Chemistry Research Journal, 2017, 2(3):198-203

Available online www.chemrj.org

ISSN: 2455-8990
Review Article CODEN(USA): CRJHA5

Influence of salt stress on water uptake, pigments system, ion-homeostasis in crop plants

Mohsen Arabshahi*, Hamid Reza Ganjali, Ahmad Mehraban

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

Abstract Increased salinization of arable land is expected to have devastating global effects, resulting in 30% land
loss within the next 25 years and up to 50% by the middle of twenty-first century. High soil salinity causes both
hyperionic and hyperosmotic stress and can lead to plant demise. Salinity in a given land area depends upon various
factors like amount of evaporation (leading to increase in salt concentration), or the amount of precipitation (leading
to decrease in salt concentration). Changes of pigment system contents under salt stress are used as parameter for
selection of tolerant and sensitive cultivars in crop plants. Chlorophyll a, chlorophyll b, chlorophyll a/b and
carotenoid contents showed increase and decrease depending on exposure time of NaCl exposure in many plants.
When K substitute by Na in biochemical reactions then Ion cytotoxicity is happening and when Na and Cl ions
interfere with non covalent interactions between their amino acid, then proteins don't function, thus in salinity
condition poisonous levels of sodium and also inadequate amount of K for enzymatic reactions and osmotic
adjustment is happened.

Keywords Toxicity, Morphological Adaptations, Salinity

Introduction
High salt depositions in the soil generate a low water potential zone in the soil, making it increasingly difficult for
the plant to acquire both water as well as nutrients. The basic physiology of high salt stress and drought stress
overlaps with each other. Salinization of soil is a serious land degradation problem and is increasing steadily in
many parts of the world, in particular in arid and semiarid areas [1-2]. Increased salinization of arable land is
expected to have devastating global effects, resulting in 30% land loss within the next 25 years and up to 50% by the
middle of twenty-first century [3]. High soil salinity causes both hyperionic and hyperosmotic stress and can lead to
plant demise. Salinity in a given land area depends upon various factors like amount of evaporation (leading to
increase in salt concentration), or the amount of precipitation (leading to decrease in salt concentration) [4].
Therefore, salt stress essentially results in a water-deficit condition in the plant and takes the form of a physiological
drought [4]. In plants, both drought and salinization are manifested primarily as osmotic stress, resulting in the
disruption of homeostasis and ion distribution in the cell [5]. Increased salinization of arable land is expected to have
devastating global effects, resulting in 30% land loss within the next 25 years and up to 50% by the middle of
twenty-first century [3].
Toxicity in plants
Toxicity in plants results mainly from high concentrations of ions Na +, CI–, [6], although most studies on salinity
effect on plants are associated with NaCI excess and few studies have focused on Na 2SO4 performance in growth [7-
8] and in physiology of plants [9]. Na+ and CI– are present in greater amounts in seawater; thus, aquatic
environments close to the sea may present high concentrations of Na + and CI–. High concentrations of SO42– may be

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found in aquatic environment where there are ores and anthropogenic activities such as agriculture [10]. Proximity
to seawater also influences sulphate concentration in coastal aquatic environments [11].

