Genome-wide characterization of WRKYs and function of SpWRKY40 and SpWRKY50 in Sesuvium portulacastrum

Genome-wide characterization of WRKYs and function of SpWRKY40 and SpWRKY50 in Sesuvium portulacastrum | 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 Genome-wide characterization of WRKYs and function of SpWRKY40 and SpWRKY50 in Sesuvium portulacastrum Yaoxiu Li, Fengyan Fang, Xiaoyou Wu, Tongjing Cui, Yingyi Yu, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7513928/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 22 Dec, 2025 Read the published version in Functional & Integrative Genomics → Version 1 posted 18 You are reading this latest preprint version Abstract WRKYs represent a large family of plant transcription factors characterized by a highly conserved WRKY domain. WRKY transcription factors are important for plant growth, development, and responses to environmental stresses. However, this family has not been previously identified in Sesuvium portulacastrum . Here, we comprehensively identified 68 SpWRKYs from the genome of S. portulacastrum by phylogenetic relationships, chromosomal localization, collinearity analysis, gene structure, duplication events, and evolutionary pressure based on nonsynonymous substitution rate/synonymous substitution rate ratios. Promoter analysis revealed the presence of multiple cis -elements associated with abiotic stress signaling, developmental regulation, phytohormone responses, light responsiveness, and tissue-specific expression. We also characterized the physicochemical and structural properties of the SpWRKY -encoded proteins, including protein property of primary structure, secondary structures, and predicted three-dimensional models, subcellular localization, conserved motifs, and conserved domains. Furthermore, transcriptome profiling under cadmium, copper, and salt stress conditions demonstrated differential expression patterns of SpWRKYs . Among them, SpWRKY40 and SpWRKY51 showed significant indued under cadmium treatment and were further validated by quantitative real-time PCR. Subcellular localization and yeast one-hybrid assays both supporting SpWRKY40 and SpWRKY51 play as transcription factors. Our study provides a detailed overview of the SpWRKY family and their potential involvement in environment tolerance. The findings offer valuable candidate genes for future applications in improving stress tolerance in S. portulacastrum and related crop species. Sesuvium portulacastrum WRKY gene family Abiotic stress SpWRKY40 SpWRKY51 Transcription factor Full Text Additional Declarations No competing interests reported. Supplementary Files S1.png Fig. S1 Prediction of the tertiary structure of SpWRKYs. S2.png Fig. S2 The conserved motifs in SpWRKYs. TableS1.TranscriptomedataofSpWRKYsunderCdtreatment.xlsx TableS2.TranscriptomedataofSpWRKYsunderCutreatment.xlsx TableS3.TranscriptomedataofSpWRKYsundersaltstress.xlsx LiYXTable.docx Cite Share Download PDF Status: Published Journal Publication published 22 Dec, 2025 Read the published version in Functional & Integrative Genomics → Version 1 posted Editorial decision: Revision requested 25 Oct, 2025 Reviews received at journal 23 Oct, 2025 Reviews received at journal 22 Oct, 2025 Reviews received at journal 21 Oct, 2025 Reviews received at journal 12 Oct, 2025 Reviewers agreed at journal 09 Oct, 2025 Reviews received at journal 08 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers agreed at journal 07 Oct, 2025 Reviewers invited by journal 07 Oct, 2025 Editor assigned by journal 12 Sep, 2025 Submission checks completed at journal 04 Sep, 2025 First submitted to journal 02 Sep, 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-7513928","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":531329392,"identity":"42edf537-9b44-4d02-a62a-82c4a02a5585","order_by":0,"name":"Yaoxiu Li","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Yaoxiu","middleName":"","lastName":"Li","suffix":""},{"id":531329394,"identity":"44abac24-367e-4cd2-9bf8-167379ac180c","order_by":1,"name":"Fengyan Fang","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Fengyan","middleName":"","lastName":"Fang","suffix":""},{"id":531329396,"identity":"6a540ef8-f765-4277-9a1c-eac7674a3d1c","order_by":2,"name":"Xiaoyou Wu","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Xiaoyou","middleName":"","lastName":"Wu","suffix":""},{"id":531329400,"identity":"9e791e5a-2e23-4756-b65b-cdd2d2abbdd5","order_by":3,"name":"Tongjing Cui","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Tongjing","middleName":"","lastName":"Cui","suffix":""},{"id":531329401,"identity":"11e08aff-1233-4246-8ca2-dd70ecdef0a5","order_by":4,"name":"Yingyi Yu","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Yingyi","middleName":"","lastName":"Yu","suffix":""},{"id":531329403,"identity":"f1ae9b32-3733-47f7-bd1b-a4fb847f80ca","order_by":5,"name":"Zhaozhen Xiang","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Zhaozhen","middleName":"","lastName":"Xiang","suffix":""},{"id":531329405,"identity":"6cd756d5-6f77-4d2e-be4c-3df03740f426","order_by":6,"name":"Guomei Zhang","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Guomei","middleName":"","lastName":"Zhang","suffix":""},{"id":531329407,"identity":"e03d6b8d-a41e-431b-a4fa-3f08061ffbac","order_by":7,"name":"Zhuangzhuang Nan","email":"","orcid":"","institution":"Guizhou University","correspondingAuthor":false,"prefix":"","firstName":"Zhuangzhuang","middleName":"","lastName":"Nan","suffix":""},{"id":531329409,"identity":"111d8fa4-4a78-4402-adf4-3ec7783bf28f","order_by":8,"name":"Shugang Hui","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAv0lEQVRIiWNgGAWjYFACHgaGhAoIU4IELWdI1sLYRooW+f61xyQezqu1NzjAfPA2D4NdHkEtBjfepUkkbjvObHCALdmahyG5mLAWiTNmQC3H2AwO8JhJ8zAcSGwg6LAZIC1zjvEYHOD/RpwWhvM9QC0NNRJAW9iI02Jwg8fYIuHYAQPJw2zGlnMMkolwWP8Zw5s/aurs+Y43P7zxpsKOCIdJJIDIwwwMzGBLCaoHAv4DILKOGKWjYBSMglEwUgEAuk04+Znc3AcAAAAASUVORK5CYII=","orcid":"","institution":"Guizhou University","correspondingAuthor":true,"prefix":"","firstName":"Shugang","middleName":"","lastName":"Hui","suffix":""}],"badges":[],"createdAt":"2025-09-02 06:23:17","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7513928/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7513928/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10142-025-01769-w","type":"published","date":"2025-12-22T15:57:22+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":93959808,"identity":"6cc47396-3692-4240-8d4b-6df20845b694","added_by":"auto","created_at":"2025-10-20 16:52:18","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":8000483,"visible":true,"origin":"","legend":"","description":"","filename":"LiYXmaniscript.docx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9cc416d3ba1bdfd6599b8c67.docx"},{"id":93960163,"identity":"405cbf39-6600-4b8f-8b20-367d747bfdd5","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":157195,"visible":true,"origin":"","legend":"","description":"","filename":"LiYXTable.docx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/18d33d588fdb84b9dc775814.docx"},{"id":93960864,"identity":"e06baa21-7459-4e20-ac1f-b742a678351a","added_by":"auto","created_at":"2025-10-20 17:00:41","extension":"json","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":9666,"visible":true,"origin":"","legend":"","description":"","filename":"0f2e3c67b2ca481496e8fc80d80d0273.json","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/465ae5180873fcdcc8d9438a.json"},{"id":93959918,"identity":"da30fec2-bf1c-4abc-8a8a-4f9d63ad9eae","added_by":"auto","created_at":"2025-10-20 16:52:25","extension":"xlsx","order_by":4,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":28307,"visible":true,"origin":"","legend":"","description":"","filename":"TableS1.TranscriptomedataofSpWRKYsunderCdtreatment.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/da49f69a43443713ab675b05.xlsx"},{"id":93961236,"identity":"fabcd294-160c-450b-98ac-16a67ec8e963","added_by":"auto","created_at":"2025-10-20 17:08:38","extension":"xlsx","order_by":5,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":19528,"visible":true,"origin":"","legend":"","description":"","filename":"TableS2.TranscriptomedataofSpWRKYsunderCutreatment.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/338e4bdde14e522b54dd1569.xlsx"},{"id":93960136,"identity":"995c1038-fde6-4ca8-a137-ff679939dc11","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"xlsx","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":17381,"visible":true,"origin":"","legend":"","description":"","filename":"TableS3.TranscriptomedataofSpWRKYsundersaltstress.