Fundamental Response and Salinity Tolerance in Brassicaceae Plants

Fundamental response and salinity tolerance in Brassicaceae
Plants
Saheli Roy1*
1
Amity Institute of Biotechnology, Amity University Uttar Pradesh, Noida – 201313, Uttar
Pradesh, India
*
Author to whom correspondence should be addressed.
Abstract:
This review comprehensively explores the response of Brassica sp. to saline conditions and its
implications for sustainable agriculture. The detrimental effects of soil salinity on agricultural land are
significant, resulting in substantial economic losses and diminished crop yields. Within this context,
salinity disrupts the normal growth and development of Brassica sp. impacting crucial physiological
and biochemical processes, particularly photosynthesis and water use efficiency. Understanding the
mechanisms underlying these responses is critical for developing effective strategies to mitigate the
adverse effects of salinity and ensure sustainable agricultural practices. Researchers have devoted
considerable effort to studying salt-tolerant genotypes of Brassica sp, which exhibit improved resilience
to saline environments. Through comprehensive investigations, they have elucidated the intricate
molecular mechanisms responsible for conferring salt tolerance in these genotypes. These mechanisms
involve the activation and/or regulation of specific salt-tolerant genes, playing a pivotal role in
mitigating the harmful effects of salinity. In addition to molecular mechanisms, plants, including
Brassica juncea, have evolved various morphological, physiological, and biochemical adaptations to
cope with abiotic stresses like salinity. These adaptations include the deposition of organic compounds,
the maintenance of ionic homeostasis, the scavenging of free radicals, and the induction of
phytohormones. These complex responses collectively contribute to the plant's ability to tolerate and
adapt to salinity, ultimately promoting its survival and productivity in challenging environments. By
unravelling the intricacies of the response of Brassica sp. to saline conditions and understanding the
underlying molecular and physiological processes, researchers and agricultural practitioners can
develop targeted strategies to improve salt tolerance in this species and others facing similar challenges.
Implementing these strategies can enhance long-term food production and ensure sustainable
agricultural practices, even in the face of increasing global population and the adverse impacts of
climate change.
Keywords: Brassica sp., Saline conditions, Sustainable agriculture, Crop yields, Growth and
Development
https://doi.org/10.26434/chemrxiv-2023-0xhnx ORCID: https://orcid.org/0009-0009-0042-2485 Content not peer-reviewed by ChemRxiv. License: CC BY-NC-ND 4.0
INTRODUCTION:
In 2050, the world’s population is expected to grow by 34%, to 9.1 billion people, necessitating
a 70% increase in food production (FAO 2050). Meeting the food needs of an ever-increasing
population is a difficult task, as climate change has put agricultural production systems’
sustainability and productivity in jeopardy (Hussain et al.2014). In the field, plants are exposed
to a range of stimulators, including salt below/above optimum levels, drought, cold, and heat.
These are the key constraints impacting crop yields in modern agriculture (Tardieu and
Tuberosa2010; Saud et al. 2013). Abiotic stressors have been blamed for a 50% loss in the
average yield of main crops (Wang et al.2001).
Soil salinity has impacted 30% of irrigated land and 6% of the total land area throughout the
world, resulting in a $12 billion loss in agricultural production (Chaves et al.2009) (Shabala
2013). Soil salinization is serious stress that affects about 831 million hectares of agricultural
areas around the world (Table; FAO2005). Gradual and cumulative salt stress exposure to arid
and semi-arid regions throughout the world is predicted to result in a 30-50 % loss of land by
2050 (Wang et al.2003). The area affected by salt will increase to 16.2 million hectares by
2050, compared to the current 6.74 million hectares in various agroecological regions of India
(Singh et al., 2019). Increased salinity on arable lands indicates the need to better understand
plant tolerance mechanisms to optimize growth conditions and preserve agricultural
productivity.
To avoid damage and ensure life, crop plants evoke a complex and distinct cellular and
molecular reactivity in response to numerous stimuli (Fahad et al.2015). Due to its high
economic and nutritional worth, Brassica sp. ranks third in the world as a source of vegetable
oil. Soil salinity stifles normal Brassica crop growth and development in two ways: first, by
disrupting the osmotic connection of tissues, and second, by particular ion impacts. Reduced
seed germination percentage, early seedling development, plant height, seed yield, and
changing oil quality are all negative consequences of salt on mustard, influencing several
physiological and biochemical processes (Singh et. al. 2014).
1. AIM OF THE REVIEW:
Salinity greatly influences photosynthesis, which is one of the most essential physiological
processes, on the other hand, it reduces the stomatal conductance and internal CO2 pressure
(Yan et al. 2012). The rate of photosynthetic activity decreases as salinity hikes. This may
include stomatal and nonstomatal constraints and photochemical process attenuation (Hichem
