The effect of various concentrations salicylic acid on mitigation of sodium chloride stress in growth parameters and photosynthetic pigments of sunflower plant (Helianthus annuus L.)
Subject Areas : Genetic
1 - Department of Biology, Faculty of Science, Payame Noor University, Tehran, Iran.
Keywords: Anthocyanin, Chlorophyll, Salicylic Acid, Sodium chloride, Helianthus annuus L,
Abstract :
S Soil salinity is a serious environmental problem that has negative effect on plants’ growth and production. On the other hand, salicylic acid plays an essential role in preventing oxidative damage in plants by detoxifying superoxide radicals produced as a result of salinity. In this research the effect of different concentrations of salicylic acid, sodium chloride, and interaction effects of both factors were studied on morphological and biochemical parameters in sunflower (Helianthus annuus L.) plants. NaCl concentrations of 0, 25, 50, 75, and 100 mM and salicylic acid concentrations of 0, 0.5, 1, and 1.5 mM were used in the form of a factorial experiment in a complete randomized design (CRD). Results showed that salinity caused significant decrease in shoot and root length, fresh and dry weights, chlorophylls, and carotenoid of plants while in salicylic acid pre-treated plants, this reduction has been moderated. On the other hand, increasing the amount of anthocyanin, chlorophylls, and carotenoid indicated the role of salicylic acid in increasing the tolerance of this plant to sodium chloride stress. These findings suggest that salicylic acid application increases plant tolerance to sodium chloride stress.
Abd-El Samad, H.M. and Shaddad, M.A.K. (1997). Salt tolerance of soybean cultivars. Biologia Plantarum. 39 (2): 263-269.
Abdul Jaleel, C., Riadh, K., Gopi, R., Manivannan, P., Ines, J., Al-Juburi, H.J., Chang-Xing, Z., Hong-Bo, S. and Panneerselvam, R. (2009). Antioxidant defense responses: Physiological plasticity in higher plants under abiotic constrains. Acta Physiologiae Plantarum. 31: 427-436.
Afzal, I. (2005). Seed enhancements to induced salt tolerance in wheat (Triticum aestivumL.). Ph.D. Thesis, Agricultural University of Faisalabad, Pakistan, pp. 266 .
Agrawal, S., Sairam, P.K., Srivasta, G.C. and Meena, R.C. (2005). Changes in antioxidant enzymes activity and oxidative stress by abscisic acid and salicylic acid in wheat genotypes. Biologia Plantarum. 49 (4): 541-550.
Anderson, J.P., Thatcher, L.F. and Singh, K.B. (2005). Plant defence responses: conservation between models and crops. Functional Plant Biology. 32: 21-34.
Arfan, M., Athar, H.R. and Ashraf, M. (2007). Does exogenous application of salicylic acid through the rooting medium modulate growth and photosynthetic capacity in two differently adapted spring wheat cultivars under salt stress? Journal of Plant Physiology. 164: 685-694.
Ashraf, M., Akram, N.A., Arteca, R.N. and Foolad, M.R. (2010). The physiological, Biochemical and molecular roles of brassinosteroids and salicylic acid in plant processes and salt tolerance. Critical Reviews in Plant Sciences. 29: 162-190.
Bartosz, G. (1997). Oxidative stress in plants . Acta Physiologiae Plantarum. 19 (1): 47-64.
Clarke, A., Mur, L.A.J., Darby, R.M. and Kenton, P. (2005). Harpin modulates the accumulation of salicylic acid by Arabidopsis cells via apoplastic alkalization. Journal of Experimental Botany. 56: 3129-3136.
Deef, H.E. (2007). Influence of salicylic acid on stress tolerance during seed germination of Triticum aestivum and Hordeum vulgare. Advances of Biological Research. 1: 40–48.
Denby, K.J., Jason, L.J.M., Murray, S.L. and Last, R.L. (2005). Ups1, an Arabidopsis thaliana camalexin accumulation mutant defective in multiple defence signaling pathways. Plant Journal. 41: 673-684.
