Journal of Integrative Agriculture 2017, 16(11): 2357–2374

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REVIEW

Effects of salt stress on rice growth, development characteristics, and

the regulating ways: A review

Sajid Hussain, ZHANG Jun-hua, ZHONG Chu, ZHU Lian-feng, CAO Xiao-chuang, YU Sheng-miao,
Allen Bohr James, HU Ji-jie, JIN Qian-yu

State Key Laboratory of Rice Biology, China National Rice Research Institute, Hangzhou 310006, P.R.China

Abstract
Rice (Oryza sativa L.) is highly susceptible to the rhizosphere salinity than other cereals. High sensitivity has been ob-
served, mainly at vegetative and reproductive stages in rice. It is the duty of plant physiologists to comprehend the growth,
development, and physiological processes of rice plants under stress. This paper includes the overview of rice growth
and developmental processes influenced by salt stress and the regulation pathways involved in these processes. It also
includes the promising salt tolerance strategies, i.e., genetic modification techniques, agronomic practices to improve rice
growth, yield, and role of phytohormones and their management, especially inhibition of ethylene biosynthesis by using
inhibitors 1-methylcyclopropene (1-MCP). Rice cultivation may be a first choice for improvement of salt tolerance through
plant growth regulators and improved cultivation techniques. This study will significantly improve the understanding toward
low rice grain yield and poor rice resistance under salt stress and will also stream scientific knowledge for effective utilization
of salt affected soils by using different regulating ways.

