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Exos are endosome-derived membrane-bound vesicles 50\u2013150 nm in size. Exos can be general or cell type-specific. Their contents enable them to function as multi-signaling and vectorized vehicles. Exos are important for maintaining cellular homeostasis. They are released into extracellular spaces, leading to uptake by neighboring or distant cells and delivering their contents to modulate cell signaling. Exos influence tissue responses to injury, infection, and disease by fusion with the target cells and transferring their cargo, including cytokines, growth and angiogenic factors, signaling molecules, lipids, DNA, mRNAs, and non-coding RNAs. They are implicated in various physiological and pathological conditions, including ocular surface events, such as corneal scarring, wound healing, and inflammation. Their biocompatibility, stability, low immunogenicity, and easy detectability in bodily fluids (blood, tears, saliva, and urine) make them promising tools for diagnosing and treating ocular diseases. The potential to engineer specific Exo cargos makes them outstanding therapeutic delivery vehicles. The objective of this review is to provide novel insights into the functions of Exo cargos and their applications as biomarkers and therapeutics, or targets in the cornea.<\/jats:p>","DOI":"10.3390\/cells14130959","type":"journal-article","created":{"date-parts":[[2025,6,24]],"date-time":"2025-06-24T10:44:41Z","timestamp":1750761881000},"page":"959","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":13,"title":["Exosomes as Future Therapeutic Tools and Targets for Corneal Diseases"],"prefix":"10.3390","volume":"14","author":[{"given":"Joshua","family":"Gamez","sequence":"first","affiliation":[{"name":"Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daxian","family":"Zha","sequence":"additional","affiliation":[{"name":"Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shaghaiegh M.","family":"Ebrahimi","sequence":"additional","affiliation":[{"name":"Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Seok","family":"White","sequence":"additional","affiliation":[{"name":"Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2398-5319","authenticated-orcid":false,"given":"Alexander V.","family":"Ljubimov","sequence":"additional","affiliation":[{"name":"Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"David Geffen School of Medicine, University of California Los Angeles, Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4956-569X","authenticated-orcid":false,"given":"Mehrnoosh","family":"Saghizadeh","sequence":"additional","affiliation":[{"name":"Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, CA 90048, USA"},{"name":"David Geffen School of Medicine, University of California Los Angeles, Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,6,23]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/j.jtos.2016.04.002","article-title":"Anatomical features and cell-cell interactions in the human limbal epithelial stem cell niche","volume":"14","author":"Dziasko","year":"2016","journal-title":"Ocul. Surf."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1046\/j.1432-0436.1997.6140251.x","article-title":"Stem cells of the corneal epithelium lack connexins and metabolite transfer capacity","volume":"61","author":"Matic","year":"1997","journal-title":"Differentiation"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Dogru, M., Chen, M., Shimmura, S., and Tsubota, K. (2009). Corneal epithelium and stem cells. Corneal Surgery, Elsevier. [4th ed.].","DOI":"10.1016\/B978-0-323-04835-4.50010-0"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"101234","DOI":"10.1016\/j.preteyeres.2023.101234","article-title":"Squishy matters\u2014Corneal mechanobiology in health and disease","volume":"99","author":"Thomasy","year":"2024","journal-title":"Prog. Retin. Eye Res."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"835","DOI":"10.7150\/ijms.16563","article-title":"The limbal epithelial progenitors in the limbal niche environment","volume":"13","author":"Zhang","year":"2016","journal-title":"Int. J. Med. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"5130","DOI":"10.1167\/iovs.03-0584","article-title":"Stromal niche controls the plasticity of limbal and corneal epithelial differentiation in a rabbit model of recombined tissue","volume":"44","author":"Espana","year":"2003","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"e130","DOI":"10.1016\/S2214-109X(20)30425-3","article-title":"Trends in prevalence of blindness and distance and near vision impairment over 30 years: An analysis for the global burden of disease study","volume":"9","author":"Bourne","year":"2021","journal-title":"Lancet Glob. Health"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Oie, Y., and Nishida, K. (2013). Regenerative medicine for the cornea. Biomed. Res. Int., 2013.","DOI":"10.1155\/2013\/428247"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1097\/IIO.0b013e3181e20d64","article-title":"Management of ocular stem cell disease","volume":"50","author":"Biber","year":"2010","journal-title":"Int. Ophthalmol. Clin."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"767","DOI":"10.2217\/rme.11.94","article-title":"Ocular epithelial transplantation: Current uses and future potential","volume":"6","author":"Mason","year":"2011","journal-title":"Regen. Med."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1047","DOI":"10.1016\/j.freeradbiomed.2008.07.021","article-title":"Oxidative stress in diseases of the human cornea","volume":"45","author":"Shoham","year":"2008","journal-title":"Free Radic. Biol. Med."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1034\/j.1600-0854.2002.30502.x","article-title":"The biogenesis and functions of exosomes","volume":"3","author":"Stoorvogel","year":"2002","journal-title":"Traffic"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"eaau6977","DOI":"10.1126\/science.aau6977","article-title":"The biology, function, and biomedical applications of exosomes","volume":"367","author":"Kalluri","year":"2020","journal-title":"Science"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/bs.acc.2015.12.005","article-title":"Tumor-derived exosomes and their role in cancer progression","volume":"74","author":"Whiteside","year":"2016","journal-title":"Adv. Clin. Chem."