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- Introduction

ity stress, reduce plant growth and yield. Salinity stress is
one of the most common stresses that cause soil and water Salinity harms crop yield, which imposes ionic and osmotic
pollution worldwide. Salicylic acid (SA) is a plant hormone stresses, which lead to a decrease in mineral uptake, thereby
that has been shown to have therapeutic effects against non- reducing plant growth and yield. Accumulation of sodium
biological stresses such as salinity. In this study, to increase ions leads to increased electrolyte leakage resulting in sec-
radish growth and tolerance to soil salinity, the effect of SA ondary stress, so that the production of reactive oxygen spe-
on reducing the effect of salinity stress on radish growth cies can damage the enzymatic activities of the cell [1].
and biochemical changes was investigated. Results showed Plant protect themselves with various defense systems,
that salinity (0, 25, 50, and 100 mM) significantly reduced including proline, glycine betaine, carotenoids, and enzy-
root and shoot growth, reduced root and shoot dry weight, matic antioxidants such as ascorbate peroxidase, catalase,
and also decreased chlorophyll and carotenoids, in contrast, superoxide dismutase, and glutathione reductase [1–3].
in plants pre-treated with SA (0, 0.5, 1, and 1.5 mM) was Some substances, such as SA and orthohydroxybenzoic acid,
reduced the response. Salinity significantly increased the decrease the adverse effects of NaCl stress on plants [2, 3].
protective metabolites in radish antioxidant system like pro- SA is a phytohormone that increases the tolerance to
line, malondialdehyde, and other aldehydes. While modulat- various abiotic stresses [4, 5], including salinity stress [5].
ing salicylic acid, this increase has increased the tolerance Decisive role of salicylic acid under stress may be due to
of this plant to salinity stress. The application of SA can increased absorption of water and minerals, photosynthetic
increase plant tolerance to salinity stress. regeneration, and membrane stability [6]. Salicylic acid is a
signal molecule and can interact with ROS signal pathways,
Keywords Lipid peroxidation · Proline · Chlorophyll · thus regulating environmental stresses against oxidative
Carotenoids · Plant species 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-
Significance Statement: In this research, salinity stress affects ish to soil salinity, the effect of SA on reducing the effect of
growth parameters and photosynthetic pigments in radish. Also, salinity stress on growth and biochemical changes of radish
the results showed that radish pretreatment with concentrations was investigated.
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
[email protected]
1
Department of Biology, Faculty of Science, Payame Noor
University, Tehran, Iran

13
Vol.:(0123456789)
810 K. Mahdavian

Material and Methods of detached fresh leaves was ground with 15 ml of 80%
acetone and filtered. The absorption was determined by
Plant Material spectrophotometer (Carry 50 Australia) at wavelengths of
646.8, 663.20 and, 470 nm and their concentrations were
This experiment was designed and performed in July 2020 calculated using the following formulas in milligrams per
in the Biology Laboratory of Payame Noor University, Cen- gram of fresh weight:
tral Kerman, in a factorial form in a completely randomized
Chla = (12.25 A663.2 − 2.79 A646.8)
design with three replications in greenhouse conditions.

Design of an Experiment Chlb = (21.21 A646.8 − 5.1 A663.2)

