Silicon

https://doi.org/10.1007/s12633-020-00797-4

ORIGINAL PAPER

Silicon Mitigates Drought Stress in Wheat (Triticum aestivum L.)

Through Improving Photosynthetic Pigments, Biochemical
and Yield Characters
Muhammad Adnan Bukhari 1 & Zahoor Ahmad 2,3 & M. Yasin Ashraf 4 & Muhammad Afzal 5 & Fahim Nawaz 6 &
Muhammad Nafees 7 & Wajid Nasim Jatoi 1 & Naveed Aslam Malghani 8 & Adnan Noor Shah 9 & Abdul Manan 10

Received: 1 September 2020 / Accepted: 20 October 2020

# Springer Nature B.V. 2020

Abstract
Feeling prone to stress differs with plant production stage, water scarcity near commencement of grain filling phase has a
significant reduced grain yield through fewer endosperm and sink cells capacity that minimize the accumulation of dry matter
and assimilates supplies. In greenhouse research was conducted to investigate the part of silicon (Si) in regulating the production,
yield and physico-chemical traits of wheat (Triticum aestivum L.) under drought situation. Two commercial good varieties of
wheat (Chakwal-50 and Sehar-06) were planted under well water (100% FC) and water stress conditions (60% FC) pooled with
Si 3 mM (seed priming), 1 mM (fertilization) and 4 mM (foliar application). In tillering and anthesis stage, yield and yield
elements, leaf pigments, gaseous exchange parameters, biochemical and nutrients were analyzed from the wheat plants. The
stress of water deficit prompted a marked reduction in agronomic parameters while biochemical attributes were increased under
conditions of water stress except for total soluble proteins. The effect of Si was found to be positive at both the stages under well-
watered and drought affected wheat plants. At anthesis stage, foliarly applied Si enhanced biochemical attributes and accumu-
lation of nutrients under the stress of water deficit and normal water plants, respectively. Under normal conditions, the most
successful method was the taking of leaf Ca by Si foliar spray at anthesis level. Applied with no Si the water stressed plants
retained the optimum concentration of Fe. Here as well, Si fertigation under well water condition was found to be better for
studied agronomic parameters at the tillering stage. It may be included that Si application improves plant biochemical attributes,
absorbs nutrients and shows higher growth rate which ultimately helps the plant mitigate the drought-stressing effects. At the
anthesis stage, foliar spray was stronger while at the tillering stage, fertigation produced good results.

Keywords Biochemical . Gaseous exchange parameters . Photosynthetic pigments . Nutrients . Wheat

* Zahoor Ahmad 6
Institute of Crop Science (340 h), University of Hohenheim,
zahoorahmadbwp@gmail.com Stuttgart, Germany

7
1 Department of Horticulture, University College of Agriculture &
Department of Agronomy, University College of Agriculture & Environmental Science, The Islamia University of Bahawalpur,
Environmental Science, The Islamia University of Bahawalpur, Bahawalpur, Pakistan
Bahawalpur, Pakistan
2 8
Department of Field Crops, Faculty of Agriculture, Cukurova Department of Plant Pathology, University College of Agriculture &
University, Adana, Turkey Environmental Science, The Islamia University of Bahawalpur,
3 Bahawalpur, Pakistan
Department of Botany, University of Central Punjab, Punjab Group
of Colleges, Bahawalpur, Pakistan 9
4
Department of Agronomy, Gomal University, Dera Ismail
Institute of Molecular Biology and Biotechnology, The University of Khan, KPK, Pakistan
Lahore, Lahore, Pakistan
5 10
College of Food and Agriculture Sciences, Plant Production Department of Horticulture, Ghazi University, Dera Ghazi
Department, King Saud University, Riyadh, Saudi Arabia Khan, Pakistan
 Silicon

