Anda di halaman 1dari 10

Silicon Alleviates Drought Stress of Rice Plants

by Improving Plant Water Status, Photosynthesis


and Mineral Nutrient Absorption
Wei Chen & Xiaoqin Yao & Kunzheng Cai & Jining Chen
Received: 21 April 2010 / Accepted: 25 May 2010 /
Published online: 8 June 2010
#
Springer Science+Business Media, LLC 2010
Abstract Drought is a major constraint for rice production in the rainfed lowlands in
China. Silicon (Si) has been verified to play an important role in enhancing plant resistance
to environmental stress. Two near-isogenic lines of rice (Oryza sativa L.), w-14 (drought
susceptible) and w-20 (drought resistant), were selected to study the effects of exogenous Si
application on the physiological traits and nutritional status of rice under drought stress. In
wet conditions, Si supply had no effects on growth and physiological parameters of rice
plants. Drought stress was found to reduce dry weight, root traits, water potential,
photosynthetic parameters, basal quantum yield (F
v
/F
0
), and maximum quantum efficiency
of PSII photochemistry (F
v
/F
m
) in rice plants, while Si application significantly increased
photosynthetic rate (Pr), transpiration rate (Tr), F
v
/F
0
, and F
v
/F
m
of rice plants under
drought stress. In addition, water stress increased K, Na, Ca, Mg, Fe content of rice plants,
but Si treatment significantly reduced these nutrient level. These results suggested that
silicon application was useful to increase drought resistance of rice through the
enhancement of photochemical efficiency and adjustment of the mineral nutrient absorption
in rice plants.
Keywords Rice
.
Silicon
.
Drought
.
Photosynthesis
.
Root activity
.
Mineral nutrient
Introduction
Drought stress is considered to be one of the most important agricultural problems. Studies
have shown that drought has many adverse impacts on crops, inhibiting growth and
photosynthesis, changing morphology, decreasing production and photosynthetic pigment
content, affecting ion balance in plant, etc. [14]. In all, drought is not beneficial for plant
Biol Trace Elem Res (2011) 142:6776
DOI 10.1007/s12011-010-8742-x
W. Chen
:
X. Yao
:
K. Cai (*)
:
J. Chen
Key Laboratory of Ecological Agriculture of Ministry of Agriculture,
South China Agricultural University, Guangzhou 510642, China
e-mail: kzcai@scau.edu.cn
X. Yao
The College of Life Sciences, Hebei University, Baoding 071002, China
growth and development, and the increase in plant resistance to drought is an important
way to overcome drought problems.
Silicon (Si) is beneficial for mostly higher plants, especially under stressful environment,
although it is not considered to be required by higher plants [5, 6]. Liang et al. [7] reported
that Si enhanced resistance of wheat to freezing stress. Physiological and cytological studies
suggested that silicon-induced defense response and cell silicification of rice leaves
altogether contribute to the silicon-induced rice resistance to blast disease [8]. Si also
alleviated the effects of salt stress, high temperature, and heavy metals toxicity on plants
[912]. In addition, there were some studies about the beneficial role of Si in plant
subjected to drought stress, mainly concentrating on water metabolism, photosynthesis, and
some physiological traits [1315]. However, the mechanism in Si-alleviated damage caused
by drought stress remains unclear.
Rice (Oryza sativa L.) is one of the most important food crops in the world. At present,
the growth of rice has been seriously influenced by drought in many regions. This paper
studies the effects of Si supply on water status, photosynthesis and mineral nutrient
absorption of rice seedlings grown under drought stress, which maybe help further elucidate
the physiological mechanism of Si in improvement of drought tolerance of plants. We
hypothesize that Si supply will alleviate drought stress of rice plants
Materials and Methods
Plant Materials and Growth Conditions
Two rice lines w-14 (drought susceptible) and w-20 (drought resistant) were used in the
experiment. Rice seeds were surface-sterilized with 10% H
2
O
2
for 10 min and rinsed
thoroughly with distilled water, then soaked with distilled water and germinated on moist
filter paper for 48 h in Petri dishes. Ten days later, seedlings were transplanted into
polyethylene plastic pot (30 cm height and 25 cm in diameter) that was filled with 5 kg
of paddy soil. The content of the soil organic matter, available N, P, K, and Si were
31.25 g/kg, 179.20, 45.4, 134.36, and 52.64 mg/kg, respectively. Plants were fertilized with
0.15 g urea and 0.15 g compound fertilizer (N/P/K fertilizer 15:15:15) per pot on the day of
transplanting.
