Investigation of improving the drought tolerance in persian petunia (Petunia sp.) by exogenous application of salicylic acid and gibberellic acid
Morteza Goldani
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, IranAli Dolatkhahi
Department of Horticultural Science and Landscape, Ferdowsi University of Mashhad, Mashhad, IranMahdi Parsa
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, IranNavid Vahdati
Department of Horticultural Science and Landscape, Ferdowsi University of Mashhad, Mashhad, IranZahra Rasouli
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, IranAbstract
Plants respond to water deficit through several mechanisms. Among which, plant hormones play an important role in induction to abiotic stress tolerance. To bring this issue closer, an experiment was conducted to examine whether salicylic acid (SA) and gibberellic acid (GA3) exogenous applications through foliar spray could alleviate detrimental effects of water deficit on Persian petunia. Treatments consisted of water stress in three levels: 100% field capacity (F.C.) as control, 50, and 25% F.C linked with different concentrations of GA3 and SA sprayed at four levels (0, 100, 200 and 300 mg L–1). Results showed that the dry and fresh biomass, leaf area, photosynthetic pigments levels, relative water content (RWC) and the total number of flowers were significantly (P≤0.01) reduced in response to water deficit. However, free proline concentration and root-to-shoot ratio significantly increased in stressed plants. Plants subjected to severe water stress (25% F.C.), exhibit about 66% stomatal conductance rate lower than the plants grown at 100% F.C. The results showed that SA-treated plants exhibited reduced negative effect of water stress on most of the qualitative and quantitative attributes compared to control, while GA3 application had little impact on studied traits. Importantly, stomatal conductance and RWC were improved in water-stressed plants treated with SA. Moreover, SA-treated plants produced more flowers at all water deficit levels compared to GA3 sprayed plants. It can be concluded that Persian petunia plants respond to water deficit through adaptive changes at biochemical and morphological levels and that adverse effects of water deficit could be alleviated by exogenous application of SA.
Keywords:
ornamental flower, photosynthetic pigments, proline, relative water content, stomatal conductance, water deficitReferences
Ahmad, S., Ahmad, R., Ashraf, M.Y., Ashraf, M. and Waraich, E.A. 2009. Sunflower (Helianthus annuus L.) response to drought stress at germination and seedling growth stages. Pak. J. Bot. 41(2): 647-654.
Ahmed, C.B., Rouina, B.B., Sensoy, S., Boukhris, M. and Abdallah, F.B. 2009. Changes in gas exchange, proline accumulation and antioxidative enzyme activities in three olive cultivars under contrasting water availability regimes. Environ. Exp. Bot. 67: 345–352.
Alhadi, F.A., Yasseen, B.T. and Jabr, M. 1999. Water stress and gibberellic acid effects on growth of fenugreek plants. Irrigation Science. 18(4):185-190.
Ali, H.M., Siddiqui, M.H., Basalah, M.O., Al-Whaibi, M.H., Sakran, A.M. and Al-Amri, A. 2012. Effects of gibberellic acid on growth and photosynthetic pigments of Hibiscus sabdariffa L. under salt stress. African Journal of Biotechnology. 11(4): 800-804.
Anjum, S.A., Xie, X.Y., Wang, L.C., Saleem, M.F., Man, C. and Lei, W. 2011. Morphological, physiological and biochemical responses of plants to drought stress. Afr. J. Agric. Res. 6: 2026–2032.
Arfan, M., Athar, H.R. and Ashraf, M. 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(6): 685-694.
Ashraf, M.F.M.R. and Foolad, M. 2007. Roles of glycinebetaine and proline in improving plant abiotic stress resistance. Environ. Exp. Bot. 59: 206–216.
Athar, H. and Ashraf, M. 2005. Photosynthesis under drought stress. Handbook of photosynthesis, 793-804.
Bahadoran, M. and Salehi, H. 2015. Growth and flowering of two tuberose (Polianthes tuberosa L.) cultivars under deficit irrigation by saline water. J. Agr. Sci. Tech., 17(2): 415-426.
Bano, A. and Yasmeen, S. 2010. Role of phytohormones under induced drought stress in wheat. Pak J Bot. 42(4): 2579-2587.
Bates, L.S., Waldern, R.P. and Teare, I.D. 1973. Rapid determination of free proline for water stress studies. Plant and Soil. 39: 205-207.
