Document Type : Original Article

Authors

1 Associate Professor, Department of Agronomy and Plant breeding, Yadegar-e-Imam Khomeini (RAH) Shahre-Rey Branch, Islamic Azad University, Tehran, Iran.

2 M.Sc in Agronomy, Eslamshahr Branch, Islamic Azad University and the member of Young researcher's and elit club, Yadegar-e-Imam Khomeini (RAH) Shahre-Rey Branch, Islamic Azad University, Tehran, Iran.

Abstract

Introduction
Summer savory (Satureja hortensis L.) is a genus of aromatic plants of the family Lamiaceae. There are about 30 different species called savories, which summer savory and winter savory are the most important in cultivation. Satureja species are native to warm temperate regions and may be annual or perennial (Hadian et al., 2008; Rechinger, 1982). Soil salinity is one of the main environmental stresses affecting the growth of plants and their yield (Allakhverdiev et al., 2000). Adaptation of plants to environmental stresses such as salinity performs by accumulation of metabolites as carbohydrates and proline (Sanito di Toppy and Gabbrielli, 1999). Ascorbic acid is a powerful antioxidant that leads to inhibition of the free radicals by reduction (Fecht Christoffers et al., 2003). Gibberellins are commercial mushroom cultures obtained from pure and natural product known that used in plants. There are four types of GA; the best known is gibberellic acid which has positive effect on metabolites related to salinity tolerance (Hedden and Proebsting, 1999).
 
Material and methods
Due to study the effect of salinity stress on some non-enzymatic mechanisms of savory under ascorbate and gibberellin application, a pot experiment was done in a greenhouse located in Pakdasht region and Islamic Azad University, Eslamshahr and Yadegar-e-Imam Khomeini (RAH) Shahre-rey Branchs in 2013. The experimental design consisted of a sixteen treatments, arranged as factorial based on completely randomized design with four replications, giving a total of 64 pots. Five-weeks old savory plants were sprayed with ascorbic acid (0 and 4 mM) and gibberellins
(0 and 4 mM) before inducing salinity stress with different concentrations of saline water
(0, 25, 50 and 75 millimolar). Control plants were sprayed with distilled water. Saline water was delivered to plants twice a week and last for three weeks. The proline by Bates et al (1973), Protein by Bradford (1976) and soluble and non soluble carbohydrates by Kochert (1978) were determined. The obtained data were subjected to analysis of variance using Duncan Multiple Range Test (P≤0.05) using SAS software.
 
 
Results and discussion
The results showed that, with the exception of ascorbate effect on shoot soluble carbohydrates, the other main effects of experimental factors were significant on all non-enzymatic mechanisms of savory tolerance. So the intense salinity showed the highest decreased in physiological traits such as shoot and root protein (304.93 mg/L and 147.71 mg/L) and shoot and root non soluble carbohydrates (11.27 and 8.25) and improved shoot and root proline (27.46 mg/L and 16.52 mg/L) and shoot and root soluble carbohydrates (18.49 mg/l and 16.32 mg/l) to decrease the harmful effects of salt stress. The results revealed that only triple interaction effects of experimental factors were significant on shoot soluble carbohydrates and shoot and root non soluble carbohydrates and protein. So, the shoot soluble carbohydrates and Shoot and root proline content has 8.23, 1.92 and 3.73 manifold increase than control in 75 mM NaCl and ascorbate and gibberellin foliar application alternatively. Therefore it can be concluded that shoot soluble carbohydrates is the most main non-enzymatic mechanisms of salt tolerance in savory especially in ascorbate and gibberellins foliar application. Some reports indicated that ascobate increased proline content in bean and pea (Alqrainy, 2007) and okra (Baghizadeh et al., 2009) that are coordinate to our findings. Hoda et al (2010) revealed that GA improved both Shoot and root soluble and non soluble carbohydrates as the best and rapid non-enzymatic mechanisms of stress tolerance in citrus.
 
Conclusion
The results showed that salinity stress increased shoot and root soluble carbohydrates and proline and reduced shoot and root non soluble carbohydrates and protein. At the same time, spraying 4 ascorbate and 2 mM GA affected all associated non-enzymatic traits to salt stress resistance, however the additive effect of GA was more than ascorbate.

