Alleviation of salt-induced photosynthesis and growth inhibition by salicylic acid involves glycinebetaine and ethylene in mungbean (Vigna radiata L.)

Glycinebetaine-induced modulation of antioxidant enzymes activities and ion accumulation in two wheat cultivars differing in salt tolerance

BackgroundApplication of exogenous glycine betaine (GB) and exogenous salicylic acid (SA) mitigates the adverse effects of salinity. Foliar spraying with exogenous GB or SA alleviates salt stress in plants by increasing leaf gas exchange and stimulating antioxidant enzyme activity. The effects of foliar application of exogenous GB and SA on the physiology and biochemistry of cotton seedlings subjected to salt stress remain unclear.ResultsResults showed that salt stress of 150 mM NaCl significantly reduced leaf gas exchange and chlorophyll fluorescence and decreased photosynthetic pigment quantities and leaf relative water content. Foliar spray concentrations of 5.0 mM exogenous GB and 1.0 mM exogenous SA promoted gas exchange and fluorescence in cotton seedlings, increased quantities of chlorophyll pigments, and stimulated the antioxidant enzyme activity. The foliar spray also increased leaf relative water content and endogenous GB and SA content in comparison with the salt-stressed only control. Despite the salt-induced increase in antioxidant enzyme content, exogenous GB and SA in experimental concentrations significantly increased the activity of glutathione reductase, ascorbate peroxidase, superoxide dismutase, catalase and peroxidase, and decreased malondialdehyde content under salt stress. Across all experimental foliar spray GB and SA concentrations, the photochemical efficiency of photosystem II (FV/FM) reached a peak at a concentration of 5.0 mM GB. The net photosynthetic rate (Pn) and FV/FM were positively correlated with chlorophyll a and chlorophyll b content in response to foliar spraying of exogenous GB and SA under salt stress.ConclusionsWe concluded, from our results, that concentrations of 5.0 mM GB or 1.0 mM SA are optimal choices for mitigating NaCl-induced damage in cotton seedlings because they promote leaf photosynthesis, increase quantities of photosynthetic pigments, and stimulate antioxidant enzyme activity. Among, 5.0 mM GB and 1.0 mM SA, the best performance in enhancing endogenous GB and SA concentrations was obtained with the foliar application of 1.0 mM SA under salt stress.

