Effects of Salinity on Seed Germination and Early Seedling Stage

*Cüneyt Uçarlı*

#### **Abstract**

Salinity is the major environmental stress source that restricts on agricultural productivity and sustainability in arid and semiarid regions by a reduction in the germination rate and a delay in the initiation of germination and subsequent seedling establishment. Salt negatively effects the crop production worldwide. Because most of the cultivated plants are salt-sensitive glycophytes. Salt stress affects the seed germination and seedling establishment through osmotic stress, ion toxicity, and oxidative stress. Salinity may adversely influence seed germination by decreasing the amounts of seed germination stimulants such as GAs, enhancing ABA amounts, and altering membrane permeability and water behavior in the seed. Rapid seed germination and subsequent seedling establishment are important factors affecting crop production under salinity conditions. Seed priming is one of the useful physiological approaches for adaptation of glycophyte species to saline conditions during germination and subsequent seedling establishment. In seed priming, seeds are exposed to an eliciting solution for a certain period that allows partial hydration without radicle protrusion. Seed priming is a simple, low cost, and powerful biotechnological tool used to overcome the salinity problem in agricultural lands.

**Keywords:** salinity, germination, glycophyte, halophyte, seed priming, plant hormones

#### **1. Introduction**

Seed dormancy and germination are distinct physiological processes, and the transition from dormancy to germination is not only a critical developmental step in the life cycle of higher plants but also determines the failure or success of the subsequent seedling establishment and plant growth [1]. Seed germination begins with the water uptake of dry seed (imbibition) and ends with radicle protrusion. Seed germination is affected by adverse environmental conditions including salinity, high temperature, and drought [2].

It is estimated that about approximately 7% of world land is affected by salinity and approximately 20% of 230 million ha irrigated land is salt-affected [3]. This number could be increased in the future due to increased land salinization as a consequence of contaminated artificial irrigation, climate change, and unsuitable land management. Salinity is a major stress responsible for the inhibition of seed germination or reduction in germination percentage and a delay in germination

time in crops. At present, around 30 crop plants provide 90% of plant-based human food and the majority of these crops are not salt tolerant, even salt-sensitive, called glycophytes [4]. There have been high yield losses in these crops under moderate salinity (EC 4–8 dS m−1, approximately 40–80 mM NaCl) [5].

High salinity leads a decrease in osmotic potential of ambient soil water, resulting with a decrease in water uptake by dry seeds (imbibition). Besides, the absorption of excess Na<sup>+</sup> and Cl− ions from soils creates ionic stress and cause toxicity which contributing to disruption in biochemical processes including nucleic and protein metabolism, energy production, and respiration [6]. Salinity also damages the nutrient and hormone balances, especially gibberellin (GA)/abscisic acid (ABA), during germination. As a result, high salinity level causes a delay in germination, even inhibition of seed germination depending on salt tolerance of plants. Dynamic balance between the generation and scavenging of reactive oxygen species (ROS) such as hydroxyl radicals, superoxide, and hydrogen peroxide could be disturbed by high salinity stress. ROS damage the macromolecules including proteins, carbohydrates, nucleic acids, and lipids, or cellular structures like membranes, resulting with inhibition of seed germination [7].

Germination has been found to be under strict regulation of plant hormones, especially GA and ABA [8]. ABA promotes seed dormancy and inhibits germination of seed, whereas GAs release dormancy and stimulate germination. Plant hormones ethylene (ET), and brassinosteroids (BRs) also have positive effect on seed germination by controlling the inhibitory effects of ABA on germination and rupturing testa and endosperm [9, 10]. The plant hormones widely took part in determining the physiological state of a seed and regulating the germination process by interacting each other [11]. Hormones are regulated by distinct transcription factors and signaling components including NO and H2O2, showing the complexity of seed germination regulation. While some plant genes control the activity of plant hormones, and the other plant genes are activated by plant hormones [10]. Signaling molecules, such as NO and H2O2, also promotes germination and reduce the dormancy by enhancing ABA catabolism and GA biosynthesis [12].

Rapid seed germination and subsequent seedling establishment are important factors determining crop production and yield under salinity stress. One of the useful physiological approaches for glycophytes to adapt saline condition is seed priming [7]. Seed priming is an easy, low cost and low risk technique. The seeds are hydrated in specific solutions including plant hormones (GA3, ET, auxins, kinetin), antioxidant compounds (ascorbic acid, glutathione, tocopherone) organic solutes (proline, glycine betaine), inorganic salts (KNO3, CaCl2, and KCl), and particular bacteria and fungi species for a certain time to allow metabolic process of germination, followed by drying the seed to inhibit occurring of radicle protrusion [13].

#### **2. Soil salinity and salinity stress**

Plants, being sessile nature, are simultaneously subjected to various adverse conditions including salinity, drought, cold, heat, excess water, and heavy metals, which limit their development and growth. Salinity is the major environmental stress source that restricts on agricultural productivity and sustainability in arid and semiarid regions [14]. Salinity is a global issue that affects about 7% of the world's total land area, including 20% total cultivated lands and 33% of irrigated land, causing estimated yield losses of 20% worldwide [15, 16]. Besides, it is estimated that every year 10 million ha of agricultural land destroyed by salinized soil [17]. This rate can be increased by global climate change, use of contaminated irrigation water, intensive farming and poor drainage [18–56]. Without proper and

**213**

germination [64].

*Effects of Salinity on Seed Germination and Early Seedling Stage*

associated with intensive farming and poor drainage [57].

−

sustainable control, salinity-affected areas will increase to more than 50% of the world's total arable land by 2050 [15]. This rate can be accelerated by increase in sea water level by climate change, excessive use of groundwater for irrigation, increasing use of low-quality water for irrigation and massive introduction of irrigation

Soil salinity is a measure of the concentration of all the soluble salts in soil water, and is usually expressed as electrical conductivity (EC) of the saturation extract (ECe) with units of deci siemens per meter (1 dS m−1) [58]. The soils were classified as saline, sodic or saline-sodic based on the total concentration of salt and the

to Ca2+ and Mg2+ in the saturated extract of the soil [59]. When the

. The pH of saline soils is usually below 8.5. Sodic

are high in these soils. Saline-sodic soils have similar salt and pH

is the

ECe exceeds 4 dS m−1 (approximately 40 mM/L NaCl) and exchangeable sodium percentage is less than 15 with sodium adsorption ratio (SAR) < 13, the soil is saline. The major problem with saline soils is the presence of soluble salts, primarily

major problem in these soils. Sodic soils have a pH between 8.5 and 10. Saline-sodic soils have an ECe > 4 dS m−1, SAR > 13, and an ESP > 15. Thus, both soluble salts and

levels as saline soils. USSL Staff [59] has described the general relationship of ECe

• very slightly saline (ECe = 2–4 dS m−1): yields of very sensitive crops may be

• slightly saline (ECe = 4–8 dS m−1): yields of many crops are restricted;

• moderately saline (ECe = 8–16 dS m−1): only salt tolerant crops yield

acidity, and nutrient also affect the seed germination [60, 61].

• strongly saline (ECe ≥ 16 dS m−1): only a few very salt tolerant crops yield

Seed germination is a complex multi-stage developmental process and regulated by internal and external factors. Internal factors include proteins, plant hormones (gibberellins/ABA balance, ethylene, and auxin), chromatin-related factors such as methylation, acetylation, histone ubiquitination, related genes (maturating genes and hormonal and epigenetics-regulating genes), non-enzymatic processes, seed age, seed size, and structural components of seed including (endosperm and seed coat). Besides, external factors containing moisture, light, salinity, temperature,

Seed germination begins with imbibition, the uptake of water by the dry mature

seed, and ends with visible protrusion of radicle through testa [62]. Successful germination requires optimum environmental conditions, including water, oxygen, and temperature to initiate this process. Germination/sprouting is regulated by plant hormones such as gibberellic acid (GA), abscisic acid (ABA), ethylene, auxins, cytokinins, and brassinosteroids [63]. Among them, ABA and GA are two important regulators, which play antagonistic roles in seed dormancy and

(alkali) soils have an ECe < 4 dS m−1, ESP > 15, and SAR > 13. Therefore, Na+

• non-saline (ECe ≤ 2 dS m−1): salinity effects mostly negligible;

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

2−, and sometimes NO3

and plant growth as the following:

ratio of Na<sup>+</sup>

exchangeable Na<sup>+</sup>

restricted;

satisfactorily; and

satisfactorily.

**3. Seed germination**

Cl<sup>−</sup> , SO4 *Effects of Salinity on Seed Germination and Early Seedling Stage DOI: http://dx.doi.org/10.5772/intechopen.93647*

*Abiotic Stress in Plants*

tion of excess Na<sup>+</sup>

time in crops. At present, around 30 crop plants provide 90% of plant-based human food and the majority of these crops are not salt tolerant, even salt-sensitive, called glycophytes [4]. There have been high yield losses in these crops under moderate

High salinity leads a decrease in osmotic potential of ambient soil water, resulting with a decrease in water uptake by dry seeds (imbibition). Besides, the absorp-

which contributing to disruption in biochemical processes including nucleic and protein metabolism, energy production, and respiration [6]. Salinity also damages the nutrient and hormone balances, especially gibberellin (GA)/abscisic acid (ABA), during germination. As a result, high salinity level causes a delay in germination, even inhibition of seed germination depending on salt tolerance of plants. Dynamic balance between the generation and scavenging of reactive oxygen species (ROS) such as hydroxyl radicals, superoxide, and hydrogen peroxide could be disturbed by high salinity stress. ROS damage the macromolecules including proteins, carbohydrates, nucleic acids, and lipids, or cellular structures like membranes,

Germination has been found to be under strict regulation of plant hormones, especially GA and ABA [8]. ABA promotes seed dormancy and inhibits germination of seed, whereas GAs release dormancy and stimulate germination. Plant hormones ethylene (ET), and brassinosteroids (BRs) also have positive effect on seed germination by controlling the inhibitory effects of ABA on germination and rupturing testa and endosperm [9, 10]. The plant hormones widely took part in determining the physiological state of a seed and regulating the germination process by interacting each other [11]. Hormones are regulated by distinct transcription factors and signaling components including NO and H2O2, showing the complexity of seed germination regulation. While some plant genes control the activity of plant hormones, and the other plant genes are activated by plant hormones [10]. Signaling molecules, such as NO and H2O2, also promotes germination and reduce

the dormancy by enhancing ABA catabolism and GA biosynthesis [12].

