{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T16:35:44Z","timestamp":1786725344620,"version":"build-2736575974"},"reference-count":200,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T00:00:00Z","timestamp":1786665600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Ministry of Science and Higher Education of the Republic of Kazakhstan","award":["AP23487182"],"award-info":[{"award-number":["AP23487182"]}]},{"award":["AP23487182"],"award-info":[{"award-number":["AP23487182"]}],"id":[{"id":"https:\/\/ror.org\/05tyne317","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>The extracellular matrix (ECM) in cancer differs from healthy tissue in structure, composition, and mechanical properties. Matricellular proteins (MCPs) play important roles in shaping ECM architecture during tissue remodeling. This narrative review, combined with a bioinformatics analysis, examines six major MCP families\u2014Fasciclins, Tenascins, Thrombospondins, Small Leucine-Rich Proteoglycans, the SPARC family, and the CCN family\u2014through a mechanobiological lens in bladder cancer. It summarizes current knowledge on the mechanical regulation of MCP expression, their effects on matrix stiffness, and their contributions to bladder cancer progression. Analyses of public datasets reveal that stromal cells are the predominant source of MCPs in the tumor microenvironment. Furthermore, mechanical upregulation and involvement in the formation of stiff ECM highlight MCPs as important players in a mechanotransduction feedback loop. While most MCPs exert pro-tumorigenic effects on bladder cancer cells, several display context-dependent or anti-tumorigenic activities. Existing studies have primarily focused on the isolated effects of MCPs on bladder cancer cell lines in two-dimensional systems or simple subcutaneous xenograft models. Both approaches fail to capture the context-dependent nature of MCPs and their involvement in ECM formation. These findings underscore the need for future studies to investigate the complex effects of MCPs on bladder cancer progression.<\/jats:p>","DOI":"10.3390\/biom16081191","type":"journal-article","created":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T15:48:36Z","timestamp":1786722516000},"page":"1191","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Mechanobiology of Matricellular Proteins in Bladder Cancer: A Narrative Review and Bioinformatics Analysis"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5905-8520","authenticated-orcid":false,"given":"Alim","family":"Turgaliyev","sequence":"first","affiliation":[{"name":"Department of Biomedical Sciences, School of Medicine, Nazarbayev University, Astana 010000, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-7950-7424","authenticated-orcid":false,"given":"Roman","family":"Konovalov","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, School of Medicine, Nazarbayev University, Astana 010000, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-0149-8680","authenticated-orcid":false,"given":"Anton","family":"Borissenko","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, School of Medicine, Nazarbayev University, Astana 010000, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4206-5529","authenticated-orcid":false,"given":"Dieter","family":"Riethmacher","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, School of Medicine, Nazarbayev University, Astana 010000, Kazakhstan"},{"name":"Human Development and Health, Faculty of Medicine, University of Southampton, Southampton SO16 6YD, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,14]]},"reference":[{"key":"ref_1","first-page":"e27330","article-title":"A Review of the Etiology and Epidemiology of Bladder Cancer: All You Need To Know","volume":"14","author":"Halaseh","year":"2022","journal-title":"Cureus"},{"key":"ref_2","first-page":"229","article-title":"Global cancer statistics 2022: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries","volume":"74","author":"Bray","year":"2024","journal-title":"CA Cancer J. Clin."},{"key":"ref_3","first-page":"51","article-title":"The Real-World Lifetime Economic Burden of Urothelial Carcinoma by Stage at Diagnosis","volume":"6","author":"Aly","year":"2020","journal-title":"J. Clin. Pathw."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/j.euf.2023.07.002","article-title":"Repeat Transurethral Resection for Non\u2013muscle-invasive Bladder Cancer: An Updated Systematic Review and Meta-analysis in the Contemporary Era","volume":"10","author":"Yanagisawa","year":"2024","journal-title":"Eur. Urol. Focus"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Bornstein, P., McPherson, J., and Sage, H. (1982). Synthesis and Secretion of Structural Macromolecules by Endothelial Cells in Culture. Pathobiology of the Endothelial Cell, Elsevier.","DOI":"10.1016\/B978-0-12-521980-8.50020-1"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1038\/nrm3904","article-title":"Remodelling the extracellular matrix in development and disease","volume":"15","author":"Bonnans","year":"2014","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"100515","DOI":"10.1016\/j.xcrp.2021.100515","article-title":"The mechanics of fibrillar collagen extracellular matrix","volume":"2","author":"Sun","year":"2021","journal-title":"Cell Rep. Phys. Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1007\/BF00266843","article-title":"Are the polarization colors of Picrosirius red-stained collagen determined only by the diameter of the fibers?","volume":"93","author":"Dayan","year":"1989","journal-title":"Histochemistry"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"930","DOI":"10.1055\/s-0042-1743145","article-title":"Polarized Microscopic Analysis of Picrosirius Red Stained Salivary Gland Pathologies: An Observational Study","volume":"16","author":"Juengsomjit","year":"2022","journal-title":"Eur. J. Dent."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Popova, N.v., and J\u00fccker, M. (2022). The Functional Role of Extracellular Matrix Proteins in Cancer. Cancers, 14.","DOI":"10.3390\/cancers14010238"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"3852","DOI":"10.1016\/j.cell.2021.06.024","article-title":"Fibroblasts: Origins, definitions, and functions in health and disease","volume":"184","author":"Plikus","year":"2021","journal-title":"Cell"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1038\/s41419-024-06697-4","article-title":"Modulating extracellular matrix stiffness: A strategic approach to boost cancer immunotherapy","volume":"15","author":"Mai","year":"2024","journal-title":"Cell Death Dis."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"36128","DOI":"10.1038\/srep36128","article-title":"Linearized texture of three-dimensional extracellular matrix is mandatory for bladder cancer cell invasion","volume":"6","author":"Alfano","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"887","DOI":"10.1038\/s41419-025-08222-7","article-title":"Matrix stiffness maintains bladder cancer stemness via integrin-nuclear skeleton axis","volume":"16","author":"Tao","year":"2025","journal-title":"Cell Death Dis."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1999","DOI":"10.1002\/jum.16216","article-title":"Contrast-enhanced Ultrasound Combined With Elastography for the Evaluation of Muscle-invasive Bladder Cancer in Rats","volume":"42","author":"Hu","year":"2023","journal-title":"J. Ultrasound Med."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1002\/jum.15381","article-title":"Shear Wave Elasticity Differentiation Between Low- and High-Grade Bladder Urothelial Carcinoma and Correlation With Collagen Fiber Content","volume":"40","author":"Huang","year":"2021","journal-title":"J. Ultrasound Med."