5059","authenticated-orcid":false,"given":"Laura Brigida","family":"Cornaghi","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, Universit\u00e0 degli Studi di Milano, via Mangiagalli 31, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9703-5303","authenticated-orcid":false,"given":"Ambra","family":"Nava","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, Universit\u00e0 degli Studi di Milano, via Mangiagalli 31, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7622-0996","authenticated-orcid":false,"given":"Michele","family":"Sommariva","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences for Health, Universit\u00e0 degli Studi di Milano, via Mangiagalli 31, 20133 Milan, Italy"},{"name":"Molecular Targeting Unit, Department of Research, Fondazione IRCCS Istituto Nazionale dei Tumori, via Amadeo 42, 20133 Milan, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emilio Giuseppe Ignazio","family":"Clementi","sequence":"additional","affiliation":[{"name":"Department of Biomedical and Clinical Sciences, Universit\u00e0 degli Studi di Milano, via G. 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Rev."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1002\/mus.22094","article-title":"Structure and function of the skeletal muscle extracellular matrix","volume":"44","author":"Gillies","year":"2011","journal-title":"Muscle Nerve"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"4089","DOI":"10.1111\/febs.12171","article-title":"Enter the matrix: Shape, signal and super-highway","volume":"280","author":"Lund","year":"2013","journal-title":"FEBS J."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1111\/j.1600-0838.2005.00464.x","article-title":"Enzymic and non-enzymic cross-linking mecha-nisms in relation to turnover of collagen: Relevance to aging and exercise","volume":"15","author":"Avery","year":"2005","journal-title":"Scand. J. Med. Sci. Sports"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1113\/jphysiol.2010.201921","article-title":"Lateral transmission of force is impaired in skeletal muscles of dystrophic mice and very old rats","volume":"589","author":"Ramaswamy","year":"2011","journal-title":"J. Physiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1007\/s12178-015-9267-x","article-title":"Normal muscle structure, growth, development, and regeneration","volume":"8","author":"Oliveira","year":"2015","journal-title":"Curr. Rev. Musculosklet. Med."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1016\/j.jbmt.2012.06.001","article-title":"The muscular force transmission system: Role of the intramuscular connective tissue","volume":"17","author":"Turrina","year":"2013","journal-title":"J. Bodyw. Mov. Ther."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"12599","DOI":"10.1074\/jbc.R300005200","article-title":"Skeletal muscle basement membrane sarcolemma-cytoskeleton interaction minireview series","volume":"278","author":"Campbell","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1097\/00003677-200201000-00005","article-title":"Intramuscular extracellular matrix: Complex environment of muscle cells","volume":"30","author":"Kovanen","year":"2002","journal-title":"Exerc. Sport Sci. Rev."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"12601","DOI":"10.1074\/jbc.R200027200","article-title":"The basement membrane\/basal lamina of skeletal muscle","volume":"278","author":"Sanes","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"C241","DOI":"10.1152\/ajpcell.00173.2013","article-title":"Cellular mechanisms of tissue fibrosis. 4. Structural and functional consequences of skeletal muscle fibrosis","volume":"305","author":"Lieber","year":"2013","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1007\/s00441-018-2955-2","article-title":"Skeletal muscle fibrosis: An overview","volume":"375","author":"Mahdy","year":"2019","journal-title":"Cell Tissue Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"2304","DOI":"10.1007\/s00018-006-6149-9","article-title":"Lysyl oxidase: An oxidative enzyme and effector of cell function","volume":"63","author":"Lucero","year":"2006","journal-title":"Cell Mol. Life Sci."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1016\/j.matbio.2004.11.002","article-title":"Tissue-specific expression and regulation of the alternatively-spliced forms of lysyl hydroxylase 2 (LH2) in human kidney cells and skin fibroblasts","volume":"23","author":"Walker","year":"2005","journal-title":"Matrix Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3109\/03008207.2014.947369","article-title":"Extracellular matrix regulation in the muscle satellite cell niche","volume":"56","author":"Thomas","year":"2015","journal-title":"Connect. Tissue Res."