Rapid seed germination and subsequent seedling establishment are important factors determining crop production and yield under salinity stress. One of the useful physiological approaches for glycophytes to adapt saline condition is seed priming [7]. Seed priming is an easy, low cost and low risk technique. The seeds are hydrated in specific solutions including plant hormones (GA3, ET, auxins, kinetin), antioxidant compounds (ascorbic acid, glutathione, tocopherone) organic solutes (proline, glycine betaine), inorganic salts (KNO3, CaCl2, and KCl), and particular bacteria and fungi species for a certain time to allow metabolic process of germination, followed by drying the seed to inhibit occurring of radicle protrusion [13].

Plants, being sessile nature, are simultaneously subjected to various adverse conditions including salinity, drought, cold, heat, excess water, and heavy metals, which limit their development and growth. Salinity is the major environmental stress source that restricts on agricultural productivity and sustainability in arid and semiarid regions [14]. Salinity is a global issue that affects about 7% of the world's total land area, including 20% total cultivated lands and 33% of irrigated land, causing estimated yield losses of 20% worldwide [15, 16]. Besides, it is estimated that every year 10 million ha of agricultural land destroyed by salinized soil [17]. This rate can be increased by global climate change, use of contaminated irrigation water, intensive farming and poor drainage [18–56]. Without proper and

ions from soils creates ionic stress and cause toxicity

salinity (EC 4–8 dS m−1, approximately 40–80 mM NaCl) [5].

and Cl−

resulting with inhibition of seed germination [7].

**2. Soil salinity and salinity stress**

**212**

sustainable control, salinity-affected areas will increase to more than 50% of the world's total arable land by 2050 [15]. This rate can be accelerated by increase in sea water level by climate change, excessive use of groundwater for irrigation, increasing use of low-quality water for irrigation and massive introduction of irrigation associated with intensive farming and poor drainage [57].

Soil salinity is a measure of the concentration of all the soluble salts in soil water, and is usually expressed as electrical conductivity (EC) of the saturation extract (ECe) with units of deci siemens per meter (1 dS m−1) [58]. The soils were classified as saline, sodic or saline-sodic based on the total concentration of salt and the ratio of Na<sup>+</sup> to Ca2+ and Mg2+ in the saturated extract of the soil [59]. When the ECe exceeds 4 dS m−1 (approximately 40 mM/L NaCl) and exchangeable sodium percentage is less than 15 with sodium adsorption ratio (SAR) < 13, the soil is saline. The major problem with saline soils is the presence of soluble salts, primarily Cl<sup>−</sup> , SO4 2−, and sometimes NO3 − . The pH of saline soils is usually below 8.5. Sodic (alkali) soils have an ECe < 4 dS m−1, ESP > 15, and SAR > 13. Therefore, Na+ is the major problem in these soils. Sodic soils have a pH between 8.5 and 10. Saline-sodic soils have an ECe > 4 dS m−1, SAR > 13, and an ESP > 15. Thus, both soluble salts and exchangeable Na<sup>+</sup> are high in these soils. Saline-sodic soils have similar salt and pH levels as saline soils. USSL Staff [59] has described the general relationship of ECe and plant growth as the following:


#### **3. Seed germination**

Seed germination is a complex multi-stage developmental process and regulated by internal and external factors. Internal factors include proteins, plant hormones (gibberellins/ABA balance, ethylene, and auxin), chromatin-related factors such as methylation, acetylation, histone ubiquitination, related genes (maturating genes and hormonal and epigenetics-regulating genes), non-enzymatic processes, seed age, seed size, and structural components of seed including (endosperm and seed coat). Besides, external factors containing moisture, light, salinity, temperature, acidity, and nutrient also affect the seed germination [60, 61].

Seed germination begins with imbibition, the uptake of water by the dry mature seed, and ends with visible protrusion of radicle through testa [62]. Successful germination requires optimum environmental conditions, including water, oxygen, and temperature to initiate this process. Germination/sprouting is regulated by plant hormones such as gibberellic acid (GA), abscisic acid (ABA), ethylene, auxins, cytokinins, and brassinosteroids [63]. Among them, ABA and GA are two important regulators, which play antagonistic roles in seed dormancy and germination [64].

The process of seed germination can be divided into three phases (**Figure 1**) [65]. Phase I begins with imbibition of dry seeds and ends with the early plateau phase of water uptake. Phase II includes reactivation of metabolisms, significant induction of hormonal and enzyme activity using surviving structures and components in the desiccated cells, genes involved in amino acid and nucleic acid synthesis, restarting of cellular respiration with genesis of mitochondria, mobilization of reserved, RNA and protein synthesis machinery [66, 67]. Phase III is postgermination stage involves establishment of seedling and the induction of genes for photosynthetic metabolism after radicle cells elongate and divide [68].

Gibberellins and ABA are two key phytohormones regulating seed germination and seedling growth [69]. While GA breaks dormancy and enhances the seed germination and seedling, ABA inhibits germination and enhances seed dormancy [10]. However, the ratio of the two hormones, rather than the absolute level of each hormone, plays a key role in regulating the breaking of seed dormancy and the onset of germination [70]. GA/ABA balance determines fate of the seed; germination or dormancy. Gibberellins induce the synthesis and production of α-amylase, proteases, and β-glucanases, resulting in the germination of seeds [71]. GAs also stimulate the genes involved in weakening of endosperm and expansion of embryo cell [10]. On the other hand, ABA suppresses expression of many hydrolytic enzyme genes to prevent viviparous germination and inhibits promoting effect of GA on radicle growth and embryo expansion by inhibiting water uptake and hence cell-wall loosening, which is a key step to start germination [72].

Ethylene is a gaseous hormone involved in various processes, including positive regulation of seed germination. Ethylene breaks the primary and secondary dormancy and promotes seed germination by reducing ABA levels or sensitivity [73]. Brassinosteroids (BRs) and auxin induce the secretion of ethylene which works in conjunction with GAs to induce germination [10]. Auxins reduce seed sensitivity to ABA by overexpressing microRNAs and interacting with GAs to counteract ABA suppression during germination [74, 75].

Low temperature decreases seed dormancy and enhances germination in many species, while high temperature has the negative effect on germination and induces secondary dormancy [70]. High temperature down-regulates the genes involved in synthesis of GA synthesis and deactivation of ABA, whereas genes involved in ABA synthesis are up-regulated by high temperature. Therefore, transcriptional changes in ABA and GA metabolism and signal pathways results with inhibition of germination or a delay in germination [76]. Light has been considered both to stimulate

**215**

*Effects of Salinity on Seed Germination and Early Seedling Stage*

germination and to terminate dormancy by increasing the expression of GA anabolic genes, GA3ox1 and GA3ox2, and repressing expression of GA catabolism

seed germination through increasing amylase activities, adjusting K<sup>+</sup>

**4. Effect of salinity on seed germination and early seedling stage**

Salinity affects seed germination process through osmotic stress, ion-specific effects and oxidative stress, shown by decreasing germination rate and extended germination time [82]. Salinity increases external osmotic potential that reduces water uptake during imbibition [83]. Salinity may affect the germination of seeds by the toxic effects of excess sodium and chloride ions on embryo viability [84, 85]. The toxic effects include disruption to the structure of enzymes and other macromolecules, damage to cell organelles and the plasma membrane, the disruption of

In general, seed germination progresses in three phases under normal conditions. Seed germination begins with the rapid water uptake by dry seed (imbibition) (Phase I). A plateau phase, known as phase II, follows this phase. The cellular metabolisms are reactivated, and water uptake is restricted in phase II. This is followed by phase III, a post-germination phase, which is characterized by continuous water uptake until germination is complete (**Figure 1**). Based on these three phases, the inhibition of seed germination or delaying in germination time under salinity stress may be generally ascribed to osmotic stress in the phase I and ionic stress in the phase II. Osmotic stress and ionic stress interact together to inhibit or

Salinity may adversely influence seed germination by decreasing the amounts of seed germination stimulants such as GAs, enhancing ABA amounts, and altering membrane permeability and water behavior in the seed [89]. In higher plants, salinity has been demonstrated to change expression profiles of the genes encoding GA metabolic enzymes, including copalyl diphosphate synthase (CPS), ent-kaurene synthase (KS), ent-kaurene oxidase (KO), ent-kaurenoic acid oxidase (KOA), GA 20-oxidase (GA20ox), GA 3-oxidase (GA3ox) and GA 2-oxidase (GA2ox), resulting

The germination of seeds is characterized by transcriptional induction of hydrolytic enzymes such as α-amylase [90]. The α-amylase is excreted into the endosperm to break the stored starch to metabolizable sugars that provide ready energy and nutrients for the growing embryo and radicle. Salinity stress may have much effect on delayed germination time than on final germination percentage for most crops. A delay of water uptake and a decrease in the activity of α-amylase with an increase in the concentration of NaCl may be main reasons for delaying of the germination time [91].

and enhancing seed respiration and ATP production [81].

respiration, photosynthesis and protein synthesis [85–87].

delay germination of seed during the phase III [88].

with change in endogenous GA levels during germination [12].

In addition to phytohormones, several signal molecules, including as nitric oxide (NO) and reactive oxygen species (ROS), also regulate seed dormancy and germination [68]. ROS is an important regulator during seed germination because of the interaction with lipids, DNA, and protein molecules, as well as phytohormones including ABA and GA in the cell [78]. The biochemical and cellular reactions stimulated by water uptake are accompanied by the generation of ROS [79]. Hydrogen peroxide (H2O2) serves as a signaling hub for the regulation of seed dormancy and germination; the accurate regulation of H2O2 accumulation by the cell antioxidant mechanism is important to achieve a balance between oxidative signaling that enhances germination and oxidative damage that inhibits germination or delays in germination time [80]. N compounds, including NO, promotes

/Na+

balance,

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

gene GA2ox2 [77].

