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Physiological Mechanism of Salt

Tolerance in Suaeda fruticosa Collected
from High Saline Fields of Khyber
Pukhtoon-Khwa, Pakistan
a b
Sami Ullah & Asghari Bano
a
Department of Botany, Bacha Khan University, Charsadda, Pakistan
b
Department of Plant Sciences, Quaid-i-Azam University, Islamabad,
Pakistan
Accepted author version posted online: 13 May 2015.Published
Click for updates online: 13 May 2015.
To cite this article: Sami Ullah & Asghari Bano (2015) Physiological Mechanism of Salt
Tolerance in Suaeda fruticosa Collected from High Saline Fields of Khyber Pukhtoon-
Khwa, Pakistan, Communications in Soil Science and Plant Analysis, 46:10, 1212-1228, DOI:
10.1080/00103624.2015.1033532
To link to this article: http://dx.doi.org/10.1080/00103624.2015.1033532
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Communications in Soil Science and Plant Analysis, 46:1212–1228, 2015
Copyright © Taylor & Francis Group, LLC
ISSN: 0010-3624 print / 1532-2416 online
DOI: 10.1080/00103624.2015.1033532
Physiological Mechanism of Salt Tolerance

in Suaeda fruticosa Collected from High Saline
Fields of Khyber Pukhtoon-Khwa, Pakistan
SAMI ULLAH1 AND ASGHARI BANO2

1
Department of Botany, Bacha Khan University, Charsadda, Pakistan
2
Department of Plant Sciences, Quaid-i-Azam University, Islamabad, Pakistan
The present investigation was aimed to evaluate the physiological mechanism of adap-
tation of salt tolerance in Suaeda fruticosa at various phenological stages with varying
levels of soil salinity of the selected districts via determining ion accumulation, growth
response, osmolyte accumulation, and antioxidant enzyme activities. Randomized com-
plete block design in the selected districts was used with plots sized 6 m × 9 m2 , having
5 rows that were 2 m long and 30 cm apart, in triplicate. Suaeda fruticosa maintained
its chlorophyll a/b ratio even at the greater electrical conductivity of rhizospheric soil
at Peshawar, which is indicative of its better adaptability. For Suaeda fruticosa it has
been concluded that the species collected from District Peshawar exhibited a significant
increase in contents of sugar, proline, and protein as compared to the species collected
from other districts, whereas superoxide dismutase, peroxidase, and carotenoid content
was maximum for the species collected from District Mardan.
Keywords Halophyte, salinity, Suaeda fruticosa
Introduction
Halophytes are considered to be those plants that have the ability to tolerate low water
potential caused by salinity, so they can survive under saline soils (Munns and Tester
2008). About 410 halophytic species from fifty-eight families has been reported by Khan
and Qaiser (2006) in Pakistan, indicating great diversity as well as great opportunity to find
cash crop halophytes (Khan 2003). The decreased in chlorophyll content under stress is a
commonly reported phenomenon and this may be due to different reasons, such as mem-
brane deterioration (Ashraf and Bhatti 2000). The reduction in leaf chlorophyll content
under sodium chloride (NaCl) stress has been attributed to the destruction of chlorophyll
pigments and the instability of the pigment protein complex (Levitt 1980). The hyperac-
cumulation of proline under stress conditions in many plant species has been correlated
with stress tolerance, and its concentration has been known to be greater in stress-tolerant
species than in stress-sensitive species (Ashraf and Foolad 2007). Accumulation of osmo-
lites such as sugars in different parts of plants is improved in response to a varying level of
environmental stress (Wang, Quebedeaux, and Stutte 1996; Prado et al. 2000; Gill et al.
2001). In the case of salt (Gill and Singh 1985) and water stress (Prado et al. 2000;
Received 19 May 2013; accepted 5 October 2014.

