Document Type : Original Article

Authors

1 MSc of Dessert Management, Faculty of Natural Resources and Deserts, Yazd University, Yazd, Iran.

2 Associate Professor. Faculty of Natural Resources and Deserts, Yazd University, Yazd, Iran.

3 Associate Professor. Faculty of Natural Resources, Yazd University, Yazd, Iran

4 Assistant Professor, National Salinity Research Center, Agricultural Research, Education and Extension Organization (AREEO), Yazd, Iran.

5 PhD student of Islamic Azad University, Science and Research Branch, Tehran, Iran.

Abstract

Introduction
Abiotic stresses adversely affect agricultural productivity worldwide. In the arid and semi-arid areas, water is one of the most basic ecological units, which determines the growth and development of plants. Therefore, efficient use of water resources should be considered in order to achieve desirable plant growth. Aside different effective factors, minerals such as silicon are the most essential affecting the growth and development of plants. Silicon has beneficial effects on many crops, especially under biotic and abiotic stresses. Silicon can affect biochemical, physiological, and photosynthetic processes, consequently ameliorates drought stress. However, the effects of silicon on stevia (Stevia rebaudiana L.) plants under drought stress has not well known. Stevia is a branched bushy shrub of the Asteraceae family and originated from South America, and its cultivation has spread worldwide. Stevia is well known for its high content of sweet components.

Materials and Methods
The purpose of this research was to determine the effects of different silicon concentrations on physiological and morphological characteristics of stevia under drought stress conditions. The treatments were four levels of drought stress including: 25%, 50%, 75%, 100% pot capacity and three levels of silicon concentrations consisted of 0, 1, 1.5 mM, which arranged in a factorial experiment based on completely randomized design with three replications. After the plants establishment, drought treatments were imposed. After 40 days of applying drought treatments, silicon was twice sprayed with 7 dyas interval. The leaves of plants were sampled one week after the last spraying and transferred to the laboratory and the amount of soluble sugars, proline and chlorophyll concentrations were measured. In order to analysis data, analysis two-way ANOVA were used and Duncan test compared means after normality test. Statistical analysis was performed using SPSS software, and Excel software was used for drawing the charts.

Results and Discussion
Applying experimental treatments had only significant effect (p<0.01) on plant height, shoot fresh weight, canopy diameter, and canopy volume, while had no significant effect on shoot dry weight and number of leaves. The effect of silicon foliar application was significant on plant height, canopy diameter and canopy volume at 1% probability level and on shoot fresh weight and shoot dry weight at 5% probability level. The interaction between stress and silicon only was significant (p<0.01) on shoot dry weight, diameter and volume canopy, but had no significant effect on other traits. The results also showed that in no spraying, 1 and 1.5 mM silicon treatments under 25% FC significantly increased soluble sugars compared to 100% PC, so that the maximum amount of soluble sugars found in 25% FC and 1.5 mM silicon. Since the sugars are made from compatible osmolytes and its accumulation lead to osmotic adjustment, keep the cellular turgor and stability of protein. Thus increasing in soluble sugars such as sucrose, glucose and fructose as a result of environmental stresses, particularly drought are considered as a strategies of plants to tolerate adverse environmental conditions. Increasing drought stress to 25% PC significantly increased proline of stevia by 74% compared to the three levels of 100%, 75% and 50% PC. The results of several studies showed the positive effect of drought stress on proline, so that proline content is greater in drought stress than normal conditions. Increasing proline content in stress conditions might be due to increasing in activity of enzymes are responsible for the synthesis of this material, reducing the proline oxidation to glutamate and/or decreasing in consumption of proline in the process of making proteins. The results of evolution of different treatments of drought stress on chlorophyll content of stevia plant showed that the effect of drought stress was significant (p<0.01) on chlorophyll b, so that the most of chlorophyll b was observed in 75% PC treatment, which had no significant difference to 50% and 25% PC treatments, and the lowest was observed in 100% PC. Increasing inconsiderably chlorophyll b in drought stress conditions showed that chlorophyll pigments are somewhat tolerance to water loss.

