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Interactive effect of salinity stress and foliar application of salicylic acid on some physiochemical traits of chicory (Cichorium intybus L.) genotypes
Scientia Horticulturae 258 (2019) 108810
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Interactive effect of salinity stress and foliar application of salicylic acid on some physiochemical traits of chicory (Cichorium intybus L.) genotypes Nazanin Poursakhi, Jamshid Razmjoo, Hassan Karimmojeni
T
⁎
Department of Agronomy and Plant Breeding, College of Agriculture, Isfahan University of Technology, Isfahan, Iran
A R T I C LE I N FO
A B S T R A C T
Keywords: Chicory Abiotic stress Plant growth regulator Oxidative stress Photosynthetic pigments Growth
Chicory (Cichorium intybus) is an important medicinal, food and feed plant. Salinity can affect its growth and quality to some extent. However, adverse effects of salinity may be alleviated upon selecting salt tolerant genotypes and application of salicylic acid. Therefore, a 2-year pot experiment was conducted with three salinity levels (control, 65 mM and 130 mM NaCl), three levels of foliar application of salicylic acid (0, 0.5 mM and 1 mM SA) and seven chicory genotypes (Ardestan, Hamedan, Shiraz, Mazandaran, Sanandaj, Kashan and Yazd). Treatment characterized with 65 and 130 mM salinity levels led to significant decrease in relative water content (RWC), membrane stability index (MSI), chlorophyll a(Chl a), chlorophyll b (Chl b) and carotenoid (Car) contents and shoot dry weight (SDW) while significant increase in leaf proline (LPC) and malondialdehyde (MDA) contents. Moreover, the highest changes in measured traits were observed at salinity level of 130 mM. Salicylic acid application at levels of 0.5 and 1 mM SA significantly ameliorated all measured traits and the greatest values were attributed to 1 mM SA. Among genotypes, Sanandaj accounted for the highest measured traits but the lowest content of MDA while Ardestan characterized with the lowest measured traits and the highest content of MDA. Salinity decreased MSI, Car and SDW while it increased LPC and MDA in all genotypes with different magnitude. Application of SA reduced MDA in all genotypes and also led to increase in RWC, MSI, Chl a, Chl b, Car, LPC and SDW and diminution in MDA under control and specially under salt stress conditions. Sanandaj with the application of 1 mM SA under control and saline conditions was the best genotype among all genotypes. The results suggested that SA may improve performance of chicory genotypes under control and saline conditions by announcing physiochemical traits.
1. Introduction Soil salinity is one of the most outstanding limiting factors for crop production, especially in arid and semi-arid parts of the world (Silveira et al., 2001; Munns, 2002). In these areas, salinity may be arised due to soil intrinsic properties, limited precipitation and poor drainage conditions, inappropriate quality of irrigation water and intense evaporation during long hot and dry seasons (González-Acevedo et al., 2016). At the same time, improper farming methods and unsuitable management of irrigation systems have exacerbated this issue seriously. (Ashraf et al., 2008). Chicory is cultivated as a forage crop in different parts of the world (Belesky et al., 1999; Kalber et al., 2012). In addition, it is utilized for its leaves, buds and roots among others (Amaducci and Pritoni, 1998). It serves as a valuable source of calcium, magnesium,
potassium, manganese, zinc, and iron (Crush and Evans, 1990). The young leaves of chicory are edible and can be excellent addition to salads (Dalara and Konczaka, 2014). The processed roots of chicory can be consumed as a coffee substitution (Baert and Bockstaele, 1992). Boyd and Rogers (2004) pointed out that chicory has a degree of salinity resistance and this makes it suitable to be cultivated in relatively moderate salinity areas. Under salinity circumstance, reducing water availability because of low osmotic potential of soil solution, creation of ion toxicity effect, disintegration of membranes, nutritional imbalance and excessive production and accumulation of ROS (Chen et al., 2000) are factors causing major changes at physiological, biochemical, and molecular processes of plant cells (Munns, 2002). These are associated with reduction in RWC, MSI, photosynthetic pigments such as Chl a, Chl b, Car and shoot dry matter and increase in LPC and MDA contents that
Abbreviations: G, Genotype; S, Salinity; SA, Salicylic acid; RWC, Relative water content; MSI, Membrane stability index; Chla, Chlorophyll a; Chlb, Chlorophyll b; Car, Carotenoid; LPC, Leaf prolin content; MDA, Malondialdehyde; SDW, Shoot dry weight; PGR, Plant growth regulator; ROS, Reactive oxygen species; FW, Fresh weight ⁎ Corresponding author. E-mail address: [email protected] (H. Karimmojeni). https://doi.org/10.1016/j.scienta.2019.108810 Received 22 March 2019; Received in revised form 18 August 2019; Accepted 25 August 2019 0304-4238/ © 2019 Elsevier B.V. All rights reserved.
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with a constant volume 7 mL plant−1 of SA solution with a manual sprayer at 6–7 leaf growth stage. Control plants were sprayed similarly with equivalent amount of water. (Methenni et al., 2018; Tahjib-Ul-Arif et al., 2018). The pot soil was covered by a plastic sheet during the SA spray in order to avoid the access of SA solution to the root system.
ultimately lead to adverse impacts on plant growth and development (El-Tayeb, 2005; Koca et al., 2007; Rasool et al., 2013; Farahbakhsh et al., 2017). The main practices to deal with salinity are soil restoration and leaching operations. These methods are not cost-effective and sometimes are impossible. Therefore, cost-effective and efficient use of other methods such as using resistant varieties, as well as PGR such as SA appears appropriate procedures to mitigate the harmful effects of salinity (Stevens et al., 2006; Shrivastava and Kumar, 2015; Elhindi et al., 2017). In literatures, it has proven that selection of tolerant genotype and application of SA in salinity conditions have important roles in increasing tolerance of plants to salinity i.e. Manaa et al. (2014) in tomato, Gurmani et al. (2018) in cucumber, Syeed et al. (2011) in mustard and Azooz (2009) in faba bean. Kaur (2012) while evaluating response of salt susceptible and tolerant mashbean genotypes to salinity stress and foliar application of SA reported that with increasing the salinity levels, RWC, MSI, photosynthetic pigments and plant dry biomass decreased, while LPC and MDA contents increased in mashbean genotypes. However, tolerant genotypes were able to tolerate higher salinity level than sensitive genotypes. In the same study (Kaur, 2012), the ameliorative effect of SA was observed as increase in RWC, MSI, photosynthetic pigments and plant dry biomass in salt-stressed plants. SA was led to increase in RWC, MSI, photosynthetic pigments and plant dry biomass in salt-stressed plants. Also, SA treatments increased the biosynthesis of LPC under salt stress while reduced the MDA content in mashbean plants. Under salinity conditions, exogenous application of SA may have considerable role in plant defense mechanisms and impacts on physiochemical and molecular processes of plant cells which lead to amelioration of plant growth and development through diminishing the noxious consequences of salt stress (Cameron, 2000; Stevens et al., 2006; Elhindi et al., 2017). This function depends on plant species, phenology and the concentration of SA. Hence, the present research was aimed to evaluate the effects of salinity stress and foliar application of SA on some physiochemical traits of chicory genotypes in order to characterize whether foliar application of SA could mitigate the destructive effects of salt stress or not.
2.3. Irrigation treatments When sown, seeds were irrigated with tap water. Within 7–10 days after sowing the seeds germinated. Until seedling be established, plants were irrigated with three days intervals and after that irrigation was continued one or twice a week until the application of salinity treatments. Irrigation with saline water (65, and 130 mM NaCl) was performed seven days after the second spraying of the plants with SA. The plants were gradually subjected to increasing incremental concentrations of 32.5 mM NaCl every three days until final concentrations of 65 and 130 mM NaCl achieved to avoid osmotic shock. Irrigation with saline water had been applied about twice a week. Control plants were irrigated with tap water. 2.4. Laboratory measurements At 50% flowering stage, RWC according to Teulat et al. (2003), MSI based on Bajji et al. (2002), Chl and Car contents as described by Lichtenthaler and Buschmann (2001), LPC as reported by Bates et al. (1973), and MDA content as mentioned by Valentovic et al. (2006) were measured. Also, at 50% flowering stage, two plants from each replicate were harvested. After that, harvested plants were air dried and then oven dried for 48 h at 70 °C for measurement of SDW. 2.5. Statistical analysis Due to lack of significant differences between the 2-yr (2015–2017), the data of two years were averaged and were subjected to analysis of variance (ANOVA) using the GLM procedure of SAS (version 9.4, SAS Institute). Treatment means were compared using the least significant difference (LSD) tests (p < 0.05). Pearson correlation coefficients for different traits were also calculated.
