{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T18:51:37Z","timestamp":1791399097454,"version":"4.3.3"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1014829","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T00:00:00Z","timestamp":1791331200000}}],"reference-count":52,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T00:00:00Z","timestamp":1790726400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100018418","name":"Graduate School of Arts and Sciences, Harvard University","doi-asserted-by":"publisher","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}],"id":[{"id":"10.13039\/100018418","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["CNS-2107078"],"award-info":[{"award-number":["CNS-2107078"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>\n                    Evolutionary graph theory (EGT) studies the effect of population structure on evolutionary dynamics. The vertices of the graph represent the\n                    <jats:italic>N<\/jats:italic>\n                    individuals. The edges denote interactions for competitive replacement. Two standard update rules are death-Birth (dB) and Birth-death (Bd). Under dB, an individual is chosen uniformly at random to die, and its neighbors\u2014the individuals on adjacent vertices of the graph\u2014 compete to fill the vacancy proportional to their fitness. Under Bd, an individual is chosen for reproduction proportional to fitness, and its offspring replaces a randomly chosen neighbor on an adjacent vertex. Here we study mixed updating between those two scenarios. In each time step, with probability\n                    <jats:inline-formula id=\"pcbi.1014829.e001\">\n                      <jats:alternatives>\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" id=\"pcbi.1014829.e001g\" mimetype=\"image\" position=\"anchor\" xlink:href=\"info:doi\/10.1371\/journal.pcbi.1014829.e001\" xlink:type=\"simple\"\/>\n                        <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" id=\"M1\">\n                          <mml:mrow>\n                            <mml:mi>\u03b4<\/mml:mi>\n                          <\/mml:mrow>\n                        <\/mml:math>\n                      <\/jats:alternatives>\n                    <\/jats:inline-formula>\n                    the update is dB and with the remaining probability it is Bd. We study fixation probabilities and times as functions of\n                    <jats:inline-formula id=\"pcbi.1014829.e002\">\n                      <jats:alternatives>\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" id=\"pcbi.1014829.e002g\" mimetype=\"image\" position=\"anchor\" xlink:href=\"info:doi\/10.1371\/journal.pcbi.1014829.e002\" xlink:type=\"simple\"\/>\n                        <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" id=\"M2\">\n                          <mml:mrow>\n                            <mml:mi>\u03b4<\/mml:mi>\n                          <\/mml:mrow>\n                        <\/mml:math>\n                      <\/jats:alternatives>\n                    <\/jats:inline-formula>\n                    under neutral evolution and constant selection. Despite the fact that fixation probabilities and times can be increasing, decreasing, or non-monotonic in\n                    <jats:inline-formula id=\"pcbi.1014829.e003\">\n                      <jats:alternatives>\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" id=\"pcbi.1014829.e003g\" mimetype=\"image\" position=\"anchor\" xlink:href=\"info:doi\/10.1371\/journal.pcbi.1014829.e003\" xlink:type=\"simple\"\/>\n                        <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" id=\"M3\">\n                          <mml:mrow>\n                            <mml:mi>\u03b4<\/mml:mi>\n                          <\/mml:mrow>\n                        <\/mml:math>\n                      <\/jats:alternatives>\n                    <\/jats:inline-formula>\n                    , we prove that nearly all unweighted undirected graphs have short fixation times and provide an efficient algorithm to estimate their fixation probabilities. Finally, we prove exact formulas for fixation probabilities on cycles, stars, and more complex structures and classify their sensitivities to\n                    <jats:inline-formula id=\"pcbi.1014829.e004\">\n                      <jats:alternatives>\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" id=\"pcbi.1014829.e004g\" mimetype=\"image\" position=\"anchor\" xlink:href=\"info:doi\/10.1371\/journal.pcbi.1014829.e004\" xlink:type=\"simple\"\/>\n                        <mml:math xmlns:mml=\"http:\/\/www.w3.org\/1998\/Math\/MathML\" display=\"inline\" id=\"M4\">\n                          <mml:mrow>\n                            <mml:mi>\u03b4<\/mml:mi>\n                          <\/mml:mrow>\n                        <\/mml:math>\n                      <\/jats:alternatives>\n                    <\/jats:inline-formula>\n                    .