Vietnam Journal of Biotechnology 19(2): 197-212, 2021

REVIEW

IMPORTANT ANALYZING PARAMETERS IN THE ASSESSMENT OF SALT

TOLERANCE IN PLANTS

Hoang Thi Lan Xuan1,2,*, Nguyen Phuong Thao1,2

1
Applied Biotechnology for Crop Development Research Unit, School of Biotechnology, International
University, Thu Duc City, Ho Chi Minh City, Vietnam
2
Vietnam National University, Ho Chi Minh City, Vietnam
*
To whom correspondence should be addressed. E-mail: htlxuan@hcmiu.edu.vn

Received: 20.11.2020
Accepted: 09.04.2021

SUMMARY

Maximal crop performance potential and land area suitable for cultivation are usually restricted
by adverse environmental conditions. Among the abiotic factors, salinity stress is considered as one
of the main threats, which causes ionic toxicity, dehydration and oxidative stresses on the plants.
Alarmingly, the impact of salinity is predicted to be more severe in the forthcoming years due to
global warming. Therefore, development of new cultivars with better salinity resistance with
mimimized yield penalty under the adverse condition, either by breeding or genetic engineering
approach, has attracted a great attention from the scientists. In this review, important parameters used
in evaluation of plant resistance ability against salinity stress are discussed, which highlights the
necessity to obtain multi-sets of biological data ranging from analyses of morphological alterations
to physiological, biochemical and molecular responses, as well as by performing -omics studies to
find out network of salinity-responsive pathways. Literature review also demonstrates that the
relevance of salinity condition setup in terms of concentration and duration is required in experimental
design. Furthermore, recent investigations on genome duplication, activities of non-coding sequence
or epigenetics also reveal their regulatory roles in shaping plant response and tolerance degree toward
salinity stress. Collection of such data not only contributes to widen scientific understanding of plant
response mechanisms and adaptation to this stress factor but also facilitates the identification of
important genes associating with plant tolerance to salinity. Therefore, the presented information
could be used as a reference for the salinity stress-related studies serving for crop innovation and
transgene function characterization.

Keywords: analyzing parameters, gene function characterization, osmotic stress, plant resistance,
salinity stress

INTRODUCTION becoming “salinized”, which contains excessive

level of Na+ ions (Chen et al., 2014). Under
In addition to drought, salinity has emerged effects of climate change and global warming,
as another major abiotic threat that agriculture area of saline soil has been critically expanded
production is currently facing with. Presence in due to aggressive intrusion of the sea water onto
high concentration of either sodium bicarbonate mainland in recent years (Li et al., 2009).
NaHCO3 (i.e. alkaline soil) or sodium chloride Furthermore, human cultivation activities such
NaCl (i.e. saline soil) is the main cause for soil as inappropriate water management and fertilizer
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 Hoang Thi Lan Xuan & Nguyen Phuong Thao

usage have worsen the situation (Pessarakli, to plant cells. Osmotic and ionic stresses can then
Huber, 1991; Wanjogu et al., 2001). Excessive trigger the oxidative stress with increased
salts in soil cause negative effects on growth and production of cellular ROS contents (Gupta,
productivity of crop plants, which mainly belong Huang, 2014) (Figure 1). It is noted that apart
to the group glycophyte with the character of low from direct effects on plants, salinity stress also
resistance to salinity (Flowers 2004; Munns, disturbs the environment in the rhizosphere (i.e.
Tester, 2008). the soil vicinity surrounding the root system),
thus preventing the plants from establishment
Under this adverse condition, plants suffer
with beneficial microorganisms and uptaking
stunted growth, leaf chlorosis, reduced
nutrients (Kulkarn et al., 2000; Rao et al., 2002).
photosynthesis, cellular water loss, disrupted
cellular ionic homeostasis and cell damage due In this review, we summarize parameters that
to accumulation of reactive oxygen species the researchers can rely on in evaluating plant
(ROS) (Kumar et al., 2013; Golldack et al., resistance capacity to salinity, which will be
2014) (Figure 1). If the stress is prolonged or too discussed in connection with current
severe, plants even cannot maintain their understanding of plant response to this adverse
survival. Following the progress events of condition and progress in advancement of
salinity stress effects on plants, in the early stage, technologies and methodologies. The
plants experience with physiological drought information presented here could be used as a
(i.e. hyperosmotic stress) due to the difficulties reference in designing a relevant and sufficient
in absorbing water from the soil with high set of assessment criteria for comparative
concentrations of ions by the root systems. studies, which serve for the selection or
Following this, accumulation of cellular Na+ and development of salinity-tolerant cultivars and for
Cl- results in ionic toxicity (i.e. hyperionic stress) gene function characterization purpose.