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/eecc4a10b1d762ebf0ce436e.xlsx"},{"id":93959997,"identity":"791c7b61-5fee-48d5-bbeb-0de416daf031","added_by":"auto","created_at":"2025-10-20 16:52:30","extension":"xml","order_by":7,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":283996,"visible":true,"origin":"","legend":"","description":"","filename":"0f2e3c67b2ca481496e8fc80d80d02731enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4624db46433b8c22093db84c.xml"},{"id":93960026,"identity":"2c396834-181b-4237-9dfb-b9c95f83201f","added_by":"auto","created_at":"2025-10-20 16:52:32","extension":"pptx","order_by":8,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":16150726,"visible":true,"origin":"","legend":"","description":"","filename":"LiYXFigures.pptx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/152916d10a535747c1a01778.pptx"},{"id":93960116,"identity":"9f7e01ad-0cda-412a-883a-0e8199383508","added_by":"auto","created_at":"2025-10-20 16:52:39","extension":"jpeg","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1091843,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/8823512a68dc854d69197c47.jpeg"},{"id":93960058,"identity":"94483dfc-28a4-4efe-9477-a00338afa828","added_by":"auto","created_at":"2025-10-20 16:52:36","extension":"jpeg","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/eda11e6293219f715e3bfdfc.jpeg"},{"id":93959909,"identity":"225007cf-0348-41fb-8bcf-9338a7b9ad22","added_by":"auto","created_at":"2025-10-20 16:52:24","extension":"jpeg","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/900a2ce3e5507042e66bc007.jpeg"},{"id":93959939,"identity":"786cf5d2-b369-47a8-b084-c7460b8b9ba8","added_by":"auto","created_at":"2025-10-20 16:52:27","extension":"jpeg","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4ebe75ac92b98b39b66cb84c.jpeg"},{"id":93960027,"identity":"810b8854-a13a-4196-9af9-3c9fde56bea4","added_by":"auto","created_at":"2025-10-20 16:52:33","extension":"jpeg","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1622113,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage13.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/96e093690698fb96a79d5129.jpeg"},{"id":93960053,"identity":"f6027d46-50e6-4018-90b6-6ec7a77443bc","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1196,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage14.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c4125807bf7faef2503c9d2b.png"},{"id":93960067,"identity":"5faa02b9-775b-4f96-adce-781cbe6fa29c","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1162,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage15.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/672c27db99b29d4234473e19.png"},{"id":93959805,"identity":"0d009798-d9e5-4232-ab74-a94b379b09ce","added_by":"auto","created_at":"2025-10-20 16:52:18","extension":"png","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1178,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage16.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/3db1988aebf88eb73339ea6f.png"},{"id":93960005,"identity":"c234e574-16eb-42a5-b249-0e5aafd55a1c","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"png","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1224,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage17.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/750f438940e9728ab0564fb4.png"},{"id":93960846,"identity":"c332ea11-a83e-48b8-94d1-c4b883a429e0","added_by":"auto","created_at":"2025-10-20 17:00:32","extension":"png","order_by":18,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1407,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage18.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/873dc292e1aa08aca64f98a6.png"},{"id":93960217,"identity":"718fb756-f83b-43df-b63f-e571fd1b638a","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":19,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1154,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage19.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e1b2200d713eb049ede99a4f.png"},{"id":93960140,"identity":"22d8efcd-0c07-41fd-b7c8-b94e4b118ef5","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"jpeg","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/32c5159eb2f09ca9d412a651.jpeg"},{"id":93960215,"identity":"20c85bbe-9b0d-48b1-83e4-a2d115ebbeab","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":21,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1510,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage20.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c53e0410cb834a4933ae1c67.png"},{"id":93960118,"identity":"ecbf684c-31be-4de1-9b8c-e46dc40ce1d5","added_by":"auto","created_at":"2025-10-20 16:52:39","extension":"png","order_by":22,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1445,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage21.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/bb4cfc0d0251bac99afc0525.png"},{"id":93960841,"identity":"94091cae-9090-4d1b-a17d-82e325bf944e","added_by":"auto","created_at":"2025-10-20 17:00:28","extension":"png","order_by":23,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1789,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage22.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4a71e3eeeed1c7643e77294f.png"},{"id":93961240,"identity":"85ba4d06-760e-47e5-b47c-5e48fc49b586","added_by":"auto","created_at":"2025-10-20 17:08:54","extension":"png","order_by":24,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1154,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage23.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/2d6894af6675f703fc796efa.png"},{"id":93960870,"identity":"432623ce-deb2-4c1a-959d-d87968c80f8d","added_by":"auto","created_at":"2025-10-20 17:00:42","extension":"png","order_by":25,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1233,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage24.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/7fd97a7b5dd1f662264d94c4.png"},{"id":93960132,"identity":"4594ed52-9e72-4d1f-8e1b-d20bbbc8e740","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"png","order_by":26,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1403,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage25.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/eac1d347e95c091dcca94c64.png"},{"id":93959828,"identity":"dcd3fec4-bf68-4a73-ad8e-d804edd266fc","added_by":"auto","created_at":"2025-10-20 16:52:22","extension":"png","order_by":27,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1506,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage26.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f798c9ca7d2b8b8ec1d5e447.png"},{"id":93961237,"identity":"a5fdd39c-477f-4213-b810-f4924e4ada5d","added_by":"auto","created_at":"2025-10-20 17:08:43","extension":"png","order_by":28,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1188,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage27.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/cca74bd8fedadf61b6c22b32.png"},{"id":93960069,"identity":"c85eec0c-dc1b-4e06-8964-0ee081b84c53","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":29,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1675,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage28.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/0b787b8d37c6286805fe0abe.png"},{"id":93960061,"identity":"dfe3f609-79a0-4010-bf66-8b9084158dc7","added_by":"auto","created_at":"2025-10-20 16:52:36","extension":"png","order_by":30,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1210,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage29.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b943df370ba2213336e0d9b7.png"},{"id":93959958,"identity":"5f79f195-70b3-46a7-9108-f581c86e12d8","added_by":"auto","created_at":"2025-10-20 16:52:28","extension":"jpeg","order_by":31,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":531890,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a0a64121cf73b74d6d2c5c30.jpeg"},{"id":93960101,"identity":"4c282ae6-9df4-4e51-b395-97ab62a16567","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":32,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1269,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage30.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/126c7774735758f36708f6ec.png"},{"id":93959917,"identity":"979fe9c7-35f4-4449-827b-b5d4e170b7bd","added_by":"auto","created_at":"2025-10-20 16:52:25","extension":"png","order_by":33,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1233,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage31.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f1ab83d8bca7796fbc501d66.png"},{"id":93961239,"identity":"e061e371-ab0b-43df-9043-aa3da13eee8b","added_by":"auto","created_at":"2025-10-20 17:08:51","extension":"png","order_by":34,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1200,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage32.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c3de9fbf72b54546f02ef2e5.png"},{"id":93960847,"identity":"1b6d970d-0f1b-4419-8925-29dc3fae7e85","added_by":"auto","created_at":"2025-10-20 17:00:33","extension":"png","order_by":35,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1408,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage33.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c6643e168737626650aac977.png"},{"id":93960843,"identity":"bd91eace-5482-4436-a4ec-e55eccb486b4","added_by":"auto","created_at":"2025-10-20 17:00:30","extension":"png","order_by":36,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1481,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage34.