Effect of salinity on biochimical and anatomical characteristics of sweet pepper (Capsicum annuum L.)

245 Bouassaba and Chougui
Int. J. Biosci. 2018
RESEARCH PAPER OPEN ACCESS
Effect of salinity on biochimical and anatomical characteristics
of sweet pepper (Capsicum annuum L.)
Karima Bouassaba1
, Saiida Chougui2
1
University Abdelhafid Boussouf Center, Mila,43000. Algeria
2
Department of Biology and Ecology Vegetal, Constantine Algeria
3
Specialty Biodiversity and plant production , Mentouri University1, Constantine, Algeria
4
Laboratory of Plant Physiology, Department of Biology Mentouri University1, Constantine, 25000.
Algeria
Key words: Tolerance, salinity, resistance, Capsicum annuum L.
http://dx.doi.org/10.12692/ijb/12.6.245-257 Article published on June 24, 2018
Abstract
Salinity is considered as the most important abiotic stress limiting crop production and plants are known to be
able continuing survive under this stress by involving many mechanisms. In this content, the present study was
carried out to evaluate the impact of NaCl on some morpho- physiological and anatomical parameters in two
sweet pepper (Capsicum annuum L.) varieties: Super marconi and Marconi. So, an experiment of eight months
was carried out underStandard room at and stress is induced by NaCl at 4 concentrations (0, 25, 50, and
150mMol/l). Results showed that increasing salinity stress, for all cultivars, decreased stems (length, fresh and
dry weights) and leaves (number and area).As the salinity increased, proline concentration and leaf total soluble
sugars also increased significantly compared with the control. The results showed that the accumulation of
proline and soluble sugars are good indicators of salinity tolerance. Results also suggest that the plant resists
against salinity through osmotic adjustment and ion absorption and sharing within its cells. This process is
essential for the survival of plants in salineMicroscopic study demonstrated that .salinity stress significantly
decreased cortex thickness because of salinity stress while xylem grown under salinity stress especially high level
of salinity .Additionally. There were changed in xylem creation and construction in stressed plants. It is
concluded that the variety Super Marconi (Sp) is more tolerant to salinity compared to the variety Marconi (M).
Hence, they have a significant role to play in agriculture, food, and economy.
* Corresponding Author: Karima Bouassaba  karima2125@ yahoo.fr.
International Journal of Biosciences | IJB |
ISSN: 2220-6655 (Print), 2222-5234 (Online)
http://www.innspub.net
Vol. 12, No. 6, p. 245-257, 2018

Introduction
Soil salinization is one of the major factors of soil
worsening. It has reached 19.5 % of the watered land
and 2.1 % of the dry-land farming existing on the
globe (FAO, 2000). Salinity effects are more easily
seen in arid and semiarid areas where 25 % of the
watered land is affected by salts. Considering that 52
% of northeast Brazil is within semiarid tropics (Lira
et al., 1982).
Irrigation has played an extremely important role in
terms of food production worldwide by increasing
crop produce and quality. However, to many crops-
watering can cause soil worsening mostly by
increasing the soil salinity (Trout, 2000). In fact, high
levels of salinity have been reported as causing of the
loss of 250,000 to 500,000 ha of irrigated land every
year. The problem happens mostly in arid and
semiarid zones where a total of 100 to 110 million ha
are reported as having problems related to
salinization which could make them unusable for
farming- based purposes (FAO, 2002). The problem
is greatest in these areas due to the high level of
evapotranspiration which increases salts, introduced
via the rising with water (secondary salinization) or as
part of the original chemical composition of the soil.
Pepper is one of the most widely grown vegetable in
the world. World production of pepper is estimated at
23.2 million tones and the largest producer is China
with 11.5 million tons, or nearly 50% (FAO, 2002).