Figure 1. Salinity effects to plant growth, development and productivity.

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ANALYZING PARAMETERS resistance associated with plant productivity,

ASSOCIATING WITH MORPHOLOGICAL, mean productivity, geometric mean productivity,
PHYSIOLOGICAL AND BIOCHEMICAL and stress tolerance index were more reliable
TRAITS parameters than others such as tolerance index,
yield index and yield stability index (read
Evaluate morphological characters and
original paper for information of each index
overall plant performance under salinity
calculation).
stress
As salinity stress inhibits early seed
Examining survival rate upon salinity stress
development, examination on germination rates
exposure is an indispensable experiment as its
as well as shoot- and/or root-associated traits
results will provide an overall evaluation on the
over a range of different salt concentrations is
salt tolerance capacity of a plant genotype before
usually conducted (Table 1). For Arabidopsis,
its tolerance mechanisms are investigated (Table
the sterilized seeds are placed on half strength
1). The assay can be divided into three main
Murashige and Skoog (1/2 MS) medium
stages, which are (i) growing plants under
(Murashige, Skoog, 1962) containing NaCl 100
normal condition, (ii) stress application, and (iii)
mM under relevant growing condition and the
recovery and survival rate calculation. For
rate of seeds with radicle emergence (i.e.
example, two-week-old Arabidopsis can be
successful germination) can be monitored every
treated with 200 mM NaCl solution daily over a
24 hours within five consecutive days since
three-week-duration, followed by three-day
seeding (Li et al., 2019). Germination test using
normal irrigation with water prior to calculating
higher salts (e.g. 250 mM) and MS medium has
the proportion of alive plants (Li et al., 2019). It
also been reported (Lee et al., 2006). In addition,
is noted that the duration and procedure of the
this test is commonly conducted over a range of
stress treatment can be modified depending on
different NaCl concentrations. For example,
plant species, plant age, plant density per
sterilized soybean (Glycine max) seeds can be
container, salt concentration and volume, as well
placed on the medium containing 0, 100, 200 and
as frequency of the salt solution application
300 mM NaCl (Li et al., 2017).
(Table 2). In study of Li et al. (2018), four-week-
old Arabidopsis seedlings were used for This in vitro assay system (100-125 mM
application of 300 mM NaCl solution every three NaCl) can also be used to assess root length and
days until the stress effects could be visualized. fresh weight of Arabidopsis seedlings that has
Meanwhile, salt treatment for rice (Oryza sativa) been grown on medium with salt
can be set up by exposing the seedlings to ten- supplementation for a week (Qin et al., 2017; Li
day-stress duration using 150 mM NaCl solution et al., 2019). For studies in tobacco (Nicotiana
(Zhu et al., 2015). Generally, dicot plants have a tabacum), NaCl solution with concentrations of
greater variation in salinity tolerance than the 100-200 mM has been applied for germination
monocot plants (Munns, Tester, 2008). and growth assays (Kobayashi et al., 2008; Yang
Furthermore, this method can be used to analyze et al., 2017). With bigger plants like soybean,
the salt effects on plant productivity by growing plants in hydroponic system using half-
investigating the reproduction-related traits strength Hoagland solution can make it easier for
rather than survival rate (Table 1). The salt treatment, simply by adding the desired salt
agronomic traits are very important in amount into the nutrition solution and immersing
agricultural and economic perspectives in the root part into this liquid (Li et al., 2017)
selecting elite cultivars not only with enhanced (Table 1). In addition, certain measurements can
stress tolerance but also with high productivity be categorized for specifically ranking the plant
(Liang et al., 2016). Singh and others (2015) tolerance capacity. For example, depending on
have demonstrated that among the tolerance the visualized damage and necrotic degree that
indices that can be used to assess the salt the studied plants can be placed in a five-point
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scale, with level 1 for no sign of necrosis and the dry weight recorded at different time points,
level 5 with the highest score of injured areas can be used to divide plants into groups of salt-
(75-100%) (Sabra et al., 2012). Similarly, degree tolerant, moderately salt-tolerant, moderately
of reduction in relative growth rate under salinity salt-sensitive and salt-sensitive species
stress condition, of which calculation is based on (Cassaniti et al., 2012).

