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Selecting Sesame Genotypes for Drought

Tolerance Based on Some Physiochemical
Traits
ARTICLE in AGRONOMY JOURNAL JANUARY 2014
Impact Factor: 1.54 DOI: 10.2134/agronj2013.0260

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Crop Economics, Production & Management

Selecting Sesame Genotypes for Drought Tolerance Based

on Some Physiochemical Traits
Azam Kadkhodaie, Jamshid Razmjoo, Morteza Zahedi, and Mohammad Pessarakli*

Abstract

To assess drought tolerance, an understanding of sesame (Sesamum indicum L.) physiological properties is needed. The objective
of this study was to determine the physiological responses of sesame accessions to drought stress. The experiment was conducted
as a two-way factorial arranged in a randomized complete block design with three replications. All of the measured traits were
accession- and drought-level-specific and discriminated the accessions into tolerant (Isfahan4, Rashtkhar, and Borazjan),
moderately tolerant (Isfahan1, Ahvaz, Shiraz, Ardestan, and Kal), and sensitive (Markezi, Ultan, Khaf2, Khaf1, and Shahreza)
groups. The results also showed that Khaf2 and Isfahan4 were the best accessions under normal and drought conditions,
respectively. In addition, based on the correlation coefficient of the traits with seed yield, root proline and leaf carotenoid
content were the superior markers with regard to drought tolerance in sesame. This study showed that accessions differences in
drought tolerance could be attributed to the ability of sesame plants to accumulate metabolites under drought conditions, and
these metabolites could be a useful tool for depicting water-deficit tolerance of sesame genotypes, which could be useful to plant
breeders for selecting and developing drought-tolerant cultivars.

Drought is a serious problem for crop production

in arid and semiarid areas of the world. Organic solutes such
as proline and carbohydrates accumulate in the plant cells to
reestablish cell turgor during drought stress. Proline plays a
major role in osmotic adjustment and also protects the cells
by scavenging reactive oxygen species (ROS) (Pinhero et al.,
2000; Mittler et al., 2004), while carbohydrates are used for
maintaining metabolism and saving energy under drought
(Khalid et al., 2010). Furthermore, the ability of a plant to
synthesize more chlorophyll under water deficit may increase
drought tolerance (Pinhero et al., 2000). The effectiveness of
these mechanisms under drought stress depends on the plant
species, the cultivar within the plant species, and the length
and intensity of the stress (Reddy et al., 2004).
Sesame, due to its high oil, antioxidant, and protein content,
is a very important crop (Koca et al., 2007). Sesame drought
tolerance is important because it is regularly subjected to
mild or severe water stress (Golestani and Pakniyat, 2007);
therefore, its quality and yield may be reduced under drought
stress if drought-tolerant genotypes are not used. Moreover,
A. Kadkhodaie, J. Razmjoo, and M. Zahedi, Dep. of Agronomy and Plant
Breeding, Isfahan Univ. of Technology, Isfahan, Iran; and M. Pessarakli,
School of Plant Sciences, Univ. of Arizona, Tucson, AZ 85721. Received 2
June 2013. *Corresponding author (pessarak@ag.arizona.edu).
Published in Agron. J. 106:111118 (2014)
doi:10.2134/agronj2013.0260
Copyright 2014 by the American Society of Agronomy, 5585 Guilford
Road, Madison, WI 53711. All rights reserved. No part of this periodical
may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, recording, or any information storage and
retrieval system, without permission in writing from the publisher.

drought tolerance mechanisms of sesame genotypes need to

be identified so that drought-tolerant genotypes can be either
planted directly in the field by farmers or used as markers in
breeding programs.
Under drought-stress conditions, carotenoids, soluble sugars,
and proline contents increased in sunflower (Helianthus
annuus L.; Nazarli et al., 2011), while chlorophyll, cartenoids,
proline, and total sugars decreased (Parida et al., 2007).
Hassanzadeh et al. (2009b) and Molaei et al. (2012) had similar
results and reported that proline can be used to assess drought
tolerance in sesame.
Consequently, the objectives of this study were to determine
the effects of drought stress on leaf relative water, chlorophyll,
carotenoid, soluble sugar, and leaf and root proline contents
and seed yield of sesame genotypes and to rank them for their
drought tolerance based on these traits.
MaterialS and Methods
Plant Material and Treatments
The experiment was a two-way factorial (3 13) arranged
in a randomized complete block design with three replicates;
each replicate contained four plants (12 plants per treatment
were used for various parameter measurements that were
sampled). The first factor was three drought levels (55% [I1],
75% [I2], and 85% [I3] depletion of the available soil water)
and the second factor was 13 sesame genotypes (Ultan, Khaf2,
Khaf1, Shahreza, Markazi, Shiraz, Kal, Ardestan, Rashtkhar,
Isfahan4, Ahvaz, Isfahan1, and Borazjan). Seeds of these
Abbreviations: ROS, reactive oxygen species; RWC, relative water content;
SSI, stress susceptibility index.