Salinity inhibition of plant growth is the result of
osmotic and ionic effects and the different plant
species have developed different mechanisms to cope
with these effects (Munns, 2002). The osmotic
adjustment, reduction of cellular osmotic potential by
net solute accumulation, has been believed an
important method to salt and drought tolerance in
plants. This reduction in osmotic potential in salt
stressed plants can be a result of inorganic ion (Na+,
Cl-, and K+) and compatible organic solute (soluble
carbohydrates, amino acids, proline, roots and leaves
contribute to the maintenance of water uptake and
cell turgor, allowing physiological processes, such as
stomatal opening, photosynthesis, and cell expansion
(Serraj and Sinclair, 2002). In addition to their role in
cell water relations, organic solute accumulation may
also help towards the maintenance of ionic
homeostasis and of the C/N ratio, removal of free
radicals,and stabilization of macromolecules and
organelles, such as proteins, protein complexes and
membranes (Bohnert and Shen, 1999, Bray et al.,
2000). These solutes may also help towards the
control of pH in the cytosol and detoxification of
excess NH4+ (Gilbert et al., 1998) Although the
relationship between osmoregulation and salt
tolerance is not clear, there is evidence that the
osmotic adjustment appears, at least partially, to be
involved in the salt tolerance of certain plant
genotypes (Richardson and Mccree, 1985). Therefore,
the objective of this investigation was to evaluate the
effects of salt stress on growth, water relations and
anatomical of different sweet pepper genotypes,
commonly grown in northeast Algeria, and at the
same time try to correlate these effects with changes
in ionic and organic solute accumulation, with a view
to a better understanding of the mechanisms of salt
tolerance in these genotypes.
Materials and methods
Study area
This study is carried out on two varieties of sweet
pepper (Capsicum annuum L.) varieties: Marconi
(M) / the origin: china) and Super marconi (Sp) / the
origin: china). The experiment was conducted in a
culture room at the Abdelhafid Boussouf University
Center (Wilaya de Mila, North-East Algeria.), During
the year 2016/2017.
The experimental design
The test is conducted according to an experimental
device in completely randomized blocks (BAC),
comprising 4 blocks (repetitions) comprising 4 blocks
(repetitions) of each variety of sweet pepper (Marconi
and Super marconi). Fo each repetition two plants.
Saline solutions
The plants of the two varieties are subjected to the
different treatments of NaCl:
S0 :NaCl ( 0 m Mol / L (control).
S1: NaCl( 25 m Mol / L.)

S2 :NaCl ( 50 m Mol / L.)
S3 :NaCl ( 150 m Mol / L.)
Measured parameters
Number of leaves NF:
We calculated the number of leaves from the beginnin
g of the growth ofthe first two papers until the rotting
of the plants for each of the two varieties separately, a
nd the averages were calculated.
Measuring leaf area SF (cm2): The leaves area was cal
culated during the seedling growth stage of both culti
vars during theremoval of seedlings for study and the
averages were calculated.
Fresh weight (PF / g): We calculated wet weight durin
g the seedling growth stage and afterremoval of the se
edlings by a sensitive balance.
Dry Weight Measurement (PS / g):
After gaining swelling weight, we wrap the balanced
seedlings individually in the aluminum foil and then
put them in the incubator for drying under 80 ° C
for48 hours to dry completely.
Dry seedlings and we make the color we get dry
weight.
Leaf proline estimation
Extraction and estimation of proline were conducted
according to the procedures described by Bates etal.
(1973).
Leaf total soluble sugars estimation
The total soluble sugars were measured using the
phenol-sulfuric acid. To measure leaves’ total soluble
sugars from the solution of5% ZnSO4, 0.3 N Ba
(OH)2, 5% (v/v) phenol solution and sulfuric acid was
used based on Stewart method. Finally, absorption
was read at 485 nm by spectrophotometry (Stewart,
1989).
Anatomical study
According to Saadoun (2005) a realization of the
cross sections of the stems of the two varieties of
pepper (Capsicum annuum L.) with a razor blade.