Table 1. Common parameters used for analyses of morphological, physiological and biochemical traits to
evaluate plant resistance capacity to salinity stress.

Assay name Analyzing parameters References

Survival rate assay post-stress survival rate Li et al., 2018; Li et al.,
2019
Germination assay germination rate Li et al., 2017
Vegetative growth assay root length, shoot length, fresh and Wang et al., 2016; Li et al.,
dry biomass 2017
Analysis of reproductive traits flowering time, number of Liang et al., 2016; Wang et
flowers/pods/fruits/fruit branches, al., 2017
yield per plant
Analysis of accumulated reactive superoxide anion and hydrogen Wang et al., 2017
oxygen species peroxide contents
Analysis of membrane damage ion leakage and malondialdehyde Orellana et al., 2010;
content Wang et al., 2016
Analysis of antioxidant enzyme superoxide dismutase, peroxidase, Li et al., 2017; Wang et al.,
activities catalase and glutathione transferase 2017
activities
Measurement of hormonal contents abscisic acid and jasmonic acid Yang et al., 2001
Measurement of osmolyte contents Proline, trehalose and soluble sugar Wang et al., 2017
contents
Measurement of intracellular ion Na+, K+ and Cl- contents Xu et al., 2016; Li et al.,
contents 2017
Evaluation of photosynthetic Chlorophyll content, stomata aperture Orellana et al., 2010;
performance and density Liang et al., 2016; Wang et
al., 2017

Table 2. Concentrations of sodium chloride that have been applied to different plant species in salinity stress-
related studies.

Plant Applied NaCl Studied

Duration Studied parameters References
species concentration system
Survival rate, chlorophyll
2 weeks
Soil and content
irrigation Cao et al.,
250 mM
2017
4, 10, 12, 14
Fv/Fm values
Arabidopsis and 16 days
thaliana 100 and 150 7 days Half-strength Root length
mM MS
He et al.,
12 days (4-d Soil and Survival rate, fresh 2019
200 mM
intervals) irrigation weights, Fv/Fm, MDA,
proline and H2O2 contents,

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antioxidant enzyme
activities

50, 100 and 5 days Half-strength

Seed germination
150 mM MS
Photosynthesis, relative
Half-strength water content, MDA and
250 mM 7 days
Hoagland proline contents,
peroxidase activity
Boehmeria An et al.,
nivea (ramie) Total plant fresh and dry 2015
300 mM 12 days Soil and
weights
irrigation
Fresh and/or dry weights of
350 mM 11 days shoot/root/bast;
transpiration
200, 250 and Half-strength
7 days Germination rate
300 mM MS medium

300 mM 48 hours Hydroponic Root characters

Glycine max Zhang et al.,
(soybean) 200 and 300 2 weeks (3 2013
Soil and Growth characters
mM times/week)
irrigation
9 days (3
300 mM Proline and sugar contents
times/week)
Hoagland
7 days Fresh and dry weights
solution
250 mM
Gossypium 2 weeks Proline and MDA contents
Liu et al.,
hirsutum
Soil and 2014
(cotton)
20 days (5- irrigation Photosynthesis, stomatal
100 and 250
or 10d- conductance and
mM
intervals) transpiration

Leaf disc assay for

100, 200 and
Musa 6 days Salt solution chlorophyll content
250 mM Tak et al.,
acuminata determination
2017
(banana) Soil and MDA, Fv/Fm, proline
250 mM 15 days
irrigation contents

Nicotiana
Half-strength Li et al.,
tabacum 100 mM 30 days Root lengths and weights
MS medium 2018
(tobacco)

Hoagland
Na+ content
solution Hong et al.,
150 mM 6 days
Half-strength Shoot height, fresh weight, 2016
Oryza sativa MS medium number of later roots
(Rice) Soil and
200 mM 12 days Survival rate
irrigation Hu et al.,
2006
100 mM 5 days Hydroponic Fresh weight

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Shoot and root growth

10 days MS medium
measurements
Solanum 100 mM Leaf disc assay for
Zhu et al.,
lycopersicum 4 days Salt solution chlorophyll content
2014
(tomato) determination
21 days (72- Soil and
400 mM Growth characters
hour interval) irrigation
Triticum Hoagland
aestivum solution Survival rate, fresh and dry Saad et al.,
2% 7 days
(bread weights 2013
wheat)

