International Journal of Farming and Allied Sciences

Available online at www.ijfas.com

©2014 IJFAS Journal-2014-3-9/980-987/ 30 September, 2014
ISSN 2322-4134 ©2014 IJFAS

Effect of salicylic acid on activities of antioxidant

enzymes, flowering and fruit yield and the role on
reduce of drought stress
Mohammad Lakzayi1, Ebrahim Sabbagh2, Khashayar Rigi2* and Abbas Keshtehgar2

1. Department of Agronomy, Zahedan Branch, Islamic Azad University, Zahedan, Iran

2. Department of plant Production, Khash Branch, Islamic Azad University, Khash, Iran

Corresponding author: Khashayar Rigi

ABSTRACT: Effect of drought is among the environmental constraints that affect crop growth and crop
production worldwide. Drought or water deficit stress elicits many different physiological responses in
plants. It causes reduction in leaf water potential, stomatal conductance, nitrate reduction and inhibits leaf
enlargement while osmolytes such as total soluble sugars and proline are increased. The decrease in
chlorophyll content under drought stress has been considered a typical symptom of oxidative stress and
may be the result of pigment photo-oxidation and chlorophyll degradation. Relative water content (RWC),
leaf water potential, stomatal resistance, the rate of transpiration, leaf temperature and canopy
temperature are important characteristics that influence plant water relations. Salicylic acid (SA) as a
potent signaling molecule in plants is involved in eliciting specific responses to biotic and abiotic stresses.
Several studies have demonstrated that exogenous SA application enhances plant growth and
development.

Keywords: Chlorophyll, nutrient uptake, proline

INTRODUCTION

Drought or water deficit stress

Effect of drought is among the environmental constraints that affect crop growth and crop production worldwide
(Ashraf and Harris, 2004; Farooq, 2009). It has been estimated that up to 45% of the world agricultural lands are
subjected to drought (Bot, 2000).

Figure 1. The impact of climate, population, and carbon dioxide on water resources
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Drought or water deficit stress elicits many different physiological responses in plants. It causes reduction in leaf
water potential, stomatal conductance, nitrate reduction and inhibits leaf enlargement while osmolytes such as total
soluble sugars and proline are increased (Jafar, 2004; Adejare and Umebese, 2007).

Figure 2. Plants are exposed to a variety of abiotic stresses

The plant response to drought consists of numerous processes that must function in coordination to alleviate
both cellular hyperosmolarity and ion disequilibrium. To cope with drought stress, plants respond with physiological
and biochemical changes. These changes aim at the retention of water in spite of the high external osmoticum and
the maintenance of photosynthetic activity, while stomatal opening is reduced to counter water loss. Accumulation of
low molecular compounds, such as glycinebetaine, sugars, sugar alcohols and proline, is a mechanism aimed at
balancing water potential following drought (Pilon-Smits . 1995).

Adapt to drought
Plants tend to adapt to drought by accumulation of cyto-compatible organic osmolytes (Rhodes and Hanson,
1993) such as polyols, proline and betaines. Seed treatment or foliar application of chemicals like glycinebetaine,
kinetin, salicylic acid (Gunes, 2007; Karlidag, 2009) may increase yield of different crops due to reduction in stress
induced inhibition of plant growth (Elwana and El-Hamahmyb, 2009), enhanced photosynthetic rates, leaf area and
plant dry matter production (Khan, 2003).

Figure 4. Influence of water stress on physiology

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Respiration
Drought tolerance is a cost-intensive phenomenon, as a considerable quantity of energy is spent to cope with it.
The fraction of carbohydrate that is lost through respiration determines the overall metabolic efficiency of the plant
(Davidson, 2000). The root is a major consumer of carbon fixed in photosynthesis and uses it for growth and
maintenance, as well as dry matter production (Lambers, 1996). Plant growth and developmental processes as well
as environmental conditions affect the size of this fraction (i.e. utilized in respiration). However, the rate of
photosynthesis often limits plant growth when soil water availability is reduced (Huang and Fu, 2000). A negative
carbon balance can occur as a result of diminished photosynthetic capacity during drought, unless simultaneous and
proportionate reductions in growth and carbon consumption take place. In wheat, depending on the growth stage,
cultivar and nutritional status, more than 50% of the daily accumulated photosynthates were transported to the root,
and around 60% of this fraction was respired (Lambers, 1996).