et al. 2009). Plants’ water use efficiency is also influenced by stomata, which control CO2 flow
and regulate water flux in plants to prevent water loss (Hentschel et al. 2016). The lower CO 2
absorption and transpiration that resulted from plant sensitivity to salinity have been linked to
a lower water use efficiency under salinity (Omamt et al. 2006; Abedinpour2017; Cruz et al.
2017). Increased water use efficiency in salinity-stressed plants suggests plant salt tolerance,
which can boost plant productivity and is crucial for long-term food supply in salt-stressed
conditions (Sikder et al. 2016). Salinity stress reduces stomatal conductance in all crop plants,
although the effect is worse in salt-sensitive genotypes (James et al. 2002). Furthermore,
salinity slowed the rate of photosynthetic CO2 absorption. This reduction might be owing to
decreased stomatal conductance and CO2 availability for carboxylation inhibition.
With this in mind, researchers looked at changes in photosynthetic features as well as ionic
parameters in salt-resistant and sensitive Indian mustard genotypes to see if they were linked
to salt tolerance. As a result, a series of interrelated molecular mechanisms that include the
activation and/or control of several salt-tolerant genes come into place (Passaia et al. 2013).
Plants use a variety of techniques in response to abiotic challenges, many of which help them
grow and produce more in adverse situations. Changes in morphological and developmental
patterns and physiological and biochemical processes in response to various stimuli are
examples of these phenomena (Tuteja2007; Saudetal.2014). In response to all of these stresses,
metabolic changes accompany the deposition of organic compounds such as sugars, polyols,
betaines, and proline, as well as the protection of cellular machinery, the maintenance of ionic
homeostasis, the scavenging of free radicals, the expression of specific proteins and up-
regulation of their genes, and the induction of phytohormones. (Parida and Das 2005;
Tuteja2007; Munns and Tester2008).
The major goal of this review is to relate the challenges that Brassica juncea faces during saline
conditions and the morphological and biochemical development of the plant under stress.
2. VITAL RESPONSES OF INDIAN MUSTARD TO SALINITY STRESS:
The effects of salinity on plants are two-fold. It all starts with osmotic stress, which reduces
plant cell water permeability and changes the plant's overall water and biological
constitution (Munns and Tester, 2008). The second reaction is linked to the build-up of
excessive sodium concentrations in higher plant parts, which eventually reach hazardous
levels and have a major detrimental impact on plant growth and development by altering
the normal status of potassium, calcium, and critical micronutrients. According to reports,
salinity has been shown to disrupt the proper operation of organelles such as chloroplast
and mitochondria. As a result of its negative effects on photosynthesis and respiration,
Munns and Tester estimated in 2008 that it produces reactive oxygen species (ROs) such
as H2O2 and other free radicals of oxygen, causing severe oxidative damage in plants,
ending in cell death and necrosis. Since salinity is a polygenic feature that interacts with
numerous genes involved in multiple biological processes, plant stress responses vary
greatly from one species to the next. Plant biomass, growth, and physiological,
biochemical, and molecular responses that are altered as a result of salt stress can all be
used to identify a species' response to salinity.
• Oxidative stress
• Antioxidant
Salinity effects
production
Rapid osmotic effect
Saline condition
Later ionic effect
• Changes in gene
expression
Recovery/adaptation • Metabolic
alterations (energy
demands)
• Protein degradation
Fig 1. Flowchart illustrating the primary effects of salinity on plant systems
Brassicaceae is one of the most significant mustard genera, with representatives such as B.
rapa, B. juncea, B. oleracea, B. napus, B. carinata, B. nigra, and B. rupestris being utilized as
vegetables, oilseeds, sauces, and biodiesel production. The majority of the members of the
brassica genus are agriculturally significant, it’s critical to know how different species of
Brassica react to diverse abiotic stressors. Salinity is one of the most critical abiotic restrictions
for agriculture, and earlier research has shown that various species of Brassica behave
differently in a saline environment (Kumar et al. 2009; Chakraborty et al. 2012). The
amphidiploid members of the family, such as B. juncea (AABB), B. carinata (BBCC), and B.
napus (AACC), demonstrated a higher degree of tolerance than their ancestral diploid species,
such as B. rapa (AA), B. nigra (BB), and B. oleracea (CC), according to an interspecific
examination of Brassica genotypes (Malik 1990; Kumar 1995; Purty et al. 2008). Because of
spontaneous hybridization and crossing over between two diploid ancestral genomes, AA (B.
rapa) and CC (B. oleracea), cytogenetic data indicated that amphidiploid species had a stronger
salt-tolerant capacity (Ashraf et al. 2001). According to Ashraf and Mcneilly (2004),
Significant inter-and intra-specific variation was discovered in several Brassica members, and
tolerant individuals maintained a high cellular potassium (K+) and calcium (Ca2+) level to
endure stress.
3. DISCUSSION IN GENERAL: THE BRASSICACEAE FAMILY'S REACTION

