{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T18:27:27Z","timestamp":1786991247063,"version":"build-2736575974"},"update-to":[{"DOI":"10.1371\/journal.ppat.1014530","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T00:00:00Z","timestamp":1786924800000}}],"reference-count":59,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T00:00:00Z","timestamp":1786320000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100007660","name":"Universiteit Antwerpen","doi-asserted-by":"publisher","award":["53274"],"award-info":[{"award-number":["53274"]}],"id":[{"id":"10.13039\/501100007660","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003130","name":"Fonds Wetenschappelijk Onderzoek","doi-asserted-by":"publisher","award":["1143426N"],"award-info":[{"award-number":["1143426N"]}],"id":[{"id":"10.13039\/501100003130","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004418","name":"Vrije Universiteit Brussel","doi-asserted-by":"publisher","award":["SRP95"],"award-info":[{"award-number":["SRP95"]}],"id":[{"id":"10.13039\/501100004418","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000865","name":"Bill and Melinda Gates Foundation","doi-asserted-by":"publisher","award":["INV031353"],"award-info":[{"award-number":["INV031353"]}],"id":[{"id":"10.13039\/100000865","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>\n                    Human African trypanosomiasis (HAT) remains a significant health burden in sub-Saharan Africa, with serological diagnosis relying heavily on parasite variant surface glycoproteins (VSGs). Experimentally validated structural studies of VSGs from\n                    <jats:italic>Trypanosoma brucei gambiense<\/jats:italic>\n                    (the main HAT causative agent) are currently lacking despite their diagnostic importance. In this study, we present a thorough characterization of LiTat 1.3, a key VSG in the diagnosis of\n                    <jats:italic>T. b. gambiense<\/jats:italic>\n                    infections, through an integrative structural biology approach encompassing AlphaFold-based structure prediction, analytical gel filtration (AGF), size exclusion chromatography with multi-angle light scattering (SEC-MALS), and small-angle X-ray scattering (SAXS). While sequence-based analyses had previously proposed LiTat 1.3 as a homotrimeric B2 class VSG, experimental evidence was yet to be presented. Here, we demonstrate that LiTat 1.3 displays a stable, concentration-independent homotrimeric architecture in solution, which is distinct from other trimeric\n                    <jats:italic>T. brucei brucei<\/jats:italic>\n                    VSGs described to date. Furthermore, the SAXS data demonstrate that the C-terminal domains of trimeric VSGs exhibit the same degree of flexibility as observed in dimeric VSGs. Hence, the biophysical characterization of LiTat 1.3 VSG adds to the growing body of knowledge that certain VSG classes occur as homotrimers instead of homodimers. Together, these results not only expand our knowledge of VSG structure and flexibility but also provide the first detailed structural insight into a major diagnostic VSG of\n                    <jats:italic>T. b. gambiense<\/jats:italic>\n                    , which holds potential for understanding antibody recognition and guiding the development of improved serological tests.\n                  <\/jats:p>","DOI":"10.1371\/journal.ppat.1014530","type":"journal-article","created":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T17:43:19Z","timestamp":1786383799000},"page":"e1014530","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["The major diagnostic VSG LiTat 1.3 of the human parasite Trypanosoma brucei gambiense is a trimer in solution"],"prefix":"10.1371","volume":"22","author":[{"given":"Niki","family":"Danel","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rob","family":"Geens","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Israel","family":"Mares-Mej\u00eda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wim","family":"Vers\u00e9es","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Van Den Abbeele","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7420-0983","authenticated-orcid":true,"given":"Yann G. J.","family":"Sterckx","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,8,10]]},"reference":[{"issue":"10110","key":"ppat.1014530.ref001","doi-asserted-by":"crossref","first-page":"2397","DOI":"10.1016\/S0140-6736(17)31510-6","article-title":"Human