Communications in Soil Science and Plant Analysis, 36: 24052419, 2005

Copyright # Taylor & Francis, Inc.

ISSN 0010-3624 print/1532-2416 online
DOI: 10.1080/00103620500253324

Salt Tolerance of Sesame Genotypes at

Germination, Vegetative, and
Maturity Stages
R. Ramrez
Centro Nacional de Investigaciones Agropecuarias (CENIAP),
Maracay, Venezuela

D. Gutierrez
Facultad de Agronoma, Universidad Nacional Experimental Francisco
de Miranda, Coro, Edo., Falcon, Venezuela

R. Villafane
Facultad de Agronoma, Universidad Central de Venezuela,
Maracay, Venezuela

J. I. Lizaso
Department of Agronomy, University of Florida,
Gainesville, Florida, USA

Abstract: Salinity and moisture stress are main limiting factors of agricultural
development on arid and semi-arid lands. The objective of this study was to evaluate
the tolerance of sesame (Sesamun indicum L.) genotypes to salinity. Salinity treatments
in rooting media were 0.5, 2.0, 3.5, 5.0, and 7.0 dS m21 using calcium chloride (CaCl2).
Germination of 50 genotypes was evaluated. Twenty seeds of each cultivar were set in
paper towels and moistened with the saline solutions for 15 days. Five germinationselected genotypes were seeded in 10-kg pots, and soil salinity was adjusted to the
same treatments. Highly significant effects were found for dry matter at 45 (DM 45)
days after planting (DAP), at 90 DAP (DM 90), and leaf area (LA). Growth differences
among genotypes were only observed for DM 90. These data suggest that sesame

Received 7 April 2004, Accepted 28 February 2005

Address correspondence to J. I. Lizaso, Department of Agronomy, University of
Florida, Gainesville, FL 32611, USA. E-mail: jlizaso@ifas.ufl.edu
2405

2406

R. Ramrez et al.

tolerance to CaCl2 salinity improved through the growing season and may be
genetically controlled.
Keywords: Nutrient uptake, root length, leaf area, calcium chloride, salinity

INTRODUCTION
A soil is affected by salts when any ion or salt, together or separately, affect soil
properties and, consequently, economic production of crops (United States
Salinity Laboratory Staff 1954). Salinity, along with water deficit, has a
strong influence on agricultural development of arid and semiarid regions.
In the past, farmers grew low intensive sesame and other crops in the flat
lands of Falcon State, in western Venezuela. These cropping systems evolved
into horticultural crops, which under intensive use of irrigation and fertilization
eventually led to soil salinization. A preliminary soil analysis study found
in the topsoil Ca concentrations from 22 to 67 mmol L21, Na concentrations
from 6 to 14 mmol L21, and Cl concentrations from 1.8 to 74 mmol L21.
Salt-tolerant sesame cultivars could be an option for these saline soils.
Plants have variable tolerance to salinity depending on species. There are
a wide spectrum of responses among cultivated species, ranging from very
sensitive to relatively tolerant (Maas and Hoffman 1977). These differences
have been attributed to the ability of certain species to adapt to osmotic
changes (Ayers and Wescot 1987). Genetic differences have also been
found within species such as alfalfa during germination, emergence, and
establishment periods (McKimmie and Dobrenz 1987).
Stress generated by salinity inhibits plant growth during the entire cycle,
but tolerance of plants to salinity varies with the growth stage. Alfalfa
tolerance to soil salinity at germination was not evident at seedling and at
physiological maturity (Rumbaugh et al. 1993). Lettuce has been found to
be more sensitive to salinity during early growth and flowering (Shannon
et al. 1983), and sugar beet exhibits higher sensitivity during germination
(Bernstein and Hayward 1958). In tomato, salinity not only reduces germination but it can also delay emergence of plants (Cuartero and Fernandez
1999).
Rice at seedling and reproductive stages showed greater sensitivity than at
germination and grain-filling stages; however, not all cultivars had the same
response (Lutts et al. 1995). There is no evidence that tolerance to salinity
during a specific growth stages is indicative of tolerance in other stages.
Sesame is generally cultivated in semiarid regions where soil salts are a
limiting factor. It has been reported that sesame is sensitive to soil salinity
(Yousif et al. 1972; Cerda et al. 1997) mostly due to the presence of
calcium chloride (Nassery et al. 1979).
The objective of this research was to evaluate the tolerance of different
sesame cultivars to salinity induced by the presence of calcium chloride in

Salt Tolerance of Sesame Genotypes

2407

the rooting environment. Future efforts will evaluate responses to sodium

chloride as a previous step to field tests.