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.tcb.2016.11.003","article-title":"Extracellular vesicles: Unique intercellular delivery vehicles","volume":"27","author":"Maas","year":"2017","journal-title":"Trends Cell Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1226","DOI":"10.1016\/j.cell.2016.01.043","article-title":"Communication by extracellular vesicles: Where we are and where we need to go","volume":"164","author":"Tkach","year":"2016","journal-title":"Cell"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Han, K.Y., Tran, J.A., Chang, J.H., Azar, D.T., and Zieske, J.D. (2017). Potential role of corneal epithelial cell-derived exosomes in corneal wound healing and neovascularization. Sci. Rep., 7.","DOI":"10.1038\/srep40548"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Leszczynska, A., Kulkarni, M., Ljubimov, A.V., and Saghizadeh, M. (2018). Exosomes from normal and diabetic human corneolimbal keratocytes differentially regulate migration, proliferation and marker expression of limbal epithelial cells. Sci. Rep., 8.","DOI":"10.1038\/s41598-018-33169-5"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"4564","DOI":"10.4049\/jimmunol.163.8.4564","article-title":"Ectosomes released by human neutrophils are specialized functional units","volume":"163","author":"Hess","year":"1999","journal-title":"J. Immunol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1146\/annurev-cellbio-101512-122326","article-title":"Biogenesis, secretion, and intercellular interactions of exosomes and other extracellular vesicles","volume":"30","author":"Colombo","year":"2014","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Wang, L., Hu, L., Zhou, X., Xiong, Z., Zhang, C., Shehada, H.M.A., Hu, B., Song, J., and Chen, L. (2017). Exosomes secreted by human adipose mesenchymal stem cells promote scarless cutaneous repair by regulating extracellular matrix remodelling. Sci. Rep., 7.","DOI":"10.1038\/s41598-017-12919-x"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"e12404","DOI":"10.1002\/jev2.12404","article-title":"Minimal information for studies of extracellular vesicles (MISEV2023): From basic to advanced approaches","volume":"13","author":"Welsh","year":"2024","journal-title":"J. Extracell. Vesicles"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Tiwari, A., Singh, A., Verma, S., Stephenson, S., Bhowmick, T., and Sangwan, V.S. (2021). Mini Review: Current trends and understanding of exosome therapeutic potential in corneal diseases. Front. Pharmacol., 12.","DOI":"10.3389\/fphar.2021.684712"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"9412","DOI":"10.1016\/S0021-9258(18)48095-7","article-title":"Vesicle formation during reticulocyte maturation. Association of plasma membrane activities with released vesicles (exosomes)","volume":"262","author":"Johnstone","year":"1987","journal-title":"J. Biol. Chem."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Schwarzenbach, H., and Gahan, P.B. (2021). Exosomes in immune regulation. Noncoding RNA, 7.","DOI":"10.3390\/ncrna7010004"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1111\/febs.15558","article-title":"Exosomes as mediators of immune regulation and immunotherapy in cancer","volume":"288","author":"Kugeratski","year":"2021","journal-title":"FEBS J."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"S\u00e1nchez-Alonso, S., Alcaraz-Serna, A., S\u00e1nchez-Madrid, F., and Alfranca, A. (2018). Extracellular vesicle-mediated immune regulation of tissue remodeling and angiogenesis after myocardial infarction. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.02799"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2038","DOI":"10.1080\/15476286.2021.1885189","article-title":"Exosomes provide unappreciated carrier effects that assist transfers of their miRNAs to targeted cells; I. They are \u2018The Elephant in the Room\u2019","volume":"18","author":"Askenase","year":"2021","journal-title":"RNA Biol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Zheng, D., Huo, M., Li, B., Wang, W., Piao, H., Wang, Y., Zhu, Z., Li, D., Wang, T., and Liu, K. (2020). The role of exosomes and exosomal microRNA in cardiovascular disease. Front. Cell Dev. Biol., 8.","DOI":"10.3389\/fcell.2020.616161"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1038\/aps.2017.178","article-title":"Exosomal cargo-loading and synthetic exosome-mimics as potential therapeutic tools","volume":"39","author":"Li","year":"2018","journal-title":"Acta Pharmacol. Sin."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1038\/ncomms1180","article-title":"Tumour microvesicles contain retrotransposon elements and amplified oncogene sequences","volume":"2","author":"Balaj","year":"2011","journal-title":"Nat. Commun."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"967","DOI":"10.1016\/0092-8674(83)90040-5","article-title":"Fate of the transferrin receptor during maturation of sheep reticulocytes in vitro: Selective externalization of the receptor","volume":"33","author":"Pan","year":"1983","journal-title":"Cell"},{"key":"ref_33","first-page":"256","article-title":"Endocytosis and intracellular processing of transferrin and colloidal gold-transferrin in rat reticulocytes: Demonstration of a pathway for receptor shedding","volume":"35","author":"Harding","year":"1984","journal-title":"Eur. J. Cell Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e12857","DOI":"10.1111\/cpr.12857","article-title":"Exosome biogenesis, secretion and function of exosomal miRNAs in skeletal muscle myogenesis","volume":"53","author":"Yue","year":"2020","journal-title":"Cell Prolif."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Jella, K.K., Nasti, T.H., Li, Z., Malla, S.R., Buchwald, Z.S., and Khan, M.K. (2018). Exosomes, their biogenesis and role in inter-cellular communication, tumor microenvironment and cancer immunotherapy. Vaccines, 6.","DOI":"10.3390\/vaccines6040069"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1744","DOI":"10.1016\/j.cmet.2021.08.006","article-title":"Exosomes as mediators of intercellular crosstalk in metabolism","volume":"33","author":"Isaac","year":"2021","journal-title":"Cell Metab."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1038\/nri855","article-title":"Exosomes: Composition, biogenesis and function","volume":"2","author":"Zitvogel","year":"2002","journal-title":"Nat. Rev. Immunol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Malenica, M., Vukomanovi\u0107, M., Kurtjak, M., Masciotti, V., Dal Zilio, S., Greco, S., Lazzarino, M., Kru\u0161i\u0107, V., Per\u010di\u0107, M., and Jelovica Badovinac, I. (2021). Perspectives of microscopy methods for morphology characterisation of extracellular vesicles from human biofluids. Biomedicines, 9.","DOI":"10.3390\/biomedicines9060603"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/0005-2736(81)90512-5","article-title":"Exfoliation of membrane ecto-enzymes in the form of micro-vesicles","volume":"645","author":"Trams","year":"1981","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1016\/j.bcmd.2005.03.002","article-title":"Revisiting the road to the discovery of exosomes","volume":"34","author":"Johnstone","year":"2005","journal-title":"Blood Cells Mol. Dis."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Xie, S., Zhang, Q., and Jiang, L. (2022). Current knowledge on exosome biogenesis, cargo-sorting mechanism and therapeutic implications. Membranes, 12.","DOI":"10.3390\/membranes12050498"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Krylova, S.V., and Feng, D. (2023). The machinery of exosomes: Biogenesis, release, and uptake. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24021337"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1016\/j.cell.2019.02.029","article-title":"Reassessment of exosome composition","volume":"177","author":"Jeppesen","year":"2019","journal-title":"Cell"},{"key":"ref_44","first-page":"5553","article-title":"Analysis of ESCRT functions in exosome biogenesis, composition and secretion highlights the heterogeneity of extracellular vesicles","volume":"126","author":"Colombo","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1038\/ncb2502","article-title":"Syndecan-syntenin-ALIX regulates the biogenesis of exosomes","volume":"14","author":"Baietti","year":"2012","journal-title":"Nat. Cell Biol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"13588","DOI":"10.1038\/ncomms13588","article-title":"ISGylation controls exosome secretion by promoting lysosomal degradation of MVB proteins","volume":"7","author":"Baixauli","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Jin, H., Tang, Y., Yang, L., Peng, X., Li, B., Fan, Q., Wei, S., Yang, S., Li, X., and Wu, B. (2021). Rab GTPases: Central coordinators of membrane trafficking in cancer. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.648384"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1126\/science.1161748","article-title":"Membrane fusion: Grappling with SNARE and SM proteins","volume":"323","author":"Rothman","year":"2009","journal-title":"Science"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1038\/s12276-024-01209-y","article-title":"Regulation of cargo selection in exosome biogenesis and its biomedical applications in cancer","volume":"56","author":"Lee","year":"2024","journal-title":"Exp. Mol. Med."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"925","DOI":"10.1080\/1061186X.2021.1894436","article-title":"An updated review on exosomes: Biosynthesis to clinical applications","volume":"29","author":"Modani","year":"2021","journal-title":"J. Drug Target."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"100029","DOI":"10.1016\/j.vesic.2023.100029","article-title":"Guidelines for clinical translation and commercialization of extracellular vesicles and exosomes based therapeutics","volume":"2","author":"Cheng","year":"2023","journal-title":"Extracell. Vesicle"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Ramos, T., Parekh, M., Kaye, S.B., and Ahmad, S. (2022). Epithelial cell-derived extracellular vesicles trigger the differentiation of two epithelial cell lines. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23031718"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Braunsperger, M.V., Martin, G., Herzig, T., Ku\u00dfberger, I., Gie\u00dfl, A., Steimle, S., Schl\u00f6tzer-Schrehardt, U., Schlunck, G., Reinhard, T., and Polisetti, N. (2025). Proteomic insights into human limbal epithelial progenitor-derived small extracellular vesicles. Stem Cell Rev. Rep.","DOI":"10.1007\/s12015-025-10877-w"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Zeng, H., Guo, S., Ren, X., Wu, Z., Liu, S., and Yao, X. (2023). Current strategies for exosome cargo loading and targeting delivery. Cells, 12.","DOI":"10.3390\/cells12101416"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1186\/s12943-019-0984-4","article-title":"The role of exosomal noncoding RNAs in cancer","volume":"18","author":"Xie","year":"2019","journal-title":"Mol. Cancer"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1016\/j.addr.2012.06.011","article-title":"Exosomes as nucleic acid nanocarriers","volume":"65","author":"Dassler","year":"2013","journal-title":"Adv. Drug Deliv. Rev."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1658","DOI":"10.1161\/CIRCRESAHA.117.309681","article-title":"Extracellular vesicles in angiogenesis","volume":"120","author":"Todorova","year":"2017","journal-title":"Circ. Res."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Jung, H.H., Kim, J.Y., Lim, J.E., and Im, Y.H. (2020). Cytokine profiling in serum-derived exosomes isolated by different methods. Sci. Rep., 10.","DOI":"10.1038\/s41598-020-70584-z"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Hyv\u00e4rinen, K., Holopainen, M., Skirdenko, V., Ruhanen, H., Lehenkari, P., Korhonen, M., K\u00e4kel\u00e4, R., Laitinen, S., and Kerkel\u00e4, E. (2018). Mesenchymal stromal cells and their extracellular vesicles enhance the anti-inflammatory phenotype of regulatory macrophages by downregulating the production of interleukin (IL)-23 and IL-22. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.00771"},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Wang, L., and Zhang, L. (2020). Circulating exosomal miRNA as diagnostic biomarkers of neurodegenerative diseases. Front. Mol. Neurosci., 13.","DOI":"10.3389\/fnmol.2020.00053"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1186\/s12943-018-0831-z","article-title":"Emerging role of exosome-derived long non-coding RNAs in tumor microenvironment","volume":"17","author":"Sun","year":"2018","journal-title":"Mol. Cancer"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1186\/s12943-019-1041-z","article-title":"Exosomal circRNAs: Biogenesis, effect and application in human diseases","volume":"18","author":"Wang","year":"2019","journal-title":"Mol. Cancer"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1016\/j.plipres.2017.03.001","article-title":"Lipids in exosomes: Current knowledge and the way forward","volume":"66","author":"Skotland","year":"2017","journal-title":"Prog. Lipid Res."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"D1241","DOI":"10.1093\/nar\/gkr828","article-title":"ExoCarta 2012: Database of exosomal proteins, RNA and lipids","volume":"40","author":"Mathivanan","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Gao, J., Li, A., Hu, J., Feng, L., Liu, L., and Shen, Z. (2023). Recent developments in isolating methods for exosomes. Front. Bioeng. Biotechnol., 10.","DOI":"10.3389\/fbioe.2022.1100892"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"70725","DOI":"10.7554\/eLife.70725","article-title":"Framework for rapid comparison of extracellular vesicle isolation methods","volume":"10","author":"Norman","year":"2021","journal-title":"eLife"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Kim, K., Park, J., Jung, J.H., Lee, R., Park, J.H., Yuk, J.M., Hwang, H., and Yeon, J.H. (2021). Cyclic tangential flow filtration system for isolation of extracellular vesicles. APL Bioeng., 5.","DOI":"10.1063\/5.0037768"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1007\/978-1-0716-1205-7_23","article-title":"Immunoaffinity-based isolation of melanoma cell-derived and T cell-derived exosomes from plasma of melanoma patients","volume":"2265","author":"Mondal","year":"2021","journal-title":"Methods Mol. Biol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"102396","DOI":"10.1016\/j.nano.2021.102396","article-title":"Comparative study of commercial protocols for high recovery of high-purity mesenchymal stem cell-derived extracellular vesicle isolation and their efficient labeling with fluorescent dyes","volume":"35","author":"Kamei","year":"2021","journal-title":"Nanomedicine"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"e12145","DOI":"10.1002\/jev2.12145","article-title":"Establishment of a simplified dichotomic size-exclusion chromatography for isolating extracellular vesicles toward clinical applications","volume":"10","author":"Guo","year":"2021","journal-title":"J. Extracell. Vesicles"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"e2103222","DOI":"10.1002\/advs.202103222","article-title":"Exosome processing and characterization approaches for research and technology development","volume":"9","author":"Lai","year":"2022","journal-title":"Adv. Sci."