Pak. j. life soc. Sci.

(2019), 17(2): 78-85 E-ISSN: 2221-7630; P-ISSN: 1727-4915

Pakistan Journal of Life and Social Sciences

www.pjlss.edu.pk

RESEARCH ARTICLE
Effects of Aerial Application of Salicylic Acid on Growth, Pigment
Concentration, Ions Uptake and Mitigation of Salinity Stress in Two
Varieties of Wheat (Triticum aestivum L.)
Sib g h a No r een * , Am in a Sh ah e en , Kau sar Hu s s ai n Sh ah an d U m m e Am m ara
In st itu te o f Pu r e & A p p lied B io lo g y (IP & A B), B ah au d d in Zak a riy a Un iv er si ty Mu lt an ,
Pak is tan

ARTICLE INFO ABSTRACT

Received: Mar 05, 2019 The agricultural crops are greatly impacted by climate change, eco-edaphic factors
Accepted: Dec 01, 2019 and salt stress. This study was conducted to assess the negative effects of salts stress
on two varieties of wheat crop i.e. S-24 and MH-97. It was aimed to determine the
Keywords proportionate extent of stress alleviation induced by salicylic acid (SA) and its
Salicylic acid impact on biological yield, relative water content, chlorophyll content, Na +/K+ ions
Salinity stress and other physiological and biochemical attributes in wheat varieties. The plants
Sustainable agriculture were challenged with salinity stress (200 mM NaCl) and the effects of foliar spray of
Triticum aestivum L. SA (200 mg L-1) were investigated. The shoot fresh weight was reduced by 30%
Wheat under saline stress, but exogenously applied SA caused an increase in shoot fresh
weight by 11.1%. The higher quantity of biological yield by 21.9% was obtained
from variety MH-97 compared to variety S-24. The varieties differed greatly in
maintenance of total soluble proteins, while a little affected by salt stress. The
varieties S-24 and MH-97 maintained the quantity of total soluble proteins by 18.7
and 11.3 mg/g, respectively. The crop treated with SA at the rate of 200 mg L-1
produced 69.6% higher total soluble proteins compared to unsprayed crop. The
varieties differed significantly in relation to total free amino acids. The variety MH-
97 contained 18.1% higher content compared to variety S-24 under saline
environment. The spray of SA enhanced the amount of total free amino acids from
12.29 to 19.42 mg/g under saline condition. The SPAD values decreased from 47.15
to 41.62 and 44.52 to 40.58 in varieties S-24 and MH-97, respectively grown on
untreated and treated soils with 200 mM salinity. Relative water contents (RWC)
were reduced by 16.1% under saline condition. The application of SA induced the
*Corresponding Author: uptake of K+ ion from soil medium. The results have revealed that salt stress in
sibgha.noreen@bzu.edu.pk wheat crop could be reduced by foliar spray of salicylic acid.

INTRODUCTION quantity and quality of the product (Hartmann et al.,

1990). There are different drastic effects of salinity
The agricultural production in arid and semi-arid land is stress on plants. The limited growth rate of plants in
significantly affected by salinity stress. The saline area is due to substantial absorption of the salts,
accumulation of soluble salts poses devastating effects particularly the Sodium ions (Na+) and Chloride ions
on the production of crop plants (Zhu, 2001). (Cl-) by plant tissues. It results in restricted water
Approximately six million hectares of the irrigated land absorption, damage to the root system, necrosis in
is affected by salinity in Pakistan (parc.gov.pk). Most leaves and progressive wilting that may lead to plant
of the cultivated land has salinity patches. The death (Husen et al., 2018; Ashraf, 2010). The uptake of
economic production of various crops in such area is a greater amounts of toxic ions lowers the turgor potential
challenging task for the farmers because they have to of plants that is main driving force for sustaining the
face problems in seedlings establishment, restricted plant growth and development (Shabala et al., 2012;
plant growth and development and substantial loss in Farooq et al., 2010). The turgidity of leaves, relative

