Boosting Salt Tolerance in Camelina cultivars through synergistic Impact of Exogenous Silicon and Selenium by modulating physiological attributes and Antioxidant activities

Boosting Salt Tolerance in Camelina cultivars through synergistic Impact of Exogenous Silicon and Selenium by modulating physiological attributes and Antioxidant activities | 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 Boosting Salt Tolerance in Camelina cultivars through synergistic Impact of Exogenous Silicon and Selenium by modulating physiological attributes and Antioxidant activities Yahya M. Alzahrani, Hesham F. Alharby, Zahoor Ahmad, Halima Nawaz, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4317048/v1 This work is licensed under a CC BY 4.0 License Status: Under Revision Version 1 posted 6 You are reading this latest preprint version Abstract Salt stress (SS) has emerged as one of the most pressing environmental threats to the production sustainability of oil-seed crops like camelina which necessitates finding out pro-environment and biologically feasible amelioration approaches. A study was performed to investigate the effects of exogenously applied silicon (Si) and selenium (Se) on two camelina varieties (Australian and Canadian) under SS conditions. The trial was comprised of two doses of Si and Se (5 and 10 ppm each) along with two co-application treatments entailing Si (5pp) + Se (5ppm) and Si (10 pp) + Se (10 ppm), whereas control treatments were kept for comparison purpose. The response variables included root-shoot length, fresh and dry weights, as well as leaf photosynthetic pigment content (chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids), leaf gas exchange attributes, biochemical characters (total proline, total soluble sugar, total soluble protein, and total free amino acid) and antioxidant activities (APX, POD, CAT, and SOD). Results showed that SS significantly reduced all growth attributes of camelina, whereas the Australian camelina variety demonstrated greater salinity tolerance in comparison to the Canadian variety. The co-application of Si (10 pp) + Se (10 ppm) outperformed the rest of the treatments by recording the highest leaf chlorophyll a (1.69 mg/g FW), chlorophyll b (1.02 mg/g FW), carotenoids (2.89 mg/g FW), photosynthetic rate (17.02 µmole CO 2 m − 2 s − 1 ), transpiration rate (3.65 µmole CO 2 m − 2 s − 1 ), stomatal conductance (0.38 µmole CO 2 m − 2 s − 1 ), total proline (199.38 µmole/g DW), total soluble sugar (120.95 µmole/g FW), total soluble protein (12.02 mg/g DW) and total free amino acid (37.15 mg/g DW) in Australian camelina as compared to Canadian camelina. The same treatment also remained effective in triggering the activity of APX (3.89 Unit/min/g FW), CAT (165.19 Unit/min/g FW), POD (178.95 Unit/min/g FW), and SOD (237.63 Unit/min/g FW). Thus, the combined exogenous application of Si and Se holds bright perspectives in alleviating the deleterious impact of SS especially for the Australian variety, and could be recommended to camelina growers after thorough field investigations. Enhancing Salt tolerance Camelina Exogenous silicon and selenium Morphological attributes Chlorophyll pigments Leaf Gas Exchange attributes antioxidants Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Camelina ( Camelina sativa L.) is an oilseed crop that has gained increasing importance due to its high oil content, nutritional value, and a potential biofuel crop (Ahmad et al. 2020a ; Waraich et al. 2021 ; Stamenković et al. 2023 ). However, like many other crops, Camelina is susceptible to various environmental stresses, including salt stress (SS), which can have detrimental effects on plant growth, photosynthesis, and induce oxidative stress (Ahmad et al. 2020b ; Hölzl et al. 2023 ). It is a recently introduced oil-producing crop belonging to the Brassica oleracea family. Camelina seeds are widely used in food production and have the potential to produce biodiesel. It is an annual plant predominantly cultivated in spring and winter seasons, primarily in European countries. Its seeds are known for their high content of vitamins, proteins, bioactive compounds, phytosterols, and polyphenols (Sydor et al. 2022 ). Camelina oil is particularly valued for its high levels of unsaturated fats, specifically linoleic acid (18:2) and linolenic acid (18:3). These fatty acids have been enhanced through agricultural engineering techniques, making its oil a rich source of omega-3 essential fatty acids (Chaudhary et al. 2023 ). The unique fatty acid composition of camelina oil has favorable nutritional and physiological effects, such as reducing blood triglyceride and cholesterol levels. As a result, camelina oil is considered suitable for direct consumption, cooking, and as an enrichment ingredient in products such as margarine, salad dressing, mayonnaise, and ice cream, providing omega-3 fatty acids (Meza et al. 2022 ; Li et al. 2023 ). Under changing climate, abiotic stresses (drought, heat, salinity, etc.) have started to threaten the sustainability of modern intensive farming systems (Iqbal et al. 2023 ). Among abiotic stresses, SS has merged as one of the most pronounced abiotic stresses that affect plant growth and productivity worldwide (Choudhary et al. 2021 ; Yasir et al. 2021 ). It occurs when the concentration of salts, particularly sodium chloride (NaCl), exceeds the tolerance threshold of plants, leading to detrimental effects on various physiological processes (Riffat and Ahmad 2020 ; Kubi et al. 2021 ; Khalid et al. 2022 ). When exposed to high salt concentrations in the soil, camelina plants experience a range of physiological and biochemical disruptions (Göre 2023 ; Hosseini Sanehkoori et al. 2023 ). The SS disrupted the osmotic balance within plant cells, resulting in water deprivation and reduced turgor pressure. As a consequence, the growth attributes of camelina, including root and shoot length, as well as fresh and dry weights were adversely affected (Mzengereza et al. 2021 ; Kukrić et al. 2023 ). Photosynthetic pigments play a crucial role in capturing light energy and converting it into chemical energy through photosynthesis. However, SS significantly restricted the photosynthetic process in camelina by reducing the synthesis of chlorophyll, the primary pigment responsible for light absorption (Hezaveh et al. 2020 ). Consequently, chlorophyll a, chlorophyll b, and total chlorophyll contents decrease, leading to a decline in photosynthetic efficiency. Additionally, SS reduced the carotenoid pigments, which serve as protective agents against oxidative damage (Yohannes et al. 2020 ; Waraich et al. 2021 ). One of the major consequences of SS was the generation of reactive oxygen species (ROS) within plant cells. These ROS, such as superoxide radicals (O 2− ) and hydrogen peroxide (H 2 O 2 ), accumulated under SS conditions and disrupted vital cellular homeostasis. This oxidative stress also damages biomolecules, including proteins, lipids, and DNA, and impairs key cellular functions. To counteract the harmful effects of ROS, plants tend to activate antioxidant defense systems. These systems include enzymes such as ascorbate peroxidase (APX), peroxidase (POD), catalase (CAT), and superoxide dismutase (SOD), which scavenge ROS and prevent oxidative damage (Nishchenko and Hasanuzzaman 2020 ; Bakyani et al. 2022 ). Silicon (Si) is a valuable element known to enhance plant resistance to SS, a phenomenon that has been observed in rice ( Oryza sativa ), wheat ( Triticum aestivum ), maize ( Zea mays ), barley ( Hordeum vulgare ), sorghum ( Sorghum bicolor ), cucumber ( Cucumis sativus ), canola ( Brassica napus ) and tomato ( Solanum lycopersicum ) (Calero Hurtado et al. 2020 ; Badawy et al. 2021 ; Feghhenabi et al. 2022 ; Mousavi et al. 2022 ; Sayed et al. 2022 ; Stadnik et al. 2022 ; Ahmad et al. 2023 ; Tobiasz-Salach et al. 2023 ). The mechanisms underlying Si-mediated alleviation of SS have been elucidated such as Si deposition in the root cortex reduces the apoplastic transport of Na + into the xylem, thereby lowering Na + transport from roots to shoots (Hajihashemi and Kazemi 2022 ; da Luz Neto et al. 2023 ). Studies suggested that Si supplementation enhanced salt tolerance by modulating hormone levels and hormone-responsive genes (Tejada-Ruiz et al. 2020 ; Stadnik et al. 2023 ). Additionally, Si application directly or indirectly mitigated salt-induced oxidative damage by enhancing antioxidant enzyme activities and defense mechanisms (Mordhi Radhi and Alkarawia 2021 ; Peña Calzada et al. 2023 ). It also alleviated SS-induced osmotic stress by regulating aquaporins and the biosynthesis and metabolism of osmotic regulatory substances (Farouk et al. 2020 ; Mordhi Radhi and Alkarawia 2021 ; Shahzad et al. 2022 ; Shen et al. 2022 ). Besides Si, selenium (Se) is another trace element that is vital to both human and animal health (Admasie et al. 2023 ; Ghanbari et al. 2023 ). The Se performs a few vital functions such as an antioxidant protector, reducing the harm that different biotic and abiotic stresses, especially SS. Recently, several research findings have revealed that the protective benefits and mechanisms of Se-mediated alleviation of SSvary on the chemical forms and dosage of Se, SS levels, plant species, and culture circumstances (Mousavi et al. 2022 ; Abdi et al. 2023 ). Exogenous application of Se alleviated SS by reducing oxidative stress through increased ROS-scavenging activity (Zahedi et al. 2019 ; Ahmad et al. 2020b , 2021 ; Lanza and Reis 2021 ; Shahraki et al. 2022 ). Foliar application of Se has been found to boost plant growth in saline conditions (Pourebrahimi et al. 2023 ). The application of Se from external sources enhanced plant tolerance to stress, improved photosynthesis by increasing the levels of photosynthetic pigments, activated antioxidant defenses, and enhanced overall plant nutrition and SS tolerance (Desoky et al. 2021 ; Göre 2023 ). To date, only a few studies have explored the individual benefits of foliar applications of Si and Se in alleviating salt-induced phytotoxicity in camelina plants. Uncertainty surrounds the roles and processes underlying the combined impacts of Si and Se on tolerance against SS. Consequently, the purpose of this study was to look into the physiological and oxidative processes behind Si and Se's advantageous effects. Therefore, we hypothesized that either Si and Se applied solely or in conjunction with each other in different doses might perform differently in mitigating the deleterious impacts of salinity in camelina cultivars under their antioxidant-modulating potential. Thus, an experiment was performed to evaluate the individual and combined effects of exogenous application of Si and Se on physiological and antioxidant activity in camelina under salt stress conditions. Moreover, another prime aim of the study was comparatively evaluate the performance of two camelina varieties (Australian and Canadian) under salt stress conditions Methodology 2.1 Experimental site A controlled experiment was conducted at the wire-house of the Department of Botany, University of Central Punjab, Constituent Punjab College Bahawalpur, Pakistan in October 2021 to investigate the role of foliar applied silicon (Si) and selenium (Se) in Camelina under SS conditions. 2.2 Experimental detail The seeds of two Camelina varieties (Australian and Canadian), were obtained from the Department of Agronomy, Stress Physiology Lab, University of Agriculture Faisalabad, Pakistan. Plastic pots were purchased from a local market in Bahawalpur. Each pot was filled with 750 g of soil obtained from the field, and before sowing the seeds, the field capacity of the soil was determined. The seeds of both Camelina varieties were sown in the pots containing the soil as the growth medium. Initially, the seedlings were watered for establishment, and once germination occurred, the entire set of seedlings was regularly watered to maintain 100% field capacity (FC), subjecting them to SS conditions. The seedlings were grown until the vegetative stage (40 DAS), and data recording was performed. Each pot contained five seeds of both Camelina varieties and SS was applied after seed germination was complete. A basic Hoagland nutrient solution was applied to promote optimal growth and development of the plants. The Camelina plants were grown until the vegetative stage, and different treatments of foliar silicon (Si) and selenium (Se) were applied. The treatments included T 0 S 0 (control), T 0 S 1 (NaCl @ 100 ppm), T 1 S 1 (Si @ 5 ppm & NaCl @ 100 ppm), T 2 S 1 (Si @ 10 ppm & NaCl @ 100 ppm), T 3 S 1 (Se @ 5 ppm & NaCl @ 100 ppm), T 4 S 1 (Se @ 10 ppm & NaCl @ 100 ppm), T 5 S 1 (Si +Se @ 5 ppm & NaCl @ 100 ppm) and T 6 S 1 (Si + Se @ 10 ppm & NaCl @ 100 ppm). Throughout the growth period, various morphological, physiological, leaf gas exchange, and antioxidant attributes of the Camelina plants were recorded using standardized procedures. These measurements aimed to evaluate the impact of different foliar Si and Se treatments on the growth and development of the Camelina plants at the vegetative stage. 