Keywords: rice, salt stress, ethylene, 1-methylcyclopropene (1-MCP), physiology

1. Introduction
Rice (Oryza sativa L.) is grown in many parts of the world
especially in Asia, Latin America, and Africa, and used as
staple food for more than 50% population of the world (Lou
et al. 2012). Almost 90% rice is cultivated in Asia, of which
Received 31 October, 2016 Accepted 28 February, 2017
Sajid Hussain, E-mail: Sajid_2077uaf@yahoo.com;
Correspondence JIN Qian-yu, Tel/Fax: +86-571-63370122,
E-mail: jinqianyu@caas.cn; ZHANG Jun-hua, Tel/Fax: +86-571-
63370122, E-mail: zhangjunhua@caas.cn
© 2017, CAAS. Publishing services by Elsevier B.V. All rights
reserved.
doi: 10.1016/S2095-3119(16)61608-8
China, India, and Pakistan contribute 30, 21, and 18%, re-
spectively, while remaining 30% is contributed by Thailand,
Indonesia, Burma, and Japan (Khush 2005; Calpe 2006).
Rice is high yielding crop, but current average yield is 10 t
ha–1 for indicia rice, 10 to 15% yield is lower than its potential
(Virmani et al. 1991; IRRI 1998). There are many reasons
for this yield gap, i.e., environmental stresses (biotic or
abiotic), management strategies, and nutrients deficiencies.
Abiotic stresses (especially salt stress) are among the major
causes of this low yield.
Salt stress has been a serious threat for crop production
in irrigated land, as expected salt stressed irrigated land is
more than 20% (Pitman and Lauchli 2002), and estimated
stressed area will expand to 50% of irrigated land by 2050
(Wang et al. 2003). Plant physiological physiognomies are
extensively susceptible to the highly saline rhizosphere.
High salt level affects seed germination, plant life, and crop
2358 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
productivity (Munns et al. 2008). Among the monocot crops,
rice is salt sensitive (Maas and Hoffman 1997; Ashraf et al.
2009), and its productivity is severely affected by the accu-
mulation of soluble salts in soils (Ashraf 2009).
There are many growth inhibiting effects of salt stress on
rice plants, such as reducing rates of net CO2 assimilation,
leaf growth, leaf cell enlargement, dry matter accumulation,
and relative growth (Cramer et al. 2001; Khan and Abdullah
2003; Amirjani 2010). Rice plant responses to salt stress
are complex and depend on duration and type of salt
stress, development stage of rice, day length, and other
factors (Bernardo et al. 2000; Cramer et al. 2001). Salinity
stress is dominated by sodium (Na+) and chloride (Cl–) ions
(Serrano et al. 1999) affect rice growth and development by
creating ionic, osmotic and oxidative stresses (Borsani et al.
2001; Tarakcioglu et al. 2002; Eraslan et al. 2007). Higher
amount Na+ in saline medium inhibits uptake and transport
of Ca2+ and might induce Ca2+ deficiency in plants because
of higher Na+/Ca2+ ratios (Lynch et al. 1985). Entrance of
high concentration of salt to rice plant will ultimately raise
toxicity level in adult leaves causing early leaf senescence
and decreasing the photosynthetic leaf area of rice (Munns
et al. 2002; Shereen et al. 2005). During early period of
salt stress, rice faces osmotic stress, and decreases leaf
development, while in long-term salinity stress, rice plants
experiences (Na+ and Cl–) ionic stress, and leads early
senescence of older leaves (Amirjani 2011).
Salt stress reduced rice growth rate, promoted metabolic
alterations, and decreased ability to uptake water and nu-
trients (Munns et al. 2002). Moreover, poor development
of rice spikelets, especially inferior spikelets caused by salt
stress significantly reduced rice grain yield (Fu et al. 2011;
Zhang et al. 2015). According to FAO (2009), 70% more
food is required for 2.3 billion people in the world by 2050.
So, it is necessary to improve food production in future es-
pecially rice yield under salt stress (Heong and Hardy 2009).
In plant life cycles, many phytohormones (especially
ethylene) play important roles in plants and environmental
interactions, i.e., salt stress, being considered as coordina-
tors between stress response and plant growth (Abeles et al.
1992), as ethylene signaling and production are essential
for plant quick response to salt stress and adaptation re-
sistance (Tao et al. 2015). Extreme ethylene productivity
tends to reduce plant growth and development, leading to
mortality. Therefore, ethylene homeostasis is important
for rice plants to survive under salt stress. Amongst the
prevailing ethylene inhibition approaches, chemical control
of ethylene biosynthesis under salt stress by using inhibitors
has become popular. The inhibitors like aminoethoxyvi-
nylglycine (AVG) and 1-methylcyclopropene (1-MCP) are
useful ethylene inhibitors for crops to maintain grain yield
(Watkins et al. 2008). Zhang et al. (2015) reported that
1-MCP played positive effect on starch content in superior
and inferior spikelets, spikelet fertility, grain yield, and har-
vest index for rice cultivars.
This review paper will cover (i) effects of salt stress on
rice growth and development, (ii) effects of salt stress on rice
physiological characters, (iii) role of ethylene in rice growth
and development, and (iv) amelioration ways for increasing
rice grain yield under salt stress.
2. Salt stress
Normal soil has pH=4.5–7.5, electrical conductivity (EC)<
4 dS m–1, exchangeable sodium percentage (ESP)<15, and
sodium absorption ration (SAR)<15, make most favorable
environment for nutrient availability and plant growth (Bohn
et al. 1985). However, salt stressed soil is that one has
a high concentration of soluble salts that may negatively
affect plant growth. According to FAO report (FAO 2009),
soils affected by salt have a high soluble salt like sodium
(Na+), magnesium (Mg2+), calcium (Ca2+), chloride (Cl–),
and sulphate (SO42–) (Bohn et al. 1985; Manchanda and
Garg 2008). Saline soil has EC>4 dS m–1, ESP<15, and
SAR<15. Sodic soil is another category of soil affected by
salt, which has EC less than 4 dS m–1, ESP≥15, and SAR≥15
(Bohn et al. 1985; SSSA 1997). Salt stress occurs due to
natural or anthropogenic activities that result in the high
accumulation of soluble salts in the underground water.
According to the FAO, more than 6% land in the world is
affected through salinity or sodicity, and more than 20% of
irrigated land has become salt stressed (Pitman and Lauchli
2002; Munns 2005).
Under salt stress, there are many explanations for poor
plant performances, i.e., agronomic management, assimilate
supply, enzyme activity, nutritional management, genotypic
traits, and hormonal imbalance (Yang et al. 2000).
2.1. Ion toxicity
The most harmful effect of salt stress is the Na+ and Cl–
accumulation in plant tissues and soil (Eraslan et al. 2007;
Nishimura et al. 2011). Entrance of Na+ and Cl– into the plant
cell causes ion imbalance in plant and soil, and uptake of Na+
and Cl– in excess by plant might cause major physiological
disorders in plant (James et al. 2011). High concentration
of salts in soil profile may cause reduction in water uptake
due to salt accumulation in root zone (physiological drought)
(Munns 2005), reduction in osmotic potential of plant, and
disruption of cell metabolic functions due to toxicity (Rozema
and Flowers 2008; Evelin et al. 2009; James et al. 2011).
According to an estimate, salt stress has reduced crop
production about 20% of irrigated lands worldwide, and the
loss of 50% arable land due to salt stress up to mid of the
 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
21st century (Mahajan and Tutejan 2005).
Generally, salt stress affects crops by osmotic, specific
ion, ion imbalance, and oxidative damage (Flowers et al.
2005; Prida and Das 2005). Excess Na+ in plant harms cell