Abbreviations conditions few mRNAs types are formed by plants. Proline

Si Silicon production is also sometimes well documented under water-
FC Field capacity limited stress [6, 9, 10].
FW Fresh weight (g) Drought affects the metabolic pathways; either to lose pho-
V Volume tosynthetic pigments or decreased production. Alteration in
E Transpiration Rate the photosynthetic pigment amounts are linked with plant bio-
A Instantaneous net CO2 mass yield [11]. Adequate humidity at or after anthesis does
(gs) Stomatal conductance not only does it help plants improve their photosynthetic pro-
BSA Bovine serum albumin duction, it also provides spare time to transfer carbohydrates
TFA Total free amino acid [12]. Carotenoids, the necessary pigment, also play quite crit-
Chla Chlorophyll a ical role in photosynthesis. Carotenoid production in plants
Chlb Chlorophyll b carried out by genetic trait but environmental factors also play
Chlt Chlorophyll t an important part under stress condition [13]. Wheat geno-
CAR Carotenoids types from anthesis to maturity under control and drought
TFA Total free amino acid conditions and said flag leaves under water-stressed condition
LP Leaf protein recorded loss in sugars contents and ultimately seed yield
TSS Total soluble sugars reduction, by increasing cell wall invertase activity and solu-
TSP Total soluble protein ble content [14]. Among morphological characters, stomata
TNOTPP Total number of tillers plant−1 are present on beneath epidermal layer that regulates the flow
NFTPP No. of fertile tillers plant−1 of moisture and CO2 among atmosphere and plants. The role
SL Spike length (cm) of stomata in comparing tolerant and sensitive to drought
NSPS No. of spikelet’s spike−1 wheat genotypes and suggested that the size of stomata was
HSW 100-grain weight (g) significantly higher in tolerant cultivar than in sensitive ones
BY Biological yield (g plant−1) [15]. The photosynthesis rate is also halted under extreme
GY Grain yield (g plant−1) drought conditions due to insufficient amount of CO2 due to
HI Harvest index the lack of stomata and mesophyll diffusion of the carbon
dioxide molecule and carbon acclimatization was also oc-
curred [16]. The decline in net photosynthetic rates (A) cause
1 Introduction closing of stomata and reduces of CO2 concentration in the
cells [17]. Metabolic factors change the Rubisco enzyme po-
Wheat (Triticum aestivum L.) is significant crop for food in tential and disturbed the activity of ribulose 1,5 bisphosphate
cereals for more than 3rd population of the worlds. It adds synthesis [1, 18].
more energy with protein contents than any other cereal crop Si is efficiently reducing the adverse effects of drought
to the world diet [1]. Yet recent crop production patterns sug- stress, increasing water table, toxicity of heavy metals,
gest that soil nutrients status and fertility are decreasing world- sodicity, boiling and freezing in plants. Plants with enough
wide due to extensive natural resource use, unsuitable man- Si are substantially derived from Si deficient plants by
agement activities and even risks to the ecosystem such as structure, crop stand, bio-chemical properties, grain and
biotic and abiotic tension. Water shortages are a universal varying yield characteristics ascribing to many stresses
issue that poses a serious threat for crop production [2, 3]. i.e. drought [5, 19]. Cao et al. (2017) [20] reported an
Water scarce condition affects plant water quality, eventually efficient way to alleviate stress effects by applying Si (soil
resulting in reduced plant growth, metabolism, physiology, or foliar). Furthermore, Si also reduced various harmful
and development [4]. Stage unique conditions of water are drought consequences in different plants under stress at
of critical importance for a successful production of the crops. various developmental levels [21]. Some evidence also
Drought resistant genotypes see more competencies than sen- suggested that silicone can be used for osmotic modifica-
sitive varieties being assimilated to fill their grains to carry tion of plants [22]. The use of silicone in stressed seedlings
pre-anthesis. Carbohydrates in rising wheat leaves are impor- has been reported to be beneficial in drought [23].
tant contributors to the osmotic change [1, 5, 6]. Chlorophyll Exogenous use of silicone has many significant impacts
contents (a and b) are significant that are involved in the on yield due in large part to its special biological functions
