physiological and biochemical response of common bean genotypes (phaseolus vulgaris l.) treated with...

Physiological and biochemical response of common bean genotypes (Phaseolus vulgaris L.) treated with salicylic acid under natural drought stress

IJPBCS


Y.A.A. Molaaldoila1,4, M. M. Al-Aqil2, A. H. A. Al-Haj3 1Department of Agronomy, Crop physiology, The Southern Highland Research Station (Taiz-Ibb), Yemen

2 Plant Breeding and Genetic. The Agricultural Research and Extension Authority (AREA), Yemen.

3 Plant Production, Horticulture Vegetables, Faculty of Agriculture and Vit. Med., Ibb Universiy, Yemen

4The Agricultural Research and Extension Authority, Khormaksar, Aden, Yemen.

A field experiment was established over three rainy seasons (2011, 2012 and 2013) at two locations (Shaban and Al-Qaidah) which represented the severe (SDS) and moderate (MDS) drought stress in the southern highlands of Yemen-Ibb. The experiment was arranged in a split plot design with three replication. The main plots were: untreated or treated seeds with 0.5 mM salicylic acid (SA), the subplot were nine CIAT bean lines and three local cultivars. The results revealed that salicylic acid improved significantly the yield and yield traits of some genotypes. Accordingly, under severe drought stress, the bean genotypes categorized into three groups; The first group (MIB-156, MIB-156, G23818B and NSL) which were high yielding and low responsiveness genotypes to SA group (HY-LSAR); The second group (BFB-139, BFB-140 and BFB-141) that perform low yielding and high responsiveness genotypes to SA (LY-HSAR) and the third group (Taiz-304, Taiz-5 and Taiz-306) that perform low yielding and low responsiveness genotypes to SA group (LY-LSAR). It is concluded that the physiological mechanism of bean cultivars response of the high tolerant lines (LY-HSAR) and the medium tolerant lines to drought (LY-HSAR) to SA was similar by causing significant increase in dry matter accumulation, photosynthetic pigments content of leaves and accumulation of high proline content, total soluble sugars, total free amino acids, and soluble proteins, and also by maintain high relative water content (RWC%) and low leaf ion leakage (LIL%) in comparison to susceptible cultivars to drought (LY-LSAR).

Key words: Soluble proteins, proline content, photosynthetic pigments, (RWC%), (LIL%), INTRODUCTION Common bean (Phaseolus vulgaris L.) is considered one of the most important grains for human alimentation and is worldwide planted on approximately 12 million hectares and is sensitive to severe environmental stress situations, such as heat and water deficiency (Bajji et al. 2001, Parry et al. 2002). This fact must be analyzed in depth, since more than half of the worldwide common bean production is grown in regions of occurrence of water deficit (Souza et al. 2003). Large genotypic differences in drought tolerance among

crops also have been reported within-species and genetic variability for tolerance to drought also has been identified in common bean (Beebe et al., 2006; Singh et al., 2003; Molaaldoila, 2016).

*Corresponding author: Y. A. A. Molaaldoila, The Agricultural Research and Extension Authority, Khormaksar, Aden, Yemen. P. O . Box. 6289. Tel: 967-777-271-041. E-mail: [email protected]

International Journal of Plant Breeding and Crop Science Vol. 4(1), pp. 152-163, January, 2017. © www.premierpublishers.org. ISSN: 2167-0449

Research Article


Molaaldoila et al. 153 However, In Yemen most of the local common bean cultivars are susceptible to drought (Molaaldoila, et al., 2016). One of the useful practice used to reduce the inhibitory effect of environment stresses was the application of SA. Significant variation in the response to SA application was observed in many crop cultivars such as in faba bean (Azooz, 2009), common bean (Machado Neto and Duraes, 2006) and sunflower (Noreen et al., 2009) genotypes that can develop different mechanisms of adaptation to stress and responded to SA application differently. The exact function of SA is however uncertain; it can be its ability to reduce the damaging effects of salt or water stress through restoration of various physiological and biochemical plant alteration. It is generally assumed that SA as stress-induced proteins might play a role in stress tolerance such stress-induced proteins might play a role in stress tolerance and this protective role may be essential for the survival of plants under extreme stress conditions (Abou Al-Hamad, 2007; Zahra et al., 2010 and Ismail et al., 2011). Sahar et al. (2011) recorded that the high soluble protein content in salt-stressed plants could play an important role in increasing the osmotic pressure of the cytoplasm and salt tolerance. In this context, Ismail et al. (2011) found that the soluble proteins have been decreased by SA treatment with the salinized wheat seedlings. There are contradictory reports about the role of salicylic acid on photosynthetic pigments. Lusia et al. (2005), reported that methyl salicylic do not have any effect on photosynthetic pigments but photosynthesis decrease under treatment salicylic acid. It is reported that salicylic acid causes increasing photosynthetic pigments in plant, under salty stress (El- Tayeb, 2005) and with salicylic acid, the leaves fill up more soluble sugar and proline (Szepesi, 2006). The objective of this study were (i) To evaluate the physiological and biochemical characteristics in common bean genotypes subjected to drought stress, (ii) To alleviate the inhibitory effect of drought stress by salicylic acid application and (iii) To understand the mechanism of SA action in alleviate the adverse effect of natural drought stress MATERIAL AND METHODS Experimental Design and Environmental Locations The response of bean cultivars (Phaseolus vulgaris L.) to the application of salicylic acid under drought stress condition was investigated. The aim of this study was to evaluate some physiological and biochemical