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1111\/nyas.14358","article-title":"Tissue stiffness contributes to YAP activation in bladder cancer patients undergoing transurethral resection","volume":"1473","author":"Ghasemi","year":"2020","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Monteiro-Reis, S., Ferreira, J.P.S., Pires, R.A., Lobo, J., Carvalho, J.A., Reis, R.L., Jorge, R.N., and Jer\u00f3nimo, C. (2023). Bladder Wall Stiffness after Cystectomy in Bladder Cancer Patients: A Preliminary Study. Cancers, 15.","DOI":"10.3390\/cancers15020359"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1177\/1535370214560973","article-title":"Metastatic bladder cancer cells distinctively sense and respond to physical cues of collagen fibril-mimetic nanotopography","volume":"240","author":"Iuliano","year":"2015","journal-title":"Exp. Biol. Med."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"100154","DOI":"10.1016\/j.mbplus.2024.100154","article-title":"Topographic modification of the extracellular matrix precedes the onset of bladder cancer","volume":"23","author":"Venegoni","year":"2024","journal-title":"Matrix Biol. Plus"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1083\/jcb.130.3.503","article-title":"Diversity of function is inherent in matricellular proteins: An appraisal of thrombospondin 1","volume":"130","author":"Bornstein","year":"1995","journal-title":"J. Cell Biol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/s12079-009-0069-z","article-title":"Matricellular proteins: An overview","volume":"3","author":"Bornstein","year":"2009","journal-title":"J. Cell Commun. Signal."},{"key":"ref_23","first-page":"1","article-title":"Revisiting the matricellular concept","volume":"37","author":"Sage","year":"2014","journal-title":"Matrix Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2705","DOI":"10.1158\/0008-5472.CAN-18-2098","article-title":"The Matrix Revolution: Matricellular Proteins and Restructuring of the Cancer Microenvironment","volume":"80","author":"Gerarduzzi","year":"2020","journal-title":"Cancer Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3043","DOI":"10.1016\/j.bbamcr.2015.05.007","article-title":"A guide to mechanobiology: Where biology and physics meet","volume":"1853","author":"Jansen","year":"2015","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"4866","DOI":"10.1038\/s41467-024-49230-z","article-title":"A YAP-centered mechanotransduction loop drives collective breast cancer cell invasion","volume":"15","author":"Khalil","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1038\/ncb2756","article-title":"Mechanotransduction and YAP-dependent matrix remodelling is required for the generation and maintenance of cancer-associated fibroblasts","volume":"15","author":"Calvo","year":"2013","journal-title":"Nat. Cell Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2506","DOI":"10.1038\/s41598-017-02689-x","article-title":"Substrate Rigidity Controls Activation and Durotaxis in Pancreatic Stellate Cells","volume":"7","author":"Lachowski","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2302327","DOI":"10.1002\/advs.202302327","article-title":"A Hierarchical Mechanotransduction System: From Macro to Micro","volume":"11","author":"Cao","year":"2024","journal-title":"Adv. Sci."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Rosselli-Murai, L.K., Almeida, L.O., Zagni, C., Galindo-Moreno, P., Padial-Molina, M., Volk, S.L., Murai, M.J., Rios, H.F., Squarize, C.H., and Castilho, R.M. (2013). Periostin Responds to Mechanical Stress and Tension by Activating the MTOR Signaling Pathway. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0083580"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"9896","DOI":"10.1073\/pnas.1919702117","article-title":"Matrix mechanotransduction mediated by thrombospondin-1\/integrin\/YAP in the vascular remodeling","volume":"117","author":"Yamashiro","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e22726","DOI":"10.1096\/fj.202200917RR","article-title":"Periostin increased by mechanical stress upregulates interleukin-6 expression in the ligamentum flavum","volume":"37","author":"Yabu","year":"2023","journal-title":"FASEB J."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1016\/j.ydbio.2012.11.023","article-title":"Fluid flow forces and rhoA regulate fibrous development of the atrioventricular valves","volume":"374","author":"Tan","year":"2013","journal-title":"Dev. Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1161\/01.HYP.0000074905.22908.A6","article-title":"Effects of Pressure Overload on Extracellular Matrix Expression in the Heart of the Atrial Natriuretic Peptide\u2013Null Mouse","volume":"42","author":"Wang","year":"2003","journal-title":"Hypertension"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1002\/cbin.11116","article-title":"Periostin plays role in force-induced stem cell potential by periodontal ligament stem cells","volume":"43","author":"Panchamanon","year":"2019","journal-title":"Cell Biol. Int."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.matbio.2020.07.002","article-title":"Periostin and matrix stiffness combine to regulate myofibroblast differentiation and fibronectin synthesis during palatal healing","volume":"94","author":"Nikoloudaki","year":"2020","journal-title":"Matrix Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1016\/S0003-9969(03)00103-1","article-title":"Mechanical stimuli enhances the expression of RGD-CAP\/\u03b2ig-h3 in the periodontal ligament","volume":"48","author":"Doi","year":"2003","journal-title":"Arch. Oral Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/j.archoralbio.2014.11.019","article-title":"Cyclic stretch induced gene expression of extracellular matrix and adhesion molecules in human periodontal ligament cells","volume":"60","author":"Ma","year":"2015","journal-title":"Arch. Oral Biol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1080\/03008200500346111","article-title":"Mechanical Regulation of Terminal Chondrocyte Differentiation via RGD-CAP\/\u03b2ig-h3 Induced by TGF-\u03b2","volume":"46","author":"Ohno","year":"2005","journal-title":"Connect. Tissue Res."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.bbamcr.2004.08.001","article-title":"Induction of tenascin-C by cyclic tensile strain versus growth factors: Distinct contributions by Rho\/ROCK and MAPK signaling pathways","volume":"1693","author":"Chiquet","year":"2004","journal-title":"Biochim. Biophys. Acta (BBA) Mol. Cell Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1150","DOI":"10.1016\/j.bbamcr.2008.01.013","article-title":"Tenascin-C induction by cyclic strain requires integrin-linked kinase","volume":"1783","author":"Maier","year":"2008","journal-title":"Biochim. Biophys. Acta (BBA) Mol. Cell Res."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"21840","DOI":"10.1074\/jbc.274.31.21840","article-title":"Induction of Tenascin-C in Cardiac Myocytes by Mechanical Deformation","volume":"274","author":"Yamamoto","year":"1999","journal-title":"J. Biol. Chem."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2857","DOI":"10.1167\/iovs.05-0075","article-title":"Changes in Gene Expression by Trabecular Meshwork Cells in Response to Mechanical Stretching","volume":"46","author":"Vittal","year":"2005","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"3354","DOI":"10.1002\/jcb.23265","article-title":"Regulation of tenascin expression in bone","volume":"112","author":"Morgan","year":"2011","journal-title":"J. Cell. Biochem."