Bezrukova, M. 2001. The role of hormonal changes in protective action of salicylic acid on growth of wheat seedlings under water deficits. Agrochemiya (Russ.) 2: 51-54.
Blum, A. 2005. Drought resistance, water-use efficiency, and yield potential are they compatible, dissonant, or mutually exclusive? Aust. J. Agric. Res. 56: 1159–1168.
Bolla, A., Voyiatzis, D., Koukourikou-Petridou, M. and Chimonidou, D. 2010. Photosynthetic parameters and cut-flower yield of rose ‘Eurored’(HT) are adversely affected by mild water stress irrespective of substrate composition. Scientia Horticulturae. 126: 390-394.
Cai, X., Starman, T., Niu, G., Hall, C. and Lombardini, L. 2012. Response of selected garden roses to drought stress. HortScience, 47(8):1050-1055.
Chaves, M.M., Flexas, J. and Pinheiro, C. 2009. Photosynthesis under drought and salt stress: regulation mechanisms from whole plant to cell. Annals of Botany. 103: 551-560.
Chaves, M.M., Pereira, J.S., Maroco, J., Rodrigues, M.L., Ricardo, C.P.P., Osório, M.L., Carvalho, I., Faria, T. and Pinheiro, C. 2002. How plants cope with water stress in the field? Photosynthesis and growth. Annals of botany. 89: 907-916.
Colebrook, E.H., Thomas, S.G., Phillips, A.L. and Hedden, P. 2014. The role of gibberellin signalling in plant responses to abiotic stress. Journal of experimental biology. 217 (1): 67-75.
Delauney, A.J. and Verma, D.P.S. 1993. Proline biosynthesis and osmoregulation in plants. Plant J. 4: 215-223.
El-Tayeb, M.A. 2005. Response of barley grains to the interactive effect of salinity and salicylic acid. Plant Growth Regul. 45:215 224.
Farooq, M., Wahid, A., Kobayashi, N., Fujita, D. and Basra, S.M.A. 2009. Plant drought stress: effects, mechanisms and management. Agronomy for Sustainable Development. 29:185-212.
Flexas, J. and Medrano, H. 2002. Energy dissipation in C3 plants under drought. Funct. Plant Biol. 29: 1209–1215.
Giorio, P., Sorrentino, G. and d’Andria, R. 1999. Stomatal behaviour leaf water status and photosynthetic response in field-grown olive trees under water deficit. Environ. Exp. Bot. 42: 95–104.
Guerfel, M., Baccouri, O., Boujnah, D., Chaïbi, W. and Zarrouk, M. 2009. Impacts of water stress on gas exchange, water relations, chlorophyll content and leaf structure in the two main Tunisian olive (Olea europaea L.) cultivars. Scientia Horticulturae. 119(3): 257-263.
Hayat, Q., Hayat, S., Irfan, M. and Ahmad, A. 2010. Effect of exogenous salicylic acid under changing environment: a review. Environmental and Experimental Botany. 68(1):14-25.
Hayat, S., Ahmad, A., and Alyemeni, M.N. 2013. Salicylic Acid: Plant Growth and Development. Springer Science & Business Media.
Hazrati, S., Tahmasebi-Sarvestani, Z., Modarres-Sanavy, S.A.M., Mokhtassi-Bidgoli, A. and Nicola, S. 2016. Effects of water stress and light intensity on chlorophyll fluorescence parameters and pigments of Aloe vera L. Plant Physiology and Biochemistry. 106: 141-148.
Huerta, L., Forment, J., Gadea, J., Fagoaga, C., Pena, L., Pérez‐Amador, M.A. and García‐Martínez, J.L. 2008. Gene expression analysis in citrus reveals the role of gibberellins on photosynthesis and stress. Plant Cell Environ. 31:1620–1633.
Hussain, M., Malik, M.A., Farooq, M., Khan, M.B., Akram, M. and Saleem, M.F. 2009. Exogenous glycine betaine and salicylic acid application improves water relations, allometry and quality of hybrid sunflower under water deficit conditions. Journal of Agronomy and Crop Science. 195: 98–109.
Idrees, M., Khan, M.M.A., Aftab, T., Naeem, M. and Hashmi, N. 2010. Salicylic acid-induced physiological and biochemical changes in lemongrass varieties under water stress. Journal of Plant Interactions. 5(4): 293-303.