2. Materials and methods 2.1. Chicory pots management
3. Results A 2-year pot experiment was considered at the Collage of Agriculture, Isfahan University of Technology, located in Isfahan, Iran during 2015–2017 under natural conditions (outdoors). The coordinates of study area is 32°40´N, 51°31´E and 1560 m above mean sea level. The experiment was a three-factor factorial arranged in a randomized complete block design with three replications. The treatments were three salinity levels of irrigation water (control, 65, and 130 mM NaCl), three levels of SA foliar application (0, 0.5, and 1 mM), and seven chicory genotypes collected from different areas in Iran (‘Ardestan’, ‘Hamadan’, ‘Shiraz’, ‘Mazandaran’, ‘Sanandaj’, ‘Kashan’ and ‘Yazd’). Chicory seeds were manually sown at 1 to 2 cm depth in the form of a clump into plastic pots (25 cm diameter and 30 cm height) in November 2015 and 2016. The plants were in the rosette conditions during autumn and winter. Early in the spring, plants were thinned at the 4–5 leaf stage and three plants were kept in each pot. Hence, given 189 experimental units (pots), totally 567 plants were remained. The soil was clay loam, with pH = 7.67 and containing 0.07% total nitrogen, 170 mg/kg available potassium, 19 mg/kg available phosphorus.
As Analysis of variance showed, genotype (G), salinity (S) and salicylic acid (SA) application had significant effects (P ≤ 0.01) on RWC, MSI, Chl a, Chl b, Car, LPC, MDA and SDW (Table 1). Salinity led to significant decrease in RWC, MSI, Chl a, Chl b, Car and SDW and significant increase in LPC and MDA compared to control condition (Table 2). As well, the decrement or increment of the measured traits in salinity level of 130 mM was more than 65 mM (Table 2). Foliar application of SA appeared that significantly (P ≤ 0.01) enhanced RWC, MSI, Chl a, Chl b, Car, LPC and SDW while significantly reduced MDA compared to control condition (Table 2). Also, the extent of increase or reduction of measured traits at level of 1 mM SA was greater than 0.5 mM (Table 2). The highest RWC, MSI, Chl a, Chl b, Car, LPC and SDW in Sanandaj and MDA in Ardestan were observed while the lowest RWC, MSI, Chl a, Chl b, Car, LPC and SDW in Ardestan and MDA in Sanandaj were noted (Table 2). As it can be seen in Tables 1 and 3, the interaction between salinity and genotype on MSI, Car, LPC, MDA and SDW implies different reactions between the genotypes in terms of these traits under various salinity levels. Salinity significantly decreased MSI, Car and SDW while significantly increased LPC and MDA rather than control condition in all genotypes (Table 3). However, the relative extent of changes in mentioned traits differed between various genotypes and different salinity levels (Table 3). In terms of MSI, Car and SDW, the greatest decreases were 23.55%, 31.23% and 30.29% in Ardestan while the smallest decreases of MSI in Mazandaran (8.19%), Car (12.50%) and
2.2. SA treatments Salicylic acid (2 hydroxybenzoic acid, molecular weight 138.1, Sigma) was dissolved in water by a magnetic stirrer and two levels of concentration including 0.5, and 1 mM were prepared individually. Foliar spray was uniformly done twice at 7 days intervals on each plant 2
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Table 1 Analysis of variance (mean squares) for relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes (G) evaluated at three levels of salinity (S) and three levels of salicylic acid (SA) in three replications (R) and two years. Trait R G S SA G×S G × SA S × SA G × S × SA Error
df 2 6 2 2 12 12 4 24 124
RWC
MSI **
5632 755.2** 1655** 603.1** 13.33ns 1.01ns 69.01** 0.156ns 16.78
Chl a **
2198 947.2** 2062** 654.3** 26.35** 3.19ns 81.52** 2.90ns 2.17
Chl b **
Car **
0.989 0.249** 0.908** 0.240** 0.001ns 0.003ns 0.018** 0.0002ns 0.003
LPC **
0.125 0.032** 0.130** 0.033** 0.0003ns 0.0005ns 0.002** 0.0001ns 0.0004
MDA **
0.017 0.012** 0.029** 0.006** 0.0002** 0.00003ns 0.0002** 0.00001ns 0.00004
101.4 6.30** 1625** 124.0** 20.49** 0.086ns 15.85** 0.003ns 0.211
SDW **
5.21** 27.88** 37.39** 5.28** 0.038** 0.006ns 0.181** 0.003ns 0.007
21.98 39.37** 26.18** 17.30** 0.54** 0.61** 1.95** 0.064** 0.017
DF, degrees of freedom. ** Significant at P ≤ 0.01. ns not significant.
higher when plants were sprayed with 1 mM SA compared to 0.5 mM SA (Table 5). Also, the amount of increase or decrease in measured traits due to the application of SA in salinity conditions was much more than control conditions (Table 5).
SDW (15.96%) in Sanandaj were observed at salinity level of 130 mM. (Table 3). Sanandaj accounted for the highest increase (231.7%) in LPC while Ardestan had the smallest increase (66.36%) in LPC (Table 3). Kashan had the highest increase (32.23%) in MDA while Sanandaj had the smallest increase (16.15%) in MDA (Table 3). Also, the amount of decline or increase in measured traits at salinity level of 130 mM was more than salinity level of 65 mM (Table 3). The interaction between genotype and foliar SA application on MDA expressed various response between the genotypes in term of this trait with different concentrations of SA application (Tables 1 and 4). Foliar application of SA significantly reduced MDA compared to control condition in all genotypes (Table 4). Ardestan had the highest decrease (21.26%) in MDA while Sanandaj had the lowest decrease (9.33%) in MDA (Table 4). Moreover, the trend of reduction of measured trait at 1 mM SA concentration was greater than the concentration of 0.5 mM SA in all genotypes (Table 4). The interaction between salinity stress and foliar SA application demonstrated in Tables 1 and 5. The data imply that the salinity stress significantly lowered the values of RWC, MSI, Chl a, Chl b, Car and SDW while significantly increased LPC and MDA contents (Table 5). The adverse effects of salinity stress on the various physiological traits and growth exacerbated with increasing salinity levels. Foliar application of SA led to increase in RWC, MSI, Chl a, Chl b, Car, LPC and SDW and decrease in MDA under both salinity and control conditions, though the values of increase and decline in measured traits were
4. Discussion Salinity stress caused significant reduction in RWC and MSI and the devaluation was intensified with increasing salinity levels in all genotypes. These results were in line with those of Khoshbakht and Asgharei (2015) and Liu et al. (2016). The foliar application of SA enhanced RWC and MSI in both saline conditions and non-saline conditions. Although the effect of SA on increasing RWC and MSI under salinity conditions was higher than control conditions. Salicylic acid treatment seems to have a share in the generation of osmolites. The production of osmolites declines the osmotic potential of the plant cells, which is effective both in maintaining intracellular water and in absorbing water from a soil solution in inappropriate environments which leads to increase in RWC. Similar results were reported by Yildirim et al. (2008); Khoshbakht and Asgharei (2015) and Elhindi et al. (2017). It seems that SA can scavenge ROS so reduce ion leakage and cell membrane injury (Gunes et al., 2007; Bastam et al., 2013; Khoshbakht and Asgharei, 2015; Liu et al., 2014). There was positive and significant correlation between RWC and MSI with SDW (Table 6) under salinity stress. This point highlights the importance of these two parameters in the
Table 2 Mean comparison for the effect of three levels of salinity (control, 65 mM and 130 mM NaCl) and salicylic acid (0, 0.5 and 1 mM) on relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes (G) in 2 years. Treatments Salinity Control 65 mM 130 mM SA 0 mM 0.5 mM 1 mM Genotype Ardestan Hamedan Mazandaran Kashan Shiraz Sanandaj Yazd
RWC (%)
MSI (%)
Chl a (mg g−1
70.63a 64.28b 60.48c
68.32a 61.44b 56.97c
0.863a 0.717b 0.625c
0.250a 0.199b 0.159c
0.204a 0.178b 0.161c
7.09c 14.84b 16.65a
4.89c 5.72b 6.16a
6.87a 5.94b 5.34c
61.69c 66.02b 67.68a
58.73c 62.94b 65.06a
0.666c 0.753b 0.786a
0.177c 0.208b 0.222a
0.170c 0.183b 0.190a
11.30c 13.27b 14.01a
6.17a 5.45b 5.15c
5.74c 6.11b 6.30a
57.99g 65.01d 70.11b 62.51e 60.24f 72.60a 67.44c
54.59g 62.49d 67.41b 58.83e 56.41f 70.61a 65.34c
0.616e 0.756c 0.828a 0.662d 0.639de 0.855a 0.790b
0.158e 0.206c 0.236b 0.180d 0.171d 0.252a 0.214c
0.153g 0.182d 0.201b 0.167e 0.160f 0.209a 0.196c
12.17e 12.75c 13.32a 12.76c 12.44d 13.55a 13.03b
7.38a 5.58d 4.35f 6.09c 6.73b 4.25g 4.75e
4.61g 6.10d 7.02b 5.56e 5.07f 7.31a 6.68c
FW)
Chl b (mg g−1
Car (mg g−1
FW)
FW)
LPC (μmol g−1
In each column, means with at least one similar letter are not significantly different according to the LSD LSD (0.05), least significant differences at P ≤ 0.05. 3
(0.05).