78
Foliarly applied salicylic acid to reduce salt stress in wheat

water contents, accumulation of proteins, biomass drought stress. Babar et al. (2014) reported the
production, and proportionate absorption and restriction mitigation of the negative effects of salinity in
of toxic ions with concurrent assimilation of K+ by Fenugreek by foliar application of SA. Likewise,
plant tissues influence greatly on crop productivity Nahrjoo and Sedaghathoor (2018) investigated the
(Ashraf, 2010). The activity of photosystem I and II is induction of salinity stress resistance in Rosemary by
negatively affected due to closure of stomata, which applying Jasmonic acid and SA and. Farouk and Arafa
results in bringing various changes in the (2018) reported the use of sodium nitroprusside for the
morphological, hormonal and physiological processes mitigation of salinity stress in Canola.
under salt stress conditions (Ashraf et al., 2011). The Wheat (Triticum aestivum L.) is a very important
The different crop plants respond differently to the food crop in the world that is cultivated on more than
salinity stress. Their response is largely dependent upon 8.66 million hectares land. Pakistan harvested 26.3
the variations at the molecular, physiological and million tonnes of Wheat grains during the year 2017-18
biochemical levels (Ashraf, 2004). It may vary in the (https://www.world-grain.com/articles/10791-pakistan-
accumulation level and appearance of toxic Na+ and Cl- wheat-production-up-in-2017-18). Some of the
ions, production of reactive oxygen species (ROS), agriculture land in Pakistan both irrigated and rainfed,
absorption level of the essential nutrients and the is affected by salinity and/or sodicity to certain extent.
balance of phytohormones required for normal growth The cultivation of Wheat in such area poses a
of plants (Hayat et al., 2005; Dat et al., 2000). The challenging task to produce significant yield of good
ability of plant species to address drought condition quality grains. Therefore, it is needed to look for
under salt stress environment is adversely affected by possible measures to alleviate the drastic effects of
the amount of generation of ROS. These factors cause salinity stress on crop plants for the sake of sustainable
enhancement in lipid peroxidation and deterioration of agriculture. Since SA is reported to be an active agent
cell membrane structures (Joseph et al., 2010). that strengthens plant tolerance level against various
The salicylic acid (SA) is a ubiquitous endogenous stresses, therefore, this study was designed to
compound in plants that can induce tolerance to salinity investigate the ability of SA to mitigate the salinity
stress (Noreen et al., 2013; Ashraf, 2010); water stress stress in two selected varieties of the Wheat crop under
(Ashraf et al., 2011; Singh and Usha, 2003); high local environment.
temperature stress (Krasensky and Jonak, 2012) and
generation of reactive oxygen species (ROS) (Noreen et MATERIALS AND METHODS
al., 2009). The SA improves plants osmotic adjustment
under stressful conditions (Jones and Turner 1980). It is
To investigate the response of two varieties of Wheat
responsible for regulating photosynthetic and
(S-24 and MH-97) towards foliar application of SA and
evapotranspiration rate (Ahmad et al., 2013). It
its efficacy to mitigate salinity stress, a pot experiment
enhances the amount of chlorophyll contents (Noreen et
was conducted in complete randomized design with
al., 2013); induces the over-production of organic
four replications at the Botanic Garden, Institute of
solutes (Ashraf, 2010) and restricts the assimilation of
Pure and Applied Biology (IP&AB), Bahuddin
Na+ and Cl- ions (Noreen and Ashraf, 2008). It is
helpful in longer retention of flowers (Hayat and Zakariya University Multan, Pakistan. Each pot (24.5
Ahmad, 2007); in greater quenching of ROS by cm diameter and 28.0 cm deep) was filled with 12 Kg
upregulating the antioxidant defense system (Sharma dry sand, irrigated with water and the seeds were sown.
and Dubey, 2005) and in attaining higher turgor Irrigation was maintained with full strength Hoagland’s
potential under stressful conditions (Mutlu et al., 2013). nutrients solution (Epstein, 1972). After ten days of
It helps in the ROS scavenging and enhances the germination, thinning was practiced leaving five
capability of plants to overcome the deterioration of cell uniform seedlings in each pot. After 18 days of
membrane leading towards improved osmotic germination, the seedlings were challenged with
adjustment in plants (Ajithkumar and Panneerselevam, salinity stress (200 mM NaCl) by administering first
2014). treatment of 50 mM NaCl along with full strength
There are several reports of the SA induced mitigation Hoagland’s nutrients solution followed by same
of abiotic stresses in crop plants. Hamada and Al- treatment for second, third and fourth time on every
Hakimi (2001) investigated alleviation of the drastic second day. The plants were regularly irrigated with
effects of salinity stress in Wheat seedlings by spraying nutrients solution every week. The plants were also
100 mg L-1 SA. El-Tayeb (2005) reported that sprayed with 100 ml deionized water to stabilize plants
increasing levels of NaCl in rooting zones resulted in under salinity stress. After 26 days of germination, the
the increased amount of soluble proteins in the roots of plants were sprayed with Aqueous solution of SA (200
barley seedlings. Ilyas et al. (2017) studied the mg L-1) supplemented with 0.1 % (w/v) Tween-20 at
influence of SA and Jasmonic Acid on Wheat under 08:00 to 10:00 AM in the morning.