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1647","DOI":"10.1161\/CIRCRESAHA.121.320694","article-title":"Tenascin-X Mediates Flow-Induced Suppression of EndMT and Atherosclerosis","volume":"130","author":"Liang","year":"2022","journal-title":"Circ. Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"F577","DOI":"10.1152\/ajprenal.00393.2004","article-title":"Mechanical strain increases SPARC levels in podocytes: Implications for glomerulosclerosis","volume":"289","author":"Durvasula","year":"2005","journal-title":"Am. J. Physiol.-Ren. Physiol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Fouladgar, F., Powell, R., Carney, E., Martinez, A.E., Jafari, A., and Habibi, N. (2026). Biomechanical Stimulation of Mesenchymal Stem Cells in 3D Peptide Nanofibers for Bone Differentiation. J. Funct. Biomater., 17.","DOI":"10.3390\/jfb17010052"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"015018","DOI":"10.1088\/1758-5090\/ae2cf2","article-title":"An in vitro model of stiffened colonic mucosa exhibits altered epithelial behavior","volume":"18","author":"Massaro","year":"2026","journal-title":"Biofabrication"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.bbrc.2023.04.058","article-title":"CCN1 expression is regulated by mechanical stimuli in tendons","volume":"663","author":"Leong","year":"2023","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"L825","DOI":"10.1152\/ajplung.00073.2020","article-title":"Alveolar CCN1 is associated with mechanical stretch and acute respiratory distress syndrome severity","volume":"319","author":"Morrell","year":"2020","journal-title":"Am. J. Physiol. Lung Cell. Mol. Physiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1007\/s11596-018-1929-7","article-title":"Stretch-induced Expression of CYR61 Increases the Secretion of IL-8 in A549 Cells via the NF-\u03ba\u03b2\/l\u03ba\u03b2 Pathway","volume":"38","author":"Zhang","year":"2018","journal-title":"Curr. Med. Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"C1524","DOI":"10.1152\/ajpcell.2001.281.5.C1524","article-title":"Regulation of Cyr61 gene expression by mechanical stretch through multiple signaling pathways","volume":"281","author":"Tamura","year":"2001","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"23125","DOI":"10.1074\/jbc.M109.019059","article-title":"Mechanical Regulation of the Proangiogenic Factor CCN1\/CYR61 Gene Requires the Combined Activities of MRTF-A and CREB-binding Protein Histone Acetyltransferase","volume":"284","author":"Hanna","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2000051","DOI":"10.1002\/adbi.202000051","article-title":"Differential Impact of Fluid Shear Stress and YAP\/TAZ on BMP\/TGF-\u03b2 Induced Osteogenic Target Genes","volume":"5","author":"Reichenbach","year":"2021","journal-title":"Adv. Biol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1508","DOI":"10.1016\/j.jbiomech.2013.03.028","article-title":"Tensile strain increases expression of CCN2 and COL2A1 by activating TGF-\u03b2-Smad2\/3 pathway in chondrocytic cells","volume":"46","author":"Furumatsu","year":"2013","journal-title":"J. Biomech."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.bbrc.2011.04.138","article-title":"Mechanical stretch increases CCN2\/CTGF expression in anterior cruciate ligament-derived cells","volume":"409","author":"Miyake","year":"2011","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1217","DOI":"10.1161\/CIRCULATIONAHA.116.026361","article-title":"Disturbed Flow Promotes Arterial Stiffening Through Thrombospondin-1","volume":"136","author":"Kim","year":"2017","journal-title":"Circulation"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"101533","DOI":"10.1016\/j.tranon.2022.101533","article-title":"Thrombospondin-1 overexpression stimulates loss of Smad4 and accelerates malignant behavior via TGF-\u03b2 signal activation in pancreatic ductal adenocarcinoma","volume":"26","author":"Matsumura","year":"2022","journal-title":"Transl. Oncol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1068","DOI":"10.1038\/s12276-021-00636-5","article-title":"WISP-1 induced by mechanical stress contributes to fibrosis and hypertrophy of the ligamentum flavum through Hedgehog-Gli1 signaling","volume":"53","author":"Sun","year":"2021","journal-title":"Exp. Mol. Med."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"3615","DOI":"10.1002\/jcp.26222","article-title":"Comprehensive transcriptome analysis of fluid shear stress altered gene expression in renal epithelial cells","volume":"233","author":"Kunnen","year":"2018","journal-title":"J. Cell. Physiol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1177\/205873920601900104","article-title":"Shear Stress Modulates the Expression of Thrombospondin-1 and CD36 in Endothelial Cells in vitro and during Shear Stress-Induced Angiogenesis in vivo","volume":"19","author":"Bongrazio","year":"2006","journal-title":"Int. J. Immunopathol. Pharmacol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1097\/HJH.0000000000001216","article-title":"Shear stress-mediated upregulation of GTP cyclohydrolase\/tetrahydrobiopterin pathway ameliorates hypertension-related decline in reendothelialization capacity of endothelial progenitor cells","volume":"35","author":"Bai","year":"2017","journal-title":"J. Hypertens."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"121185","DOI":"10.1016\/j.biomaterials.2021.121185","article-title":"ECM stiffness-tuned exosomes drive breast cancer motility through thrombospondin-1","volume":"279","author":"Patwardhan","year":"2021","journal-title":"Biomaterials"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"109624","DOI":"10.1016\/j.abb.2023.109624","article-title":"ECM stiffness-regulated exosomal thrombospondin-1 promotes tunneling nanotubes-based cellular networking in breast cancer cells","volume":"742","author":"Mahadik","year":"2023","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"e71275","DOI":"10.1002\/hsr2.71275","article-title":"Optimal Mechanical Stretch Promotes TSP-1 Expression Through Akt and GSK-3\u03b2\/\u03b2-Catenin Signaling Pathways in Keloid Formation","volume":"8","author":"Xu","year":"2025","journal-title":"Health Sci. Rep."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.matbio.2024.01.005","article-title":"Thrombospondin-1 promotes mechanical stress-mediated ligamentum flavum hypertrophy through the TGF\u03b21\/Smad3 signaling pathway","volume":"127","author":"Zhao","year":"2024","journal-title":"Matrix Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1165\/rcmb.2007-0283OC","article-title":"Cyclic Stretch Affects Pulmonary Endothelial Cell Control of Pulmonary Smooth Muscle Cell Growth","volume":"39","author":"Ochoa","year":"2008","journal-title":"Am. J. Respir. Cell Mol. Biol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"777","DOI":"10.2353\/ajpath.2007.070056","article-title":"A Thrombospondin-1 Antagonist of Transforming Growth Factor-\u03b2 Activation Blocks Cardiomyopathy in Rats with Diabetes and Elevated Angiotensin II","volume":"171","author":"Belmadani","year":"2007","journal-title":"Am. J. Pathol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"2103343","DOI":"10.1002\/advs.202103343","article-title":"Periosteal CD68 + F4\/80 + Macrophages Are Mechanosensitive for Cortical Bone Formation by Secretion and Activation of TGF-\u03b21","volume":"9","author":"Deng","year":"2022","journal-title":"Adv. Sci."