photosynthetic phase. Water shortage can similarly delay the [24]. Si also enhanced the exchange of gases, water trans-
development of different proteins meanwhile boosts protein portation, leaf water potential and chlorophyll contents
synthesis [7]. The drought stress of coleoptiles in avena [25]. Si use in the soil area control water loss through
changed the protein production rates [8]. However, stress from stomata, SC, net photosynthesis and CO2 movement in
drought can stop protein production and under water-limited the cells was reported in leguminous plants stressed by
Silicon
drought [26]. It was also suggested that Si enhance source
and sink relationship to produce carbohydrates in many
crops such as cucumber and sugar cane improve the activ-
ity of leaf ultra-diagram, chlorophyll content, and Rubisco
activity [27]. Earlier studies suggested that Si enhanced net
photosynthetic production in different wheat cultivars un-
der water deficit and maintained the enzyme activity and
improve the biological pathway in plant cell [28]. In addi-
tion, the introduction of Si to the medium growing root
caused a substantial growing the rate of photosynthesis
and CO 2 fixation in various maize genotypes [29].
Transpiration decreased with silicone fertilization in differ-
ent plants [27]. The reduction in evapotranspiration can
attributed to the accumulation of Si below the cuticle,
which limit water loss from the stomata. The significant
influence of silicone has been associated with reducing
high water loss when transpiring [30] or with accumulation
of Si crystals beneath the leaves and stem cells [31], which
can minimize loss of water through the cuticular. Previous
studies in (Zea mays L.) and Oryza sativa L. suggested
minimum water loss by evapo-transpiration was primarily
correlated by limiting stomatal aperture [32], signifying Si
connection with opening and closing of stomata. It was
also noticed that Si minimize oxidative membrane damage
and increase the maize output of water usage up to 35%
[29]. Under water stress conditions the plants are closed
their stomata and unable to uptake nutrients and water so
the growth, physiology as well as the yield of plants re-
duced under water deficit conditions. Under such circum-
stance the foliar method of application of nutrients suc-
cessfully for the better growth and helpful in improving
the yield of plant under drought stress conditions. Based
on the situation the current research work was performed
under controlled conditions with aim and objective, foliar
silicon improving the yield of wheat through alteration its
physiological process under drought stress conditions.
2 Materials and Methods
Present control experiment was performed in the
Department of Agronomy, University of Agriculture
Faisalabad under greenhouse conditions. Seed of two ge-
notypes high yielded local wheat varieties, viz. Chakwal-
50 and Sehar-06, with two degrees of water tension with
following treatments i.e. control and water stress with
silicon apply in the form of seed forming (3 mM),
fertigation 1 mM and foliar application @ 4 mM in a wire
pots size (25 d × 22 l cm) were loaded with 7 kg of soil
and retained their respective field potential was main-
tained at the start of the experiment. The physiochemical
characteristics of soil which are used for this experiment
was presented in Table 1. Ten seeds have been grown and
rinsed with distilled water in each pot on a regular basis.
The plants were thinned out after 1 week of emergence
and 3 healthy plants were selected. The drought stress was
applied after the compilation of germination with two
levels such as 100% FC (control) and 60% FC (water
stress). Pots were weight on daily basis and maintain their
weight in both levels such as control and 60% FC. When
the soil is being prepared, the required dose of N urea
(110 kg N ha−1), P diammonium phosphate (70 kg P205
ha−1) and K potassium sulphate (50 kg SOP ha−1), was
poured in the soil, however, three separate doses of N
were added. All other agronomics procedure is maintained
during the whole season of wheat plant. Plants were
allowed to grow until matured, and the following methods
noted data on various bio-chemicals, plant nutrient analy-
sis and other agronomic parameters:
2.1 Pigments
[33] Methods were used to determine the chlorophyll and
carotenoid contents. The sample was centrifuged at 14000 x
g for 5 min and the supernatant absorbance was measured at
645, 652, 663 and 480 nm on a spectrophotometer (Hitachi,
U-2800). The following calculation was used to determine the
contents.