Evaluate physiological and biochemical traits Qin et al., 2017; Yang et al., 2017). In other
of plants under salinity stress papers, measurement of Na+ and K+ contents is
addressed as the plant growth is negatively
It has been known that oxidative stress is the
affected by the high level of Na+ in the cytosol
secondary stress induced by osmotic and ionic
but supported by the presence of K+ (Munns et
disturbance, with increased production of ROS
al., 2006; Chen et al., 2014; Xu et al., 2016; Li
contents (Gupta, Huang, 2014). Accumulation of
et al., 2017) (Table 1). Therefore, under salinity
species such as superoxide (O2•−) and hydrogen
stress conditions, maintaining a low cytosolic
peroxide (H2O2) in plants can cause damage of
Na+/K+ ratio is important for normal metabolic
cellular membrane and molecules, as well as
activities to take place (Munns et al., 2006; Chen
interruption of metabolic activities (Gill, Tuteja,
et al., 2014). In certain species that are able to
2010). Knowing accumulation degree of these
effectively prevent the Na+ accumulation on
ROS can be used as indicators for the estimation
leaves, measurement of Cl- should be conducted
of cellular oxidative stress level. Detection of
as its concentration might be enhanced to a toxic
ROS in the leaf tissues can be achieved by
dose level along with potassium ions (Munns,
staining methods using nitro blue tetrazolium
Tester, 2008).
(NBT) for O2•− (Shi et al., 2010) and 3,3’-
diaminobenzidine (DAB) for H2O2 (Liu et al., Plants use both enzymatic and non-
2014). Although total ROS production in intact enzymatic pathways to protect themselves from
cells can be visualized by staining with 2,7- oxidative stress effects, mainly by scavenging
dichlorofluorescin diacetate (H2DCF-DA) ROS or by using molecules functioning as
(Zhang et al., 2011; Yang et al., 2017), this antioxidants. Regards to the enzyme-mediated
method has been claimed not to be accurate due defense, superoxide dismutase (SOD) plays in
to non-specificity in substrate binding of the the first line by converting superoxide into H2O2.
chemical reagent (Jakubowski, Bartosz, 2000; The generated H2O2 will be further detoxified by
Chen et al., 2010). Apart from these peroxidase (POD) and catalase (CAT) enzymes.
histochemical staining assays, H2O2 content can In the non-enzymatic defense pathway, certain
be quantified by spectrophotometric approach molecules such as proline, soluble sugars (e.g.
(Patterson et al., 1984) (Table 1). In addition, trehalose, glucose and fructose) and glycine
oxidative stress-induced damage of cellular betaine will play a role in antioxidative
membrane can be estimated based on the protection (Ashraf, Foolad, 2007; Gupta, Huang,
measurement of electrolyte leakage or 2014; Qin et al., 2017; Wang et al., 2017). The
malondialdehyde (MDA) contents (Campos et main functions of these compounds are to
al., 2003; Li et al., 2015) (Table 1). In some enhance water retention capacity by lowering
studies, examination on cell death by Evans blue cellular water potential under osmotic stress as
staining is also conducted (Zhang et al., 2011; well as stabilize cellular environment to maintain

