JOURNAL OF AGRICULTURE AND BASIC SCIENCES

ISSN Online: 2518-4210

http://www.jabsjournal.com/

Full Length Review Article

Salinity Stress in Crop Plants: Effects of stress, Tolerance Mechanisms and
Breeding Strategies for Improvement
Muhammad Aslam*, Khalil Ahmad, Muhammad Arslan Akhtar and Muhammad Amir Maqbool
Department of Plant Breeding and Genetics, University of Agriculture Faisalabad, Pakistan
*
Corresponding author: [email protected]

Abstract
Salinity, whether primary or secondary, is among the most destructive abiotic stresses that disturb the
plants from germination to physiological maturity. This problem is more severe in arid areas that get low
yearly rainfall and are prone to high evapo-transpiration. Land under salinity stress is increasing on daily
basis and it is thought that about half of the fertile land would become saline by the year 2050. The
effects of salinity are highly diverse and depends on large number of factors like amount, intensity and
duration of salinity and crop growth stages. Increased uptake of toxic ions couples with limited uptake of
essential minerals resulting in significant reduction in enzymatic activity and disturbance in cell
metabolism. Moreover, increased solute potential and reactive oxygen species inside the cells act as
secondary damaging factors to physiological process and plant anatomy. Plants struggle to avoid or
escape the stress by closing the stomata which results in stunted growth, wilting of plants and reduced
productivity. Germplasm can be evaluated using different methods to access the salinity tolerance and
their further administration in different breeding programs could develop tolerant cultivars.
Key Words: Drought Tolerance Indices, PCA, Radar Diagram, Yield and Yield Components

Introduction been drastically reduced. It has been estimated that

Abiotic stress can be referred to as the about 20% yield losses are caused by salt stress.
environmental conditions that decrease plant About 17% crop reduction is by drought stress,
growth and yield from optimum level (Skirycz and 40% by elevated temperature, 20% by low
Inze, 2010). There are a number of types of biotic temperature, and other factors contribute to 8%
and abiotic stresses which cause significant reduction in crop yield (Rehman et al., 2005;
reduction in plant productivity (Aslam et al., 2013 Ashraf et al., 2008). Overall, various abiotic
a, b, c; Aslam et al., 2014 a, b; Aslam et al., 2015 stresses may cause reduction in annual yield of
a, b, c; Aslam et al., 2016; Maqbool et al., 2015 a, crops by 51-82 % at global level (Bray et al.,
b; Maqbool et al., 2016; Naveed et al., 2014). 2000).
Among the abiotic stresses, the most common are
drought, heat, salinity, water logging etc. (Nishida Salinity Stress
and Murata, 1996). Subsoil constraints can be Salt stress explains the presence of soluble
physical (like compaction or gravel layers), salts in excessive amount in the soil which alters
chemical (like salinity, sodicity, acidity, nutrient plant’s normal physiological processes and hinders
deficiencies and toxicities) or biological (high or plant growth. Irrigation water or soil solution with
low microbial activity) and have the ability to limit electric conductivity of 4 dsm-1 or greater (Cramer,
the availability of water and nutrients, thus having 1993) which is equal to 40 mM NaCI is considered
a damaging effect on plant growth (Dang et al., as saline one. Environmental contaminants
2006). Plants respond to abiotic stresses in various specifically those containing salt radicals affect
complex ways (Cramer, 2010). This depends upon crops adversely (Kijne, 2006). Building up of
the intensity of stress i.e. whether it is acute or soluble salts in soil column causes serious damage
chronic as well as duration to which plant is to agricultural production and environmental health
exposed to stress (Tattersall et al., 2007; Pinheiro (Rengasamy, 2006). It has been estimated that up
and Chaves, 2011). Reduction in arable land is to 2050, about 50% of fertile land may be lost due
another factor contributing to yield reduction of to environmental constraints (Manchanda and
agricultural crops caused by limited water Garg, 2008). In order to cope with the increasing
resources, climate change and global warming demands of growing population, 38% increase in
(Lobell et al., 2011). Due to increasing global food production is needed by the year 2025 that
population and excessive exploitation of natural would further rise to 50% by the year 2050. There
resources, a number of problems have been are many factors which are limiting the global
emerged. These include water scarcity, salt stress, demands mainly by physical and chemical
acidity of soil, high temperature, flooding and degradation of soil due to erosion and salinity
diseases by different pathogens. Due to such (Wild, 2003).
environmental stresses, productivity of crops has

