Global consequences of long-term mineral fertilization on soil microbiomes | 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 Article Global consequences of long-term mineral fertilization on soil microbiomes Huaiying Yao, Shengwen Xu, Manuel Delgado-Baquerizo, Yongxiang Yu, and 10 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7587066/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Mineral fertilization has sustained food security for decades, yet its long-term impacts on soil microbial communities underpinning soil health remain virtually unknown. We combined standardized field surveys and literature synthesis to assemble a global dataset of 501 long-term agricultural experiments (median 25 years) to evaluate the impacts of sustained mineral fertilization on soil properties, microbes and functions. A biogeochemical trade-off between organic carbon accumulation and acidification was observed under long-term mineral fertilization. Microbial biomass increased with soil organic carbon accumulation, and we found major shifts in dominant soil taxa with the relative abundance of Proteobacteria increasing and that of Firmicutes declining with acidification. Virulent bacteriophages increased, with host associations consistent with these bacterial shifts. Microbial activities revealed a decoupling of increased nitrogen- and phosphorus-acquisition enzymes without effects on carbon-mineralization enzymes. Microbial communities were taxonomically reorganized without reducing richness or promoting fungal pathogens. Our findings provide global-scale evidence on the consequences of long-term mineral fertilization for soil health, which is integral to guiding fertilizer management for sustainable agriculture. Earth and environmental sciences/Ecology/Microbial ecology Biological sciences/Microbiology/Environmental microbiology Earth and environmental sciences/Ecology/Agroecology Earth and environmental sciences/Environmental sciences/Environmental impact Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Full Text Additional Declarations There is NO Competing Interest. Supplementary Files SupportingInformationXufin.docx Supplementary Information for “Global consequences of long-term mineral fertilization on soil microbiomes” ExtendedDataFig1Xu.tif Extended Data Fig. 1. Conceptual figure of globally consistent soil microbial responses to long-term mineral fertilization. Cite Share Download PDF Status: Posted Version 1 posted 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-7587066","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":517212254,"identity":"2d2c8ea4-9331-4000-ae89-0318be7db5fa","order_by":0,"name":"Huaiying Yao","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAsklEQVRIiWNgGAWjYBACAzBZIUGyljMka2FsI0EHg7lEjuGHj/MsEvsZmB9+YKi5Q1iL5YwcY8mZ2yQSZzawGUswHHtGhMNu5BhI8wK1bDjAYMbA2HCYKC3Gv3nnSCTuP8D+jWgtZtK8DUBbGHiIteXMszLLGcckjGcc5imWSDhGjJbjyZtvfKipk+1vb9/44UMNEVoYBDIgscnADMQJRGhgYOA//oAodaNgFIyCUTCCAQCK0jhL3JT5wAAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-1932-8765","institution":"Wuhan Institute of Technology","correspondingAuthor":true,"prefix":"","firstName":"Huaiying","middleName":"","lastName":"Yao","suffix":""},{"id":517212255,"identity":"2473da5c-fad9-4b13-80dc-23dd5369cef8","order_by":1,"name":"Shengwen Xu","email":"","orcid":"https://orcid.org/0000-0002-0534-0835","institution":"Wuhan Institute of Technology","correspondingAuthor":false,"prefix":"","firstName":"Shengwen","middleName":"","lastName":"Xu","suffix":""},{"id":517212256,"identity":"a4c20693-43a7-4089-9bac-2c94b4db5f02","order_by":2,"name":"Manuel Delgado-Baquerizo","email":"","orcid":"https://orcid.org/0000-0002-6499-576X","institution":"Instituto de Recursos Naturales y Agrobiología de Sevilla (IRNAS)","correspondingAuthor":false,"prefix":"","firstName":"Manuel","middleName":"","lastName":"Delgado-Baquerizo","suffix":""},{"id":517212257,"identity":"798b87ca-2fda-4793-8aef-996fb415a5c0","order_by":3,"name":"Yongxiang Yu","email":"","orcid":"","institution":"Wuhan Institute of Technology","correspondingAuthor":false,"prefix":"","firstName":"Yongxiang","middleName":"","lastName":"Yu","suffix":""},{"id":517212258,"identity":"4c160c95-ddc1-417f-83ae-2ef66ba7b177","order_by":4,"name":"Haoxin Fan","email":"","orcid":"","institution":"Wuhan Institute of Technology","correspondingAuthor":false,"prefix":"","firstName":"Haoxin","middleName":"","lastName":"Fan","suffix":""},{"id":517212259,"identity":"b586adb1-dac3-4b05-b66b-62c54fb96621","order_by":5,"name":"Limei Zhang","email":"","orcid":"https://orcid.org/0000-0002-7383-8475","institution":"Research Center for Eco-Environmental Sciences, Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Limei","middleName":"","lastName":"Zhang","suffix":""},{"id":517212260,"identity":"70040ce7-be5c-4864-b320-10bdf505369f","order_by":6,"name":"Xiaoyan Tang","email":"","orcid":"","institution":"Sichuan Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Xiaoyan","middleName":"","lastName":"Tang","suffix":""},{"id":517212261,"identity":"fcce3725-b994-448e-9ef7-5501c3cc100e","order_by":7,"name":"Zhi