Preliminary determination of early growth response of wheat to potassium iodate priming

Preliminary determination of early growth response of wheat to potassium iodate priming | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Preliminary determination of early growth response of wheat to potassium iodate priming Albert Thembinkosi Modi This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4626171/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 The objective of the study was to determine the efficacy of iodine priming in combination with water availability levels on the physiological attributes of wheat (Triticum aestivum ) as determined at stage Z14.7/22 of plant growth. Two levels of potassium iodate halopriming (100 mg l − 1 and 200 mg l − 1 ) were compared with distilled for priming seeds before planting to grow seedlings under conditions of controlled water availability determined by field capacity (FC). Adequate water availability (100% FC) throughout seedling growth was compared with declining water availability from planting to 25% FC before harvesting plants to determine biomass in terms of root: shoot ratio (RSR). Crop physiological response was determined using membrane stability index (MSI) and leaf chlorophyll and carotenoid contents. Results confirmed the positive role of halopriming wheat seeds to improve resistance to water stress at the early stage of crop development. The use of potassium iodate negligibly improved plant biomass accumulation, but leaf membrane stability index was significantly improved. These changes were associated with enhancement of chlorophyll (a, b) and carotenoid contents. This study provides insight into the potential effect of seed treatment on wheat response to soil water availability at the early stages of development. Biological sciences/Physiology Biological sciences/Plant sciences Carotenoids Chlorophyll Halopriming Water deficit Wheat Seed Figures Figure 1 Introduction Seed quality is multidimensional in that it requires understating of physiological responses to a variety of factors that affect seed hormones and protein response singularly or in combination. Seed germination is the best indicator of seed quality, but it is triggered by different environmental treatments during development and after physiological maturity, including storage. Hence, it is not only important to improve seed germination, but the ability to minimise the risk of misdiagnosis of seed quality linked to it is becoming more important (Nehoshtan, 2021). Seed priming is an old empirical technique that has been widely adopted by the seed industry. The original imperative was to reduce physiological activities of tri-phasic seed germination pattern in order to enhance seedling establishment. Improvement in seed science and technology over time has increased the scope of priming definition and implementation. Seed imbibition with water to enhance seed germination has been superseded by methods that are known to have direct effects on a wider range of physiological and biochemical processes associated with seed germination to include early crop establishment processes (Mansour et al. 2022 ). Halopriming is the use of specific salts, which can have nutritional value – nutripriming. Hormopriming relies on application of plant growth regulators. The use of osmotic agents is osmopriming. The significance of crop response to salinity is well-documented (Akhtar et al. 2000, Kiferle et al. 2022 ). Response of nutripriming varies in different plants and is dependent on the concentration of the micronutrient used (Farooq et al. 2012). Favourable priming treatment activates avoidance response such as protective enzymes and accumulating osmoprotectants, which improve seed germination and seedling vigour in different crops (Ali et al. 2018, Nadeem and Farooq 2019). Using iodine as a priming agent could ameliorate the adversities of oxidative stress by detoxification of reactive oxygen species (ROS) both directly or by enhancing the activities of antioxidants. For instance, several studies have reported that iodine in plants increases the accumulation of antioxidants and osmoprotectants, which enhance stress tolerance (Blasco et al. 2013, Leyva et al. 2011, Gupta et al. 2015 , Medrano-Macías 2016). Previous studies were carried out in hydroponics, using foliar spraying trials. Little information is available regarding priming seeds with iodine. This invokes a significant question as to whether iodine priming can be used as a tool to influence tolerance of wheat genotypes at early stages of growth under simulated water stress. Drought tolerance is a major area of crop improvement because the negative effects of water stress can lead to crop loss at initiation, developmental stages and even yield (Hadebe and Modi 2017 ). Germination and early establishment, which have been proven to respond to priming, are