3.3 Measurement of growth traits of camelina The procedure involved in measuring the root-shoot length, as well as the fresh and dry weight of Camelina plants, was conducted with two plants from each replication. The Camelina plants were allowed to grow for 40 days under salt-stress conditions. At the end of the growth period, the root-shoot length was measured using a ruler or measuring tape, carefully noting the length of both the root and shoot for each plant in the replication. This measurement indicated the overall growth and development of the plants under different treatments. Subsequently, the fresh weight of the Camelina plants was determined by weighing each plant using a digital balance immediately after harvest. The plants were carefully separated from the soil to obtain accurate measurements of their fresh weight. To determine the dry weight, the plants were then dried in an oven at a specific temperature (usually around 60-70°C) for a predetermined period, typically until a constant weight was achieved. This ensured that all moisture content was removed from the plant tissue, allowing for an accurate measurement of the dry weight. 3.4 Determination of photosynthetic pigments in camelina leaf The procedure for measuring leaf chlorophyll pigments, including chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid contents in Camelina leaves, involved several steps. Firstly, healthy and fully expanded leaves from the Camelina plants were selected for the analysis. These leaves were carefully collected from each treatment group, ensuring representative samples from the different foliar Si and Se concentrations. The leaf samples were then washed gently with distilled water to remove any dust or debris that might interfere with the pigment measurements. Care was taken to handle the leaves delicately to prevent any damage that could affect the accuracy of the results. Next, the leaf chlorophyll pigments were extracted using an appropriate solvent, typically acetone or ethanol. The leaves were placed in a test tube or vial, and a sufficient amount of solvent was added to cover the leaves completely. The tubes were then tightly sealed or covered to prevent any evaporation during the extraction process. To extract the pigments effectively, the tubes were placed in a dark environment or wrapped in aluminum foil to minimize exposure to light. The samples were left to soak in the solvent for a specific duration, allowing the pigments to dissolve and separate from the leaf tissue. After the extraction period, the tubes were gently agitated or mixed to ensure thorough mixing of the pigments with the solvent. This step aimed to achieve a homogeneous solution that would facilitate accurate measurements. Once the pigment extraction was complete, the absorbance of the solution was measured using a spectrophotometer. The spectrophotometer was set to appropriate wavelengths, typically around 663 nm and 645 nm for chlorophyll a and chlorophyll b, respectively, and around 470 nm for carotenoids. The absorbance readings indicated the concentration of the respective pigments in the leaf extract. Based on the absorbance measurements, the concentrations of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids were calculated using standard equations or formulas (Lichtenthaler and Wellburn, 1983). These calculations involved converting the absorbance values into pigment concentrations, considering factors such as dilution and sample volume. 3.5 Measurement of gas exchange attributes in camelina At the 40 DAS vegetative growth stage, samples for measuring gas exchange properties were taken between 9:00 and 11:00 am in direct sunlight. Using an infrared gas analyzer (CI-340 portable, Hoddesdon, England), gas exchange attributes such as transpiration rate (Tr), stomatal conductance (Gs), photosynthetic rate (Pn), and intercellular CO2 concentration (Ci) were determined (Waraich et al. 2015). 3.6 Measurement of assay of antioxidants in camelina leaf The procedure for measuring the assay of antioxidants, including catalase (CAT), peroxidase (POD), superoxide dismutase (SOD), and ascorbate peroxidase (APX), in Camelina leaves was conducted to assess the antioxidant defense mechanisms under SS conditions. Firstly, Camelina leaves from each treatment group were carefully collected at the designated time point, typically after 40 days of growth under salt-stress conditions. The leaves were selected from multiple plants within each replication to ensure representative samples. The collected Camelina leaves were homogenized in a suitable extraction buffer using a mortar and pestle or a homogenizer. The extraction buffer typically contained a phosphate buffer or a Tris-HCl buffer, along with additives to stabilize the enzyme activity and maintain pH conditions. Once the leaves were thoroughly homogenized, the resulting leaf extract was centrifuged to separate the soluble fraction from the cellular debris. The supernatant containing the enzyme extract was then collected for further analysis. 3.5.1 Activity of catalase The activity of catalase (CAT) was determined by measuring the decomposition of hydrogen peroxide (H 2 O 2 ) spectrophotometrically at a specific wavelength. The rate of H 2 O 2 decomposition indicated CAT activity in the leaf extract (Maehly and Chance, 1954). 3.5.2 Activity of peroxidase By observing the oxidation of a suitable substrate such as guaiacol in the presence of hydrogen peroxide, peroxidase (POD) activity was measured. The color development resulting from the oxidation reaction was measured spectrophotometrically, reflecting the POD activity in the leaf extract (Maehly and Chance, 1954). 3.5.3 Activity of superoxide dismutase The suppression of the photochemical reduction of nitro blue tetrazolium (NBT) in the presence of riboflavin and light was used to evaluate the activity of superoxide dismutase (SOD). The reduction in NBT photo reduction indicated the SOD activity in the leaf extract (Giannopolitis and Ries 1977). 3.5.4 Activity of ascorbate peroxidase Ascorbate peroxidase (APX) activity was determined by monitoring the oxidation of ascorbate in the presence of hydrogen peroxide. The decrease in absorbance at a specific wavelength, corresponding to ascorbate oxidation, provided a measure of APX activity in the leaf extract (Nakano and Asada 1981). 3.6 Statistical analysis The recorded data were organized and analyzed using the Statistics 8.1 programming software. The data were subjected to a one-way analysis of variance (ANOVA) technique for determining the overall significance of employed treatments, thereafter, the least significant difference (LSD) test was conducted at a significance level of 5% to assess the significance of the treatment means (Steel and Torrie 1962). Results 3.1 Growth attributes of camelina responses under the separate and combined effects of Si and Se under salinity stress Recorded results of growth attributes, such as root-shoot length and their fresh as well as dry weight, indicate that the different treatments of silicon (Si) and selenium (Se) separately or in combination significantly (P ≥ 0.005) affected all growth attributes under SS conditions (Table 1). A graphical representation illustrates that SS significantly reduced the measured attributes of growth, such as shoot length, shoot fresh weight, shoot dry weight, root length, root fresh weight, and root dry weight in both camelina genotypes, Australian camelina, and Canadian camelina. The highest reduction of shoot length (9.00 cm), shoot fresh weight (11.2 g), shoot dry weight (3.01 g), root length (15.01 cm), root fresh weight (13.89 g), and root dry weight (2.87 g) were noted in Canadian camelina compared to Australian camelina under SS conditions ((Figure 1 a-f). The different doses of Si and Se also significantly improved the measured growth characteristics of both camelina genotypes. The maximum shoot length (32.55 cm), shoot fresh weight (31.24 g), shoot dry weight (6.93 g), root length (38.91 cm), root fresh weight (33.15 g), and root dry weight (7.63 g) were recorded in the treatment where Si and Se were applied in combination (Si + Se @ 10 ppm and NaCl @ 100 ppm) compared to all other treatments in Australian camelina, while the lowest values of shoot length, shoot fresh weight, shoot dry weight, root length, root fresh weight, and root dry weight were noted in Canadian camelina under SS conditions (Figure 1 a-f). Table 1: Analysis of variance table for growth parameters, leaf chlorophyll pigments, infrared gas analyzer attributes, biochemical, and antioxidant assays of camelina under s7alinity stress conditions Measured attributes Source of variance Treatments (T) Varieties (V) T x V Shoot length 369.77 *** 34.748 ** 1.768 NS Shoot fresh weight 11.915 *** 0.691 NS 0.115 NS Shoot dry weight 248.401*** 26.940** 0.497 NS Root length 333.464 *** 42.563 ** 0.348 NS Root fresh weight 270.231 *** 19.520 ** 0.974 NS Root dry weight 15.294 *** 1.166 * 0.016 NS Chlorophyll a content 0.361 *** 0.029 * 0.0010 NS Chlorophyll b content 0.447 *** 0.030 * 0.0007 NS Total chlorophyll content 1.603 *** 0.119 * 0.0024 NS Carotenoids content 1.282 *** 0.102 ** 0.0012 NS Photosynthetic rate 118.31 *** 11.55 ** 0.414 NS Transpiration rate 6.2588 *** 0.8453 *** 0.0314 NS Stomatal conductance 0.5262 *** 0.0060 ** 0.00008 NS Total proline 10155 *** 899.2 *** 15.6 NS Total soluble sugar 5952.7*** 525.29 ** 23.09 NS Total soluble protein 70.497 *** 3.763 ** 0.162 NS Total free amino-acid 555.09 *** 40.223 ** 1.177 NS APX 2.433 *** 0.243 ** 0.0038 NS CAT 11799 *** 1190.5 *** 36.2 NS POD 13567 *** 1373.5 *** 29.2 NS SOD 9258 *** 598.2 ** 18.38 NS *** P ≤ 0.001, ** P ≤ 0.01, * P ≤ 0.05; NS , non-signifcant 3.2 Leaf chlorophyll pigments of camelina responses under the separate and combined effects of Si and Se under salinity stress Recorded results of leaf chlorophyll pigments, including chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid content, indicate that the different treatments of silicon (Si) and selenium (Se), separately or in combination, significantly (P ≥ 0.005) affected all leaf chlorophyll pigments under SS conditions (Table 1). A graphical representation illustrates that SS significantly reduced the measured attributes of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid content in both camelina genotypes, (Australian camelina, and Canadian camelina). The highest reduction of chlorophyll a (0.89 mg/g FW), chlorophyll b (0.18 mg/g FW), total chlorophyll (1.07 mg/g FW), and carotenoid content (1.41 mg/g FW) was noted in Canadian camelina compared to Australian camelina under SS conditions (Figure 2 a-d). The different doses of Si and Se also significantly improved both camelina genotypes' measured leaf chlorophyll pigments. The maximum chlorophyll a (1.69 mg/g FW), chlorophyll b (1.02 mg/g FW), total chlorophyll (2.71 mg/g FW), and carotenoid content (2.89 mg/g FW) were recorded in the treatment where Si and Se were applied in combination (Si + Se @ 10 ppm and NaCl @ 100 ppm) compared to all other treatments in Australian camelina. In contrast, the lowest values of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid content were noted in Canadian camelina under SS conditions (Figure 2 a-d). 3.3 Gas exchange attributes of camelina responses under the separate and combined effect of Si and Se under salinity stress All leaf gas exchange attributes under SS conditions were significantly (P ≥ 0.005) affected by the various treatments of silicon (Si) and selenium (Se), either separately or in combination, according to recorded results of gas exchange attributes, including photosynthetic rate, transpiration rate, and stomatal conductance (Table 1). A graphical depiction shows that in both camelina genotypes (Australian and Canadian), SS dramatically decreased the measured characteristics of photosynthetic rate, transpiration rate, and stomatal conductance. Under SS conditions, Canadian camelina showed the greatest drop in photosynthetic rate (3.54 µmol CO 2 m -2 s -1 ), transpiration rate (0.48 µmol CO 2 m -2 s -1 ), and stomatal conductance (0.09 µmol CO 2 m -2 s -1 ) when compared to Australian camelina (Figure 3 a-c). The assessed gas exchange properties of both camelina genotypes were significantly improved by the varied dosages of Si and Se. When Si and Se were applied together (Si + Se @ 10 ppm and NaCl @ 100 ppm), the maximum photosynthetic rate (17.02 µmol CO 2 m -2 s -1 ), transpiration rate (3.65 µmol CO 2 m -2 s -1 ), and stomatal conductance (0.38 µmol CO 2 m -2 s -1 ) were recorded in Australian camelina as compared to all other treatments. On the other hand, under SS circumstances, Canadian camelina showed the lowest values of transpiration rate, stomatal conductance, and photosynthetic rate (Figure 3 a-c). 3.4 Biochemical characters of camelina responses under the separate and combined effect of Si and Se under salinity stress Under SS conditions, all biochemical characters, including total proline, total soluble sugar, total soluble protein, and total free amino acid, showed significant (P ≥ 0.005) effects from the different treatments of silicon (Si) and selenium (Se), either separately or in combination (Table 1). A graphical representation demonstrates that SS significantly reduced the observed features of total proline, total soluble sugar, total soluble protein, and total free amino acid in both Australian and Canadian camelina genotypes. Comparing Canadian and Australian camelina under SS, the former displayed the largest declines in total proline (71.32 µmol/g DW), total soluble sugar (31.32 µmole/g FW), total soluble protein (1.79 mg/g DW), and total free amino acid (8.41 mg/g DW) were observed in Canadian camelina (Figure 4 a-d). The different doses of Si and Se resulted in a significant improvement in the measured gas exchange characteristics of both camelina genotypes. In comparison to all other treatments, the maximum levels of total proline (195.15 µmol/g DW), total soluble sugar (120.95 µmole/g FW), total soluble protein (12.02 mg/g DW), and total free amino acid (37.15 mg/g DW) were recorded in Australian camelina when Si and Se were applied together (Si + Se @ 10 ppm and NaCl @ 100 ppm). Conversely, under SS conditions, total proline, total soluble sugar, total soluble protein, and total free amino acid values in Canadian camelina were the lowest (Figure 4 a-d). 