Zhang","email":"","orcid":"","institution":"Shaanxi Normal University","correspondingAuthor":false,"prefix":"","firstName":"Zhi","middleName":"","lastName":"Zhang","suffix":""},{"id":517212262,"identity":"99685cae-dd40-419d-a367-3c4e1ed5b9bb","order_by":8,"name":"Shuai Wu","email":"","orcid":"https://orcid.org/0009-0009-4563-5708","institution":"Hohai University","correspondingAuthor":false,"prefix":"","firstName":"Shuai","middleName":"","lastName":"Wu","suffix":""},{"id":517212263,"identity":"9a2d8b8d-0ad9-494b-9ad2-39ae277b331e","order_by":9,"name":"Nataliya Bilyera","email":"","orcid":"","institution":"University of Tübingen","correspondingAuthor":false,"prefix":"","firstName":"Nataliya","middleName":"","lastName":"Bilyera","suffix":""},{"id":517212264,"identity":"6103870f-5278-491d-a67a-3a4d7dfe2b6d","order_by":10,"name":"Yakov Kuzyakov","email":"","orcid":"https://orcid.org/0000-0002-9863-8461","institution":"Georg-August University of Göttingen","correspondingAuthor":false,"prefix":"","firstName":"Yakov","middleName":"","lastName":"Kuzyakov","suffix":""},{"id":517212265,"identity":"91b24e79-fd17-44fe-b6e5-a61d96fe6789","order_by":11,"name":"Christoph Tebbe","email":"","orcid":"https://orcid.org/0000-0003-4861-0214","institution":"Thünen Institute - Federal Research Institute für Rural Areas, Forestry and Fisheries","correspondingAuthor":false,"prefix":"","firstName":"Christoph","middleName":"","lastName":"Tebbe","suffix":""},{"id":517212266,"identity":"e1a89fde-3cbf-4f77-885d-a483cf73a7af","order_by":12,"name":"Graeme Nicol","email":"","orcid":"","institution":"Université Claude Bernard Lyon 1","correspondingAuthor":false,"prefix":"","firstName":"Graeme","middleName":"","lastName":"Nicol","suffix":""},{"id":517212267,"identity":"0332d6b2-744b-4b12-8602-76b5055b150a","order_by":13,"name":"Joann Whalen","email":"","orcid":"https://orcid.org/0000-0001-8774-0594","institution":"McGill University","correspondingAuthor":false,"prefix":"","firstName":"Joann","middleName":"","lastName":"Whalen","suffix":""}],"badges":[],"createdAt":"2025-09-11 02:50:38","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7587066/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7587066/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":91671550,"identity":"17dde049-a4fd-4780-91aa-5ebaa6f0498e","added_by":"auto","created_at":"2025-09-19 03:49:17","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":820733,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eGlobal distribution and characteristics of long-term mineral fertilization experiments. \u003c/strong\u003e(\u003cstrong\u003ea\u003c/strong\u003e)\u003cstrong\u003e \u003c/strong\u003eLocations of 501 long-term experiments (LTEs), each comprising paired plots with mineral N (minN) fertilizer and unfertilized control. Symbol size is proportional to fertilization duration. Lower panels show proportions of site-level metadata categories derived from literature synthesis and field sampling. (\u003cstrong\u003eb\u003c/strong\u003e) Mean annual temperature (MAT) and precipitation (MAP), with the five experiments of longest duration highlighted. (\u003cstrong\u003ec\u003c/strong\u003e) Boxplots of elemental N, P, and K application rates under single and compound minN fertilization (with P and/or K) in paddy and upland systems. Numbers above boxes denote means.\u003c/p\u003e","description":"","filename":"Figure1Xu.png","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/d5da0e5e8efb59f33ca39f4e.png"},{"id":91672077,"identity":"86dbe933-e021-4fd9-a374-322e65a267d9","added_by":"auto","created_at":"2025-09-19 03:57:17","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":572583,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eEffects of long-term mineral fertilization on soil physicochemical properties, crop productivity and microbial responses. \u003c/strong\u003eBias-robust estimates (±95% confidence intervals) derived from multilevel meta-analysis models (see Methods). Solid red circles indicate significant negative effects, solid blue circles indicate significant positive effects, and open circles denote non-significant effects. Asterisks denote significance (*\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05; **\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01; ***\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001). Numbers of effect sizes (\u003cem\u003ek\u003c/em\u003e) are shown in parentheses.