Downton, W.J.S., Grant, W.J.R. and Robinson, S.P. (1985). Photosynthetic and stomatal responses of spinach leaves to salt stress. Plant Physiology. 77: 85-88.
El-Tayeb, M.A. (2005). Response of barley grains to the interactive effect of salinity and salicylic acid. Plant Growth Regulation. 45: 215–224.
Gunes, A., Inal, A., Alpaslam, M., Erslan, F., Bagsi, E.G. and Cicek, N. (2007). Salicylic acid induced changes on some physiological parameters symptomatic for oxidative stress and mineral nutrition in maize (Zea mays L.) grown under salinity. Journal of Plant Phsiology. 164: 728-736.
Hamid, M., Rehman, K.H. and Ashraf, M. (2010). Salicylic acid–induced growth and biochemical changes in salt-stressed wheat. Communications in Soil Science and Plant Analysis. 41:373–389.
Heidari, A., Toorchi, M., Bandehagh, A and Shakiba, M.R. (2011). Effect of Nacl stress on growth, water relations, organic and inorganic osmolytes accumulation in Sunflower (Helianthus annuus L.) lines. Universal Journal of Environmental Research and Technology. 1: 351-362.
Chartzoulakis, K.S. (1994). Photosynthesis, water relations and leaf growth of cucumber exposed to salt stress. Scientia Horticulturae. 59: 27–35.
Jabeen, S., Shahbaz, M. and Akram, N.A. (2007). Influence of exogenous application of salicylic acid on growth and gas exchange characteristics of wheat (Triticum aestivum L.) under control or saline conditions. International Journal of Life Sciences. 1: 425– 431.
Juan, M., Rivero, R.M., Romero, L., Rviz, J.M. (2005). Evaluation of some nutritional and biochemical indicators in selecting salt-resistant tomato cultivars. Environmental and Experimental Botany. 54: 193-201.
Kaliamoorthy, S. and Rao, A.S. (1994). Effect of salinity on anthocyanin accumulation in the root of maize. Indian Journal of Plant Physiology. 37: 169-170.
Kao, W.Y., Tsai, T.T., Tsai, H.C. and Shih, C.N. (2006). Response of three glycine species to salt stress. Environmental and Experimental Botany. 56: 120-125.
Khan, M.I.R., Syeed, S., Nazar, R. and Anjum, N.A. (2012). An insight into the role of salicylic acid and jasmonic acid in salt stress tolerance. In: Phytohormones and abiotic stress tolerance in plants (Eds. Khan, N.A., Nazar, R., Iqbal, N. and Anjum, N.A.) pp. 277-300. Springer, New York.
Khodary, S.E.A. (2004). Effect of salicylic acid on the growth, photosynthesis and carbohydrate metabolism in salt-stressed maize plants. Journal of Agriculture and Biology. 6: 5–8.
Lichtenthaler, H.K. (1987). Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. Methods in Enzymology. 148: 350-382.
Manchanda, G. and Garg, N. (2008). Salinity and its effects on the functional biology of legumes. Acta Physiologiae Plantarum. 30: 595-618.
Masood, A., Shab, A., Zeeshan, M. and Abraham, G. (2006). Differential response of antioxidant enzymes to salinity stress in two varieties of Azolla (Azolla pinnata and Azolla filiculcides). Environmental and Experimental Botany. 58: 216-222.
Murray, Y. (1994). Ca2+ regulation of outward rectifying K+ channel in the plasma membrane of tobacco cultured cells in suspension: a role of the K+ channel in mitigation of salt-stress effects by external Ca+. Plant Cell Physiology. 39: 1039-1044
Nazar, R., Iqbal, N., Syeed, S. and Khan, N.A. (2011). Salicylic acid alleviates decreases in photosynthesis under salt stress by enhancing nitrogen and sulfur assimilation and antioxidant metabolism differentially in two mungbean cultivars. Journal of Plant Physiology. 168: 807-815.