Address correspondence to Sami Ullah, Department of Botany, Bacha Khan University,
Charsadda, Pakistan. E-mail: sami_jan69@yahoo.com
1212
Soil Salinity and Halophytes 1213
Siddique, Hamid, and Islam 2000), adaptation to the stresses has been attributed to the
stress-induced increase in sugar accumulation in plants.
Salt stress, like other abiotic stresses, can also lead to oxidative stress through the
increase in reactive oxygen species (ROS), such as superoxide (O2 − ), hydrogen peroxide
(H2 O2 ), and hydroxyl radicals (OH), which are highly reactive and may cause cellular
damage through oxidation of lipids, proteins, and nucleic acids (Apel and Hirt 2004).
To minimize the effects of oxidative stress, plant cells have evolved a complex antioxidant
system, which is composed of low-molecular-mass antioxidants (glutathione, ascorbate,
and carotenoids) as well as ROS-scavenging enzymes, such as superoxide dismutase
(SOD), catalase (CAT), ascorbate peroxidase (APX), guaiacol peroxidase (GPX), and glu-
tathione reductase (GR) (Alscher, Donahue, and Cramer 1997; Apel and Hirt, 2004). It has
been reported that antioxidant enzyme activities decrease in plant cells under salt stress
(Shalata et al. 2001; Khedr et al. 2003; Mishra and Das 2003) and that antioxidant enzyme
activities increase in the presence of proline (Chen and Dickman 2005). Some researchers
also suggest that salt stress leads not to a decrease in SOD activity in salt-sensitive plants
but to an increase in salt-tolerant ones (Shalata and Tal 1998; Sreenivasulu et al. 2000).
The present investigation was aimed to evaluate the physiological mechanism of adap-
tation of salt tolerance in Suaeda fruticosa at germination, vegetative, and flowering stages
with varying levels of soil salinity (EC 4.3–8.24) at four districts of Khyber Pukhtoon-
khwa, Pakistan. The responses of Suaeda fruticosa to the salt stress have been evaluated on
the basis of selectivity of ion uptake, osmoregulation, and antioxidants enzyme activities.
Materials and Methods
Experimental Design
The experiment was conducted under natural conditions in the saline soil of District
Mardan (EC 4.3–5.5), District Charsadda (EC 5.6–6.5), District Nowshera (EC 8.2–9.02),
and District Peshawar (EC 6.8–8.0). A randomized complete block design was used, with
plot sizes of 6 m × 9 m2 , 5 rows that were 2 m long and 30 cm apart, in triplicate. Seeds
were obtained from National Agricultural Research Center, Islamabad. Seeds were sur-
face sterilized, followed by sterilization with 10 percent chlorox for 3 min, washed with
95 percent ethanol for 3 min with constant shaking, and finally washed with sterile water.
Collection of Halophyte Species

In the present study Suaeda fruticosa at various phenological stages were collected from
their natural habitat of high saline fields of the selected districts. The study was conducted
for 2 consecutive years, 2010 and 2011.
Physicochemical Analysis of Rhizospheric Soil
Soil pH and Electrical Conductivity (EC)

The rhizospheric soil at a depth of 6 inches from the upper surface has been collected
from five places in the selected areas and made into a composite soil sample. The pH of
rhizospheric soil was measured by preparing a 1:1 (soil/water) suspension (McKeague
1978; McLean 1982). An air-dried soil sample (10 g) was mixed in 10 ml distilled water
and stirred for 1 h on a magnetic stirrer for homogenous mixing. Then the suspension was
1214 S. Ullah and A. Bano
filtered with Whatman no. 42 filter paper. The pH of filtrate was determined with a pH
meter, while the electrical conductivity meter recorded the EC of the extract. Readings
were measured in microsiemens per centimeter (µS/cm).
Nutrient Analysis of Rhizospheric Soil

The rhizospheric soil was analyzed for macronutrients Na+ , Ca+2 , Mg+2 , and K+ fol-
lowing the ammonium bicarbonate–DTPA method developed by Soltanpour and Schwab
(1977).
Extraction Solution Preparation

The 0.005 M solution was obtained by adding 1.97 g DTPA to 800 ml distilled water.
Approximately 2 ml (1:1) ammonium hydroxide (NH4 OH) was added to facilitate disso-
lution and to prevent effervescence when bicarbonate was added. When most of DTPA
was dissolved, 79.06 g ammonium bicarbonate (NH4 HCO3 ) was added and stirred gently
until dissolved. The pH was adjusted to 7.6 with ammonium hydroxide. The solution was
diluted to 1 L with distilled water.
Extraction Method
The 10 ml extracting solution was added into 10 g of air-dried rhizospheric soil and shaken
on a reciprocal shaker for 15 min at 180 cycles/min. The rhizospheric soil extract was then
filtered through filter paper, Whatman no. 42. This rhizospheric soil extract was used to
analyze the following macro- and micronutrients.
Macronutrient Analysis of Hizospheric Soil

One ml of rhizospheric soil extract and 9 ml of distill water were taken in a test tube
and analyzed for Na+ , Ca+2 , Mg+2 , and K+ on a Shimadzu AA-670 atomic absorption
spectrophotometer.
Physiological and Biochemical Analysis of Suaeda fruticosa
Chlorophyll Content of Leaves

