Proc. Natl. Acad. Sci., India, Sect. B Biol. Sci.
(Oct–Dec 2023) 93(4):809–818
https://doi.org/10.1007/s40011-023-01484-1
RESEARCH ARTICLE
Application of Salicylic Acid on Chlorophyll, Carotenoids,
and Proline in Radish Under Salinity Stress
Kobra Mahdavian1
Received: 11 May 2022 / Revised: 3 June 2022 / Accepted: 26 March 2023 / Published online: 28 April 2023
© The Author(s), under exclusive licence to The National Academy of Sciences, India 2023
Abstract Various environmental stresses, including salin-
ity stress, reduce plant growth and yield. Salinity stress is
one of the most common stresses that cause soil and water
pollution worldwide. Salicylic acid (SA) is a plant hormone
that has been shown to have therapeutic effects against non-
biological stresses such as salinity. In this study, to increase
radish growth and tolerance to soil salinity, the effect of SA
on reducing the effect of salinity stress on radish growth
and biochemical changes was investigated. Results showed
that salinity (0, 25, 50, and 100 mM) significantly reduced
root and shoot growth, reduced root and shoot dry weight,
and also decreased chlorophyll and carotenoids, in contrast,
in plants pre-treated with SA (0, 0.5, 1, and 1.5 mM) was
reduced the response. Salinity significantly increased the
protective metabolites in radish antioxidant system like pro-
line, malondialdehyde, and other aldehydes. While modulat-
ing salicylic acid, this increase has increased the tolerance
of this plant to salinity stress. The application of SA can
increase plant tolerance to salinity stress.
Keywords Lipid peroxidation · Proline · Chlorophyll ·
Carotenoids · Plant species
Significance Statement: In this research, salinity stress affects
growth parameters and photosynthetic pigments in radish. Also,
the results showed that radish pretreatment with concentrations
of 0.5, 1, and 1.5 mM SA increases plant tolerance to NaCl
stress. Therefore, the use of SA in reducing the destructive
effects of salinity stress can be suggested for radish.
* Kobra Mahdavian
k.mahdavian@pnu.ac.ir
1
Department of Biology, Faculty of Science, Payame Noor
University, Tehran, Iran
Introduction
Salinity harms crop yield, which imposes ionic and osmotic
stresses, which lead to a decrease in mineral uptake, thereby
reducing plant growth and yield. Accumulation of sodium
ions leads to increased electrolyte leakage resulting in sec-
ondary stress, so that the production of reactive oxygen spe-
cies can damage the enzymatic activities of the cell [1].
Plant protect themselves with various defense systems,
including proline, glycine betaine, carotenoids, and enzy-
matic antioxidants such as ascorbate peroxidase, catalase,
superoxide dismutase, and glutathione reductase [1–3].
Some substances, such as SA and orthohydroxybenzoic acid,
decrease the adverse effects of NaCl stress on plants [2, 3].
SA is a phytohormone that increases the tolerance to
various abiotic stresses [4, 5], including salinity stress [5].
Decisive role of salicylic acid under stress may be due to
increased absorption of water and minerals, photosynthetic
regeneration, and membrane stability [6]. Salicylic acid is a
signal molecule and can interact with ROS signal pathways,
thus regulating environmental stresses against oxidative
stress [7]. It has been found that SA increases the activity
of antioxidant enzymes in various plants and reduces lipid
peroxidation [8].
In this study, to increase the growth and tolerance of rad-
ish to soil salinity, the effect of SA on reducing the effect of
salinity stress on growth and biochemical changes of radish
was investigated.
13
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Material and Methods
Plant Material
This experiment was designed and performed in July 2020
in the Biology Laboratory of Payame Noor University, Cen-
tral Kerman, in a factorial form in a completely randomized
design with three replications in greenhouse conditions.
Design of an Experiment
The radish seeds were first disinfected with 5% sodium
hypochlorite for two minutes and then washed three times
with distilled water. Five seeds were planted in pots with
field soil, washed sand, and leaf fertilizer in a ratio of 1:1:2