characteristics in common bean cultivars subjected to natural drought stress and to alleviate the adverse effect of drought by salicylic application. A field experiment was established over three rainy seasons (2011, 2012 and 2013) at two locations (Shaban and Al-Qaidah) which represented the severe (SDS) and moderate (MDS) drought stress in the southern highlands of Yemen-Ibb. The experiment was arranged in a split plot design with three replication. The main plots were: untreated or treated seeds with 0.5 mM salicylic acid (SA), the subplot were bean cultivars; (MIB-156), (G23818B), (BFB-140), (BFB-143), (BFB-144), ( Taiz-304), (Taiz-305) , (Taiz-308), ( Taiz-309). In SA treatments seeds were soaked in 0.5 mM SA for 6 hrs where SA was dissolved in absolute ethanol and then added drop wise to water (ethanol:water, 1:1000, v/v) (Williams et al., 2003) and after that seed were planted. Plant Phenology and Production After three weeks of drought stresses the plants were harvested and the dry weight obtained by oven-drying at 65°C for 48h, dry weight of the roots drought weight g plant

-1(RDW) and shoots drought weight g plant

-1(SDW)

were determined and the root/shoot ratio of plant was calculated for dry weights at the sampling stage. At harvest: Seed yield (kg ha

-1), biological yield, harvest

index, pod numbers/plant, seeds numbers/plant and 100 seed weight (g plant

-1) were recorded and values of the

two formers were adjusted to 14% moisture by weight. Harvest index (HI) that is seed biomass dry weight at harvest/total shoot biomass dry weight at mid-pod filling × 100 also recorded. Determination of Photosynthetic Pigments: For chlorophyll and carotenoids we used method of Lichtenther (1987) and Welfare et al. (1996). Hence, chla, chlb, chlT and car show the concentration of chlorophyll a, chlorophyll b, the total chlorophyll and carotenoides (include carotene and xanthophylls), respectively. Accordingly, the results of measuring photosynthetic pigments content was calculated and presented in fresh weight in gram Determination of Leaf Water Content and Leaf Ion Leakage %: Relative leaf water content (LRWC%) is a useful measure of the physiological water status of plants was determined according to the method of (Teran, and Singh, 2002). Leaf ion leakage % for measuring leakage of cell membrane was evaluated by the method of Marty's et al. (2005).. Protein Metabolism and Soluble Sugars Tissue powder samples of shoots (50 mg) were extracted


Int. J. Plant Breeding Crop Sci. 154 twice in distilled water with continuous stirring for 30 min at 60 °C. After cooling, the water extract was centrifuged and the supernatant was decanted and completed to a definite volume using distilled water. The soluble proteins were then determined in the supernatant by folin reagent according to the method adopted by Lowry et al. (1951). For evaluating proline content in leaf tissue, we use method of Bates et al. (1973) and The results of measuring proline content was calculated and

presented with µg mg -1

DW. Free amino acids were

extracted from plant tissues and determined according to the method of Moore and Stein (1948). The water-soluble sugars were estimated by the method of anthrone sulphuric acid method described by Badour (1959). Statistical Analysis Statistical analysis was carried out with the aid of S.A.S. statistical package (SAS institute Inc., USA) and mean comparison according to Duncan Multiple Range Test (DMRT) at P < 0.05. For data analysis, the cropping seasons and replications were considered as random effects and (SA

+) versus (SA

-) treatments and common

bean genotypes as fixed effects (Mcintosh, 1983). In this paper we are representing the overall average of the three seasons for all the parameters and also we are focusing on the action of salicylic (SA) treatment in ameliorating the adverse effect of natural drought stresses. RESULTS AND DISCUSSION Yield and Yield Attributes The results revealed that the SDS had inhibitory effect on the seed yield and biological yield of bean genotypes more than MDS. The seed yield and biological yield of SDS reduced significantly to the extent of 27.9, and 17.1% in comparison with SDS, respectively. However, SA application reduced the deleterious effect of SDS and improved the seed yield and biological yield to about 23.3, and 22.2%, respectively. In contrast, the results didn’t show significant changes of SA application in harvest index although SDS increased HI to the extent of 18.2% in comparison with MDS (Table 1). On the other hand, pods number per plant, seeds number and 100 seed weight of common bean genotypes were substantially affected by severe drought in comparison with moderate drought. The pods number/plant, seeds number and 100 seed weight were reduced genotypes grown under SDS to about 35.9, 26.6and 33.2%, respectively. However, when SA applied under SDS, it improved pods number/plant, seeds number and 100 seed weight to the extent of 32.0, 24.6 and 25.6%, respectively (Table 2). These results are in accordance

to some earlier studies in which it has been observed that exogenous application of SA promotes the growth and counteracts the stress-induced growth inhibition due to abiotic stresses in different crop species (Metwally et al., 2003; Shakirova et al., 2003; Singh & Usha, 2003). In contrast, working with maize, Nemeth et al., (2002) reported that exogenously applied SA through the rooting medium caused an increase in growth inhibition. Genotypes responded to SDS and MDS differently in seed yield, and other yield traits. However, application of SA not only mitigated the inhibitory effect of drought stress on some of these genotypes, but also in some cases induced a stimulatory effect on greater than that estimated in the control plants. Accordingly, under severe drought stress, the bean genotypes can be categorized into three groups; The first group (MIB-156, MIB-156, G23818B and NSL) that were high yielding and low responsiveness genotypes to SA (HY-LSAR); The second group (BFB-139, BFB-140 and BFB-141) that perform low yielding and high responsiveness genotypes to SA (LY-HSAR) and the third group (Taiz-304,Taiz-305and Taiz-306) that perform low yielding and low responsiveness genotypes to SA group (LY-LSAR). However, these yield traits didn’t changes significantly on HY-LSAR or LY-LSAR. On the other hand, effect of SDS was more drastic than the MDS Interestingly, the LY-LSAR responded to SA application significantly under both SDS and MDS in seed yield, and other yield traits. Moderate to high drought stress can reduce biomass, number of seeds and pods, harvest index, seed yield, and seed weight in common bean (Acosta-Gallegos and Adams, 1991; Ramirez-Vallejo and Kelly, 1998). However, exogenously applied SA through the rooting medium caused an increase in photosynthesis, plant growth and yield under non-stress or drought stress conditions (Natr & Lawlor, 2005).