Chlorophyll content of Suaeda fruticosa leaves was determined by the method of Aronson
(1985).
Procedure. Crude preparation 1 mL was mixed with 4 ml of 80 percent (w/v) acetone and
allowed to stand in dark at room temperature. It was centrifuged at 2000 rpm for 5 min to
clear the suspension. Supernatant was used for chlorophyll determination. Absorbance of
solution was read at 645 nm (chlorophyll a) and at 663 nm (chlorophyll b) on spectropho-
tometer against 80 percent (v/v) acetone blank. Total chlorophyll was determined for the
equation given by Aronson (1985):
Total chlorophyll (mg/l) = (20.2 × A 645) + (8.02 × B 663)

Leaf Protein Contents

Protein content of Suaeda fruticosa leaves was determined following the method of Lowry
et al. (1951) using BSA as standard.
Phosphate Buffer (Stock Solution)

To create monobasic sodium phosphate, 27.6 g was dissolved in distilled water (1000 ml).
To create dibasic sodium phosphate, 53.6 g was dissolved in 1000 ml. Monobasic sodium
phosphate (16 ml) and dibasic sodium phosphate (84 ml) were mixed together to get the
desired pH (7.5) of the phosphate buffer.
Reagents
To create reagent A, 2 g sodium carbonate (Na2 CO3 ), 0.4 g NaOH (0.1 N), and 1 g Na-
K tartrate were dissolved in 100 ml of distilled water. To create reagent B, CuSO4 ·5H2 O
(0.5 g) was dissolved in 100 ml of distilled water. To create reagent C, solution A (50 ml)
and solution B (1 ml) were mixed. To create reagent D, folin phenol reagent was diluted
with distilled water in a 1:1 ratio.
Procedure. Fresh leaves 0.1 g were ground with the help of mortar and pestle in 1 ml of
phosphate buffer pH 7.5 and centrifuged for 10 min at 3000 rpm. The supernatant (0.1 ml)
of the given sample containing an unknown amount of protein was poured in the test tubes
and the total volume of 1 ml was made by distilled water. One ml of reagent C was added.
After shaking for 10 min, 0.1 ml of reagent D was added. The absorbance of each sample
was recorded at 650 nm after 30 min of incubation. The concentration of protein contents
was determined with the reference to standard curve made by using standard BSA (bovine
serum albumen). The BSA of different concentrations (20, 40, 60, 80, 320, and 640 mg)
was prepared. The absorbance of BSA was recorded at 650 nm.
Sugar Estimation
Sugar estimation of fresh leaves of Suaeda fruticosa was done following method of Dubois
et al. (1956).
Procedure. Fresh plant material (0.5 g) was homogenized with 10 ml of distilled water
in a clean mortar and then centrifuged at 3000 rpm for 5 min. To 0.1 ml of supernatant,
1 ml of 80 percent (w/v) phenol was added, and after incubation at room temperature,
5 ml concentrated sulfuric acid was added. The sample was incubated for 4 h and ten
absorbances of each sample were recorded at 420 nm. The concentration of unknown
sample was calculated with reference to standard curve made by using glucose.
Proline C
The proline contents of Suaeda fruticosa leaves were measured by the method of Bates,
Waldren, and Teare (1973). Fully expanded fresh leaves were sampled. Purified praline was
used to standardize the procedure for quantifying sample values. Reagent acid ninhydrin
was prepared by warming 1.25 g ninhydrin in 30 ml glacial acetic acid and 20 ml 6 M
phosphoric acid, with agitation, until dissolved. It was kept cool (4 ◦ C), and the reagent
remained stable for 24 h. Approximately 0.5 g of plant material was homogenized in 10 ml

of 3 percent aqueous sulfosalicylic acid and homogenate filtered with Whatman no. 42 filter
paper. Two ml of filtrate was reacted with 2 ml acid ninhydrin and 2 ml of glacial acetic
acid in a test tube for 1 h at 100 ◦ C and reaction was terminated in an ice bath. The reaction
mixture was extracted with 4 ml toluene and mixed vigorously with a test tube stirrer for
15–20 s. The chromophore containing toluene was aspirated from the aqueous phase and
warmed to room temperature, and the absorbance was read at 520 nm against toluene as
blank. The proline concentration was determined from a standard curve and calculated on
a fresh-weight basis as follows:

(µ g proline/ml × ml toluene) /115.5 µ g/umol / (g sample) /5
= µ mol proline/g of fresh weight material.

