{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,19]],"date-time":"2026-03-19T00:46:18Z","timestamp":1773881178503,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1014077","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T00:00:00Z","timestamp":1773792000000}}],"reference-count":57,"publisher":"Public Library of Science (PLoS)","issue":"3","license":[{"start":{"date-parts":[[2026,3,11]],"date-time":"2026-03-11T00:00:00Z","timestamp":1773187200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Key Research and Development Program of China","award":["2023YFF1204200"],"award-info":[{"award-number":["2023YFF1204200"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32470688"],"award-info":[{"award-number":["32470688"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32000468"],"award-info":[{"award-number":["32000468"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100010229","name":"Natural Science Foundation of Tianjin Municipal Science and Technology Commission","doi-asserted-by":"publisher","award":["24JCZDJC01400"],"award-info":[{"award-number":["24JCZDJC01400"]}],"id":[{"id":"10.13039\/501100010229","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>A fundamental question in immunology is how the adaptive immune system encodes antigen specificity while maintaining repertoire diversity. B cell receptor (BCR) or antibody clonal families, defined by groups of B cells descending from a common ancestor, are key to deciphering this encoding. Although paired heavy and light chains jointly determine antibody specificity, most repertoire analyses have historically relied on heavy-chain-only data due to the loss of native pairing information in bulk BCR sequencing. This reliance introduces potential biases in computational clonal cluster inference, which may complicate efforts to resolve disease-associated immune signatures. Here, we leverage large-scale paired-chain BCR sequencing data to demonstrate that heavy-chain-based clustering may misrepresent true clonal architecture, and identify two major artifacts: chain-mixed clusters, in which similar heavy chains are paired with distinct light chains, and naive-like pseudo-clonal clusters, which are detected in an individual\u2019s naive B cell repertoire and exhibit highly similar heavy and light chains without reflecting true clonal expansion. To address these limitations, we present fastBCR-p, an optimized framework that integrates light-chain-informed subclustering, with public sequence aware refinement to improve clonal family inference. By resolving both technical artifacts and biological convergence, fastBCR-p improves the chain concordance and overall clustering quality of clonal inference in real-world datasets. This enables more accurate tracking of immune dynamics in health and disease and facilitates the identification of clinically relevant antibody lineages.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1014077","type":"journal-article","created":{"date-parts":[[2026,3,11]],"date-time":"2026-03-11T17:35:45Z","timestamp":1773250545000},"page":"e1014077","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Large-scale paired chain BCR analysis reveals antibody clonal family inference bias and enhances resolution with machine learning"],"prefix":"10.1371","volume":"22","author":[{"given":"Hao","family":"Wang","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kaixuan","family":"Wang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Qihang","family":"Xu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Linru","family":"Cai","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Chuanxiang","family":"Huang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Linlin","family":"Chen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yunliang","family":"Zang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xihao","family":"Hu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4966-2086","authenticated-orcid":true,"given":"Jian","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2026,3,11]]},"reference":[{"key":"pcbi.1014077.ref001","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1016\/0167-5699(92)90043-7","article-title":"VDJ recombination","volume":"13","author":"FW Alt","year":"1992","journal-title":"Immunology