Asian Journal of Agricultural Sciences 2(3): 99-105, 2010
ISSN: 2041-3890
© M axwell Scientific Organization, 2010
Submitted Date: April 12, 2010
Accepted Date: April 27, 2010
Published Date: July 05, 2010
Biological and Physiological Perspectives of Specificity in Abiotic Salt Stress
Response from Various Rice Plants
Baby Joseph, D. Jini and S. Sujatha
International Centre for Bioresources Managem ent, M alankara Cath olic C ollege,
Mariagiri, Kaliakkavilai - 629153 , Kanyakumari district, T amilNadu, In dia
Abstract: Abiotic pressures like salt stress and chemical insultance can impose limitations on crop p roductivity
and also limit land available for farming, often in regions that can ill afford such constraints, thus highlighting
a greater need for understanding how plants respond to adverse conditions with the hope of improving tolerance
of plan ts to environmental stress. More is becoming known about the physiological and molecular effects of
environmental stress. Salt-tolerant transgenic rice plants have been produced using a host of different genes and
transcript profiling by micro- and ma croarray-based methods has opened the gates for the disco very of novel
salt stress m echa nisms in rice, and co mpa rative genomics is turning out to be a critical input in this respect.
Despite, Rice is mod erately sensitive to salt in the field as alm ost all the other crop sp ecies. A lthoug h rice is
considered as a sensitive crop to salinity, it is one o f the most w idely grown crop s in coastal areas frequently
inundated with saline seawater during high tidal period. In this review, highlighted the effects of salinity on the
morphological, physiological, biochemical and genetic characters in mono cotyledon plant of rice were
discussed along with recent dev elopm ent in sa lt stress research . Since rice is moderately resistant to salinity
and great degree of genotypic variation is found, hen ce, it is possible to deve lop va rieties w ith enhance d salt
tolerance if appropriate strateg ies and adva nced techniques are adopted .
Key w ords: Abiotic-stress, Oryza sativa, salinity, salt tolerance
INTRODUCTION
Environmental stresses such as salinity and drought
reduce the growth and agricultural productivity more than
other factors (Karakas et al., 1997 ). Because of the
increases in global pop ulation, world agriculture must
produce a greater yield per unit area than ever before.
How ever, worldwide one-half of all irrigated lands are
seriously affected by salinity or water logging. Currently,
more land is not being irrigated due to salinity problems
than
there
is new land coming under irrigation
(Wilson et al., 2005). Higher salinity levels caused
significant reduction in grow th parame ters like leaf area,
leaf
length and root and shoot dry weight
(Ashrafuzzaman et al., 2002).
Rice (Oryza sativa L.) is one of the most important
stable food crops in the world. In Asia, more than two
billion people are getting 60-70% of their energy
requirement from rice and its derived products. In India,
rice occupies an area of 44 million hectare with an
average production of 90 million tone s with productivity
of 2.0 tonnes/ha. Demand for rice is growing every year
and it is estimated that in 2010 and 2025 AD the
requirement would be 100 and 140 million tones
respectively. To sustain present food self-sufficiency and
to meet future food requirements, India has to increase its
rice productivity by 3% per annum (Thiyagarajan and
Selva raju, 2001).
Rice (Oryza sativa L.) has been cultivated as a major
crop for 11,500 years, and it currently sustains nearly onehalf of the world population (W u et al., 2004 ), and its
native is in tropical and subtropical Southeastern Asia and
Africa. Rice is the principal source of food for more than
one third of the world’s population.
Salinity is the most serious threats to agriculture and
for more important globally (Sahi, 2006) water consisted
minerals and salt materials are m ajor harmful factors in
arid and semi-arid region of worldwide. Salt stress causes
the reduction of rice yield, and some times-severe salt
stress may even threaten surv ival. Rice is moderately
sensitive to salt in the field as almost all the other crop
species. Although rice is considered as a sensitive crop to
salinity, it is one of the most w idely grown crop s in
coastal areas frequently inundated with saline seawater
during high tidal period (A kbar et al., 1972 ; Mori and
Kinoshita, 1987).