Silicon and Drought Treatments
Our preliminary experiment using different Si concentration (0, 0.5, 1.0, 1.5, 2.0 mM)
showed that 1.5 mM of silicon has the best effects to improve the negative influence of
drought stress, so in this experiment 1.5 mM of silicon was used. There were four
treatments for both rice lines: (1) no drought and no silicon added (wet, Si); (2) no drought
and 1.5 mM silicon added (wet, Si+); (3) drought and no silicon added (dry, Si); (4)
drought and 1.5 mM silicon added (dry, Si+). The silicon was added as potassium silicate
(K
2
SiO
3
) to the soil before sowing. In the silicon-deficient treatment, potassium chloride
was used to replenish potassium. The rice seedlings contained in pots were submerged and
maintained as 12 cm water layer. Drought treatment was imposed at 28 days after sowing
for a period of 15 days for both two rice lines. During the period of dry treatment, the water
content of the pot soil was adjusted gravimetrically every day to 25.1%. Treatments for
each rice line were arranged factorially in a randomized complete block design with four
replicates per treatment. Plants were collected at 43 days (after 15 days of drought
68 Chen et al.
treatment) to determine plant biomass, root system traits and water utilization efficiency, the
content of chlorophyll and inorganic element in leaves. Chlorophyll fluorescence
parameters and gas exchange parameters were also investigated.
Determination of Plant Biomass, Leaf Water Potential, and Water Utilization Efficiency
The treated rice plants were sampled to determine biomass, leaf water potential and water
utilization efficiency. For dry weight (DW) determination, shoots and roots were dried at
70C for 48 h and weighed. The leaf water potential was measured by using water potential
instrument (WP4, Decagon, USA). Water utilization efficiency (g.cm
2
) was calculated as
the plant dry weight divided by irrigated water amount.
Determination of Root Traits
Root samples were taken and scanned to measure total root length, root surface area, root
average diameter and total root volume by using WinRHIZO software (v. 3.1, Regent
Instruments, Inc., Quebec, QC, Canada). Rice root activity was determined by measuring
oxidation of alpha-naphthylamine (-NA) [16]. Briefly, one gram of fresh roots was
transferred into a 150 mL flask containing 50 mL of 50 g. mL
1
-NA. The flasks were
incubated for 2 h at room temperature in an end-over-end shaker. After incubation, the
aliquots were filtered and 2 mL of aliquot was mixed with 1 mL NaNO
3
(1.18 mmol L
1
)
and 1 mL sulphanilic acid (0.06 mol L
1
), when the color of the mixed solution became red,
the solution was finally diluted to a volume of 25 mL with distilled demonized water and
measured by spectrophotometer.
Determination of Chlorophyll Content
About 0.2 g rice leaves were ground to a fine powder and extracted with 10 mL of 80%
acetone (v/v). The homogenate was centrifuged at 4,000g at 4C for 10 min, and the
supernatant was separated and used for the chlorophyll assay. The amounts of chlorophyll a
and b were determined spectrophotometrically, by reading the absorbance at 663 and
645 nm. The chlorophyll content results are expressed as units mg per gram-fresh weight
(mg/g FW) and calculated by using the extinction coefficients and the equations given by
[17].
Measurement of Chlorophyll Fluorescence
Fluorescence parameters variable (F
v
)/minimal (F
0
) and F
v
/maximal (F
m
) of rice leaves
were measured using chlorophyll fluorometer (OS-30P, Opti-Sciences, Inc., Tyngsboro,
MA, USA) after a 20-min dark period in ambient conditions in the laboratory. F
0
, F
m
, and
F
v
level were recorded, and then the maximum quantum efficiency of PSII photochemistry
(F
v
/F
m
) and the basal quantum yield (F
v
/F
0
) were calculated.
