A pendant awn phenotype linked to the EMBRYONIC FLOWER 1 LIKE (EMF1L) gene in barley | 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 A pendant awn phenotype linked to the EMBRYONIC FLOWER 1 LIKE (EMF1L) gene in barley Srijan Jhingan, Zsa Zsa Friederique Boyny, Twan Rutten, Axel Himmelbach, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7764158/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 19 Dec, 2025 Read the published version in Theoretical and Applied Genetics → Version 1 posted 4 You are reading this latest preprint version Abstract Awns are apical extensions of the lemma that are prevalent in many wild and cultivated grass species. In cultivated cereals they are relevant, given their influence on grain yield through contributions to photosynthesis, transpiration, carbohydrate accumulation and drought stress mitigation. While several genetic factors controlling barley awn traits have been identified, their complex network and interactions are not fully understood. This study characterizes a “pendant awn” mutant derived from an ethyl methanesulfonate (EMS)-mutagenized population of the two-rowed winter barley cultivar ‘Igri’ which is marked by shorter, thinner, and less upright awns. Segregation analyses in an F₂ population of 149 individuals indicated a monogenic recessive inheritance of the mutant trait. Molecular genetic mapping identified a 356.8 Mbp region on chromosome 3H associated with the pendant awn mutant trait. Whole-genome re-sequencing of mutant and wild-type individuals led to discovery of a premature stop codon mutation specific to the pendant awn mutant in the EMBRYONIC FLOWER 1 LIKE (HvEMF1L) gene, predicted to truncate the protein by 62.3%. Pan-genomic and -transcriptomic analyses revealed a high level of sequence conservation of HvEMF1L in global barley germplasm and an inflorescence-specific gene expression profile. The study highlights HvEMF1L as a putative regulator of barley awn development and underscores the scope of combining mutagenesis, genetic mapping, and genome sequencing for trait dissection. Hordeum vulgare EMS mutagenesis awn development trait mapping Full Text Supplementary Files PendantawnSuppldata01.10.2025.pdf Cite Share Download PDF Status: Published Journal Publication published 19 Dec, 2025 Read the published version in Theoretical and Applied Genetics → Version 1 posted Reviewers agreed at journal 14 Oct, 2025 Reviewers invited by journal 14 Oct, 2025 Editor assigned by journal 03 Oct, 2025 First submitted to journal 01 Oct, 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-7764158","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":529341184,"identity":"9089ea90-823f-443f-95b7-4a12d4111ccb","order_by":0,"name":"Srijan Jhingan","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABHUlEQVRIie2QsWrDMBCGJQSa7GSVsbFfQcZDX8da6imQsYOhKgZ5c1dB+jA2AmfxA2QItKXgPTQEQ4qpnThLUU3HDvqWOw593P0CwGD4h0AJ+Vh9DPBthji61HJeiSaFjrN5BZBrYfzPCtpk4v2Y7pPCFTE8pl/+3XP19LF+2IPFlusPe6nyyKvblfDqEnk1jbwdyyLZtMBp9GugZMIlWK0ESfIzwZRJAoVrCwXoLp5RepVgknBEevool1V+tvtBeX37VXEOQsWY3JfwIGhMABPI5uMWffzLFliocMgSI1hEoSQsc2WtLKfRHxbKpHW6kwqCjaCwO/kBWarqc50qf7HVxw/58NHW2JFbM2Fp3w8E43HdpFwbg8FgMPzgG+XdY+YWGqxDAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-9746-8659","institution":"Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung Gatersleben: Leibniz-Institut fur Pflanzengenetik und Kulturpflanzenforschung (IPK)","correspondingAuthor":true,"prefix":"","firstName":"Srijan","middleName":"","lastName":"Jhingan","suffix":""},{"id":529341185,"identity":"0622227b-1b3e-4c0d-8a1e-523b9678c767","order_by":1,"name":"Zsa Zsa Friederique Boyny","email":"","orcid":"","institution":"Justus Liebig University Giessen: Justus-Liebig-Universitat Giessen","correspondingAuthor":false,"prefix":"","firstName":"Zsa","middleName":"Zsa Friederique","lastName":"Boyny","suffix":""},{"id":529341186,"identity":"16956920-5482-434c-99f6-8be984fbe8b0","order_by":2,"name":"Twan Rutten","email":"","orcid":"","institution":"Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung Gatersleben: Leibniz-Institut fur Pflanzengenetik und Kulturpflanzenforschung (IPK)","correspondingAuthor":false,"prefix":"","firstName":"Twan","middleName":"","lastName":"Rutten","suffix":""},{"id":529341187,"identity":"68fc493a-5bd7-4a6f-a605-354f6957ae56","order_by":3,"name":"Axel Himmelbach","email":"","orcid":"","institution":"Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung Gatersleben: Leibniz-Institut fur Pflanzengenetik und Kulturpflanzenforschung (IPK)","correspondingAuthor":false,"prefix":"","firstName":"Axel","middleName":"","lastName":"Himmelbach","suffix":""},{"id":529341188,"identity":"0f5cc910-596b-49cb-8ea2-4465392ea5b2","order_by":4,"name":"Nils Stein","email":"","orcid":"https://orcid.org/0000-0003-3011-8731","institution":"Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung Gatersleben: Leibniz-Institut fur Pflanzengenetik und Kulturpflanzenforschung (IPK)","correspondingAuthor":false,"prefix":"","firstName":"Nils","middleName":"","lastName":"Stein","suffix":""}],"badges":[],"createdAt":"2025-10-02 06:31:07","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7764158/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7764158/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00122-025-05105-5","type":"published","date":"2025-12-19T15:58:14+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":94576355,"identity":"b77473db-18c7-4c39-978b-0ba4adbdd913","added_by":"auto","created_at":"2025-10-28 18:10:01","extension":"xml","order_by":1,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":9618,"visible":true,"origin":"","legend":"","description":"","filename":"taagTAAGD2500893.xml","url":"https://assets-eu.researchsquare.com/files/rs-7764158/v1/07c46c8981adaa7c4dedb0bd.xml"},{"id":94575181,"identity":"c1990c96-678d-42b6-8636-560154b76fcc","added_by":"auto","created_at":"2025-10-28 