Background: As sessile organisms, plants cannot
physically move away from environmental stresses that
can negatively affect growth. Therefore, plants have had
to evolve strategies to cope with abiotic stress. Indeed, the
ability to respond and ultimately adapt to abiotic stress
Corresponding Author: S. Sujatha, International Centre for Bioresources Management, Malankara Catholic College, Mariagiri,
Kaliakkavilai - 629153, Kanyakumari district, TamilNadu, India
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Asian J. Agr. Sci., 2(3): 99-105, 2010
may be a driving force in speciation (Lexer and Fay,
2005; Aleel and Grennan, 20 06). Plants m ust be able to
“sense” the enviro nme ntal cues before be ing ab le to
respond appropriately to the abiotic stress. Due to the
complex nature of stress, multiple sensors, rather than a
single senso r, are more likely to be responsible for
perception of the stress. After the initial recognition of the
stress, a signal transduction cascade is invoked.
Secon dary messengers relay the signal, ultimately
activating stress-responsive genes generating the initial
stress response. Stress-induced gene products can be
divided into two m ajor groups: those involved in stress
tolerance and those involved in sign al transduction.
Stress-tolerance genes enable plants to cope with the
stress situation, in terms of both sh ort- and long-term
responses. These can include the synthesis of chaperones
and enzy mes for osm oly te biosynthesis and
detoxification, to a change in the composition of
membrane lipids as is found with cold stress. Gene
products can also act as transcription regulators
controlling sets of stress-specific genes or be involv ed in
the production of regulatory molecules, such as the plant
hormone ABA.
Salt tolerance in rice plant: Rice is moderately sensitive
to salinity. Salinity affects virtually all aspects of rice
grow th in varying degree at all stages from germination
through maturity. Tolerance to salinity is genetically and
physiologically complicated and inherited quantitatively.
Lee et al. (2003 ) tested 10 varieties of rice in order to
identify the degree o f salinity tolerance of the indica and
japonica rice groups, under saline and non-saline
conditions. There exists tremendous variation for salt
tolerance within species in rice, provid ing op portunities to
improve crop salt-stress tolerance through genetic means.
Some attemp ts to dev elop salt-tolerant genotype s were
based on highly tolerant traditional rice cultivars i.e.
Pokkali and N ona-B okra (Akbar et al., 1985 ; Greg orio
and Senadhira, 199 3).
M any Available reports show ed that salt tolerance
classification in rice lines (Oryza sativa L.) was based on
bioch emical, physiological characteristic and molecular
markers as well as the relationship be tween bo th
performances
for
rapid
screening
techniques
(Zeng et al., 2004). Ten advanced rice lines were screened
for salinity tolerance at seedling stage u sing a rapid
screening techn ique. Ou t of the lines tested, five were
found tolerant, four were graded as mo derately
tolerant and o ne as susce ptible (A li et al., 2004).
Theerakulpisut et al. (2005) studied on three local Thai
cultivars, Pokkali and IR29 which found that the shoot
and root dry weight and net photosynthesis rate showed
high positive correlation w ith level of salinity tolerance.
Krishnamurthy (1991) investigated amelioration of NaCl
toxicity in the salt tolerant rice cultivar Co 43 on 25th and
45th day old plants by foliar application of diamine
putrescine (10 :M) in a pot culture e xperimen t.
Generally, the salt tolerant varieties of rice maintain
low concentration of Na + in their leaves than those of salt
sensitive lines, when exposed to salt stress (Lutts and
Guerrir, 1995; Lutts et al., 1995), therefore Na+ in the
leaves of crop plants can be used as an important indicator
of salinity tolerance; and breeding for low ion
accumulation could be a sim ple w ay to im prove salt
tolerance. Selection w ithin varieties or lines with low Na+
transport has been made in rice (Yeo et al., 1988 ).