MATERIALS AND METHODS

Germination Experiment
Seeds used in this research were supplied by the National Center of Agricultural
Research (CENIAP) from cultivars of a wide geographic origin. Fifty genotypes
were evaluated: 12 from Africa, 6 from India, 5 from USA, 3 from Venezuela,
3 from Sudan, 3 from China, 2 from Mexico, 2 from Argentina, 2 from Japan,
2 from Greece, and 1 from each of the following countries: Italy, Russia,
Colombia, Nicaragua, Peru, Guatemala, Israel, Syria, Korea, and Brazil.
Salinity treatments were prepared by using 25% strength Hoagland and
Arnon II nutrient solution (Hoagland and Arnon 1950). Reagent grade
calcium chloride was added to obtain saline solutions of four concentrations:
0.5 (control), 3.0, 6.0, and 9.0 dS m21. Twenty individually selected seeds
(i.e., rounded seeds longer that 3 mm) of each genotype, treated with Captan
and Vitavax, were placed between two paper germination towels in
21
25 cm2 aluminum trays. Each tray was supplied with the corresponding
saline solution and the amount of water required for germination. Trays were
placed in germination chambers with temperatures between 21 and 258C and
about 90% relative humidity. A completely randomized design with three replications, in a factorial arrangement of genotype by salinity treatment, was used.
Germination was evaluated 5 days after the experiment was initiated.
Some seedlings exhibited abnormal growth. Germinated normal plants
(GNP) were considered only those plants with radicles at least 5 mm long
and well-developed hypocotyls and cotyledonary leaves.
A toxicity index (TI) was calculated to determine relative tolerance to
salinity:
TI

T  Ts
T

where T and Ts are percentage of germination of control and salinity treatments. TI for dry matter, leaf area, and seed yield was calculated by
following the same procedure.

Pot Experiment
To study the effect of calcium chloride-induced salinity on sesame development, an experiment was conducted in a greenhouse by using a sandy loam
soil, pH 6.4 (1 : 2.5 soil to water ratio), electric conductivity 0.51 dS m21
(saturated soil paste), and rich in Ca (1110 mg kg21).

2408

R. Ramrez et al.

Four salt-tolerant cultivars and one salt-sensitive cultivar in the germination experiment were chosen. Tolerant genotypes were Single Stem (G22),
Ajimo Star S-13 (G41), White China (G42), and White Sindos (G12).
Sensitive germplasm was Teras (G5).
To determine the required amount of salt in the solution to produce the
required soil salinity of each treatment, a soil incubation experiment was set
up by using analytical grade CaCl2 with saline solutions in the range of
0.5 9.0 dS m21. A regression analysis was used to relate required amount
of salt in the solution with 0.5, 2.0, 3.5, 5.0, and 7.0 dS m21 salinity in the soil.
Ten kilograms of air-dried soil passing through a 3-mm sieve was placed
in 12-L plastic pots. Each pot was supplied with 150 mg N kg21 soil,
100 mg P kg21 soil, and 40 mg K kg21 soil in the form of urea, triple super
phosphate, and potassium sulfate, respectively. Ten seeds of each cultivar
were planted in each pot and thinned 10 days after germination to leave the
four best plants.
Deionized water was used to keep the soil moisture between 40 and 80%
of available water. A soil water retention curve was established for each
treatment. To reduce salt accumulation in the top of soil, watering was done
alternatively at the top of the soil or the base of the pot through a 2.5-cm
diameter plastic tube placed before planting in the center of the pot.
Salinity (5) and genotype (5) treatments were distributed in a completely
randomized design with three replications and a factorial arraignment of treatments. Two plants were randomly harvested from each pot 45 days after
planting. They were washed with deionized water and oven dried for 48 h
at 808C to determine dry matter (DM). Plant tissue was ground by using a
1.0-mm sieve and digested with sulfuric acid and hydrogen peroxide
(Thomas et al. 1967). Extracted solutions were analyzed for potassium (K),
calcium (Ca), and magnesium (Mg) by atomic absorption spectroscopy and
for Cl by colorimetry.
Leaf area of a complete plant per pot was measured with a Li-Cor 3000 A
meter, 45 days after planting (DAP). Ninety DAP, the two remaining plants of
each pot were harvested; tops and seeds were separated and dried as described
before.

