{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T06:00:35Z","timestamp":1785477635372,"version":"3.56.0"},"reference-count":61,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T00:00:00Z","timestamp":1785456000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:p>\n                    Invasive candidiasis remains a life-threatening infection with mortality rates reaching 40-55%, yet current gold-standard diagnostics such as blood culture are often slow and insensitive. This review evaluates fungal extracellular vesicles (EVs) as both mechanistic drivers of pathogenesis and clinical utility as biomarkers. We first outline EV biogenesis and translocation across the fungal cell wall, emphasizing the roles of turgor pressure and enzymatic remodeling. We then discuss how EVs act as protected carriers of virulence factors, such as candidalysin and Als3, which modulate tissue invasion and host responses. While profiling these circulating fungal EVs provides a promising and sensitive window into systemic invasive candidiasis, this approach faces unique hurdles in cases of central nervous system (CNS) involvement. In the CNS, fungal signatures are sequestered behind the blood-brain barrier (BBB) and heavily diluted by the vast pool of peripheral EVs, rendering total plasma analysis inadequate. To overcome these barriers, we propose a specialized diagnostic framework that combines dual-marker immunocapture of neuron-derived EVs (NDEVs), using neural cell adhesion molecule (NCAM) and intercellular adhesion molecule 5 (ICAM-5), with the detection of internal\n                    <jats:italic>Candida<\/jats:italic>\n                    -associated cargo. We emphasize that while this host-vesicle enrichment strategy is mechanistically grounded, it requires standardized isolation workflows and prospective clinical validation before deployment as a non-invasive platform for early CNS diagnosis.\n                  <\/jats:p>","DOI":"10.3389\/fcell.2026.1854037","type":"journal-article","created":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T05:37:27Z","timestamp":1785476247000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Fungal extracellular vesicles as biomarkers in candidiasis: biogenesis, virulence, and CNS diagnostic applications"],"prefix":"10.3389","volume":"14","author":[{"given":"Jeongin","family":"Choi","sequence":"first","affiliation":[{"name":"Department of Life Sciences, School of Life Sciences and Biotechnology, Korea University","place":["Seoul, Republic of Korea"]},{"name":"National Research Laboratory for Convergence Degradation Biology, Korea University","place":["Seoul, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuejin","family":"Wang","sequence":"additional","affiliation":[{"name":"Department of Life Sciences, School of Life Sciences and Biotechnology, Korea University","place":["Seoul, Republic of Korea"]},{"name":"National Research Laboratory for Convergence Degradation Biology, Korea University","place":["Seoul, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jihyun","family":"Moon","sequence":"additional","affiliation":[{"name":"Department of Medical and Biological Science, The Catholic University of Korea","place":["Bucheon, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dae-Kyum","family":"Kim","sequence":"additional","affiliation":[{"name":"Cancer Research Program, The Research Institute of the McGill University Health Centre","place":["Montreal, QC, Canada"]},{"name":"Department of Surgery, Surgical and Interventional Sciences, McGill University","place":["Montreal, QC, Canada"]},{"name":"Department of Surgery, Division of Thoracic and Upper Gastrointestinal Surgery, Faculty of Medicine and Health Sciences, McGill University","place":["Montreal, QC, Canada"]},{"name":"Montreal General Hospital Foundation, McGill University Health Centre","place":["Montreal, QC, Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hunsang","family":"Lee","sequence":"additional","affiliation":[{"name":"Department of Life Sciences, School of Life Sciences and Biotechnology, Korea University","place":["Seoul, Republic of Korea"]},{"name":"National Research Laboratory for Convergence Degradation Biology, Korea University","place":["Seoul, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ji-Eun","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Medical and Biological Science, The Catholic University of Korea","place":["Bucheon, Republic of Korea"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,7,31]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"844194","DOI":"10.3389\/fnmol.2022.844194","article-title":"Role of extracellular vesicles in glia-neuron intercellular communication","volume":"15","author":"Ahmad","year":"2022","journal-title":"Front. Mol. Neurosci."