Reduction of rice yield by salinity: Growth and y ield
reduction of crops is a serious issue in salinity prone areas
of the world (Ashraf et al., 1994; Ashraf, 2009). Among
the various factors limiting rice yield, salinity is one of the
oldest and most serious environmental problems in the
world (McW illiam, 1986). In Pakistan one million
hectares of the rice growing area is salt-affected
(Qureshi et al., 1991) and in case of redu ctions for salinity
accounts abou t 64% in crop yield on these soils
recognized by A fzal et al. (2005). In Bangladesh, over
thirty percent of the net cultivable area is in the coastal
region. Out of 2.85 million hectares of the coastal and offshore areas, about 0.833 million hec tares are arable lands,
which constitute about 52.8% of the net cultivable area in
13 districts (Karim et al., 1990 ). Since , the rice is
recognized as a salt-sensitive crop he nce there is a serious
concern that plant stand (i.e., seedling survival) and the
development of yield components are affected by water
salinity.
Rice is one of the most widely grown crops in coastal
areas inundated with seawater during high tidal period,
although it is usually considered mod erately susce ptible
to
salinity (Akbar
et al., 1972; Korbe and
Abdel-A al, 1974; Mori and Kinoshita, 1987). At the onset
of the growing season, salts accumulated by capillary rise
in the topsoil are released into the soil solution and
floodw ater. Rice fields often lack drainage facilities, or
drain from one field to the other, thus building up salt
levels during the season. Salt stress may, therefore, occur
throughout the gro wing season and m ay co incide with
susceptible growth stages of the rice crop (Asch and
W opereis, 2001).
Need for salt tolerant cultivar: Rice is a major cereal
crop in Asia, providing fo od to m ore than ha lf of the
world population (Ma et al., 2007 ). Salinity happens to be
a major constraint to the sustainability and expansion of
rice cultivation in areas where rice production has not
kept up with increasing demand from a growing
population. The development of salt tolerant cultivars of
rice through conventional and modern molecular
techniques wou ld help solve the problem of world food
security. The cultivation of tolerant varieties on saline
fields may also reduce salinity by the process of
biological reclamation. The selection of salt tolerant
100
Asian J. Agr. Sci., 2(3): 99-105, 2010
In vitro salt tolerant rice plants established by step up
treatment with 0.5, 1.0, 1.5 and 2.0% NaCl at 3-week
intervals were examined by M iki et al. (2001). They
proposed that the in vitro step up salt selection induces the
capa bility to maintain no lethal concentration of NaCl in
the leaves. For overcoming salt stress, plants have
evolved complex mechanisms that contribute to the
adaptation to osmotic and ionic stress caused by high
salinity. These mechanisms include osmotic adjustment
that is usually accomplished by uptake of inorganic ions
as well as the ac cum ulation of com patible solutes
(osmop rotectants). Inorganic ions are sequestered in the
vacuoles (Yeo, 1998), while organic solutes are
compartmentalized in the cytoplasm to balance the low
osmotic potential in the vacuole (Rontein et al., 2002 ).
plants based on morp holog ical ma rkers has met with
limited success (A shraf et al., 2008 ). In the present,
salinity is the second most widespread soil problem in rice
growing countries next to drought and considers as a
serious constraint to increased rice production worldwide
(Gregorio, 1997). U nder this cond ition, rice is
perm anen tly challenged by drought, salinity stress along
with nutrient deficiency of the soil and other kinds of
biotic and abiotic stresses. This challenge results, if not
always but frequently, in a partial to drastic reduction of
yield. It was reported that breedin g for qu antative traits
like salinity, drought and heat tolerance. Chemical
insulting would be facilitated by the development of
procedu re to be used in Marker-Asisted Selection (MAS)
that is capable of identifying high performance genotypes
in early gene rations (Bohnert et al., 1995).
Genes liable for salt tolerance: There are some research
that showed, the salt tolerance in rice was controlled by
polygenes with the additive and dominant effects, the
former playing a major role (Moeljopawiro et al., 1981;
Gregorio and Senadhira, 1993; Gu et al., 1999). Akbar
et al. (1985) reported that the dry matter weight of rice
seed ling under salt stress was affected by at least two
groups of genes with additive effect, and no epistatic
effect was detected. Gregorio and Senadhira (1993)
observed that there were two groups of genes involv ed in
the sodium and potassium uptake in rice, one group for
sodium exclusion and the other for potassium absorption.