Root, Shoot Dry Weight and Shoot/ Root Ratio

In this study, the RDW, SDW and SRR were decreased significantly under SDS as compared with the MDS. The reduction were to the extent of 29.4, 38.6 and 13.1%, respectively. On the other hand, SA treatment caused a significant increase in root and shoot dry weight. The extent of increase in RDW, SDW and SRR under SDS were about 30.9, 43.5 and 18.0%, respectively. However, the accumulation of RDW, SDW and SRR improved significantly in HY-HSAR and LY-LSAR bean genotypes due to SA application under SDS environments in comparison with MDS. The increase of SRR indicated that SA application induced accumulation dry matter in shoot more than root in comparison with HY-LSAR. SDW caused a significant improvement in root and shoot dry weights of bean genotypes (Table 3). These results correspond with


Molaaldoila et al. 155

Table 1. The action of salicylic (SA) treatment in improving the adverse effect of SDS and MDS on seed yield (t/ha), Biological yield (t/ha) and Harvest index %

Traits/

Genotypes

Seed Yield Biological yield Harvest index %

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

MD

S

M-155 2.424 2.538 2.481 4.459 4.720 4.590 45.7 46.3 46.0

M-156 2.475 2.464 2.470 4.390 4.638 4.514 43.6 46.8 45.2

G23818B 2.462 2.548 2.505 4.514 4.792 4.653 45.3 46.7 46.0

NSL 2.483 2.604 2.544 4.469 4.772 4.621 44.5 45.4 45.0

BFB-139 2.343 2.482 2.413 4.460 4.766 4.614 47.3 47.9 47.6

BFB-140 2.381 2.537 2.459 3.865 4.439 4.153 38.4 42.8 40.6

BFB-141 2.270 2.407 2.339 4.067 4.616 4.342 44.1 47.9 46.0

Taiz-304 1.676 1.948 1.812 3.322 3.958 3.640 49.7 50.7 50.2

Taiz-305 1.788 2.146 1.967 3.229 4.011 3.620 44.6 46.5 45.6

Taiz-306 1.852 2.306 2.079 3.436 4.088 3.762 46.1 43.7 44.9

Average 2.121 2.286 2.307 4.021 4.385 4.250 44.9 46.5 45.7

DMRT at 0.05 0.261 0.321 0.402 0.517 0.576 0.561 NS NS NS

CV% 18.6 19.9 23.4 23.2 21.7 25.0 18.3 21.5 19.1

SD

S

M-155 1.889 2.315 2.102 3.735 4.498 4.116 49.4 48.5 49.0

M-156 1.825 2.244 2.035 3.784 4.492 4.138 51.8 50.0 50.9

G23818B 1.767 2.253 2.010 3.765 4.384 4.074 53.1 48.6 50.8

NSL 1.601 2.260 1.931 3.505 4.742 4.123 54.3 52.3 53.3

BFB-139 1.771 2.153 1.962 3.511 4.892 4.201 49.6 56.0 52.8

BFB-140 1.808 1.747 1.777 3.960 4.717 4.338 54.3 63.0 58.7

BFB-141 1.793 1.768 1.780 3.495 4.852 4.174 48.7 63.6 56.1

Taiz-304 0.910 1.627 1.268 2.634 3.171 2.902 65.4 48.7 57.1

Taiz-305 0.977 1.783 1.380 2.416 3.176 2.796 59.6 43.9 51.7

Taiz-306 0.938 1.783 1.360 2.513 3.916 3.215 62.7 54.5 58.6

Average 1.528 1.993 1.760 3.332 4.284 3.808 54.9 52.9 53.9

DMRT at 0.05 0.320 0.296 0.332 0.377 0.372 0.379 9.3 7.7 7.7

CV% 20.5 22.7 25.5 25.3 20.6 21.5 16.7 20.0 24.9


Int. J. Plant Breeding Crop Sci. 156

Table 2. The action of salicylic (SA) treatment in improving the adverse effect of SDS and MDS on pods number/plant, seeds number/plant and 100 seed weight (gm)