*Effects of Salinity on Seed Germination and Early Seedling Stage DOI: http://dx.doi.org/10.5772/intechopen.93647*

*Abiotic Stress in Plants*

The process of seed germination can be divided into three phases (**Figure 1**) [65]. Phase I begins with imbibition of dry seeds and ends with the early plateau phase of water uptake. Phase II includes reactivation of metabolisms, significant induction of hormonal and enzyme activity using surviving structures and components in the desiccated cells, genes involved in amino acid and nucleic acid synthesis, restarting of cellular respiration with genesis of mitochondria, mobilization of reserved, RNA and protein synthesis machinery [66, 67]. Phase III is postgermination stage involves establishment of seedling and the induction of genes for

Gibberellins and ABA are two key phytohormones regulating seed germination and seedling growth [69]. While GA breaks dormancy and enhances the seed germination and seedling, ABA inhibits germination and enhances seed dormancy [10]. However, the ratio of the two hormones, rather than the absolute level of each hormone, plays a key role in regulating the breaking of seed dormancy and the onset of germination [70]. GA/ABA balance determines fate of the seed; germination or dormancy. Gibberellins induce the synthesis and production of α-amylase, proteases, and β-glucanases, resulting in the germination of seeds [71]. GAs also stimulate the genes involved in weakening of endosperm and expansion of embryo cell [10]. On the other hand, ABA suppresses expression of many hydrolytic enzyme genes to prevent viviparous germination and inhibits promoting effect of GA on radicle growth and embryo expansion by inhibiting water uptake and hence

Ethylene is a gaseous hormone involved in various processes, including positive regulation of seed germination. Ethylene breaks the primary and secondary dormancy and promotes seed germination by reducing ABA levels or sensitivity [73]. Brassinosteroids (BRs) and auxin induce the secretion of ethylene which works in conjunction with GAs to induce germination [10]. Auxins reduce seed sensitivity to ABA by overexpressing microRNAs and interacting with GAs to counteract ABA

Low temperature decreases seed dormancy and enhances germination in many species, while high temperature has the negative effect on germination and induces secondary dormancy [70]. High temperature down-regulates the genes involved in synthesis of GA synthesis and deactivation of ABA, whereas genes involved in ABA synthesis are up-regulated by high temperature. Therefore, transcriptional changes in ABA and GA metabolism and signal pathways results with inhibition of germination or a delay in germination [76]. Light has been considered both to stimulate

*Major events associated with germination and subsequent post-germinative growth (based on [13, 65]).*

photosynthetic metabolism after radicle cells elongate and divide [68].

cell-wall loosening, which is a key step to start germination [72].

suppression during germination [74, 75].

**214**

**Figure 1.**

germination and to terminate dormancy by increasing the expression of GA anabolic genes, GA3ox1 and GA3ox2, and repressing expression of GA catabolism gene GA2ox2 [77].

In addition to phytohormones, several signal molecules, including as nitric oxide (NO) and reactive oxygen species (ROS), also regulate seed dormancy and germination [68]. ROS is an important regulator during seed germination because of the interaction with lipids, DNA, and protein molecules, as well as phytohormones including ABA and GA in the cell [78]. The biochemical and cellular reactions stimulated by water uptake are accompanied by the generation of ROS [79]. Hydrogen peroxide (H2O2) serves as a signaling hub for the regulation of seed dormancy and germination; the accurate regulation of H2O2 accumulation by the cell antioxidant mechanism is important to achieve a balance between oxidative signaling that enhances germination and oxidative damage that inhibits germination or delays in germination time [80]. N compounds, including NO, promotes seed germination through increasing amylase activities, adjusting K<sup>+</sup> /Na+ balance, and enhancing seed respiration and ATP production [81].

#### **4. Effect of salinity on seed germination and early seedling stage**

Salinity affects seed germination process through osmotic stress, ion-specific effects and oxidative stress, shown by decreasing germination rate and extended germination time [82]. Salinity increases external osmotic potential that reduces water uptake during imbibition [83]. Salinity may affect the germination of seeds by the toxic effects of excess sodium and chloride ions on embryo viability [84, 85]. The toxic effects include disruption to the structure of enzymes and other macromolecules, damage to cell organelles and the plasma membrane, the disruption of respiration, photosynthesis and protein synthesis [85–87].

In general, seed germination progresses in three phases under normal conditions. Seed germination begins with the rapid water uptake by dry seed (imbibition) (Phase I). A plateau phase, known as phase II, follows this phase. The cellular metabolisms are reactivated, and water uptake is restricted in phase II. This is followed by phase III, a post-germination phase, which is characterized by continuous water uptake until germination is complete (**Figure 1**). Based on these three phases, the inhibition of seed germination or delaying in germination time under salinity stress may be generally ascribed to osmotic stress in the phase I and ionic stress in the phase II. Osmotic stress and ionic stress interact together to inhibit or delay germination of seed during the phase III [88].

Salinity may adversely influence seed germination by decreasing the amounts of seed germination stimulants such as GAs, enhancing ABA amounts, and altering membrane permeability and water behavior in the seed [89]. In higher plants, salinity has been demonstrated to change expression profiles of the genes encoding GA metabolic enzymes, including copalyl diphosphate synthase (CPS), ent-kaurene synthase (KS), ent-kaurene oxidase (KO), ent-kaurenoic acid oxidase (KOA), GA 20-oxidase (GA20ox), GA 3-oxidase (GA3ox) and GA 2-oxidase (GA2ox), resulting with change in endogenous GA levels during germination [12].

The germination of seeds is characterized by transcriptional induction of hydrolytic enzymes such as α-amylase [90]. The α-amylase is excreted into the endosperm to break the stored starch to metabolizable sugars that provide ready energy and nutrients for the growing embryo and radicle. Salinity stress may have much effect on delayed germination time than on final germination percentage for most crops. A delay of water uptake and a decrease in the activity of α-amylase with an increase in the concentration of NaCl may be main reasons for delaying of the germination time [91].

#### *Abiotic Stress in Plants*

The decrease in the α-amylase activity have been reported to be higher in the saltsensitive genotypes than in the salt-tolerant genotypes. This reduction in the α-amylase activity results with a significant reduction in the translocation of sugars, essential for the developing embryo. Besides, decreasing sugar concentrations also change the osmotic potential of growing cells, resulting in a decrease in water uptake [88].

Both osmotic and ionic effects of salt stress leads to generation of excess reactive oxygen species (ROS) and oxidative damage, which disrupts proteins, lipids, and nucleic acids or the cellular structure including lipid membrane [83].

Plants can be divided into two main groups based on their response to saline stress; salt-tolerant halophytes and salt-sensitive glycophytes (non-halophytes) [6]. The halophytes are plants that are able to grow in the presence of high salt concentrations that generate a low water potential of the soil and kill 99% of other


**217**

*Effects of Salinity on Seed Germination and Early Seedling Stage*

affected by osmotic stress during germination [12].

**5. Alleviation salinity stress on germination by seed priming**

Most crops are highly susceptible to saline soil, even when soil has electrical conductivity (ECe) as low as 3 dS m−1 [119]. Therefore, salinity stress appears to be a major limitation factor for crop productivity. Seed germination and seedling establishments are the two critical stages in plant growth. These stages are the most sensitive to environmental conditions including salinity [120]. Plants are usually seeded within the top layer of the soil which is more saline than lower layers [121]. Salinity stress may delay or prevent germination of germination of high quality seeds, resulting with crop loss. Rapid seed germination and subsequent seedling establishment are important factors affecting crop production under salinity conditions. Therefore, to decrease the negative effects of salinity stress on seed germination, it is important to know to what extent the genotypic variation in the water uptake pattern during these phases is associated with the salt tolerance of genotypes

Seed priming is one of the useful physiological approaches for adaptation of glycophyte species to saline conditions during germination and subsequent seedling establishment. Seed priming is a simple, low cost and powerful biotechnological tool used to overcome the salinity problem by promoting seed germination and seedling establishment in agricultural lands [122]. Seed are exposed to an eliciting solution for a constant period that allows partial hydration, but radicle emergence does not occur by re-drying of seed. Seed germination occurs three distinct phases: (i) imbibition, (ii) lag phase (reactivation of metabolisms) and (iii) protrusion of the radicle through the testa. The goal of seed priming is to extend the lag phase, which allows pre-germinative physiological and biochemical processes, but prevent the seed transition towards full germination [123]. Enhanced and uniformed germination of primed seeds occurs by reduction in the lag time of imbibition, activation of enzyme involved in seed germination, initiation of biochemical mechanisms of cell repair, increase in the RNA content and DNA replication, decrease in ROS and lipid peroxidation with increased activity of antioxidant enzymes including as superoxide dismutase, catalase, and glutathione reductase, and increase in osmotic

Several methods of seed priming have been developed in order to revive seeds under salt stress conditions. Some of these methods are hydro-priming, osmopriming, solid matrix priming, hormonal-priming, bio-priming, chemical priming, and nutripriming [13]. In recent years, many studies have been reported to exhibit the

species. They are adapted to survive and complete their life cycle under saline levels of higher than 200 mM NaCl. However, seed germination was also affected under salt stress and germination percentage was reduced to less than 10% under 1.7 M NaCl [92, 93]. In halophytes, maximum salt tolerance for seed germination has been reported to vary from 1.7 to 0.26 M NaCl depending on halophyte species and other environment conditions such as temperature, moisture, and light

A majority of the common crops, such as tomato, bean, rice, corn, etc., are salinity sensitive or even hypersensitive and they are described as glycophytes [5]. The glycophytes contain 99% of the world's flora and are susceptible to even low levels of salinity (ECe < 4 dS m−1, approximately 40 mM NaCl) [92]. Under conditions of moderate salinity (EC 4–8 dS m−1), all important glycophytic crops reduce average yields by 50–80% [118]. Seed germination in glycophytes is severely inhibited under salinity due to both osmotic stress and ionic toxicity stress, while halophytes are less

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

(**Table 1**).

at the germination stage.

adjustment and starch metabolism [124, 125].