Physiological traits associated with salt tolerance in
rice were complex and controlled by a few major
QTLs (Yeo and Flowers, 1990). Previously, Xinjian
et al. (2002) identified some salt-stress responsive genes
in rice by cDNA array. T sai et al. (2005) demonstrated
that OsGR gene expression was increased in response to
NaCl and H 2 O 2 in roots of etiolated rice seedlings.
Eventhough, rice is a salt sensitive crop (Maas, 1990) yet
considerable genetic variability has been observed among
and even within rice cultivars (Akbar et al., 1972; Flow ers
and Yeo, 1981), which is helpful in selection and
development of tolerant rice to salinity stress through
genetic means (Nejad et al., 2008 ).
Saline soils and crop field: Generally, salt affected areas
are of two categories, the sodic and the saline. The major
differences between these two types of salinity are the
nature of anions and the pH of the soil. In sodic soils
carbonate or bicarbonate are the major ions wh ereas in
saline soil chloride or sulphate dominate. The pH of the
sodic soil is mostly above 8.5 while in saline soils it is
<8.5. Soil salinity is, generally, measured in units of
electrical conductivity (dS m-1) of a saturated so il paste
extract (ECe). If the ECe is > 4 dS m-1, the exchangeab le
sodium percentage is <15% and pH <8.5 the soil can be
considered as saline (Szabolcs, 1994). Although most of
the agriculturally important crops cannot grow in saline
soils, it is entirely not inimical to growth of all plant
species. Some plants grow well in salt affected coastal
areas, shores of backwaters lakes and marshy lands.
Those plants that can survive and grow well on high
concentrations of salt in the rhizosphere are called
halophytes. However, some other plants cannot even
tolerate a salinity caused by 10 % of seawater. Such plan ts
are called glycophytes or non-halophytes. Despite of these
expediency salinity responses of glycophytes w ill only be
discussed giving special reference to mulberry and other
woody plants, w herev er possible. H alophytes and their
response to salinity have been reviewed by Cherian
et al. (1999) and Gorha m (1995).
Salt stress proteins in rice plants: Kong -ngern
et al. (2005 ) investigated the salt-induced ch anges in
protein synthesis of the Thai rice using on e-dim ensional
SDS-PAGE and two-dimensional PAGE (2DE). The
protein was strongly induced by salinity in leaf sheaths of
treated tolerant Pokkali and Leuang Anan, compare d to
control plants. In contrast, its level in the sensitive rice
remained unchanged with salt treatment. Salt stress
proteins have also been reported in the roots of rice
plant (de-Souza et al., 2003; Claes et al., 1990;
Salekdeh et al., 2002).
Cap ability of salt tolerance in rice: Application of
molecular-marker aided selection technique fo r
improvement of salinity tolera nce w ould a ccelerate
breeding progress by increasing selection efficiency of
rice plant (Fotokian et al., 2005). With advents in plant
biotechnology as an aiding tool to conventional breeding,
molecular markers technique has become a powerful tool
to improve salt tolerance of rice by mapping and tagging
of genes involved in the control of growth and yield under
stressful environments (Haq et al., 2009).
Accumu lation of H 2 O 2 by salt stress: Tsai et al. (2005)
observed the accum ulation of H 2 O 2 by N aCl in the roo ts
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Asian J. Agr. Sci., 2(3): 99-105, 2010
systems in roots of rice seedlings. Sh ankhd har et al.
(2000) studied the embryogenic callus growth, plant
regeneration, and proline and total protein contents under
salt stress in six cultivars of rice (Oryza sativa L).
