Abiotic stresses in rice production: Impacts and management

Abstract

Rice, a key staple food crop in the world and India, offers food and nutrition security to millions of the global population. Abiotic (water, soil, atmospheric) stresses affect yield and quality of rice. This necessitates stress-resilient rice production technologies sufficiently fortified by novel stress mitigation and adaptation strategies. Recent crop improvement strategy has partially managed to resolve the challenges presented by abiotic stresses such as high temperature, drought, salinity, alkalinity, waterlogging and mineral deficiency. The complication and multiplicity of abiotic stresses necessitate the use of extensive, integrative and multidisciplinary techniques to achieve resilience. Crop improvement, along with the agronomic interventions, is essential to stabilise the productivity and profitability of rice production. This article gives an overview of the potential impacts of abiotic stress on rice and suggests the adaptation and mitigation strategies.

Full text

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Abiotic stresses in rice production: Impacts and management
Himanshu Pathak1*, Mahesh Kumar1, Kutubuddin A Molla2* and Koushik Chakraborty2
1ICAR-National Institute of Abiotic Stress Management, Baramati, Maharashtra, India
2ICAR-National Rice Research Institute, Cuttack, Odisha, India
*Corresponding Author e-mail: hpathak.iari@gmail.com; Kutubuddin.Molla@icar.gov.in
ABSTRACT
Rice, a key staple food crop in the world and India, offers food and nutrition security to millions of the global
population. Abiotic (water, soil, atmospheric) stresses affect yield and quality of rice. This necessitates stress-
resilient rice production technologies sufficiently fortified by novel stress mitigation and adaptation strategies.
Recent crop improvement strategy has partially managed to resolve the challenges presented by abiotic stresses
such as high temperature, drought, salinity, alkalinity, waterlogging and mineral deficiency. The complication
and multiplicity of abiotic stresses necessitate the use of extensive, integrative and multi-disciplinary techniques
to achieve resilience. Crop improvement, along with the agronomic interventions, is essential to stabilise the
productivity and profitability of rice production. This article gives an overview of the potential impacts of
abiotic stress on rice and suggests the adaptation and mitigation strategies.
Key words: Submergence, drought, salinity, climate change, crop residue burning, management strategies
Received : 5 April 2021 Accepted: 15 April 2021 Published : 22 April 2021
Oryza Vol. 58 (Special Issue) 2021 (103-125)
DOI https://doi.org/10.35709/ory.2021.58.spl.4
1. INTRODUCTION
Rice (Oryza sativa L.) is the world's most important
cereal crop and is a staple food for about half of the
global population. Rice fields cover around 155 million
hectares (Mha) in a wide range of climatic conditions
spanning from 44oN in North Korea to 35oS in Australia.
It is cultivated from 6 feet below sea level (such as in
Kerala, India) to 2700 feet above sea level in the
Himalayas. In India, the crop is grown under varying
climatic and soil conditions under diverse ecologies
spread over about 43 Mha. Rice is cultivated round the
year in one or the other parts of the country.
Significant achievements in rice production
have been made in the last few decades. Production of
rice has increased about six times since 1950-51 and
made India self-reliant in rice (Fig. 1). The source of
growth was increase in yield. Population growth is the
major driving force for increasing rice demand in India.
In addition, the low-income segment of the population
will also demand more rice with increase in income. It
is estimated that about 130 and 140 Mt of rice would
be required by 2030 and 2050, respectively in India. In
addition, India is exporting about 10 Mt of rice per year,
which earns valuable foreign exchange for the country.
This increased production has to necessarily come from
increased productivity rather than increase in area under
rice and that too under deteriorating soil, water and
other natural resources. Therefore, to sustain present
food self-sufficiency and to meet future demand of food
and export, the production has to increase by about 1.5
Mt yr-1 and the productivity to 3.25 t ha-1 by 2050 from
the current level of 2.56 t ha-1 i.e., an increase of about
30% (Pathak et al., 2018).
In the backdrop of all these successes, rice
farmers and researchers however, face threats of
climate change, temperature fluctuations, low water
availability, poor soil quality and low nutrient availability
(Fig. 2). These abiotic stresses cause significant
problems by reducing crop growth and productivity of
rice (Table 1). Although active suppression of growth
is a strategy helpful for maximizing plant survival in

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Abiotic Stresses in Rice Pathak et al.
stressed condition, it is often negatively impact crop
productivity (Zhang et al., 2020). The challenge is to
combine productivity and profitability enhancement of
rice while improving the climate resilience and
excellence of the environment on which production
depends. In this article, we assess the impacts of major
abiotic stresses on rice production and productivity and
suggest the potential management options.
2. Abiotic Stresses Affecting Rice Production
2.1. Water Stress
About 76% of world's rice is produced from 55% of
the world's irrigated rice. Rice is cultivated in a rainfed
(both upland or lowland) environment, which contributes
about 25% to the total rice production from 45% rice-
growing area. The productivity of water for rice is
significantly less. It is considered to be drought-
susceptible crop due to its shallow root system, quick
stomatal closure and leaf senescence during mild water
stress (Bernier, 2007). Rice takes about two times as
much water as wheat or maize. Unlike other crops,
even above field capacity rice suffers from water stress.
Thus, adequate water supply is crucial to better growth
and high rice yields.
In many parts of South Asia, the sustainability
of irrigated rice production systems is already being
affected by lower water availability and poor use
efficiency, increased competition from the domestic and
industrial sectors, and rising costs. For instance, rice
cultivation in the upper transect of the Indo-Gangetic
plains (IGP) resulted in declining water tables and also
the water quality. A steadily declining water level is
also a primary concern for potential productivity growth
in many northern and southern states. Waterlogging in
some areas is the other side of the water issue. In some
districts of Haryana, the water level rises to 0.14 to 1.0
m per year and over 0.4 Mha of land has a water table
within 3 meters from the surface. In addition to water
shortage, growing demand for mechanisation,
urbanisation and cash cropland lead to a decline in the
rice field.
Drought is a recurrent phenomenon of rice
farming, particularly in rainfed areas. India has an
annual rechargeable groundwater capacity of around
Fig. 1. Trends in area, production and productivity of rice in India.

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430 billion cubic metres, and about 30% of its potential
has been tapped for irrigation and domestic use so far.