RESULTS AND DISCUSSION

Germination Experiment
Analysis of variance for percentage of germinated normal plants (GNP) was
highly significant for salinity treatments, genotype, and salinity by genotype
interaction. These results indicate that the germination of sesame seeds
depends on the individual performance of each genetic material, as well as
on the level of salinity of the nutrient solution.

Salt Tolerance of Sesame Genotypes

2409

The values of GNP in the control (0.50 dS m21) ranged from 67% for
genotype G24 up to a maximum of 99% for G6 (Table 1). GNP of 14 of
the 50 genotypes was 95% or higher, 15 genotypes within 90 93%, 16
80 88%, and only 5 genotypes felt below 70%. These differences in GNP
between genotypes are due to seed quality.
The first increase in salinity up to 3.0 dS m21 in the nutrient solution
reduced the overall seed germination. Only nine genotypes had 95 or more
GNP, and 14 had 90 94 GNP. However, the germination percentage of
14 genotypes increased when treated with a low salinity stress of 3.0 dS m21,
suggesting that such a level of salinity in the soil could stimulate germination.
As salinity increased to 6.0 dS m21, only eight genotypes reached germination
levels above 90%. With the application of the highest dose of calcium chloride,
considerable differences were found between genotypes. The most outstanding
genotypes were G42, G41, G40, G35, G12, G22, 37, and 38, which had
62 80% germination. Other cultivars such as G3, G4, G5, G6, and G13
exhibited germination values below 10%.
These results indicated high variability in tolerance to salinity during germination among the sesame genotypes studied. Reports in the literature reveal
similar results for different species (McKimmie and Dobrenz 1987; Carlson
et al. 1983; Murillo-Amador et al. 2001; Daniells et al. 2001; Greenway
and Munns 1980; Maas 1986).
A toxicity index (TI) was calculated to ascertain the relative salinity
tolerance of genotypes. TI values around zero indicate similar germination to
the control. Negative TI values reveal higher germination under salinity, suggesting a promoting effect of the calcium chloride treatment on germination.
Values higher than zero occur when germination was reduced because of the
toxicity of calcium chloride. TI for all cultivars progressively increased with
higher salinity of the nutrient solution (Table 1). With the lowest level of
salinity (3.0 dS m21) TI ranged between 20.35 and 0.18. These values illustrate
the existence of a wide range of variation in the response of the genetic
materials to salinity. TI was less than zero in 14 of the 50 genotypes, which
is an indication that at low levels of soil salinity seed germination of some
genotypes could be stimulated. TI was equal to zero in 12 genotypes, indicating
that salinity did not affect germination of such genotypes.
When the level of salinity was increased to 6.0 dS m21, the range of TI
was between 20.1 and 0.48, but only five genotypes showed TI values
lower than zero. Three of these five genotypes exhibited a very low GNP
when assayed with the nutrient solution that did not contain calcium
chloride (67% for G24, 73% for G32, and 80% for G38). Germination of
these three genotypes was stimulated when treated with 3.0 and 6.0 dS m21.
Toxicity of the 9.0 dS m21 saline solution was severe for all tested
genotypes. TI values were in all cases greater than that found for saline
solutions of lower electric conductivity (Table 1). Only G42 consistently
had low TI values of 0.0 for 3.0 dS m21, 20.02 for 6.0 dS m21, and 0.09 for
9.0 dS m21.