FW)
MDA (nmol g−1
FW)
SDW (g plant−1)
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could possibly be due to changes in stability of the pigment-protein complex or increased chlorophyllase activity (Levitt, 1980) which lead to negative effects on components of photosynthetic apparatus of chicory genotypes. Our results were in consistence with several reports that noted reduction of Chl and Car contents with increasing salinity levels (Qu et al., 2012; Rasool et al., 2013; Tabatabaei and Ehsanzadeh, 2016; Farahbakhsh et al., 2017). Salicylic acid treatments ameliorated the detrimental effects of salinity on photosynthetic pigments. Salicylic acid seems to improve the function of the photosynthetic apparatus by increasing the biosynthesis of chloroplast pigments and the activity of certain enzymes that lead to increased net photosynthesis under salinity conditions. The results obtained in this study were in concurrence with Idrees et al. (2012) in lemongrass and El-Tayeb (2005) in barley. Shoot dry weight had positive and significant correlation with Chl and Car contents (Table 6) under both control and salinity conditions. These results showed that Chl and Car contents had higher correlation with SDW under salinity stress in comparison with control condition. With increasing salinity stress, all photosynthetic pigments significantly decreased which indicated the destructive effect of salinity on photosynthetic pigments, reduction in photosynthetic capacity and thus decline in SDW production. Osmotic adjustment is found to be one of the most important mechanisms for plant adaptation to salt stress. Proline serves as an effective component in osmotic regulation in the plants that its concentration increases under salinity stress conditions (El-Tayeb, 2005; Aftab et al., 2011; Idrees et al., 2012; Pirasteh-anosheh et al., 2014; Dong et al., 2015). In this study, salt stress caused significant increment in LPC. Prolin concentration enhanced with increasing salinity levels in all genotypes. Proline declines the cytoplasmic osmotic potential that facilitates water absorption under salt stress condition. In addition to osmotic adjustment, proline has a major role in maintaining the integrity of the plasma membrane, preventing destruction of enzymes and proteins, and scavenging ROS (Hare and Cress, 1997). The foliar application of SA in plants under salt stress increased LPC, which reduced deleterious effects of salinity stress on plants. There was a positive and significant correlation between LPC and SDW (Table 6) under salinity stress. Also, positive and significant correlation between LPC with RWC and MSI was observed under salinity condition (Table 6). Significant and positive correlation between RWC and LPC was indicative of osmotic adjustment in chicory under salt stress conditions. In addition to its osmotic effect, proline by maintaining the cell membrane stability and protecting the enzymes, plays a major role in counteracting the plant cells against salt stress that caused to an increase in SDW. In the present research, MDA concentration significantly was elevated at two levels of salinity treatments rather than control condition and its increase was exacerbated by higher salinity levels in all genotypes. Our results were in agreement with the findings of Rasool et al. (2013) in chickpea, Ashrafi et al. (2015) in alfalfa and Methenni et al. (2018) in olive. Salinity adversely affects function of plasma membrane of the plant cells that causes the membrane lipid peroxidation and production of MDA. The membrane damage is associated either by generation of ROS or by the direct degradation of polyunsaturated fatty acids (Sairam et al., 2000; Darvizheh et al., 2019). The SA treatments under salinity condition prevented lipid peroxidation and so alleviated the cell membrane injury. In line with our results, Gunes et al. (2007) in maize; Liu et al. (2016) in Nitraria tangutorum and Dong et al. (2015) in cotton demonstrated that SA decreased the level of lipid peroxidation under salt stress. The concentration of MDA expresses degradation rate of the cell membrane because this compound is released by peroxidation of the cell membrane lipids. Shoot dry weight had a negative and significant correlation with MDA (Table 6) under salinity stress which indicated the negative effect of cell membrane degradation and MDA generation on plant growth and shoot dry matter production. Salinity stress significantly inhibited the plant growth, inducing a reduction in SDW of the genotypes at two levels of NaCl treatments
Table 3 Mean comparisons for interaction effects of genotype * salinity on membrane stability index (MSI), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes. Genotype
Ardestan
Hamedan
Mazandaran
Kashan
Shiraz
Sanandaj
Yazd
Salinity
MSI (%)
Car (mg g−1 FW)
LPC (μmol g−1 FW)
MDA (nmol g−1 FW)
SDW (g plant−1)
Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM
62.50h 53.50l 47.78n 69.10cd 61.54jk 56.84i 70.13bc 67.73de 64.38g 66.11f 57.79j 52.61l 63.98g 55.49k 49.78m 75.28a 69.57c 66.97ef 71.17b 64.44g 60.42i
0.1847ghi 0.148l 0.127n 0.206c 0.1788hi 0.162jk 0.219ab 0.199d 0.1848gh 0.192ef 0.164j 0.146l 0.188gf 0.156k 0.136m 0.224a 0.207c 0.196de 0.217b 0.192ef 0.1786i
8.80k 13.07j 14.64h 6.68m 14.92h 16.64d 5.96no 16.15f 17.86b 7.90l 14.18i 16.20ef 8.43k 13.48j 15.40g 5.57° 16.61de 18.48a 6.28mn 15.50g 17.31c
6.33d 7.57b 8.23a 4.87i 5.70g 6.16e 3.92n 4.46k 4.68j 5.18h 6.23de 6.85c 5.85f 6.84c 7.49b 3.90n 4.33l 4.53k 4.19m 4.88i 5.18h
5.48k 4.53° 3.82q 6.97e 5.97i 5.37l 7.80b 6.92e 6.33h 6.40h 5.45kl 4.83n 5.89j 4.98m 4.36p 8.02a 7.18d 6.74f 7.54c 6.54g 5.97ij
In each column, means with at least one similar letter are not significantly different according to the LSD (0.05). LSD (0.05), least significant differences at P ≤ 0.05. Table 4 Mean comparisons for interaction effects of genotype* salicylic acid on malondialdehyde content (MDA) of seven chicory genotypes. Genotype
Ardestan
Hamedan
Mazandaran
Kashan
Shiraz
Sanandaj
Yazd
SA
MDA (nmol g−1 FW)
Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM
8.37a 7.17c 6.59e 6.10f 5.46h 5.18i 4.68j 4.26l 4.13m 6.74d 5.96g 5.56h 7.62b 6.52e 6.05fg 4.50k 4.17lm 4.08m 5.16i 4.65j 4.45k
In each column, means with at least one similar letter are not significantly different according to the LSD (0.05). LSD (0.05), least significant differences at P ≤ 0.05.
production of shoot dry matter under salt stress conditions. One of the most important factors in the plants growth and development is photosynthesis rate. The concentration of photosynthetic pigments plays a substantial role in enhancing photosynthetic efficiency (Hopkins and Huner, 2008). In the present research, Chl a, Chl b and Car contents of the chicory genotypes significantly lessened with elevating salinity levels, which resulted in diminished photosynthetic capacity of the plants and eventually decreased growth and dry matter productions. The decline in Chl contents by increasing salinity levels 4
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Table 5 Mean comparisons for interaction effects of salinity * salicylic acid on relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes. Salinity
SA
RWC (%)
MSI (%)
Chl a (mg g−1
Control Control Control 65 mM 65 mM 65 mM 130 mM 130 mM 130 mM
0 mM 0.5 mM 1 mM 0 mM 0.5 mM 1 mM 0 mM 0.5 mM 1 mM
69.53ab 70.80a 71.55a 59.83f 65.55cd 67.46bc 55.70g 61.70ef 64.04de
67.40b 68.10b 69.47a 56.71g 62.64d 64.96c 52.07h 58.09f 60.74e
0.832b 0.868a 0.889a 0.622f 0.744d 0.786c 0.545g 0.648ef 0.682e
FW)
Chl b (mg g−1
Car (mg g−1
FW)
0.238b 0.251a 0.259a 0.163e 0.207c 0.228b 0.130f 0.167de 0.179d
LPC (μmol g−1
FW)
0.197c 0.206b 0.210a 0.165g 0.180e 0.188d 0.148h 0.163g 0.172f
FW)
6.62h 7.22g 7.42g 12.92f 15.36d 16.25c 14.36e 17.22b 18.36a
In each column, means with at least one similar letter are not significantly different according to the LSD LSD (0.05), least significant differences at P ≤ 0.05.
MDA (nmol g−1
FW)
5.12g 4.84h 4.71i 6.33b 5.55d 5.27f 7.06a 5.97c 5.46e
SDW (g plant−1) 6.67c 6.92b 7.03a 5.56g 6.02e 6.23d 4.98i 5.39h 5.66f
(0.05).
Table 6 Correlation coefficients among relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes under salinity stress (lower triangle, bold type) and control conditions (upper triangle). Traits RWC MSI Chl a Chl b Car LPC MDA SDW
RWC 1 0.93** 0.694** 0.691** 0.89** 0.90** −0.35** 0.46**
MSI
Chl a **
0.90 1 0.772** 0.773** 0.96** 0.94** −0.58** 0.67**
Chl b **
0.561 0.631** 1 0.99** 0.729** 0.752** −0.512** 0.574**
Car **
0.566 0.632** 0.99** 1 0.72** 0.75** −0.515** 0.579**
LPC **
0.85 0.92** 0.655** 0.650** 1 0.88** −0.65** 0.74**
MDA ns
0.08 −0.14ns −0.33** −0.31* −0.24* 1 −0.51** 0.55**
ns
0.05 −0.18ns −0.37** −0.37** −0.33** 0.87** 1 −0.95**
SDW 0.09ns 0.33** 0.463** 0.45** 0.464** −0.88** −0.97** 1
** Significant at P ≤ 0.01. * Significant at P ≤ 0.05. ns Not significant.
Acknowledgment
compared to non-salt-treated plants. Salt stress adversely restricts plant growth and development through its influence on the photosynthesis, antioxidant phenomena, proline metabolism, osmolyte accumulation and other physiochemical process of plant cells (Aftab et al., 2010; Idrees et al., 2012). Application of SA alleviated the destructive effects of salinity that led to increase in growth parameters and total biomass. Salicylic acid has a key impress in regulation of plant growth and development under stress conditions that causes the function of the photosynthetic apparatus to improve and increase plant biomass (Senaratna et al., 2000). The results of this research were in accordance with the reports of El-Tayeb (2005); Idrees et al. (2012) and Pirastehanosheh et al. (2014). Based on measured traits under salinity condition, SDW had the highest correlation (Table 6) with MDA (r=-0.95**) followed by Car (r = 0.74**) and MSI (r = 0.67**), respectively.