and in greenhouse conditions at 25–27 °C for 16 h (using
a combination of ordinary and incandescent bulbs) were
maintained.
The three- to the four-leaf stage plants were used in the
experiment. After two weeks, treatment began. Four con-
centrations of sodium chloride, including 0, 25, 50, and
100 mM, were used for irrigation. Four concentrations of
SA, including 0, 0.5, 1, and 1.5 mM, were used for foliar
application. The factorial experiment was conducted in a
completely randomized design with three replications in
greenhouse conditions. Salicylic acid was first sprayed on
the leaves every day for a week, always early morning with a
constant pressure spray and completely uniform. The amount
of foliar application was such that soluble droplets were vis-
ible on the leaves.
Preparation of SA
Also, for salicylic acid treatment, Triton X-100 in the
amount of one hundred percent was used as a surfactant,
and then salinity treatments were used every day for a week.
In addition to pot drainage, to prevent salt accumulation,
water washing was performed uniformly for all treatments
since salinity treatment.
To measure dry weight, stems and roots were isolated
from the plant 35 days after the start of the experiment and
dried at 70 °C for 48 h. After complete drying of the sam-
ples, their dry weight was measured.
The root and shoot length were measured one week after
the treatment. The length of the shoot from the collar to the
end of the shoot and the length of the root from the collar to
the root were measured.
Photosynthetic Pigment Determination
The carotenoids, chlorophyll a, b, and total chlorophyll
were measured by Lichtenthaler [9]. The amount of 0.2 g
13
K. Mahdavian
of detached fresh leaves was ground with 15 ml of 80%
acetone and filtered. The absorption was determined by
spectrophotometer (Carry 50 Australia) at wavelengths of
646.8, 663.20 and, 470 nm and their concentrations were
calculated using the following formulas in milligrams per
gram of fresh weight:
Chla = (12.25 A663.2 − 2.79 A646.8)
Chlb = (21.21 A646.8 − 5.1 A663.2)
ChlT = Chla + Chlb
Car = [1000 A470 − 1.8 Chla − 85.02 Chlb]∕198
In this formula, Chla, Chlb, ChlT, and Car are the concen-
trations of chlorophyll a, chlorophyll b, total chlorophyll, and
carotenoids (including carotenes and xanthophylls), respec-
tively [9].
Reactive Oxygen Scavenging Enzymes Determination
Malondialdehyde concentration was measured by the
method of Heath and Packer [10]. In this method, 0.2 g of
fresh leaf tissue was weighed and centrifuged in a mortar
containing 5 ml of 0.1% trichloroacetic. The resulting extract
was centrifuged at 1000 g for 5 min. 20 g of trichloroacetic
acid and 0.5 g of thiobarbituric acid was dissolved in 100 ml
of water, and then 4 ml of this solution (20% solution of
trichloroacetic acid containing 0.5% thiobarbituric acid) was
dissolved in 1 ml of the supernatant. The resulting solution
was heated in a hot water bath at 95° C for 30 min and imme-
diately cooled in ice. The solution was centrifuged at 1000 g
for 10 min and then placed on ice and absorbed at 532 nm
with a spectrophotometer. In this wavelength, the adsorp-
tion of the red thiobarbituric acid-malondialdehyde complex
was measured. Adsorption of other non-specific pigments
was determined and subtracted at 600 nm. The quenching
coefficient of 155 ­mmol−1 ­cm−1 was used to calculate the
concentration of malondialdehyde [10], and at 455 nm for
other aldehydes. Adsorption of other non-specific pigments
was also read at 600 nm and subtracted from this value.
For other aldehydes calculation, a quenching coefficient of
458 × ­102 ­mmol−1 ­cm−1 was used [11]. The extinction coef-
ficient is the average extinction coefficient of five aldehydes.
Proline was measured by Bates et al. [12] method. The
first 0.05 g of fresh leaves of the plants were weighed on
ice with three replications in each concentration. Samples
completely dissolve with 1.7 ml of sulfosalicylic acid and
centrifuge for 20 min. A 0.5 mL aliquot of the supernatant
was transferred to a tube containing 0.5 mL acetic acid
Application of Salicylic Acid on Chlorophyll, Carotenoids, and Proline in Radish Under Salinity… 811