Keywords

Akbari, N., Barani, M., Ahmadi, H., 2008. Effect of Gibberelic Acid (GA3) on Agronomic thaits of Green Gram (Vigna radiata L. wilczek) irrigated with different levels of saline water. World Applied Science Journal. 5(2), 199-203.
Alian, A., Altman, A., Heuer, B., 2000. Genotypic difference in salinity and water stress tolerance of fresh market tomato cultivars. Plant Science. 152, 59-65.
Allakhverdiev, S.L., Sakamoto, A., Nishiyama, Y., Inaba, M., Murata, N., 2000.Ionic and osmotic effects of Nacl-induced in activation of photo system І and ІІ in Synechococcus sp. Journal of Plant Physiology. 123, 1047-56.
Alqrainy, F., 2007. Responses of bean and pea to vitamin C under salinity stress. Journal of Agriculture and Biological Science. 3(6), 714-722.
Asada, K., 1999. The water–water cycle in chloroplasts: scavenging of active oxygens and dissipation of excess photons. Annual Review of Plant Physiology and Plant Molecular Biology. 50, 601–639.
Baghizadeh, A., Ghorbanli, M., Rezaei, H.M., Mozafri, H., 2009. Evaluation of Interaction effect of drought stress with ascorbate and salicylic acid on some of physiological and Biochemical parameters in okra (Hibiscus esculentus L.). Journal of Biological sciences. 4 (4), 380-387.
Bates, L., Waldern, R.P., Teare, I.D., 1973. Rapid determination of free Proline for water stress Studies. Plant and Soil, pp. 205-207.
Beltagi, M.S., 2008. Exogenous ascorbic acid (Vitamin C) induced anabolic changes for salt tolerance in chick pea (Cicer arietinum L.) plants. African Journal of Plant Science. 2(10), 118-123.
Bradford, M.M., 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Annal Biochemistry, 72, 248-254.
Buhnert, H.J., Nelson, D.E., Jensen, R.G., 1995. Adaptations to environmental stresses. The Plant Cell. 7(7), 1099-1111.
Dolatabadian, A., Sanavy, S.A.M.M., Chashmi, N.A., 2008. The effects of foliar application of ascorbic acid (vitamin C) on antioxidant enzymes activities, lipid peroxidation and proline accumulation of Canola (Brassica napus L.) under conditions of salt stress. Journal of Agronomy and Crop Science. 194(3), 206-213.
Fecht Christoffers, M.M., Maier, P., Horst, W.J., 2003. Apoplastic peroxidases and ascorbate are involved in manganese toxicity and tolerance of Vigna unguiculata. Journal of Plant Physiology. 117, 237-244
Foyer, C.H., Lelandais, M., Kunert, K.J., 1994. Photooxidative stress in plants. Plant Physiology. 92, 696-717.
Hadian, J., 2008. Assessment of genetic diversity of savory native species fromIran. Horticultural Science thesis. Tehran University. 180pp. [In Persian with English Summary].
Hadian, J., Tabatabaei, S.M.F., Naghavi, M.R., Jamzad, Z., Ramak-Masoumi, R., 2008. Genetic diversity of Iranian accessions of Satureja hortensis L. based on horticultural traits and RAPD markers. Scientia Horticulture. 115, 196-202.
Hedden, P., Proebsting, W.M., 1999. Genetic analysis of gibberellin Biosynthes is,Plant Physiology. 119, 365-370.
Heidari, M., Mesri, F., 2008. Salinity effects on compatible solutes, antioxidants enzymes and ion content in three wheat cultivars. Pakistan Journal of Biological Sciences. 11(10), 1385-1389.
Hoda, M., Abd EL-Rahman, G.F., Abd EL-Raheem, M.E., 2010.Impact of gibberellic acid enhancing treatments on shortening time to budding of citrus nursery stocks. Journal of American Science. 6(12), 410- 422.
Inanloofar, M., Omidi, H., Pazoki, A.R., 2013. Morphological, agronomical changes and oil content in purslane (Portulaca oleracea L.) under drought stress and biological/ chemical fertilizer of nitrogen. Journal of Medicinal Plants. 12 (48), 170-184. [In Persian with English Summary].