critical stages with respect to crop vulnerability to biotic and abiotic stress conditions (Fuentes et al. 2022 ). The vulnerability is linked to the physiological and biochemical processes. It has been reported that water stress causes an increase in reactive oxygen species (ROS) and oxidative stress in plant cells, which further induces membrane deterioration, protein damage and lipid peroxidation (Kerchev and van Breusegem 2022 ). Wheat is the major cereal for global food security. Understanding the response of this crop to any potential changes in environmental conditions during growth is essential. The objective of this study was to determine the early growth stage response of wheat to halopriming and simulated water stress in terms of physiological indicators associated with leaf membrane stability index and potential photosynthetic activity. Results and discussion Water deficit reduced both the morphological [root to shoot biomass ratio (RSR) and membrane stability index (MSI)] and physiological (carotenoids and chlorophyll contents) traits of wheat seedlings in a manner that was directly related to relative water content (Table 1 and Fig. 1 ). The accumulation of the shoot biomass was limited under water deficit conditions and the seedlings grew more roots in order to compensate for the water stress. However, the accumulation of both root and shoot biomass was not significantly affected by iodate halopriming, although, judging by the root to shoot ratio there was a clear trend of better shoot accumulation with increasing concentration of halopriming (Table 1 ). Membrane stability index also improved with potassium iodate halopriming concentration (Table 1 ). This is an indication that cell membrane leakage was minimised by KIO 3 halopriming (Table 1 ). Table 1 Table 1 . Wheat root shoot ratio (RSR), membrane stability index (MSI) and relative water content (RWC) in response to water availability (FC = Field capacity; WD = Water deficit to 25% FC) and KIO3 halopriming concentrations. Values sharing the same letter (a, b, c) are not significantly different (p ≤ 0.05). Source of variance RSR MSI (%) RWC (%) Water availability FC 0.190a ≠ 80.8a 74.4a WD 0.262b 63.9b 47.3b Halopriming (mg l − 1 ) 0 0.246a 57.2a 57a 100 0.227a 62.9b 66b 200 0.205a 77.2c 69b Wheat genotypes SST806 0.215a 85a 61a SST8135 0.252a 79a 60a PAN3111 0.311ab 81a 62a CV (%) 36.0 26.4 18.1 Potassium iodate is known for its ability to improve the antioxidant enzymatic system of plants (Lawson et al., 2015 ). Therefore, it can also play a role in protecting plant cells from reactive oxygen species (ROS) which cause oxidation of nucleic acids and cause cellular dysfunction. In this study, accumulation of carotenoids and chlorophyll content were used as indicators of plant health (Fig. 1 ). The response of the three wheat cultivars used in this study was similar with respect to the effects of halopriming and water availability (Fig. 1 ). Halopriming caused a consistent improvement of both chlorophyll a and chlorophyll b in wheat, irrespective of water availability status during seedling growth (Fig. 1 ). However, the change effect of halopriming and water availability was highly significant for chlorophyll b compared to chlorophyll a (Fig. 1 ). Water stress compromises leaf and root membrane stability (Deskoy et al., 2021, Recchia et al. 2018 ). The findings of this study show that there is a positive role of KIO 3 halopriming beyond seed vigour improvement. Based on previous studies, the effect of KIO 3 could be explained in the context of protection from reactive oxidative species (ROS) which damage plant cells (Chen et al., 2017 ). This finding is confirmed by the significant improvement of membrane stability index (MSI) in response to KIO3 halopriming (Table 1 ). This observation is an indication of protection of cells from leakage of electrolytes, because increased MSI is an indication of electrical conductivity reduction. It is suggested that the increase in chlorophyll content due to KIO 3 halopriming is an indication of the positive physiological performance of wheat seedlings with respect to photosynthesis. Enhancement of the light-harvesting antenna in plants due to accelerated increase in the photosynthetic pigments (chlorophyll and carotenoids) has been shown in previous studies. While the current data focussed on one critical wheat plant development stage, Z14.7/22, it can be suggested that the effect of KIO 3 halopriming is an indicator of effective crop manipulation to resist water stress (Desoky et al. 2021 ). This role can be extended to the potential of wheat to resist heat stress, because carotenoids are useful for the harvesting of toxic oxygen species and protecting chlorophyll from light damage. The study offers the importance of the chemo-physiology of KIO3 iodate halopriming in the context of the green gap during