However, some areas of India have utilised over 90%
of their groundwater resources. The groundwater table
is depleting in Punjab, Haryana, parts of Rajasthan,
Gujarat, the western part of Uttar Pradesh and in the
Deccan states. India's limited water resources thus are
stressed while demands from various sectors are
increasing (Pathak and Ayyappan, 2020).
Increased water use efficiency (WUE) must
reduce water use in rice production through decreasing
losses due to seepage, percolation, and evaporation,
laser land levelling, crack ploughing to reduce bypass
flow and bund maintenance. Management options to
improve the productive use of rainwater include crop
scheduling, sustainable cropping, and development of
small ponds that can be used for on-farm water
harvesting reservoirs. Various crop and water
management systems, such as water-saving irrigation
techniques, periodic soil drying, growing rice with
reduced or no-tillage on either flat or raised beds, and
moving away from regularly flooded irrigation
(anaerobic) to partially or fully aerobic rice, can
considerably improve water use efficiency in rice
(Pathak et al., 2019).
Approximately 53% of rice harvesting area is
expected to experience the effect of climate variability
on yield at a rate of about 0.1 tons/ha/year and about
32% of rice yield variability is explained by year-to-
year climate variability (Ray et al., 2015). With water
resources diminishing worldwide for agriculture, there
is a need to boost the drought adaptation of rice for
food security.
A complex network of responses that combines
different altered morphological, biochemical and
molecular characteristics is involved in water stress
tolerance. Water stress affects diverse plant processes
and induces numerous plant reactions that help them to
acclimatise. Water deficit affects photosynthesis,
stomatal conductance, transpiration rate, water use
efficiency, PSII activity, relative water content (RWC),
membrane stability index (MSI) etc. in rice plant. These
parameters are reduced with the plant experiences to
soil moisture stress. Water is required for cell turgor
and cell expansion, two crucial physiological
characteristics for plant growth. Loss of grain yield due
to water stress also occurs possibly through the curbing
of grain filling period (Shahryari et al., 2008), disrupting
leaf gas exchange, reduced phloem loading and
translocation of assimilates.
Further, morphological characteristics such as
restricted germination, number of tillers, plant height,
biomass production, specific root and shoot traits could
also be influenced by water stress. Plants also undergo
metabolic shifts like aggregation of osmo-protectant
such as proline, carbohydrates, antioxidants and
polyamines. Innumerable molecular fine tuning including
altered expression of genes encoding transcription
factors and defence-related proteins occur in response
to water stress, ultimately affecting the yield of rice.
Genetic manipulation of rice regulatory and functional
elements has improved crop productivity. Several
transcription factors (TFs) in rice have recently been
documented, which are responsible for moisture stress
tolerance and enhanced yield under stressful conditions.
Water stress at vegetative and reproductive
stage, can affect flower initiation, reduce spikelet
fertility and hamper grain filling, which leads to lower
grain weight and ultimately poor paddy yield (Botwright
Acuña et al., 2008). Reduced grain size, grain weight,
and seed setting rate are typical features of rice under
water stress. Reduced WUE, altered plant water
relations, reduced sucrose and starch synthesis related
enzymes activities and condensed assimilate
partitioning, lead to a decline in plant growth and
productivity in response to water stress in rice. The
extent of the loss of grain yield depends on the period
and level of the drought stress and the stage of crop
growth (Gana, 2011; Kumar et al., 2014).
Improving tolerance to drought in rice is a
challenging task because of its complex and dynamic
nature. A detailed understanding of the various morpho-
physiological characteristics that regulate the production
of rice in conditions of water stress is a prerequisite
for facilitating the development of tolerant varieties that
may survive and provide good yield under drought
conditions. The ideal trait is the minimal loss of grain
yield during the drought.
2.2 Excess Water Stress
Rice cultivation in the rainfed agro-ecology faces multi-
faceted challenges. Erratic rainfall pattern may cause
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water deficit as well as excess water stress, both of
which leads to significant damage to crop growth and
finally yield (Sarkar et al., 2019). With the changing
climatic conditions, there could be even more uneven
distribution of rainfall, although the expected mean
annual precipitation may remain the same (Kumar et
al., 2011). Such erratic rainfall pattern makes the rice
crop vulnerable to excess water stress. If it happens
during germination stage in direct seeded rice, it leads
to germination stage oxygen deficiency (GSOD) stress
(Ray et al., 2016). Occurrence of excess water in the
form of flash flood or stagnant flooding (may be for
more than a month) leads to complete and partial
submergence stress depending upon the crop growth
stage at which it occurs. Submergence stress refers to
a condition, where the entire plant canopy (in case of
complete submergence) remains under water for a
period of a few days to couple of weeks. Under
submergence, plants face a number of external
challenges simultaneously or sequentially, which results
in multiple internal stresses, affecting plant growth and
survival. Submergence substantially reduces the gas
diffusion rate in the leaf tissue, restricting oxygen uptake
and forcing carbon inefficient carbohydrate metabolism
to occure via anaerobic route (Panda et al. 2016).
Flooding with turbid water often aggravate the problems
associated with submergence stress. Das et al. (2009)
reported that survival of rice plants was significantly
reduced when submergence stress was imposed with
turbid water as compared to fresh water. Reduced light
availability in this case restricted underwater
photosynthesis and gas exchange, which possibly has
impeding effects on transpiration and, absorption and
transport of nutrients (Colmer and Pedersen, 2008).
Presence of leaf gas film is another important
morpho-physiological factors contributing to
submergence tolerance ability of rice. Raskin and
Kende (1983) first reported that among the major
cereals, rice possesses thickest layer of leaf gas film
upon its immersion under water. Later, it was found
that presence of leaf gas film facilitates underwater
photosynthesis and respiration and basipetal movement
of O2 from shoot to root (Pedersen et al., 2009; Winkel
et al., 2011). Recently, Chakraborty et al. (2021)
reported that thickness of leaf gas film is influenced by
presence of SUB1 QTL in the genetic background of
rice. In nature, there are few rice genotypes, which
possess the inherent capacity to tolerate complete
submergence of about two weeks. These genotypes
can tolerate complete submergence usually by exhibiting
very limited elongation during submergence, a strategy
known as quiescence or low-oxygen syndrome (Bailey-
Serres, 2012; Fukao, 2013). One such genotype is
FR13A, in which detailed functional characterization
revealed the molecular mechanism of true
submergence tolerance in rice (Xu et al. 2006).
