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Research Paper
Research Journal of Agricultural Sciences
8(5): 1081-1086, September-October (2017)
ISSN: 0976-1675 https:
//
www.rjas.org DI: 4494-2605-2017-247
Comparative Study of Relative Water, Chlorophyll and Proline Content in
Drought Tolerant and Susceptible Genotypes of Lentil (Lens culinaris Medik.)
Asit Prasad Dash, *Dilip Kumar De, **Rajib Nath, ***Ashutosh Sarkar and Soumitra Mohanty
Department of Plant Breeding and Genetics,
College of Agriculture, Orissa University of Agriculture and Technology, Bhubaneswar - 751 003, Odisha, India
*Department of Genetics and Plant Breeding, **Department of Agronomy,
Faculty of Agriculture, Bidhan Chandra Krishi Viswavidyalaya, Mohanpur - 741 252, West Bengal, India
***NASC Complex, DPS Marg, Pusa, New Delhi - 110 012
e-mail: dashpraasit@gmail.com
Received: 26 May 2017; Revised accepted: 31 July 2017
A B S T R A C T
The present experiment was carried out at the Department of Genetics and Plant Breeding, Bidhan Chandra Krishi
Viswavidyalaya, West Bengal, India to study the effect of imposed drought on relative water content, chlorophyll
content and proline accumulation in five drought tolerant (PL-406, IPL-324, LL-1146, IPL-325, K-75) and five
drought sensitive (L1112-20, LP-112, ILL-10803, KLS-113, KLS-107) genotypes of lentil (Lens culinaris Medik.). For
this purpose, the genotypes were grown under both control (0.0 bar) and drought stress (-9 bar) condition by using
PEG- 6000 as osmoticum. The experiment was laid out in an asymmetrical factorial completely randomized design
with three replications. Roots, shoots and leaves of ten days old seedlings were harvested and physiological and
biochemical analyses were carried out for the above parameters. From the study it was revealed that relative
water content and chlorophyll content (chl a, Chl b, total Chl) were reduced, while proline content was increased
due to imposition of drought stress with compared to respective control. However, the tolerant genotypes
exhibited lower relative reduction for RWC and chlorophyll content and proline content increased more as
compared to susceptible ones under drought stress condition. The results suggested that higher levels of RWC,
chlorophyll content and proline accumulation in tolerant variety of lentil could play an important role in drought
stress tolerance.
Key words: Lens culinaris, Drought, RWC, Chlorophyll, Proline
limate change is a global phenomenon that renders
adverse impact on agriculture. Drought has been
considered as one of the most devastating abiotic stresses
(Vinocur and Altman 2005). During domestication of crop
plants and their improvement, much emphasis was given for
selection on different desirable characters including yield
and tolerance to drought was unlikely to be one of such
criteria for selection. Taking into consideration global water
scarcity and increases in demand for non-agricultural uses of
water, expansion of the area under irrigation in developing
countries does not appear to be a realistic scenario to address
the challenge of food security. Thus, the scenario as a whole
reveals that when the global food requirement is increasing
rapidly, the water stress is also increasing alarmingly.
Therefore, food production in the twenty-first century will
have to rely more on the release of varieties with higher
potentiality to combat drought and with high yield stability
(Borlaug 2007). In this respect understanding of the
functioning capacity of drought tolerant plants under
drought stress appears inevitable. Different parameters like
RWC, proline, chlorophyll content have been considered as
markers of such stress.
Lentil generally possesses an inbuilt mechanism to
tolerate drought but there are known degrees of differences
between varieties. During the course of evolution, plants
have developed several mechanisms that help in perceiving
the stresses and rapidly regulating their physiological and
metabolic activities to combat them. An understanding of
plants’ physiological and biochemical mechanism to combat
water stress can aid in identifying cultivars that can
profitably be used to produce sustainably high yield.
However, studies on different physiological and biochemical
factors during drought stress in a drought tolerant crop like
lentil still remain elusive. Therefore, the present study was
taken up to compare the changing status of chlorophyll,
proline and relative water content in the seedlings of drought
C
1081
tolerant and sensitive genotypes due to imposition of
drought stress induced by PEG-6000.
MATERIALS AND METHODS
Seeds of forty-eight genotypes of lentil (Lens culinaris
Medik.) procured from AICRP on MULLaRP, Kalyani
Centre, Bidhan Chandra Krishi Viswavidyalaya (BCKV),
Mohanpur, West Bengal, India were screened for drought
tolerance during the period from first week of November to
last week of January in two consecutive years (2013-14 and
2014-15) in the laboratory of the Department of Genetics
and Plant Breeding, BCKV using PEG-6000 as osmoticum.