\u003c/p\u003e","description":"","filename":"Figure2Xu.png","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/a772cbc27197082bb15d4dd5.png"},{"id":91671552,"identity":"7acce0d7-bd50-42e2-82a9-3b365e4e1688","added_by":"auto","created_at":"2025-09-19 03:49:18","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":218575,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eEffects of long-term mineral fertilization on microbial taxa and functional guilds. \u003c/strong\u003eBias-robust estimates (±95% confidence intervals) for each variable, derived from multilevel meta-analysis models (see Methods). Solid red circles indicate significant negative effects, solid blue circles indicate significant positive effects, and open circles denote non-significant effects. Asterisks denote significance (*\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05; **\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.01). Number of effect sizes (\u003cem\u003ek\u003c/em\u003e) is shown in parentheses.\u003c/p\u003e","description":"","filename":"Figure3Xu.png","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/174c9699ce3773e754b45206.png"},{"id":91671551,"identity":"23d64b12-b709-4cfa-bdba-c52f13cd3168","added_by":"auto","created_at":"2025-09-19 03:49:17","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":610986,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSoil organic carbon and pH explain microbial responses to long-term mineral fertilization, with virulent phage patterns consistent with bacterial host shifts. \u003c/strong\u003e(\u003cstrong\u003ea\u003c/strong\u003e, \u003cstrong\u003eb\u003c/strong\u003e) Inverse-variance weighted regressions show higher soil organic carbon was associated with increased microbial biomass, whereas acidification was associated with increased \u003cem\u003eProteobacteria\u003c/em\u003e and decreased \u003cem\u003eFirmicutes\u003c/em\u003e. Regression lines and shaded areas represent slopes and 95% confidence intervals for all sites (grey), sites with fertilization duration ≤25 years (green), and \u0026gt;25 years (blue). (\u003cstrong\u003ec\u003c/strong\u003e) Heatmap from standardized field surveys shows that most sites exhibited increases in abundance of \u003cem\u003eProteobacteria\u003c/em\u003e and virulent phages and decreases in \u003cem\u003eFirmicutes\u003c/em\u003e at the class level. Colors denote log2 fold change (mineral-fertilized relative to unfertilized), truncated at ±0.5. (\u003cstrong\u003ed\u003c/strong\u003e) Co-occurrence networks of host-associated virulent phage vOTUs (present in at least 5% of samples and among the 300 most abundant taxa) show that \u003cem\u003eActinobacteria\u003c/em\u003ewere the most common predicted hosts. Long-term mineral fertilization increased the connectivity of virulent phages with \u003cem\u003eProteobacteria\u003c/em\u003e and reduced their connectivity with \u003cem\u003eFirmicutes\u003c/em\u003e. Nodes are colored by predicted bacterial host phylum and scaled to degree, and edges represent significant correlations (|r| \u0026gt; 0.5, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"Figure4Xu.png","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/9ecedc1b267a6af50ac98753.png"},{"id":91671553,"identity":"36c6c340-2e29-4f54-9a0f-abea5880a835","added_by":"auto","created_at":"2025-09-19 03:49:18","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":554184,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSoil acidification and soil organic carbon accumulation broadly explain microbial responses to long-term mineral fertilization. \u003c/strong\u003e(\u003cstrong\u003ea\u003c/strong\u003e) The magnitude of soil acidification (△pH) and soil organic carbon (SOC) accumulation (lnRR) increased with higher fertilization intensity, peaking at neutral pH and low baseline SOC, respectively. Red diamonds denote bin-averaged means, and curves with shaded bands represent generalized additive model fits to all data points. (\u003cstrong\u003eb\u003c/strong\u003e) Correlation heatmap showing weighted regressions between microbial responses and fertilizer-induced changes in soil properties, geographic variables, and baseline edaphic factors. Circle size and color indicate correlation coefficient (r). Asterisks denote significance (*\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05, **\u003cem\u003ep\u003c/em\u003e\u0026lt; 0.01, ***\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001). Note that △pH is negative under acidification; correlation signs reflect responses to pH decline. Variables shown in bold blue or red font respectively indicate overall increases or decreases in the meta-analysis.\u003c/p\u003e","description":"","filename":"Figure5Xu.png","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/69654c39ad328e261d996764.png"},{"id":91671554,"identity":"3536e58a-052d-4c31-a34c-fa70e91ba17f","added_by":"auto","created_at":"2025-09-19 03:49:18","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":830343,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eGlobal predictions of microbial responses to long-term mineral fertilization. \u003c/strong\u003eGeneralized additive models were trained using meta-analytic records linking microbial responses (lnRR) with edaphic and climatic predictors, including annual mineral nitrogen application rate. Predictions were extrapolated to global croplands with available predictors. Spatial smoothing terms were included to account for geographic autocorrelation (see Methods). Bivariate maps display predicted effects and associated certainty (1/standard error). Model performance was evaluated using the coefficient of determination (R\u003csup\u003e2\u003c/sup\u003e), root mean square error (RMSE), and mean absolute error (MAE).