Discovery of SUB1 QTL and subsequent identification
of SUB1A-1 gene in this QTL region, helped in
understanding the ethylene mediated regulatory
mechanism of submergence tolerance in rice (Jackson
2008; Tamang and Fukao, 2015). An active SUB1A-1
locus located in chromosome 9 of a very specific rice
group, reported to encode a variable cluster of 2-3
tandem repeated group of protein which suppresses
the action of ethylene by affecting ERF-VII (Ethylene
Responsive Factors-VII) restricting stem elongation and
growth under submergence (Xu et al., 2006; Mackill et
al., 2012). In general, most of the rice accessions were
reported to have SUB1B and SUB1C genes in the SUB1
QTL region, while only a few of the indica and aus
ecotype accessions possess active form SUB1A gene
which reported to contribute as much as 70% of
submergence tolerance ability in rice (Singh et al.,
2010).
Besides, complete submergence which lasts for
about one to two weeks, prolonged stagnation of water
resulting in partial immersion of plant canopy is another
important abiotic stress in rice, particularly in lowland
ecologies. Under such circumstances, no other cereals
besides rice can survive and produce. This unique
ability in rice is attributed to its ability to elongate rapidly
with the onset of water stagnation. Under both the
situations, ethylene responsive factors genes control the
elongation but in opposite direction i.e. quiescence and
elongation (expansin) (Hattori et al., 2009). Presence
of genes like Snorkel 1 (SK1) and Snorkel 2 (SK2) in
the genomic background allow rice to elongate fast when
comes in contact to rising water level, whereas
Submergence1A-1 (SUB1A-1) restricts internode
elongation and minimal growth for adaptation to water
stagnation and flash floods conditions, respectively. Both
SKs genes and Sub1A encode ethylene-responsive
factor, a specific group of transcription factor related
to gibberellin biosynthesis or signal transduction.
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Several submergence sensitive mega rice varieties,
being converted to submergence tolerant types through
introgression of SUB1A-1 gene through marker assisted
back crossing and released for commercial cultivation
in different submergence prone area of Asia and Africa
(Singh et al., 2016; Rupasinghe, 2016; Dar et al., 2017).
In case of direct seeded rice, heavy rainfall
just after sowing creates severe hypoxic conditions (3%
oxygen level) during germination and seedling
establishment stage. Some of the rice genotypes have
unique ability to germinate underwater and extend its
coleoptile even under complete anoxic condition, which
is referred as anaerobic germination process
(Magneschi and Perata, 2009). Usually, the genotypes
having high anaerobic germination potential (AGP) are
able to elongate its coleoptile at a rate of 1 mm per
hour to come out of water by rapid elongation of basal
cells of emerging embryo (Narsai et al., 2015).
However, the AGP varies greatly from one genotype
to other, which ultimately determine the GSOD
tolerance in rice. Interestingly, the genes and QTLs
identified for complete submergence at vegetative stage
do not work during AG process, rather in many cases
like carbohydrate metabolism, ethylene production and
signalling a completely opposite mechanism works in
GSOD tolerant rice genotypes (Vijayan et al., 2018).
2.3. Soil Stress
Soil stress may be classified as chemical and physical.
Although these two are related, we focus here on the
chemical stress of soil that significantly influence crop
growth and yield. Unfavourable pH, presence of toxic
chemical, and nutrient imbalances cause chemical stress
(Tattar, 2012). Sodium, an integral constituent of our
earth crust is naturally present in all soil types. At lower
concentration, Na+ may promote growth in rice but
eventually it becomes phyto-toxic when present in high
concentration in the growing medium (Flowers and
Colmer 2008). Although both Na+ and K+ have highly
similar ionic as well as physicochemical properties, but
unlike Na+, K+ act as an essential element and play
important part in cellular integrity, enzyme functions and
growth (Schachtman and Liu, 1999). The basic
physiological processes viz., maintenance of membrane
potentials, stomatal opening and closing, pollen tube
growth which are particularly K+-dependent are
interrupted due to hindrances in absorption, uptake and
in-planta transport of K+ under high influence of Na+
(Britto and Kronzucker, 2008).
The salinity is a prevalent problem in dryland
where rainfall is very poor, and evaporation is too high.
The intrusion of sea-water can also result in salinity in
coastal areas. Twenty-three per cent of the world's
total cultivated land is considered saline, while 37 per
cent are sodic. At the same time, salinity and
waterlogging are estimated to affect half of the world's
irrigated soil. Salinity is considered to be the second
important abiotic stress in rice production after the
drought (Gregorio, 1997) although, the cereal that has
been suggested as a desalinisation crop. In their
response to salinity, cereal crops vary, with barley being
the most resistant and rice being the most sensitive
(Munns and Tester, 2008). Having a threshold salinity
level of only 3 dS m-1, there are significantly different
salt-specific response observed in rice genotypes.
Studies across difference species of Oryza suggested
that except halophytic species of O. coarctata, O.
sativa seems to be most tolerant to salt stress (Menguer
et al. 2017). Among the two predominant ecotypes of
cultivated Asian rice, indica ecotype found to possess
greater salt-tolerance ability than japonica. Tolerant
indica cultivars are reported to be better Na+ excluders
with high K+ uptake and tissue K+-retention ability for
which they can maintain a low Na+/K+ ratio in the upper
plant parts (Chakraborty et al., 2019).
Interestingly, rice crop is more vulnerable to
salinity stress at early seedling and reproductive stages,
while it was reported to be relatively tolerant during
germination, active tillering and maturity stages (Zeng
et al., 2000). Exposure to salt stress at the reproductive
stage, reduces grain yield significantly more than
seedling stage (Negrao et al., 2011; Mohammadi et al.,
2014). High seedling vigour, salt exclusion at the root
level, ion compartmentation in structural and older
tissues, high tissue tolerance, receptive stomata that
close within minutes of exposure to salt stress but
partially reopen after a time of acclimatisation, and up-
regulation of antioxidant systems are key features for
improved performance under salinity. Tolerant
genotypes often tend to exclude salt from flag leaves
and panicles (Yeo and Flowers, 1986; Chakraborty et
al., 2019). Several pieces of research indicate that the
physiological and molecular mechanism of salt tolerance
during the early seedling stage and reproductive stage
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differs considerably in rice (Ismail and Horie, 2017;
Mohammadi et al. (2014).