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1078","DOI":"10.1126\/science.1191035","article-title":"Substrate elasticity regulates skeletal muscle stem cell self-renewal in culture","volume":"329","author":"Gilbert","year":"2010","journal-title":"Science"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"E900","DOI":"10.1152\/ajpendo.00444.2005","article-title":"Effects of streptozotocin-induced diabetes and physical training on gene expression of extracellular matrix proteins in mouse skeletal muscle","volume":"290","author":"Lehti","year":"2006","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"740","DOI":"10.1002\/mus.26845","article-title":"Conference report on contractures in musculoskeletal and neurological conditions","volume":"61","author":"Nuckolls","year":"2020","journal-title":"Muscle Nerve"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1080\/03008207.2019.1694011","article-title":"Contribution of extracellular matrix components to the stiffness of skeletal muscle contractures in patients with cerebral palsy","volume":"62","author":"Smith","year":"2021","journal-title":"Connect. Tissue Res."},{"key":"ref_20","unstructured":"Engel, A.G., and Franzini-Armstrong, C. (1994). Myology, McGraw-Hill. [2nd ed.]."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"332","DOI":"10.1034\/j.1600-0838.2000.010006332.x","article-title":"Relation between myofibers and connective tissue during muscle injury repair","volume":"10","author":"Kaariainen","year":"2000","journal-title":"Scand. J. Med. Sci. Sports"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1397","DOI":"10.1091\/mbc.e06-08-0693","article-title":"Muscle satellite cells and endothelial cells: Close neighbors and privileged partners","volume":"18","author":"Christov","year":"2007","journal-title":"Mol. Biol. Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1016\/0092-8674(87)90579-4","article-title":"Dystrophin: The protein product of the Duchenne muscular dystrophy locus","volume":"51","author":"Hoffman","year":"1987","journal-title":"Cell"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/0960-8966(91)90039-U","article-title":"Population frequencies of inherited neuromuscular diseases\u2014A world survey","volume":"1","author":"Emery","year":"1991","journal-title":"Neuromuscul. Disord."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1016\/j.nmd.2014.03.008","article-title":"A systematic review and meta-analysis on the epidemiology of Duchenne and Becker muscular dystrophy","volume":"24","author":"Mah","year":"2014","journal-title":"Neuromuscul. Disord."},{"key":"ref_26","first-page":"210797","article-title":"The dystrophin-glycoprotein complex in the prevention of muscle damage","volume":"2011","author":"Gumerson","year":"2011","journal-title":"J. Biomed. Biotechnol."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Emery, A.E.H., Muntoni, F., and Quinlivan, R.C.M. (2015). Duchenne Muscular Dystrophy, OUP Oxford.","DOI":"10.1093\/med\/9780199681488.001.0001"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.pediatrneurol.2006.09.016","article-title":"Pathophysiology of duchenne muscular dystrophy: Current hypotheses","volume":"36","author":"Deconinck","year":"2007","journal-title":"Pediatr. Neurol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Kharraz, Y., Guerra, J., Pessina, P., Serrano, A.L., and Mu\u00f1oz-C\u00e1noves, P. (2014). Understanding the process of fibrosis in Duchenne muscular dystrophy. Biomed. Res. Int., 2014.","DOI":"10.1155\/2014\/965631"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1080\/14656566.2020.1732350","article-title":"New and emerging pharmacotherapy for duchenne muscular dystrophy: A focus on synthetic therapeutics","volume":"21","author":"Grages","year":"2020","journal-title":"Expert Opin. Pharmacother."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1038\/s41434-020-0156-6","article-title":"Optimization of antisense-mediated exon skipping for Duchenne muscular dystrophy","volume":"27","author":"Dzierlega","year":"2020","journal-title":"Gene Ther."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1189","DOI":"10.1073\/pnas.81.4.1189","article-title":"X chromosome-linked muscular dystrophy","volume":"81","author":"Bulfield","year":"1984","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1242\/dmm.018424","article-title":"Animal models of Duchenne muscular dystrophy: From basic mechanisms to gene therapy","volume":"8","author":"McGreevy","year":"2015","journal-title":"Dis. Models Mech."