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metabolic activities (Ashraf, Foolad, 2007). analyses. In addition, gene expression data
Therefore, analyzing enzymatic activities or would provide complementary evidence for
contents of these antioxidant/osmoprotectant supporting the phenotypic, physiological or
molecules would provide important information biochemical results, making the conclusion more
on plant defense capacity to salinity (Li et al., reliable. For example, transgenic Arabidopsis
2018; Li et al., 2019) (Table 1). ectopically expressing sweet potato (Ipomoea
batatas) IbRAP2-12 acquired better salt
As salinity stress also causes adverse effects
tolerance, with higher proline content and in
on photosynthetic molecules and performance,
consistency with higher expression of pyrroline-
measurement of chlorophyll content is usually
5-carboxylate synthase 2 (P5CS2) (Li et al.,
included in the study. To do this, in small plants
2019). This gene encodes the key enzyme in
like Arabidopsis, aerial part of different plants
biosynthesis of proline, a molecule functioning
can be pooled together for being used as a
as an osmolyte used for osmotic adjustment and
biological replicate (Li et al., 2018) and in bigger
as an antioxidant in protecting bio-
plants, individual leaf samples can be analysed
macromolecules and scavenging ROS (Ashraf,
separately (Liang et al., 2016). In addition,
Foolad, 2007; Li et al., 2019). In another
investigation of stomata-related traits such as
example, increased trehalose content coupled
aperture size and density also reveal useful
with up-regulation of genes ThTPS1-3 and
information for evaluation of photosynthetic
ThTPPA encoding the key enzymes 3 trehalose-
activity potential (Orellana et al., 2010; Liang et
6-phosphate synthase (TPS) and 1 trehalose-6-
al., 2016; Wang et al., 2017).
phosphate phosphatase (TPP), respectively, in
Hormone-mediated plant response to salinity the biosynthetic pathway of trehalose is observed
stress, including abscisic acid (ABA) and in the transgenic Tamarix hispida
jasmonate acid (JA), has also been well overexpressing cytokinin response factor 1
documented (Tuteja, 2007; Zhang et al., 2017). (CRF1) (Qin et al., 2017).
A number of transcription factors regulating
Table 3 presents important pathways and
plant response to salinity has been found to work
functional groups whose gene expression could
in ABA-dependent manner (e.g. tomato
be regulated in mediating plant response to
(Solanum lycopersicum) JERF1), or in both
salinity stress. In general, expression of genes
pathways (e.g. Arabidopsis ERF1 and AtMYC2)
encoding the enzymes working in the
(Cheng et al., 2013; Zhao et al., 2014).
biosynthesis of hormones (e.g. ABA and JA),
Therefore, quantification of ABA and JA
osmoprotectant (e.g. proline, trehalose), as well
contents by enzyme-linked immunosorbent
as in ROS removal (e.g. CATs and PODs) is
assays (ELISAs) can be considered (Yang et al.,
induced upon salinity stress challenging. For
2001) (Table 1).
example, increase in expression of Arabidopsis
TARGET GENES FOR EXPRESSION dehydroascorbate reductase 1-encoding gene
ANALYSIS BY QUANTITATIVE REVERSE (DHAR1) under this adverse condition, was
TRANSCRIPTION PCR (RT-qPCR) METHOD reported (Li et al., 2019). DHAR1 is an enzyme
belongs to glutathione S-transferase superfamily
Over the last decade, RT-qPCR has become and responsible for the regeneration of
a more widely used method than RNA gel ascorbate, an antioxidant molecule (Ding et al.,
blotting in detecting differential gene expression 2020). Therefore, activity of this enzyme also
between conditions (e.g. stressed versus normal plays an important role in plant defense.
conditions) or genotypes, from which important Dehydrin proteins such as late embryogenesis
gene activities in connection with salt tolerance abundant (LEA) proteins, responsive-to-ABA
capacity can be identified. RT-qPCR is also (RAB) proteins and cold-regulated (COR)
employed to validate the transcriptomic proteins are well-known members functioning in

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cellular protein protection and membrane reside on the plasma membrane and tonoplast
stabilization under osmotic stress (Verslues et (vacuole) membrane, respectively, and
al., 2006; Jia et al., 2014; Shinde et al., 2019). responsible for the prevention of intracellular
Therefore, expression study of their Na+ accumulation, either by Na+ exclusion or
corresponding encoding genes is an interest. It is compartmentalization (Apse et al., 2003; Shi et
found out that the transgenic tomato (Solanum al., 2000).
lycopersicum) overexpressing SlAREB1 had
For transgenic studies using transcription
increased expression in two dehydrin encoding
factor-encoding genes as the transgenes,
genes TAS14 and LE25, suggesting their
analyzing the cis-motifs present in the promoter
contribution to the enhanced tolerance of the
region of target genes could help compiling the
transgenic tomato under salinity stress (Orellana
list of potential genes whose expression should
et al., 2010).
be prioritized for investigation. Of course, it is
As saline conditions cause ionic and osmotic possible that genes without the cis-acting
imbalance, it is important to study the expression elements for the transcription factor binding are
levels of genes encoding transporter proteins in also its downstream target genes, as an outcome
the root tissue. Particularly, attention should be of indirect regulation/interaction. Salinity stress-
paid to genes coding for Na+ transporters [e.g. related studies have identified participation of
salt overly sensitive 1 (SOS1), cation/H+ various transcription factors that belong to
exchanger (CHX1) and Na+/H+ antiporters 1 different families, such as dehydration-
(NHX1), and K+ transporters [e.g. CHX1 and responsive-element (DRE)-binding proteins
high-affinity potassium transporter 1;4 (DREBs), zinc finger proteins (ZFPs), ethylene
(HKT1;4)], as well as water channels (known as response factor proteins (ERFs) and
“aquaporin”) [e.g. plasma membrane intrinsic myeloblastosis proteins (MYBs) (Xu et al.,
protein 1;6 (GmPIP1;6)]. SOS1 is a well-known 2016; Wang et al., 2017). As drought and salinity
Na+/H+ antiporter working in the SOS-signaling stresses cause osmotic stress, similar strategies
pathway for regulating cellular Na+ efflux and components are used by plants in response to
(Cellier et al., 2004; Chen et al., 2014; Gupta, drought and high salinity conditions (Ashraf,
Huang, 2014; Qi et al., 2014; Zhou et al., 2014; Foolad, 2007; Gill, Tuteja, 2010; Golldack et al.,
Li et al., 2017). SOS1 and NHX1 are known to 2014; Li et al., 2017).