Based on the performance of forty-eight genotypes in the
laboratory with respect to different morphological characters
at seedling stage and their respective drought tolerance
efficiency, five most tolerant genotypes viz., PL-406, IPL-
324, LL-1146, IPL-325 and K-75 and five most susceptible
genotypes viz. L1112-20, LP-112, ILL-10803, KLS-113 and
KLS-107 were identified for their study on physiological
and biochemical status to drought stress.
Seventy two healthy, viable seeds of each lentil
genotype were surface sterilized by immersing them in 70%
ethanol for 2 minutes followed by thorough washing with
distilled water. Twelve such seeds of a genotype were
arranged in a row with even space over a glass plate (20 cm
× 30 cm) wrapped with a blotting paper. To prevent the
seeds from sliding down when the set was kept in a slanted
position in a stand, another glass strip (20 cm × 2 cm) was
placed over the seeds with the help of a piece of thermocol
at the two ends and guarder in such a way that the seeds
remained in their position and the seedlings grew without
any hindrance. The whole set was then placed in a
transparent polythene bag. There were six such sets for each
genotype; three sets represented three replications for
control and the remaining as three replications for treatment.
In the treatment plates, PEG solution of -9 bar water
potential was used as germinating medium, while in the
control plates pure distilled water was used for the purpose.
The seeds were then allowed to germinate and grow for 10
days under indoor laboratory condition under sufficient light
of 3000 lux, 70-80% relative humidity (RH) and at a
temperature range of 20-25ºC. Six competitive seedlings
were randomly selected from each plate and further analysis
was carried out through destructive sampling for relative
water content, chlorophyll content and proline content
following Barr and Weatherley (1962), Arnon (1949), Bates
et al. (1973) respectively.
RESULTS AND DISCUSSION
Relative water content (RWC)
The data on relative water content in different plant
parts of ten days old seedling of the above mentioned
susceptible and tolerant genotypes grown under control (0.0
bar) and water stress (-9.0 bar) conditions have been
presented in (Table 1). All genotypes exhibited reduced
relative water content in root due to imposition of water
stress but such reduction was comparatively more in
susceptible genotypes than in tolerant ones. Under control
condition, only one genotype from tolerant group (LL-1146)
and two genotypes (KLS-113 and (L-1112-20) from
susceptible group exhibited significantly higher mean RWC
while under drought stress all the genotypes except PL-403
from tolerant group and none from the susceptible group
exhibited significantly higher mean as revealed by CD
(critical difference) value. The highest reduction with
compared to respective control was observed in a
susceptible genotype, KLS-107 (29.257%) and the lowest
was in a tolerant one, LL-1146 (11.054%).
Table 1 Relative water content in different parts of lentil seedlings among tolerant and susceptible genotypes under control
and drought condition
Character
Root
Shoot
Leaf
Genotype
Control
PEG
Soln
RR (%)
Control
PEG
Soln
RR (%)
Control
PEG
Soln
RR (%)
PL-406 (T)
81.784
63.271
21.645
84.392
76.015
9.926
82.357
67.777
17.703
IPL-324 (T)
83.452
65.389
11.054
81.380
72.985
10.316
83.253
71.212
14.463
LL-1146 (T)
85.474
76.026
20.062
86.471
80.221
7.228
84.141
75.420
10.365
IPL-325 (T)
83.493
66.743
12.738
81.949
73.052
10.857
81.077
69.108
14.763
K-75 (T)
83.170
72.576
29.257
80.130
73.137
8.727
83.697
73.600
12.064
KLS-107 (S)
83.467
59.047
29.196
81.730
71.044
13.075
84.097
67.811
19.366
KLS-113 (S)
85.657
60.649
23.113
83.253
72.014
13.500
84.477
66.940
20.759
ILL-10803 (S)
83.164
63.942
24.259
80.156
68.073
15.074
83.220
64.775
22.164
LP-112 (S)
80.173
60.724
27.111
78.920
70.080
11.201
80.310
70.060
12.763
L1112-20 (S)
84.091
61.293
21.645
81.126
68.715
15.298
85.842
65.335
23.889
Mean B
83.393
64.966
74.179
81.951
72.534
77.242
83.247
69.204
76.226
Factors
SE(m)
SE(d)
C.D.(0.05)
SE(m)
SE(d)
C.D.(0.05)
SE(m)
SE(d)
C.D.(0.05)
Genotype (G)
0.035
0.049
0.103
0.035
0.049
0.103
0.042
0.059
0.125
Treatment (T)
0.015
0.022
0.046
0.015
0.022
0.046
0.019
0.027
0.056
G × T
0.049
0.069
0.145
0.049
0.069
0.145
0.059
0.084
0.177
T= Morphologically tolerant S= Morphologically susceptible RR=Relative reduction
Dash et al. 2017
Research Journal of Agricultural Sciences 8(5)
1082
As observed in root in case of shoot also all the ten
genotypes witnessed a reduction of RWC due to imposition
of drought and it was much pronounced in susceptible
genotypes. However, under control condition, two
genotypes from tolerant group (PL-406 and LL-1146) and
only one genotype (KLS-113) from susceptible group
exhibited significantly higher mean RWC but under water
stress condition all the tolerant genotypes exhibited
significantly higher mean but none from the susceptible
group could do so. Interestingly, root was found to be more
influenced by imposed drought than shoot.