We sincerely thank to the anonymous reviewers for the helpful and constructive comments on the earlier version of our manuscript. We would also like to thank for Isfahan University of Technology for financing this research. References Aftab, T., Khan, M.M.A., Idrees, M., Naeem, M., Moinuddin, A.S., 2010. Salicylic acid acts as potent enhancer of growth, photosynthesis and artemisinin production in Artemisia annua L. J. Crop Sci. Biotech. 13, 183–188. Aftab, T., Khan, M.M.A., Silva, J.A.T., Idrees, M., Naeem, M., Moinuddin, A.S., 2011. Role of salicylic acid in promoting salt stress tolerance and enhanced artemisinin production in Artemisia annua L. J. Plant Growth Regul. 30, 425–435. Amaducci, S., Pritoni, G., 1998. Effect of harvest date and cultivar on Cichorium intybus yield components in north Italy. Ind. Crop. Prod. 7, 345–349. Ashraf, M., Athar, H.R., Harris, P.J.C., Kwon, T.R., 2008. Some prospective strategies for improving crop salt tolerance. Adv. Agron. 97, 45–110. Ashrafi, E., Razmjoo, J., Zahedi, M., Pessarakli, M., 2015. Screening alfalfa for salt tolerance based on lipid peroxidation and antioxidant enzymes. Agron. J. 107, 1–7. Azooz, M.M., 2009. Salt stress mitigation by seed priming with salicylic acid in two faba bean genotypes differing in salt tolerance. Int. J. Agric. Biol. 11, 343–350. Baert, J.R.A., Bockstaele, E.J.V., 1992. Cultivation and breeding of root chicory for inulin production. Ind. Crop. Prod. 1, 229–234. Bajji, M., Kinet, J.M., Lutts, S., 2002. The use of the electrolyte leakage method for assessing cell membrane stability as a water stress tolerance test in durum wheat. Plant Growth Regul. 36, 61–70. Bastam, N., Baninasab, B., Ghobadi, C., 2013. Improving salt tolerance by exogenous application of salicylic acid in seedlings of pistachio. Plant Growth Regul. 69, 275–284. Bates, L.S., Waldran, R.P., Teare, I.D., 1973. Rapid determination of free proline for water studies. Plant Soil 39, 205–208. Belesky, D.P., Fedders, J.M., Turner, K.E., Ruckle, J.M., 1999. Productivity, botanical composition, and nutritive value of swards including forage chicory. Agron. J. 91, 450–456. Boyd, D.C., Rogers, M.E., 2004. Effect of salinity on the growth of chicory (Cichorium intybus cv. Puna) - A potential dairy forage species for irrigation areas. Aust. J. Exp. Agric. 44, 189–192.
5. Conclusion In light of above discussion, it can be concluded that salt stress led to significant decline in RWC, MSI, Chl a, Chl b, Car and SDW and significant increase in LPC and MDA compared to control condition. Also, increasing salinity levels, intensified extent of reduction or enhancement of the measured traits. Foliar application of SA significantly ameliorated all measured traits and the greatest improvements were obtained in the highest concentration of SA. Among genotypes, Sanandaj had the highest RWC, MSI, Chl a, Chl b, Car, LPC and the lowest content of MDA that caused to the highest SDW in comparison with other genotypes while Ardestan had the lowest RWC, MSI, Chl a, Chl b, Car, LPC and the highest content of MDA that caused to the lowest SDW. Drawing upon such findings, foliar application of SA can play a role in diminishing deleterious effects of salinity stress and reinforcing chicory plant tolerance to salt stress. 5
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431–438. Liu, S., Dong, Y., Xu, L., 2014. Effects of foliar applications of nitric oxide and salicylic acid on salt-induced changes in photosynthesis and antioxidative metabolism of cotton seedlings. Plant Growth Regul. 73, 67–78. Liu, W., Zhang, Y., Yuan, X., Xuan, Y., Gao, Y., Yan, Y., 2016. Exogenous salicylic acid improves salinity tolerance of nitraria tangutorum. Russ. J. Plant Physiol. 63, 132–142. Manaa, A., Gharbi, E., Mimouni, H., Wasti, S., Aschi-Smiti, S., Lutts, S., Ben Ahmed, H., 2014. Simultaneous application of salicylic acid and calcium improves salt tolerance in two contrasting tomato (Solanum lycopersicum) cultivars. S. Afr. J. Bot. 95, 32–39. Methenni, K., Abdallah, M.B., Nouairi, I., Smaoui, A., Ammar, W.B., Zarrouk, M., Youssef, N.B., 2018. Salicylic acid and calcium pretreatments alleviate the toxic effect of salinity in the Oueslati olive variety. Sci. Hortic. 233, 349–358. Munns, R., 2002. Comparative physiology of salt and water stress. Plant Cell Envir. 25, 239–250. Pirasteh-anosheh, H., Ranjbar, Gh., Emam, Y., Ashraf, M., 2014. Salicylic acid induced recovery ability in salt-stressed Hordeum vulgare plants. Turk. J. Bot. 38, 112–121. Qu, C., Liu, C., Gong, X., Li, C., Hong, M., Wang, L., Hong, F., 2012. Impairment of maize seedling photosynthesis d by a combination of potassium deficiency and salt stress. Environ. Exp. Bot. 75, 134–141. Rasool, S., Ahmad, A., Siddiqi, T.O., Ahmad, P., 2013. Changes in growth, lipid peroxidation and some key antioxidant enzymes in chickpea genotypes under salt stress. Acta Physiol. Plant. 35, 1039–1050. Sairam, R.K., Srivastava, G.C., Saxena, D.C., 2000. Increased antioxidant activity under elevated temperatures: a mechanism of heat stress tolerance in wheat genotypes. Biol. Plantarum. 43, 245–251. Senaratna, T., Touchell, D., Bunn, E., Dixon, K., 2000. Acetyl salicylic acid (aspirin) and salicylic acid induce multiple stress tolerance in bean and tomato plants. Plant Growth Regul. 30, 157–161. Shrivastava, P., Kumar, R., 2015. Soil salinity: a serious environmental issue and plant growth promoting bacteria as one of the tools for its alleviation. Saudi J. Biol. Sci. 22, 123–131. Silveira, J.A.G., Melo, A.R.B., Viégas, R.A., Oliveira, J.T.A., 2001. Salinity-induced effects on nitrogen assimilation related to growth in cowpea plants. Environ. Exp. Bot. 46, 171–179. Stevens, J., Senaratna, T., Sivasithamparam, K., 2006. Salicylic acid induces salinity tolerance in tomato (Lycopersicon esculentum cv. Roma): associated changes in gas exchange, water relations and membrane stabilization. Plant Growth Regul. 49, 77–83. Syeed, Sh., Anjum, N.A., Nazar, R., Iqbal, N., Masood, A., Khan, N.A., 2011. Salicylic acidmediated changes in photosynthesis, nutrients content and antioxidant metabolism in two mustard (Brassica juncea L.) cultivars differing in salt tolerance. Acta Physiol. Plant. 33, 877–886. Tabatabaei, S., Ehsanzadeh, P., 2016. Photosynthetic pigments, ionic and antioxidative behaviour of hulled tetraploid wheat in response to NaCl. Photosynthetica. 54, 340–350. Tahjib‑Ul‑Arif, M., Siddiqui, M.N., Sohag, A.A.M., Sakil, M.A., Rahman, M.M., Polash, M.A.S., Mostofa, M.G., Tran, L.S.P., 2018. Salicylic acid-mediated enhancement of photosynthesis attributes and antioxidant capacity contributes to yield improvement of maize plants under salt stress. J. Plant Growth Regul. 37, 1318–1330. Teulat, B., Zoumarou-Wallis, N., Rotter, B., Salem, M.B., Bahri, H., This, D., 2003. QTL for relative water content in field-grown barley and their stability across Mediterranean environments. Theor. Appl. Genet. 108, 181–188. Valentovic, P., Luxova, M., Kolarovi, L., Gasparikora, O., 2006. Effect of osmotic stress on compatible solutes content, memberane stability and water relation in two maize. Plant Soil Environ. 52, 186–191. Yildirim, E., Turan, M., Guvenc, I., 2008. Effect of foliar salicylic acid applications on growth, chlorophyll, and mineral content of cucumber grown under salt stress. J. Plant Nutr. 31, 593–612.