membrane and organelles of plant, consequently reduction
in plant physiological mechanisms leading to plant cell
death (Quintero et al. 2007; Siringam et al. 2011). These
physiological changes in plant include the membrane dis-
ruption, inability to detoxify the reactive oxygen species
(ROS), reduced photosynthetic rate, and transformations
of the antioxidant enzymes (Rahnama et al. 2010; James
et al. 2011). These oxidative systems can interrupt the
routine functions of various plant cellular components such
as proteins, DNA, and lipids, interfering dynamic cellular
functions in plant (Demiral et al. 2005).
2.2. Nutrient imbalance
Salt stress in soil can also cause nutrient imbalance in rice
plants. Root has direct contact with the growth medium and
the rest of the plant. It is considered as a bridge between
nutrients and plants in salt stress. The most important
harmful effect of salt stress is the Na+ and Cl– accumulation
in plant tissues and soil (Nishimura et al. 2011). High Na+
concentration has antagonistic effect on potassium (K+) ion
which is an essential plant nutrient for growth and plant
development (Jung et al. 2009). Moreover, nitrogen uptake
reduction by plant has also been observed under high salt
conditions (Abdelgadir et al. 2005). Further studies showed
that salt stress has antagonistic effect on P, K+, Zn, Fe, Ca2+,
and Mn but has synergistic effect on nitrogen (N) and Mg in
rice crop (Jung et al. 2009; Garcia et al. 2010).
3. Effects of salt stress on rice growth and
development
Rice is considered as a salinity sensitive crop in the field as
compared to other main cereal crops (Joseph et al. 2010).
Salt stress severely influences the morphology traits of rice
plants (Jamil et al. 2006; Zhang et al. 2012).
3.1. Effects on rice growth
Rice is more sensitive to salt stress at early seedling stage as
compared with tillering stage (Grattan et al. 2002; Shereen
et al. 2005). Seedling growth and fresh weight decreased
with increased salt stress from 5 to 7.5 dS m–1 (Kazemi and
Eskandari 2011). Various green house and field trials also
revealed that salt stress decrease rice stand density and
seedling biomass production (Zeng and Shannon 2000).
Root has direct contact with the biotic and abiotic fac-
tors in the soil environment and the rest of the plant (Smet
2359
et al. 2012). H2O and solutes enter by the roots through
a combination of different pathways such as symplastic,
apoplastic, and transcellular, however, transport of water
and solutes via apoplastic transpiration pathway is very
important in rice (Ochiai and Matoh 2002; Kronzucker and
Britto 2011). Mostly, Na+ passages in shoots of rice through
the apoplastic pathway where Na+ transports via apoplast
through solvent drag and Casparian bands (Ranathunge
et al. 2005; Gong et al. 2006). Significant reduction in mean
root length, mean root numbers per plant, and shoot length
occurred under increased salt stress (Jamil et al. 2006; Jiang
2010). Thus, root and shoot lengths are two indicators of
rice plant response to salt stress. Rice cell division and cell
elongation are seriously affected by salt stress, which lead
to reduction of root, leaf growth, and yield (Munns 2002). Tu
et al. (2014) studied that genome duplication also effect on
rice root resistance against salt stress. They observed that
tetraploid rice (HN2026-4x and Nipponbare-4x) improved
root growth, proline, membrane, organelles, stability of nu-
clei, normal epidermis cell frequency, H+ efflux on root tip,
and decreased Na+ in rice roots, malondialdehyde (MDA)
contents than diploid rice (HN2026-2x and Nipponbare-2x).
The reason behind this phenomenon is the exposure of pro-
tective gap between cortex and pericycle cells in tetraploid
rice (HN2026-4x and Nipponbare-4x), which increases root
resistance to salt stress, improves H+ transport to the root
surface, and plays a role in decreased entry of Na+ into the
rice roots (Tu et al. 2014).
Rice leaf mortality increased with increased salt stress in all
rice cultivars at early seedling stage (Shereen et al. 2005). It
is about 0 to 300% leaf mortality in salt stress exposure after
1 wk. Later in a few months, salt stress shows reduction of
growth and development (Munns 2005). It may cause death
of leaves and decrease in leaf area and ultimately reduce
photosynthesis rate of plant (Amirjani 2011). Salt stress has
specific effects on plant cell metabolism, particularly on leaf
senescence. It can also injure the cells in transpiring leaves,
and cause rice plant growth inhibition. The salt concentrate
in the old leaves cause the leaves death, which is crucial for
the survival of a plant (Munns et al. 2006).
3.2. Effects on rice grain development
Panicle sterility is a major issue in rice grain yield under salt
stress (Flowers and Yeo 1981). In some rice cultivars, salt
stress caused panicle sterility, especially at pollination and
fertilization stages due to some genetic mechanisms and
nutrient deficiencies resulting from salinity effect (Khatun
and Flowers 1995; Hasamuzzaman et al. 2009). Various
studies showed that during fertilization, salt stress may
cause sterility of panicle, which leads to a decline in grain
setting, pollen bearing capacity, and decrease of the stig-
2360 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
matic surface, or both (Abdullah et al. 2001).
The main cause of decreased grain yield under salt stress
is lack of transformation of carbohydrates to vegetative
growth and spikelets development. Zeng and Shannon
(2000) observed negative linear relationships in many yield
components with increased salt stress such as number of
tillers per plant, number of spikelets per panicle, and percent
of sterile florets (Khatun and Flowers 1995; Zeng and Shan-
non 2000). Moreover, significant reduction in translocation
of soluble sugar contents to superior and inferior spikelets
and inhibition of starch synthetase activity during grain
development are the main reasons behind lower rice grain
yield under salt stress (Abdullah et al. 2001).
Yield components such as spikelets per panicle, panicle
length, number of tillers per plant, number of florets per
panicle, and 1 000-grain weight are all severely affected due
to increasing salt stress (Khatun and Flowers 1995; Farshid
and Hassan 2012). Zeng and Shannon (2000) studied the
interrelationship among yield parameters of rice under salt
stress. Significant reduction in seedling growth was ob-
served with increased salt stress from 1.9 to 6.1 dS m–1. In
another study, reduction in number of filled rice grains per
panicle decreased with increased salt stress from 2 to 8 dS
m–1 (Farshid and Hassan 2012). All these yield components
are linked to each other regarding final grain yield. There
is a need to understand this phenomenon about salt stress
regarding rice yield loss.
4. Effects of salt stress on rice physiolog-
ical characteristics
Plant physiological characters are susceptible to the high
salt level in its rhizosphere. Salt stress showed numerous
physiological effects on rice plants, such as decrease in
photosynthetically active radiation (PAR), net photosynthe-
sis (Pn), stomatal conductance (Gs), transpiration rate (Tr),
degradation of pigment, and relative water content (RWC)
(Cattivelli et al. 2008; Tuna et al. 2010). Salt stress also
has significant effect on water use efficiency (WUE) of rice
plant (Ramezani et al. 2012). Biomass and seed yield have
positive correlation with WUE. WUE of chickpea varieties
improved in 2 dS m–1, but WUE of rice plant reduced with
increased salt stress (Gholipoor et al. 2002). All these
factors cause negative pleiotropic effects on rice physiology