#### **Table 1.**

*Maximum salt tolerance of halophytes and glycophytes at the germination stage.*

#### *Effects of Salinity on Seed Germination and Early Seedling Stage DOI: http://dx.doi.org/10.5772/intechopen.93647*

*Abiotic Stress in Plants*

The decrease in the α-amylase activity have been reported to be higher in the saltsensitive genotypes than in the salt-tolerant genotypes. This reduction in the α-amylase activity results with a significant reduction in the translocation of sugars, essential for the developing embryo. Besides, decreasing sugar concentrations also change the osmotic potential of growing cells, resulting in a decrease in water uptake [88].

nucleic acids or the cellular structure including lipid membrane [83].

*Maximum NaCl concentration at which seed germination percentage reduced to 10–20%.*

*Maximum salt tolerance of halophytes and glycophytes at the germination stage.*

**Plant species \***

Both osmotic and ionic effects of salt stress leads to generation of excess reactive oxygen species (ROS) and oxidative damage, which disrupts proteins, lipids, and

Plants can be divided into two main groups based on their response to saline stress; salt-tolerant halophytes and salt-sensitive glycophytes (non-halophytes) [6]. The halophytes are plants that are able to grow in the presence of high salt concentrations that generate a low water potential of the soil and kill 99% of other

*Salicornia herbacea* 1.7 M NaCl Halophyte [93] *Suaeda aralocapsica* 1.5 M NaCl Halophyte [94] *Limonium vulgare* 1.5 M NaCl Halophyte [95] *Sarcocornia perennis* 1.3 M NaCl Halophyte [96] *Haloxylon ammodendron* 1.3 M NaCl Halophyte [97] *Kochia scoparia* 1.0 M NaCl Halophyte [98] *Kochia prostrata* 0.85 M NaCl Halophyte [99] *Haloxylon salicornicum* 0.8 M NaCl Halophyte [100] *Prosopis juliflora* 0.6 M NaCl Halophyte [100] *Limonium mansanetianum* 0.5 M NaCl Halophyte [101] *Limonium stocksi* 0.4 M NaCl Halophyte [102] *Limonium lilacinum* 0.3 M NaCl Halophyte [103] *Tanacetum cinerariifolium* 0.26 M NaCl Halophyte [104] Quinoa (*Chenopodium quinoa* Willd.) 0.3 M NaCl Halophyte [105] Barley (*Hordeum vulgare* L.). 0.25 M NaCl Glycophyte [106] Maize (Zea mays) 0.24 M NaCl Glycophyte [107] Chicory (*Cichorium intybus* L.) 0.21 M NaCl Glycophyte [108] Lentil (*Lens culinaris* Medik.) 0.2 M NaCl Glycophyte [14] *Brassica napus* 0.2 M NaCl Glycophyte [109] Peanut (*Arachis hypogaea*) 0.2 M NaCl Glycophyte [110] Rice (*Oryza sativa*) 0.16 M NaCl Glycophyte [111] Fig (*Ficus carica* L.) 0.17 M NaCl Glycophyte [112] Button grass (*Dactyloctenium radulans*) 0.1 M NaCl Glycophyte [113] Sorghum (*Sorghum bicolor* Moench) 0.1 M NaCl Glycophyte [114] Ryegrass (*Lolium rigidum*) 0.1 M NaCl Glycophyte [115] Chickpea (*Cicer arietinum* L.) 0.09 M NaCl Glycophyte [116] Tomato (*Solanum lycopersicum*) 0.05 M NaCl Glycophyte [117]

**Maximum salt tolerance**

**Salt tolerance type**

**Reference**

**216**

**Table 1.**

*\**

species. They are adapted to survive and complete their life cycle under saline levels of higher than 200 mM NaCl. However, seed germination was also affected under salt stress and germination percentage was reduced to less than 10% under 1.7 M NaCl [92, 93]. In halophytes, maximum salt tolerance for seed germination has been reported to vary from 1.7 to 0.26 M NaCl depending on halophyte species and other environment conditions such as temperature, moisture, and light (**Table 1**).

A majority of the common crops, such as tomato, bean, rice, corn, etc., are salinity sensitive or even hypersensitive and they are described as glycophytes [5]. The glycophytes contain 99% of the world's flora and are susceptible to even low levels of salinity (ECe < 4 dS m−1, approximately 40 mM NaCl) [92]. Under conditions of moderate salinity (EC 4–8 dS m−1), all important glycophytic crops reduce average yields by 50–80% [118]. Seed germination in glycophytes is severely inhibited under salinity due to both osmotic stress and ionic toxicity stress, while halophytes are less affected by osmotic stress during germination [12].

#### **5. Alleviation salinity stress on germination by seed priming**

Most crops are highly susceptible to saline soil, even when soil has electrical conductivity (ECe) as low as 3 dS m−1 [119]. Therefore, salinity stress appears to be a major limitation factor for crop productivity. Seed germination and seedling establishments are the two critical stages in plant growth. These stages are the most sensitive to environmental conditions including salinity [120]. Plants are usually seeded within the top layer of the soil which is more saline than lower layers [121]. Salinity stress may delay or prevent germination of germination of high quality seeds, resulting with crop loss. Rapid seed germination and subsequent seedling establishment are important factors affecting crop production under salinity conditions. Therefore, to decrease the negative effects of salinity stress on seed germination, it is important to know to what extent the genotypic variation in the water uptake pattern during these phases is associated with the salt tolerance of genotypes at the germination stage.

Seed priming is one of the useful physiological approaches for adaptation of glycophyte species to saline conditions during germination and subsequent seedling establishment. Seed priming is a simple, low cost and powerful biotechnological tool used to overcome the salinity problem by promoting seed germination and seedling establishment in agricultural lands [122]. Seed are exposed to an eliciting solution for a constant period that allows partial hydration, but radicle emergence does not occur by re-drying of seed. Seed germination occurs three distinct phases: (i) imbibition, (ii) lag phase (reactivation of metabolisms) and (iii) protrusion of the radicle through the testa. The goal of seed priming is to extend the lag phase, which allows pre-germinative physiological and biochemical processes, but prevent the seed transition towards full germination [123]. Enhanced and uniformed germination of primed seeds occurs by reduction in the lag time of imbibition, activation of enzyme involved in seed germination, initiation of biochemical mechanisms of cell repair, increase in the RNA content and DNA replication, decrease in ROS and lipid peroxidation with increased activity of antioxidant enzymes including as superoxide dismutase, catalase, and glutathione reductase, and increase in osmotic adjustment and starch metabolism [124, 125].

Several methods of seed priming have been developed in order to revive seeds under salt stress conditions. Some of these methods are hydro-priming, osmopriming, solid matrix priming, hormonal-priming, bio-priming, chemical priming, and nutripriming [13]. In recent years, many studies have been reported to exhibit the

positive effects of seed priming on germination under salinity conditions in many crops (**Table 2**).

Hydro-priming is the simplest and one of the mostly used seed priming method. Hydro-priming depends on seed soaking in pure water without chemical substances for 6–24 h and re-drying to original moisture content prior to sowing without emergence of radicle [144]. This method is a low-cost and environmentally friendly due to no use of additional chemicals. The uncontrolled water uptake by seeds is major disadvantage of this technique. Rapid hydration may cause leakage of


**219**

**Table 2.**

*Effects of Salinity on Seed Germination and Early Seedling Stage*

Priming with 288 μM Gibberellin (GA3) for 32–48 h

Priming with 2 mM silicon (K2SiO3) for 7 days at 25°C in

Priming with 150 ppm gibberellin (GA3) for 24 h

(NaSi) for 36 h

50 μM kinetin

Priming with 200 μM gibberellins (GA4 + 7), 200 μM fluridone (FLU), 200 μM cytokinin (CK), 100 μM sodium nitroprusside (SNP), or 100 μM thiourea (TH) in the dark or light

Inoculation with *Bacillus aryabhattai* SP1016-20

*The functions of seed priming in plant at the germination stage under salinity condition***.**

*Limonium bicolor* Priming with 200 μM melatonin

*Ceratoides lanata* Priming with 10 mM

Priming with 0.3 mM silicon

ethephon, 5 μM fusicoccin or

Soaking with 0.5 mM salicylic acid or 0.1 mM H2O2 at 25°C in

**Plant Treatment Alleviating effect Reference**

Salicylic acid and H2O2 enhanced the germination percentage from 71 to 86 and

SA significantly increased germination rate, germination potential, and germination index of the seeds under 200 mM NaCl

GA3 significantly increased the water uptake, resulting with increased cumulative germination percentage and germination index under

Silicon significantly enhanced the germination rate and percentage, as well as vitality index under 90 mM

[135]

[136]

[122]

[137]

[138]

[139]

[140].

[141]

[142]

[143]

87%, respectively

100 mM NaCl

GA3 enhanced the germination percentage from 56.64 to 76.03% under 100 mM

Silicon enhanced the germination percentage and index, and seedling vigor index

under 200 mM NaCl

200 mM NaCl

NaCl

NaCl

Melatonin significantly increased germination rate, potential and index under

Fusicoccin, kinetin, and ethephon increased the germination percentage from 10 to 40, 50, and 84%, respectively under 900 mM

GA and FLU significantly increased the germination percentage from 7 to 23 and 59% in the light, respectively, while SNP, CK and TH increased the germination percentage from 9 to 54, 55, and 30%, respectively, in the dark under 200 mM NaCl

Inoculation with *B. aryabhattai* enhanced the final germination percentage and mean daily germination from 21.3 to 46.7%, and from 1.6 to 4.5%, respectively, under 510 mM

NaCl

NaCl

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

the dark

*Limonium bicolor* Priming with 80 μM salicylic acid (SA)

the dark

Lentil (*Lens culinaris* cv. Ncir)

Sweet sorghum (*Sorghum bicolor* cv. Chuntian 1)

Maize (*Zea mays*)

Oat (*Avena sativa* cv. NDO-2)

Cucumber (*Cucumis sativus* cv. Jinyou 1)

*Leymus chinensis* cv. Jisheng 3

*Salicornia ramosissima*


*Abiotic Stress in Plants*

crops (**Table 2**).