Cameron, R.K., 2000. Salicylic acid and its role in plant defense responses: what do we really know? Physiol. Mol. Plant Pathol. 56, 91–93. Chen, W.P., Li, P.H., Chen, T.H.H., 2000. Glycinebetaine increases chilling tolerance and reduces chilling-induced lipid peroxidation in Zea mays L. Plant Cell Environ. 23, 609–618. Crush, J.R., Evans, J.P.M., 1990. Shoot growth and herbage element concentrations of ‘Grasslands Puna’ chicory (Cichorium intybus L.) under varying soil pH. Proceedings of the New Zealand Grassland Association 51, 163–166. Dalara, A., Konczaka, I., 2014. Cichorium intybus from eastern Anatolia: phenolic composition, antioxidant and enzyme inhibitory activities. Ind. Crop. Prod. 60, 79–85. Darvizheh, H., Zahedi, M., Abbaszadeh, B., Razmjoo, J., 2019. Changes in some antioxidant enzymes and physiological indices of purple coneflower (Echinacea purpurea L.) in response to water deficit and foliar application of salicylic acid and spermine under field condition. Sci. Hortic. 247, 390–399. Dong, Y.J., Wang, Z.L., Zhang, J.W., Liu, S., He, Z.L., He, M.R., 2015. Interaction effects of nitric oxide and salicylic acid in alleviating salt stress of Gossypium hirsutum L. J. Soil Sci. Plant Nut. 15, 561–573. Elhindi, K.M., Al-Amri, S.M., Abdel-Salam, E.M., Al-Suhaibani, N.A., 2017. Effectiveness of salicylic acid in mitigating salt-induced adverse effects on different physio-biochemical attributes in sweet basil (Ocimum basilicum L.). J. Plant Nutr. 6, 908–919. El-Tayeb, M.A., 2005. Response of barley grains to the interactive effect of salinity and salicylic acid. Plant Growth Regul. 45, 215–224. Farahbakhsh, H., Pasandi Pour, A., Reiahi, N., 2017. Physiological response of henna (Lawsonia inermis L.) to salicylic acid and salinity. Plant Prod. Sci. 20, 237–247. González-Acevedo, Z.I., Padilla-Reyes, D.A., Ramos-Leal, J.A., 2016. Quality assessment of irrigation water related to soil salinization in. Tierra Nueva, San Luis Potosí, Mexico. Rev. Mex. Clenc. Geol. 3, 271–285. Gunes, A., Inal, A., Alpaslan, M., Eraslan, F., Bagci, E.G., 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. J. Plant Physiol. 164, 728–736. Gurmani, A.R., Ullah Khan, S., Ali, A., Rubab, T., Schwinghamer, T., Jilani, Gh., Farid, A., Zhang, J., 2018. Salicylic acid and kinetin mediated stimulation of salt tolerance in cucumber (Cucumis sativus L.) genotypes varying in salinity tolerance. Hortic. Environ. Biote. 59, 461–471. Hare, P.D., Cress, W.A., 1997. Metabolic implications of stress-induced proline accumulation in plants. Plant Growth Regul. 21, 79–102. Hopkins, W.G., Huner, N.P.A., 2008. Introduction to Plant Physiology, fourth edition. The University of Western Ontario. Idrees, M., Naeem, M., Khan, M.N., Aftab, T., Masroor, M., Khan, A., Moinuddin, A.S., 2012. Alleviation of salt stress in lemongrass by salicylic acid. Protoplasma. 249, 709–720. Kalber, T., Kreuzer, M., Leiber, F., 2012. Silages containing buckwheat and chicory: quality, digestibility and nitrogen utilization by lactating cows. Arch. Anim. Nutr. 66, 50–65. Kaur, M., 2012. Salicylic Acid Induced Changes in Some Physiological and Biochemical Parameters in Mashbean (Vigna Mungo L. Hepper) Genotypes Grown Under Salinity. MSc Thesis. Department of Botany. College of Basic Sciences and Humanities. Punjab Agricultural University, Ludhiana. Khoshbakht, D., Asgharei, M.R., 2015. Influence of foliar-applied salicylic acid on growth, gas-exchange characteristics, and chlorophyll fluorescence in citrus under saline conditions. Photosynthetica. 53, 1–10. Koca, H., Bor, M., Ozdemir, F., Turkan, I., 2007. The effect of salt stress on lipid peroxidation, antioxidative enzymes and proline content of sesame cultivars. Environ. Exp. Bot. 60, 344–351. Levitt, J., 1980. Responses of Plant to Environmental Stress: Water, Radiation, Salt and Other Stresses. Academic Press, New York 365. Lichtenthaler, H.K., Buschmann, C., 2001. Chlorophylls and carotenoids: measurement and characterization by UV–vis spectroscopy. Curr. Protoc. Food Anal. Chem. 1,
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Interactive effect of salinity stress and foliar application of salicylic acid on some physiochemical traits of chicory (Cichorium intybus L.) genotypes Nazanin Poursakhi, Jamshid Razmjoo, Hassan Karimmojeni
T
⁎
Department of Agronomy and Plant Breeding, College of Agriculture, Isfahan University of Technology, Isfahan, Iran
A R T I C LE I N FO
A B S T R A C T
Keywords: Chicory Abiotic stress Plant growth regulator Oxidative stress Photosynthetic pigments Growth
Chicory (Cichorium intybus) is an important medicinal, food and feed plant. Salinity can affect its growth and quality to some extent. However, adverse effects of salinity may be alleviated upon selecting salt tolerant genotypes and application of salicylic acid. Therefore, a 2-year pot experiment was conducted with three salinity levels (control, 65 mM and 130 mM NaCl), three levels of foliar application of salicylic acid (0, 0.5 mM and 1 mM SA) and seven chicory genotypes (Ardestan, Hamedan, Shiraz, Mazandaran, Sanandaj, Kashan and Yazd). Treatment characterized with 65 and 130 mM salinity levels led to significant decrease in relative water content (RWC), membrane stability index (MSI), chlorophyll a(Chl a), chlorophyll b (Chl b) and carotenoid (Car) contents and shoot dry weight (SDW) while significant increase in leaf proline (LPC) and malondialdehyde (MDA) contents. Moreover, the highest changes in measured traits were observed at salinity level of 130 mM. Salicylic acid application at levels of 0.5 and 1 mM SA significantly ameliorated all measured traits and the greatest values were attributed to 1 mM SA. Among genotypes, Sanandaj accounted for the highest measured traits but the lowest content of MDA while Ardestan characterized with the lowest measured traits and the highest content of MDA. Salinity decreased MSI, Car and SDW while it increased LPC and MDA in all genotypes with different magnitude. Application of SA reduced MDA in all genotypes and also led to increase in RWC, MSI, Chl a, Chl b, Car, LPC and SDW and diminution in MDA under control and specially under salt stress conditions. Sanandaj with the application of 1 mM SA under control and saline conditions was the best genotype among all genotypes. The results suggested that SA may improve performance of chicory genotypes under control and saline conditions by announcing physiochemical traits.
1. Introduction Soil salinity is one of the most outstanding limiting factors for crop production, especially in arid and semi-arid parts of the world (Silveira et al., 2001; Munns, 2002). In these areas, salinity may be arised due to soil intrinsic properties, limited precipitation and poor drainage conditions, inappropriate quality of irrigation water and intense evaporation during long hot and dry seasons (González-Acevedo et al., 2016). At the same time, improper farming methods and unsuitable management of irrigation systems have exacerbated this issue seriously. (Ashraf et al., 2008). Chicory is cultivated as a forage crop in different parts of the world (Belesky et al., 1999; Kalber et al., 2012). In addition, it is utilized for its leaves, buds and roots among others (Amaducci and Pritoni, 1998). It serves as a valuable source of calcium, magnesium,
potassium, manganese, zinc, and iron (Crush and Evans, 1990). The young leaves of chicory are edible and can be excellent addition to salads (Dalara and Konczaka, 2014). The processed roots of chicory can be consumed as a coffee substitution (Baert and Bockstaele, 1992). Boyd and Rogers (2004) pointed out that chicory has a degree of salinity resistance and this makes it suitable to be cultivated in relatively moderate salinity areas. Under salinity circumstance, reducing water availability because of low osmotic potential of soil solution, creation of ion toxicity effect, disintegration of membranes, nutritional imbalance and excessive production and accumulation of ROS (Chen et al., 2000) are factors causing major changes at physiological, biochemical, and molecular processes of plant cells (Munns, 2002). These are associated with reduction in RWC, MSI, photosynthetic pigments such as Chl a, Chl b, Car and shoot dry matter and increase in LPC and MDA contents that
Abbreviations: G, Genotype; S, Salinity; SA, Salicylic acid; RWC, Relative water content; MSI, Membrane stability index; Chla, Chlorophyll a; Chlb, Chlorophyll b; Car, Carotenoid; LPC, Leaf prolin content; MDA, Malondialdehyde; SDW, Shoot dry weight; PGR, Plant growth regulator; ROS, Reactive oxygen species; FW, Fresh weight ⁎ Corresponding author. E-mail address: [email protected] (H. Karimmojeni). https://doi.org/10.1016/j.scienta.2019.108810 Received 22 March 2019; Received in revised form 18 August 2019; Accepted 25 August 2019 0304-4238/ © 2019 Elsevier B.V. All rights reserved.