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4bc1318673768cf01c2fae0e.png"},{"id":93960897,"identity":"0b2ae5f3-ba60-4298-8197-3ba833926088","added_by":"auto","created_at":"2025-10-20 17:00:44","extension":"png","order_by":37,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1506,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage35.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9f73a6168a5685c7c016054b.png"},{"id":93960148,"identity":"c7d0a3f2-f320-44ef-87c6-e76c52d8f0fe","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"png","order_by":38,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1505,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage36.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/fc8393b948dc301edf1e589d.png"},{"id":93960117,"identity":"9c0d43b5-5f4b-47ed-ab5d-157577b2a2cc","added_by":"auto","created_at":"2025-10-20 16:52:39","extension":"png","order_by":39,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1361,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage37.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c46c75f87f2a0579896ccd9f.png"},{"id":93960859,"identity":"86a2f6e0-db09-4636-b4df-d635605fc463","added_by":"auto","created_at":"2025-10-20 17:00:39","extension":"png","order_by":40,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1392,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage38.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/cfd60d96bfc1678231018fe5.png"},{"id":93959991,"identity":"6b15cbc2-3b04-4ef3-bb0c-983bd01ea33d","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"png","order_by":41,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1273,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage39.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b78524d2e36587039da6edac.png"},{"id":93959928,"identity":"61b9e7bd-99f8-4e6c-a561-3d5ce0d53c34","added_by":"auto","created_at":"2025-10-20 16:52:26","extension":"jpeg","order_by":42,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/87921cc5d167337e5a46b57b.jpeg"},{"id":93960057,"identity":"2fd4d111-300f-4656-9bac-47960330cc67","added_by":"auto","created_at":"2025-10-20 16:52:36","extension":"png","order_by":43,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1278,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage40.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/fd72bd5c7d0c6b5c85acff88.png"},{"id":93960006,"identity":"0cef08d3-cd21-4c6b-bf29-007ed9a62b58","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"png","order_by":44,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1202,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage41.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/db4239e3cdac6241dcafcb5a.png"},{"id":93959693,"identity":"7df32f16-9fc5-4079-89d1-de35d8faf38a","added_by":"auto","created_at":"2025-10-20 16:52:10","extension":"png","order_by":45,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1300,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage42.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c6f73b4c0387d3e97d6b902f.png"},{"id":93960839,"identity":"7969535e-5d35-4072-8d16-bcc3dcfb2cda","added_by":"auto","created_at":"2025-10-20 17:00:28","extension":"png","order_by":46,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1082,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage43.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c5c00e5f26887bc59a80acb9.png"},{"id":93960899,"identity":"90f93fef-c4e0-4385-b52b-c72a3a83444b","added_by":"auto","created_at":"2025-10-20 17:00:46","extension":"png","order_by":47,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1236,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage44.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a9010d821bf6c0eb4e124c60.png"},{"id":93960039,"identity":"d2294351-2c33-4a3a-b749-3b994c85e365","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":48,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1342,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage45.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/86d5a8c64a6438cfc78e803f.png"},{"id":93960150,"identity":"fd76874a-8346-4a25-8c1d-fa7b6d8efbb7","added_by":"auto","created_at":"2025-10-20 16:52:42","extension":"png","order_by":49,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1376,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage46.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/7a7c3af219085ac45e9fc02c.png"},{"id":93960060,"identity":"9e83ffb0-4a63-4144-a0a9-fe14e10c768a","added_by":"auto","created_at":"2025-10-20 16:52:36","extension":"png","order_by":50,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1280,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage47.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/27f382c0ed94746c12c6fbdb.png"},{"id":93960022,"identity":"0279efa5-4a7e-4fba-a2c4-f7447ac637e3","added_by":"auto","created_at":"2025-10-20 16:52:32","extension":"png","order_by":51,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1199,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage48.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/43120211d8281394a28351aa.png"},{"id":93960076,"identity":"1c04f849-0dc5-4769-a23e-705bce0bbcf3","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":52,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1284,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage49.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/073ce84ecd527b4f9de041f2.png"},{"id":93960042,"identity":"e0efedde-4c17-4fbc-ae9a-47091e4fbd70","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"jpeg","order_by":53,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1146604,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a09eb9ac2025b8e6baa924d9.jpeg"},{"id":93959881,"identity":"1bb5836e-5bee-4997-90bf-86635a2345a7","added_by":"auto","created_at":"2025-10-20 16:52:24","extension":"png","order_by":54,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1333,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage50.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/62b42782d083a38a6672c16f.png"},{"id":93960143,"identity":"3854c5aa-50c5-463b-93cd-ea45fb8b9db8","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"png","order_by":55,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1261,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage51.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/569452fc65899b1155997ad0.png"},{"id":93960001,"identity":"578419d8-9094-44b3-bbda-2b2802d57bd2","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"png","order_by":56,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1213,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage52.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/faae4d243fbe1b76c48e542b.png"},{"id":93960030,"identity":"003b7aec-5cf5-44ce-a36e-4344b85d9552","added_by":"auto","created_at":"2025-10-20 16:52:34","extension":"png","order_by":57,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1170,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage53.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e97d9020396a6cc2629127c8.png"},{"id":93960102,"identity":"3bdf0cbb-06da-4a2c-af34-a17b8fca2155","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":58,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1489,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage54.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a601f48763bf6bb53e1dae7b.png"},{"id":93960121,"identity":"f52ee2db-4f6d-4375-853d-60f845715361","added_by":"auto","created_at":"2025-10-20 16:52:39","extension":"png","order_by":59,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1246,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage55.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a68bae719c4533c17654d761.png"},{"id":93960209,"identity":"bc912f51-beb5-4fbd-bacc-7192ccf5ed3a","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":60,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1210,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage56.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4b206510c70b6422d8f98b66.png"},{"id":93960874,"identity":"3260fe74-4a0a-4082-8a40-3f47f8cc405c","added_by":"auto","created_at":"2025-10-20 17:00:43","extension":"png","order_by":61,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1441,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage57.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f78325043cf69c680551f837.png"},{"id":93959719,"identity":"9d2c1273-e3de-4d8c-a595-e08ecaab5348","added_by":"auto","created_at":"2025-10-20 16:52:12","extension":"png","order_by":62,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1227,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage58.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/456fc303c860bbf92d6e4082.png"},{"id":93960018,"identity":"9c8437db-e619-4345-b1cf-d46214338016","added_by":"auto","created_at":"2025-10-20 16:52:32","extension":"png","order_by":63,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1253,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage59.