photosynthesis (Martignago et al. 2020 ). Understanding crop response to environment is a multidimensional element of plant science in that it requires aspects of morphology, physiology, biochemistry and genetics. A combination of these aspects is not easy. This study showed that the common approach to seed science and technology, priming, can be extended in its value to explain wheat seed response beyond the traditional germination capacity index. The study purposefully excluded germination data for its predictability and focussed on seedling response. Results indirectly observations that were previously made in a variety of vegetable crops and cereals, including wheat regarding the effect of potassium iodate on plant growth and regulation. The innovative aspect of the study was the ability to relate halopriming of wheat seeds with seedling performance under conditions of variable water availability. Relative water content is an important indicator of water stress tolerance by plants. Membrane stability index is a critical indicator of resistance to damage due to ROS. The positive response of chlorophyll and carotenoids is an indication of potential enhancement of productivity in the long term. This study should inform research on wheat yield determination in relation to seed treatment. Methods Plant and material Seeds of three wheat genotypes (SST806, SST8135 and PAN3111 cultivars), from MacDonald Seed Co, Pietermaritzburg, South, were subjected to halopriming followed by simulated water availability during seedling growth. Seed treatment Halopriming of seeds was performed for a period of 24 hours at 25 o C (Golob et al., 2020 ; Mejía-Ramírez et al., 2023 ). Three priming solutions were prepared from a stock solution of potassium iodate (1000 ppm KIO 3 , 99%, Sigma Aldrich, St Louis, MO, USA). Pure salt (168.59 mg) was dissolved in 100 ml of distilled water. Distilled water was used to gauge the stock solution to 25 ml. Dilutions of 4 and 8 were used to obtain 100 mg l − 1 and 200 mg l − 1 , respectively. Distilled water was used as a control (0 mg l − 1 ). Water stress and biomass accumulation Using a pot trial, three seeds of wheat were planted in 1kg of loam soil (van Zyl et al. 2020 ). Fertliser application was based on recommendation of soil analysis results (Zhang et al 2019). Simulated water stress was determined according to El-Sanatawy (2021). Briefly, water availability was maintained at field capacity (100% FC) from planting until the seedling harvest stage Z14.7/22 (Zadoks et al. 1974 ) by applying the required water content twice a week, whereas water deficit treatment was based on no irrigation from planting to harvest, at which time water availability was at 25% FC. At harvest, seedling biomass accumulation indication was determined using the root: shoot ratio (RSR) (Bláha, 2019 ). Plant relative water content (RWC) was determined as previously published (Soltys-Kalina et al. 2016 ) as: RWC = (Fresh Weight - Dry weight)/ (Turgid Weight-Dry Weight) x 100 Eq. 1 Membrane stability index Membrane stability index (MSI) was determined according to the modification of the published methods (Dastborhan and Ghassemi-Golezani, 2015 , Kumar et al. 2015). For this study, electrical conductivity leakage from wheat seedling leaves was determined as: MSI = [1 - (C 1 /C 2 )] x 100 Eq. 2 where, C 1 = Leaf conductivity in distilled water at 40 o C, 30 minutes, returning to 25 o C C 2 = Leaf conductivity in distilled water at 100 o C, 30 minutes, returning to 25 o C Chlorophyll and carotenoid determination Spectrophotometric determinations of chlorophyll a, b and carotenoids were performed according to Chappelle et al. ( 1992 ). Statistical analysis Three levels of halopriming and two levels of water availability were blocked within each one of the three wheat genotypes. Analysis of variance (ANOVA) was performed using Genstat®. The null hypothesis was that there is no difference between halopriming and water availability treatments with respect to wheat cultivar response in terms of the measured variables, namely, RSR, MSI, Chlorophyll and carotenoids. Significant differences were determined using standard error (SE, p ≤ 0.05). Declarations Conflict of interest There are no conflicts of interest to declare. Ethical approval The study did not require ethical clearance. Author Contribution Albert Modi collected data, analyzed and wrote the manuscript for review. Acknowledgement The support of research assistants and field workers of the Controlled Environment facility is acknowledged. Data Availability Data is provided within the manuscript References Akhtar, J., Qureshi, M.A., Naseem, A., Iqbal, M.S. & Massod, M.A. Differential response of fragrant rice cultivars to salinity and hydrogen rich water in relation to growth and antioxidative defense mechanisms. Int. J. Phytoremediation 23(11):1–9 (2021). DOI: 10.1080/15226514.2021.1889963 Bláha, L. Importance of Root - Shoot Ratio for Crops Production. J. Agron. Agric. Sci. 2: 012 (2019). doi: 10.24966/AAS-8292/100012 Blasco, B. Ríos J.J., Leyva R, Cervilla L.M., Sánchez-Rodríguez E. Rubio-Wilhelmi M.M., Rosales M.A., Ruiz J..M & Romero L. 