3.5 Antioxidants assay of camelina responses under the separate and combined effects of Si and Se under salinity stress Under SS circumstances, all antioxidant enzymes were significantly (P > 0.005) impacted by the various treatments of silicon (Si) and selenium (Se), either independently or in combination, according to recorded results of antioxidant activities, including APX, CAT, POD, and SOD (Table 1). A graphical depiction shows that in both camelina genotypes (Australian and Canadian), the SS dramatically decreased the observed characteristics of APX, CAT, POD, and SOD. When comparing Canadian camelina to Australian camelina under SS conditions, the greatest reductions in APX (1.95 Unit/min/g FW), CAT (29.36 Unit/min/g FW), POD (34.77 Unit/min/g FW), and SOD (126.42 Unit/min/g FW) were observed (Figure 5 a-d). The measured leaf chlorophyll pigments of both camelina genotypes were significantly enhanced by the various dosages of Si and Se. When Si and Se were treated in combination (Si + Se @ 10 ppm and NaCl @ 100 ppm), the greatest APX (3.89 Unit/min/g FW), CAT (165.19 Unit/min/g FW), POD (178.95 Unit/min/g FW), and SOD (237.63 Unit/min/g FW) were reported in Australian camelina as compared to the other treatments. On the other hand, under SS circumstances, Canadian camelina showed the lowest values of APX, CAT, POD, and SOD (Figure 5 a-d). Discussion Due to the susceptibility of crop plants to varying levels of SS, salinity has posed a daunting challenge to oil-seed crop productivity (Abdi et al. 2023). The deleterious impacts of SS on multiple physiological systems, including photosynthesis, ion homeostasis, and antioxidant enzyme activity, have been reported in a wide array of crop plants. The increase in Na+ and Cl-ions in many plants has frequently been linked to the detrimental effects of salt; as a result, these ions provide essential circumstances for plant life by interfering with various plant processes (Pourebrahimi et al. 2023). The current study showed that all measured growth attributes such as root-shoot length and their fresh as well as dry weight (Figure 1 a-f) were reduced under SS (NaCl was applied at the rate of 100 ppm). The same results were observed in the previous work, they stated that the SS significantly reduced the growth characteristics such as root-shoot length and their fresh as well as dry weight in plants (Hosseinpour et al. 2020; Noreen et al. 2021). The reduction in the growth attributes of the plant might be attributed to more accumulation of salt in the soil, which led to reduced water potential nutrient uptake, and ultimately growth attributes were seriously reduced under SS conditions (Al-Farsi et al. 2020a; Ahmad et al. 2023). However, in this study, exogenous application of Si and Se either separately or in combination remained effective in ameliorating the SS effects by boosting growth attributes in both camelina genotypes (Australian camelina and Canadian camelina). The height improvement was observed in all growth parameters where Si + Se was applied at the rate of 10 ppm under 100 pm NaCl stress conditions (Figure 1 a-f). Similar findings have previously been reported whereby the exogenous application of Si and Se imparted tolerance against SS by modulating physiological processes (Abdi et al. 2023). The role of Si in the improvement of growth parameters in different crops was noted and recorded in the previous work which showed that the foliar application of Si improved the growth and development of plants under SS conditions (Farhangi-Abriz and Torabian 2018; Mordhi Radhi and Alkarawia 2021). Similarly, the exogenous application of Se also played an important role in the enhancement of growth attributes of plants by restricting the uptake of salts (Hossain et al. 2021). Moreover, Si and Se when applied in combination showed more pronounced positive responses in the enhancement of the growth as well as the physiological process of plants under SS conditions (Rajput et al. 2021; Taha et al. 2021). Under SS, Plants experienced oxidative stress due to increased ROS generation, which disrupted the normal metabolism of plant cells which led to decreased growth attributes and crop productivity (da Luz Neto et al. 2023). Under these circumstances, ROS causes oxidative stress to proteins, chlorophyll, and cell membranes; in response, plants increase their antioxidant capacity (Zhao et al. 2021). In the current study, some detrimental effects of NaCl-induced SS were noted on the physiological functions of the lemon verbena plant. In the environment around the roots of lemon plants, adding more NaCl increased proline, soluble sugar, and antioxidant enzymes while decreasing chlorophyll and relative water contents (RWC)(Ciriello et al. 2022). The findings demonstrated that under varying degrees of SS, foliar spraying lemon verbena with 10 µM Se enhanced the plant's growth metrics and RWC. Furthermore, plants treated with Se exhibited a decreased leaf-level accumulation (LEL) and accumulation of MDA and H 2 O 2 in comparison to control plants. This suggests that the plants were less susceptible to oxidative damage induced by salt. Oxidative damage has been linked to the build-up of MDA and H 2 O 2 under stressful settings(Admasie et al. 2023). To impart tolerance against abiotic stresses, the antioxidant defense system consisting of both enzymatic and non-enzymatic components performed key role in scavenging the ROS (Elsheery et al. 2020). However, under SS, the results of this trial demonstrated that the exogenous administration of Se boosted the proline, soluble carbohydrates, and total protein of lemon verbena plants. There have been prior reports on the build-up of proline and other osmolytes in stressed plants when exogenous Se is applied (Xu et al. 2021; El-Badri et al. 2022). Proline plays a unique role in osmotic stress circumstances, and helps plants to withstand stress conditions by regulating osmotic pressure, eliminating free radicals, conserving energy, and protecting macromolecules (Noreen et al. 2021). It was also deduced that by lowering oxidative damage and enhancing plant development in lemon verbena under SS, the exogenous administration of Se triggered the defense mechanisms. This was explained by the accumulation of osmolytes. According to reports in this regard, Se treatment changed the physiological processes of the plant, including the accumulation of protein and osmolytes, the enhancement of the antioxidant capacity, the maintenance of ion homeostasis, and the enhancement of the photosynthetic capacity, thereby reducing the effects of stress on the plant and promoting plant growth under SS condition (Zahedi et al. 2019; Xu et al. 2021; Ghanbari et al. 2023). According to Regni et al. (2021), the administration of Se enhanced olive plants' resistance to abiotic stress by preventing oxidative damage to cells and controlling their water status. Furthermore, it has been documented that the foliar application of selenium (Se) remained effective in boosting the concentration of chlorophyll in plants through the preservation of chloroplast enzymes and the augmentation of photosynthetic pigment production (Ghanbari et al. 2023). The current investigation found that the SS significantly affected the leaf gas exchange attributes such as net photosynthetic rate (P N ), transpiration rate (E), and stomatal conductance (g s ) while the application of foliar Si and Se either separately or in combination assisted cameline plants in the improving leaf gas exchange attributes in both camelina genotypes (Figure 3 a-c). In the studies conducted on oats by (Qin et al. 2018) and (Shah et al. 2020), revealed that SS promoted stomatal closure, which reduced the amount of carbon dioxide available in the leaves and prevented carbon fixation. This raised the amount of excitation energy that the chloroplasts were exposed to, increasing the production of reactive oxygen species (ROS) and oxidative stress. Furthermore, it was observed that the presence of salt led to a reduction in the gas exchange characteristics (Ondrasek et al. 2022). By altering the PSII system and the chloroplast ultrastructure, salinity reduced stomatal conductance and inhibited transpiration and gaseous exchange (Pan et al. 2021). A reduced supply of CO 2 resulting from partial stomata closure and an altered biochemical binding mechanism for CO 2 could be the cause of the inhibition of plant absorption during SS (Shah et al. 2020; Kukrić et al. 2023). This resulted in changes to cellular metabolism, a decrease in CO2 photosynthetic binding, and an increase in ROS production in chloroplasts. ROS harmed the photosynthetic apparatus, especially PSII, which resulted in photoinhibition. This was caused by an imbalance in the photosynthetic redox signaling pathways and suppression of PSII repair (Yan et al. 2021; Liang et al. 2024). In a stressful environment, plants typically tend to close their stomata to preserve water, which also lowers the stomatal conductance and photosynthesis (Zahra et al. 2022). Se and/or Si-induced improvements in wheat growth features might be due to their roles in controlling physiological processes, such as photosynthesis, plant tissue water status, and antioxidant machinery (Taha et al. 2021). Wheat growth and productivity were found to be positively impacted by the stimulating roles that Se and Si play in protein synthesis, chlorophyll synthesis, and the absorption of essential nutrients (Sattar et al. 2017; Qin et al. 2024). Plants under SS benefitted from Si's beneficial effects on their metabolic processes (Mordhi Radhi and Alkarawia 2021). Additionally, it caused a reduction in the stomata lumen and silicifies the leaf surface, which both affected the rate of plant transpiration (Sayed et al. 2022). The transport and absorption of several nutrients were enhanced by the exogenous application of Si (Raza et al. 2019; Zia et al. 2023). In the same setting, applying Se resulted in an amino acid metabolism imbalance; as a result, stressed plants exhibited higher levels of soluble protein and lower nitrate activity (Ahmad et al. 2021; Ghanbari et al. 2023). Similar to our findings, the Se application also remained instrumental in protecting plant cells, increased the formation of starch in chloroplasts, and supported plant growth in salt-stressed environments (Regni et al. 2021). Organic osmolytes like proline and total soluble sugars help plants regulate their cell osmoregulation and lessen the effects of stress (uz Zaman et al. 2022). Osmoprotectant concentrations and stress tolerance are closely correlated, and they are regarded as one of the markers for the potential for stress tolerance (Al-Farsi et al. 2020b). Additionally, these decreased the negative effects of stress by controlling the synthesis of vital proteins like Rubisco, shielding the photosynthetic system, improving membrane stability, preserving the redox equilibrium, and eliminating ROS (Ahmad et al. 2020a). Plants responded to internal Si stress by boosting the production of metabolites through soluble sugars and proline. Through the osmotic balance, which preserved cell turgor and guaranteed the stability and safety of cellular membranes, these metabolites promoted tolerance against SS. This reduced photo-oxidation and oxidative damage (Farouk et al. 2020; Sayed et al. 2022). According to our research findings, applying Se and/or Si increased the levels of organic molecules such as proline and TSS that could be attributed to increased amylase activity and the hydrolysis of starch by selenium; fructose 1, 6-bisphosphatase, which is involved in the metabolism of carbohydrates. When compared to unstressed plants, wheat plants subjected to SS showed increased TSS and proline buildup in leaf tissue. The method of adaptation to salinity stress is reflected in these increases in proline and TSS concentrations in wheat leaves during SS. Spraying the wheat plants with Se, Si, and their mixtures resulted in a further increase in the accumulation of proline and TSS. Salinity in the soil causes osmotic stress and can reduce the amount of water that roots take in, which can eventually disrupt the activities of stomata and roots (Badawy et al. 2021; Xu et al. 2021; Abdi et al. 2023). This could be the cause of the decrease in stomatal conductance in wheat plants during SS. In the present study, plant water conditions and hydraulic conductivity under SS may have been mediated by an increase in TSS and proline buildup in response to both Se and Si, as well as their mixtures. Furthermore, wheat plants treated with Se/Si exhibited a substantial accumulation of proline, which may have contributed to ROS scavenging in the current study to reduce oxidative damage (Manzoor et al. 2024; Qin et al. 2024). Conclusion In conclusion, the research findings revealed the deleterious