Neocleous, D. and Nasilakakis, M. (2007). Effects of Nacl stress on red raspberry (Rubus idaeus L. “Autumn Bliss”). Scientia Horticulturae. 112: 282-289.
Parida, A.K. and Das, A.B. (2005). Salt tolerance and salinity effects on plants:a review. Ecotoxicology and Environmental Safety. 60: 324–349.
Popova, L., Pancheva, T. and Uzunova, A. (1997). Salicylic acid: properties, biosynthesis and physiological role. Bulgarian Journal of Plant Physiology. 23 (1-2): 85-93.
Popova, L.P., Maslenkova, L.T., Yordanova, R.Y., Ivanova, A.P., Krantev, A.P. and Szalai, G. (2009). Exogenous treatment with salicylic acid attenuates cadmium toxicity in Pea seedlings. Plant Physiology and Biochemistry. 47:224-231.
Sakhabutdinova, A.R., Fatkhutdinova, D.R., Bezrukova, M.V. and Shakirova, F.M. (2003). Salicylic acid prevents the damaging action of stress factors on wheat plants. Bulgarian Journal of Plant Physiology. 1: 314-319.
Sakhabutdinova, A.R., Fatkhutdinova, D.R., Bezrukova, M.V. and Shakirova, F.M. (2003). Salicylic acid prevents the damaging action of stress factors on wheat plants. Bulgarian Journal of Plant Physiology. 1: 314-319.
Shakirova, F.M., Sakhabutdinova, A.R., Bezrukova, M.V., Fatkhutdinova, R.A. and Fatkhutdinova, D.R. (2003). Changes in the hormonal status of wheat seedlings induced by salicylic acid and salinity. Plant Science. 164: 317–322.
Shannon, M.C. and Grieve, C.M. (1999). Tolerance of vegetables to salinity. Scientia Horticulturae. 78: 5–38.
Shibli, R.A., Kushad, M., Yousef, G.G. and Lila, M.A. (2007). Physiological and biochemical responses of tomato micro shoots to induced salinity stress with associated ethylene accumulation. Plant Growth Regulation. 51: 159-169.
Shim, Ie-S., Momose, Y., Yamamoto, A., Kim, D.W. and Usui, K. (2003). Inhibition of catalase activity by oxidative stress and its relationship salicylic acid accumulation in plants. Plant Growth Regulation. 39: 285–292.
Shirasu, K., Nakajima, H., Rajshekar, K., Dixon, R.A., and Lamb, C. (1997). Salicylic acid potentiates an agonist-dependent gain control that amplifies pathogen signal in the activation of defence mechanism. Plant Cell. 9: 261-270.
Stevens, J., Seneratna, T. and Sivasithamparam, K. (2006). Salicylic acid induces salinity tolerance in tomato (Lycopersicon esculentum cv. Roma): Associated changes in gas exchange, water relations and membrane stabilisation. Plant Growth Regulation. 49: 77-83.
Sudhakar, C., Lakshmi, A. and Giridarakumar, S. (2001). Chages in the antioxidant enzyme efficacy in two high yielding genotypes of mulberry (Morus alba L.) under Nacl salinity. Plant Science. 141: 613-619.
Syeed, S., Anjum, N.A., Nazar, R., Iqbal, N., Masood, A. and Khan, N.A. (2011). Salicylic acid mediated changes in photosynthesis, nutrients content and antioxidant metabolism in two mustard (Brassica juncea L.) cultivars differing in salt tolerance. Acta Physiologiae Plantarum. 33: 877-886.
Szalai, G., Tari, I., Janda, T., Pestenacz, A. and Paldi, E. (2000). Effects of cold acclimation and salicylic acid on changes in ACC and MACC contents in maize during chilling. Biologia Plantarum. 43: 637-640.
Szepesi, A. (2006). Salicylic acid improves the acclimation of Lycopersicon esculentum Mill. L. to high salinity by approximating its salt-stress response to that of the wild species L. pennellii. Acta Biologica Szegediensis. 50: 177.