In general, salinity stress poses two-
dimensional threat to the plants: osmotic and ionic
stresses. The genetic basis of tolerance to ionic stress
is relatively better understood than to osmotic stress
(Roy et al., 2014). Initially osmotic stress causes more
harm, but with increased time of exposure more salt is
absorbed by the plant and ionic stress plays the key
role. To achieve salt-tolerance, adaptation to both
osmotic and ionic stresses are of prime importance
(Chakraborty et al., 2018). Salt stress mediated
reactions in plant is cumulative and increases with time.
So, restricting the movements of phyto-toxic ions such
as Na+ or Cl- to growing meristematic tissues and young
photosynthetic organs are crucial for survival. Tolerant
rice cultivars like Pokkali or some of the mangrove
region rice landraces either absorbs low levels of Na+
or restricts the movement of Na+ in comparison to K+
and thereby maintains low Na+:K+ ratio in shoot / leaf
and protects the vital tissues (Singh and Sarkar, 2014;
Chakraborty et al., 2020). High Na+ in the cell
cytoplasm impairs several physiological and biochemical
courses of action (Bendaly et al., 2016), which restricts
plant growth.
Higher temperature aggravates the situation by
excessive salt accumulation on the surface due to
capillary action, which is remarkably difficult to leave
below the rooting zone. Hence, leaching is necessary
for the sustainability of crop production. However,
unless there is adequate drainage beyond or below the
root zone, this practice leads to rising water table and
waterlogged conditions. Indeed, high water tables
increase salt transport to the root zone due to capillary
rise from the water table.
In saline conditions, water-use efficiency
potentially could lower the uptake of salt by the plants.
Under these environmental conditions, partial stomatal
closure could be an ideal salt tolerance mechanism
(Rozema et al., 1991). Salt can also cause an imbalance
of nutrients in rice plants, and the root is regarded as a
crossroad between nutrients and plants under salt
stress. The high concentration of Na+ has an inverse
effect on the uptake of potassium (K+) ion, which is a
significant plant nutrient for plant growth and yield (Jung
et al., 2009). Similarly, reduction in plant nitrogen uptake
was also shown under high salinity (Abdelgadir et al.,
2005).
Besides, salt stress affects P, K+, Ca2+, Fe, Zn
and Mn uptake but has interactive effects on N and
Mg (Jung et al., 2009; Garcia et al., 2010). Boron (B),
silicon (Si), zinc (Zn) availability decreased under high
salt in plants (Wimmer et al., 2001; Currie and Perry,
2007). Although, increased toxicity of cadmium (Cd)
has also been reported in response to high salinity in
paddy (Amanullah, 2016).
In addition to salinity discussed above, soil
sodicity is another kind of stress in the soil, owing to
the high concentration of salt, the low rate of penetration
and weak hydraulic conductivity. This forces water
stagnation on the soil surface, which in turn allows no
crop to thrive but rice. Hence, it is recommended as
the first crop to be grown during reclamation of these
types of soil. Unlike other crops, many rice germplasms
can survive and adapt to salt affected areas and as a
result rice became favoured crop in these unfavourable
environments (Singh et al., 2015).
2.4. Temperature stress
Rice is highly sensitive to both cold and heat during the
reproductive stage. Like other plants, rice has an
optimum range of temperature for growth and
reproduction. Rice yield is negatively impacted if the
temperature reaches above or below the optimum range.
Cold stress, comprising freezing and chilling stress,
causes pollen sterility due to hypertrophy of tapetal cells
and resultant nutrient imbalances (Arshad et al., 2017).
Further, it also increases floret sterility, grain abortion,
and lowers yield. A temperature below 13°C for 15
days has been reported to cause 19-29% yield loss in
rice (Ghadirnezhad and Fallah, 2014).
On the other hand, heat stress results in poor
anther dehiscence, causing reduced pollen dispersal and
hampered pollination. Among other impacts of heat
stress, reduced pollen viability and pollen tube growth,
increased spikelet sterility, delayed heading and
increased chalkiness are notable (Arshad et al., 2017).
Heat stress is also known to hamper starch biosynthesis
and subsequent starch accumulation in developing grain
(Ito et al., 2009). At a mean day temperature of more
than 33°C at the heading stage has been shown to
reduce rice yield by 24-27% (Cao et al. 2009) (Table
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1). In the scenarios of IPCC projected climate change,
asian rice production would likely decline by 3.8% by
the 21st century end (Murdiyarso, 2000).
2.5. Nutrient Stress
Soil erosion is caused by improper agricultural practices,
deforestation and overgrazing. Physical and chemical
problems in the soil are considered as significant
restrictions on wheat and rice crops, such as excessive
tillage, low soil carbon, K and Zn as well as low nutrient
use efficiency (NUE). Asia, which is undergoing high
soil depletion, produces 90 percent of the rice
worldwide. Rice yields are also decreasing as a result
of poor nutrient balance and poor soil and crop
management. A negative balance of nutrients and
inadequate management of soil and plant have restricted
the rice production.
Indian soils are low in organic carbon. Along
with the intensification of production, emerging nutrient
deficiencies are increasingly reported. Annually 8-10
Mt of NPK is mined from the soil, and 93, 91, 51 and
43% soils are ranked low in N, P, K and Zn, respectively
(Minhas et al., 2017). Rice plants are able to use about
30-50% of the applied N fertiliser, while more than 50%
is lost from the soil-plant system through leaching,
denitrification and volatilisation. Though farmers apply
some of the macronutrients like N, P and K, they usually
neglect the application of micronutrients. Even the N,
P, K fertilisers are not applied proportionately. In the
long run, this may lead to an imbalance in soil and plant
nutrition resulting in yield decline. It has been reported
that during 1950s, there was only one nutrient deficiency,
which has increased to eight (N, Fe, S, Mn, B, P, Zn, K,
Mo) during 1990s. Moreover, the soil quality is
deteriorating from the loss of organic carbon, erosion,
soil compaction, salinisation, introgression of heavy
metal from industries and pesticides, and also due to
other anthropogenic activities. Uptake of nutrients like
Table 1. Effect of different abiotic stresses on rice yield.