Today"},{"issue":"13","key":"pcbi.1014077.ref002","doi-asserted-by":"crossref","first-page":"4932","DOI":"10.1073\/pnas.87.13.4932","article-title":"Stochastic pairing of heavy-chain and kappa light-chain variable gene families occurs in polyclonally activated B cells","volume":"87","author":"A Kaushik","year":"1990","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"4","key":"pcbi.1014077.ref003","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1038\/s41435-024-00279-2","article-title":"Resolving haplotype variation and complex genetic architecture in the human immunoglobulin kappa chain locus in individuals of diverse ancestry","volume":"25","author":"E Engelbrecht","year":"2024","journal-title":"Genes Immun"},{"issue":"1","key":"pcbi.1014077.ref004","doi-asserted-by":"crossref","first-page":"4419","DOI":"10.1038\/s41467-023-40070-x","article-title":"Genetic variation in the immunoglobulin heavy chain locus shapes the human antibody repertoire","volume":"14","author":"OL Rodriguez","year":"2023","journal-title":"Nat Commun"},{"issue":"1","key":"pcbi.1014077.ref005","doi-asserted-by":"crossref","first-page":"1729683","DOI":"10.1080\/19420862.2020.1729683","article-title":"Understanding the human antibody repertoire","volume":"12","author":"AR Rees","year":"2020","journal-title":"MAbs"},{"issue":"7744","key":"pcbi.1014077.ref006","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1038\/s41586-019-0879-y","article-title":"Commonality despite exceptional diversity in the baseline human antibody repertoire","volume":"566","author":"B Briney","year":"2019","journal-title":"Nature"},{"key":"pcbi.1014077.ref007","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1146\/annurev.biochem.76.061705.090740","article-title":"Molecular mechanisms of antibody somatic hypermutation","volume":"76","author":"JM Di Noia","year":"2007","journal-title":"Annu Rev Biochem"},{"key":"pcbi.1014077.ref008","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.coi.2014.02.010","article-title":"Clonal and cellular dynamics in germinal centers","volume":"28","author":"GD Victora","year":"2014","journal-title":"Curr Opin Immunol"},{"issue":"11","key":"pcbi.1014077.ref009","doi-asserted-by":"crossref","first-page":"101267","DOI":"10.1016\/j.xcrm.2023.101267","article-title":"SARS-CoV-2 antibodies from children exhibit broad neutralization and belong to adult public clonotypes","volume":"4","author":"SC Wall","year":"2023","journal-title":"Cell Rep Med"},{"issue":"28","key":"pcbi.1014077.ref010","doi-asserted-by":"crossref","first-page":"10275","DOI":"10.1073\/pnas.1409954111","article-title":"Affinity maturation in an HIV broadly neutralizing B-cell lineage through reorientation of variable domains","volume":"111","author":"D Fera","year":"2014","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"8","key":"pcbi.1014077.ref011","doi-asserted-by":"crossref","DOI":"10.1016\/j.immuni.2021.07.008","article-title":"Affinity maturation of SARS-CoV-2 neutralizing antibodies confers potency, breadth, and resilience to viral escape mutations","volume":"54","author":"F Muecksch","year":"2021","journal-title":"Immunity"},{"issue":"11","key":"pcbi.1014077.ref012","doi-asserted-by":"crossref","first-page":"2002","DOI":"10.1016\/j.bbapap.2014.07.006","article-title":"Antibody informatics for drug discovery","volume":"1844","author":"H Shirai","year":"2014","journal-title":"Biochim Biophys Acta"},{"issue":"5","key":"pcbi.1014077.ref013","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1038\/nrd.2016.268","article-title":"Strategies and challenges for the next generation of antibody-drug conjugates","volume":"16","author":"A Beck","year":"2017","journal-title":"Nat Rev Drug Discov"},{"issue":"4","key":"pcbi.1014077.ref014","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1016\/j.jaut.2010.07.004","article-title":"Somatic hypermutation and antigen-driven selection of B cells are altered in autoimmune diseases","volume":"35","author":"NS Zuckerman","year":"2010","journal-title":"J Autoimmun"},{"issue":"7776","key":"pcbi.1014077.ref015","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1038\/s41586-019-1595-3","article-title":"Analysis of the B cell receptor repertoire in six immune-mediated diseases","volume":"574","author":"RJM