Tari, I., Csiszar, J., Gabriella, S., Horvath, F., Pecsvaradi, A., Kiss, G., Szepsi, A., Szabo, M. and Erdei, L. (2002). Acclimation of tomato plants to salinity stress after a salicylic acid pre-treatment. Acta Biologica Szegediensis. 46: 55–56.
Tuna, A.L., Kaya, C., Dilkilitas, M., Yokas, I., Buruni, B. and Altunlu, H. (2007). Comparative effects of various salicylic acid derivatives on key growth parameters and some enzyme activities in salinity stressed maize (Zea mays L.) plants. Pakistan Journal of Botany. 39: 787–798.
Wahid, A. and Ghazanfar, A. (2006). Possible involvement of some secondary metabolites in salt tolerance of sugarcane. Journal of Plant Physiology 163: 723-730.
Wang, Y., Yang, Z.M., Zhang, Q.F. and Li, J.L. (2009). Enhanced chilling tolerance in Zoysia matrella by pre-treatment with salicylic acid, calcium chloride, hydrogen peroxide or 6-benzylaminopurine. Biologia Plantarum. 53: 179-182.
Wanger, G.J. (1979). Content and vacuole/extra vacuole distribution of neutral sugars, free amino acids, and anthocyanins in protoplasts. Plant Physiology. 64: 88-93.
Yalpani, N., Enyedi, A..J., Leon, J., Raskin, I. (1994). Ultraviolet light and ozone stimulate accumulation of salicylic acid and pathogenesis related proteins and virus resistance in tobacco. Planta. 193: 373-376.
Yildirim, E., Turan, M. and Guvenc, I. (2008). Effect of foliar salicylic acid applications on growth, chlorophyll, and mineral content of cucumber grown under salt stress. Journal of Plant Nutrition. 31: 593–612.
Yusuf, M., Hasan, S.A., Ali, B., Hayat, S., Fariduddin, Q. and Ahmad, A. (2008). Effect of salicylic acid on salinity induced changes in Brassica juncea. Journal of Integrative Plant Biology. 50: 1096–1102.
Zahra, S., Amin, B., Mohamad Ali, V.S., Ali, Y. and Mehdi, Y. (2010). The salicylic acid effect on the tomato (Lycopersicum esculentum Mill.) sugar, protein and proline contents under salinity stress (NaCl). Journal of Biophysical Struture Biology. 2: 35-41.
Zhu, Z., Wei, G., Li, J., Qian, Q. and Yu, J. (2004). Silicon alleviates salt stress and increases antioxidant enzymes activity in leaves of salt-stressed cucumber (Cucumis sativus L.). Plant Science. 167: 527–533.
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Abd-El Samad, H.M. and Shaddad, M.A.K. (1997). Salt tolerance of soybean cultivars. Biologia Plantarum. 39 (2): 263-269.
Abdul Jaleel, C., Riadh, K., Gopi, R., Manivannan, P., Ines, J., Al-Juburi, H.J., Chang-Xing, Z., Hong-Bo, S. and Panneerselvam, R. (2009). Antioxidant defense responses: Physiological plasticity in higher plants under abiotic constrains. Acta Physiologiae Plantarum. 31: 427-436.
Afzal, I. (2005). Seed enhancements to induced salt tolerance in wheat (Triticum aestivumL.). Ph.D. Thesis, Agricultural University of Faisalabad, Pakistan, pp. 266 .
Agrawal, S., Sairam, P.K., Srivasta, G.C. and Meena, R.C. (2005). Changes in antioxidant enzymes activity and oxidative stress by abscisic acid and salicylic acid in wheat genotypes. Biologia Plantarum. 49 (4): 541-550.
Anderson, J.P., Thatcher, L.F. and Singh, K.B. (2005). Plant defence responses: conservation between models and crops. Functional Plant Biology. 32: 21-34.