Figure 5. Plant responses to water stress

Droughtsensitive spring wheat (Longchun, 8139–2) used a relatively greater amount of glucose to absorb water,
especially in severe drought stress (Liu, 2004).

Chlorophyll content
Chlorophyll which is one of the major chloroplast components for photosynthesis, and relative chlorophyll content
has a positive relationship with photosynthetic rate. The decrease in chlorophyll content under drought stress has
been considered a typical symptom of oxidative stress and may be the result of pigment photo-oxidation and
chlorophyll degradation (Anjum , 2011). Limited water supply usually causes a reduction in chlorophyll content
(Paknejad ., 2007). Being positively correlated with yield (Zaharieva ., 2001) relatively high chlorophyll content may
contribute to the plant productivity under stress conditions. Photosynthesis is one of the most sensitive processes to
drought stress (Chaves , 2009). The inhibitory effects of drought on photosynthesis may be associated with low CO2
availability due to low stomatal and mesophyll conductance (Flexas, 2008) and/or impairments in carbon assimilation
metabolism (Peeva and Cornic, 2009). Stomatal closure is an early response to drought and an efficient way to
reduce water loss in water-limiting environments. Biochemical limitation of photosynthesis also plays an important
role under prolonged periods of drought stress (Flexas, 2008).

Figure 6. Plant global response to water stress, high temperature and salinity

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Relative water content

Relative water content (RWC), leaf water potential, stomatal resistance, the rate of transpiration, leaf temperature
and canopy temperature are important characteristics that influence plant water relations. Relative water content is
considered a measure of plant water status, reflecting the metabolic activity in tissues and used as a most meaningful
index for dehydration tolerance. RWC of leaves is higher in the initial stages of leaf development and declines as the
dry matter accumulates and leaf matures. RWC related to water uptake by the roots as well as water loss by
transpiration (Anjum , 2011). RWC tend to decline when transpiration exceeds water absorption under drought
condition (Tas and Tas, 2007) leading to decrease in cell turgor. Maintenance of high RWC under drought due to
relatively more growth of the roots than shoots and/or abscisic acid induced reduction in stomatal opening (Makoto
., 1990) tends to maintain cell turgidity, chlorophyll content (Keyvan, 2010).

Figure 7. Water and salt stress tolerance mechanisms in plants

Effect of drought stress on glycine betaine content

Drought stressed shallot plants showed an increase in glycine betaine content when compared to control. The
glycine betaine content increased under drought stress in Radix astragali (Tan, 2006), in barley (Nakamura, 2001)
and in higher plants (Jun, 2000).

Figure 8. Leaf rolling in maize

Glycine betaine is considered to be one of the most abundant quaternary ammonium compounds produced in
higher plants under stressful environment (Yang, 2003). Glycine betaine has been shown to protect the enzymes
and membranes and also to stabilize PSII protein pigment complexes under stressful conditions (Papageorgiou and
Morata, 1995).

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Figure 9. Physiological mechanisms induced by water stress

Accumulation of proline
The accumulation of proline, primarily in the cytosol, often occurs in plants under stress witha strong correlation
between stress tolerance and proline accumulation, but the relationship is not universal and may be species
dependent (Ashraf and Foolad, 2007).There are other roles were proposed for proline besides osmotic adjustment
in stressed plants include acting as hydroxyl scavenger, stabilization of membrane and protein structure, as a sink
for carbon and nitrogen for stress recovery, and buffering cellular redox potential under stress (Lee, 2008).

Fig 10. Illustration of the response of plants to water stress. Stomatal response, ROS scavenging, metabolic changes, and
photosynthesis are all affected when plants are subjected to water stress

Nutrient relations
Decreasing water availability under drought generally results in limited total nutrient uptake and their diminished
tissue concentrations in crop plants. An important effect of water deficit is on the acquisition of nutrients by the root
and their transport to shoots. Lowered absorption of the inorganic nutrients can result from interference in nutrient
uptake and the unloading mechanism, and reduced transpirational flow (Garg, 2003; McWilliams, 2003). However,
plant species and genotypes of a species may vary in their response to mineral uptake under water stress. In general,
moisture stress induces an increase in N, a definitive decline in P and no definitive effects on K (Garg, 2003).
Transpiration is inhibited by drought, as shown for beech (Peuke, 2002), but this may not necessarily affect nutrient
uptake in a similar manner. Influence of drought on plant nutrition may also be related to limited availability of energy
for assimilation of NO−3 /NH+4 , PO3−4 and SO2−4 : they must be converted in energy-dependent processes before
these ions can be used for growth and development of plants (Grossman and Takahashi, 2001). As nutrient and
water requirements are closely related, fertilizer application is likely to increase the efficiency of crops in utilizing
available water. This indicates a significant interaction between soil moisture deficits and nutrient acquisition. Studies
show a positive response of crops to improved soil fertility under arid and semi-arid conditions. Currently, it is evident
that crop yields can be substantially improved by enhancing the plant nutrient efficiency under limited moisture supply
(Garg, 2003). It was shown that N and K uptake was hampered under drought stress in cotton (McWilliams, 2003).