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Livshits, M.A., Khomyakova, E., Evtushenko, E.G., Lazarev, V.N., Kulemin, N.A., Semina, S.E., Generozov, E.V., and Govorun, V.M. (2015). Isolation of exosomes by differential centrifugation: Theoretical analysis of a commonly used protocol. Sci. Rep., 5.","DOI":"10.1038\/srep17319"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1007\/978-1-0716-1205-7_22","article-title":"A rapid exosome isolation using ultrafiltration and size exclusion chromatography (REIUS) method for exosome isolation from melanoma cell lines","volume":"2265","author":"Shu","year":"2021","journal-title":"Methods Mol. Biol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"2217","DOI":"10.1021\/acs.jproteome.9b00693","article-title":"Characterization of urinary exosomes purified with size exclusion chromatography and ultracentrifugation","volume":"19","author":"Guan","year":"2020","journal-title":"J. Proteome Res."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"e13904","DOI":"10.1111\/cts.13904","article-title":"Regulation of exosomes as biologic medicines: Regulatory challenges faced in exosome development and manufacturing processes","volume":"17","author":"Wang","year":"2024","journal-title":"Clin. Transl. Sci."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"e1008347","DOI":"10.1080\/2162402X.2015.1008347","article-title":"Exosomes isolated from plasma of glioma patients enrolled in a vaccination trial reflect antitumor immune activity and might predict survival","volume":"4","author":"Muller","year":"2015","journal-title":"Oncoimmunology"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Ludwig, N., Whiteside, T.L., and Reichert, T.E. (2019). Challenges in exosome isolation and analysis in health and disease. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20194684"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Wu, Y., Wang, Y., Lu, Y., Luo, X., Huang, Y., Xie, T., Pilarsky, C., Dang, Y., and Zhang, J. (2022). Microfluidic technology for the isolation and analysis of exosomes. Micromachines, 13.","DOI":"10.3390\/mi13101571"},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Kang, Y.T., Hadlock, T., Jolly, S., and Nagrath, S. (2020). Extracellular vesicles on demand (EVOD) chip for screening and quantification of cancer-associated extracellular vesicles. Biosens. Bioelectron., 168.","DOI":"10.1016\/j.bios.2020.112535"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"834","DOI":"10.3892\/ijmm.2017.3080","article-title":"Comparison of isolation methods of exosomes and exosomal RNA from cell culture medium and serum","volume":"40","author":"Tang","year":"2017","journal-title":"Int. J. Mol. Med."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"32945","DOI":"10.3402\/jev.v5.32945","article-title":"Techniques used for the isolation and characterization of extracellular vesicles: Results of a worldwide survey","volume":"5","author":"Gardiner","year":"2016","journal-title":"J. Extracell. Vesicles"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"212","DOI":"10.1038\/s41592-020-01034-x","article-title":"Exosome detection via the ultrafast-isolation system: EXODUS","volume":"18","author":"Chen","year":"2021","journal-title":"Nat. Methods"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Serrano-Pertierra, E., Oliveira-Rodr\u00edguez, M., Rivas, M., Oliva, P., Villafani, J., Navarro, A., Blanco-L\u00f3pez, M.C., and Cernuda-Moroll\u00f3n, E. (2019). Characterization of plasma-derived extracellular vesicles isolated by different methods: A comparison study. Bioengineering, 6.","DOI":"10.3390\/bioengineering6010008"},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Logozzi, M., Mizzoni, D., Di Raimo, R., and Fais, S. (2020). Exosomes: A source for new and old biomarkers in cancer. Cancers, 12.","DOI":"10.3390\/cancers12092566"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1038\/s12276-024-01201-6","article-title":"Recent advances in extracellular vesicles for therapeutic cargo delivery","volume":"56","author":"Kim","year":"2024","journal-title":"Exp. Mol. Med."},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Hamzah, R.N., Alghazali, K.M., Biris, A.S., and Griffin, R.J. (2021). Exosome traceability and cell source dependence on composition and cell-cell cross talk. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22105346"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1007\/s10238-023-01282-z","article-title":"Therapeutic potential of mesenchymal stem cell-derived exosomes for regenerative medicine applications","volume":"24","author":"Roszkowski","year":"2024","journal-title":"Clin. Exp. Med."},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Olejarz, W., Kubiak-Tomaszewska, G., Chrzanowska, A., and Lorenc, T. (2020). Exosomes in angiogenesis and anti-angiogenic therapy in cancers. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21165840"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"532","DOI":"10.1002\/ana.22615","article-title":"Pathogenic protein seeding in Alzheimer disease and other neurodegenerative disorders","volume":"70","author":"Jucker","year":"2011","journal-title":"Ann. Neurol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1167\/tvst.10.2.12","article-title":"The role of deregulated microRNAs in age-related macular degeneration pathology","volume":"10","author":"ElShelmani","year":"2021","journal-title":"Transl. Vis. Sci. Technol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"877","DOI":"10.2174\/1386207324666210216092917","article-title":"Extracellular vesicle-derived miR-26b-5p is up-regulated in the serum of patients with diabetic retinopathy","volume":"25","author":"Zhang","year":"2022","journal-title":"Comb. Chem. High. Throughput Screen."},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Chen, C., Wu, Y., Li, M., Cui, C., Zhao, Y., Sun, X., Wang, Y., Liu, C., Wu, H., and Zhong, X. (2020). Different Exosomal microRNA Profile in Aquaporin-4 Antibody Positive Neuromyelitis Optica Spectrum Disorders. Front. Immunol., 11.","DOI":"10.3389\/fimmu.2020.01064"},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Wang, L., and Wei, X. (2023). Exosome-based crosstalk in glaucoma pathogenesis: A focus on oxidative stress and neuroinflammation. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1202704"},{"key":"ref_94","doi-asserted-by":"crossref","unstructured":"Yeung, V., Boychev, N., Farhat, W., Ntentakis, D.P., Hutcheon, A.E.K., Ross, A.E., and Ciolino, J.B. (2022). Extracellular vesicles in corneal fibrosis\/scarring. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23115921"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"5194","DOI":"10.1167\/iovs.18-24803","article-title":"Effect of human corneal mesenchymal stromal cell-derived exosomes on corneal epithelial wound healing","volume":"59","author":"Samaeekia","year":"2018","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Verma, N., Khare, D., Poe, A.J., Amador, C., Ghiam, S., Fealy, A., Ebrahimi, S., Shadrokh, O., Song, X.Y., and Santiskulvong, C. (2023). MicroRNA and protein cargos of human limbal epithelial cell-derived exosomes and their regulatory roles in limbal stromal cells of diabetic and non-diabetic corneas. Cells, 12.","DOI":"10.3390\/cells12212524"},{"key":"ref_97","doi-asserted-by":"crossref","unstructured":"Desjardins, P., Berthiaume, R., Couture, C., Le-Bel, G., Roy, V., Gros-Louis, F., Moulin, V.J., Proulx, S., Chemtob, S., and Germain, L. (2022). Impact of exosomes released by different corneal cell types on the wound healing properties of human corneal epithelial cells. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232012201"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"149898","DOI":"10.1016\/j.cej.2024.149898","article-title":"Milk-derived exosome as delivery system for lutein encapsulation in alleviating dry eye disease","volume":"486","author":"Wu","year":"2024","journal-title":"Chem. Eng. J."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1016\/j.jtos.2024.09.006","article-title":"Development of human amniotic epithelial cell-derived extracellular vesicles as cell-free therapy for dry eye disease","volume":"34","author":"Yi","year":"2024","journal-title":"Ocul. Surf."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1167\/iovs.65.6.16","article-title":"Mesenchymal stem cell-derived exosomes as drug carriers for delivering miRNA-29b to ameliorate inflammation in corneal injury via activating autophagy","volume":"65","author":"Liu","year":"2024","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/j.jtos.2025.01.012","article-title":"Application of stem cell-derived exosomes in anterior segment eye diseases: A comprehensive update review","volume":"36","author":"Hashemian","year":"2025","journal-title":"Ocul. Surf."},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Chan, S.M., Tsai, C., Lee, T.P., Huang, Z.R., Huang, W.H., and Lin, C.T. (2025). Therapeutic potential of umbilical cord MSC-derived exosomes in a severe dry eye rat model: Enhancing corneal protection and modulating inflammation. Biomedicines, 13.","DOI":"10.3390\/biomedicines13051174"},{"key":"ref_103","doi-asserted-by":"crossref","unstructured":"Massoumi, H., Amin, S., Soleimani, M., Momenaei, B., Ashraf, M.J., Guaiquil, V.H., Hematti, P., Rosenblatt, M.I., Djalilian, A.R., and Jalilian, E. (2023). Extracellular-vesicle-based therapeutics in neuro-ophthalmic disorders. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24109006"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1016\/j.jtos.2024.10.005","article-title":"Target specification and therapeutic potential of extracellular vesicles for regulating corneal angiogenesis, lymphangiogenesis, and nerve repair","volume":"34","author":"Pedersen","year":"2024","journal-title":"Ocul. Surf."},{"key":"ref_105","doi-asserted-by":"crossref","unstructured":"Ciferri, M.C., Quarto, R., and Tasso, R. (2021). Extracellular vesicles as biomarkers and therapeutic tools: From pre-clinical to clinical applications. Biology, 10.","DOI":"10.3390\/biology10050359"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"11720","DOI":"10.1021\/acsnano.2c02531","article-title":"Discovering the secret of diseases by incorporated tear exosomes analysis","volume":"16","author":"Hu","year":"2022","journal-title":"ACS Nano"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1186\/s13075-017-1228-x","article-title":"Identification of potential saliva and tear biomarkers in primary Sj\u00f6gren\u2019s syndrome, utilising the extraction of extracellular vesicles and proteomics analysis","volume":"19","author":"Aqrawi","year":"2017","journal-title":"Arthritis Res. Ther."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Amorim, M., Martins, B., Caramelo, F., Gon\u00e7alves, C., Trindade, G., Sim\u00e3o, J., Barreto, P., Marques, I., Leal, E.C., and Carvalho, E. (2022). Putative biomarkers in tears for diabetic retinopathy diagnosis. Front. Med., 9.","DOI":"10.3389\/fmed.2022.873483"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"109242","DOI":"10.1016\/j.exer.2022.109242","article-title":"Revealing the presence of tear extracellular vesicles in keratoconus","volume":"224","author":"Hefley","year":"2022","journal-title":"Exp. Eye Res."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1186\/s41544-019-0025-x","article-title":"Exosomal miRNA: An alternative mediator of cell-to-cell communication","volume":"1","author":"Li","year":"2019","journal-title":"ExRNA"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.gpb.2015.02.001","article-title":"Exosome and exosomal microRNA: Trafficking, sorting, and function","volume":"13","author":"Zhang","year":"2015","journal-title":"Genom. Proteom. Bioinform."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"685","DOI":"10.1002\/bit.28029","article-title":"Biogenesis and mechanisms of microRNA-mediated gene regulation","volume":"119","author":"Rani","year":"2022","journal-title":"Biotechnol. Bioeng."},{"key":"ref_113","doi-asserted-by":"crossref","unstructured":"Dexheimer, P.J., and Cochella, L. (2020). MicroRNAs: From mechanism to organism. Front. Cell Dev. Biol., 8.","DOI":"10.3389\/fcell.2020.00409"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1038\/s41392-021-00779-x","article-title":"Roles and mechanisms of exosomal non-coding RNAs in human health and diseases","volume":"6","author":"Li","year":"2021","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"2111","DOI":"10.1007\/s11010-021-04053-0","article-title":"The role of exosomal microRNAs in central nervous system diseases","volume":"476","author":"Yu","year":"2021","journal-title":"Mol. Cell Biochem."},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Peng, J., Liang, Q., Xu, Z., Cai, Y., Peng, B., Li, J., Zhang, W., Kang, F., Hong, Q., and Yan, Y. (2021). Current understanding of exosomal microRNAs in glioma immune regulation and therapeutic responses. Front. Immunol., 12.","DOI":"10.3389\/fimmu.2021.813747"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.molimm.2017.11.008","article-title":"BMSCs-derived miR-223-containing exosomes contribute to liver protection in experimental autoimmune hepatitis","volume":"93","author":"Chen","year":"2018","journal-title":"Mol. Immunol."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1016\/j.preteyeres.2017.04.004","article-title":"Roles of exosomes in the normal and diseased eye","volume":"59","author":"Klingeborn","year":"2017","journal-title":"Prog. Retin. Eye Res."},{"key":"ref_119","doi-asserted-by":"crossref","unstructured":"Winkler, M.A., Dib, C., Ljubimov, A.V., and Saghizadeh, M. (2014). Targeting miR-146a to treat delayed wound healing in human diabetic organ-cultured corneas. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0114692"},{"key":"ref_120","doi-asserted-by":"crossref","unstructured":"Stunf Pukl, S. (2022). MicroRNA of epithelial to mesenchymal transition in Fuchs\u2019 endothelial corneal dystrophy. Genes, 13.","DOI":"10.3390\/genes13101711"},{"key":"ref_121","doi-asserted-by":"crossref","unstructured":"Kulkarni, M., Leszczynska, A., Wei, G., Winkler, M.A., Tang, J., Funari, V.A., Deng, N., Liu, Z., Punj, V., and Deng, S.X. (2017). Genome-wide analysis suggests a differential microRNA signature associated with normal and diabetic human corneal limbus. Sci. Rep., 7.","DOI":"10.1038\/s41598-017-03449-7"},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"7939","DOI":"10.1167\/iovs.15-17619","article-title":"Human corneal microRNA expression profile in fungal keratitis","volume":"56","author":"Boomiraj","year":"2015","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1167\/iovs.13-12689","article-title":"Endothelial cell microRNA expression in human late-onset Fuchs\u2019 dystrophy","volume":"55","author":"Matthaei","year":"2014","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"10519","DOI":"10.2147\/IJN.S277190","article-title":"Roles of exosomes in ocular diseases","volume":"15","author":"Liu","year":"2020","journal-title":"Int. J. Nanomed."