and then 1 mL acid ninhydrin was added. The mixture was
boiled for one hour. The reaction was stopped in an asso-
ciate degree ice bath. The reaction mixture was extracted
with 1 mL toluene, mixed completely by the vortex. The
optical density of the upper toluene phase was determined
at a wavelength of 520 nm, using toluene as the blank, and
L-proline as a standard [12].
by the Duncan method at the level of 5% probability using
SPSS software. To evaluate the interaction of the two fac-
tors on the measured traits, data were analyzed by SPSS
software and a two-way analysis of variance.
Results and Discussion

The results of the analysis of variance 1 showed that the

Statistical Analysis effect of salinity and salicylic acid stress at the level of
1% probability on the shoot and root length was signifi-
Statistical analysis in this study was performed based on a cant. Also, their interaction at the level of 1% probabil-
completely randomized design. The means were compared ity on shoot length was significant. The results showed

Fig. 1  Effect of salicylic acid at

different salinity levels on shoot
a b a a
c d e
length (a) and root (b) radish. f
Columns with different letters 10 h i g
have a significant difference at j k
the 5% probability level l
Shoot length (cm)

8 m
n
0
6 o
SA 0.5
4 SA 1
SA 1.5
2

0
control 25 50 100
Salinity treatment (mM)

b18 a
b b a
16
14 c
d d e
e
Root length (cm)

12 f
g
10 i h 0
k j
8 l SA 0.5
6 SA 1
4 SA 1.5
2
0
control 25 50 100
Salinity treatment (mM)