impacts of salinity stress on the growth, photosynthetic pigments, leaf gas exchange, and antioxidant activities of Camelina, as well as the potential benefits of foliar Si and Se application in mitigating these effects. The findings demonstrated that SS negatively affected Camelina's growth attributes photosynthetic pigments and leaf gas exchange attributes indicating its vulnerability to salt-induced limitations. However, the foliar application of Si and Se in combination, particularly at higher concentrations (10 ppm), significantly improved the measured parameters compared to the control. The positive effects of Si and Se on Camelina under SS conditions can be attributed to their antioxidant properties and their ability to enhance the activity of antioxidant enzymes. Si and Se supplementation helped to scavenge reactive oxygen species (ROS) and minimize oxidative damage, thereby improving the overall tolerance of Camelina to salinity stress. The results also revealed genotype-specific variations in SS tolerance, with the Australian variety of Camelina displaying superior performance to the Canadian variety. This highlights the importance of considering genetic factors when studying plant responses to environmental stressors. The results of this investigation advance our comprehension of the systems underlying SS tolerance in Camelina and provide valuable insights for the development of strategies to enhance its productivity in saline environments. The application of Si and Se as foliar supplements shows promise as a practical approach to improving SS tolerance in Camelina crops. Declarations Author Contributions ; Hesham F. Alharby and Zahoor Ahmad planned the research, Halima Nawaz and Zahoor Ahmad conducted the research work; Hameed Alsamadany and Muhammad Aamir Iqbal wrote the introduction and helped in the improvement of the manuscript's quality; Zahoor Ahmad, Yahya M. Alzahrani, Muhammad Aamir Iqbal, and Awatif M. Abdulmajeed did the static analysis and graphical representation; Zahoor Ahmad read the manuscript as proofreading and arranged it according to the journal style; Zahoor Ahmad provided reagents, assisted in the analytical work; and Zahoor Ahmad, Muhammad Aamir Iqbal and Hesham F. Alharby improved the English language quality of the manuscript. All authors participate in the improvement of the quality of the manuscript. Funding: This work was funded by Institutional Fund Projects under grant no. (IFPIP: 737-130-1443), Ministry of Education in Saudi Arabia. Acknowledgment: This research work was funded by Institutional Fund Projects under grant no. (IFPIP: 737-130-1443). The authors gratefully acknowledge the technical and financial support from the Ministry of Education and King Abdulaziz University, DSR, Jeddah, Saudi Arabia. Ethical Approval: All authors approve this manuscript for submission to Environmental Science and Pollution Research. 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Environ Sci Pollut Res. https://doi.org/10.1007/s11356-023-26902-9 Cite Share Download PDF Status: Under Revision Version 1 posted Editorial decision: Major Revision 10 Jul, 2024 Reviewers agreed at journal 01 Jun, 2024 Reviewers invited by journal 30 May, 2024 Editor invited by journal 21 May, 2024 Editor assigned by journal 29 Apr, 2024 First submitted to journal 24 Apr, 2024 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-4317048","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":308860714,"identity":"30c4729f-db31-40d7-b1df-046dc7bd1a02","order_by":0,"name":"Yahya M. Alzahrani","email":"","orcid":"","institution":"King Abdulaziz University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yahya","middleName":"M.","lastName":"Alzahrani","suffix":""},{"id":308860715,"identity":"d6900f46-aadd-4978-9a44-d90b09d6192d","order_by":1,"name":"Hesham F. Alharby","email":"","orcid":"","institution":"King Abdulaziz University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Hesham","middleName":"F.","lastName":"Alharby","suffix":""},{"id":308860716,"identity":"a856be43-e2b0-40de-9f89-7c4de34aae0f","order_by":2,"name":"Zahoor Ahmad","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9klEQVRIiWNgGAWjYFACHjCZAMRmQGwDxIyNB0jRkgbS0kCSlsNgHl4t8u1nDz74uMcuT7f98LbHvDvO261tPwy0pcYmGpcWxp68ZMMZz5KLzc6klRvznrmdvO1MIlDLsbTcBhxamBlyzKR5DjAnbjsAZOS23U42OwDUwthwGKcWNv43IC31idvOvwFpOZdsdv4hfi08EmBbDiduuwG25YCd2Q0CtkhIvDE2nHHgOFDLs3Ljv2eSE8xuAG1JwOMX+f4cwwcfDlQDHZa87eHMHXb2ZufTHz74UGODUwsqYGxgSASrTCBKOVSLPdGKR8EoGAWjYMQAAFmDaE5AsTxIAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0001-8453-5130","institution":"University of Central Punjab","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Zahoor","middleName":"","lastName":"Ahmad","suffix":""},{"id":308860717,"identity":"8c0dd078-eca2-4e19-b7da-39e7472102a2","order_by":3,"name":"Halima Nawaz","email":"","orcid":"","institution":"University of Central Punjab","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Halima","middleName":"","lastName":"Nawaz","suffix":""},{"id":308860718,"identity":"f970fcfb-9baf-4bbe-8344-1c9747ec3d70","order_by":4,"name":"Muhammad Aamir Iqbal","email":"","orcid":"","institution":"Louisiana Tech University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Muhammad","middleName":"Aamir","lastName":"Iqbal","suffix":""},{"id":308860719,"identity":"9379e91e-a20c-4c09-9a3f-c662e300b844","order_by":5,"name":"Hameed Alsamadany","email":"","orcid":"","institution":"King Abdulaziz University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Hameed","middleName":"","lastName":"Alsamadany","suffix":""},{"id":308860720,"identity":"abe2d3dc-fbd1-49ff-b43f-c471bc6154fd","order_by":6,"name":"Awatif M. Abdulmajeed","email":"","orcid":"","institution":"University of Tabuk","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Awatif","middleName":"M.","lastName":"Abdulmajeed","suffix":""}],"badges":[],"createdAt":"2024-04-24 09:23:10","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4317048/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4317048/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":58290567,"identity":"70f300b6-2f5a-4a4e-a53e-99244fba4a24","added_by":"auto","created_at":"2024-06-13 13:28:41","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":267745,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSynergistic effect of foliar silicon and selenium on the morphological \u0026nbsp;\u0026nbsp;characteristics of camelina under salinity-induced stress\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"F1.png","url":"https://assets-eu.researchsquare.com/files/rs-4317048/v1/bb0b8d2531d9d6e9cc24096f.png"},{"id":58290570,"identity":"69bcb402-5982-4832-9751-a6fafa248f3d","added_by":"auto","created_at":"2024-06-13 13:28:41","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":91867,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSynergistic effect of foliar silicon and selenium on the leaf chlorophyll \u0026nbsp;\u0026nbsp;pigments of camelina under salinity-induced stress\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"F2.png","url":"https://assets-eu.researchsquare.com/files/rs-4317048/v1/6d7f5de4a7fcf3f846178b61.png"},{"id":58290568,"identity":"b2a88eab-65a0-42d4-b5e2-dcac78456551","added_by":"auto","created_at":"2024-06-13 13:28:41","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":70401,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSynergistic effect of foliar silicon and selenium on the leaf gas exchange \u0026nbsp;\u0026nbsp;attributes of camelina under salinity-induced stress\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"F3.png","url":"https://assets-eu.researchsquare.com/files/rs-4317048/v1/0c6dad22ad1ffefeeff1cd48.png"},{"id":58290571,"identity":"fee93a12-6197-4ab1-9caf-c66750190ea4","added_by":"auto","created_at":"2024-06-13 13:28:41","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":171755,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSynergistic effect of foliar silicon and selenium on the biochemical \u0026nbsp;\u0026nbsp;characters of camelina under salinity-induced stress\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"F4.png","url":"https://assets-eu.researchsquare.com/files/rs-4317048/v1/e384a9b0af736ffba26a413e.png"},{"id":58290569,"identity":"c8623967-5812-4cd6-995b-b19a02971529","added_by":"auto","created_at":"2024-06-13 13:28:41","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":188735,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSynergistic effect of foliar silicon and selenium on the antioxidants \u0026nbsp;\u0026nbsp;assay of camelina under salinity-induced stress\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"F5.png","url":"https://assets-eu.researchsquare.com/files/rs-4317048/v1/6dda97a0ceb1d936f573ce69.png"},{"id":58290969,"identity":"c9658489-52cb-4d87-82aa-6fa9baa6ded8","added_by":"auto","created_at":"2024-06-13 13:36:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1708951,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4317048/v1/5111cd2a-4611-49c0-b867-3c9e187b66a0.pdf"}],"financialInterests":"","formattedTitle":"Boosting Salt Tolerance in Camelina cultivars through synergistic Impact of Exogenous Silicon and Selenium by modulating physiological attributes and Antioxidant activities","fulltext":[{"header":"Introduction","content":"\u003cp\u003eCamelina (\u003cem\u003eCamelina sativa\u003c/em\u003e L.) is an oilseed crop that has gained increasing importance due to its high oil content, nutritional value, and a potential biofuel crop (Ahmad et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e; Waraich et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Stamenković et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, like many other crops, Camelina is susceptible to various environmental stresses, including salt stress (SS), which can have detrimental effects on plant growth, photosynthesis, and induce oxidative stress (Ahmad et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e; H\u0026ouml;lzl et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). It is a recently introduced oil-producing crop belonging to the Brassica oleracea family. Camelina seeds are widely used in food production and have the potential to produce biodiesel. It is an annual plant predominantly cultivated in spring and winter seasons, primarily in European countries. Its seeds are known for their high content of vitamins, proteins, bioactive compounds, phytosterols, and polyphenols (Sydor et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Camelina oil is particularly valued for its high levels of unsaturated fats, specifically linoleic acid (18:2) and linolenic acid (18:3). These fatty acids have been enhanced through agricultural engineering techniques, making its oil a rich source of omega-3 essential fatty acids (Chaudhary et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The unique fatty acid composition of camelina oil has favorable nutritional and physiological effects, such as reducing blood triglyceride and cholesterol levels. As a result, camelina oil is considered suitable for direct consumption, cooking, and as an enrichment ingredient in products such as margarine, salad dressing, mayonnaise, and ice cream, providing omega-3 fatty acids (Meza et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eUnder changing climate, abiotic stresses (drought, heat, salinity, etc.) have started to threaten the sustainability of modern intensive farming systems (Iqbal et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Among abiotic stresses, SS has merged as one of the most pronounced abiotic stresses that affect plant growth and productivity worldwide (Choudhary et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Yasir et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). It occurs when the concentration of salts, particularly sodium chloride (NaCl), exceeds the tolerance threshold of plants, leading to detrimental effects on various physiological processes (Riffat and Ahmad \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Kubi et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Khalid et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). When exposed to high salt concentrations in the soil, camelina plants experience a range of physiological and biochemical disruptions (G\u0026ouml;re \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Hosseini Sanehkoori et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The SS disrupted the osmotic balance within plant cells, resulting in water deprivation and reduced turgor pressure. As a consequence, the growth attributes of camelina, including root and shoot length, as well as fresh and dry weights were adversely affected (Mzengereza et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kukrić et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Photosynthetic pigments play a crucial role in capturing light energy and converting it into chemical energy through photosynthesis. However, SS significantly restricted the photosynthetic process in camelina by reducing the synthesis of chlorophyll, the primary pigment responsible for light absorption (Hezaveh et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Consequently, chlorophyll a, chlorophyll b, and total chlorophyll contents decrease, leading to a decline in photosynthetic efficiency. Additionally, SS reduced the carotenoid pigments, which serve as protective agents against oxidative damage (Yohannes