Conclusions
The results of this experiment showed that height, shoot fresh weight of, diameter and volume of canopy, root volume, dry and root fresh weight of, and chlorophyll b were significantly decreased with increasing in drought stress level, however, using of both different silicon concentrations, especially 1 mM modulated damages of drought stress on stevia. Generally, the results of this research revealed the positive role of silicon in decreasing the negative effects of drought stress in stevia.

Keywords

Abaszadeh, B., Sharifiashorabadi, A., Lebaschi, M.H., Naderi Haji Bagherkandi, M., Moghadami, F., 2008. The effect of drought stress on proline contents, soluble sugars, chlorophyll and relative water contents of balm (Melissa officinalis L.).         Iranian Journal of Medicinal and Aromatic Plants. 23(4), 504-513. [In Persian with English summary].
Ahmed, M., Hassen, F., Khurshid. Y., 2011. Does silicon and irrigation have impact on drought tolerance mechanism of sorghum? Agricultural Water Management. 98(12), 1808-1812.
Ali, A., Gull, I., Nas, S., Afghan, S., 2010. Biochemical investigation during different stages of in vitro propagation of Stevia rebaudiana. Pakistan Journal of Botany. 42(4), 2827-2837.
Alizadeh, A., 2005. Soil, Water and Plant Relationship. Astane qodse razavi publication. 222p. [In Persian].
Aranjuelo, I., Molero, G., Erice, G., Christophe Avice, J., Nogues, S., 2011. Plant physiology and proteomics reveals the leaf response to drought in alfalfa (Medicago sativa L.). Experimental Botany. 62,111-123.
Amini, S., Ghobadi, C., Yamchi, A., 2015. Proline accumulation and osmotic stress, an overview of p5cs gens in plants. Journal of Plant Molecular Breeding. 3(2), 44-55.
Aromatic Plants. 23(4), 504-513. [In Persian with English summary].
Armizatul, S.A.H.R., Mohd Aziz, A.B., Azhar, k., 2010. Effects of night break on Stevia rebaudiana Poster presented at 21th Malaysian Society of Plant Physiology Conference, Cameron Highland.
Bansal, K.C., Nagarajan, S., Sukumaran, N.P., 1991. A rapid screening technique for drought resistance in potato (Solanum tuberosum L.). Potato Research. 34, 241-248.
Bates, L., Waldren, R.P., Teare, I.D. 1973. Rapid determination of free proline for water stress studies. Plant and Soil. 39, 205-207.
Cao, B. I., Ma, Q., Zhao, Q., Wang, L., Xu, k., 2015. Effects of silicon on absorbed light allocation, antioxidant enzymesand ultrastructure of chloroplasts in tomato leaves under simulated drought stress. Scientia Horticulturae. 194, 53-62.
Chanbdracar, B.L., Sechar, N., Tuteja, S.S., Tripathi, R.S., 1994. Effect of irrigation and nitrogen of growth and yield of summer sesame (Sesamum indicum) Indean Journal Agronomy. 39, 701-702.
Chandra Obul Reddy, P., Sairanganayakulu, G., Thippeswamy, M., Sudhakar Reddy, P., Reddy M.K., Chinta Sudhakar, H., 2008. Identification of stress-induced genes from the drought tolerant semi-arid legume crop horsegram. (Macrotyloma uniflorum (Lam) Verdc) through analysis of subtracted expressed sequence tags. Plant Science. 175, 372-384.
Connor, D.G., Jones, T.K., 1985. Response of sunflower to strategies of irrigation: II. Morphological and physiological responses to water stress. Field Crops Research. 12, 91-103.
