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Biology and Biotechnology of Environmental Stress Tolerance in Plants, Volume 3
as saline stress. This ultimately results in the inhibition of plant growth and
leads to death, thereby resulting in a reduced yield of crop plants. Plants
may be categorized into two major classes: (i) halophytes; and (ii) glyco
phytes. Halophytes are the plants that can tolerate salinity and survive on
the other hand, glycophytes are the group of plants that cannot withstand
saline stress, and as a result, they die eventually. Glycophytic plants account
for a large portion of agricultural plants (Gupta & Huang, 2014). Thus,
salinity is among the most deleterious environmental stressors that hinder
agricultural yield across the world (Flowers, 2004; Munns & Tester, 2008).
Salinity stress contributes to the alteration of different metabolic as well
as cellular activities, the intensity and period of the stress, hamper crop
production (James et al., 2011; Rozema & Flowers, 2008). Abnormal intake
of sodium (Na+) and chloride (Cl–) ions causing cytotoxicity and creates
an imbalance of nutrition, which results in a decrease in plant growth as
well as development through water stress. Salinity goes hand in hand with
oxidative stress since reactive oxygen species (ROS) are formed along with
increased salinity (Isayenkov & Maathuis, 2019). The effect of salinity in
plants has been divided into two phases of plant growth. Stomatal openings
close and the leaf expansion is repressed in phase I, which takes place in a
short time, i.e., minutes or few days. While in phase II, cytotoxic ions build
up especially in mature foliages producing premature senescence, which
downtrend metabolic processes, decrease production eventually kills the
plant (Kotagiri & Kolluru, 2017). The growth and development in agricul
tural plants get hindered due to toxicity caused by ions (especially Na+ ions),
hyper osmotic stress and oxidative damage in a highly saline environment.
Cellular Na+ ion homeostasis is maintained by the secondary messenger
Ca2+ along with Salt Overly Sensitive proteins. Ca2+ signals are decoded
by Ca2+ sensors. The signal is interpreted into physiological, metabolic,
and molecular adaptations by calcineurin B-like protein 10 (CBL10) (Zhu,
2016). Na+ influx is mediated by high‐affinity K+ transporters 1 (HKT1). In
Arabidopsis, considerable methylation is shown at 2.6 kb upstream of the
start site of small RNA which represses the expression of AtHKT1 in shoots
(Baek et al., 2011). The expression of HKT1 is increased due to a decrease
of DNA methylation level in RdDM mutant rdr2; indicating AtHKT1 gene
expression is negatively regulated by RdDM, this control process is even
shown for wheat (Kumar et al., 2017). In the roots and shoots of salt-tolerant
and salt-sensitive plants, the expression of TaHKT2;1 and TaHKT2;3 gets
repressed. This repression is due to cytosine methylation as a result of salt
stress, although the modulation of DNA methylation is not the reason for