Stress Growth stage Details of abiotic stress Decrease in yield (%) Reference
Low Flowering <13 oC for 15 days 19-29 Ghadirnezhad and Fallah
temperature (2014)
Reproductive and <13 oC for 15 days 9.2-26 Lee (2001)
grain filling
High Heading stage Mean daytime temperature >33 oC 24-27 Cao et al. (2009)
temperature Heading stage Mean daytime temperature >30 oC 20-30 Zheng et al. (2005)
for 15 days
Early grain filling Mean daytime temperature above 6.6-13.0 Cao et al. (2009)
33oC
Growing season 1oC increase in minimum temperature 10 Peng et al. (2004)
Drought Flowering Withholding water at 60 DAT 23-24 Yang et al. (2019)
( -30±5 kPa)
Vegetative Withholding water for 24 days from 50.6 Guan et al. (2010)
21 DAT
Vegetative/ Drying soil (dried beyond -20 kPa) 22.6 Carrijo et al. (2017)
reproductive
Flowering Sever stress >70 Shamsudin et al. (2016)
Reproductive stress Moderate to severe stress (Withholding 51-60 Swamy et al. (2017)
water-70 kPa at 0.2-m depth)
Reproductive stress Moderate stress 51-57 Dixit et al. (2014)
Salinity Over growing period 1.0, 3.9, 6.5 dS m-1 > 50 % Zeng and Shannon (2000)
Over growing period 7.4 dS m-1 50% Grattan (2002)
Reproductive stage 8.0 dS m-1 >60% Chattopadhyay et al. (2018)
Reproductive stage 8.0 dS m-1 50-75% Chakraborty et al. (2019)
Submergence Seedling stage Complete submergence with 80 cm 30-60% Bhaduri et al. (2020)
(Complete and of water for 12 days
partial) Vegetative to 50 cm of water depth by gradual 30-70% Kuanar et al. (2019)
anthesis Stage incraese of 10 cm day-1 rise in
water level
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N in rice is influenced by salinity status in soil
(Abdelgadir et al., 2005). Potassium, Calcium and
Manganese are the main elements that has been
observed with lower concentrations and imbalanced
ratios in plants due to competition (Hussain et al., 2017).
Iron toxicity is frequent in Ultisols and Oxisols and in
acid sulphate soils rich in reducible Fe. Fe toxicity may
cause complex nutrient disorder and deficiencies of
other plant nutrients, particularly P, K and Zn
(Sahrawat, 2004).
2.6. Climatic Stress
The rising concentration of greenhouse gases in the
atmosphere is expected to impact the climate and the
global average air temperature, which is estimated to
increase between 1990 and 2100 by 1.4 - 5.8oC. The
varying atmospheric temperature may be low or high,
cause damage to the rice plant. Rice production leads
to global climate change (through methane and nitrous
oxide emissions) and in turn, suffers from the effects.
Climate conditions including temperature, atmospheric
CO2 precipitation and solar radiation are essential
requirements for rice production (Nath and Raju, 2018).
Higher maximum temperature decreasing spikelet
fertility and higher minimum temperature increasing
respiration in the rice. The increase in temperature,
particularly the mean minimum night-time temperature,
has harmful effects on productivity. The high
temperature might reduce crop cycle duration, increases
respiration rates, alters source-sink balance, hastens
nutrient mineralisation in soils, decreases NUE and
increases evapotranspiration (Wassmann et al., 2009a,
b). Rice is highly vulnerable to heat stress, particularly
during the reproductive stage. The overall influence of
changing climate on yield depends on both the yield
vulnerability and the magnitude of changes in the
climatic variables, which are of critical importance for
rice growth and finally, productivity (Tao et al., 2012).
An increase in the concentration of CO2 for
rice induces Rubisco and decreases photorespiration.
As a result, rising temperatures may lead to higher CO2
assimilation through photosynthesis. Furthermore, rice
is a vital sink for assimilates, and the restriction of this
carbon sink would not exploit the high CO2 due to the
insensitivity of photosynthesis. With the substantial
increase in the root/shoot ratio along with high CO2
and increased temperature, it was suggested that
alternative sinks become active recipients with reduced
grain carbon sink capability due to spikelet sterility in
response to temperature (Ziska et al., 1996). Increased
CO2 concentration and elevated temperature may also
affect different nutritional and grain quality parameters
in rice.
The timing of rice flowering in a season is
primarily controlled by temperature and photoperiod.
During the floral meristem initiation process, rice
cultivars with improved tolerance to combined heat and
drought stress will be critical to ensure the continued
adoption of water-saving technologies in future hotter
climates to complement the progress made through other
sensitive developmental stages such as flowering to
overcome the adverse impact (Jagadish et al., 2015).
A rise in atmospheric carbon dioxide has a
fertilizing effect on rice, encouraging its growth and
productivity, but the beneficial effects can be negated
as photosynthetically active radiation (PAR) is likely to
decline by 1% (Hume et al., 1990). Elevated CO2 might
influence physiological traits such as leaf water content,
relative water content (RWC), membrane stability index
(MSI), chlorophyll content, carbon assimilation rate and
Total Soluble Sugar (TSS) content, but responses of
these traits were opposite with rising in temperature
(Dwivedi et al., 2015). Recent studies, however,
suggested that the global warming implications will be
largely negative for rice production due to increased
respiration and shorter vegetative and grain-filling
periods. It is also supposed that climate change would
impact crop quality, particularly important aromatic
crops such as basmati rice. In addition to these direct
impacts on rice crop, climate change might influence
other organisms linked with rice and thus, alter the
occurrence and severity of different rice pests. It is
also important to address the challenge of rising climate-
related severe weather events, such as more recurrent
floods and droughts, as well as more frequent
hurricanes, and their impact on the development of rice.
A variety of changes in physiological and molecular
processes are triggered by low temperature stress and
often result in the formation of free radicles in the cells
of crop plant, including rice.