13
812
that the salinity treatment at concentrations of 25, 50, and
100 mM reduced shoot and root lengths. The results of
combined treatment of SA and salinity show that com-
bined treatment of SA with concentrations of 0.5, 1 and,
1.5 mM and salinity with concentrations of 25, 50 and,
100 mM significantly increased shoot length and roots for
plants that have been treated only with salinity (Fig. 1). It
has been shown that in salinity-resistant species, growth
parameters decrease. However, in semi-susceptible and
susceptible species, it does not decrease [13]. Inhibition
of cell division expansion, reduction of leaf area, damage
to photosynthetic apparatus, change in photosynthetic pig-
ments, oxidative stress, and lipid peroxidation, are among
the reasons that have reduced plant growth under NaCl
stress [8, 14, 15]. Application of SA modulates the effects
of NaCl stress. Studies conducted by Mahdavian [14] also
showed that SA in plants under NaCl stress significantly
increases growth [14, 15]. In this study, it was determined
that salicylic acid increased the growth parameters in rad-
ish in comparison with control plants. Similar findings
have been reported by Mahdavian (2017) for barley and
sunflower, which shows that SA has a positive effect on
root and stem growth parameters [14, 15].
The results of the analysis of variance showed that the
effect of salinity stress and salicylic acid foliar application
at the level of 1% probability, as well as the effect of their
interaction at the level of 5% probability on the shoot and
dry root weight, were significant (Table 1). The results of
this study indicated that salinity stress (25, 50, and 100 mM)
decreased the dry weight of shoots and roots of radish plants.
However, foliar application of SA considerably improved
the dry weight of shoots and roots in radish plants by alter-
ing the biochemical and physiological processes (Fig. 2).
The hormones auxin and cytokinin decrease under stress,
their reduction slows growth. Therefore, salicylic acid can
improve growth in stressed plants by affecting the hormones
Table 1  Analysis of variance of trait size in radishes treated with salicylic acid at different salinity levels
Other aldehydes Malondialdehyde Shoot dry weight Root dry weight Shoot length Root length Degrees
001/0** 00,013/0** 006/0** 001/0**
001/0** 000,035/0** 008/0** 000,251/0**
000,007/0** 000,000/0* 000,129/0* 000,014/0*
66/3 36/4 45/4 05/4
*, ** and ns: are significant at the level of ≥ 5%, 1%, non-significant, respectively
13
K. Mahdavian
auxin and cytokinin [16]. It has also been reported that root
and shoot length have increased in sunflower [15], barley
[14], and bean [17] with the use of salicylic acid.
Salinity treatment has been reported to reduce some
growth parameters such as dry weight of sunflower roots
[3]. In this study, SA pretreatment improved the growth
parameters of stressed plants. Research results on tomatoes
[8], sunflower [15], barley [14] and, beans [17] have shown
that SA has reduced the harmful effects of salinity stress
on the dry and fresh weight of plants. The positive effect
of SA is additionally attributed as a plant growth regulator
because of its ability to increase nutrient uptake, enzymatic
activities, protein synthesis, photosynthetic activity, protect
against biotic and abiotic stresses, and increase in antioxi-
dant capacity of plant [18].
The results showed the analysis of variance Table 2 that
the effect of salinity and SA stress on the level of one percent
probability on chlorophyll and carotenoid concentration was
significant. The results of the effect of different NaCl con-
centrations show that chlorophyll a, b, total and carotenoid in
treatment with different NaCl concentrations of 25, 50, and
100 mM show a significant decrease compared with control
plants. Also, the combined application of SA and salinity
treatment show that salicylic acid foliar application improves
the chlorophyll concentration of radish leaves under salinity
stress compared to plants that were not affected by salicylic
acid treatment (Figs. 3, 4). Studies have shown that applica-
tion of SA improved the photosynthetic pigments of plants
under stress conditions, which may provide the ability of the
plant to tolerate environmental stresses by osmotic stress.
In this study, Chl (a, b and total) and carotenoids contents
were significantly decreased in plants grown in salinity stress
as compared to control. Similar results were found in sun-
flower, barley, and bean [17]. Salinity will induce changes in
the levels of pigments like Chl and carotenoids so that carot-
enoids are necessary for the integrity of the photosystem
Sources of variance
of free-
dom
277/25** 512/51** 3 Salinity
193/10** 866/32** 3 Salicylic acid
181/0** 039/0ns 9 Salicylic acid × Salinity
13/2 49/2 – Coefficient of variation (%)
Application of Salicylic Acid on Chlorophyll, Carotenoids, and Proline in Radish Under Salinity… 813

Fig. 2  Effect of combined sali- 0.2

a
cylic acid and salinity treatment a
on shoot dry weight (a) and root 0.18 bc ab bc
bc
(b) radish. Columns with dif-
0.16 c c

Shoot dry weight (g)

ferent letters have a significant d
difference at the 5% probability 0.14 e f
f f
level g
0.12 h
0
0.1 i
j SA 0.5
0.08
SA 1
0.06
0.04 SA 1.5
0.02
0
control 25 50 100
Salinity treatment (mM)

b 0.08 a
b a
0.07 b

0.06
cd c d cd
Root dry weight (g)

df
0.05 f e f f
g g 0
0.04
h SA 0.5
0.03
SA 1
0.02 SA 1.5
0.01
0
control 25 50 100
Salinity treatment (mM)

Table 2  Analysis of variance of trait size in radishes treated with salicylic acid at different salinity levels
Proline Total chlorophyll Chlorophyll b Chlorophyll a Carotenoids Degrees of Sources of variance
freedom

311/30** 386/11** 675/2** 557/4** 241/3** 3 Salinity

230/28** 221/17** 250/3** 740/3** 632/8** 3 Salicylic acid
303/0** 130ns/0 140/0* 031/0ns 128/0** 9 Salicylic acid × salinity
20/2 95/1 43/2 50/2 99/2 Coefficient of variation (%)

*, ** and ns: are significant at the level of five percent, one percent, non-significant, respectively