\n                  <\/jats:p>","DOI":"10.1371\/journal.pcbi.1014829","type":"journal-article","created":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T17:49:03Z","timestamp":1790790543000},"page":"e1014829","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Mixed updating in structured populations"],"prefix":"10.1371","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9776-4343","authenticated-orcid":true,"given":"David A.","family":"Brewster","sequence":"first","affiliation":[{"name":"Harvard University","place":["Boston, Massachusetts, United States of America"],"department":["John A. Paulson School of Engineering and Applied Sciences"]},{"name":"Harvard University","place":["Cambridge, Massachusetts, United States of America"],"department":["Department of Molecular and Cellular Biology"]},{"name":"Harvard University","place":["Cambridge, Massachusetts, United States of America"],"department":["Department of Organismic and Evolutionary Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yichen","family":"Huang","sequence":"additional","affiliation":[{"name":"Harvard University","place":["Boston, Massachusetts, United States of America"],"department":["John A. Paulson School of Engineering and Applied Sciences"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Mitzenmacher","sequence":"additional","affiliation":[{"name":"Harvard University","place":["Boston, Massachusetts, United States of America"],"department":["John A. Paulson School of Engineering and Applied Sciences"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin A.","family":"Nowak","sequence":"additional","affiliation":[{"name":"Harvard University","place":["Cambridge, Massachusetts, United States of America"],"department":["Department of Organismic and Evolutionary Biology"]},{"name":"Harvard University","place":["Cambridge, Massachusetts, United States of America"],"department":["Department of Mathematics"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,9,30]]},"reference":[{"issue":"1","key":"pcbi.1014829.ref001","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1007\/s00453-012-9722-7","article-title":"Approximating Fixation Probabilities in the Generalized Moran Process","volume":"69","author":"J D\u00edaz","year":"2014","journal-title":"Algorithmica"},{"issue":"215","key":"pcbi.1014829.ref002","doi-asserted-by":"crossref","first-page":"20230594","DOI":"10.1098\/rsif.2023.0594","article-title":"The speed of neutral evolution on graphs","volume":"21","author":"S Gao","year":"2024","journal-title":"J R Soc Interface"},{"key":"pcbi.1014829.ref003","article-title":"Maintaining diversity in structured populations","author":"DA Brewster","year":"2025","journal-title":"PNAS Nexus"},{"issue":"2098","key":"pcbi.1014829.ref004","doi-asserted-by":"crossref","first-page":"2609","DOI":"10.1098\/rspa.2008.0058","article-title":"An analysis of the fixation probability of a mutant on special classes of non-directed graphs","volume":"464","author":"M Broom","year":"2008","journal-title":"Proc R Soc A"},{"issue":"2156","key":"pcbi.1014829.ref005","doi-asserted-by":"crossref","first-page":"20130193","DOI":"10.1098\/rspa.2013.0193","article-title":"On the fixation probability of superstars","volume":"469","author":"J D\u00edaz","year":"2013","journal-title":"Proc R Soc A"},{"issue":"11","key":"pcbi.1014829.ref006","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1004437","article-title":"Most Undirected Random Graphs Are Amplifiers of Selection for Birth-Death Dynamics, but Suppressors of Selection for Death-Birth Dynamics","volume":"11","author":"L Hindersin","year":"2015","journal-title":"PLoS Comput Biol"},{"issue":"2181","key":"pcbi.1014829.ref007","doi-asserted-by":"crossref","first-page":"20150114","DOI":"10.1098\/rspa.2015.0114","article-title":"Amplifiers of selection","volume":"471","author":"B Adlam","year":"2015","journal-title":"Proceedings of the Royal Society A: Mathematical, Physical