and development at the molecular and biochemical levels
(Munns 2002; Tester and Davenport 2003), and cause ab-
normal rice growth, development, and ultimately plant death
(Parida and Das 2005; Nishimura et al. 2011).
4.1. Effects on rice photosynthesis
Rice plants under salt stress have decreased photosyn-
thetic efficiency by the complex of photosystem II (PSII).
Moreover, chlorophyll contents in rice leaves are damaged
by the addition of Na+ and Cl–, which might hinder the major
electron transport in PSII (Sudhir and Murthy 2004; Munns
et al. 2006). Cha-umi et al. (2009) found that chlorophyll
and carotenoids contents in rice leaves were significantly
decreased after imposition of salt stress. High salt stress
has also been reported to reduce the quantum yield of
the complex PSII, and to lower K+/Na+ ratio (Garcia et al.
2012). Dionisio-Sese and Tobita (2000) observed higher
non-photochemical quenching (qN) salt sensitive cultivars
with increased salt levels while, reduction in quantum yield
of PSII (Fv/Fm) and photochemical quenching coefficient (qP)
(Netondo et al. 2004). Therefore, higher photosynthesis rate
is better for integrate partitioning and sink strength of supe-
rior and inferior spikelets (Li et al. 2012; Chen et al. 2013).
4.2. Effects on nutrient accumulation and transport
Salt stress may cause ion imbalance, due to severe compe-
tition of Na+ and Cl– with K+, Ca2+, and NO3–. Generally, high
NaCl lead to specific ion (Na+ and Cl–) toxicity in plants. It
decreases the N, P, K, Ca, Mg, and increases Na+/K+, Na+/
Ca2+, and Ca2+/Mg2+, and Cl–/NO3– in plants which cause
imbalance of nutrients (Grattan and Grieve 1999; Abd El-
Wahab 2006; Razzaque et al. 2011; Zeinolabedin 2012).
Salinity is also associated with a reduction in nutrient uptake
by plants, such as N in plant tissues observed in various
crops like bean (Chakrabarti and Mukherji 2003) and rice
(Abdelgadir et al. 2005). Ionic imbalance occurs due to
high accumulation of Na+ and Cl–, and antagonistically it
decreased the uptake of other nutrients in plant cell and
tissues (Table 1). K+, Ca2+, and Mn2+ are the main elements
observed with lower contents and imbalanced ratios in plants
due to competition (Sudhir and Murthy 2004; Soledad et al.
2012). Abdur et al. (2011) reported that high salt stress
significantly affect Na+, Ca2+, K+, and Mg2+ concentration in
root and shoot in rice plant. While boron (B) and silicone (Si)
availability decreased under high salt levels in plants (Wim-
mer et al. 2001; Currie and Perry 2007). High concentration
of NaCl in many cereal crops including rice, decreased zinc
(Zn) availability and increased cadmium (Cd) toxicity (Nor-
evell et al. 2000; Amanullah and Inamullah 2016).
Effects on hormone production: Plant hormones play their
roles as chemical messenger within the plant’s parts, where
they play important roles in plant growth and development,
and response to biotic or abiotic stress at very minute con-
centration, so they are known as plant growth regulators
(PGRs) (Javid et al. 2011). Plant hormones play key roles
in the plants to adapt to changing biotic and abiotic environ-
ments, by facilitating growth and development, source and
sink transformation, and nutrient balance in plant (Fahad
 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
Table 1 Nutrient deficiencies under salt stress in rice
Na+/Cl– toxicity Antagonistic
Potassium ion (K+) High sodium (Na+) and less potassium (K+) in
cytoplasm of the cell
Na+ concentration increases as
Magnesium (Mg) – Mg increased as salt stress
Zinc (Zn) Zn deficiency occur when salt stress increase
Nitrogen (N) N deficiency occurs under salt stress
Phosphorous (P) P decrease in rice grain under salt stress
Calcium ion (Ca2+) Ca2+ decrease under high Na+
Iron (Fe) Fe deficiency occurs as Na+ increase
et al. 2014). Hormonal imbalance can occur in rice plants
under severe environmental stresses, such as drought,
heat, cold, and high salt stress (Hojin and Yong-Gu 2015).
This imbalance varies from abscisic acid (ABA) to gibber-
ellins (GAs), cytokinins (CKs) to auxins (IAA), jasmonate
(JA), salicyclic acid, brassinosteriods (BRs), and triazoles
under stress. But the involvement of phytohormones in the
abiotic stress tolerance is still not clear (Maetsri et al. 2002;
Fahad et al. 2014). The CKs and ABA play regulatory roles
between leaf senescence and assimilate remobilization in
wheat and rice (Yang et al. 2002, 2003, 2004, 2006). These
plant hormones are considered as the most vital elements
that are critical in controlling physiological responses in
plant under salt stress environment (Khan and Khan 2013).
ABA ABA is considered as the stress hormone because
it effect internal signaling of plant against adverse environ-
ment, and plays a vital role throughout the plant life cycle
and facilitates plant reactions to different abiotic and biotic
stresses (Keskin et al. 2010; Devinar et al. 2013). ABA bio-
synthesis increased in rice under osmotic stress due to salt
accumulation (Kumar et al. 2013). Especially, in rice roots,
abiotic stress significantly increases ABA then decreases
the zeatin riboside concentrations and zeatin in the leaves
and root exudates (Yang et al. 2002; Hong et al. 2007). ABA
has direct effect on grain filling rate. Higher concentrations
of ABA such as 0.1 mol L–1, hinder the transport of sucrose
to the grains and decrease the synthesis ability of starch
of intact grains, consequently, lower grain filling rate and
grain weight (Ahmadi and Baker 1999; Yang et al. 2006).
Exogenous ABA remobilizes the carbon (CO2) from the
leaves and stems, far away from superior to cytokinins (Yang
et al. 2003). It is evidence that application of ABA improved
expression of OsAPx8 in rice roots, rice grain production,
and 20% survival rate (100 µmol L–1, application of ABA to
indica rice) through accumulation of soluble sugars, proline
by expression of OsP5CS1 gene, enhancement of K+/Na+
ratio, and homeostasis in K+ and Ca2+ was useful in avoiding
2361
Synergistic References
– Jung et al. (2009);
Razaque et al. (2011)
salt stress increases
Razaque et al. (2011)
increase
– Amanullah and Inamullah (2016)
– Singh et al. (2005)
– Naheed et al. (2007);
Amanullah and Inamullah (2016)
– Amanullah and Inamullah (2016)
– Garcia et al. (2010)
imbalance of Na+, Cl–, and Na+/K+ ratio in rice under salt
stress (Hong 2007; Gurmani et al. 2013; Sripinyowanich
et al. 2013).
GAs GAs as a plant growth hormone are involved in plant’s
growth and development, and biosynthesis of GAs was
rapidly reduced in rice plants under salt stress (Javid et al.
2011; Daviere and Achard 2013; Colebrook et al. 2014). Re-
cent researches have discovered that the key GA negative
regulator, DELLA family proteins, are essentially involved in
controlling environmental signals and other plant hormone
signaling pathways (Achard et al. 2006). The dominant
roles of GA for regulation of plant growth is due to DELLA
proteins facilitated growth restrain on exposure to salt stress
(Achard et al. 2008). The plant growth parameters and
salt tolerance in a range of the DELLA mutants indicated a
strong correlation among plant height, flowering time, and
exposure to heavy salt stress. It is suggested that the ac-
cumulation of DELLA protein reduces growth and improves
salt tolerance by decreasing GA signaling action (Achard
et al. 2008). GA hormone is specifically important in rice as
compared to other plant hormones, concerning in alleviation
of harmful effect on salt stress (Colebrook et al. 2014). In
this situation, application of GA3 can improve rice growth
and development in salt stress (Javid et al. 2011).
IAA Auxin (IAA) is considered the first ever discovered plant