Assay for Antioxidant Enzyme Activity
Superoxide Dismutase Activity (SOD)

The SOD activity was determined by measuring inhibition of photochemical reduction
of nitroblue tetrazolium (NBT) using method of Beauchamp and Fridovich (1971). The
reaction mixture (3 ml) was composed of 13 mM methionine, 0.075 mM NBT, 0.1 mM
EDTA, 0.002 mM riboflavin, and 0.1 ml of enzyme extract in 50 mM phosphate buffer (pH
7.8). The mixture in tube was placed below a light chamber for 15 min. The absorbance
was read at 560 nm with a spectrophotometer. One unit of enzyme activity was taken as
that quantity of enzyme, which reduced the absorbance reading to fifty in comparison with
tube lacking enzyme.
Peroxidase Activity (POD)

The peroxidase activity was determined by the method of Vetter, Steinberg, and Nelson
(1958) as modified by Gorin and Heidema (1976). The POD activity was measured follow-
ing the assay mixture contained 0.1 ml enzyme extract, 1.35 ml of 100 mM MES buffer (PH
5.5), 0.05 percent H2 O2 , and 0.1 percent phenylene diamine. Change in absorbance was
recorded at 485 nm for 3 min with spectrophotometer. The activity of POD was presented
as OD 485 nm / min mg protein.
Ascorbate Peroxidase Activity (APX)

Ascorbate peroxidase (APX) activity was determined according to Asada and Takahashi
(1987). The reaction mixture (1 ml) contained 50 mM of potassium phosphate buffer
(pH 7.0), 0.5 mM of ascorbic acid, 0.1 mM of H2 O2 , and 200 µl of enzyme extract.The
absorbance was read as the decrease at 290 nm against the blank, and correction was done
for the low, nonenzymatic oxidation of ascorbic acid by H2 O2 . The enzymatic activity was
expressed as OD/min/g FW.
Catalase Activity (CAT)

Catalase (CAT) was measured according to Chandlee and Scandalios (1984) with mod-
ification. The essay mixture contained 2.6 ml of 50 mM potassium potassium phosphate
buffer (pH 7.0), 0.4 ml of 15 mM H2 O2 , and 0.04 ml of enzyme extract. The decomposition
of H2 O2 was followed by the decline in absorbance at 240 nm. The enzyme activity was
expressed as OD/min/g FW.
Statistical Analysis
The data were analyzed statistically by an analysis of variance (ANOVA) technique using
Costate software (Cohort software, Monterey, USA). Mean comparisons were conducted
using ANOVA protected least significance difference (LSD) (P < 0.05) test.
Results
Physicochemical Analysis of Rhizospheric Soil of Suaeda fruticosa

Comparison of Na+ content in rhizospheric soil of Suaeda fruticosa at various pheno-

logical stages were made for the samples collected from their natural biotope of the four
District of KP, Pakistan with a varying level of soil salinity (Table 1). Results showed that
maximum Na+ content has been reported in rhizospheric soil of Suaeda fruticosa col-
lected from District Nowshera at the germination stage, whereas the lowest Na+ content
in rhizospheric soil of Suaeda fruticosa has been reported at the same stage collected from
District Nowshera. The greatest Ca+2 content in rhizospheric soil of Suaeda fruticosa at
germination stage collected from District Peshawar has been reported in Table 1 followed
by a gradual decrease in Ca+2 content in rhizospheric soil of Suaeda fruticosa collected
from District Nowshera and Mardan at the same stage. The lowest mean Ca+2 content
in rhizospheric soil of Suaeda fruticosa have been reported at all the three phenological
stages collected from District Charsadda. The greatest Mg+2 content of rhizospheric soil
of Suaeda fruticosa at germination stage collected from District Nowshera followed by a
considerable decrease in the samples collected from District Charsadda at the same stage
has been reported in Table 1. The lowest Mg+2 content have been reported in rhizospheric
soil of Suaeda fruticosa at all the three growth stages collected from District Peshawar.
The highest content of K+ content in rhizospheric soil of Suaeda fruticosa at germination
stage collected from District Charsadda has been reported. The difference in K+ content in
rhizospheric soil of Suaeda fruticosa at germination stage collected from District Mardan
is considerable with the greatest content, as mentioned previously.
Physiological and Biochemical Analysis of Suaeda fruticosa