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The effect of drought stress on leaf relative water content and tuber yield of a half-sib family of “Katahdin’-derived potato cultivars. Breed. Sci. 66 (2): 328–331(2016). doi: 10.1270/jsbbs.66.328 Van Zyl, G. Turner, D., Paterson, G, Koch, J., van Tol, J., Barichiey, K., Clarke, C., du Plessis M. & van der Venter P. The new soil classification system in South Africa, its history, important changes made and implications for users. South African Journal of Plant and Soil 37(5): 331–342 (2020). https://doi.org/10.1080/02571862.2020.1815244 Zadoks, J.C., Chang, T.T. & Konzak, C.F. (1974). A decimal code for the growth stages of cereals. Weed Res. 14: 415–421 (1974). https://doi.org/10.1111/j.1365-3180.1974.tb01084.x Zhang, W., Liu, D., Liu, Y., Chen, X., Zou, C. Overuse of phosphorus fertilizer reduces the grain and flour protein contents and zinc bioavailability of winter wheat (Triticum aestivum L.). J. Agric. Food Chem. 65(8): 1473–1482 (2017) http://dx.doi.org/10.1021/acs.jafc.6b04778 (2017) Additional Declarations No competing interests reported. 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-4626171","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":327139265,"identity":"5b807876-9dfc-464d-9e29-6188bc32fcd7","order_by":0,"name":"Albert Thembinkosi Modi","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5ElEQVRIiWNgGAWjYDACdh4o4zDzwQdAioePoBZmkJYEkBa2ZAOQFjbitRzgUZMACRDUwt/Me/Bx5Y9tcnzHedgqv+bYybAxMD98dAOPFonDfMmGZxJuG0se5j12W3ZbMtBhbMbGOfisOcxjJtmQcDtxw2G+tNuS25iBWnjYpPFpkT/MY/4TooXHrFhyWz1hLQZAlYwwLYwftx0mrMUQ6BfJhjSQX9iSpRm3AQOBmYBf5I73HvzYYHNbju/84YMff26rtudnb374GK/3kQE4jhiYiVUOAow/SFE9CkbBKBgFIwYAALUDRi/WUGzLAAAAAElFTkSuQmCC","orcid":"","institution":"Walter Sisulu University","correspondingAuthor":true,"prefix":"","firstName":"Albert","middleName":"Thembinkosi","lastName":"Modi","suffix":""}],"badges":[],"createdAt":"2024-06-23 17:38:20","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4626171/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4626171/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":60518099,"identity":"7c6d06e7-464e-4ed8-89a3-6bc0655c8793","added_by":"auto","created_at":"2024-07-17 15:54:37","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":25249,"visible":true,"origin":"","legend":"\u003cp\u003eChlorophyll (A) and carotenoid (B) contents in in response of three wheat genotypes (SST806, SST8135, PAN3111) to water availability (FC = Field capacity; WD = Water deficit to 25% FC) and KIO\u003csub\u003e3\u003c/sub\u003e halopriming concentrations (0, 100 and 200 mgl\u003csup\u003e-1\u003c/sup\u003e). Note: Chl a means chlorophyll a and\u0026nbsp; Chl b means chlorophyll b.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4626171/v1/702ce536909fdc624ef98a30.png"},{"id":66168675,"identity":"dd4ed9e4-13d5-469b-9147-b50b06b6e43e","added_by":"auto","created_at":"2024-10-08 10:17:18","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":357857,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4626171/v1/6ed0b4fe-a94a-4689-84d8-ba07d24ca8d2.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Preliminary determination of early growth response of wheat to potassium iodate priming","fulltext":[{"header":"Introduction","content":"\u003cp\u003eSeed quality is multidimensional in that it requires understating of physiological responses to a variety of factors that affect seed hormones and protein response singularly or in combination. Seed germination is the best indicator of seed quality, but it is triggered by different environmental treatments during development and after physiological maturity, including storage. Hence, it is not only important to improve seed germination, but the ability to minimise the risk of misdiagnosis of seed quality linked to it is becoming more important (Nehoshtan, 2021). Seed priming is an old empirical technique that has been widely adopted by the seed industry. The original imperative was to reduce physiological activities of tri-phasic seed germination pattern in order to enhance seedling establishment. Improvement in seed science and technology over time has increased the scope of priming definition and implementation. Seed imbibition with water to enhance seed germination has been superseded by methods that are known to have direct effects on a wider range of physiological and biochemical processes associated with seed germination to include early crop establishment processes (Mansour et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Halopriming is the use of specific salts, which can have nutritional value \u0026ndash; nutripriming. Hormopriming relies on application of plant growth regulators. The use of osmotic agents is osmopriming. The significance of crop response to salinity is well-documented (Akhtar et al. 2000, Kiferle et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eResponse of nutripriming varies in different plants and is dependent on the concentration of the micronutrient used (Farooq et al. 2012). Favourable priming treatment activates avoidance response such as protective enzymes and accumulating osmoprotectants, which improve seed germination and seedling vigour in different crops (Ali et al. 2018, Nadeem and Farooq 2019). Using iodine as a priming agent could ameliorate the adversities of oxidative stress by detoxification of reactive oxygen species (ROS) both directly or by enhancing the activities of antioxidants. For instance, several studies have reported that iodine in plants increases the accumulation of antioxidants and osmoprotectants, which enhance stress tolerance (Blasco et al. 2013, Leyva et al. 2011, Gupta et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, Medrano-Mac\u0026iacute;as 2016). Previous studies were carried out in hydroponics, using foliar spraying trials. Little information is available regarding priming seeds with iodine. This invokes a significant question as to whether iodine priming can be used as a tool to influence tolerance of wheat genotypes at early stages of growth under simulated water stress.\u003c/p\u003e \u003cp\u003eDrought tolerance is a major area of crop improvement because the negative effects of water stress can lead to crop loss at initiation, developmental stages and even yield (Hadebe and Modi \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Germination and early establishment, which have been proven to respond to priming, are critical stages with respect to crop vulnerability to biotic and abiotic stress conditions (Fuentes et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The vulnerability is linked to the physiological and biochemical processes. It has been reported that water stress causes an increase in reactive oxygen species (ROS) and oxidative stress in plant cells, which further induces membrane deterioration, protein damage and lipid peroxidation (Kerchev and van Breusegem \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Wheat is the major cereal for global food security. Understanding the response of this crop to any potential changes in environmental conditions during growth is essential. The objective of this study was to determine the early growth stage response of wheat to halopriming and simulated water stress in terms of physiological indicators associated with leaf membrane stability index and potential photosynthetic activity.\u003c/p\u003e"},{"header":"Results and discussion","content":"\u003cp\u003eWater deficit reduced both the morphological [root to shoot biomass ratio (RSR) and membrane stability index (MSI)] and physiological (carotenoids and chlorophyll contents) traits of wheat seedlings in a manner that was directly related to relative water content (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e and Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The accumulation of the shoot biomass was limited under water deficit conditions and the seedlings grew more roots in order to compensate for the water stress. However, the accumulation of both root and shoot biomass was not significantly affected by iodate halopriming, although, judging by the root to shoot ratio there was a clear trend of better shoot accumulation with increasing concentration of halopriming (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Membrane stability index also improved with potassium iodate halopriming concentration (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). This is an indication that cell membrane leakage was minimised by KIO\u003csub\u003e3\u003c/sub\u003e halopriming (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Wheat root shoot ratio (RSR), membrane stability index (MSI) and relative water content (RWC) in response to water availability (FC\u0026thinsp;=\u0026thinsp;Field capacity; WD\u0026thinsp;=\u0026thinsp;Water deficit to 25% FC) and KIO3 halopriming concentrations. Values sharing the same letter (a, b, c) are not significantly different (p\u0026thinsp;\u0026le;\u0026thinsp;0.05).