},{"key":"B2","doi-asserted-by":"publisher","first-page":"127","DOI":"10.3389\/fnins.2016.00127","article-title":"Extracellular vesicles and a novel form of communication in the brain","volume":"10","author":"Basso","year":"2016","journal-title":"Front. Neurosci."},{"key":"B3","doi-asserted-by":"publisher","first-page":"620","DOI":"10.1038\/nrmicro3480","article-title":"Through the wall: extracellular vesicles in Gram-positive bacteria, mycobacteria and fungi","volume":"13","author":"Brown","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"266","DOI":"10.1093\/mmy\/myaa051","article-title":"Central nervous system candidiasis beyond neonates: lessons from a nationwide study","volume":"59","author":"Chaussade","year":"2021","journal-title":"Med. Mycol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1186\/s44330-025-00034-7","article-title":"Examination of the enrichment of neuronal extracellular vesicles from cell conditioned media and human plasma using an anti-NCAM immunocapture bead approach","volume":"2","author":"Collier","year":"2025","journal-title":"BMC Methods"},{"key":"B6","doi-asserted-by":"publisher","first-page":"578","DOI":"10.1126\/science.285.5427.578","article-title":"An adhesin of the yeast pathogen Candida glabrata mediating adherence to human epithelial cells","volume":"285","author":"Cormack","year":"1999","journal-title":"Science"},{"key":"B7","doi-asserted-by":"publisher","first-page":"1750810","DOI":"10.1080\/20013078.2020.1750810","article-title":"Protein markers for Candida albicans EVs include claudin-like Sur7 family proteins","volume":"9","author":"Dawson","year":"2020","journal-title":"J. Extracell. Vesicles"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1101\/2025.11.07.686794","article-title":"ICAM-5: a novel marker for neuronally derived extracellular vesicles","author":"Deleon","year":"2025","journal-title":"bioRxiv"},{"key":"B8","doi-asserted-by":"publisher","first-page":"e70065","DOI":"10.1002\/jex2.70065","article-title":"Extracellular vesicles from fungal infection in humans: a key player in immunological responses","volume":"4","author":"de Rezende","year":"2025","journal-title":"J. Extracell. Biol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"e428","DOI":"10.1016\/S1473-3099(23)00692-8","article-title":"Global incidence and mortality of severe fungal disease","volume":"24","author":"Denning","year":"2024","journal-title":"Lancet Infect. Dis."},{"key":"B11","doi-asserted-by":"publisher","first-page":"817844","DOI":"10.3389\/fmicb.2022.817844","article-title":"Biogenesis and biological functions of extracellular vesicles in cellular and organismal communication with microbes","volume":"13","author":"Fang","year":"2022","journal-title":"Front. Microbiol."},{"key":"B12","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1186\/s12929-023-00926-2","article-title":"Diagnosis of invasive fungal infections: challenges and recent developments","volume":"30","author":"Fang","year":"2023","journal-title":"J. Biomed. Sci."},{"key":"B13","doi-asserted-by":"publisher","first-page":"3689","DOI":"10.1128\/IAI.70.7.3689-3700.2002","article-title":"Candida albicans hyphal formation and the expression of the Efg1-regulated proteinases Sap4 to Sap6 are required for the invasion of parenchymal organs","volume":"70","author":"Felk","year":"2002","journal-title":"Infect. Immun."},{"key":"B14","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.fbr.2016.03.002","article-title":"The importance of subclasses of chitin synthase enzymes with myosin-like domains for the fitness of fungi","volume":"30","author":"Fernandes","year":"2016","journal-title":"Fungal Biol. Rev."},{"key":"B15","doi-asserted-by":"publisher","first-page":"600","DOI":"10.1016\/j.jalz.2014.06.008","article-title":"Identification of preclinical alzheimer's disease by a profile of pathogenic proteins in neurally derived blood exosomes: a case-control study","volume":"11","author":"Fiandaca","year":"2015","journal-title":"Alzheimers Dement."