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On the basis of source, salinity has been erosion. Almost 14 hundred thousand hectares of
divided into primary and secondary salinity. The agricultural area in Pakistan has been skipped out
main source of primary salinity is the decay of of cultivation which is about 25% irrigated land of
rocks that releases variety of salts. Secondary the country (World Bank, 2006). Subsoil salinity
salinity is caused by anthropogenic activities i.e. had significantly limited crop growth and yield in
irrigation, overgrazing, deforestation and intensive Pakistan (Dang et al., 2006). The soil of Pakistan is
cropping etc. (Ashraf, 1994). generally calcareous in nature, alkaline in chemical
composition and have low organic matter
Global scenario of salinity stress (Sillanpaa, 1982 and Khattak, 1991).
Salinity is major issue of irrigated areas. Approximately, 0.2 to 0.4 percent of the total
Irrigation is practiced on approximately 17% of the cultivable area is being skipped of cultivation each
cultivated land that adds to 30% of global year because of water logging and salinity
agricultural production (Hillel, 2000). Globally, (Qureshi, 1978) and only due to salinity problem,
salinity is severe abiotic stress that causes huge 33% (6.8 Mha) area of cultivation has been
reduction in the growth and development of plant. affected (Anonymous, 2008).
It has been estimated that salt stress has affected
approximately 6% of land area of the world that Mechanism of salt stress on plants
constitutes above 800 million hectares of dry land Osmotic effect
(FAO, 2008). Out of 230 million hectares of Plants are stressed under high salt
irrigated area affected by different abiotic stresses concentrations either by increased osmotic
45 million hectares is affected by salinity potential or by toxic effects of high ionic
worldwide (Ashraf, 2010). According to another concentrations (Brady and Weil, 2002). In osmotic
estimate, globally, 831 x 106 hectares area has been or H2O deficit environments, soluble salts reduce
affected by salts (Beltran and Manzur, 2005). the water potential and make water not freely
Moreover, it is estimated that salinity has affected available to plants for uptake which is the major
one-half of the irrigated land (that is about 2.5 x reason for stunted growth under salinity. It is very
108 ha; Rhoades and Loveday, 1990). About 50% difficult to distinguish between either water
of the arable land will be prone to damages by salt deficiency is due to salinity or drought (Nawaz et
stress up to year 2050 (Manchanda and Garg, al., 2010). The water potential of soil controls new
2008). Saboora et al. (2006) stated that the salinity Galley proof
leaf formation. Rapidly growing cells have the
is degrading about 10% of land area every year in capacity to store higher levels of salts in their
the world. expanding vacuoles, so the growth of the new
The problem of salinity is more adverse in leaves is not restricted due to gathering of salts in
dried areas especially the areas receiving little the cytoplasm (Munns, 2005a). Root and shoot
rainfall with high evapo-transpiration rate and growth is more disturbed because of water stress
temperature (Neto et al., 2006). The problem of the than salt specific effect during the early days of
soil salinity is elevating throughout the world due stress (Munns, 2002). At moderate osmotic stress,
to the use of low quality water resulting in root growth is not much affected whereas the
accumulation of some soluble salts in the root zone reduction of shoot growth is maximum (Hsiao and
of the plants. The problem is further increasing by Xu, 2000). Damage due to osmotic effect is
poor soil management practices (Misra et al., 1997; governed by plant species, time period of stress,
Pitman and Lauchli, 2002). types of cells and tissues and the method of stress
application (Munns et al., 2000).
Scenario of salinity stress in Pakistan Specific ion effect
Pakistan is predominately an agricultural Ion specific toxicity, generally, is because
country and thus, its development depends upon of certain ions like sodium, chloride and sulphate
the improvement of the agriculture sector. Pakistan which are taken up in larger quantities than routine.
comprises of 79.61 million hectares of total It affects the crop right from emergence to
geographical area (Khan et al., 2004) out of which physiological maturity. Crops fail especially when
80% (62,400 km long) land is irrigated with canal specific ions affect at lateral growth stages.
water. Sustainable agriculture of Pakistan is in Regarding tolerance against salt stress different
constant threat by a number of factors like drought, crops have different levels of responses. Most of
soil salinity, change in climate, low and high water the higher plants especially agricultural crops are
stress and abiotic stresses. Among these problems, highly susceptible to this stress (Abrol et al., 1988).
sodicity and salinity are the major ones to the soil Under saline or sodic environments, high
degradation (Khan et al., 2004). concentrations of sodium and chloride ions coupled
Salt stress has declined soil potential for with low concentration of potassium ions was
crop productivity in Pakistan. Improper cultivation observed in leaves of wheat varieties (Maas et al.,
practices and mismanagement of agricultural 1986). Mostly the salts are accumulated in the older
resources are serious factors for the cause of soil leaves of plants. With higher concentration of salt