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4466d73060474bfa6c4f9461.png"},{"id":93960110,"identity":"58743f1f-4b46-4406-acc1-d72e7a043a77","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"jpeg","order_by":64,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage10.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/0ad4c0cb7e43ebc63db2ffe1.jpeg"},{"id":93960860,"identity":"e5771706-8bec-4578-b8c7-93f1839a2a4a","added_by":"auto","created_at":"2025-10-20 17:00:39","extension":"png","order_by":65,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1291,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage60.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/ab18bb9ec9c24f9e8bed1872.png"},{"id":93960228,"identity":"da9f119c-ca3d-45e7-861d-5f705a3158cd","added_by":"auto","created_at":"2025-10-20 16:52:46","extension":"png","order_by":66,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1221,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage61.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/74c6b0a3f445a503b702c7bc.png"},{"id":93960070,"identity":"c03bc410-fab4-45ba-abfa-39c44e64a57f","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":67,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1329,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage62.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/99500ee109ae450188abda7a.png"},{"id":93961241,"identity":"b873b21a-9f78-46c5-a35e-de6bcc51949b","added_by":"auto","created_at":"2025-10-20 17:08:56","extension":"png","order_by":68,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1460,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage63.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9d745d7f2e78e73ac5e904e2.png"},{"id":93960220,"identity":"7efe8922-bd29-4e00-b112-262655f31673","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":69,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1219,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage64.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/957e18ed32ac63a75c3fb9ea.png"},{"id":93959923,"identity":"9b103b78-1231-428d-96c9-fee0cfa5efc8","added_by":"auto","created_at":"2025-10-20 16:52:26","extension":"png","order_by":70,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1269,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage65.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a50ae50371f4f8ce770b28b0.png"},{"id":93960851,"identity":"39485ae9-09b5-40bf-947c-5f33d1f4fff8","added_by":"auto","created_at":"2025-10-20 17:00:38","extension":"png","order_by":71,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1214,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage66.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/75a4da1820246c4e93e63fe7.png"},{"id":93959943,"identity":"02663476-1e04-4439-a318-3dac0af8e79d","added_by":"auto","created_at":"2025-10-20 16:52:27","extension":"png","order_by":72,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1276,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage67.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9da376423a1889f67abf8c3d.png"},{"id":93960207,"identity":"c4d96720-2fa0-432a-ae3a-1be81dcc1198","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":73,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1403,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage68.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f66ff4f3bfac6f088d7540b1.png"},{"id":93960865,"identity":"57d05d53-bca1-43e0-bde9-8da3b8136784","added_by":"auto","created_at":"2025-10-20 17:00:41","extension":"png","order_by":74,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1479,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage69.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c6f194274efcd8b4f75c66db.png"},{"id":93960007,"identity":"8e862fc8-2c3a-4b05-bbb9-268e63692991","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"jpeg","order_by":75,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":499276,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage7.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/0c8944698c06acd6a7b454e7.jpeg"},{"id":93960062,"identity":"714e2581-195c-4964-8da0-b11d5d584233","added_by":"auto","created_at":"2025-10-20 16:52:36","extension":"png","order_by":76,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1285,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage70.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/c64f407a1ebba0670e03ae6c.png"},{"id":93959818,"identity":"8abca9a3-169a-4b21-a1b4-962b11ff7f39","added_by":"auto","created_at":"2025-10-20 16:52:19","extension":"png","order_by":77,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1140,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage71.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/22e3aeda98fc69c4ca5b9c12.png"},{"id":93961238,"identity":"1f9b27e0-6254-49ed-8680-38723e269efa","added_by":"auto","created_at":"2025-10-20 17:08:43","extension":"png","order_by":78,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1304,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage72.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/0a9c8d3d1e48e9ef1a078828.png"},{"id":93960112,"identity":"b547de0e-42d5-4d40-8027-333a5cf1ce05","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":79,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1285,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage70.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/538bd7fcf3f532c09816e120.png"},{"id":93959945,"identity":"404d35dd-2278-46b4-ae1f-659f4f1b33c3","added_by":"auto","created_at":"2025-10-20 16:52:28","extension":"png","order_by":80,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1060,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage74.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/df79657eaddb0a67c50f94c6.png"},{"id":93959996,"identity":"95c9f086-4032-4642-aba1-939ed5a1712b","added_by":"auto","created_at":"2025-10-20 16:52:30","extension":"png","order_by":81,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1405,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage75.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f451513da8917923c17ac559.png"},{"id":93960869,"identity":"6e77267b-e9e5-4bfb-986a-a3f6fa6e021c","added_by":"auto","created_at":"2025-10-20 17:00:42","extension":"png","order_by":82,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1199,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage76.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/1fe0427447b882da385ec3a5.png"},{"id":93960129,"identity":"6a51ca9b-56db-4890-9184-977839243d61","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"png","order_by":83,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1308,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage77.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/be5efcd164432b338018fe59.png"},{"id":93960863,"identity":"ea2e05d0-7f14-4686-9e6f-e40009e9096a","added_by":"auto","created_at":"2025-10-20 17:00:41","extension":"png","order_by":84,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1216,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage78.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/8ce320d7758d1de8852a4f15.png"},{"id":93960032,"identity":"a44b1124-b57e-4e15-bd6a-8cbd7ead5e3e","added_by":"auto","created_at":"2025-10-20 16:52:34","extension":"png","order_by":85,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1312,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage79.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e418960ad832d2f4c0e2a994.png"},{"id":93960011,"identity":"96e87a3a-7989-4bfb-b49f-3c965db8aef8","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"jpeg","order_by":86,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1503880,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage8.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e00dbe117f5626beb7753b61.jpeg"},{"id":93960064,"identity":"d1274e7a-6b2d-4925-95e8-75355d649bbb","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":87,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1298,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage80.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b1099a094e0fb4de4bd7516c.png"},{"id":93960130,"identity":"89e0235f-4b6d-4ffe-b990-b11bd6b9a6c1","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"png","order_by":88,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1315,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage81.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/1d4aee0338cca3d9b30b77ba.png"},{"id":93960025,"identity":"e207ba01-7209-43f5-a5cb-3edef71b4ae4","added_by":"auto","created_at":"2025-10-20 16:52:32","extension":"jpeg","order_by":89,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":2108305,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage9.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/fd83159e7d32adb09249283d.jpeg"},{"id":93960159,"identity":"d1ebe1e3-0ff1-484a-b0d9-1ef0a2175991","added_by":"auto","created_at":"2025-10-20 16:52:42","extension":"png","order_by":90,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":236198,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/21153ba8f0105e14b692a870.png"},{"id":93960844,"identity":"25d44a5c-d32e-4dbb-b95b-b64c137e135f","added_by":"auto","created_at":"2025-10-20 17:00:31","extension":"png","order_by":91,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage10.