A g ro n o my J o u r n a l Vo l u m e 10 6 , I s s u e 1 2 014

111

sesame genotypes were sown into pots (20-cm diameter and

60-cm height) under natural conditions at the College of
Agriculture, Isfahan University of Technology, Iran, during
two growing seasons (2011 and 2012).
The soil was a clay loam Typical Haplargid, with a bulk
density of 1.34 g cm3, a pH of 7.5 (measured in a 1:2 soil/
water suspension), water-holding capacity at field capacity of
240 g kg1 (determined gravimetrically), organic C content
of 2.7 g kg1 (measured by wet-digestion analysis), total N of
300 mg kg1 (determined by the Kjeldahl digestion method),
and available P and available K of 14.9 and 250 mg kg1
(measured by the Olsen method and extraction with
NH4OAc, respectively) (Soil Survey Laboratory Staff, 2004).
Drought treatments started 30 d after planting. Drought
treatments were determined based on managed allowed
depletion (MAD) (Allen et al., 2000) after 55% (I1), 75% (I2),
or 85% (I3) depletion of the available soil water; 55% MAD was
used as the control or non-drought-stress treatment. The soil
water potential based on depletion of the available soil water
was determined by a soil moisture release curve. The amount of
soil water was defined as (Allen et al., 2000)

q irrig = q FC -(q FC -q pwp )P

where q irrig (g kg1) is the average water in the root
development depth at the time of irrigation, q FC is the soil
water content at field capacity (g kg1), q pwp is the soil water
content at the permanent wilting point (g kg1), and P is the
fraction of available water that can be extracted from the soil
by plants without showing water stress symptoms. The volume
of irrigation water was determined as (Allen et al., 2000)

V irrig = (q FC -qavg )r bV pot

where Virrig is the volume of irrigation water, which at the end
for each irrigation treatment (I1, I2 , and I3) was 0.060, 0.048,
and 0.045 m3, respectively, q FC is soil water content at field
capacity (%), q avg is the average water in the root development
depth, Vpot is the volume of the pot, and r b is the bulk density
of the soil. The time of irrigation was determined based on the
soil water potential measured by gypsum blocks. Leaf relative
water, chlorophylls, carotenoid, soluble sugars, and proline
contents, root proline content, and seed yield were measured.
All of the physiological parameters were determined at the full
flowering stage.
Total leaf (the third fully developed leaf from the apex of the
plant) and root proline contents were determined according to
the method of Bates et al. (1973). The soluble sugars content
in the leaves was determined based on the phenolsulfuric acid
method (Irigoyen et al., 1992). Leaf chlorophyll and carotenoid
contents were determined based on the method of Gao (2006).
Leaf relative water content was determined according to the
method of Smart and Bingham (1974).
Stress Susceptibility Index
The stress susceptibility index (SSI) was calculated according
to Fischer and Maurer (1978):
112

SSI =

1-Y si Y pi

SI = 1-

SI

Ys
Yp

where Ypi is the yield of the genotype under non-stress

conditions, Ysi is the yield of the genotype under stress
conditions, Ys is the mean yield of all genotypes under stress
conditions, and Yp is the mean yield of all genotypes under nonstress conditions. A lower SSI indicates higher drought tolerance.
Statistical Analysis
The data from the 2 yr were combined, and the combined
data were subjected to analysis of variance (ANOVA) using the
GLM procedure of SAS (version 9.2, SAS Institute). Treatment
means were separated using the LSD test (p < 0.05).
Results and Discussion
Leaf proline, chlorophylls, carotenoid, and soluble
carbohydrate contents, relative water contents, root proline
content, and seed yield were significantly affected by genotype,
drought level, and their interaction (Table 1). Similar results
were also reported by others (Sharifi et al., 2012; Shahandeh et
al., 2013).
Leaf Relative Water Content
The highest relative water content (RWC) was noted in
Khaf1 under I1, in Shahreza under I2 (with Rashtkhar), and
in Rashtkhar under I3 and on average, while Isfahan4 under I1
and Ultan under I2 and I3 had the lowest RWC (Table 2). The
largest increase in RWC was recorded in Shahreza under I2 and
in Isfahan4 under I3, whereas the largest decrease was obtained
in Khaf1 under I2 and I3 (Table 2). Ramos et al. (2003) in bean
(Phaseolus vulgaris L.) and Fazeli et al. (2006) in sesame showed
a reduction in RWC under drought stress. Ganji Arjenaki et
al. (2012) and Talebi et al. (2013) had similar results in wheat
(Triticum aestivum L.) and chickpea (Cicer arietinum L.) and
reported that drought-tolerant cultivars maintained higher
RWC under drought than drought-sensitive cultivars.
Water stress causes water loss within the plant cells and,
therefore, a reduction in RWC. Consequently, leaf water
content indicates the extent of cell dehydration and thus
may be used to assess cellular damage. The RWC is one of
the most reliable and widely used indicators for defining
both the sensitivity and the tolerance to water stress in
plants (Sanchez-Rodriguez et al., 2010). The RWC may also
indicate the balance between water absorbed by the plant
and consumed through transpiration. Osmoregulation is one
of the mechanisms of preserving turgor pressure in plants
against water loss under water stress. It causes the plant to
continue water absorption and thus to retain its metabolic
activity (Gunasekera and Berkowitz, 1992). This suggests that
RWC in our study was regulated by other parameters such as
proline, carbohydrate, and carotenoid contents and antioxidant
enzymes (Yousfi et al., 2010; Arabzadeh, 2012).
Agronomy Journal Volume 106, Issue 1 2014

Table 1. Analysis of variance for the effect of different drought levels on

relative water content (RWC), leaf soluble sugars (LSS), leaf proline content (LPC), root proline content (RPC), chlorophyll a, chlorophyll b, total
chlorophyll, carotenoid content, and seed yield of sesame genotypes.
Genotype Drought
Block
(G)
(D)
DG
df
2
12
2
24
RWC
59.33 ns
843.8*
112.8** 211.8**
LSS
567.3 ns 18380.9** 9806.7** 3351.9**
LPC
1.2 ns
102.0**
269.9**
19.5**
RPC
2.6 ns
861.3**
761.4** 150.2**
Chlorophyll a
0.09 ns
1.6**
1.3**
0.76**
Chlorophyll b
0.003 ns
0.16**
0.07*
0.06**
Total chlorophyll 0.0012 ns
2.6**
0.97**
1.2**
Carotenoid
0.0003 ns 0.008**
0.01**
0.005**
Seed yield
0.10 ns
3.7**
31.6**
1.5**
Trait

Error
76
35.41
400.4
1.4
7.9
0.16
0.01
0.22
0.001
0.19

* Significant at the 0.05 probability level; ns, not significant.