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with a constant volume 7 mL plant−1 of SA solution with a manual sprayer at 6–7 leaf growth stage. Control plants were sprayed similarly with equivalent amount of water. (Methenni et al., 2018; Tahjib-Ul-Arif et al., 2018). The pot soil was covered by a plastic sheet during the SA spray in order to avoid the access of SA solution to the root system.
ultimately lead to adverse impacts on plant growth and development (El-Tayeb, 2005; Koca et al., 2007; Rasool et al., 2013; Farahbakhsh et al., 2017). The main practices to deal with salinity are soil restoration and leaching operations. These methods are not cost-effective and sometimes are impossible. Therefore, cost-effective and efficient use of other methods such as using resistant varieties, as well as PGR such as SA appears appropriate procedures to mitigate the harmful effects of salinity (Stevens et al., 2006; Shrivastava and Kumar, 2015; Elhindi et al., 2017). In literatures, it has proven that selection of tolerant genotype and application of SA in salinity conditions have important roles in increasing tolerance of plants to salinity i.e. Manaa et al. (2014) in tomato, Gurmani et al. (2018) in cucumber, Syeed et al. (2011) in mustard and Azooz (2009) in faba bean. Kaur (2012) while evaluating response of salt susceptible and tolerant mashbean genotypes to salinity stress and foliar application of SA reported that with increasing the salinity levels, RWC, MSI, photosynthetic pigments and plant dry biomass decreased, while LPC and MDA contents increased in mashbean genotypes. However, tolerant genotypes were able to tolerate higher salinity level than sensitive genotypes. In the same study (Kaur, 2012), the ameliorative effect of SA was observed as increase in RWC, MSI, photosynthetic pigments and plant dry biomass in salt-stressed plants. SA was led to increase in RWC, MSI, photosynthetic pigments and plant dry biomass in salt-stressed plants. Also, SA treatments increased the biosynthesis of LPC under salt stress while reduced the MDA content in mashbean plants. Under salinity conditions, exogenous application of SA may have considerable role in plant defense mechanisms and impacts on physiochemical and molecular processes of plant cells which lead to amelioration of plant growth and development through diminishing the noxious consequences of salt stress (Cameron, 2000; Stevens et al., 2006; Elhindi et al., 2017). This function depends on plant species, phenology and the concentration of SA. Hence, the present research was aimed to evaluate the effects of salinity stress and foliar application of SA on some physiochemical traits of chicory genotypes in order to characterize whether foliar application of SA could mitigate the destructive effects of salt stress or not.
2.3. Irrigation treatments When sown, seeds were irrigated with tap water. Within 7–10 days after sowing the seeds germinated. Until seedling be established, plants were irrigated with three days intervals and after that irrigation was continued one or twice a week until the application of salinity treatments. Irrigation with saline water (65, and 130 mM NaCl) was performed seven days after the second spraying of the plants with SA. The plants were gradually subjected to increasing incremental concentrations of 32.5 mM NaCl every three days until final concentrations of 65 and 130 mM NaCl achieved to avoid osmotic shock. Irrigation with saline water had been applied about twice a week. Control plants were irrigated with tap water. 2.4. Laboratory measurements At 50% flowering stage, RWC according to Teulat et al. (2003), MSI based on Bajji et al. (2002), Chl and Car contents as described by Lichtenthaler and Buschmann (2001), LPC as reported by Bates et al. (1973), and MDA content as mentioned by Valentovic et al. (2006) were measured. Also, at 50% flowering stage, two plants from each replicate were harvested. After that, harvested plants were air dried and then oven dried for 48 h at 70 °C for measurement of SDW. 2.5. Statistical analysis Due to lack of significant differences between the 2-yr (2015–2017), the data of two years were averaged and were subjected to analysis of variance (ANOVA) using the GLM procedure of SAS (version 9.4, SAS Institute). Treatment means were compared using the least significant difference (LSD) tests (p < 0.05). Pearson correlation coefficients for different traits were also calculated.
2. Materials and methods 2.1. Chicory pots management
3. Results A 2-year pot experiment was considered at the Collage of Agriculture, Isfahan University of Technology, located in Isfahan, Iran during 2015–2017 under natural conditions (outdoors). The coordinates of study area is 32°40´N, 51°31´E and 1560 m above mean sea level. The experiment was a three-factor factorial arranged in a randomized complete block design with three replications. The treatments were three salinity levels of irrigation water (control, 65, and 130 mM NaCl), three levels of SA foliar application (0, 0.5, and 1 mM), and seven chicory genotypes collected from different areas in Iran (‘Ardestan’, ‘Hamadan’, ‘Shiraz’, ‘Mazandaran’, ‘Sanandaj’, ‘Kashan’ and ‘Yazd’). Chicory seeds were manually sown at 1 to 2 cm depth in the form of a clump into plastic pots (25 cm diameter and 30 cm height) in November 2015 and 2016. The plants were in the rosette conditions during autumn and winter. Early in the spring, plants were thinned at the 4–5 leaf stage and three plants were kept in each pot. Hence, given 189 experimental units (pots), totally 567 plants were remained. The soil was clay loam, with pH = 7.67 and containing 0.07% total nitrogen, 170 mg/kg available potassium, 19 mg/kg available phosphorus.
As Analysis of variance showed, genotype (G), salinity (S) and salicylic acid (SA) application had significant effects (P ≤ 0.01) on RWC, MSI, Chl a, Chl b, Car, LPC, MDA and SDW (Table 1). Salinity led to significant decrease in RWC, MSI, Chl a, Chl b, Car and SDW and significant increase in LPC and MDA compared to control condition (Table 2). As well, the decrement or increment of the measured traits in salinity level of 130 mM was more than 65 mM (Table 2). Foliar application of SA appeared that significantly (P ≤ 0.01) enhanced RWC, MSI, Chl a, Chl b, Car, LPC and SDW while significantly reduced MDA compared to control condition (Table 2). Also, the extent of increase or reduction of measured traits at level of 1 mM SA was greater than 0.5 mM (Table 2). The highest RWC, MSI, Chl a, Chl b, Car, LPC and SDW in Sanandaj and MDA in Ardestan were observed while the lowest RWC, MSI, Chl a, Chl b, Car, LPC and SDW in Ardestan and MDA in Sanandaj were noted (Table 2). As it can be seen in Tables 1 and 3, the interaction between salinity and genotype on MSI, Car, LPC, MDA and SDW implies different reactions between the genotypes in terms of these traits under various salinity levels. Salinity significantly decreased MSI, Car and SDW while significantly increased LPC and MDA rather than control condition in all genotypes (Table 3). However, the relative extent of changes in mentioned traits differed between various genotypes and different salinity levels (Table 3). In terms of MSI, Car and SDW, the greatest decreases were 23.55%, 31.23% and 30.29% in Ardestan while the smallest decreases of MSI in Mazandaran (8.19%), Car (12.50%) and
2.2. SA treatments Salicylic acid (2 hydroxybenzoic acid, molecular weight 138.1, Sigma) was dissolved in water by a magnetic stirrer and two levels of concentration including 0.5, and 1 mM were prepared individually. Foliar spray was uniformly done twice at 7 days intervals on each plant 2
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Table 1 Analysis of variance (mean squares) for relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes (G) evaluated at three levels of salinity (S) and three levels of salicylic acid (SA) in three replications (R) and two years. Trait R G S SA G×S G × SA S × SA G × S × SA Error
df 2 6 2 2 12 12 4 24 124
RWC
MSI **
5632 755.2** 1655** 603.1** 13.33ns 1.01ns 69.01** 0.156ns 16.78
Chl a **
2198 947.2** 2062** 654.3** 26.35** 3.19ns 81.52** 2.90ns 2.17
Chl b **
Car **
0.989 0.249** 0.908** 0.240** 0.001ns 0.003ns 0.018** 0.0002ns 0.003
LPC **
0.125 0.032** 0.130** 0.033** 0.0003ns 0.0005ns 0.002** 0.0001ns 0.0004
MDA **
0.017 0.012** 0.029** 0.006** 0.0002** 0.00003ns 0.0002** 0.00001ns 0.00004
101.4 6.30** 1625** 124.0** 20.49** 0.086ns 15.85** 0.003ns 0.211
SDW **
5.21** 27.88** 37.39** 5.28** 0.038** 0.006ns 0.181** 0.003ns 0.007
21.98 39.37** 26.18** 17.30** 0.54** 0.61** 1.95** 0.064** 0.017
DF, degrees of freedom. ** Significant at P ≤ 0.01. ns not significant.
higher when plants were sprayed with 1 mM SA compared to 0.5 mM SA (Table 5). Also, the amount of increase or decrease in measured traits due to the application of SA in salinity conditions was much more than control conditions (Table 5).
