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

Physiological and biochemical response of common bean genotypes (Phaseolus vulgaris L.) treated with salicylic acid under natural
drought stress
IJPBCS
Physiological and biochemical response of
common bean genotypes (Phaseolus vulgaris L.)
treated with salicylic acid under natural
drought stress
Y.A.A. Molaaldoila1,4
, M. M. Al-Aqil2
, A. H. A. Al-Haj3
1
Department 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
4
The 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: yaldoila@yahoo.com
International Journal of Plant Breeding and Crop Science
Vol. 4(1), pp. 152-163, January, 2017. © www.premierpublishers.org. ISSN: 2167-0449
Research Article

drought stress
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

drought stress
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

drought stress
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
MDS
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
SDS
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

drought stress
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
MDS
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
SDS
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

drought stress
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
MDS
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
SDS
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

drought stress
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

drought stress
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
+
AverageMDS
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
SDS
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

drought stress
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
MDS
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
SDS
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

drought stress
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
MDS
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
SDS
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

drought stress
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
MDS
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
SDS
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

drought stress
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.

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

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