79
Noreen et al

After three weeks of the SA treatment, the total varieties alone and/or interaction with salinity regimes
chlorophyll contents were estimated with the help of a did not show any significant differences amongst
portable SPAD chlorophyll meter by randomly themselves in this parameter. Averaged across SA
selecting the fully expanded leaf from the apex. The levels and salinity regimes 21.99% higher biological
relative water contents were measured by following the yield was obtained by variety MH-97 in comparison
method of Jones and Turner (1980) by collecting fully with variety S-24. The data further revealed that
expanded leaves from the top. Thereafter, the plants biological yield was reduced by 34.23% in crop grown
were harvested and the roots were washed with water. under salt stress (Fig. 1). The salt stress and interaction
The shoots and roots samples were weighed by using a of varieties x SA levels produced significant effect in
digital balance to record fresh weight. Afterwards these reducing the biological yield, however, a little
samples were oven dried at 70°C for 48 hours and the influenced by other interactive treatments. A higher
shoot and root dry mass was measured to determine the quantity of biological yield by 15.16% was resulted
biological yield. The fresh plant material was used to from variety MH-97 in comparison with variety S-24.
determine the total free amino acids and total soluble The spray of 200 mg L-1 SA resulted in the
proteins contents by employing the method of Bradford improvement of biological yield by 13.7% in
(1976). The plant material was used for the analysis of comparison with unsprayed crop. Overall, the wheat
Sodium and Potassium ions concentration by following varieties grown under salinity stress produced lower
the methods of Ryan et al., (2001). The data were biological yield compared with under normal condition.
subjected to statistical analysis by using MSTAT-C The amount of total soluble proteins (TSPs) was
computer Package (Cohart Software, Berkley, CA). The significantly decreased due to different varieties and SA
least significant differences (LSD) were calculated by levels, however, salts stress did not have any positive
using technique of Snedecor and Cochran (1980). effect. The various interactive treatments (salinity
regimes x varieties and salinity regimes x SA levels x
RESULTS varieties) caused significant effect in maintaining
differential quantities of total soluble protein in leaf
The results showed that the Wheat variety S-24 was tissues (Fig. 2). The varieties produced pronounced
relatively better in growth as compared to the variety effects in maintaining the differential quantities in the
MH-97 under normal condition. The salinity stress has leaf tissues. An amount of 18.7 mg g-1 of the TSPs was
significantly decreased the growth of both varieties accumulated in variety MH-97, while it was 11.25 mg
however this decrease was relatively more in variety S- g-1 in variety S-24. Thereby, variety S-24 accumulated
24. The fresh shoot weight was significantly affected by 67.11% higher amount of TSPs in comparison with
foliar spray of SA and salinity levels respectively. variety MH-97. Furthermore, the spray of SA proved to
However, there was a little difference among wheat be good in maintaining greater amount of TSPs in
varieties, MH-97 and S-24. However, it was little wheat varieties. An improvement of 69.58% TSPs was
affected by the interactive effects of varieties x SA observed by exogenous application of SA over the
levels, varieties x salinity regimes, SA levels x salinity unsprayed crop. The interactive treatment of salinity
and varieties x SA levels and salinity regimes (Fig. 1). regimes and SA levels also produced significant
The salinity stress reduced the fresh shoot weight by (P<0.01) effects in maintaining higher amount of TSPs.
40% and 28% in varieties S-24 and MH-97, The varieties S-24 and MH-97 maintained 44.66% and
respectively. Overall, the salt stress caused reduction in 40.59% higher quantities of TSPs under salinity stress
fresh shoot weight by 34%. However, the variety S-24 compared to non-saline conditions.
and MH-97 sprayed with SA produced 10% and 05% There were significant differences in total free amino
higher fresh shoot weight respectively under saline acids (FAAs) due to effects produced by interaction of
conditions, whereas SA spray resulted in 13% and 17% varieties to salinity regimes, while a little affected by
higher fresh weight in variety S-24 and MH-97 spraying of SA. The differences in total FAAs were
respectively under normal conditions (Fig. 1). The statistically non-significant by other interactive
salinity stress has significantly affected (P<0.001) the treatments. However, varieties behaved differently in
dry shoot weight. The SA spray enhanced 15% and response to various treatments. A higher value of FAAs
09% shoot dry weight in variety S-24 and MH-97 by 18.1% was recorded in variety MH-97 compared
respectively under saline condition. The various with variety S-24. Overall the salinity stress caused
interactive combinations (varieties x SA levels, significant (P<0.001) effect enhancing the number of
varieties x salinity regimes, SA levels x salinity regimes total FAAs in both varieties. The total FAAs were
and varieties x SA levels and salinity regimes) also increased from 12.29 to 19.42 mg g -1, in crop grown on
produced a little effect in improving the fresh shoot saline soil, showing an increase by 58.1% over non-
weight (Fig. 1). Furthermore, various SA level and saline soil (Fig. 2).