SDW (15.96%) in Sanandaj were observed at salinity level of 130 mM. (Table 3). Sanandaj accounted for the highest increase (231.7%) in LPC while Ardestan had the smallest increase (66.36%) in LPC (Table 3). Kashan had the highest increase (32.23%) in MDA while Sanandaj had the smallest increase (16.15%) in MDA (Table 3). Also, the amount of decline or increase in measured traits at salinity level of 130 mM was more than salinity level of 65 mM (Table 3). The interaction between genotype and foliar SA application on MDA expressed various response between the genotypes in term of this trait with different concentrations of SA application (Tables 1 and 4). Foliar application of SA significantly reduced MDA compared to control condition in all genotypes (Table 4). Ardestan had the highest decrease (21.26%) in MDA while Sanandaj had the lowest decrease (9.33%) in MDA (Table 4). Moreover, the trend of reduction of measured trait at 1 mM SA concentration was greater than the concentration of 0.5 mM SA in all genotypes (Table 4). The interaction between salinity stress and foliar SA application demonstrated in Tables 1 and 5. The data imply that the salinity stress significantly lowered the values of RWC, MSI, Chl a, Chl b, Car and SDW while significantly increased LPC and MDA contents (Table 5). The adverse effects of salinity stress on the various physiological traits and growth exacerbated with increasing salinity levels. Foliar application of SA led to increase in RWC, MSI, Chl a, Chl b, Car, LPC and SDW and decrease in MDA under both salinity and control conditions, though the values of increase and decline in measured traits were
4. Discussion Salinity stress caused significant reduction in RWC and MSI and the devaluation was intensified with increasing salinity levels in all genotypes. These results were in line with those of Khoshbakht and Asgharei (2015) and Liu et al. (2016). The foliar application of SA enhanced RWC and MSI in both saline conditions and non-saline conditions. Although the effect of SA on increasing RWC and MSI under salinity conditions was higher than control conditions. Salicylic acid treatment seems to have a share in the generation of osmolites. The production of osmolites declines the osmotic potential of the plant cells, which is effective both in maintaining intracellular water and in absorbing water from a soil solution in inappropriate environments which leads to increase in RWC. Similar results were reported by Yildirim et al. (2008); Khoshbakht and Asgharei (2015) and Elhindi et al. (2017). It seems that SA can scavenge ROS so reduce ion leakage and cell membrane injury (Gunes et al., 2007; Bastam et al., 2013; Khoshbakht and Asgharei, 2015; Liu et al., 2014). There was positive and significant correlation between RWC and MSI with SDW (Table 6) under salinity stress. This point highlights the importance of these two parameters in the
Table 2 Mean comparison for the effect of three levels of salinity (control, 65 mM and 130 mM NaCl) and salicylic acid (0, 0.5 and 1 mM) on relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes (G) in 2 years. Treatments Salinity Control 65 mM 130 mM SA 0 mM 0.5 mM 1 mM Genotype Ardestan Hamedan Mazandaran Kashan Shiraz Sanandaj Yazd
RWC (%)
MSI (%)
Chl a (mg g−1
70.63a 64.28b 60.48c
68.32a 61.44b 56.97c
0.863a 0.717b 0.625c
0.250a 0.199b 0.159c
0.204a 0.178b 0.161c
7.09c 14.84b 16.65a
4.89c 5.72b 6.16a
6.87a 5.94b 5.34c
61.69c 66.02b 67.68a
58.73c 62.94b 65.06a
0.666c 0.753b 0.786a
0.177c 0.208b 0.222a
0.170c 0.183b 0.190a
11.30c 13.27b 14.01a
6.17a 5.45b 5.15c
5.74c 6.11b 6.30a
57.99g 65.01d 70.11b 62.51e 60.24f 72.60a 67.44c
54.59g 62.49d 67.41b 58.83e 56.41f 70.61a 65.34c
0.616e 0.756c 0.828a 0.662d 0.639de 0.855a 0.790b
0.158e 0.206c 0.236b 0.180d 0.171d 0.252a 0.214c
0.153g 0.182d 0.201b 0.167e 0.160f 0.209a 0.196c
12.17e 12.75c 13.32a 12.76c 12.44d 13.55a 13.03b
7.38a 5.58d 4.35f 6.09c 6.73b 4.25g 4.75e
4.61g 6.10d 7.02b 5.56e 5.07f 7.31a 6.68c
FW)
Chl b (mg g−1
Car (mg g−1
FW)
FW)
LPC (μmol g−1
In each column, means with at least one similar letter are not significantly different according to the LSD LSD (0.05), least significant differences at P ≤ 0.05. 3
(0.05).
FW)
MDA (nmol g−1
FW)
SDW (g plant−1)
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could possibly be due to changes in stability of the pigment-protein complex or increased chlorophyllase activity (Levitt, 1980) which lead to negative effects on components of photosynthetic apparatus of chicory genotypes. Our results were in consistence with several reports that noted reduction of Chl and Car contents with increasing salinity levels (Qu et al., 2012; Rasool et al., 2013; Tabatabaei and Ehsanzadeh, 2016; Farahbakhsh et al., 2017). Salicylic acid treatments ameliorated the detrimental effects of salinity on photosynthetic pigments. Salicylic acid seems to improve the function of the photosynthetic apparatus by increasing the biosynthesis of chloroplast pigments and the activity of certain enzymes that lead to increased net photosynthesis under salinity conditions. The results obtained in this study were in concurrence with Idrees et al. (2012) in lemongrass and El-Tayeb (2005) in barley. Shoot dry weight had positive and significant correlation with Chl and Car contents (Table 6) under both control and salinity conditions. These results showed that Chl and Car contents had higher correlation with SDW under salinity stress in comparison with control condition. With increasing salinity stress, all photosynthetic pigments significantly decreased which indicated the destructive effect of salinity on photosynthetic pigments, reduction in photosynthetic capacity and thus decline in SDW production. Osmotic adjustment is found to be one of the most important mechanisms for plant adaptation to salt stress. Proline serves as an effective component in osmotic regulation in the plants that its concentration increases under salinity stress conditions (El-Tayeb, 2005; Aftab et al., 2011; Idrees et al., 2012; Pirasteh-anosheh et al., 2014; Dong et al., 2015). In this study, salt stress caused significant increment in LPC. Prolin concentration enhanced with increasing salinity levels in all genotypes. Proline declines the cytoplasmic osmotic potential that facilitates water absorption under salt stress condition. In addition to osmotic adjustment, proline has a major role in maintaining the integrity of the plasma membrane, preventing destruction of enzymes and proteins, and scavenging ROS (Hare and Cress, 1997). The foliar application of SA in plants under salt stress increased LPC, which reduced deleterious effects of salinity stress on plants. There was a positive and significant correlation between LPC and SDW (Table 6) under salinity stress. Also, positive and significant correlation between LPC with RWC and MSI was observed under salinity condition (Table 6). Significant and positive correlation between RWC and LPC was indicative of osmotic adjustment in chicory under salt stress conditions. In addition to its osmotic effect, proline by maintaining the cell membrane stability and protecting the enzymes, plays a major role in counteracting the plant cells against salt stress that caused to an increase in SDW. In the present research, MDA concentration significantly was elevated at two levels of salinity treatments rather than control condition and its increase was exacerbated by higher salinity levels in all genotypes. Our results were in agreement with the findings of Rasool et al. (2013) in chickpea, Ashrafi et al. (2015) in alfalfa and Methenni et al. (2018) in olive. Salinity adversely affects function of plasma membrane of the plant cells that causes the membrane lipid peroxidation and production of MDA. The membrane damage is associated either by generation of ROS or by the direct degradation of polyunsaturated fatty acids (Sairam et al., 2000; Darvizheh et al., 2019). The SA treatments under salinity condition prevented lipid peroxidation and so alleviated the cell membrane injury. In line with our results, Gunes et al. (2007) in maize; Liu et al. (2016) in Nitraria tangutorum and Dong et al. (2015) in cotton demonstrated that SA decreased the level of lipid peroxidation under salt stress. The concentration of MDA expresses degradation rate of the cell membrane because this compound is released by peroxidation of the cell membrane lipids. Shoot dry weight had a negative and significant correlation with MDA (Table 6) under salinity stress which indicated the negative effect of cell membrane degradation and MDA generation on plant growth and shoot dry matter production. Salinity stress significantly inhibited the plant growth, inducing a reduction in SDW of the genotypes at two levels of NaCl treatments
Table 3 Mean comparisons for interaction effects of genotype * salinity on membrane stability index (MSI), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes. Genotype
Ardestan
Hamedan
Mazandaran
Kashan
Shiraz
Sanandaj
Yazd
Salinity
MSI (%)
Car (mg g−1 FW)
LPC (μmol g−1 FW)
MDA (nmol g−1 FW)
SDW (g plant−1)
Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM Control 65 mM 130 mM
62.50h 53.50l 47.78n 69.10cd 61.54jk 56.84i 70.13bc 67.73de 64.38g 66.11f 57.79j 52.61l 63.98g 55.49k 49.78m 75.28a 69.57c 66.97ef 71.17b 64.44g 60.42i
0.1847ghi 0.148l 0.127n 0.206c 0.1788hi 0.162jk 0.219ab 0.199d 0.1848gh 0.192ef 0.164j 0.146l 0.188gf 0.156k 0.136m 0.224a 0.207c 0.196de 0.217b 0.192ef 0.1786i
8.80k 13.07j 14.64h 6.68m 14.92h 16.64d 5.96no 16.15f 17.86b 7.90l 14.18i 16.20ef 8.43k 13.48j 15.40g 5.57° 16.61de 18.48a 6.28mn 15.50g 17.31c
6.33d 7.57b 8.23a 4.87i 5.70g 6.16e 3.92n 4.46k 4.68j 5.18h 6.23de 6.85c 5.85f 6.84c 7.49b 3.90n 4.33l 4.53k 4.19m 4.88i 5.18h
5.48k 4.53° 3.82q 6.97e 5.97i 5.37l 7.80b 6.92e 6.33h 6.40h 5.45kl 4.83n 5.89j 4.98m 4.36p 8.02a 7.18d 6.74f 7.54c 6.54g 5.97ij
In each column, means with at least one similar letter are not significantly different according to the LSD (0.05). LSD (0.05), least significant differences at P ≤ 0.05. Table 4 Mean comparisons for interaction effects of genotype* salicylic acid on malondialdehyde content (MDA) of seven chicory genotypes. Genotype
Ardestan
Hamedan
Mazandaran
Kashan
Shiraz
Sanandaj
Yazd
SA
MDA (nmol g−1 FW)
Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM Control 0.5 mM 1 mM
8.37a 7.17c 6.59e 6.10f 5.46h 5.18i 4.68j 4.26l 4.13m 6.74d 5.96g 5.56h 7.62b 6.52e 6.05fg 4.50k 4.17lm 4.08m 5.16i 4.65j 4.45k
In each column, means with at least one similar letter are not significantly different according to the LSD (0.05). LSD (0.05), least significant differences at P ≤ 0.05.