Kaur, S., Gupta, A.K. and Kaur, N. 2000. Effect of GA3, kinetin and indole acetic acid on carbohydrate metabolism in chickpea seedlings germinating under water stress. Plant Growth Regulation. 30 (1): 61-70.
Kawano T. and Muto, S. 2000 – Mechanism of peroxidase actions for salicylic acid-induced generation of active oxygen species and an increase in cytosolic calcium in tobacco cell suspension culture. J. Exp. Bot. 51: 685-693.
Kaya, C., Kirnak, H., Higgs, D. and Saltali, K. 2002. Supplementary calcium enhances plant growth and fruit yield in strawberry cultivars grown at high salinity. Sci. Hortic. 93: 65-74.
Khan, W., Prithiviraj, B. and Smith, D.L. 2003. Photosynthetic responses of corn and soybean to foliar application of salicylates. Journal of plant physiology. 160 (5):485-492.
Lichtenthaler, H.K. and Wellburn, A.R. 1983. Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents.
Liu, C., Liu, Y., Guo, K., Fan, D., Li, G., Zheng, Y., Yu, L. and Yang, R. 2011. Effect of drought on pigments, osmotic adjustment and antioxidant enzymes in six woody plant species in karst habitats of southwestern China. Environmental and Experimental Botany. 71(2):174-183.
Liu, F. and Stützel, H. 2004. Biomass partitioning, specific leaf area, and water use efficiency of vegetable amaranth (Amaranthus spp.) in response to drought stress. Scientia Horticulturae. 102:15-27.
Martin-Mex, R., Villanueva-Couoh, E., Herrera-Campos, T. and Larque-Saavedra, A. 2005. Positive effect of salicylates on the flowering of African violet. Science Horticulture, 103: 499–502.
Martín-Mex, R., Villanueva-Couoh, E. and Uicab-Quijano, V. 2003. Positive effect of salicylic acid on the flowering of gloxinia. In: Proceedings 31st Annual Meeting. Plant Growth Regulatory Society America, Canada: Vancouver (pp 149–151). August 3–6 2003.
Medrano, H., Escalona, J.M., Bota, J., Gulías, J. and Flexas, J. 2002. Regulation of photosynthesis of C3 plants in response to progressive drought: stomatal conductance as a reference parameter. Annals of Botany. 89:895-905.
Patel, M.S. and Golakiya, B.A. 1988. Effect of Water-Stress on Yield Attributes and Yield of Groundnut (Arachis-Hypogaea). Indian Journal of Agricultural Sciences. 58(9):701-703.
Pirasteh-Anosheh, H., Emam, Y. and Pessarakli, M. 2013. Changes in endogenous hormonal status in corn (Zea mays) hybrids under drought stress. Journal of plant nutrition. 36(11):1695-1707.
Raskin, I. 1992. Role of salicylic acid in plants. Annual Review Plant Physiology. 43: 439–463.
Saruhan, N., Saglam, A. and Kadioglu, A. 2012. Salicylic acid pretreatment induces drought
tolerance and delays leaf rolling by inducing antioxidant systems in maize genotypes. Acta
Physiologiae Plantarum. 34: 97–106.
Shalhevet, J. 1993. Plants under salt and water stress. Plant Adaptation to Environmental Stress. 133-154.
Shamshiri, M.H., Mozafari, V., Sedaghati, E. and Bagheri, V. 2011. Response of petunia plants (Petunia hybrida cv. Mix) inoculated with Glomus mosseae and Glomus intraradices to phosphorous and drought stress. Journal of agricultural science and technology. 13: 929-942.
Shao, H.B., Chu, L.Y., Jaleel, C.A. and Zhao, C.X. 2008. Water-deficit stress-induced anatomical changes in higher plants. Comptes Rendus Biologies. 331:215-225.
Singh, B. and Usha, K. 2003. Salicylic acid induced physiological and biochemical changes in wheat seedlings under water stress. Plant Growth Regulation. 39 (2):137-141.
Kim, S.T., Kang, S.Y., Wang, Y., Kim, S.G., Hwang, D.H. and Kang, K.Y. 2008. Analysis of embryonic proteome modulation by GA and ABA from germinating rice seeds. Proteomics 8:3577–3587.
Tahir, M.H.N. and Mehid, S.S. 2001. Evaluation of open pollinated sunflower (Helianthus annuus L.) populations under water stress and normal conditions. Int. J. Agric. Biol. 3:236-238.