Farooq, M., Wahid, A., Kobayashi, N., Fujita, D., Basrs, S.M.A., 2009. Planr drought stress, effects, mechanisms and management. Agronomy for Sustainable Development. 29, 185-212.
Gagoonani, S., Enteshari, S., Delavar, K., Behyar, M., 2011. Interactive effects off silicon and aluminum on the malondialdehyde (MDA), proline, protein and phenolic compounds in Borago officinalis L. Journal of Medicinal Plants Research. 5(24), 5818-5827.
Geuns, J.M.C., 2003. Molecules of interest: Stevioside. Phytochemistry. 64(5), 913-921.
Ghorbanli, M., Gafarabad, M., Amirkian, T., Allahverdi Mamaghani, B., 2013. Investigation of proline, total protein, chlorophyll, ascorbate and dehydro ascorbate changes under drought stress in Akria and Mobil tomato cultivars. Journal of Plant Physiology. 3(2), 651- 658. [In Persian with English summary].
Gong, H., Zhu, X., Chen, K., Wang, S., Zhang, C., 2005. Silicon alleviates oxidative damage of wheat plants in pots under drought. Plant Science. 169, 313-321.
Emad, M., 2007. 108p. Identification of Medicinal, Industrial, Pasture and Forest Plants and their Use. Tosee Rostaee Publication. [In Persian].
Hanson, A.D., Nelson, C.E., Pederson, A.R., 1999. Capacity for proline accumulation during water stress in barley and implications for breeding for drought stress. Crop Science. 19, 489-493
Hassanzadeh-Delouei, M., Vazin, F., Nadaf, J., 2013. Effect of salt stress in different stages of growth on qualitative and quantitative characteristics of cumin (Cuminum Cyminum L.). Cercetari Agronomice in Moldova. 1153, 89-97.
Hidari, M., Shahpesandy, S., Mosavi nik, S.M., Bijani, M., 2015. Effect of Nano-silicon on the germination and seedling growth of native and bread cultivars of wheat (Triticum aestivum L.). Journal of Seed Ecophysiology. 1(1), 1-16. [In Persian with English summary].
Hu, C.A., Delauney, A.J., Verma, D.P., 1992. A bifunctional enzyme (al-pyrroline-5-carboxylate synthetase) catalyzes the first two steps in proline biosynthesis I plants. Proceeding of the National Academy of Science. 89, 9354-9358.
Huany, B., Johnson J.W., 1995. Root respiration and carbohydrate status of two wheat genotypes in response to hypoxia. Annals of Botany. 75, 427-432.
Kage, H., Kochler, M., Stuzel, H., 2004. Root growth and dry matter partitioning of cauliflower under drought stress conditions, measurement and simulation. European Journal of Agronomy. 20, 379-394.
Khajeh, M., Mosavinik, M., Siroosmehr, A., Yadoalahi Dehcheshmeh, P., Amiri, A. 2015. Effects of water stress and spraying silicone on wheat yield and photosynthetic pigments in Sistan. Crop Physiology. 26(7), 5-19. [In Persian with English summary].
Kim, Y.H., Khan, A.L., Hamayun, M., Kang, S.M., 2011. Influeence of short-term silicon application on endogenous physiohormonal levels of Oryza sativa L. under wounding stress. Biological Trace Element Research Journal. 144, 1175-1185.
Kochert, G., 1978. Method Phycologia and Biochemical Methods: Carbohydrate Determination by the Phenol Sulfuric Acid. Cambridge University Press, London.
Lehmann, S., Funck, D., Szabdos, L., Rentsch, D., 2010. Proline metabolism and transport in plant development. Amino Acids. 39, 949-962.
Lichtenthaler, H.K., 1987. Chlorophylls and carotenoids pigments photosynthetic membranes. Methods in Enzymology. 148, 350-382.
Ma, J.F., Yamaji, N., 2006. Silicon uptake 6and accumulation in higher plants. Plant Science. 11, 392-397.