The greatest chlorophyll a/b ratio in SA at the germination stage collected from District
Charsadda has been reported in Figure 1. The lowest chlorophyll a/b ratio has been
reported in Suaeda fruticosa at all the three growth stages collected from Districts Mardan,
Nowshera, and Peshawar. Figure 1 revealed a significant difference in chlorophyll a/b ratio
in Suaeda fruticosa at various stages for the two consecutive years at P < 0.05. Comparison
of carotenoid content in Suaeda fruticosa at various phenological stages were made for the
species collected from their natural habitat of the selected areas with a varying level of
soil salinity showed that maximum carotenoid content has been reported in Suaeda fruti-
cosa collected from District Mardan at flowering stage (Figure 2). The lowest carotenoid
content in Suaeda fruticosa has been reported at the same stage collected from District
Table 1
Physicochemical analysis of rhizospheric soil of Suaeda fruticosa
Na+ content (µg/g) Ca+2 content (µg/g) Mg+2 content (µg/g) K+ content (µg/g)
Study area EC (dS/m) pH Ger Veg Flo Ger Veg Flo Ger Veg Flo Ger Veg Flo
Mardan 5.1 ± 0.48 7.8 ± 0.01 18.5 ± 0.145 17.81 ± 0.166 16.9 ± 0.115 46.93 ± 0.067 46.21 ± 0.167 44.41 ± 0.054 4.32 ± 0.034 3.49 ± 0.054 2.83 ± 0.056 0.213 ± 0.006 0.183 ± 0.005 0.1145 ± 0.003
(34◦ 20 N,
72◦ 0 E)
Charsadda 5.7 ± 0.28 5.0 ± 0.01 14.83 ± 0.214 14.11 ± 0.421 13.66 ± 0.548 44.93 ± 0.043 43.01 ± 0.153 41.5 ± 0.058 5.045 ± 0.069 3.81 ± 0.051 3.11 ± 0.081 0.344 ± 0.004 0.297 ± 0.007 0.2335 ± 0.005
(34◦ 07 N,
1218
71◦ 45 E)
Nowshera 8.24 ± 0.75 9.2 ± 0.02 20.55 ± 0.360 19.56 ± 0.448 18.9 ± 0.742 49.77 ± 0.466 47.54 ± 0.348 46.8 ± 0.920 5.82 ± 0.059 4.27 ± 0.052 3.72 ± 0.049 0.084 ± 0.0039 0.061 ± 0.002 0.03 ± 0.004
(34◦ 0 N,
72◦ 0 E)
Peshawar 8.0 ± 0.27 4.5 ± 0.01 14.66 ± 0.709 13.1 ± 0.081 12.29 ± 0.101 52.47 ± 0.635 50.62 ± 0.409 48.5 ± 0.726 4.07 ± 0.043 3.16 ± 0.040 2.65 ± 0.046 0.067 ± 0.0029 0.045 ± 0.004 0.0195 ± 0.004
(34◦ 02 N,
71◦ 37 E)
Notes. Comparison of Na+ , Ca+2 , Mg+2 , and K+ contents (µg/g) in rhizospheric soil of Suaeda fruticosa for the mean value of two consecutive years at various phenological stages collected from their natural biotope of the four
districts of KP, Pakistan, with varying levels of soil salinity. Data are expressed as mean ± SE (n = 5) of five composite samplings. Ger, germination stage; Veg, vegetative stage; and Flo, flowering stage.
Germination Vegetative Flowering

a
2.5
b
chlorophyll a/b ratio (mg/g)

2 c
1.5 d
e f
g g g
h
1 i i
0.5
0
Figure 1. Comparison of chlorophyll a/b ratio (mg/g) of Suaeda fruticosa for the mean value of
two consecutive years at various phenological stages collected from their natural biotope of the four
districts of KP, Pakistan, with varying levels of soil salinity: SEM (Suaeda fruticosa collected from
District Mardan), SFC (Suaeda fruticosa collected from District Charsadda), SFN (Suaeda fruticosa
collected from District Nowshera), and SFP (Suaeda fruticosa collected from District Peshawar). All
bars that share the same letters are not significantly different at the P < 0.05 level of significance.
LSD value: 0.0444.
a Germination Vegetative Flowering

14
Carotenoid content (mg/g)
12 b
c
10
e
d d d
e
8 f
g
6 h
i
4
Figure 2. Comparison of carotenoid content (mg/g) of Suaeda fruticosa for the mean values of
bars that share same letters are not significantly different at the P < 0.05 level of significance. LSD
value: 0.2499.
Charsadda. Figure 2 revealed significant difference for carotenoid content in Suaeda fru-
ticosa at various phenological stages at P < 0.05. A linear increase in carotenoid content
has been observed from germination into flowering stage.