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1007\/s13770-023-00548-7","article-title":"Thrombospondin-2 Couples Pressure-Promoted Chondrogenesis through NF-\u03ba\u03b2 Signaling","volume":"20","author":"Niu","year":"2023","journal-title":"Tissue Eng. Regen. Med."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Wang, S., Hu, T., Wang, Z., Li, N., Zhou, L., Liao, L., Wang, M., Liao, L., Wang, H., and Zeng, L. (2017). Macroglia-derived thrombospondin 2 regulates alterations of presynaptic proteins of retinal neurons following elevated hydrostatic pressure. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0185388"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1410","DOI":"10.1161\/CIRCRESAHA.111.256743","article-title":"Thrombospondin-4 Is Required for Stretch-Mediated Contractility Augmentation in Cardiac Muscle","volume":"109","author":"Cingolani","year":"2011","journal-title":"Circ. Res."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/j.actbio.2017.05.058","article-title":"Effects of substrate stiffness on the tenoinduction of human mesenchymal stem cells","volume":"58","author":"Islam","year":"2017","journal-title":"Acta Biomater."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1111\/j.1365-2222.2004.01980.x","article-title":"Mechanical strain enhances proteoglycan message in fibroblasts from asthmatic subjects","volume":"34","author":"Ludwig","year":"2004","journal-title":"Clin. Exp. Allergy"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1111\/j.1600-0765.2004.00782.x","article-title":"Modulation of extracellular matrix synthesis and alkaline phosphatase activity of periodontal ligament cells by mechanical stress","volume":"40","author":"Ozaki","year":"2005","journal-title":"J. Periodontal Res."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Wang, S., L\u00fc, D., Zhang, Z., Jia, X., and Yang, L. (2018). Effects of mechanical stretching on the morphology of extracellular polymers and the mRNA expression of collagens and small leucine-rich repeat proteoglycans in vaginal fibroblasts from women with pelvic organ prolapse. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0193456"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"2382","DOI":"10.1111\/febs.12235","article-title":"Lumican is increased in experimental and clinical heart failure, and its production by cardiac fibroblasts is induced by mechanical and proinflammatory stimuli","volume":"280","author":"Engebretsen","year":"2013","journal-title":"FEBS J."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1136\/bjophthalmol-2017-311150","article-title":"Mechanical stress potentiates the differentiation of periodontal ligament stem cells into keratocytes","volume":"102","author":"Chen","year":"2018","journal-title":"Br. J. Ophthalmol."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Li, L.-F., Chen, B.-X., Tsai, Y.-H., Kao, W.W.-Y., Yang, C.-T., and Chu, P.-H. (2011). Lumican Expression in Diaphragm Induced by Mechanical Ventilation. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0024692"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1016\/j.archoralbio.2015.11.010","article-title":"Asporin in compressed periodontal ligament cells inhibits bone formation","volume":"62","author":"Ueda","year":"2016","journal-title":"Arch. Oral Biol."},{"key":"ref_81","first-page":"477","article-title":"Collagen fiber arrangement in the normal bladder lamina propria and their potential impact on the pathological substaging of bladder cancer stage T1","volume":"40","author":"Gofrit","year":"2025","journal-title":"Histol. Histopathol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"4313","DOI":"10.1364\/BOE.5.004313","article-title":"Quantitative elasticity measurement of urinary bladder wall using laser-induced surface acoustic waves","volume":"5","author":"Li","year":"2014","journal-title":"Biomed. Opt. Express."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1038\/s41598-023-51006-2","article-title":"Progressive alteration of murine bladder elasticity in actinic cystitis detected by Brillouin microscopy","volume":"14","author":"Testi","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"418","DOI":"10.1186\/s12967-026-07855-2","article-title":"Hydrostatic pressure regulates glutamine metabolism to promote bladder fibroblast activation via the Piezo1\/YAP1\/GLS1 axis","volume":"24","author":"Li","year":"2026","journal-title":"J. Transl. Med."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Ekman, M., Bhattachariya, A., Dahan, D., Uvelius, B., Albinsson, S., and Sw\u00e4rd, K. (2013). Mir-29 Repression in Bladder Outlet Obstruction Contributes to Matrix Remodeling and Altered Stiffness. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0082308"},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Gao, Y., Liu, P., He, F., Yang, X., Wu, R., Chen, W., Li, L., and Yang, Z. (2021). Fibroblast Growth Factor 2 Promotes Bladder Hypertrophy Caused by Partial Bladder Outlet Obstruction. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.630228"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"695","DOI":"10.1002\/jcb.21224","article-title":"Periostin regulates collagen fibrillogenesis and the biomechanical properties of connective tissues","volume":"101","author":"Norris","year":"2007","journal-title":"J. Cell. Biochem."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"100124","DOI":"10.1016\/j.mbplus.2022.100124","article-title":"Periostin modulates extracellular matrix behavior in tendons","volume":"16","author":"Rolnick","year":"2022","journal-title":"Matrix Biol. Plus"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"13294","DOI":"10.1074\/jbc.M109.088864","article-title":"Interaction between Periostin and BMP-1 Promotes Proteolytic Activation of Lysyl Oxidase","volume":"285","author":"Maruhashi","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1084\/jem.20071297","article-title":"Periostin is essential for cardiac healingafter acute myocardial infarction","volume":"205","author":"Shimazaki","year":"2008","journal-title":"J. Exp. Med."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1136\/gutjnl-2018-317570","article-title":"Stromal protein \u03b2ig-h3 reprogrammes tumour microenvironment in pancreatic cancer","volume":"68","author":"Goehrig","year":"2019","journal-title":"Gut"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"103758","DOI":"10.1016\/j.isci.2022.103758","article-title":"\u03b2ig-h3-structured collagen alters macrophage phenotype and function in pancreatic cancer","volume":"25","author":"Bachy","year":"2022","journal-title":"iScience"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1111\/febs.14321","article-title":"Proteomic profiling of TGFBI-null mouse corneas reveals only minor changes in matrix composition supportive of TGFBI knockdown as therapy against TGFBI-linked corneal dystrophies","volume":"285","author":"Poulsen","year":"2018","journal-title":"FEBS J."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"H1072","DOI":"10.1152\/ajpheart.00255.2009","article-title":"Tenascin-C may aggravate left ventricular remodeling and function after myocardial infarction in mice","volume":"298","author":"Nishioka","year":"2010","journal-title":"Am. J. -Physiol.-Heart Circ. Physiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"12941","DOI":"10.1038\/s41598-018-31126-w","article-title":"Targeted matrisome analysis identifies thrombospondin-2 and tenascin-C in aligned collagen stroma from invasive breast carcinoma","volume":"8","author":"Tomko","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1038\/ng850","article-title":"Tenascin-X deficiency mimics Ehlers-Danlos syndrome in mice through alteration of collagen deposition","volume":"30","author":"Mao","year":"2002","journal-title":"Nat. Genet."