Table 3. Main pathways that might be under regulation in mediating plant response to salinity stress, based on
studies of transgenic plants with improved salinity tolerance and analyzed by RT-qPCR method. Examples for
genes with altered expression in each pathway are included.

Function Transgenic Transgene Responsive genes References

plants
Abscisic acid Arabidopsis Ipomoea batatas abscisic aldehyde oxidase 3 (AAO3) Li et al.,
biosynthesis- IbRAP2-12 2019
related Gossypium Zea mays ABP9 nine-cis-epoxycarotenoid Wang et al.,
pathway hirsutum dioxygensase 2 (GhNCED2) 2017
Jasmonic lipoxygenase 2 (LOX2)
acid Arabidopsis I. batatas IbRAP2- allene oxide synthase Li et al.,
biosynthesis- 12 (AOS) 2019
related
pathway
Proline Arabidopsis Tamarix hispida pyrroline-5-carboxylate synthase 1 Qin et al.,
biosynthesis- ThCRF1 (P5CS1) 2017
related
pathway

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Trehalose T. hispida T. hispida ThCRF1 3 trehalose-6-phosphate synthase Qin et al.,

biosynthesis- (ThTPS1-3); 1 trehalose-6- 2017
related phosphate phosphatase (ThTPPA)
pathway
Arabidopsis I. batatas IbRAP2- glutathione peroxidase 7 (GPX7), Li et al.,
12 ascorbate peroxidase 1 (APX1), 2019
Reactive dehydroascorbate reductase 1
oxygen (DHAR1), catalase 1 (CAT1)
species T. hispida T. hispida ThCRF1 superoxide dismutase-encoding Qin et al.,
removal genes (ThSOD1, ThSOD2, 2017
ThSOD3)
G. hirsutum Z. mays ABP9 Superoxide dismutase (GhSOD), Wang et al.,
peroxidase (GhPOD), catalase 2017
(GhCAT), Glutathione-S-transferase
(GhGST)
Transporter Glycine max G. max GmFDL19 Cation/H+ exchanger (GmCHX1), Li et al.,
proteins plasma membrane intrinsic protein 2017
1;6 (GmPIP1;6); Na+/H+ antiporters 1
(GmNHX1), GmHKT1;4; salt overly
sensitive (GmSOS1)
Dehydrin Solanum S. lycopersicum TAS14, LE25 Orellana et
proteins (e.g. lycopersicum al., 2010
LEA, RAB
and COR
subfamilies)
G. max G. max GmFDL19 GmbZIP1, GmNAC11, Li et al.,
GmNAC29, GmDERB2A;2, 2017
Transcription GmWRKY27, GmERF5,
factors GmMYB174
G. hirsutum Z. mays ABP9 dehydration-responsive-element Wang et al.,
(DRE)-binding protein 2 (GhDBP2), 2017
zinc finger protein 1 (GhZFP1),
ethylene response factor 1
(GhERF1)

