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However, radiation therapy is a nonspecific method and associated with significant challenges such as radioresistance and non-targeted effects. The radiation-induced non-targeted effects on nonirradiated cells nearby are known as bystander effects, while effects far from the ionising radiation-exposed cells are known as abscopal effects. These effects are presented as a consequence of intercellular communications. Therefore, a better understanding of the involved intercellular signals may bring promising new strategies for radiation risk assessment and potential targets for developing novel radiotherapy strategies. Recent studies indicate that radiation-derived extracellular vesicles, particularly exosomes, play a vital role in intercellular communications and may result in radioresistance and non-targeted effects. This review describes exosome biology, intercellular interactions, and response to different environmental stressors and diseases, and focuses on their role as functional mediators in inducing radiation-induced bystander effect (RIBE).<\/jats:p>","DOI":"10.3390\/bioengineering9060243","type":"journal-article","created":{"date-parts":[[2022,5,31]],"date-time":"2022-05-31T05:25:42Z","timestamp":1653974742000},"page":"243","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":28,"title":["The Footprint of Exosomes in the Radiation-Induced Bystander Effects"],"prefix":"10.3390","volume":"9","author":[{"given":"Safura","family":"Jokar","sequence":"first","affiliation":[{"name":"Department of Nuclear Pharmacy, Faculty of Pharmacy, Tehran University of Medical Sciences, Tehran P94V+927, Iran"},{"name":"Institute of Biophysics, Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Coimbra Institute for Clinical and Biomedical Research (iCBR), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Center for Innovative Biomedicine and Biotechnology (CIBB), University of Coimbra, 3000-548 Coimbra, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5690-3020","authenticated-orcid":false,"given":"In\u00eas A.","family":"Marques","sequence":"additional","affiliation":[{"name":"Institute of Biophysics, Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Coimbra Institute for Clinical and Biomedical Research (iCBR), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Center for Innovative Biomedicine and Biotechnology (CIBB), University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Centre of Investigation in Environment, Genetics and Oncobiology (CIMAGO), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Faculty of Pharmacy, University of Coimbra, 3000-548 Coimbra, Portugal"}]},{"given":"Saeedeh","family":"Khazaei","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical Biomaterials, Faculty of Pharmacy, Tehran University of Medical Sciences, Tehran P94V+927, Iran"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5798-3923","authenticated-orcid":false,"given":"Tania","family":"Martins-Marques","sequence":"additional","affiliation":[{"name":"Coimbra Institute for Clinical and Biomedical Research (iCBR), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Center for Innovative Biomedicine and Biotechnology (CIBB), University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Clinical and Academic Centre of Coimbra (CACC), 3004-561 Coimbra, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5786-8447","authenticated-orcid":false,"given":"Henrique","family":"Girao","sequence":"additional","affiliation":[{"name":"Coimbra Institute for Clinical and Biomedical Research (iCBR), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Center for Innovative Biomedicine and Biotechnology (CIBB), University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Clinical and Academic Centre of Coimbra (CACC), 3004-561 Coimbra, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0689-6007","authenticated-orcid":false,"given":"Mafalda","family":"Laranjo","sequence":"additional","affiliation":[{"name":"Institute of Biophysics, Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Coimbra Institute for Clinical and Biomedical Research (iCBR), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Center for Innovative Biomedicine and Biotechnology (CIBB), University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Centre of Investigation in Environment, Genetics and Oncobiology (CIMAGO), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Clinical and Academic Centre of Coimbra (CACC), 3004-561 Coimbra, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7202-1650","authenticated-orcid":false,"given":"Maria Filomena","family":"Botelho","sequence":"additional","affiliation":[{"name":"Institute of Biophysics, Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Coimbra Institute for Clinical and Biomedical Research (iCBR), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Center for Innovative Biomedicine and Biotechnology (CIBB), University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Centre of Investigation in Environment, Genetics and Oncobiology (CIMAGO), Faculty of Medicine, University of Coimbra, 3000-548 Coimbra, Portugal"},{"name":"Clinical and Academic Centre of Coimbra (CACC), 3004-561 Coimbra, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2022,5,31]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"209","DOI":"10.3322\/caac.21660","article-title":"Global cancer statistics 2020: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries","volume":"71","author":"Sung","year":"2021","journal-title":"CA Cancer J. Clin."