TO SALT
3.1 PERFORMANCE OF PLANT BIOMASS AND GROWTH IN THE

PRESENCE OF SALINITY:
Several characteristics are there for determining a plant’s salt sensitivity, amongst which
total plant biomass and growth are crucial. In presence of saline conditions, more than a
few families of plants show the greatest drop in growth and biomass in general. Although
preserving a large amount of plant biomass is an indication of tolerance. Chakraborty et al.
in 2016 disclosed that amidst three distinct Brassicaceae species- B. napus, B. juncea, and
B. oleracea, salinity causes a reduction in fresh weight and dry weight, and the size of the
loss was linked to the quantity of salt build-up and overall salinity tolerance.
3.2 SALINITY INDUCED CHANGES IN LEAF CHLOROPHYLL CONTENT,
PHOTOSYNTHETIC EFFICIENCY, TRANSPIRATION RATE, AND
STOMATAL PARAMETER:
Plants are affected by salt stress in two ways: osmotic (cell dehydration) and lethal ion
accumulation (Flowers 2004). A substantial shift in plant water relations happens in saline
conditions, resulting in the partial closure of stomata to prevent water loss (Kaymakanova
and Stoeva 2008, Gama et al. 2009). This lowers CO2 delivery to leaves, resulting in a
reduced CO2/O2 ratio in chloroplasts that is unfavorable (Remorini et al. 2009). As a result
of the accumulation of photo-reducing power, there is a surplus of electrochemical energy
in the membranes (Zhu 2001). This additional energy is funneled by the Mehler process,
which produces reactive oxygen species (ROs) which include superoxide anion (O2-) and
hydrogen peroxide (H2O2), resulting in oxidative stress syndrome (Hernandez et al. 2000,
Baltruschat et al. 2008). ROs may interact with a variety of cellular components, resulting
in damage to membranes and other cellular structures.
4.2.1. IN A SALINE ENVIRONMENT, THERE IS AN EFFECT ON

PHOTOSYNTHETIC ACTIVITY:
A considerable drop in photosynthesis is regarded as one of the most critical factors

constraining plant growth and production, among the many other negative or inhibitory
impacts of salt stress (Yan et al. 2012). The loss in photosynthetic activity in saline
conditions is mostly leading to a decline in stomatal conductance due to partial stomatal
closure or the breakdown of photosynthetic pigment systems in the mesophyll tissue due to
toxic ion deposition through the xylem sap (Munns 2002, Parida et al. 2004). Hasegawa et
al. (2000) stated that the accumulation of sugars and other suitable solutes is a regular
occurrence under saline circumstances. However, according to Saha et al. (2010),
experienced that using ions to alter osmotic pressure is more energy-efficient, many plants
collect organic osmolytes to cope with osmotic stressors such as proline, betaine, polyols,
sugar alcohols, and soluble sugars. These organic osmolytes act as Osmo protectants by
stabilizing both macromolecular protein structures and highly structured cell membrane
structures (Wassmann et al. 2009). Hayat et al. (2011) demonstrated that proline can work
as a carbon and nitrogen sink, a free radical scavenger, a stabilizer of subcellular structures
(membranes and proteins), and a cellular redox potential buffer. In Brassica species,
increased activity of the proline biosynthesis pathway gene was linked to greater salt
tolerance (Chakraborty et al. 2012a). Chakraborty et al. (2012b) found that stress reduced
leaf chlorophyll content, photosynthetic efficiency, transpiration rate, and stomatal
conductance in three distinct species of Brassica. According to the findings, tolerant
B.napus had the least chlorophyll degradation, followed by moderately tolerant B.juncea,
and susceptible B.oleracea had the highest. During salinity stress, transpiration,
photosynthesis, and stomatal conductance were all reduced; the degree of reduction is the
most critical factor in determining the intra-specific differences in tolerance levels seen in
various Brassica sp members. A lectin gene was isolated from a Leguminosae chickpea and
effectively transformed the lectin gene in B.juncea plants to observe the tolerance in saline
conditions. Kumar et al. (2015) explained that genetically modified plants over-expressing
the lectin gene demonstrated augmented resistance to the Alternaria brassicae leaf necrosis
disease. Further research with the transgenic plants revealed that the gene also provides salt
resistance by maintaining a high level of chlorophyll in the face of stress.
4.2.2. STOMATAL REGULATION IN A SALINE ENVIRONMENT:
Stomata, which assist plants exchange CO2 and water vapor between leaves and the
atmosphere, govern photosynthesis and transpiration. In general, the rate of stomatal
transpiration decreases in saline conditions. Excessive water loss is highly essential under
the influence of osmotic stress; hence stomata closure is the most rapid reaction by plants.
With the high saline condition, net photosynthesis normally declines because the stomata
are also a significant location for CO2 influx in addition a comparison amongst sensitive
and tolerant genotypes showed the exclusive difference between the number of guard cells
and stomatal aperture size under saline conditions. According to Ulfat et al. (2017), salt
stress caused a reduction in net CO2 absorption in B.napus. Furthermore, experiments with
halophytes revealed that guard cells of halophytic members may sustain function even
when grown in high salt concentrations (Robinson et al. 1997). Only the resistant
accessions of B. rapa were able to sustain gas exchange parameters like photosynthetic
efficiency and stomatal conductance at high concentrations of salt stress, according to
Noreen et al. (2010).
3.3 FOR SALINITY TOLERANCE, A LOW SODIUM-POTASSIUM
PROPORTION MUST BE MAINTAINED
ROs have also been shown to directly affect the activity of a variety of Na+, Ca2+, and K+
permeability channels (Demidchik et al. 2007, 2010, Zepeda-Jazo et al. 2011), producing
substantial ionic homeostasis disruptions.
The ionic impact of salt stress on many agricultural plants may be linked to two key
components: Na+ toxicity and K+ retention in the roots and leaves. The buildup of Na+ in
the shoot causes Na+ toxicity (Munns 2002, Tester and Davenport 2003, Mller and Tester
2007). Tester and Davenport (2003) again conveyed that the plant’s root system absorbs
Na+ from the soil and transports it to the shoot via the transpiration stream. The ionic action
of salt stress occurs in both root and shoot tissues, although at a distinct period after the
stress is imposed. Only after extended exposure to salinity treatment does Na+ toxicity in
above-ground plant sections appear to be critical (Munns and Tester 2008). Shabala and
Pottosin & Anschutz et al. (2014) also experienced that another factor that has the
deleterious influence of salinity on root and leaf is the K+ retention, which leads to
decreased metabolic activity and in certain cases, programmed cell death in plant tissues.
Shabala and Pottosin in 2014 reviewed that for many species, there has been a substantial
positive link between K+ retention capacity and overall salinity stress tolerance. However,
a better selectivity towards K+ and reduced absorption of Na+ appears to impart resistance
to salt stress in Brassica (Kumar 1995).
An intricate and extremely effective network of enzymatic and non-enzymatic antioxidants