},{"key":"ref_125","first-page":"640","article-title":"MiRNA 24-3p-rich exosomes functionalized DEGMA-modified hyaluronic acid hydrogels for corneal epithelial healing","volume":"25","author":"Sun","year":"2023","journal-title":"Bioact. Mater."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"1091","DOI":"10.1007\/s13577-024-01082-x","article-title":"DNMT1 driven by mouse amniotic fluid mesenchymal stem cell exosomes improved corneal cryoinjury via inducing microRNA-33 promoter DNA hypermethylation modification in corneal epithelium cells","volume":"37","author":"Xu","year":"2024","journal-title":"Hum. Cell"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1002\/biof.2085","article-title":"Crosstalk among miR-29 \u03b1-SMA, and TGF\u03b21\/\u03b23 in melatonin-induced exosome (Mel-prExo) treated human limbal mesenchymal stem cells (hLMSCs): An insight into scarless healing of the cornea","volume":"50","author":"Altug","year":"2024","journal-title":"Biofactors"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"904","DOI":"10.1080\/02713683.2019.1707229","article-title":"MiRNA-155-5p reduces corneal epithelial permeability by remodeling epithelial tight junctions during corneal wound healing","volume":"45","author":"Wang","year":"2020","journal-title":"Curr. Eye Res."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"4093","DOI":"10.18632\/aging.102802","article-title":"Exosomal miR-19a from adipose-derived stem cells suppresses differentiation of corneal keratocytes into myofibroblasts","volume":"12","author":"Shen","year":"2020","journal-title":"Aging"},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1167\/iovs.65.12.1","article-title":"Mesenchymal stem cells-derived exosomal miR-223-3p alleviates ocular surface damage and inflammation by downregulating Fbxw7 in dry eye models","volume":"65","author":"Wang","year":"2024","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"1252557","DOI":"10.1155\/2022\/1252557","article-title":"Umbilical cord mesenchymal stem cell-derived small extracellular vesicles deliver miR-21 to promote corneal epithelial wound healing through PTEN\/PI3K\/Akt pathway","volume":"2022","author":"Liu","year":"2022","journal-title":"Stem Cells Int."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"5043","DOI":"10.1002\/jcb.26706","article-title":"The exosomes released from different cell types and their effects in wound healing","volume":"119","author":"Golchin","year":"2018","journal-title":"J. Cell Biochem."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1021\/acs.molpharmaceut.2c00414","article-title":"Extracellular vesicles as biomarkers and therapeutics for inflammatory eye diseases","volume":"20","author":"Louie","year":"2023","journal-title":"Mol. Pharm."},{"key":"ref_134","doi-asserted-by":"crossref","unstructured":"Marote, A., Teixeira, F.G., Mendes-Pinheiro, B., and Salgado, A.J. (2016). MSCs-derived exosomes: Cell-secreted nanovesicles with regenerative potential. Front. Pharmacol., 7.","DOI":"10.3389\/fphar.2016.00231"},{"key":"ref_135","doi-asserted-by":"crossref","unstructured":"Jin, Q.H., Kim, H.K., Na, J.Y., Jin, C., and Seon, J.K. (2022). Anti-inflammatory effects of mesenchymal stem cell-conditioned media inhibited macrophages activation in vitro. Sci. Rep., 12.","DOI":"10.1038\/s41598-022-08398-4"},{"key":"ref_136","doi-asserted-by":"crossref","unstructured":"Ong, H.S., Riau, A.K., Yam, G.H., Yusoff, N.Z.B.M., Han, E.J.Y., Goh, T.W., Lai, R.C., Lim, S.K., and Mehta, J.S. (2023). Mesenchymal stem cell exosomes as immunomodulatory therapy for corneal scarring. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24087456"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1186\/s13287-020-02129-0","article-title":"Subconjunctival injection of mesenchymal stem cells for corneal failure due to limbal stem cell deficiency: State of the art","volume":"12","author":"Galindo","year":"2021","journal-title":"Stem Cell Res. Ther."},{"key":"ref_138","doi-asserted-by":"crossref","unstructured":"Buono, L., Scalabrin, S., De Iuliis, M., Tanzi, A., Grange, C., Tapparo, M., Nuzzi, R., and Bussolati, B. (2021). Mesenchymal stem cell-derived extracellular vesicles protect human corneal endothelial cells from endoplasmic reticulum stress-mediated apoptosis. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22094930"},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"1192","DOI":"10.1002\/sctm.18-0297","article-title":"Mesenchymal stem cells reduce corneal fibrosis and inflammation via extracellular vesicle-mediated delivery of miRNA","volume":"8","author":"Shojaati","year":"2019","journal-title":"Stem Cells Transl. Med."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"3142","DOI":"10.1182\/blood-2011-11-391144","article-title":"Allogeneic mesenchymal stem cell treatment alleviates experimental and clinical Sj\u00f6gren syndrome","volume":"120","author":"Xu","year":"2012","journal-title":"Blood"},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"2347","DOI":"10.1038\/mt.2012.208","article-title":"Mesenchymal stromal cells treatment attenuates dry eye in patients with chronic graft-versus-host disease","volume":"20","author":"Weng","year":"2012","journal-title":"Mol. Ther."},{"key":"ref_142","doi-asserted-by":"crossref","unstructured":"Kistenmacher, S., Schw\u00e4mmle, M., Martin, G., Ulrich, E., Tholen, S., Schilling, O., Gie\u00dfl, A., Schl\u00f6tzer-Schrehardt, U., Bucher, F., and Schlunck, G. (2024). Enrichment, characterization, and proteomic profiling of small extracellular vesicles derived from human limbal mesenchymal stromal cells and melanocytes. Cells, 13.","DOI":"10.3390\/cells13070623"},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"Ortiz-Melo, M.T., Campos, J.E., S\u00e1nchez-Guzm\u00e1n, E., Herrera-Aguirre, M.E., and Castro-Mu\u00f1ozledo, F. (2024). Regulation of corneal epithelial differentiation: miR-141-3p promotes the arrest of cell proliferation and enhances the expression of terminal phenotype. PLoS ONE, 19.","DOI":"10.1371\/journal.pone.0315296"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"e2304381","DOI":"10.1002\/adhm.202304381","article-title":"Tetrahedral DNA-based functional microRNA-21 delivery system: Application to corneal epithelial wound healing","volume":"13","author":"Li","year":"2024","journal-title":"Adv. Health Mater."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"7137","DOI":"10.2147\/IJN.S464249","article-title":"Engineering exosomes for therapeutic applications: Decoding biogenesis, content modification, and cargo loading strategies","volume":"19","author":"Bahadorani","year":"2024","journal-title":"Int. J. Nanomed."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"1453","DOI":"10.1038\/s41551-024-01214-6","article-title":"Antibody-displaying extracellular vesicles for targeted cancer therapy","volume":"8","author":"Wiklander","year":"2024","journal-title":"Nat. Biomed. Eng."