** Significant at the 0.01 probability level.

Leaf Soluble Carbohydrates

The highest leaf soluble carbohydrates were produced in
Khaf2 under I1 and in Borazjan under I2 and I3, while the
largest increase in leaf soluble carbohydrates was detected in
Borazjan under both I2 and I3 and the largest decrease was
noted in Khaf1 and Shahreza under I2 and in Shahreza under
I3 (Table 2). Increased soluble carbohydrate contents under
drought stress could be due to inhibition of growth and the
hydrolysis of complex storage carbohydrates such as starch
by the decreasing water potential (Moradshahi et al., 2004).
Soluble sugars, however, depend on species, genotype within
species, drought level and duration, developmental stage, and
experimental conditions and may increase (Mostajeran and
Rahimi-Eichi, 2009; Yousfi et al., 2010), decrease (Silvente
et al., 2012), or even stay constant (Pirzad et al., 2011) under
drought. Soluble sugars are sources of C and energy in cells
and they are important signaling molecules, playing important
roles in the mechanisms of stress tolerance in plants through
osmotic adjustment (Mostajeran and Rahimi-Eichi, 2009). In
addition, Pinhero et al. (2000) reported that carbohydrates

may feed the oxidative pentose phosphate pathway that can

contribute to ROS scavenging. Consequently, fluctuations in
sugar content that accompany alterations in the environment
will, therefore, influence ROS production, placing sugars as key
players in the redox balance in plants. Furthermore, Arabzadeh
(2012) concluded that plants actively accumulate soluble sugars
that can enhance the plants water-holding capacity and reduce
drought stress via regulation of the plant osmotic potential.
Leaf and Root Proline Contents
The highest leaf proline content was recorded in Ardestan
under I1 and in Isfahan4 under I2 and I3, while the lowest
was noted in Rashtkhar under I1 and in Kal under I2 and
I3 (Table 3). The largest increase in leaf proline content was
detected in Borazjan under I2 and I3, while the lowest was
noted in Khaf1 under I2 and I3 (Table 3). Root proline content
was also increased under drought; however, the increase was
greater under I2 (Table 3). The highest root proline content was
recorded in Khaf2 under I1 and in Isfahan4 under I2 and I3
and on average, while the lowest was measured in Ahvaz under
I1, in Ardestan under I2 , and in Shahreza under I3 (Table 3).
The largest root proline content increase was noted in Isfahan4
under I2 and I3, whereas the smallest increase and the largest
decrease were noted in Khaf2 under I2 and in Shahreza under
I3, respectively (Table 3).
Moradshahi et al. (2004), Fazeli et al. (2006), and Omidi
(2010) had similar results and reported that proline content
increased with yield in various crops, including sesame and
canola (Brassica napus L.) under drought stress conditions.
Pirzad et al. (2011), however, showed that the proline content
of Matricaria chamomilla L. was not affected, and Silvente et
al. (2012) reported that the proline content of soybean [Glycine
max (L.) Merr.] cultivars was not changed in early stages but
increased in the flowering stage under drought stress. These
contrasting results could be due to differences in irrigation
regimes, stages of growth, environmental conditions, and plant
genotypes. In addition, Fazeli et al. (2006) concluded that

Table 2. Effects of drought level (D), genotype (G), and their interaction on leaf relative water content (RWC) and soluble sugars of sesame under three
levels of drought (I1, I2 , and I3 = 55, 75, and 85% depletion of available soil water).
RWC
Accession
Rashtkhar
Khaf2
Khaf1
Isfahan4
Shahreza
Borazjan
Ahvaz
Kal
Shiraz
Markazi
Ardestan
Ultan
Isfahan1
Avg.

I1

I2

Leaf soluble sugars

I3

Avg.

%
51.6 cde
66.9 a (+29.5)
72.0 a (+39.4)
63.5 A
42.2 efghi
39.9 fghij (5.5)
27.3 lmn (32)
36.5 E
56.2 bc
37.9 fghijk (32.6) 32.2 jklm (42.6) 42.1 D
21.8 n
28.6 klmn (+31)
31.6 jklm (+45)
27.3 H
42.8 efgh
67.7 a (+58)
34.6 hijkl (19)
48.4C
53.4 bcd
39.9 fghij (25)
62.6 ab (+17)
52.0 B
32.9 ijklm
38.8 fghij (+18)
36.1 ghijkl (+9.5) 35.9E
31.9 jklm
35.6 ghijkl (+12)
30.9 jklmn (3)
32.8 F
39.2 fghij
46.2 def (+18)
42.9 efgh (+9.6) 42.8 D
44.4 defgh
43.2 efgh (2.5)
39.4 fghij (11)
42.3D
37.8 fghijk
46.0 def (+21.6)
42.7efgh (+13)
42.2D
36.7 fghijkl
27.1 lmn (26)
24.6 mn (33)
29.5 G
40.1 fghij
45.2 defg (+13)
43.7 defgh (+9)
43.0 D
40.8B
43.3 A
40.1 B