This method is based on the use of certain dyes:
methyl green, Congo red or carmino-green Mirande,
or methyl green and carmine alum. It makes it
possible to exactly color the cell walls according to
their chemical composition. The staining principle of
the raised sections is based on the following steps:
First, Making cuts by hand using a simple razor, the
organ is delivered in slices of a few micrometers thick
so that the light rays can cross it.
Second, The sections are immersed, before staining,
in a 12% sodium hypochlorite bath for a quarter of an
hour to evacuate the contents of the cells.
Third, Rinsing with distilled water.
Fourth, Immersion in 2% acetic acid for five minutes
to eventually fix the dyes on the cells and remove all
traces of sodium hypochlorite.
Fifth, Rinse briefly with distilled water.
Sixth, Immersion in the double dye (methyl green /
Congo red) for 10 to 15 minutes.
Finally, Rinsing with distilled water, then mounting
the finest cuts between blade and coverslip.
Observation under a microscope equipped with a
digital camera (Motic).
Statistical analysis
The Excel Stat (2017) software was used to perform
all variance analyzes as well as the Newman keuils
test (α = 0.05) to compare the averages between the
control and processed samples for each parameter
analyzed.
Results and discussion
Leaf number
The different saline concentrations resulted in a
significant decrease in the number of leaves for both
cultivars. There was also a difference in the average
number of leaves between the two cultivars, especially
in the concentrations (25-50 mM / L).

Table 1. Analysis of variance on sweet pepper leaf number affected by salt stress.
Source DDL Sum of Squares Averag Squares F Pr> F
Model 7 151,889 21,698 6,873 0,000
Error 24 75,766 3,157
Total corrected 31 227,654
The highest mean number of leaves (12.25) 11.5) in
Super marconi in the control plants. In the
concentration (50 mM), Super marconi surpasses the
average number of leaves (9.43) on Marconi, which
has an average of 6.5 sheets.(50-150 mmol / L). The
average number of leaves ranged between(7.25-
7.81)inthe concentration (50 mM / L) and
between(6.06-6.26) in the concentration Saline (150
mM / L).
Table 2. Analysis of variance on sweet pepper leaf area affected by salt stress.
Model 7 148,969 21,281 17,462 < 0,0001
Error 24 29,250 1,219
The analysis of variance showed a significant effect (p
<0.01) on all traits due to salinity stress.
The results are shown in Table 1.The New man
Keuilstest confirms these differences between
cultured plants in the different saline media and
between the two cultivars. Super marconi recorded
the highest values of mean number of leaves in all
saline = concentrations compared to Marconi.
Which confirms the results of Unlükara et al. (2008).
Inhibition of the formation of leaf primordia under
salinity stress could be the probable reason for low
leaf number. Iyenger et al. (1977) reported that saline
irrigation water reduced the number of leaves.
Table 3. Analysis of variance on sweet pepper dry weight affected by salt stress.
Model 7 0,180 0,026 657,223 <0,0001
Error 24 0,001 0,000
Leafs area
According to the results shown in Fig. 2 the effect of
salt stress negatively affects the paper area of the
studied species. The highest concentration of paper
area in the control plants of Super marconi (9.75
cm2) and average values (6.75-8 cm2) in the
concentrations (25-50 mmol / L) The lowest value of
paper area (4 cm2) was recorded in the high salt
concentration (150 mM / L).(S = 0) and saline
concentration (1S = 25 mM / L) and the paper area
(6.5 cm2) and the mean values in the concentration
(S2 = 50 mM / l) ) And the lowest (2.25 cm2) in the
high salt concentration (S3 = 150 mmol / L).
The two varieties generally showed a different
response in the paper area in all saline treatments,
where the Super marconi exceeded Marconi. The
ANOVA study showed that the paper area recorded
very significant differences as shown in Table 2.