Barley (*Hordeum vulgare* cv. Bülbül 89)

Wheat (*Triticum aestivum* cv. Chamran)

Pakchoi (*Brassica chinensis* L. cv Tiancuiqing)

*Melilotus officinalis*

*melo*)

Melon (*Cucumis* 

Wheat (*Triticum aestivum* cv. Khirman)

Grain sorghum (*Sorghum bicolor* Moench)

*Zea mays*, *Pisum sativum, Lathyrus sativus*

Tomato (*Solanum lycopersicum* cv. Hezuo 903)

positive effects of seed priming on germination under salinity conditions in many

**Plant Treatment Alleviating effect Reference**

H2O2 increased the germination index from 16.71 to 25.07%, and from 8.19 to 14.65% under 250 mM and 300 mM NaCl,

EGCG increased germination rate and index from 84.7 to 97.0%, and from 29.4 to 35.2%,

Spermidine, proline, and K2SiO3 enhanced the germination rate by 32, 18, and 17%, respectively, under salinity stress (20 dS m−1)

GA3 enhanced germination percentage from 16.67, 26.67, and 50 to 60, 73.3, and 86.67% in *Z. mays*, *P. sativum,* and *L. sativus,* respectively, and resulted in 20% reduction in mean germination time under salinity stress (12 dS m−1)

Germination potential, germination index, and vitality index were increased by 7.67%, 14.20% and 74.51% after 10 μM SNP pre-treatment under

the germination percentage and recovery germination percentage under 200 mM NaCl

Melatonin increase the germination percentage from 50 to 80% under salinity stress

Priming treatments significantly enhanced germination index and final germination percentage, and reduced mean germination time under salinity stress (12 dS m−1)

Treatments improved the speed and percent of germination under 150 mM NaCl

100 mM NaCl

(14 dS m−1)

Soaking with 10 mM Ca2+ Ca2+ significantly increased

[126]

[127]

[128]

[129]

[130]

[131]

[132]

[133]

[134]

respectively

respectively

Priming with aqueous solution of 30 μM H2O2 for 24 h at room

temperature

Priming with 100 μM Epigallocatechin-3-Gallate (EGCG) at 28 ± 3°C

Priming with 0.5 mM spermidine for 24 h, 25 mM proline for 2 days, or 1.5 mM silicon (K2SiO3) for 6 h

Priming with 0.2 g/L GA3 solution for 12 h at room temperature without light.

Priming with sodium nitroprusside (SNP) for 2 h in

Priming with 10–50 μM melatonin for 6 h

Priming with 50 mg L−1 ascorbate, 50 mM proline, 25 μM triacontanol, or 100 μM indole acetic acid for 12 h

Priming with 100–500 mg L−1 nano-iron oxide (n-Fe2O3) for 10 h and soaking with 10 mg L−1 n-Fe2O3 for 3 days

dark at 25 ± 1°C

Hydro-priming is the simplest and one of the mostly used seed priming method. Hydro-priming depends on seed soaking in pure water without chemical substances for 6–24 h and re-drying to original moisture content prior to sowing without emergence of radicle [144]. This method is a low-cost and environmentally friendly due to no use of additional chemicals. The uncontrolled water uptake by seeds is major disadvantage of this technique. Rapid hydration may cause leakage of

**218**

#### **Table 2.**

*The functions of seed priming in plant at the germination stage under salinity condition***.**

essential nutrients out of the seed during germination, resulting in seed damage in some species [145].

Osmo-priming, also known as osmotic conditioning, involves soaking seeds in aerated low water potential solution including sugar, polyethylene glycol (PEG), glycerol, sorbitol, or mannitol with low water potential instead of pure water, followed by air drying before sowing. Due to low water potential of osmotic solutions, water is absorbed slowly by dry seed, which allows gradual seed imbibition [146]. While osmo-priming promotes activation of early phases of germination, inhibiting radicle emergence. Osmo-priming improves seed germination and enhances general crop performance under salt conditions. Water potential of osmotic agent is critical factor since main purpose is to restrict oxidative damage caused by ROS by inhibiting excess water from entering [147]. If inorganic salts such as NaCl, KCl, KNO3, K3PO4, MgSO4, and CaCl2 are used as an osmo-priming agent, the method is generally referred as halopriming.

In hormonal priming, seed imbibition occurs in the presence of plan hormones such as GA3, ethylene, auxins, and salicylic acid, which can gave effect on seed metabolism. Chemical priming is a promising seed priming technique to enhance germination under high salinity stress. Seeds were pre-treated with different chemical solutions used as priming agents. Chemical agents includes a wide range of both natural and synthetic compounds such as antioxidants (ascorbic acid, glutathione, tocopherol, and melatonin), sodium hydrosulfide, polyamines hydrogen peroxide, sodium nitroprusside, urea, selenium, chitosan, fungicide, etc. [13].

Biopriming involves seed imbibition together with particular bacteria or fungi. These microorganisms are able to create endophytic connections with the plant. As other priming method, this treatment increases rate and uniformity of germination under salt conditions, as well as protects seeds against the soil and seed-borne pathogens [147]. The most frequently used biopriming species are *Bacillus* spp., *Enterobacter* spp., *Pseudomonas* spp., and *Trichoderma* spp. [148].

Seed priming efficiency is influence by many factors and strongly depends on treated plant species and chosen priming technique. Physical and chemical factors including osmotica and water potential, priming agent, duration, temperature, presence or absence of light, aeration, and seed condition also influence priming success and determine germination rate and time, seedling vigor, and further plant development [13, 144].

**221**

**Author details**

Department of Molecular Biology and Genetics, Istanbul University, Istanbul,

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

\*Address all correspondence to: ucarlicu@istanbul.edu.tr

provided the original work is properly cited.

Cüneyt Uçarlı

Turkey

*Effects of Salinity on Seed Germination and Early Seedling Stage*

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

#### **Conflict of interest**

No conflict of interest.

*Effects of Salinity on Seed Germination and Early Seedling Stage DOI: http://dx.doi.org/10.5772/intechopen.93647*

*Abiotic Stress in Plants*

some species [145].

generally referred as halopriming.

development [13, 144].

**Conflict of interest**

No conflict of interest.

essential nutrients out of the seed during germination, resulting in seed damage in

Osmo-priming, also known as osmotic conditioning, involves soaking seeds in aerated low water potential solution including sugar, polyethylene glycol (PEG), glycerol, sorbitol, or mannitol with low water potential instead of pure water, followed by air drying before sowing. Due to low water potential of osmotic solutions, water is absorbed slowly by dry seed, which allows gradual seed imbibition [146]. While osmo-priming promotes activation of early phases of germination, inhibiting radicle emergence. Osmo-priming improves seed germination and enhances general crop performance under salt conditions. Water potential of osmotic agent is critical factor since main purpose is to restrict oxidative damage caused by ROS by inhibiting excess water from entering [147]. If inorganic salts such as NaCl, KCl, KNO3, K3PO4, MgSO4, and CaCl2 are used as an osmo-priming agent, the method is

In hormonal priming, seed imbibition occurs in the presence of plan hormones such as GA3, ethylene, auxins, and salicylic acid, which can gave effect on seed metabolism. Chemical priming is a promising seed priming technique to enhance germination under high salinity stress. Seeds were pre-treated with different chemical solutions used as priming agents. Chemical agents includes a wide range of both natural and synthetic compounds such as antioxidants (ascorbic acid, glutathione, tocopherol, and melatonin), sodium hydrosulfide, polyamines hydrogen peroxide,

Biopriming involves seed imbibition together with particular bacteria or fungi. These microorganisms are able to create endophytic connections with the plant. As other priming method, this treatment increases rate and uniformity of germination under salt conditions, as well as protects seeds against the soil and seed-borne pathogens [147]. The most frequently used biopriming species are *Bacillus* spp.,

Seed priming efficiency is influence by many factors and strongly depends on treated plant species and chosen priming technique. Physical and chemical factors including osmotica and water potential, priming agent, duration, temperature, presence or absence of light, aeration, and seed condition also influence priming success and determine germination rate and time, seedling vigor, and further plant

sodium nitroprusside, urea, selenium, chitosan, fungicide, etc. [13].

*Enterobacter* spp., *Pseudomonas* spp., and *Trichoderma* spp. [148].

**220**

## **Author details**

Cüneyt Uçarlı Department of Molecular Biology and Genetics, Istanbul University, Istanbul, Turkey

\*Address all correspondence to: ucarlicu@istanbul.edu.tr

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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**References**

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[85] Daszkowska-Golec A. Arabidopsis seed germination under abiotic stress as a concert of action of phytohormones. OMICS. 2011;**15**:763-774

[86] Parida AK, Das AB. Salt tolerance and salinity effects on plants: A review. Ecotoxicology and Environmental Safety. 2005;**60**(3):324-349

[87] Panda SK, Khan MHG. Growth, oxidative damage and antioxidant responses in Greengram (*Vigna radiata* L.) under short-term salinity stress and its recovery. Journal of Agronomy and Crop Science. 2009;**195**:442-454

[88] El-Hendawy S, Elshafei A, Al-Suhaibani N, Alotabi M, Hassan W, Dewir YH, et al. Assessment of the salt tolerance of wheat genotypes during the germination stage based on germination ability parameters and associated SSR markers. Journal of Plant Interactions. 2019;**14**(1):151-163

[89] Lee SC, Luan S. ABA signal transduction at the crossroad of biotic and abiotic stress responses. Plant, Cell & Environment. 2012;**35**:53-60

[90] Weiss D, Ori N. Mechanisms of cross talk between gibberellin and other hormones. Plant Physiology. 2007;**144**:1240-1246

[91] Kaneko M, Itoh H, Ueguchi-Tanaka M, Ashikari M, Matsuoka M. The α-amylase induction in endosperm during rice seed germination is caused by gibberellin synthesized in epithelium. Plant Physiology. 2002;**128**:1264-1270

[92] Flowers TJ, Colmer TD. Salinity tolerance in halophytes. The New Phytologist. 2008;**179**(4):945-963

[93] Chapman VJ. Salt Marshes and Salt Deserts of the World. New York: Interscience Publishers; 1960

[94] Wang L, Huang Z, Baskin CC, Baskin JM, Dong M. Germination of dimorphic seeds of the desert annual halophyte *Suaeda aralocaspica* (Chenopodiaceae), a C4 plant without kranz anatomy. Annals of Botany. 2008;**102**:757-776