growth hormone and known as IAA maxima (Lau et al. 2008;
Fahad et al. 2015). It plays an important role in cell elonga-
tion, apical dominance, and development of vascular tissue
of plant (Wang et al. 2001). It is also reported the response
of IAA under salt stress and correlation between salt stress
and IAA signaling in may crops. These studies proposed that
sprouting of seed is modified by a membranous bounded
transcript (NTM2) factor that integrates IAA signal in ger-
mination of seed. Moreover, signal pathway in salt stress
is intermediated by overexpression of transcription factor
NTM2 of IAA30 in plant (Jung and Park 2011; Park et al.
2011; Iqbla et al. 2014). Reduction of IAA was observed
2362 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
in rice under salt stress (Nilsen and Orcutt 1996; Kazan
2013). Many primary IAA response genes are stimulated
under unfavorable environment in Arabidopsis, soybean,
and rice plants (Hangen and Guilfoyle 2002). These auxin
in IAA responsive genes are categories into three families,
for example: (i) GH3, (ii) small auxin-up RNA (SAUR), and
(iii) auxin/indoleacetic acid (Aux/IAA) gene families. IAA
hinder the development of tillering in rice by the expression
of down regulating gene (OsIPT) and cytokinin production
in buds and nodes (Liu et al. 2011). The reduction in rice
plant growth and development under salt stress might be
the consequence of accumulation and recognition of auxin.
So, exogenous application of IAA or NAA can provide an
effective method to counter the salt stress conditions in
rice cultivation.
CKs CKs (adenine derivatives with aromatic or isoprenoid
side chain) play key roles in cell division, apical dominance,
biogenesis of chloroplast, differentiation of vascular tissues
and shoot, leaf senescence, production of anthocyanin, and
development of photo-morphogenesis in plant (Davies 2004;
Fahad et al. 2014). CKs have resistant effect against salt
stress in plants (Barcizewski et al. 2000; Fahad et al. 2014).
CKs have antagonistic effect on ABA and synergistic/antag-
onistic effect on IAA and help plants to reduce salt stress
(Iqbal et al. 2006, 2014). CKs synthesis in rice plant was
also reduced under salt stress (Nishiyama et al. 2012). It
plays negative role in plants by delaying closure of stomata
and leaf senescence (Pospisilova et al. 2005). It is reported
that exogenous application of CKs or seed priming reduced
the effect of salt stress and increased plant tolerance to salt
stress in cereal crops and improved growth and yield in rice
(Zahir et al. 2001; Iqbal et al. 2006; Javid et al. 2011).
JA The JA is an important cellular plant growth regulator.
It plays vital role in seed germination, cell and root growth,
leaf senescence, and flowering (Tani et al. 2008). The main
sites for JA production in plant are peroxisomes and chloro-
plast as well as root and leaf, and act as defense response
to unfavorable environment (Cheong and Choi 2003; Mei
et al. 2006). JA was reported more in salt tolerant cultivars
than in salt sensitive cultivars (Kang et al. 2005). Recent
researches proposed some evidences of link between JA
and ABA for salt stress regulation (Shinozaki and Yamagu-
chi-Shinozaki 2007). A defense receptor protein SCFCOI1
through JAZ (jasmonate ZIM-domain) transcription factors
of JA plays important roles in plant development in abiotic
stress (Kazan and Manners 2012). Tani et al. (2008) re-
ported that with increased salt stress, level of JA increased
in roots and leaves which results in induced proteins and
JA production genes in rice. JA persuades proteolysis of
JAZ through SCFCOI1 facilitated 26s proteasome pathway,
which functions in direct suppression of MYC2. MYC2 could
trigger the downstream target gene expressions related to
JA, ABA, and salt stress (Kazan and Manners 2012). Many
researchers suggested that JA-activated MYC2 may func-
tion in controlling the ABA-mediated regulation of salt stress
responsive genes. Along with these, several JAZ proteins
have been identified such as OsTIFY1, 6, 9, 10, and 11 as salt
resistant genes in rice (Ye et al. 2009). JA plays a important
role in antagonistic effect with ABA in rice plant, helping in Na+
reduction in rice root (Kang et al. 2005). External application
of JA decreased concentration of Na+, caused balance with
ABA, and amended salt affected rice seedlings especially in
salt sensitive rice cultivar (Kang et al. 2005).
BRs BRs are modern discovered plant hormone contained
castasetrone (CS) and brassiniloide (BL) and their deriva-
tives that play important roles in plant growth regulation,
development, and other physiological mechanisms (Kartal
et al. 2009). Key roles of BRs are, germination of seed,
vegetative growth, ethylene, nucleic acids, production of
protein, net photosynthesis (Pn), reproductive growth, and
maturity (Hayat et al. 2010; Fahad et al. 2015). BRs also
reduced the harmful effects of salt stress on plant growth,
development, and production (Wang et al. 2011). Post appli-
cation of BRs improved the germination of seed, root growth,
and rice production through recovering chlorophyll contents
and enhancing activity of nitrate reductase (Anuradha and
Rao 2001). Treatment of BRs (24-epibrassinolide, 24-EBL)
in rice increased germination of seeds, growth, proline,
antioxidant system, and decreased lipid peroxidation under
salt stress, and modulate enzymatic and non-enzymatic
antioxidant in Arabidopsis, Brassica, and rice under salt
stress (Ozdemir et al. 2004; Sharma et al. 2013). In salt
stress conditions BRs has positive features by evidence of
some researches that focused on treatment of 24-EBL in
Arabidopsis and rice (Ozdemir et al. 2004; Divi et al. 2010).
BRs has synergistic effect on GA and IAA during hypocotyl
elongation in plant, while ABA has antagonistic effect and
repressed the BRs enhanced expression proteins (BEE1,
BEE2, and BEE3) (Friedrichsen et al. 2002; Tanaka et al.
2003). Still BRs response in carbon assimilation, way of
biosynthesis, involvement of enzymes, response against
stress, and microorganisms need further studies.
4.3. Effects of polyamines (PAs) on hormones under
salt stress
PAs play key roles in plant growth and development pro-
cesses, i.e., germination of seed, lignification of tissue,
development of vegetative, and reproductive growth, ab-
scission, senescence, endosperm cell division, and grain
development in rice and response of biotic and abiotic
stresses (Jang et al. 2012; Wang et al. 2012). PAs (pu-
 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
trescine Put, spermine Spm, and spermidine Spd) have
synergistic and antagonistic effects on production and
signaling of plant hormones. Putrescine (Put) showed
positive effect to the biosynthesis of ABA regulating gene,
however, Put had passive effects on production of JA, GA,
and ethylene under stress conditions. Spermidine (Spd)
showed opposite effects on hormones than that of Put. On
the other hand, spermine (Spm) increased the expression of
genes for production of JA and ethylene, and decreased for
GA and ABA in rice under stress. PAs also play important
roles in pathways of hormone signaling. Spd, Spm, and Put
positively linked with singaling gene of salicylic (SA), IAA
and CKs, and impartial to positive in JA, respectively. PAs
look to be unbiased in production and signaling pathways
to BRs in rice under stress condition (Anwar et al. 2013).
PAs showed antagonistic effect on ethylene biosynthesis
in rice under stress and improve spikelet’s development
especially inferior spikelets of rice. Application of Put, Spm,
Spd or an inhibitor (AVG) to rice biosynthesis of ethylene