Traits/

Genotypes

Pod number/plant seed number/plant 100 seed weight

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

MD

S

M-155 27.1 27.1 27.1 121.2 124.2 122.7 27.9 29.3 28.6

M-156 25.7 26.1 25.9 122.5 124.6 123.6 28.2 31.3 29.8

G23818B 27.5 28.3 27.9 125.5 129.5 127.5 26.8 28.3 27.6

NSL 25.0 27.0 26.0 121.5 128.8 125.2 27.0 29.4 28.2

BFB-139 23.8 27.2 25.5 119.9 125.3 122.6 29.6 32.6 31.1

BFB-140 25.9 27.5 26.7 122.7 124.7 116.7 25.6 27.0 26.3

BFB-141 26.2 27.6 27.0 120.8 126.6 118.7 27.3 29.7 28.5

Taiz-304 18.2 24.4 20.1 95.0 113.0 101.5 20.1 23.4 22.3

Taiz-305 18.9 24.9 21.6 99.9 119.8 109.9 18.8 24.1 23.2

Taiz-306 20.6 26.4 29.7 111.7 128.6 120.1 20.6 25.1 23.3

Average 23.9 26.7 25.7 116.1 124.5 118.9 25.2 28.0 26.9

DMRT at 0.05 5.8 6.8 6.5 9.8 10.8 12.1 NS NS NS

CV% 15.4 16.8 16.9 15.1 18.3 18.1 16.5 17.8 17.1

SD

S

M-155 18.7 23.4 21.0 93.7 113.2 108.7 19.4 24.0 21.7

M-156 18.2 24.2 21.2 95.2 113.5 109.7 20.8 25.0 22.9

G23818B 17.6 24.7 21.2 88.3 117.4 107.8 17.9 23.3 20.6

NSL 17.0 23.0 20.0 93.0 117.1 110.2 17.6 23.9 20.7

BFB-139 16.5 23.7 20.1 93.0 119.4 111.3 18.4 24.9 21.6

BFB-140 14.2 22.0 18.1 84.7 118.4 106.3 15.1 21.8 18.4

BFB-141 13.4 22.0 17.7 76.4 113.7 96.8 16.1 20.8 18.5

Taiz-304 11.8 20.7 16.3 69.5 95.7 84.0 13.1 20.6 16.8

Taiz-305 12.7 20.5 16.6 75.2 108.3 96.0 14.2 20.7 17.4

Taiz-306 12.8 20.8 16.8 83.3 113.8 105.7 15.4 20.9 19.6

Average 15.3 22.5 18.9 85.2 113.0 103.6 16.8 22.6 19.8

DMRT at 0.05 3.6 4.0 4.6 6.6 7.6 6.8 4.6 4.7 4.7

CV% 19.3 21.4 21.8 16.5 17.5 17.3 15.5 17.6 17.6



Table 3. The action of salicylic (SA) treatment in improving the adverse effect of SDS and MDS on root (RSW), shoot (SDW) dry weight (gm/plant-1) and root/shoot ratio (SRR)

