The impact of weeds on performance of cereal-legume intercropping systems

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Managing species dominance in cereal-legume intercrop systems | 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 Managing species dominance in cereal-legume intercrop systems David Kottelenberg, Jochem Evers, Niels Anten, Lammert Bastiaans This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5930680/v2 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 31 May, 2026 Read the published version in European Journal of Agronomy → Version 2 posted You are reading this latest preprint version Show more versions Abstract Weeds threaten crop productivity, while reliance on herbicides raises environmental and health concerns. Intercropping offers an alternative for sustainable weed suppression, particularly when component crops differ in competitive ability. However, such differences can also lead to competitive dominance by the stronger crop species, reducing the yield of its companion. To design intercrops for enhanced weed suppression without compromising yield, it is important to understand how species selection and spatial arrangements influence competitive dynamics. We conducted four field experiments (2022–2024) evaluating how species combination, mixing ratio, and spatial design influence dominance in cereal–legume intercrops under herbicide-treated and weed-infested conditions. Cereals showed superior light capture compared to legumes, which aligned with higher weed suppression but not necessarily with competitive dominance, indicating that strong weed suppression can be achieved without excessive legume suppression. Species choice strongly influenced competitive balance: rye severely suppressed legume yields, while other species combinations showed more balanced competitive relationships. In triticale-faba bean intercrops, reducing triticale proportion improved faba bean’s relative performance, with the 1:3 triticale:faba bean ratio achieving balanced yield contributions from both species. Wider row spacing reduced triticale’s competitive dominance but decreased total intercrop productivity. Under weed-infested conditions competitive dynamics shifted, with reduced triticale dominance and improved faba bean performance, suggesting that cereal–weed competition indirectly facilitates legume growth. These patterns demonstrate that thoughtful intercrop design can balance weed suppression with legume productivity, providing a viable approach to reducing chemical weed control while maintaining crop yields. Agronomy Agroecology Cereal-legume intercropping herbicide weed yield facilitation competition light interception Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Full Text Additional Declarations The authors declare no competing interests. Supplementary Files Supplement1.pdf Supplement 1 Supplement2.pdf Supplement 2 Supplement3.pdf Supplement 3 Supplement4.pdf Supplement 4 Cite Share Download PDF Status: Published Journal Publication published 31 May, 2026 Read the published version in European Journal of Agronomy → Version 2 posted You are reading this latest preprint version Show more versions 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. 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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-5930680","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":409401289,"identity":"73db5d44-36e5-4594-aa09-b52457fafaea","order_by":0,"name":"David 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19:57:34","extension":"html","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":239859,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/685278976d97fa3a1bc64643.html"},{"id":95242918,"identity":"4f626159-c126-46ce-8f81-ce7e08e117d7","added_by":"auto","created_at":"2025-11-05 19:57:33","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":294998,"visible":true,"origin":"","legend":"\u003cp\u003e2022 light interception. Proportion of photosynthetically active radiation (PAR) intercepted over time (in cumulative daily average temperature, or °Cd) \u0026nbsp;for sole crops (a; rye, barley, triticale, wheat, pea, and faba bean) and intercrops (b-i; Rye-Pea, etc.) in the 2022 experiment. The values in the legend of subplot a are the sole crop weed biomasses in g m\u003csup\u003e-2\u003c/sup\u003e that were measured at 21 June 2022, indicated by the dashed line at 873.3 °Cd. Coloured lines are fitted logistic functions assuming a normal distribution.