Occasional Hermaphroditism and Plasticity in the Flowering Habit of Persian Walnut (Juglans regia L.)- New Report

Occasional Hermaphroditism and Plasticity in the Flowering Habit of Persian Walnut (Juglans regia L.)- New Report | 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 Short Report Occasional Hermaphroditism and Plasticity in the Flowering Habit of Persian Walnut ( Juglans regia L.)- New Report Om Chand Sharma, Javid Iqbal Mir, Mohd Abas Shah, Mahendra Kumar Verma This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9265533/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 Walnut ( Juglans regia L.) is traditionally characterized as a monoecious plant, bearing staminate catkins on one-year-old wood and pistillate flowers on current-season growth. However, recent observations at ICAR-CITH, Srinagar, reveal significant plasticity in this bearing habit. Out of 219 bearing plants observed, instances of atypical flowering were recorded, including the development of male catkins on current-season growth and, most notably, the occurrence of hermaphrodite flowers. These hermaphrodite flowers contain both functional stamens and a receptive stigma, a phenomenon that could potentially bypass the limitations of dichogamy in commercial plantations. This report documents these rare floral transitions and suggests that environmental stressors or hormonal shifts may trigger these deviations from the standard monoecious model. Horticulture Walnut Juglans regia L Hermaphrodite flowers Floral Plasticity Dichogamy Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The Persian walnut ( Juglans regia L.) is the premier temperate nut crop in India, with primary production concentrated in Jammu & Kashmir, Himachal Pradesh, and Uttarakhand. It is successfully cultivated at altitudes ranging from 1,200 to 2,200 meters, though it has been observed at elevations exceeding 3,000 meters in the Ladakh region (Sharma et al. 2016 ). As a monoecious species of the Juglandaceae family, walnuts typically separate male and female reproductive structures on the same tree. Staminate catkins are borne laterally on one-year-old branches, while pistillate inflorescences (usually 1–3 flowers) appear terminally on current-season shoots (Polito 1998 ). A major challenge in walnut production is dichogamy-the asynchronous maturation of male and female flowers which necessitates the interplanting of specific cultivars to ensure pollen shedding matches stigma receptivity (Polito 1998 ; Solar and Stampar 20011). While most Indian plantations were historically of seedling origin, the transition toward vegetative propagation highlights the need for a deeper understanding of floral biology to ensure stable yields in solid-block plantations. Materials and Methods Observations were conducted at the existing walnut germplasm block of ICAR-CITH, Srinagar (Latitude 33°58’58.71” to 33°59’00.16” N; Longitude 74°80’06.10” to 74°40’18.09” E; Altitude 1610m). A total of 219 bearing plants were monitored for flowering behavior. Phenological changes and atypical floral structures were recorded through critical visual inspection and photographic documentation. Results and Discussion While the standard bearing habit was prevalent (Fig. 1 ), walnut trees exhibited occasional, localized changes in their reproductive patterns. The following atypical behaviors were documented: Reversed Spatial Positioning: Male catkins were observed developing on current-season growth following fruit set (Fig. 2 ). Floral Reversion: In some instances, current-season growth initially produced female flowers, but subsequently converted into catkin-like structures that eventually desiccated (Fig. 3 ). Hermaphroditism: The most significant observation was the presence of hermaphrodite flowers containing both undehisced/dehisced anthers and a receptive stigma (Fig. 4 ). The emergence of hermaphrodite flowers is a novel observation that opens new avenues for walnut research. If this trait can be stabilized or exploited, it could resolve the inherent difficulties of dichogamy by allowing for self-pollinating, homogamous cultivars. These anomalies frequently occurred during late or "offseason" flowering. Such shifts in floral identity are likely linked to environmental triggers such as high temperature or moisture stress which influence internal hormonal balances. Previous studies by Hassankhah et al. ( 2018 ; 2020 ) support the idea that walnut flowering patterns are environmentally and genetically sensitive; for instance, GA 3 application has been shown to significantly alter the male-to-female flower ratio. At a molecular level, the transition from dormancy to flowering is governed by the activation of genes such as FT, SOC1, and LFY, while TFL1 acts as a floral inhibitor. The localized "breakdown" of monoecy observed here suggests that specific micro-climatic conditions may override standard genetic suppression, leading to the