Traits/

Genotypes

RDW SDW SRR

MD

S

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

M-155 7.11 7.62 7.40 77.6 89.4 83.5 11.0 11.7 11.3

M-156 7.10 7.96 7.55 81.0 89.6 85.3 11.4 11.4 11.4

G23818B 7.12 7.66 7.40 70.7 77.9 74.3 10.0 10.2 10.1

NSL 6.95 7.92 7.45 66.3 72.7 69.5 9.5 9.2 9.4

BFB-139 6.43 6.99 6.50 64.0 72.2 68.1 10.0 10.3 10.2

BFB-140 7.03 7.18 6.85 58.4 73.1 65.8 8.3 10.4 9.3

BFB-141 6.66 6.76 6.50 62.0 70.1 66.1 9.3 10.4 9.8

Taiz-304 5.19 6.63 5.90 57.9 67.1 62.5 11.2 10.2 10.7

Taiz-305 5.58 8.07 6.85 63.3 78.9 71.1 11.4 9.8 10.6

Taiz-306 5.75 8.31 7.00 64.8 89.1 77.0 11.3 10.8 11.0

Average 6.49 7.51 6.94 66.6 78.0 72.3 10.3 10.4 10.4

DMRT at 0.05 2.00 2.30 2.40 4.2 4.7 4.5 NS NS NS

CV% 18.3 19.0 18.75 14.7 20.1 23.0 14.2 17.3 19.3

SD

S

M-155 5.24 7.60 7.10 53.7 76.9 74.4 10.2 10.1 10.2

M-156 5.11 7.28 7.05 50.3 85.0 79.4 9.8 11.7 10.8

G23818B 3.95 7.52 6.20 42.0 81.8 75.4 10.6 10.9 10.7

NSL 4.29 7.62 6.50 45.0 75.4 66.5 10.5 9.9 10.2

BFB-139 4.98 6.49 6.15 37.8 75.3 64.5 7.6 11.6 9.6

BFB-140 4.90 6.18 5.55 34.7 62.1 57.0 7.1 10.0 8.6

BFB-141 4.97 6.06 5.20 32.4 65.8 51.0 6.5 10.9 8.7

Taiz-304 4.30 5.77 4.90 35.0 55.7 50.1 8.1 9.7 8.9

Taiz-305 4.07 5.75 6.40 35.9 69.4 58.7 8.8 12.1 10.5

Taiz-306 4.02 6.03 6.50 41.9 76.4 62.8 10.4 12.7 10.7

Average 4.58 6.63 6.16 40.9 72.4 64.0 9.0 10.9 9.9

DMRT at 0.05 1.23 1.42 1.50 4.3 4.9 5.3 1.2 1.1 1.1

CV% 19.9 18.5 18.5 22.8 23.4 19.9 17.8 16.1 16.3


Int. J. Plant Breeding Crop Sci. 158 the finding of some researchers. For example, Herralde et al., (1998) believed that drought causes decreasing biomass in argyranthemum plant. Working with maize, Nemeth et al., (2002) reported that exogenously applied SA through the rooting medium caused an increase in growth inhibition. In some previous studies, it was found that salt tolerant (S-24) genotype (Pritchard et al., 2001) and moderately salt sensitive (MH-97) genotype (Iqbal & Ashraf, 2005) had high root/shoot ratio. Photosynthetic Pigments SDS caused decrease in photosynthetic pigments of leaves bean genotypes in comparison with moderate drought. The extent of reduction in Chl a, Chl b, under SDS was about 36.4 and 32.5%, respectively. Interestingly, the effect of SDS on Chl a/ Chl b ratio in comparison with MDS was not significant (Table 4). The results also indicated that these bean genotypes can develop different mechanisms of adaptation to drought stress. On the other hand, SA treatment caused a significant increase in Chl a, Chl b. The extent of increase in Chl a, Chl b, under SDS were about 33.2, 29.9 and 14.8%, respectively. The effect of SA application on Chl a, Chl b, were significantly high in HY-HSAR and LY-LSAR bean genotypes due to SA application under SDS environments in comparison with MDS bean genotypes and remained unchanged in HY-HSAR bean genotypes (Table 4). These results revealed beneficial effect of SA and one of the mechanisms of beneficial effect of SA to drought stress is the restoration in photosynthetic pigments of leaves. Similarly, the results showed a decrease in total Chl, Carotenes contents and total Chl : Carotenes ratio in leaves with increasing drought environments. The extent of reduction was about 26.7, 20.4 and 25.2.8%, respectively. However, the application of SA mitigate the adverse effects of drought stress. It increased the total Chl, Carotenes contents and Chl : Carotenes ratio to 28.1, 29.7 and 28.5%, respectively. HY-HSAR and LY-LSAR bean genotypes showed higher total Chl, Carotenes contents and total Chl : Carotenes ratio than HY-HSAR under MDS environment (Table 5). Evidently, the results showed that drought stress alone causes decreasing in chlorophyll a, b, total and Carotenoides in compare with check plants. The decrease of these pigments content on bean cultivars improved with the treatment salicylic acid. These results were in accordance with the results of De Lacerda et al. (2003), Khodary (2004), Parida and Das (2005), Al-Sobahi et al. (2006), Khan et al. (2007), Almodares et al. (2008), Khan et al. (2009) and Carpici et al. (2010). There are contradictory reports about the role of salicylic acid on photosynthetic pigments. Lusia et al. (2005), reported that methyl salicylic do not have any effect on

photosynthetic pigments but photosynthesis decrease under treatment salicylic acid. The results also revealed that the ratio of chlorophyll a/b was decreased with increasing drought. Similar results was obtained by Al-Hakimi (2001) who found adverse effect of salt stress on chlorophyll a/b ratio. On the contrary, chl a/b ratio increased significantly with an increase in NaCl concentration (Al-Sobahi et al., 2006). SA generally, effective in antagonizing partially the inhibitory effect of drought stress on chl. a/b ratio. In this respect, our results are in agreement with these of Mady (2009), they found that application of SA increased chl. a/b ratio in tomato plants. Raafat et al. (2011) reported increase in the chl. a/b ratio of wheat leaves plants in response to SA treatment. In contrast, chl a/b ratio decrease significantly with an increase in SA concentration indicating that SA affected light-harvesting antenna size (Moharekar et al., 2003). The change in the chl. a/b ratio used as an indicator for relative photosystem stoichiomtry (Pfannschmidt et al., 1999). Leaf Water Content % and Leaf ion Leakage %: The genotypes had significantly difference in RWC% and LIL% as well under both MDS and SDS environments. The LRWC exhibited significant decrease under SDS environments in comparison with MDS environments. The reduction in RWC% reached to 23.9% when the plants subjected to SDS environments in comparison with MDS environments. In contrast, LIL% exhibited significant increase under SDS environments in comparison with MDS environments. The increase in LIL% reached to 25.2% when the plants subjected to SDS environments in comparison with MDS environments. However, the application of SA significantly alleviate the adverse effects of MDS stress on RWC% and LIL%. Application of SA caused a significant increase in LRWC and decrease in LIL% bean genotypes under drought environments. The increase in LRWC was to about 25.2% and the reduction in LIL% was to about 39.0 % when the plants subjected to SDS environments in comparison with MDS environments. Furthermore, genotypes differed very markedly in their response to this level of drought stress. HY-HSAR and LY-LSAR bean genotypes maintain high RWC% and low LIL% than HY-HSAR under MDS environment in comparison with HY-LSAR (Table 6). Thus, the resultant limited supply of water in plant would naturally decrease LRWC. The aforementioned results are conceding with those of El-Tayeb (2005) and Yildirim et al. (2008). Parida and Das (2005) reported that the LRWC of plants become more negative with an increase in salinity. The reduction in LRWC leads to flaccidity responsible for stopping of cell division as a cell has to occupy a requisite size before entering in the cell cycle



Table 4. The action of salicylic (SA) treatment in ameliorating the adverse effect of SDS and MDS on Chlorophyll a, Chlorophyll b (µg mg -1 FW) and Chl. a/ Chl. b ratio