The radish seeds were first disinfected with 5% sodium ChlT = Chla + Chlb
hypochlorite for two minutes and then washed three times
with distilled water. Five seeds were planted in pots with Car = [1000 A470 − 1.8 Chla − 85.02 Chlb]∕198
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 In this formula, Chla, Chlb, ChlT, and Car are the concen-
a combination of ordinary and incandescent bulbs) were trations of chlorophyll a, chlorophyll b, total chlorophyll, and
maintained. carotenoids (including carotenes and xanthophylls), respec-
The three- to the four-leaf stage plants were used in the tively [9].
experiment. After two weeks, treatment began. Four con-
centrations of sodium chloride, including 0, 25, 50, and Reactive Oxygen Scavenging Enzymes Determination
100 mM, were used for irrigation. Four concentrations of
SA, including 0, 0.5, 1, and 1.5 mM, were used for foliar Malondialdehyde concentration was measured by the
application. The factorial experiment was conducted in a method of Heath and Packer [10]. In this method, 0.2 g of
completely randomized design with three replications in fresh leaf tissue was weighed and centrifuged in a mortar
greenhouse conditions. Salicylic acid was first sprayed on containing 5 ml of 0.1% trichloroacetic. The resulting extract
the leaves every day for a week, always early morning with a was centrifuged at 1000 g for 5 min. 20 g of trichloroacetic
constant pressure spray and completely uniform. The amount acid and 0.5 g of thiobarbituric acid was dissolved in 100 ml
of foliar application was such that soluble droplets were vis- of water, and then 4 ml of this solution (20% solution of
ible on the leaves. trichloroacetic acid containing 0.5% thiobarbituric acid) was
dissolved in 1 ml of the supernatant. The resulting solution
Preparation of SA 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
Also, for salicylic acid treatment, Triton X-100 in the for 10 min and then placed on ice and absorbed at 532 nm
amount of one hundred percent was used as a surfactant, with a spectrophotometer. In this wavelength, the adsorp-
and then salinity treatments were used every day for a week. tion of the red thiobarbituric acid-malondialdehyde complex
In addition to pot drainage, to prevent salt accumulation, was measured. Adsorption of other non-specific pigments
water washing was performed uniformly for all treatments was determined and subtracted at 600 nm. The quenching
since salinity treatment. coefficient of 155 ­mmol−1 ­cm−1 was used to calculate the
To measure dry weight, stems and roots were isolated concentration of malondialdehyde [10], and at 455 nm for
from the plant 35 days after the start of the experiment and other aldehydes. Adsorption of other non-specific pigments
dried at 70 °C for 48 h. After complete drying of the sam- was also read at 600 nm and subtracted from this value.
ples, their dry weight was measured. For other aldehydes calculation, a quenching coefficient of
The root and shoot length were measured one week after 458 × ­102 ­mmol−1 ­cm−1 was used [11]. The extinction coef-
the treatment. The length of the shoot from the collar to the ficient is the average extinction coefficient of five aldehydes.
end of the shoot and the length of the root from the collar to Proline was measured by Bates et al. [12] method. The
the root were measured. first 0.05 g of fresh leaves of the plants were weighed on
ice with three replications in each concentration. Samples
Photosynthetic Pigment Determination completely dissolve with 1.7 ml of sulfosalicylic acid and
centrifuge for 20 min. A 0.5 mL aliquot of the supernatant
The carotenoids, chlorophyll a, b, and total chlorophyll was transferred to a tube containing 0.5 mL acetic acid
were measured by Lichtenthaler [9]. The amount of 0.2 g

13
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 by the Duncan method at the level of 5% probability using
boiled for one hour. The reaction was stopped in an asso- SPSS software. To evaluate the interaction of the two fac-
ciate degree ice bath. The reaction mixture was extracted tors on the measured traits, data were analyzed by SPSS
with 1 mL toluene, mixed completely by the vortex. The software and a two-way analysis of variance.
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]. 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 K. Mahdavian

