{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T00:01:16Z","timestamp":1784851276234,"version":"3.55.0"},"reference-count":145,"publisher":"The Microbiological Society of Korea","issue":"7","license":[{"start":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T00:00:00Z","timestamp":1785456000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100011705","name":"Korea Institute of Marine Science and Technology promotion","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100011705","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003566","name":"Ministry of Oceans and Fisheries","doi-asserted-by":"publisher","award":["RS-2024-00405273"],"award-info":[{"award-number":["RS-2024-00405273"]}],"id":[{"id":"10.13039\/501100003566","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100014188","name":"Ministry of Science and ICT","doi-asserted-by":"crossref","award":["RS-2025-00523171"],"award-info":[{"award-number":["RS-2025-00523171"]}],"id":[{"id":"10.13039\/501100014188","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100002460","name":"Chung-Ang University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100002460","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["jmicrobiol.or.kr"],"crossmark-restriction":true},"short-container-title":["Journal of Microbiology"],"published-print":{"date-parts":[[2026,7,31]]},"abstract":"<jats:p>Fungal extracellular vesicles (EVs) have emerged as critical mediators of fungal physiology, virulence, and host\u2013pathogen interactions. Since their first description in Cryptococcus neoformans, EVs have been identified in several fungal species and shown to transport a broad repertoire of bioactive cargo. Increasing evidence indicates that fungal EVs participate in multiple biological processes, including cell wall remodeling, stress adaptation, biofilm formation, antifungal resistance, and modulation of host immune responses. Recent advances in cryo-electron microscopy, multi-omics approaches, and functional genetics have substantially expanded our understanding of the molecular mechanisms governing EV biogenesis, cargo selection, and extracellular trafficking. These studies have further revealed that EV cargo loading is a highly regulated process linked to intracellular proteostasis, glycosylation, lipid homeostasis, and environmental adaptation. In parallel, the intrinsic immunogenicity and structural stability of fungal EVs have highlighted their translational potential as diagnostic biomarkers, vaccine platforms, therapeutic targets, and nanoscale delivery systems. Given the increasing global burden of invasive fungal infections, this review focuses on EVs derived from clinically relevant human fungal pathogens. We summarize recent advances in EV biogenesis, cargo regulation, their roles in pathogenesis, highlight emerging translational applications, and discuss key unresolved questions and future research directions in the field.<\/jats:p>","DOI":"10.71150\/jm.2606008","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T23:11:33Z","timestamp":1784848293000},"page":"e2606008","update-policy":"https:\/\/doi.org\/10.71150\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Extracellular vesicles in human fungal pathogens: Biogenesis, functions, and translational applications"],"prefix":"10.71150","volume":"64","author":[{"given":"Catia","family":"Mota","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0124-2533","authenticated-orcid":true,"given":"Heeyoun","family":"Hwang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3722-525X","authenticated-orcid":true,"given":"Hyun