\u003cstrong\u003e \u003c/strong\u003eMap lines do not necessarily depict accepted national boundaries.\u003cstrong\u003e \u003c/strong\u003e(\u003cstrong\u003ea\u003c/strong\u003e) Predicted increases in microbial biomass carbon.\u003cstrong\u003e \u003c/strong\u003e(\u003cstrong\u003eb\u003c/strong\u003e) Predicted increases in the relative abundance of \u003cem\u003eProteobacteria\u003c/em\u003e and decreases in \u003cem\u003eFirmicutes\u003c/em\u003e. In both panels, upper-right tiles indicate high-certainty increases, and upper-left tiles (outlined in red) indicate high-certainty decreases. Partial effects show the independent influence of soil organic carbon (SOC) and pH on lnRR, where positive values indicate stronger fertilization effects and negative values indicate weaker effects. Higher SOC was associated with greater MBC increases, whereas lower pH was associated with \u003cem\u003eProteobacteria\u003c/em\u003e increases and \u003cem\u003eFirmicutes\u003c/em\u003e decreases.\u003c/p\u003e","description":"","filename":"Figure6Xu.png","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/f09a759055ee0e5c249decb1.png"},{"id":91672540,"identity":"1fe8580a-1451-434f-8403-43e44174d6b2","added_by":"auto","created_at":"2025-09-19 04:13:26","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1744068,"visible":true,"origin":"","legend":"","description":"","filename":"ManuscriptXufin.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1_covered_3d51fb9a-1136-4d69-b90a-d8d50ab7bcac.pdf"},{"id":91671557,"identity":"ce03d5aa-c6af-4791-acd7-0cd818171c0d","added_by":"auto","created_at":"2025-09-19 03:49:18","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":30976650,"visible":true,"origin":"","legend":"Supplementary Information for \u0026#x201C;Global consequences of long-term mineral fertilization on soil microbiomes\u0026#x201D;","description":"","filename":"SupportingInformationXufin.docx","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/8169a8e0f845751f31ff3b79.docx"},{"id":91671556,"identity":"3b9d2cb3-d695-42d2-92b4-5a0a1febb303","added_by":"auto","created_at":"2025-09-19 03:49:18","extension":"tif","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":2416797,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eExtended Data Fig. 1. Conceptual figure of globally consistent soil microbial responses to long-term mineral fertilization.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"ExtendedDataFig1Xu.tif","url":"https://assets-eu.researchsquare.com/files/rs-7587066/v1/9edaf2ac9b08a216b00682c0.tif"}],"financialInterests":"There is \u003cb\u003eNO\u003c/b\u003e Competing Interest.","formattedTitle":"Global consequences of long-term mineral fertilization on soil microbiomes","fulltext":[],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":true,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":true,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"
[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-7587066/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7587066/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Mineral fertilization has sustained food security for decades, yet its long-term impacts on soil microbial communities underpinning soil health remain virtually unknown. We combined standardized field surveys and literature synthesis to assemble a global dataset of 501 long-term agricultural experiments (median 25 years) to evaluate the impacts of sustained mineral fertilization on soil properties, microbes and functions. A biogeochemical trade-off between organic carbon accumulation and acidification was observed under long-term mineral fertilization. Microbial biomass increased with soil organic carbon accumulation, and we found major shifts in dominant soil taxa with the relative abundance of Proteobacteria increasing and that of Firmicutes declining with acidification. Virulent bacteriophages increased, with host associations consistent with these bacterial shifts. Microbial activities revealed a decoupling of increased nitrogen- and phosphorus-acquisition enzymes without effects on carbon-mineralization enzymes. Microbial communities were taxonomically reorganized without reducing richness or promoting fungal pathogens. Our findings provide global-scale evidence on the consequences of long-term mineral fertilization for soil health, which is integral to guiding fertilizer management for sustainable agriculture.","manuscriptTitle":"Global consequences of long-term mineral fertilization on soil microbiomes","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-19 03:49:13","doi":"10.21203/rs.3.rs-7587066/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"
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