Chla mg g−1 FW
¼ ½12:7 ðOD 663Þ−2:69 ðOD 645Þ x V=1000 x W ð1Þ


Chlb mg g−1 FW
¼ ½22:9 ðOD 645Þ−4:68 ðOD 663Þ x V=1000 x W ð2Þ


Total Chlaþb mg g−1 FW
¼ ½20:2 ðOD 645Þ þ 8:02 ðOD 663Þ x V=100 x W ð3Þ


Carotenoids μg g−1 FW ¼ Acar =Emx 100 ð4Þ
where V denotes the sample extract volume, and W represents
weight.
2.2 Gas Exchange Characteristics
Each plant completely expanded youngest leaf (the second
leaf from the top) used for measurement of transpiration rate
(E), instantaneous net CO2 assimilation rate (A), and stomatal
conductance (gs) with the help of the CI-340 portable infrared
gas analyzer for photosynthesis (Analytical Development
Company, Hoddesdon, England). The data recording was tak-
en from 09 to 11.00 am with the following requirements
403.3 mmol m−2 S−1 (air leaf area), 99.9kPA (atmospheric
pressure), 1711 molm−2 s−1 (leaf PAR value), 28.4–32.4 °C
(leaf temperature), 22.4–27.9 °C (ambient temperature),
352 mol mol−1 (ambient CO2) procedure describe by the [34].
 Silicon

Table 1 Physiochemical
characteristics of the soil used for Soil Characteristics Values Soil Characteristics Values
the experiments
Chemical
ECe (dS m−1) 0.77–0.97 HCO3 (meq L−1) 4.0–4.2
Soil pH 7.6–7.9 NO3-N (mg kg−1) 11.08–14.3
Organic matter (%) 0.36–0.61 Available K (mg kg−1) 102–200
Ca + Mg (meq L−1) 3.88–5.73 Available P (mg kg−1) 7.8–10.9
CO3 (meq L−1) NIL Available Si (mg kg−1) 39–47
Physical
Soil texture Sandy loam Saturation percentage 39.5

2.3 Biochemical Parameters
The system used by [1] to assess the total soluble proteins
(TSP). Spectrophotometer (Hitachi-2800) read the optical
density (OD) at 620 nm. The amount of protein was deter-
mined using the standard curve established by varying Bovine
serum albumin (BSA) concentration. Total free amino acid
(TFA) was calculated using the Ahmad et al. (2017) methods.
Using spectrophotometer (Hitatchi, U-2800) to read the OD of
the colored solution at 570 nm. The following formula for
total free amino acid was used for estimation.


Total free amino acids μmol g−1 FW ¼ ðGraph reading of sampleÞ X ðDilution factorÞ
ð5Þ
Dilution Factor ¼ ðVolume of the sample=weight of the sampleÞ

[35] Method was used to determine the total soluble sugar. concentrations according to the above procedure. [36]
Spectrophotometer (Hitatchi, U-2800) read the absorbance at Method was used to determine the proline content and con-
625 nm. Centered on the standard curve, the concentration of centration was calculated using standard curve based on FW
soluble sugars was determined using different glucose basis by Analar grade proline.


  
m mole proline g−1 FW ¼ mg of proline mL−1 x ðmL of tolueneÞ =ðwt:of sample=5Þ =115 ð6Þ