keeps ROs levels under tight control (Gao et al. 2008). As a result, cellular detoxification
from damaging ROs is a key tolerance technique used by plants to deal with salt stress.
3.4 RESPONSES IN BIOCHEMISTRY:
The availability of ions and nutrients significantly impacts normal cellular activity. Because
it competes with potassium and calcium ion absorption, salinity may have altered the cell’s
ionic and nutritional equilibrium. Dozens of new pieces of evidence suggested that salt was
to blame for the K+ efflux or leakage. As a result, the actual quantity of sodium, potassium,
and calcium in a stressed cell is another essential component in determining the plant’s
tolerance. The low quantity of sodium in the cell and the preservation of a high K+/Na+
ratio has been identified as a characteristic trait for detecting salt tolerance genotypes in
earlier investigations (Shabala and Pottosin, 2014). According to Chakraborty et al.
(2016)’s report, the greatest tissue Na+ level was found in B. oleracea plants. The findings
might be interpreted as evidence of a compensatory mechanism designed to protect this
species from Na+ absorption into the shoot due to its inability to do so effortlessly. This
suggests that they are inefficient in one or more key processes that govern Na+
accumulation in the shoot, such as excluding Na+ from uptake, controlling Na+ loading into
the xylem, or retrieving Na+ from the shoot (Tester and Davenport 2003, Sunarpi et al.
2005, Horie et al. 2009, Shabala 2013, Bose et al. 2014a). Therefore, it can be depicted that
B. oleracea typically develops a highly efficient tissue tolerance mechanism to thrive in
saline circumstances.
3.5 CHANGES IN IONIC BALANCE:
Studies on rapeseed indicated that showing a consistent K+ level in the cell was important
for plant life (Rameeh et al. 2012). In B. campestris, Das et al. (1994) found a substantial
association between Na+ influx and K+ efflux. Under the stressed condition, the ratio of
K+/Na+ was just as critical as the quantity of sodium and potassium. Under stress,
amphidiploid Brassica species were able to sustain a high K+/Na+ ratio when compared to
diploid Brassica species (Ashraf and McNeilly 2004). In tolerant cultivars of Brassica
juncea have exhibited reduced tissue Na+ uptake and decreased Na+/ K+ in both roots and
shoot tissues (Chakraborty et al. 2012). Therefore, salt stress caused significant Na+ uptake
in Brassica plants, increasing [Na+] and reducing [K+] in the leaf sap. In B.napus, the lowest
rise in [Na+] and the lowest drop in [K+] were recorded, two parameters that ultimately
influenced their overall performance. According to Chakraborty et al. (2012), the tolerant
character of B.juncea cultivars CS52 and CS54 was due to the preservation of a low Na+/K+
ratio. In several species, the capacity of a plant to maintain a high cytosolic K+/Na+ ratio
has been frequently cited as a fundamental driver of plant salt tolerance (Dubcovsky et al.
1996, Maathuis and Amtmann 1999, Shabal and Cuin 2008). Many plant species, including
barley (Chen et al. 2005, 2007, Wu et al. 2014), wheat (Cuin et al. 2008, 2012), lucerne
(Smethurst et al. 2008), and poplar (Sun et al. 2009) have been discovered to have a
substantial association between tissue K+ retention and salt tolerance.
3.6 CHLORIDE EXCLUSION IN SALINITY TOLERANCE
Micronutrients such as chloride serve as one of the important micronutrients intended for
ideal growth for most plants (Marschener 1986). Osmoregulation and development are
other significant functions of chloride (Barbier-Brygoo et al. 2000). An excessive amount
of Cl- in a saline environment, on the other hand, can be hazardous to plants (Mengel and
Kirkby 1987). Chakraborty et al. (2016) investigated that harmful Cl- accumulation was
much lower in B. napus, B. juncea, and B. oleracea and was closely linked with overall
plant performance. Furthermore, both under normal and stress conditions, the Cl-
concentration in the leaf tissue was greater than that of [Na+], implying a higher
permeability of Cl- in the tissue and its subsequent buildup during stress. In plants, García
and Medina (2013) have previously investigated that Cl- has a higher tissue permeability
than Na+ in both normal and salt-treated conditions. Some species have been shown to be
more vulnerable to Cl- toxicity than to Na+ toxicity and it is uncertain if this is true with
Brassica (Teakle and Tyerman 2010).
4. UNDER SALINITY STRESS, ORGANIC MOLECULES ARE DEPOSITED AS

