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Application of Salicylic Acid Increases Contents of Nutrients And International Journal of Plant Biology 2010; volume 1:e1 Application of salicylic acid water and osmotic potential. 4 The increasing increases contents of nutrients concentration of Na + and Cl – in the rooting Correspondence: Nafees A. Khan, Department of medium suppresses the uptake of essential Botany, Aligarh Muslim University, Aligarh and antioxidative metabolism nutrients N, P, K, and Ca, 5 and alters ionic rela - 202002, India. E-mail: [email protected] in mungbean and alleviates tionships. 6 Salt stress also generates reactive adverse effects of salinity oxygen species (ROS) causing oxidative stress Key words: antioxidant enzymes, mungbean, photo - – stress on plants. These ROS such as superoxide (O 2 ), synthesis, salicylic acid, salinity. hydrogen peroxide (H 2O2), hydroxyl radical – 1 Contributions: each author contributed equally to Nafees A. Khan, Shabina Syeed, Asim (OH ), and singlet oxygen ( O2) can damage 7 the manuscript. Masood, Rahat Nazar, Noushina Iqbal chloroplasts and reduce photosynthesis, involv - ing stomatal behavior and inhibition of photo - Department of Botany, Aligarh Muslim Received for publication: 26 August 2009. chemical processes, 8 and disturbance in homeo- University, Aligarh, India Revision received: 3 December 2009. stasis of Na + and Cl – ions and essential miner - Accepted for publication: 3 December 2009. al nutrients. 9 The decrease in photosynthesis may lead to reduction in growth and yield. This work is licensed under a Creative Commons Attribution 3.0 License (by-nc 3.0). Abstract Plants operate several mechanisms to counter - act the adverse effects of salt. These mechan- ©Copyright N.A. Khan et al.,2010 isms may be enhanced by the application of Salicylic acid (SA), a naturally occurring Licensee PAGEPress, Italy 4 International Journal of Plant Biology 2010; 1:e1 plant hormone, is an important signal molecule chemicals to the plants. One such mechanism doi:10.4081/pb.2010.e1 known to have diverse effects on biotic and abi - is the activation of an antioxidant enzyme sys - otic stress tolerance. Its growth-promoting tem, which may be influenced by the interaction effect on various plants has been shown, but the of plant growth regulators and salt. Plants con - taining high activities of antioxidant enzymes information on the response of mungbean, an Materials and Methods important leguminous plant, to SA application have shown considerable resistance to the 7,10-11 under salt stress is limited. Mungbean ( Vigna oxidative damage caused by ROS. The radiata L.) cultivar Pusa Vishal plants grown antioxidant enzyme system constitutes super - Plant material and growth condi - with 50 mM NaCl were sprayed with 0.1, 0.5, or oxide dismutase (SOD, EC 1.15.1.1) as the pri - tions 1.0 mM SA and basic physiological processes mary step of cellular defense. It dismutates Plants of mungbean [ Vigna radiata L. – were studied to substantiate our understanding superoxide ions (O 2 ) to H 2O2 and O 2. Further, (Wilczek)] cultivar Pusa Vishal were raised of their role in tolerance to salinity-induced the accumulation of H 2O2 is restricted by the from seeds planted in 15 cm-diameter earthen oxidative stress and how much such processes action of the ascorbate-glutathione cycle, where pots filled with acid-washed sand purified are induced by SA application. Treatment of ascorbate peroxidase (APX, EC 1.11.1.11) according to Hewitt. 31 Two plants were grown plants with 0.5 mM SA resulted in a maximum reduces it to H 2O. The final step is catalyzed by in a pot kept in a naturally illuminated green - + – decrease in the content of Na , Cl , H 2O2, and glutathione reductase (GR, EC 1.6.4.2), which house (photosynthetically active radiation thiobarbituric acid reactive substances catalyzes the NADPH-dependent reaction of oxi - (PAR) >900±22 µmol/m 2/s and average tem - (TBARS), and electrolyte leakage under saline dized glutathione (GSSG) to reduced glu - perature of 33±2ºC) of the Department of conditions compared to the control. In contrast, tathione (GSH). 12 Botany, Aligarh Muslim University, Aligarh, this treatment increased N, P, K, and Ca con - Salicylic acid (SA), a naturally occurring India. Plants in each pot were fed with 300 ml tent, activity of antioxidant enzymes, glu - plant hormone, acts as an important signaling Hoagland nutrient solution containing 50 mM tathione content, photosynthesis, and yield molecule in plants, and has diverse effects on NaCl every alternate day and 200 mL of maximally under nonsaline and saline condi - tolerance to abiotic stress. 13-15 Exogenous appli - deionised water daily. The nutrient solution tions. The application of higher concentration cation of SA may participate in the regulation of was replaced weekly. Salicylic acid was dis - of SA (1.0 mM) either proved inhibitory or was physiological processes in plants, such as stom - solved in absolute ethanol then added drop- of no additional benefit. It was concluded that atal closure, ion uptake and transport, 16 mem - wise to water (ethanol/water: 1/1000, v/v). 