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e3e66e147cb9409b4124ba7e.png"},{"id":93960063,"identity":"6bb09cf6-eb1a-4a7a-9d74-c725a0f78e23","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":92,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage10.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f779be41f49efe3a01da097f.png"},{"id":93960113,"identity":"011d18ea-ff06-4acc-98ff-208147cd446b","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":93,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage10.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/cb74e1750ec7dd36c5db7c6c.png"},{"id":93960019,"identity":"99ea4cab-d1f9-4226-8ecb-f88479a16580","added_by":"auto","created_at":"2025-10-20 16:52:32","extension":"png","order_by":94,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":364182,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage13.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/3ee4f66a9bc0aa038174bc7d.png"},{"id":93960223,"identity":"4d91d2af-a009-49db-8103-6c644c25c067","added_by":"auto","created_at":"2025-10-20 16:52:46","extension":"png","order_by":95,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1173,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage14.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9a72af8f8e3976882b9ff2e5.png"},{"id":93960071,"identity":"2e57e5b2-35c3-4784-a57c-84c8e82c6ab7","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":96,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1177,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage15.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/8dc1eb3d87435d80141b22fc.png"},{"id":93960124,"identity":"431c2a76-6782-45f7-b01e-d44e0f73c156","added_by":"auto","created_at":"2025-10-20 16:52:39","extension":"png","order_by":97,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1085,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage16.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f1858a7a222e08463b4fb99a.png"},{"id":93960836,"identity":"3586cb7e-7a8d-40e6-b3f1-4260d55d7afd","added_by":"auto","created_at":"2025-10-20 17:00:26","extension":"png","order_by":98,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1070,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage17.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/3368042a207136a788673cbb.png"},{"id":93959936,"identity":"53856039-359b-4590-a9d2-8c9015697481","added_by":"auto","created_at":"2025-10-20 16:52:27","extension":"png","order_by":99,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1134,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage18.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/870d091bd3b12969ac064ba7.png"},{"id":93960045,"identity":"87af2b1e-c50f-472e-a555-47427ca540f8","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":100,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1039,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage19.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/1f082dd37b8ebc7ad234d958.png"},{"id":93960047,"identity":"79bcf2ae-0994-4e59-99c5-08acb6f37d69","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":101,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage10.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/ee537906329e3e6da67d836b.png"},{"id":93959995,"identity":"9c81d6ea-bce0-47b0-82a1-cced019d3550","added_by":"auto","created_at":"2025-10-20 16:52:30","extension":"png","order_by":102,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1408,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage20.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/25c8ff3b57d0f337ae2302c2.png"},{"id":93960049,"identity":"d0c02ea7-0883-40c6-b041-62a37357c679","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":103,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1213,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage21.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/03f99b13565f16bcb3c37b2b.png"},{"id":93960205,"identity":"d53e1f88-01c5-47df-a90f-6b74ab0cfb7e","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":104,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1420,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage22.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/84d2ce120250a4d27041cd99.png"},{"id":93959927,"identity":"5d3acde7-b864-4223-84e2-eea68b9940dc","added_by":"auto","created_at":"2025-10-20 16:52:26","extension":"png","order_by":105,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1036,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage23.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/3c57c23805a8181d73c6b99c.png"},{"id":93960107,"identity":"454cf3dc-a3b4-477d-9479-b643d4c8ed1f","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":106,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1088,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage24.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/10947bc66316cecc7235483b.png"},{"id":93959988,"identity":"4aae4721-d30c-445e-a170-69771d9d6ba2","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"png","order_by":107,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1455,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage25.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/bd428491a9a66a35cb3c8e1d.png"},{"id":93960125,"identity":"73fedefa-4470-4608-8417-3a76096a32f0","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"png","order_by":108,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1145,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage26.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/cdf2e71b8adec327a3fda57c.png"},{"id":93960174,"identity":"54a39b68-6488-41e8-9547-e650fa97157b","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"png","order_by":109,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1180,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage27.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/cad70e250f4893cf6ce2e25e.png"},{"id":93960203,"identity":"eec0bf49-2359-4dcb-b4bb-9376ca6bc0d2","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":110,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1546,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage28.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/bbc76310199dd2a015fb9705.png"},{"id":93959979,"identity":"8918a1b1-7c31-4ae0-b336-48ca4ec421c2","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"png","order_by":111,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1141,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage29.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/dc56de7628e8e2e6f0303d34.png"},{"id":93960868,"identity":"e0f05449-1d6f-4583-8be8-b452ebd50010","added_by":"auto","created_at":"2025-10-20 17:00:42","extension":"png","order_by":112,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":153786,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/fe0224c4a712ba1fd8b5dc00.png"},{"id":93960199,"identity":"93962204-de76-4dcb-8bed-667e7a272f25","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"png","order_by":113,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1318,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage30.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4cf9a59a3fd44e44b0ecf398.png"},{"id":93960041,"identity":"cae8544f-feaf-4182-a529-5117e26e75f9","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":114,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1112,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage31.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/8576690a7b86d549193930b0.png"},{"id":93959911,"identity":"d485be16-8fc3-4376-add7-4181128672d4","added_by":"auto","created_at":"2025-10-20 16:52:25","extension":"png","order_by":115,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1145,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage32.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b3cfb734ad2f5c6384edc4e4.png"},{"id":93960202,"identity":"35fed9b7-accb-4dd7-9744-4098e7a4c513","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"png","order_by":116,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1202,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage33.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/8acf0e306270f869f2c3643a.png"},{"id":93959924,"identity":"c09f1783-15a1-4633-a7a6-239de7d406bf","added_by":"auto","created_at":"2025-10-20 16:52:26","extension":"png","order_by":117,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1320,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage34.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/2ff4864e14b5edc9d28f261d.png"},{"id":93960009,"identity":"79d48ca4-b490-4904-93e0-45a805069907","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"png","order_by":118,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1447,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage35.