and ACC significantly decreased, however, improved Put,
Spm, and Spd contents, grain filling rate, and grain weight
of inferior spikelets in rice under osmotic stress (Chen et al.
2013). Excessive expression of PAs gene in transgenic
plants improved stress tolerance in rice under stress (Al-
cazar et al. 2010).
5. Role of ethylene in rice growth and
development
Ethylene (C2H4) is a very important plant growth regulator
Methionine
(C 5H11 NO 2S)
S-AdoMet synthase
S-Adenosyle-methionine (SAM)
C15H22N6O 5S
ACC synthase
1-Amino-cyclopropane-
ACC-oxidase
carboxylic acid (ACC)
C4H7NO 2
Fig. 1 Biosynthesis of ethylene pathway, extracted from Lin et al. (2009) and Yang et al. (1984).
2363
(PGR), and is gaseous organic compound. Ethylene is
important in flowering, sexual diversity, senescence of leaf,
and numerous abiotic stress responses in plants (Naik and
Mohapatra 2000; Li and Solomon 2003). In the plant life
cycle, biosynthesis of ethylene increased during seed ger-
mination, ripening of fruits, leaves abscission, and flower’s
senescence of plant. When rice plant is under salt stress,
ethylene production is increased, so named as stress hor-
mone (Zhao et al. 2004; Dimitra and Derrick 2013).
5.1. Ethylene biosynthesis under stress
Biosynthesis of ethylene in plants comprises of three
enzymatic reactions. The first step is that methionine is
changed into S-Adenosyl-methionine (S-AdoMet) through
S-AdoMet synthetase (Fig. 1). During the second step, the
straight ethylene precursor ACC is produced from S-AdoMet
through ACC synthase (ACS). In final step, ethylene is
synthesized by the oxidation reaction of ACC through ACC
oxidase (ACO) (Lin et al. 2009). ACS as limiting enzyme,
its main target is aimed at regulation of ethylene produc-
tivity in abiotic stress (Bleecker and Kende 2000; Wang
et al. 2002). Ethylene production in rice plant increased
generally by enhancing ACOs and ACSs under salt stress
(Peng et al. 2014).
ACO is a main ethylene biosynthesis enzyme controlled
by salt stress. Under high salt stress, both ACO and ACC
production increases (Kukreja et al. 2005). We can suggest
through these studies that plants influenced by salt stress
and other environmental factors incline toward increased
Methylethioadenosine
(MTA)
Effect plant
Ethyelne (C 2H4) growth and yield
CO 2+CNH+H2O
1/2O 2
2364 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
ethylene production generally by enhancing ACOs and
ACSs (Peng et al. 2014). Recently it has been found in
rice that a kinase salt intolerance 1 (SIT1) lectin receptor
facilitated sensitivity of salt by homeostasis of ethylene
regulation. In existence of salt stress, SIT1 was stimulated
quickly to phosphorylate MPK3 or MPK6 and then enhanced
biosynthesis of ethylene. More expression of SIT1 in rice
enhanced sensitivity to salt stress (Li et al. 2014).
Ethylene stimulates leaf senescence, reduces chlorophyll
contents and Pn in leaves, decreases the level of starch and
protein, and enhances the function of numerous hydrolytic
enzymes (Weaver et al. 1998; Antonia and Alessandra
2006). It also involves the activation of some genes linked
with senescence like SAGs (Lers et al. 1998). In case of
rice plant, salt stress activated mitogen-activated protein
kinases (MAPK) forces to steady ACSs, that led to increased
ethylene signaling and production of ethylene, the reactive
oxygen species (ROS) accumulation, and growth inhibition
(Vandenbussche et al. 2012; Li et al. 2014; Steffens et al.
2014).
5.2. Role of ethylene in rice growth and development
Several studies found key positive ethylene signaling com-
pounds, such as MHZ7/OsEIN2, MHZ6/OsEIL1, and OsEIL2
(Tao et al. 2015). All these compounds play negative role
in rice seedlings in salt stress. Recent studies showed
that ACS2, ACS6, ACS7, and ACS8 in Arabidopsis were
encouraged through high salt stress; however pretreatment
of moderately low salt stress is helpful for salt acclimation
(Shen et al. 2014). These results give a clue that increasing
of ethylene production by a high and abrupt salt stresses
may be causing various negative impact on plant (Shen
et al. 2014).
Generally, ethylene is considered as a bridge between
salt stress and plant growth. However, extreme ethylene
production under severe salt stress mainly reduces plant
growth and development. It affects an extensive range
of physiological mechanisms in rice. These physiologi-
cal mechanisms cover senescence, ripening, chlorophyll
content loss, and other physiological disorders including
browning, sprouting, deterioration, and stimulus of defense
mechanism (Saltveit et al. 1999). Ethylene is not the major
factor in salt stress prompted inhibition of rice growth. The
situation is quite possible that ethylene might be playing
an indirect adverse effect in rice plant under salt stress at
different growth stages. Therefore, it is essential to know
the actual role of ethylene in growth and development of rice
response under salt stress (Shibli et al. 2007).
According to Mohapatra and Mohapatra (2006), com-
pact-panicle rice cultivars did not gain high yield potential
because of poor spikelets development, particularly inferior
spikelets. Consequently, inferior spikelets developmental
modification is very important for new hybrid rice genotypes
to get optimal yields. Spikelets produced high ethylene at
rice anthesis stage, and inferior spikelets synthesize more
ethylene at anthesis stage in rice (Yang et al. 2007). There
is a negative correlation among the ethylene biosynthesis
rate and cell division, growth, and grain filling rate in inferior
spikeltes contributing poor spikeltes development (Yang
et al. 2005). Therefore, developmental modification of infe-
rior spikelets is very important for new hybrid rice genotypes
with high yield potential. Ethylene production in spikelets
is one of the major yield reducing factors in rice. So, it
needs to adopt ethylene inhibiting management strategies
for increasing rice yield along with genetic improvement
under salt stress.
6. Adoptation responses of rice to salt
stress
The ability of rice plant to tolerate salt stress is an important
feature in rice production (Momayezi et al. 2009; Siringam
et al. 2011). Tolerance or resistance of rice plant to salt
stress involves many adaptive responses at molecular,
cellular, and physiological levels.
6.1. Osmotic adjustment
Osmotic adjustment is the most important physiological
aspect, as it determines the tolerance limit of rice plant to
bear toxic ions concentration. The most favorable strategy
is to reduce the accumulation of NaCl in the cytoplasm of
the plant cell or compartmentalization in vacuoles (Asch and
Wopereis 2001). Moreover, accumulation of compatible
compounds like organic solutes, free sugar, glycinebetaine,
and proline in the cytoplasm of plant cell is an important
strategy and gain attention of physiologists and genetic
engineers (Jampeetong and Brix 2009). It indicates that
under osmotic stress, carbohydrates or trehalose might be
more important for rice than proline (Nounjan et al. 2012).
Salt stress induces osmotic and ionic stresses, which inhibits
the normal cell growth and division of rice plant. To meet
the antagonistic biotic and abiotic stresses, plants maintain
homeostasis in osmotic/ion, and with quick osmotic signaling
and ionic signaling as shown in Fig. 2 (kumar et al. 2013).
Leaf area of rice plant is another important characteristic
that can lower effects of high Na+ concentration in rice
leaves by dilution effects and the transpiration force (Akita
and Cabuslay 1990).
6.2. Closure of stomata
Rapid closure of stomata is an initial response to salt stress
 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374 2365