African trypanosomiasis","volume":"390","author":"P B\u00fcscher","year":"2017","journal-title":"The Lancet"},{"key":"ppat.1014530.ref002"},{"issue":"1","key":"ppat.1014530.ref003","doi-asserted-by":"crossref","first-page":"7083","DOI":"10.1038\/s41467-022-34757-w","article-title":"Impact of pulmonary African trypanosomes on the immunology and function of the lung","volume":"13","author":"D Mabille","year":"2022","journal-title":"Nat Commun"},{"issue":"12","key":"ppat.1014530.ref004","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.1016\/j.pt.2018.08.002","article-title":"Beyond Blood: African Trypanosomes on the Move","volume":"34","author":"T Kr\u00fcger","year":"2018","journal-title":"Trends Parasitol"},{"key":"ppat.1014530.ref005","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.17716","article-title":"The skin is a significant but overlooked anatomical reservoir for vector-borne African trypanosomes","volume":"5","author":"P Capewell","year":"2016","journal-title":"Elife"},{"issue":"6","key":"ppat.1014530.ref006","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1016\/j.chom.2016.05.002","article-title":"Trypanosoma brucei Parasites Occupy and Functionally Adapt to the Adipose Tissue in Mice","volume":"19","author":"S Trindade","year":"2016","journal-title":"Cell Host Microbe"},{"issue":"7","key":"ppat.1014530.ref007","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1005744","article-title":"The Dermis as a Delivery Site of Trypanosoma brucei for Tsetse Flies","volume":"12","author":"G Caljon","year":"2016","journal-title":"PLoS Pathog"},{"issue":"1131","key":"ppat.1014530.ref008","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1098\/rstb.1984.0106","article-title":"Antigenic variation in its biological context","volume":"307","author":"MJ Turner","year":"1984","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ppat.1014530.ref009","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1146\/annurev.iy.08.040190.000503","article-title":"Cellular and genetic aspects of antigenic variation in trypanosomes","volume":"8","author":"GA Cross","year":"1990","journal-title":"Annu Rev Immunol"},{"issue":"6229","key":"ppat.1014530.ref010","doi-asserted-by":"crossref","first-page":"1470","DOI":"10.1126\/science.aaa4502","article-title":"The in vivo dynamics of antigenic variation in Trypanosoma brucei","volume":"347","author":"MR Mugnier","year":"2015","journal-title":"Science"},{"key":"ppat.1014530.ref011","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1007\/978-3-319-20819-0_2","article-title":"A Host-Pathogen Interaction Reduced to First Principles: Antigenic Variation in T. brucei","volume":"57","author":"G Hovel-Miner","year":"2015","journal-title":"Results Probl Cell Differ"},{"key":"ppat.1014530.ref012","doi-asserted-by":"crossref","first-page":"233","DOI":"10.3389\/fimmu.2016.00233","article-title":"Immune Evasion Strategies of Trypanosoma brucei within the Mammalian Host: Progression to Pathogenicity","volume":"7","author":"B Stijlemans","year":"2016","journal-title":"Front Immunol"},{"issue":"9","key":"ppat.1014530.ref013","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1005784","article-title":"Masters of Disguise: Antigenic Variation and the VSG Coat in Trypanosoma brucei","volume":"12","author":"MR Mugnier","year":"2016","journal-title":"PLoS Pathog"},{"issue":"9","key":"ppat.1014530.ref014","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0011621","article-title":"A structural classification of the variant surface glycoproteins of the African trypanosome","volume":"17","author":"S \u0110akovi\u0107","year":"2023","journal-title":"PLoS Negl Trop Dis"},{"issue":"8","key":"ppat.1014530.ref015","doi-asserted-by":"crossref","first-page":"932","DOI":"10.1038\/s41564-018-0187-6","article-title":"African trypanosomes evade immune clearance by O-glycosylation of the VSG surface coat","volume":"3","author":"J Pinger","year":"2018","journal-title":"Nat Microbiol"},{"issue":"2","key":"ppat.1014530.ref016","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0011093","article-title":"Structural similarities between the metacyclic and bloodstream form variant surface glycoproteins of the African trypanosome","volume":"17","author":"M Chandra","year":"2023","journal-title":"PLoS Negl Trop