Khosravinejad, F., Heydari,R., Farboodnia,T., 2009. Effect of salinity on organic solutes contents in barley. Pakistan Journal of Biological Sciences. 12(12), 158-162.
Kochert, G., 1978. Carbohydrate determinationby the phenol sulfuric acid method. In: Helebust.J.A.Craig J.s. (ed): Hand book of phycological Method 56-97. Cambridge Univ. Press. Cambridge.
Moghadam, S., Mehrafarin, A., Naghdi Badi, H., Pazoki, A.R., Ghavami, N., 2012., Evaluation of phytochemical yield of thyme (Thymus vulgaris L.) under foliar application of hydroalcohols. Journal of Medicinal Plants. 11(44), 130-139. [In Persian with English Summary].
Maghsoudi Moud, A., Maghsoudi, K., 2008. Salt stress effects on respiration and growth of germinated seed of different wheat (Triticum aestivum L.) cultivars. World Journal of Agricultural Sciences. 4(3), 351-358.
Molassiotis, A., Sotiropoulos, T., Tanou, G., Diamantidis, G., Therios, I., 2006. Boron induced oxidative damage and antioxidant and nucleolytic responses in shoot tips culture of the apple rootstock EM9 (Malus domestica Borkh). Environmental and Experimental Botany. 56, 54-62.
Mostafa, G,G., Abou Alhamd, M.F., 2011. Effect of Gibberellic acid and indole 3- acetic acid on improving growth and accumulation of phytochemical composition in Balanites aegyptiaca plants. American Journal of Plant Physiology. 6(1), 36-43.
Nasir Khan, M., Siddiqui, M.H., Mohammad, F., Masroor, M., Khan, A., Naeem, M., 2007. Salinity induced changes in growth, enzyme activities, photosynthesis, proline accumulation and yield in linseed genotypes. World Journal of Agricultural Sciences. 3, 685-695.
Nedjimi, B., Daoud, Y., Touati, M., 2006. Growth, water relations, proline and ion content of in vitrocultured Atriplex halimus subsp. schweinfurthii as affected by CaCl2. Communications in biometry and crop science. 1(2), 79-89.
Noctor, G., Foyer, CH., 1998. Ascorbate and glutathione: Keeping active oxygen under control. Annual Review of plant physiology and plant Molecular Biology. 49, 249-279.
Novak, J., Bahoo, L., Mitteregger, U., Franz, C., 2006. Composition of individual essential oil glands of savory (Satureja hortensis L., Lamiaceae) from Syria. Flavor and Fragrance Journal. 21(4), 731-734.
Rechinger, K.H., 1982. Satureja. In Flora desiranischen hoclandes and derumrahmenden gebirge. Akademische druku verlags antalt graz Austria. 150, 495–504.
Sanito di Toppy, L., Gabbrielli, R., 1999. Response to cadmium in higher plants. Environmental and Experimental Botany. 41, 105-130.
Shahsevand Hassani. H., 2000. The process of production new allopeloeid tritipyrum. 6th Iranian Crop Science Congress, Babolsar, pp: 22-24. [In Persian with English Summary].
Shalata., A., Neumann, P.M., 2001. Exogenous ascorbic acid (Vitamin C) increases resistance to salt stress and reduces lipid peroxidation. Journal of Experimental Botany. 52, 2207–2211.
Sheteawi, S.A., 2007. Improving growth and yield of salt-stressed soybean by exogenous application of jasmonic acid and ascobin. International Journal of Agriculture and Biology. 9(3), 473-478.
Singh, L., Pal, B., 1995. Effect of water salinity on yield and yield attributing characters of blond psyllium (Plantago ovata). Indian Journal of Agricultural Sciences. 65(7), 503-505.
Smirnoff, N., Wheeler, G.L., 2000. Ascorbic acid in plants: biosynthesis and function. CRC Critical Review in Plant Sciences. 19, 267-290.