The approach used by [36] was for calculating the
total soluble protein. The spectrophotometer (Hitachi,
U-2800) read the optical density (OD) at 620 nm. The
concentration of proteins was measured using standard
curve formed by using Bovine serum albumin (BSA)
concentrations.
2.4 Nutrients Analyses
Analysis of nutrients measured by using the procedure of [37].
The above phase was repetitive till colorless was refrigerated
digested materials. The extract volume was maintained in vol-
umetric flasks to up to 50 mL.
2.5 Yield and Yield Components
1 m square steel quadrant had been used to count tillers
number m−2. At harvest randomly designated quadrant in
allocated area was calculated and averaged the tillers counts
from each plot was observed. Productive tillers in plot were
counted from randomly selected subtracts the non-
productive tillers from the total tillers. For growing quad-
rate, 5 plants were selected to record the spike length, and
the spike length of these was determined distinctly, and
calculated the spike length average. In each quadrate five
spikes have been removed from randomly selected plant.
The spikelet’s were counted in each spike and calculated on
average.
The grains were picked at random/counted from each
quadrate and their weight was calculated using an electrical
balance. By weighing plants on an electrical balance, bio-
logical yield m−2 was determined and converted into tha−1.
After manually threshed and measuring of grains on an
electrical balance, the grain yield m−2 was measured and
translated into tha−1. For every quadrate it was reported
using formula:
Silicon
h i
HI ¼ Economic yield*=Biological yield**  100*Economic yield ¼ grain yield;**Biological yield ¼ grain þ straw
2.6 Statistical Analysis
The data collected were statistically analyzed using vari-
ance technique analysis and the program centered on
MSTAT-C Computer. The Least Significant Difference
(LSD) test was conducted to determine relevant mean at
5% probability.
3 Results
The physio chemical characteristics of the soil were presented
in the Table 1. The data was presenting based on mean square
(MS) (ANOVA) table for the effect of time and method of
silicon on gas exchange and biochemical analysis of 2 wheat
genotypes planted under control and drought was presented in
the Table 2. The individual effect i.e. stages, Si and FC re-
corded significant effect on the studied characters i.e. A, E, gs,
Chla, chlb, Chlt, CAR, TFA, LP, TSS and TSP while geno-
types recorded non-significant effect was recorded for A
(μ mol CO 2 m −2 s − 1 ), gs (μmol H 2 O m − 2 s −1 ), LP
(mmolprolineg−1 FW), TSS (mg g−1 FW) and TSP (mg g−1
FW) (Table 2).
The effect of stage (S) and silicon (Si) observed significant
effect except gs (μmol H2O m−2 s−1) and E (mmol H2O
m−2 s−1) recorded insiginificant effect of different stages and
silicon application (Table 2). The effect of different stages and
field capacity (S × FC) also recorded significant for all the
studied parameters except TSS (mg g−1 FW). The impact of
Si × FC was recorded significant for most of the characters
except A (μ mol CO2 m−2 s−1) and E (mmol H2O m−2 s−1).
The combine effect of (Si × G) was observed insignificant for
most of the studied characters except Chla (mg g−1 FW), Chlb
(mg g−1 FW) and TSP (mg g−1 FW). The interaction regarding
field capacity and genotypes (FC x G) observed significant
effect on A, gs, cha, chb, cht, CAR (μg g−1 FW), TFA (mg g−1
FW), LP (mmolprolineg−1 FW), TSS (mg g−1 FW) e, TSP
(mg g−1 FW) except E (mmol H2O m−2 s−1). The effect of
different stages, silicon application and field capacity interac-
tion (S × Si × FC) also recorded significant effect on the stud-
ied characters except E (mmol H2O m−2 s−1) and gs (μmol
H2O m−2 s−1). Similarly, the effect of different stages, silicon
application along with different wheat genotypes (S × Si × G)
recorded insignificant effect except CAR (μg g−1 FW), TFA
(mg g−1 FW) and TSP (mg g−1 FW) (Table 2). The combined
effect of (S × Si × FC × G) recorded insignificant effect for A
ð7Þ
(μ mol CO2 m−2 s−1), E (mmol H2O m−2 s−1), gs (μmol H2O
m −2 s −1 ), Chl b (mg g −1 FW), Chl t (mg g −1 FW), LP
(mmolprolineg−1 FW) and significant was recorded for Chla
(mg g−1 FW), CAR (μg g−1 FW), TFA (mg g−1 FW), TSS
(mg g−1 FW) and TSP (mg g−1 FW) (Table 2).
The yield and yield components data were presented based
on MS value from analysis of variance (ANOVA) table on
effect of time and method of silicon application of two wheat
genotypes grown under control and drought conditions were
presented in the Table 3. The individual effect of field capacity
(FC), silicon (S) and genotypes (g) recorded significant effect
for tillers number plant−1, fertile tillers plant−1, spike length
(cm), number of spikelet’s spike−1, 100-grain weight (g),
biological yield (g plant−1), grain yield (g plant−1) and HI
(%) except 100-grain weight (g) for genotypes recorded
insignificant (Table 3). The combine effect of FC × Si,
FC × G and Si × G also recorded impact on the yield and
yield components. Furthermore, FC × Si × G had been re-
corded high impact on components of yield and yield
(Table 3).
3.1 Pigments
The analysis of variance (ANOVA) data regarding impact of
time and silicone application on Ca (mg g−1 DW) and iron
(mg g−1 DW) noted significant for two genotypes for wheat
grown under natural conditions and of water stress has been
listed in the Table 2. The effect of field capacity (highly sig-
nificant), silicon application (highly significant), genotypes
(insignificant) and their interaction (FC x Si) (highly signifi-
cant), (FC x G) (highly significant), and (Si x G) (highly
significant) was recorded (Table 2).
Plants which were showing minimal to water stress condi-
tions much lower Chla (mg g−1 FW) and Chlt (mg g−1 FW)
than control while higher Chlb (mg g−1 FW) and CAR (μg g−1
FW) under well-watered conditions than drought stressed
plants. Wheat plants maintained higher Chl a contents
(mg g−1 FW), Chlb contents (mg g−1 FW), Chlt (2.49 mg g−1
FW) and CAR (μg g−1 FW) at tillering than anthesis stage
(Fig. 1). The plants were sprayed foliarly with Si and showed
optimum Chla (g−1 FW), Chlb (g−1 FW), Chlt (g−1 FW) and
CAR (μg g−1 FW) then all other treatments. The plants ex-
posed no Si exhibited minimum Chla (g−1 FW), Chlb (mg g−1
FW), Chlt (mg g−1 FW) and CAR (μg g−1 FW). The cultivars
often differed expressively (P ≤ 0.001). The genotype
Chakwal-50 was recorded maximum Chla (mg g−1 FW),
Table 2 Mean squares from analysis of variance (ANOVA) of the data for effect of time and method of silicon application on net CO2 assimilation rate (A), transpiration rate (E), stomatal conductance
(gs), chlorophyll a (mg g−1 FW), chlorophyll b (mg g−1 FW), total chlorophyll (mg g−1 FW), carotenoids (μg g−1 FW), total free amino acid (mg g−1 FW), leaf proline (mmolprolineg−1 FW), total soluble
sugars (mg g−1 FW) and total soluble proteins (mg g−1 FW) of two wheat genotypes grown under normal (100% FC) and water stress (60% FC) conditions