Dis"},{"issue":"11","key":"ppat.1014530.ref017","doi-asserted-by":"crossref","first-page":"1523","DOI":"10.1038\/s41564-017-0013-6","article-title":"Structural basis for the shielding function of the dynamic trypanosome variant surface glycoprotein coat","volume":"2","author":"T Bartossek","year":"2017","journal-title":"Nat Microbiol"},{"issue":"4","key":"ppat.1014530.ref018","doi-asserted-by":"crossref","DOI":"10.1371\/journal.ppat.1012186","article-title":"Beyond the VSG layer: Exploring the role of intrinsic disorder in the invariant surface glycoproteins of African trypanosomes","volume":"20","author":"H S\u00fclzen","year":"2024","journal-title":"PLoS Pathog"},{"key":"ppat.1014530.ref019","doi-asserted-by":"crossref","first-page":"1037094","DOI":"10.3389\/fmed.2022.1037094","article-title":"Recent progress in diagnosis and treatment of Human African Trypanosomiasis has made the elimination of this disease a realistic target by 2030","volume":"9","author":"A \u00c1lvarez-Rodr\u00edguez","year":"2022","journal-title":"Front Med (Lausanne)"},{"issue":"984","key":"ppat.1014530.ref020","first-page":"1","article-title":"Control and surveillance of human African trypanosomiasis","author":"World Health Organization","year":"2013","journal-title":"World Health Organ Tech Rep Ser"},{"issue":"2","key":"ppat.1014530.ref021","doi-asserted-by":"crossref","first-page":"246","DOI":"10.3390\/diagnostics12020246","article-title":"Two-Year Follow-Up of Trypanosoma brucei gambiense Serology after Successful Treatment of Human African Trypanosomiasis: Results of Four Different Sero-Diagnostic Tests","volume":"12","author":"R Inocencio da Luz","year":"2022","journal-title":"Diagnostics (Basel)"},{"issue":"6","key":"ppat.1014530.ref022","article-title":"VSGs expressed during natural T. b. gambiense infection exhibit extensive sequence divergence and a subspecies-specific bias towards type B N-terminal domains","volume":"13","author":"J So","year":"2022","journal-title":"mBio"},{"issue":"5","key":"ppat.1014530.ref023","first-page":"761","article-title":"Affinity of concanavalin A for \u201ctrypanosoma equiperdum\u201d: purification of the antigenic type specific glycoprotein fraction (author\u2019s transl)","volume":"127","author":"T Baltz","year":"1976","journal-title":"Ann Immunol (Paris)"},{"issue":"6","key":"ppat.1014530.ref024","doi-asserted-by":"crossref","first-page":"1698","DOI":"10.1107\/S1600576721008736","article-title":"BioSAXS on the SWING beamline at Synchrotron SOLEIL","volume":"54","author":"A Thureau","year":"2021","journal-title":"J Appl Crystallogr"},{"issue":"2","key":"ppat.1014530.ref025","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1107\/S0021889899015216","article-title":"SAXS experiments on absolute scale with Kratky systems using water as a secondary standard","volume":"33","author":"D Orthaber","year":"2000","journal-title":"J Appl Crystallogr"},{"issue":"11","key":"ppat.1014530.ref026","doi-asserted-by":"crossref","first-page":"1944","DOI":"10.1093\/bioinformatics\/btx846","article-title":"CHROMIXS: automatic and interactive analysis of chromatography-coupled small-angle X-ray scattering data","volume":"34","author":"A Panjkovich","year":"2018","journal-title":"Bioinformatics"},{"key":"ppat.1014530.ref027","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1107\/S1600576720013412","article-title":"ATSAS 3.0: expanded functionality and new tools for small-angle scattering data analysis","volume":"54","author":"K Manalastas-Cantos","year":"2021","journal-title":"J Appl Crystallogr"},{"key":"ppat.1014530.ref028","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1107\/S1600576723011019","article-title":"BioXTAS RAW 2: new developments for a free open-source program for small-angle scattering data reduction and analysis","volume":"57","author":"JB Hopkins","year":"2024","journal-title":"J Appl Crystallogr"},{"key":"ppat.1014530.ref029","article-title":"Protein complex prediction with AlphaFold-Multimer","author":"R Evans","year":"2022","journal-title":"bioRxiv"},{"issue":"7","key":"ppat.1014530.ref030","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pntd.0003006","article-title":"Recombinant antigens expressed in Pichia pastoris for the diagnosis of sleeping sickness caused by Trypanosoma brucei gambiense","volume":"8","author":"S Rog\u00e9","year":"2014","journal-title":"PLoS Negl Trop