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accumulation there may be death of leaves; this during the normal body and cell growth (Polle,
happens when the salt concentration is too high, 2001) but increased production occurs during
hence cannot be retained inside the vacuoles. In stressed conditions (Laloi et al., 2004). Osmotic
such cases the excessive salts go to the cytoplasm effect inhibits the stomata opening and decreases
where they affect the normal mechanisms of the CO2 supply for photosynthesis which stimulates
enzyme action. On the other hand, excessive salts the deposition of super oxides in chloroplast. This
cause cell dehydration by being accumulated in the deposition of super oxides promotes the
cell walls (Munns, 2005b). In defense against this photoinhibation and photooxidation in plant cells
effect, plants either try to restrict the salt entry in (Ashraf, 2009). Plants have unique appliances to
their bodies or reduce the amount of salts in their salvage these ROS such as stimulation of the
cytoplasm. Concentration of sodium in the enzymes of antioxidative pathway (Smirnoff,
cytoplasm of the root cells is from 10-30 mM 2005).
(Tester and Davenport, 2003). Due to high
concentrations of sodium and chloride ions inside Effects of salinity on plant growth
leaf sap, root and shoot fresh weight reduces up to Germination stage
50% (Parveen and Qureshi, 1992). Salinity severely affects both glycophytes
Nutritional imbalance and halophytes especially at germination stage
Ions discrepancy is caused by higher (Sosa et al., 2005). Seed germination is very
accretion of sodium and chloride and consequently important stage for the successful establishment of
less absorption of the other minerals such as healthy seedlings which are very sensitive to
calcium, manganese and potassium (Karimi et al., salinity as compared to other vegetative stages.
2005). Elevated Na+: K+ ratio causes enzyme Salinity accumulates the toxic ion in plants causing
inactivation and affects normal metabolic functions a mineral imbalance. The essential ions are reduced
of the plants (Booth and Beardall, 1991). Building and do not meet the demand resulting in hindrance
up of salt deposition disturbs water relations of the in normal physiological activities of plant. High
plants; this results in limited uptake and utilization salt stress retards seed germination process while
of important nutrients. As a result, metabolic low salt stress causes seed dormancy (Khan and
activities of the cell and functioning of the enzymes Weber, 2008). To cope with such nutritional
is disturbed (Lacerda et al., 2003). limitation, seeds develop a mechanism of
Nutrients and salt interaction cause maintaining low water potential (Allen, 1994), or
deficiencies and imbalances of the major nutrients Galley proof