Arfan, M., Athar, H.R. and Ashraf, M. (2007). Does exogenous application of salicylic acid through the rooting medium modulate growth and photosynthetic capacity in two differently adapted spring wheat cultivars under salt stress? Journal of Plant Physiology. 164: 685-694.
Ashraf, M., Akram, N.A., Arteca, R.N. and Foolad, M.R. (2010). The physiological, Biochemical and molecular roles of brassinosteroids and salicylic acid in plant processes and salt tolerance. Critical Reviews in Plant Sciences. 29: 162-190.
Bartosz, G. (1997). Oxidative stress in plants . Acta Physiologiae Plantarum. 19 (1): 47-64.
Clarke, A., Mur, L.A.J., Darby, R.M. and Kenton, P. (2005). Harpin modulates the accumulation of salicylic acid by Arabidopsis cells via apoplastic alkalization. Journal of Experimental Botany. 56: 3129-3136.
Deef, H.E. (2007). Influence of salicylic acid on stress tolerance during seed germination of Triticum aestivum and Hordeum vulgare. Advances of Biological Research. 1: 40–48.
Denby, K.J., Jason, L.J.M., Murray, S.L. and Last, R.L. (2005). Ups1, an Arabidopsis thaliana camalexin accumulation mutant defective in multiple defence signaling pathways. Plant Journal. 41: 673-684.
Downton, W.J.S., Grant, W.J.R. and Robinson, S.P. (1985). Photosynthetic and stomatal responses of spinach leaves to salt stress. Plant Physiology. 77: 85-88.
El-Tayeb, M.A. (2005). Response of barley grains to the interactive effect of salinity and salicylic acid. Plant Growth Regulation. 45: 215–224.
Gunes, A., Inal, A., Alpaslam, M., Erslan, F., Bagsi, E.G. and Cicek, N. (2007). Salicylic acid induced changes on some physiological parameters symptomatic for oxidative stress and mineral nutrition in maize (Zea mays L.) grown under salinity. Journal of Plant Phsiology. 164: 728-736.
Hamid, M., Rehman, K.H. and Ashraf, M. (2010). Salicylic acid–induced growth and biochemical changes in salt-stressed wheat. Communications in Soil Science and Plant Analysis. 41:373–389.
Heidari, A., Toorchi, M., Bandehagh, A and Shakiba, M.R. (2011). Effect of Nacl stress on growth, water relations, organic and inorganic osmolytes accumulation in Sunflower (Helianthus annuus L.) lines. Universal Journal of Environmental Research and Technology. 1: 351-362.
Chartzoulakis, K.S. (1994). Photosynthesis, water relations and leaf growth of cucumber exposed to salt stress. Scientia Horticulturae. 59: 27–35.
Jabeen, S., Shahbaz, M. and Akram, N.A. (2007). Influence of exogenous application of salicylic acid on growth and gas exchange characteristics of wheat (Triticum aestivum L.) under control or saline conditions. International Journal of Life Sciences. 1: 425– 431.
Juan, M., Rivero, R.M., Romero, L., Rviz, J.M. (2005). Evaluation of some nutritional and biochemical indicators in selecting salt-resistant tomato cultivars. Environmental and Experimental Botany. 54: 193-201.
Kaliamoorthy, S. and Rao, A.S. (1994). Effect of salinity on anthocyanin accumulation in the root of maize. Indian Journal of Plant Physiology. 37: 169-170.
Kao, W.Y., Tsai, T.T., Tsai, H.C. and Shih, C.N. (2006). Response of three glycine species to salt stress. Environmental and Experimental Botany. 56: 120-125.
Khan, M.I.R., Syeed, S., Nazar, R. and Anjum, N.A. (2012). An insight into the role of salicylic acid and jasmonic acid in salt stress tolerance. In: Phytohormones and abiotic stress tolerance in plants (Eds. Khan, N.A., Nazar, R., Iqbal, N. and Anjum, N.A.) pp. 277-300. Springer, New York.