that the salinity treatment at concentrations of 25, 50, and auxin and cytokinin [16]. It has also been reported that root
100 mM reduced shoot and root lengths. The results of and shoot length have increased in sunflower [15], barley
combined treatment of SA and salinity show that com- [14], and bean [17] with the use of salicylic acid.
bined treatment of SA with concentrations of 0.5, 1 and, Salinity treatment has been reported to reduce some
1.5 mM and salinity with concentrations of 25, 50 and, growth parameters such as dry weight of sunflower roots
100 mM significantly increased shoot length and roots for [3]. In this study, SA pretreatment improved the growth
plants that have been treated only with salinity (Fig. 1). It parameters of stressed plants. Research results on tomatoes
has been shown that in salinity-resistant species, growth [8], sunflower [15], barley [14] and, beans [17] have shown
parameters decrease. However, in semi-susceptible and that SA has reduced the harmful effects of salinity stress
susceptible species, it does not decrease [13]. Inhibition on the dry and fresh weight of plants. The positive effect
of cell division expansion, reduction of leaf area, damage of SA is additionally attributed as a plant growth regulator
to photosynthetic apparatus, change in photosynthetic pig- because of its ability to increase nutrient uptake, enzymatic
ments, oxidative stress, and lipid peroxidation, are among activities, protein synthesis, photosynthetic activity, protect
the reasons that have reduced plant growth under NaCl against biotic and abiotic stresses, and increase in antioxi-
stress [8, 14, 15]. Application of SA modulates the effects dant capacity of plant [18].
of NaCl stress. Studies conducted by Mahdavian [14] also The results showed the analysis of variance Table 2 that
showed that SA in plants under NaCl stress significantly the effect of salinity and SA stress on the level of one percent
increases growth [14, 15]. In this study, it was determined probability on chlorophyll and carotenoid concentration was
that salicylic acid increased the growth parameters in rad- significant. The results of the effect of different NaCl con-
ish in comparison with control plants. Similar findings centrations show that chlorophyll a, b, total and carotenoid in
have been reported by Mahdavian (2017) for barley and treatment with different NaCl concentrations of 25, 50, and
sunflower, which shows that SA has a positive effect on 100 mM show a significant decrease compared with control
root and stem growth parameters [14, 15]. plants. Also, the combined application of SA and salinity
The results of the analysis of variance showed that the treatment show that salicylic acid foliar application improves
effect of salinity stress and salicylic acid foliar application the chlorophyll concentration of radish leaves under salinity
at the level of 1% probability, as well as the effect of their stress compared to plants that were not affected by salicylic
interaction at the level of 5% probability on the shoot and acid treatment (Figs. 3, 4). Studies have shown that applica-
dry root weight, were significant (Table 1). The results of tion of SA improved the photosynthetic pigments of plants
this study indicated that salinity stress (25, 50, and 100 mM) under stress conditions, which may provide the ability of the
decreased the dry weight of shoots and roots of radish plants. plant to tolerate environmental stresses by osmotic stress.
However, foliar application of SA considerably improved In this study, Chl (a, b and total) and carotenoids contents
the dry weight of shoots and roots in radish plants by alter- were significantly decreased in plants grown in salinity stress
ing the biochemical and physiological processes (Fig. 2). as compared to control. Similar results were found in sun-
The hormones auxin and cytokinin decrease under stress, flower, barley, and bean [17]. Salinity will induce changes in
their reduction slows growth. Therefore, salicylic acid can the levels of pigments like Chl and carotenoids so that carot-
improve growth in stressed plants by affecting the hormones enoids are necessary for the integrity of the photosystem

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 Sources of variance
of free-
dom

001/0** 00,013/0** 006/0** 001/0** 277/25** 512/51** 3 Salinity

001/0** 000,035/0** 008/0** 000,251/0** 193/10** 866/32** 3 Salicylic acid
000,007/0** 000,000/0* 000,129/0* 000,014/0* 181/0** 039/0ns 9 Salicylic acid × Salinity
66/3 36/4 45/4 05/4 13/2 49/2 – Coefficient of variation (%)

*, ** and ns: are significant at the level of ≥ 5%, 1%, non-significant, respectively

13
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… 815

Fig. 4  Effect of combined sali- 7

cylic acid and salinity treatment

Leaf carotenoid content (mg/g fresh

b a a
on carotenoid content in radish.
Columns with different letters 6
c c d
have a significant difference at f e
the 5% probability level 5 g f g
h
4 i 0

weight)
i
j
k SA 0.5
3
SA 1
2
SA 1.5
1

0
control 25 50 100
Salinity treatment (mM)