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"7","DOI":"10.3322\/caac.21590","article-title":"Cancer statistics, 2020","volume":"70","author":"Siegel","year":"2020","journal-title":"CA Cancer J. Clin."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"193","DOI":"10.7150\/ijms.3635","article-title":"Cancer and radiation therapy: Current advances and future directions","volume":"9","author":"Baskar","year":"2012","journal-title":"Int. J. Med. Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1016\/S1470-2045(15)00222-3","article-title":"Expanding global access to radiotherapy","volume":"16","author":"Atun","year":"2015","journal-title":"Lancet Oncol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1038\/s41572-019-0064-5","article-title":"Radiotherapy toxicity","volume":"5","author":"Niedermann","year":"2019","journal-title":"Nat. Rev. Dis. Prim."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1016\/j.molonc.2012.01.007","article-title":"Individualization of cancer treatment from radiotherapy perspective","volume":"6","author":"Yaromina","year":"2012","journal-title":"Mol. Oncol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.1056\/NEJMra1608986","article-title":"Recent developments in radiotherapy","volume":"377","author":"Citrin","year":"2017","journal-title":"N. Engl. J. Med."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"30165","DOI":"10.1038\/srep30165","article-title":"Bystander autophagy mediated by radiation-induced exosomal miR-7-5p in non-targeted human bronchial epithelial cells","volume":"6","author":"Song","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Mortezaee, K., Najafi, M., Farhood, B., Ahmadi, A., Shabeeb, D., and Eleojo Musa, A. (2019). Genomic instability and carcinogenesis of heavy charged particles radiation: Clinical and environmental implications. Medicina, 55.","DOI":"10.3390\/medicina55090591"},{"key":"ref_10","first-page":"99","article-title":"Changes in gene expression as one of the key mechanisms involved in radiation-induced bystander effect","volume":"9","author":"Sokolov","year":"2018","journal-title":"Biomed. Rep."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1038\/s41416-020-0942-3","article-title":"Radiation-induced bystander and abscopal effects: Important lessons from preclinical models","volume":"123","author":"Daguenet","year":"2020","journal-title":"Br. J. Cancer"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1038\/nrc1277","article-title":"Radiation-induced bystander effects\u2014implications for cancer","volume":"4","author":"Seymour","year":"2004","journal-title":"Nat. Rev. Cancer"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"62","DOI":"10.3109\/09553002.2014.950715","article-title":"Genetic changes in progeny of bystander human fibroblasts after microbeam irradiation with X-rays, protons or carbon ions: The relevance to cancer risk","volume":"91","author":"Autsavapromporn","year":"2015","journal-title":"Int. J. Radiat. Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.canlet.2013.12.017","article-title":"Bystander effects as manifestation of intercellular communication of DNA damage and of the cellular oxidative status","volume":"356","author":"Klammer","year":"2015","journal-title":"Cancer Lett."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.mrfmmm.2013.02.001","article-title":"Apoptosis is signalled early by low doses of ionising radiation in a radiation-induced bystander effect","volume":"741","author":"Furlong","year":"2013","journal-title":"Mutat. Res. Fundam. Mol. Mech. Mutagenesis"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1667\/RR13736.1","article-title":"Irradiated U937 cells trigger inflammatory bystander responses in human umbilical vein endothelial cells through the p38 pathway","volume":"182","author":"Xiao","year":"2014","journal-title":"Radiat. Res."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"406","DOI":"10.1667\/RR3158.2","article-title":"The influence of p53 functions on radiation-induced inflammatory bystander-type signaling in murine bone marrow","volume":"179","author":"Lorimore","year":"2013","journal-title":"Radiat. Res."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"869","DOI":"10.3389\/fonc.2019.00869","article-title":"Exosomes in Cancer Radioresistance","volume":"9","author":"Ni","year":"2019","journal-title":"Front. Oncol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"894","DOI":"10.3389\/fonc.2019.00894","article-title":"Exosomes in Head and Neck Squamous Cell Carcinoma","volume":"9","author":"Xiao","year":"2019","journal-title":"Front. Oncol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1907","DOI":"10.1016\/j.jprot.2010.06.006","article-title":"Exosomes: Extracellular organelles important in intercellular communication","volume":"73","author":"Mathivanan","year":"2010","journal-title":"J. Proteom."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/s12953-019-0154-z","article-title":"Exploring the key communicator role of exosomes in cancer microenvironment through proteomics","volume":"17","author":"Kim","year":"2019","journal-title":"Proteome Sci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1593\/tlo.13640","article-title":"Ionizing radiation and glioblastoma exosomes: Implications in tumor biology and cell migration","volume":"6","author":"Arscott","year":"2013","journal-title":"Transl. Oncol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1016\/j.devcel.2019.04.011","article-title":"Exosome-mediated metastasis: Communication from a distance","volume":"49","author":"Wortzel","year":"2019","journal-title":"Dev. Cell"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Gurunathan, S., Kang, M.