other specific tolerance mechanism to prevent the

(McCue and Hanson, 1990). More uptake of Na + damage due to salt stress (Rumbaugh et al., 1993).
causes reduction in the uptake of potassium and Salinity disturbs germination in a number
symptoms like potassium deficiency are observed of ways. From reducing the osmotic potential of
(Gopal and Dube, 2003). The regulation of calcium soil which makes decline in water imbibition by
within the plant under saline condition is a crucial seed (Khan and Weber, 2008) to the creation of
parameter of plant salt tolerance (Soussi et al., ionic toxicity which alters enzymes action involved
2001). in nucleic acid metabolism. Other impacts of salt
Potassium is main component for protein stress on seed germination include change in
formation, osmoregulation, photosynthesis and metabolism of protein (Rasheed, 2009).
maintenance of cell turgor pressure (Ashraf, 2004). Seeds are more susceptible to salt stress
Decrease in potassium ion uptake in due to salinity due to close association to surface of the soil (Dodd
stress was observed (Marcar et al., 1991). K+ along and Donovan, 1999). With sodium chloride
with Ca+2 are necessary for maintaining the accumulation to a toxic level in soil, ionic stress
integrity and proper working of the cell membranes decreases the rate of germination (Murillo-Amador
(Wenxue et al., 2003). Sufficient amount of K + in et al., 2002). Water absorption by the seed is
plant cell under salinity depends on the uptake on reduced because of lower water potential caused by
selection basis of the potassium ions and salt stress thus posing toxic effects to the
discriminatory compartmentalization of K+ and Na+ developing embryo, resulting in delay in
ions in the shoots (Munns et al., 2000). germination process (Khan and Ungar, 1984).
Oxidative stress The average time of seed germination is
A major effect of salinity is elevation in dependent on salinity stress strength and genotypes.
production of ROS (reactive oxygen species) e.g. With increasing trend of salinity stress there is
H2O2, O2-, and OH- (Mittler, 2002; Munns, 2002). always decreasing rate of germination (Ditommaso,
Proteins, lipids and nucleic acids are damaged 2004). Carpici et al. (2009) estimated the reduction
oxidatively by ROS and hence negatively affect the in germination index of maize cultivars because of
normal cellular metabolism (Imlay, 2003). salt stress. Salinity exerts very pronounced effect
Reduction of oxygen causes formation of these on the germination index and seed size of chickpea
ROS that disturb plant metabolic routes (Asada, (Kaya et al., 2008). Small sized seeds show high
1999). ROS production occurs at minute level value of germination index as compared to large

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size seeds under salinity stress. Salinity delays the affected biological processes ultimately reduces
time to achieve 50% germination in citrus (Zerki, crop yield.
1993). Plant anatomy
According to Farooq et al. (2006) toxicity Salinity has significant effects on plant
of salts on rice seedlings and time to 50% anatomy. Plants adopt various strategies in
germination are reduced if seedlings are treated response of salinity, that allow them to deal with
with ethanol treatment. Salinity has negative effect the problem. Plants with growth in high salt
on the vigor index by raising salt concentration in concentration, have more thickness of leaves
the growing medium (Djanaguiraman et al., 2003). (Waisel, 1991), epidermis, cell walls and cuticles.
Bordi (2010) reported reduction in germination The high salt concentration, increases mesophyll
percentage, rate of germination and germination cell layers and cell size (Zekri and Parsons, 1990),
speed due to salinity. Salinity caused due to more extension in cell wall at high turgor
approximately 32% reduction in germination rate, pressure (Munns and Termaat, 1986). Plants grown
80% in root length, 78% in plumule length, 78% in in salt stress conditions have large in number but
shoot length and 95% in seed vigor of Zea mays narrow xylem vessels as compared to plant grown
(Khodarahmpour et al., 2012). in salt free media (Walker et al., 1985). Salinity
Plant physiology increases the density of stomata of lower side of
Increased amounts of salt in the soil poses leaves and leaf thickness (Raafat et al., 1991) with
a serious threat to different processes of plants increased palisade tissues (Hussein et al., 2012);
which results in reduction of crop productivity. however, it reduces number of cells per leaf.
Epstein (1980) reported reduction in the uptake of Salinity reduces the number of stomata on the
essential ions in the plants due to salinity; this surface of epidermis (Cavisoglu et al., 2007), the
causes alteration in metabolic rates leading to total leaf area, (Awang et al., 1993), leaf
reduction in growth rate. Excessive salt plastochron index (Bray and Reid, 2002). Vascular
concentration in root zone of plant causes change bundle length, xylem rows, number of vessels have
in plant water relations. To deal with the increased also been reported to decline due to salinity
amounts of salinity, the osmotic potential decreases (Hussein et al., 2012). Salinity stress causes
(Rodriguez et al., 1997; Gama et al., 2007; increase in subrin amounts inside the roots (Walker
Kaymakanova and Stoeva, 2008). Salinity causes et al., 1985).
reduction in turgor in plant cells due to reduction in Salinity lowered the xylem development
water uptake by the plant. Low water uptake Galley proof