R. Ramrez et al.

2410

Table 1. Germination percentage of sesame seeds under salinity stress and toxicity
index. Seeds were germinated embedded in solutions with electrical conductivity of
0.5, 3.0, 6.0, or 9.0 dS m21
Electrical conductivity (dS m21)
Germination (%)
Genotypes
6
9
12
5
14
46
3
39
40
25
29
10
22
50
19
4
13
26
35
37
41
18
44
34
15
16
17
7
33
42
43
1
2
49
20
23
30

Toxicity index

0.5

3.0

6.0

9.0

3.0

6.0

9.0

99.33
96.67
96.67
96.67
96.67
96.67
95.00
95.00
95.00
95.00
95.00
95.00
95.00
95.00
93.33
93.33
93.33
93.33
93.33
93.33
91.67
91.67
91.67
91.67
90.00
90.00
90.00
90.00
90.00
88.33
88.33
86.67
86.67
86.67
86.67
85.00
85.00

88.33
86.67
90.00
96.67
100.00
96.67
81.65
95.00
95.00
95.00
98.33
91.67
93.33
88.33
88.33
88.33
91.67
93.33
90.00
93.33
91.67
91.67
86.67
96.67
95.00
86.67
85.00
88.33
90.00
88.33
86.67
80.00
83.33
85.00
90.00
81.67
85.00

68.33
78.33
90.00
53.33
91.67
93.33
65.00
83.33
86.67
86.67
83.33
80.00
93.33
50.00
65.00
60.00
80.00
78.33
90.00
83.33
91.67
88.33
73.33
80.00
83.33
75.00
76.67
91.67
70.00
90.00
65.00
45.00
76.67
71.67
71.67
71.67
60.00

6.67
56.67
68.38
8.33
28.33
31.67
0.10
56.67
73.33
48.33
30.00
33.33
61.67
13.33
45.00
6.67
1.67
46.67
70.00
65.00
73.33
38.33
38.33
46.67
60.00
36.67
50.00
48.33
21.67
80.00
31.67
15.00
13.33
26.67
50.00
30.00
26.67

0.11
0.10
0.07
0.00
20.03
0.00
0.14
0.00
0.00
0.00
20.04
0.04
0.02
0.07
0.05
0.05
0.02
0.00
0.04
0.00
0.00
0.00
0.05
20.05
20.06
0.04
0.06
0.02
0.00
0.00
0.02
0.08
0.04
0.02
20.04
0.04
0.00

0.31
0.19
0.07
0.45
0.05
0.03
0.32
0.12
0.09
0.09
0.12
0.16
0.02
0.47
0.30
0.36
0.14
0.16
0.04
0.11
0.00
0.04
0.20
0.13
0.07
0.17
0.15
20.02
0.22
20.02
0.26
0.48
0.12
0.17
0.17
0.16
0.29

0.93
0.41
0.29
0.91
0.71
0.67
1.00
0.40
0.23
0.49
0.68
0.65
0.35
0.86
0.52
0.93
0.98
0.50
0.25
0.30
0.20
0.58
0.58
0.49
0.33
0.59
0.44
0.46
0.76
0.09
0.64
0.83
0.85
0.69
0.42
0.65
0.69

(continued)

Salt Tolerance of Sesame Genotypes

2411

Table 1. Continued
Electrical conductivity (dS m21)
Germination (%)
Genotypes
36
27
8
45
28
48
38
31
11
47
32
21
24

Toxicity index

0.5

3.0

6.0

9.0

3.0

6.0

9.0

85.00
85.00
85.00
81.67
81.67
81.67
80.00
80.00
78.33
78.33
73.33
71.67
66.67

93.33
76.67
70.00
83.33
86.67
71.67
81.67
85.00
81.67
76.67
90.00
80.00
90.00

76.67
71.67
76.67
63.33
71.67
48.33
83.33
76.67
68.33
56.67
80.00
70.00
73.33

33.33
35.00
43.33
13.33
26.67
21.67
61.67
31.67
43.33
36.67
41.67
30.00
35.00

20.10
0.10
0.18
20.02
20.06
0.12
20.02
20.06
20.04
0.02
20.23
20.12
20.35

0.10
0.16
0.10
0.22
0.12
0.41
20.04
0.04
0.13
0.28
20.09
0.02
20.10

0.61
0.59
0.49
0.84
0.67
0.73
0.23
0.60
0.45
0.53
0.43
0.58
0.48

The information from the germination experiment suggests that

tolerance of sesame to salinity during germination is apparently genetically
controlled.

