The key role of cheaters in the persistence of cooperation

The key role of cheaters in the persistence of cooperation | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article The key role of cheaters in the persistence of cooperation Sanasar Babajanyan, Yuri Wolf, Eugene Koonin, Nash Rochman This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6280939/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 13 You are reading this latest preprint version Abstract Evolution of cooperation is a major, extensively studied problem in evolutionary biology. Cooperation is beneficial for a population as a whole but costly for the bearers of social traits such that cheaters enjoy a selective advantage over cooperators. Here we focus on coevolution of cooperators and cheaters in a multi-level selection framework, by modeling competition among groups composed of cooperators and cheaters. Cheaters enjoy a reproductive advantage over cooperators at the individual level, independent of the presence of cooperators in the group. Cooperators carry a social trait that provides a fitness advantage to the respective groups. In the case of absolute fitness advantage, where the survival probability of a group is independent of the composition of other groups, the survival of cooperators does not correlate with the presence of cheaters. By contrast, in the case of relative fitness advantage, where the survival probability of a group depends on the composition of all groups, the survival of cooperators positively correlates with the presence of cheaters. Increasing the strength of the social trait alone fails to ensure survival of cooperators, and the increase of the reproduction advantage of the cheaters is necessary to avoid population extinction. This unexpected effect comes from multilevel selection whereby cheaters at the individual level become altruists at the group level, enabling overall growth of the population that is essential for the persistence of cooperators. We validate these theoretical results with an agent-based model of a bacterial biofilm where emergence of the cooperative trait is facilitated by the presence of cheaters, leading to evolution of spatial organization. This finding contrasts the classical view that spatial organization facilitates cooperation. Our results suggest that, counterintuitively, cheaters often promote, not destabilize, evolution of cooperation. Full Text Additional Declarations No competing interests reported. Supplementary Files BMCbioSI.pdf Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 09 Jun, 2025 Reviews received at journal 03 Jun, 2025 Reviews received at journal 30 May, 2025 Reviewers agreed at journal 25 May, 2025 Reviewers agreed at journal 19 May, 2025 Reviewers agreed at journal 19 May, 2025 Reviews received at journal 26 Apr, 2025 Reviewers agreed at journal 03 Apr, 2025 Reviewers agreed at journal 02 Apr, 2025 Reviewers invited by journal 02 Apr, 2025 Editor assigned by journal 24 Mar, 2025 Submission checks completed at journal 24 Mar, 2025 First submitted to journal 21 Mar, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6280939","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":442049418,"identity":"199a061f-e1a7-4947-aae7-05d4ea7f35cf","order_by":0,"name":"Sanasar Babajanyan","email":"","orcid":"","institution":"United States National Library of Medicine","correspondingAuthor":false,"prefix":"","firstName":"Sanasar","middleName":"","lastName":"Babajanyan","suffix":""},{"id":442049419,"identity":"35376ed2-1af0-4356-955f-9d14a0c7517d","order_by":1,"name":"Yuri Wolf","email":"","orcid":"","institution":"United States National Library of Medicine","correspondingAuthor":false,"prefix":"","firstName":"Yuri","middleName":"","lastName":"Wolf","suffix":""},{"id":442049421,"identity":"6af9cc66-5ba5-469c-bee9-8e0e6489429c","order_by":2,"name":"Eugene Koonin","email":"","orcid":"","institution":"United States National Library of Medicine","correspondingAuthor":false,"prefix":"","firstName":"Eugene","middleName":"","lastName":"Koonin","suffix":""},{"id":442049424,"identity":"5650a4eb-d2ae-43f7-893c-a5b13d276b30","order_by":3,"name":"Nash Rochman","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA/ElEQVRIiWNgGAWjYHACNjBpACI+MDAkwIQZG4jRwjiDZC3MPMRoMWc//uwB4x4beXOxw88e2/y5k8fAfsbsMQ+DjeyGA9i1WPbkmBswPEsz3Dk7zdw4t+1ZMQNPjrkxD0OaMS4tBgdy2CQYDhxOMLidYCad23A4sUGCx0yah+FwIk4t558/g2pJ/yZt8Qeu5T9uLTcSzKBacsykGdjgWg7g0fLGTCLhQJrhhts5ZZK9bc8S23jSyiTnGCQbz8TpsPRnEh8O2MgDHbZN4sefO4n97Ie3SbypsJPtw6EFDBIQzAPgaGLiMcCjHA1ATGb8QbyOUTAKRsEoGP4AAIRvXfSPmUMPAAAAAElFTkSuQmCC","orcid":"","institution":"City University of New York","correspondingAuthor":true,"prefix":"","firstName":"Nash","middleName":"","lastName":"Rochman","suffix":""}],"badges":[],"createdAt":"2025-03-22 02:23:05","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6280939/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6280939/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":81181358,"identity":"228690dc-3b9b-4e3d-953c-63f43afe03fd","added_by":"auto","created_at":"2025-04-23 07:29:41","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4033118,"visible":true,"origin":"","legend":"","description":"","filename":"BMCbio.