13
814 K. Mahdavian

Fig. 3  Effect of combined treat- 9 a

ment of salicylic acid and salin- a d b b e
c
f
ity on the amount of chlorophyll 8 g
a (a), chlorophyll b (b), and hi h h

Leaf chlorophyll a content

total chlorophyll (c) in radish. 7 j i
k k

(mg/g fresh weight)

Columns with different letters
have a significant difference at 6 l
the 5% probability level 0
5
4 SA 0.5
3 SA 1
2 SA 1.5
1
0
control 25 50 100
Salinity treatment (mM)

b 10 a
b
9 d cd c de d
e fg f fg
Leaf chlorophyll b content

8 g
h j j i
(mg/g fresh weight)

7
6
0
5
SA 0.5
4
SA 1
3
2 SA 1.5
1
0
control 25 50 100
Salinity treatment (mM)

c 20
18 a a
cd bc b d c
e d e
16
Total chlorophyll content

g f
h h
(mg/g fresh weight)

14 i j
12
0
10
SA 0.5
8
SA 1
6
4 SA 1.5
2
0
control 25 50 100
Salinity treatment (mM)

13
Application of Salicylic Acid on Chlorophyll, Carotenoids, and Proline in Radish Under Salinity…
Fig. 4  Effect of combined sali- 7
cylic acid and salinity treatment
Leaf carotenoid content (mg/g fresh
on carotenoid content in radish.
Columns with different letters 6
have a significant difference at
the 5% probability level 5 g
4 i
weight)
3
2
1
0
and will scavenge reactive oxygen species generated under
salinity conditions [16]. The application of salicylic acid
in chickpea [19], sunflower [15], barley [14] and bean [17]
plants increased the amount of chlorophyll in salinity con-
ditions. Salinity stress harms growth and chlorophyll con-
tent in the bean plant. However, plants treated with salicylic
acid had a higher dry and fresh weight of stems and roots
and higher chlorophyll content under salinity stress [17]. It
has been found that SA treatment increased carotenoids and
chlorophyll a, b under NaCl stress [17].
Carotenoids can protect chloroplast membranes because
they can release large amounts of energy in the form of heat
through photosystems I and II [20].
Based on the analysis of variance 1, the effect of salinity
stress, foliar application of SA, and their interaction on the
concentration of malondialdehyde and other aldehydes were
significant at the level of 1% probability. Leaves showed
higher values of MDA and other aldehydes under NaCl
stress, as compared to their respective controls. The addi-
tion of SA considerably reduced the production of MDA
and other aldehydes, in leaves at all levels of NaCl (Fig. 5).
In this study salinity stress increased MDA, which agrees
with previous research [17]. MDA is usually used to indicate
oxidative damage in fatty acids and its accumulation under
salinity stress has been determined in plants like cotton.
Increased lipid peroxidation, which measured by MDA accu-
mulation, could result from the generation of ROS under
salinity conditions. It can be hypothesized that the applica-
tion of SA includes a role in the induction of antioxidant
815
b a a
c c d
f e
f g
h
0
i
j
k SA 0.5
SA 1
SA 1.5
control 25 50 100
Salinity treatment (mM)
enzyme activity resulting in a decrease in ROS and lipid
peroxidation on a long term basis [17].
The analysis of variance showed that salinity stress, sali-
cylic acid, and their interaction were significant at the level
of 1% probability on proline concentration (Table 2). Proline
content in radish leaves increased, in 25, 50, and 100 mM of
NaCl exposed plants as compared to the control. The level of
proline in plants treated with SA improved in radish plants
under salinity (Fig. 6). Proline accumulation increased with
increasing salinity concentration in the present study. Simi-
lar reports have been reported for other plants [4, 17]. The
accumulation of proline was a typical response to salt stress
and was shown to be associated with increasing NaCl con-
centrations. It has been reported as a biochemical marker
for enhanced NaCl tolerance in potato plants [21, 22]. Pro-
line is involved in osmotic regulation and helps stabilize
membranes and maintain stressed protein structures [4, 5].
Proline has also been reported to have antioxidant proper-
ties that quench ROS, reduce damage to thylakoid mem-
branes, act as an energy store and enhance nitrogen fixation
in plants. SA increased proline content under salinity stress
in plants, and similar results have been reported in Vinga
radiate [4].
Conclusion
In this research, salinity stress affects growth parameters
and photosynthetic pigments in radish. The plants have
13
816 K. Mahdavian