In case of leaf, two genotypes from tolerant group (LL-
1146 and K-75) and three (KLS-107, KLS-113 and L1112-
20) from susceptible group under control, while three
genotypes (IPL-324, LL-1146 and K-75) from control group
and only one (LP-112) from susceptible group exhibited
significantly higher mean as revealed by CD value. Though
in all genotypes leaf RWC decreased, it was more in case of
susceptible genotypes as compared to tolerant ones. Among
the three characters the leaves were moderately affected as
revealed by the relative reduction percentage. Therefore, it
may be concluded that imposition of water stress reduced
the relative water content of all the morphological parts of
the seedlings (root, shoot, leaf), which has been reported
earlier by Merah (2001) in case of wheat. The higher
relative reduction of RWC in sensitive genotypes of lentil
suggests more profound effect of drought stress on them as
compared to tolerant ones. The possible cause of this
reduction may be the loss of turgidity of cells under water
stress. According to Ritchie et al. (1990) high relative water
content is a resistant mechanism to drought, and that high
relative water content is the result of more osmotic
regulation or less elasticity of tissue cell wall. Similar
mechanism might have played role in the tolerant genotypes
expressing significantly higher RWC for all the parts under
drought stress in lentil. Earlier Bhardwaj and Yadav (2012)
found significantly higher RWC in the tolerant variety under
drought stress.
Table 2 Chlorophyll-a, Chlorophyll-b, Total chlorophyll and Chl. a/b in the leaves of 10 days old lentil seedlings of
tolerant and susceptible genotypes under control and drought treatment
Characters
Chlorophyll-a (mg/g
tissue)
Chlorophyll-b(mg/g tissue)
Total Chlorophyll mg/g
tissue)
Chlorophyll a/b
Genotype
Control
PEG
Soln.
RR (%)
Control
PEG
Soln.
RR (%)
Control
PEG
Soln.
RR (%)
Control
PEG
Soln.
Mean
PL-406 (T)
0.425
0.363
14.588
0.250
0.182
27.200
0.673
0.544
19.168
1.703
1.998
1.850
IPL-324 (T)
0.280
0.248
11.429
0.158
0.112
29.114
0.436
0.359
17.661
1.777
2.225
2.001
LL-1146 (T)
0.325
0.283
12.923
0.173
0.131
24.277
0.497
0.413
16.901
1.888
2.164
2.026
IPL-325 (T)
0.387
0.339
12.403
0.224
0.174
22.321
0.610
0.512
16.066
1.732
1.956
1.844
K-75 (T)
0.290
0.264
8.966
0.132
0.100
24.242
0.421
0.363
13.777
2.212
2.639
2.425
KLS-107 (S)
0.323
0.201
37.771
0.180
0.117
35.000
0.502
0.316
37.052
1.797
1.724
1.761
KLS-113 (S)
0.451
0.368
18.404
0.342
0.223
34.795
0.791
0.589
25.537
1.320
1.653
1.486
ILL-10803 (S)
0.268
0.196
26.866
0.145
0.082
43.448
0.412
0.277
32.767
1.850
2.394
2.122
LP-112 (S)
0.464
0.389
16.164
0.305
0.221
27.541
0.768
0.609
20.703
1.520
1.761
1.640
L1112-20 (S)
0.376
0.259
31.117
0.214
0.143
33.178
0.589
0.401
31.919
1.764
1.813
1.788
Mean
0.359
0.291
0.325
0.212
0.149
0.180
0.570
0.438
0.504
1.756
2.033
1.894
Factors
SE(m)
SE(d)
C.D(0.05)
SE(m)
SE(d)
C.D.(0.05)
SE(m)
SE(d)
C.D.(0.05)
SE(m)
SE(d)
C.D(0.05)
Genotype (G)
0.003
0.005
0.009
0.003
0.005
0.009
0.006
0.009
0.019
0.024
0.034
0.071
Treatment (T)
0.001
0.002
0.004
0.001
0.002
0.004
0.003
0.004
0.008
0.011
0.015
0.032
G × T
0.005
0.006
0.013
0.005
0.006
0.013
0.009
0.013
0.027
0.034
0.048
0.101
Chlorophyll content
The results obtained with respect to the content of
chlorophyll ‘a’, chlorophyll ‘b’ and total chlorophyll per
unit quantity of fresh leaf tissue from 10 days old seedling
of the above mentioned genotypes grown under control (0.0
bar) and water stress (-9.0 bar) condition presented in (Table
2) reveal significant inter-genotypic differences as indicated
by the CD value. Such differential response might have
arisen due to differential genetic makeup of different