The New man Keuils test confirms these differences
between plants grown in different saline cultures and

between the two cultivars. Super marconi recorded
the highest values of the leaf evolution index in all
saline concentrations compared to Marconi. Sagi et
al. (1997) also found the adverse effects ofsalinity
stress on leaf area. It has been reported that, decline
in plant biomass may be due to excessive
accumulation of NaCl in chloroplasts of sweet pepper,
which affects growth rate, and is often associated with
a decrease in the electron transport activities of
photosynthesis (Kirst, 1989) and inhibition of PSII
activity (Kao et al., 2003).
Table 3. Analysis of variance on sweet pepper dry weight affected by salt stress.
Model 7 0,180 0,026 657,223 <0,0001
Error 24 0,001 0,000
In general, salinity reduces leaf number leaf area,
shoot and root dry weight, leading to low yields (Essa,
2002; Hamdy et al., 1993; Li et al., 2006; Sharifi et
al., 2007).Our results are inconsistence with Gosset
and Lucas (1996) who reported that NaCl highly
reduced total leaf area; it seems that plant height was
more sensitive to salinity than leaf number or leaf
area expansion.
Dry weigth and fresh weigths
Results shown in Fig. 3 show salinity has a negative
effect that resulted in a clear decrease in average dry
weight by increasing saline concentrations. The
highest mean dry weight (0.28 g) was recorded in
saline concentration (S0 = 0) in control plants and
(0.14 g) in saline concentration (S1 = 25 mM / L) for
Marconi species, while lowest values were recorded
for the same cultivar Saline (50-150 mM / L) ranged
between 0.05 g and -0.08 g.
Table 4. Analysis of variance on sweet pepper fresh weigth affected by salt stress.
Model 7 7,653 1,093 7,556 < 0,0001
Error 24 3,473 0,145
Super marconi had the highest mean dry weight
(0.23 g) in the control plants, and in the
concentrations (25-50 mM / L), the mean wet weight
values ranged from 0.11 g to 0.12 g while the lowest
mean Wet weight (0.07 g) in high salt concentration
(150 mM / L).
Salinity stress was diminished plant growth and
significantly decreased total dry weight. There was
downward decrease in shoot weight because of
deterrent effect of salinity on plant height (Table 3).
Salt stress adversely affects the growth and
development of crops, and the results of our study
confirm that all growth variables of sweet pepper
drastically decreased with NaCl treatment.
Fresh weigth
The results shown in Fig. 4 showed that the negative
salinity effect resulted in a clear decrease in average
wet weight by increasing saline concentrations. The
highest mean wet weight (2.16 g) was recorded in
saline concentration (S0 = 0) in the control plants
and (1.37 g) in the saline concentration (S1 = 25 mM /
L) of the Marconi species. Saline (50-150 mM / L)
ranged between (0.38 g - 0.94 g). Super marconi had
the highest mean wet weight (1.54 g) in the control
plants and in the concentrations (25-50 mM / L). The
mean wet weight values ranged between (1.19 g - 1.22
g) while the lowest mean Wet weight (0.88 g) in high
salt concentration (150 mM / L).

Table 5. Analysis of variance on sweet pepper proline affected by salt stress.
Analysis of the variance (Variable Pro L)
Model 8 2101,371 262,671 336,591 < 0,0001
Error 24 18,729 0,780
Table 6. Analysis of variance on sweet pepper soluble sugars affected by salt stress.
Analysis of the variance (Variable Sug L) :
Model 8 2990,653 373,832 6219,405 < 0,0001
Error 24 1,443 0,060
The ANOVA study showed that the average wet
weight recorded very significant differences. As
shown in Table 4 The New man Keuils test confirms
these differences between cultured plants in the
different saline media and between the two cultivars.