and will scavenge reactive oxygen species generated under enzyme activity resulting in a decrease in ROS and lipid
salinity conditions [16]. The application of salicylic acid peroxidation on a long term basis [17].
in chickpea [19], sunflower [15], barley [14] and bean [17] The analysis of variance showed that salinity stress, sali-
plants increased the amount of chlorophyll in salinity con- cylic acid, and their interaction were significant at the level
ditions. Salinity stress harms growth and chlorophyll con- of 1% probability on proline concentration (Table 2). Proline
tent in the bean plant. However, plants treated with salicylic content in radish leaves increased, in 25, 50, and 100 mM of
acid had a higher dry and fresh weight of stems and roots NaCl exposed plants as compared to the control. The level of
and higher chlorophyll content under salinity stress [17]. It proline in plants treated with SA improved in radish plants
has been found that SA treatment increased carotenoids and under salinity (Fig. 6). Proline accumulation increased with
chlorophyll a, b under NaCl stress [17]. increasing salinity concentration in the present study. Simi-
Carotenoids can protect chloroplast membranes because lar reports have been reported for other plants [4, 17]. The
they can release large amounts of energy in the form of heat accumulation of proline was a typical response to salt stress
through photosystems I and II [20]. and was shown to be associated with increasing NaCl con-
Based on the analysis of variance 1, the effect of salinity centrations. It has been reported as a biochemical marker
stress, foliar application of SA, and their interaction on the for enhanced NaCl tolerance in potato plants [21, 22]. Pro-
concentration of malondialdehyde and other aldehydes were line is involved in osmotic regulation and helps stabilize
significant at the level of 1% probability. Leaves showed membranes and maintain stressed protein structures [4, 5].
higher values of MDA and other aldehydes under NaCl Proline has also been reported to have antioxidant proper-
stress, as compared to their respective controls. The addi- ties that quench ROS, reduce damage to thylakoid mem-
tion of SA considerably reduced the production of MDA branes, act as an energy store and enhance nitrogen fixation
and other aldehydes, in leaves at all levels of NaCl (Fig. 5). in plants. SA increased proline content under salinity stress
In this study salinity stress increased MDA, which agrees in plants, and similar results have been reported in Vinga
with previous research [17]. MDA is usually used to indicate radiate [4].
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- Conclusion
mulation, could result from the generation of ROS under
salinity conditions. It can be hypothesized that the applica- In this research, salinity stress affects growth parameters
tion of SA includes a role in the induction of antioxidant 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 aldehydes; while salicylic acid modulates plant’s tolerance
stresses. Our results showed that salinity reduces dry weight, to salinity stress. This research showed that radish pretreat-
root and shoot growth, carotenoids, and chlorophyll; how- ment with concentrations of 0.5, 1, and 1.5 mM SA increases
ever, according to the results mentioned in this study, in plant tolerance to salinity stress. In general, the use of SA
pre-treated plants with salicylic acid, this reduction has in reducing the destructive effects of salinity stress can be
been modified. On the other hand, salinity significantly suggested for radish.
increased the amount of proline, malondialdehyde, and other

13
Application of Salicylic Acid on Chlorophyll, Carotenoids, and Proline in Radish Under Salinity… 817

Fig. 6  The effect of combined 8

treatment of salicylic acid a
and salinity on the amount of 7 b

Proline content (µmol g-1 FW)

proline in radish. Columns with
different letters have a signifi- c
6
cant difference at the level of d
five percent probability e
5
0
4 f g
f
SA 0.5
3 h
i j SA 1
k l
2 m n SA 1.5
1 o
0
control 25 50 100
Salinity treatment (mM)

Acknowledgements I wish to thank Payame Noor University induced salinity stress with associated ethylene accumulation.
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Conflict of interest The authors declare that they have no conflict
Arch Biochem Biophys 125:189–198
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21. Khodary SEA (2004) Effect of salicylic acid on the growth, pho- Springer Nature or its licensor (e.g. a society or other partner) holds
tosynthesis and carbohydrate metabolism in salt-stressed maize exclusive rights to this article under a publishing agreement with the
plants. J Agric Biol 6:5–8 author(s) or other rightsholder(s); author self-archiving of the accepted
22. Khan MIR, Syeed S, Nazar R, Anjum NA (2012) An insight into manuscript version of this article is solely governed by the terms of
the role of salicylic acid and jasmonic acid in salt stress tolerance. such publishing agreement and applicable law.
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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