Traits/

Genotypes

Chl a Chl b Chla/Chl b

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

MD

S

M-155 0.91 0.93 0.92 0.69 0.78 0.73 1.32 1.19 1.26

M-156 0.88 0.92 0.90 0.68 0.77 0.72 1.30 1.19 1.25

G23818B 0.82 0.89 0.85 0.61 0.74 0.68 1.34 1.20 1.27

NSL 0.82 0.89 0.86 0.58 0.66 0.62 1.43 1.36 1.39

BFB-139 0.80 0.89 0.84 0.53 0.65 0.59 1.50 1.36 1.43

BFB-140 0.77 0.82 0.79 0.52 0.60 0.56 1.50 1.37 1.43

BFB-141 0.72 0.83 0.78 0.52 0.52 0.52 1.40 1.61 1.50

Taiz-304 0.69 0.75 0.72 0.51 0.53 0.52 1.35 1.41 1.38

Taiz-305 0.70 0.76 0.73 0.51 0.56 0.53 1.39 1.36 1.37

Taiz-306 0.71 0.76 0.73 0.49 0.63 0.56 1.45 1.21 1.33

Average 0.78 0.84 0.81 0.56 0.64 0.60 1.40 1.33 1.36

DMRT at 0.05 0.11 0.13 0.12 0.12 0.13 0.12 0.14 0.12 0.10

CV% 16.2 18.2 21.0 18.0 18.8 17.7 24.1 20.4 19.6

SD

S

M-155 0.56 0.72 0.64 0.44 0.63 0.53 1.27 1.15 1.21

M-156 0.58 0.72 0.65 0.42 0.62 0.52 1.36 1.16 1.26

G23818B 0.51 0.71 0.61 0.41 0.59 0.50 1.24 1.21 1.22

NSL 0.51 0.78 0.64 0.43 0.59 0.51 1.20 1.32 1.26

BFB-139 0.51 0.87 0.69 0.36 0.53 0.45 1.41 1.64 1.53

BFB-140 0.47 0.76 0.61 0.38 0.51 0.44 1.25 1.48 1.36

BFB-141 0.46 0.73 0.59 0.41 0.48 0.44 1.15 1.53 1.34

Taiz-304 0.44 0.71 0.57 0.29 0.46 0.37 1.51 1.53 1.52

Taiz-305 0.46 0.72 0.59 0.32 0.53 0.42 1.43 1.37 1.40

Taiz-306 0.47 0.74 0.60 0.33 0.49 0.41 1.41 1.51 1.46

Average 0.50 0.74 0.62 0.38 0.54 0.46 1.32 1.39 1.36

DMRT at 0.05 0.11 0.11 0.13 0.13 0.13 7.57 1.21 1.29 1.26

CV% 18.0 17.5 19.9 17.9 20.9 21.1 16.4 19.6 21.6

(Khan et al., 2007). Our results showed that SA treatments induced an increase in LRWC of the drought stressed plants. Increases in LRWC of bean genotypes treated with SA were also reported for other crops grown under salt stress including tomato (Tari et al., 2002), barley (El-Tayeb, 2005 and Khosravinejad et al., 2008), cucumber (Yildirim et al., 2008), Ocimum basilicum (Delavari et al., 2010) and banana (Bidabadi et al., 2012). The increase of LIL% under drought stress and the alleviation effect of SA on drought stress damages by reducing ion leakage also observed by several workers . It is reported that in maize doesn't have a great change than a check sample lonely in ion leakage, but when the plant placed under drought stress, salicylic causes enough changes in ion leakage (Nemeth et aI., 2002). The studies show that salicylic acid causes preventing from damage to the reduction of membrane leakage and preventing from tilacoide membrane in the time of salty stress in Arabidopsis (Borsanio et al., 2001). SDS also caused significant increase in soluble sugar content in comparison with MDS stress. The increase was about 27.2% when plants were subjected to SDS in comparison with MDS stress. Treatment with

salicylic caused reduction in soluble sugar content to the extent of 18.4%. Furthermore, genotypes differed very markedly in their response to this level of drought stress. LY-HSAR bean genotypes responded significantly to SA application in restoring soluble sugar content than HY-HSAR and LY-LSAR under MDS environment in comparison with HY-LSAR (Table 6). It is reported that increasing proline and sugar causes protecting turgidity and reducing of membrane damage on plants. Thus, osmo-regulation is an adaptation that increase the tolerance toward drought stress (Inze and Montago, 2000). On the contrary, Khodary (2004) reported that the decrease in soluble carbohydrates content by reason of SA treatment might be assumed to inhibit polycarbohydrates-hydrolysing enzyme system or one hand and/or accelerate the incorporation of soluble sugar into polycarbohydrates. Treatment with salicylic caused improving resistance of plant on stress and as a result sugar approach to its normal (Miguel et al., 2006) condition. Protein Metabolism The soluble protein content, proline content and total



Table 5. The action of salicylic (SA) treatment in improving the adverse effect of SDS and MDS on total chlorophyll (Chl T), carotenoids (µg mg -1 FW) and Chl / Car ratio