Taiz, L. and Zeiger, E. 2006. Plant Physiology Fourth Edition Sinauer Associates. Inc. Publishers. Sunderland, Massachusetts.
Tarchevskii, L.A. 2002 - Signal'nye sistemy kletok rastenii (Signal Systems of Plant Cells). Nauka, Moscow, 294 p.
Umebese, C.E., Olatimilehin, T.O. and Ogunsusi, T.A. 2009. Salicylic acid protects nitrate reductase activity, growth and proline in amaranth and tomato plants during water deficit. American Journal of Agricultural and Biological Sciences. 4 (3): 224-229.
Van Iersel, M.W., Dove, S., Kang, J.G. and Burnett, S.E. 2010. Growth and water use of petunia as affected by substrate water content and daily light integral. HortScience, 45(2):277-282.
Verbruggen, N. and Hermans, C. 2008. Proline accumulation in plants: a review. Amino acids. 35(4), pp.753-759.
Verma, V., Ravindran, P. and Kumar, P.P. 2016. Plant hormone-mediated regulation of stress responses. BMC plant biology, 16(1), p.86.
Wang, C., Yang, A., Yin, H. and Zhang, J. 2008. Influence of water stress on endogenous hormone contents and cell damage of maize seedlings. Journal of Integrative Plant Biology.50 (4): 427-434.
Waseem, M., Athar, H.U.R. and Asahrafi, M. 2006. Effect of salicylic acid applied through
rooting medium on drought tolerance of wheat. Pakistan Journal of Botany. 38:1127–1136.
Waterland, N.L., Campbell, C.A., Finer, J.J. and Jones, M.L. 2010. Abscisic acid application enhances drought stress tolerance in bedding plants. HortScience. 45(3):409-413.
Yamada, M., Morishita, H., Urano, K., Shiozaki, N., Yamaguchi-Shinozaki, K., Shinozaki, K. and Yoshiba, Y. 2005. Effects of free proline accumulation in petunias under drought stress. Journal of Experimental Botany. 56 (417): 1975-1981.
Yamasaki, S. and Dillenburg, L.R. 1999. Measurements of leaf relative water content in Araucaria angustifolia. Revista Brasilleira de Fisiologia Vegetal. 11: 69-75.
Yasseen, B.T. 1983. An analysis of the effects of salinity on leaf growth in Mexican wheats. PhD thesis, University of Leeds.
Yasseen, B.T. 1992. Physiology of water stress in plants. University of Mosul. University Press, Iraq.
Zarghami Moghaddam, M., Shoor, M., Ganjeali, A., Moshtaghi, N. and Tehranifar, A. 2014. Effect of salicylic acid on morphological and Ornamental characteristics of petunia hybrida at drought stress. Indian Journal of Fundamental and Applied Life Sciences, 4.
Zhang, S. and Liu, Y. 2001. Activation of salicylic acid-induced protein kinase, a mitogen-activated protein kinase, induces multiple defense responses in tobacco. Plant Cell. 13:1877 1889.
Ahmed, C.B., Rouina, B.B., Sensoy, S., Boukhris, M. and Abdallah, F.B. 2009. Changes in gas exchange, proline accumulation and antioxidative enzyme activities in three olive cultivars under contrasting water availability regimes. Environ. Exp. Bot. 67: 345–352.
Alhadi, F.A., Yasseen, B.T. and Jabr, M. 1999. Water stress and gibberellic acid effects on growth of fenugreek plants. Irrigation Science. 18(4):185-190.
Ali, H.M., Siddiqui, M.H., Basalah, M.O., Al-Whaibi, M.H., Sakran, A.M. and Al-Amri, A. 2012. Effects of gibberellic acid on growth and photosynthetic pigments of Hibiscus sabdariffa L. under salt stress. African Journal of Biotechnology. 11(4): 800-804.
Anjum, S.A., Xie, X.Y., Wang, L.C., Saleem, M.F., Man, C. and Lei, W. 2011. Morphological, physiological and biochemical responses of plants to drought stress. Afr. J. Agric. Res. 6: 2026–2032.
Arfan, M., Athar, H.R. and Ashraf, M. 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(6): 685-694.
Ashraf, M.F.M.R. and Foolad, M. 2007. Roles of glycinebetaine and proline in improving plant abiotic stress resistance. Environ. Exp. Bot. 59: 206–216.