Fig. 5  Effect of combined sali- a

0.009
cylic acid and salinity treatment a
on the amount of malondialde- 0.008 b

MDA content (µmol g-1 FW)

hyde (a) and other aldehydes
(b) in radish. Columns with dif- 0.007 c
ferent letters have a significant d d
difference at the 5% probability 0.006 d
e
level
0.005 f fg g 0
g h
0.004 i SA 0.5
j
0.003 k SA 1
0.002 l SA1.5
0.001
0
control 25 50 100
Salinity treatment (mM)

b 0.06

0.05 a
Other aldehyde content

0.04
(µmol g-1 FW)

b
c c 0
0.03 d
e
SA 0.5
f
0.02 g SA 1
h i h
j k
k
l SA 1.5
0.01 m

0
control 25 50 100
Salinity treatment (mM)

mechanisms to deal with these types of environmental
stresses. Our results showed that salinity reduces dry weight,
root and shoot growth, carotenoids, and chlorophyll; how-
ever, according to the results mentioned in this study, in
pre-treated plants with salicylic acid, this reduction has
been modified. On the other hand, salinity significantly
increased the amount of proline, malondialdehyde, and other
aldehydes; while salicylic acid modulates plant’s tolerance
to salinity stress. This research showed that radish pretreat-
ment with concentrations of 0.5, 1, and 1.5 mM SA increases
plant tolerance to salinity stress. In general, the use of SA
in reducing the destructive effects of salinity stress can be
suggested for radish.