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/a5908065d23f9db1e801f3b1.png"},{"id":93960097,"identity":"6c766881-85f5-406a-b9ed-265453c9667b","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":119,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1430,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage36.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/d852be6f54640e5dff84e3f2.png"},{"id":93960158,"identity":"d488e94e-b101-4f43-bad4-97359e19ffdd","added_by":"auto","created_at":"2025-10-20 16:52:42","extension":"png","order_by":120,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1409,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage37.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b625bce411623908b8b5ad34.png"},{"id":93960214,"identity":"656bd8fc-d39e-4929-8fa5-075e04d957c5","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":121,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1333,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage38.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/15440532ba2834f990a52868.png"},{"id":93960126,"identity":"f016cda5-54d8-45d8-86a5-b2bbc22416dc","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"png","order_by":122,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1229,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage39.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/611bd1115db0e6d8fab12e64.png"},{"id":93960044,"identity":"0d3deeed-a881-4e0f-8729-c7d05b687a08","added_by":"auto","created_at":"2025-10-20 16:52:35","extension":"png","order_by":123,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b9549ece04416781431b470e.png"},{"id":93960213,"identity":"267d9dbe-8b91-445e-bb21-de265b9d8d63","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":124,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1250,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage40.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/69a0d86b32330b66ded1391f.png"},{"id":93960138,"identity":"8d5132ab-a599-4e13-a9ca-7e1fe8412349","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"png","order_by":125,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1152,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage41.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/6a2c5e60fe2dff266999b9b5.png"},{"id":93960858,"identity":"d625affd-bb07-4777-8988-f53262517410","added_by":"auto","created_at":"2025-10-20 17:00:39","extension":"png","order_by":126,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1251,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage42.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/15af6cc6e481b7722ec861df.png"},{"id":93960856,"identity":"5ae3bfc1-ad0c-42a5-ab89-d274b5c49a25","added_by":"auto","created_at":"2025-10-20 17:00:39","extension":"png","order_by":127,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":951,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage43.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/17fc81d89c7d8bce096c5134.png"},{"id":93959709,"identity":"065ac885-7458-4805-98b2-4e45c8d371bd","added_by":"auto","created_at":"2025-10-20 16:52:11","extension":"png","order_by":128,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1063,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage44.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/7c4a41ecc462664761343903.png"},{"id":93960153,"identity":"12275038-9caf-452f-ab90-bf47c3793baf","added_by":"auto","created_at":"2025-10-20 16:52:42","extension":"png","order_by":129,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1155,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage45.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/af9065d7020e14f1263b8ab3.png"},{"id":93960000,"identity":"3abe33c5-0c77-4b68-8889-389f61ddc5ea","added_by":"auto","created_at":"2025-10-20 16:52:31","extension":"png","order_by":130,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1291,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage46.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/74c6dd6e7e74138c4a7f0487.png"},{"id":93960838,"identity":"5f466b03-70c8-4f25-acb8-b8c7badb102c","added_by":"auto","created_at":"2025-10-20 17:00:28","extension":"png","order_by":131,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1220,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage47.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/82bab0e273a1c37f67a0c2c0.png"},{"id":93959993,"identity":"0547319a-5d73-4cf1-8ee5-b23f026da44e","added_by":"auto","created_at":"2025-10-20 16:52:30","extension":"png","order_by":132,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1192,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage48.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9679d0ae4ba75f7f540e108f.png"},{"id":93960204,"identity":"dd960578-d8eb-4aed-9cb9-14950c604144","added_by":"auto","created_at":"2025-10-20 16:52:44","extension":"png","order_by":133,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1179,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage49.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/96c30bba163e848396550076.png"},{"id":93960231,"identity":"a67389b2-e061-419a-a240-1e4abf561147","added_by":"auto","created_at":"2025-10-20 16:52:47","extension":"png","order_by":134,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":249425,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/19aa56eaaaedfb5f5fa85e8c.png"},{"id":93960850,"identity":"69901313-f205-4360-b8df-446b2ddcb722","added_by":"auto","created_at":"2025-10-20 17:00:36","extension":"png","order_by":135,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1236,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage50.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/23cc0edf2a529dd66d5ee8dc.png"},{"id":93959825,"identity":"6d9c05b8-03ff-470e-b321-dea22fd6c98a","added_by":"auto","created_at":"2025-10-20 16:52:22","extension":"png","order_by":136,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1181,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage51.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/da4a30de876eca656996df54.png"},{"id":93960862,"identity":"1ae768b4-5257-4436-89af-44791cba7960","added_by":"auto","created_at":"2025-10-20 17:00:40","extension":"png","order_by":137,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1167,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage52.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/1e28741fa45439202683b4b8.png"},{"id":93959983,"identity":"ce3b6177-0372-40f3-b540-448ce05a599f","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"png","order_by":138,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1175,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage53.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/92a06925c04e099524dff8a8.png"},{"id":93960145,"identity":"5e95607a-2cf3-4b56-a00a-75343ea5c7d0","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"png","order_by":139,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1434,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage54.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/840872ddd079ccbe2b7a2f4e.png"},{"id":93960872,"identity":"795c324b-2a36-486d-974f-541ba7d6fdb3","added_by":"auto","created_at":"2025-10-20 17:00:42","extension":"png","order_by":140,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1154,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage55.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/9b622c21c7c0b58b8fc56b30.png"},{"id":93960198,"identity":"0035b815-0d9c-4c70-bd95-f53d3a0ed1e1","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"png","order_by":141,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1135,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage56.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/296366b022dd0bd9c342eb45.png"},{"id":93960095,"identity":"f3f53e96-98f8-4c17-86ef-ee441da9020e","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":142,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1359,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage57.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/02c18aef1b2cc10db0d32fc7.png"},{"id":93960854,"identity":"32e05c15-195f-42f5-918a-1f0f0d595624","added_by":"auto","created_at":"2025-10-20 17:00:38","extension":"png","order_by":143,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1194,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage58.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/0103a97307e911f843c4b3af.png"},{"id":93960149,"identity":"05aaa3a5-9d3e-4dfc-b1dd-17a423bc949c","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"png","order_by":144,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1136,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage59.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/47a1f1e8762c2c3a82072da2.png"},{"id":93960068,"identity":"98ec1d95-61ee-4e37-83ad-b5a1326efa12","added_by":"auto","created_at":"2025-10-20 16:52:37","extension":"png","order_by":145,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/1135958445c7679f7256d36d.png"},{"id":93959722,"identity":"60176f72-3297-4a10-8135-2c23283e6eb3","added_by":"auto","created_at":"2025-10-20 16:52:12","extension":"png","order_by":146,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1334,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage60.