18:09:13","extension":"xml","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1002,"visible":true,"origin":"","legend":"","description":"","filename":"TAAGD250089319452.go.xml","url":"https://assets-eu.researchsquare.com/files/rs-7764158/v1/7c42dc73f83419499851703f.xml"},{"id":94575472,"identity":"f5cc8f10-ff4d-432c-884f-06847ff5714e","added_by":"auto","created_at":"2025-10-28 18:09:22","extension":"xml","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":790,"visible":true,"origin":"","legend":"","description":"","filename":"TAAGD2500893Import.xml","url":"https://assets-eu.researchsquare.com/files/rs-7764158/v1/2a9ca534c17b8e8c4e6aa5ed.xml"},{"id":98813971,"identity":"70c00019-f162-4867-b319-a4a58d37e61a","added_by":"auto","created_at":"2025-12-22 16:08:45","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1885832,"visible":true,"origin":"","legend":"","description":"","filename":"Pendantawndraft01.10.2025.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7764158/v1_covered_999d8356-c3a8-475e-b73e-a230d49587b0.pdf"},{"id":94575689,"identity":"d68fec90-e79a-4f0b-97b2-2b1baa904831","added_by":"auto","created_at":"2025-10-28 18:09:31","extension":"pdf","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":2515279,"visible":true,"origin":"","legend":"","description":"","filename":"PendantawnSuppldata01.10.2025.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7764158/v1/c3c833315b19a7e9a8f229bb.pdf"}],"financialInterests":"","formattedTitle":"A pendant awn phenotype linked to the EMBRYONIC FLOWER 1 LIKE (EMF1L) gene in barley","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":"theoretical-and-applied-genetics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"taag","sideBox":"Learn more about [Theoretical and Applied Genetics](https://www.springer.com/journal/122)","snPcode":"122","submissionUrl":"https://submission.nature.com/new-submission/122/3","title":"Theoretical and Applied Genetics","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Hordeum vulgare, EMS mutagenesis, awn development, trait mapping","lastPublishedDoi":"10.21203/rs.3.rs-7764158/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7764158/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Awns are apical extensions of the lemma that are prevalent in many wild and cultivated grass species. In cultivated cereals they are relevant, given their influence on grain yield through contributions to photosynthesis, transpiration, carbohydrate accumulation and drought stress mitigation. While several genetic factors controlling barley awn traits have been identified, their complex network and interactions are not fully understood. This study characterizes a “pendant awn” mutant derived from an ethyl methanesulfonate (EMS)-mutagenized population of the two-rowed winter barley cultivar ‘Igri’ which is marked by shorter, thinner, and less upright awns. Segregation analyses in an F₂ population of 149 individuals indicated a monogenic recessive inheritance of the mutant trait. Molecular genetic mapping identified a 356.8 Mbp region on chromosome 3H associated with the pendant awn mutant trait. Whole-genome re-sequencing of mutant and wild-type individuals led to discovery of a premature stop codon mutation specific to the pendant awn mutant in the EMBRYONIC FLOWER 1 LIKE (HvEMF1L) gene, predicted to truncate the protein by 62.3%. Pan-genomic and -transcriptomic analyses revealed a high level of sequence conservation of HvEMF1L in global barley germplasm and an inflorescence-specific gene expression profile. The study highlights HvEMF1L as a putative regulator of barley awn development and underscores the scope of combining mutagenesis, genetic mapping, and genome sequencing for trait dissection.","manuscriptTitle":"A pendant awn phenotype linked to the EMBRYONIC FLOWER 1 LIKE (EMF1L) gene in barley","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-28 13:50:26","doi":"10.21203/rs.3.rs-7764158/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-10-14T09:20:17+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-10-14T09:10:43+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-10-03T16:16:19+00:00","index":"","fulltext":""},{"type":"submitted","content":"Theoretical and Applied Genetics","date":"2025-10-02T02:30:27+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"
[email protected]","identity":"theoretical-and-applied-genetics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"taag","sideBox":"Learn more about [Theoretical and Applied Genetics](https://www.springer.com/journal/122)","snPcode":"122","submissionUrl":"https://submission.nature.com/new-submission/122/3","title":"Theoretical and Applied Genetics","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"7549c1c6-e5d3-41bd-bda3-7a56c0331cb7","owner":[],"postedDate":"October 28th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-12-22T16:02:45+00:00","versionOfRecord":{"articleIdentity":"rs-7764158","link":"https://doi.org/10.1007/s00122-025-05105-5","journal":{"identity":"theoretical-and-applied-genetics","isVorOnly":false,"title":"Theoretical and Applied Genetics"},"publishedOn":"2025-12-19 15:58:14","publishedOnDateReadable":"December 19th, 2025"},"versionCreatedAt":"2025-10-28 13:50:26","video":"","vorDoi":"10.1007/s00122-025-05105-5","vorDoiUrl":"https://doi.org/10.1007/s00122-025-05105-5","workflowStages":[]},"version":"v1","identity":"rs-7764158","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7764158","identity":"rs-7764158","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}
Text is read by the "Ask this paper" AI Q&A widget below.
Extraction quality varies by source — PMC NXML preserves structure
cleanly, OA-HTML may include some navigation residue, and OA-PDF can
have broken hyphenation. The publisher copy
(via DOI)
is the canonical version.