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4177","DOI":"10.1111\/febs.12267","article-title":"The mdx mouse model as a surrogate for Duchenne muscular dystrophy","volume":"280","author":"Partridge","year":"2013","journal-title":"FEBS J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1089\/hum.2019.255","article-title":"Dose-escalation study of systemically selivered rAAVrh74.MHCK7.micro-dystrophin in the mdx mouse model of Duchenne Muscular Dystrophy","volume":"32","author":"Potter","year":"2021","journal-title":"Hum. Gene Ther."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1186\/2044-5040-1-21","article-title":"Aberrant repair and fibrosis development in skeletal muscle","volume":"1","author":"Mann","year":"2011","journal-title":"Skelet. Muscle"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"105751","DOI":"10.1016\/j.phrs.2021.105751","article-title":"Givinostat as metabolic enhancer reverting mitochondrial biogenesis deficit in Duchenne Muscular Dystrophy","volume":"170","author":"Giovarelli","year":"2021","journal-title":"Pharmacol. Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1139\/Y09-058","article-title":"Calcium and the damage pathways in muscular dystrophy","volume":"88","author":"Allen","year":"2010","journal-title":"Can. J. Physiol. Pharmacol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1007\/BF00314217","article-title":"Muscle hypertrophy in duchenne muscular dystrophy","volume":"236","author":"Cros","year":"1989","journal-title":"J. Neurol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1007\/s00441-009-0889-4","article-title":"Altered production of extracellular matrix components by muscle-derived Duchenne muscular dystrophy fibroblasts before and after TGF- beta1 treatment","volume":"339","author":"Zanotti","year":"2010","journal-title":"Cell Tissue Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"38371","DOI":"10.1038\/srep38371","article-title":"Impaired regenerative capacity and lower revertant fibre expansion in dystrophin-deficient mdx muscles on DBA\/2 background","volume":"6","author":"Rodrigues","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/0022-510X(93)90130-Q","article-title":"Age-related changes in replication of myogenic cells in rndx mice: Quantitative autoradiographic studies","volume":"119","author":"McGeachie","year":"1993","journal-title":"J. Neurol. Sci."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1096\/fj.03-1024com","article-title":"Anti-TNFalpha (Remicade) therapy protects dystrophic skeletal muscle from necrosis","volume":"18","author":"Grounds","year":"2004","journal-title":"FASEB J."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1016\/j.expneurol.2005.11.021","article-title":"Nuclear factor kappa-B blockade reduces skeletal muscle degeneration and enhances muscle function in Mdx mice","volume":"198","author":"Messina","year":"2006","journal-title":"Exp. Neurol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1002\/ar.1092420109","article-title":"Phenotype of dystrophinopathy in old mdx mice","volume":"242","author":"Lefaucheur","year":"1995","journal-title":"Anat. Rec."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1038\/mt.2008.264","article-title":"Cardiac expression of a mini-dystrophin that normalizes skeletal muscle force only partially restores heart function in aged Mdx mice","volume":"17","author":"Bostick","year":"2009","journal-title":"Mol. Ther."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"536","DOI":"10.1038\/352536a0","article-title":"The mdx mouse diaphragm reproduces the degenerative changes of Duchenne muscular dystrophy","volume":"352","author":"Stedman","year":"1991","journal-title":"Nature"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"820","DOI":"10.1002\/mus.20523","article-title":"Pentoxifylline fails to attenuate fibrosis in dystrophic (mdx) diaphragm muscle","volume":"33","author":"Gosselin","year":"2006","journal-title":"Muscle Nerve"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1152\/japplphysiol.00248.2021","article-title":"Diaphragm muscle fibrosis involves changes in collagen organization with mechanical implications in Duchenne muscular dystrophy","volume":"132","author":"Sahani","year":"2022","journal-title":"J. Appl. Physiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"7683","DOI":"10.1073\/pnas.83.20.7683","article-title":"Primary structure of an extracellular matrix proteoglycan core protein deduced from cloned cDNA","volume":"83","author":"Krusius","year":"1986","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"963","DOI":"10.1369\/0022155412456380","article-title":"Biglycan: A multivalent proteoglycan providing structure and signals","volume":"60","author":"Nastase","year":"2012","journal-title":"J. Histochem. Cytochem."