{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,10]],"date-time":"2026-11-10T21:11:07Z","timestamp":1794345067671,"version":"4.9.0"},"reference-count":81,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2024,5,21]],"date-time":"2024-05-21T00:00:00Z","timestamp":1716249600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100006769","name":"Russian Science Foundation","doi-asserted-by":"publisher","award":["21-75-10082"],"award-info":[{"award-number":["21-75-10082"]}],"id":[{"id":"10.13039\/501100006769","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Joint-resident chondrogenic precursor cells have become a significant therapeutic option due to the lack of regenerative capacity in articular cartilage. Progenitor cells are located in the superficial zone of the articular cartilage, producing lubricin\/Prg4 to decrease friction of cartilage surfaces during joint movement. Prg4-positive progenitors are crucial in maintaining the joint\u2019s structure and functionality. The disappearance of progenitor cells leads to changes in articular hyaline cartilage over time, subchondral bone abnormalities, and the formation of ectopic ossification. Genetic labeling cell technology has been the main tool used to characterize Prg4-expressing progenitor cells of articular cartilage in vivo through drug injection at different time points. This technology allows for the determination of the origin of progenitor cells and the tracking of their progeny during joint development and cartilage damage. We endeavored to highlight the currently known information about the Prg4-producing cell population in the joint to underline the significance of the role of these cells in the development of articular cartilage and its homeostasis. This review focuses on superficial progenitors in the joint, how they contribute to postnatal articular cartilage formation, their capacity for regeneration, and the consequences of Prg4 deficiency in these cells. We have accumulated information about the Prg4+ cell population of articular cartilage obtained through various elegantly designed experiments using transgenic technologies to identify potential opportunities for further research.<\/jats:p>","DOI":"10.3390\/ijms25115605","type":"journal-article","created":{"date-parts":[[2024,5,22]],"date-time":"2024-05-22T07:56:11Z","timestamp":1716364571000},"page":"5605","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["Prg4-Expressing Chondroprogenitor Cells in the Superficial Zone of Articular Cartilage"],"prefix":"10.3390","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-4421-0495","authenticated-orcid":false,"given":"Nadezda","family":"Ignatyeva","sequence":"first","affiliation":[{"name":"Institute for Regenerative Medicine, Sechenov First Moscow State Medical University (Sechenov University), 8-2 Trubetskaya St., Moscow 119048, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nikita","family":"Gavrilov","sequence":"additional","affiliation":[{"name":"Institute for Regenerative Medicine, Sechenov First Moscow State Medical University (Sechenov University), 8-2 Trubetskaya St., Moscow 119048, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter S.","family":"Timashev","sequence":"additional","affiliation":[{"name":"Institute for Regenerative Medicine, Sechenov First Moscow State Medical University (Sechenov University), 8-2 Trubetskaya St., Moscow 119048, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ekaterina V.","family":"Medvedeva","sequence":"additional","affiliation":[{"name":"Institute for Regenerative Medicine, Sechenov First Moscow State Medical University (Sechenov University), 8-2 Trubetskaya St., Moscow 119048, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2024,5,21]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1007\/s11914-015-0290-z","article-title":"Articular Cartilage: Structural and Developmental Intricacies and Questions","volume":"13","author":"Decker","year":"2015","journal-title":"Curr. Osteoporos. Rep."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Medvedeva, E.V., Grebenik, E.A., Gornostaeva, S.N., Telpuhov, V.I., Lychagin, A.V., Timashev, P.S., and Chagin, A.S. (2018). Repair of Damaged Articular Cartilage: Current Approaches and Future Directions. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19082366"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Kurenkova, A.D., Romanova, I.A., Kibirskiy, P.D., Timashev, P., and Medvedeva, E.V. (2022). Strategies to Convert Cells into Hyaline Cartilage: Magic Spells for Adult Stem Cells. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms231911169"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1186\/s41232-022-00202-0","article-title":"The superficial zone of articular cartilage","volume":"42","author":"Saito","year":"2022","journal-title":"Inflamm. Regen."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1097\/00004424-200010000-00003","article-title":"Anatomy, biochemistry, and physiology of articular cartilage","volume":"35","author":"Huber","year":"2000","journal-title":"Investig. Radiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1016\/j.joca.2006.09.010","article-title":"The structural architecture of adult mammalian articular cartilage evolves by a synchronized process of tissue resorption and neoformation during postnatal development","volume":"15","author":"Hunziker","year":"2007","journal-title":"Osteoarthr. Cartil."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.semcdb.2016.10.005","article-title":"Articular cartilage and joint development from embryogenesis to adulthood","volume":"62","author":"Decker","year":"2017","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Mienaltowski, M.J., Huang, L., Frisbie, D.D., McIlwraith, C.W., Stromberg, A.J., Bathke, A.C., and Macleod, J.N. (2009). Transcriptional profiling differences for articular cartilage and repair tissue in equine joint surface lesions. BMC Med. Genom., 2.","DOI":"10.1186\/1755-8794-2-60"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"346ra90","DOI":"10.1126\/scitranslmed.aad8335","article-title":"Radiocarbon dating reveals minimal collagen turnover in both healthy and osteoarthritic human cartilage","volume":"8","author":"Heinemeier","year":"2016","journal-title":"Sci. Transl. Med."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1016\/j.addr.2018.12.015","article-title":"Articular fibrocartilage\u2014Why does hyaline cartilage fail to repair?","volume":"146","author":"Armiento","year":"2019","journal-title":"Adv. Drug Deliv. Rev."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1007\/s004290100178","article-title":"The development of articular cartilage: Evidence for an appositional growth mechanism","volume":"203","author":"Hayes","year":"2001","journal-title":"Anat. Embryol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1242\/jcs.00912","article-title":"The surface of articular cartilage contains a progenitor cell population","volume":"117","author":"Dowthwaite","year":"2004","journal-title":"J. Cell Sci."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1016\/j.joca.2008.08.002","article-title":"Clonal chondroprogenitors maintain telomerase activity and Sox9 expression during extended monolayer culture and retain chondrogenic potential","volume":"17","author":"Khan","year":"2009","journal-title":"Osteoarthr. Cartil."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1261","DOI":"10.1002\/art.39030","article-title":"Identification of a Prg4-expressing articular cartilage progenitor cell population in mice","volume":"67","author":"Kozhemyakina","year":"2015","journal-title":"Arthritis Rheumatol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"3939","DOI":"10.1007\/s00018-019-03191-5","article-title":"Mechanisms of synovial joint and articular cartilage development","volume":"76","author":"Chijimatsu","year":"2019","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/bs.ctdb.2018.11.002","article-title":"Joints in the appendicular skeleton: Developmental mechanisms and evolu-tionary influences","volume":"133","author":"Rux","year":"2019","journal-title":"Curr. Top. Dev. Biol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"e12998","DOI":"10.1111\/cpr.12998","article-title":"Growth differentiation factor 5 in cartilage and osteoarthritis: A possible therapeutic candidate","volume":"54","author":"Sun","year":"2021","journal-title":"Cell Prolif."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1016\/j.ydbio.2008.01.012","article-title":"A distinct cohort of progenitor cells participates in synovial joint and articular cartilage formation during mouse limb skeletogenesis","volume":"316","author":"Koyama","year":"2008","journal-title":"Dev. Biol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2577","DOI":"10.1016\/j.celrep.2016.05.055","article-title":"Joint Development Involves a Continuous Influx of Gdf5-Positive Cells","volume":"15","author":"Shwartz","year":"2016","journal-title":"Cell Rep."