[95] Woodell SRJ. Salinity and seed germination patterns in coastal plants. Vegetation. 1985;**61**:223-229

[96] Redondo S, Rubio-Casal AE, Castillo JM, Luque CJ, Álvarez AA, Luque T, et al. Influences of salinity and light on germination of three Sarcocornia taxa with contrasted habitats. Aquatic Botany. 2004;**78**:255-264

[97] Huang Z, Zhang X, Zheng G, Gutterman Y. Influence of light, temperature, salinity and storage on seed germination of *Haloxylon ammodendron*. Journal of Arid Environments. 2003;**55**:453-464

[98] Khan MA, Gul B, Weber DJ. Effect of salinity and temperature on the germination of Kochia scoparia. Wetlands Ecology and Management. 2001;**9**:483-489

[99] Orlovsky NS, Japakova UN, Shulgina I, Volis S. Comparative study of seed germination and growth of *Kochia prostrata* and *Kochia scoparia* (Chenopodiaceae) under salinity. Journal of Arid Environments. 2011;**75**:532-537

[100] El-Keblawy A, Al-Shamsi N. Effects of salinity, temperature and light on seed germination of *Haloxylon salicornicum*, a common perennial shrub of the Arabian deserts. Seed Science and Technology. 2008;**36**:679-688

[101] Fos M, Alfonso L, Ferrer-Gallego PP, Laguna E. Effect of salinity, temperature and hypersaline conditions on the seed germination in *Limonium mansanetianum* an endemic and threatened Mediterranean species. Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology. 2020;**0**(0):1-7

[102] Zia S, Khan MA. Effect of light, salinity and temperature on the germination of Limonium stocksii. Canadian Journal of Botany. 2004;**82**:151-157

[103] Yildiz M, Cenkci S, Kargioglu M. Effects of salinity, temperature, and light on seed germination in two Turkish endemic halophytes, *Limonium iconicum* and *L. lilacinum* (Plumbaginaceae). Seed Science and Technology. 2008;**36**:646-656

[104] Li J, Yin LY, Jongsma MA, Wang CY. Effect of light, hydro-priming and abiotic stress on seed germination, and shoot and root growth pyrethrum (*Tanacetum cinerariifolium*). Industrial Crops and Products. 2011;**34**:1543-1549

[105] Causin HF, Bordón DAE, Burrieza H. Salinity tolerance mechanisms during germination and early seedling growth in *Chenopodium quinoa* Wild. genotypes with different sensitivity to saline stress. Environmental and Experimental Botany. 2020;**172**:103995

[106] El Goumi Y, Fakiri M, Lamsaouri O, Benchekroun M. Salt stress effect on seed germination and some physiological traits in three Moroccan barley (*Hordeum vulgare* L.) cultivars. Journal of

**229**

*Effects of Salinity on Seed Germination and Early Seedling Stage*

(R.Br.) P.Beauv. Weed Biology and Management. 2019;**19**(3):85-92

[114] Zhu G, An L, Jiao X, Chen X, Zhou G, McLaughlin N. Effects of gibberellic acid on water uptake and germination of sweet sorghum seeds under salinity stress. Chilean Journal of Agricultural Research.

[115] Rahman A, Asaduzzaman M. Statistical modelling of seed

[116] Lavrenko SO, Lavrenko NM, Lykhovyd PV. Effect of degree of salinity on seed germination and initial growth of chickpea (*Cicer arietinum*). Biosystems Diversity.

[117] Moles TM, Guglielminet L, Reyes TH. Differential effects of sodium chloride on germination and post-germination stages of two tomato genotypes. Scientia Horticulturae.

[118] Panta S, Flowers T, Lane P, Doyle R, Haros G, Shabala S.

[119] Francois LE, Maas EV. Crop response and management on saltaffected soils. In: Pessarakli M, editor. Handbook of Plant and Crop Stress. New York: Marcel Dekker; 1994. pp.

[120] Ali E, Hussain N, Shamsi IH, Jabeen Z, Siddiqui MH, Jiang L. Role of jasmonic acid in improving tolerance of rapeseed (*Brassica napus* L.) to Cd toxicity. Journal of Zhejiang University: Science B (Biomedicine & Biotechnology). 2018;**19**(2):130-146

Halophyte agriculture: Success stories. Environmental and Experimental

germination and seedlings root response of annual ryegrass (*Lolium rigidum*) to different stress. Agricultural Research.

2019;**79**(3):415-424

2019;**8**(2):262-269

2019;**27**(2):101-105

2019;**257**:108730

Botany. 2014;**107**:71-83

149-181

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

Materials and Environmental Science.

Motamedi M. Effects of NaCl salinity on maize (*Zea mays* L.) at germination and early seedling stage. African Journal of Biotechnology. 2012;**11**(2):298-304

[107] Khodarahmpour Z, Ifar M,

[108] Vahabinia F, Pirdashti H, Bakhshandeh E. Environmental factors' effect on seed germination and seedling growth of chicory (*Cichorium intybus* L.) as an important medicinal plant. Acta Physiologiae Plantarum.

[109] Wu H, Guo J, Wang C, Li K, Zhang X, Yang Z, et al. An effective screening method and a reliable screening trait for salt tolerance of *Brassica napus* at the germination stage. Frontiers in Plant Science. 2019;**10**:530

[110] Chakraborty K, Bishi SK, Goswami N, Singh AL, Bhaduri D, Zala PV. Salinity-induced changes in seed germination and the expression profile of antioxidant enzymes in peanut as early and late responses in emerging radicles. Acta Physiologiae

Plantarum. 2019;**41**(8):1-16

[111] Hakim MA, Juraimi AS, Begum M, Hanafi MM, Ismail MR, Selamat A. Effect of salt stress on germination and early seedling growth of rice (*Oryza sativa* L.). African Journal of Biotechnology. 2010;**9**(13):1911-1918

[112] Pakyürek M, Dumanoğlu H. Effects of saline stress on in vitro seed germination and seedling growth of some Turkish fig cultivars (*Ficus carica* L.). Applied Ecology and Environmental Research. 2019;**17**(6):13485-13492

[113] Asaduzzaman M, Koetz E, Rahman A. Factors affecting seed germination and emergence of button grass (*Dactyloctenium radulans*)

2014;**5**(2):625-632

2019;**41**(2):1-13

*Effects of Salinity on Seed Germination and Early Seedling Stage DOI: http://dx.doi.org/10.5772/intechopen.93647*

Materials and Environmental Science. 2014;**5**(2):625-632

*Abiotic Stress in Plants*

2008;**102**:757-776

2004;**78**:255-264

2001;**9**:483-489

2011;**75**:532-537

[92] Flowers TJ, Colmer TD. Salinity tolerance in halophytes. The New Phytologist. 2008;**179**(4):945-963

light on seed germination of *Haloxylon salicornicum*, a common perennial shrub of the Arabian deserts. Seed Science and

Technology. 2008;**36**:679-688

[101] Fos M, Alfonso L, Ferrer-Gallego PP, Laguna E. Effect of salinity, temperature and hypersaline conditions on the seed germination in *Limonium mansanetianum* an endemic and threatened Mediterranean species. Plant Biosystems - An International Journal Dealing with all Aspects of Plant

Biology. 2020;**0**(0):1-7

2004;**82**:151-157

2008;**36**:646-656

[102] Zia S, Khan MA. Effect of light, salinity and temperature on the germination of Limonium stocksii. Canadian Journal of Botany.

[104] Li J, Yin LY, Jongsma MA,

[105] Causin HF, Bordón DAE, Burrieza H. Salinity tolerance mechanisms during germination and early seedling growth in *Chenopodium* 

*quinoa* Wild. genotypes with different sensitivity to saline stress. Environmental and Experimental

Botany. 2020;**172**:103995

[106] El Goumi Y, Fakiri M,

Lamsaouri O, Benchekroun M. Salt stress effect on seed germination and some physiological traits in three Moroccan barley (*Hordeum vulgare* L.) cultivars. Journal of

Wang CY. Effect of light, hydro-priming and abiotic stress on seed germination, and shoot and root growth pyrethrum (*Tanacetum cinerariifolium*). Industrial Crops and Products. 2011;**34**:1543-1549

[103] Yildiz M, Cenkci S, Kargioglu M. Effects of salinity, temperature, and light on seed germination in two Turkish endemic halophytes, *Limonium iconicum* and *L. lilacinum* (Plumbaginaceae). Seed Science and Technology.

[93] Chapman VJ. Salt Marshes and Salt Deserts of the World. New York:

[94] Wang L, Huang Z, Baskin CC, Baskin JM, Dong M. Germination of dimorphic seeds of the desert annual halophyte *Suaeda aralocaspica* (Chenopodiaceae), a C4 plant without kranz anatomy. Annals of Botany.

[95] Woodell SRJ. Salinity and seed germination patterns in coastal plants.

[96] Redondo S, Rubio-Casal AE, Castillo JM, Luque CJ, Álvarez AA, Luque T, et al. Influences of salinity and light on germination of three Sarcocornia taxa with contrasted habitats. Aquatic Botany.

[97] Huang Z, Zhang X, Zheng G, Gutterman Y. Influence of light, temperature, salinity and storage on seed germination of *Haloxylon ammodendron*. Journal of Arid Environments. 2003;**55**:453-464

[98] Khan MA, Gul B, Weber DJ. Effect of salinity and temperature on the germination of Kochia scoparia. Wetlands Ecology and Management.

[99] Orlovsky NS, Japakova UN, Shulgina I, Volis S. Comparative study of seed germination and growth of *Kochia prostrata* and *Kochia scoparia* (Chenopodiaceae) under salinity. Journal of Arid Environments.