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"e12248","DOI":"10.1002\/jev2.12248","article-title":"Extracellular vesicles engineered to bind albumin demonstrate extended circulation time and lymph node accumulation in mouse models","volume":"11","author":"Liang","year":"2022","journal-title":"J. Extracell. Vesicles"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1016\/j.cell.2021.12.021","article-title":"Engineered virus-like particles for efficient in vivo delivery of therapeutic proteins","volume":"185","author":"Banskota","year":"2022","journal-title":"Cell"},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"4028","DOI":"10.1038\/s41467-025-59377-y","article-title":"Engineering of extracellular vesicles for efficient intracellular delivery of multimodal therapeutics including genome editors","volume":"16","author":"Liang","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1007\/s13346-024-01710-x","article-title":"Exosome-based drug delivery systems for enhanced neurological therapeutics","volume":"15","author":"Vahab","year":"2025","journal-title":"Drug Deliv. Transl. Res."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"e130","DOI":"10.1093\/nar\/gks463","article-title":"Plasma exosomes can deliver exogenous short interfering RNA to monocytes and lymphocytes","volume":"40","author":"Wahlgren","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_152","doi-asserted-by":"crossref","unstructured":"Lozano, V., Mart\u00edn, C., Blanco, N., Alcalde, I., Fernandez-Vega Cueto, L., Merayo-Lloves, J., and Quir\u00f3s, L.M. (2022). Exosomes released by corneal stromal cells show molecular alterations in keratoconus patients and induce different cellular behavior. Biomedicines, 10.","DOI":"10.3390\/biomedicines10102348"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1186\/s41232-023-00260-y","article-title":"Accelerating corneal wound healing using exosome-mediated targeting of NF-\u03baB c-Rel","volume":"43","author":"Zhao","year":"2023","journal-title":"Inflamm. Regen."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"e26800","DOI":"10.1016\/j.heliyon.2024.e26800","article-title":"EGF-conditioned M1 macrophages convey reduced inflammation into corneal endothelial cells through exosomes","volume":"10","author":"Lee","year":"2024","journal-title":"Heliyon"},{"key":"ref_155","doi-asserted-by":"crossref","unstructured":"Choudhery, M.S., Arif, T., Mahmood, R., and Harris, D.T. (2024). Stem cell-based acellular therapy: Insight into biogenesis, bioengineering and therapeutic applications of exosomes. Biomolecules, 14.","DOI":"10.3390\/biom14070792"},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"1430","DOI":"10.7150\/ijbs.80430","article-title":"Exosomes and their bioengineering strategies in the cutaneous wound healing and related complications: Current knowledge and future perspectives","volume":"19","author":"Yang","year":"2023","journal-title":"Int. J. Biol. Sci."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"19546","DOI":"10.18632\/aging.103904","article-title":"Comparison of exosomes derived from induced pluripotent stem cells and mesenchymal stem cells as therapeutic nanoparticles for treatment of corneal epithelial defects","volume":"12","author":"Wang","year":"2020","journal-title":"Aging"},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"7384","DOI":"10.1039\/D4TB00849A","article-title":"Cationic-motif-modified exosomes for mRNA delivery to retinal photoreceptors","volume":"12","author":"Pathrikar","year":"2024","journal-title":"J. Mater. Chem. B"},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Chen, X., Li, K., Chen, J., and Tan, S. (2023). Breakthrough in large-scale production of iPSCs-derived exosomes to promote clinical applications. Front. Bioeng. Biotechnol., 11.","DOI":"10.3389\/fbioe.2023.1257186"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"875","DOI":"10.1021\/acs.analchem.1c03644","article-title":"Precise molecular profiling of circulating exosomes using a metal-organic framework-based sensing interface and an enzyme-based electrochemical logic platform","volume":"94","author":"Wang","year":"2022","journal-title":"Anal. Chem."},{"key":"ref_161","doi-asserted-by":"crossref","unstructured":"Mitchell, M.I., Ma, J., Carter, C.L., and Loudig, O. (2022). Circulating exosome cargoes contain functionally diverse cancer biomarkers: From biogenesis and function to purification and potential translational utility. Cancers, 14.","DOI":"10.3390\/cancers14143350"},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.coph.2016.06.003","article-title":"Vesicle-MaNiA: Extracellular vesicles in liquid biopsy and cancer","volume":"29","author":"Torrano","year":"2016","journal-title":"Curr. Opin. Pharmacol."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"30867","DOI":"10.18632\/oncotarget.9018","article-title":"Tumour-associated circulating microparticles: A novel liquid biopsy tool for screening and therapy monitoring of colorectal carcinoma and other epithelial neoplasia","volume":"7","author":"Willms","year":"2016","journal-title":"Oncotarget"},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"3180","DOI":"10.1038\/s41433-023-02473-4","article-title":"Role of tear exosomes in the spread of herpes simplex virus type 1 in recurrent herpes simplex keratitis","volume":"37","author":"Huang","year":"2023","journal-title":"Eye"},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.blre.2012.12.002","article-title":"Extracellular vesicles in physiological and pathological conditions","volume":"27","author":"Yuana","year":"2013","journal-title":"Blood Rev."},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"1703244","DOI":"10.1080\/20013078.2019.1703244","article-title":"Inhibiting extracellular vesicles formation and release: A review of EV inhibitors","volume":"9","author":"Catalano","year":"2020","journal-title":"J. Extracell. Vesicles"},{"key":"ref_167","doi-asserted-by":"crossref","unstructured":"Jorfi, S., Ansa-Addo, E.A., Kholia, S., Stratton, D., Valley, S., Lange, S., and Inal, J. (2015). Inhibition of microvesiculation sensitizes prostate cancer cells to chemotherapy and reduces docetaxel dose required to limit tumor growth in vivo. Sci. Rep., 5.","DOI":"10.1038\/srep13006"},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"1803","DOI":"10.1111\/cas.13310","article-title":"Accelerated growth of B16BL6 tumor in mice through efficient uptake of their own exosomes by B16BL6 cells","volume":"108","author":"Matsumoto","year":"2017","journal-title":"Cancer Sci."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.bbrc.2004.05.127","article-title":"Statin decreases endothelial microparticle release from human coronary artery endothelial cells: Implication for the Rho-kinase pathway","volume":"320","author":"Tramontano","year":"2004","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.canlet.2017.08.020","article-title":"Manumycin A suppresses exosome biogenesis and secretion via targeted inhibition of Ras\/Raf\/ERK1\/2 signaling and hnRNP H1 in castration-resistant prostate cancer cells","volume":"408","author":"Datta","year":"2017","journal-title":"Cancer Lett."