Ah","family":"Kang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4586","reference":[{"key":"ref1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1462-5822.2008.01160.x"},{"key":"ref2","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2019.00214"},{"key":"ref3","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-017-02126-7"},{"key":"ref4","doi-asserted-by":"publisher","DOI":"10.1038\/nbt.1807"},{"key":"ref5","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00176-19"},{"key":"ref6","doi-asserted-by":"publisher","DOI":"10.3390\/jof8100990"},{"key":"ref7","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms8439"},{"key":"ref8","doi-asserted-by":"publisher","DOI":"10.1038\/nri2868"},{"key":"ref9","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00085-15"},{"key":"ref10","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-25665-5"},{"key":"ref11","doi-asserted-by":"publisher","DOI":"10.1128\/ec.4.11.1902-1912.2005"},{"key":"ref12","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-03991-6"},{"key":"ref13","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.02343"},{"key":"ref14","doi-asserted-by":"publisher","DOI":"10.1002\/pmic.201800232"},{"key":"ref15","doi-asserted-by":"publisher","DOI":"10.1093\/jimmun\/vkaf302"},{"key":"ref16","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.03045-23"},{"key":"ref17","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00190-20"},{"key":"ref18","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.3004404"},{"key":"ref19","doi-asserted-by":"publisher","DOI":"10.1038\/s41564-023-01546-0"},{"key":"ref20","doi-asserted-by":"publisher","DOI":"10.1126\/science.aar4142"},{"key":"ref21","doi-asserted-by":"publisher","DOI":"10.15698\/mic2018.01.607"},{"key":"ref22","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.01397-19"},{"key":"ref23","doi-asserted-by":"publisher","DOI":"10.1128\/iai.00037-24"},{"key":"ref24","doi-asserted-by":"publisher","DOI":"10.1080\/22221751.2022.2098058"},{"key":"ref25","doi-asserted-by":"publisher","DOI":"10.1111\/cmi.13217"},{"key":"ref26","doi-asserted-by":"publisher","DOI":"10.1016\/b978-0-12-800261-2.00001-3"},{"key":"ref27","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00190-20"},{"key":"ref28","doi-asserted-by":"publisher","DOI":"10.1080\/20013078.2020.1750810"},{"key":"ref29","doi-asserted-by":"publisher","DOI":"10.1016\/s1473-3099(23)00692-8"},{"key":"ref30","doi-asserted-by":"publisher","DOI":"10.1016\/j.micpath.2024.106606"},{"key":"ref31","doi-asserted-by":"publisher","DOI":"10.1099\/mic.0.032854-0"},{"key":"ref32","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.00986-15"},{"key":"ref33","doi-asserted-by":"publisher","DOI":"10.1038\/nrmicro.2015.21"},{"key":"ref34","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2958.2008.06193.x"},{"key":"ref35","doi-asserted-by":"publisher","DOI":"10.1038\/s41579-023-00861-x"},{"key":"ref36","doi-asserted-by":"publisher","DOI":"10.1126\/science.aap7999"},{"key":"ref37","doi-asserted-by":"publisher","DOI":"10.1016\/b978-0-12-407677-8.00002-6"},{"key":"ref38","doi-asserted-by":"publisher","DOI":"10.3390\/jof9050541"},{"key":"ref39","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00747-19"},{"key":"ref40","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0021480"},{"key":"ref41","doi-asserted-by":"publisher","DOI":"10.1093\/femsre\/fuy042"},{"key":"ref42","doi-asserted-by":"publisher","DOI":"10.1021\/pr5007944"},{"key":"ref43","doi-asserted-by":"publisher","DOI":"10.1038\/srep06198"},{"key":"ref44","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00209-19"},{"key":"ref45","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2011.05.015"},{"key":"ref46","doi-asserted-by":"publisher","DOI":"10.1016\/j.xphs.2022.08.032"},{"key":"ref47","doi-asserted-by":"publisher","DOI":"10.1128\/aac.00463-08"},{"key":"ref48","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-030-83391-6_10"},{"key":"ref49","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00467-24"},{"key":"ref50","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.00301-22"},{"key":"ref51","doi-asserted-by":"publisher","DOI":"10.1111\/mmi.12985"},{"key":"ref52","doi-asserted-by":"publisher","DOI":"10.1128\/iai.01037-12"},{"key":"ref53","doi-asserted-by":"publisher","DOI":"10.1128\/iai.70.10.5485-5493.2002"},{"key":"ref54","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0048570"},{"key":"ref55","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2018.02286"},{"key":"ref56","doi-asserted-by":"publisher","DOI":"10.1080\/13693780412331271081"},{"key":"ref57","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-27451-9"},{"key":"ref58","doi-asserted-by":"publisher","DOI":"10.3390\/cells9071722"},{"key":"ref59","