Athar, H. and Ashraf, M. 2005. Photosynthesis under drought stress. Handbook of photosynthesis, 793-804.
Bahadoran, M. and Salehi, H. 2015. Growth and flowering of two tuberose (Polianthes tuberosa L.) cultivars under deficit irrigation by saline water. J. Agr. Sci. Tech., 17(2): 415-426.
Bano, A. and Yasmeen, S. 2010. Role of phytohormones under induced drought stress in wheat. Pak J Bot. 42(4): 2579-2587.
Bates, L.S., Waldern, R.P. and Teare, I.D. 1973. Rapid determination of free proline for water stress studies. Plant and Soil. 39: 205-207.
Bezrukova, M. 2001. The role of hormonal changes in protective action of salicylic acid on growth of wheat seedlings under water deficits. Agrochemiya (Russ.) 2: 51-54.
Blum, A. 2005. Drought resistance, water-use efficiency, and yield potential are they compatible, dissonant, or mutually exclusive? Aust. J. Agric. Res. 56: 1159–1168.
Bolla, A., Voyiatzis, D., Koukourikou-Petridou, M. and Chimonidou, D. 2010. Photosynthetic parameters and cut-flower yield of rose ‘Eurored’(HT) are adversely affected by mild water stress irrespective of substrate composition. Scientia Horticulturae. 126: 390-394.
Cai, X., Starman, T., Niu, G., Hall, C. and Lombardini, L. 2012. Response of selected garden roses to drought stress. HortScience, 47(8):1050-1055.
Chaves, M.M., Flexas, J. and Pinheiro, C. 2009. Photosynthesis under drought and salt stress: regulation mechanisms from whole plant to cell. Annals of Botany. 103: 551-560.
Chaves, M.M., Pereira, J.S., Maroco, J., Rodrigues, M.L., Ricardo, C.P.P., Osório, M.L., Carvalho, I., Faria, T. and Pinheiro, C. 2002. How plants cope with water stress in the field? Photosynthesis and growth. Annals of botany. 89: 907-916.
Colebrook, E.H., Thomas, S.G., Phillips, A.L. and Hedden, P. 2014. The role of gibberellin signalling in plant responses to abiotic stress. Journal of experimental biology. 217 (1): 67-75.
Delauney, A.J. and Verma, D.P.S. 1993. Proline biosynthesis and osmoregulation in plants. Plant J. 4: 215-223.
El-Tayeb, M.A. 2005. Response of barley grains to the interactive effect of salinity and salicylic acid. Plant Growth Regul. 45:215 224.
Farooq, M., Wahid, A., Kobayashi, N., Fujita, D. and Basra, S.M.A. 2009. Plant drought stress: effects, mechanisms and management. Agronomy for Sustainable Development. 29:185-212.
Flexas, J. and Medrano, H. 2002. Energy dissipation in C3 plants under drought. Funct. Plant Biol. 29: 1209–1215.
Giorio, P., Sorrentino, G. and d’Andria, R. 1999. Stomatal behaviour leaf water status and photosynthetic response in field-grown olive trees under water deficit. Environ. Exp. Bot. 42: 95–104.
Guerfel, M., Baccouri, O., Boujnah, D., Chaïbi, W. and Zarrouk, M. 2009. Impacts of water stress on gas exchange, water relations, chlorophyll content and leaf structure in the two main Tunisian olive (Olea europaea L.) cultivars. Scientia Horticulturae. 119(3): 257-263.
Hayat, Q., Hayat, S., Irfan, M. and Ahmad, A. 2010. Effect of exogenous salicylic acid under changing environment: a review. Environmental and Experimental Botany. 68(1):14-25.
Hayat, S., Ahmad, A., and Alyemeni, M.N. 2013. Salicylic Acid: Plant Growth and Development. Springer Science & Business Media.
Hazrati, S., Tahmasebi-Sarvestani, Z., Modarres-Sanavy, S.A.M., Mokhtassi-Bidgoli, A. and Nicola, S. 2016. Effects of water stress and light intensity on chlorophyll fluorescence parameters and pigments of Aloe vera L. Plant Physiology and Biochemistry. 106: 141-148.
Huerta, L., Forment, J., Gadea, J., Fagoaga, C., Pena, L., Pérez‐Amador, M.A. and García‐Martínez, J.L. 2008. Gene expression analysis in citrus reveals the role of gibberellins on photosynthesis and stress. Plant Cell Environ. 31:1620–1633.
