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Physiology of Salt Stress in Plants


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turns the lands nonproductive or unfertile.

      The impact of salt stress is found to be most severe on agricultural crops. The primary issues involve the non‐germination of seeds, reduced leaf surface area, retarded plant growth, strength, hampered yield, etc. Elevated soil salinity hampers the plants in various ways such as osmotic stress (OS), ionic toxicity, retarded cell division, reduced photosynthesis, to name a few. The inclusive impact of all the above factors boosts the mortality rate (Lauchli and Grattan 1970).

      Immediate exposure to higher saline medium primarily increases the OS, causing reduced leaf surface area (i.e. due to repressed cell division and growth). Whereas, prolonged exposure imparts ionic stress leading to stomatal closure, immature senescence of mature leaves, chlorosis, necrosis, etc. The reduced biomass negatively affects photosynthesis and plant growth (Darko et al. 2019). In contrast, exposure to elevated sodicity, especially NaCl,affects the enzymatic system and augments cell swelling. The mutual impact leads to suppressed energy synthesis. Furthermore, excess exposure hinders all the growth‐oriented processes like metabolism and protein synthesis (Acosta‐Motos et al. 2017).

      Therefore, prolonged exposure provoked the development of a defense mechanism in some species against salt stress and toxicity either by excluding through cells or by enhancing the salt tolerance. Additionally, synthetic species with transgenic properties are also synthesized by genetic engineering by altering the levels of gene expression (Carillo et al. 2011).

      Accumulation of excessive salt content in the soil causing direct and indirect adverse effects on flora and fauna is termed as salt stress (Shrivastava and Kumar 2015). The above situation can inhibit plant growth, and prolonged exposure may lead to a decrease. Higher saline level impacts the plants in various ways such as genotoxicity, alteration of metabolic processes, oxidative stress, water stress, ion toxicity, nutritional disorders, reduction of cell division and expansion, and membrane disorganization (Hasegawa et al. 2000; Munns 2002). The preliminary exposure to salt stress causes leaf surface area reduction. The immediate impacts include suppressed cell expansion and cell division and closure of stomata due to osmotic influence (Munns 2002; Flowers 2004). Furthermore, prolonged exposure imparts ionic stress leading to early senescence of mature leaves and thereby reducing the leaf surface area responsible for photosynthesis and plant growth.

      Based on the origin and root cause, there are two different categories of salinity, namely, primary and secondary. Primary salinity is a natural phenomenon and mostly occurs due to the former presence of salt lakes, slat clads, tidal swamp, etc., at a particular location. It is majorly a kind of sodicity. At the same time, secondary salinity is imposed due to man‐made activities such as urbanization, saline irrigation, etc. (Shahid and Rahman 2011). Detailed reasons are delineated below.

      Primary salinity:

      1 Spreading from the saline artesian well.

      2 Capillary rise from saline groundwater.

      3 Seawater intrusion.

      4 Canopy formation due to the movement of fine sea sand by the sea breeze.

      5 Waterlogging.

      Secondary salinity:

      1 Irrigation with impeded drainage

      2 Effluent discharge

      3 Excess fertilizer dosing

      4 Deforestation

      5 Saline irrigation

      Furthermore, based on the predominance of the type of anions present and the pH value, salt‐affected soils are categorized as saline soil and sodic soil. Sodic soil typically comprises sodium carbonate and or bicarbonate ions with a pH value beyond 8.5, but contrarily, saline soil majorly incorporates chloride and sulphate ions with pH value below 8.5. Certain plant species manage to compensate the imparted stress through its metabolism and survive in the severe salt conditions known as halophytes. Remaining plant species are termed as glycophytes with a higher mortality rate overexposure to 10% or more concentration of saline water (Gorham 1995; Parida and Das 2005; Mane et al. 2011; Gupta and Huang 2014).

      Primarily, hydro‐geological activities contribute in escalating soil salinity and sodicity. Moreover, the soil is generated because of the weathering actions on intermediate and basic igneous rocks; sandstones already carry salt as a primary constituent. In the regions with moderate to low rainfall, a greater rate of evapotranspiration induces higher salinity and sodicity. Furthermore, coastal regions with tidal exposure may also develop salinity problems. A study conducted by Sultana et al. (2001) depicted that rice yield in coastal Asia gets often impaired due to the intrusion of saline Indian Ocean water. Inland precipitation also surprisingly elevates the soil sodicity. It is evidenced that rainwater can constitute up to a few milligrams of salt against each kilogram of a downpour with an electrical conductance (EC) value of 0.01 dS/m (Cucci et al. 2016; Corwin and Yemoto 2017; Hossain, 2019).

      However, the deteriorating impacts of artificially induced salinity are more predominant. Over‐irrigation or saline water irrigation is cited as one of the prima facie reason for human‐induced salinity. Roughly, it is estimated that globally half of the irrigated lands are anyhow saltaffected. Other than irrigation, probable sources of inland salinity are the following:

      1 Salt accumulation: Effluent and waste discharged into the surface water bodies from the industries and effluent treatment plants (ETPs) beyond absolute concentrations can accumulate and form salt films downstream to cause acute saline toxicity (Naidoo and Olaniran 2013).

      2 Reduction of greenbelt: Deforestation accelerates the salinization process by facilitating salt movement both through upper and lower soil layers. It further results in depleted annual precipitation and elevated soil temperature. Subsequent heating and cooling promote wear and tear, higher runoff, and substantial sedimentation to cause flooding and salt