[100] El-Keblawy A, Al-Shamsi N. Effects of salinity, temperature and

Vegetation. 1985;**61**:223-229

Interscience Publishers; 1960

**228**

[107] Khodarahmpour Z, Ifar M, Motamedi M. Effects of NaCl salinity on maize (*Zea mays* L.) at germination and early seedling stage. African Journal of Biotechnology. 2012;**11**(2):298-304

[108] Vahabinia F, Pirdashti H, Bakhshandeh E. Environmental factors' effect on seed germination and seedling growth of chicory (*Cichorium intybus* L.) as an important medicinal plant. Acta Physiologiae Plantarum. 2019;**41**(2):1-13

[109] Wu H, Guo J, Wang C, Li K, Zhang X, Yang Z, et al. An effective screening method and a reliable screening trait for salt tolerance of *Brassica napus* at the germination stage. Frontiers in Plant Science. 2019;**10**:530

[110] Chakraborty K, Bishi SK, Goswami N, Singh AL, Bhaduri D, Zala PV. Salinity-induced changes in seed germination and the expression profile of antioxidant enzymes in peanut as early and late responses in emerging radicles. Acta Physiologiae Plantarum. 2019;**41**(8):1-16

[111] Hakim MA, Juraimi AS, Begum M, Hanafi MM, Ismail MR, Selamat A. Effect of salt stress on germination and early seedling growth of rice (*Oryza sativa* L.). African Journal of Biotechnology. 2010;**9**(13):1911-1918

[112] Pakyürek M, Dumanoğlu H. Effects of saline stress on in vitro seed germination and seedling growth of some Turkish fig cultivars (*Ficus carica* L.). Applied Ecology and Environmental Research. 2019;**17**(6):13485-13492

[113] Asaduzzaman M, Koetz E, Rahman A. Factors affecting seed germination and emergence of button grass (*Dactyloctenium radulans*)

(R.Br.) P.Beauv. Weed Biology and Management. 2019;**19**(3):85-92

[114] Zhu G, An L, Jiao X, Chen X, Zhou G, McLaughlin N. Effects of gibberellic acid on water uptake and germination of sweet sorghum seeds under salinity stress. Chilean Journal of Agricultural Research. 2019;**79**(3):415-424

[115] Rahman A, Asaduzzaman M. Statistical modelling of seed germination and seedlings root response of annual ryegrass (*Lolium rigidum*) to different stress. Agricultural Research. 2019;**8**(2):262-269

[116] Lavrenko SO, Lavrenko NM, Lykhovyd PV. Effect of degree of salinity on seed germination and initial growth of chickpea (*Cicer arietinum*). Biosystems Diversity. 2019;**27**(2):101-105

[117] Moles TM, Guglielminet L, Reyes TH. Differential effects of sodium chloride on germination and post-germination stages of two tomato genotypes. Scientia Horticulturae. 2019;**257**:108730

[118] Panta S, Flowers T, Lane P, Doyle R, Haros G, Shabala S. Halophyte agriculture: Success stories. Environmental and Experimental Botany. 2014;**107**:71-83

[119] Francois LE, Maas EV. Crop response and management on saltaffected soils. In: Pessarakli M, editor. Handbook of Plant and Crop Stress. New York: Marcel Dekker; 1994. pp. 149-181

[120] Ali E, Hussain N, Shamsi IH, Jabeen Z, Siddiqui MH, Jiang L. Role of jasmonic acid in improving tolerance of rapeseed (*Brassica napus* L.) to Cd toxicity. Journal of Zhejiang University: Science B (Biomedicine & Biotechnology). 2018;**19**(2):130-146

[121] Esechie HA, Al-Saidi A, Al-Khanjari S. Effect of sodium chloride salinity on seedling emergence in chickpea. Journal of Agronomy and Crop Science. 2002;**188**:155-160

[122] Maiti R, Pramanik K. Vegetable seed priming: A low cost, simple and powerful techniques for farmers' livelihood. International Journal of Bio-resource and Stress Management. 2013;**4**:475-481

[123] Hussain T, Koyro HW, Huchzermeyer B, Khan MA. Ecophysiological adaptations of *Panicum antidotale* to hyperosmotic salinity: Water and ion relations and antioxidant feedback. Flora. 2015;**212**:30-37

[124] Lemmens E, Deleu LJ, De Brier N, De Man WL, De Proft M, Prinsen E, et al. The impact of hydro-priming and Osmopriming on seedling characteristics, plant hormone concentrations, activity of selected hydrolytic enzymes, and Cell Wall and Phytate hydrolysis in sprouted wheat (*Triticum aestivum* L.). ACS Omega. 2019;**4**(26):22089-22100

[125] del Carmen Martínez-Ballesta M, Egea-Gilabert C, Conesa E, Ochoa J, Vicente MJ, Franco JA, et al. The importance of ion homeostasis and nutrient status in seed development and germination. Agronomy. 2020;**10**(4):504

[126] Kilic S, Kahraman A. The mitigation effects of exogenous hydrogen peroxide when alleviating seed germination and seedling growth inhibition on salinity-induced stress in barley. Polish Journal of Environmental Studies. 2016;**25**(3):1053-1059

[127] Ahammed GJ, Li Y, Li X, Han WY, Chen S. Epigallocatechin-3-Gallate alleviates salinity-retarded seed germination and oxidative stress in tomato. Journal of Plant Growth Regulation. 2018;**37**(4):1349-1356

[128] Feghhenabi F, Hadi H, Khodaverdiloo H, van Genuchten MT. Seed priming alleviated salinity stress during germination and emergence of wheat (Triticum aestivum L.). Agricultural Water Management. 2020;**231**:106022

[129] Tsegay BA, Andargie M. Seed priming with gibberellic acid (GA 3) alleviates salinity induced inhibition of germination and seedling growth of *Zea mays* L., *Pisum sativum* Var. abyssinicum A. Braun and *Lathyrus sativus* L. Journal of Crop Science and Biotechnology. 2018;**21**(3):261-267

[130] Ren Y, Wang W, He J, Zhang L, Wei Y, Yang M. Nitric oxide alleviates salt stress in seed germination and early seedling growth of pakchoi (*Brassica chinensis* L.) by enhancing physiological and biochemical parameters. Ecotoxicology and Environmental Safety. 2020;**187**:109785

[131] Zhang DW, Vu TS, Huang J, Chi CY, Xing Y, Fu DD, et al. Effects of calcium on germination and seedling growth in *melilotus officinalis* L. (Fabaceae) under salt stress. Pakistan Journal of Botany. 2019;**51**:1-9

[132] Luis Castañares J, Alberto Bouzo C. Effect of exogenous melatonin on seed germination and seedling growth in melon (*Cucumis melo* L.) under salt stress. Horticultural Plant Journal. 2019;**5**(2):79-87

[133] Mahboob W, Khan A. Seed priming modulates germination potential, osmoprotectants accumulation and ionic uptake in wheat seedlings under salt stress screening for abiotic stress view project screening of salt tolerant wheat genotypes view project. International Journal of Agricultural and Biology. 2019;**22**:594-600

[134] Maswada HF, Djanaguiraman M, Prasad PVV. Seed treatment with nanoiron (III) oxide enhances germination, seeding growth and salinity tolerance

**231**

2004;**82**:37-42

*Effects of Salinity on Seed Germination and Early Seedling Stage*

[142] Wu YP, Chen F, Hu XW, Baskin CC, Baskin JM. Alleviation of salinity stress on germination of *Leymus chinensis* seeds by plant growth regulators and nitrogenous compounds

under contrasting light/dark

2016;**71**(3):497-506

conditions. Grass and Forage Science.

[143] Figueira C, Ferreira MJ, Silva H, Cunha A. Improved germination efficiency of *Salicornia ramosissima* seeds inoculated with *Bacillus* 

*aryabhattai* SP1016-20. The Annals of Applied Biology. 2019;**174**(3):319-328

[144] Waqas M, Korres NE, Khan MD, Nizami A, Deeba F, Ali I, et al. Advances in the concept and methods of seed priming. In: Hasanuzzaman M, Fotopoulos V, editors. Priming and Pretreatment of Seeds and Seedlings. Singapore: Springer; 2019. pp. 11-41

[145] Nawaz J, Hussain M, Jabbar A, Nadeem GA, Sajid M, Subtain M, et al. Seed priming a technique. International

Journal of Agriculture and Crop Sciences. 2013;**6**(20):1373-1381

[146] Di Girolamo G, Barbanti L. Treatment conditions and biochemical processes influencing seed priming effectiveness. Italian Journal of

[147] Abdelhamid MT, El-masry RR, Darwish DS, Abdalla MMF, Oba S, Ragab R, et al. Mechanisms of seed priming involved in salt stress amelioration. In: Hasanuzzaman M, Fotopoulos V, editors. Priming and Pretreatment of Seeds and Seedlings. Singapore: Springer; 2019. pp. 219-251

[148] Niranjan Raj S, Shetty NP, Shetty HS. Seed bio-priming with Pseudomonas fluorescens isolates enhances growth of pearl millet plants and induces resistance against downy mildew. International Journal of Pest

Management. 2004;**50**:41-48

Agronomy. 2012;**7**:8-18

*DOI: http://dx.doi.org/10.5772/intechopen.93647*

of sorghum. Journal of Agronomy and Crop Science. 2018;**204**(6):577-587

[135] Bouallègue A, Souissi F, Nouairi I, Souibgui M, Abbes Z, Mhadhbi H. Physiological and biochemicals changes modulated by seeds' priming of lentil (*Lens culinaris* L.) under salt stress at germination stage. Acta Scientiarum Polonorum Hortorum Cultus.

[136] Liu J, Li L, Yuan F, Chen M. Exogenous salicylic acid improves the germination of Limonium bicolor seeds under salt stress. Plant Signaling &

[137] Khan WUD, Aziz T, Waraich EA, Khalid M. Silicon application improves germination and vegetative growth in maize grown under salt stress. Pakistan Journal of Agricultural Sciences.

[138] Chauhan A, Abuamarah BA, Kumar A, Verma JS, Ghramh HA, Ali K, et al. Influence of gibberellic acid and different salt concentrations on germination percentage and physiological parameters of oat cultivars. Saudi Journal of Biological Sciences. 2019;**26**(6):1298-1304

[139] Gou T, Chen X, Han R, Liu J, Zhu Y, Gong H. Silicon can improve seed germination and ameliorate oxidative damage of bud seedlings in cucumber under salt stress. Acta Physiologiae

[140] Li J, Zhao C, Zhang M, Yuan F, Chen M. Exogenous melatonin improves seed germination in Limonium bicolor under salt stress. Plant Signaling & Behavior. 2019;**14**(11):1659705

[141] Khan MA, Gul B, Weber DJ. Action of plant growth regulators and salinity on the seed germination of *Ceratoides lanata*. Canadian Journal of Botany.