et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Waraich et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). One of the major consequences of SS was the generation of reactive oxygen species (ROS) within plant cells. These ROS, such as superoxide radicals (O\u003csup\u003e2\u0026minus;\u003c/sup\u003e) and hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e), accumulated under SS conditions and disrupted vital cellular homeostasis. This oxidative stress also damages biomolecules, including proteins, lipids, and DNA, and impairs key cellular functions. To counteract the harmful effects of ROS, plants tend to activate antioxidant defense systems. These systems include enzymes such as ascorbate peroxidase (APX), peroxidase (POD), catalase (CAT), and superoxide dismutase (SOD), which scavenge ROS and prevent oxidative damage (Nishchenko and Hasanuzzaman \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bakyani et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSilicon (Si) is a valuable element known to enhance plant resistance to SS, a phenomenon that has been observed in rice (\u003cem\u003eOryza sativa\u003c/em\u003e), wheat (\u003cem\u003eTriticum aestivum\u003c/em\u003e), maize (\u003cem\u003eZea mays\u003c/em\u003e), barley (\u003cem\u003eHordeum vulgare\u003c/em\u003e), sorghum (\u003cem\u003eSorghum bicolor\u003c/em\u003e), cucumber (\u003cem\u003eCucumis sativus\u003c/em\u003e), canola (\u003cem\u003eBrassica napus\u003c/em\u003e) and tomato (\u003cem\u003eSolanum lycopersicum\u003c/em\u003e) (Calero Hurtado et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Badawy et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Feghhenabi et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Mousavi et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Sayed et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Stadnik et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Ahmad et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Tobiasz-Salach et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The mechanisms underlying Si-mediated alleviation of SS have been elucidated such as Si deposition in the root cortex reduces the apoplastic transport of Na\u0026thinsp;+\u0026thinsp;into the xylem, thereby lowering Na\u0026thinsp;+\u0026thinsp;transport from roots to shoots (Hajihashemi and Kazemi \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; da Luz Neto et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Studies suggested that Si supplementation enhanced salt tolerance by modulating hormone levels and hormone-responsive genes (Tejada-Ruiz et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Stadnik et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Additionally, Si application directly or indirectly mitigated salt-induced oxidative damage by enhancing antioxidant enzyme activities and defense mechanisms (Mordhi Radhi and Alkarawia \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Pe\u0026ntilde;a Calzada et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). It also alleviated SS-induced osmotic stress by regulating aquaporins and the biosynthesis and metabolism of osmotic regulatory substances (Farouk et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mordhi Radhi and Alkarawia \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Shahzad et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Shen et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBesides Si, selenium (Se) is another trace element that is vital to both human and animal health (Admasie et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Ghanbari et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The Se performs a few vital functions such as an antioxidant protector, reducing the harm that different biotic and abiotic stresses, especially SS. Recently, several research findings have revealed that the protective benefits and mechanisms of Se-mediated alleviation of SSvary on the chemical forms and dosage of Se, SS levels, plant species, and culture circumstances (Mousavi et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Abdi et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Exogenous application of Se alleviated SS by reducing oxidative stress through increased ROS-scavenging activity (Zahedi et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Ahmad et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Lanza and Reis \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Shahraki et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Foliar application of Se has been found to boost plant growth in saline conditions (Pourebrahimi et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The application of Se from external sources enhanced plant tolerance to stress, improved photosynthesis by increasing the levels of photosynthetic pigments, activated antioxidant defenses, and enhanced overall plant nutrition and SS tolerance (Desoky et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; G\u0026ouml;re \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo date, only a few studies have explored the individual benefits of foliar applications of Si and Se in alleviating salt-induced phytotoxicity in camelina plants. Uncertainty surrounds the roles and processes underlying the combined impacts of Si and Se on tolerance against SS. Consequently, the purpose of this study was to look into the physiological and oxidative processes behind Si and Se's advantageous effects.\u003c/p\u003e \u003cp\u003eTherefore, we hypothesized that either Si and Se applied solely or in conjunction with each other in different doses might perform differently in mitigating the deleterious impacts of salinity in camelina cultivars under their antioxidant-modulating potential. Thus, an experiment was performed to evaluate the individual and combined effects of exogenous application of Si and Se on physiological and antioxidant activity in camelina under salt stress conditions. Moreover, another prime aim of the study was comparatively evaluate the performance of two camelina varieties (Australian and Canadian) under salt stress conditions\u003c/p\u003e"},{"header":"Methodology","content":"\u003cp\u003e\u003cstrong\u003e2.1 Experimental site\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eA controlled experiment was conducted at the wire-house of the Department of Botany, University of Central Punjab, Constituent Punjab College Bahawalpur, Pakistan in October 2021 to investigate the role of foliar applied silicon (Si) and selenium (Se) in Camelina under SS conditions.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e2.2 Experimental detail\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe seeds of two Camelina varieties (Australian and Canadian), were obtained from the Department of Agronomy, Stress Physiology Lab, University of Agriculture Faisalabad, Pakistan. Plastic pots were purchased from a local market in Bahawalpur. Each pot was filled with 750 g of soil obtained from the field, and before sowing the seeds, the field capacity of the soil was determined. The seeds of both Camelina varieties were sown in the pots containing the soil as the growth medium.\u003c/p\u003e\n\u003cp\u003eInitially, the seedlings were watered for establishment, and once germination occurred, the entire set of seedlings was regularly watered to maintain 100% field capacity (FC), subjecting them to SS conditions. The seedlings were grown until the vegetative stage (40 DAS), and data recording was performed. Each pot contained five seeds of both Camelina varieties and SS was applied after seed germination was complete. A basic Hoagland nutrient solution was applied to promote optimal growth and development of the plants. The Camelina plants were grown until the vegetative stage, and different treatments of foliar silicon (Si) and selenium (Se) were applied. The treatments included\u0026nbsp;T\u003csub\u003e0\u003c/sub\u003eS\u003csub\u003e0\u003c/sub\u003e (control), T\u003csub\u003e0\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (NaCl @ 100 ppm), T\u003csub\u003e1\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (Si @ 5 ppm \u0026amp; NaCl @ 100 ppm), T\u003csub\u003e2\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (Si @ 10 ppm \u0026amp; NaCl @ 100 ppm), T\u003csub\u003e3\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (Se @ 5 ppm \u0026amp; NaCl @ 100 ppm), T\u003csub\u003e4\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (Se @ 10 ppm \u0026amp; NaCl @ 100 ppm), T\u003csub\u003e5\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (Si +Se @ 5 ppm \u0026amp; NaCl @ 100 ppm) and T\u003csub\u003e6\u003c/sub\u003eS\u003csub\u003e1\u003c/sub\u003e (Si + Se @ 10 ppm \u0026amp; NaCl @ 100 ppm). Throughout the growth period, various morphological, physiological, leaf gas exchange, and antioxidant attributes of the Camelina plants were recorded using standardized procedures. These measurements aimed to evaluate the impact of different foliar Si and Se treatments on the growth and development of the Camelina plants at the vegetative stage.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.3 Measurement of growth traits of camelina\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe procedure involved in measuring the root-shoot length, as well as the fresh and dry weight of Camelina plants, was conducted with two plants from each replication. The Camelina plants were allowed to grow for 40 days under salt-stress conditions. At the end of the growth period, the root-shoot length was measured using a ruler or measuring tape, carefully noting the length of both the root and shoot for each plant in the replication. This measurement indicated the overall growth and development of the plants under different treatments. Subsequently, the fresh weight of the Camelina plants was determined by weighing each plant using a digital balance immediately after harvest. The plants were carefully separated from the soil to obtain accurate measurements of their fresh weight. To determine the dry weight, the plants were then dried in an oven at a specific temperature (usually around 60-70\u0026deg;C) for a predetermined period, typically until a constant weight was achieved. This ensured that all moisture content was removed from the plant tissue, allowing for an accurate measurement of the dry weight.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.4 Determination of photosynthetic pigments in camelina leaf\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe procedure for measuring leaf chlorophyll pigments, including chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid contents in Camelina leaves, involved several steps. Firstly, healthy and fully expanded leaves from the Camelina plants were selected for the analysis. These leaves were carefully collected from each treatment group, ensuring representative samples from the different foliar Si and Se concentrations. The leaf samples were then washed gently with distilled water to remove any dust or debris that might interfere with the pigment measurements. Care was taken to handle the leaves delicately to prevent any damage that could affect the accuracy of the results. Next, the leaf chlorophyll pigments were extracted using an appropriate solvent, typically acetone or ethanol. The leaves were placed in a test tube or vial, and a sufficient amount of solvent was added to cover the leaves completely. The tubes were then tightly sealed or covered to prevent any evaporation during the extraction process. To extract the pigments effectively, the tubes were placed in a dark environment or wrapped in aluminum foil to minimize exposure to light. The samples were left to soak in the solvent for a specific duration, allowing the pigments to dissolve and separate from the leaf tissue. After the extraction period, the tubes were gently agitated or mixed to ensure thorough mixing of the pigments with the solvent. This step aimed to achieve a homogeneous solution that would facilitate accurate measurements. Once the pigment extraction was complete, the absorbance of the solution was measured using a spectrophotometer. The spectrophotometer was set to appropriate wavelengths, typically around 663 nm and 645 nm for chlorophyll a and chlorophyll b, respectively, and around 470 nm for carotenoids. The absorbance readings indicated the concentration of the respective pigments in the leaf extract. Based on the absorbance measurements, the concentrations of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids were calculated using standard equations or formulas\u0026nbsp;(Lichtenthaler and Wellburn, 1983). These calculations involved converting the absorbance values into pigment concentrations, considering factors such as dilution and sample volume.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.5 Measurement of gas exchange attributes in camelina\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAt the 40 DAS vegetative growth stage, samples for measuring gas exchange properties were taken between 9:00 and 11:00 am in direct sunlight. Using an infrared gas analyzer (CI-340 portable, Hoddesdon, England), gas exchange attributes such as transpiration rate (Tr), stomatal conductance (Gs), photosynthetic rate (Pn), and intercellular CO2 concentration (Ci) were determined\u0026nbsp;(Waraich et al. 2015).