Salt stress in rice

Osmotic effect Ionic effect

Osmotice signaling Homeostasis in Ionic signaling

osmotic/ionic

Cell division and growth

Salt tolerance in rice

Fig. 2 Salt stress response in rice plant, extracted from Kundan et al. (2013).

in plant (Foad and Abdelbagi 2007). Increase in salt level
(0 to 20 dS m–1) reduces plant growth by decreasing CO2
assimilation, which cause stomatal closure causing lower
intracellular CO2 partial pressure leading to decreased pho-
tosynthetic rate (Amirjani 2010). Transpiration rate declines
with increased NaCl concentration in rhizosphere, lowering
of water potential in roots, and the transport of ABA from
root to shoot, consequently, tend to induce stomatal closure
and prevent the dehydration of leave tissues (Zheng et al.
2001; Zhang et al. 2006). Salt stress induces biosynthesis
of ABA that closes stomata when reached to guard cells
(Zeinolabedin 2012). The redox state of ascorbate controls
stomatal action by regulating concentration of H2O2 as a
signal in guard cell movement (Chen et al. 2004). Generally,
salt stress inhibits photosynthesis rate in numerous plant
species (Dionisio-Sese and Tobita 2000). But in case of
monocots crops including rice, the credible reasons for de-
clining photosynthesis rate are closure of stomata, reduction
of sink activity, reduced efficiency of rubisco, dislocation of
vital cations from the membrane structure of leaf which lead
to changes in permeability, and swelling and inefficiency of
the grana (Flowers and Yeo 1981), or may be due to the
direct effects of salt on conductance of stomata through a de-
crease in guard cell turgidity and CO2 partial pressure within
cell (Dionisio-Sese and Tobita 2000). Closure of stomata
plays vital role to survive with salt stress. Chen et al. (2004)
studied that the ascorbate or ascorbic acid (AsA) redox state
controls the transpiration rate and conductance of stomata.
They noticed that a higher AsA redox state significantly in-
creases the area of the total open stomata of a leaf which
results higher conductance of stomata and a greater rate
of transpiration. Stomatal guard cells control through Na+
which control transpiration rate according to concentration
of salt presented in the environment (Ashour et al. 2002).
7. Management strategies for increasing
rice grain yield under salt stress
7.1. Genetic modification in rice cultivars
To gain potential production of rice crop, genetic modification
in rice against environmental stresses is very important. In
this approach, we can modify salt resistance genes in our
desired rice cultivar. We can also use advanced techniques
to improve rice tolerance and resistance against stress,
like gene sequencing, QTL mapping, and marker-assisted
breading (Sing and Sengar 2014). We should develop salt
tolerant genotypes of rice such as Oryza rufipogon and
Oryza nivara. We should develop salt tolerant genotypes of
rice such as O. rufipogon and O. nivara which have essential
genetic characteristics. Genotypic development of these
lines with O. sativa and modification of introgression lines
(ILs) showed yield improvement in rice under salt stress
(Brar and Khush 2002; Swamy and Sarla 2008; Fu and Xue
2010; Pushpalatha et al. 2013).
QTL mapping can select good salt tolerant genotype
which is not influenced by salt under field condition (Rich-
ards et al. 1996). Many salt resistance genes can be found
through this approach (Flowers et al. 2004). QTL mapping
assists researchers to sort out salt or other abiotic stresses
2366 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
tolerant, resistance genes and transformation. For exam-
ple, QTL linked with salt tolerance at germination stage in
Arabidopsis and barley with vegetative growth (Gregario
et al. 2002).
In molecular biology, DNA marker development plays
a crucial role to identify QTLs. Using these markers in
breading improves efficiency of particular gene selection.
High density DNA microsatellite marker, such as amplified
fragment length polymorphisms (AFLP), restriction fragment
length polymorphisms (RFLP), and the latest marker selec-
tion will help researchers to sort out traits of interest in crop
salt tolerance (Sing and Sengar 2014).
7.2. Gene identification, sequencing, micro array
analysis, and plant transformation
Many salt tolerant genes have been discovered by using
traditional breeding techniques, such as subtractive hy-
bridization, differential hybridization, and through genetic
information from model organism. Furthermore, protein
crystallography, proteomic study has enabled researchers
to explore the protein structure and function for salt tolerant
genes. After gene identification, many latest techniques
for foreign gene transformation to desired plant can help to
improve rice crop production. Such as PEG mediated gene
transfer, electroporation, partial or micro projectile bombard-
ment, microinjection, and agrobacterium-mediated gene
transfer. These techniques are available for many crops
including rice (Grover et al. 2003; Sing and Sengar 2014).
7.3. Agronomic strategies to reduce salt stress in rice
Salt stress develops due to accumulation of salt through
irrigated water and soil chemical properties. Under salt
stress condition due to Na+ and Cl– ion toxicity, the ratios of
Na+/K+, Na+/Ca2+, Ca2+/Mg2+, and Cl–/NO3– increased in soil.
This ion imbalance decreased plant growth, yield, and effect
physiological and metabolic components in plant (Grattan
and Grieve 1999; Hu and Schmidhalter 2005). We can
manage plant growth (rice) by adopting some agronomic
strategies (water management and nutrient management)
to improve soil health, plant growth, and input use efficiency
(IUE) under salt stress (USSLS 1954).
7.4. Reclamation of salt affected soils by water
management
Reduction of crop yield depends on climate conditions, crop
growth, and concentration of salts in soil and irrigated water.
The method used to remove soluble salts from root zone
is called reclamation of salt affected soils. Irrigated water
with high EC, pH value, sodium adsorption ratio (SAR), and
residual sodium corbonate (RSC) also cause of increased
salt stress and reduction of plant growth (Murtaza et al.
2006). A wise water management strategy is necessary to
cope with salt stress issue for better agriculture production.
Application of excessive amount of good quality irrigated wa-
ter for drainage and leaching of salt concentration from root
zone in rice is very important for survival of crop and grain
yield (Mitchell 2000; Ezeaku et al. 2015). Standing water
in lowland rice along the season is a best option to remove
salt from root zone through leaching or dilution (Ezeaku et al.
2015). It is reported that if underground water is brackish,
we can replace with canal water for irrigation in rice crop.