Traits/

genotypes

Chl T Carotenoids

Total chl/Car

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

MD

S

M-155 1.36 1.53 1.44 0.52 0.58 0.55 2.65 2.67 2.66

M-156 1.38 1.46 1.42 0.45 0.57 0.51 3.10 2.56 2.83

G23818B 1.29 1.40 1.35 0.40 0.52 0.46 3.28 2.77 3.03

BFB-139 1.35 1.50 1.42 0.38 0.57 0.47 3.75 2.67 3.21

BFB-140 1.28 1.49 1.38 0.45 0.57 0.50 2.87 2.63 2.75

BFB-141 1.23 1.36 1.30 0.47 0.52 0.50 2.62 2.61 2.61

NSL 1.19 1.30 1.24 0.36 0.49 0.42 3.37 2.68 3.03

Taiz-304 1.30 1.38 1.34 0.37 0.51 0.43 3.56 2.73 3.15

Taiz-305 1.23 1.46 1.35 0.39 0.53 0.46 3.19 2.76 2.97

Taiz-306 1.19 1.47 1.33 0.38 0.51 0.47 3.18 2.93 3.05

Average 1.28 1.43 1.36 0.41 0.53 0.47 3.16 2.70 2.93

DMRT (P=0.05) 0.17 0.16 0.16 0.15 0.15 0.14 NS NS NS

CV% 17.7 14.6 20.9 18.2 17.0 20.6 22.1 18.3 20.18

SD

S

M-155 1.14 1.35 1.24 0.46 0.50 0.48 2.47 2.70 2.58

M-156 1.19 1.38 1.29 0.39 0.52 0.46 3.06 2.68 2.87

G23818B 1.09 1.39 1.24 0.42 0.47 0.44 2.64 3.00 2.82

BFB-139 1.12 1.36 1.24 0.35 0.52 0.44 3.19 2.67 2.93

BFB-140 1.04 1.39 1.21 0.35 0.51 0.43 2.93 2.75 2.84

BFB-141 1.01 1.18 1.09 0.34 0.43 0.39 2.95 2.74 2.85

NSL 1.03 1.25 1.14 0.37 0.48 0.42 2.82 2.59 2.70

Taiz-304 0.92 1.17 1.04 0.33 0.41 0.37 2.77 2.84 2.80

Taiz-305 0.93 1.30 1.11 0.34 0.43 0.38 2.76 3.04 2.90

Taiz-306 0.96 1.28 1.12 0.31 0.43 0.37 3.10 3.00 3.05

Average 1.04 1.29 1.17 0.36 0.48 0.42 2.87 2.80 2.84

DMRT (P=0.05) 0.18 0.16 0.15 0.12 0.14 0.13 0.18 0.16 0.16

CV% 19.0 20.5 19.2 18.4 20.2 22.1 16.9 17.6 19.6



Table 6. The action of salicylic (SA) treatment in ameliorating the adverse effect of SDS and MDS on relative water content (RWC %), leaf ion leakage % (LIL %) and soluble sugar content (µg mg -1 DW)

Traits/

genotypes

RWC% LIL% Soluble sugar content

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

MD

S

M-155 76.6 79.4 78.0 31.6 27.7 29.7 632.0 691.5 691.8

M-156 78.6 81.8 80.2 33.5 31.0 32.2 627.5 661.5 694.5

G23818B 76.8 81.7 79.2 38.4 25.8 32.1 625.0 656.0 690.5

NSL 76.2 67.3 71.7 37.4 28.2 32.8 603.5 631.0 667.3

BFB-139 78.0 79.4 78.7 35.4 28.0 31.7 674.0 692.0 643.0

BFB-140 74.7 79.0 76.8 38.3 27.3 32.8 670.5 713.0 741.8

BFB-141 77.5 79.0 78.2 36.0 29.9 33.0 608.0 686.0 747.0

Taiz-304 72.6 76.2 74.4 35.5 29.5 32.5 697.0 663.0 730.0

Taiz-305 72.8 75.7 74.3 35.8 28.5 32.1 629.0 654.0 741.5

Taiz-306 72.6 76.3 74.4 36.4 28.3 32.4 632.0 674.0 753.0

Average 75.6 77.6 76.6 35.8 28.4 32.1 639.9 672.2 710.0

DMRT at 0.05 3.2 4.2 5.2 3.3 2.8 2.9 42.3 36.6 36.9

CV% 17.6 17.6 17.7 17.1 16.9 17.7 19.7 18.8 20.2

SD

S

M-155 61.4 77.9 69.7 46.7 25.6 36.1 879.1 757.5 818.3

M-156 61.4 76.8 69.1 46.7 29.9 38.3 892.1 721.0 806.5

G23818B 56.8 77.0 66.9 42.4 28.6 35.5 818.1 649.5 733.8

NSL 58.3 76.7 67.5 43.8 29.2 36.5 851.1 700.0 775.5

BFB-139 55.6 77.2 66.4 51.3 27.8 39.5 918.0 680.5 799.3

BFB-140 55.2 72.1 63.6 50.9 28.7 39.8 929.9 753.0 841.4

BFB-141 55.3 71.9 63.6 51.0 31.9 41.4 831.0 746.0 788.5

Taiz-304 54.8 69.4 62.1 50.6 29.9 40.2 851.5 764.5 808.0

Taiz-305 53.4 72.1 62.7 49.3 31.9 40.6 865.0 696.5 780.7

Taiz-306 54.0 72.5 63.2 49.8 31.0 40.4 947.7 695.5 821.6

Average 56.6 74.3 65.5 48.2 29.4 38.8 878.3 716.4 797.4

DMRT at 0.05 2.0 4.0 3.0 3.3 3.6 2.6 30.9 44.0 37.3

CV% 19.0 20.8 19.0 18.8 20.2 20.3 14.8 17.4 18.5



Table 7. The action of salicylic (SA) treatment in improving the adverse effect of SDS and MDS on soluble protein, proline and free amino acids contents (µg mg -1 DW)