Determination of Gas Exchange
The photosynthetic rate (Pr), transpiration rate (Tr), stomatal conductance (gs), and
intercellular CO
2
concentration (Ci) of the sampled leaves were measured under clear and
sunny weather using a portable photosynthesis system (LI-6400, LI-COR, Inc., Lincoln,
NE, USA).
Silicon Alleviates Drought Stress of Rice Plants 69
Determination of Si Content
The Si contents of rice leaves were determined by the colorimetric molybdenum blue
method [18]. Briefly, 0.3 g samples of rice leaves were ashed in porcelain crucibles for 3 h
at 550C, the ash was dissolved in 1.3% hydrogen fluoride, then the Si concentrations in
the solutions were measured by the colorimetric molybdenum blue method at 811 nm with
a spectrophotometer (PGENERAL TU-1901 UV-VIS, Beijing, China). A control (no
sample) was added to eliminate porcelain crucible errors.
Measurement of Mineral Nutrient Content
Fully developed leaf samples for each replication with Si and drought treatment were sampled
to determine inorganic element content, including Na, K, Ca, Mg, and Fe. 0.3 g leaf samples
were dried at 70C for 48 h and DW was determined. Dried samples were ground by hand with
a glass rod in test tubes. 5 mL perchloric acid and 15 mL nitric acid were added and incubated
overnight at roomtemperature, then digested at 180Cusing a heating block, and finally diluted
to a volume of 50 mLwith distilled demonized water. The concentrations of Na, K, Ca, Mg, and
Fe was determined by atomic absorption spectrometer (SpectrAA220FS, Varian, USA). A
control (no leaf sample) was added to eliminate errors
Statistical Analysis of the Data
All the data in the figures and table were expressed as the meansstandard error of four
replicates and analyzed by ANOVAS using SPSS software(version 13.0; SPSS Inc.,
Chicago, USA) computer program, statistical differences among treatments within the same
rice line was determined using the least significant difference at a 0.05 probability level.
Results
Dry Matter Accumulation
Silicon application did not influence the dry weight of rice plants in the wet treatment.
Upon expose to drought stress, rice growth decreased significantly, while exogenously
applied 1.5 mM silicon significantly increased plant dry matter. The shoot dry weight in the
dry/Si+ treatment increased by 103.2% for w-14 and 97.4% for w-20 compared with dry /Si
treatment (Table 1). Root and total dry weight were similarly affected by Si and drought
treatment in both lines.
Si Content
Drought significantly reduced silicon content in roots but not in leaves. Silicon content in leaves
and roots was increased by silicon application regardless of the soil water regime (Table 1).
There were no significant differences in the silicon content between two rice lines.
Root Traits
The present study showed that drought treatment significantly inhibited root system growth,
total root length, root surface, root volume and root activity, but Si application significantly
70 Chen et al.
improved these traits (Table 2). There was no significant difference in root traits in the wet
treatments regardless of silicon application. Root traits were similarly affected in both rice
lines.
Leaf Water Status, Chlorophyll Fluorescence Parameter, and Chlorophyll Content
As shown in Table 3, leaf water potential and water utilization efficiency (WUE) were
significantly decreased after drought treatment, while application of 1.5 mM Si
concentration significantly increased leaf water potential by 14.69% and 26.56%, WUE
by 118.8% and 116.7% for line w-14 and w-20, respectively.