13
Application of Salicylic Acid on Chlorophyll, Carotenoids, and Proline in Radish Under Salinity…
Fig. 6  The effect of combined 8
treatment of salicylic acid
and salinity on the amount of 7 b
Proline content (µmol g-1 FW)
proline in radish. Columns with
different letters have a signifi-
6
cant difference at the level of
five percent probability
5
4 f
3 h
i j
k l
2 m
1 o
0
control
Salinity treatment (mM)
Acknowledgements I wish to thank Payame Noor University
Research Council for approval and providing financial support.
Declarations
Conflict of interest The authors declare that they have no conflict
of interest.
References
1. Ahmad P, Hashem A, Abd-Allah EF, Alqarawi AA, John R,
Egamberdieva D (2015) Role of Trichoderma harzianum in miti-
gating NaCl stress in Indian mustard (Brassica juncea L.) through
antioxidative defense system. Front Plant Sci 6:868
2. Ahmad P, Latef AA, Hashem AE, Abd-Allah F, Gucel S, Tran
LSP (2016) Nitric oxide mitigates salt stress by regulating levels
of osmolytes and antioxidant enzymes in chickpea. Front Plant
Sci 7:347
3. Ahanger M, Agarwal ARM (2017) Potassium up-regulates anti-
oxidant metabolism and alleviates growth inhibition under water
and osmotic stress in wheat (Triticum aestivum L.). Protoplasma
254(4):1471–1486
4. Khan MIR, Asgher M, Khan NA (2014) Alleviation of salt-
induced photosynthesis and growth inhibition by salicylic acid
involves glycine betaine and ethylene in mung bean (Vigna
radiata L.). Plant Physiol Biochem 80:67–74
5. Kaya C, Kirnak H, Higgs D, Saltali K (2002) Supplementary
calcium enhances plant growth and fruit yield in strawberry
cultivars grown at high salinity. Sci Hortic 93:65–74
6. Ahmad P, Nabi G, Ashraf M (2011) Cadmiuminduced oxidative
damage in mustard [Brassica juncea L. Czern. & Coss.] plants
can be alleviated by salicylic acid. S Afr J Bot 77:36–44
7. Borsani O, Valpuesta V, Botella MA (2001) Evidence for a role
of salicylic acid in the oxidative damage generated by NaCl
and osmotic stress in Arabidopsis seedlings. Plant Physiol
126:1024–1030
8. Shibli RA, Kushad M, Yousef GG, Lila MA (2007) Physio-
logical and biochemical responses of tomato micro shoots to
817
a
c
d
e
0
g
f
SA 0.5
SA 1
n SA 1.5
25 50 100
induced salinity stress with associated ethylene accumulation.
Plant Growth Regul 51:159–169
9. Lichtenthaler HK (1987) Chlorophylls and carotenoids: pigments
of photosynthetic biomembranes. Methods Enzymol 148:350–382
10. Heath RL, Packer L (1968) Photoperoxidation in isolated cholo-
roplast. I. Kinetics and stoichiometry of fatty acid peroxidation.
Arch Biochem Biophys 125:189–198
11. Meirs S, Aharoni N (1992) Ethylene increased accumulation of
fluorescent lipid-peroxidation products detected during parsley by
a newly developed method. J Am Soc Hortic Sci 117:128–132
12. Bates LS, Waldren RP, Tear ID (1973) Rapid determination of
free proline for water stress studies. Plant Soil 39:205–207
13. Kao WY, Tsai TT, Tsai HC, Shih CN (2006) Response of three
glycine species to salt stress. Environ Exp Bot 56:120–125
14. Mahdavian K (2017) Effect of different concentrations of salicylic
acid on salt tolerance of barley (Hordeum vulgare L.). J Crop
Physiol 36:121–136
15. Mahdavian K (2017) The effect of different concentrations of sali-
cylic acid on adjustment of the effects of sodium chloride stress
on growth parameters and photosynthetic pigments in sunflower
plant (Helianthus annuus L.). J Plant Environ Physiol 47:93–106
16. Sakhabutdinova AR, Fatkhutdinova DR, Bezrukova MV, Shaki-
rova FM (2003) Salicylic acid prevents the damaging action of
stress factors on wheat plants. Bulg J Plant Physiol 1:314–319
17. Ahmad P, Alyemeni MN, Ahanger MA, Egamberdieva D, Wijaya
L, Alam P (2018) Salicylic acid (SA) induced alterations in
growth, biochemical attributes and antioxidant enzyme activity
in Faba Bean (Vicia faba L.) seedlings under NaCl toxicity. Russ
J Plant Physiol 65(1):104–114
18. Gunes A, Inal A, Alpaslam M, Erslan F, Bagsi EG, Cicek N
(2007) Salicylic acid induced changes on some physiological
parameters symptomatic for oxidative stress and mineral nutri-
tion in maize (Zea mays L.) grown under salinity. J Plant Physiol
164:728–736
19. Popova LP, Maslenkova LT, Yordanova RY, Ivanova AP, Krantev
AP, Szalai G (2009) Exogenous treatment with salicylic acid
attenuates cadmium toxicity in Pea seedlings. Plant Physiol Bio-
chem 47:224–231
20. Juan M, Rivero RM, Romero L, Rviz JM (2005) Evaluation of
some nutritional and biochemical indicators in selecting salt-
resistant tomato cultivars. Environ Exp Bot 54:193–201
13
818 K. Mahdavian

21. Khodary SEA (2004) Effect of salicylic acid on the growth, pho-
tosynthesis and carbohydrate metabolism in salt-stressed maize
plants. J Agric Biol 6:5–8
22. Khan MIR, Syeed S, Nazar R, Anjum NA (2012) An insight into
the role of salicylic acid and jasmonic acid in salt stress tolerance.
In: Khan NA, Nazar R, Iqbal N, Anjum NA (eds) Phytohormones
and abiotic stress tolerance in plants. Springer, New York, pp
277–300
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