and Engineering Sciences"},{"issue":"1","key":"pcbi.1014829.ref008","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1145\/3019609","article-title":"Amplifiers for the Moran Process","volume":"64","author":"A Galanis","year":"2017","journal-title":"J ACM"},{"issue":"1","key":"pcbi.1014829.ref009","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1038\/s41598-017-00107-w","article-title":"Amplification on Undirected Population Structures: Comets Beat Stars","volume":"7","author":"A Pavlogiannis","year":"2017","journal-title":"Sci Rep"},{"key":"pcbi.1014829.ref010","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1038\/s42003-018-0078-7","article-title":"Construction of arbitrarily strong amplifiers of natural selection using evolutionary graph theory","volume":"1","author":"A Pavlogiannis","year":"2018","journal-title":"Commun Biol"},{"key":"pcbi.1014829.ref011","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1038\/s42003-019-0374-x","article-title":"Exploring and mapping the universe of evolutionary graphs identifies structural properties affecting fixation probability and time","volume":"2","author":"M M\u00f6ller","year":"2019","journal-title":"Commun Biol"},{"key":"pcbi.1014829.ref012","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.tcs.2018.08.005","article-title":"Asymptotically optimal amplifiers for the Moran process","volume":"758","author":"LA Goldberg","year":"2019","journal-title":"Theoretical Computer Science"},{"issue":"1","key":"pcbi.1014829.ref013","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1007494","article-title":"Limits on amplifiers of natural selection under death-Birth updating","volume":"16","author":"J Tkadlec","year":"2020","journal-title":"PLoS Comput Biol"},{"issue":"2","key":"pcbi.1014829.ref014","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1008695","article-title":"Fixation probabilities in graph-structured populations under weak selection","volume":"17","author":"B Allen","year":"2021","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1014829.ref015","article-title":"Constant-selection evolutionary dynamics on weighted networks","author":"J Bhaumik","year":"2024","journal-title":"arXiv preprint arXiv:2403.17208"},{"issue":"2","key":"pcbi.1014829.ref016","article-title":"Evolutionary graph theory beyond single mutation dynamics: on how network-structured populations cross fitness landscapes","volume":"227","author":"YP Kuo","year":"2024","journal-title":"Genetics"},{"issue":"1","key":"pcbi.1014829.ref017","doi-asserted-by":"crossref","first-page":"4666","DOI":"10.1038\/s41467-024-48617-2","article-title":"A theory of evolutionary dynamics on any complex population structure reveals stem cell niche architecture as a spatial suppressor of selection","volume":"15","author":"YP Kuo","year":"2024","journal-title":"Nat Commun"},{"issue":"7","key":"pcbi.1014829.ref018","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1012299","article-title":"Fixation times on directed graphs","volume":"20","author":"DA Brewster","year":"2024","journal-title":"PLOS Computational Biology"},{"issue":"11","key":"pcbi.1014829.ref019","doi-asserted-by":"crossref","first-page":"2667","DOI":"10.1111\/2041-210X.70121","article-title":"Ecological graph theory: Simulating competition and coexistence on graphs","volume":"16","author":"A Kalirad","year":"2025","journal-title":"Methods Ecol Evol"},{"issue":"3","key":"pcbi.1014829.ref020","doi-asserted-by":"crossref","DOI":"10.1002\/rsa.70003","article-title":"The Moran Process on a Random Graph","volume":"66","author":"A Frieze","year":"2025","journal-title":"Random Struct Algorithms"},{"issue":"1598","key":"pcbi.1014829.ref021","doi-asserted-by":"crossref","first-page":"2249","DOI":"10.1098\/rspb.2006.3576","article-title":"Evolutionary games on cycles","volume":"273","author":"H Ohtsuki","year":"2006","journal-title":"Proceedings of the Royal Society B: Biological Sciences"},{"issue":"10","key":"pcbi.1014829.ref022","doi-asserted-by":"crossref","first-page":"108103","DOI":"10.1103\/PhysRevLett.98.108103","article-title":"Dynamical organization of cooperation in complex topologies","volume":"98","author":"J G\u00f3mez-Garde\u00f1es","year":"2007","journal-title":"Phys Rev Lett"},{"issue":"2117","key":"pcbi.1014829.ref023","doi-asserted-by":"crossref","first-page":"1327","DOI":"10.1098\/rspa.2009.0487","article-title":"Evolutionary games on graphs and the speed of the