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.6 Measurement of assay of antioxidants in camelina leaf\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe procedure for measuring the assay of antioxidants, including catalase (CAT), peroxidase (POD), superoxide dismutase (SOD), and ascorbate peroxidase (APX), in Camelina leaves was conducted to assess the antioxidant defense mechanisms under SS conditions. Firstly, Camelina leaves from each treatment group were carefully collected at the designated time point, typically after 40 days of growth under salt-stress conditions. The leaves were selected from multiple plants within each replication to ensure representative samples. The collected Camelina leaves were homogenized in a suitable extraction buffer using a mortar and pestle or a homogenizer. The extraction buffer typically contained a phosphate buffer or a Tris-HCl buffer, along with additives to stabilize the enzyme activity and maintain pH conditions. Once the leaves were thoroughly homogenized, the resulting leaf extract was centrifuged to separate the soluble fraction from the cellular debris. The supernatant containing the enzyme extract was then collected for further analysis.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.5.1 Activity of catalase\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe activity of catalase (CAT) was determined by measuring the decomposition of hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e) spectrophotometrically at a specific wavelength. The rate of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u0026nbsp;\u003c/sub\u003edecomposition indicated CAT activity in the leaf extract\u0026nbsp;(Maehly and Chance, 1954).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.5.2 Activity of peroxidase\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBy observing the oxidation of a suitable substrate such as guaiacol in the presence of hydrogen peroxide, peroxidase (POD) activity was measured. The color development resulting from the oxidation reaction was measured spectrophotometrically, reflecting the POD activity in the leaf extract\u0026nbsp;(Maehly and Chance, 1954).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.5.3 Activity of superoxide dismutase\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe suppression of the photochemical reduction of nitro blue tetrazolium (NBT) in the presence of riboflavin and light was used to evaluate the activity of superoxide dismutase (SOD). The reduction in NBT photo reduction indicated the SOD activity in the leaf extract\u0026nbsp;(Giannopolitis and Ries 1977).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.5.4 Activity of ascorbate peroxidase\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAscorbate peroxidase (APX) activity was determined by monitoring the oxidation of ascorbate in the presence of hydrogen peroxide. The decrease in absorbance at a specific wavelength, corresponding to ascorbate oxidation, provided a measure of APX activity in the leaf extract\u0026nbsp;(Nakano and Asada 1981).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.6 Statistical analysis\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe recorded data were organized and analyzed using the Statistics 8.1 programming software. The data were subjected to a one-way analysis of variance (ANOVA) technique for determining the overall significance of employed treatments, thereafter, the least significant difference (LSD) test was conducted at a significance level of 5% to assess the significance of the treatment means (Steel and Torrie 1962).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003e3.1 Growth attributes of camelina responses under the separate and combined effects of Si and Se under salinity stress\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eRecorded results of growth attributes, such as root-shoot length and their fresh as well as dry weight, indicate that the different treatments of silicon (Si) and selenium (Se) separately or in combination significantly (P \u0026ge; 0.005) affected all growth attributes under SS conditions (Table 1). A graphical representation illustrates that SS significantly reduced the measured attributes of growth, such as shoot length, shoot fresh weight, shoot dry weight, root length, root fresh weight, and root dry weight in both camelina genotypes, Australian camelina, and Canadian camelina. The highest reduction of shoot length (9.00 cm), shoot fresh weight (11.2 g), shoot dry weight (3.01 g), root length (15.01 cm), root fresh weight (13.89 g), and root dry weight (2.87 g) were noted in Canadian camelina compared to Australian camelina under SS conditions ((Figure 1 a-f).\u003c/p\u003e\n\u003cp\u003eThe different doses of Si and Se also significantly improved the measured growth characteristics of both camelina genotypes. The maximum shoot length (32.55 cm), shoot fresh weight (31.24 g), shoot dry weight (6.93 g), root length (38.91 cm), root fresh weight (33.15 g), and root dry weight (7.63 g) were recorded in the treatment where Si and Se were applied in combination (Si + Se @ 10 ppm and NaCl @ 100 ppm) compared to all other treatments in Australian camelina, while the lowest values of shoot length, shoot fresh weight, shoot dry weight, root length, root fresh weight, and root dry weight were noted in Canadian camelina under SS conditions (Figure 1 a-f).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 1: Analysis of variance table for growth parameters, leaf chlorophyll pigments, infrared gas analyzer attributes, biochemical, and antioxidant assays of camelina under s7alinity stress conditions\u003c/strong\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" rowspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMeasured attributes\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"70.70707070707071%\" colspan=\"3\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSource of variance\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"32.857142857142854%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTreatments (T)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.285714285714285%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eVarieties (V)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"32.857142857142854%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eT x V\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eShoot length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e369.77 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e34.748 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e1.768 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eShoot fresh weight\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e11.915 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.691 NS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.115 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eShoot dry weight\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e248.401***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e26.940**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.497 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eRoot length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e333.464 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e42.563 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.348 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eRoot fresh weight\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e270.231 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e19.520 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.974 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eRoot dry weight\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e15.294 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e1.166 *\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.016 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eChlorophyll \u003cem\u003ea\u003c/em\u003e content\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.361 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.029 *\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.0010 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eChlorophyll \u003cem\u003eb\u003c/em\u003e content\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.447 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.030 *\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.0007 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eTotal chlorophyll content\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e1.603 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.119 *\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.0024 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eCarotenoids content\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e1.282 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.102 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.0012 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003ePhotosynthetic rate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e118.31 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e11.55 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.414 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eTranspiration rate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e6.2588 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.8453 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.0314 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eStomatal conductance\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.5262 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.0060 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.00008 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eTotal proline\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e10155 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e899.2 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e15.6 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eTotal soluble sugar\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e5952.7***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e525.29 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e23.09 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eTotal soluble protein\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e70.497 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e3.763 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.162 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eTotal free amino-acid\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e555.09 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e40.223 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e1.177 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eAPX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e2.433 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e0.243 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e0.0038 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eCAT\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e11799 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e1190.5 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e36.2 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003ePOD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e13567 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e1373.5 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e29.2 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"29.292929292929294%\" valign=\"top\"\u003e\n \u003cp\u003eSOD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e9258 ***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e598.2 **\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.232323232323232%\" valign=\"top\"\u003e\n \u003cp\u003e18.38 \u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e***\u003cem\u003eP\u0026nbsp;\u003c/em\u003e\u0026le; 0.001, **\u003cem\u003eP\u0026nbsp;\u003c/em\u003e\u0026le; 0.01, *\u003cem\u003eP\u0026nbsp;\u003c/em\u003e\u0026le; 0.05; \u003cem\u003eNS\u003c/em\u003e, non-signifcant\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.2 Leaf chlorophyll pigments of camelina responses under the separate and combined effects of Si and Se under salinity stress\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eRecorded results of leaf chlorophyll pigments, including chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid content, indicate that the different treatments of silicon (Si) and selenium (Se), separately or in combination, significantly (P \u0026ge; 0.005) affected all leaf chlorophyll pigments under SS conditions (Table 1). A graphical representation illustrates that SS significantly reduced the measured attributes of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid content in both camelina genotypes, (Australian camelina, and Canadian camelina). The highest reduction of chlorophyll a (0.89 mg/g FW), chlorophyll b (0.18 mg/g FW), total chlorophyll (1.07 mg/g FW), and carotenoid content (1.41 mg/g FW) was noted in Canadian camelina compared to Australian camelina under SS conditions (Figure 2 a-d).\u003c/p\u003e\n\u003cp\u003eThe different doses of Si and Se also significantly improved both camelina genotypes\u0026apos; measured leaf chlorophyll pigments. The maximum chlorophyll a (1.69 mg/g FW), chlorophyll b (1.02 mg/g FW), total chlorophyll (2.71 mg/g FW), and carotenoid content (2.89 mg/g FW) were recorded in the treatment where Si and Se were applied in combination (Si + Se @ 10 ppm and NaCl @ 100 ppm) compared to all other treatments in Australian camelina. In contrast, the lowest values of chlorophyll a, chlorophyll b, total chlorophyll, and carotenoid content were noted in Canadian camelina under SS conditions (Figure 2 a-d).