Agric Res J 58 (5) : 783-788, October 2021

DOI No. 10.5958/2395-146X.2021.00111.3

COMBINED EFFECT OF HIGH TEMPERATURE AND SALINITY ON GROWTH

AND PHYSIOLOGY OF RICE (Oryza sativa L.)
Alif Ali B S, Beena R* and Stephen K
Department of Plant Physiology, College of Agriculture, Vellayani,
Kerala Agricultural University, Thiruvananthapuram- 695522, Kerala

Abstract
High temperature and salinity are the major climatic factors affecting the productivity of rice.
Combined effect of both the stresses can cause severe yield reduction in rice. Therefore, objective of
this study was to assess the combined effect of high temperature and salinity on three rice genotypes
(MO-16 - popular high yielding variety; N-22 - tolerant to high temperature and drought and NL-44 -
tolerant to high temperature). The experiment was laid out in completely randomized design with
three replications. The three genotypes of rice were grown in non-saline conditions at optimum
temperature until booting stage. At booting stage, plants were irrigated with 250mM NaCl for three
days and exposed to high temperature (38ºC) for 10 days. High temperature combined with salinity
significantly affected the physiological processes and yield parameters. Tolerant genotype, N-22
showed higher proline content, cell membrane stability index, superoxide dismutase activity, pollen
viability, spikelet fertility percentage, yield per plant and less malondialdehyde content and Na+/K+
ratio compared to other genotypes.
Keywords: High temperature, Interactive effect, Physiological traits, Rice, Salinity

R ice (Oryza sativa L.) is a widely consumed food crop,

which is grown worldwide (Beena et al., 2021). The
global population is expected to be nine billion by 2050,
combined effect of high temperature and salinity in rice.