ORGANIC OSMOLYTES:
The buildup of greater amounts of inorganic ions (Na+ and Cl-) in the leaf tissue, along
with salt-induced water loss, has resulted in a dramatic rise in leaf sap osmolarity in plants
under salt stress (Chen et al. 2005). The most typical salinity stress reactions are a decrease
in water content, chlorophyll pigments, fresh weight, and dry weight. Plants generate many
forms of osmolyte, such as proline, sugars, and glycine betaine, to maintain cellular
homeostasis and proper water levels in the cells. As a result of the accumulation of these
molecules, the cellular water balance is maintained, and various proteins’ activities in the
cell are preserved. The increased synthesis of osmolytes also helped to restore plasma
membrane integrity and prevent electrolyte leakage. According to Chakraborty et al.
(2016), the overall osmolarity of leaf sap increased the most in B. oleracea, followed by
B. juncea, and the least in B. napus. B. oleracea had the largest contribution of inorganic
osmolytes to osmotic adjustment, whereas B. napus had the lowest, indicating a more
effective mechanism to reject Na+ and its consequent accumulation in the shoot. It also
suggests that when under stress, B. napus rely heavily on the generation of organic
osmolytes to maintain osmotic balance. However, the quantity of proline in the cell is a
significant determinant in deciding the size of the tolerance in B. juncea genotypes among
different forms of osmoprotectants. Furthermore, Murakeozy et al. (2003) established the
fact that an increased cytoplasmic amount of proline can diminish the cell’s osmotic
potential. According to research, glycine betaine (a quaternary ammonium compound
formed in plants from choline) and trehalose levels in the cell are also important in
determining the tolerance threshold (Chakraborty et al. 2012). The sensitive genotypes of
B. juncea, CS52, and CS54, were able to synthesize substantial quantities of proline,
glycine betaine, and trehalose under stress, and the enhanced activity of these compounds
is required for their salt tolerance. Betaine aldehyde dehydrogenase (BADH) and
Trehalose 6 phosphate synthase (T6PS): are the two key enzymes involved in the glycine
betaine and trehalose osmolyte biosynthetic pathway. They are excessively present in the
tolerant genotypes of B. juncea, indicating that the induced activity of these genes is also
important (Chakraborty and Sairam 2017). According to some experts, glycine betaine is
the most efficient osmoprotectant in terms of energy management; it helps plants survive
osmotic and oxidative stress and is created by the action of two enzymes, choline
monooxygenase, and betaine aldehyde dehydrogenase. When compared to control
Brassica plants, the transgenic plants were able to synthesize more glycine betaine and had
improved germination ability under salt stress (Huang et al. 2000). According to Wang et
al. (2010), the choline monooxygenase from Anthrobacter tumefacient enhanced the
amount of glycine betaine in B. campestris, which improved its salt tolerance. A bacterial
Cod A gene can successfully convert B. juncea plants to increase transgenic B. juncea
plants (Prasad et al. 2000). Bhattacharya et al. (2004) effectively transformed the Bet A
gene extracted from bacteria into B. oleracea utilizing the Agrobacterium-mediated
transformation approach; biologically produced Bet A gene that produces more glycine
betaine constantly was shown to be more resistant to salt stress.
5. THE ROLE OF PHYTOHORMONES IN SALINE-TOLERANT BRASSICA:
Endogenous phytohormones such as ABA (Abscisic acid), IAA (Indole 3 acetic acid), GA
(Gibberellic acid), JA (Jasmonic acid), SA (Salicylic acid), BR (Brassinosteroids), and
Triazoles govern plant growth, development, and every mechanism. Pavlovic et al. (2019)
found that phytohormones have a function in environmental reactions. The ABA content
of stomatal movement is well known, and a higher amount of ABA during stress can
provide salt tolerance in Brassica plants (He and Cramer 1996). It's also linked to
improving stomatal closure and reducing water loss in stressful conditions (Ryu and Cho
2015). There are many links between ABA and salt tolerance in the plant kingdom, but
they are extremely dependent on genotype, developmental stage, and stress circumstances.
Increases in ABA have been found in sensitive tomato (Duan et al., 2012), basil
(Mancarella et al., 2016), and barley cultivars in response to salt (Witzel et al., 2014). Early
reactions of a tolerant barley variety (Kamboj et al., 2015) and persistent responses of a
tolerant tomato cultivar have both shown considerable increases in ABA (Gharbi et al.,
2017). Under severe salinity, the halophytes Cakile maritima and Thellungiella salsuginea
may collect ABA more effectively than the glycophyte A. thaliana, according to Ellouzi
et al. (2014). Pavlovic et al. (2019) showed that important hormones like ABA were found
to be positively connected with tolerance levels, and they are plentiful in B. rapa sp.
pekinensis, B. oleracea var. capitata, and B. oleracea var. acephala was positively
connected, whereas JA was adversely correlated. Auxin levels in the cell are also critical
for detail development, and ABA.
Fig 2: Summary of the functions of plant hormones in controlling a plant's ability to