Bashford-Rogers","year":"2019","journal-title":"Nature"},{"issue":"7","key":"pcbi.1014077.ref016","doi-asserted-by":"crossref","first-page":"112722","DOI":"10.1016\/j.celrep.2023.112722","article-title":"Dietary restriction mitigates the age-associated decline in mouse B cell receptor repertoire diversity","volume":"42","author":"C Monz\u00f3","year":"2023","journal-title":"Cell Rep"},{"issue":"6695","key":"pcbi.1014077.ref017","article-title":"A blueprint for tumor-infiltrating B cells across human cancers","volume":"384","author":"J Ma","year":"2024","journal-title":"Science"},{"key":"pcbi.1014077.ref018","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.92718","article-title":"The physiological landscape and specificity of antibody repertoires are consolidated by multiple immunizations","volume":"13","author":"L Csepregi","year":"2024","journal-title":"Elife"},{"key":"pcbi.1014077.ref019","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1016\/j.coi.2016.06.001","article-title":"Antibody-guided vaccine design: identification of protective epitopes","volume":"41","author":"A Lanzavecchia","year":"2016","journal-title":"Curr Opin Immunol"},{"issue":"7864","key":"pcbi.1014077.ref020","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1038\/s41586-021-03594-0","article-title":"Neutralizing antibody vaccine for pandemic and pre-emergent coronaviruses","volume":"594","author":"KO Saunders","year":"2021","journal-title":"Nature"},{"key":"pcbi.1014077.ref021","doi-asserted-by":"crossref","first-page":"118","DOI":"10.3389\/fimmu.2018.00118","article-title":"Next-Generation Sequencing of Antibody Display Repertoires","volume":"9","author":"R Rouet","year":"2018","journal-title":"Front Immunol"},{"issue":"12","key":"pcbi.1014077.ref022","article-title":"Evaluating predictive patterns of antigen-specific B cells by single-cell transcriptome and antibody repertoire sequencing","volume":"15","author":"L Erlach","year":"2024","journal-title":"Cell Syst"},{"issue":"10","key":"pcbi.1014077.ref023","doi-asserted-by":"crossref","first-page":"885","DOI":"10.1038\/icb.2015.57","article-title":"BCR repertoire sequencing: different patterns of B-cell activation after two Meningococcal vaccines","volume":"93","author":"JD Galson","year":"2015","journal-title":"Immunol Cell Biol"},{"key":"pcbi.1014077.ref024","doi-asserted-by":"crossref","unstructured":"Angermueller C, Mariet Z, Jester B, Engelhart E, Emerson R, Alipanahi B. High-throughput ML-guided design of diverse single-domain antibodies against SARS-CoV-2. 2023. https:\/\/doi.org\/10.1101\/2023.12.01.569227","DOI":"10.1101\/2023.12.01.569227"},{"issue":"9","key":"pcbi.1014077.ref025","doi-asserted-by":"crossref","first-page":"540","DOI":"10.5483\/BMBRep.2019.52.9.192","article-title":"Deep sequencing of B cell receptor repertoire","volume":"52","author":"D Kim","year":"2019","journal-title":"BMB Rep"},{"issue":"10","key":"pcbi.1014077.ref026","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1005086","article-title":"Likelihood-Based Inference of B Cell Clonal Families","volume":"12","author":"DK Ralph","year":"2016","journal-title":"PLoS Comput Biol"},{"issue":"8","key":"pcbi.1014077.ref027","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1010411","article-title":"A multi-objective based clustering for inferring BCR clonal lineages from high-throughput B cell repertoire data","volume":"18","author":"N Abdollahi","year":"2022","journal-title":"PLoS Comput Biol"},{"issue":"13","key":"pcbi.1014077.ref028","doi-asserted-by":"crossref","DOI":"10.1093\/bioinformatics\/bty235","article-title":"A spectral clustering-based method for identifying clones from high-throughput B cell repertoire sequencing data","volume":"34","author":"N Nouri","year":"2018","journal-title":"Bioinformatics"},{"key":"pcbi.1014077.ref029","doi-asserted-by":"crossref","first-page":"2489","DOI":"10.4049\/jimmunol.1601850","article-title":"Hierarchical Clustering Can Identify B Cell Clones with High Confidence in Ig Repertoire Sequencing Data","volume":"198","author":"NT Gupta","year":"2017","journal-title":"The Journal of Immunology"},{"key":"pcbi.1014077.ref030","doi-asserted-by":"crossref","first-page":"2249","DOI":"10.3389\/fimmu.2018.02249","article-title":"Immunoglobulin Light Chain Gene Rearrangements, Receptor Editing and the Development of a Self-Tolerant Antibody Repertoire","volume":"9","author":"AM