International Journal of Agriculture and Crop Sciences. Available online at www.ijagcs.com IJACS/2012/4-11/685-690 ISSN 2227-670X ©2012 IJACS Journal
Impact of exogenous salicylic acid application on some traits of common bean (Phaseolus vulgaris l.) under water stress conditions Omid Sadeghipour*, Parviz Aghaei Department of Agronomy, Shahre-Rey Branch, Islamic Azad University, Tehran, Iran Corresponding Author email:
[email protected] ABSTRACT: Salicylic acid (SA) is an endogenous plant growth regulator of phenolic nature that plays a key role in the regulation of plant growth, development, interaction with other organisms and in the responses to environmental stresses such as drought. An experiment was therefore, conducted to evaluation the influence of exogenous SA application on some traits of common bean under water stress conditions in Iran during 2011. Results showed that drought decreased plant height, leaf area index (LAI) and protein yield but increased seed protein content. Nevertheless, seeds soaking in SA (especially 0.5 mM) diminished drought damages and increased plant height, LAI and protein yield in both water stress and optimum conditions. SA application also decreased seed protein content. Results indicate that exogenous application of this phytohormone can act as an effective tool in improving the growth and production of common bean under water stress conditions. Keywords: Salicylic acid; drought; plant height; leaf area index; seed protein content; protein yield. INTRODUCTION Food productivity is decreasing due to detrimental effects of various biotic and abiotic stresses; therefore minimizing these losses is a major area of concern to ensure food security under changing climate. Environmental abiotic stresses, such as drought, extreme temperature, cold, heavy metals or high salinity, severely impair plant growth and productivity worldwide (Anjum et al., 2011). Drought being the most important environmental stress severely impairs plant growth and development, limits plant production and the performance of crop plants more than any other environmental factor (Shao et al., 2009). Available water resources for successful crop productions have been decreasing in recent years. Furthermore, in view of various climatic change models scientists suggested that in many regions of the world, crop losses due to increasing water shortage will further aggravate its impacts (Anjum et al., 2011). The susceptibility of plants to drought stress varies in dependence of stress degree, different accompanying stress factors, plant species and their developmental stages (Demirevska et al., 2009). Acclimation of plants to water deficit is the result of different events, which lead to adaptive changes in plant growth and physio-biochemical processes such as changes in plant structure, growth rate, tissue osmotic potential and antioxidant defenses (Duan et al., 2007). The common bean (Phaseolus vulgaris L.) is considered the most important legume for human nutrition in the world. However, this crop can be affected by several environmental stresses, and the drought is one of the most important causes of yield reduction. Common bean has low tolerance to water stress. This conditions is very important considering that as much as 60% of bean production in the world occurs under conditions of drought stress (Franca et al., 2000). The effect of drought is complex in its mode of action, highly variable in response, accentuated by interacting factors and localized within environmental regions. Yield and yield components of common bean are negatively affected by drought (Ramirez-Vallejo and Kelly, 1998). Various chemicals such as osmoprotectants, growth regulators and stress signaling molecules are being successfully used to induce the tolerance against several biotic and abiotic stresses (Farooq et al., 2010). Salicylic acid (SA) is an endogenous growth regulator of phenolic nature, which participates in the regulation of physiological processes in plants and it is also important in disease resistance (Raskin, 1992). It has been known for many years that exogenous SA is involved in the defense against pathogen attack and more recently its role has been widely investigated in both biotic and abiotic stresses (Shi et al., 2006). Many studies report the role of SA in inducing stress tolerance in plants. For example, SA has been found to induce drought tolerance in wheat (Singh and Usha, 2003), salinity tolerance in barley (El-Tayeb, 2005), heat tolerance in mustard (Dat et al., 1998), chilling tolerance in maize (Janda et al., 1999) and heavy metal stress tolerance in barley (Metwally et al., 2003). The results reported by Singh and Usha (2003) revealed that the wheat seedlings