withstand salt stress.
Nishiyama et al. (2011) already established the fact that a molecular interaction between
ABA and CK phytohormones is a negative regulator for salt stress and begins a response
that causes stomatal closure and movement to be delayed. Studies on JA found that
resistant cultivars maintained a low level of jasmonates under stress, which helped to
reduce the salt level inside cells. Krishna (2003), for instance, studied BR and revealed
that increasing the BR level in Arabidopsis might potentially improve overall salt
tolerance. Pavlovic et al. (2019) discovered that salt tolerance was recently revealed to be
favourably linked with the quantity of BR precursor chemical Typhasterol in Brassica
family members. Exogenous administration of active BRs, which enhances their tolerance,
principally through stimulation of H2O2 signalling and antioxidant machinery, has been
used to investigate the role of BRs in salt stress (Sharma et al., 2017). It can also help to
counteract some of the harmful effects of salt on photosynthesis (Siddiqui et al., 2017).
However, the functions of endogenous BRs in salinity stress responses are poorly
understood. BR insensitive BIN 2-1 and BR deficient DET 2-1 are a few examples of
Arabidopsis BR signalling mutants, which are more sensitive to salt stress during seed
germination and seedling growth than wild-type counterparts. However, salt
hypersensitive BR mutants have lower levels of proline and transcripts of salt and ABA-
induced genes (Zeng et al., 2010). The accumulation of BRs in roots was also observed.
In the root-growth bioassay, the significant accumulation of BL in this salt-sensitive
cultivar coincides with its higher suppression of root development. Experiments using
Arabidopsisbes1-D mutants with increased BR Provisional signalling, as well as treatment
of wild-type plants with high levels of BL, have indicated that BRs induce general root
growth inhibition (González-Garca et al., 2011). Because BR signaling is temporarily
blocked in the early stages of root development responses to salt, maintaining appropriate
amounts of active BRs in roots may contribute to tolerance mechanisms (Geng et al. 2013).
6. MOLECULAR INTERACTION IN SALINE ENVIRONMENT:
Typically, plants employ three distinct types of defense mechanisms to counteract the
damaging effects of salt stress.
I. Restricting the inflow of sodium ions.
II. Sodium exclusion approach that directly eliminates sodium from the soil.
III. Sodium compartmentation strategy: Cells typically store sodium in vacuoles to
keep the level of sodium in the cytoplasm non-toxic. The SOS gene family
members SOS1, SOS2, SOS3, HKT1, and 2, and H+ pumps found in the plasma
membrane are the primary players among the several processes that cause Na+
exclusion.
SOS gene family prevents sodium from entering or leaving the cytoplasm:
A. thaliana's plasma membrane has the AtSOS1 gene, which Shi et al. (2000) initially
identified as a possible Na+/H+ transporter. This gene aids in maintaining low amounts
of sodium inside the cell by excluding sodium during salt stress. The SOS2-SOS3
complex actively regulates the activity of the SOS1 gene. SnRK3 stands for sugar non-
fermenting-1-related protein kinase-3, and SOS2 is a protein kinase that is a member of
that family. SOS3 encodes a calcium-binding protein that detects the Ca2+ signal. Due
to an increase in the amount of sodium in the cell, the calcium concentration rises and
activates calcium-binding protein 8 (also known as calcineurin B-like CBL10), which
then binds to SOS2 to activate the SOS2-SOS3 complex and phosphorylates SOS1 to
activate it. Using SOS1 mutant lines of A. thaliana, it was functionally demonstrated that
SOS1 contributes to the maintenance of cellular homeostasis and ion exclusion (Shi et
al. 2002).
According to reports, SOS1 is involved in excluding sodium from the cytosol to the
surrounding medium of epidermal cells and from vascular tissues to the surrounding
parenchyma cells when expressed excessively (Na+/H+antiportergene-AtSOS1) in
Arabidopsis in a constitutive way. The SOS1 gene was cloned for this purpose while
being controlled by the constitutive promoter CaMV35S. The AtSOS1 gene was
overexpressed in transgenic plants, which showed better salt tolerance and the ability to
maintain low Na+ concentrations under high salt concentrations. Thus, by restricting the
accumulation of sodium in cells, it is also feasible to increase the level of salt tolerance.
Kumar et al. (2009) investigated the expression patterns of several SOS family genes in
B. juncea cv CS52 and B. napus. According to the findings to endure stress, amphidiploid
B. juncea and diploid B. napus retained a distinctive transcript abundance pattern of SOS.
The expression of SOS was high in the salt-tolerant genotype of B. juncea, as
demonstrated by Chakraborty et al. (2012b), and induction of three distinct SOS family
members (SOS1, SOS2, and SOS3) was different in B. juncea and B. campestris.
The sodium enters the cell through NSCC (Non-Specific Cation Channel) channels at
high external sodium concentrations, which raises the sodium concentration inside the
cell. The expression of the vacuolar CAX and ACA gene families is linked to the rapid
release of calcium when sodium levels are elevated. Additionally, the inactive SOS3
protein is activated by calcium, and the active SOS3 combines with the dormant SOS2
protein to activate the H+-mediated Na+ antiporter SOS1 protein, which prevents sodium
from entering the cell.
Family Member of the High-Affinity K+ Transporter Gene helps to provide