Super marconi recorded the lowest values for mean
wet weight in saline concentrations (0-25 mM / L)
and highest values In saline concentrations (50-150
mM / L) compared to Marconi. At the lower salinities,
cell expansion may have been sufficient to diluted the
ion concentration within the plant; thus avoiding
toxic accumulation (Munns, 2002). Plants irrigated
with salinities of 3.5 dS∙m−1 and greater may have
had a limited water uptake due to high osmotic
potential in the root zone. Salt-stressed plants tend to
intrinsically save water; thus restricting growth as an
adaptation to low water availability (Binzel and
Reuveni, 1994).
Table 7.Pearson correlation between growth parameters affected by salt stress.
Variable Leaf number Leaf dry weigtht Leaf fresh weigth Leaf area Leaf proline Leaf suger
Leaf number 1
Leaf dry weigth 0,740** 1
Leaf fresh weigth 0,585 0,766** 1
Leaf area 0,570 0,591 0,546 1
Leaf proline -0,622* -0,765** -0,577 -0,687* 1
Leaf suger -0,421 -0,655* -0,462 -0,318 0,576 1
*** Significant at the 0.05 and 0.01probbility levels, respectively.
Plants may have higher concentrations of Na+ and Cl−
in the leaves. Both Bethke and Drew (1992) and
Chartzoulakis and Klapaki (2000) reported higher Cl−
content than Na+ in the leaves of peppers at high
salinities, whereas Blom-Zandstra et al.,(1998)
reported the reverse. Both processes which ultimately
limit nutrient uptake, causing nutrient imbalances
(Grattan and Grieve, 1999). Based on our results, it
could be speculated that Cl− concentration in the
leaves might have been more harmful than Na+ based
on the fact that peppers are incapable of Cl− exclusion
(Chartzoulakis and Klapaki, 2000) and that chloride
was the most abundant ion in our saline irrigation
water.
Kirnak et al., (2003) found that mulched plants under
water stress had significantly greater water content in
leaves, shoot dry weight and stem diameters than
plants grown in bare soil. Under non-saline
conditions mulches have been found to significantly

increase plant height and stem diameter (Locher et
al., 2005), shoot fresh weight (Aziz, 1994) and
number of leaves (Siwek et al., 1994).
Proline
Results of the measurement of the amino acid proline
content, as shown in Figure (5), is the most studied
amino acid among the other amino acids involved in
the formation of proteins as a measure of the
physiological condition of the plant (salt stress).
Fig. 1. Effect of NaCl on leaves numbers of two sweet pepper cultivars during 60 days.
In the normal case (S0 = 0), the ratio of the amino
acid proline in the leaves of the pepper varieties was
weak, ranging from (3.05-4.76 μg / 100mg / MF).
While the amount of amino acid proline in the leaves
increased by saline concentrations ranged between
(4.89-6.06 μg / 100mg / MF) in the concentrations
(25-50 mM / L) of sodium chloride salt and the
maximum values were recorded in the high salt
concentration Mol / l) for both studied cultivars
(9.39-11.10 μg / 100mg / MF).
Fig. 2. Effect of NaCl on leaves area of two sweet pepper cultivars during 60 days.
The ANOVA study showed that the amino acid
content of proline in the leaves of the two varieties of
paprika has recorded very significant differences. As
shown in Table (5). The New man Keuils test
confirms these differences between plants grown in
different saline cultures and between the two
cultivars. Super marconi recorded the highest values

of the leaf evolution index in all saline concentrations
compared to Marconi.
Solubles sugars
Figure 6 shows that the content of sugars decreased
significantly in non- salted plants. While the effect of
salinity varied on sugar content between the two
studied cultivars with the highest values of (9.82-
10.15 μg / 100mg / MF) were recorded in the Super
marconi cultivars (25-50-150 mM / L), while the
lowest values were recorded in the Marconi cultivars
in the control plants and the low salinity
concentration (25 mM / L) ranged between (9.16-9.37
μg / 100mg / MF) compared with high salinity
concentrations with the highest values ranging from
(9.50-9.94 μg / 100mg / MF).
