{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T00:17:40Z","timestamp":1784247460893,"version":"3.55.0"},"update-to":[{"DOI":"10.1371\/journal.pntd.0013401","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,8,21]],"date-time":"2025-08-21T00:00:00Z","timestamp":1755734400000}}],"reference-count":64,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2025,8,13]],"date-time":"2025-08-13T00:00:00Z","timestamp":1755043200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004587","name":"Instituto de Salud Carlos III","doi-asserted-by":"publisher","award":["PI20CIII-00037"],"award-info":[{"award-number":["PI20CIII-00037"]}],"id":[{"id":"10.13039\/501100004587","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100012818","name":"Comunidad de Madrid","doi-asserted-by":"publisher","award":["TEC-2024\/BIO-66"],"award-info":[{"award-number":["TEC-2024\/BIO-66"]}],"id":[{"id":"10.13039\/100012818","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosntds.org"],"crossmark-restriction":false},"short-container-title":["PLoS Negl Trop Dis"],"abstract":"<jats:p><jats:italic><jats:italic>Babesia divergens<\/jats:italic><\/jats:italic> is a blood-borne parasite that invades, replicates within and destroys red blood cells (RBCs) during its asexual life cycle, causing babesiosis in humans and cattle. This study focuses on BdP50, a putative <jats:italic><jats:italic>B. divergens<\/jats:italic><\/jats:italic> glycosylphosphatidylinositol-anchored protein involved in the parasite life cycle. BdP50 is found on the surface of <jats:italic><jats:italic>B. divergens<\/jats:italic><\/jats:italic> invasive parasites (merozoites) as well as on extracellular vesicles (<jats:italic><jats:italic>Bd<\/jats:italic><\/jats:italic>-derived EVs). These EVs are secreted by parasites cultured in fresh human RBCs and, in addition to BdP50, are enriched in human and parasite proteins, including proteins related to the parasite invasion process. BdP50 binds to RBCs and could mediate interactions of free merozoites and <jats:italic><jats:italic>Bd<\/jats:italic><\/jats:italic>-derived EVs with the host cell. Anti-BdP50 antibodies support this by blocking the BdP50 protein and inhibiting up to 88% of merozoite entry into na\u00efve RBCs. This reinforces the role of BdP50 in parasite-host cell interactions and invasion. However, the inhibitory effect of BdP50 antibodies begins to gradually decrease slightly several hours after invasion, leading to a progressive increase in <jats:italic><jats:italic>B. divergens<\/jats:italic><\/jats:italic> infected RBCs over time. Consistent with these findings, our in vitro <jats:italic><jats:italic>de novo<\/jats:italic><\/jats:italic> infection assays showed that <jats:italic><jats:italic>Bd<\/jats:italic><\/jats:italic>-derived EVs, in addition to promoting parasite propagation, display proteins such as BdP50 that mimic the merozoite surface to likely attenuate the blocking effect of antibodies, thereby ensuring the parasite survival during subsequent rounds of invasion and growth. Given the role of BdP50 and <jats:italic><jats:italic>Bd<\/jats:italic><\/jats:italic>-derived EVs in the <jats:italic><jats:italic>B. divergens<\/jats:italic><\/jats:italic> life cycle<jats:italic>,<\/jats:italic> this study could have future implications for developing new approaches to interfere with parasite invasion proteins and <jats:italic><jats:italic>Bd<\/jats:italic><\/jats:italic>-derived EVs functions.<\/jats:p>","DOI":"10.1371\/journal.pntd.0013401","type":"journal-article","created":{"date-parts":[[2025,8,13]],"date-time":"2025-08-13T18:31:53Z","timestamp":1755109913000},"page":"e0013401","update-policy":"https:\/\/doi.org\/10.1371\/journal.pntd.corrections_policy","source":"Crossref","is-referenced-by-count":2,"title":["Elucidating the functional role of the novel BdP50 protein and extracellular vesicles in the human erythrocyte infection by Babesia divergens"],"prefix":"10.1371","volume":"19","author":[{"given":"Luis Miguel","family":"Gonzalez","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Belen","family":"Revuelta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aitor","family":"Gil","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda C.","family":"Terr\u00f3n","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin Christoph","family":"Sachse","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Javier","family":"Sotillo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel","family":"Luque","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S M Raihan","family":"Rahman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Reginaldo G.","family":"Bastos","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carlos Esteban","family":"Suarez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3852-960X","authenticated-orcid":true,"given":"Estrella","family":"Montero","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,8,13]]},"reference":[{"issue":"11","key":"pntd.0013401.ref001","doi-asserted-by":"crossref","first-page":"1447","DOI":"10.3390\/pathogens10111447","article-title":"The Global Emergence of Human Babesiosis","volume":"10","author":"A Kumar","year":"2021","journal-title":"Pathogens"},{"issue":"9","key":"pntd.0013401.ref002","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.3390\/pathogens10091165","article-title":"Human Babesiosis in Europe","volume":"10","author":"A Hildebrandt","year":"2021","journal-title":"Pathogens"},{"issue":"4","key":"pntd.0013401.ref003","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1128\/CMR.16.4.622-636.2003","article-title":"Babesia divergens, a bovine blood parasite of veterinary and zoonotic importance","volume":"16","author":"A Zintl","year":"2003","journal-title":"Clin Microbiol Rev"},{"issue":"11","key":"pntd.0013401.ref004","doi-asserted-by":"crossref","first-page":"1430","DOI":"10.3390\/pathogens10111430","article-title":"Ticks, Human Babesiosis and Climate Change","volume":"10","author":"JS Gray","year":"2021","journal-title":"Pathogens"},{"issue":"5","key":"pntd.0013401.ref005","doi-asserted-by":"crossref","DOI":"10.1128\/mSphere.00928-20","article-title":"Four-Dimensional Characterization of the Babesia divergens Asexual Life Cycle, from the Trophozoite to the Multiparasite Stage","volume":"5","author":"JJ Conesa","year":"2020","journal-title":"mSphere"},{"issue":"6","key":"pntd.0013401.ref006","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1111\/cmi.12555","article-title":"Babesia divergens builds a complex population structure composed of specific ratios of infected cells to ensure a prompt response to changing environmental conditions","volume":"18","author":"JR Cursino-Santos","year":"2016","journal-title":"Cell Microbiol"},{"issue":"1","key":"pntd.0013401.ref007","doi-asserted-by":"crossref","first-page":"14116","DOI":"10.1038\/s41598-018-32349-7","article-title":"Kinetics of the invasion and egress processes of Babesia divergens, observed by time-lapse video microscopy","volume":"8","author":"E Sevilla","year":"2018","journal-title":"Sci Rep"},{"issue":"7","key":"pntd.0013401.ref008","doi-asserted-by":"crossref","first-page":"1052","DOI":"10.1111\/cmi.12422","article-title":"The apicomplexan parasite Babesia divergens internalizes band 3, glycophorin A and spectrin during invasion of human red blood cells","volume":"17","author":"U Repnik","year":"2015","journal-title":"Cell Microbiol"},{"issue":"8","key":"pntd.0013401.ref009","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0007680","article-title":"Comparative and functional genomics of the protozoan parasite Babesia divergens highlighting the invasion and egress processes","volume":"13","author":"LM Gonz\u00e1lez","year":"2019","journal-title":"PLoS Negl Trop Dis"},{"issue":"6","key":"pntd.0013401.ref010","doi-asserted-by":"crossref","first-page":"797","DOI":"10.1111\/j.1462-5822.2011.01597.x","article-title":"The moving junction of apicomplexan parasites: a key structure for invasion","volume":"13","author":"S Besteiro","year":"2011","journal-title":"Cell Microbiol"},{"issue":"11","key":"pntd.0013401.ref011","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1038\/nrmicro.2017.86","article-title":"Gliding motility powers invasion and egress in Apicomplexa","volume":"15","author":"K Fr\u00e9nal","year":"2017","journal-title":"Nat Rev Microbiol"},{"issue":"47","key":"pntd.0013401.ref012","doi-asserted-by":"crossref","first-page":"35717","DOI":"10.1074\/jbc.M604344200","article-title":"A conserved subtilisin protease identified in Babesia divergens