production of shoot dry matter under salt stress conditions. One of the most important factors in the plants growth and development is photosynthesis rate. The concentration of photosynthetic pigments plays a substantial role in enhancing photosynthetic efficiency (Hopkins and Huner, 2008). In the present research, Chl a, Chl b and Car contents of the chicory genotypes significantly lessened with elevating salinity levels, which resulted in diminished photosynthetic capacity of the plants and eventually decreased growth and dry matter productions. The decline in Chl contents by increasing salinity levels 4
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Table 5 Mean comparisons for interaction effects of salinity * salicylic acid on relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes. Salinity
SA
RWC (%)
MSI (%)
Chl a (mg g−1
Control Control Control 65 mM 65 mM 65 mM 130 mM 130 mM 130 mM
0 mM 0.5 mM 1 mM 0 mM 0.5 mM 1 mM 0 mM 0.5 mM 1 mM
69.53ab 70.80a 71.55a 59.83f 65.55cd 67.46bc 55.70g 61.70ef 64.04de
67.40b 68.10b 69.47a 56.71g 62.64d 64.96c 52.07h 58.09f 60.74e
0.832b 0.868a 0.889a 0.622f 0.744d 0.786c 0.545g 0.648ef 0.682e
FW)
Chl b (mg g−1
Car (mg g−1
FW)
0.238b 0.251a 0.259a 0.163e 0.207c 0.228b 0.130f 0.167de 0.179d
LPC (μmol g−1
FW)
0.197c 0.206b 0.210a 0.165g 0.180e 0.188d 0.148h 0.163g 0.172f
FW)
6.62h 7.22g 7.42g 12.92f 15.36d 16.25c 14.36e 17.22b 18.36a
In each column, means with at least one similar letter are not significantly different according to the LSD LSD (0.05), least significant differences at P ≤ 0.05.
MDA (nmol g−1
FW)
5.12g 4.84h 4.71i 6.33b 5.55d 5.27f 7.06a 5.97c 5.46e
SDW (g plant−1) 6.67c 6.92b 7.03a 5.56g 6.02e 6.23d 4.98i 5.39h 5.66f
(0.05).
Table 6 Correlation coefficients among relative water content (RWC), membrane stability index (MSI), chlorophyll acontent (Chl a), chlorophyll b content (Chl b), carotenoid content (Car), leaf prolin content (LPC), malondialdehyde content (MDA) and shoot dry weight (SDW) of seven chicory genotypes under salinity stress (lower triangle, bold type) and control conditions (upper triangle). Traits RWC MSI Chl a Chl b Car LPC MDA SDW
RWC 1 0.93** 0.694** 0.691** 0.89** 0.90** −0.35** 0.46**
MSI
Chl a **
0.90 1 0.772** 0.773** 0.96** 0.94** −0.58** 0.67**
Chl b **
0.561 0.631** 1 0.99** 0.729** 0.752** −0.512** 0.574**
Car **
0.566 0.632** 0.99** 1 0.72** 0.75** −0.515** 0.579**
LPC **
0.85 0.92** 0.655** 0.650** 1 0.88** −0.65** 0.74**
MDA ns
0.08 −0.14ns −0.33** −0.31* −0.24* 1 −0.51** 0.55**
ns
0.05 −0.18ns −0.37** −0.37** −0.33** 0.87** 1 −0.95**
SDW 0.09ns 0.33** 0.463** 0.45** 0.464** −0.88** −0.97** 1
** Significant at P ≤ 0.01. * Significant at P ≤ 0.05. ns Not significant.
Acknowledgment
compared to non-salt-treated plants. Salt stress adversely restricts plant growth and development through its influence on the photosynthesis, antioxidant phenomena, proline metabolism, osmolyte accumulation and other physiochemical process of plant cells (Aftab et al., 2010; Idrees et al., 2012). Application of SA alleviated the destructive effects of salinity that led to increase in growth parameters and total biomass. Salicylic acid has a key impress in regulation of plant growth and development under stress conditions that causes the function of the photosynthetic apparatus to improve and increase plant biomass (Senaratna et al., 2000). The results of this research were in accordance with the reports of El-Tayeb (2005); Idrees et al. (2012) and Pirastehanosheh et al. (2014). Based on measured traits under salinity condition, SDW had the highest correlation (Table 6) with MDA (r=-0.95**) followed by Car (r = 0.74**) and MSI (r = 0.67**), respectively.
We sincerely thank to the anonymous reviewers for the helpful and constructive comments on the earlier version of our manuscript. We would also like to thank for Isfahan University of Technology for financing this research. References Aftab, T., Khan, M.M.A., Idrees, M., Naeem, M., Moinuddin, A.S., 2010. Salicylic acid acts as potent enhancer of growth, photosynthesis and artemisinin production in Artemisia annua L. J. Crop Sci. Biotech. 13, 183–188. Aftab, T., Khan, M.M.A., Silva, J.A.T., Idrees, M., Naeem, M., Moinuddin, A.S., 2011. Role of salicylic acid in promoting salt stress tolerance and enhanced artemisinin production in Artemisia annua L. J. Plant Growth Regul. 30, 425–435. Amaducci, S., Pritoni, G., 1998. Effect of harvest date and cultivar on Cichorium intybus yield components in north Italy. Ind. Crop. Prod. 7, 345–349. Ashraf, M., Athar, H.R., Harris, P.J.C., Kwon, T.R., 2008. Some prospective strategies for improving crop salt tolerance. Adv. Agron. 97, 45–110. Ashrafi, E., Razmjoo, J., Zahedi, M., Pessarakli, M., 2015. Screening alfalfa for salt tolerance based on lipid peroxidation and antioxidant enzymes. Agron. J. 107, 1–7. Azooz, M.M., 2009. Salt stress mitigation by seed priming with salicylic acid in two faba bean genotypes differing in salt tolerance. Int. J. Agric. Biol. 11, 343–350. Baert, J.R.A., Bockstaele, E.J.V., 1992. Cultivation and breeding of root chicory for inulin production. Ind. Crop. Prod. 1, 229–234. Bajji, M., Kinet, J.M., Lutts, S., 2002. The use of the electrolyte leakage method for assessing cell membrane stability as a water stress tolerance test in durum wheat. Plant Growth Regul. 36, 61–70. Bastam, N., Baninasab, B., Ghobadi, C., 2013. Improving salt tolerance by exogenous application of salicylic acid in seedlings of pistachio. Plant Growth Regul. 69, 275–284. Bates, L.S., Waldran, R.P., Teare, I.D., 1973. Rapid determination of free proline for water studies. Plant Soil 39, 205–208. Belesky, D.P., Fedders, J.M., Turner, K.E., Ruckle, J.M., 1999. Productivity, botanical composition, and nutritive value of swards including forage chicory. Agron. J. 91, 450–456. Boyd, D.C., Rogers, M.E., 2004. Effect of salinity on the growth of chicory (Cichorium intybus cv. Puna) - A potential dairy forage species for irrigation areas. Aust. J. Exp. Agric. 44, 189–192.
5. Conclusion In light of above discussion, it can be concluded that salt stress led to significant decline in RWC, MSI, Chl a, Chl b, Car and SDW and significant increase in LPC and MDA compared to control condition. Also, increasing salinity levels, intensified extent of reduction or enhancement of the measured traits. Foliar application of SA significantly ameliorated all measured traits and the greatest improvements were obtained in the highest concentration of SA. Among genotypes, Sanandaj had the highest RWC, MSI, Chl a, Chl b, Car, LPC and the lowest content of MDA that caused to the highest SDW in comparison with other genotypes while Ardestan had the lowest RWC, MSI, Chl a, Chl b, Car, LPC and the highest content of MDA that caused to the lowest SDW. Drawing upon such findings, foliar application of SA can play a role in diminishing deleterious effects of salinity stress and reinforcing chicory plant tolerance to salt stress. 5
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