400 a
b
350
d
e g e
Sugar content (µg/g)

h f c
300 i j
k
250
200
150
100
50
0
Figure 3. Comparison of sugar content (µg/g) of Suaeda fruticosa for the mean values of two con-
secutive years at various phenological stages collected from their natural biotope of the four districts
of KP, Pakistan, with varying levels of soil salinity: SEM (Suaeda fruticosa collected from District
Mardan), SFC (Suaeda fruticosa collected from District Charsadda), SFN (Suaeda fruticosa col-
lected from District Nowshera), and SFP (Suaeda fruticosa collected from District Peshawar). All
bars that share same letters are not significantly different at the P < 0.05 level of significance. LSD
value: 0.734.
The greatest sugar content in Suaeda fruticosa at the flowering stage collected from
District Peshawar has been reported in Figure 3. The lowest sugar content in Suaeda fru-
ticosa has been reported at the same phenological stage collected from District Mardan
and Charsadda with a nonsignificant difference at P < 0.05 from each other. The greatest
proline content in Suaeda fruticosa at flowering stage collected from District Peshawar has
been reported in Figure 4. The lowest proline content has been reported in Suaeda fru-
ticosa at all the three growth stages collected from Districts Nowshera and Mardan. The
greatest content of protein in Suaeda fruticosa at germination stage collected from District
Peshawar has been reported in Figure 5. A significant decrease in protein content has been
reported in Suaeda fruticosa at germination stage collected from Districts Nowshera and
Charsadda at P < 0.05 in comparison with the one collected from District Peshawar at the
same stage. The lowest content of protein content at germination stages has been reported
in Suaeda fruticosa collected from District Mardan.
Assay for Antioxidant Enzyme Activity

Results in Figure 6 show that maximum SOD content has been reported in Suaeda fru-
ticosa collected from District Mardan at the germination stage. A significant decrease at
P < 0.05 has been reported in Suaeda fruticosa collected from District Nowshera followed
by Peshawar at the same stage. The lowest SOD content in Suaeda fruticosa has been
reported at the same stage collected from District Charsadda. Figure 6 reveals that sig-
nificant difference occurred for SOD content in Suaeda fruticosa at various phenological
stages at P < 0.05. The greatest POD content in Suaeda fruticosa at flowering stage col-
lected from District Nowshera has been reported in Figure 7 followed by a nonsignificant
decrease at P < 0.05 in the species collected from District Charsadda at the same pheno-
logical stage. The lowest POD content in Suaeda fruticosa has been reported at flowering
stage collected from District Peshawar. The greatest CAT content in Suaeda fruticosa at
Germination Vegetative Flowering a

400
b
350
d c
Proline content (µg/g)

300 e
f
250
g
200 h
i j
k j
150
100
50
0
Figure 4. Comparison of proline content (µg/g) of Suaeda fruticosa for the mean values of two
consecutive years at various phenological stages collected from their natural biotope of the four
LSD value: 0.934.

a
400 b c
350
d
Protein content (µg/g)
300
250 e f
g
h h i
i j
200
150
100
50
Figure 5. Comparison of protein content (µg/g) of Suaeda fruticosa for the mean values of two
consecutive years at various phenological stages collected from their natural biotope of the four
LSD value: 2.627.
germination stage collected from District Charsadda has been reported in Figure 8 fol-
lowed by a nonsignificant decrease at P < 0.05 reported in the species collected from
District Nowshera at the same phenological stage. The lowest CAT content at germination
stages has been reported in Suaeda fruticosa collected from District Peshawar. The greatest

a
0.7
b b
0.6
SOD content (unit g–1 f.w)