-H., Jeyaraj, M., Qasim, M., and Kim, J.-H. (2019). Review of the isolation, characterization, biological function, and multifarious therapeutic approaches of exosomes. Cells, 8.","DOI":"10.3390\/cells8040307"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Doyle, L.M., and Wang, M.Z. (2019). Overview of extracellular vesicles, their origin, composition, purpose, and methods for exosome isolation and analysis. Cells, 8.","DOI":"10.3390\/cells8070727"},{"key":"ref_26","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"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"152","DOI":"10.3390\/genes4020152","article-title":"Signaling pathways in exosomes biogenesis, secretion and fate","volume":"4","author":"Urbanelli","year":"2013","journal-title":"Genes"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1111\/j.1365-2141.1967.tb08741.x","article-title":"The nature and significance of platelet products in human plasma","volume":"13","author":"Wolf","year":"1967","journal-title":"Br. J. Haematol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"656","DOI":"10.2174\/0929866523666160427105138","article-title":"The influence of ionizing radiation on exosome composition, secretion and intercellular communication","volume":"23","author":"Jelonek","year":"2016","journal-title":"Protein Pept. Lett."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"e1413","DOI":"10.1002\/wrna.1413","article-title":"RNA in extracellular vesicles","volume":"8","author":"Kim","year":"2017","journal-title":"Wiley Interdiscip. Rev. RNA"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Fanale, D., Taverna, S., Russo, A., and Bazan, V. (2018). Circular RNA in Exosomes. Circular RNAs, Springer.","DOI":"10.1007\/978-981-13-1426-1_9"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Szatm\u00e1ri, T., Hargitai, R., S\u00e1fr\u00e1ny, G., and Lumniczky, K. (2019). Extracellular vesicles in modifying the effects of ionizing radiation. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20225527"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1038\/nri3622","article-title":"Regulation of immune responses by extracellular vesicles","volume":"14","author":"Robbins","year":"2014","journal-title":"Nat. Rev. Immunol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1016\/j.ceb.2009.03.007","article-title":"Exosomes\u2013vesicular carriers for intercellular communication","volume":"21","author":"Simons","year":"2009","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"2060","DOI":"10.1016\/j.biocel.2012.08.007","article-title":"Exosomes: New players in cell\u2013cell communication","volume":"44","author":"Bang","year":"2012","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1007\/s00109-013-1020-6","article-title":"Exosomes in tumor microenvironment influence cancer progression and metastasis","volume":"91","author":"Kahlert","year":"2013","journal-title":"J. Mol. Med."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1038\/s41392-020-00261-0","article-title":"Exosomes: Key players in cancer and potential therapeutic strategy","volume":"5","author":"Dai","year":"2020","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1038\/nrd3978","article-title":"Extracellular vesicles: Biology and emerging therapeutic opportunities","volume":"12","author":"Andaloussi","year":"2013","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1016\/j.nano.2017.09.011","article-title":"Engineering macrophage-derived exosomes for targeted paclitaxel delivery to pulmonary metastases: In vitro and in vivo evaluations","volume":"14","author":"Kim","year":"2018","journal-title":"Nanomed. Nanotechnol. Biol. Med."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2383","DOI":"10.1016\/j.biomaterials.2013.11.083","article-title":"A doxorubicin delivery platform using engineered natural membrane vesicle exosomes for targeted tumor therapy","volume":"35","author":"Tian","year":"2014","journal-title":"Biomaterials"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1186\/1478-811X-11-88","article-title":"Exosomes are natural carriers of exogenous siRNA to human cells in vitro","volume":"11","author":"Shtam","year":"2013","journal-title":"Cell Commun. Signal."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.apsb.2016.02.001","article-title":"Exosomes as therapeutic drug carriers and delivery vehicles across biological membranes: Current perspectives and future challenges","volume":"6","author":"Ha","year":"2016","journal-title":"Acta Pharm. Sin. B"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"32538","DOI":"10.3402\/jev.v5.32538","article-title":"Presence of Cx43 in extracellular vesicles reduces the cardiotoxicity of the anti-tumour therapeutic approach with doxorubicin","volume":"5","author":"Pinho","year":"2016","journal-title":"J. Extracell. Vesicles"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"387","DOI":"10.2174\/1566524019666190429144310","article-title":"Exosome: An emerging source of biomarkers for human diseases","volume":"19","author":"Xu","year":"2019","journal-title":"Curr. Mol. Med."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"3689","DOI":"10.1096\/fj.201700149","article-title":"Altered cargo proteins of human plasma endothelial cell\u2013derived exosomes in atherosclerotic cerebrovascular disease","volume":"31","author":"Goetzl","year":"2017","journal-title":"FASEB J."