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSource of variance\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eRSR\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMSI\u003c/p\u003e \u003cp\u003e(%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRWC\u003c/p\u003e \u003cp\u003e(%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eWater availability\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.190a\u003csup\u003e\u0026ne;\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e80.8a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e74.4a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.262b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e63.9b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e47.3b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eHalopriming (mg l\u003c/b\u003e\u003csup\u003e\u003cb\u003e\u0026minus;\u0026thinsp;1\u003c/b\u003e\u003c/sup\u003e\u003cb\u003e)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.246a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e57.2a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e57a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.227a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e62.9b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e66b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.205a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e77.2c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e69b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eWheat genotypes\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSST806\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.215a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e85a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e61a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSST8135\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.252a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e79a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e60a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePAN3111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.311ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e81a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e62a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCV (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e36.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e26.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003ePotassium iodate is known for its ability to improve the antioxidant enzymatic system of plants (Lawson et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Therefore, it can also play a role in protecting plant cells from reactive oxygen species (ROS) which cause oxidation of nucleic acids and cause cellular dysfunction. In this study, accumulation of carotenoids and chlorophyll content were used as indicators of plant health (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The response of the three wheat cultivars used in this study was similar with respect to the effects of halopriming and water availability (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Halopriming caused a consistent improvement of both chlorophyll a and chlorophyll b in wheat, irrespective of water availability status during seedling growth (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). However, the change effect of halopriming and water availability was highly significant for chlorophyll b compared to chlorophyll a (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWater stress compromises leaf and root membrane stability (Deskoy et al., 2021, Recchia et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The findings of this study show that there is a positive role of KIO\u003csub\u003e3\u003c/sub\u003e halopriming beyond seed vigour improvement. Based on previous studies, the effect of KIO\u003csub\u003e3\u003c/sub\u003e could be explained in the context of protection from reactive oxidative species (ROS) which damage plant cells (Chen et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). This finding is confirmed by the significant improvement of membrane stability index (MSI) in response to KIO3 halopriming (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). This observation is an indication of protection of cells from leakage of electrolytes, because increased MSI is an indication of electrical conductivity reduction. It is suggested that the increase in chlorophyll content due to KIO\u003csub\u003e3\u003c/sub\u003e halopriming is an indication of the positive physiological performance of wheat seedlings with respect to photosynthesis. Enhancement of the light-harvesting antenna in plants due to accelerated increase in the photosynthetic pigments (chlorophyll and carotenoids) has been shown in previous studies. While the current data focussed on one critical wheat plant development stage, Z14.7/22, it can be suggested that the effect of KIO\u003csub\u003e3\u003c/sub\u003e halopriming is an indicator of effective crop manipulation to resist water stress (Desoky et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This role can be extended to the potential of wheat to resist heat stress, because carotenoids are useful for the harvesting of toxic oxygen species and protecting chlorophyll from light damage. The study offers the importance of the chemo-physiology of KIO3 iodate halopriming in the context of