merozoites","volume":"281","author":"E Montero","year":"2006","journal-title":"J Biol Chem"},{"issue":"1","key":"pntd.0013401.ref013","doi-asserted-by":"crossref","first-page":"99","DOI":"10.3390\/pathogens11010099","article-title":"Major Surface Antigens in Zoonotic Babesia","volume":"11","author":"S Delbecq","year":"2022","journal-title":"Pathogens"},{"issue":"11","key":"pntd.0013401.ref014","doi-asserted-by":"crossref","first-page":"4783","DOI":"10.1128\/IAI.00969-08","article-title":"Babesia divergens apical membrane antigen 1 and its interaction with the human red blood cell","volume":"77","author":"E Montero","year":"2009","journal-title":"Infect Immun"},{"issue":"5","key":"pntd.0013401.ref015","doi-asserted-by":"crossref","first-page":"1574","DOI":"10.1128\/IAI.00107-16","article-title":"Identification and Characterization of the Rhoptry Neck Protein 2 in Babesia divergens and B. microti","volume":"84","author":"RL Ord","year":"2016","journal-title":"Infect Immun"},{"issue":"25","key":"pntd.0013401.ref016","doi-asserted-by":"crossref","first-page":"2397","DOI":"10.1056\/NEJMra1202018","article-title":"Human babesiosis","volume":"366","author":"E Vannier","year":"2012","journal-title":"N Engl J Med"},{"issue":"2","key":"pntd.0013401.ref017","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1007\/s12104-012-9418-6","article-title":"\u00b9H, \u00b9\u2075N and \u00b9\u00b3C Backbone resonance assignments of a conformational mutant of the adhesion protein \u2206-Bd37 from Babesia divergens","volume":"7","author":"P Barthe","year":"2013","journal-title":"Biomol NMR Assign"},{"key":"pntd.0013401.ref018","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1017\/S0031182002002160","article-title":"Babesia divergens: cloning and biochemical characterization of Bd37","volume":"125","author":"S Delbecq","year":"2002","journal-title":"Parasitology"},{"issue":"5","key":"pntd.0013401.ref019","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1016\/j.vaccine.2005.08.073","article-title":"Hydrophobic moeties in recombinant proteins are crucial to generate efficient saponin-based vaccine against Apicomplexan Babesia divergens","volume":"24","author":"S Delbecq","year":"2006","journal-title":"Vaccine"},{"issue":"12","key":"pntd.0013401.ref020","doi-asserted-by":"crossref","first-page":"9495","DOI":"10.1074\/jbc.M111.260745","article-title":"Structural and functional characterization of Bc28.1, major erythrocyte-binding protein from Babesia canis merozoite surface","volume":"287","author":"Y-S Yang","year":"2012","journal-title":"J Biol Chem"},{"key":"pntd.0013401.ref021","doi-asserted-by":"crossref","first-page":"3046","DOI":"10.3389\/fmicb.2019.03046","article-title":"Surface Antigen 1 Is a Crucial Secreted Protein That Mediates Babesia microti Invasion Into Host Cells","volume":"10","author":"M Li","year":"2020","journal-title":"Front Microbiol"},{"key":"pntd.0013401.ref022","doi-asserted-by":"crossref","first-page":"720536","DOI":"10.3389\/fcell.2021.720536","article-title":"The Glycosylphosphatidylinositol Anchor: A Linchpin for Cell Surface Versatility of Trypanosomatids","volume":"9","author":"AR Borges","year":"2021","journal-title":"Front Cell Dev Biol"},{"key":"pntd.0013401.ref023","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.biochi.2019.09.007","article-title":"Babesia divergens glycosylphosphatidylinositols modulate blood coagulation and induce Th2-biased cytokine profiles in antigen presenting cells","volume":"167","author":"F Debierre-Grockiego","year":"2019","journal-title":"Biochimie"},{"issue":"1","key":"pntd.0013401.ref024","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/BF00934377","article-title":"Cytological and immunological responses to Babesia divergens in different hosts: ox, gerbil, man","volume":"77","author":"A Gorenflot","year":"1991","journal-title":"Parasitol Res"},{"issue":"7","key":"pntd.0013401.ref025","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1038\/s41587-021-01156-3","article-title":"SignalP 6.0 predicts all five types of signal peptides using protein language models","volume":"40","author":"F Teufel","year":"2022","journal-title":"Nat Biotechnol"},{"key":"pntd.0013401.ref026","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1186\/1471-2105-9-392","article-title":"PredGPI: a GPI-anchor predictor","volume":"9","author":"A Pierleoni","year":"2008","journal-title":"BMC Bioinformatics"},{"issue":"1","key":"pntd.0013401.ref027","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1186\/s12936-022-04430-0","article-title":"FT-GPI, a highly sensitive and accurate predictor of GPI-anchored proteins, reveals the composition and evolution of the GPI proteome in Plasmodium species","volume":"22","author":"LM Sauer","year":"2023","journal-title":"Malar J"},{"key":"pntd.0013401.ref028","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkab1061","article-title":"AlphaFold Protein Structure Database: massively expanding the structural coverage of protein-sequence space with high-accuracy models","volume":"50","author":"M Varadi","year":"2022","journal-title":"Nucleic Acids Res"},{"issue":"7873","key":"pntd.0013401.ref029","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with AlphaFold","volume":"596","author":"J Jumper","year":"2021","journal-title":"Nature"},{"issue":"1","key":"pntd.0013401.ref030","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1002\/pro.3235","article-title":"UCSF ChimeraX: Meeting modern challenges in visualization and analysis","volume":"27","author":"TD Goddard","year":"2018","journal-title":"Protein Sci"},{"key":"pntd.0013401.ref031","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1007\/978-1-62703-646-7_6","article-title":"Clustal Omega, accurate alignment of very large numbers of sequences","volume":"1079","author":"F Sievers","year":"2014","journal-title":"Methods Mol Biol"},{"issue":"1","key":"pntd.0013401.ref032","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1007\/BF02473201","article-title":"Improving structural integrity of cryosections for immunogold labeling","volume":"106","author":"W Liou","year":"1996","journal-title":"Histochem Cell Biol"},{"issue":"5","key":"pntd.0013401.ref033","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1038\/nmeth.4256","article-title":"MSFragger: ultrafast and comprehensive peptide identification in mass spectrometry-based proteomics","volume":"14","author":"AT Kong","year":"2017","journal-title":"Nat Methods"},{"issue":"9","key":"pntd.0013401.ref034","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1038\/s41592-020-0912-y","article-title":"Philosopher: a versatile toolkit for shotgun proteomics data analysis","volume":"17","author":"F da Veiga Leprevost","year":"2020","journal-title":"Nat Methods"},{"issue":"1","key":"pntd.0013401.ref035","doi-asserted-by":"crossref","first-page":"4539","DOI":"10.1038\/s41467-023-40129-9","article-title":"MSBooster: improving peptide identification rates using deep learning-based features","volume":"14","author":"KL Yang","year":"2023","journal-title":"Nat Commun"},{"issue":"11","key":"pntd.0013401.ref036","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.1007\/s13361-016-1460-7","article-title":"Fast and Accurate Protein False Discovery Rates on Large-Scale Proteomics Data Sets with Percolator 3.0","volume":"27","author":"M The","year":"2016","journal-title":"J Am Soc Mass Spectrom"},{"issue":"17","key":"pntd.0013401.ref037","doi-asserted-by":"crossref","first-page":"4646","DOI":"10.1021\/ac0341261","article-title":"A statistical model for identifying proteins by tandem mass spectrometry","volume":"75","author":"AI Nesvizhskii","year":"2003","journal-title":"Anal Chem"},{"key":"pntd.0013401.ref038","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkac1040","article-title":"The ProteomeXchange consortium at 10 years: 2023 update","volume":"51","author":"EW Deutsch","year":"2023","journal-title":"Nucleic Acids Res"},{"key":"pntd.0013401.ref039","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkab1038","article-title":"The PRIDE database resources in 2022: a hub for mass spectrometry-based proteomics evidences","volume":"50","author":"Y Perez-Riverol","year":"2022","journal-title":"Nucleic Acids Res"},{"key":"pntd.0013401.ref040","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/S1570-0232(02)00909-1","article-title":"Taenia solium cDNA sequence encoding a putative immunodiagnostic antigen for human cysticercosis","volume":"786","author":"E Montero","year":"2003","journal-title":"J Chromatogr B Analyt Technol Biomed Life