and width of vascular bundle in mungbean (Beida

reduces cell division and regulation of stomata and Ho, 1993; Rashid et al., 2004). In rice, stem
aperture which ultimately lead to low diameter was reported to be reduced (Pimmongkol
photosynthesis and finally death of plant tissues et al., 2002), while trichome and stomata density
(Marschner, 1995; Munns et al., 2002). Reduction increased. Junghans et al. (2006) reported that
in turgor pressure results in stomata closure which cambial activity is also disturbed by raising the salt
causes reduction in gaseous exchange through concentration in Populus euphratica. A reduction
transpiration (Munns, 1993; Munns and Tester, in thickness and area of mesophyll tissue around
2008). Other physiological activities under the the axis of leaves of kallar grass was found by
influence of salinity include changes in membrane increasing salinity (Ola et al., 2012). Salt stress
permeability leading to destabilization of reduced cell size, epidermal thickness of leaves,
membrane proteins (Gupta et al., 2002; Grattan and apical meristem, diameter of cortex and central
Grieve, 1992) and reduction in the process of cylinder (Reinhardt and Rost, 1995; Javed et al.,
photosynthesis (Sayed, 2003; Kao et al., 2003; 2001). Salinity caused thickening of endo- as well
Ashraf and Shahbaz, 2003). Lowering of as exodermis (Gomes et al., 2011; Degenhardt and
photosynthesis rate happens due to reduction in Gimmler, 2000) and increased development of
enzymes and pigments carrying out photosynthesis sclerenchymatous tissues (Javed et al., 2001). In
(Ashraf and Harris, 2013; Misra et al., 1997; other studies, lignification of intercellular spaces in
Saravanavel et al., 2011). In a study on mungbean exodermis in Vracbiaria decumbens was also
grown under saline conditions, it was found that the observed (Degenhardt and Gimmler, 2000; Gomes
additional increase of leaf Na+ and Cl- also causes et al., 2011).
the production ROS followed by reduced Plant morphology
photosynthetic capacity leading to low plant Once the seed has germinated, next goal
growth (Nazar et al., 2011). Many processes which for plant growth is crop establishment. Salinity
are related with plant physiology and biochemistry cause reduction in crop establishment by reducing
are affected by salinity like photosynthesis (Hayat shoot growth, blocking leaf development and
et al., 2010), water conductance through stomata expansion, reducing growth of internodes and
(Perez-Perez et al., 2009), various biomolecules promoting abscission of leaf (Ziska et al., 1990;
and plant-water relations. All these adversely Zekri, 1991). While studying morphological
attributes of Suaeda salsa, Guan et al. (2011)

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observed that that salt stress decreased shoot decreased by 64% with increase in the root zone
length, diameter and number of branches due to salinity.
increased Na+ and Cl- contents. Salinity accelerates Reddy and Vora (1986) proposed that salt
a number of factors in plants like osmotic stress, stress decreases the yield components as a result of
ion toxicity and nutrient imbalance; these are change in the normal plant metabolism. As already
identified as most prominent causes of reduction in discussed, salinity affects various physiological
crop growth which finally lead to crop failure. growth parameters. The considerable effects are
However, different stages like germination, observed on the grain yield per panicle, seed
vegetative growth, flowering, seed establishment growth. Nahar and Hasanuzzaman (2009) came
and grain filling of crops behave differently with with a result that that salt stress decreased different
salinity. components of yield in V. radiate and rice (Gain et
Salinity affects plant morphology in al., 2004). Kafi and Goldam (2000) determined the
different ways. These modifications depend on response of plants against salinity stress. They
cultivars used, intensity and duration of the stress concluded that salinity poses a serious problem in
(Khan et al., 2003b; Munns and James, 2003). vegetative and reproductive stage in the plants.
Ashraf et al. (2002) studied the morphogenetic
parameters of germplasm of 15 guar ecotypes Mechanism of salt tolerance in plants
under varying degrees of salinity. They concluded Osmotic adjustment
that salinity poses a significant reduction in various Osmotic adjustment or osmoregulation is
vegetative parameters. In different plants the toxic generally considered a significant adaptation to salt
ions effects were observed like burning of the and water stress. The plant with a better osmotic
leaves, chlorosis, reduction in the leaf area and the adjustment mechanism maintains growth and yield
necrosis. The plants affected by salinity have dark and survives better in dry or saline soils because it
green, thicker and succulent leaves. Salt reduced improves cell turgor as well as cell volume
leaf area and volume (Bray and Reid, 2002). The (Munns, 1988). Osmoregulation is considered a
main harmful effects of salinity are reduced key mechanism to ameliorate toxic effects of
germination and emergence, stand and salinity as it causes reduction in osmotic potential
establishment of seedlings (Wahid et al., 1999), because of gathering of some compatible solutes
and enhanced chlorosis and senescence of leaves (Hasegawa et al., 2000). Ions outside the root cause
(Lutts et al., 1996; Wahid et al., 1997; Curtis and osmotic stress (Flowers and Colmer, 2008).
Lauchli, 1987). To cope with osmotic stress, plants Galley proof