Collins","year":"2018","journal-title":"Front Immunol"},{"issue":"7935","key":"pcbi.1014077.ref031","doi-asserted-by":"crossref","first-page":"352","DOI":"10.1038\/s41586-022-05371-z","article-title":"Functional antibodies exhibit light chain coherence","volume":"611","author":"DB Jaffe","year":"2022","journal-title":"Nature"},{"issue":"4","key":"pcbi.1014077.ref032","article-title":"Human B Cell Clonal Expansion and Convergent Antibody Responses to SARS-CoV-2","volume":"28","author":"SCA Nielsen","year":"2020","journal-title":"Cell Host Microbe"},{"issue":"1676","key":"pcbi.1014077.ref033","doi-asserted-by":"crossref","first-page":"20140239","DOI":"10.1098\/rstb.2014.0239","article-title":"The analysis of clonal expansions in normal and autoimmune B cell repertoires","volume":"370","author":"U Hershberg","year":"2015","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"issue":"7","key":"pcbi.1014077.ref034","doi-asserted-by":"crossref","first-page":"1467","DOI":"10.1016\/j.celrep.2017.04.054","article-title":"Systems Analysis Reveals High Genetic and Antigen-Driven Predetermination of Antibody Repertoires throughout B Cell Development","volume":"19","author":"V Greiff","year":"2017","journal-title":"Cell Rep"},{"issue":"1","key":"pcbi.1014077.ref035","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1038\/s41467-017-00645-x","article-title":"Accurate immune repertoire sequencing reveals malaria infection driven antibody lineage diversification in young children","volume":"8","author":"BS Wendel","year":"2017","journal-title":"Nat Commun"},{"issue":"6","key":"pcbi.1014077.ref036","doi-asserted-by":"crossref","DOI":"10.1016\/j.chom.2018.05.001","article-title":"Multi-Donor Longitudinal Antibody Repertoire Sequencing Reveals the Existence of Public Antibody Clonotypes in HIV-1 Infection","volume":"23","author":"I Setliff","year":"2018","journal-title":"Cell Host Microbe"},{"issue":"7744","key":"pcbi.1014077.ref037","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1038\/s41586-019-0934-8","article-title":"High frequency of shared clonotypes in human B cell receptor repertoires","volume":"566","author":"C Soto","year":"2019","journal-title":"Nature"},{"issue":"1","key":"pcbi.1014077.ref038","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1002\/pro.4205","article-title":"Observed Antibody Space: A diverse database of cleaned, annotated, and translated unpaired and paired antibody sequences","volume":"31","author":"TH Olsen","year":"2022","journal-title":"Protein Sci"},{"issue":"2","key":"pcbi.1014077.ref039","doi-asserted-by":"crossref","first-page":"102969","DOI":"10.1016\/j.xpro.2024.102969","article-title":"Protocol for fast clonal family inference and analysis from large-scale B cell receptor repertoire sequencing data","volume":"5","author":"K Wang","year":"2024","journal-title":"STAR Protoc"},{"issue":"10","key":"pcbi.1014077.ref040","doi-asserted-by":"crossref","first-page":"100601","DOI":"10.1016\/j.crmeth.2023.100601","article-title":"Fast clonal family inference from large-scale B cell repertoire sequencing data","volume":"3","author":"K Wang","year":"2023","journal-title":"Cell Rep Methods"},{"issue":"2","key":"pcbi.1014077.ref041","doi-asserted-by":"crossref","first-page":"378","DOI":"10.32614\/RJ-2017-066","article-title":"glmmTMB Balances Speed and Flexibility Among Packages for Zero-inflated Generalized Linear Mixed Modeling","volume":"9","year":"2017","journal-title":"The R Journal"},{"key":"pcbi.1014077.ref042","article-title":"BERT: Pre-training of Deep Bidirectional Transformers for Language Understanding","author":"J Devlin","year":"2018"},{"issue":"7","key":"pcbi.1014077.ref043","doi-asserted-by":"crossref","first-page":"100513","DOI":"10.1016\/j.patter.2022.100513","article-title":"Deciphering the language of antibodies using self-supervised learning","volume":"3","author":"J Leem","year":"2022","journal-title":"Patterns (N Y)"},{"issue":"17","key":"pcbi.1014077.ref044","doi-asserted-by":"crossref","first-page":"2974","DOI":"10.1093\/bioinformatics\/btz035","article-title":"OLGA: fast computation of generation probabilities of B- and T-cell receptor amino acid sequences and motifs","volume":"35","author":"Z Sethna","year":"2019","journal-title":"Bioinformatics"},{"issue":"1","key":"pcbi.1014077.ref045","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1038\/s41467-018-02832-w","article-title":"High-throughput