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Xu, W., Wang, W., Liu, D., and Liao, D. (2022). Roles of Cartilage-Resident Stem\/Progenitor Cells in Cartilage Physiology, Development, Repair and Osteoarthritis. Cells, 11.","DOI":"10.3390\/cells11152305"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1016\/j.stemcr.2019.08.006","article-title":"Lgr5 and Col22a1 Mark Progenitor Cells in the Lineage toward Juvenile Articular Chondrocytes","volume":"13","author":"Feng","year":"2019","journal-title":"Stem Cell Rep."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1179","DOI":"10.1016\/j.stem.2023.08.004","article-title":"Lgr5-expressing secretory cells form a Wnt inhibitory niche in cartilage critical for chondrocyte identity","volume":"30","author":"Ruscitto","year":"2023","journal-title":"Cell Stem Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"e81569","DOI":"10.7554\/eLife.81569","article-title":"NFATc1 marks articular cartilage progenitors and negatively determines articular chondrocyte differentiation","volume":"12","author":"Zhang","year":"2023","journal-title":"eLife"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1739","DOI":"10.1038\/labinvest.2011.144","article-title":"Roles of \u03b2-catenin signaling in phenotypic expression and proliferation of articular cartilage superficial zone cells","volume":"91","author":"Yasuhara","year":"2011","journal-title":"Lab. Investig."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Zhang, C.-H., Gao, Y., Jadhav, U., Hung, H.-H., Holton, K.M., Grodzinsky, A.J., Shivdasani, R.A., and Lassar, A.B. (2021). Creb5 establishes the competence for Prg4 expression in articular cartilage. Commun. Biol., 4.","DOI":"10.1038\/s42003-021-01857-0"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"418","DOI":"10.1002\/art.37760","article-title":"Zone-specific gene expression patterns in articular cartilage","volume":"65","author":"Grogan","year":"2013","journal-title":"Arthritis Rheum."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"71","DOI":"10.3109\/03008209209014228","article-title":"Characterization of a bovine synovial fluid lubricating factor. I. Chemical, surface activity and lubricating properties","volume":"28","author":"Jay","year":"1992","journal-title":"Connect. Tissue Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1016\/S0736-0266(00)00040-1","article-title":"Homology of lubricin and superficial zone protein (SZP): Products of megakaryocyte stimulating factor (MSF) gene expression by human synovial fibroblasts and articular chondrocytes localized to chromosome 1q25","volume":"19","author":"Jay","year":"2001","journal-title":"J. Orthop. Res."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2065","DOI":"10.1002\/art.23548","article-title":"Biomechanics of cartilage articulation: Effects of lubrication and degeneration on shear deformation","volume":"58","author":"Wong","year":"2008","journal-title":"Arthritis Rheum."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"6497","DOI":"10.1038\/ncomms7497","article-title":"Supramolecular synergy in the boundary lubrication of synovial joints","volume":"6","author":"Seror","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1159\/000056791","article-title":"Isolation, characterization and mapping of the mouse and human PRG4 (proteoglycan 4) genes","volume":"90","author":"Ikegawa","year":"2000","journal-title":"Cytogenet. Cell Genet."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1172\/JCI200522263","article-title":"The secreted glycoprotein lubricin protects cartilage surfaces and inhibits synovial cell overgrowth","volume":"115","author":"Rhee","year":"2005","journal-title":"J. Clin. Investig."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1002\/jbmr.508","article-title":"Proteoglycan 4: A dynamic regulator of skeletogenesis and parathyroid hormone skeletal anabolism","volume":"27","author":"Novince","year":"2012","journal-title":"J. Bone Miner. Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2538","DOI":"10.7150\/thno.39386","article-title":"Proteoglycan 4 predicts tribological properties of repaired cartilage tissue","volume":"10","author":"Qiao","year":"2020","journal-title":"Theranostics"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"An, H., Liu, Y., Yi, J., Xie, H., Li, C., Wang, X., and Chai, W. (2022). Research progress of cartilage lubrication and biomimetic cartilage lubrication materials. Front. Bioeng. Biotechnol., 10.","DOI":"10.3389\/fbioe.2022.1012653"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"De Luca, P., Kouroupis, D., Vigan\u00f2, M., Perucca-Orfei, C., Kaplan, L., Zagra, L., de Girolamo, L., Correa, D., and Colombini, A. (2019). Human Diseased Articular Cartilage Contains a Mesenchymal Stem Cell-Like Population of Chondroprogenitors with Strong Immunomodulatory Responses. J. Clin. Med., 8.","DOI":"10.3390\/jcm8040423"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1512","DOI":"10.1177\/0363546516686965","article-title":"Intra-articular Recombinant Human Proteoglycan 4 Mitigates Cartilage Damage After Destabilization of the Medial Meniscus in the Yucatan Minipig","volume":"45","author":"Waller","year":"2017","journal-title":"Am. J. Sports Med."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"6909","DOI":"10.1038\/s41467-023-42199-1","article-title":"Loss of Grem1-lineage chondrogenic progenitor cells causes osteoarthritis","volume":"14","author":"Ng","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1016\/j.orthres.2004.11.011","article-title":"Proteoglycan 4 (PRG4) synthesis and immunolocalization in bovine meniscus","volume":"23","author":"Schumacher","year":"2005","journal-title":"J. Orthop. Res."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1916","DOI":"10.1002\/jor.21473","article-title":"Lubricin distribution in the torn human anterior cruciate ligament and meniscus","volume":"29","author":"Zhang","year":"2011","journal-title":"J. Orthop. Res."},{"key":"ref_41","first-page":"594","article-title":"Lubricin is a product of megakaryocyte stimulating factor gene expression by human synovial fibroblasts","volume":"27","author":"Jay","year":"2000","journal-title":"J. Rheumatol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"882","DOI":"10.1002\/art.22446","article-title":"Boundary lubrication of articular cartilage: Role of synovial fluid constituents","volume":"56","author":"Schmidt","year":"2007","journal-title":"Arthritis Rheum."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1016\/S0945-053X(02)00056-2","article-title":"Immunolocalisation and expression of proteoglycan 4 (cartilage superficial zone proteoglycan) in tendon","volume":"21","author":"Rees","year":"2002","journal-title":"Matrix Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e79925","DOI":"10.7554\/eLife.79925","article-title":"Lineage-specific differences and regulatory networks governing human chondrocyte development","volume":"12","author":"Richard","year":"2023","journal-title":"eLife"},{"key":"ref_45","first-page":"e167858","article-title":"Adult Prg4+ progenitors repair long-term articular cartilage wounds in vivo","volume":"8","author":"Massengale","year":"2023","journal-title":"J. Clin. Investig."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1806","DOI":"10.3382\/ps.0701806","article-title":"The role of the growth plate in longitudinal bone growth","volume":"70","author":"Pines","year":"1991","journal-title":"Poult. Sci."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1038\/s41586-019-0989-6","article-title":"A radical switch in clonality reveals a stem cell niche in the epiphyseal growth plate","volume":"567","author":"Newton","year":"2019","journal-title":"Nature"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1002\/1529-0131(199804)41:4<730::AID-ART22>3.0.CO;2-Y","article-title":"The camptodactyly-arthropathy-coxa vara-pericarditis syndrome: Clinical features and genetic mapping to human chromosome 1","volume":"41","author":"Bahabri","year":"1998","journal-title":"Arthritis Rheum."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1038\/15496","article-title":"CACP, encoding a secreted proteoglycan, is mutated in camptodactyly-arthropathy-coxa vara-pericarditis syndrome","volume":"23","author":"Marcelino","year":"1999","journal-title":"Nat. Genet."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1002\/1096-8628(20001127)95:3<233::AID-AJMG9>3.0.CO;2-3","article-title":"Clinical variability and genetic homogeneity of the camptodactyly-arthropathy-coxa vara-pericarditis syndrome","volume":"95","author":"Faivre","year":"2000","journal-title":"Am. J. Med. Genet."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"3662","DOI":"10.1002\/art.22974","article-title":"Association between friction and wear in diarthrodial joints lacking lubricin","volume":"56","author":"Jay","year":"2007","journal-title":"Arthritis Rheum."