80
Foliarly applied salicylic acid to reduce salt stress in wheat

Fig. 1: The effects of foliar spray of salicylic acid on shoot and root fresh and dry weight of two wheat
varieties grown under saline and non-saline conditions.

Fig. 2: The effects of foliar spray of salicylic acid on total soluble proteins, total free amino acids, total
chlorophyll content and relative water contents of two wheat varieties grown under saline and non-
saline conditions.

81
Noreen et al

Fig. 3: The effects of foliar spray of salicylic acid on the Na+ and K+ ions concentration in shoot and root of
two wheat varieties grown under saline and non-saline conditions.

The total chlorophyll content were affected The pattern of Na+ in root tissues differed significantly
significantly by various factors. However, it was a little (P<0.01) by various varieties. The root tissues of
influenced by various interaction treatments (varieties x variety S-24 absorbed 16.96 mg Na+ g-1 compared to
SA levels, salinity regimes x SA levels and salinity variety MH-97 having 15.18 mg Na+ g-1 (Fig. 3).
regimes x SA levels x varieties). In variety S-24, the
chlorophyll content was decreased from 47.15 to 41.62, DISCUSSION
while it was increased from 40.58 to 44.52 in variety
MH-97 under normal and salt-treated soils. The relative The adverse effects of salinity on agricultural
water content (RWC) was affected significantly by production exceed far a greater proportion than any
various factors. The RWC was found higher by 3.34% environmental factors. The agricultural production
in variety MH-97 compared to variety S-24. The spray areas either irrigated and/or rainfed are equally
of SA increased RWC by 19.57% over unsprayed crop. vulnerable to salinity stress. The crop husbandry
Contrarily, RWC was reduced by 16.1% under saline practices, including growing of salt tolerant crop
condition in comparison with salt treated soils (Fig. 2). species, soil amendments and spraying of plant growth
The varieties significantly differed in the concentration regulators are of common practice in most of the areas.
of Na+ and K+ in roots and shoots in response to SA The exogenous spray of SA could alleviate stress in
levels and salinity. However, both varieties maintained crop species caused by salinity (El- Tayeb, 2005).
similar values of K+ content in their root tissues. The The results of the present study indicated that various
spray of SA caused improvement in K+ concentration attributes of growth and accumulation of Sodium and
from 13.45 to 15.26 mg g-1 compared to untreated crop. Potassium ions were substantially affected by salinity.
There was decrease in K+ concentration from 14.42 to The varieties showed differential pattern of response to
12.26 mg g-1 under non saline condition (Fig. 3). The salt stress. The variation in response to salts stress was
Na+ in root tissues were affected significantly due to associated with genetic make-up of different crop
varieties and salinity levels. While, it was a little species, composition of salt in the growth medium and
influenced by foliar spray of SA. Moreover, it was also management practices (Iqbal and Ashraf, 2005).
unaffected by various interactive treatments (varieties x Thereby, the production of biological yield varied
SA levels, salinity regimes x varieties, salinity regimes greatly in response of salinity levels and treating the
x SA levels and salinity regimes x SA levels x varieties. crop with salicylic acid. The maintenance of low