Table 2 Effects of Si Application and Drought on Root Traits in Rice Lines w-14 and w-20
Treatment Total root length
(cm.pot
1
)
Total surface area
(cm
2
.pot
1
)
Total root
volume
(cm
3
.pot
1
)
Average root
diameter (cm)
root activity
(g.g
1
.h
1
FW)
W-14
Wet, Si 1768.58114.69 b 602.8770.32 a, b 19.631.09 a, b 0.240.01 a 51.285.49 a, a
Wet, Si+ 2790.11189.72 a 789.2368.37 a 22.271.97 a 0.240.01 a 53.137.40 a
Dry, Si 1367.8335.65 b 529.4266.92 c 15.281.18 b 0.220.01 a 21.964.11 c
Dry, Si+ 2263.73109.86 a 627.2659.3 a, b 21.341.23 a 0.240.01 a 35.523.43 b
W-20
Wet, Si 1931.16157.58 a, b 532.6169.42 b 15.851.00 a, b 0.210.01 a 41.977.38 a
Wet,Si+ 2539.55209.34 a 726.8964.29 a 17.470.82 a 0.230.01 a 42.784.99 a
Dry, Si 1529.66157.76 b 472.1449.42 c 13.710.95 b 0.210.01 a 17.290.92 c
Dry, Si+ 2142.49130.35 a 652.3684.94 a, b 16.731.25 a 0.220.01 a 28.191.20 b
Si+ and Si indicate treatment with or without 1.50 mM silicon. Dry and wet indicate treatments with and
without drought, respectively. Values are meansstandard error from four replicates (n=4). Different
lowercase letters denotes statistical difference using a least significant difference test (P<0.05)
Table 1 Effects of Si and Drought on Dry Matter Accumulation and Si Content in Rice Lines w-14 and w-20
Treatment Dry matter accumulation (g.pot
1
) Si content (g/kg)
Root weight Shoot weight Total weight Leaf Root
W-14
Wet 1.310.14 a, b 13.200.79 a, b 16.110.74 a, b 38.741.66 c 13.042.11 b
Wet 1.560.27 a 15.190.78 a 18.350.60 a 45.881.40 b 28.914.61 a
Dry 0.530.05 c 5.630.44 c 7.760.43 c 40.951.12 c 9.091.72 d
Dry 1.020.09 b 11.441.05 b 14.051.07 b 50.901.05 a 22.513.45 a, b
W-20
Wet 1.820.29 a 14.530.51 a, b 17.960.56 a, b 40.961.32 b 20.181.12 b
Wet 1.730.12 a 15.951.03 a 19.281.07 a 53.611.37 a 31.982.14 a
Dry 0.800.03 b 6.610.82 c 9.010.84 c 44.571.00 b 14.240.49 c
Dry 1.090.18 b 13.050.84 b 15.741.02 b 54.122.11 a 25.552.49 b
Si+ and Si indicate treatment with or without 1.50 mM silicon. Dry and wet indicate treatments with and
without drought, respectively. Values are meansstandard error from four replicates (n=4). Different
lowercase letters denotes statistical difference using a least significant difference test (P<0.05)
Silicon Alleviates Drought Stress of Rice Plants 71
The basal quantum yield (F
v
/F
0
) and maximal quantum yield of PS II (F
v
/F
m
) of rice
leaves showed no significant difference for well-watered rice plants regardless of silicon
application (Table 3). Drought treatment significantly decreased the ratio of F
v
/F
0
and
F
v
/F
m
. In the drought conditions, the value of F
v
/F
0
and F
v
/F
m
were decreased by 13.07%
and 2.65% for w-14, respectively, 14.92% and 2.35% for w-20. In the wet treatment,
chlorophyll fluorescence parameters F
v
/F
0
, F
v
/F
m
were not affected by Si application.
Similar results were seen for chlorophyll content.