immune repertoire analysis with IGoR","volume":"9","author":"Q Marcou","year":"2018","journal-title":"Nat Commun"},{"issue":"10","key":"pcbi.1014077.ref046","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1005086","article-title":"Likelihood-Based Inference of B Cell Clonal Families","volume":"12","author":"DK Ralph","year":"2016","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1014077.ref047","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.86181","article-title":"Combining mutation and recombination statistics to infer clonal families in antibody repertoires","volume":"13","author":"N Spisak","year":"2024","journal-title":"Elife"},{"issue":"3","key":"pcbi.1014077.ref048","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1038\/nri2508","article-title":"B-cell responses to vaccination at the extremes of age","volume":"9","author":"C-A Siegrist","year":"2009","journal-title":"Nat Rev Immunol"},{"issue":"1","key":"pcbi.1014077.ref049","doi-asserted-by":"crossref","first-page":"1996732","DOI":"10.1080\/19420862.2021.1996732","article-title":"Current strategies for detecting functional convergence across B-cell receptor repertoires","volume":"13","author":"MIJ Raybould","year":"2021","journal-title":"MAbs"},{"key":"pcbi.1014077.ref050","doi-asserted-by":"crossref","unstructured":"Friedensohn S, Neumeier D, Khan TA, Csepregi L, Parola C, De Vries ARG, et al. Convergent selection in antibody repertoires is revealed by deep learning. 2020. https:\/\/doi.org\/10.1101\/2020.02.25.965673","DOI":"10.1101\/2020.02.25.965673"},{"issue":"1","key":"pcbi.1014077.ref051","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1186\/s12865-024-00600-8","article-title":"Systematic evaluation of B-cell clonal family inference approaches","volume":"25","author":"D Balashova","year":"2024","journal-title":"BMC Immunol"},{"issue":"11","key":"pcbi.1014077.ref052","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1010723","article-title":"Inference of B cell clonal families using heavy\/light chain pairing information","volume":"18","author":"DK Ralph","year":"2022","journal-title":"PLoS Comput Biol"},{"issue":"4","key":"pcbi.1014077.ref053","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1093\/intimm\/dxh214","article-title":"Stochastic pairing of Ig heavy and light chains frequently generates B cell antigen receptors that are subject to editing in vivo","volume":"17","author":"T Novobrantseva","year":"2005","journal-title":"Int Immunol"},{"issue":"1","key":"pcbi.1014077.ref054","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1016\/j.smim.2005.11.001","article-title":"Pre-BCR signals and the control of Ig gene rearrangements","volume":"18","author":"JK Geier","year":"2006","journal-title":"Semin Immunol"},{"issue":"3","key":"pcbi.1014077.ref055","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1016\/j.bbrc.2019.05.149","article-title":"Artificial immunoglobulin light chain with potential to associate with a wide variety of immunoglobulin heavy chains","volume":"515","author":"H Xue","year":"2019","journal-title":"Biochem Biophys Res Commun"},{"key":"pcbi.1014077.ref056","article-title":"IgBLAST: an immunoglobulin variable domain sequence analysis tool","volume":"41","author":"J Ye","year":"2013","journal-title":"Nucleic Acids Res"},{"key":"pcbi.1014077.ref057","article-title":"IMGT\u00ae, the international ImMunoGeneTics information system\u00ae 25 years on","volume":"43","author":"M-P Lefranc","year":"2015","journal-title":"Nucleic Acids Res"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1014077","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T00:00:00Z","timestamp":1773792000000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014077","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T17:48:01Z","timestamp":1773856081000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014077"}},"subtitle":[],"editor":[{"given":"Brittany","family":"Rife Magalis","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2026,3,11]]},"references-count":57,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2026,3,11]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1014077","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,11]]}}}