82
Foliarly applied salicylic acid to reduce salt stress in wheat

osmotic potential under saline conditions reduces the reported that wheat cultivars absorb higher quantities of
absorption of water from rooting zone, thereby osmotic K+ due to foliar spray of salicylic acid. However, the
adjustment is not sustained (Ashraf, 1994). The plant absorption and translocation of K+ and Na+ by barley
species strive to adjust the situation for continuation of crop was reduced in response to foliar spray of SA
their life cycle. The stressful conditions induce anti- under saline conditions (El-Tayeb, 2005). Zhang et al.
oxidant defense system and accumulate the organic (1998) observed that spray of salicylic acid favored in
osmolytes to counter the negative effects of external absorption and translocation of higher quantities of K +,
stress (Noreen et al., 2018; Li et al., 2018). The with simultaneous reduction in Na+ and Cl- ions by
accumulation and over production of organic solutes plant tissues in response to salt stress. In conclusion, the
maintain the osmotic adjustment for continuation of exogenous application of SA proved to be highly
growth (Munns et al., 2000). effective in mitigating the drastic effects of salinity
The results showed that exogenous application of SA stress in both wheat varieties i.e. S-24 and MH-97. The
resulted in improving the biological yield under saline exogenous application of SA improved antioxidant
as well as non-saline conditions. Various researchers defensive mechanism and induced the tolerance
reported that grain yield of wheat was improved by mechanism of the plants under abiotic stress conditions.
spraying of SA under salt stress conditions (El-Tayeb, Thus, wheat crop can be sprayed with SA (200 mg L-1)
2005; Singh and Usha, 2003). Other than spraying of to reduce the salinity stress under arid environment.
SA, the spray of glycine–betaine has also been found Authors’ contribution
effective in reducing the negative effects on salinity SN conceived the idea, designed the project and wrote
Brassica species and spring cereals (Makela et al., the manuscript; AS performed the experiments. UA
1996). The results of this study are collaborated with helped in Lab-work; KHS participated in the design of
those of Khodary (2004) and El-Tayeb (2005) that SA the study and helped in conducting the experiments,
enhanced growth rate and also counteracted the data analysis and writing the manuscript. All authors
negative effects caused by salinity stress in various crop read and approved the final manuscript.
species. The response of crop species to SA vary
greatly due to time of application, stage of crop growth, REFERENCES
crop species and concentration of hormone being
applied (Noreen et al., 2013). Various researchers Ahmad I, SMA Basra, I Afzal, M Farooq and A Wahid,
(Singhvi et al., 2002; Jayachandran et al., 2001; 2013. Growth improvement in spring maize
Sivakumar et al., 2001;) reported that arable crops, through exogenous application of ascorbic
namely rice and pearl millet responded positively in acid, salicylic acid and hydrogen peroxide.
mitigating the deleterious effects of salinity by spray of International Journal of Agriculture and
SA ranging from 100 to 150 mg L -1 during vegetative Biology, 15: 95-100.
growth. Ajithkumar IP and R Panneerselevam, 2014. ROS
The similar results were reported by Chandra et al. scavenging system, osmotic maintenance,
(2007) that deleterious effects of salts could be off-set pigment and growth status of Panicum
through exogenous application of SA on various crop sumatrense Roth under drought stress. Cell
species. The amount of total soluble sugars and soluble Biochemistry and Biophysics, 68: 587-595.
proteins were increased in cowpeas plants, which Ashraf M, 1994. Breeding for salinity tolerance in
accrued in safeguarding the cell membrane structures plants. Critical Reviews in Plant Sciences, 13:
and maintaining the various functions involved in 17-42.
growth and development. It was attributed that Ashraf M, 2004. Some important physiological
maintenance of differential quantities of total soluble selection criteria for salt tolerance in plants.
proteins and free amino acids were associated with Flora, 199: 361-376.
inherited characteristics and eco-edaphic factors in Ashraf M, 2010. Inducing drought tolerance in plants:
production areas. Moreover, the varieties maintained Recent Advances. Biotechnology Advances,
similar quantities of chlorophyll content in their leaf 28: 169-183.
tissues under both non-saline and salt-stressed Ashraf M, NA Akram, F Al-Qurainyed and MR Foolad,
environments. Contrarily, chlorophyll content was 2011. Drought tolerance. Roles of organic
impacted to a greater extent in response to salt-stress or osmolytes, growth regulators, and mineral
water-stress (Tas and Tas, 2007). The effects of water- nutrients. Advances in Agronomy, 111: 249-
stress and/or salt-stress could be reduced to a 296.
proportionate level by spraying SA (Zhang et al., 2004). Babar S, EH Siddiqi, I Hussain, KH Bhatti and R
It was further revealed that absorption of Na + by both Rasheed, 2014. Mitigating the effects of
crop varieties was increased with concurrent reduction salinity by foliar application of Salicylic acid
in K+ content under saline environment. It was in Fenugreek. Physiology Journal, 2014: 869058.