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1016\/j.yexcr.2004.04.030","article-title":"Modulation of collagen fibrillogenesis by tenascin-X and type VI collagen","volume":"298","author":"Minamitani","year":"2004","journal-title":"Exp. Cell Res."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/j.bpc.2009.12.011","article-title":"Tenascin-X increases the stiffness of collagen gels without affecting fibrillogenesis","volume":"147","author":"Margaron","year":"2010","journal-title":"Biophys. Chem."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1167\/iovs.16-20261","article-title":"Altered Anterior Segment Biometric Parameters in Mice Deficient in SPARC","volume":"58","author":"Ho","year":"2017","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1242\/dmm.003228","article-title":"Lack of host SPARC enhances vascular function and tumor spread in an orthotopic murine model of pancreatic carcinoma","volume":"3","author":"Arnold","year":"2010","journal-title":"Dis. Model Mech."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"1649","DOI":"10.1292\/jvms.19-0485","article-title":"Reduced fibrillar collagen accumulation in skeletal muscle of secreted protein acidic and rich in cysteine (SPARC)-null mice","volume":"81","author":"Omi","year":"2019","journal-title":"J. Vet. Med. Sci."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"32635","DOI":"10.1038\/srep32635","article-title":"Pleiotropic roles of the matricellular protein Sparc in tendon maturation and ageing","volume":"6","author":"Gehwolf","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1046\/j.1523-1747.2003.12241.x","article-title":"SPARC-Null Mice Display Abnormalities in the Dermis Characterized by Decreased Collagen Fibril Diameter and Reduced Tensile Strength","volume":"120","author":"Bradshaw","year":"2003","journal-title":"J. Investig. Dermatol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"27621","DOI":"10.1074\/jbc.M510507200","article-title":"Matricellular Hevin Regulates Decorin Production and Collagen Assembly","volume":"281","author":"Sullivan","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1050","DOI":"10.1002\/1878-0261.12073","article-title":"Stromal SPOCK 1 supports invasive pancreatic cancer growth","volume":"11","author":"Veenstra","year":"2017","journal-title":"Mol. Oncol."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"9785","DOI":"10.1038\/s41598-020-66660-z","article-title":"SPOCK1 is a novel inducer of epithelial to mesenchymal transition in drug-induced gingival overgrowth","volume":"10","author":"Alshargabi","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.matbio.2024.08.001","article-title":"Matricellular protein CCN1 promotes collagen alignment and scar integrity after myocardial infarction","volume":"133","author":"Fischer","year":"2024","journal-title":"Matrix Biol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"100009","DOI":"10.1016\/j.mbplus.2019.100009","article-title":"CCN1 expression by fibroblasts is required for bleomycin-induced skin fibrosis","volume":"3","author":"Quesnel","year":"2019","journal-title":"Matrix Biol. Plus"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"2047","DOI":"10.1096\/fj.201800622RR","article-title":"Genetic manipulation of CCN2\/CTGF unveils cell-specific ECM-remodeling effects in injured skeletal muscle","volume":"33","author":"Petrosino","year":"2019","journal-title":"FASEB J."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"1555","DOI":"10.1002\/jor.24287","article-title":"Effect of dose and release rate of CTGF and TGF\u03b23 on avascular meniscus healing","volume":"37","author":"Tarafder","year":"2019","journal-title":"J. Orthop. Res."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1007\/s13770-018-0166-2","article-title":"Local Silencing of Connective Tissue Growth Factor by siRNA\/Peptide Improves Dermal Collagen Arrangements","volume":"15","author":"Woo","year":"2018","journal-title":"Tissue Eng. Regen. Med."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"2052","DOI":"10.1002\/jor.23842","article-title":"Effect of connective tissue growth factor delivered via porous sutures on the proliferative stage of intrasynovial tendon repair","volume":"36","author":"Linderman","year":"2018","journal-title":"J. Orthop. Res."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1016\/j.joca.2014.12.012","article-title":"Targeted mutation of NOV\/CCN3 in mice disrupts joint homeostasis and causes osteoarthritis-like disease","volume":"23","author":"Roddy","year":"2015","journal-title":"Osteoarthr. Cartil."},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Marchal, P.-O., Kavvadas, P., Abed, A., Kazazian, C., Authier, F., Koseki, H., Hiraoka, S., Boffa, J.-J., Martinerie, C., and Chadjichristos, C.E. (2015). Reduced NOV\/CCN3 Expression Limits Inflammation and Interstitial Renal Fibrosis after Obstructive Nephropathy in Mice. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0137876"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"EMBJ2018101302","DOI":"10.15252\/embj.2018101302","article-title":"The tumor cell-secreted matricellular protein WISP1 drives pro-metastatic collagen linearization","volume":"38","author":"Jia","year":"2019","journal-title":"EMBO J."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"5666","DOI":"10.1158\/0008-5472.CAN-20-3982","article-title":"Matricellular Protein WISP2 Is an Endogenous Inhibitor of Collagen Linearization and Cancer Metastasis","volume":"81","author":"Janjanam","year":"2021","journal-title":"Cancer Res."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"eaar2566","DOI":"10.1126\/scisignal.aar2566","article-title":"Thrombospondin-1 promotes matrix homeostasis by interacting with collagen and lysyl oxidase precursors and collagen cross-linking sites","volume":"11","author":"Rosini","year":"2018","journal-title":"Sci. Signal."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"2640","DOI":"10.1091\/mbc.E14-05-0996","article-title":"Intermolecular interactions of thrombospondins drive their accumulation in extracellular matrix","volume":"26","author":"Kim","year":"2015","journal-title":"Mol. Biol. Cell"},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"6708","DOI":"10.1167\/iovs.11-9013","article-title":"Thrombospondin-1 (TSP1)\u2013Null and TSP2-Null Mice Exhibit Lower Intraocular Pressures","volume":"53","author":"Haddadin","year":"2012","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1083\/jcb.140.2.419","article-title":"Mice That Lack Thrombospondin 2 Display Connective Tissue Abnormalities That Are Associated with Disordered Collagen Fibrillogenesis, an Increased Vascular Density, and a Bleeding Diathesis","volume":"140","author":"Kyriakides","year":"1998","journal-title":"J. Cell Biol."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"872","DOI":"10.1152\/japplphysiol.00340.2015","article-title":"Thrombospondin-2 deficiency in growing mice alters bone collagen ultrastructure and leads to a brittle bone phenotype","volume":"119","author":"Manley","year":"2015","journal-title":"J. Appl. Physiol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"275","DOI":"10.3109\/03008207.2013.811236","article-title":"Thrombospondin-2 facilitates assembly of a type-I collagen-rich matrix in marrow stromal cells undergoing osteoblastic differentiation","volume":"54","author":"Alford","year":"2013","journal-title":"Connect. Tissue Res."