Pot Experiment
Four of the most salt-tolerant genotypes (G22, G41, G42, G12) in the germination tests and one of the most susceptible (G5) were chosen for the pot experiments. Results of the analysis of variance for dry matter at 45 days (DM45)
and at 90 days after planting (DM90), seed weight (SW), and leaf area
(LA), showed highly significant effects (p . 0.01) for salinity and
germplasm treatments. These results suggest that differences between means
are related to the dose of salt applied and the specific calcium chloride
tolerance of individual genotypes. The significant interaction genotypesalinity was only evident for plants sampled at 45 days after planting. In all
cases, the salt-susceptible cultivar during germination (G5) was also the
most sensitive in growth and seed yield.
Average of DM45 progressively decreased as soil salinity increased
(Table 2) until it reached 34.7% of control when salinity was at the
highest level of 7.0 dS m21. However, DM was not affected by 2.0 dS m21
salinity level. DM of G12 was significantly higher than that of other
genotypes, indicating a higher tolerance to salinity at 45 DAP. G12

R. Ramrez et al.

2412

Table 2. Sesame dry matter at 45 and 90 days after planting (DAP) under the effect of
soil salinity
Dry matter (g plant21), electrical conductivity (dS m21)
Genotypes

0.5

2.0

3.5

5.0

7.0

Average

45 DAP

22
41
42
12
5
Average

4.78 a
4.86 a
4.02 b
5.04 a
3.93 b
4.53 a

3.67c
3.61 c
4.29 ab
5.02 a
4.28 b
4.17 ab

3.10 b
3.42 b
2.90 b
5.55 a
2.84 c
3.56 b

2.84
2.86
1.97
3.72
1.83
2.64

b
b
c
a
c
c

2.04 a
1.47 a
1.46 a
1.40 a
1.47 a
1.57 c

3.29 b
3.24 b
3.93 b
4.35 a
2.87 b

90 DAP

22
41
42
12
5
Average

4.84
7.18
5.63
5.45
4.35
5.49 a

5.49
6.85
6.12
5.27
4.49
4.64 a

5.89
6.41
5.17
5.10
4.36
5.44 a

4.85
5.74
5.51
5.52
2.86
4.90 a

3.50
3.55
3.78
3.18
2.11
3.22 b

4.91 b
5.94 a
5.24 ab
4.90 b
3.69 c

Means in the same column followed by different letters are significantly different at
p , 0.05 by Tukeys test.

consistently accumulated more biomass up to a salinity level of 5.0 dS m21.

G22 and G41 exhibited high capacity to produce DM when no calcium
chloride was applied to the soils. However, when these two genotypes
were grown in saline soils, plant growth was reduced. No differences in
DM were found for genetic materials at the highest level of salinity
(Table 2).
Comparing the biomass accumulated by the genotypes tested, we found
different responses at 45 and 90 DAP (Table 2). At 90 DAP, no significant
differences were found among genotypes at electrical conductivities of 0.5,
2.0, 3.5, and 5.0 dS m21; however, early in the season at 45 DAP, such differences were considerable. It is known that species susceptibility to salinity
varies with stage of development (Lunin et al. 1963; Lutts et al. 1995;
Abbar and Neue 1987; Shannon et al. 1983; Francois 1984).
On average, the most outstanding genotypes at 90 DAP were G41 and
G42, whereas at 45 DAP, G12 showed the highest level of DM production.
Only G5 exhibited a consistent low DM production at both evaluation
periods (Table 2).
TI estimates the relative tolerance of genotypes to different salinity levels
in the soil, by standardizing the responses to those evaluated under nonstress
conditions. Higher TI values correspond to decreased growth and seed production due to the lethal effect of salinity. On the other hand, negative TI
values indicate an enhanced plant growth and yield in response to moderate
levels of soil salinity (Figure 1).