83
Noreen et al

Bradford MM, 1976. A rapid and sensitive method for spring wheat. Plant Growth Regulation, 60:
the quantification of microgram quantities of 41-52.
protein utilizing the principle of protein dye Jayachandran M, P Rajendran and M Thangaraj, 2001.
binding. Analytical Biochemistry, 72: 248-253. Effect of growth regulators on growth and
Chandra A, A Anand and A Dubey, 2007. Effect of yield of wet season rice. Madras Agriculture
salicylic acid on morphological and Journal, 87: 340-342.
biochemical attributes in cowpeas. Journal of Jones MM and NC Turner, 1980. Osmotic adjustment
Environmental Biology, 28: 193-196. in expanding and fully expanded leaves of
Dat JF, H Lopez-Delgado, CH Fayer and IM Scott, sunflower in response to water deficit.
2000. Effects of salicylic acid on oxidative Australian Journal of Plant Physiology, 7: 181-
stress and thermo-tolerance in tobacco. Journal 192.
of Plant Physiology, 156: 659-665. Joseph B, D Jini and S Sujatha, 2010. Insight into the
El-Tayeb MA, 2005. Response of barley plants to the role of exogenous salicylic acid on plants
interactive effect of salinity and salicylic acid. grown under salt environment. Asian Journal
Plant Growth Regulation, 45: 215-224. of Crop Science, 2: 226-235.
Epstein E, 1972. Mineral Nutrition of Plants; Principles Khodary SEA, 2004. Effect of salicylic acid on growth,
and Perspectives. John Wiley & Sons photosynthesis and carbohydrate metabolism
Publishers, New York, USA. in salt stressed maize plants. International
Farooq M, A Wahid, DJ Lee, S Cheema and T Aziz, Journal of Agriculture and Biology, 6: 5-8.
2010. Drought stress: Comparative time course Krasensky J and C Jonak, 2012. Drought, salt and
action of the foliar applied glycine betaine, temperature stress-induced metabolic
salicylic acid, nitrous oxide, brassinosteroids rearrangements and regulatory networks.
and spermine in improving drought resistance Journal of Experimental Botany, 63: 1593-
of rice. Journal of Agronomy and Crop 1608.
Science, 196: 336-345. Li X, M Ke, M Zhang, WJGM Peijnenburg, X Fan, J
Farouk S and SA Arafa, 2018. Mitigation of salinity Xu, Z Zhang, T Lu, Z Fu and H Qian, 2018.
stress in Canola plants by Sodium The interactive effects of diclofop-methyl and
nitroprusside application. Spanish Journal of silver nanoparticles on Arabidopsis thaliana:
Agricultural Research, 16: e0802. Growth, photosynthesis and antioxidant
Hamada AM and AM Al-Hakimi, 2001. Salicylic acid system. Environmental Pollution, 232: 212-
versus salinity drought induced stress on wheat 219.
seedling. Rostlinna Vyroba, 47: 444-450. Makela P, J Mantila, R Hinkanen, E Pehu and P Sainio,
Hartmann HT, DE Kester and FT Davies, 1990. Plant 1996. Effect of foliar application of glycine
Propagation Principles and Practices. 5th betaine on stress tolerance, growth and yield of
Edition, Prentice Hall, Englewood Cliffs, New spring cereals and summer turnip rape in
Jersey, USA. Finland. Journal Agronomy and Crop Science,
Hayat S and A Ahmad, 2007. Salicylic Acid: A plant 176: 223-234.
hormone. Springer Publishers, Dordrecht, The Munns R, JB Passioura, J Gua, O Chazen and GR
Netherlands. Cramer, 2000. Water relations and leaf
Hayat S, Q Fariduddin, B Ali and A Ahmad, 2005. expansion: importance of time scale. Journal
Effect of salicylic acid on growth and enzyme of Experimental Botany, 51: 1495-1504.
activities of wheat seedlings. Acta Mutlu S, O Karadogolu, O Atici and B Nalbantoglu,
Agronomica Hungarica, 53: 433–437. 2013. Protective role of salicylic acid applied
Husen A, M Iqbal, SS Sohrab and MKA Ansari, 2018. before cold stress on antioxidative system and
Salicylic acid alleviates salinity-caused proteins patterns in barley apoplast. Biologia
damage to foliar functions, plant growth and Plantarum, 57: 507-513.
antioxidant system in Ethiopian mustard Nahrjoo M and S Sedaghathoor, 2018. The induction of
(Brassica carinata A. Br.). Agriculture and salinity stress resistance in rosemary as
Food Security, 7: 44. influenced by salicylic acid and jasmonic acid.
Ilyas N, R Gull, R Mazhar, M Saeed, S Kanwal, S Communications in Soil Science and Plant
Shabir and F Bibi, 2017. Influence of salicylic Analysis, 49: 1761-1773.
acid and jasmonic acid on wheat under Noreen S, M Ashraf, M Hussain, and A Jamil, 2009.
drought stress. Communications in Soil Exogenous application of salicylic acid
Science and Plant Analysis, 48: 2715-2723. enhances antioxidative capacity in salt-stressed
Iqbal M and M Ashraf, 2005. Changes in growth, sunflower (Helianthus annuus L.) plants.
photosynthetic capacity and ionic relations in Pakistan Journal of Botany, 41: 473-479.