Physiological Mechanism of Salicylic Acid for Alleviation of Salt Stress in Rice

Green gram (Vigna radiata) is considered the chief legume in Pakistan. Thus, current study was conducted to examine the ameliorating effect of phytohormones pre-treatments under salt stress on proteome profile of green gram by sodium-dodecyl-sulphate-polyacrylamide gel electrophoresis (SDS-PAGE). The soluble green gram seedlings proteins were resolved on 4% stacking and 12% resolving gels. The SDS-PAGE resolved 24 polypeptide bands ranging from 200 to 17kDa. Among these, 12 out of 24 bands of proteins were essentials house-keeping or growth proteins of green grams. While, 120, 114.6, 51.8, 29.1, and 22.8 kDa bands were over-expressed under 50 to 350mM salt with phytohormones treatments. The others 104.5 kDa, 99.8 kDa, 95.3 kDa, 91.0 kDa, 55 kDa, 46 kDa, and 17kDa bands were related to the GAᴣ, IAA, and SA induced tolerance. Overall 120 kDa, 114.6 kDa, 104.5 kDa, 99.8, 95.3 kDa, 51.8 kDa, 29.1 kDa and 22.8kDa bands were first time identified in the current study. The information retrieved from NCBI protein database, the resolved peptides were principally belonging to 7S and 8S vicilin, 2S, 8S, 11S, and 16.5S globulins. It is determined that seed priming with SA enhanced tolerance in green gram by rapidly synthesizing stress alleviating peptides.Key word: Cluster analysis, dendrogram, mungbean, salt stress, SDS-PAGEINTRODUCTIONVarious world-wide health concerning organization recommended the use of high graded plant protein such as legumes to prevent the risk of metabolic disorder (Hou et al., 2019). Legumes are most important protein crop on the earth. Among the legumes, the green gram is the major pulses. Its seeds are rich in superior quality storage protein, which account 85% of the total protein while, another 15% have not been broadly studied (Yi-Shen et al., 2018). The soluble storage protein comprises of 60% globulins, 25% albumin and 15% prolamins. Globulins are further divided into 3.4% basic-type (7S), 7.6% legumin-type (11S), and 89% vicilin-type (8S) (Mendoza et al., 2001; Itoh et al., 2006). Other than proteins, the green gram seeds also contain starch, fiber, phenolic compound, saponins, vitamins, calcium zinc, potassium, folate, magnesium, manganese and very low in fat that made it meager man’s meat (Hou et al., 2019). It is also a good source of green manure and fodder (El-Kafafi et al., 2015). Its root has ability to fix 30 to 50 Kg/ha atmospheric nitrogen in the soil which is essential for maintaining soil fertility (Chadha, 2010). The green gram is the valuable and the major Rabi pulse crop of Pakistan. Its cultivation area in 2016-2017 was about 179,000 hectares with seed yield of 130,000 tones. In comparison during 2017-2018, it was cultivated on 161,800 hectares land with 118,800 tones seed yield (GOP, 2018). One of the reasons of this 9% decrease in both land and productivity is the shortage of irrigated land due to soil salinity. The salinity induce oxidative bust in the mungbean cells, caused by responsive oxygen species (ROS) such as hydrogen peroxide, singlet oxygen, hydroxyl radical and superoxide radical. The ROS create hindrance in various metabolic processes of plant via interacting with macromolecules like proteins (Alharby et al., 2016). However, phytohormones like gibberellic acid (GAᴣ), indole acetic acid (IAA), and salicylic acid (SA) take part in the biosynthesis of salt tolerance proteins under salinity. These salt tolerance proteins acclimate plants under salinity stress. Application of biotechnology plays a significant role in agriculture (Khan et al., 2017). Therefore, production of particular proteins under salt stress is a specific response of cell which can be analyzed by sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS-PAGE). SDS-PAGE is the simple, valid, and cost-effective biochemical marker (Mushtaq et al., 2018). This marker has been widely used to determine the extent of evolutionary variations in crops (El-Kafafi et al., 2015).OBJECTIVES The present study was directed first time with the aim to investigate the toxic effect of sodium chloride (0-350 mM) and stress acclimation by pre-treatment of GAᴣ, IAA, and SA on the proteome profile of NM-92 cultivar of a Pakistani green gram.MATERIALS AND METHODSThe present study was replicated thrice in the plant laboratory of Department of Genetics, Faculty of Science, and University of Karachi. The seeds of mung bean cultivar NM-92 were acquired from National Agricultural Research Centre (NARC), Islamabad. These freshly collected 15 seedsˉ1 treatment / replication were divided into two sets. The first was named as sodium chloride (SC) stress treatments were imbibed in sterile distilled water (DW) whereas, second set soaked in gibberellic acid (GAᴣ) (BDH Chemicals, England), indole acetic acid (IAA) (Fluka, Switzerland), and salicylic acid (SA) (J.T. Baker, Holland) in the separate beaker for 24 hours under dark condition. After 24 hours, given ample time to