genotypes. The range of chlorophyll ‘a’ content in the
seedlings grown under control in the susceptible group from
0.268 mg/g tissue to 0.464 mg/g tissue and between 0.280
mg/g tissue and 0.425 mg/g tissue in case of tolerant ones
indicated no remarkable difference in content of chlorophyll
‘a’ in the two groups of genotypes. But when the seedlings
were subjected to water stress, there was differential
behavior in the genotypes belonging to the two groups. Both
tolerant as well as susceptible genotypes expressed a
reduction in chlorophyll ‘a’ content due to treatment.
Considering the mean values it was observed that in case of
control, two genotypes from tolerant group viz. PL-406 and
IPL-325 and three genotypes from susceptible group viz.
KLS-113, LP-112 and L1112-20 exhibited significantly
higher mean. While in case of treatment though the same
two genotypes from tolerant group exhibited significantly
higher mean, but two out of three genotypes (KLS-113, LP-
112) from susceptible group could do so. However, the
extent of reduction of chlorophyll ‘a’ content due to drought
Drought Tolerant and Susceptible Genotypes of Lentil
1083
was higher in susceptible genotypes than in the tolerant
ones. While the highest and the lowest reduction in the
content of chlorophyll ‘a’ among the susceptible genotypes
were from KLS-107 (37.771%) and LP-112 (16.164%), the
same from tolerant genotypes were from PL-406 (14.588%)
and K-75 (8.966%) respectively. Therefore, the tolerant
genotypes revealed to be less affected due to imposition of
drought with compared to the susceptible ones. Similar
results in Soybean (Hossaina et al. 2014), Faba bean
(Siddiqui et al. 2015) have been reported earlier.
In case of chlorophyll ‘b’ the mean value under control
varied between 0.145 mg/g and 0.342 mg/g in the
susceptible group and between 0.132 mg/g and 0.250 mg/g
in case of tolerant ones. The genotypes that expressed
significantly higher mean for chlorophyll ‘a’ under control
as well as under water stress condition of both the groups
could do so for chlorophyll ‘b’ as well. As observed in case
of chlorophyll ‘a’, both tolerant and susceptible genotypes
expressed a reduction in chlorophyll ‘b’ content due to
treatment and the extent of such reduction due to drought
stress was higher in susceptible genotypes than in the
tolerant ones. Highest reduction of chlorophyll ‘b’ content
was observed in a susceptible genotype, ILL-10803
(43.448%), while lowest was in a tolerant genotype, IPL-
325 (22.321%). Such decrease in chlorophyll content under
drought stress may be considered a typical symptom of
oxidative stress and may be the result of pigment photo-
oxidation and chlorophyll degradation.
Generally, the total chlorophyll content reflects the
additive result of the two types of pigments studied,
therefore, as expected, similar result was also observed for
total chlorophyll content, where the susceptible genotypes
expressed a higher relative reduction compared to tolerant
ones. It may, therefore, be stated that chlorophyll ‘a’ and ‘b’
as well as total chlorophyll content reduced due to
imposition of drought condition. However, in both the
groups, the content of chlorophyll ‘b’ was more affected due
to treatment than that of chlorophyll ‘a’. The present
findings were in conformity with Kumar et al. (2011) who
reported that drought stress reduced concentration of
chlorophyll ‘b’ more than chlorophyll ‘a’. Due to the
imposition of water stress the ratio of two types of pigments
(chlorophyll-a/b) was increased in all genotypes except
KLS-107 indicating a higher reduction of chlorophyll ‘b’ as
compared to chlorophyll ‘a’. Such increase was observed to
be more in tolerant genotypes than in susceptible ones.