Khodary, S.E.A. (2004). Effect of salicylic acid on the growth, photosynthesis and carbohydrate metabolism in salt-stressed maize plants. Journal of Agriculture and Biology. 6: 5–8.
Lichtenthaler, H.K. (1987). Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. Methods in Enzymology. 148: 350-382.
Manchanda, G. and Garg, N. (2008). Salinity and its effects on the functional biology of legumes. Acta Physiologiae Plantarum. 30: 595-618.
Masood, A., Shab, A., Zeeshan, M. and Abraham, G. (2006). Differential response of antioxidant enzymes to salinity stress in two varieties of Azolla (Azolla pinnata and Azolla filiculcides). Environmental and Experimental Botany. 58: 216-222.
Murray, Y. (1994). Ca2+ regulation of outward rectifying K+ channel in the plasma membrane of tobacco cultured cells in suspension: a role of the K+ channel in mitigation of salt-stress effects by external Ca+. Plant Cell Physiology. 39: 1039-1044
Nazar, R., Iqbal, N., Syeed, S. and Khan, N.A. (2011). Salicylic acid alleviates decreases in photosynthesis under salt stress by enhancing nitrogen and sulfur assimilation and antioxidant metabolism differentially in two mungbean cultivars. Journal of Plant Physiology. 168: 807-815.
Neocleous, D. and Nasilakakis, M. (2007). Effects of Nacl stress on red raspberry (Rubus idaeus L. “Autumn Bliss”). Scientia Horticulturae. 112: 282-289.
Parida, A.K. and Das, A.B. (2005). Salt tolerance and salinity effects on plants:a review. Ecotoxicology and Environmental Safety. 60: 324–349.
Popova, L., Pancheva, T. and Uzunova, A. (1997). Salicylic acid: properties, biosynthesis and physiological role. Bulgarian Journal of Plant Physiology. 23 (1-2): 85-93.
Popova, L.P., Maslenkova, L.T., Yordanova, R.Y., Ivanova, A.P., Krantev, A.P. and Szalai, G. (2009). Exogenous treatment with salicylic acid attenuates cadmium toxicity in Pea seedlings. Plant Physiology and Biochemistry. 47:224-231.
Sakhabutdinova, A.R., Fatkhutdinova, D.R., Bezrukova, M.V. and Shakirova, F.M. (2003). Salicylic acid prevents the damaging action of stress factors on wheat plants. Bulgarian Journal of Plant Physiology. 1: 314-319.
Sakhabutdinova, A.R., Fatkhutdinova, D.R., Bezrukova, M.V. and Shakirova, F.M. (2003). Salicylic acid prevents the damaging action of stress factors on wheat plants. Bulgarian Journal of Plant Physiology. 1: 314-319.
Shakirova, F.M., Sakhabutdinova, A.R., Bezrukova, M.V., Fatkhutdinova, R.A. and Fatkhutdinova, D.R. (2003). Changes in the hormonal status of wheat seedlings induced by salicylic acid and salinity. Plant Science. 164: 317–322.
Shannon, M.C. and Grieve, C.M. (1999). Tolerance of vegetables to salinity. Scientia Horticulturae. 78: 5–38.
Shibli, R.A., Kushad, M., Yousef, G.G. and Lila, M.A. (2007). Physiological and biochemical responses of tomato micro shoots to induced salinity stress with associated ethylene accumulation. Plant Growth Regulation. 51: 159-169.
Shim, Ie-S., Momose, Y., Yamamoto, A., Kim, D.W. and Usui, K. (2003). Inhibition of catalase activity by oxidative stress and its relationship salicylic acid accumulation in plants. Plant Growth Regulation. 39: 285–292.
Shirasu, K., Nakajima, H., Rajshekar, K., Dixon, R.A., and Lamb, C. (1997). Salicylic acid potentiates an agonist-dependent gain control that amplifies pathogen signal in the activation of defence mechanism. Plant Cell. 9: 261-270.