Salt Tolerance of Sesame Genotypes

2413

Figure 1. Toxicity index for five sesame genotypes. DM 45 and leaf area were
sampled 45 days after planting. DM 90 and seed yield were sampled 90 days after
planting.

TI calculated from DM was different at each evaluation time. At 45 DAP,

the lowest level of salinity did not affect G5, G12, and G42. Genotype G12
performed better when salinity was increased up to 5.0 dS m21 (Figure 1).
This cultivar produced more DM in a soil without salt as well as in a soil
treated with salts. The TI curve of G12 indicated a better capacity to
tolerate salinity up to 5.0 dS m21 during early growth than the other
evaluated genotypes (Figure 1). TI values calculated from DM at 90 DAP
showed that growth of G12 and G41 was slightly reduced by the lowest

R. Ramrez et al.

2414

level of salinity. However, DM production of G5, G22, and G42 was stimulated (Figure 1). The curve for G22 showed that growth was 22% larger
than the control at a salinity level of 3.5 dS m21 and that growth was still
similar to the control at a salinity level as high as 5.0 dS m21.
By comparing DM evaluated at 45 and at 90 DAP (Figure 1), it becomes
apparent that TI values for DM90 were considerably lower than the ones found
for DM45, indicating that the effect of salinity on plant growth was less
harmful during the reproductive stage than during the vegetative stage. This
suggested that sesame plants adapted to soil salinity through the growing
season and that some genotypes adapted earlier than others.
Salinity levels above 2.0 dS m21 significantly reduced average seed yield
(Table 3). The percentage of reduction in seed production was 16%
at 3.5 dS m21, 41% at 5.0 dS m21, and 71% at 7.0 dS m21. No significant
differences were found between G22, G41, G42, and G12 means;
however, G5 seed yield was about 30% lower than the seed yield of the
other cultivars.
Shoot biomass production (DM90) did not follow the same trend as seed
production (Figure 2). Salinity treatments ranging from 0.5 to 5.0 dS m21 did
not produce significant differences on DM90; however, seed production was
highly affected by salinity levels above 3.5 dS m21. DM45 responses to
salinity followed a similar trend as seed production. This relation requires confirmation in future investigations because it may be adopted as a selection
criterion for tolerance to salinity.
TI values calculated from seed yield exhibited similar trends for all
genotypes (Figure 1). The only difference was G12, which had reduced
yield and thus high TI at low salinity levels (2.0 and 3.5 dS m21). These
results suggested that seed yield is more sensitive to salinity than DM.
TI curves indicated differences to tolerate the effects of soil salinity
induced by calcium chloride among genotypes. It also indicated a better
growth at higher salinity levels later in the season, suggesting a process

Table 3.

Sesame seed yield as affected by soil salinity

Seed yield (g plant21), electrical conductivity (dS m21)

Genotypes
22
41
42
12
5
Average

0.5

2.0

3.5

5.0

7.0

3.02
3.42
2.93
3.06
2.14
2.91 a

3.06
3.61
3.08
2.53
2.24
2.81 ab

2.47
2.90
2.73
2.17
1.97
2.45 b

1.62
2.11
1.90
1.91
1.06
1.72 c

1.11
0.87
0.75
0.95
0.60
0.85 d

Average
2.26
2.49
2.27
2.13
1.60

a
a
a
a
b

Means within genotypes or electrical conductivity followed by different letters are

significantly different at p , 0.05 by Tukeys test.