India is known to be one of the regions most
vulnerable to large-scale climate change. Atmospheric
temperature, which has already risen by about 1oC, is
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111
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rising consistently. Global warming reports on rice
productivity also indicate that as global temperatures
rise, the productivity of rice and other tropical crops
will decline. The amount, intensity, variability and
extreme events of rainfall (unseasonal rain, drought and
flood) are rising while the duration of rainfall is reducing
(IPCC, 2019). However, there could be some positive
impacts of climate change. As mentioned earlier,
increased CO2 level will have beneficial effects on the
yield of rice. More rain, particularly in water-deficit,
rainfed areas and increased temperature in cool,
temperate regions may have the congenial temperature
for crop growth and reduce cold injury. Various studies
indicate a decrease of up to ~12% in rice yield with
time (Naresh Kumar et al., 2019). The climate suitability
for rainfed rice is projected to decline in the range of
15-40% by the year 2050 (Singh, 2017). Rice also
contributes to climate change. Out of 417 Mt CO2e of
total greenhouse gas (GHG) emissions from Indian
agriculture, 18% is emitted from rice cultivation
(MoEFCC, 2018).
Vast tracts in the Ganges's low-lying deltas and
other major river systems in the major rice regions were
hit by tidal waves. Enterprising farmers around the
world have strengthened rice production from tidal
affected lands. They have used flooded rice systems
to reclaim land to cultivate other crops for their
livelihood. However, due to the salinity in the soil, rice
yields in tidal-affected lands are usually lower than
lowlands not impacted by tides. In low-lying areas,
flooding is the most important restriction on the
production of rice. For rainfed lowlands, irrigated and
deep-water habitats, most rice varieties can withstand
full submergence for at least six days until 50% of them
die. The mortality rate, however, is 100% when
submergence lasts for 14 days (Depledge, 2002).
2.7. Burning of Rice Residues
Crop residue burning is one of the foremost reasons
for declining productivity in India, particularly in rice-
wheat cropping system. Increased removal and burning
of the crop residue will lead to a net loss of nutrients
according to conventional fertilisation methods,
eventually leading to higher short-term nutrient costs
and a decrease of soil quality as well as productivity.
Rice straw burning could result in 70% CO2, 7% CO,
0.66% CH4 and 2.09% N2O gas emissions in the
atmosphere (Gupta et al., 2004). Rice straws and husks
are also not eco-friendly disposed of. In a recent survey,
it has been observed that in the Punjab and north-
western states of Haryana, 82% and 60% of the rice
straw produced is burnt in the field (Pathak et al., 2012).
Out of a total of 37 M tonnes of residues, approximately
20 Mt of rice and wheat residues are burned in situ
every year in Punjab (India). This burning results in
the loss of approximately 8 Mt C equivalent to a CO2
load of approximately 29 Mt per year and a loss of
approximately 1 x 105 tonnes N, plus loss of S and
depletion of beneficial soil microflora (Singh et al., 2010).
Burning rice straw is harmful to the atmosphere as it
leads to (1) The release of soot particles and smoke,
causing asthma or other respiratory problems; (2)
release of GHGs, which cause global warming; and
(3) loss of N, P, K, and S and other plant nutrients.
Crop residues are a great source of plant
nutrient and are a significant component of farm
ecosystem stability. Nearly all C and N, 25% of P, half
S and 20% of K present with straw are lost because of
burning. Retained crop residue can enrich the soil C,
N, K and P as well in the soil. However, only a few
farmers have followed in situ method of the
incorporation of rice straw as an alternative to burning
due to high incorporation costs and energy and time-
intensiveness. The heat generated by cereal straw
burning can penetrate up to 1 cm of soil and raise the
temperature to 33.8-42.2 oC, even to 15 cm of the
topsoil. Because of these, soil microbes populations in
the top 2.5 cm of soil are reduced instantly and
significantly on burning. Frequent burning may lead to
permanent loss of these microbial population by more
than 50%, but fungi appear to have recovered and also
decreased soil respiration. Residue burning increases
exchangeable NH4+-N and extractable phosphorus bi-
carbonate, but there is no nutrient build up in the profile
(Hobbs et al., 2002). Burning for longer periods
decreases total N as well as C and potentially
mineralised N to the top oil layer. Burning may also
lead to loss of soil organic matter (Gupta et al., 2004).
3. Abiotic stress management
3.1. Rice crop improvement efforts for managing
abiotic stresses
Rice genetic improvement is considered one of the top
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112
priority areas to increase yield overcoming abiotic
stresses to meet the future demand (Molla et al., 2015).
Developing varieties tolerant to multiple stresses using
stress-tolerant QTLs, genes and alleles in elite cultivars
is an efficient way of achieving higher yield and
sustainability (Fig. 2). Crop varieties such as CR-Dhan
801 and CR-Dhan 802 for rice and several for other
crops, which are tolerant to multiple stresses, i.e.,
submergence, salinity, drought, heat and pest and
diseases have been developed. Methods of conserving,
storing and enhancing water use efficiency such as
pressurised, low cost and demand-driven micro-
irrigation methods have been developed with substantial
success. Rice is the most water demanding crop;
technology has been developed to minimise the use of
water and mitigate gas emissions, for example,
alternative weathering and drying and dry direct
showing of seeds (DSR). Drought-related quantitative
trait loci (QTLs) is defined using QTL mapping. The
chromosome areas responsible for drought tolerance
are also labelled with the aid of molecular markers.
The Sahbhagi Dhan variety, released and notified in
India in 2010, showed strong performance consistently
under rain-fed direct-seeded upland and transplanted
low land conditions. New drought-tolerant rice varieties,
such as RD12, a glutinous rice and RD33, anon-
glutinous rice have been developed. Oryza glaberrima
is a wild rice plant and a valuable genetic resource for
heat stress resistant trait because of its habit of early
flowering in the morning and high transpiration with
sufficient water tolerance (Markam, 2013).
Improving crops helps to develop high-
temperature resistant cultivars. Based on crop-growing
time modulation, early or late cultivars should be
adopted. In addition, crop models were used to predict
crop yields at scales from individual fields to regions or
countries (Yang, 2010). This will replace the need for
years of costly multi-location, station and farm trials to
classify rice cultivars.
Fig. 2. Schematic showing different abiotic stresses rice plant encounters (upper panel) and their management strategies
(lower panel).
Abiotic Stresses in Rice Pathak et al.

113

Table 2. Approaches to better understand and management of abiotic stresses in rice.