Stevens, J., Seneratna, T. and Sivasithamparam, K. (2006). Salicylic acid induces salinity tolerance in tomato (Lycopersicon esculentum cv. Roma): Associated changes in gas exchange, water relations and membrane stabilisation. Plant Growth Regulation. 49: 77-83.
Sudhakar, C., Lakshmi, A. and Giridarakumar, S. (2001). Chages in the antioxidant enzyme efficacy in two high yielding genotypes of mulberry (Morus alba L.) under Nacl salinity. Plant Science. 141: 613-619.
Syeed, S., Anjum, N.A., Nazar, R., Iqbal, N., Masood, A. and Khan, N.A. (2011). Salicylic acid mediated changes in photosynthesis, nutrients content and antioxidant metabolism in two mustard (Brassica juncea L.) cultivars differing in salt tolerance. Acta Physiologiae Plantarum. 33: 877-886.
Szalai, G., Tari, I., Janda, T., Pestenacz, A. and Paldi, E. (2000). Effects of cold acclimation and salicylic acid on changes in ACC and MACC contents in maize during chilling. Biologia Plantarum. 43: 637-640.
Szepesi, A. (2006). Salicylic acid improves the acclimation of Lycopersicon esculentum Mill. L. to high salinity by approximating its salt-stress response to that of the wild species L. pennellii. Acta Biologica Szegediensis. 50: 177.
Tari, I., Csiszar, J., Gabriella, S., Horvath, F., Pecsvaradi, A., Kiss, G., Szepsi, A., Szabo, M. and Erdei, L. (2002). Acclimation of tomato plants to salinity stress after a salicylic acid pre-treatment. Acta Biologica Szegediensis. 46: 55–56.
Tuna, A.L., Kaya, C., Dilkilitas, M., Yokas, I., Buruni, B. and Altunlu, H. (2007). Comparative effects of various salicylic acid derivatives on key growth parameters and some enzyme activities in salinity stressed maize (Zea mays L.) plants. Pakistan Journal of Botany. 39: 787–798.
Wahid, A. and Ghazanfar, A. (2006). Possible involvement of some secondary metabolites in salt tolerance of sugarcane. Journal of Plant Physiology 163: 723-730.
Wang, Y., Yang, Z.M., Zhang, Q.F. and Li, J.L. (2009). Enhanced chilling tolerance in Zoysia matrella by pre-treatment with salicylic acid, calcium chloride, hydrogen peroxide or 6-benzylaminopurine. Biologia Plantarum. 53: 179-182.
Wanger, G.J. (1979). Content and vacuole/extra vacuole distribution of neutral sugars, free amino acids, and anthocyanins in protoplasts. Plant Physiology. 64: 88-93.
Yalpani, N., Enyedi, A..J., Leon, J., Raskin, I. (1994). Ultraviolet light and ozone stimulate accumulation of salicylic acid and pathogenesis related proteins and virus resistance in tobacco. Planta. 193: 373-376.
Yildirim, E., Turan, M. and Guvenc, I. (2008). Effect of foliar salicylic acid applications on growth, chlorophyll, and mineral content of cucumber grown under salt stress. Journal of Plant Nutrition. 31: 593–612.
Yusuf, M., Hasan, S.A., Ali, B., Hayat, S., Fariduddin, Q. and Ahmad, A. (2008). Effect of salicylic acid on salinity induced changes in Brassica juncea. Journal of Integrative Plant Biology. 50: 1096–1102.
Zahra, S., Amin, B., Mohamad Ali, V.S., Ali, Y. and Mehdi, Y. (2010). The salicylic acid effect on the tomato (Lycopersicum esculentum Mill.) sugar, protein and proline contents under salinity stress (NaCl). Journal of Biophysical Struture Biology. 2: 35-41.
Zhu, Z., Wei, G., Li, J., Qian, Q. and Yu, J. (2004). Silicon alleviates salt stress and increases antioxidant enzymes activity in leaves of salt-stressed cucumber (Cucumis sativus L.). Plant Science. 167: 527–533.