Salt Tolerance of Sesame Genotypes

2415

Figure 2. Effect of soil salinity on sesame dry matter, seed yield, and leaf area. Each
value is the average of five cultivars.

of adaptation. Unfortunately, the adaptation of growth to soil salinity

was not paralleled by a similar adaptation of seed yield to soil salinity
(Figure 1).
Leaf area was evaluated at midseason (45 DAP) to explore the effect of
soil salinity on expansion growth. G12 and G42 (Table 4) had higher
average values (454 and 420 cm2), whereas G5 had the lowest LA
(310 cm2). On average, leaf area was not affected by the lowest level of
salinity but was drastically reduced by higher levels of salinity. Average
leaf area in nonsaline soil was 453 cm2 and 459 cm2 at 2.0 dS m21, but at
higher levels of salinity leaf area exhibited significant decreases of 14% at
3.5 dS m21, 21% at 5.0 dS m21, and 37% at 7.0 dS m21. No significant differences were found between leaf area values of genotypes in nonsaline soil as
well as in the most saline soil. Leaf area of G12 stood out as being the least
affected by salinity compared with the other genotypes. TI values were
below zero up to 3.5 dS m21 level expanding 40% more leaf surface than
the control at low salinity (2.0 dS m21, Figure 1). This response may be attributed to the stimulation of leaf expansion due to the presence of low levels of
salts in the soil.
Leaf area is an indicator of plant photosynthetic capacity. A greater leaf
area per plant will correspond with greater assimilate production and possibly

R. Ramrez et al.

2416
Table 4.

Sesame leaf area under soil salinity stress at 45 days after planting
Leaf area (cm2 plant21), electrical conductivity (dS m21)

Genotypes
22
41
42
12
5
Average

0.5
459
492
515
438
362
453 a

2.0
345
405
547
616
380
459

c
b
ab
a
b
ab

3.5
434
360
341
549
271
391

5.0
b
c
c
a
c
b

345
373
392
420
267
359

7.0
b
c
a
a
c
c

321
279
306
248
268
285 d

Average
378
381
420
454
310

ab
ab
a
a
b

Means in the same column followed by different letters are significantly different at
p , 0.05 by Tukeys test.

greater biomass. If this premise holds, leaf area should be related with dry
matter production. When genotypes leaf area is compared with DM45 and
seed weight, a similar trend can be observed (Figure 2). However, the trend
was different when the leaf area is compared with DM90 and total DM
weight (DM seed weight). Such response of leaf area and DM at plant
maturity may be attributed to different partitioning of photosynthates among
roots, shoots, and seeds, or maybe to differences in carbon use for respiration
on each genotype. Leaf area decrease due to soil salinity has been reported
previously (Khan et al. 1998; Lutts et al. 1995). The toxic effect of salinity
on plant growth may be partially explained by the reduction of leaf area
that causes lower photosynthesis (Wahid et al. 1997). This lower photosynthesis may occur because of decreased gas exchange through stomata, with
a reduction in CO2 availability for carboxilation (Sharma 1996; Li-Wen
et al. 1997).
Calcium concentration in leaves at 45 DAP increased with the level
of CaCl2 applied to the soil. Similar trend was observed for Cl (Figure 3),
but K concentration curves showed a decreasing trend as Ca and Cl
levels in plant tissue increased. Little change was observed in Mg
concentration.
The decrease of DM90 was closely related with an increase of Ca and
Cl concentration and with K concentration decrease (Figure 3). This
harmful effect of Ca and Cl is possibly due to the accumulation of these
ions in leaf tissue to the detriment of K status. Tolerance to salinity has
been associated with the ability of more vigorous plants to maintain
lower concentrations of ions, especially in leaves, and to sustain higher
growth rates, thus creating a dilution effect (McKimmie and Dobrenz
1987).
The information reported here indicates that there were no salinityinduced differences among genotypes in DM45, but such differences were

Salt Tolerance of Sesame Genotypes

2417

Figure 3. Dry matter at 90 DAP and nutrient concentration in leaves at 45 DAP of

five sesame genotypes.

considerable in DM90. It also suggested that plants adapt to soil salinity

throughout the growing season. However, responses to soil salinity were
variable across genotypes. Sesame tolerance to soil salinity seems to be
related to plant stage of growth and is under genetic control.

2418

R. Ramrez et al.

ACKNOWLEDGMENTS
We thank Dr. Rosemary Warnock for her assistance with the English version
of this paper.

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