Treatment with NaCl caused an increase in the activities
of Ascorbate Peroxidase (APX) and Glutathione
Reductase (GR) and the expression of OsAPX and OsGR
in rice roots. Exogenously applied H 2 O 2 also enhanced the
activities of APX and GR and the expression of OsAPX
and OsG R in rice roots (Tsai et al., 2005).
of rice seedlings. The accum ulation of H 2 O 2 in rice roo ts
in response to NaCl w as inhibited by the NA DPH oxidase
inhibitors, diphenyleneiodonium chloride (DPI) and
imida zole (IMD ). How ever, DP I, IM D, and
dimethylthiourea, a H 2 O 2 trap, did not reduce N aClenhanced activities of APX and GR and expression of
OsAPX and O sGR . It appears that H 2 O 2 is not involv ed in
the regulation of NaCl-induced APX and GR activities
and OsA PX and O sGR expression in rice roots
(Tsai et al., 2005). The reduction of root growth by NaCl
is closely correlated with the increase in H 2 O 2 level.
Exogenous H 2 O 2 was found to inhibit root growth of rice
seedlings (Lin and Kao, 2001). Lee et al., (2001)
suggested that SOD leads to the overproduction of
hydrogen peroxide in the leav es of rice plants subjec ted to
salt stress. The overproduction of hydrogen peroxide
functions as the signal of salt stress, which induces the
induction of specific APX isoforms but not specific GR
isoforms under catalase deactivation.
Effect of salt offensive and its significance on rice
plant: Heenan et al. (1988) studied the salt tolerance of
several Australian and overseas rice varieties at
germination, early vegetative growth, and reproductive
development in a temperature controlled glasshouse to
determine the reliability of screening at any particular
stage. The Indica rice (Oryza sativa L.) varieties Pokkali
and Non a Bok ra are well-know n salt tolerance donors in
classical breed ing. In an attem pt to understand the
molecular basis of their tolerance, physiological and gene
expression studies were initiated (Moons et al., 1995 ).
Pokkali is a well-known salt tolerant dono r in
classical breeding and is commonly grown in coastal area
of Kerala, India. It is a trasditional, tall, photoperiodsensitive rice cultivar that is susceptible to lodging and
has low tillering capacity with long, broad, dark, and
droopy leaves. M oreover, the leav es senesc ence oc curs
quick ly after flowering. The grain has pericarp and
poor cooking quality (Gregorio et al., 2002).
Kawasaki et al. (2001) showed that Pokkali continued
growing at a low photosynthetic rate after 7 days of salt
stress, plant biomass approximately doubled. It was
known that Pokkali maintained water content in the shoot
during a 6-week stress under the conditions of 150 mM
NaCl. Lee et al., (2003) reported that there were no
significant differences between Japonica and Indica rice
types for shoot K+ concentration. Lee and
Senadhira (1999) also reported that sh oot K + conc. in
Japo nica rice showed no relation to salt toleran ce.
Role of antioxidant system in salt tolerance: Reactive
oxygen species are thought to play an importan t role in
NaCl stress. Plants tolerant to N aCl stress may evolve
certain strategies to remove these ROS, thus reducing
their toxic effects. Therefore, the expression patterns of
the gene family encoding the H 2 O 2 -scavenging enzyme
ascorbate peroxidase and glutathione reductase were
analyzed in roots of etiolated rice (Oryza sativa L.)
seedlings in response to NaCl stress (Hong et al., 2007;
Hong et al., 2009 ). Tsai et al. (2004) examined the
response of antioxidan t systems to NaCl stress and the
relative importance of Na+ and Cl– in Na Cl-induced
antioxidant systems in roots of rice seedlings. NaCl
treatment caused an increase in the activities o f Asc orbate
Peroxidase (APX) and G lutathione R eductase (G R) in
roots of rice seedlings, but had no effect on the activities
of Superoxide Dismutase (SOD) and Catalase (CAT)
(Tsai et al., 2004).
Lin and Kao (2001) investigated the chan ges in cellwall peroxidase (POD) activity and H 2 O 2 level in roots of
NaCl-stressed rice seedlings and their correlation with
root grow th. In the leaves of the rice plant, salt stress
prefere ntially enhanc ed the con tent of H 2 O 2 as well as the
activities of the Superoxide Dismutase (SOD), A scorb ate
Peroxidase (APX), an d pero xidase specific to gu aiacol,
whereas it induce d the decrease of catalase activity. On
the other hand, salt stress had little effec t on the activity
levels of Glutathione Reductase (GR) (Lee et al., 2001).