Table 3 Effects of Si and Drought on Leaf Water Potential, Water Utilization Efficiency, Chlorophyll
Content, and Chlorophyll Fluorescence Parameter Fv/F0, Fv/Fm of Leaves in Rice Lines w-14 and w-20
Treatment Leaf water
potential (Mpa)
Water utilization
efficiency (g.cm
2
)
Chlorophyll
content (mg.g
1
)
Fv/F0 Fv/Fm
W-14
Wet, Si 1.190.05 a, b 0.290.01 b, c 2.210.06 a 3.730.11 a, b 0.7890.02 a
Wet, Si+ 1.040.04 a 0.420.02 a 2.240.11 a 4.230.32 a 0.7930.01 a
Dry, Si 1.770.13 c 0.160.01 c 1.600.03 c 3.060.09 c 0.7540.01 c
Dry, Si+ 1.510.18 b 0.350.01 b 1.910.11 b 3.460.03 b, c 0.7740.01 b
W-20
Wet, Si 1.330.23 a 0.330.01 b 1.980.13 a 3.960.08 a 0.7950.01 a, b
Wet, Si+ 1.270.02 a 0.440.02 a 1.950.17 a 4.040.12 a 0.7990.01 a
Dry, Si 1.920.19 b 0.180.01 c 1.410.04 c 3.150.10 c 0.7660.02 c
Dry, Si+ 1.410.08 a 0.390.01 a, b 1.640.08 b 3.620.09 b 0.7840.01 b
Si+ and Si indicate treatment with or without 1.50 mM silicon. Dry and wet indicate treatments with and
without drought, respectively. Values are meansstandard error from four replicates (n=4). Different
lowercase letters denotes statistical difference using a least significant difference test (P<0.05)
Table 4 Effects of Si and Drought on Photosynthetic rate (Pr), Transpiration rate (Tr), Stomatal
Conductance (gs), and Intercellular CO2 Concentration (Ci) of Leaves in Rice Lines w-14 and w-20
Treatment Pr (mol CO
2
m
2
s
1
) Tr (mmol H
2
O m
2
s
1
) g
s
(mmol H
2
O m
2
s
1
) Ci (mol CO
2
mol
1
)
W-14
Wet, Si 8.920.28 a 8.870.26 a, b 0.430.01 a 355.650.56 a
Wet, Si+ 9.170.38 a 9.250.08 a 0.390.01 a 345.614.63 a
Dry, Si 5.780.06 c 6.420.87 c 0.280.03 b 335.061.21 a, b
Dry, Si+ 7.890.28 b 7.610.23 b 0.140.02 c 323.713.26 b
W-20
Wet, Si 7.000.25 a, b 5.470.17 a 0.260.02 a 332.095.38 a
Wet, Si+ 7.350.40 a 5.640.21 a 0.240.02 a 333.832.28 a
Dry, Si 4.851.09 c 3.380.42 b 0.180.01 b 310.842.25 b
Dry, Si+ 6.280.12 b 5.180.26 a 0.120.01 c 301.794.20 b
Si+ and Si indicate treatment with or without 1.50 mM silicon. Dry and wet indicate treatments with and
without drought, respectively. Values are meansstandard error from four replicates (n=4). Different
lowercase letters denotes statistical difference using a least significant difference test (P<0.05)
72 Chen et al.
Gas Exchange
Under drought conditions, Pr, Tr, and gs of rice lines were decreased significantly by 27.6
38.2% than those in the wet treatment (Table 4). Compared with the dry/Si treatment, the
dry/Si+ treatment significantly increased Pr, Tr, but the value of gs was decreased. The
influence of silicon application on Ci was not significant regardless of drought treatment.
Gas exchange was similarly affected in both rice lines.
Mineral Nutrient Content
Silicon application and drought treatment had significant effects on mineral nutrient content
of rice leaves (Table 5). No significant change was observed in K, Na, Ca, Mg, Fe content
of rice leaves between plants supplied or not with Si in the wet conditions. Drought stress
resulted in a significant increase of mineral nutrient content in rice leaves compared with
that in wet treatments. Line w-20 accumulated higher level of K, Na, Ca, Mg but lower
level of Fe than that in line w-14. Compared with the dry/Si treatment, plants treated with
added Si significantly reduced K, Na, Ca, Mg, and Fe concentrations in rice lines to a level
close to wet/Si treatment.
Discussion
Drought is an important abiotic factor that could influence the growth and physiological
traits of plants [19, 20]. Some reports showed that Si could increase the tolerance of plants to
stressful environment [7, 10, 21]. In the two studied rice lines, drought treatment (Dry/Si)
caused significant decreases in biomass accumulation, and Si supply promoted biomass
accumulation of seedlings under drought stress, but it did not significantly affect on biomass
under wet condition (Table 1).