MATERIALS AND METHODS

which demands 60–110% more rice production than
the present-day (Ray et al., 2013). However, it is very This experiment was laid out as a pot culture
difficult to combat the challenges like change in climatic study in the Department of Plant Physiology, College
factors, less availability of land area for cultivation , of Agriculture, Vellayani, Kerala Agricultural University
lack of high yielding stress tolerant varieties. Studies during the year 2017-2018. Three varieties viz., MO-
show that average temperature is expected to rise by 16- popular high yielding variety; Nagina-22- an aus
2–3◦C over the next 30– 50 years (Hatfield and Prueger, rice type tolerant to high temperature and drought; and
2015). Rice growth and development is affected above NL-44 - tolerant to high temperature were selected
a threshold temperature of 32◦C: the most critical for this study. Seeds were sown in portrays filled with
temperature was found to be 33◦C during the flowering potting mixture (coir pith compost and vermicompost
stage (Jagadish et al., 2007). High temperature is @ 2:1 ratio) and labelling was done properly. Irrigation
detrimental to most physiological processes including was provided regularly. Eighteen days after sowing,
stomatal opening, photosynthesis, growth, and grain seedlings were transplanted to pots with potting mixture
yield in rice (Beena et al., 2018). made from soil, sand and cow dung on equal volume
by volume basis. Six replications were maintained for
Salinity is the second most devastating constraint each variety as control and treatment until booting
in rice production after drought, affecting approximately stage. The experiment was laid out in completely
1 billion ha of land globally (Fageria et al., 2012). This is randomized design with two treatment levels i.e. control
equal to more than 6% of the world’s total farming area and treatment with three replications each. Five plants
(Ismail and Horie, 2017) and nearly 20% of the globally per each replication were maintained. During booting
irrigated area. Most abiotic stress studies concentrated stage, a set of three genotypes with three replicates was
only a single stress condition, whereas, under field transferred to stress condition. At booting stage, plants
conditions, a combination of stresses may occur. The were irrigated with 250mM NaCl for three days and
effects of combined heat and salinity stress have not exposed to high temperature (38ºC) for 10 days. After
been studied extensively. Hence, in the present study, a three days regular irrigation was given. Physiological
pot culture experiment was designed to understand the observations were taken ten days after stress induction
from the fully expanded third leaf and yield data
*Corresponding author : [email protected] were taken at harvest. Physiological parameters and
Date of receipt: 28.05.2021, Date of acceptance: 10.09.2021 yield traits were taken as per standard procedures;
783
Chlorophyll content (Hiscox and Israelstam, 1979); Cell reduced under high temperature stress in rice (Beena
membrane stability (Blum and Ebercon, 1981); Proline et al., 2018; Pravallika et al., 2020). Heat stress hastens
content (Bates et al., 1983); Na+/K+ ratio (Zasoski and the rate of grain filling, but reduces the duration of grain
Buraum,1977); Malondialdehyde content (Wang et al., filling, as reported in wheat ( Farooq et al., 2011),
2013); Superoxide dismutase activity (Beauchamp and rice (Beena et al., 2012) which may be due to direct
Fridovich, (1971); and pollen viability test (Baker and effects of heat stress on the source–sink relationship
Baker, 1979) were also recorded. that reduce photoassimilate supply to developing seeds
Cell membrane stability index was estimated as per (Calderini et al., 2006).
the procedure described by Blum and Ebercon (1981). The pollen viability of plants grown under the
Samples collected from both control and stress imposed combined stress conditions (Th×Sh) was significantly
plants were washed three times in deionised water to reduced with a mean of 73.2%, whereas the control
remove electrolytes adhered on the surface. Samples grown plants showed a higher pollen viability percentage
were kept in a capped vial (20ml) containing 10ml of of 93.01%. Among the genotypes, NL-44 exhibited the
deionised water and incubated in the dark for 24 hours highest pollen viability percentage (79.12%) under the
at room temperature. The conductance was measured combined stress treatment followed by N-22 (74.52%).
with a conductivity meter. After the first measurement, Pollen viability and germination are generally adversely
the vials were autoclaved for 15 minutes to kill the affected under saline stress conditions. Pollen viability
leaf tissue and release the electrolytes. After cooling, was reduced in all varieties following treatments
the second conductivity reading was taken. These compared to their corresponding controls. This may
two measurements were carried out individually for be due to the tapetum, the innermost cell layer of the
both control and stress treated plants. Cell membrane anther wall which plays a crucial role in supplying
stability index was calculated by using following formula nutrients to these microspores and in regulating their
and expressed as per cent. CMS (%) = [1-(T1/T2)/1- release. Temperature and salinity stress during tapetal
(C1/C2)] x 100 36 Where, T and C refer to the stress development (such as early degeneration, hypertrophy,
and control samples, respectively. Where, T and C or mutations in the archesporial cell) leads to aborted
refer to the stress and control samples, respectively. micro gametogenesis and male sterility (Chaudhury et
The subscripts 1 and 2 refer to the initial and final al., 1993).
conductance readings, respectively. Plants grown under the combined stress of highest
tolerated level of temperature and salinity (Th×Sh)
Statistical analysis exhibited spikelet fertility percentage of 51.1% which was
The overall effects of treatment and variety and their significantly lower than the control grown plants which
interaction were analyzed by means of two-way ANOVA had a higher mean of 82.3%. Among the genotypes,
with combined effect of heat treatment with salinity and N-22 had the highest spikelet fertility percentage
genotypes taken as fixed factors. Genotypes were (62.82%) among the combined stress treatment (Th×Sh)
treated as fixed factors because we were interested whereas under the control conditions, the spikelet
in the response of the specific genotypes used in this fertility percentage was highest for MO-16 (86.57%). In
experiment. The statistical analysis was done using this study, spikelet fertility percentage was significantly
OPSTAT software (CRD with two factors). reduced under combined stress of high temperature
and salinity. This result agrees with other studies, which
RESULTS AND DISCUSSION concluded that there was a reduction of grain number
The results of the combined effect of high per plant under high temperature stress (Pradhan et al.,
temperature and salinity with control are presented in 2012; Pravaliika et al., 2020), under salinity condition
Table 1. (Sairam et al., 2002), under combined salinity and high
temperature stress (Anjum et al., 2008)). Another study
The combined stress of high temperature and
showed that the actual number of spikelet is determined
salinity (Th×Sh) caused a significant reduction in mean
by the length of the reproductive phase (Rahman et al.,
plant yield (5.66 g/plant) compared to control conditions
1978). Stress induction during the critical reproductive
(14.83 g/plant). Among the genotypes, the highest yield
stage may have affected the spikelet formation in rice.
per plant was expressed by N-22 (6.75 g/plant) under
the stress conditions, while MO-16 showed the highest There is no significant difference between
yield (21.5 g/plant) followed by N-22 (12.5 g/plant) under treatments for plant height. Mean height for both the
the control conditions. In this study, grain yield per plant treatments was 111cm. Among the genotypes, N-22
was significantly affected in all genotypes and higher recorded the highest plant height in both the treatments.
reduction was reported in high yielding variety (MO-16). Since the stress was imposed during the booting stage,
Studies reported that grain yield and harvest index were there was no significant difference in plant height.