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"18396","DOI":"10.3402\/jev.v1i0.18396","article-title":"Cellular stress conditions are reflected in the protein and RNA content of endothelial cell-derived exosomes","volume":"1","author":"Verhaar","year":"2012","journal-title":"J. Extracell. Vesicles"},{"key":"ref_47","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_48","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.mrfmmm.2014.12.007","article-title":"The non-targeted effects of radiation are perpetuated by exosomes","volume":"772","author":"Bright","year":"2015","journal-title":"Mutat. Res. Fundam. Mol. Mech. Mutagenesis"},{"key":"ref_49","first-page":"294","article-title":"Oncogenic extracellular vesicles in brain tumor progression","volume":"3","author":"Garnier","year":"2012","journal-title":"Front. Physiol."},{"key":"ref_50","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_51","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1016\/j.bbrc.2010.07.083","article-title":"Exosome secretion of dendritic cells is regulated by Hrs, an ESCRT-0 protein","volume":"399","author":"Tamai","year":"2010","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/S0092-8674(01)00434-2","article-title":"Ubiquitin-dependent sorting into the multivesicular body pathway requires the function of a conserved endosomal protein sorting complex, ESCRT-I","volume":"106","author":"Katzmann","year":"2001","journal-title":"Cell"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1016\/S0955-0674(03)00080-2","article-title":"Protein sorting into multivesicular endosomes","volume":"15","author":"Raiborg","year":"2003","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"556","DOI":"10.1038\/nrm2937","article-title":"Membrane budding and scission by the ESCRT machinery: It\u2019s all in the neck","volume":"11","author":"Hurley","year":"2010","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"7309","DOI":"10.4049\/jimmunol.166.12.7309","article-title":"Proteomic analysis of dendritic cell-derived exosomes: A secreted subcellular compartment distinct from apoptotic vesicles","volume":"166","author":"Boussac","year":"2001","journal-title":"J. Immunol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"e202000821","DOI":"10.26508\/lsa.202000821","article-title":"Myocardial infarction affects Cx43 content of extracellular vesicles secreted by cardiomyocytes","volume":"3","author":"Zuzarte","year":"2020","journal-title":"Life Sci. Alliance"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1244","DOI":"10.1126\/science.1153124","article-title":"Ceramide triggers budding of exosome vesicles into multivesicular endosomes","volume":"319","author":"Trajkovic","year":"2008","journal-title":"Science"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"11649","DOI":"10.1074\/jbc.M112.445304","article-title":"The intracellular interactome of tetraspanin-enriched microdomains reveals their function as sorting machineries toward exosomes","volume":"288","author":"Jorge","year":"2013","journal-title":"J. Biol. Chem."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.plipres.2014.01.004","article-title":"Ceramide: A simple sphingolipid with unique biophysical properties","volume":"54","author":"Castro","year":"2014","journal-title":"Prog. Lipid Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1528","DOI":"10.1111\/j.1600-0854.2009.00963.x","article-title":"MHC II in dendritic cells is targeted to lysosomes or T cell-induced exosomes via distinct multivesicular body pathways","volume":"10","author":"Buschow","year":"2009","journal-title":"Traffic"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"eabm1140","DOI":"10.1126\/sciadv.abm1140","article-title":"LAMP2A regulates the loading of proteins into exosomes","volume":"8","author":"Ferreira","year":"2021","journal-title":"Sci. Adv."},{"key":"ref_62","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_63","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1586\/epr.09.17","article-title":"Exosomes: Proteomic insights and diagnostic potential","volume":"6","author":"Simpson","year":"2009","journal-title":"Expert Rev. Proteom."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"654","DOI":"10.1038\/ncb1596","article-title":"Exosome-mediated transfer of mRNAs and microRNAs is a novel mechanism of genetic exchange between cells","volume":"9","author":"Valadi","year":"2007","journal-title":"Nat. Cell Biol."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Statello, L., Maugeri, M., Garre, E., Nawaz, M., Wahlgren, J., Papadimitriou, A., Lundqvist, C., Lindfors, L., Collen, A., and Sunnerhagen, P. (2018). Identification of RNA-binding proteins in exosomes capable of interacting with different types of RNA: RBP-facilitated transport of RNAs into exosomes. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0195969"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"478","DOI":"10.4161\/cib.3.5.12693","article-title":"Secretory microRNAs as a versatile communication tool","volume":"3","author":"Iguchi","year":"2010","journal-title":"Commun. Integr. Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"6328","DOI":"10.1073\/pnas.0914843107","article-title":"Functional delivery of viral miRNAs via exosomes","volume":"107","author":"Pegtel","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1038\/ncomms1285","article-title":"Unidirectional transfer of microRNA-loaded exosomes from T cells to antigen-presenting cells","volume":"2","author":"Mittelbrunn","year":"2011","journal-title":"Nat. Commun."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s00702-009-0288-8","article-title":"Astrocytes and Glioblastoma cells release exosomes carrying mtDNA","volume":"117","author":"Guescini","year":"2010","journal-title":"J. Neural Transm."