c c c c c
0.5 d
e e
0.4 f
0.3
0.2
0.1
0
Figure 6. Comparison of SOD content (unit g−1 FW) of Suaeda fruticosa for the mean values of
districts of KP, Pakistan, with varying levels of soil salinity. SEM (Suaeda fruticosa collected from
LSD value: 0.0271.
APX content in Suaeda fruticosa at the germination stage collected from District Mardan
is reported in Figure 9, followed by nonsignificant decrease at P < 0.05 in the species col-
lected from District Nowshera at the same phenological stage. The lowest APX content at
germination stages has been reported in Suaeda fruticosa collected from District Peshawar.
Figure 9 revealed that a significant difference occurs in APX content in Suaeda fruticosa
at various stages for the two consecutive years at P < 0.05.
Discussion
In the present study Suaeda fruticosa maintained a K+ /Na+ ratio greater at the vegetative
phase, perhaps to cope with greater metabolic activity during the vegetative phase of plant
growth. The sensitivity of cytosolic enzymes to salt is similar in both glycophytes and
halophytes, indicating that the maintenance of high cytosolic K+ /Na+ ratio and Ca+2 /Na+
ratio is a key requirement for plant growth under salt stress conditions (Glenn and Brown
1999; Ebrahimian, Roshdi, and Bybordi 2011). At the vegetative phase the K+ /Na+ ratio
of rhizospheric soil of Suaeda fruticosa from lower EC (District Mardan) was greater than
1. The K+ content has been found reduced in all the selected soil samples as compared
to Na+ , Ca+2 , and Mg+2 is supported by Shabala and Cuin (2008), who reported that
availability of K+ content under saline condition is hindered by an antagonist effect of
Na+ due to similar physicochemical properties and their competition for the uptake sites
at the cell surface membrane. It has been previously reported that Ca+2 content usually
increased as compared to Mg+2 content (Ruiz, Martinez, and Cerda 1997) is in support of
the present findings.
Concentrations of chlorophyll components of the photosynthetic apparatus are nor-
mally used to quantify leaf senescence in salt-stressed plants. Salinity adversely affects
a a
2.5
b b
b
POD content (OD/min/g f.w)

c
2 cd d cd
e
f
1.5 f
0.5
0
Figure 7. Comparison of POD content (OD / min / g FW) of Suaeda fruticosa for the mean value of
LSD value: 0.118.

1.6
a a
1.4
CAT content (OD/min/g f.w)
1.2 b
bc
1 c cd cd
d
0.8 de
e e
0.6
f
0.4
0.2
Figure 8. Comparison of CAT contents (OD / min / g FW) of Suaeda fruticosa for the mean value
of two consecutive years at various phenological stages collected from their natural biotope of the
four districts of KP, Pakistan, with varying levels of soil salinity: SEM (Suaeda fruticosa collected
from District Mardan), SFC (Suaeda fruticosa collected from District Charsadda), SFN (Suaeda
fruticosa collected from District Nowshera), and SFP (Suaeda fruticosa collected from District
Peshawar). All bars that share the same letters are not significantly different at the P < 0.05 level of
significance. LSD value: 0.143.
chlorophyll content in plants grown under salt stress (Meloni et al. 2003). The greater
chlorophyll a/b ratio considered salt tolerance index been reported in plants grown under
high soil salinity as compared to that in the lower saline conditions (St˛epień and Kłbus
2006). The chlorophyll a/b ratio was found maximum in Suaeda fruticosa collected from

1.8 a
a
b
APX content (OD/min/g f.w)