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"5852","DOI":"10.1073\/pnas.1219289110","article-title":"Role of lubricin and boundary lubrication in the prevention of chondrocyte apoptosis","volume":"110","author":"Waller","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Yang, D.S., Dickerson, E.E., Zhang, L.X., Richendrfer, H., Karamchedu, P.N., Badger, G.J., Schmidt, T.A., Fredericks, A.M., Elsaid, K.A., and Jay, G.D. (2022). Quadruped Gait and Regulation of Apoptotic Factors in Tibiofemoral Joints following Intra-Articular rhPRG4 Injection in Prg4 Null Mice. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23084245"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1186\/s13075-016-0967-4","article-title":"Superficial zone cellularity is deficient in mice lacking lubricin: A stereoscopic analysis","volume":"18","author":"Karamchedu","year":"2016","journal-title":"Arthritis Res. Ther."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2077","DOI":"10.1002\/jor.24378","article-title":"Absence of Proteoglycan 4 (Prg4) Leads to Increased Subchondral Bone Porosity Which Can Be Mitigated Through Intra-Articular Injection of PRG4","volume":"37","author":"Abubacker","year":"2019","journal-title":"J. Orthop. Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1002\/jbmr.4226","article-title":"Lubricin Contributes to Homeostasis of Articular Cartilage by Modulating Differentiation of Superficial Zone Cells","volume":"36","author":"Maenohara","year":"2020","journal-title":"J. Bone Miner. Res."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Takahata, Y., Hagino, H., Kimura, A., Urushizaki, M., Yamamoto, S., Wakamori, K., Murakami, T., Hata, K., and Nishimura, R. (2022). Regulatory Mechanisms of Prg4 and Gdf5 Expression in Articular Cartilage and Functions in Osteoarthritis. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23094672"},{"key":"ref_58","first-page":"191","article-title":"Static and dynamic compression regulate cartilage metabolism of PRoteoGlycan 4 (PRG4)","volume":"43","author":"Nugent","year":"2006","journal-title":"Biorheology"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1002\/art.24143","article-title":"Modulation of lubricin biosynthesis and tissue surface properties following cartilage mechanical injury","volume":"60","author":"Jones","year":"2008","journal-title":"Arthritis Rheum."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1101\/gad.231969.113","article-title":"Mechanical motion promotes expression of Prg4 in articular cartilage via multiple CREB-dependent, fluid flow shear stress-induced signaling pathways","volume":"28","author":"Ogawa","year":"2014","journal-title":"Genes Dev."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Yin, Y., Zhang, Y., Guo, L., Li, P., Wang, D., Huang, L., Zhao, X., Wu, G., Li, L., and Wei, X. (2023). Effect of Moderate Exercise on the Superficial Zone of Articular Cartilage in Age-Related Osteoarthritis. Diagnostics, 13.","DOI":"10.3390\/diagnostics13203193"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Ladner, Y.D., and Stoddart, M.J. (2024). Tribological loading of cartilage and chondrogenic cells. Biomater. Biosyst., 13.","DOI":"10.1016\/j.bbiosy.2024.100088"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1186\/s13075-019-2041-5","article-title":"Wnt\/\u03b2-catenin signaling contributes to articular cartilage homeostasis through lubricin induction in the superficial zone","volume":"21","author":"Xuan","year":"2019","journal-title":"Arthritis Res. Ther."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1136\/annrheumdis-2015-208577","article-title":"WNT16 antagonises excessive canonical WNT activation and protects cartilage in osteoarthritis","volume":"76","author":"Nalesso","year":"2017","journal-title":"Ann. Rheum. Dis."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Satake, T., Komura, S., Aoki, H., Hirakawa, A., Imai, Y., and Akiyama, H. (2022). Induction of iPSC-derived Prg4-positive cells with characteristics of superficial zone chondrocytes and fibroblast-like synovial cells. BMC Cell Biol., 23.","DOI":"10.1186\/s12860-022-00431-8"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"18910","DOI":"10.1038\/srep18910","article-title":"Lubricin\/Proteoglycan 4 binds to and regulates the activity of Toll-Like Receptors In Vitro","volume":"6","author":"Iqbal","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1186\/s13075-017-1301-5","article-title":"The autocrine role of proteoglycan-4 (PRG4) in modulating osteoarthritic synoviocyte proliferation and expression of matrix degrading enzymes","volume":"19","author":"Alquraini","year":"2017","journal-title":"Arthritis Res. Ther."