I1

I2

I3

Avg.

mg g1
137.6 jklmno
199.0 c (+44.6)
179.4 cdef (+30)
172.0 D
197.9 cd
175.7 cdefg (14)
165.2 efghijk (16.5)
179.6 C
146.2 ghijklmno
124.0 no (15)
119.6 o (18)
130.0 I
129.5 mno
182.4 cdef (+41)
166.17defghijk (+28.6)
159.3 E
167.5 cdefghij
141.9 ijklmno (15)
126.8 mno (24)
145.4 FG
174.1 cdefgh
350.0 a (+101)
321.0 a (+84.4)
281.7 A
123.3 o
163.7 efghijkl (+33)
141.5 ijklmno (+15) 142.8 FGH
130.6 mno
156.6 efgn (+20)
125.8 no (3.6)
137.7 H
135.6 jklmno
167.7 cdefghij (+23.6)
135.8 jklmno (0.3)
146.4 F
188.3 cde
164.6 efghijk (12.6)
177.6 cdefg (6)
176.8 CD
131.3 lmno
158.74 efghijklm (+21)
129.1 mno (2)
139.7 GH
152.6 fghijklmno 133.6 klmno (12.4)
130.6 mno (14.4)
138.9 H
174.5 cdefgh
283.3 b (+62.4)
272.0 b (+60)
243.3 B
153.0 C
184.7 A
168.5 B

Means in columns and row (interaction) for each treatment followed by the same lowercase letter(s) are not significantly different at the 5% probability level.
Values in the parentheses are the percentage reduction (-) or increase (+) compared with the control (I1).
Means in each column (main effect of genotype) or row (main effect of drought level) for each treatment followed by the same uppercase letter(s) are not significantly different at the 5% probability level.

Agronomy Journal Volume 106, Issue 1 2014

113

Table 3. Effects of drought level (D), genotype (G), and their interaction on leaf and root proline contents of sesame under three levels of drought
(I1, I2 , and I3 = 55, 75, and 85% depletion of available soil water).
Leaf proline content
Accession

I1

I2

Root proline content

I3

Avg.

I1

I2

I3

Avg.

g1

Rashtkhar
Khaf2
Khaf1
Isfahan4
Shahreza
Borazjan
Ahvaz
kal
Shiraz
Markazi
Ardestan
Ultan
Isfahan1
Avg.

mmol

1.18 o
2.43 lmno (+105) 5.24 hijk (+346) 2.95 H
10.52 efghijklm
34.72 c (+230)
37.10 c (+252.5)
27.4B
3.89 klm
4.34 kl (+11.4)
4.56 ijk (+17)
4.27 G
16.01 defgh
17.81 def (+11)
12.98 ghijk (19)
15.6C
5.27 hijk
5.09 hijk (3.5)
5.52 ghijk (+5)
5.29 F
10.02 jklm
11.18 ijkl (+11.5)
7.33 lmnopq (27)
9.5F
5.06 hijk
12.0 c (+139)
23.62 a (+360)
13.5 A
14.51 fghij
49.15 b (+243)
55.44 a (+284.4)
39.7A
5.75 ghijk
6.43 fghij (+12)
6.50 efghi (+13)
6.23 E
4.73 opq
5.38 lmnopq (+14)
3.29 q (30)
4.4I
1.90 no
4.68 ijk (+145.5)
9.00 d (+372)
5.19 F
6.33 mnopq
19.68 de (+211)
20.42 d (+221)
15.4C
2.01 mno
4.73 ijk (+135)
8.11 def (+311)
4.95 F
3.31 q
8.79 klmno (+165)
8.47 klmnop (+158)
6.8H
1.63 o
2.34 mno (+43.5) 3.81 klmn (+133) 2.59 H
9.24 klmno
12.15 hijk (+31)
10.02 jklm (+8.5)
10.4E
4.33 kl
6.74 efgh (+56)
11.20 c (+159)
7.42 D
6.91 lmnopq
17.68 def (+156)
16.92 defg (+146)
13.8D
3.85 klmn
4.49 jk (+16.6)
6.47 fghi (+68)
4.94 F
15.86 efgh
17.69 def (+13)
12.81 ghijk (18.6)
15.4C
8.05 def
11.76 c (+46)
18.29 b (+127)
12.70 B
3.90 q
5.05 opq (+30)
4.24 pq (+9)
4.3I
7.39 defg
8.44 de (+14)
8.94 d (+21)
8.26 C
14.75 fghi
16.85 defg (+15)
10.66 ijklm (27.5)
14.0D
4.70 ijkk
8.21 def (+75)
11.63 c (+147.4) 8.18 C
3.79 q
10.80 ijklm (+185)
9.81 klmn (+160.6)
8.10G
4.23 C
6.29 B
9.54 A
9.22 C
17.45 A
16.11 B

Means in columns and row (interaction) for each treatment followed by the same lowercase letter(s) are not significantly different at the 5% probability level.
Values in the parentheses are the percentage reduction (-) or increase (+) compared with the control (I1).
Means in each column (main effect of genotype) or row (main effect of drought level) for each treatment followed by the same uppercase letter(s) are not significantly
different at the 5% probability level.

Table 4. Effects of drought level (D), genotype (G), and their interaction on the chlorophyll a and b contents of sesame under three levels of drought
(I1, I2 , and I3 = 55, 75, and 85% depletion of available soil water).
Chlorophyll a
Accession
Rashtkhar
Khaf2
Khaf1
Isfahan4
Shahreza
Borazjan
Ahvaz
kal
Shiraz
Markazi
Ardestan
Ultan
Isfahan1
Avg.