Hussain, M., Malik, M.A., Farooq, M., Khan, M.B., Akram, M. and Saleem, M.F. 2009. Exogenous glycine betaine and salicylic acid application improves water relations, allometry and quality of hybrid sunflower under water deficit conditions. Journal of Agronomy and Crop Science. 195: 98–109.
Idrees, M., Khan, M.M.A., Aftab, T., Naeem, M. and Hashmi, N. 2010. Salicylic acid-induced physiological and biochemical changes in lemongrass varieties under water stress. Journal of Plant Interactions. 5(4): 293-303.
Kaur, S., Gupta, A.K. and Kaur, N. 2000. Effect of GA3, kinetin and indole acetic acid on carbohydrate metabolism in chickpea seedlings germinating under water stress. Plant Growth Regulation. 30 (1): 61-70.
Kawano T. and Muto, S. 2000 – Mechanism of peroxidase actions for salicylic acid-induced generation of active oxygen species and an increase in cytosolic calcium in tobacco cell suspension culture. J. Exp. Bot. 51: 685-693.
Kaya, C., Kirnak, H., Higgs, D. and Saltali, K. 2002. Supplementary calcium enhances plant growth and fruit yield in strawberry cultivars grown at high salinity. Sci. Hortic. 93: 65-74.
Khan, W., Prithiviraj, B. and Smith, D.L. 2003. Photosynthetic responses of corn and soybean to foliar application of salicylates. Journal of plant physiology. 160 (5):485-492.
Lichtenthaler, H.K. and Wellburn, A.R. 1983. Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents.
Liu, C., Liu, Y., Guo, K., Fan, D., Li, G., Zheng, Y., Yu, L. and Yang, R. 2011. Effect of drought on pigments, osmotic adjustment and antioxidant enzymes in six woody plant species in karst habitats of southwestern China. Environmental and Experimental Botany. 71(2):174-183.
Liu, F. and Stützel, H. 2004. Biomass partitioning, specific leaf area, and water use efficiency of vegetable amaranth (Amaranthus spp.) in response to drought stress. Scientia Horticulturae. 102:15-27.
Martin-Mex, R., Villanueva-Couoh, E., Herrera-Campos, T. and Larque-Saavedra, A. 2005. Positive effect of salicylates on the flowering of African violet. Science Horticulture, 103: 499–502.
Martín-Mex, R., Villanueva-Couoh, E. and Uicab-Quijano, V. 2003. Positive effect of salicylic acid on the flowering of gloxinia. In: Proceedings 31st Annual Meeting. Plant Growth Regulatory Society America, Canada: Vancouver (pp 149–151). August 3–6 2003.
Medrano, H., Escalona, J.M., Bota, J., Gulías, J. and Flexas, J. 2002. Regulation of photosynthesis of C3 plants in response to progressive drought: stomatal conductance as a reference parameter. Annals of Botany. 89:895-905.
Patel, M.S. and Golakiya, B.A. 1988. Effect of Water-Stress on Yield Attributes and Yield of Groundnut (Arachis-Hypogaea). Indian Journal of Agricultural Sciences. 58(9):701-703.
Pirasteh-Anosheh, H., Emam, Y. and Pessarakli, M. 2013. Changes in endogenous hormonal status in corn (Zea mays) hybrids under drought stress. Journal of plant nutrition. 36(11):1695-1707.
Raskin, I. 1992. Role of salicylic acid in plants. Annual Review Plant Physiology. 43: 439–463.
Saruhan, N., Saglam, A. and Kadioglu, A. 2012. Salicylic acid pretreatment induces drought
tolerance and delays leaf rolling by inducing antioxidant systems in maize genotypes. Acta
Physiologiae Plantarum. 34: 97–106.
Shalhevet, J. 1993. Plants under salt and water stress. Plant Adaptation to Environmental Stress. 133-154.
Shamshiri, M.H., Mozafari, V., Sedaghati, E. and Bagheri, V. 2011. Response of petunia plants (Petunia hybrida cv. Mix) inoculated with Glomus mosseae and Glomus intraradices to phosphorous and drought stress. Journal of agricultural science and technology. 13: 929-942.
Shao, H.B., Chu, L.Y., Jaleel, C.A. and Zhao, C.X. 2008. Water-deficit stress-induced anatomical changes in higher plants. Comptes Rendus Biologies. 331:215-225.