Stress Approaches Details Reference
High Genomic approaches Multiple stress tolerant rice varieties Chen et al. (2020)
temperature QTLs for heat tolerance by crossing heat-tolerant and heat Shanmugavadivel et al.
-sensitive genotype (2017)
Agronomic Advance the flowering time up to 2 hours with chemicals Khan et al. (2019)
management/ Modification in plant architecture Shah et al. (2011)
practice
Application of Salicylic acid mitigates the adverse effect by enhancing the Zhang et al. (2017);
bioregulator proline, sugar, antioxidative enzymes Chandrakala et al. (2013)
Methyl jasmonate advances the flower opening time two hours Kobayasi and Atsuta (2010)
PGR combination improves heat under temperature stress Fahad et al. (2016)
Osmo-protectant: Glycine betaine protects enzymes from Mohammed and Tarpley
heat degradation (2009)
Identification of Cooler canopy temperature Zhang et al. (2015) a, b
superior traits/ Higher non-structural carbohydrate Khan et al. (2019)
genotypes Protected thermal degradation of enzymes Scafaro et al. (2012)
More HSPs production for protection Khan et al. (2019)
Better chlorophyll Fluorescence and cell membrane stability Sailaja et al. (2015)
Low Genomic approaches QTL identification through Genome wide association studies Pandit et al. (2017)
temperature QTL identification and mapping for cold tolerance at Saito et al. (2001)
reproductive stage Andaya and Mackill (2003),
Suh et al. (2010),
Shirasawa et al. (2012)
QTL identification and mapping for cold tolerance at Baruah et al. (2009)
germination stage
QTL identification and mapping for cold tolerance at Koseki et al. (2010),
seedling stage Suh et al. (2012)
Identification of Chilling stress tolerant genotypes Pradhan et al. 2016
superior genotypes
and molecular
markers
Drought Drought protective LEA proteins Xiao et al. (2007)
genes Duan and Cai (2012)
Todaka et al. (2015)
Osmo-protectant synthesis You et al. (2012)
Protein kinase Saijo et al. (2001)
Ramegowda et al. (2014)
Phytohormone/ ABA: Improves drought tolerance; Improves aquaporin Tardieu et al. (2010)
biochemicals/ functioning and osmotic regulation; improve sink strength Yang and Zhang (2010)
Nutrients Teng et al. (2014)
Application of Abscinazole-E3M improved the drought Takeuchi et al. (2016)
tolerance in rice by targeting abscisic acid 8' hydroxylase
CK: regulate grain development, increased GY under pre- Yang et al. (2001a);
and post-anthesis drought Peleg et al. (2011);
Khan et al. (2019)
Proline application (30 mM) Hanif et al. (2020)
GA: Rapid embryo development Yang et al. (2001b)
Application of potassium (K+) for improving drought Zain et al. (2014)
tolerance of rice
Exogenous application of silicon can improved the drought Chen et al. (2011)
tolerance in rice
Exogenous application of nitric oxide enhanced the performance Farooq et al. (2009)
of rice in case of drought stress
Exogenous application of glutathione (GSH) improved the Sohag et al. (2020)
drought tolerance by minimizing the oxidative damage
Continued......
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114
Agronomic Aerobic rice cultivar Parthasarathi et al. (2012)
management/
practice
Salinity Cultivar and genomic Salinity tolerant cultivars and markers for breeding Chattopadhyay et al. 2016;
approaches Chattopadhyay et al. 2018;
Chattopadhyay et al. 2014;
Markers for salt tolerance breeding Molla et al. 2015;
Ganie et al. 2019;
Application of Gypsum application reduced soil pH, EC, and increased Shaaban et al. (2013)
bioregulator/ root length and yield.
chemical Polyamines defend against salt damage by maintaining K+/Na+Roychoudhury et al. (2011)
balance, activating osmolyte levels and antioxidant enzyme
activity
Salt-tolerant PGP bacteria improves the protection mechanism Prittesh et al. (2020)
Exogenous proline Siddique et al. (2015)
Apigenin pretreatment improved growth and salinity tolerance Mekawy et al. (2018)
through Na+ transporter-encoding genes
Application of Zn and Mn increases sodium and chloride Nadeem et al. (2020)
concentrations and increased potassium (K+) concentration
Exogenous application of ABA and silicon improved salinity Gurmani et al. (2013)
tolerance of rice by reducing the transport of sodium (Na+)
Pretreatment of rice plants with hydrogen sulfide can improved Mostofa et al. (2015)
the overall drought tolerance
Exogenous application of proline improved the salinity tolerance Sane et al. 2019
in transplanted aman rice
Application of boron for improving the productivity of rice Mehmood et al. 2009
under salinity stress
Agronomic Banning of summer rice, delayed transplanting and better Chhabra (2002)
management/practice management of irrigation water
Submergence Cultivar Introgression of SUB1 QTL in popular and mega rice varieties Singh et al. (2009)
Kuanar et al. (2019)
Application of Glutathione (GSH) increase chlorophylls, carotenoids, soluble Siddiqui et al. (2021)
bioregulator/chem- proteins and proline content in rice
ical/ Nutrients Exogenous application of K+ improves submergence tolerance Gautam et al. (2016)
of rice Dwivedi et al. (2017)
Exogenous application of protocatechuic acid and vanillic Xuan and Khang (2018)
acid improves the submergence tolerance of rice
Application of N and P for improving the performance of rice Gautam et al. (2014)
under submergence was investigated at different reproductive
stages
Application of 20% more phosphorous in soil can improve Lal et al. (2018)
the productivity of submergence tolerant rice genotypes
Nitrification inhibitors reduce nitrous oxide production under Kumar et al.(2000)
submergence that beneficial for limiting methane emission from Sahrawat(2004)
submerged rice soils
Manipulating plant Manipulating the conventional spacing techniques, submergence Bhaduri et al. (2020)
spacing, Agronomic tolerance can be improved in rice
Management
Greenhouse Methane emission Direct seeding, Mid-season drainage and intermittent irrigation, Pathak et al. (2013)
gas Composting organic amendments, Supply of N, P, and K nutrients
through sulfate-containing fertilisers
Rice straw and urea (1:1) application Bhattacharyya et al. (2012)
Zero tillage Dash et al. (2017)
Short duration rice variety, switching from urea to ammonium Hasan, (2013)
sulphate, and midseason drainage
Use of methanotrophs Davamani et al. (2020)
Continued......