Table 5 Effects of Si and Drought on Mineral Nutrient Content of Leaves in Rice Near-isogenic
Lines w-14 and w-20
Treatment g kg
1
DW
K Na Ca Mg Fe
W-14
Wet, Si 24.370.75 c 1.560.05 b 3.240.15 b 1.890.08 b 0.130.01 b
Wet, Si+ 23.400.60 c 1.510.04 b 2.890.08 b 1.580.04 b 0.130.02 b
Dry, Si 28.680.58 a 2.240.21 a 4.340.32 a 2.370.13 a 0.200.02 a
Dry, Si+ 26.380.78 b 1.710.11 b 3.240.12 b 1.860.13 b 0.170.02 b
W-20
Wet, Si 22.000.53 b 1.850.10 b 3.610.39 c 1.710.12 b, c 0.120.02 b
Wet, Si+ 21.130.29 b 1.740.19 b 3.510.21 c 1.580.05 c 0.120.01 b
Dry, Si 27.541.10 a 3.630.43 a 6.280.53 a 2.630.25 a 0.170.01 a
Dry, Si+ 22.821.05 b 2.090.14 b 4.790.17 b 2.070.06 b 0.140.01 b
Si+ and Si indicate treatment with or without 1.50 mM silicon. Dry and wet indicate treatments with and
without drought, respectively. Values are meansstandard error from four replicates (n=4). Different
lowercase letters denotes statistical difference using a least significant difference test (P<0.05)
Silicon Alleviates Drought Stress of Rice Plants 73
Root is not only a vital organ of plants for absorbing nutrient and water, but also is very
important for assimilation, synthesis, and transportation of many materials. Therefore root
traits directly affected growth and development of plants [22]. Drought stress not only
significantly decreased root growth of two rice lines, but also inhibited biomass
accumulation of seedlings (Table 2). Under drought stress, Si supply significantly increased
total root length, total surface area, total root volume, and root activity of both lines, silicon
content of roots was also increased. These results indicated that Si supply could improve
root traits and enhance water absorption, therefore increased the resistance of rice plants to
drought stress.
Water metabolism is an important index in measuring plant physiological function and
growth and development. In the present study, Si could increase resistance of plant to
drought stress by improving water status of plant for both two rice lines (Table 3). Similar
results were reported for some other plant species including wheat (Triticum aestivum L.)
[23, 24], sorghum [13], and maize [21]. The positive impacts of Si on plants under drought
conditions were associated with transpiration reduction [25, 26]. However, our study
showed that transpiration rate in leaves of drought-stressed rice was significantly increased
by Si addition (Table 4). The similar phenomena were observed in sorghum [13] and wheat
[23] under drought stress. Recent studies by Hattori et al. [27] showed that the silicon-
induced alleviation was independent of leaf water status. This leads to the conclusion that
the resistance mechanisms induced by Si in response to drought stress are more
complicated, it is worthy of further study.
The reduced growth was also closely related to a strong reduction in photosynthesis
under drought stress. Studies showed that Si supply could increase photosynthetic rate of
plants subjected to drought stress [6, 13]. Similar results were found in this study. Ci can be
used to discriminate between changes in photosynthetic rate resulting from stomatal
limitation or non-stomatal limitations [28]. From our results, the decrease in photosynthetic
rate caused by drought stress maybe resulted from non-stomatal and stomatal limitations
(Table 4). However, under drought stress, it was not clear that the roles of Si in the increase
of photosynthetic rate were due to non-stomatal or stomatal limitations.