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.matbio.2019.03.002","article-title":"Thrombospondin-2 regulates extracellular matrix production, LOX levels, and cross-linking via downregulation of miR-29","volume":"82","author":"Calabro","year":"2019","journal-title":"Matrix Biol."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.matbio.2014.02.003","article-title":"Control of organization and function of muscle and tendon by thrombospondin-4","volume":"37","author":"Frolova","year":"2014","journal-title":"Matrix Biol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"2363","DOI":"10.1096\/fj.11-190728","article-title":"Thrombospondin-4 regulates fibrosis and remodeling of the myocardium in response to pressure overload","volume":"26","author":"Frolova","year":"2012","journal-title":"FASEB J."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"H1486","DOI":"10.1152\/ajpheart.00046.2016","article-title":"Thrombospondin-4 knockout in hypertension protects small-artery endothelial function but induces aortic aneurysms","volume":"310","author":"Palao","year":"2016","journal-title":"Am. J. Physiol.-Heart Circ. Physiol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1177\/1947603510361238","article-title":"Elevated Levels of Cartilage Oligomeric Matrix Protein during In Vitro Cartilage Matrix Generation Decrease Collagen Fibril Diameter","volume":"1","author":"Koevoet","year":"2010","journal-title":"Cartilage"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1083\/jcb.136.3.729","article-title":"Targeted Disruption of Decorin Leads to Abnormal Collagen Fibril Morphology and Skin Fragility","volume":"136","author":"Danielson","year":"1997","journal-title":"J. Cell Biol."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/j.matbio.2013.04.004","article-title":"Effects of decorin proteoglycan on fibrillogenesis, ultrastructure, and mechanics of type I collagen gels","volume":"32","author":"Reese","year":"2013","journal-title":"Matrix Biol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"108933","DOI":"10.1016\/j.exer.2022.108933","article-title":"Decorin regulates collagen fibrillogenesis during corneal wound healing in mouse in vivo","volume":"216","author":"Gupta","year":"2022","journal-title":"Exp. Eye Res."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.matbio.2017.08.004","article-title":"Decorin and biglycan are necessary for maintaining collagen fibril structure, fiber realignment, and mechanical properties of mature tendons","volume":"64","author":"Robinson","year":"2017","journal-title":"Matrix Biol."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"4101","DOI":"10.1021\/acsbiomaterials.5c00732","article-title":"Recombinant Small Leucine-Rich Proteoglycans Modulate Fiber Structure and Mechanical Properties of Collagen Gels","volume":"11","author":"Lopez","year":"2025","journal-title":"ACS Biomater. Sci. Eng."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"2546","DOI":"10.1002\/jor.25299","article-title":"Biglycan has a major role in maintenance of mature tendon mechanics","volume":"40","author":"Beach","year":"2022","journal-title":"J. Orthop. Res."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"1180","DOI":"10.1359\/jbmr.2002.17.7.1180","article-title":"Phenotypic Effects of Biglycan Deficiency Are Linked to Collagen Fibril Abnormalities, Are Synergized by Decorin Deficiency, and Mimic Ehlers-Danlos-Like Changes in Bone and Other Connective Tissues","volume":"17","author":"Corsi","year":"2002","journal-title":"J. Bone Miner. Res."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1083\/jcb.151.4.779","article-title":"Differential Expression of Lumican and Fibromodulin Regulate Collagen Fibrillogenesis in Developing Mouse Tendons","volume":"151","author":"Ezura","year":"2000","journal-title":"J. Cell Biol."},{"key":"ref_136","first-page":"3365","article-title":"Corneal opacity in lumican-null mice: Defects in collagen fibril structure and packing in the posterior stroma","volume":"41","author":"Chakravarti","year":"2000","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"1277","DOI":"10.1083\/jcb.141.5.1277","article-title":"Lumican Regulates Collagen Fibril Assembly: Skin Fragility and Corneal Opacity in the Absence of Lumican","volume":"141","author":"Chakravarti","year":"1998","journal-title":"J. Cell Biol."},{"key":"ref_138","doi-asserted-by":"crossref","unstructured":"Maccarana, M., Svensson, R.B., Knutsson, A., Giannopoulos, A., Pelkonen, M., Weis, M., Eyre, D., Warman, M., and Kalamajski, S. (2017). Asporin-deficient mice have tougher skin and altered skin glycosaminoglycan content and structure. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0184028"},{"key":"ref_139","doi-asserted-by":"crossref","unstructured":"Kinoshita, M., Yamada, S., Sasaki, J., Suzuki, S., Kajikawa, T., Iwayama, T., Fujihara, C., Imazato, S., and Murakami, S. (2023). Mice Lacking PLAP-1\/Asporin Show Alteration of Periodontal Ligament Structures and Acceleration of Bone Loss in Periodontitis. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms242115989"},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"19065","DOI":"10.1038\/s41598-020-76107-0","article-title":"Distinct effects of different matrix proteoglycans on collagen fibrillogenesis and cell-mediated collagen reorganization","volume":"10","author":"Chen","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_141","doi-asserted-by":"crossref","unstructured":"de Weck, A., Bitter, H., and Kauffmann, A. (2017). Fibroblasts cell lines misclassified as cancer cell lines. bioRxiv.","DOI":"10.1101\/166199"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1038\/s41389-021-00302-5","article-title":"TGF-\u03b21-mediated transition of resident fibroblasts to cancer-associated fibroblasts promotes cancer metastasis in gastrointestinal stromal tumor","volume":"10","author":"Yoon","year":"2021","journal-title":"Oncogenesis"},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1016\/j.ccr.2014.04.005","article-title":"Depletion of Carcinoma-Associated Fibroblasts and Fibrosis Induces Immunosuppression and Accelerates Pancreas Cancer with Reduced Survival","volume":"25","author":"Carstens","year":"2014","journal-title":"Cancer Cell"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1084\/jem.20162024","article-title":"Distinct populations of inflammatory fibroblasts and myofibroblasts in pancreatic cancer","volume":"214","author":"Biffi","year":"2017","journal-title":"J. Exp. Med."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"5077","DOI":"10.1038\/s41467-020-18916-5","article-title":"Single-cell RNA sequencing highlights the role of inflammatory cancer-associated fibroblasts in bladder urothelial carcinoma","volume":"11","author":"Chen","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"1346","DOI":"10.1016\/j.ccell.2026.06.001","article-title":"Molecular phenotypes and spatial archetypes: A new framework for cancer-associated fibroblasts","volume":"44","author":"Liu","year":"2026","journal-title":"Cancer Cell"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"2592","DOI":"10.1158\/0008-5472.CAN-24-3954","article-title":"The Hippo Pathway Is Dysregulated in Cancer-Associated Fibroblasts in Anti\u2013PD-L1\u2013Resistant Cancer","volume":"86","author":"Roels","year":"2026","journal-title":"Cancer Res."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1158\/1541-7786.MCR-17-0784","article-title":"Exosomes Induce Fibroblast Differentiation into Cancer-Associated Fibroblasts through TGF\u03b2 Signaling","volume":"16","author":"Bernard","year":"2018","journal-title":"Mol. Cancer Res."