development of bisexual organs. Declarations Author Contribution statement OCS conceptualized the study, developed the methodology, and conducted the primary field observations and data collection. JIM performed data curation, formal analysis, and critical manuscript review. MAS contributed to the investigation, provided photographic documentation, and assisted with technical validation. MKV provided overall supervision, facilitated resource acquisition, and performed the final editing and proofreading of the report. All authors have read and approved the final version of the manuscript for submission. Acknowledgement The authors are grateful to the Indian Council of Agricultural Research (ICAR), New Delhi, for providing the necessary financial support and facilities to carry out this research under the institutional programs at ICAR-Central Institute of Temperate Horticulture (CITH), Srinagar. References Hassankhah A, Rahemi M, Mozafari MR, Vahdati K (2018) Flower Development in Walnut: Altering the Flowering Pattern by Gibberellic Acid Application. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 46(2):700–706 Hassankhah A, Rahemi M, Ramshini H, Sarikhani S, Vahdati K (2020) Flowering in Persian walnut: patterns of gene expression during flower development. BMC Plant Biol 20(1):136 Polito VS (1998) Floral Biology: Flower Development, Pollination, and Fertilization in Walnut. Walnut Production Manual, University of California Sharma OC, Singh DB, Zahoor S, Padder BA, Haji SA (2016) Gynodioecious behaviour in some walnut genotypes-a new report. J Hill Agric 7(2):283–285 Solar A, Stampar F (2011) Characterisation of selected walnut (Juglans regia L.) cultivars by phenological and morphological parameters. Genet Resour Crop Evol 58:1155–1174 Additional Declarations The authors declare no competing interests. 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-9265533","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Short Report","associatedPublications":[],"authors":[{"id":614496938,"identity":"ac422324-a119-4aa4-8d8d-770824054e8f","order_by":0,"name":"Om Chand Sharma","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAtklEQVRIiWNgGAWjYNACNgk5BgYeErUYk6yFIbGBaC38YqcTPxeUWaRvOH724IMPDHZyug0EtEjOzt0sPeOcRO6GM3nJhjMYko3NDhDQYnA7d4M0bxtQy4EcM2kehgOJ24jQsvk3UEu6wfk3xGvZBrIlweAGsbYA/bLNmuechOHMG2+MDWcYEOEXfunczbd5yurk+c7nGD74UGEnR1ALHCiAVRoQqxwE5BtIUT0KRsEoGAUjCgAAhas/CwBrQhIAAAAASUVORK5CYII=","orcid":"","institution":"ICAR-Central Institute of Temperate Horticulture, Srinagar (J\u0026K)-191132","correspondingAuthor":true,"prefix":"","firstName":"Om","middleName":"Chand","lastName":"Sharma","suffix":""},{"id":614496939,"identity":"629a9a87-58b5-4c71-83ae-e29cd2c2a75c","order_by":1,"name":"Javid Iqbal Mir","email":"","orcid":"","institution":"ICAR-Central Institute of Temperate Horticulture, Srinagar (J\u0026K)-191132","correspondingAuthor":false,"prefix":"","firstName":"Javid","middleName":"Iqbal","lastName":"Mir","suffix":""},{"id":614496940,"identity":"3e529125-87e6-4b08-b74a-03691742c759","order_by":2,"name":"Mohd Abas Shah","email":"","orcid":"","institution":"ICAR-Central Institute of Temperate Horticulture, Srinagar (J\u0026K)-191132","correspondingAuthor":false,"prefix":"","firstName":"Mohd","middleName":"Abas","lastName":"Shah","suffix":""},{"id":614496941,"identity":"07ab9386-8065-4a06-98d5-cc456f1156c7","order_by":3,"name":"Mahendra Kumar Verma","email":"","orcid":"","institution":"ICAR-Central Institute of Temperate Horticulture, Srinagar (J\u0026K)-191132","correspondingAuthor":false,"prefix":"","firstName":"Mahendra","middleName":"Kumar","lastName":"Verma","suffix":""}],"badges":[],"createdAt":"2026-03-30 10:38:25","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":true,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":true},"doi":"10.21203/rs.3.rs-9265533/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9265533/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105812040,"identity":"c48a62a1-b661-446a-b459-01c1faa1a53a","added_by":"auto","created_at":"2026-03-31 11:37:19","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":715254,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eNormal bearing habit: (a) male catkins on previous year's growth; (b) female flowers on current season growth.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9265533/v1/2b1fd8a8989a0af4dd36fe68.jpeg"},{"id":105812041,"identity":"91fc0dd9-d501-4328-a6ac-eaf944119dce","added_by":"auto","created_at":"2026-03-31 11:37:19","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":690620,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eAtypical male catkin born on current season growth after fruit set. Catkin born on current season growth after bearing fruit (a \u0026amp; b) and catkin born on current season growth (c)\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9265533/v1/019df93fcb9a3c1d3c716ddd.jpeg"},{"id":105812042,"identity":"e10edddc-b604-4bbe-8b6c-42bd6b31ddd8","added_by":"auto","created_at":"2026-03-31 11:37:19","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":468770,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eCurrent season growth showing initial female bearing (a) followed by conversion into catkin-like structures (b).