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1977","DOI":"10.1016\/j.yexcr.2010.04.006","article-title":"C2C12 myoblasts release micro-vesicles containing mtDNA and proteins involved in signal transduction","volume":"316","author":"Guescini","year":"2010","journal-title":"Exp. Cell Res."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Hofmann, L., Ludwig, S., Vahl, J.M., Brunner, C., Hoffmann, T.K., and Theodoraki, M.-N. (2020). The Emerging Role of Exosomes in Diagnosis, Prognosis, and Therapy in Head and Neck Cancer. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21114072"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.7150\/ijbs.18890","article-title":"Exosomal miR-7 mediates bystander autophagy in lung after focal brain irradiation in mice","volume":"13","author":"Cai","year":"2017","journal-title":"Int. J. Biol. Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"492","DOI":"10.1038\/s41416-018-0192-9","article-title":"Exosome-packaged miR-1246 contributes to bystander DNA damage by targeting LIG4","volume":"119","author":"Mo","year":"2018","journal-title":"Br. J. Cancer"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/j.biochi.2006.10.014","article-title":"Exosome lipidomics unravels lipid sorting at the level of multivesicular bodies","volume":"89","author":"Subra","year":"2007","journal-title":"Biochimie"},{"key":"ref_75","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_76","doi-asserted-by":"crossref","first-page":"2105","DOI":"10.1194\/jlr.M003657","article-title":"Exosomes account for vesicle-mediated transcellular transport of activatable phospholipases and prostaglandins","volume":"51","author":"Subra","year":"2010","journal-title":"J. Lipid Res."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1977","DOI":"10.1182\/blood-2008-08-174094","article-title":"Activated T cells recruit exosomes secreted by dendritic cells via LFA-1","volume":"113","author":"Buschow","year":"2009","journal-title":"Blood"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1111\/j.1600-0854.2010.01041.x","article-title":"Cellular internalization of exosomes occurs through phagocytosis","volume":"11","author":"Feng","year":"2010","journal-title":"Traffic"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1297","DOI":"10.1073\/pnas.0307981100","article-title":"Release of full-length 55-kDa TNF receptor 1 in exosome-like vesicles: A mechanism for generation of soluble cytokine receptors","volume":"101","author":"Hawari","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"2620","DOI":"10.1002\/eji.200424969","article-title":"Complement inhibitor membrane cofactor protein (MCP; CD46) is constitutively shed from cancer cell membranes in vesicles and converted by a metalloproteinase to a functionally active soluble form","volume":"34","author":"Hakulinen","year":"2004","journal-title":"Eur. J. Immunol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1042\/BJ20051013","article-title":"A role for exosomes in the constitutive and stimulus-induced ectodomain cleavage of L1 and CD44","volume":"393","author":"Stoeck","year":"2006","journal-title":"Biochem. J."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"13243","DOI":"10.1038\/srep13243","article-title":"Gap junctional protein Cx43 is involved in the communication between extracellular vesicles and mammalian cells","volume":"5","author":"Soares","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"24641","DOI":"10.3402\/jev.v3.24641","article-title":"Routes and mechanisms of extracellular vesicle uptake","volume":"3","author":"Mulcahy","year":"2014","journal-title":"J. Extracell. Vesicles"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"768","DOI":"10.1016\/j.bbrc.2014.12.015","article-title":"Effective internalization of U251-MG-secreted exosomes into cancer cells and characterization of their lipid components","volume":"456","author":"Toda","year":"2015","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/j.canlet.2015.04.026","article-title":"Cell-specific uptake of mantle cell lymphoma-derived exosomes by malignant and non-malignant B-lymphocytes","volume":"364","author":"Rosenblum","year":"2015","journal-title":"Cancer Lett."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.semcdb.2017.01.002","article-title":"Extracellular vesicle docking at the cellular port: Extracellular vesicle binding and uptake","volume":"67","author":"French","year":"2017","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1083\/jcb.146.2.477","article-title":"Characterization of integrin\u2013tetraspanin adhesion complexes: Role of tetraspanins in integrin signaling","volume":"146","author":"Berditchevski","year":"1999","journal-title":"J. Cell Biol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1574","DOI":"10.1016\/j.biocel.2012.06.018","article-title":"Toward tailored exosomes: The exosomal tetraspanin web contributes to target cell selection","volume":"44","author":"Rana","year":"2012","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_89","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_90","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1007\/s40572-019-00233-9","article-title":"Exosomes as Mediators of Chemical-Induced Toxicity","volume":"6","author":"Rokad","year":"2019","journal-title":"Curr. Env. Health Rep."},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"Gai, C., Camussi, F., Broccoletti, R., Gambino, A., Cabras, M., Molinaro, L., Carossa, S., Camussi, G., and Arduino, P.G. (2018). Salivary extracellular vesicle-associated miRNAs as potential biomarkers in oral squamous cell carcinoma. BMC Cancer, 18.","DOI":"10.1186\/s12885-018-4364-z"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"25","DOI":"10.2147\/CMAR.S272140","article-title":"Identification of Urinary Exosomal miRNAs for the Non-Invasive Diagnosis of Prostate Cancer","volume":"13","author":"Li","year":"2021","journal-title":"Cancer Manag. Res."