Singh, B. and Usha, K. 2003. Salicylic acid induced physiological and biochemical changes in wheat seedlings under water stress. Plant Growth Regulation. 39 (2):137-141.
Kim, S.T., Kang, S.Y., Wang, Y., Kim, S.G., Hwang, D.H. and Kang, K.Y. 2008. Analysis of embryonic proteome modulation by GA and ABA from germinating rice seeds. Proteomics 8:3577–3587.
Tahir, M.H.N. and Mehid, S.S. 2001. Evaluation of open pollinated sunflower (Helianthus annuus L.) populations under water stress and normal conditions. Int. J. Agric. Biol. 3:236-238.
Taiz, L. and Zeiger, E. 2006. Plant Physiology Fourth Edition Sinauer Associates. Inc. Publishers. Sunderland, Massachusetts.
Tarchevskii, L.A. 2002 - Signal'nye sistemy kletok rastenii (Signal Systems of Plant Cells). Nauka, Moscow, 294 p.
Umebese, C.E., Olatimilehin, T.O. and Ogunsusi, T.A. 2009. Salicylic acid protects nitrate reductase activity, growth and proline in amaranth and tomato plants during water deficit. American Journal of Agricultural and Biological Sciences. 4 (3): 224-229.
Van Iersel, M.W., Dove, S., Kang, J.G. and Burnett, S.E. 2010. Growth and water use of petunia as affected by substrate water content and daily light integral. HortScience, 45(2):277-282.
Verbruggen, N. and Hermans, C. 2008. Proline accumulation in plants: a review. Amino acids. 35(4), pp.753-759.
Verma, V., Ravindran, P. and Kumar, P.P. 2016. Plant hormone-mediated regulation of stress responses. BMC plant biology, 16(1), p.86.
Wang, C., Yang, A., Yin, H. and Zhang, J. 2008. Influence of water stress on endogenous hormone contents and cell damage of maize seedlings. Journal of Integrative Plant Biology.50 (4): 427-434.
Waseem, M., Athar, H.U.R. and Asahrafi, M. 2006. Effect of salicylic acid applied through
rooting medium on drought tolerance of wheat. Pakistan Journal of Botany. 38:1127–1136.
Waterland, N.L., Campbell, C.A., Finer, J.J. and Jones, M.L. 2010. Abscisic acid application enhances drought stress tolerance in bedding plants. HortScience. 45(3):409-413.
Yamada, M., Morishita, H., Urano, K., Shiozaki, N., Yamaguchi-Shinozaki, K., Shinozaki, K. and Yoshiba, Y. 2005. Effects of free proline accumulation in petunias under drought stress. Journal of Experimental Botany. 56 (417): 1975-1981.
Yamasaki, S. and Dillenburg, L.R. 1999. Measurements of leaf relative water content in Araucaria angustifolia. Revista Brasilleira de Fisiologia Vegetal. 11: 69-75.
Yasseen, B.T. 1983. An analysis of the effects of salinity on leaf growth in Mexican wheats. PhD thesis, University of Leeds.
Yasseen, B.T. 1992. Physiology of water stress in plants. University of Mosul. University Press, Iraq.
Zarghami Moghaddam, M., Shoor, M., Ganjeali, A., Moshtaghi, N. and Tehranifar, A. 2014. Effect of salicylic acid on morphological and Ornamental characteristics of petunia hybrida at drought stress. Indian Journal of Fundamental and Applied Life Sciences, 4.
Zhang, S. and Liu, Y. 2001. Activation of salicylic acid-induced protein kinase, a mitogen-activated protein kinase, induces multiple defense responses in tobacco. Plant Cell. 13:1877 1889.
Morteza Goldani
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, Iran
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, Iran
Ali Dolatkhahi
Department of Horticultural Science and Landscape, Ferdowsi University of Mashhad, Mashhad, Iran
Department of Horticultural Science and Landscape, Ferdowsi University of Mashhad, Mashhad, Iran
Mahdi Parsa
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, Iran
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, Iran
Navid Vahdati
Department of Horticultural Science and Landscape, Ferdowsi University of Mashhad, Mashhad, Iran
Department of Horticultural Science and Landscape, Ferdowsi University of Mashhad, Mashhad, Iran
Zahra Rasouli
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, Iran
Department of Agrotechnology, Ferdowsi University of Mashhad, Mashhad, Iran
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