Dis"},{"issue":"7","key":"ppat.1014530.ref031","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":"ppat.1014530.ref032","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1016\/j.crbiot.2021.01.001","article-title":"Prediction of GPI-anchored proteins with pointer neural networks","volume":"3","author":"MH G\u00edslason","year":"2021","journal-title":"Curr Res Biotechnol"},{"key":"ppat.1014530.ref033","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkq461","article-title":"FoXS: a web server for rapid computation and fitting of SAXS profiles","volume":"38","author":"D Schneidman-Duhovny","year":"2010","journal-title":"Nucleic Acids Res"},{"issue":"2","key":"ppat.1014530.ref034","doi-asserted-by":"crossref","first-page":"174","DOI":"10.4149\/gpb_2009_02_174","article-title":"Structure and flexibility within proteins as identified through small angle X-ray scattering","volume":"28","author":"M Pelikan","year":"2009","journal-title":"Gen Physiol Biophys"},{"key":"ppat.1014530.ref035","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkw389","article-title":"FoXS, FoXSDock and MultiFoXS: Single-state and multi-state structural modeling of proteins and their complexes based on SAXS profiles","volume":"44","author":"D Schneidman-Duhovny","year":"2016","journal-title":"Nucleic Acids Res"},{"issue":"4","key":"ppat.1014530.ref036","doi-asserted-by":"crossref","first-page":"962","DOI":"10.1016\/j.bpj.2013.07.020","article-title":"Accurate SAXS profile computation and its assessment by contrast variation experiments","volume":"105","author":"D Schneidman-Duhovny","year":"2013","journal-title":"Biophys J"},{"issue":"1","key":"ppat.1014530.ref037","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1002\/pro.3731","article-title":"SASBDB: Towards an automatically curated and validated repository for biological scattering data","volume":"29","author":"AG Kikhney","year":"2019","journal-title":"Protein Science"},{"key":"ppat.1014530.ref038","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkad947","article-title":"PED in 2024: improving the community deposition of structural ensembles for intrinsically disordered proteins","volume":"52","author":"H Ghafouri","year":"2024","journal-title":"Nucleic Acids Res"},{"key":"ppat.1014530.ref039","article-title":"Metapredict Enables Accurate Disorder Prediction Across the Tree of Life","author":"JM Lotthammer","year":"2024"},{"key":"ppat.1014530.ref040","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkab408","article-title":"IUPred3: prediction of protein disorder enhanced with unambiguous experimental annotation and visualization of evolutionary conservation","volume":"49","author":"G Erd\u0151s","year":"2021","journal-title":"Nucleic Acids Res"},{"key":"ppat.1014530.ref041","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkae385","article-title":"AIUPred: combining energy estimation with deep learning for the enhanced prediction of protein disorder","volume":"52","author":"G Erd\u0151s","year":"2024","journal-title":"Nucleic Acids Res"},{"key":"ppat.1014530.ref042","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gkm363","article-title":"PrDOS: prediction of disordered protein regions from amino acid sequence","volume":"35","author":"T Ishida","year":"2007","journal-title":"Nucleic Acids Res"},{"issue":"11","key":"ppat.1014530.ref043","article-title":"UCSF ChimeraX: Tools for structure building and analysis","volume":"32","author":"EC Meng","year":"2023","journal-title":"Protein Sci"},{"key":"ppat.1014530.ref044"},{"issue":"7873","key":"ppat.1014530.ref045","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1038\/s41586-021-03828-1","article-title":"Highly accurate protein structure prediction for the human proteome","volume":"596","author":"K Tunyasuvunakool","year":"2021","journal-title":"Nature"},{"key":"ppat.1014530.ref046","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1016\/j.crstbi.2020.08.004","article-title":"Methods, development and applications of small-angle X-ray scattering to characterize biological macromolecules in solution","volume":"2","author":"S Da Vela","year":"2020","journal-title":"Curr Res Struct Biol"},{"key":"ppat.1014530.ref047","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/j.sbi.2019.04.004","article-title":"Evolving SAXS versatility: solution X-ray scattering for macromolecular architecture, functional landscapes, and integrative structural biology","volume":"58","author":"CA Brosey","year":"2019","journal-title":"Curr Opin Struct