I1

I2

Chlorophyll b
I3

Avg.

I1

I2

I3

Avg.

mg g1
2.11 klmno
3.87 a (+83)
3.72 ab (+77)
3.23 A
0.593 no
1.20 abc (+103)
1.00 cdefghi (+69)
0.934 C
3.07 bcdefg 2.36 ijklmn (23) 2.10 klmnop (31)
2.51 E
0.933 efghijk
0.853 fghijkl (9)
0.820 ghijklmn (13) 0.868 D
3.37 abcde
2.56 fghijkl (24)
2.31 ijklmn (32) 2.75 CD
1.11 abcde
0.923 efghijk (17)
0.890 efghijkl (20) 0.976 BC
1.89 nop
3.57 abc (+86)
3.18 bcdef (+68)
2.88 BC
0.796 ghijklmn
1.24 ab (+56.5)
1.24 ab (+57)
1.09 A
3.37 abcde 2.83 defghij (16) 2.54 fghijklmn (25) 2.91 B
1.19 abcd
1.04 bcdefg (12.5) 0.976 defghij (18.5)
1.07 A
1.45 p
2.18 jklmno (+50) 1.93 mmop (+33)
1.85 G
0.566 o
0.783 ijklmno (+38) 0.820 ghijklmn (+43.5) 0.723 F
2.04 lmnop 2.61 efghijkl (+28) 2.22 jklmno (+9)
2.29 F
0.610 mno
0.773 jklmno (+27) 0.783 hijklmno (+29) 0.722 F
2.83 cdefghi 3.38 abcde (+19)
2.79 efghij (2)
3.00 B
0.913 efghijk 1.08 abcdef (+18.5)
0.903 efghijk (1.2) 0.965 BC
1.89 nop
2.72 defghijk (+44) 2.10 klmnop (+12)
2.24 F
0.796 hijklmn
0.730 klmno (8)
0.720 klmno (10)
0.748 F
3.07 bcdefg 2.67 fghijkl (13)
2.39 hijkl (22)
2.71 D
0.893 efghijkl
0.830 ghijklm (7)
0.823 ghijklm (9)
0.848 DE
2.93 cdefghi
3.04 cdefg (+4)
2.95 cdefghi (1)
2.98 B
0.963 efghij
1.02 cdefgh (+6)
0.963 efghij (1.5)
0.983 B
3.47 abcd
2.96 cdefghi (15) 2.45 ghijklmn (29) 2.96 B
1.29 a
1.07 abcdef (17)
0.976 defghij (24.4)
1.11 A
1.59 op
2.33 ijklmn (+46) 2.03 lmnop (+28)
1.98 G
0.666 lmno
0.906 efghijk (+36)
0.893 efghijkl (+33)
0.822 E
2.54 B
2.85 A
2.51 B
0.87 C
0.95 A
0.90 B

Means in columns and row (interaction) for each treatment followed by the same lowercase letter(s) are not significantly different at the 5% probability level.
Values in the parentheses are the percentage reduction (-) or increase (+) compared with the control (I1).
Means in each column (main effect of genotype) or row (main effect of drought level) for each treatment followed by the same uppercase letter(s) are not significantly
different at the 5% probability level.

plants actively accumulate proline that can enhance the plants

water-holding capacity and reduce drought stress by regulating
the osmotic potential of the plant. In contrast, IbarraCaballero et al. (1988) showed that proline accumulation was
not directly related to drought resistance in maize (Zea mays
L.). Increases in proline content under drought could be due
to induction of proline biosynthesis as well as inhibition of its
oxidation under water stress (Fazeli et al., 2006). Results from
these studies suggest that proline content may be related to
drought tolerance.
Chlorophyll Content
Chlorophyll a content was increased under I2 but remained
unchanged under I3 compared with I1; however, the response
of the genotypes was dependent on the irrigation level and
114

chlorophyll a content was highest in Ultan under I1 and in

Rashkhar under I2 and I3 and lowest in Borazjan under I1, I2 ,
and I3 (Table 4). The largest increase in chlorophyll a content
was detected in Isfahan4 under I2 and in Rashtkhar under I3,
while the largest decrease was noted in Khaf1 under both I2
and I3 (Table 4).
Chlorophyll b was increased under drought, but the
increase was greater under I 2 than I 3; however, changes in
chlorophyll b content was genotype- and drought-levelspecific and the largest increase was noted in Rashtkhar
under I 2 and I 3, while the largest decrease was obtained
in Khaf1 under I 2 and in Ultan under I3 (Table 4). Ultan
contained the most chlorophyll a under I1, while Isfahan4
contained the most chlorophyll b under both I 2 and I 3 (Table
4). On the other hand, the chlorophyll b content was lowest
Agronomy Journal Volume 106, Issue 1 2014

Table 5. Effects of drought level (D), genotype (G), and their interaction on total chlorophyll and carotenoid content of sesame under three levels of
drought (I1, I2 , and I3 = 55, 75, and 85% depletion of available soil water).
Total chlorophyll
Accession
Rashtkhar
Khaf2
Khaf1
Isfahan4
Shahreza
Borazjan
Ahvaz
kal
Shiraz
Markazi
Ardestan
Ulan
Isfahan1
Avg.

I1

I2

Carotenoid content
I3

Avg.