784
These differences may be due to the genetic control

Table 1. Variation in physio-morphological and yield traits under combined stress of highest tolerated level of high temperature (Th) and salinity (Sh) levels

G =0.800; T=
0.979; G×T =
of the trait. On contrary, reduction in plant height under

Yield per
plant (g/

14.833
21.50a
12.50b
10.50c
plant)

1.385
5.666
4.75b
6.75a
5.50b
temperature stress is due to decreased cell growth
especially cell elongation (Vijayakumar et al., 2021).
Zinn et al. (2010) reported that high temperature stress
mainly affect the reproductive stage.

G = 2.939; T=
3.599; G×T =
viability (%)

73.205

93.016
Pollen

74.52b
a

90.91b
93.66a
94.48a
The highest number of productive tillers under

65.97c

5.090
79.12
control conditions were found in MO-16 (15.6) whereas,
N-22 plants had the highest productive tiller number
(5.5) under combined stress treatments. The difference

G =2.714; T=
3.324; G×T =
percentage
in the productive tiller number between the genotypes

Spikelet
fertility

51.075

82.256
62.82a
b

86.57a
82.61b
31.87c

77.58c

4.701
58.52
(%)
under the control conditions was greater as compared
to the plants grown under combined stress treatments.
Salinity adversely affects a number of yield elements
including the panicles, tillers and spikelets per plant,

G = N/S ; T=
Superoxide

activity (g-1

0.036; G×T
dismutase

=0.063
floret sterility, individual size of grain, and even delayed

min-1)

0.37
0.42
0.34
0.38

0.23
0.24
0.29
0.26
heading (Grattan et al., 2002). Increasing temperatures
from 24 to 32°C resulted in reductions in tiller number
in rice (Harsant et al., 2013).

Na+ - K+ ratio

G =0.07 ; T=
0.057 ; G×T
Under combined stress condition, highest proline

=0.099
0.72a

0.69a
0.30b
0.29b
0.46c

0.57

0.42
0.54
content was recorded by N-22 (27.33 µg/g), followed
by NL-44 (26.12 µg/g) and MO-16 (24.76 µg/g).
Under control condition, highest proline content was

Control condition
recorded by N-22 (24.90 µg/g), followed by NL-44
Chlorophyll

0.201 ; G×T
G =N/S ; T=
a/b ratio

Th x Sh

= N/S
(23.48 µg/g) and MO-16 (19.97 µg/g). Average proline

1.23
1.07
1.01
1.10

2.07
2.08
2.01
2.05
content under control condition was 22.79 µg/g and
under stress condition was 26.07 µg/g. Salinity and
high temperature induced an accumulation of proline

T=0.111 ; G×T
Malondial-

(mmol g-1)

G = 0.136;
in rice leaves. In fact, the accumulation of amino acids
content
dehyde

= 0.192
a

b
2.13c

2.39

0.96
0.94
1.04
0.98
2.63

2.42
in plants has been widely reported as a response to
salinity and temperature (Dadkhah and Rassam, 2016).
The increase in proline content may be due to osmotic
regulation. Some plants are able to stand salinity by
membrane

index (%)
stability

10.556
b

69.17a
60.197

reducing the cellular osmotic potential as a result of

59.62
49.52
Cell

a net increase in inorganic and solute accumulation.

In the present study, an increase of proline was seen
in all genotypes tested. The present result was in

G = 0.970 ; T=
0.792 ; G×T
agreement with other results on wheat seedling treated
content
Proline

=1.372
(µg/g)

27.33a
b

24.90a
23.48b
c

19.97c
26.07

22.79
24.76

26.12

with high temperature and salinity, which also found

that high temperature and salinity treatments resulted
in a significant increase in proline in some genotypes
G = 0.545; T=
0.668 ; G×T =

(Hamada and Khulaef, 1995). Proline has various

tillers/plant
productive

roles, such as stabilizing proteins, membranes and

No. of

15.60a
4.500

8.933

0.944
5.50a
b

6.60b
c

4.60c
3.50

4.50

sub cellular structures, and protecting cellular functions

by scavenging reactive oxygen species under abiotic
stress conditions (Kishor et al., 2005). Proline acts as
T= N/S ; G×T
G = 16.718 ;
height(cm)

an osmoprotectant and helps plants to maintain cellular

112.00ab
125.00a

100.00b
122.00a
b

111.00

111.00
c

110.00
Plant

=N/S
98.00

homeostasis under saline stress condition (Huang et

al., 2013).
N-22 plants expressed the highest level of cell
C.D. (p≤0.05)