Plantarum. 2020;**42**(1):1-11

2019;**18**(5):27-38

Behavior. 2019;**14**:10

2015;**52**(4):937-944

*Effects of Salinity on Seed Germination and Early Seedling Stage DOI: http://dx.doi.org/10.5772/intechopen.93647*

of sorghum. Journal of Agronomy and Crop Science. 2018;**204**(6):577-587

*Abiotic Stress in Plants*

2013;**4**:475-481

[123] Hussain T, Koyro HW, Huchzermeyer B, Khan MA. Ecophysiological adaptations of *Panicum antidotale* to hyperosmotic salinity: Water and ion relations and antioxidant feedback. Flora. 2015;**212**:30-37

[121] Esechie HA, Al-Saidi A,

Al-Khanjari S. Effect of sodium chloride salinity on seedling emergence in chickpea. Journal of Agronomy and Crop Science. 2002;**188**:155-160

Seed priming alleviated salinity stress during germination and emergence of wheat (Triticum aestivum L.). Agricultural Water Management.

[129] Tsegay BA, Andargie M. Seed priming with gibberellic acid (GA 3) alleviates salinity induced inhibition of germination and seedling growth of *Zea mays* L., *Pisum sativum* Var. abyssinicum A. Braun and *Lathyrus sativus* L. Journal of Crop Science and Biotechnology.

[130] Ren Y, Wang W, He J, Zhang L, Wei Y, Yang M. Nitric oxide alleviates salt stress in seed germination and early seedling growth of pakchoi (*Brassica chinensis* L.) by enhancing physiological

and biochemical parameters. Ecotoxicology and Environmental

[131] Zhang DW, Vu TS, Huang J, Chi CY, Xing Y, Fu DD, et al. Effects of calcium on germination and seedling growth in *melilotus officinalis* L. (Fabaceae) under salt stress. Pakistan

Journal of Botany. 2019;**51**:1-9

[133] Mahboob W, Khan A. Seed priming modulates germination potential, osmoprotectants accumulation and ionic uptake in wheat seedlings under salt stress screening for abiotic stress view project screening of salt tolerant wheat genotypes view project. International Journal of Agricultural and Biology.

[134] Maswada HF, Djanaguiraman M, Prasad PVV. Seed treatment with nanoiron (III) oxide enhances germination, seeding growth and salinity tolerance

2019;**5**(2):79-87

2019;**22**:594-600

[132] Luis Castañares J, Alberto Bouzo C. Effect of exogenous melatonin on seed germination and seedling growth in melon (*Cucumis melo* L.) under salt stress. Horticultural Plant Journal.

Safety. 2020;**187**:109785

2020;**231**:106022

2018;**21**(3):261-267

[122] Maiti R, Pramanik K. Vegetable seed priming: A low cost, simple and powerful techniques for farmers' livelihood. International Journal of Bio-resource and Stress Management.

[124] Lemmens E, Deleu LJ, De Brier N, De Man WL, De Proft M, Prinsen E, et al. The impact of hydro-priming and Osmopriming on seedling characteristics, plant hormone concentrations, activity of selected hydrolytic enzymes, and Cell Wall and Phytate hydrolysis in sprouted wheat (*Triticum aestivum* L.). ACS Omega. 2019;**4**(26):22089-22100

[125] del Carmen Martínez-Ballesta M, Egea-Gilabert C, Conesa E, Ochoa J, Vicente MJ, Franco JA, et al. The importance of ion homeostasis and nutrient status in seed development and germination. Agronomy. 2020;**10**(4):504

[126] Kilic S, Kahraman A. The mitigation effects of exogenous hydrogen peroxide when alleviating seed germination and seedling growth inhibition on salinity-induced stress in barley. Polish Journal of Environmental

Studies. 2016;**25**(3):1053-1059

[128] Feghhenabi F, Hadi H,

[127] Ahammed GJ, Li Y, Li X, Han WY, Chen S. Epigallocatechin-3-Gallate alleviates salinity-retarded seed germination and oxidative stress in tomato. Journal of Plant Growth Regulation. 2018;**37**(4):1349-1356

Khodaverdiloo H, van Genuchten MT.

**230**

[135] Bouallègue A, Souissi F, Nouairi I, Souibgui M, Abbes Z, Mhadhbi H. Physiological and biochemicals changes modulated by seeds' priming of lentil (*Lens culinaris* L.) under salt stress at germination stage. Acta Scientiarum Polonorum Hortorum Cultus. 2019;**18**(5):27-38

[136] Liu J, Li L, Yuan F, Chen M. Exogenous salicylic acid improves the germination of Limonium bicolor seeds under salt stress. Plant Signaling & Behavior. 2019;**14**:10

[137] Khan WUD, Aziz T, Waraich EA, Khalid M. Silicon application improves germination and vegetative growth in maize grown under salt stress. Pakistan Journal of Agricultural Sciences. 2015;**52**(4):937-944

[138] Chauhan A, Abuamarah BA, Kumar A, Verma JS, Ghramh HA, Ali K, et al. Influence of gibberellic acid and different salt concentrations on germination percentage and physiological parameters of oat cultivars. Saudi Journal of Biological Sciences. 2019;**26**(6):1298-1304

[139] Gou T, Chen X, Han R, Liu J, Zhu Y, Gong H. Silicon can improve seed germination and ameliorate oxidative damage of bud seedlings in cucumber under salt stress. Acta Physiologiae Plantarum. 2020;**42**(1):1-11

[140] Li J, Zhao C, Zhang M, Yuan F, Chen M. Exogenous melatonin improves seed germination in Limonium bicolor under salt stress. Plant Signaling & Behavior. 2019;**14**(11):1659705

[141] Khan MA, Gul B, Weber DJ. Action of plant growth regulators and salinity on the seed germination of *Ceratoides lanata*. Canadian Journal of Botany. 2004;**82**:37-42

[142] Wu YP, Chen F, Hu XW, Baskin CC, Baskin JM. Alleviation of salinity stress on germination of *Leymus chinensis* seeds by plant growth regulators and nitrogenous compounds under contrasting light/dark conditions. Grass and Forage Science. 2016;**71**(3):497-506

[143] Figueira C, Ferreira MJ, Silva H, Cunha A. Improved germination efficiency of *Salicornia ramosissima* seeds inoculated with *Bacillus aryabhattai* SP1016-20. The Annals of Applied Biology. 2019;**174**(3):319-328

[144] Waqas M, Korres NE, Khan MD, Nizami A, Deeba F, Ali I, et al. Advances in the concept and methods of seed priming. In: Hasanuzzaman M, Fotopoulos V, editors. Priming and Pretreatment of Seeds and Seedlings. Singapore: Springer; 2019. pp. 11-41

[145] Nawaz J, Hussain M, Jabbar A, Nadeem GA, Sajid M, Subtain M, et al. Seed priming a technique. International Journal of Agriculture and Crop Sciences. 2013;**6**(20):1373-1381

[146] Di Girolamo G, Barbanti L. Treatment conditions and biochemical processes influencing seed priming effectiveness. Italian Journal of Agronomy. 2012;**7**:8-18

[147] Abdelhamid MT, El-masry RR, Darwish DS, Abdalla MMF, Oba S, Ragab R, et al. Mechanisms of seed priming involved in salt stress amelioration. In: Hasanuzzaman M, Fotopoulos V, editors. Priming and Pretreatment of Seeds and Seedlings. Singapore: Springer; 2019. pp. 219-251

[148] Niranjan Raj S, Shetty NP, Shetty HS. Seed bio-priming with Pseudomonas fluorescens isolates enhances growth of pearl millet plants and induces resistance against downy mildew. International Journal of Pest Management. 2004;**50**:41-48

**233**

**Chapter 12**

**Abstract**

crop plants.

**1. Introduction**

Abiotic Stress Tolerance in Crop

Crop plants are encountered by various abiotic pressures which limit their growth and development. Stresses such as drought, heat, pathogen attack, heavy metal, salinity, and radiations impose negative effect on crop plants. The reduction in crop productivity in the current era of climate change is compromising the efforts/strategies used for sustainable agricultural practices. Therefore, plant stress physiologists are engineering plants with suitable exogenous signaling elicitors to engineer tolerance to various stresses. In the present chapter, an appraisal has been made in the "Introduction" section to first assess the damages caused by various abiotic stresses in crop plants. In the second section, we attempt to summarize the role of various plant hormones, namely, salicylic acid (SA), brassinosteroids (BRs), ethylene (ET), and methyl jasmonate (MJ) in enhancing abiotic stress tolerance. The current concept may lead to the development of strategies for unraveling the underlying mechanisms of plant hormone-mediated abiotic stress tolerance in

Plants: Role of Phytohormones

*Abdelghafar M. Abu-Elsaoud and Mona H. Soliman*

**Keywords:** abiotic stress, plant hormones, crop plants, stress tolerance

Due to overexploitation of natural resources for catering to basic human needs

coupled with hunger, the aim of Food and Agricultural Organization's [1] of making world agriculture for enhancing 70% food production by 2050 for 2.3 billion newborn individual is facing a lot of challenges. As a result of changing climate and global warming, these threats are now called as "global warning" due to their effects on plant productivity [2]. According to a report published by IPCC [3], global climate change will increase the mean temperature of the Earth by 2–4°C. Due to increased anthropogenic activities, the change in climate will further increase and will definitely affect the agriculture sector as this is being more prone to climate changes [4]. A change in climate scenarios is the main reason of biotic and abiotic pressures of a particular region. Various abiotic stresses all over the world have now been adjudged as the most potential threat for sustainable agricultural productivity [5, 6]. Myriads of anthropogenic emissions in the developmental era have further intensified the abiotic stress-induced negative impacts on agricultural productivity. The major abiotic stresses such as metals/metalloids (hereafter termed as "metal/s"), salt, ozone, UV-B radiation, temperature extremes, nutrient (deficiency and excess), flooding, and water deficit are jeopardizing the agriculture system

*Haifa Abdulaziz S. Alhaithloul,* 

#### **Chapter 12**