I1

I2

I3

Avg.

mg g1
2.70 lmno
5.07 a (+88)
4.73 ab (+75)
4.17 A
0.167 nopq
0.248 defghij (+46.4)
0.299 abcd (+79)
0.238 DE
4.00 bcdefgh 3.24 h-m (19.6)
2.94 jklmn (27)
3.39 E
0.263 abcdefg
0.190 lmnopq (28)
0.177 mnopq (33)
0.210 G
4.49 abcd 3.47 fghijk (22.4) 3.24 hijklm (28.6) 3.73 C
0.252 cdefghi 0.209 ghijklmn (17) 0.197 ijklmnop (22) 0.220 FG
2.69 lmno
4.76 ab (+77.4)
4.42 abcde (+65) 3.95 B
0.183 lmnopq 0.248 defghij (+35.6)
0.312 ab (+72)
0.247 CD
4.57 abc
3.87 cdefgh (15) 3.49 fghijk (23.5) 3.98 B
0.202 hijklmno
0.171 nopq (15)
0.140 q (31)
0.171 I
2.02 o
2.97 jklmn (+47) 2.72 klmno (+36) 2.57 H
0.145 pq
0.202 hijklmno (+39) 0.239 efghijk (+64.4) 0.195 H
2.65 mno 3.39 fghijklm (+28) 3.01 ijklmn (+13.4) 3.02 F
0.176 mnopq
0.233 fghijkl (+32)
0.275 abcdef (+55.6) 0.228 EF
3.75 defghi
4.49 abcd (+19)
3.69 efghij (1.6) 3.79 B
0.232 fghijkl
0.245 defghij (+6)
0.289 abcde (+24)
0.255 C
2.69 lmno 3.45 fghijkl (+28.4) 2.85 klmn (+5.4)
2.99 F
0.244 efghijk
0.259 bcdefg (+6)
0.306 abc (+29)
0.269 B
3.96 cdefgh 3.49 fghijk (12)
3.29 hijklm (19) 3.51 E
0.255 defgh
0.227 ghijklm (13)
0.227 fghijklm (13)
0.236 E
3.90 cdefgh 4.07 bcdef (+4.3)
3.87 cdefgh (1) 3.57 D
0.257 bcdefg
0.270 abcdef (+5.5)
0.315 a (+23)
0.281 A
4.76 ab
4.03 bcdefg (15) 3.43 fghijkl (28) 4.07 AB
0.279 abcdef 0.236 efghijkl (15.5) 0.196 jklmnop (29.5) 0.237 DE
2.25 no
3.29 ghijklm (+43) 2.95 jklmn (+29.5) 2.83 G
0.149 opq
0.184 lmnopq (+23.6) 0.246 defghij (+64)
0.193 H
3.42 B
3.81 A
3.43 B
0.216 C
0.225 B
0.247 A

Means in columns and row (interaction) for each treatment followed by the same lowercase letter(s) are not significantly different at the 5% probability level.
Values in the parentheses are the percentage reduction (-) or increase (+) compared with the control (I1).
Means in each column (main effect of genotype) or row (main effect of drought level) for each treatment followed by the same uppercase letter(s) are not significantly
different at the 5% probability level.

in Borazjan under I1, in Shiraz under I 2 , and in Borazjan and

Ahvaz under I 3.
Total chlorophyll contents were increased the most in
Rashtkhar under I2 and I3 but were reduced the most in
Khaf1 under I2 and I3 (Table 5). Total chlorophyll contents
were highest in Ultan under I1 and in Rashtkhar under I2
and I3, but lowest in Borazjan under I1, I2 , and I3 (Table 5).
Among the accessions tested, chlorophyll a, chlorophyll b, and
total chlorophyll contents of Rashtkhar, Isfahan4, Borazjan,
Ahvaz, and Isfahan1 remained stable and higher than the
control under both I2 and I3 stresses. Drought stress reduced or
increased the chlorophyll contents of the genotypes, indicating
that the effects were accession and drought-level specific.
Decreased or unchanged chlorophyll levels under drought have
been reported in several species, depending on the duration and
severity of the drought (Kyparissis et al., 1995). Chlorophyll
is one of the most important chloroplast components for
photosynthesis because it harvests the light and produces
reducing powers; however, both chlorophyll a and chlorophyll
b are susceptible to drought and may affect plant yield and
quality. Higher chlorophyll content and stability have been
reported to be associated with the drought tolerance of
plants. Thus, the selection of genotypes based on increased
or stable chlorophyll content may prevent yield losses under
drought stress. In our study, chlorophyll a, chlorophyll b, and
total chlorophyll contents of Rashtkhar, Isfahan4, Borazjan,
Ahvaz, and Isfahan1 were higher than control, suggesting that
these accessions may be used under drought stress. Increased
chlorophyll content in these accessions under drought could
be attributed to increased RWC of the leaves, leaf soluble
carbohydrate contents, leaf and root proline, and carotenoid,
as indicated in Tables 2, 3, and 5, respectively. Reduction in
chlorophyll a, chlorophyll b, and total chlorophyll contents
of the other accessions could be due to reduction in the same
traits, as indicated in Tables 4 and 5.
In addition, reduction or loss of chlorophyll content under
drought could be due to pigment photooxidation (Anjum et