Morphological Adaptations
Munns (1992) [12] concluded that the salts absorbed by plants do not control growth directly, but that they do
influence turgor, photosynthesis and/or the activity of specific enzymes. Demonstrating the complexity of salt stress,
this author developed a model showing a two-phase effect of salinity on plant growth. Growth is first reduced by a
decrease in the soil water potential (osmotic phase) and, later, a specific effect appears as salt injury in leaves, which
die because of a rapid increase in salt in the cell walls or cytoplasm when the vacuoles can no longer sequester
incoming salts (ionic phase). Munns (1992) [12] found that this salt accumulation in the old leaves accelerates their
death and thus decreases the supply of carbohydrates and/or growth hormones to the meristematic regions, thereby
inhibiting growth. The fact that plant growth is limited by a reduction in the photosynthesis rate and by an excessive
uptake of salts affects the production of specific metabolites that directly inhibit growth (Azza et al., 2007). Salinity
reduces plant growth through osmotic and toxic effects, and high sodium uptake ratio values cause sodicity, which
increases soil resistance, reduces root growth, and reduces water movement through the root with a decrease in
hydraulic conductivity [13]. Root hydraulic conductivity in field crops may vary in response to the salt content of
the irrigation water applied [14]. In general, the root hydraulic conductivity in plants irrigated with poor quality
water tends to decrease. In fact, the effect on root hydraulic conductivity is one of the main factors determining the
sustainability of a reclaimed water irrigation system. Normally, root hydraulic conductance is expressed in terms of
the whole root dry weight, without taking into account the role of root architecture in the water uptake capacity.
Nonetheless, for any given root dry weight value, the amount of fine roots, which determine the root length and
surface area, may vary greatly, thus, affecting the water absorption level [15]. Moreover, in assays where the grafted
technique has been applied, the rootstock properties can affect the plant response to salinity. Salt-tolerant rootstocks
alleviate the negative effects of abiotic stress to a greater extent than salt-sensitive rootstocks. Navarro et al. (2010)
[16] described worse fruit yield and quality in Clemenules mandarin trees grafted on Carrizo (salt-sensitive
rootstock) compared with Cleopatra (salt-tolerant rootstock), both irrigated with a NaCl solution (30 mM). The
anatomy of the root system (length, root diameter, etc.) determines root performance, enabling plants to acquire
water and nutrients and thereby increase the replacement rate of plant water lost [17]. Optimum root systems can
support shoot growth and improve plant yields, since roots serve as an interface between plants and the soil [18]. A
proliferated root system would therefore appear to be better for plants, for it allows them to penetrate deeper layers
of soil to acquire water and nutrients [19]. Recent studies, however, have shown that species with other root
features, including small roots, can be more advantageous for shoot development [20]. For example, just a few roots
in moist soil can provide amounts of water independent of the root number. Other root characteristics, such as the
number and diameter of xylem vessels, width of the root cortex, number of root hairs, and the suberin deposition in
both the root exodermis and endodermis, also determine the permeability of roots to water [21]. Furthermore,
environmental factors in the soil (changes in temperature, lack of O 2, mechanical impedance, salinity) can also
produce marked impacts on root anatomy. The cell walls of root cells of salinized plants are often unevenly
thickened and convoluted [22]. Salts often promote the suberisation of the hypodermis and endodermis in woody
tree roots, resulting in the formation of a well-developed casparian strip closer to the root apex, different to that
found in non-salinized roots [23]. Furthermore, the morphology of some plants shows their sensitivity to salinity.
For example, in avocado trees, the root system is quite superficial and presents low ramification, thus reducing the
water and nutrient absorption capacity [24], resulting in a higher sensitivity to soil salinity [25]. These
morphological features limit the distribution of this crop to areas where irrigation water is of good quality. Saline
water irrigation has also been found to alter the root system morphology of Callistemon citrinus plants [26].

Pigments system
Net photosynthesis and stomatal conductance are significantly affected by salt stress due to changes in chlorophyll
content and chlorophyll fluoresance, damage of photosynthetic apparatus and chloroplast structure [27]. Plant

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pigments content were determined in different tolerant and sensitive plant varieties at wide range of salt
concentrations [28]. Chlorophyll a, chlorophyll b and carotenoid are main photosynthetic pigments and they play
important role in photosynthesis. The changes in the amount of pigments system were evaluated as the changes in
photosynthesis. Changes of pigment system contents under salt stress are used as parameter for selection of tolerant
and sensitive cultivars in crop plants [29]. Chlorophyll a, chlorophyll b, chlorophyll a/b and carotenoid contents
showed increase and decrease depending on exposure time of NaCl exposure in many plants [30]. The reduction of
chlorophyll a and chlorophyll b amounts with NaCl application was reported in many plants such as Zea mays,
Carthamus tinctorius, Bean and Paulownia imperiallis that this due to increasing of destructive enzymes called
chlorophyllase [31]. Pigments system reduction is attributed to a salt induced weakening of protein-pigment-lipid
complex or increased chlorophyllase enzyme activity [32]. Also, increase in pigment content was observed in
salinity stressed plant such as Rice [27] and Purslane [31], that this increment may be due to increase in the number
of chloroplast in the stressed plant leaves [33]. In other hand, the changes in pigment system were affected by
exposure time and salt concentration [27].