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1186\/s12967-025-06325-5","article-title":"CAFs activated by YAP1 upregulate cancer matrix stiffness to mediate hepatocellular carcinoma progression","volume":"23","author":"Yan","year":"2025","journal-title":"J. Transl. Med."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"6966","DOI":"10.1038\/s41467-023-42382-4","article-title":"Cancer-associated fibroblasts actively compress cancer cells and modulate mechanotransduction","volume":"14","author":"Barbazan","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"23335","DOI":"10.18632\/oncotarget.8024","article-title":"Identification of extracellular vesicle-borne periostin as a feature of muscle-invasive bladder cancer","volume":"7","author":"Silvers","year":"2016","journal-title":"Oncotarget"},{"key":"ref_152","doi-asserted-by":"crossref","unstructured":"Lang, K., Kahveci, S., Bonberg, N., Wichert, K., Behrens, T., Hovanec, J., Roghmann, F., Noldus, J., Tam, Y.C., and Tannapfel, A. (2019). TGFBI Protein Is Increased in the Urine of Patients with High-Grade Urothelial Carcinomas, and Promotes Cell Proliferation and Migration. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20184483"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"1641","DOI":"10.1016\/j.cca.2012.05.005","article-title":"Expression of CD147, BIGH3 and Stathmin and their potential role as diagnostic marker in patients with urothelial carcinoma of the bladder","volume":"413","author":"Bhagirath","year":"2012","journal-title":"Clin. Chim. Acta"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"961","DOI":"10.3892\/mmr.2013.1873","article-title":"Urine tenascin-C is an independent risk factor for bladder cancer patients","volume":"9","author":"Guan","year":"2014","journal-title":"Mol. Med. Rep."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"1351","DOI":"10.1007\/s00432-009-0576-6","article-title":"B and C domain containing tenascin-C: Urinary markers for invasiveness of urothelial carcinoma of the urinary bladder?","volume":"135","author":"Richter","year":"2009","journal-title":"J. Cancer Res. Clin. Oncol."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1177\/1010428317699112","article-title":"Altered expression of small leucine-rich proteoglycans (Decorin, Biglycan and Lumican): Plausible diagnostic marker in urothelial carcinoma of bladder","volume":"39","author":"Appunni","year":"2017","journal-title":"Tumor Biol."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"1800148","DOI":"10.1002\/prca.201800148","article-title":"Diagnostic and Prognostic Performance of Secreted Protein Acidic and Rich in Cysteine (SPARC) Assay for Detecting Primary and Recurrent Urinary Bladder Cancer","volume":"13","author":"Critselis","year":"2019","journal-title":"Proteom. Clin. Appl."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1006\/excr.2002.5590","article-title":"Suppression of Anchorage-Independent Growth of Human Cancer Cell Lines by the TRIF52\/Periostin\/OSF-2 Gene","volume":"279","author":"Yoshioka","year":"2002","journal-title":"Exp. Cell Res."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1002\/ijc.21120","article-title":"Periostin is down-regulated in high grade human bladder cancers and suppresses in vitro cell invasiveness and in vivo metastasis of cancer cells","volume":"117","author":"Kim","year":"2005","journal-title":"Int. J. Cancer"},{"key":"ref_160","first-page":"161","article-title":"Role of alternative splicing of periostin in human bladder carcinogenesis","volume":"32","author":"Kim","year":"2008","journal-title":"Int. J. Oncol."},{"key":"ref_161","first-page":"1759","article-title":"Opposite regulation of epithelial-to-mesenchymal transition and cell invasiveness by periostin between prostate and bladder cancer cells","volume":"38","author":"Kim","year":"2011","journal-title":"Int. J. Oncol."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"4276","DOI":"10.3892\/ol.2017.6004","article-title":"Periostin suppresses in vivo invasiveness via PDK1\/Akt\/mTOR signaling pathway in a mouse orthotopic model of bladder cancer","volume":"13","author":"Kim","year":"2017","journal-title":"Oncol. Lett."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"zyag001","DOI":"10.1093\/intbio\/zyag001","article-title":"POSTN+ cancer-associated fibroblasts promote bladder cancer progression via angiogenesis and immune modulation: An analysis based on single-cell Transcriptomics","volume":"18","author":"Zhang","year":"2026","journal-title":"Integr. Biol."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"980","DOI":"10.1111\/cas.13403","article-title":"Epirubicin suppresses proliferative and metastatic potential by downregulating transforming growth factor-\u03b2-induced expression in urothelial carcinoma","volume":"109","author":"Shang","year":"2018","journal-title":"Cancer Sci."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"1134","DOI":"10.34133\/research.1134","article-title":"Hypoxia-Induced TGFBI Promotes Bladder Cancer Progression by Creating a Stemness Regulation Loop through Stabilizing the Disulfide Bonds of GDF15","volume":"9","author":"Zhang","year":"2026","journal-title":"Research"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"6995","DOI":"10.2147\/CMAR.S208984","article-title":"Secreted TGF-beta-induced protein promotes aggressive progression in bladder cancer cells","volume":"11","author":"Zou","year":"2019","journal-title":"Cancer Manag. Res."},{"key":"ref_167","doi-asserted-by":"crossref","unstructured":"Guan, Z., Sun, Y., Mu, L., Jiang, Y., and Fan, J. (2022). Tenascin-C promotes bladder cancer progression and its action depends on syndecan-4 and involves NF-\u03baB signaling activation. BMC Cancer, 22.","DOI":"10.1186\/s12885-022-09285-x"},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1172\/JCI71241","article-title":"Loss of SPARC in bladder cancer enhances carcinogenesis and progression","volume":"123","author":"Said","year":"2013","journal-title":"J. Clin. Investig."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"3243","DOI":"10.1021\/pr100189d","article-title":"Analysis of Secreted Proteins for the Study of Bladder Cancer Cell Aggressiveness","volume":"9","author":"Makridakis","year":"2010","journal-title":"J. Proteome Res."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"e70196","DOI":"10.1111\/jcmm.70196","article-title":"The oncogenic functions of SPARCL1 in bladder cancer","volume":"28","author":"Li","year":"2024","journal-title":"J. Cell. Mol. Med."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"66316","DOI":"10.18632\/oncotarget.19987","article-title":"Targeting of CCN2 suppresses tumor progression and improves chemo-sensitivity in urothelial bladder cancer","volume":"8","author":"Wang","year":"2017","journal-title":"Oncotarget"},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"2386753","DOI":"10.1080\/15384047.2024.2386753","article-title":"Amino-truncated NOV expression and its correlation with clinicopathologic features, prognosis, metastasis, and chemoresistance in bladder cancer","volume":"25","author":"Xiong","year":"2024","journal-title":"Cancer Biol. Ther."},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"102680","DOI":"10.1016\/j.tranon.2026.102680","article-title":"WISP1 is the stromal-secreting oncoprotein via paracrine downregulation of NDRG1, KAI1, and Maspin in human bladder cancer cells","volume":"65","author":"Chen","year":"2026","journal-title":"Transl. Oncol."