},{"key":"B16","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1111\/myc.13451","article-title":"Serologic biomarkers in candida and aspergillus infections of the central nervous system: a comparison of galactomannan, mannan and \u03b2-1,3-D-glucan testing from serum and cerebrospinal fluid","volume":"65","author":"Forster","year":"2022","journal-title":"Mycoses"},{"key":"B17","doi-asserted-by":"publisher","first-page":"e1001604","DOI":"10.1371\/journal.pbio.1001604","article-title":"Neurotransmitter-triggered transfer of exosomes mediates oligodendrocyte-neuron communication","volume":"11","author":"Fr\u00fchbeis","year":"2013","journal-title":"PLoS Biol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1021\/pr5007944","article-title":"Proteomics unravels extracellular vesicles as carriers of classical cytoplasmic proteins in Candida albicans","volume":"14","author":"Gil-Bona","year":"2015","journal-title":"J. Proteome Res."},{"key":"B19","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1093\/femsre\/fuy042","article-title":"Extracellular membrane vesicles in the three domains of life and beyond","volume":"43","author":"Gill","year":"2019","journal-title":"FEMS Microbiol. Rev."},{"key":"B20","doi-asserted-by":"publisher","first-page":"3190","DOI":"10.1002\/pmic.200900840","article-title":"Proteomic analysis identifies highly antigenic proteins in exosomes from M. Tuberculosis-infected and culture filtrate protein-treated macrophages","volume":"10","author":"Giri","year":"2010","journal-title":"Proteomics"},{"key":"B21","doi-asserted-by":"publisher","first-page":"1781365","DOI":"10.3389\/fimmu.2026.1781365","article-title":"The role and prospects of extracellular vesicles in advanced drug and vaccine delivery","volume":"17","author":"Huang","year":"2026","journal-title":"Front. Immunol."},{"key":"B22","doi-asserted-by":"publisher","first-page":"e00404","DOI":"10.1128\/iai.00404-24","article-title":"Candida albicans biofilm extracellular vesicles deliver candidalysin to epithelial cell membranes and induce host cell responses","volume":"93","author":"Lee","year":"2025","journal-title":"Infect. Immun."},{"key":"B23","doi-asserted-by":"publisher","first-page":"7221","DOI":"10.3390\/ijms22137221","article-title":"Extracellular vesicles in the fungi kingdom","volume":"22","author":"Liebana-Jordan","year":"2021","journal-title":"Int. J. Mol. Sci."},{"key":"B24","doi-asserted-by":"publisher","first-page":"1127","DOI":"10.1093\/mmy\/myab053","article-title":"Opportunistic yeast pathogen candida spp.: secreted and membrane-bound virulence factors","volume":"59","author":"Lim","year":"2021","journal-title":"Med. Mycol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"1205477","DOI":"10.3389\/fmicb.2023.1205477","article-title":"Fungal extracellular vesicle-mediated regulation: from virulence factor to clinical application","volume":"14","author":"Liu","year":"2023","journal-title":"Front. Microbiol."},{"key":"B26","doi-asserted-by":"publisher","first-page":"e1002305","DOI":"10.1371\/journal.ppat.1002305","article-title":"Mechanisms of Candida albicans trafficking to the brain","volume":"7","author":"Liu","year":"2011","journal-title":"PLoS Pathog."},{"key":"B27","doi-asserted-by":"publisher","first-page":"e0069822","DOI":"10.1128\/spectrum.00698-22","article-title":"Candida albicans hyphal extracellular vesicles are different from yeast ones, carrying an active proteasome complex and showing a different role in host immune response","volume":"10","author":"Mart\u00ednez-L\u00f3pez","year":"2022","journal-title":"Microbiol. Spectr."},{"key":"B28","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1038\/nature17625","article-title":"Candidalysin is a fungal peptide toxin critical for mucosal infection","volume":"532","author":"Moyes","year":"2016","journal-title":"Nature"},{"key":"B29","doi-asserted-by":"publisher","first-page":"e01538\u201321","DOI":"10.1128\/Spectrum.01538-21","article-title":"Cellular and extracellular vesicle RNA analysis in the global threat fungus Candida auris","volume":"9","author":"Munhoz da Rocha","year":"2021","journal-title":"Microbiol. Spectr."},{"key":"B30","doi-asserted-by":"publisher","first-page":"278","DOI":"10.3389\/fnins.2017.00278","article-title":"Plasma extracellular vesicles enriched for neuronal origin: a potential window into brain pathologic processes","volume":"11","author":"Mustapic","year":"2017","journal-title":"Front. Neurosci."