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"20170065","DOI":"10.1098\/rstb.2017.0065","article-title":"Cisplatin induces the release of extracellular vesicles from ovarian cancer cells that can induce invasiveness and drug resistance in bystander cells","volume":"372","author":"Samuel","year":"2018","journal-title":"Philos. Trans. R. Soc. B"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1157","DOI":"10.18632\/oncotarget.27531","article-title":"Exosome mediated miR-155 delivery confers cisplatin chemoresistance in oral cancer cells via epithelial-mesenchymal transition","volume":"11","author":"Kirave","year":"2020","journal-title":"Oncotarget"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"8382","DOI":"10.1038\/s41419-020-02962-4","article-title":"Bone marrow mesenchymal stem cells-derived exosomal microRNA-193a reduces cisplatin resistance of non-small cell lung cancer cells via targeting LRRC1","volume":"11","author":"Wu","year":"2020","journal-title":"Cell Death Dis."},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Kabakov, A.E., and Yakimova, A.O. (2021). Hypoxia-induced cancer cell responses driving radioresistance of hypoxic tumors: Approaches to targeting and radiosensitizing. Cancers, 13.","DOI":"10.3390\/cancers13051102"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.canlet.2020.03.017","article-title":"Hypoxia in tumor microenvironment regulates exosome biogenesis: Molecular mechanisms and translational opportunities","volume":"479","author":"Kumar","year":"2020","journal-title":"Cancer Lett."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1667\/RR15555.1","article-title":"Effects of Hypoxia and Radiation-Induced Exosomes on Migration of Lung Cancer Cells and Angiogenesis of Umbilical Vein Endothelial Cells","volume":"194","author":"Mo","year":"2020","journal-title":"Radiat. Res."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"7312","DOI":"10.1073\/pnas.1220998110","article-title":"Exosomes reflect the hypoxic status of glioma cells and mediate hypoxia-dependent activation of vascular cells during tumor development","volume":"110","author":"Kucharzewska","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Ouellette, M.M., Zhou, S., and Yan, Y. (2022). Cell Signaling Pathways That Promote Radioresistance of Cancer Cells. Diagnostics, 12.","DOI":"10.3390\/diagnostics12030656"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1007\/s12079-019-00515-9","article-title":"Bystander effectors of chondrosarcoma cells irradiated at different LET impair proliferation of chondrocytes","volume":"13","author":"Lepleux","year":"2019","journal-title":"J. Cell Commun. Signal."},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Elbakrawy, E.M., Mayah, A., Hill, M.A., and Kadhim, M. (2020). Induction of genomic instability in a primary human fibroblast cell line following low-dose alpha-particle exposure and the potential role of exosomes. Biology, 10.","DOI":"10.3390\/biology10010011"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1667\/RR13337.1","article-title":"Exosomes are involved in mediating radiation induced bystander signaling in human keratinocyte cells","volume":"181","author":"flJella","year":"2014","journal-title":"Radiat. Res."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"265","DOI":"10.18388\/abp.2015_970","article-title":"Ionizing radiation affects protein composition of exosomes secreted in vitro from head and neck squamous cell carcinoma","volume":"62","author":"Jelonek","year":"2015","journal-title":"Acta Biochim. Pol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"34","DOI":"10.2174\/1874471011666171229123130","article-title":"Mechanisms of Radiation Bystander and Non-Targeted Effects: Implications to Radiation Carcinogenesis and Radiotherapy","volume":"11","author":"Yahyapour","year":"2018","journal-title":"Curr. Radiopharm."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"7864","DOI":"10.1158\/0008-5472.CAN-07-6538","article-title":"Senescence-Associated Exosome Release from Human Prostate Cancer Cells","volume":"68","author":"Lehmann","year":"2008","journal-title":"Cancer Res."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"12423","DOI":"10.1038\/s41598-017-12403-6","article-title":"Radiation alters the cargo of exosomes released from squamous head and neck cancer cells to promote migration of recipient cells","volume":"7","author":"Mutschelknaus","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/s13046-016-0285-3","article-title":"Radiation-induced miR-208a increases the proliferation and radioresistance by targeting p21 in human lung cancer cells","volume":"35","author":"Tang","year":"2016","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s10495-010-0534-4","article-title":"Survivin is released from cancer cells via exosomes","volume":"16","author":"Khan","year":"2011","journal-title":"Apoptosis"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"4795","DOI":"10.1158\/0008-5472.CAN-05-4579","article-title":"The Regulation of Exosome Secretion: A Novel Function of the p53 Protein","volume":"66","author":"Yu","year":"2006","journal-title":"Cancer Res."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1186\/s13014-016-0636-4","article-title":"Circulating miR-29a and miR-150 correlate with delivered dose during thoracic radiation therapy for non-small cell lung cancer","volume":"11","author":"Dinh","year":"2016","journal-title":"Radiat. Oncol."