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2007.06.029"},{"key":"ref60","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2958.2011.07686.x"},{"key":"ref61","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2003700117"},{"key":"ref62","doi-asserted-by":"publisher","DOI":"10.3389\/fphar.2018.01199"},{"key":"ref63","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1012879"},{"key":"ref64","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2022.879018"},{"key":"ref65","doi-asserted-by":"publisher","DOI":"10.3390\/cells9040963"},{"key":"ref66","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.cellbio.20.010403.105307"},{"key":"ref67","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.181331398"},{"key":"ref68","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22137221"},{"key":"ref69","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2020.00256"},{"key":"ref70","doi-asserted-by":"publisher","DOI":"10.1128\/aac.04239-14"},{"key":"ref71","doi-asserted-by":"publisher","DOI":"10.3390\/microorganisms8111815"},{"key":"ref72","doi-asserted-by":"publisher","DOI":"10.1080\/20013078.2017.1359478"},{"key":"ref73","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.00698-22"},{"key":"ref74","doi-asserted-by":"publisher","DOI":"10.1080\/20013078.2017.1286095"},{"key":"ref75","doi-asserted-by":"publisher","DOI":"10.1002\/jex2.82"},{"key":"ref76","doi-asserted-by":"publisher","DOI":"10.1016\/j.resmic.2021.103915"},{"key":"ref77","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1421437112"},{"key":"ref78","doi-asserted-by":"publisher","DOI":"10.1111\/j.1567-1364.2006.00072.x"},{"key":"ref79","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.174.6.3461"},{"key":"ref80","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.103729"},{"key":"ref81","doi-asserted-by":"publisher","DOI":"10.1128\/Spectrum.01538-21"},{"key":"ref82","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2018.10.075"},{"key":"ref83","doi-asserted-by":"publisher","DOI":"10.1128\/mmbr.00081-22"},{"key":"ref84","doi-asserted-by":"publisher","DOI":"10.1016\/j.pharmthera.2018.10.008"},{"key":"ref85","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00044-09"},{"key":"ref86","doi-asserted-by":"publisher","DOI":"10.1038\/ng.605"},{"key":"ref87","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2022.834653"},{"key":"ref88","doi-asserted-by":"publisher","DOI":"10.1016\/j.csbj.2025.04.014"},{"key":"ref89","doi-asserted-by":"publisher","DOI":"10.3390\/jof9050562"},{"key":"ref90","doi-asserted-by":"publisher","DOI":"10.2217\/fmb-2016-0090"},{"key":"ref91","doi-asserted-by":"publisher","DOI":"10.1128\/iai.01171-09"},{"key":"ref92","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0011113"},{"key":"ref93","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2958.2008.06588.x"},{"key":"ref94","doi-asserted-by":"publisher","DOI":"10.1128\/iai.00954-19"},{"key":"ref95","doi-asserted-by":"publisher","DOI":"10.1038\/srep14213"},{"key":"ref96","doi-asserted-by":"publisher","DOI":"10.3390\/cells8070765"},{"key":"ref97","doi-asserted-by":"publisher","DOI":"10.3390\/genes9080400"},{"key":"ref98","doi-asserted-by":"publisher","DOI":"10.1038\/srep07763"},{"key":"ref99","doi-asserted-by":"publisher","DOI":"10.1038\/nrd.2017.46"},{"key":"ref100","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.03469-25"},{"key":"ref101","doi-asserted-by":"publisher","DOI":"10.1016\/s1473-3099(22)00499-6"},{"key":"ref102","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201211138"},{"key":"ref103","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00080-19"},{"key":"ref104","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.00125-21"},{"key":"ref105","doi-asserted-by":"publisher","DOI":"10.1172\/jci27890"},{"key":"ref106","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00476-20"},{"key":"ref107","doi-asserted-by":"publisher","DOI":"10.26226\/morressier.5ac39996d462b8028d89a04d"},{"key":"ref108","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00273-13"},{"key":"ref109","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.00870-23"},{"key":"ref110","doi-asserted-by":"publisher","DOI":"10.1002\/jev2.12129