membrane stability index (69.17%) under the combined

stress treatment (Th×Sh). The mean cell membrane
Varieties
MO-16

MO-16
NL-44

NL-44

stability index was 59.62% with MO-16 expressing the

Mean

Mean
N-22

N-22

least value (49.52%) followed by NL-44 (60.19%). Cell

785
membrane stability was reduced under high temperature genotypes, MO-16 expressed higher Na+-K+ ratio of
condition in rice (Beena et al., 2018). Heat stress 0.72 and 0.69 compared to other genotypes under the
primarily affects the stability of plasma membranes, stressed and control conditions, respectively. Under
several proteins, cytoskeleton organization, and the control condition, lowest Na+-K+ was recorded by NL-
efficiency of cell enzymatic reactions and creating 44 (0.29). But under combined stress condition, lowest
metabolic disparity (Xu et al., 2006). Heat-stress- Na+-K+ was maintained by N-22 (0.46). This is mainly
induced oxidative stress causes peroxidation of attributed to the higher ionic imbalances resulting in
membrane lipids, proteins, and nucleic acids. Due passive accumulation of Na in root and shoot during
to reduced membrane stability, electrolyte leakage salinity stress. Na+ is the principal poisonous ion in
increases, which intensifies the membrane injuries salinized soil and low Na+ /K+ ratio in the cytoplasm is
(Wahid et al., 2007). However, under temperature and essential to maintain a number of enzymatic reactions
salinity stress conditions antioxidant concentration is (Reshna and Beena, 2021).
decreased leading to cell harm (Kreiner et al., 2002). The superoxide dismutase activity of combined
Increased cell damage due to ROS can reduce the stress treatment (Th×Sh) was 0.38 g-1 min-1 which was
thermal stability of the membrane, thereby disrupting significantly higher than 0.26 g-1 min-1under the control
the motion of water, ion and organic-solutes. conditions. Among the genotypes, N-22 expressed
Plants grown under the combined stress of highest significantly higher superoxide dismutase activity (0.42
tolerated level of drought and salinity (Th×Sh) exhibited a g-1 min-1) followed by MO-16 (0.37 g-1 min-1) under
mean malondialdehyde content of 0.98 mmol g-1 which the stress conditions whereas NL-44 expressed the
was significantly lower than the control grown plants highest superoxide dismutase activity (0.29 g-1 min-1)
which had a higher mean of 2.39 mmol g-1. Among the under the control conditions. However, the difference
genotypes, MO-16 had the highest malondialdehyde in the superoxide dismutase activity was found to be
content (2.63 mmol g-1) among the combined stress non-significant between the genotypes. The severe
treatments (Th×Sh) followed by NL-44 (2.42 mmol g-1). effects of heat stress are oxidative damage to cells by
The MDA content is linked with the oxidization of the reactive oxygen species, as found with low temperature,
cell membrane and the content of MDA is often used as drought, and salinity stress. Plants have developed
an indicator of lipid peroxidation resulting from oxidative enzymatic and nonenzymatic scavenging systems to
stress. MDA has been considered an indicator of salt- quench active oxygen, and to eliminate the detrimental
induced oxidation in cell membranes and a tool for effects of active oxygen (Bowler et al., 1992).
determining salt tolerance in plants (Ghafiyehsanj et al., From this study, we found that N-22 had a high
2013; Radi et al., 2013). In this study, MDA content were pollen viability, chlorophyll content, proline, cell
significantly increased by combined stress condition. membrane stability index, superoxide dismutase
The chlorophyll a/b ratio of the plants under control activity; less MDA content and Na+/K+ ratio. This is
conditions was higher with a mean of 2.05 while the possible due to accumulation of osmo-protectant that
mean ratio of plants grown under the combined stress may have enhanced the turgor by osmotic adjustment
of high temperature and salinity (Th×Sh) was 1.10 leading to stomatal opening and maintaining CO2 level
which was significantly lower. The chlorophyll a/b for efficient photosynthesis.
ratio among the genotypes as well as the interaction
between the genotype and treatments was found to be Authors’ contribution
non-significant. As the most significant photosynthetic Conceptualization and designing of the research
pigment, chlorophyll plays an important role in work (BR); Execution of field/lab experiments and data
controlling crop yields. Chlorophyll, however, is quite collection (ABS); Analysis of data and interpretation
fragile, not very stable and is readily influenced by (ABS, BR, SK); Preparation of manuscript (BR,SK).
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