Fig. 3. Effect of NaCl on dry weigth of two sweet pepper cultivars during 60 days.
The ANOVA study showed that the sugar content of
the leaves showed very significant differences. As
shown in Table (6). The New man Keuils test
confirms these differences between plants grown in
different saline media and between the two cultivars.
Super marconi recorded the highest sugar content in
all saline concentrations compared to Marconi.
Fig. 4. Effect of NaCl on fresh weight of two sweet pepper cultivars during 60 day.

In this study, proline concentration in the leaf of
Capsicum annuum L. was increased with increasing
salinity, particularly at the highest external salt level,
thus, showing the positive role of proline in the salt
tolerance of this crop. Proline, as a
signaling/regulatory molecule, can activate multiple
responses, which are component to the process of
adaptation to abiotic stresses including salt stress
(Ashraf and Orooj, 2006).
Fig. 5. Effect of NaCl on proline of two sweet pepper cultivars during 60 day.
Increase in proline under salinity has also been
reported in some medicinal plants (Hajar et al., 1996;
Munns., 2002; Ashraf and Orooj, 2006; Abdul et al.,
2007). Leaf soluble sugars considerably increased in
response to the increase in salinity. Accumulation of
soluble carbohydrates in response to environmental
stress has been widely reported despite specific
reduction in net CO2 assimilation levels (Chaves et
al., 2003; Meloni et al., 2008). Sugars, in addition to
the role of regulating osmotic balance, also act as the
metabolic signals in the stress conditions (Chaves et
al., 2003).
Fig. 6. Effect of NaCl on soluble sugars of two sweet pepper cultivars during 60 day.

Anatomical results
A comparison of the anatomical cuts made on the
pepper stems (Capsicum annuum L.) of the control
and stressed plants shows that the stem of the control
plants consists of two zones: the bark and the central
cylinder (Fig. 5). In stressed pepper plants, an
increase in parenchymal cell size is observed under
saline treatment of 25 mMol/l, followed by an
increase in parenchyma thickness. However, under
treatment of 50-150 mMol/l, cell thickness decreases.
Furthermore, we observed that, in salt stressed
plants, number of trichomes was increased from
epidermal stem cells.
Fig. 7. Anatomical structur of sweet pepper stem under salinity stress.1 :Exodermis,2 : Endodermis,3 : central
cylinder.

In the other word, increase of salinity level led to
more trichomes on epidermal layer in compare with
control plants (Data are not shown) (Fig. 5). There are
several reports on increased trichome density under
environmental stresses such as drought and salinity
(Abernethy et al., 1998; Aguirre-Medina et al., 2002).
Increase of trichome density may be a mechanism to
increase of tolerance to salt stress.
It was recently suggested that leaf glandular
trichomes could contribute to the high salt tolerance
by the excretion of ions (Gucci et al., 1997). Salinity
induced structural changes in xylems in stems.
In salt stressed plants, stems vascular cell thickness
was much larger than control treatment; the salinity
effect was concentration dependent. Generally, plants
grown in saline solution showed higher thickness in
cuticle, vascular tissues and vessel than unstressed
plant while cortex zone thickness was decreased (Fig.
5). when the cell is under stress and when
itdifferentiates to particular specialized tissues,
notably the xylem (Christensen et al., 1998).
Salinity stress has been associated Witha greater
deposition of lignin in vascular tissues and/caused
earlier and stronger lignifications, which has been or
xylem development.
In bean- root vascular tissue, NaCl suggested to be a
factor that inhibits root growth and, consequently,
represents an adaptation mechanism in resisting
salinity-imposed stress (Cachorro et al., 1993).
Conclusion
The results show differences and similarities between
the two varieties during saline treatments. Super
marconi (Sup) is more tolerant to salinity compared
to Marconi (M).
In conclusion, this study shows that salt stress
decreases sweet pepper growth and induces changes
in anatomical characteristics such as increment of cut
in synthesis on epidermal stem cells and also changes
in xylem structure and lignification of them in
soybean stems.