tolerance:
The members of the class of integral membrane proteins known as high-affinity
potassium transporters are found in the plasma membrane of plants and enable ion
transport across the membrane in an energy-dependent way. Studies using the HKT gene
family have demonstrated that it contributes to salt tolerance. There are several
indications that HKT members actively transport Na+ and K+ to preserve cellular
homeostasis. AtHKT1.1 is largely found in the plasma membrane of xylem parenchyma
cells, where it enables Na+ removal from xylem vessels and loads sodium into the cells.
Sunarpi et al. (2005) demonstrated this. Similarly, AtHKT1.1 plays a significant role in
safeguarding the photosynthesis system in Arabidopsis by removing sodium ions from
the leaves (Horie et al., 2009). Moller et al. (2009) demonstrated that plants over-
expressing the AtHKT1.1gene were more tolerant and able to sustain low sodium in the
upper plant parts, validating the role of the AtHKT1.1gene in salt tolerance. Evidence
showed that AtHKT1.1 plays a crucial role in Na+ exclusion from leaves in Arabidopsis
thaliana and aids in maintaining K+ homeostasis in leaves during salt stress. Accordingly,
it is evident from the earlier results that the AtHKT1;1 gene played multiple roles in cells
by removing Na+ along with releasing K+ from xylem parenchyma cells to xylem
vessels.
Plasma Membrane-Related H+ Pumps:

The ions must be transported across the plasma membrane of plant cells via the proton
gradient. Evidence suggested that two distinct proton pumps, known as P-type H+-
ATPase and P-type H+-PPase, were present in the plasma membrane of plants. According
to Fuglsang et al. (2011), both of these pumps aid in the removal of protons from cells
by forming phosphorylated aspartyls, which are used to generate energy and phosphate
groups from ATP. According to reports, 11 distinct kinds of AHA genes exist in the A.
thaliana genome. AHA1 and AHA2 are the most prevalent kinds of AHA isoforms in
plants and are mostly expressed in their leaves and roots (Fuglsang et al. 2011).
Vacuolar-related H+ pumps:
According to Popova and Golldack (2007) and Krebs et al. (2010), the activity of the
vacuolar H+pumps is closely related to the activities of the genes found in the vacuolar
membrane, such as NHX1. The proton pumps vacuolar H+-ATPase (V-ATPase) and
vacuolar H+ pyrophosphatase (V-ATPase) were found in the vacuoles of A. thaliana.
There are several instances when increased salinity tolerance is provided by vacuolar H+-
ATPase overexpression. The Arabidopsis AVP1 gene was cloned and constitutively
overexpressed by Gaxiola et al. in 1999. According to reports, plants that overexpress
the AVP1 construct have better salt tolerance. Similar improvements in salt tolerance
were seen in yeast cells that overexpressed the AVP1 gene. Vacuolar ATPase Gene
(AVP1) from Arabidopsis was successfully converted into sugarcane by Kumar et al.
(2013). The Arabidopsis AVP1 gene was overexpressed in transgenic sugarcane plants,
and they displayed increased drought and salt tolerance.
7. CONCLUSION:
Following a thorough examination of the literature on the salt tolerance of Brassicas, it

is clear that species respond to saline growth media in different ways at different growth
stages. Firstly, plants are often more sensitive at the germination and seedling stages than
at the mature state. The second parameter would be, that the criteria for distinguishing
between salt-tolerant and salt-sensitive species cannot be based on a single plant
characteristic. Finally, there are conflicting data on the relationship between ion
exclusion and salt tolerance in brassicas, although salt tolerance is usually linked to ion
exclusion.
8. FUTURE PERSPECTIVE:
According to studies, the diploid members of this family are more vulnerable to salt than
amphidiploid species. Various morphological, physiological, biochemical, and
molecular studies with glycophytic and halophytic members of the Brassicaceae family
have been conducted to date, revealing that retention of growth and biomass (shoot
length, root length, dry weight, and fresh weight) under stress was the most legitimate
and differentiating factor for salt tolerance in the Brassicaceae family. Other
physiological responses, including water potential retention, relative water content, total
chlorophyll, and membrane integrity, have also been proven to be beneficial in
identifying salt-tolerant genotypes. Osmotic adjustment occurs in plants exposed to a
saline substrate, but it occurs to a greater extent in salt-tolerant Brassica species or
cultivars, owing to the accumulation of various types of organic osmotica such as total
soluble sugars, free amino acids, and free proline in most salt-tolerant Brassica species.
Under salt stress, tolerant members of the family might produce more proline, glycine
betaine, and other organic chemicals, allowing them to preserve cellular homeostasis.
Different phytohormones such as ABA, IAA, and BR were found to be enhanced under
stress, whereas CK and JA were found to be lowered, preventing water loss and
maintaining development in Brassica under stress. High ion selectivity (K+ over Na+)
during upward transportation, as well as a low Na+/K+ ratio supported by the activation
of SOS, HKT, and PM-ATPase genes, were revealed to be critical for varied salt
tolerance in distinct members of the Brassicaceae family. Although a significant amount
of research has been done to understand the salinity-specific responses of different
members of the Brassicaceae family, the majority of it has focused on glycophytic
species, and there is still room for more research into salt tolerance strategies in
halophytic members of the family. Furthermore, more research into the involvement of
several phytohormones in imparting salt tolerance is required for an efficient stress
adaption approach in these crops.
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Fundamental Response and Salinity Tolerance in Brassicaceae Plants

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Fundamental Response and Salinity Tolerance in Brassicaceae Plants

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Fundamental response and salinity tolerance in Brassicaceae

Rapid osmotic effect

Later ionic effect

Fig 1. Flowchart illustrating the primary effects of salinity on plant systems

3. DISCUSSION IN GENERAL: THE BRASSICACEAE FAMILY'S REACTION

3.1 PERFORMANCE OF PLANT BIOMASS AND GROWTH IN THE

4.2.1. IN A SALINE ENVIRONMENT, THERE IS AN EFFECT ON

A considerable drop in photosynthesis is regarded as one of the most critical factors

4.2.2. STOMATAL REGULATION IN A SALINE ENVIRONMENT:

An intricate and extremely effective network of enzymatic and non-enzymatic antioxidants

3.4 RESPONSES IN BIOCHEMISTRY:

3.5 CHANGES IN IONIC BALANCE:

4. UNDER SALINITY STRESS, ORGANIC MOLECULES ARE DEPOSITED AS

5. THE ROLE OF PHYTOHORMONES IN SALINE-TOLERANT BRASSICA:

Fig 2: Summary of the functions of plant hormones in controlling a plant's ability to

6. MOLECULAR INTERACTION IN SALINE ENVIRONMENT:

Family Member of the High-Affinity K+ Transporter Gene helps to provide

Plasma Membrane-Related H+ Pumps:

Following a thorough examination of the literature on the salt tolerance of Brassicas, it

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