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.ydbio.2017.04.006","article-title":"Cell origin, volume and arrangement are drivers of articular cartilage formation, morphogenesis and response to injury in mouse limbs","volume":"426","author":"Decker","year":"2017","journal-title":"Dev. Biol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1096\/fj.201600918R","article-title":"Superficial cells are self-renewing chondrocyte progenitors, which form the articular cartilage in juvenile mice","volume":"31","author":"Li","year":"2017","journal-title":"FASEB J."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1006\/excr.2002.5527","article-title":"Growth plate chondrocyte maturation is regulated by basal intracellular calcium","volume":"276","author":"Zuscik","year":"2002","journal-title":"Exp. Cell Res."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1002\/jor.23768","article-title":"Calcium signaling of in situ chondrocytes in articular cartilage under compressive loading: Roles of calcium sources and cell membrane ion channels","volume":"36","author":"Lv","year":"2017","journal-title":"J. Orthop. Res."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"726","DOI":"10.1002\/jbmr.3643","article-title":"Prevention of Injury-Induced Osteoarthritis in Rodent Temporomandibular Joint by Targeting Chondrocyte CaSR","volume":"34","author":"Zhang","year":"2019","journal-title":"J. Bone Miner. Res."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.semcdb.2015.12.004","article-title":"Interplay between CaSR and PTH1R signaling in skeletal development and osteoanabolism","volume":"49","author":"Maria","year":"2016","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"e23004","DOI":"10.1096\/fj.202300037RR","article-title":"CaSR modulates proliferation of the superficial zone cells in temporomandibular joint cartilage via the PTHrP nuclear localization sequence","volume":"37","author":"Zhou","year":"2023","journal-title":"FASEB J."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"522","DOI":"10.1038\/nrrheum.2017.127","article-title":"Regenerative medicine: Cartilage stem cells identified, but can they heal?","volume":"13","author":"Chagin","year":"2017","journal-title":"Nat. Rev. Rheumatol."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Liu, W., Feng, M., and Xu, P. (2022). From regeneration to osteoarthritis in the knee joint: The role shift of cartilage-derived progenitor cells. Front. Cell Dev. Biol., 10.","DOI":"10.3389\/fcell.2022.1010818"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"15040","DOI":"10.1038\/ncomms15040","article-title":"Joint morphogenetic cells in the adult mammalian synovium","volume":"8","author":"Roelofs","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"2893","DOI":"10.1172\/JCI83676","article-title":"Induced superficial chondrocyte death reduces catabolic cartilage damage in murine posttraumatic osteoarthritis","volume":"126","author":"Zhang","year":"2016","journal-title":"J. Clin. Investig."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1002\/jor.24812","article-title":"Effects of in vivo cyclic compressive loading on the distribution of local Col2 and superficial lubricin in rat knee cartilage","volume":"39","author":"Ji","year":"2021","journal-title":"J. Orthop. Res."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"2997","DOI":"10.1002\/art.24800","article-title":"The impact of anterior cruciate ligament injury on lubricin metabolism and the effect of inhibiting tumor necrosis factor \u03b1 on chondroprotection in an animal model","volume":"60","author":"Elsaid","year":"2009","journal-title":"Arthritis Rheum."},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Shiromoto, Y., Niki, Y., Kikuchi, T., Yoshihara, Y., Oguma, T., Nemoto, K., Chiba, K., Kanaji, A., Matsumoto, M., and Nakamura, M. (2022). Increased migratory activity and cartilage regeneration by superficial-zone chondrocytes in enzymatically treated cartilage explants. BMC Musculoskelet. Disord., 23.","DOI":"10.1186\/s12891-022-05210-2"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/25\/11\/5605\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T14:45:59Z","timestamp":1760107559000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/25\/11\/5605"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,5,21]]},"references-count":81,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2024,6]]}},"alternative-id":["ijms25115605"],"URL":"https:\/\/doi.org\/10.3390\/ijms25115605","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,5,21]]}}}