RECENT ANALYTIC APPROACHES Other studies have indicated that the salt
CONTRIBUTING TO COMPREHENSIVE tolerance of plants can be also affected by
UNDERSTANDING OF PLANT epigenetic changes such as post-translational
TOLERANCE TO SALINITY modification via activity of ubiquitin ligases or
With rapid progress in developing novel degree of DNA methylation (Park et al., 2010;
technologies and advanced instruments, in Feng et al., 2012) (Table 4). With the reduction
addition to transgenic/mutant-based systems, in cost and time as well as improved instrument
other approaches can be utilized to versality, analysis at systemic scale can provide
comprehensively understand the plant tolerance global information for the salinity stress-induced
to salinity. It has been shown that microRNA changes in transcript profile, protein profile or
(miRNA) molecules also play a role in metabolite profile (Hernández, 2019). It must be
determining the plant tolerance capacity (Genie emphasized that genome-wide studies remain as
et al., 2019) (Table 4). For example, increasing an important approach as the tolerance capacity
transcript abundance of certain miRNAs could among different cultivars can be compared and
make the transgenic rice more vulnerable to assessed based on genetic variants or polyploidy
salinity stress (Gao et al., 2010; Gao et al., 2011). status (Tu et al., 2014; Ganie et al., 2019).
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Furthermore, using molecular markers also (Ky et al., 2018; Lang et al., 2019; Le et al.,
contributes to the identification of important 2021) (Table 4). Previously, an important QTL
salinity-related genes and establishment of QTL for salinity tolerance in rice, known as Saltol, is
(quantitative trait locus) mapping of these genes reported (Vu et al., 2012).

Table 4. Other data for comprehensive understanding on plant tolerance capacity toward salinity.

Targets Examples Added information value References

Non-protein- miRNA Novel mechanism of plant Gao et al., 2010; Gao et
coding genes responses to salinity al., 2011
Epigenetics Ubiquitination genes, Affecting protein stability and Park et al., 2010; Feng et
DNA methylation expression degree al., 2012
Genome Genome-wide To examine polyploidy status in Tu et al., 2014
duplication analysis association with salt tolerance
capacity
-Omic studies Transcriptomic To obtain global salt responsive- Ganie et al., 2019;
profiling, proteomic network and identify important Hernández, 2019
profiling, metabolic participants
profiling
DNA markers Simple sequence Locate important salinity-related Ky et al., 2018; Lang et
repeats, expressed genes and quantitative trait loci al., 2019; Le et al., 2021
sequence tag (QTL mapping)
markers, SNPs

SALINITY STRESS TOLERANCE STUDIES this breeding process (Lang et al., 2019).
IN VIETNAM Similarly, marker-assisted backcrossing
(MABC) is also adopted to speed up the
In Vietnam, salinity has not been considered
development of salt-tolerant rice varieties in
a major threat to agricultural production until
comparison with the traditional backcrossing
recent years, when a higher rate of seawater
method (Vu et al., 2012). Following this,
intrusion to the coastal region and river has been
introgression lines with improved salt tolerance
observed. Particularly, the rise in sea level due to
were generated by introduction of Saltol QTL
climate change makes the agricultural production
into BT7, a rice variety carrying certain desired
in Mekong River Delta become vulnerable more
agronomic traits, by crossing this with a salt-
than ever. To cope with this, various measures
tolerant donor variety, FL478-Saltol (Linh et al.,
have been suggested or deployed, including
2012).
infrastructural establishment to prevent the
invasion of seawater into the mainland, changes Other studies focus on evaluating the salinity
in agronomic practices and cropping pattern tolerance of different rice cultivars (Ky et al.,
(Dam et al., 2021). Regarding development of 2018; Lang et al., 2019), including the mutant
elite salinity-tolerant culivars, so far this has rice lines (Huong et al., 2020). For example,
been an interest for rice only. This is easily twelve different rice varieties in Tra Vinh have
understood as Vietnam is one of the main global been analyzed for their salinity tolerance
rice suppliers and its economy heavily depends capacity to NaCl 6‰ based on three SSR
on the rice productivity. In fact, research on (Simple sequence repeat) markers (RM336,
improvement of rice tolerance to salinity has RM10793 and RM10825) and ratio of K+/Na+
been conducted many years ago using uptake (Ky et al., 2018). Phenotype-based
conventional breeding and the application of parameters, including plant height, root length,
marker-assisted selection (MAS) has accelerated survival rate and biomass, have been also
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Vietnam Journal of Biotechnology 19(2): 197-212, 2021