32 0.5 mM SA alleviates salinity-inhibited photo - brane permeability, 17 and photosynthesis and Fifteen days after sowing (DAS), SA was synthesis and yield through a decrease in Na +, growth. 18 Its role in abiotic stress tolerance applied on the foliage of 0 or 50 mM NaCl – Cl , H 2O2, and TBARS content, and electrolyte such as ozone, UV-B, heat, heavy metal, and grown plants at a concentration of 0.1, 0.5, or leakage, and an increase in N, P, K, and Ca con - osmotic stress 19-22 has been reported. Studies 1.0 mM with a hand sprayer. A control group of tent, activity of antioxidant enzymes, and glu - on barley ( Hordeum vulgare ), maize ( Zea plants was grown without NaCl and sprayed tathione content. mays ), wheat ( Triticum aestivum ), bean with deionized water. A surfactant teepol (Phaseolus vulgaris ), lentil ( Lens culinaris ), (0.5%) was added with the control and SA and sunflower ( Helianthus annuus ) suggest treatment solutions. The volume of the spray that SA may be used to alleviate salt stress. 21,23- was 30 mL per pot. The treatments were Introduction 28 In contrast, studies on Zea mays 29 and arranged in a completely randomized block design and replicated three times. Arabidopsis 30 have shown an inhibitory effect of Salinity is one of the major stresses in arid SA. In view of the contrasting reports on the and semi-arid regions causing adverse effects effect of SA and limited literature available on Measurements at physiological, biochemical, 1 and molecular leguminous crops, the research was undertaken At 15 days after salicylic acid application levels, 2 limiting crop productivity. 1,3 Salt stress on mungbean ( Vigna radiata ), an important (i.e. 30 DAS), the content of Na + and Cl –, can disturb growth and photosynthetic process - crop grown in Asia and other parts of the world, nutrients, H 2O2, and thiobarbituric acid reac - es by causing changes in the accumulation of to study the influence of SA application in the tive substances (TBARS), electrolyte leakage, Na +, Cl –, and nutrients, and disturbance in alleviation of salinity stress. activity of antioxidant enzymes, and photosyn - [International Journal of Plant Biology 2010; 1:e1] [page 1] Article thetic traits were determined from each treat - Glutathione content USA). The measurements were done between ment and replicate. At harvest (60 DAS), yield The leaf tissue (200 mg) was homogenized 11h00-12h00 at light-saturating intensity and traits were recorded. in an ice bath in 1.0 mL of 5% 5-sulphosalicylic at atmospheric CO 2 concentration 365±5 acid and the homogenate was centrifuged for µmol/mol. Water use efficiency was calculated Content of Na + and Cl – and nutrients 10,000 ¥ g for 10 min. The supernatant was as the ratio of photosynthesis to transpiration. The determination of Na +, Cl –, N, P, K, and used for glutathione (GSH) estimation using Ca was done in acid-peroxide digested oven- 5,5’-dithio-2,2’-dinitrobenzoic acid (DTNB) by Yield traits reading the absorbance at 412 nm, following dried leaf samples. Na +, K, and Ca were meas - At harvest (60 DAS), pods were collected and 41 ured using a flame photometer (Khera-391: the method of Anderson. counted. Pod length was measured on a meter Khera Instruments, New Delhi) whereas Cl- scale. The number of seeds from each pod was counted. Seeds from a plant in each treatment was determined by potentiometric titration Photosynthetic traits were cleared, sun-dried, and weighed to com - with AgNO . N and P were determined by The activity of CA in leaf was estimated in 3 pute seed yield per plant. using the methods of Lindner, 33 and Fiske and the leaves used for photosynthesis measure - ment according to the method described by Subba Row, 34 respectively. Makino et al., 42 determined following time- Data analysis Data were statistically analyzed using Content of H O and thiobarbituric dependent reduction in pH 8.25-7.45 at 0-3°C. 2 2 Net photosynthetic rate, stomatal conduct- SPSS, 10.0 for Windows. Standard error was acid reactive substances, and elec - ance, intercellular CO 2 concentration, and calculated and analysis of variance (ANOVA) trolyte leakage transpiration rate were measured in fully was performed on the data to determine the least significant difference (LSD) between The content of H 2O2 was determined follow - expanded, uppermost leaves of plants using treatment means with the level of significance ing the method of Okuda et al. 35 Fresh leaf tis - an infrared gas analyzer (LiCOR-6200, at P<0.05. sues (50 mg) were ground in ice-cold 200 mM Portable Photosynthesis System, Nebraska, perchloric acid. The reaction was started by the addition of peroxidase and increases in the absorbance were recorded at 590 nm for 3 min. The content of TBARS was measured according to the method described by Dhindsa Figure 1.
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