1.6 c
d
1.4 ef e
g fg
1.2
1
0.8 h
0.6 i
0.4 j
0.2
0
Figure 9. Comparison of APX content (OD / min / g FW) of Suaeda fruticosa for the mean value of
LSD value: 0.0765.
District Peshawar (EC: 6.8–8.0). Maximum accumulation of sugar and proline content
at P < 0.05 in Suaeda fruticosa at the flowering stage has been reported from District
Peshawar (EC 6.8–8.0). This increase in sugar content at flowering stage indicated the pos-
sible increase in the hydrolysis of stored carbohydrates contents in the leaves, which might
contribute to speed up the rate of translocation of prestored reserves used for maturation
of fruiting body (Yang and Zhang 2006). The correlation between sugar accumulation and
osmotic stress tolerance has been previously reported for plants grown under salinity stress
condition (Ramanjulu, Veeranjaneyulu, and Sudhakar 1994; Baki et al. 2000; Streeter,
Lohnes, and Fioritto 2001; Taji et al. 2002). Proline accumulation is considered important
for osmotic adjustment under salt- and drought-stress conditions (Singh and Dara 1973;
Riazi, Matsuda, and Arslan 1985; Navari, Quartacci, and Izzo 1990; Ober and Sharp 1994;
Hyun et al. 2003). The increase in the level of soluble proteins content under varying
levels of soil salinity in some halophytes and decrease in others was concurrent with the
previous work of Ge et al. (2006). Significantly lower protein content has been reported in
Suaeda fruticosa collected from District Mardan (EC 4.3–5.5) as compared with those col-
lected from other districts, whereas Suaeda fruticosa maintain greater protein content when
collected from District Peshawar (EC 6.8–8.0) only. The greater protein content at germi-
nation stage followed by a gradual decrease in vegetative and flowering stage has been
supported by the work of Niazi and Ahmed (1986), who reported a progressive decrease in
the soluble protein content from vegetative into flowering stage in kallar grass. According
to Ashraf, Kausar, and Ashraf (2003), accumulation of proteins in leaves under water stress
conditions might be an adaptive mechanism in plants growing under salt stress as some of
the soluble proteins are up-regulated under osmotic stress condition, whereas others are
degraded.
Under stressed conditions plant membranes are subject to changes often associ-
ated with the increases in permeability and loss of integrity (Blokhina, Virolainen, and
Fagerstedt 2003). Changes of antioxidants reflect the impact of environmental stresses on
plant metabolism (Herbinger et al. 2002). The SOD content has been found maximum at
P < 0.05 in Suaeda fruticosa, at vegetative stage collected from soil having lower EC of
District Mardan. A significant decrease at P < 0.05 has been reported from vegetative into
flowering stage for the selected halophyte species. Different plants counteract stress dif-
ferentially as a result of variations in their antioxidant enzyme systems (Pastori and Trippi
1992; Turcsanyi et al. 1994; Kraus, McKersie, and Fletcher 1995). It is known that plants
have well organized defensive systems against reactive oxygen species under osmotic
stress conditions and SOD constitutes the first line of defense through detoxification of
superoxide radicals (Sairam et al. 2000). Increase in the antioxidant enzymes potential act
as a scavenging agent and thus provides defense against oxidative stress, which otherwise
could cause lipid-peroxidation resulting in injuries to the cell membrane, organelles, pro-
tein, and DNA structure and inhibit photosynthesis and other enzyme activities (Sairam
and Saxena 2000).
Conclusion
Halophytes differ in their physiological mechanism of adaptation to high salinity. Some
have better systems for regulation of osmotic potential using both sugar and proline as
osmoprotectants, whereas others have more efficient systems of alleviating oxidative stress
(created as secondary stress of salinity) by enhancing the activities of antioxidant enzymes.
Greater accumulation of osmolytes including sugar and proline along with antioxidant
enzyme activity including SOD and POD are the best physiological markers studied in
the present work. However, research regarding the progress in metabolomics, genomics,
gene regulation, and proteomics has to be started in diverse halophytic species including
Suaeda fruticosa with the use of advanced techniques to gain detailed knowledge of abiotic
stress tolerance.
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Physiological Mechanism of Salt

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Physiological Mechanism of Salt Tolerance

SAMI ULLAH1 AND ASGHARI BANO2

Keywords Halophyte, salinity, Suaeda fruticosa

Received 19 May 2013; accepted 5 October 2014.

Materials and Methods

Collection of Halophyte Species

Physicochemical Analysis of Rhizospheric Soil

Soil pH and Electrical Conductivity (EC)

Nutrient Analysis of Rhizospheric Soil

Extraction Solution Preparation

Macronutrient Analysis of Hizospheric Soil

Physiological and Biochemical Analysis of Suaeda fruticosa

Chlorophyll Content of Leaves

Total chlorophyll (mg/l) = (20.2 × A 645) + (8.02 × B 663)

Leaf Protein Contents

Phosphate Buffer (Stock Solution)

remained stable for 24 h. Approximately 0.5 g of plant material was homogenized in 10 ml

= µ mol proline/g of fresh weight material.

Assay for Antioxidant Enzyme Activity

Superoxide Dismutase Activity (SOD)

Peroxidase Activity (POD)

Ascorbate Peroxidase Activity (APX)

Catalase Activity (CAT)

Physicochemical Analysis of Rhizospheric Soil of Suaeda fruticosa

Comparison of Na+ content in rhizospheric soil of Suaeda fruticosa at various pheno-

Physiological and Biochemical Analysis of Suaeda fruticosa

Germination Vegetative Flowering

chlorophyll a/b ratio (mg/g)

a Germination Vegetative Flowering

Germination Vegetative Flowering

Sugar content (µg/g)

Assay for Antioxidant Enzyme Activity

Germination Vegetative Flowering a

Proline content (µg/g)

Germination Vegetative Flowering

Germination Vegetative Flowering

SOD content (unit g–1 f.w)

Germination Vegetative Flowering

POD content (OD/min/g f.w)

Germination Vegetative Flowering

Germination Vegetative Flowering

APX content (OD/min/g f.w)

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