both the sets at room temperature. After recovery, all 20 treatments were sown in the 150 X 30 mm sized petri-dishes containing 0, 50, 150, 250, and 350 millimolar (mM) sodium chloride solution (Fisher Scientific, UK) for 72 hours.Protein extraction: Protein extraction was done by taking 0.3g of seedlings in an ice chilled mortar and crushed by adding 600µL 0.2 M Tris-HCl buffer having pH 7.5 contained 5% SDS (w/v) and 5% 2-mercaptoethanol (v/v). The homogenate was incubated at 0oC for 30 min., boiled in the water bath for 3 min. at 100oC. Samples were centrifuged in Heraeus Biofuge D-37520, Germany for 30 min. at 8000 rpm. The protein supernatant was saved at below 0°C for quantitative and qualitative determination with minor modifications. The total soluble protein content of the samples was estimated via “Bovine Serum Albumin (BSA) standard curve” and explicit in µg protein milligramˉ1 fresh weight of mung seedlings.Bovine serum albumin standard curve (2000 μg/mL): Total protein standard curve was made by dissolving 0.05g of Bovine Serum Albumin (BSA) in 25mL of distilled water. Ten serial dilutions were made from 0.1 mL to 1mL by BSA solution then performed Lowry. A standard curve of total proteins was plotted by taking BSA absorbance at Y-axis and 2000 μg BSA / mL at X-axisSample preparation for SDS-PAGE: For qualitative assessment of total proteins; the 35μL of saved protein supernatant was combined with 15μL of sample diluting buffer (SDB). The SDB was made up of 0.0625 M Tris-HCl pH 6.8 with 2% of SDS, 10% of glycerol, 0.003% of bromophenol blue dye and 5% of 2-mercaptoethanol. Boil the 50μL protein SDB supernatant at 100oC in water bath for 3 min., centrifuged at 6000 rpm for 4 min. The supernatant was loaded on SDS-PAGE gel with the given formulae. The SDS- PAGE: Total proteins were fractionated via SDS-PAGE with 4% stacking and 12% resolving gel. The resolving gel of 12% was made by taking 6mL solution A, 1.8 mL 3 M Tris 1 M HCl buffer pH 8.8, 144μL 10% SDS, 5.74 mL sterile distilled water, 720μL 1.5% ammonium persulphate (APS) in deionized water and 10μL TEMED. While, stacking was composed of 1.25mL of solution A, 2.5mL of 0.5M Tris 1M HCl buffer pH 6.8, 100μL 10% SDS, 1.8 mL of distilled water, 500μL 1.5% APS and 12μL TEMED. Solution A was prepared by conjoining 30% acrylamide and 0.8% N, N’-methylene-bisacrylamide in deionized water. To avoid polymerization in the beaker; the prepared solution was quickly poured into the 3 mm thick gel plates after adding TEMED. The stacking was lined over resolving gel, then combs were inserted between the gel plates of SCIE-PLAS TV-100 separation system, UK, and allowed to polymerize for ½ an hour. After polymerization gel was placed in the tank which were filled with Tris-Glycine buffer (electrode buffer) pH 8.4 then combs were removed. The electrode buffer contained 0.3% Tris, 1.41% Glycine and 0.1% SDS in 2000mL d/w. The gel was pre-run for 15 min. at 60 volts and 120 mA currents. The prepared SDS-PAGE samples were loaded in wells with BlueStepTM Broad Range Protein Marker, AMRESCO, USA as standard and run at 60 volts & 120 mA for about 45 min. When samples entered in resolving gel, and then gave 100 volts and 200 mA currents for around 2.5 hours. Furthermore, electrophoresis was carried out at a constant watt.The Gel was washed with 30% ethanol on Uni Thermo Shaker NTS-1300 EYELA, Japan at the constant shaking for 30 min. Then gels were placed in 10% glacial acetic acid in 50% methanol solution (Fixative) for 24 hours. SDS Gel was stained until protein bands were visible thereat placed as 5% of Methanol in 7.5% acetic acid glacial solution to destain the bands background. SDS-PAGE stain composed of 0.125% coomassie brilliant blue R-250 dissolved in 40% of Methanol and 7% acetic acid glacial solution. The stain was stirred on Magnetic stirrer & hot plate M6/1, Germany for 6-10 hours before used. Photographs were taken by Sanyo digital camera VPC-T1284BL and bands were scored through numbering pattern. Gels preserved in 10% acetic acid solution at 4°C.Interpretation of bands and data analysis: The total soluble protein bands relative mobility calculated by below formulae and Dendrogram was constructed via SPSS v. 20Where,F=(Migrated distance of protein band)/(Migrated distance of dye front)Slop=(Log MW of protein marker lower limit band–log〖MW of protein marker upper limit band )/(RF protein marker lower limit band –RF of protein marker upper limit band)RESULTS:The total soluble proteins extracted from green gram were perceived by SDS-PAGE Blue StepTm broad range biochemical markers. The protein-based marker was used to evaluate the toxic effect of sodium chloride along with pre-treatments of GAᴣ, IAA, and SA on proteome assay. In the current work, seedlings total soluble proteome resolved 24 polypeptide bands ranging from 200 to 17.1 kDa were recognized by using SDS-PAGE. The figure 1 showed Dendrogram assay, which classified the 20 treatments of SC, GAᴣ, IAA and SA into two major cl