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1083\/jcb.148.4.801","article-title":"The small leucine-rich repeat proteoglycan biglycan binds to alpha-dystroglycan and is upregulated in dystrophic muscle","volume":"148","author":"Bowe","year":"2000","journal-title":"J. Cell Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1002\/jcp.20740","article-title":"Biglycan binds to alpha- and gamma-sarcoglycan and regulates their expression during development","volume":"20","author":"Rafii","year":"2006","journal-title":"J. Cell Physiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"15000","DOI":"10.1073\/pnas.192571199","article-title":"Gene expression comparison of biopsies from Duchenne muscular dystrophy (DMD) and normal skeletal muscle","volume":"99","author":"Haslett","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2546","DOI":"10.1093\/brain\/awh635","article-title":"Decorin and biglycan expression is differentially altered in several muscular dystrophies","volume":"128","author":"Zanotti","year":"2005","journal-title":"Brain"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1111\/j.1582-4934.2006.tb00435.x","article-title":"Increase in decorin and biglycan in Duchenne Muscular Dystrophy: Role of fibroblasts as cell source of these proteoglycans in the disease","volume":"10","author":"Fadic","year":"2006","journal-title":"J. Cell Mol. Med."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1186\/s13578-021-00579-4","article-title":"Extracellular matrix: An important regulator of cell functions and skeletal muscle development","volume":"11","author":"Zhang","year":"2021","journal-title":"Cell Biosci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.stem.2016.09.010","article-title":"Myogenic progenitor cells control extracellular matrix production by fibroblasts during skeletal muscle hypertrophy","volume":"20","author":"Fry","year":"2017","journal-title":"Cell Stem Cell"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1021\/bi00854a021","article-title":"Some properties of the products of reaction of tadpole collagenase with collagen","volume":"6","author":"Sakai","year":"1967","journal-title":"Biochemistry"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1111\/j.1749-6632.1994.tb24722.x","article-title":"Regulation of matrix metalloproteinase activity","volume":"732","author":"Murphy","year":"1994","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_61","unstructured":"(2010, November 01). DMD_M.2.2.006 Whole Body Tension Measurements. Available online: http:\/\/www.treat-nmd.eu\/downloads\/file\/sops\/dmd\/MDX\/dmd_m.2.2.006.pdf."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1152\/japplphysiol.90985.2008","article-title":"Multiple pathological events in exercised dystrophic mdx mice are targeted by pentoxifylline: Outcome of a large array of in vivo and ex vivo tests","volume":"106","author":"Burdi","year":"2009","journal-title":"J. Appl. Physiol."},{"key":"ref_63","unstructured":"(2015, May 23). DMD_M.2.1.003 Use of Treadmill and Wheel Exercise to Assess Dystrophic State. Available online: https:\/\/treat-nmd.org\/wp-content\/uploads\/2016\/08\/MDX-DMD_M.2.1.003-34.pdf."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1007\/BF01002772","article-title":"Picrosirius staining plus polarization microscopy, a specific method for collagen detection in tissue sections","volume":"11","author":"Junqueira","year":"1979","journal-title":"Histochem. J."},{"key":"ref_65","first-page":"88","article-title":"Picropolychrome: Histological staining technic intended for the study of normal and pathological connective tissue","volume":"8","author":"Herovici","year":"1963","journal-title":"Rev. Fr. D\u2019etudes Clin. Biol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1007\/BF00427227","article-title":"Differential staining of collagen types in paraffin sections: A color change in degraded forms","volume":"386","author":"Lillie","year":"1980","journal-title":"Virchows Arch. A Pathol. Anat. Histol."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Randelli, F., Menon, A., Giai Via, A., Mazzoleni, M.G., Sciancalepore, F., Brioschi, M., and Gagliano, N. (2018). Effect of a Collagen-Based Compound on Morpho-Functional Properties of Cultured Human Tenocytes. Cells, 7.","DOI":"10.3390\/cells7120246"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/15\/8735\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T00:04:49Z","timestamp":1760141089000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/15\/8735"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,8,5]]},"references-count":67,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2022,8]]}},"alternative-id":["ijms23158735"],"URL":"https:\/\/doi.org\/10.3390\/ijms23158735","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,8,5]]}}}