employed to screen for the rice germplasms with CONCLUSION

higher salt tolerance (Lang et al., 2019).
This review demonstrated complex
Meanwhile, another study unraveled the salinity
responses that the plants employ to cope with
tolerance of different rice varieties based on
salinity stress. This also means that there are
yield-related properties including productiviy,
various potential target pathways or genes for a
amylose and protein contents (Quan, Vo, 2017).
researcher to manipulate in developing crop
Recently, a transcriptomic analysis has been
varieties with improved salinity stress resistance.
conducted for two rice varieties with contrasting
Clearly, in-depth understanding of mechanisms
salinity tolerance, in order to identify pathways
and performance of plants under salinity stress
and genes associating with the plant tolerance
conditions requires a combined data set, which is
(Ky et al., 2021).
derived from phenotypic, physiological,
In other plant species, the gained biochemical and molecular analyses. In terms of
information and outcomes remain limited as economic and agricultural perspectives,
only a handful studies have been conducted in productivity ability of a studied genotype should
relation to salinity stress in Vietnam. Among of be examined along with its salinity stress
these studies, it has been demonstrated that resistance potential. In the future, investigation
exogenous application of salicylic acid and/or on how salt-resistant plants (i.e. halophytes) can
calcium can enhance the salinity tolerance of withstand high salinity conditions might
amaranth (Amaranthus tricolor) via promotion beneficially provide new strategies for
of Na+ exclusion from roots, accumulation of development of crop cultivars with better salt
phenolic and flavonoid compounds as well as resistance.
increased antioxidant activities (Hoang et al.,
2020). Tolerance of various chili pepper Acknowledgements: This research is funded by
genotypes against different concentrations of Vietnam National University Ho Chi Minh City
NaCl or CaCl2 (ranging from 0-300 mM) has (VNU-HCM) under grant number C2018-28-04.
been also explored, which was based on
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CÁC CHỈ SỐ PHÂN TÍCH QUAN TRỌNG DÙNG TRONG NGHIÊN CỨU ĐÁNH
GIÁ KHẢ NĂNG CHỊU MẶN Ở THỰC VẬT

Hoàng Thị Lan Xuân1,2, Nguyễn Phương Thảo1,2

1
Phòng thí nghiệm Ứng dụng Công nghệ Sinh học trong Phát triển giống cây trồng, Khoa Công nghệ
Sinh học, Trường Đại học Quốc tế, Thành phố Thủ Đức, Thành phố Hồ Chí Minh, Việt Nam
2
Đại học Quốc gia, Thành phố Hồ Chí Minh, Việt Nam

TÓM TẮT

Tiềm năng sản lượng tối đa của cây trồng và diện tích đất phù hợp cho trồng trọt thường bị hạn
chế bởi các yếu tố bất lợi từ môi trường. Trong số các nhân tố stress phi sinh học, stress mặn là một
trong những mối đe dọa chính, gây ra độc ion nội bào, stress mất nước và stress ôxy hóa. Tác động

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 Hoang Thi Lan Xuan & Nguyen Phuong Thao

của stress mặn được dự báo là ngày càng nghiêm trọng hơn do biến đổi khí hậu. Vì vậy, phát triển
các giống cây trồng mới có khả năng chịu mặn tốt hơn bằng phương pháp lai tạo truyền thống hay
bằng kỹ thuật di truyền luôn là mối quan tâm của các nhà khoa học. Trong bài viết này, chúng tôi
thảo luận những chỉ số quan trọng dùng trong việc đánh giá về khả năng chịu mặn của cây để thu thập
bộ dữ liệu đầy đủ liên quan đến thay đổi hình thái và điều chỉnh sinh lý, sinh hóa và phân tử; hoặc từ
các phân tích ở quy mô -omics để có cái nhìn tổng quan về mạng lưới các con đường tham gia đáp
ứng mặn. Các nghiên cứu cũng cho thấy rằng việc thiết lập điều kiện stress mặn phù hợp về mặt nồng
độ và thời gian là rất cần thiết trong thí nghiệm. Hơn nữa, các nghiên cứu gần đây cũng chứng minh
rằng số lượng gen trong genome, hoạt động từ các phân tử không mã hóa protein và điều hòa ngoài
gen cũng ảnh hưởng đến khả năng chống chịu của cây. Tập hợp các thông tin này không chỉ mở rộng
mức độ hiểu biết khoa học về các cơ chế đáp ứng thích nghi của thực vật mà còn giúp tìm ra các gen
quan trọng trong đáp ứng stress mặn. Do đó, bài viết này có thể dùng để tham khảo trong các nghiên
cứu về stress mặn phục vụ công tác cải tạo giống và phân tích chức năng gen.

Từ khóa: các chỉ số phân tích, khả năng chống chịu stress của thực vật, phân tích chức năng gen,
stress mặn, stress thẩm thấu

212