Traits/

genotypes

Soluble protein Proline content Total amino acids

SA- SA

+ Average SA

- SA

+ Average SA

- SA

+ Average

MD

S

M-155 169.3 183.9 176.6 79.0 96.7 87.8 67.3 81.0 74.2

M-156 169.9 180.5 175.2 85.3 95.7 90.5 68.9 82.3 75.6

G23818B 143.5 160.1 151.8 66.3 79.8 73.1 57.8 70.5 64.1

NSL 165.7 183.3 174.5 67.6 90.2 78.9 67.5 82.7 75.1

BFB-139 152.4 168.4 160.4 60.1 89.1 74.6 63.2 75.6 69.4

BFB-140 157.4 173.4 165.4 72.1 84.0 78.0 59.3 72.4 65.9

BFB-141 143.9 163.5 153.7 59.5 69.5 64.5 67.2 77.4 72.3

Taiz-304 152.7 171.9 162.3 44.1 57.3 50.7 54.4 78.8 66.6

Taiz-305 141.5 164.6 153.0 49.0 58.5 53.7 49.1 69.6 59.3

Taiz-306 149.3 172.5 160.9 40.7 52.2 46.4 54.1 76.4 65.2

Average 155.0 172.2 163.4 61.5 77.3 69.8 60.9 77.0 68.8

DMRT at 0.05 22.7 30.8 31.3 20.4 33.7 33.1 18.8 18.6 26.1

CV% 18.9 20.0 21.6 16.1 18.8 21.3 20.9 19.0 17.9

SD

S

M-155 236.4 174.4 205.4 60.6 80.0 73.7 64.7 82.9 73.8

M-156 226.6 174.2 200.4 63.3 83.5 76.9 71.6 82.2 76.9

G23818B 227.2 162.3 194.7 49.7 67.2 61.2 55.9 73.3 64.6

NSL 244.9 185.2 215.1 43.5 59.4 53.9 43.8 82.0 62.9

BFB-139 232.1 176.4 204.2 59.2 82.6 74.2 42.4 74.9 58.6

BFB-140 231.6 173.5 202.5 50.9 79.0 67.8 35.7 70.7 53.2

BFB-141 224.4 166.9 195.6 46.4 75.7 63.6 58.3 78.6 68.4

Taiz-304 210.7 155.7 183.2 45.0 57.7 53.8 38.9 69.9 54.4

Taiz-305 202.1 154.6 178.3 34.6 52.5 45.5 40.1 73.1 56.6

Taiz-306 194.5 153.2 173.9 29.8 43.3 38.2 35.8 64.1 49.9

Average 223.1 167.6 195.3 48.3 68.1 60.9 48.7 75.1 61.9

DMRT at 0.05 24.4 28.1 20.1 12.2 28.0 24.4 26.3 22.6 28.6

CV% 21.4 19.0 19.4 19.9 20.5 23.6 16.1 18.8 21.3


Molaaldoila et al. 163 amino acids of shoot increased by drought stress considerably; the magnitude of increase in soluble protein content, proline content and total amino acids were in the extent of 30.5%, 21.4% and 20.0%, respectively. However, the salicylic acid application minimize the stress effects, by adjusting soluble protein content, proline content and total amino acids of shoot to about 24.9%, 29.0% and 35.2%. Moreover, genotypes differed very markedly in their response to this level of drought stress. The magnitude of restoration on soluble protein content, proline content and total amino acids of shoot were observed in the HY-HSAR bean genotypes than LY-LSAR under MDS environment in comparison with HY-LSAR (Table 7). This can mean that in spite of the low responsiveness of genotypes group LY-LSAR bean genotypes to SA, responded with a positive stimulus in an attempt to minimize the stress effects, an adjustment for which proline is responsible; and that SA application could activate other defense systems or inhibit soluble protein content, proline content and total amino acids accumulation. It is generally assumed that SA as stress-induced proteins might play a role in stress tolerance such stress-induced proteins might play a role in stress tolerance and this protective role may be essential for the survival of plants under extreme stress conditions (Abou Al-Hamad, 2007; Zahra et al., 2010 and Ismail et al., 2011). Sahar et al. (2011) also recorded that the high soluble protein content in salt-stressed plants could play an important role in increasing the osmotic pressure of the cytoplasm and salt tolerance. In this context, Ismail et al. (2011) found that the soluble proteins have been decreased by SA treatment with the salinized wheat seedlings. However, proline and amino acids accumulation has been suggested as the result of degradation or synthesis (Sudhakar et al. 1993), inhibition of the protein synthesis while in common bean it can be related to degradation mechanisms (Andrade et al. 1995). Bates et al. (1973) and Stewart and Larher (1980) pointed out the role of proline as solute during stress, where an increase in the proline content would indicate resistance or tolerance to water deficit, serving as parameter for the selection of highly resistant cultivars. But Maggio et al. (2002) demonstrated that proline accumulating genotypes were susceptible to this type of stress. However, the SA application caused a diminution in proline accumulation, but raised the soluble protein content in the tolerant variety Guarumbe. Compared with the control plants, the proline content increased in plants not treated with SA and dropped with 0.05mM of salicylic acid (Yokota 2003).

CONCLUSION From the above discussion, it can be concluded that

yield and yield traits as well as some biochemical constituents and physiological traits of the investigated bean genotypes were severely deteriorated by drought stress SDS in comparison with MDS. The accumulation of RDW, SDW and SRR, photosynthetic pigments, RWC% and sugar content as well as soluble proteins, amino acids and proline content has been considered a tool for the determination of the drought adaptation and as an indicator of drought stress. However, The use of salicylic acid to alleviate the adverse effect of drought is achieved by the accumulation dry matter and proline, the maintaining high RWC%, sugar content as well as soluble proteins, amino acids and lowering LIL or ion leakage. The beneficial effect of SA could be used for improving their drought tolerance. The results also indicated that these bean genotypes can develop different mechanisms of adaptation to drought stress. Accordingly, under severe drought stress, the bean genotypes categorized into three groups; The HY-LSAR (MIB-156, MIB-156, G23818B and NSL) that were high yielding and low responsiveness genotypes to SA; LY-HSAR (BFB-139, BFB-140 and BFB-141) that perform low yielding and high responsiveness genotypes to SA and LY-LSAR (Taiz-304, Taiz-305and Taiz-306) that perform low yielding and low responsiveness genotypes to SA group. Thus, further studies are required to explicitly elucidate the mechanism of SA influx through different ways and the target enzymes or metabolites involved in plants respond to SA application. ACKNOWLEDGEMENTS The authors would like to thank Dr. Steve Beebe (CIAT) for providing us bean lines samples. We also appreciate the help of Ibb extension experts in locations and farmer fields selection for conducting the experiments.

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Accepted 19 October, 2016. Citation: Molaaldoila YAA, Al-Aqil MM, Al-Haj AHA (2017). Physiological and biochemical response of common bean genotypes (Phaseolus vulgaris L.) treated with salicylic acid under natural drought stress. International Journal of Plant Breeding and Crop Science, 4(1): 152-165.

Copyright: © 2017 Molaaldoila et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are cited.

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