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1016\/j.radonc.2010.03.024","article-title":"Radiation therapy induces circulating serum Hsp72 in patients with prostate cancer","volume":"95","author":"Hurwitz","year":"2010","journal-title":"Radiother. Oncol. J. Eur. Soc. Ther. Radiol. Oncol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1016\/j.bbrc.2014.03.067","article-title":"Radiation increases the cellular uptake of exosomes through CD29\/CD81 complex formation","volume":"446","author":"Hazawa","year":"2014","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1186\/s12943-018-0867-0","article-title":"Exosomes derived from mesenchymal stem cells enhance radiotherapy-induced cell death in tumor and metastatic tumor foci","volume":"17","author":"Anderson","year":"2018","journal-title":"Mol. Cancer"},{"key":"ref_115","first-page":"229","article-title":"Autocrine secretions enhance radioresistance in an exosome-independent manner in NSCLC cells","volume":"54","author":"Wang","year":"2019","journal-title":"Int. J. Oncol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/s13246-010-0030-8","article-title":"Non-targeted effects of ionising radiation and radiotherapy","volume":"33","author":"Sjostedt","year":"2010","journal-title":"Australas. Phys. Eng. Sci. Med."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.canlet.2013.09.009","article-title":"Non-targeted effects induced by ionizing radiation: Mechanisms and potential impact on radiation induced health effects","volume":"356","author":"Morgan","year":"2015","journal-title":"Cancer Lett."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1089\/ars.2013.5489","article-title":"Effects of ionizing radiation on biological molecules\u2014mechanisms of damage and emerging methods of detection","volume":"21","author":"Reisz","year":"2014","journal-title":"Antioxid. Redox Signal."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"586","DOI":"10.1016\/j.clon.2013.06.005","article-title":"Bystander signalling: Exploring clinical relevance through new approaches and new models","volume":"25","author":"Butterworth","year":"2013","journal-title":"Clin. Oncol."},{"key":"ref_120","first-page":"6394","article-title":"Induction of sister chromatid exchanges by extremely low doses of \u03b1-particles","volume":"52","author":"Nagasawa","year":"1992","journal-title":"Cancer Res."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"12445","DOI":"10.1073\/pnas.0804186105","article-title":"Oncogenic bystander radiation effects in Patched heterozygous mouse cerebellum","volume":"105","author":"Mancuso","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1615\/CritRevOncog.v18.i1-2.40","article-title":"On the origin of cancer metastasis","volume":"18","author":"Seyfried","year":"2013","journal-title":"Crit. Rev. Oncog."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1016\/j.ejphar.2009.07.028","article-title":"Ionizing radiation-induced bystander effects, potential targets for modulation of radiotherapy","volume":"625","author":"Przybyszewski","year":"2009","journal-title":"Eur. J. Pharmacol."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"1","DOI":"10.4236\/ijmpcero.2016.51001","article-title":"Radiation induced bystander effect: From in vitro studies to clinical application","volume":"5","author":"Widel","year":"2016","journal-title":"Int. J. Med. Phys. Clin. Eng. Radiat. Oncol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1093\/jrr\/rrz001","article-title":"Ionizing radiation affects the composition of the proteome of extracellular vesicles released by head-and-neck cancer cells in vitro","volume":"60","author":"Abramowicz","year":"2019","journal-title":"J. Radiat. Res."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1080\/09553002.2021.1980630","article-title":"Non-targeted effects of radiation: A personal perspective on the role of exosomes in an evolving paradigm","volume":"98","author":"Kadhim","year":"2022","journal-title":"Int. J. Radiat. Biol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.lssr.2020.11.001","article-title":"Characterization of exosome release and extracellular vesicle-associated miRNAs for human bronchial epithelial cells irradiated with high charge and energy ions","volume":"28","author":"Li","year":"2021","journal-title":"Life Sci. Space Res."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"9103","DOI":"10.1038\/s41598-019-45669-z","article-title":"Radiation-induced bystander effect is mediated by mitochondrial DNA in exosome-like vesicles","volume":"9","author":"Ariyoshi","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_129","doi-asserted-by":"crossref","unstructured":"Tortolici, F., Vumbaca, S., Incocciati, B., Dayal, R., Aquilano, K., Giovanetti, A., and Rufini, S. (2021). Ionizing Radiation-Induced Extracellular Vesicle Release Promotes AKT-Associated Survival Response in SH-SY5Y Neuroblastoma Cells. Cells, 10.","DOI":"10.3390\/cells10010107"},{"key":"ref_130","first-page":"347","article-title":"Extracellular vesicles mediate radiation-induced systemic bystander signals in the bone marrow and spleen","volume":"8","author":"Kis","year":"2017","journal-title":"Front. Immunol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1080\/09553002.2018.1450533","article-title":"Extracellular vesicles mediate low dose ionizing radiation-induced immune and inflammatory responses in the blood","volume":"95","author":"Persa","year":"2019","journal-title":"Int. J. Radiat. Biol."