Measurements of chlorophyll fluorescence parameters provides useful information about
photosystem II (PSII) activity and changes in photosynthetic metabolism of stress plants
[29]. Both of maximum/potential quantum efficiency of PS II (F
v
/F
m
) and the maximum
primary yield of photochemistry of PS II (Fv/F
0
) are related with photosynthetic efficiency
of plant leaves [30]. F
v
/F
m
has been widely used to detect stress-induced perturbations in
the photosynthetic apparatus. In the literature, there were no available data on the
relationship between Si and photochemical efficiency of PSII. As Table 3 shown, drought
stress resulted in the reduction of F
v
/F
m
, which may be associated with decreased efficiency
of energy transfer from the antenna to the reaction centers and/or inactivation of PSII
reaction centers [31]. The decrease of F
v
/F
m
might impair photosynthesis, and ultimately
affect plant growth and development. It should be stressed that Si supply enhanced Fv/Fm
under drought stress, but not under well-water condition, which further suggested that Si
supply was able to alleviate the harmful effects of drought on the photochemical reactions.
Uptake and translocation of mineral nutrients within the plant could be affected by
drought stress [32]. Accumulation of inorganic ions is one way for plants to adapt to
environmental stresses [3335]. Kaya et al. [21] found that Si application could increase
Ca, K, and Si content of maize leaves under drought stress. However, in the present
investigation, water stress stimulated mineral nutrient absorption of rice plant especially for
drought lines (w-20), Si supply only increased Si content in rice leaves, whereas reduced K,
Na, Ca, Mg, and Fe content under drought stress (Table 5). Pei et al. [24] reported the
74 Chen et al.
similar results that Si could decrease the absorption of plant to K, Ca, Mg under water-
deficit stress induced by polyethylene glycol in wheat seedlings. From the above analysis to
know, Si had some influence on mineral element in plants, but it still need further
investigation.
In conclusion, the paper illustrated that Si alleviated drought stress of both rice
seedlings, mainly by improving water use efficiency, photosynthesis, and photochemical
efficiency, and by adjustment of mineral nutrient level in rice plants subjected to drought
stress.