Omidbaigi, R., 1997. Production and Processing of Medicinal Plants. Tehran Nashr Publisher. 196p. [in Persian].
Raj, H.A., Thakral, K.K., 2008. Effect of chemical fertilizers on growth, yield and quality of fennel. Weed Technology. 17, 134-139.
Ramesh, K.V., Singh, N.W., Megeji, K., 2006. Cultivation of stevia (Stevia rebaudiana Bertoni): A comprehensive review. Advances in Agronomy Journal. 89, 137-177.
Richmond, K.E., Sussman, M., 2003. Got silicon the non-essential beneficial plant nutrient. Current Opinions in Plant Biology. 6, 268-272.
Saadatmand, M., Enteshari, Sh., 2012. Effects of pretreatment with silicon on salt tolerance in Iranian borage (Echium amoenum Fisch& C.A. mey). Science and Technology of Greenhouse Culture. 3(12), 45-56. [In Persian with English summary].
Safikhani, F., HidariShariabadi, H., SharifiAshorabadi, A., Siadat, A., SeyedNejad, M., Abaszadeh B. 2008. The effect of stress water on the function of various organs Badrashbu (Dracocephalum moldavical) under greenhouse conditions. Pajouhesh & Sazandgi. 21, 2-12. . [In Persian with English summary].
Sarmadnia, G.H., Koocheki, A., 1997. Crop Physiology. Jahad Daneshgahi Mashhad Press [In Persian].
Shariat, A., Asare, M.H., 2008. Effects of drought stress on pigments, prolin, soluble sugars and growth parameters on four eucalyptus species. Pajouhesh & Sazandegi, 78, 139-148. . [In Persian with English summary].
Shi, Y., Zhang, Y., Yao, h., Wu, J., Sun, H., Gong, H., 2014. Silicon improves seed germination and alleviates oxidative stress of bud seedlings in tomato under water deficit stress. Plant Physiology and Biochemistry. 78, 27-36.
Siddique, M.R.B., Hamid, A., Islam, M.S., 1999. Drought stress effects on photosynthetic rate and leaf gas exchange of wheat. Botanical Bulletin of the Academia Sinica. 40, 141-145.
Szabdos, L., Savoure, A., 2009. Proline amultifuntional amino acid. Trends in Plant Science. 15, 89-97.
Taiz, L., Zeiger, E. 2006. Plant physiology. Fourth Edition. Sinauer Associates. Sunderland, MA. 764 p.
Tajamolian, M., 2011. Assessing water relationships of Fortune bingo in dry climate. MSc. Dissertation, Faculty of Natural Resources, University of Yazd, Iran. [In Persian with English summary].
Tajamolian, M., Iran Nezhad Parizi, M.H., Malekinezhad, H., Rad, M.H., Sodaei Zadeh, H., 2013. Effects of different drought Stress treatments on some morphological characteristics of Fortuynia bungei Boiss. Journal of Rangeland. 9(3), 294-303.
Torabi, F., Majd, A., Enteshari, Sh., Irian, S., 2013. Study of Effect of Silicon on Some Anatomical and Physiological Characteristics of Borage (Borago officinalis L.) in Hydroponic Conditions. Journal of Cell & Tissue. 4(3), 275-285.
Verma, S., Dubeym, S., 2001. Effect of cadmium on soluble and enzymes of their metabolism in rice. Biology Plantarum. 1,117-123.
Yanqiong, L., Xingliang, L., Shaowei, Z., Hong, C., Yongjie, Y., Changlong, M., Jun, L., 2007. Drought-resistant physiological characteristics of four shrub species in arid valley of Minjiang River, China. Acta Ecologica Sinica. 27(3), 870-877.
Yin, L., Wang, S., Liu, P., Wang, W., Cao, D., Deng, X., 2014. Silicon-mediated changes in polyamine and 1-aminocyclopropane-1-carboxylic acid are involved in silicon-induced drought resistance in Sorghum bicolorL. Plant Physiology and Biochemistry. 80, 268-277