SOV DF A E gs Chla Chlb Chlt CAR TFA LP TSS TSP

(μ mol CO2 mˉ2 sˉ1) (mmol H2O mˉ2 sˉ1) (μmol H2O mˉ2 sˉ1) (mg g−1 FW) (mg g−1 FW) (mg g−1 FW) (μg g−1 FW) (mg g−1 FW) (mmolprolineg−1 FW) (mg g−1 FW) (mg g−1 FW)

S 1 99.166*** 19.3501*** 16.8589*** 12.0063*** 4.68608*** 31.3336*** 19.8108*** 81.7772*** 45.3200*** 61.5681*** 134.616***
Si 3 48.802*** 11.8866*** 12.5825*** 1.2508*** 0.32593*** 3.2498*** 1.2972*** 41.6592*** 55.6564*** 9.0407*** 31.188***
FC 1 524.675*** 33.8913*** 56.4727*** 8.7061*** 1.19483*** 16.2914*** 3.7170*** 56.1862*** 38.1528*** 1.4751*** 37.951***
G 1 1.116ns 2.1420** 0.0072ns 0.0473*** 0.02975*** 0.1453*** 0.1609*** 0.7183*** 0.0353ns 0.0975ns 0.032ns
S × Si 3 1.661* 0.3520ns 0.1374ns 0.0941*** 0.07619*** 0.2717*** 0.1684*** 4.0182*** 1.6874*** 1.2863*** 2.271***
S × FC 1 5.525*** 3.1176*** 2.5448*** 0.4174*** 0.15440*** 0.0664*** 0.0823** 2.0543*** 1.3680** 0.0160ns 0.313***
S×G 1 1.342ns 0.0267ns 0.2430ns 0.0011ns 0.00143ns 0.0005ns 0.0006ns 0.6789*** 0.0084ns 0.1380* 0.115***
Si × FC 3 0.710ns 0.0594ns 0.4244* 0.0091*** 0.00621*** 0.0125* 0.0592*** 1.5383*** 0.8214** 0.1128** 7.205***
Si × G 3 0.057ns 0.1490ns 0.0284ns 0.0079*** 0.01141*** 0.0040ns 0.0187ns 0.0332ns 0.1195ns 0.0291ns 0.509***
FC × G 1 14.735*** 0.0392ns 2.6103*** 0.3071*** 0.03046*** 0.5697*** 1.7147*** 10.4766*** 19.2067*** 1.2376*** 24.321***
S × Si 3 2.078** 0.1135ns 0.0896ns 0.0138*** 0.00533*** 0.0206** 0.0614*** 0.1995*** 0.9106*** 0.1252** 3.166***
× FC
S × Si 3 0.972ns 0.0902ns 0.0593ns 0.0010ns 0.00151ns 0.0051ns 0.0319* 0.3683*** 0.0686ns 0.0616ns 0.764***
×G
S × FC 1 1.356ns 0.0817ns 0.2636ns 0.0102*** 0.00143ns 0.0248* 0.0059ns 4.7763*** 0.6080* 0.0024ns 0.756***
×G
Si × FC 3 0.958ns 0.0095ns 0.1657ns 0.0037*** 0.00702*** 0.0110* 0.0507** 1.2962*** 0.1188ns 0.0157ns 1.503***
×G
S × Si 3 0.433ns 0.0888ns 0.1012ns 0.0027** 0.00088ns 0.0026ns 0.0541** 0.5644*** 0.2228ns 0.0809* 0.420***
× FC
×G
Error 62 0.442 0.1451 0.1350 0.0006 0.00078 0.0039 0.0089 0.0316 0.1376 0.0274 0.008