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.3 Gas exchange attributes of camelina responses under the separate and combined effect of Si and Se under salinity stress\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll leaf gas exchange attributes under SS conditions were significantly (P \u0026ge; 0.005) affected by the various treatments of silicon (Si) and selenium (Se), either separately or in combination, according to recorded results of gas exchange attributes, including photosynthetic rate, transpiration rate, and stomatal conductance (Table 1). A graphical depiction shows that in both camelina genotypes (Australian and Canadian), SS dramatically decreased the measured characteristics of photosynthetic rate, transpiration rate, and stomatal conductance. Under SS conditions, Canadian camelina showed the greatest drop in photosynthetic rate (3.54 \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e), transpiration rate (0.48 \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e), and stomatal conductance (0.09 \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e) when compared to Australian camelina (Figure 3 a-c).\u003c/p\u003e\n\u003cp\u003eThe assessed gas exchange properties of both camelina genotypes were significantly improved by the varied dosages of Si and Se. When Si and Se were applied together (Si + Se @ 10 ppm and NaCl @ 100 ppm), the maximum photosynthetic rate (17.02 \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e), transpiration rate (3.65 \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e), and stomatal conductance (0.38 \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e) were recorded in Australian camelina as compared to all other treatments. On the other hand, under SS circumstances, Canadian camelina showed the lowest values of transpiration rate, stomatal conductance, and photosynthetic rate (Figure 3 a-c).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.4 Biochemical characters of camelina responses under the separate and combined effect of Si and Se under salinity stress\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eUnder SS conditions, all biochemical characters, including total proline, total soluble sugar, total soluble protein, and total free amino acid, showed significant (P \u0026ge; 0.005) effects from the different treatments of silicon (Si) and selenium (Se), either separately or in combination (Table 1). A graphical representation demonstrates that SS significantly reduced the observed features of total proline, total soluble sugar, total soluble protein, and total free amino acid in both Australian and Canadian camelina genotypes. Comparing Canadian and Australian camelina under SS, the former displayed the largest declines in total proline (71.32 \u0026micro;mol/g DW), total soluble sugar (31.32 \u0026micro;mole/g FW), total soluble protein (1.79 mg/g DW), and total free amino acid (8.41 mg/g DW) were observed in Canadian camelina (Figure 4 a-d).\u003c/p\u003e\n\u003cp\u003eThe different doses of Si and Se resulted in a significant improvement in the measured gas exchange characteristics of both camelina genotypes. In comparison to all other treatments, the maximum levels of total proline (195.15 \u0026micro;mol/g DW), total soluble sugar (120.95 \u0026micro;mole/g FW), total soluble protein (12.02 mg/g DW), and total free amino acid (37.15 mg/g DW) were recorded in Australian camelina when Si and Se were applied together (Si + Se @ 10 ppm and NaCl @ 100 ppm). Conversely, under SS conditions, total proline, total soluble sugar, total soluble protein, and total free amino acid values in Canadian camelina were the lowest (Figure 4 a-d).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.5 Antioxidants assay of camelina responses under the separate and combined effects of Si and Se under salinity stress\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eUnder SS circumstances, all antioxidant enzymes were significantly (P \u0026gt; 0.005) impacted by the various treatments of silicon (Si) and selenium (Se), either independently or in combination, according to recorded results of antioxidant activities, including APX, CAT, POD, and SOD (Table 1). A graphical depiction shows that in both camelina genotypes (Australian and Canadian), the SS dramatically decreased the observed characteristics of APX, CAT, POD, and SOD. When comparing Canadian camelina to Australian camelina under SS conditions, the greatest reductions in APX (1.95 Unit/min/g FW), CAT (29.36 Unit/min/g FW), POD (34.77 Unit/min/g FW), and SOD (126.42 Unit/min/g FW) were observed (Figure 5 a-d).\u003c/p\u003e\n\u003cp\u003eThe measured leaf chlorophyll pigments of both camelina genotypes were significantly enhanced by the various dosages of Si and Se. When Si and Se were treated in combination (Si + Se @ 10 ppm and NaCl @ 100 ppm), the greatest APX (3.89 Unit/min/g FW), CAT (165.19 Unit/min/g FW), POD (178.95 Unit/min/g FW), and SOD (237.63 Unit/min/g FW) were reported in Australian camelina as compared to the other treatments. On the other hand, under SS circumstances, Canadian camelina showed the lowest values of APX, CAT, POD, and SOD (Figure 5 a-d).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eDue to the susceptibility of crop plants to varying levels of SS, salinity has posed a daunting challenge to oil-seed crop productivity\u0026nbsp;(Abdi et al. 2023). The deleterious impacts of SS on multiple physiological systems, including photosynthesis, ion homeostasis, and antioxidant enzyme activity, have been reported in a wide array of crop plants. The increase in Na+ and Cl-ions in many plants has frequently been linked to the detrimental effects of salt; as a result, these ions provide essential circumstances for plant life by interfering with various plant processes\u0026nbsp;(Pourebrahimi et al. 2023). The current study showed that all measured growth attributes such as root-shoot length and their fresh as well as dry weight (Figure 1 a-f) were reduced under SS (NaCl was applied at the rate of 100 ppm). The same results were observed in the previous work, they stated that the SS significantly reduced the growth characteristics such as root-shoot length and their fresh as well as dry weight in plants\u0026nbsp;(Hosseinpour et al. 2020; Noreen et al. 2021). The reduction in the growth attributes of the plant might be attributed to more accumulation of salt in the soil, which led to reduced water potential nutrient uptake, and ultimately growth attributes were seriously reduced under SS conditions\u0026nbsp;(Al-Farsi et al. 2020a; Ahmad et al. 2023). However, in this study, exogenous application of Si and Se either separately or in combination remained effective in ameliorating the SS effects by boosting growth attributes in both camelina genotypes (Australian camelina and Canadian camelina). The height improvement was observed in all growth parameters where Si + Se was applied at the rate of 10 ppm under 100 pm NaCl stress conditions (Figure 1 a-f). Similar findings have previously been reported whereby the exogenous application of Si and Se imparted tolerance against SS by modulating physiological processes\u0026nbsp;(Abdi et al. 2023). The role of Si in the improvement of growth parameters in different crops was noted and recorded in the previous work which showed that the foliar application of Si improved the growth and development of plants under SS conditions\u0026nbsp;(Farhangi-Abriz and Torabian 2018; Mordhi Radhi and Alkarawia 2021). Similarly, the exogenous application of Se also played an important role in the enhancement of growth attributes of plants by restricting the uptake of salts\u0026nbsp;(Hossain et al. 2021). Moreover, \u0026nbsp;Si and Se when applied in combination showed more pronounced positive responses in the enhancement of the growth as well as the physiological process of plants under SS conditions\u0026nbsp;(Rajput et al. 2021; Taha et al. 2021).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eUnder SS, Plants experienced oxidative stress due to increased ROS generation, which disrupted the normal metabolism of plant cells which led to decreased growth attributes and crop productivity\u0026nbsp;(da Luz Neto et al. 2023). Under these circumstances, ROS causes oxidative stress to proteins, chlorophyll, and cell membranes; in response, plants increase their antioxidant capacity\u0026nbsp;(Zhao et al. 2021). In the current study, some detrimental effects of NaCl-induced SS were noted on the physiological functions of the lemon verbena plant. In the environment around the roots of lemon plants, adding more NaCl increased proline, soluble sugar, and antioxidant enzymes while decreasing chlorophyll and relative water contents (RWC)(Ciriello et al. 2022). The findings demonstrated that under varying degrees of SS, foliar spraying lemon verbena with 10 \u0026micro;M Se enhanced the plant\u0026apos;s growth metrics and RWC. Furthermore, plants treated with Se exhibited a decreased leaf-level accumulation (LEL) and accumulation of MDA and H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e in comparison to control plants. This suggests that the plants were less susceptible to oxidative damage induced by salt. Oxidative damage has been linked to the build-up of MDA and H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e under stressful settings(Admasie et al. 2023). To impart tolerance against abiotic stresses, the antioxidant defense system consisting of both enzymatic and non-enzymatic components performed key role in scavenging the ROS\u0026nbsp;(Elsheery et al. 2020).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eHowever, under SS, the results of this trial demonstrated that the exogenous administration of Se boosted the proline, soluble carbohydrates, and total protein of lemon verbena plants. There have been prior reports on the build-up of proline and other osmolytes in stressed plants when exogenous Se is applied\u0026nbsp;(Xu et al. 2021; El-Badri et al. 2022). Proline plays a unique role in osmotic stress circumstances, and helps plants to withstand stress conditions by regulating osmotic pressure, eliminating free radicals, conserving energy, and protecting macromolecules\u0026nbsp;(Noreen et al. 2021). It was also deduced that by lowering oxidative damage and enhancing plant development in lemon verbena under SS, the exogenous administration of Se triggered the defense mechanisms. This was explained by the accumulation of osmolytes. According to reports in this regard, Se treatment changed the physiological processes of the plant, including the accumulation of protein and osmolytes, the enhancement of the antioxidant capacity, the maintenance of ion homeostasis, and the enhancement of the photosynthetic capacity, thereby reducing the effects of stress on the plant and promoting plant growth under SS condition\u0026nbsp;(Zahedi et al. 2019; Xu et al. 2021; Ghanbari et al. 2023). According to\u0026nbsp;Regni \u003cem\u003eet al.\u003c/em\u003e (2021), the administration of Se enhanced olive plants\u0026apos; resistance to abiotic stress by preventing oxidative damage to cells and controlling their water status. Furthermore, it has been documented that the foliar application of selenium (Se) remained effective in boosting the concentration of chlorophyll in plants through the preservation of chloroplast enzymes and the augmentation of photosynthetic pigment production\u0026nbsp;(Ghanbari et al. 2023).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe current investigation found that the SS significantly affected the leaf gas exchange attributes such as net photosynthetic rate (P\u003csub\u003eN\u003c/sub\u003e), transpiration rate (E), and stomatal conductance (g\u003csub\u003es\u003c/sub\u003e) while the application of foliar Si and Se either separately or in combination assisted cameline plants in the improving leaf gas exchange attributes in both camelina genotypes (Figure 3 a-c). In the studies conducted on oats by\u0026nbsp;(Qin et al. 2018)\u0026nbsp;and\u0026nbsp;(Shah et al. 2020), \u0026nbsp;revealed that SS promoted stomatal closure, which reduced the amount of carbon dioxide available in the leaves and prevented carbon fixation. This raised the amount of excitation energy that the chloroplasts were exposed to, increasing the production of reactive oxygen species (ROS) and oxidative stress. Furthermore, it was observed that the presence of salt led to a reduction in the gas exchange characteristics\u0026nbsp;(Ondrasek et al. 2022). By altering the PSII system and the chloroplast ultrastructure, salinity reduced stomatal conductance and inhibited transpiration and gaseous exchange\u0026nbsp;(Pan et al. 2021). A reduced supply of CO\u003csub\u003e2\u0026nbsp;\u003c/sub\u003eresulting from partial stomata closure and an altered biochemical binding mechanism for CO\u003csub\u003e2\u003c/sub\u003e could be the cause of the inhibition of plant absorption during SS\u0026nbsp;(Shah et al. 2020; Kukrić et al. 2023). This resulted in changes to cellular metabolism, a decrease in CO2 photosynthetic binding, and an increase in ROS production in chloroplasts. ROS harmed the photosynthetic apparatus, especially PSII, which resulted in photoinhibition. This was caused by an imbalance in the photosynthetic redox signaling pathways and suppression of PSII repair\u0026nbsp;(Yan et al. 2021; Liang et al. 2024). In a stressful environment, plants typically tend to close their stomata to preserve water, which also lowers the stomatal conductance and photosynthesis\u0026nbsp;(Zahra et al. 2022).