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6280939/v1_covered_3e2b24dd-29ab-4709-9230-57d41f24b9b2.pdf"},{"id":81180651,"identity":"015eadec-2e51-46da-a7d7-a19359961914","added_by":"auto","created_at":"2025-04-23 07:21:30","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":920855,"visible":true,"origin":"","legend":"","description":"","filename":"BMCbioSI.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6280939/v1/8187a890369126a19ea4e919.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"The key role of cheaters in the persistence of cooperation","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":true,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"bmc-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"Learn more about [BMC Biology](https://bmcbiol.biomedcentral.com/)","snPcode":"12915","submissionUrl":"https://submission.springernature.com/new-submission/12915/3","title":"BMC Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-6280939/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6280939/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Evolution of cooperation is a major, extensively studied problem in evolutionary biology. Cooperation is beneficial for a population as a whole but costly for the bearers of social traits such that cheaters enjoy a selective advantage over cooperators. Here we focus on coevolution of cooperators and cheaters in a multi-level selection framework, by modeling competition among groups composed of cooperators and cheaters. Cheaters enjoy a reproductive advantage over cooperators at the individual level, independent of the presence of cooperators in the group. Cooperators carry a social trait that provides a fitness advantage to the respective groups. In the case of absolute fitness advantage, where the survival probability of a group is independent of the composition of other groups, the survival of cooperators does not correlate with the presence of cheaters. By contrast, in the case of relative fitness advantage, where the survival probability of a group depends on the composition of all groups, the survival of cooperators positively correlates with the presence of cheaters. Increasing the strength of the social trait alone fails to ensure survival of cooperators, and the increase of the reproduction advantage of the cheaters is necessary to avoid population extinction. This unexpected effect comes from multilevel selection whereby cheaters at the individual level become altruists at the group level, enabling overall growth of the population that is essential for the persistence of cooperators. We validate these theoretical results with an agent-based model of a bacterial biofilm where emergence of the cooperative trait is facilitated by the presence of cheaters, leading to evolution of spatial organization. This finding contrasts the classical view that spatial organization facilitates cooperation. Our results suggest that, counterintuitively, cheaters often promote, not destabilize, evolution of cooperation.","manuscriptTitle":"The key role of cheaters in the persistence of cooperation","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-04-23 07:21:19","doi":"10.21203/rs.3.rs-6280939/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-06-09T12:56:34+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-03T08:40:34+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-05-30T07:09:00+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"58723451709361969611813343904163978354","date":"2025-05-25T20:42:01+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"191397277366495695956528943907800312246","date":"2025-05-20T02:07:25+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"314918111820352704560207987187649070329","date":"2025-05-19T16:41:11+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-26T15:29:03+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"246458129769064113440462071084077130558","date":"2025-04-03T07:18:58+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"153800270553782433750364558635771108749","date":"2025-04-02T16:10:37+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-04-02T12:54:38+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-24T14:24:56+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-03-24T09:51:22+00:00","index":"","fulltext":""},{"type":"submitted","content":"BMC Biology","date":"2025-03-22T02:07:51+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"bmc-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"Learn more about [BMC Biology](https://bmcbiol.biomedcentral.com/)","snPcode":"12915","submissionUrl":"https://submission.springernature.com/new-submission/12915/3","title":"BMC Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"fc62da11-827d-4905-93e9-e6690092a71a","owner":[],"postedDate":"April 23rd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2025-12-10T17:08:36+00:00","versionOfRecord":[],"versionCreatedAt":"2025-04-23 07:21:19","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6280939","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6280939","identity":"rs-6280939","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}