84
Foliarly applied salicylic acid to reduce salt stress in wheat

Noreen S, MS Akhter, T Yaamin and M Arfan, 2018. Singh B and K Usha, 2003. Salicylic acid induced
The ameliorative effects of exogenously physiological and biochemical changes in
applied proline on physiological and wheat seedlings under water stress. Plant
biochemical parameters of wheat (Triticum Growth Regulation, 39: 137-141.
aestivum L.) crop under copper stress Singhvi NR, J Kodandaramiah, M Rekha, A Sarkar and
condition. International Journal of Plant RK Datta, 2002. Influence of salicylic acid on
Sciences, 13: 221-230 mulberry growth and leaf yield. Advances in
Noreen S and M Ashraf, 2008. Alleviation of adverse Plant Science, 15: 463-466.
effects of salt stress on sunflower (Helianthus Sivakumar R, MK Kalarani, V Malika and KB Sujatha,
annuus L.) by exogenous application of 2001. Effect of growth regulators on
salicylic acid: growth and photosynthesis. biochemical attributes, grain yield and quality
Pakistan Journal of Botany, 40: 1657-1663. in pearl millet. Madras Agriculture Journal,
Noreen S, HR Athar and M Ashraf, 2013. Interactive 88: 256-259.
effects of watering regimes and exogenously Snedecor GW and WG Cochran, 1980. Statistical
applied osmoprotectants on earliness indices Methods. 7th Edition, The Iowa State
and leaf area index in cotton (Gossypium University Press, Ames, Iowa, USA.
hirsutum L.) crop. Pakistan Journal of Botany, Tas S and B Tas, 2007. Some physiological response of
45: 1873-1881. drought stress in wheat genotypes with
Ryan J, G Estefan and A Rashid, 2001. Soil Plant different ploidity in Turkey. World Journal of
Analysis Laboratory Manual. 2nd Edition, Agriculture Science, 3: 178-183.
International Centre for Agriculture Research Zhang ML, Z Zhai, J Li, X Tian, B Wang, Z He and Z
in Dry Area (ICARDA), Syria, pp: 172. Li, 2004. Effect of plant growth regulators on
Shabala L, A Mackay, Y Tian, SE Jacobsen, D Zhou water deficit induced yield loss in soybean.
and S Shabala, 2012. Oxidative stress, Proceedings of 4th International Crop Science
protection and stomatal patterning as Congress, Brisbane, Australia.
components of salinity tolerance mechanism in Zhang SG, JY Gao and JZ Song, 1998. The effects of
quinoa (Chenopodium quinoa). Physiologia salicylic acid and aspirin on the Na+, K+ and
Plantarum, 146: 26-38. Cl- contents and distribution in the wheat
Sharma P and RS Dubey, 2005. Drought induced seedlings under salinity stress. Beijing
oxidative stress enhances activities of Agricultural Sciences, 16: 1-5.
antioxidants systems in growing rice seedling. Zhu JK, 2001. Plant Salt Tolerance. Trends in Plant
Plant Growth Regulation, 46: 209-221. Science, 6: 66-71.

85