\u003c/p\u003e\n\u003cp\u003eSe and/or Si-induced improvements in wheat growth features might be due to their roles in controlling physiological processes, such as photosynthesis, plant tissue water status, and antioxidant machinery\u0026nbsp;(Taha et al. 2021). Wheat growth and productivity were found to be positively impacted by the stimulating roles that Se and Si play in protein synthesis, chlorophyll synthesis, and the absorption of essential nutrients\u0026nbsp;(Sattar et al. 2017; Qin et al. 2024). Plants under SS benefitted from Si\u0026apos;s beneficial effects on their metabolic processes\u0026nbsp;(Mordhi Radhi and Alkarawia 2021). Additionally, it caused a reduction in the stomata lumen and silicifies the leaf surface, which both affected the rate of plant transpiration\u0026nbsp;(Sayed et al. 2022). The transport and absorption of several nutrients were enhanced by the exogenous application of Si\u0026nbsp;(Raza et al. 2019; Zia et al. 2023). In the same setting, applying Se resulted in an amino acid metabolism imbalance; as a result, stressed plants exhibited higher levels of soluble protein and lower nitrate activity\u0026nbsp;(Ahmad et al. 2021; Ghanbari et al. 2023). Similar to our findings, the Se application also remained instrumental in protecting plant cells, increased the formation of starch in chloroplasts, and supported plant growth in salt-stressed environments\u0026nbsp;(Regni et al. 2021).\u003c/p\u003e\n\u003cp\u003eOrganic osmolytes like proline and total soluble sugars help plants regulate their cell osmoregulation and lessen the effects of stress (uz Zaman et al. 2022). Osmoprotectant concentrations and stress tolerance are closely correlated, and they are regarded as one of the markers for the potential for stress tolerance (Al-Farsi et al. 2020b). Additionally, these decreased the negative effects of stress by controlling the synthesis of vital proteins like Rubisco, shielding the photosynthetic system, improving membrane stability, preserving the redox equilibrium, and eliminating ROS (Ahmad et al. 2020a). Plants responded to internal Si stress by boosting the production of metabolites through soluble sugars and proline. Through the osmotic balance, which preserved cell turgor and guaranteed the stability and safety of cellular membranes, these metabolites promoted tolerance against SS. This reduced photo-oxidation and oxidative damage (Farouk et al. 2020; Sayed et al. 2022). According to our research findings, applying Se and/or Si increased the levels of organic molecules such as proline and TSS that could be attributed to increased amylase activity and the hydrolysis of starch by selenium; fructose 1, 6-bisphosphatase, which is involved in the metabolism of carbohydrates. When compared to unstressed plants, wheat plants subjected to SS showed increased TSS and proline buildup in leaf tissue. The method of adaptation to salinity stress is reflected in these increases in proline and TSS concentrations in wheat leaves during SS. Spraying the wheat plants with Se, Si, and their mixtures resulted in a further increase in the accumulation of proline and TSS. Salinity in the soil causes osmotic stress and can reduce the amount of water that roots take in, which can eventually disrupt the activities of stomata and roots (Badawy et al. 2021; Xu et al. 2021; Abdi et al. 2023). This could be the cause of the decrease in stomatal conductance in wheat plants during SS. In the present study, plant water conditions and hydraulic conductivity under SS may have been mediated by an increase in TSS and proline buildup in response to both Se and Si, as well as their mixtures. Furthermore, wheat plants treated with Se/Si exhibited a substantial accumulation of proline, which may have contributed to ROS scavenging in the current study to reduce oxidative damage (Manzoor et al. 2024; Qin et al. 2024).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn conclusion, the research findings revealed the deleterious impacts of salinity stress on the growth, photosynthetic pigments, leaf gas exchange, and antioxidant activities of Camelina, as well as the potential benefits of foliar Si and Se application in mitigating these effects. The findings demonstrated that SS negatively affected Camelina\u0026apos;s growth attributes photosynthetic pigments and leaf gas exchange attributes indicating its vulnerability to salt-induced limitations. However, the foliar application of Si and Se in combination, particularly at higher concentrations (10 ppm), significantly improved the measured parameters compared to the control. The positive effects of Si and Se on Camelina under SS conditions can be attributed to their antioxidant properties and their ability to enhance the activity of antioxidant enzymes. Si and Se supplementation helped to scavenge reactive oxygen species (ROS) and minimize oxidative damage, thereby improving the overall tolerance of Camelina to salinity stress. The results also revealed genotype-specific variations in SS tolerance, with the Australian variety of Camelina displaying superior performance to the Canadian variety. This highlights the importance of considering genetic factors when studying plant responses to environmental stressors. The results of this investigation advance our comprehension of the systems underlying SS tolerance in Camelina and provide valuable insights for the development of strategies to enhance its productivity in saline environments. The application of Si and Se as foliar supplements shows promise as a practical approach to improving SS tolerance in Camelina crops.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e; Hesham F. Alharby and Zahoor Ahmad planned the research, Halima Nawaz and Zahoor Ahmad conducted the research work; Hameed Alsamadany and Muhammad Aamir Iqbal wrote the introduction and helped in the improvement of the manuscript\u0026apos;s quality; Zahoor Ahmad, Yahya M. Alzahrani, Muhammad Aamir Iqbal, and Awatif M. Abdulmajeed did the static analysis and graphical representation; Zahoor Ahmad read the manuscript as proofreading and arranged it according to the journal style; Zahoor Ahmad provided reagents, assisted in the analytical work; and Zahoor Ahmad, Muhammad Aamir Iqbal and Hesham F. Alharby improved the English language quality of the manuscript. All authors participate in the improvement of the quality of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was funded by Institutional Fund Projects under grant no. (IFPIP: 737-130-1443), Ministry of Education in Saudi Arabia.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgment:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research work was funded by Institutional Fund Projects under grant no. (IFPIP: 737-130-1443). The authors gratefully acknowledge the technical and financial support from the Ministry of Education and King Abdulaziz University, DSR, Jeddah, Saudi Arabia. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical Approval:\u0026nbsp;\u003c/strong\u003eAll authors approve this manuscript for submission to Environmental Science and Pollution Research.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to Participate:\u0026nbsp;\u003c/strong\u003eAll authors participate in the preparation of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to Publish:\u0026nbsp;\u003c/strong\u003eAll authors consent to the manuscript published in Environmental Science and Pollution Research.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interests:\u0026nbsp;\u003c/strong\u003eThe authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbdi MJ, Ghanbari Jahromi M, Mortazavi SN et al (2023) Foliar-applied silicon and selenium nanoparticles modulated salinity stress through modifying yield, biochemical attribute, and fatty acid profile of Physalis alkekengi L. 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Environ Sci Pollut Res. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11356-023-26902-9\u003c/span\u003e\u003cspan address=\"10.1007/s11356-023-26902-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"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":"environmental-science-and-pollution-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"espr","sideBox":"Learn more about [Environmental Science and Pollution Research](https://www.springer.com/journal/11356)","snPcode":"11356","submissionUrl":"https://submission.nature.com/new-submission/11356/3","title":"Environmental Science and Pollution Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Enhancing Salt tolerance, Camelina, Exogenous silicon and selenium, Morphological attributes, Chlorophyll pigments, Leaf Gas Exchange attributes, antioxidants","lastPublishedDoi":"10.21203/rs.3.rs-4317048/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4317048/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSalt stress (SS) has emerged as one of the most pressing environmental threats to the production sustainability of oil-seed crops like camelina which necessitates finding out pro-environment and biologically feasible amelioration approaches. A study was performed to investigate the effects of exogenously applied silicon (Si) and selenium (Se) on two camelina varieties (Australian and Canadian) under SS conditions. The trial was comprised of two doses of Si and Se (5 and 10 ppm each) along with two co-application treatments entailing Si (5pp)\u0026thinsp;+\u0026thinsp;Se (5ppm) and Si (10 pp)\u0026thinsp;+\u0026thinsp;Se (10 ppm), whereas control treatments were kept for comparison purpose. The response variables included root-shoot length, fresh and dry weights, as well as leaf photosynthetic pigment content (chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids), leaf gas exchange attributes, biochemical characters (total proline, total soluble sugar, total soluble protein, and total free amino acid) and antioxidant activities (APX, POD, CAT, and SOD). Results showed that SS significantly reduced all growth attributes of camelina, whereas the Australian camelina variety demonstrated greater salinity tolerance in comparison to the Canadian variety. The co-application of Si (10 pp)\u0026thinsp;+\u0026thinsp;Se (10 ppm) outperformed the rest of the treatments by recording the highest leaf chlorophyll a (1.69 mg/g FW), chlorophyll b (1.02 mg/g FW), carotenoids (2.89 mg/g FW), photosynthetic rate (17.02 \u0026micro;mole CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), transpiration rate (3.65 \u0026micro;mole CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), stomatal conductance (0.38 \u0026micro;mole CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), total proline (199.38 \u0026micro;mole/g DW), total soluble sugar (120.95 \u0026micro;mole/g FW), total soluble protein (12.02 mg/g DW) and total free amino acid (37.15 mg/g DW) in Australian camelina as compared to Canadian camelina. The same treatment also remained effective in triggering the activity of APX (3.89 Unit/min/g FW), CAT (165.19 Unit/min/g FW), POD (178.95 Unit/min/g FW), and SOD (237.63 Unit/min/g FW). Thus, the combined exogenous application of Si and Se holds bright perspectives in alleviating the deleterious impact of SS especially for the Australian variety, and could be recommended to camelina growers after thorough field investigations.\u003c/p\u003e","manuscriptTitle":"Boosting Salt Tolerance in Camelina cultivars through synergistic Impact of Exogenous Silicon and Selenium by modulating physiological attributes and Antioxidant activities","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-06-13 13:28:36","doi":"10.21203/rs.3.rs-4317048/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major Revision","date":"2024-07-10T19:17:01+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2024-06-01T13:37:57+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-05-31T01:16:48+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Environmental Science and Pollution Research","date":"2024-05-21T19:18:30+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-04-29T04:24:31+00:00","index":"","fulltext":""},{"type":"submitted","content":"Environmental Science and Pollution Research","date":"2024-04-25T03:29:19+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"environmental-science-and-pollution-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"espr","sideBox":"Learn more about [Environmental Science and Pollution Research](https://www.springer.com/journal/11356)","snPcode":"11356","submissionUrl":"https://submission.nature.com/new-submission/11356/3","title":"Environmental Science and Pollution Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"538608a9-4b8c-40e2-b0c9-394878ce944d","owner":[],"postedDate":"June 13th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"in-revision","subjectAreas":[],"tags":[],"updatedAt":"2024-07-10T23:17:32+00:00","versionOfRecord":[],"versionCreatedAt":"2024-06-13 13:28:36","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4317048","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4317048","identity":"rs-4317048","version":["v1"]},"buildId":"WrCJVZZCHTDjtuVLN7oU0","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}