In the process of osmosis, a solvent moves

reduce the leaf area and increase the rooting from low to high solute concentration. Roots have
density (Guo et al., 2002; Han and Wang, 2005). high concentration of solutes in the form of sugars
Qureshi et al. (2000) and (Sanadgol, 2002) and various organic compounds as compared to the
reported that in Eucalyptus camaldulens and soil solution outside the roots. Water moves
Agropyron elongatom respectively. Salinity through selectively permeable membrane from cell
reduces leaf bio-mass of stem of maize (Evlagon et walls to the root cells. However, this balance in
al., 1992). Other parameters like area of leaves, solute concentration is disturbed when solute
root and shoot dry weights were also reduced by concentration in soil rises, thereby reduces water
increasing salt levels (Ashrafuzzaman et al., 2002). uptake by plants. To compensate the high
Salinity reduced relative growth and net concentration of solutes outside the root media,
assimilation rate (Khan, 2001; Ahmad, 2010). plants accumulate different osmotica in the form of
Ashraf and Bhatti (2000) stated that in rice, salinity sugars and organic compounds. Among the amino
decreased biomass and leaf area. In sugar beet, acids, proline is necessary for osmotic adjustment
growth and leaf area expansion were reduced even (Zhao and Harris, 1992) by lowering cell water
at very low NaCl concentration (Terry and potential. This needs energy which is provided at
Waldren, 1984). the cost of cut of energy supply to other metabolic
activities. So due to diversion of energy and
Reproductive growth stage resources, overall growth of plants is reduced
Salinity causes about 50% downfall in (Hanson et al., 1999). The osmotic problem
crop growth, productivity and yield throughout the stimulates the decrease in efficiency of utility and
globe. Isla et al. (1998) observed that high level of uptake of other essential nutrients which results in
salinity caused reduction in the growth and yield of an imbalance of nutrients in the plant. This is
barley by 65% and increased ash content. Ahmad et mainly because of competition of sodium with
al. (1995) indicated that with increased salinity other essential elements for availability and uptake
levels, dry matter and seed cotton yield was in soil solutions and in plant roots. The enzymatic
significantly reduced. Khan et al. (1999) evaluated activity and metabolic processes are also
different wheat cultivars against salinity stress for susceptible to the high salt concentration (Lacerda
their intra-varietal variability. They concluded that et al., 2003).
the grain yield decreased 69% and straw yield Synthesis of compatible solutes

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 Genetics of salinity stress in maize / J. Agric. Basic Sci. Vol. 02, No. 01, 2017

One method to avoid toxic effects of growth under high salt concentration (Hasegawa et
salinity is the presence of metabolites which serve al., 2000; Blumwald et al., 2000).
the purpose of compatible solutes. Plants Synthesis of antioxidant
synthesize low molecular weight solutes to The disturbance of cellular homeostasis in
acclimate the toxic effects of salinity; these can be response to different stressed conditions such as
termed as compatible solutes (Hasegawa et al., drought, salinity and cold produce reactive oxygen
2000). Organic osmolytes do not disturb the normal species (ROS) (Dubey, 2011). These are created as
metabolic processes of plants, even at high an additional or due to escape of electrons from
concentration (Rhodes and Hanson, 1993); their electron transport processes carried out in
high concentration stabilizes membrane integrity, chloroplast and mitochondria (Rishi and Sneha,
other macromolecules assemblies and control 2013; Joseph and Jini, 2010). The cellular
enzyme functions (Genard et al., 1991; Schwab and metabolism causes the production of free radical.
Gaff, 1990). These solutes play a major part in
order to maintain the pH of cells, scavenging of Breeding strategies to enhance salt
free radicles produced during oxidation process, tolerance in plants
detoxification of harmful chemicals and serve as Screening and selection of existing germplasm
stored form of nitrogen (Mansour et al., 2000); Scientists have developed different
these molecules guard the cell from the toxic methods to cope the salinity problem. One way is
effects of salt stress and drought (Genard et al., to exploit variations in genomes of the available
1991; Krishnamurthy and Bhagwat, 1991). germplasm for identification of a salt tolerant
Other metabolites that serve as compatible genotype that can enhance crop yield (Ashraf et al.,
solute include polyols, betains (Sakamoto and 2006). Since salinity reduces the crop growth and
Murata, 2000) and soluble sugars (Wahid, 2004) productivity, a proper selection criterion is needed
etc. Among these, osmotic adjustment is to determine the relative salt tolerance (Francois
established by amino acid (Bohnert et al., 1992). and Mass, 1994). Different levels of tolerance
Now it is well established that polyamines, that against salinity exist within plants (Francois and
maintains the solute potential between cells relative Mass, 1994). Many scientists have developed
to its surroundings (Pollard and Wyn Jones, 1979). screening criteria for different crops like wheat (Ali
Ionic compartmentalization et al., 2002), maize (Khan et al., 2003a) and rice
Among different ways to tolerate high (Shannon, 1998).
salinity, one is to compartmentalize inside the cells Galley proof