References
Abernethy GA, Fountain DW, McManus
MT.1998. Observations on the leaf anatomy of
Festuca novae-zelandiae and biochemical responses
to a water deficit. N Z J Bot36:113-123
Aguirre-Medina JF, Acosta GallegosJA, del.
Ruiz Posadas L, Shibata JK, Trejo Lopez C
.2002. Morphological differences on the leaf
epidermis of common bean and their relationship to
drought tolerance. Agricultura technical en Mexico
28,53-64.
Abdul JC, Gopi R, Sankar B, Manivannan P,
Kishorekumar A, Sridharan R,
Panneerselvam R. 2007.Studies on germination,
seeding vigour lipid peroxidation and proline
metabolism in Catharanthus roseus, seedings under
salt stress.Journal.Botanic.73,190-195.
Ashraf M, Orooj A. 2006. Salt stress effects on
growth, ion accumulation and seed oil concentration
in an arid zone traditional medicinal plant ajwain
(Trachyspermum ammi L.) Sprague. Journal. Arid
Environ. 64,209-220.
Bethke PC, DrewMC. 1992. Stomatal and
nonstomatal components to inhibition of
photosynthesis in leaves of Capsicum annuumduring
progressive exposure to NaCl salinity. Plant
Physiology 99,219-226.
Bates LS, Waldren RP, Treare ID.1973. Rapid
determination of free proline for water stress studies.
Plant Soil 39,205-207.
Bethke PC, Drew MC. 1992. Stomatal and
nonstomatal components to inhibition of
photosynthesis in leaves of Capsicum annuumduring
progressive exposure to NaCl salinity. Plant
Physiology 99,219-226.

Binzel ML, Reuveni M. 1994. Cellular mechanisms
of salt tolerance in plant cells. Horticultural Reviews
16,33-69.
Bray EA, Bailey-Serres, Weretilnyk E. 2000.
Responses to abiotic stress. In: Buchanan B,
Gruissem W and Jones R (eds.), Biochemistry and
Molecular Biology of Plants. American Society of
Plant Physiology , Rockville, 1158-1203.
Bohnert HJ, Shen B.1999. Transformation and
compatible solutes. Scientia Horticulturaeourn.
78,237-260.
Chartzoulakis K, Klapaki G. 2000. Response of
two greenhouse pepper hybrids to NaCl salinity
during different growth stages. Scientia Horticulturae
86, 247-260.
BohnertHJ, Shen B. 1999. Transformation and
compatible solutes.Sci. Hortic.78, 237-260.
Chartzoulakis K, Klapaki G. 2000. Response of
two greenhouse pepper hybrids to NaCl salinity
during different growth stages. Scientia Horticulturae
86,247-260.
CachorroP, Ortiz A, Barcelo AR, Cerda A.1993.
Lignin deposition in vascular tissues of Phaseolus
vulgaris roots in response to salt stress. Phyton-Ann
Rei Bot 33,33-40.
Chaves MM, Maroco JP, Pereira JS.2003.
Understanding plant response to drought: from genes
to the whole plant. Funct. Plant Biology. 30: 239-264.
Christensen JH, Bauw G, Welinder KG,Van
Montagu M, Boerjan W. 1998. Purification and
characterization of peroxidases correlated with
lignification in poplar xylem. Plant Physiol 118:125-
135.
Essa TA.2002. Effect of salinity stress on growth and
nutrient composition of three soybeans (Glycine max
(L.) Merrill) cultivars. J Agronomy and Crop
Sci188,86-93.
FAO.2000. Irrigation in Latin America and The
Caribbean in figures. Water Reports 20. FAO, Rome.
348 p.
FAO.2002. Crops and drops: Making the best use of
water for agriculture. FAO, Rome. 22 p.