Plants growing in saline soil

Plants growing in saline soil are subjected to three distinct physiological stresses. First, the toxic effects of specific
ions such as sodium and chloride, prevalent in saline soils, disrupt the structure of enzymes and other
macromolecules, damage cell organelles, disrupt photosynthesis and respiration, inhibit protein synthesis, and
induce ion deficiencies [34]. Second, plants exposed to the low osmotic potentials of saline soil are at risk of
physiological drought because they must maintain lower internal osmotic potentials to prevent water moving from
the roots into the soil. Finally, salinity also produces nutrient imbalance in the plant caused by decreased nutrient
uptake and/or transport to the shoot [35-36]. As a consequence, salt stress affects all the major processes, such as
growth, photosynthesis, protein synthesis, and energy and lipid metabolisms [37].

Ion-homeostasis
Salinity interrupts homeostasis ion distribution and water potential at cellular and whole plant levels [38-39]. When
K substitute by Na in biochemical reactions then Ion cytotoxicity is happening and when Na + and Cl- ions interfere
with non covalent interactions between their amino acid, then proteins don't function, thus in salinity condition
poisonous levels of sodium and also inadequate amount of K for enzymatic reactions and osmotic adjustment is
happened [40-42]. Potassium is necessary in high concentrations, and functions in the cell through direct interacting
in enzyme activation, stabilization of protein synthesis, and neutralization of negative charges on proteins [43-45]
also the positive correlation between potassium content and final yield in different crops was reported [41, 46-47].
When potassium shortage is happened, acidification and sodium transport are increased [48-49].

Water uptake
Salt has two major effects on plants: osmotic stress and ionic toxicity, both of which affect all major plant processes
[50]. Plants are able to take up water and essential minerals because they have a higher water pressure than the soil
under normal conditions. When salt stress occurs, the osmotic pressure of the soil solution is greater than that in
plant cells. Thus, the plant cannot get enough water [51]. In addition, its cells will have decreased turgor and its
stomata will close to conserve water. Stomatal closing can lead to less carbon fixation and the production of
Reactive Oxygen Species (ROS) such as superoxide and singlet oxygen. ROS disrupts cell processes through
damage to lipids, proteins, and nucleic acids [52]. Ionic toxicity occurs when concentrations of salts are imbalanced
inside cells and inhibit cellular metabolism and processes. Sodium ions at the root surface disrupt plant nutrition of
the similar cation potassium by inhibiting both potassium uptake and enzymatic activities within the cell. Potassium
is an important nutrient in a plant, regulating over 50 enzymes (Kader, 2010). Essential for maintaining cell turgor
pressure, creating membrane potential, and regulating enzymatic activities, potassium must be maintained at 100-
200mM in the cytosol. Sodium, on the other hand, causes stress at concentrations higher than 10mM in the cytosol

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[51]. Na+ is a cation similar to K+ and easily crosses the cell membrane. It also acts as an inhibitor to many enzymes,
affecting metabolic processes. Calcium cations, however, protect some plants through signaling pathways that
regulate potassium sodium transporters [52]. When a plant senses salt stress through transmembrane proteins or
enzymes in the cytosol, the amount of calcium in the cytosol increases [51]. Calcium is a second messenger
important to many biochemical pathways and can aid plants in responding to salt stress. The osmotic and ionic stress
induced by salinity can halt plant growth as the plant focuses its energy on conserving water and improving ionic
balance. In order for plants to return to normal functioning and photosynthesis, the plant must facilitate its own
detoxification – damage must be prevented or lessened, homeostasis must be reestablished, and growth must resume
[39].

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