Sci"},{"issue":"11","key":"pntd.0013401.ref041","doi-asserted-by":"crossref","first-page":"7113","DOI":"10.1093\/nar\/gku322","article-title":"The evolutionary dynamics of variant antigen genes in Babesia reveal a history of genomic innovation underlying host-parasite interaction","volume":"42","author":"AP Jackson","year":"2014","journal-title":"Nucleic Acids Res"},{"issue":"7","key":"pntd.0013401.ref042","doi-asserted-by":"crossref","first-page":"2603","DOI":"10.1128\/JCM.39.7.2603-2609.2001","article-title":"Identification and expression of a 50-kilodalton surface antigen of Babesia gibsoni and evaluation of its diagnostic potential in an enzyme-linked immunosorbent assay","volume":"39","author":"S Fukumoto","year":"2001","journal-title":"J Clin Microbiol"},{"key":"pntd.0013401.ref043","unstructured":"Goo Y-K, Jia H, Aboge GO, Terkawi MA, Yamagishi J, Nishikawa Y, et al. Identification of an immunodominant Babesia gibsoni 47-kDa antigen. 2009 [cited 20 Jul 2023]. Available: http:\/\/erepository.uonbi.ac.ke\/handle\/11295\/13368"},{"issue":"2","key":"pntd.0013401.ref044","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/j.ijpara.2018.12.002","article-title":"Vaccination against babesiosis using recombinant GPI-anchored proteins","volume":"49","author":"S Nathaly Wieser","year":"2019","journal-title":"Int J Parasitol"},{"issue":"1","key":"pntd.0013401.ref045","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1006\/expr.1993.1065","article-title":"Babesia bovis: characterization of the T helper cell response against the 42-kDa merozoite surface antigen (MSA-1) in cattle","volume":"77","author":"WC Brown","year":"1993","journal-title":"Exp Parasitol"},{"issue":"5","key":"pntd.0013401.ref046","doi-asserted-by":"crossref","first-page":"591","DOI":"10.3390\/pathogens11050591","article-title":"The GP-45 Protein, a Highly Variable Antigen from Babesia bigemina, Contains Conserved B-Cell Epitopes in Geographically Distant Isolates","volume":"11","author":"MA Mercado-Uriostegui","year":"2022","journal-title":"Pathogens"},{"issue":"6","key":"pntd.0013401.ref047","doi-asserted-by":"crossref","first-page":"3782","DOI":"10.1128\/IAI.69.6.3782-3790.2001","article-title":"Molecular basis for variable expression of merozoite surface antigen gp45 among American isolates of Babesia bigemina","volume":"69","author":"TG Fisher","year":"2001","journal-title":"Infect Immun"},{"key":"pntd.0013401.ref048","doi-asserted-by":"crossref","first-page":"962944","DOI":"10.3389\/fcimb.2022.962944","article-title":"Identification and characterization of extracellular vesicles from red cells infected with Babesia divergens and Babesia microti","volume":"12","author":"D Beri","year":"2022","journal-title":"Front Cell Infect Microbiol"},{"issue":"3","key":"pntd.0013401.ref049","doi-asserted-by":"crossref","DOI":"10.26508\/lsa.201900382","article-title":"Evidence for vesicle-mediated antigen export by the human pathogen Babesia microti","volume":"2","author":"J Thekkiniath","year":"2019","journal-title":"Life Sci Alliance"},{"issue":"8","key":"pntd.0013401.ref050","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1038\/nri855","article-title":"Exosomes: composition, biogenesis and function","volume":"2","author":"C Th\u00e9ry","year":"2002","journal-title":"Nat Rev Immunol"},{"issue":"10","key":"pntd.0013401.ref051","doi-asserted-by":"crossref","first-page":"4160","DOI":"10.1096\/fj.11-202077","article-title":"Prion-infected cells regulate the release of exosomes with distinct ultrastructural features","volume":"26","author":"BM Coleman","year":"2012","journal-title":"FASEB J"},{"issue":"24","key":"pntd.0013401.ref052","doi-asserted-by":"crossref","first-page":"17713","DOI":"10.1074\/jbc.M112.445403","article-title":"Exosome uptake depends on ERK1\/2-heat shock protein 27 signaling and lipid Raft-mediated endocytosis negatively regulated by caveolin-1","volume":"288","author":"KJ Svensson","year":"2013","journal-title":"J Biol Chem"},{"issue":"1","key":"pntd.0013401.ref053","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1038\/s41467-021-21344-8","article-title":"20S proteasomes secreted by the malaria parasite promote its growth","volume":"12","author":"E Dekel","year":"2021","journal-title":"Nat