},{"key":"B31","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1038\/s41592-021-01174-8","article-title":"L1CAM is not associated with extracellular vesicles in human cerebrospinal fluid or plasma","volume":"18","author":"Norman","year":"2021","journal-title":"Nat. Methods"},{"key":"B32","doi-asserted-by":"publisher","first-page":"e11113","DOI":"10.1371\/journal.pone.0011113","article-title":"Characterization of yeast extracellular vesicles: evidence for the participation of different pathways of cellular traffic in vesicle biogenesis","volume":"5","author":"Oliveira","year":"2010","journal-title":"PLoS One"},{"key":"B33","doi-asserted-by":"publisher","first-page":"1887","DOI":"10.1016\/j.csbj.2025.04.014","article-title":"Deciphering cargo contents in extracellular vesicles of Candida haemulonii Var. vulnera","volume":"27","author":"Oliveira","year":"2025","journal-title":"Comput. Struct. Biotechnol. J."},{"key":"B34","doi-asserted-by":"publisher","first-page":"1165","DOI":"10.1111\/j.1365-2958.2008.06588.x","article-title":"Sec6-dependent sorting of fungal extracellular exosomes and laccase of Cryptococcus neoformans","volume":"71","author":"Panepinto","year":"2009","journal-title":"Mol. Microbiol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"2286","DOI":"10.1016\/j.csbj.2021.04.031","article-title":"ExVe: the knowledge base of orthologous proteins identified in fungal extracellular vesicles","volume":"19","author":"Parreira","year":"2021","journal-title":"Comput. Struct. Biotechnol. J."},{"key":"B36","doi-asserted-by":"publisher","first-page":"7763","DOI":"10.1038\/srep07763","article-title":"Extracellular vesicle-mediated export of fungal RNA","volume":"5","author":"Peres da Silva","year":"2015","journal-title":"Sci. Rep."},{"key":"B37","doi-asserted-by":"publisher","first-page":"346","DOI":"10.3389\/fcimb.2020.00346","article-title":"Extracellular vesicles in fungi: past, present, and future perspectives","volume":"10","author":"Rizzo","year":"2020","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"e12129","DOI":"10.1002\/jev2.12129","article-title":"Cryptococcus extracellular vesicles properties and their use as vaccine platforms","volume":"10","author":"Rizzo","year":"2021","journal-title":"J. Extracell. Vesicles"},{"key":"B39","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1128\/EC.00318-06","article-title":"Vesicular polysaccharide export in Cryptococcus neoformans is a eukaryotic solution to the problem of fungal trans-cell wall transport","volume":"6","author":"Rodrigues","year":"2007","journal-title":"Eukaryot. Cell."},{"key":"B40","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1128\/EC.00370-07","article-title":"Extracellular vesicles produced by Cryptococcus neoformans contain protein components associated with virulence","volume":"7","author":"Rodrigues","year":"2008","journal-title":"Eukaryot. Cell."},{"key":"B41","doi-asserted-by":"publisher","first-page":"825","DOI":"10.1038\/s41564-025-01962-4","article-title":"Characterizing extracellular vesicles of human fungal pathogens","volume":"10","author":"Rodrigues","year":"2025","journal-title":"Nat. Microbiol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"myaf037","DOI":"10.1093\/mmy\/myaf037","article-title":"Optimization of cutoff values for (1\u21923)-\u03b2-D-glucan and galactomannan assays in cerebrospinal fluid for the diagnosis of non-cryptococcal fungal infections of the central nervous system","volume":"63","author":"Samaddar","year":"2025","journal-title":"Med. Mycol."},{"key":"B43","doi-asserted-by":"publisher","first-page":"32115","DOI":"10.1038\/s41598-025-15577-6","article-title":"Functional roles of purified yapsins from Candida glabrata (Nakaseomyces glabratus) in immune modulation and cross-species biofilm formation","volume":"15","author":"Satala","year":"2025","journal-title":"Sci. Rep."},{"key":"B44","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1186\/s12987-025-00644-x","article-title":"Hyphal penetration is the major pathway of translocation of Candida albicans across the blood\u2013cerebrospinal fluid barrier","volume":"22","author":"Schmidt","year":"2025","journal-title":"Fluids Barriers CNS"},{"key":"B45","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1111\/tra.12558","article-title":"The biology of extracellular microvesicles","volume":"19","author":"Sedgwick","year":"2018","journal-title":"Traffic"},{"key":"B46","doi-asserted-by":"publisher","first-page":"e18564","DOI":"10.1371\/journal.pone.0018564","article-title":"Exosomes released from M. tuberculosis