The influence of exogenously applied glycinebetaine (0, 50 and 100 mm) as a foliar spray at different growth stages, i.e. vegetative, reproductive or both at the vegetative and reproductive stages on gas exchange characteristics, glycinebetaine (GB) and the activities of antioxidant enzymes, such as superoxide dismutase (SOD), catalase (CAT) and peroxidase (POD) was examined in plants of two maize cultivars, Golden and C‐20 grown under saline conditions. Salt stress caused a marked decrease in photosynthetic capacity, chlorophyll contents and SOD activity in both maize cultivars. However, activities of CAT and POD remained almost unchanged in both maize cultivars under salt stress. Accumulation of GB increased with an increase in exogenous level of GB, i.e. 100 mm GB spray caused a greater accumulation of GB in the leaves of maize plants than did 0 or 50 mm. Although exogenously applied GB enhanced photosynthetic capacity of salt‐stressed plants of both cultivars, it enhanced the activities of antioxidant enzymes, SOD, CAT and POD, in salt‐stressed plants of cv. C‐20 only. Overall, the adverse effects of salt stress on maize plants were alleviated by the exogenous application of GB at different growth stages, which up‐regulated photosynthetic capacity and the activities of some antioxidant enzymes.

Halotolerant species are of interest since they occur naturally in environments with excess toxic ions. The cyanobacterium Halothece sp. PCC 7418 (hereafter referred to as Halothece) exhibits remarkable halotolerance and was used to examine stress-responsive regulatory mechanisms. The effects of five different stimuli on Halothece transcriptomes were examined using RNA sequencing. In response to diverse stresses, there were both common and stress-specific transcriptional responses. A common upregulated gene set under all stresses consisted of nine differentially expressed genes (DEGs). We also found that osmotic stress elicited the largest set of DEGs. Salt- and osmotic-responsive regulatory mechanisms shared common pathways. DEGs that were upregulated under salt stress encoded proteins involved in photosynthesis and related machineries. Furthermore, DEGs encoding two-component system (TCS) factors, transcriptional factors, scaffolds for protein-protein interactions, transporters, protein turnover factors, and lipid biosynthesis enzymes were also identified under salt stress. Notably, one-carbon (1C) metabolism factors, glycine betaine (GB) synthesis enzymes, and GB transporters were upregulated under salt stress. Metabolic analyses revealed that GB accumulated under salt stress, while mycosporine-2-glycine (M2G) accumulated under salt or osmotic stress. None of the nutrient starvations induced GB nor M2G accumulation. These results suggested that GB and M2G are two osmoprotectants that contribute to halotolerance. Based on our results, we proposed regulatory mechanisms that are crucial for halotolerance, which are coordinated with the GB, M2G, 1C, amino acid, and central carbon interlinking metabolic pathways. 1C metabolism directly fulfills the high metabolite requirements for halotolerance together with the ancillary role of several metabolic pathways.Key Points• Global transcriptome surveys together with molecular and metabolite analyses provide insights into regulatory networks that are crucial for halotolerance• Regulatory networks that are crucial for halotolerance coordinated with the two key osmoprotectants, one carbon, amino acid, and central carbon interlinking metabolic pathways• The findings have translational relevance in genomic and transcriptomic mechanisms of halotolerance.

Level of osmolytes accumulation to a variable extent in bacteria and plants is associated with the salt tolerance.We hypothesized that two previously isolated salt tolerant strains, Halomonas meridiana (PAa6) and Halomonas aquamarina (RT2), found to stimulate the growth of wheat also have the ability to accumulate osmolytes.These strains can favour plant growth due to their endogenous osmolyte accumulation.To ascertain this, strains were checked for osmolyte accumulation under salt stress.Growth and osmolyte accumulation of inoculated and non-inoculated seeds of the economically significant plant, Zea mays Var.EV.90, was recorded.Seeds were sown in soil supplemented with and without exogenous proline and glycine betaine and salt stress for 15 days.The results show that strains can accumulate osmolytes (preferably glycine betaine) at higher salt stress.Inoculation and exogenous osmolytes (10 mM) improved the plant growth at higher salt stress.However, concurrent application had an amplified effect.Endogenous level of osmolyte (preferably glycine betaine) on the fresh weight basis was significantly higher and improved bacterial and plant growth under stress.We conclude that accumulation of osmolytes in bacteria and plants significantly stimulated growth and protect them from the adverse effect of salt stress.This can be an alternate economic approach to increase crop productivity.