the green gap during photosynthesis (Martignago et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eUnderstanding crop response to environment is a multidimensional element of plant science in that it requires aspects of morphology, physiology, biochemistry and genetics. A combination of these aspects is not easy. This study showed that the common approach to seed science and technology, priming, can be extended in its value to explain wheat seed response beyond the traditional germination capacity index. The study purposefully excluded germination data for its predictability and focussed on seedling response. Results indirectly observations that were previously made in a variety of vegetable crops and cereals, including wheat regarding the effect of potassium iodate on plant growth and regulation. The innovative aspect of the study was the ability to relate halopriming of wheat seeds with seedling performance under conditions of variable water availability. Relative water content is an important indicator of water stress tolerance by plants. Membrane stability index is a critical indicator of resistance to damage due to ROS. The positive response of chlorophyll and carotenoids is an indication of potential enhancement of productivity in the long term. This study should inform research on wheat yield determination in relation to seed treatment.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003ePlant and material\u003c/h2\u003e \u003cp\u003eSeeds of three wheat genotypes (SST806, SST8135 and PAN3111 cultivars), from MacDonald Seed Co, Pietermaritzburg, South, were subjected to halopriming followed by simulated water availability during seedling growth.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eSeed treatment\u003c/h2\u003e \u003cp\u003eHalopriming of seeds was performed for a period of 24 hours at 25\u003csup\u003eo\u003c/sup\u003eC (Golob et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mej\u0026iacute;a-Ram\u0026iacute;rez et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Three priming solutions were prepared from a stock solution of potassium iodate (1000 ppm KIO\u003csub\u003e3\u003c/sub\u003e, 99%, Sigma Aldrich, St Louis, MO, USA). Pure salt (168.59 mg) was dissolved in 100 ml of distilled water. Distilled water was used to gauge the stock solution to 25 ml. Dilutions of 4 and 8 were used to obtain 100 mg l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e and 200 mg l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively. Distilled water was used as a control (0 mg l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eWater stress and biomass accumulation\u003c/h2\u003e \u003cp\u003eUsing a pot trial, three seeds of wheat were planted in 1kg of loam soil (van Zyl et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Fertliser application was based on recommendation of soil analysis results (Zhang et al 2019). Simulated water stress was determined according to El-Sanatawy (2021). Briefly, water availability was maintained at field capacity (100% FC) from planting until the seedling harvest stage Z14.7/22 (Zadoks et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1974\u003c/span\u003e) by applying the required water content twice a week, whereas water deficit treatment was based on no irrigation from planting to harvest, at which time water availability was at 25% FC. At harvest, seedling biomass accumulation indication was determined using the root: shoot ratio (RSR) (Bl\u0026aacute;ha, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Plant relative water content (RWC) was determined as previously published (Soltys-Kalina et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) as:\u003c/p\u003e \u003cp\u003e \u003cem\u003eRWC = (Fresh Weight - Dry weight)/ (Turgid Weight-Dry Weight) x 100\u003c/em\u003e Eq.\u0026nbsp;1\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eMembrane stability index\u003c/h2\u003e \u003cp\u003eMembrane stability index (MSI) was determined according to the modification of the published methods (Dastborhan and Ghassemi-Golezani, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, Kumar et al. 2015). For this study, electrical conductivity leakage from wheat seedling leaves was determined as:\u003c/p\u003e \u003cp\u003e \u003cem\u003eMSI = [1 - (C\u003c/em\u003e \u003csub\u003e \u003cem\u003e1\u003c/em\u003e \u003c/sub\u003e \u003cem\u003e/C\u003c/em\u003e \u003csub\u003e \u003cem\u003e2\u003c/em\u003e \u003c/sub\u003e \u003cem\u003e)] x 100\u003c/em\u003e Eq.