Dionisio-Sese and Tobita (1998) studied the possible
involvement of activated oxygen species in the
mechanism of damage by NaCl stress in leaves of four
varieties of rice (Oryza sativa L.) exhibiting different
sensitivities to N aCl.
Tsai et al. (2004) examined the response of
antioxidant systems to NaCl stress and the relative
importance of Na + and Cl– in Na Cl-induced antioxidant
Salinity induced limitations in rice plant: Rice is rated
as a salt sensitive crop (Shannon et al., 1998). Although,
salinity affects all stages of the growth and development
of rice plant and the crop respon ses to salinity varies w ith
grow th stages, concentration and duration of exp osure to
salt. In the most commonly cultivated rice young
seedlings were very sensitive to salinity (Flowers and
Yeo, 1981; Lutts et al., 1995). Water culture studies of
short (seedling stage) an d long term (maturity) stage w ere
conducted to evaluate the effect of different levels of
salinity on the grow th, yield a nd yield compo nents of
different inbred rice lines (Shereen et al., 2005).
Zeng et al. (2001) analyzed the effects of salinity on plant
grow th and yield components of rice by composing 20day periods of salinization at different growth stages.
102
Asian J. Agr. Sci., 2(3): 99-105, 2010
Asch, F. and M.C .S. Wopereis, 2001. Responses of fieldgrown irrigated rice cultiva rs to varying levels of
floodwater salinity in a sem i-arid enviro nme nt. Field
Crop. Res., 70(2): 127-137.
Ashraf, M., 1994. Breeding for salinity toleran ce in
plants. Crit. Rev. Plant Sci., 13: 17-42.
Ashraf, M., 2009. Biotechnological approach of
improving plant sa lt tolerance using antioxidan ts as
markers. Biotechnol. Adv., 27: 84-93.
Ashraf, M., H.R. Athar, P.J.C. Harris and T.R. Kwon,
2008. Som e prospective strategies for improving crop
salt tolerance. Adv. Agron., 97: 45-110.
Ashrafuzzaman, M., M.H. Khan and S.M. Shahidullah,
2002. Vegetative growth of maize (Zea mays) as
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Dionisio-Sese, M.L. and S. Tobita, 1998. Antioxidant
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chloride resistance within rice (Oryza sativa L.)
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Fotokian, M., A. Taleie, B. G harey azie, K . Postini,
A.A. Shahnejat Bushehri and Z.K. Li, 2005. QTL
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Gregorio, G.B . and D . Sena dhira, 1993. Gen etic ana lysis
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Gregorio, G.B., D. Senadhira and R.D. Mendoza, 1997.
Screening rice for salinity tolerance. IRRI discussion
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Gregorio, G.B ., D. Sen adhira, R.D . Mendoza,
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+CONCLUSION
Rice (Oryza sativa L.) is one of the most important
stable food crops in the world. Salinity is the major
constraint to rice cultivation that causes reduction of
yield. Thus in short, reviews previously explained on
various aspects of salinity tolerance in rice plants also
been expo sed numb er of mecha nism of responsible for
salinity tolerance in monocotyledon plants like different
varieties in rice plants. As this field is further advanced,
the accuracy of transcript profiling will become
increasingly exact, allowing the pattern of the complex
web of signaling to be uncovered. The response of rice
plants to salt and other environmental stresses have been
exten sively investigated for many decades, still we have
not been able to understand fully the mechanism which
imparts tolerance to som e plants and sensitivity to others
due to the complexity of the mechan ism. Emph asis shou ld
also be given to explore the natural genetic variations in
salt tolerance am ong rice plants and their wild relatives.
Regarding the rice plant, recent developments in genetics,
tissue culture, transgenesis, and linkage mapping show
that research in the coming years will definitely change
the salt tolerant capac ity of rice p lant.
ACKNOWLEDGMENT
The authors gratefully acknowledged to our
Malankara Catholic College Correspondent Fr. Prem
Kumar (M.S.W.) given encouragement and support for
preparation of this research man uscript. We wish to
express the sincere thank s to Mr. K. Suresh (MC C) for
updated article collection and all the efficien t supports of
this review preparation.
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