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"3455","DOI":"10.1074\/jbc.M114.605253","article-title":"The neutral sphingomyelinase pathway regulates packaging of the prion protein into exosomes","volume":"290","author":"Guo","year":"2015","journal-title":"J. Biol. Chem."},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.jbior.2014.10.002","article-title":"Roles and regulation of neutral sphingomyelinase-2 in cellular and pathological processes","volume":"57","author":"Shamseddine","year":"2015","journal-title":"Adv. Biol. Regul."},{"key":"ref_173","doi-asserted-by":"crossref","unstructured":"Hagemann, A., Altrogge, P.K., Kehrenberg, M.C.A., Diehl, D., Jung, D., Weber, L., and Bachmann, H.S. (2022). Analyzing the postulated inhibitory effect of Manumycin A on farnesyltransferase. Front. Chem., 10.","DOI":"10.3389\/fchem.2022.967947"},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"F963","DOI":"10.1152\/ajprenal.00078.2017","article-title":"Exosome production and its regulation of EGFR during wound healing in renal tubular cells","volume":"312","author":"Zhou","year":"2017","journal-title":"Am. J. Physiol. Ren. Physiol."},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/j.biomaterials.2016.10.006","article-title":"Convective exosome-tracing microfluidics for analysis of cell-non-autonomous neurogenesis","volume":"112","author":"Oh","year":"2017","journal-title":"Biomaterials"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1096\/fj.12-216531","article-title":"Cooperation between \u03b2- and \u03b3-cytoplasmic actions in the mechanical regulation of endothelial microparticle formation","volume":"27","author":"Latham","year":"2013","journal-title":"FASEB J."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"e18138","DOI":"10.1111\/jcmm.18138","article-title":"Exosome inhibition improves response to first-line therapy in small cell lung cancer","volume":"28","author":"Irep","year":"2024","journal-title":"J. Cell Mol. Med."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"110260","DOI":"10.1016\/j.exer.2025.110260","article-title":"Corneal stromal cells from patients with keratoconus exhibit alterations in the ESCRT-dependent machinery responsible for multivesicular body formation","volume":"252","author":"Ye","year":"2025","journal-title":"Exp. Eye Res."},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"1066","DOI":"10.1016\/j.tibtech.2020.05.012","article-title":"Technologies and standardization in research on extracellular vesicles","volume":"38","author":"Gandham","year":"2020","journal-title":"Trends Biotechnol."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"101161","DOI":"10.1016\/j.preteyeres.2022.101161","article-title":"The pre-Descemet\u2019s layer (Dua\u2019s layer, also known as the Dua-Fine layer and the pre-posterior limiting lamina layer): Discovery, characterisation, clinical and surgical applications, and the controversy","volume":"97","author":"Dua","year":"2023","journal-title":"Prog. Retin. Eye Res."},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1038\/nrd3978","article-title":"Extracellular vesicles: Biology and emerging therapeutic opportunities","volume":"12","author":"Breakefield","year":"2013","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_182","doi-asserted-by":"crossref","unstructured":"Robbins, B.T., Montreuil, K.A., Kundu, N., Kumar, P., and Agrahari, V. (2024). Corneal treatment, repair, and regeneration: Exosomes at rescue. Pharmaceutics, 16.","DOI":"10.3390\/pharmaceutics16111424"},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.addr.2018.06.017","article-title":"Strategic design of extracellular vesicle drug delivery systems","volume":"130","author":"Armstrong","year":"2018","journal-title":"Adv. Drug Deliv. Rev."},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1021\/acs.nanolett.8b02689","article-title":"In vitro and in vivo RNA inhibition by CD9-HuR functionalized exosomes encapsulated with miRNA or CRISPR\/dCas9","volume":"19","author":"Li","year":"2019","journal-title":"Nano Lett."},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"3232869","DOI":"10.1155\/2020\/3232869","article-title":"The role of exosomal microRNAs and oxidative stress in neurodegenerative diseases","volume":"2020","author":"Wang","year":"2020","journal-title":"Oxid. Med. Cell Longev."},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"134970","DOI":"10.1016\/j.cej.2022.134970","article-title":"Synthesis, bioactive properties, and biomedical applications of intrinsically therapeutic nanoparticles for disease treatment","volume":"435","author":"Nguyen","year":"2022","journal-title":"Chem. Eng. J."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"3650","DOI":"10.1021\/acs.molpharmaceut.5b00364","article-title":"Exogenous DNA loading into extracellular vesicles via electroporation is size-dependent and enables limited gene delivery","volume":"12","author":"Lamichhane","year":"2015","journal-title":"Mol. Pharm."},{"key":"ref_188","doi-asserted-by":"crossref","unstructured":"Xu, M., Yang, Q., Sun, X., and Wang, Y. (2020). Recent advancements in the loading and modification of therapeutic exosomes. Front. Bioeng. Biotechnol., 8.","DOI":"10.3389\/fbioe.2020.586130"},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"1517","DOI":"10.1016\/j.stemcr.2024.09.008","article-title":"The urgent need for clear and concise regulations on exosome-based interventions","volume":"19","author":"Fujita","year":"2024","journal-title":"Stem Cell Rep."},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"2010","DOI":"10.1016\/j.addr.2005.09.004","article-title":"Topical and systemic drug delivery to the posterior segments","volume":"57","author":"Hughes","year":"2005","journal-title":"Adv. Drug Deliv. Rev."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1038\/nrneph.2017.98","article-title":"Extracellular vesicles in renal disease","volume":"13","author":"Karpman","year":"2017","journal-title":"Nat. Rev. Nephrol."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"348","DOI":"10.1208\/s12248-010-9183-3","article-title":"Ocular drug delivery","volume":"12","author":"Gaudana","year":"2010","journal-title":"AAPS J."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"30087","DOI":"10.3402\/jev.v4.30087","article-title":"Applying extracellular vesicles based therapeutics in clinical trials\u2014An ISEV position paper","volume":"4","author":"Lener","year":"2015","journal-title":"J. Extracell. Vesicles"},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"1535750","DOI":"10.1080\/20013078.2018.1535750","article-title":"Minimal information for studies of extracellular vesicles 2018 (MISEV2018): A position statement of the International Society for Extracellular Vesicles and update of the MISEV2014 guidelines","volume":"7","author":"Witwer","year":"2018","journal-title":"J. Extracell. Vesicles"}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/14\/13\/959\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:57:11Z","timestamp":1760032631000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/14\/13\/959"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,6,23]]},"references-count":194,"journal-issue":{"issue":"13","published-online":{"date-parts":[[2025,7]]}},"alternative-id":["cells14130959"],"URL":"https:\/\/doi.org\/10.3390\/cells14130959","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,6,23]]}}}