},{"key":"ref_132","doi-asserted-by":"crossref","unstructured":"Hargitai, R., Kis, D., Persa, E., Szatm\u00e1ri, T., S\u00e1fr\u00e1ny, G., and Lumniczky, K. (2021). Oxidative Stress and Gene Expression Modifications Mediated by Extracellular Vesicles: An In Vivo Study of the Radiation-Induced Bystander Effect. Antioxidants, 10.","DOI":"10.3390\/antiox10020156"},{"key":"ref_133","doi-asserted-by":"crossref","unstructured":"Kis, D., Csord\u00e1s, I.B., Persa, E., Jezs\u00f3, B., Hargitai, R., Szatm\u00e1ri, T., S\u00e1ndor, N., Kis, E., Bal\u00e1zs, K., and S\u00e1fr\u00e1ny, G. (2022). Extracellular Vesicles Derived from Bone Marrow in an Early Stage of Ionizing Radiation Damage Are Able to Induce Bystander Responses in the Bone Marrow. Cells, 11.","DOI":"10.3390\/cells11010155"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"2228","DOI":"10.1091\/mbc.E18-02-0130","article-title":"Extracellular vesicles transfer nuclear Abl-dependent and radiation-induced miR-34c into unirradiated cells to cause bystander effects","volume":"29","author":"Rastogi","year":"2018","journal-title":"Mol. Biol. Cell"},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"36083","DOI":"10.18632\/oncotarget.26300","article-title":"Exosomes impact survival to radiation exposure in cell line models of nervous system cancer","volume":"9","author":"Mrowczynski","year":"2018","journal-title":"Oncotarget"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1093\/jrr\/rraa106","article-title":"Extracellular vesicles released from irradiated neonatal mouse cheek tissue increased cell survival after radiation","volume":"62","author":"Ariyoshi","year":"2021","journal-title":"J. Radiat. Res."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"2359","DOI":"10.1002\/lary.22180","article-title":"Disruption of the AKT pathway inhibits metastasis in an orthotopic model of head and neck squamous cell carcinoma","volume":"121","author":"Knowles","year":"2011","journal-title":"Laryngoscope"},{"key":"ref_138","doi-asserted-by":"crossref","unstructured":"Mutschelknaus, L., Peters, C., Winkler, K., Yentrapalli, R., Heider, T., Atkinson, M.J., and Moertl, S. (2016). Exosomes derived from squamous head and neck cancer promote cell survival after ionizing radiation. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0152213"},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1667\/RR2868.1","article-title":"Possible role of exosomes containing RNA in mediating nontargeted effect of ionizing radiation","volume":"177","author":"Irons","year":"2012","journal-title":"Radiat. Res."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1002\/path.4299","article-title":"Responses to ionizing radiation mediated by inflammatory mechanisms","volume":"232","author":"Mukherjee","year":"2014","journal-title":"J. Pathol."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1007\/s00411-011-0386-5","article-title":"The role of miRNA in the direct and indirect effects of ionizing radiation","volume":"50","author":"Dickey","year":"2011","journal-title":"Radiat. Environ. Biophys."},{"key":"ref_142","first-page":"634","article-title":"Differential regulation of microRNA expression in irradiated and bystander cells","volume":"46","author":"Chaudhry","year":"2012","journal-title":"Mol Biol Mosk."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"1355","DOI":"10.1080\/15476286.2015.1100795","article-title":"Exosome-mediated microRNA transfer plays a role in radiation-induced bystander effect","volume":"12","author":"Xu","year":"2015","journal-title":"RNA Biol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"1882","DOI":"10.1093\/carcin\/bgq119","article-title":"microRNAome changes in bystander three-dimensional human tissue models suggest priming of apoptotic pathways","volume":"31","author":"Kovalchuk","year":"2010","journal-title":"Carcinogenesis"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/j.mrfmmm.2015.08.003","article-title":"The key role of miR-21-regulated SOD2 in the medium-mediated bystander responses in human fibroblasts induced by \u03b1-irradiated keratinocytes","volume":"780","author":"Tian","year":"2015","journal-title":"Mutat. Res. Fundam. Mol. Mech. Mutagenesis"},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"11373","DOI":"10.1038\/srep11373","article-title":"Radiation quality-dependence of bystander effect in unirradiated fibroblasts is associated with TGF-\u03b21-Smad2 pathway and miR-21 in irradiated keratinocytes","volume":"5","author":"Yin","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"1161","DOI":"10.4161\/rna.34380","article-title":"MiR-21 is involved in radiation-induced bystander effects","volume":"11","author":"Xu","year":"2014","journal-title":"RNA Biol."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1080\/09553002.2017.1294772","article-title":"Quantitative changes in the protein and miRNA cargo of plasma exosome-like vesicles after exposure to ionizing radiation","volume":"93","author":"Yentrapalli","year":"2017","journal-title":"Int. J. Radiat. Biol."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1186\/s13045-020-00987-y","article-title":"Isolation and characterization of exosomes for cancer research","volume":"13","author":"Zhu","year":"2020","journal-title":"J. Hematol. Oncol."},{"key":"ref_150","doi-asserted-by":"crossref","unstructured":"Logozzi, M., Mizzoni, D., Angelini, D.F., Di Raimo, R., Falchi, M., Battistini, L., and Fais, S. (2018). Microenvironmental pH and Exosome Levels Interplay in Human Cancer Cell Lines of Different Histotypes. 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