},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"6046","DOI":"10.1111\/jcmm.16463","article-title":"Thrombospondin-4 promotes bladder cancer cell migration and invasion via MMP2 production","volume":"25","author":"Chou","year":"2021","journal-title":"J. Cell. Mol. Med."},{"key":"ref_175","doi-asserted-by":"crossref","unstructured":"Sainio, A., Nyman, M., Lund, R., Vuorikoski, S., Bostr\u00f6m, P., Laato, M., Bostr\u00f6m, P.J., and J\u00e4rvel\u00e4inen, H. (2013). Lack of Decorin Expression by Human Bladder Cancer Cells Offers New Tools in the Therapy of Urothelial Malignancies. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0076190"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"1835","DOI":"10.1002\/emmm.201302655","article-title":"An essential role for decorin in bladder cancer invasiveness","volume":"5","author":"Krumeich","year":"2013","journal-title":"EMBO Mol. Med."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"2163","DOI":"10.1007\/s13402-024-00998-8","article-title":"Non-glycanated \u0394DCN isoform in muscle invasive bladder cancer mediates cancer stemness and gemcitabine resistance","volume":"47","author":"Wu","year":"2024","journal-title":"Cell. Oncol."},{"key":"ref_178","doi-asserted-by":"crossref","unstructured":"Niedworok, C., R\u00f6ck, K., Kretschmer, I., Freudenberger, T., Nagy, N., Szarvas, T., vom Dorp, F., Reis, H., R\u00fcbben, H., and Fischer, J.W. (2013). Inhibitory Role of the Small Leucine-Rich Proteoglycan Biglycan in Bladder Cancer. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0080084"},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1016\/j.tranon.2019.05.014","article-title":"Knockdown of Lumican Inhibits Proliferation and Migration of Bladder Cancer","volume":"12","author":"Mao","year":"2019","journal-title":"Transl. Oncol."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"147573","DOI":"10.1016\/j.gene.2023.147573","article-title":"Prognostic and immunological role of Asporin across cancers and exploration in bladder cancer","volume":"878","author":"Wei","year":"2023","journal-title":"Gene"},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"2573","DOI":"10.1038\/s41388-018-0129-z","article-title":"Alternative transcription of a shorter, non-anti-angiogenic thrombospondin-2 variant in cancer-associated blood vessels","volume":"37","author":"Roudnicky","year":"2018","journal-title":"Oncogene"},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"204","DOI":"10.7150\/ijms.122895","article-title":"Thrombospondin-4 drives lymphangiogenesis through cooperation with VEGF-C in human bladder cancer","volume":"23","author":"Hwang","year":"2026","journal-title":"Int. J. Med. Sci."},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1016\/j.toxlet.2010.08.020","article-title":"SPARC gene expression is repressed in human urothelial cells (UROtsa) exposed to or malignantly transformed by cadmium or arsenite","volume":"199","author":"Larson","year":"2010","journal-title":"Toxicol. Lett."},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1007\/s00428-006-0347-6","article-title":"CD34+ fibrocytes in chronic cystitis and noninvasive and invasive urothelial carcinomas of the urinary bladder","volume":"450","author":"Nimphius","year":"2007","journal-title":"Virchows Arch."},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1038\/s41585-024-00885-9","article-title":"Preclinical models of bladder cancer: BBN and beyond","volume":"21","author":"Matye","year":"2024","journal-title":"Nat. Rev. Urol."},{"key":"ref_186","doi-asserted-by":"crossref","unstructured":"Millet, M., Bollmann, E., Ringuette Goulet, C., Bernard, G., Chabaud, S., Huot, M.-\u00c9., Pouliot, F., Bolduc, S., and Bordeleau, F. (2022). Cancer-Associated Fibroblasts in a 3D Engineered Tissue Model Induce Tumor-like Matrix Stiffening and EMT Transition. Cancers, 14.","DOI":"10.3390\/cancers14153810"},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1016\/j.cell.2017.09.007","article-title":"Comprehensive Molecular Characterization of Muscle-Invasive Bladder Cancer","volume":"171","author":"Robertson","year":"2017","journal-title":"Cell"},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"1318","DOI":"10.1126\/science.aaz1776","article-title":"The GTEx Consortium atlas of genetic regulatory effects across human tissues","volume":"369","author":"Aguet","year":"2020","journal-title":"Science"},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1038\/s41586-019-1186-3","article-title":"Next-generation characterization of the Cancer Cell Line Encyclopedia","volume":"569","author":"Ghandi","year":"2019","journal-title":"Nature"},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"D638","DOI":"10.1093\/nar\/gkac1000","article-title":"The STRING database in 2023: Protein\u2013protein association networks and functional enrichment analyses for any sequenced genome of interest","volume":"51","author":"Szklarczyk","year":"2023","journal-title":"Nucleic Acids Res."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"2612","DOI":"10.1038\/ncomms3612","article-title":"Inferring tumour purity and stromal and immune cell admixture from expression data","volume":"4","author":"Yoshihara","year":"2013","journal-title":"Nat. Commun."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"btad547","DOI":"10.1093\/bioinformatics\/btad547","article-title":"PyDESeq2: A python package for bulk RNA-seq differential expression analysis","volume":"39","author":"Muzellec","year":"2023","journal-title":"Bioinformatics"},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"D991","DOI":"10.1093\/nar\/gks1193","article-title":"NCBI GEO: Archive for functional genomics data sets\u2014update","volume":"41","author":"Barrett","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"W83","DOI":"10.1093\/nar\/gkae410","article-title":"The Galaxy platform for accessible, reproducible, and collaborative data analyses: 2024 update","volume":"52","author":"Abueg","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: A flexible trimmer for Illumina sequence data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1038\/s41587-019-0201-4","article-title":"Graph-based genome alignment and genotyping with HISAT2 and HISAT-genotype","volume":"37","author":"Kim","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1186\/s13059-019-1910-1","article-title":"Transcriptome assembly from long-read RNA-seq alignments with StringTie2","volume":"20","author":"Kovaka","year":"2019","journal-title":"Genome Biol."},{"key":"ref_198","doi-asserted-by":"crossref","first-page":"eabl4896","DOI":"10.1126\/science.abl4896","article-title":"The Tabula Sapiens: A multiple-organ, single-cell transcriptomic atlas of humans","volume":"376","author":"Jones","year":"2022","journal-title":"Science"},{"key":"ref_199","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s13059-017-1382-0","article-title":"SCANPY: Large-scale single-cell gene expression data analysis","volume":"19","author":"Wolf","year":"2018","journal-title":"Genome Biol."},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"1317","DOI":"10.21105\/joss.01317","article-title":"Lifelines: Survival analysis in Python","volume":"4","year":"2019","journal-title":"J. Open Source Softw."}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/16\/8\/1191\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T16:01:24Z","timestamp":1786723284000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/16\/8\/1191"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,14]]},"references-count":200,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["biom16081191"],"URL":"https:\/\/doi.org\/10.3390\/biom16081191","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,14]]}}}