\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9265533/v1/5fb5551ff11393859e415145.jpeg"},{"id":105904165,"identity":"518b307c-d530-4bba-a7ef-026f9a496aa2","added_by":"auto","created_at":"2026-04-01 10:05:34","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":560495,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eHermaphroditic traits: (a) shoot with hermaphrodite flower; (b) individual flower with stigma and anthers; (c) dehisced anthers.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9265533/v1/6649f2ec1b9129076e69e147.jpeg"},{"id":106093007,"identity":"bf61d99f-2320-451d-a20f-664de551440f","added_by":"auto","created_at":"2026-04-03 11:32:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2905816,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9265533/v1/26e1bfd5-50a7-4add-aad4-731df0b6f4c9.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eOccasional Hermaphroditism and Plasticity in the Flowering Habit of Persian Walnut (\u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eJuglans regia\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e L.)- New Report\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe Persian walnut (\u003cem\u003eJuglans regia\u003c/em\u003e L.) is the premier temperate nut crop in India, with primary production concentrated in Jammu \u0026amp; Kashmir, Himachal Pradesh, and Uttarakhand. It is successfully cultivated at altitudes ranging from 1,200 to 2,200 meters, though it has been observed at elevations exceeding 3,000 meters in the Ladakh region (Sharma et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAs a monoecious species of the Juglandaceae family, walnuts typically separate male and female reproductive structures on the same tree. Staminate catkins are borne laterally on one-year-old branches, while pistillate inflorescences (usually 1\u0026ndash;3 flowers) appear terminally on current-season shoots (Polito \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). A major challenge in walnut production is dichogamy-the asynchronous maturation of male and female flowers which necessitates the interplanting of specific cultivars to ensure pollen shedding matches stigma receptivity (Polito \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Solar and Stampar 20011). While most Indian plantations were historically of seedling origin, the transition toward vegetative propagation highlights the need for a deeper understanding of floral biology to ensure stable yields in solid-block plantations.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eObservations were conducted at the existing walnut germplasm block of ICAR-CITH, Srinagar (Latitude 33\u0026deg;58\u0026rsquo;58.71\u0026rdquo; to 33\u0026deg;59\u0026rsquo;00.16\u0026rdquo; N; Longitude 74\u0026deg;80\u0026rsquo;06.10\u0026rdquo; to 74\u0026deg;40\u0026rsquo;18.09\u0026rdquo; E; Altitude 1610m). A total of 219 bearing plants were monitored for flowering behavior. Phenological changes and atypical floral structures were recorded through critical visual inspection and photographic documentation.\u003c/p\u003e"},{"header":"Results and Discussion","content":"\u003cp\u003eWhile the standard bearing habit was prevalent (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), walnut trees exhibited occasional, localized changes in their reproductive patterns. The following atypical behaviors were documented:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003eReversed Spatial Positioning: Male catkins were observed developing on current-season growth following fruit set (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eFloral Reversion: In some instances, current-season growth initially produced female flowers, but subsequently converted into catkin-like structures that eventually desiccated (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eHermaphroditism: The most significant observation was the presence of hermaphrodite flowers containing both undehisced/dehisced anthers and a receptive stigma (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e \u003cp\u003eThe emergence of hermaphrodite flowers is a novel observation that opens new avenues for walnut research. If this trait can be stabilized or exploited, it could resolve the inherent difficulties of dichogamy by allowing for self-pollinating, homogamous cultivars.