\u003c/p\u003e","description":"","filename":"figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/f9fcd35f22221af6ea760dc5.png"},{"id":95242919,"identity":"6cb8886d-c6e5-40b3-affc-7c66bb8b2fcf","added_by":"auto","created_at":"2025-11-05 19:57:33","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":147383,"visible":true,"origin":"","legend":"\u003cp\u003eYield of the 2022 experiment. Yield (g m\u003csup\u003e-2\u003c/sup\u003e) of cereal (a) and legume (b) sole crops (rye, barley, triticale, wheat, pea, faba bean) and 1:1 ratio row intercrops (rye-faba bean: 1R:1F, etc.). Letters indicate significant differences between treatments within cereal or legume groups at P \u0026lt; 0.05.\u003c/p\u003e","description":"","filename":"figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/2b9ce8cd76635cc9981b98e5.png"},{"id":95312395,"identity":"c90e1edc-9a8f-4c26-a467-ffee29049c15","added_by":"auto","created_at":"2025-11-06 15:49:10","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":145433,"visible":true,"origin":"","legend":"\u003cp\u003e2023-SP light interception. Proportion of photosynthetically active radiation (PAR) intercepted over time (in cumulative daily average temperature, or °Cd)\u0026nbsp; for triticale and faba bean sole crops (high density triticale: T+; normal density triticale: T; high density faba bean: F+, normal density faba bean: F) and intercrop treatments, including 3:1, 1:1, and 1:3 ratio triticale-faba bean row intercrops (3T:1F, 1T:1F, 1T:3F, respectively) and 1:1 ratio mixed intercrop (1T:1F-M), in the 2023-SP experiment. The values in the legend of subplot a are the weed biomasses in g m\u003csup\u003e-2\u003c/sup\u003e that were measured at 23 May 2023, indicated by the dashed line at 780.3 °Cd. Coloured lines are fitted logistic functions assuming a normal distribution. The grey lines in (b) are the lines of T (top grey line) and F (bottom) from (a).\u003c/p\u003e","description":"","filename":"figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/7069430fb5d9e7801780c82c.png"},{"id":95312969,"identity":"9c80fd40-b0bd-4e88-96e0-5e7ef6a845fa","added_by":"auto","created_at":"2025-11-06 15:50:39","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":86704,"visible":true,"origin":"","legend":"\u003cp\u003eYield of the 2023-SP experiment. Yield (g m\u003csup\u003e-2\u003c/sup\u003e) of triticale (a) and faba bean (b) sole crops (high density triticale: T+; normal density triticale: T; high density faba bean: F+, normal density faba bean: F) and intercrop treatments, including 3:1, 1:1, and 1:3 ratio triticale-faba bean row intercrops (3T:1F, 1T:1F, 1T:3F, respectively) and 1:1 ratio mixed intercrop (1T:1F-M). Letters indicate significant differences at P \u0026lt; 0.05.\u003c/p\u003e","description":"","filename":"figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/9ea9d8668fee33fd1e8417a8.png"},{"id":95312772,"identity":"a13fe0dc-c7ff-4bfd-a948-8d8fb88acecc","added_by":"auto","created_at":"2025-11-06 15:50:16","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":214580,"visible":true,"origin":"","legend":"\u003cp\u003e2023-RD light interception. Proportion of photosynthetically active radiation (PAR) intercepted over time (in cumulative daily average temperature, or °Cd)\u0026nbsp; for triticale and faba bean sole crops (triticale: T; faba bean: F) and intercrop treatments, including 1:1 ratio row and mixed intercrops (1T:1F and 1T:1F-M, respectively), at 12.5 cm or 37.5 cm row distance (37.5 cm row distance indicated with ‘-375’), in the 2023-RD experiment. The values in the legend of subplot a are the weed biomasses in g m\u003csup\u003e-2\u003c/sup\u003e that were measured at 23 May 2023, indicated by the dashed line at 780.3 °Cd. Coloured lines are fitted logistic functions assuming a normal distribution.\u003c/p\u003e","description":"","filename":"figure5.