Some plants show significant tolerance to

(Carlos et al., 2009). The salt stress results in an salinity in a remarkable way that indicates the
imbalance of the nutrients required for plant genetic potential in plants to tolerate the toxic
growth. Ionic stress causes increase of Na+, Cl- and effects of salinity that is required trait in agriculture
decrease of K+ ions. The imbalance of these ions is (Mahmood et al., 2000). Plants accumulate salts
harmful for growth of plants (Abdel Kader et al., according to their capacity to salt tolerance. Plants
2011). Increased Na+ under salinity decreases Ca+2, with capability to exclude more Na+ ions out of the
K+, Mg+2 within the cytosol and disturbs the ionic cell are considered more salt tolerant as compared
homeostasis (Aleman et al., 2011). Na+ is very to plants that accumulate Na+ ions (Schachtman
toxic at high concentration that causes membrane and Munns, 1992). So the salt tolerant species of
destabilization, loss of functional proteins, slows plants accumulate higher quantities of K + and low
down cell division and expansion, lower plant quantities of Na+ ions as compared to salt sensitive
metabolic rates and alters the homeostasis of the species (Tipirdamaz and Cakirlar, 1989). However,
mineral nutrients (Munns and Tester, 2008). Na+ it was proposed that there does not seem any strong
also competes with potassium for cellular uptake relationship in removal of Na+ with tolerance to
when its external concentration is very high (Niu et salinity in wheat; hence both are independent
al., 1995). With increase in Na+ more than 100 mM features of crops. Moreno et al. (2000) determined
in the cytosol, it becomes toxic (Serrano et al., variation in genotypes in cultivars of Phaseolus
1999). vulgaris L. They reported that some bean cultivars
Na+ in high concentration causes had higher root growth and mineral nutrient
reduction in K+ concentration accompanied with accumulation as compared to others. Giaveno et al.
imbalance of K+/Na+ ratio and other electrolytes (2007) performed a screening experiment of
vital for optimum homeostasis of cytosol (Shabala tropical maize for salt tolerance. The experiment
and Cuin, 2008). Plants maintain ionic homeostasis was conducted at germination and seedling stages.
by regular control of Na+ entry in the cell and its They concluded that significant genetic variability
compartmentalization into the vacuole (Zhu, 2003). existed at germination stage while the early
This strategy is very effective in both halophytes seedling growth had no association with the
and glycophytes since it helps in lowering the Na + germination under salinity stress. Akram et al.
and Cl- concentration within the cell and maintains (2010) performed an experiment for screening at
early stages of maize. The genetic variations for

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 Genetics of salinity stress in maize / J. Agric. Basic Sci. Vol. 02, No. 01, 2017