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/cd0c114c553ca83e76b35691.png"},{"id":93960896,"identity":"fb6b0a91-80e5-43d9-bc10-257e3552d0ba","added_by":"auto","created_at":"2025-10-20 17:00:44","extension":"png","order_by":147,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1090,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage61.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/2cf8522b44d7b871c86ec5b1.png"},{"id":93960857,"identity":"3658533f-434d-41d9-a002-8c60aa1e39aa","added_by":"auto","created_at":"2025-10-20 17:00:39","extension":"png","order_by":148,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1217,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage62.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/5af30d60be087e20bf45a46c.png"},{"id":93960227,"identity":"4cdb2367-3dce-4f1f-a86e-1d8c894ff670","added_by":"auto","created_at":"2025-10-20 16:52:46","extension":"png","order_by":149,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1350,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage63.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/37c13b6baace3a2684958a96.png"},{"id":93959990,"identity":"d96ebd99-e5fd-471a-9c97-6c38a44e5c82","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"png","order_by":150,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1391,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage64.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b5031980d26cab813a394642.png"},{"id":93960216,"identity":"b3eec231-916a-4a67-8505-2a9d695db860","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":151,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1141,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage65.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/d6d4446670ec4b83b27d376a.png"},{"id":93960114,"identity":"35e2dc30-ceac-4660-92c2-4225d8d7f2be","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":152,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1148,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage66.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/02b27abb6e9a7e71ac503a5d.png"},{"id":93960160,"identity":"f2bb481b-8496-4031-8caa-eb7759da157b","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"png","order_by":153,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1155,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage67.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/eac7e21a6095c57305ddb7c6.png"},{"id":93960875,"identity":"350f60a0-bfd3-40c6-9547-c1e579ffe06f","added_by":"auto","created_at":"2025-10-20 17:00:43","extension":"png","order_by":154,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1288,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage68.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/8007177a1d74681f62a7f96d.png"},{"id":93960898,"identity":"3d8718e1-c0ef-4754-a476-2d3f4bfee348","added_by":"auto","created_at":"2025-10-20 17:00:44","extension":"png","order_by":155,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1598,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage69.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/28dd77365f84911c8072303b.png"},{"id":93960161,"identity":"8fffe41d-7c53-45da-8243-26bff029bcde","added_by":"auto","created_at":"2025-10-20 16:52:43","extension":"png","order_by":156,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":158037,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/309f9011c7c050f514b5c626.png"},{"id":93959941,"identity":"e8d5f42d-30aa-41b4-a7d4-7b801b61d531","added_by":"auto","created_at":"2025-10-20 16:52:27","extension":"png","order_by":157,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1196,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage70.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e5e3e3cd8693a7af292749ef.png"},{"id":93960219,"identity":"c07dcb24-8acb-4556-a84b-ba3d5ceb2fad","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":158,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1071,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage71.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/fcc204bcccbb31b0b3c874f5.png"},{"id":93959819,"identity":"257a2684-7f78-48a1-a5d5-bc3d12dc15a1","added_by":"auto","created_at":"2025-10-20 16:52:19","extension":"png","order_by":159,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1147,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage72.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/166f978e6b6aed2933eacb29.png"},{"id":93960853,"identity":"5476dbe1-5d1e-4e8f-ad9f-b5aee6d9452e","added_by":"auto","created_at":"2025-10-20 17:00:38","extension":"png","order_by":160,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1196,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage73.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/740332f74a836d929a5177e0.png"},{"id":93960218,"identity":"54d0b86b-26f6-45b7-b6b3-4f89c9f8cf50","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":161,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":930,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage74.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/3429ae65d6f1c43b06b161a2.png"},{"id":93960845,"identity":"ad032874-79b9-4f41-8e36-45779de0f619","added_by":"auto","created_at":"2025-10-20 17:00:31","extension":"png","order_by":162,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1335,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage75.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/0e72a73f283d89d921719794.png"},{"id":93960239,"identity":"f5e539fa-111d-4872-b088-9e9b51149f04","added_by":"auto","created_at":"2025-10-20 16:52:47","extension":"png","order_by":163,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1173,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage76.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/098bfc620adc1b9d017d4873.png"},{"id":93960901,"identity":"2e72be64-6fd2-4b91-8373-f8a56bea86a4","added_by":"auto","created_at":"2025-10-20 17:00:48","extension":"png","order_by":164,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1165,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage77.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/29538ebf108d6265e111a4ba.png"},{"id":93960221,"identity":"9c506399-cec3-40a6-913c-6d0e9c7efad9","added_by":"auto","created_at":"2025-10-20 16:52:45","extension":"png","order_by":165,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1213,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage78.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/08cad6b625572efb47b923d1.png"},{"id":93960142,"identity":"83baa71e-a80e-436a-8993-340e40ac08ed","added_by":"auto","created_at":"2025-10-20 16:52:41","extension":"png","order_by":166,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1419,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage79.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/b7cd0256dfdb732575e4d287.png"},{"id":93961242,"identity":"1adbe669-7344-42c5-978d-a4f202b324df","added_by":"auto","created_at":"2025-10-20 17:08:57","extension":"png","order_by":167,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":159684,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage8.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/5e3e840612c82a7619ed7d66.png"},{"id":93960104,"identity":"edc6c8b6-6965-4f71-a167-4e2faa8cfab9","added_by":"auto","created_at":"2025-10-20 16:52:38","extension":"png","order_by":168,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1310,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage80.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/dd85e45cb24b443428ab880f.png"},{"id":93960152,"identity":"a04c2987-bcd6-4336-a5ac-d70866c05a1a","added_by":"auto","created_at":"2025-10-20 16:52:42","extension":"png","order_by":169,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1269,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage81.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/486aca6e01d776f93b28a94c.png"},{"id":93960849,"identity":"be993e28-26cc-46f5-8c29-04bf9f079d2c","added_by":"auto","created_at":"2025-10-20 17:00:36","extension":"png","order_by":170,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":320407,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage9.