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subjected to drought stress when treated with salicylic acid, generally exhibited higher moisture content and also higher dry matter accumulation, carboxylase activity of rubisco, superoxide dismutase and total chlorophyll content compared to the untreated plants. Further, the treatment also provided a considerable protection to the enzyme nitrate reductase thereby maintained the normal level of various proteins in the leaves. Therefore, the present study evaluates the role of exogenous SA application in improving of common bean drought tolerance, based on changes in some traits such as, plant height, leaf area index (LAI), seed protein content and seed protein yield MATERIALS AND METHODS This field trial was conducted during summer 2011 at the research farm of the Islamic Azad University of Shahre-Rey (51° 28´ E, 35° 35´ N and 1000 m above sea level) in Tehran, Iran. The climate of region is characterized as arid with the mean annual rainfall and temperature 201.7 mm and 20.4° C respectively. The soil of the experimental farm was sandy clay loam having pH 7.8, EC 2.8 ds m-1, nitrogen 0.091%, phosphorous 9.1 ppm and potassium 350 ppm. This study was laid out in a split plot on the basis of randomized complete block design with four replications. Each replication had two main plots as irrigation levels viz. I0: Irrigation after 50 mm evaporation from class A pan and I1: Irrigation after 100 mm evaporation from class A pan, as control and water stress conditions, respectively. Each main plot consisted of five sub plots as common bean (cv. Derakhshan) seeds were soaked for 6 h in salicylic acid solutions (0, 0.25, 0.5, 0.75 and 1 mM). Seeds before treatment were sterilized with sodium hypochlorite solution (1%) for 5 min and then washed thoroughly with distilled water. Seeds were treated with Bavistin and then were sown by hand on 12 June 2011 in 4 cm depth of soil. At the same time plots were fertilized with 100 kg ha-1 ammonium phosphate. Each sub plot had four planting rows with length and distance of 5 m and 0.5 m respectively, thus; size of each plot was 10 m2. Distances of plants on rows were 10 cm. All plots were irrigated immediately after sowing, but subsequent irrigations were carried out according to the treatments. Crop management practices such as hand weeding and thinning were done as required. At the flowering stage, all leaves of 10 plants from each plot were detached and leaf area was determined by using Handheld Laser Leaf Area Meter CI-203, CID, Bio-Science, USA. At physiological maturity, 10 plants from each plot were harvested to determine plant height and seed protein content by kjeldahl method (6.25 × N). Finally, plants in 2 m2 of each plot were harvested and sundried, then; seeds were detached from the pods and seed yield per unit area was recorded. Protein yield in unit area was measured by product of seed yield and seed protein percentage. Collected data were analyzed by MSTATC statistical software and the means were compared by Duncan's Multiple Range Test (DMRT) at the 5% probability level. RESULTS Results showed that plant height was decreased significantly under water stress conditions (21% as compared to control) but SA application (especially 0.5 mM) increased plant height in both drought and control conditions. Seeds soaking in 0.5 mM SA as compared to no application increased plant height by 6% and 5% under drought and control conditions, respectively. The highest plant height was recorded in control + SA application (0.5 mM) treatment, while the lowest plant height was produced in water stress + no application of SA treatment (Fig. 1). We observed that drought significantly decreased leaf area index (LAI) of common bean (31% as compared to control) nonetheless seeds soaking in SA increased leaf area index in the water stress conditions as well as control. Application of 0.5 mM SA as compared to no application, increased leaf area index by 29% under drought and optimum conditions. The maximum LAI was recorded in control + SA application (0.5 mM) treatment, against the minimum LAI was produced in water stress + no application of SA treatment (Fig. 2). Our results showed that seed protein content of common bean increased under drought conditions (5% as compared to control). On the other hand SA treatment decreased seed protein content not only in the water stressed plants but also in the well watered plants. Seeds soaking in 0.5 mM SA as compared to no application, decreased seed protein content by 5% and 4% under drought and control conditions, respectively. The highest seed protein content was achieved in water stress + no application of SA treatment, while the lowest seed protein content was observed in control + SA application (0.5 mM) treatment (Fig. 3). Water stress significantly decreased seed protein yield of common bean (33% as compared to control) nevertheless seeds soaking in SA (especially 0.5 mM) ameliorated seed protein yield in both drought and control conditions. Application of 0.5 mM SA as compared to no application, increased seed protein yield by 26% and 38% under drought and optimum conditions, respectively. The maximum seed protein yield was produced in control + SA application (0.5 mM) treatment, against the minimum seed protein yield was recorded in water stress + no application of SA treatment (Fig. 4).
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Figure 1. Effect of seeds soaking in salicylic acid (SA) on plant height of common bean under water stress and control conditions. Means with the same letter are not significantly different (P 0.05).
Figure 2. Effect of seeds soaking in salicylic acid (SA) on leaf area index of common bean under water stress and control conditions. Means with the same letter(s) are not significantly different (P 0.05).
Figure 3. Effect of seeds soaking in salicylic acid (SA) on seed protein content of common bean under water stress and control conditions. Means with the same letter are not significantly different (P 0.05).
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Figure 4. Effect of seeds soaking in salicylic acid (SA) on protein yield of common bean under water stress and control conditions. Means with the same letter are not significantly different (P 0.05).