infected cells can suppress IFN-\u03b3 mediated activation of na\u00efve macrophages","volume":"6","author":"Singh","year":"2011","journal-title":"PLoS One"},{"key":"B47","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1007\/s11046-025-01035-4","article-title":"Diagnostic approaches to invasive candidiasis: challenges and new perspectives","volume":"191","author":"Slep\u010danov\u00e1","year":"2026","journal-title":"Mycopathologia"},{"key":"B48","doi-asserted-by":"publisher","first-page":"55","DOI":"10.3390\/jof12010055","article-title":"Diagnostic biomarkers for invasive candidiasis: a clinician-oriented review","volume":"12","author":"Smadu","year":"2026","journal-title":"J. Fungi"},{"key":"B49","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1038\/nrneurol.2016.68","article-title":"Extracellular vesicles in neurodegenerative disease \u2014 pathogenesis to biomarkers","volume":"12","author":"Thompson","year":"2016","journal-title":"Nat. Rev. Neurol."},{"key":"B50","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1111\/cmi.12374","article-title":"Compositional and immunobiological analyses of extracellular vesicles released by Candida albicans","volume":"17","author":"Vargas","year":"2015","journal-title":"Cell. Microbiol."},{"key":"B51","doi-asserted-by":"publisher","first-page":"e02383-17","DOI":"10.1128\/mBio.02383-17","article-title":"The viscoelastic properties of the fungal cell wall allow traffic of AmBisome as intact liposome vesicles","volume":"9","author":"Walker","year":"2018","journal-title":"mBio"},{"key":"B52","doi-asserted-by":"publisher","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":"B53","doi-asserted-by":"publisher","first-page":"1484","DOI":"10.1128\/EC.00111-14","article-title":"Interaction of Cryptococcus neoformans extracellular vesicles with the cell wall","volume":"13","author":"Wolf","year":"2014","journal-title":"Eukaryot. Cell."},{"key":"B54","volume-title":"WHO Fungal Priority Pathogens List to Guide Research, Development and Public Health Action","year":"2022"},{"key":"B55","volume-title":"Landscape Analysis of Commercially Available and Pipeline in Vitro Diagnostics for Fungal Priority Pathogens","year":"2025"},{"key":"B56","doi-asserted-by":"publisher","first-page":"2455508","DOI":"10.1080\/19490976.2025.2455508","article-title":"PMA1-containing extracellular vesicles of Candida albicans triggers immune responses and colitis progression","volume":"17","author":"Xu","year":"2025","journal-title":"Gut Microbes"},{"key":"B57","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1007\/978-1-0716-2417-3_1","article-title":"Isolation of Candida auris in clinical specimens","volume":"2517","author":"Yadav","year":"2022","journal-title":"Methods Mol. Biol."},{"key":"B58","doi-asserted-by":"publisher","first-page":"e00822-21","DOI":"10.1128\/mSystems.00822-21","article-title":"Comparative molecular and immunoregulatory analysis of extracellular vesicles from Candida albicans and Candida auris","volume":"6","author":"Zamith-Miranda","year":"2021","journal-title":"mSystems"},{"key":"B59","doi-asserted-by":"publisher","first-page":"e2006872","DOI":"10.1371\/journal.pbio.2006872","article-title":"Candida albicans biofilm-induced vesicles confer drug resistance through matrix biogenesis","volume":"16","author":"Zarnowski","year":"2018","journal-title":"PLoS Biol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1038\/s41556-018-0040-4","article-title":"Identification of distinct nanoparticles and subsets of extracellular vesicles by asymmetric flow field-flow fractionation","volume":"20","author":"Zhang","year":"2018","journal-title":"Nat. Cell. Biol."},{"key":"B61","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1038\/s42003-019-0538-8","article-title":"Extracellular vesicles secreted by Saccharomyces cerevisiae are involved in cell wall remodelling","volume":"2","author":"Zhao","year":"2019","journal-title":"Commun. Biol."}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2026.1854037\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T05:37:28Z","timestamp":1785476248000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2026.1854037\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,31]]},"references-count":61,"alternative-id":["10.3389\/fcell.2026.1854037"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2026.1854037","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,31]]},"article-number":"1854037"}}