Biol"},{"issue":"1","key":"ppat.1014530.ref048","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.jsb.2009.08.009","article-title":"NADPH oxidase activator p67(phox) behaves in solution as a multidomain protein with semi-flexible linkers","volume":"169","author":"D Durand","year":"2010","journal-title":"J Struct Biol"},{"key":"ppat.1014530.ref049","doi-asserted-by":"crossref","first-page":"1412","DOI":"10.1107\/S1600576716010906","article-title":"Guinier peak analysis for visual and automated inspection of small-angle X-ray scattering data","volume":"49","author":"CD Putnam","year":"2016","journal-title":"J Appl Crystallogr"},{"key":"ppat.1014530.ref050","volume-title":"Report of the fifth WHO stakeholders meeting on gambiense and rhodesiense human African trypanosomiasis elimination","author":"World Health Organization","year":"2024"},{"issue":"3","key":"ppat.1014530.ref051","first-page":"169","article-title":"A card-agglutination test with stained trypanosomes (C.A.T.T.) for the serological diagnosis of T. b. gambiense trypanosomiasis","volume":"58","author":"E Magnus","year":"1978","journal-title":"Ann Soc Belg Med Trop"},{"issue":"6","key":"ppat.1014530.ref052","doi-asserted-by":"crossref","DOI":"10.1016\/S2214-109X(14)70203-7","article-title":"Sensitivity and specificity of HAT Sero-K-SeT, a rapid diagnostic test for serodiagnosis of sleeping sickness caused by Trypanosoma brucei gambiense: a case-control study","volume":"2","author":"P B\u00fcscher","year":"2014","journal-title":"Lancet Glob Health"},{"issue":"8","key":"ppat.1014530.ref053","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1111\/tra.12806","article-title":"Dynamic, variable oligomerization and the trafficking of variant surface glycoproteins of Trypanosoma brucei","volume":"22","author":"K Umaer","year":"2021","journal-title":"Traffic"},{"key":"ppat.1014530.ref054","article-title":"Ensemblify: a user-friendly tool for generating ensembles of intrinsically disordered regions of AlphaFold and user-defined models","volume":"2025","author":"NP Fernandes","year":"2025","journal-title":"bioRxiv"},{"issue":"15","key":"ppat.1014530.ref055","doi-asserted-by":"crossref","first-page":"169003","DOI":"10.1016\/j.jmb.2025.169003","article-title":"AFflecto: A web server to generate conformational ensembles of flexible proteins from AlphaFold models","volume":"437","author":"M Pajkos","year":"2025","journal-title":"J Mol Biol"},{"issue":"1","key":"ppat.1014530.ref056","doi-asserted-by":"crossref","first-page":"10394","DOI":"10.1038\/srep10394","article-title":"The molecular size of the extra-membrane domain influences the diffusion of the GPI-anchored VSG on the trypanosome plasma membrane","volume":"5","author":"AJW Hartel","year":"2015","journal-title":"Sci Rep"},{"issue":"3","key":"ppat.1014530.ref057","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1016\/j.cell.2007.08.046","article-title":"Hydrodynamic flow-mediated protein sorting on the cell surface of trypanosomes","volume":"131","author":"M Engstler","year":"2007","journal-title":"Cell"},{"issue":"3","key":"ppat.1014530.ref058","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/0001-706X(95)00127-Z","article-title":"Evaluation of variant specific trypanolysis tests for serodiagnosis of human infections with Trypanosoma brucei gambiense","volume":"60","author":"N Van Meirvenne","year":"1995","journal-title":"Acta Trop"},{"issue":"2","key":"ppat.1014530.ref059","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1111\/tmi.14077","article-title":"Retrospective clinical performance evaluation of the Abbott Bioline HAT 2.0, a rapid diagnostic test for human African trypanosomiasis based on recombinant antigens","volume":"30","author":"S Tablado Alonso","year":"2025","journal-title":"Trop Med Int Health"}],"container-title":["PLOS Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1014530","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T17:40:15Z","timestamp":1786988415000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1014530"}},"subtitle":[],"editor":[{"given":"Dominique","family":"Soldati-Favre","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,8,10]]},"references-count":59,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8,10]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.ppat.1014530","relation":{},"ISSN":["1553-7374"],"issn-type":[{"value":"1553-7374","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,10]]}}}