Salinization of agricultural lands, particularly rice paddies, results in the drastic decline of crop yields. Soil salinization impacts the plant physiology by inducing salt stress which may leads to osmotic stress, ionic stress and water-related nutrient imbalance. These imbalances necessitate the need for plants to produce osmolytes including proline and glycine betaine. This study aimed to elucidate the dynamic changes in proline and glycine betaine accumulation modulated by the inoculation of Brevibacterium linens RS16 in salt-sensitive and moderately salt-tolerant rice plants under salt stress conditions. This study showed the interaction of four major factors including rice genotypes with differing tolerance to salt stress, length of exposure to salt stress, level of salt stress and effects of inoculation. Salt stress resulted in significant reduction in plant growth parameters with the salt-sensitive rice genotype (IR29) having a more significant growth reduction. Both the salt-sensitive and salt-tolerant rice genotypes increased in total proline and glycine betaine accumulation at 3 days and 10 days after subjecting under 50 mM and 150 mM salt stress conditions. A significant increase in proline and glycine betaine was observed in the salt-sensitive genotype after 10 days under 50 mM and 150 mM salt stress conditions. Inoculation of the rice genotypes with B. linens RS16 resulted in the improvement of plant growth parameters in both rice genotypes, and total proline and glycine betaine accumulation, especially in IR29. This study showed that proline and glycine betaine are compatible osmolytes of rice under salt stress, and that inoculation of rice genotypes with B. linens RS16 mediated salt tolerance through improvement of plant growth parameters and proline and glycine betaine accumulation in rice plants.

As an antioxidant, alpha-tocopherol (α-Toc) protects plants from salinity-induced oxidative bursts. This study was conducted twice to determine the effect of α-Toc as a foliar spray (at 0 (no spray), 100, 200, and 300 mg L−1) to improve the yield and biochemical constituents of fresh green capsules of okra (Abelmoschus esculentus L. Moench) under salt stress (0 and 100 mM). Salt stress significantly reduced K+ and Ca2+ ion concentration and yield, whereas it increased H2O2, malondialdehyde (MDA), Na+, glycine betaine (GB), total free proline, total phenolics, and the activities of catalase (CAT), guaiacol peroxidase (GPX), and protease in both okra varieties (Noori and Sabzpari). Foliar application of α-Toc significantly improved the yield in tested okra varieties by increasing the activity of antioxidants (CAT, GPX, SOD, and ascorbic acid), accumulation of GB, and total free proline in fruit tissues under saline and non-saline conditions. Moreover, α-Toc application as a foliar spray alleviated the adverse effects of salt stress by reducing Na+ concentration, MDA, and H2O2 levels and improving the uptake of K+ and Ca2+. Among the tested okra varieties, Noori performed better than Sabzpari across all physio-biochemical attributes. Of all the foliar-applied α-Toc levels, 200 mg L−1 and 300 mg L−1 were more effective in the amelioration of salinity-induced adverse effects in okra. Thus, we concluded that higher levels of α-Toc (200 mg L−1 and 300 mg L−1) combat salinity stress more effectively by boosting the antioxidant potential of okra plants.

Salicylic acid (SA) is a common, plant-produced signal molecule that is responsible for inducing tolerance to a number of biotic and abiotic stresses. Our experiment was therefore conducted to test whether the application of SA at various concentrations (0, 0.10, 0.50, and 1.00 mM) as a foliar spray would protect citrus seedlings (Valencia orange/Bakraii) subjected to salt stress (0, 25, 50, and 75 mM NaCl). Growth parameters, leaf chlorophyll (Chl) content, relative water content (RWC), maximal quantum yield of PSII photochemistry (Fv/Fm), and gas-exchange variables were negatively affected by salinity. In addition, leaf electrolyte leakage (EL) and proline content increased by salinity treatments. Application of SA increased net photosynthetic rate and proline content in salt stressed plants and may have contributed to the enhanced growth parameters. SA treated plants had greater Chl content and RWC compared with untreated plants when exposed to salt stress. Fv/Fm ratio and stomatal conductance were also significantly higher in SA treated plants under saline stress conditions. SA application reduced EL compared to untreated plants, indicating possible protection of integrity of the cellular membrane. It appeared that the best ameliorative remedies of SA were obtained when Valencia orange/Bakraii seedlings were sprayed by 0.50 and 1.00 mM solutions. Overall, the adverse effects of salt stress could be alleviated by exogenous application of SA.