\u0026nbsp;2\u003c/p\u003e \u003cp\u003ewhere,\u003c/p\u003e \u003cp\u003eC\u003csub\u003e1\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;Leaf conductivity in distilled water at 40\u003csup\u003eo\u003c/sup\u003eC, 30 minutes, returning to 25\u003csup\u003eo\u003c/sup\u003eC\u003c/p\u003e \u003cp\u003eC\u003csub\u003e2\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;Leaf conductivity in distilled water at 100\u003csup\u003eo\u003c/sup\u003eC, 30 minutes, returning to 25\u003csup\u003eo\u003c/sup\u003eC\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eChlorophyll and carotenoid determination\u003c/h2\u003e \u003cp\u003eSpectrophotometric determinations of chlorophyll a, b and carotenoids were performed according to Chappelle et al. (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1992\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eThree levels of halopriming and two levels of water availability were blocked within each one of the three wheat genotypes. Analysis of variance (ANOVA) was performed using Genstat\u0026reg;. The null hypothesis was that there is no difference between halopriming and water availability treatments with respect to wheat cultivar response in terms of the measured variables, namely, RSR, MSI, Chlorophyll and carotenoids. Significant differences were determined using standard error (SE, p\u0026thinsp;\u0026le;\u0026thinsp;0.05).\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eConflict of interest\u003c/h2\u003e \u003cp\u003eThere are no conflicts of interest to declare.\u003c/p\u003e \u003c/p\u003e\u003cp\u003e \u003ch2\u003eEthical approval\u003c/h2\u003e \u003cp\u003eThe study did not require ethical clearance.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAlbert Modi collected data, analyzed and wrote the manuscript for review.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eThe support of research assistants and field workers of the Controlled Environment facility is acknowledged.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData is provided within the manuscript\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAkhtar, J., Qureshi, M.A., Naseem, A., Iqbal, M.S. \u0026amp; Massod, M.A. Differential response of fragrant rice cultivars to salinity and hydrogen rich water in relation to growth and antioxidative defense mechanisms. 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Food Chem. 65(8): 1473\u0026ndash;1482 (2017)\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.1021/acs.jafc.6b04778\u003c/span\u003e\u003cspan address=\"10.1021/acs.jafc.6b04778\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2017)\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","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":"Carotenoids, Chlorophyll, Halopriming, Water deficit, Wheat, Seed","lastPublishedDoi":"10.21203/rs.3.rs-4626171/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4626171/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe objective of the study was to determine the efficacy of iodine priming in combination with water availability levels on the physiological attributes of wheat \u003cem\u003e(Triticum aestivum\u003c/em\u003e) as determined at stage Z14.7/22 of plant growth. Two levels of potassium iodate halopriming (100 mg l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e and 200 mg l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e ) were compared with distilled for priming seeds before planting to grow seedlings under conditions of controlled water availability determined by field capacity (FC). Adequate water availability (100% FC) throughout seedling growth was compared with declining water availability from planting to 25% FC before harvesting plants to determine biomass in terms of root: shoot ratio (RSR). Crop physiological response was determined using membrane stability index (MSI) and leaf chlorophyll and carotenoid contents. Results confirmed the positive role of halopriming wheat seeds to improve resistance to water stress at the early stage of crop development. The use of potassium iodate negligibly improved plant biomass accumulation, but leaf membrane stability index was significantly improved. These changes were associated with enhancement of chlorophyll (a, b) and carotenoid contents. This study provides insight into the potential effect of seed treatment on wheat response to soil water availability at the early stages of development.\u003c/p\u003e","manuscriptTitle":"Preliminary determination of early growth response of wheat to potassium iodate priming","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-07-17 15:54:33","doi":"10.21203/rs.3.rs-4626171/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":"5f4f4b72-2a28-460f-804b-e4a918de3f36","owner":[],"postedDate":"July 17th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":34618832,"name":"Biological sciences/Physiology"},{"id":34618833,"name":"Biological sciences/Plant sciences"}],"tags":[],"updatedAt":"2024-10-08T10:09:02+00:00","versionOfRecord":[],"versionCreatedAt":"2024-07-17 15:54:33","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4626171","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4626171","identity":"rs-4626171","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}