Grattan SR, Grieve CM. 1999.Salinity mineral
nutrient relations in horticultural crops. Scientia
Horticulturae 78,127-157.
Gregory PJ. 2004. Agronomic approaches to
increasing water use efficiency.
Gilbert GA, Gadush MV, Wilson C, Madore
MA.1998. Amino acid accumulation in sink and
source tissues of Coleus blumei Benth. during salinity
stress. Journal. Exp. Bot. 49,107-114.
Gossett DR, Lucas MC.1994. Antioxidant response
to NaCl stress in salt tolerant and salt sensitive
cultivars of cotton.Crop Sciences.34,706-714.
Hamdy A. 1999.Saline irrigation and management
for a sustainable use. In: Advanced Short Course on
Saline Irrigation Proceeding, Agadir: 152-227.
Hajar AS, Zidan MA, Al-Zahrani HS.1996. Effect
of salinity stress on the germination, growth and
some physiological activities of black cumin (Nigella
sativa L.). Arab Gulf. Journal. Scence i. Reserche.
14,445-454.
Kirnak H, Kaya C, Higgs D, Tas I. 2003.
Responses of drip irrigated bell pepper to water stress
and different nitrogen levels with or without mulch
cover. Journal of Plant Nutrition 26,263-277.
Kao WY, Tsai T, Shih CN.2003.Photosynthetic gas
exchange and chlorophyll a fluorescence of three wild
soybean species in response to NaCl treatments.
Photosynthetica .41,415-419.

Kirst GO.1989.Salinity tolerance of eukaryotic
marine algae. Plant Molecular Biology . 40:21-53
Li X, An P,Inanaga S, Eneji AE, Tanabe K.
2006.Salinity and Defoliation Effects on Soybean
Growth. J Plant Nutr 29,1499-1508.
Locher J, Ombodi A, Kassai T, Dimeny J. 2005.
Influence of coloured mulches on soil temperature
and yield of sweet pepper. European Journal of
Horticultural Science 70, 135-141.
Iyengar ERR, Patolia JS, Kurian T. 1977.
Varietal difference of barley to salinity. Z Pflanzen
Physio. l84,355-362.
Munns R. 2002. Comparative physiology of salt and
water stress. Plant Cell and Environment 25,239-
250.
Munns R.2002. Comparative physiology of salt and
water stress. Plant Cell.Environ.25,239-250.
Meloni DA, Gulotta MR, Martinez CA. 2008.
Salinity tolerance inSchinopsis quebracho colorado:
Seed germination, growth, ion relations and
metabolic responses. Journal. Arid Environ. 72, 1785-
1792.
Richardson SG, Mc Cree KJ. 1985. Carbon
balance and water relations of sorghum exposed to
salt and water stress. Plant Physiol. 79,1015-1020.
SagiM, Savidov NA, Vov NPL, Lips SH.1997.
Nitrate reductase and molybdenum cofactor in
annual ryegrass as affected by salinity and nitrogen
source. Physiol Plant. 99,546-553.
SharifiM, Ghorbanli M, Ebrahimzadeh H.
2007. Improved growth of salinity-stressed soybean
after inoculation with salt pre-treated mycorrhizal
fungi. J Plant Physiol164,1144-1151.
Siwek P, Cebula S, Libik A, Mydlarz J. 1994.
The effect of mulching on changes in microclimate
and on the growth of yield of sweet pepper grown in
plastic tunnels. Acta Horticulture 366,161-167.
Stewart EA.1989. Analysis of vegetation and other
organic material. In: Acad. Press, New York. 46-60 p.
Trout TJ.2000. Environmental effects of irrigated
agriculture. Acta Horticulturae 537,605-610. Van
Derwerken, J.E., and D.
U‫ـ‬nlükara A,Cemek B, Karaman S, Erşahin S.
2008. Response of lettuce (Lactuca sativa var. crispa)
to salinity of irrigation water. New Zealand J Crop
Hort Sci. 36,265-273.

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