Salinity stress may involve the accumulation of glycine betaine (GB). The objective of this study was to examine whether exogenous GB would ameliorate the detrimental effect of salinity stress on perennial ryegrass ( Lolium perenne ). The grass was subjected to two salinity levels (0 and 250 m m NaCl) and three GB levels (0, 20, and 50 m m ). Salinity resulted in a remarkable decrease in vertical shoot growth rate (VSGR), shoot and root fresh weight, relative water content (RWC), relative transpiration rate (Tr), and chlorophyll (Chl) content, superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX) activities. Plants subjected to salt exhibited an increase in leaf electrolyte leakage (EL), lipid peroxidation (MDA), and proline content. Application of GB reduced EL, MDA, and proline content in salt-stressed plants. Perennial ryegrass subjected to salt stress plus GB had a greater level of VSGR, RWC, relative Tr, Chl content, and activities of SOD, CAT, and APX when compared with salt-stressed without GB. Salt stress increased Na + and decreased K + content, which resulted in a higher Na + /K + ratio in perennial ryegrass. Application of 20 m m GB suppressed Na + accumulation, whereas the K + content was significantly increased in shoot, which led to a higher K + /Na + ratio under saline conditions. These results suggested that GB-enhanced salt tolerance in perennial ryegrass was mainly related to the elevated SOD, CAT, and APX activity and alleviation of cell membrane damage by reducing oxidation of membrane lipid and improving the ion homeostasis under salt stress.

The study was conducted to evaluate the effect of Salicylic acid (SA) as a foliar spray to mitigate the deleterious effects of salt stress on mungbean plants. Two mungbean genotypes viz; Shikha and Heera were grown under induced saline conditions (100, 150 and 200 mM NaCl) and compared with untreated control. Salt exposure decreased the morpho-physiological, biochemical and yield traits of both the genotypes, which was responded in the same way under induced salt stress. SA found to be essential in mitigating the adverse effects of salt stress in both tested mungbean genotypes. At the threshold combination of SA and NaCl (100 ppm SA + 100 mM NaCl), SA led to maximum plant height, number of leaves per plant, relative water content (RWC), chlorophyll content, number of pods and seed yield. Its foliar application was also expressed positive response in modulating the total proline content under different salt stress conditions. It was also concluded that foliar application of SA @100 ppm was highly effective in reducing the deleterious effect of salt at the combination with 100 mM NaCl and 150 mM NaCl and strengthened the ability of survival of both the mungbean genotypes under salt stress.

Physiological and transcriptional variations inducing complex adaptive mechanisms in grapevine by salt stress

Homeobox 2 promotes betaine aldehyde dehydrogenase gene expression and biosynthesis of glycine betaine in 'Baxijiao' banana under salt stress.

The objective of this study was to determine the effect of foliar salicylic acid (SA) applications on growth, chlorophyll, and mineral content of cucumber grown under salt stress. The study was conducted in pot experiments under greenhouse conditions. Cucumber seedlings were treated with foliar SA applications at different concentrations (0.0, 0.25, 0.50, and 1.00 mM). Salinity treatments were established by adding 0, 60, and 120 mM of sodium chloride (NaCl) to a base complete nutrient solution. The SA was applied with spraying two times as before and after transplanting. Salt stress negatively affected the growth, chlorophyll content and mineral uptake of cucumber plants. However, foliar applications of SA resulted in greater shoot fresh weight, shoot dry weight, root fresh weight, and root dry weight as well as higher plants under salt stress. Shoot diameter and leaf number per plant increased with SA treatments under salt stress. The greatest chlorophyll content was obtained with 1.00 mM SA treatment in both saline and non-saline conditions. Leaf water relative content (LWRC) reduced in response to salt stress while SA raised LWRC of salt stressed cucumber plants. Salinity treatments induced significant increases in electrolyte leakage. Plants treated with foliar SA had lower values of electrolyte leakage than non-treated ones. In regard to nutrient content, it can be interfered that foliar SA applications increased almost all nutrient content in leaves and roots of cucumber plants under salt stress. Generally, the greatest values were obtained from 1.00 mM SA application. Based on these findings, the SA treatments may help alleviate the negative effect of salinity on the growth of cucumber.