References
1. Flowers TJ, Yeo AR (1986) Ion relations of plants under drought and salinity. Aust J Plant Physiol 1:75
91
2. Jamieson PD, Martin RJ, Francis GS (1995) Drought influences on grain yield of barley, wheat, and
maize. New Zeal J Crop Hort 23:5556
3. Iturbe-Ormaetxe I, Escuredo PR, Arrese-Igor C et al (1998) Oxidative damage in pea plants exposed to
water deficit or paraquat. Plant Physiol 116:173181
4. Xu H, Biswas DK, Li WD (2007) Photosynthesis and yield responses of ozone-polluted winter wheat to
drought. Photosynthetica 45:582588
5. Ma JF (2004) Role of silicon in enhancing the resistance of plants to biotic and abiotic stresses. Soil Sci
Plant Nutr 50:1118
6. Li QF, Ma CC, Ji J (2009) Effect of silicon on water metabolism in maize plants under drought stress.
Acta Ecologica Sinica 29:841634168
7. Liang YC, Zhu J, Li ZJ et al (2008) Role of silicon in enhancing resistance to freezing stress in two
contrasting winter wheat cultivars. Environ Exp Bot 64:286294
8. Cai KZ, Gao D, Luo SM et al (2008) Physiological and cytological mechanisms of silicon-induced
resistance in rice against blast disease. Physiol Plant 134:324333
9. Ahmad R, Zaheer SH, Ismail S (1992) Role of silicon in salt tolerance of wheat (Triticum aestivum L.).
Plant Sci 85:4350
10. Matsui T, Omasa K, Horie T (2001) The difference in sterility due to high temperature during the
flowering period among Japanese-rice varieties. Plant Prod Sci 2:9093
11. Hammond KE, Evans DE, Hodson MJ (1995) Aluminium/silicon interactions in barley (Hordeumvul-
gare L.) seedlings. Plant Soil 173:8995
12. Shi XH, Zhang CC, Wang H et al (2005) Effect of Si on the distribution of Cd in rice seedlings. Plant
Soil 272:5360
13. Hattori T, Inanagaa S, Arakib H et al (2005) Application of silicon enhanced drought tolerance in
sorghum bicolor. Physiol Plant 123:459466
14. Liang YC, Sun WC, Zhu YG et al (2007) Mechanisms of silicon-mediated alleviation of abiotic stresses
in higher plants: a review. Environ Pollut 147:422428
15. Crusciol CAC, Pulza AL, Lemosb LB et al (2009) Effects of silicon and drought stress on tuber yield
and leaf biochemical characteristics in potato. Crop Sci 49:949954
16. Zhang X, Tan G, Huang Y (1994) Experimental technology of plant physiology. Liaoning Science and
Technology Press, Shenyang, China
17. Porra RJ, Thompson WA, Kriedemann PE (1989) Determination of accurate extinction coefficients and
simultaneous equations for assaying chlorophyll a and b extracted with four different solvents:
verification of the concentration of chlorophyll standards by atomic absorption spectroscopy. Biochim
Biophys Acta 975:384394
18. Van der Vorm PDJ (1987) Dry ashing of plant material and dissolution of the ash in HF for the
colorimetric determination of silicon. Commun Soil Sci Plant Anal 18:11811189
19. Farooq M, Wahid A, Lee DJ et al (2009) Advance in drought resistance of rice. Crit Rev Plant Sci
4:199217
20. Yao XQ, Chu JZ, Wang GY (2009) Effects of drought stress and selenium supply on growth and
physiological characteristics of wheat seedlings. Acta Physiol Plant 5:10311036
21. Kaya C, Tuna L, Higgs D (2006) Effect of silicon on plant growth and mineral nutrition of maize grown
under water-stress conditions. J Plant Nutr 29:14691480
22. Aiken RM (1996) Root system regulation of whole plant growth. Annu Rev Phytopathol 34:32546
Silicon Alleviates Drought Stress of Rice Plants 75
23. Gong H, Chen K, Chen G et al (2003) Effects of silicon on growth of wheat under drought. J Plant Nutr
26:10551063
24. Pei ZF, Ming DF, Liu D et al (2010) Silicon Improves the tolerance to water-deficit stress induced by
polyethylene plycol in wheat (Triticum aestivum L.) seedlings. J Plant Growth Regul 29:106115
25. Agarie S, Agata W, Kubota F et al (1998) Effects of silicon on transpiration and leaf conductance in rice
plants (Oryza sativa L.). Plant Prod Sci 2:8995
26. Gao X, Zou C, Wang L et al (2006) Silicon decreases transpiration rate and conductance from stomata of
maize plants. J Plant Nutr 29:16371647
27. Hattori T, Sonobe K, Inanaga S et al (2008) Effects of silicon on photosynthesis of young cucumber
seedlings under osmotic stress. J Plant Nutr 31:10461058
28. Farquhar GD, Sharkey TD (1982) Stomatal conductance and photosynthesis. J Ann Rev Plant Physiol
33:317345
29. Schnabel G, Strittmatter G, Noga G (1998) Changes in photosynthetic electron transport in potato
cultivars with different field resistance after infection with Phytophthora infestans. J Phytopathol
146:205210
30. Shangguan ZP, Shao MA, Dyckmans J (2000) Effects of nitrogen nutrition and water deficit on net
photosynthetic rate and chlorophyll fluorescence in winter wheat. J Plant Physiol 156:4651
31. Kooten O, Snel JFH (1990) The use of chlorophyll fluorescence nomenclature in plant stress physiology.
Photosynth Res 3:147150
32. Hu YC, Schmidhalter U (2005) Drought and salinity: a comparison of their effects on mineral nutrition
of plants. J Plant Nutr Soil Sci 168:541549
33. Patakas A, Nikolaou N, Zioziou E et al (2002) The role of organic solute and ion accumulation in
osmotic adjustment in drought-stressed grapevines. Plant Sci 163:361367
34. Zhu X, Gong H, Chen G et al (2005) Different solute levels in two spring wheat cultivars induced by
progressive field water stress at different developmental stages. J Arid Environ 62:114
35. Chaves MM, Oliveira MM (2005) Mechanisms underlying plant resilience to water deficits: prospects
for water-saving agriculture. J Exp Bot 407:23652384
76 Chen et al.