"},{"key":"ref111","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1005240"},{"key":"ref112","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00370-07"},{"key":"ref113","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00318-06"},{"key":"ref114","doi-asserted-by":"publisher","DOI":"10.2217\/fmb.11.112"},{"key":"ref115","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4939-3804-9_12"},{"key":"ref116","doi-asserted-by":"publisher","DOI":"10.1016\/j.tcsw.2018.05.002"},{"key":"ref117","doi-asserted-by":"publisher","DOI":"10.1038\/nm769"},{"key":"ref118","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2019.02008"},{"key":"ref119","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-0854.2009.00920.x"},{"key":"ref120","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2025.146290"},{"key":"ref121","doi-asserted-by":"publisher","DOI":"10.1080\/20013078.2018.1535750"},{"key":"ref122","doi-asserted-by":"publisher","DOI":"10.1080\/21541248.2016.1215283"},{"key":"ref123","doi-asserted-by":"publisher","DOI":"10.1038\/ncb1596"},{"key":"ref124","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00227-10"},{"key":"ref125","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0039463"},{"key":"ref126","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2020.00006"},{"key":"ref127","doi-asserted-by":"publisher","DOI":"10.1038\/nrm.2017.125"},{"key":"ref128","doi-asserted-by":"publisher","DOI":"10.1111\/cmi.13238"},{"key":"ref129","doi-asserted-by":"publisher","DOI":"10.1111\/cmi.12374"},{"key":"ref130","doi-asserted-by":"publisher","DOI":"10.1111\/j.1462-5822.2008.01125.x"},{"key":"ref131","doi-asserted-by":"publisher","DOI":"10.1002\/jev2.12404"},{"key":"ref132","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.aav8521"},{"key":"ref133","doi-asserted-by":"publisher","DOI":"10.1038\/nrm2162"},{"key":"ref134","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00054-15"},{"key":"ref135","doi-asserted-by":"publisher","DOI":"10.1128\/ec.00111-14"},{"key":"ref136","doi-asserted-by":"publisher","DOI":"10.1111\/j.1462-5822.2012.01757.x"},{"key":"ref137","doi-asserted-by":"publisher","DOI":"10.1080\/10717544.2020.1869866"},{"key":"ref138","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2020.603183"},{"key":"ref139","doi-asserted-by":"publisher","DOI":"10.1128\/msystems.00822-21"},{"key":"ref140","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2021.648524"},{"key":"ref141","doi-asserted-by":"publisher","DOI":"10.1016\/S0065-2164(09)01204-0"},{"key":"ref142","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.02988-22"},{"key":"ref143","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-26525-z"},{"key":"ref144","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.2006872"},{"key":"ref145","doi-asserted-by":"publisher","DOI":"10.1186\/s40249-023-01073-4"}],"container-title":["Journal of Microbiology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/jmicrobiol.or.kr\/upload\/pdf\/jm-2606008.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/jmicrobiol.or.kr\/journal\/view.php?doi=10.71150\/jm.2606008","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/jmicrobiol.or.kr\/upload\/pdf\/jm-2606008.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T23:11:42Z","timestamp":1784848302000},"score":1,"resource":{"primary":{"URL":"http:\/\/jmicrobiol.or.kr\/journal\/view.php?doi=10.71150\/jm.2606008"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,31]]},"references-count":145,"journal-issue":{"issue":"7","published-print":{"date-parts":[[2026,7]]}},"URL":"https:\/\/doi.org\/10.71150\/jm.2606008","relation":{},"ISSN":["1225-8873","1976-3794"],"issn-type":[{"value":"1225-8873","type":"print"},{"value":"1976-3794","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,31]]},"assertion":[{"value":"2026-06-09","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-06-24","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-31","order":2,"name":"published","label":"Published online","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-31","order":3,"name":"published","label":"Print publication","group":{"name":"publication_history","label":"Publication History"}}]}}