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/350e77c98a1e60a696b9f8e1.png"},{"id":95312464,"identity":"e841bf52-88fe-427d-8fe3-6ef17bf6c72d","added_by":"auto","created_at":"2025-11-06 15:49:28","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":90802,"visible":true,"origin":"","legend":"\u003cp\u003eYield of the 2023-RD experiment. Yield (g m\u003csup\u003e-2\u003c/sup\u003e) of triticale (a) and faba bean (b) sole crops (triticale: T; faba bean: F) and intercrop treatments, including 1:1 ratio row and mixed intercrops (1T:1F and 1T:1F-M, respectively), at 12.5 cm or 37.5 cm row distance (37.5 cm row distance indicated with ‘-375’). Letters indicate significant differences at P \u0026lt; 0.05.\u003c/p\u003e","description":"","filename":"figure6.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/8967475dbc39638953f02089.png"},{"id":95242934,"identity":"47568213-ed52-4ee5-b941-f55ac64cc653","added_by":"auto","created_at":"2025-11-05 19:57:33","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":174173,"visible":true,"origin":"","legend":"\u003cp\u003eLight interception across years. Proportion of photosynthetically active radiation (PAR) intercepted over time (in cumulative daily average temperature, or °Cd)\u0026nbsp; for triticale and faba bean sole crops (triticale: T; faba bean: F) and 1:1 row intercrop treatments, across the 2022, 2023-SP, 2023-RD, and 2024 experiments. The values in the legend of subplot a are the weed biomasses in g m\u003csup\u003e-2\u003c/sup\u003e that were measured at 21 June 2022 (873.3 °Cd), 23 May 2023 (780.3 °Cd), or 26 June 2024 (1045.4 °Cd), indicated by the dashed lines Coloured lines are fitted logistic functions assuming a normal distribution.\u003c/p\u003e","description":"","filename":"figure7.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/32f2417b07673e66adcb7e21.png"},{"id":95312747,"identity":"d5f7b4e1-48c6-4652-b4b6-b155b3a07a69","added_by":"auto","created_at":"2025-11-06 15:50:12","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":130881,"visible":true,"origin":"","legend":"\u003cp\u003eCrop yields across years. Triticale (a) and faba bean (b) sole crop (triticale: T; faba bean: F) and 1:1 ratio row intercrop (1T:1F) yields in the 2022, 2023-SP, 2023-RD, and 2024 experiments. Letters indicate significant differences between sole crop and intercrop per experiment at P \u0026lt; 0.05.\u003c/p\u003e","description":"","filename":"figure8.png","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/b1181fddba5ef172633106cf.png"},{"id":104266470,"identity":"d267fab2-3bd9-4a0f-bdb9-df5f2ceb66f9","added_by":"auto","created_at":"2026-03-09 20:25:25","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3146029,"visible":true,"origin":"","legend":"","description":"","filename":"Manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2_covered_6ac3127a-1e38-4a47-855e-2970f16491c8.pdf"},{"id":95312625,"identity":"e6ba869f-4233-4a0f-bb05-38427eecf2e9","added_by":"auto","created_at":"2025-11-06 15:49:50","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":1327165,"visible":true,"origin":"","legend":"\u003cp\u003eSupplement 1\u003c/p\u003e","description":"","filename":"Supplement1.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/ecb3ad877bcbf9a05e7bcc50.pdf"},{"id":95242924,"identity":"fa27df22-73b9-4f8d-9d99-c9a6c88a44a4","added_by":"auto","created_at":"2025-11-05 19:57:33","extension":"pdf","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":1871190,"visible":true,"origin":"","legend":"\u003cp\u003eSupplement 2\u003c/p\u003e","description":"","filename":"Supplement2.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/3660ecd1b0d69f37524329ea.pdf"},{"id":95312852,"identity":"fba14771-1840-4bb2-84c2-5fdcb886a33f","added_by":"auto","created_at":"2025-11-06 15:50:27","extension":"pdf","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":143327,"visible":true,"origin":"","legend":"\u003cp\u003eSupplement 3\u003c/p\u003e","description":"","filename":"Supplement3.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/746ba9c17491af174a8ba865.pdf"},{"id":95312481,"identity":"e90db08e-2c40-4019-b45e-98b27982eb2e","added_by":"auto","created_at":"2025-11-06 15:49:30","extension":"pdf","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":166275,"visible":true,"origin":"","legend":"\u003cp\u003eSupplement 4\u003c/p\u003e","description":"","filename":"Supplement4.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5930680/v2/11ed6227bf30b7f0c96a51c4.