Addition of gypsum is the best option for usage of unfit
underground water because it increased 25 to 294% rice
yield and 182% wheat production than control (Zaka 2009).
ECe, pH value, and SAR of soil (at 0–15 and 15–30 cm
depths) decreased with application of canal water with 100%
gypsum requirement (GR) as compared to saline water with
100% GR in rice (Mehdi 2013). The availability of canal
water is the best option to leach down salt from the root
zone. If availability of fit water is less than 25%, gypsum
amendment with unfit water is the best option. Canal water
should use irrigation scheduling and never mix with brackish
tube well or underground water (Zaka 2009).
7.5. Reclamation of salt affected soils by nutrients
management
Under salt stress, nutrients imbalance was occured by tox-
icity of Na+ and Cl– and deficiency of N, P, K+, Ca2+, S, Mn,
and Zn2+ in various crops including rice (Dara 2004; Pandeyl
2013; Ezeaku et al. 2015). For nutrient management in salt
stressed soils, the common method is application of gypsum
(CaSO4·2H2O) as calcium (Ca2+) source for reclamation of
Na+ toxic soils and to improve soil water infiltration. Applica-
tion of 100% gypsum, pyrite, combination of gypsum+farm
yard manure (FYM)+H2SO4, CaSO4·2H2O+FYM+chiseling,
and humic acid (HA), have improved soil characteristics,
plant biomass, and grain productivity in rice, wheat, and
millet (Hussain 2001; Manzoor 2014; Ezeaku et al. 2015).
Pandeyl (2013) and Dash (2015) studied that application of
N, P, K, S, Zn, B, and Mn separately or with different combi-
nation increased rice total above ground biomass and grain
production under salt affected soils. Humic acid improve
nutrients availability by chelating with unavailable nutrients
(P, K, Ca, Fe, Zn, Cu) and buffered pH value, and enhanced
soil microbial, enzymatic and physiological activities, and
plant growth under salt stress (Barron and Willson 1981;
Julie and Bugbee 2006; Khattak et al. 2013; Manzoor et al.
2014). Combined effect of HA with gypsum (24 and 48 kg
ha–1) in rice was higher than alone effect of HA on EC and
SAR due to its chelating effect with other nutrients under salt
 Sajid Hussain et al. Journal of Integrative Agriculture 2017, 16(11): 2357–2374
stress (Shaaban et al. 2013). We can also use by-product
of sugarcane (pressmud), green manure, poultry manure,
and Sesbania as cover crop for amendment of soil to reduce
the effect of salt stress which is source of macro and micro
nutrients specially Zn and S in rice crop (Ismail 2009).
We can also use some other agronomic practices to
reduce the effect of salt stress on plant such as periodic
use of fresh water, sub soiling (50+5 cm crosswise and 20–
150 cm apart, furrows), deep tillage, sanding, and application
of organic and inorganic fertilizers, and adopting rice wheat
crop rotation (Mclean 1982; Manzoor et al. 2014; Ezeaku
et al. 2015).
7.6. Role of 1-methylecyclopropene (1-MCP) as an
ethylene inhibitor
The discovery of 1-MCP (a cyclopropane member) as a
high affinity binder to ethylene receptor is a significant key
innovation in monitoring ethylene action (Blankenship and
Dole 2003; Yang et al. 2007). It is active even at lower con-
centrations than ethylene and holds 10 times higher affinity
to occupy ethylene receptors (Blankenship and Dole 2003).
1-MCP blocks protein production and enzymes released for
ethylene biosynthesis (Su and Finlayson 2012).
1-MCP availability has provided a great opportunity for
researchers to investigate ethylene actions during senes-
cence and maturity. Normally, ethylene in rice plant binds to
the receptor site under stress. 1-MCP has ability to compete
with ethylene for binding site in the receptor. Generally,
ethylene acts as key and receptor as a lock. In this way,
ethylene cannot bind to the receptor site, consequently
ethylene function is inhibited (Blankenship et al. 2001).
Many researchers reported that 1-MCP inhibited major
biosynthetic enzyme, such as ACC synthase, ACC oxidase,
and linked mRNA accumulation enzymes (Dong et al. 2001;
Owino et al. 2002). The physiological and molecular re-
sponses of 1-MCP involved the positive ACO, ACS, cell wall
transforming enzyme activity, and GhCel1 gene expression
in plant leaf abscission region, and reduced breaking tissue
due to ethylene reduction (Mishra et al. 2008a, b). Further,
1-MCP application enhanced photosystem II quantum effi-
ciency, greater membrane integrity and delayed senescence
(Chen et al. 2014).
Various researchers have exploited use of 1-MCP on
agronomic crops. The positive effects on cotton and wheat
yield under abiotic stresses have been reported (Kawakami
et al. 2006; Hays et al. 2007). In another study, 1-MCP
played positive role on rice biomass and grain productivity.
1-MCP can play variety of special effects on ethylene
production, respiration, chlorophyll contents, and sugar
contents in plants (Blankenship and Dole 2003). All these
studies give us a strong clue that 1-MCP has potential to
2367
inhibit ethylene biosynthesis in plants under salt stress.
This review will give new directions to scientist that 1-MCP
is useful inhibitor for rice plants under salt stress.
8. Conclusion and suggestions
Impact mechanism of salt stress on rice photosynthetic
performance and development of superior and inferior
spikelets has not been clear. This review paper not only
give knowledge about effect of salt stress on rice plant
growth and development but also give us information about
the roles PGRs and management strategies to improve rice
grain yield. Many studies reported that 1-MCP is a good
ethylene inhibitor in fruits and other horticultural crops, but
effects of 1-MCP on superior and inferior spikelets devel-
opment and physiological mechanism of rice under salt
stress is not clear. So this review paper will also give us
new direction about the effects of 1-MCP on rice and growth
development mechanism under salt stress. Therefore, im-
proving tolerance through breading or cultivation techniques
such as use of 1-MCP as ethylene inhibitor is crucial to rice
growth and yield in the regions that experience salt stress.
Scientist should focus on the biotic as well as abiotic factors
responsible for low rice yield due to poor development of
superior and inferior spikelets. Future research activities
should be aimed at identification and breeding of tolerance
germplasms exploiting the variation in both genotypic and
morphological characteristics.
Acknowledgements
This work was supported by the Natural Science Foun-
dation of Zhejiang Province, China (LY13C130007), the
National Key Research and Development Program of
China (2016YFD0200801), and the Basic Research Foun-
dation of National Commonweal Research Institute, China
(2014RG004-5).
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