al., 2011) or loss of chloroplast membranes, distortion of the

lamellae vesiculation, and excessive swelling (Kaiser et al.,
1981). Damage to chloroplasts by active oxygen species under
drought might be another reason for chlorophyll reduction
(Smirnoff, 1993). In addition, reduction in chlorophyll
contents under drought could be due to their slower synthesis
and/or faster breakdown or dissociation of the chlorophyll
molecules (Majumdar et al., 1991).
Carotenoid Content
The highest carotenoid content was observed in Ultan under
I1 and in Ardestan under I2 , I3, and on average, whereas the
lowest was noted in Borazjan under I1 and in Shahreza under
I2 , I3, and on average (Table 5); however, the largest increase in
carotenoid content was detected in Rashtkhar under I2 and I3,
and the largest reduction was noted in Khaf2 under I2 and I3
(Table 5). Carotenoid plays a fundamental role and could help
plants to tolerate drought stress (Jaleel et al., 2009). Reduction
in carotenoid content in Ctenanthe setosa (Terzi and Kadioglu,
2006), cotton (Gossypium spp.; Parida et al., 2007), sunflower
(Poormohammad Kiani et al., 2008), and pearl millet
[Pennisetum glaucum (L.) R. Br.; Ashraf et al., 2001; Kholova
et al., 2011] has been reported; however, increases in carotenoid
content in maize cultivars (Mohammadkhani and Heidari,
2007) and sunflower (Nazarli et al., 2011) has also been
reported. In addition, Kalefetoglu Macar and Ekmekci (2009)
showed that carotenoid contents in chickpea were not affected
by drought. In contrast to these reports, our results showed
that the carotenoid content increased in some and decreased
in other accessions, and carotenoid contents were species and
drought-level specific.
Seed Yield
Khaf2 under I1 and Isfahan4 under I2 and I3 produced the
highest, while Isfahan1 under I1 and on average and Ultan
under I2 and I3 produced the lowest seed yields (Table 6);
however, Rashtkhar under I2 and Isfahan4 under I3 had the

Agronomy Journal Volume 106, Issue 1 2014

115

Table 6. Effects of drought level (D), genotype (G), and their interaction
on seed yield of sesame under three levels of drought (I1, I2 , and I3 =
55, 75, and 85% depletion of available soil water).
Seed yield
Accession

I1

I2

I3

Avg.

g
Rashtkhar 3.17 fgh
2.94 ghi (7)
2.80 hijk (12)
2.97 E
Khaf2
5.86 a
2.60 hijkl (55.6) 2.41 ijklmn (59)
3.62 B
Khaf1
5.28 ab 2.60 hijklm (51) 2.13 klmno (59.6) 3.34 CD
Isfahan4
4.32 cd
3.99 de (7.6)
3.88 def (10)
4.06 A
Shahreza
3.57 efg 2.21 jklmn (38.4)
1.35 p (62)
2.37 H
Borazjan
3.60 efg
3.27 fgh (9)
2.90 ghij (19)
3.25 D
Ahvaz
3.77 def
3.21 fgh (15)
2.84 hijk (24.7)
3.27 D
Kal
3.71 def 2.72 hijkl (26.5) 2.30 ijklmn (38)
2.91 E
Shiraz
3.19 fgh
2.74 hijkl (14)
2.18 klmn (31.6) 2.70 F
Markazi
5.84 a
3.75 def (36)
2.71 hijkl (53.5)
4.01 A
Ardestan
4.38 cd
3.22 fgh (26.4)
2.82 hijk (35.6)
3.48 C
Ultan
2.34 bc
1.45 op (38)
1.19 p (49)
2.52 G
Isfahan1
2.07 lmno 1.89 mnop (9)
1.76 nop (15)
1.90 I
Avg.
4.13 A
2.81 B
2.40 C
Means in columns and row (interaction) for each treatment followed by the
same lowercase letter(s) are not significantly different at the 5% probability level.
Values in the parentheses are the percentage reduction (-) or increase (+)
compared with the control (I1).
Means in each column (main effect of genotype) or row (main effect of drought
level) for each treatment followed by the same uppercase letter(s) are not significantly different at the 5% probability level.

Table 7. Rank of accessions from 1 to 13 (1 = highest) based on the

measured traits of leaf relative water content (RWC), leaf soluble
sugars (LSS), leaf proline (LP), root proline (RP), total chlorophyll (chl),
carotenoid (Carot), and seed yield (SY) under the I3 drought level (85%
depletion of available soil water).
Accession
Rashtkhar
Khaf2
Khaf1
Isfahan4
Shahreza
Borazjan
Ahvaz
Kal
Shiraz
Markazi
Ardestan
Ultan
Isfahan1

RWC
2
12
13
1
10
3
6
8
5
9
4
11
7

LSS
3
11
12
4
13
1
5
8
5
9
9
10
2

LP
3
11
13
2
12
1
4
7
5
9
8
10
6

RP
2
10
11
1
13
3
5
8
6
9
7
12
4

Total
chl
1
11
13
2
10
3
5
8
6
9
7
12
4

Carot
1
13
10
2
12
3
4
6
5
8
8
11
3

SY
2
11
12
1
13
4
5
8
6
10
7
9
3

Table 8. Correlation coefficients among measured traits in sesame under stress conditions (upper triangle, bold type) and normal conditions
(lower triangle).
Trait RWC
SY
Chl Carot LP
LSS
RP
SSI
RWC 1.00 0.36
0.21 0.36 0.05 0.57* 0.3 0.61*
SY
0.07 1.00
0.37 0.64* 0.5 0.18 0.68* 0.64*
Chl
0.07 0.65* 1.00 0.41 0.31 0.30 0.60* 0.24
Carot 0.05 0.72** 0.80** 1.00 0.52 0.02 0.47 0.72**
LP
0.18 0.27
0.55* 0.52 1.00 0.06 0.49 0.45
LSS
0.41 0.323 0.16
0.07 0.02 1.00 0.2
0.48
RP
0.07 0.71** 0.36
0.48 0.02 0.33 1.00 0.53*
SSI
0.46 0.58* 0.83** 0.63* 0.19 0.35 0.35
1.00
RWC, leaf relative water content; SY, seed yield; Chl, total chlorophyll; Carot,
carotenoid; LP, leaf proline; LSS, leaf soluble sugars; RP, root proline; SSI, Stress
Susceptibility Index.
* Significant at the 0.05 probability level.
** Significant at the 0.01 probability level.