evolutionary process","volume":"466","author":"M Broom","year":"2010","journal-title":"Proceedings of the Royal Society A: Mathematical, Physical and Engineering Sciences"},{"issue":"03","key":"pcbi.1014829.ref024","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1142\/S0218127410026137","article-title":"Cooperation in the prisoner\u2019s dilemma game in random scale-free graphs","volume":"20","author":"J Poncela","year":"2010","journal-title":"International Journal of Bifurcation and Chaos"},{"issue":"3","key":"pcbi.1014829.ref025","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1007\/s13235-011-0022-7","article-title":"Evolutionary Games on Star Graphs Under Various Updating Rules","volume":"1","author":"C Hadjichrysanthou","year":"2011","journal-title":"Dyn Games Appl"},{"issue":"1","key":"pcbi.1014829.ref026","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0054639","article-title":"Consolidating birth-death and death-birth processes in structured populations","volume":"8","author":"J Zukewich","year":"2013","journal-title":"PLoS One"},{"key":"pcbi.1014829.ref027","doi-asserted-by":"crossref","first-page":"3055","DOI":"10.1038\/srep03055","article-title":"Impact of social punishment on cooperative behavior in complex networks","volume":"3","author":"Z Wang","year":"2013","journal-title":"Sci Rep"},{"issue":"7649","key":"pcbi.1014829.ref028","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1038\/nature21723","article-title":"Evolutionary dynamics on any population structure","volume":"544","author":"B Allen","year":"2017","journal-title":"Nature"},{"key":"pcbi.1014829.ref029","doi-asserted-by":"crossref","first-page":"100347","DOI":"10.1016\/j.cosrev.2020.100347","article-title":"A survey of the modified Moran process and evolutionary graph theory","volume":"39","author":"J D\u00edaz","year":"2021","journal-title":"Computer Science Review"},{"issue":"4","key":"pcbi.1014829.ref030","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1038\/s43588-024-00621-8","article-title":"Strategy evolution on higher-order networks","volume":"4","author":"A Sheng","year":"2024","journal-title":"Nature Computational Science"},{"key":"pcbi.1014829.ref031","doi-asserted-by":"crossref","first-page":"024307","DOI":"10.1103\/PhysRevE.109.024307","article-title":"Evolution of cooperation in deme-structured populations on graphs","volume":"109","author":"A Moawad","year":"2024","journal-title":"Phys Rev E"},{"key":"pcbi.1014829.ref032","doi-asserted-by":"crossref","DOI":"10.2307\/j.ctvjghw98","volume-title":"Evolutionary Dynamics: Exploring the Equations of Life","author":"A Nowak","year":"2006"},{"issue":"25","key":"pcbi.1014829.ref033","doi-asserted-by":"crossref","first-page":"14966","DOI":"10.1073\/pnas.2535419100","article-title":"The linear process of somatic evolution","volume":"100","author":"MA Nowak","year":"2003","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"7023","key":"pcbi.1014829.ref034","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1038\/nature03204","article-title":"Evolutionary dynamics on graphs","volume":"433","author":"E Lieberman","year":"2005","journal-title":"Nature"},{"issue":"1","key":"pcbi.1014829.ref035","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1007529","article-title":"Transient amplifiers of selection and reducers of fixation for death-Birth updating on graphs","volume":"16","author":"B Allen","year":"2020","journal-title":"PLoS Comput Biol"},{"issue":"37","key":"pcbi.1014829.ref036","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2205424119","article-title":"Suppressors of fixation can increase average fitness beyond amplifiers of selection","volume":"119","author":"N Sharma","year":"2022","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"5","key":"pcbi.1014829.ref037","doi-asserted-by":"crossref","first-page":"220011","DOI":"10.1098\/rsos.220011","article-title":"Martingales and the fixation time of evolutionary graphs with arbitrary dimensionality","volume":"9","author":"T Monk","year":"2022","journal-title":"R Soc Open Sci"},{"issue":"50","key":"pcbi.1014829.ref038","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2405605121","article-title":"Density amplifiers of cooperation for spatial games","volume":"121","author":"J Svoboda","year":"2024","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"pcbi.1014829.ref039","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1038\/s42003-019-0373-y","article-title":"Population