salinity tolerance were determined in maize by divided into primary (caused by decaying of rocks),
hydroponic technique. They used ten hybrids of and secondary (caused by anthropogenic activities).
maize against four levels of salinity i.e. 0, 40, 80 The problem of salinity is more severe in dry areas
and 120 mM. It was observed that salinity had having low rainfall and high evapo-transpiration
significant effect in reducing biomass and rates because the water in the soil is lost to
production of maize. atmosphere and salts cannot leach down causing
Genetic variability for salt tolerance increase in salinity. About 800 Mha of dry land has
To breed crop plants tolerant to salinity, already fallen to salt stress. The rate of the soil
there should be significant variation available in becoming saline is so fast that it is assumed that by
salinity tolerance. However, Noble and Rogers 2050, aabout half of the fertile soil of the globe
(1992) have pointed out that there was very little would become saline. In Pakistan, approximately
variation between varieties for salt tolerance. Plant- 0.2 to 0.4 percent of cultivable area is being put out
plant variations for tolerance against salinity of cultivation each year and almost 25% of the
among different varieties was reported by different irrigated land has already been skipped out because
researchers mostly in cross-pollinated species, e.g. of water logging and salinity by the year 2006.
in alfalfa (Al-Khatib et al., 1993) and maize (Khan Increased solute potential in soil coupled
et al., 2003a). In recent years, Ashraf (2002) has with toxic concentrations of the salt ions affect
screened out plant material from F3 population, growth, development and yield of plants. Highly
developed by cross between LU26S (salt tolerant hypertonic conditions in the soil make water less
wheat cultivar) with cv. Kharchia from India, likely to be available to plants for uptake causing
which grew successfully under S24dsm-1 and S36 dsm-1. stunted growth and even wilting. So it is very
The success of development of salinity tolerant difficult to determine whether water deficiency in
germplasm depends upon the selection criterion to plant is due to drought or salinity. As plant cells
be followed while screening the segregating have ability to store excessive amounts of salts in
generations under saline conditions, and is also their vacuoles, so water stress seems to affect more
subject to heritability of the character. than salt stress.
The potential to select and breed for plants Regarding salinity, Na+. Cl- and SO4-2
with increased salt tolerance may be good if the most commonly affect plants right from
variation shown for that specific trait is genetically germination till physiological maturity of the crop.
determined. Information on the genetic basis of The response against salinity varies from plant to
tolerance would assist the breeder in two ways; Galley proof

plant, but on the average, when the salt

firstly, it helps him to devise appropriate selection concentration increases inside plant, it tends to shift
protocols for screening tolerant plant, and testing it into older leaves to be stored in the valoules of
their progenies, and secondly, it provides estimates the cells. That’s why high levels of sodium and
of heritability of the character. In sorghum, genetic chloride ions and reduced amount of essential
variation demonstrated that salinity tolerance is elements like calcium, potassium and manganese
controlled by both additive and dominant gene can be observed in salinity stressed plants. When
actions (Ashraf et al., 1987). Adetimirin et al., the amount of salinity exceeds plants limit, the salts
2001 reported high interaction of epistatic effects go into cytoplasm of the cells causing full fledge
with environment rather than other gene actions. disturbance in the enzymatic activity and
Revilla et al., 2000 revealed the role of additive eventually death of cells leading to senescence.
and dominance × dominance gene action to govern Another result of salt stress is the over production
most of the traits in control condition studies. In of ROS which cause photoinhibition and
addition to these studies, the reports available on photooxidation.
rice (Shannon et al., 1998), maize (Rao and The amount, duration and intensity of salts
McNeilly, 1999), Triticeae (Xing et al., 2002), in the vicinity coupled with the growth stage of
cotton (Akhtar and Azhar, 2001; Azhar and plant collectively determine plant’s fate. Generally,
Ahmad, 2000; Noor et al., 2001), Aegilops ovala every stage has its own importance regarding
(Farooq, 2002), and lucerne (Al-Khatib et al., growth and production, but salinity at germination
1994) revealed that salt tolerance was heritable in is quite dangerous for plant stature while that at
nature and we could greatly improve these species reproductive stage is importance in lowering the
through direct selection or breeding techniques. production. Plants may adjust their osmotic
potential to minimize salt uptake but this creates
Summary moisture stress. Another important strategy is the
Salinity, as we know that it is one of the production of compatible solutes that play critical
most damaging abiotic stresses, which causes role in maintaining cell pH and lowering toxicity of
serious decrease in growth, development and salts. Cell compartmentalization, and production of
production of almost all biota. If soil has water the antioxidants are also quite acceptable strategies to
electric conductivity of which is greater than 4 save as much as possible. Hence, best strategy
dsm-1, it can be declared as saline. Salinity can be against salinity stress so far is to develop a

76
 Genetics of salinity stress in maize / J. Agric. Basic Sci. Vol. 02, No. 01, 2017

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