116

smallest reduction in the seed yield (Table 6). In line with our
results, Mensah et al. (2006), Hassanzadeh et al. (2009a), and
Saeidi et al. (2012) also reported that the seed yield of sesame
genotypes was reduced under drought conditions, but with
differences among the genotypes.
Severe reduction in the seed yield of some accessions could be
due to reductions in RWC, leaf soluble carbohydrate contents,
leaf and root proline, chlorophyll, and carotenoid, as indicated
in Tables 2, 3, and 5, respectively. In other words, plants
invested more in stress protection mechanisms than in growth
when subjected to drought stress. Moreover, a reduction in
seed yield may also be due to less availability of nutrients along
with lower translocation of photosynthates from source to sink
areas under drought stress, as suggested by Manivannan et al.
(2007), Sankar et al. (2007), and Farooq et al. (2009).
Based on the overall results under I3 drought stress, we
were able to classify the accessions into tolerant (Isfahan4,
Rashtkhar, and Borazjan), moderately tolerant (Isfahan1,
Ahvaz, Shiraz, Ardestan, and Kal), and sensitive (Markezi,
Ultan, Khaf2, Khaf1, and Shahreza) groups. The accessions
were also ranked from 1 to 13 (1 = highest and 13 = lowest)
according to the measured traits under the I3 drought level
(Table 7). Isfahan4 had the first rank based on its RWC,
root proline content, and seed yield and second rank among
the other traits, except leaf soluble carbohydrate contents,
while Rashtkhar was ranked first based on its carotenoid and
chlorophyll contents and ranked second based on other traits
except soluble carbohydrates and leaf proline content.
A significant and positive correlation was also found
between seed yield and carotenoid (r = 0.64) and root proline
(r = 0.68) under drought (Table 8). These results again showed
the importance of carotenoid and root proline as markers for
resistance to drought stress. There were also significant and
positive correlations between soluble sugars and RWC (r =
0.57, p < 0.05) and root proline content and total chlorophyll
(r = 0.60, p < 0.05).
The SSI was calculated to assess the drought tolerance of
the sesame genotypes; the values of SSI for the accessions
were 0.19 (Isfahan4), 0.23 (Rashtkhar), 0.27 (Isfahan1), 0.35
(Borazjan), 0.45 (Ahvaz), 0.57 (Shiraz), (0.64) Ardestan,
0.68 (Kal), 0.89 (Ultan), 0.97 (Markezi), 1.06 (Khaf2), 1.08
(Khaf1), and 1.12 (Shahreza). Accessions estimated as drought
tolerant on the basis of lower yield reduction under drought
and other physiological characteristics generally had lower
SSI values. A lower SSI indicates a higher drought tolerance
(Fischer and Maurer, 1978). To explore relationships among
potentially useful traits to be used in breeding programs for
drought tolerance, the correlation coefficients between SSI
and the estimated parameters are given in Table 8. There was a
negative correlation between SSI and RWC (0.61, p < 0.05),
carotenoid (0.72, p < 0.01), and root proline (0.53, p < 0.05).
The negative correlation (more parameters, accessions with
lower sensitivity) between SSI and the mentioned parameters
suggest that these parameters may be used as selection criteria
for sesame genotypes in drought-stressed environments. These
results are in agreement with Gunes et al. (2008), Sairam and
Saxena (2000), and Talebi et al. (2013). In general agreement
with our results, Talebi et al. (2013) reported a significant
correlation between SSI and RWC in 35 chickpea genotypes
Agronomy Journal Volume 106, Issue 1 2014

and concluded that RWC may be used for selecting genotypes

of this species for drought tolerance. The RWC was also
reported to be correlated with chlorophyll, proline, and
soluble sugar contents in Aeluropus lagopoides (Mohsenzadeh
et al., 2006). Moreover, the observed correlations among
physiological parameters showed that most of traits were closely
correlated with each other, and these correlations confirm the
relative importance of these physiological parameters that may
be used as selection criteria for sesame genotypes in droughtstressed environments (Table 8).
Conclusions
The measured traits were accession and drought-level
dependent and discriminated the accessions based on their
drought tolerance. Based on the overall results under I3
drought stress, Isfahan4 was the most drought-tolerant
accession, followed by Rashtkhar, Borazjan, Isfahan1, Ahvaz,
Shiraz, Ardestan, Kal, Markezi, Ultan, Khaf2, Khaf1, and
Shahreza, in that order. The results also showed that Khaf2
was the best accession under normal conditions and Isfahan4
was the best accession under drought conditions. In addition,
based on correlation coefficients for the traits with seed
yield, root proline content and carotenoid content were the
superior markers for drought tolerance in sesame. This study
showed that accession differences in drought tolerance could
be attributed to the ability of sesame plants to acclimate
physiochemicals under drought conditions, and metabolites
could be a useful tool for depicting water-deficit tolerance of
sesame genotypes, which could be useful to plant breeders for
selecting and developing drought-tolerant cultivars.
Acknowledgments
This work was financially supported by the Isfahan University of
Technology.
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