structure determines the tradeoff between fixation probability and fixation time","volume":"2","author":"J Tkadlec","year":"2019","journal-title":"Communications biology"},{"key":"pcbi.1014829.ref040","article-title":"Population structure amplifies mutation load","author":"N Sharma","year":"2025","journal-title":"bioRxiv"},{"issue":"1","key":"pcbi.1014829.ref041","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1017\/S0305004100033193","article-title":"Random processes in genetics","volume":"54","author":"PAP Moran","year":"1958","journal-title":"Math Proc Camb Phil Soc"},{"issue":"4","key":"pcbi.1014829.ref042","doi-asserted-by":"crossref","first-page":"140465","DOI":"10.1098\/rsos.140465","article-title":"The duality of spatial death-birth and birth-death processes and limitations of the isothermal theorem","volume":"2","author":"K Kaveh","year":"2015","journal-title":"R Soc Open Sci"},{"issue":"200","key":"pcbi.1014829.ref043","doi-asserted-by":"crossref","first-page":"20220769","DOI":"10.1098\/rsif.2022.0769","article-title":"Categorizing update mechanisms for graph-structured metapopulations","volume":"20","author":"S Yagoobi","year":"2023","journal-title":"J R Soc Interface"},{"issue":"3","key":"pcbi.1014829.ref044","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1012008","article-title":"Amplifiers of selection for the Moran process with both Birth-death and death-Birth updating","volume":"20","author":"J Svoboda","year":"2024","journal-title":"PLoS Comput Biol"},{"issue":"7092","key":"pcbi.1014829.ref045","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1038\/nature04605","article-title":"A simple rule for the evolution of cooperation on graphs and social networks","volume":"441","author":"H Ohtsuki","year":"2006","journal-title":"Nature"},{"key":"pcbi.1014829.ref046","article-title":"Understanding the rift between update rules in evolutionary graph theory: The intrinsic death rate drives star graphs from amplifying to suppressing natural selection","author":"M Dew","year":"2025","journal-title":"arXiv preprint"},{"issue":"2121","key":"pcbi.1014829.ref047","doi-asserted-by":"crossref","first-page":"2795","DOI":"10.1098\/rspa.2010.0067","article-title":"Two results on evolutionary processes on general non-directed graphs","volume":"466","author":"M Broom","year":"2010","journal-title":"Proc R Soc A"},{"key":"pcbi.1014829.ref048","unstructured":"Cooper C, Rivera N. The linear voting model. In: 43rd International Colloquium on Automata, Languages, and Programming (ICALP 2016), 2016."},{"issue":"1","key":"pcbi.1014829.ref049","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1002\/rsa.20617","article-title":"Absorption time of the Moran process","volume":"49","author":"J D\u00edaz","year":"2016","journal-title":"Random Struct Alg"},{"issue":"3","key":"pcbi.1014829.ref050","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1002\/rsa.20890","article-title":"Phase transitions of the Moran process and algorithmic consequences","volume":"56","author":"L Ann Goldberg","year":"2019","journal-title":"Random Struct Algorithms"},{"key":"pcbi.1014829.ref051","unstructured":"Chatterjee K, Ibsen-Jensen R, Nowak MA. Faster Monte-Carlo Algorithms for Fixation Probability of the Moran Process on Undirected Graphs. In: 42nd International Symposium on Mathematical Foundations of Computer Science (MFCS 2017), Dagstuhl, Germany, 2017. 61:1-61:13. https:\/\/doi.org\/10.4230\/LIPIcs.MFCS.2017.61"},{"key":"pcbi.1014829.ref052","unstructured":"Brewster DA, Huang Y, Mitzenmacher M, Nowak MA. The mixed birth-death\/death-birth Moran process. 2025. https:\/\/doi.org\/arXiv:2511.18252"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1014829","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T00:00:00Z","timestamp":1791331200000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014829","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T18:06:19Z","timestamp":1791396379000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014829"}},"subtitle":[],"editor":[{"given":"Jordan","family":"Douglas","sequence":"first","affiliation":[{"name":"University of Auckland","place":["NEW ZEALAND"]}],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,9,30]]},"references-count":52,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9,30]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1014829","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,30]]}}}