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003eManaging species dominance in cereal-legume intercrop systems\u003c/p\u003e","fulltext":[],"fulltextSource":"","fullText":"","funders":[{"identity":"b18a96ba-05af-448e-b03b-f83cdc8d0dae","identifier":"10.13039/501100003246","name":"Nederlandse Organisatie voor Wetenschappelijk Onderzoek","awardNumber":"17626","order_by":0}],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":false,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":false,"highlight":"","institution":"Wageningen University \u0026 Research","isAcceptedByJournal":true,"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":"Cereal-legume intercropping, herbicide, weed, yield, facilitation, competition, light interception","lastPublishedDoi":"10.21203/rs.3.rs-5930680/v2","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5930680/v2","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWeeds threaten crop productivity, while reliance on herbicides raises environmental and health concerns. Intercropping offers an alternative for sustainable weed suppression, particularly when component crops differ in competitive ability. However, such differences can also lead to competitive dominance by the stronger crop species, reducing the yield of its companion. To design intercrops for enhanced weed suppression without compromising yield, it is important to understand how species selection and spatial arrangements influence competitive dynamics. We conducted four field experiments (2022\u0026ndash;2024) evaluating how species combination, mixing ratio, and spatial design influence dominance in cereal\u0026ndash;legume intercrops under herbicide-treated and weed-infested conditions. Cereals showed superior light capture compared to legumes, which aligned with higher weed suppression but not necessarily with competitive dominance, indicating that strong weed suppression can be achieved without excessive legume suppression. Species choice strongly influenced competitive balance: rye severely suppressed legume yields, while other species combinations showed more balanced competitive relationships. In triticale-faba bean intercrops, reducing triticale proportion improved faba bean\u0026rsquo;s relative performance, with the 1:3 triticale:faba bean ratio achieving balanced yield contributions from both species. Wider row spacing reduced triticale\u0026rsquo;s competitive dominance but decreased total intercrop productivity. Under weed-infested conditions competitive dynamics shifted, with reduced triticale dominance and improved faba bean performance, suggesting that cereal\u0026ndash;weed competition indirectly facilitates legume growth. These patterns demonstrate that thoughtful intercrop design can balance weed suppression with legume productivity, providing a viable approach to reducing chemical weed control while maintaining crop yields.\u003c/p\u003e","manuscriptTitle":"Managing species dominance in cereal-legume intercrop systems","msid":"","msnumber":"","nonDraftVersions":[{"code":2,"date":"2025-11-05 19:57:28","doi":"10.21203/rs.3.rs-5930680/v2","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","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}},{"code":1,"date":"2025-01-31 06:47:41","doi":"10.21203/rs.3.rs-5930680/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","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}}],"origin":"","ownerIdentity":"e4737655-f45d-448b-9879-686890835f72","owner":[],"postedDate":"November 5th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":56644490,"name":"Agronomy"},{"id":56644491,"name":"Agroecology"}],"tags":[],"updatedAt":"2026-03-09T20:24:55+00:00","versionOfRecord":{"articleIdentity":"rs-5930680","link":"https://doi.org/10.1016/j.eja.2026.128065","journal":{"identity":"european-journal-of-agronomy","isVorOnly":true,"title":"European Journal of Agronomy"},"publishedOn":"2026-06-01 00:00:00","publishedOnDateReadable":"June 1st, 2026"},"versionCreatedAt":"2025-11-05 19:57:28","video":"","vorDoi":"10.1016/j.eja.2026.128065","vorDoiUrl":"https://doi.org/10.1016/j.eja.2026.128065","workflowStages":[]},"version":"v2","identity":"rs-5930680","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5930680","identity":"rs-5930680","version":["v2"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}

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