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/089901d6e78830c6f0709af1.png"},{"id":93960014,"identity":"7d5f9c47-45b5-4bb3-a0b4-19755e5d3528","added_by":"auto","created_at":"2025-10-20 16:52:32","extension":"xml","order_by":171,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":281823,"visible":true,"origin":"","legend":"","description":"","filename":"0f2e3c67b2ca481496e8fc80d80d02731structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/46d7058866d553a273e4a33d.xml"},{"id":93959985,"identity":"3d3426a2-d301-4959-ab67-2ae18e5f931a","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"html","order_by":172,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":305056,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e6c5ec3c1f99f4ae2ab61066.html"},{"id":99172493,"identity":"5f2eee0d-3b1c-4c23-8755-bfb22d999bcc","added_by":"auto","created_at":"2025-12-29 16:10:28","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2418329,"visible":true,"origin":"","legend":"","description":"","filename":"LiYXmaniscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1_covered_4f84c6f7-54aa-41a5-b813-327d61c38586.pdf"},{"id":93960131,"identity":"b5f0eb7c-ae63-469b-bc66-18b6b12e98de","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"png","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":268853,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S1 \u003c/strong\u003ePrediction of the tertiary structure of SpWRKYs.\u003c/p\u003e","description":"","filename":"S1.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/f82f1c1dd6dab406daa2e26b.png"},{"id":93959930,"identity":"86e13d44-e171-4011-82e7-b13741ae1d26","added_by":"auto","created_at":"2025-10-20 16:52:26","extension":"png","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":106062,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFig. S2 \u003c/strong\u003eThe conserved motifs in SpWRKYs.\u003c/p\u003e","description":"","filename":"S2.png","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/1a507cb0b7017635244c2ca2.png"},{"id":93960134,"identity":"581b997d-7778-4bf1-bfd8-5312d72c9a55","added_by":"auto","created_at":"2025-10-20 16:52:40","extension":"xlsx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":28307,"visible":true,"origin":"","legend":"","description":"","filename":"TableS1.TranscriptomedataofSpWRKYsunderCdtreatment.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/33dc220584d94c5c9607f379.xlsx"},{"id":93959594,"identity":"385c8965-5afa-485a-aeb3-e104b546487a","added_by":"auto","created_at":"2025-10-20 16:51:54","extension":"xlsx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":19528,"visible":true,"origin":"","legend":"","description":"","filename":"TableS2.TranscriptomedataofSpWRKYsunderCutreatment.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/4f18f20b98b7875c6a1fadac.xlsx"},{"id":93960123,"identity":"73a2c48f-4969-4529-8333-e0624b505fcf","added_by":"auto","created_at":"2025-10-20 16:52:39","extension":"xlsx","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":17381,"visible":true,"origin":"","legend":"","description":"","filename":"TableS3.TranscriptomedataofSpWRKYsundersaltstress.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/e6b7f94ea72e1d1efeab211d.xlsx"},{"id":93959980,"identity":"d05ebca4-b3de-4c86-8491-129a43f17379","added_by":"auto","created_at":"2025-10-20 16:52:29","extension":"docx","order_by":6,"title":"","display":"","copyAsset":false,"role":"supplement","size":157195,"visible":true,"origin":"","legend":"","description":"","filename":"LiYXTable.docx","url":"https://assets-eu.researchsquare.com/files/rs-7513928/v1/69874cf419166a1242b28246.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Genome-wide characterization of WRKYs and function of SpWRKY40 and SpWRKY50 in Sesuvium portulacastrum","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":"functional-and-integrative-genomics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"fige","sideBox":"Learn more about [Functional \u0026 Integrative Genomics](http://link.springer.com/journal/10142)","snPcode":"10142","submissionUrl":"https://submission.nature.com/new-submission/10142/3","title":"Functional \u0026 Integrative Genomics","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Sesuvium portulacastrum, WRKY gene family, Abiotic stress, SpWRKY40, SpWRKY51, Transcription factor","lastPublishedDoi":"10.21203/rs.3.rs-7513928/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7513928/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWRKYs represent a large family of plant transcription factors characterized by a highly conserved WRKY domain. WRKY transcription factors are important for plant growth, development, and responses to environmental stresses. However, this family has not been previously identified in \u003cem\u003eSesuvium portulacastrum\u003c/em\u003e. Here, we comprehensively identified 68 \u003cem\u003eSpWRKYs \u003c/em\u003efrom the genome of \u003cem\u003eS. portulacastrum\u003c/em\u003e by phylogenetic relationships, chromosomal localization, collinearity analysis, gene structure, duplication events, and evolutionary pressure based on nonsynonymous substitution rate/synonymous substitution rate ratios. Promoter analysis revealed the presence of multiple \u003cem\u003ecis\u003c/em\u003e-elements associated with abiotic stress signaling, developmental regulation, phytohormone responses, light responsiveness, and tissue-specific expression. We also characterized the physicochemical and structural properties of the \u003cem\u003eSpWRKY\u003c/em\u003e-encoded proteins, including protein property of primary structure, secondary structures, and predicted three-dimensional models, subcellular localization, conserved motifs, and conserved domains. Furthermore, transcriptome profiling under cadmium, copper, and salt stress conditions demonstrated differential expression patterns of \u003cem\u003eSpWRKYs\u003c/em\u003e. Among them, \u003cem\u003eSpWRKY40\u003c/em\u003e and \u003cem\u003eSpWRKY51\u003c/em\u003eshowed significant indued under cadmium treatment and were further validated by quantitative real-time PCR. Subcellular localization and yeast one-hybrid assays both supporting SpWRKY40 and SpWRKY51 play as transcription factors. Our study provides a detailed overview of the \u003cem\u003eSpWRKY\u003c/em\u003e family and their potential involvement in environment tolerance. The findings offer valuable candidate genes for future applications in improving stress tolerance in \u003cem\u003eS. portulacastrum\u003c/em\u003e and related crop species.\u003c/p\u003e","manuscriptTitle":"Genome-wide characterization of WRKYs and function of SpWRKY40 and SpWRKY50 in Sesuvium portulacastrum","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-20 16:49:16","doi":"10.21203/rs.3.rs-7513928/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-10-25T23:25:39+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-23T07:37:28+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-22T14:52:50+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-21T11:23:00+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-12T17:46:32+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"207224461759626345503117790165247915217","date":"2025-10-09T16:26:22+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-08T08:30:25+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"87931886210608153920482984096010852150","date":"2025-10-07T18:24:56+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"55511146401682045258903861883771352739","date":"2025-10-07T17:52:56+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"136357074266779303881072284600643207851","date":"2025-10-07T16:45:34+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"302210791085429452129409207026513724268","date":"2025-10-07T16:32:31+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"269427946102373298238006811883471910890","date":"2025-10-07T16:32:23+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"167188960943805044657717764599936502105","date":"2025-10-07T16:16:37+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"89704558132204253748411479907565784355","date":"2025-10-07T16:06:41+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-10-07T16:00:29+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-09-12T16:31:46+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-09-04T13:06:58+00:00","index":"","fulltext":""},{"type":"submitted","content":"Functional \u0026 Integrative Genomics","date":"2025-09-02T06:10:27+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"functional-and-integrative-genomics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"fige","sideBox":"Learn more about [Functional \u0026 Integrative Genomics](http://link.springer.com/journal/10142)","snPcode":"10142","submissionUrl":"https://submission.nature.com/new-submission/10142/3","title":"Functional \u0026 Integrative Genomics","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"3ca2b482-b1ef-4637-b939-00dae05ed7cd","owner":[],"postedDate":"October 20th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-12-29T16:06:12+00:00","versionOfRecord":{"articleIdentity":"rs-7513928","link":"https://doi.org/10.1007/s10142-025-01769-w","journal":{"identity":"functional-and-integrative-genomics","isVorOnly":false,"title":"Functional \u0026 Integrative Genomics"},"publishedOn":"2025-12-22 15:57:22","publishedOnDateReadable":"December 22nd, 2025"},"versionCreatedAt":"2025-10-20 16:49:16","video":"","vorDoi":"10.1007/s10142-025-01769-w","vorDoiUrl":"https://doi.org/10.1007/s10142-025-01769-w","workflowStages":[]},"version":"v1","identity":"rs-7513928","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7513928","identity":"rs-7513928","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}