Salicylic acid
Salicylic acid (SA) or ortho-hydroxy benzoic acid and other salicylates are known to affect various physiological
and biochemical activities of plants and may play a key role in regulating their growth and productivity (Hayat ., 2010).
Salicylic acid is considered to be an endogenous growth regulator of phenolic nature that enhanced the leaf area
and dry mass production in corn and soybean (Khan, 2003).
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Figure 11. The difference between structure of aspirin and salicylic acid

Relation between abiotic stress and Salicylic acid

Salicylic acid (SA) as a potent signaling molecule in plants is involved in eliciting specific responses to biotic and
abiotic stresses. It has been shown that SA provides protection in maize (Janda, 1999) and winter wheat plants
(Tasgin, 2003) against low-temperature stress, induces termotolerence in mustard seedlings (Chen, 1997; Dat, 1998)
or modulates plant responses to salt and osmotic stresses (Borsani, 2001) ozone or UV light (Sharma, 1996) drought
(Senaratna, 2002) and herbicides (Ananieva, 2004).

Enhance the activities of antioxidant enzymes

Salicylic acid was found to enhance the activities of antioxidant enzymes such as peroxidase (POD), SOD and
CAT, when sprayed exogenously to the drought stressed plants of tomato (Hayat, 2008) or to the salinity stressed
plants (Szepesi, 2008; Yusuf, 2008).

Plant growth
Several studies have demonstrated that exogenous SA application enhances plant growth and development.
Fariduddin, (2003) showed that mustard plants sprayed with low concentrations of SA produced larger amounts of
dry matter and had higher photosynthetic rate in comparison with control plants. SA application to corn and soybean
promoted leaf area and dry weight of plants (Khan, 2003). In another study Hussein, (2007) revealed that growth
traits of wheat plants were improved as a result of SA spraying on the plants. In addition, Hayat, (2005) reported that
soaking of wheat grains in low concentrations of SA significantly promoted growth of wheat seedlings.

Flowering and fruit yield

Another important parameter that is directly related to yield and productivity of plants. Salicylic acid has been
reported to induce flowering in a number of plants. In cucumber and tomato, the fruit yield enhanced significantly
when the plants were sprayed with lower concentrations of salicylic acid (Larque-Saavedra and Martin-Mex 2007). It
was reported that the foliar application of salicylic acid to soybean also enhanced the flowering and pod formation
(Hayat, 2010). Flowering is another important parameter that is directly related to yield and productivity of plants.
Salicylic acid has been reported to induce flowering in a number of plants. In cucumber and tomato, the fruit yield
enhanced significantly when the plants were sprayed with lower concentrations of salicylic acid (Larque-Saavedra
and Martin-Mex 2007). It was reported that the foliar application of salicylic acid to soybean also enhanced the
flowering and pod formation(Hayat, 2010).

Exogenous SA application on accumulation of glycine betaine

Exogenous SA application to the drought stressed plants increased glycine betaine content as compared to
drought stressed plants as well as control plants. The accumulation of glycine betaine might serve as an intercellular
osmoticum and it can be closely correlated with the elevation of osmotic pressure (Kavikishore ., 1995).

Seed germination
SA significantly stimulated the activities of enzymes involved in germination such as transkelolase, enolase,
malate dehydrogenase, phosphoglycerate kinase, glyceraldehyde 3-phosphate, dehydrogenase, fructose 1,6-
diphosphatase, and pyruvate decarboxylase. In addition to it seeds germinated in SA supplemented media showed
abundant levels of isocitrate lyase and malate synthase (key enzymes of glyoxylate cycle) (Eastmond and Graham,
2001 and Rajjou, 2006). Rajjou, (2006) hypothesized that detoxification mechanism in germinating seeds counteract
by exogenous SA treatments. Thus the present work has been conducted to investigate the response of different
groundnut cultivars to salicylic acid.

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