According to Farooq et al. (2009) both the chlorophyll ‘a’
and ‘b’ are prone to soil dehydration. Reduction in total
chlorophyll content was also reported by Aniat-ul-Haq et al.
(2012); Al-Quraan et al. (2014) in lentil and Saglam et al.
(2014) in chickpea. Manivannan et al. (2007) has reported a
large decline in chlorophyll ‘a’, chlorophyll ‘b’ and the total
chlorophyll content in different sunflower varieties due to
water stress. According to Smirnoff (1995) reduction in
chlorophyll under drought stress is mainly the result of
damage to chloroplasts caused by reactive oxygen species.
Therefore, in the present experiment imposition of drought
might have caused damage to chloroplasts through reactive
oxygen species.
Table 3 Proline content in lentil seedlings of tolerant and susceptible genotypes under control and drought conditions
Genotypes
Proline content (micromole/g of fresh tissue)
Mean
Control
PEG soln
PL-406 (T)
110.520
249.070
179.795
IPL-324 (T)
97.970
237.930
167.950
LL-1146 (T)
166.100
361.810
263.955
IPL-325 (T)
154.170
370.440
262.305
K-75 (T)
113.540
320.660
217.100
KLS-107 (S)
130.140
203.310
166.725
KLS-113 (S)
108.320
211.320
159.820
ILL-10803 (S)
115.220
190.860
153.040
LP-112 (S)
80.170
139.760
109.965
L1112-20 (S)
73.120
120.520
96.820
Mean
114.927
240.568
177.748
Factors
SE(m)
SE(d)
C.D.(0.05)
Genotype(G)
0.190
0.268
0.564
Treatment(T)
0.085
0.120
0.252
G × T
0.268
0.380
0.798
Proline content
It is well described that under stress conditions many
plant species accumulate proline as an adaptive response to
adverse conditions. Although a clear-cut relationship
between proline accumulation and stress adaptation has been
questioned by some authors, it is generally believed that the
increase in proline content following stress injury is
beneficial for the plant cell (Mattioli et al. 2009). In the
present experiment, analysis of proline content revealed a
significant inter-genotypic difference in the seedlings grown
under control as well as under water stress condition (Table
3). Among the susceptible genotypes grown under control,
only KLS-107 recorded significantly higher mean, whereas
two genotypes viz. LL-1146 and IPL-325 from the tolerant
group exhibited significantly higher mean proline content. It
was observed that the proline content increased in all the
Dash et al. 2017
Research Journal of Agricultural Sciences 8(5)
1084
genotypes due to treatment but such increase was significant
in four of the five tolerant genotypes viz. PL-406, LL-1146,
IPL-325 and K-75 but none among the susceptible
genotypes. Earlier Talukdar (2013), Muscolo et al. (2014) in
lentil and Rizvi et al. (2014) in chickpea reported proline
accumulation under drought stress. Highest increase (182.42
%) in proline accumulation observed in K-75 is a tolerant
genotype and the lowest increase (56.224%) was in KLS-
107, a susceptible genotype. Interestingly, K-75 and IPL-
325 revealed very low relative reduction for RWC and
chlorophyll content also which indicated their consistency in
tolerance to drought. According to Kavi-Kishor et al. (2005)
Proline is one of the most important organic solutes that has
been reported in plants to maintain the water content under
stressful conditions by acting as osmoprotectant for
membrane stabilization. Increased level of proline in PEG
induced water stress in plants may be an adaptation for the
purpose of overcoming the stress conditions. Generally,
proline accumulates under stress condition to supplies
energy for survival and growth and thereby to help the plant
to tolerate stress (Sankar et al. 2007, Aktas et al. 2007).
According to Kumar et al. (2011), increase in the free
proline content during water stress condition would suggest
that proline is one of the common compatible osmolytes
under water stress condition that helps to combat drought.
Therefore, from the present findings it may be concluded
that all genotypes have potential to withstand drought
through production of higher amount of proline under stress.
But generally the tolerant genotypes have higher potential to
withstand drought due to more accumulation of the amino
acid. For further study on the inheritance pattern of drought
tolerance, the genotypes like K-75, IPL-324, and IPL-325
from tolerant group and KLS-113 and LP-112 from
susceptible group may be used in future.
In the present study it was observed that both relative
water content and chlorophyll content reduced, while proline
content increased in both susceptible and tolerant genotypes
due to imposition of water stress. Though the difference in
the three parameters under control condition was not
remarkable among the genotypes within the two groups
separately, a conspicuously higher value was observed for
all parameters in tolerant genotypes compared to susceptible
genotypes under water stress condition could be noticed.
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