DISCUSSION In the present study we observed that water stress reduced plant height of common bean but seeds soaking in SA ameliorated this trait in both water stress and control conditions. Under severe water deficiency, cell elongation of higher plants can be inhibited by interruption of water flow from the xylem to the surrounding elongating cells (Nonami, 1998). Drought caused impaired mitosis; cell elongation and expansion resulted in reduced growth and yield traits (Hussain et al., 2008). Similar to our results Bideshki and Arvin (2010) reported that, drought stress decreased plant height, leaf area and yield of garlic but application of SA improved these traits in both drought and control conditions. In another research drought stress substantially reduced the rice stem height; nevertheless application of various compounds such as glycinebetaine (GB), SA, nitric oxide, brassinosteroid and spermine assuaged the damaging effects of drought. Foliar spray of all chemicals improved growth possibly because of the improved carbon assimilation, enhanced synthesis of metabolites and maintenance of tissue water status (Farooq et al., 2010). Alike to our results, Umebese et al., (2009) found that water stress reduced tomato and amaranth stem height significantly at the vegetative stages and 3 mM application of SA was effective in keeping plant height similar to the control which was related to the ability of SA to induce antioxidant responses that protect them from damage. Role of SA in amelioration of plant height under water stress may be related to improve mitosis and cell elongation. Our findings are in agreement with the results of other researches (Gutiérrez-Coronado et al., 1998; Hussein et al., 2007). We found that drought decreased common bean LAI, nevertheless application of SA improved LAI not only in water stressed plants but also in well watered plants. Development of optimal leaf area is important to photosynthesis and dry matter yield. Water deficit stress mostly reduces leaf growth and in turns the leaf areas in many species of plant (Jaleel et al., 2009). Water deficit reduces the number of leaves per plant and individual leaf size, leaf longevity by decreasing the soil’s water potential. Leaf area expansion depends on leaf turgor, temperature and assimilating supply for growth. Drought-induced reduction in leaf area is ascribed to suppression of leaf expansion through reduction in photosynthesis (Anjum et al., 2011). Reduction of dry bean leaf area under drought stress conditions during vegetative growth stage has been reported in many studies (Emam et al., 2010; Nielsen and Nelson, 1998; Markhart, 1985). Indeed, loss of leaf area, which could be resulted from reduced size of younger leaves and inhibition of the expansion of developing foliage, is also considered as an adaptation mechanism to moisture deficit. Hussain et al., (2009) revealed that in sunflower; water stress reduced the LAI, leaf area duration (LAD), crop growth rate (CGR), relative water content (RWC), water potential, osmotic potential, turgor pressure, achene yield and water use efficiency. Nevertheless, exogenous GB and SA application appreciably improved these attributes under water stress. Under drought conditions SA treatment cause maintenance of RWC and photosynthesis thus improves leaf area index. Similar to our results, increasing of leaf area under treatment with SA has been reported in pearl millet (Mathur and Vyas, 2007), wheat (Hayat et al., 2005) and corn and soybean (Khan et al., 2003). Results of our experiment showed that, drought conditions increased seed protein content of common bean. Nonetheless, exogenous application of SA decreased this characteristic in control and drought conditions. A possible explanation for the increased protein content under drought conditions could be the de novo synthesis of drought proteins (De-Mejia et al., 2003). Hussain et al., (2009) reported that the protein contents of sunflower seeds were appreciably increased by increasing water stress at different growth stages, but were reduced by exogenous GB and SA application. One of the interesting findings of the study was that achene protein was 688
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reduced as a result of GB application and increasing water application, which might be due to dilution because yield was increased. This result is similar to our findings for SA application. Contrary to our observations Kumar et al., (1999) reported that the total protein content was increased in soybean plants sprayed with SA and this increase might be due to enhanced activity of nitrate reductase following the SA treatment. Further, in another study treatment with SA provided a considerable protection to the enzyme nitrate reductase thereby maintained the normal level of various proteins in the wheat leaves (Singh and Usha, 2003). It seems that influence of exogenous SA on protein content is related to plant species, stress intensity, SA concentration, application method and plant organ. We observed that drought decreased protein yield of common bean but seeds soaking with SA increased this attribute not only in water stress but also in optimum conditions. Flowering is one of the important parameter that is directly related to yield and productivity of plants. Salicylic acid has been reported to induce flowering in a number of plants (Hayat et al., 2010). Hussain et al., (2008) found that exogenous application of SA can improve yield and yield related traits in sunflower under drought stress. It may be the result of maintenance of photosynthetic activity owing to SA application. By means of osmotic adjustment, the organelles and cytoplasmic activities take place at about a normal pace and help plants to perform better in terms of growth, photosynthesis and assimilate partitioning to grain filling (Ludlow and Muchow, 1990; Subbarao et al., 2000). In agreement with our findings another researchers also reported that exogenous SA improved yield and yield components of garlic and sunflower under water stress conditions (Bideshki and Arvin, 2010; Hussain et al., 2008). Increase in parameters of water stressed plants in response to SA may be related to the induction of antioxidant responses that protect the plant from oxidative damage, accumulation of proline and maintenance of RWC and photosynthesis. In future, the exogenous application of this phytohormone might act as a powerful tool in enhancing the growth, productivity and also in combating the ill effects generated by various abiotic stresses in plants. The future applications of this plant hormone hold a great promise as a management tool for providing tolerance to our agricultural crops against the constrains consequently aiding to accelerate potential crop yield in near future (Hayat et al., 2010). In conclusion, our results showed that although common bean is a sensitive plant to water stress and plant height, LAI and protein yield are reduced with water deficit, it appears that seeds soaking in SA (especially 0.5 mM) can help to drought tolerance of this crop. 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