\u003c/p\u003e \u003cp\u003eThese anomalies frequently occurred during late or \"offseason\" flowering. Such shifts in floral identity are likely linked to environmental triggers such as high temperature or moisture stress which influence internal hormonal balances. Previous studies by Hassankhah et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) support the idea that walnut flowering patterns are environmentally and genetically sensitive; for instance, GA\u003csub\u003e3\u003c/sub\u003e application has been shown to significantly alter the male-to-female flower ratio. At a molecular level, the transition from dormancy to flowering is governed by the activation of genes such as FT, SOC1, and LFY, while TFL1 acts as a floral inhibitor. The localized \"breakdown\" of monoecy observed here suggests that specific micro-climatic conditions may override standard genetic suppression, leading to the development of bisexual organs.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eAuthor Contribution statement\u003c/h2\u003e \u003cp\u003eOCS conceptualized the study, developed the methodology, and conducted the primary field observations and data collection. JIM performed data curation, formal analysis, and critical manuscript review. MAS contributed to the investigation, provided photographic documentation, and assisted with technical validation. MKV provided overall supervision, facilitated resource acquisition, and performed the final editing and proofreading of the report. All authors have read and approved the final version of the manuscript for submission.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e \u003cp\u003eThe authors are grateful to the Indian Council of Agricultural Research (ICAR), New Delhi, for providing the necessary financial support and facilities to carry out this research under the institutional programs at ICAR-Central Institute of Temperate Horticulture (CITH), Srinagar.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eHassankhah A, Rahemi M, Mozafari MR, Vahdati K (2018) Flower Development in Walnut: Altering the Flowering Pattern by Gibberellic Acid Application. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 46(2):700\u0026ndash;706\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHassankhah A, Rahemi M, Ramshini H, Sarikhani S, Vahdati K (2020) Flowering in Persian walnut: patterns of gene expression during flower development. BMC Plant Biol 20(1):136\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePolito VS (1998) Floral Biology: Flower Development, Pollination, and Fertilization in Walnut. Walnut Production Manual, University of California\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSharma OC, Singh DB, Zahoor S, Padder BA, Haji SA (2016) Gynodioecious behaviour in some walnut genotypes-a new report. J Hill Agric 7(2):283\u0026ndash;285\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSolar A, Stampar F (2011) Characterisation of selected walnut (Juglans regia L.) cultivars by phenological and morphological parameters. Genet Resour Crop Evol 58:1155\u0026ndash;1174\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"ICAR-Central Institute of Temperate Horticulture, Srinagar (J\u0026K)-191132","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"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":"Walnut, Juglans regia L, Hermaphrodite flowers, Floral Plasticity, Dichogamy","lastPublishedDoi":"10.21203/rs.3.rs-9265533/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9265533/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWalnut (\u003cem\u003eJuglans regia\u003c/em\u003e L.) is traditionally characterized as a monoecious plant, bearing staminate catkins on one-year-old wood and pistillate flowers on current-season growth. However, recent observations at ICAR-CITH, Srinagar, reveal significant plasticity in this bearing habit. Out of 219 bearing plants observed, instances of atypical flowering were recorded, including the development of male catkins on current-season growth and, most notably, the occurrence of hermaphrodite flowers. These hermaphrodite flowers contain both functional stamens and a receptive stigma, a phenomenon that could potentially bypass the limitations of dichogamy in commercial plantations. This report documents these rare floral transitions and suggests that environmental stressors or hormonal shifts may trigger these deviations from the standard monoecious model.\u003c/p\u003e","manuscriptTitle":"Occasional Hermaphroditism and Plasticity in the Flowering Habit of Persian Walnut (Juglans regia L.)- New Report","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-31 11:37:14","doi":"10.21203/rs.3.rs-9265533/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":"65115739-5292-4463-8133-5601e6c67cd9","owner":[],"postedDate":"March 31st, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":65387422,"name":"Horticulture"}],"tags":[],"updatedAt":"2026-03-31T11:37:14+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-31 11:37:14","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9265533","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9265533","identity":"rs-9265533","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}