Commun"},{"issue":"3","key":"pntd.0013401.ref054","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/0248-4900(93)90126-Y","article-title":"Structural analysis of glycosyl-phosphatidylinositol membrane anchor of the Toxoplasma gondii tachyzoite surface glycoprotein gp23","volume":"78","author":"S Tomavo","year":"1993","journal-title":"Biol Cell"},{"issue":"9","key":"pntd.0013401.ref055","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0107727","article-title":"Identification and characterization of the RouenBd1987 Babesia divergens Rhopty-Associated Protein 1","volume":"9","author":"M Rodriguez","year":"2014","journal-title":"PLoS One"},{"issue":"7","key":"pntd.0013401.ref056","doi-asserted-by":"crossref","DOI":"10.15252\/embr.202255499","article-title":"Extracellular vesicles from malaria-infected red blood cells: not all are secreted equal","volume":"23","author":"F Blow","year":"2022","journal-title":"EMBO Rep"},{"issue":"4","key":"pntd.0013401.ref057","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1016\/j.chom.2015.09.007","article-title":"Processing of Plasmodium falciparum Merozoite Surface Protein MSP1 Activates a Spectrin-Binding Function Enabling Parasite Egress from RBCs","volume":"18","author":"S Das","year":"2015","journal-title":"Cell Host Microbe"},{"issue":"1","key":"pntd.0013401.ref058","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1084\/jem.20040989","article-title":"The Plasmodium circumsporozoite protein is proteolytically processed during cell invasion","volume":"201","author":"A Coppi","year":"2005","journal-title":"J Exp Med"},{"issue":"12","key":"pntd.0013401.ref059","doi-asserted-by":"crossref","first-page":"8599","DOI":"10.1016\/S0021-9258(18)42485-4","article-title":"Inositol acylation of a potential glycosyl phosphoinositol anchor precursor from yeast requires acyl coenzyme A","volume":"267","author":"LC Costello","year":"1992","journal-title":"Journal of Biological Chemistry"},{"key":"pntd.0013401.ref060","article-title":"Phosphatidylinositol-specific phospholipase C (PI-PLC) treatment of glycosylphosphatidylinositol (GPI)-anchored proteins on the cell surface of intact cells.","volume-title":"Glycoscience Protocols (GlycoPODv2)","author":"M Fujita","year":"2021"},{"issue":"11","key":"pntd.0013401.ref061","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1016\/j.ijpara.2007.03.013","article-title":"Plasmodium falciparum Pf34, a novel GPI-anchored rhoptry protein found in detergent-resistant microdomains","volume":"37","author":"NI Proellocks","year":"2007","journal-title":"Int J Parasitol"},{"issue":"3","key":"pntd.0013401.ref062","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1016\/j.ipha.2024.02.004","article-title":"Extracellular vesicles: The future of therapeutics and drug delivery systems","volume":"2","author":"MJ Uddin","year":"2024","journal-title":"Intelligent Pharmacy"},{"issue":"7","key":"pntd.0013401.ref063","doi-asserted-by":"crossref","DOI":"10.15252\/embr.202254755","article-title":"Malaria parasites release vesicle subpopulations with signatures of different destinations","volume":"23","author":"P Abou Karam","year":"2022","journal-title":"EMBO Rep"},{"issue":"7","key":"pntd.0013401.ref064","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.3000363","article-title":"The biology of extracellular vesicles: The known unknowns","volume":"17","author":"L Margolis","year":"2019","journal-title":"PLoS Biol"}],"updated-by":[{"DOI":"10.1371\/journal.pntd.0013401","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,8,21]],"date-time":"2025-08-21T00:00:00Z","timestamp":1755734400000}}],"container-title":["PLOS Neglected Tropical Diseases"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pntd.0013401","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,8,21]],"date-time":"2025-08-21T17:59:28Z","timestamp":1755799168000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pntd.0013401"}},"subtitle":[],"editor":[{"given":"Kyoko","family":"Hayashida","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2025,8,13]]},"references-count":64,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2025,8,13]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pntd.0013401","relation":{},"ISSN":["1935-2735"],"issn-type":[{"value":"1935-2735","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,13]]}}}