Abiotic Stresses in Rice Pathak et al.

115

A major QTL, Saltol, located on Pokkali
Chromosomes 1 and SKC1 (OsHKT 1:5) identified, is
responsible for salinity tolerance in rice. Breeders
attempted to combine Sub1 gene and Saltol QTL in the
same genetic background, increasing the resistance of
rice against stresses like salinity and submergence
(Mackill et al., 2010). Molecular genetic approaches
have been used to boost chilling resistance in rice to
sustain rice production in existing regions, extending it
to northern areas with lesser annual temperatures. Acid
soils have elevated levels of Al, Mn and Fe and low
levels of phosphorus. In upland soil, P was restricted
by P-fixation as the low pH of the soil and the removal
of additional P from lowland conditions. Is is also well
documented that elevated CO2 concentration, mitigates
toxicity by decreasing cell wall hemicellulose in rice.
Phosphorus uptake1 (PUP1), an important quantitative
locus derived from Kasalath shows 78.8% P-uptake
phenotypic variance and the only available QTL for
marker-assisted rice selection (Dabi and Khanna, 2018).
Genetic variation in adaptation and resistance to iron
toxicity has been used for the production of ferro-toxic
resistant cultivars. The genetic variation between the
different accessions needs to be exploited. A rare QTL,
qFRSDW11 associated with tolerance to iron and zinc
toxicity has recently been also reported (Liu et al., 2016).
3.2. Production strategies to minimize the abiotic
stress
A multi-pronged strategy with integration,
diversification, clustering, customised farm
mechanization, value addition and market access is
required for sustainable rice farming (Fig. 2). Some
approaches are presented in Table 2 for improving
abiotic stress tolerance in rice.
Use of neem-coated urea, soil health card and
leaf colour chart for enhancing fertiliser use efficiency
have been successfully utilised in India. Microbe-based
technologies for N fixation, nutrient recycling, bio-
residue management and alleviation of abiotic and biotic
stresses have also been found useful. Another significant
development is conservation agriculture to reduce the
carbon footprint of the production system, improves
productivity and enhances adaptability by modulating
soil moisture and temperature regimes. These practices
need to be scaled up with due refinements and
incentives (Pathak and Ayyappan, 2020). Mechanisation
in agriculture with renewable energy sources such as
solar-powered machinery such as water pumps,
sprayers and weeders are better alternatives to diesel-
powered machines. Such machines are economical,
eco-friendly, i.e., do not release the GHGs and other
polluting gases. Information and communication
technologies (ICTs) for short-term weather forecast
and advisories at block level could greatly help in
addressing climatic extremes and develop contingency
plans. Sustainable insurance system needs to be
developed, while the rural poor need to be informed
about taking advantage of these opportunities.
The preservation on the surface of the land
would be a possible solution to the issue of rice straw
burning. It prevents soil moisture loss, manages weeds,
regulate soil temperature to enhance crop growth and
increases soil organic matter content. Now seeds can
be planted in fields kept with residues through the
Nitrous oxide and Use of nitrification inhibitors and urease inhibitors Malla et al. (2005)
methane emissions Direct seeded rice (DSR) and Real time N management through Dash et al. (2017)
Leaf Colour Chart (LCC)
Use of 'system of rice intensification' (SRI) technique Gathorne-Hardy et al. 2016
Use of 'system of rice intensification' (SRI) technique with a Hasanah et al. (2019)
water table level of 5 cm from soil surface
Neem Coated Urea application using LCC Mohanty et al. (2018)
Nutritional Tolerant cultivar Develop P and Zn responsive cultivars Rose et al. (2013)
deficiencies Crop production Crop residue management Goswami et al. (2019);
Kumari et al. (2018)
Biochar soil amendment Liu et al. (2019),
Munda et al. (2018)
Integrated application of rice husk biochar (BC) and Munda et al. (2016)
coal fly ash (FA)
Boron fertilization for boron deficiency Rehman et al. (2018)
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116
advance and modern machines like Happy Seeder. The
production of biochar by means of pyrolysis, that is,
burning waste residue at high temperature (300-600oC)
with partial or total elimination of oxygen and its field
application as a way of moving waste biomass carbon
from a fast to a slow carbon-cycling pool in soil. When
applied to soil its highly porous nature results in increased
water retention and increases soil surface area. It
primarily interacts with soil matrix, microorganisms, and
roots of plants, helps to maintain nutrients, and initiates
a broad range of biochemical cycles. It is well
documented that an increase in pH give rise to an
increase in the population of earthworms and a
subsequent decrease in fertiliser use (Bhuvaneshwari
et al., 2019). The high organic content of crop residues
makes it similar to another substitute, the perfect raw
material for compost. The compost is rich in carbon
and organic matter comprised of 45% of total solids,
26.7% of organic matter, 15.3% of carbon and 1.36%
of total nitrogen. Added soil compost improves the
physio-chemical and biological properties of the soil and
can reduce the use of agricultural chemicals like
fertilisers and pesticides (Singh and Prabha, 2017).
Adaptation strategies for climate change
include developing cultivars tolerant to different
stresses, amending crop management practices,
improving water management, adopting new techniques
such as conservation agriculture (CA), improved pest
management, better weather predictions and crop
insurance. Advanced solutions to reduce methane
emissions could be utilising a changed water system
(mid-season runoff, alternative flooding), modified
residue management (straw sequestration), additive use
(phosphogypsum, nitrification inhibitors) and modified
land management (modified land management) (direct
seeding, reduced tillage and site-specific nutrient
management) (Pathak et al., 2015). Similarly, novel
approaches of demand-driven N supply using leaf
colour charts and site-specific N management minimise
the excessive nitrogen pool in the soil and thus reduce
nitrous oxide emissions.
Way forward
Rice productivity can be increased as, we are currently
harvesting only 50-60% of the genetic potential of most
crops. Institutions, however, need to be reshaped
towards resource conservation and climate-resilience.
Creating awareness among farmers, policymakers and
extension personnel on the impacts of technologies for
climate risk mitigation is equally important. Finally, to
make Indian agriculture climate-smart, we need smart
farmers. We need to start from the grass-root with
strong policy support for sustainable production,
processing, pricing, procurement and promotion of
climate-smart technologies.
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