*, **, *** = Significant at 0.05, 0.01, 0.001 probability levels, respectively, ns = Non-significant at 5% probability level, S = Stages, Si = Silicon, G = Genotypes, FC = Field capacity levels
Silicon
Silicon

Table 3 Mean squares from analysis of variance (ANOVA) of the data biological yield (g plant−1), grain yield (g plant−1) and harvest index (%)
for effect of time and method of silicon application on plant height (cm), of two wheat genotypes grown under normal (100% FC) and water stress
dry weight plant−1 (g), total number of tillers plant−1, fertile tillers plant−1, (60% FC) conditions
spike length (cm), spikelets spike−1, grains spike−1, 100-grain weight (g),

SOV DF Total number of No. of fertile Spike No. of spikelets 100-grain Biological yield Grain yield Harvest
tillers plant−1 tillers plant−1 length (cm) spike−1 weight (g) (g plant−1) (g plant−1) index (%)

FC 1 18.2710*** 9.08090*** 18.4832*** 84.1537*** 25.6089*** 226.632*** 55.7920*** 404.275***

Si 5 4.1318*** 3.23931*** 4.2480*** 2.1427*** 0.5647*** 1.790*** 0.5183*** 6.333***
G 1 0.0369*** 0.00911*** 0.7938* 23.3017*** 0.0003ns 1.027*** 0.1013*** 68.309***
FC ×Si 5 0.0586*** 0.02516*** 0.1779ns 0.5895*** 0.0548*** 0.053*** 0.0418*** 5.592***
FC ×G 1 2.9000*** 1.55761*** 22.7813*** 78.8349*** 0.4140*** 4.764*** 1.4678*** 43.292***
Si × G 5 0.1294*** 0.00508*** 0.1130ns 0.1811*** 0.0258*** 0.096*** 0.0838*** 8.233***
FC × Si ×G 5 0.0473*** 0.04873*** 1.0086*** 0.2378*** 0.0134** 0.115*** 0.0421*** 0.718***
Error 46 0.0006 0.00070 0.1795 0.0011 0.0031 0.001 0.0015 0.047

**, *** = Significant at 0.01, 0.001 probability levels, respectively, NS Non-significant at 5% probability level, Si Silicon, G Genotypes, FC Field
capacity levels

Chlb Chlt (mg g−1 FW), and CAR (μg g−1 FW) relative to
Sehar-06 wheat genotype (Fig. 1).
3.2 Gaseous Exchange
The higher A (μmol CO2 m−2 s−1), E (mmol H2O m−2 s−1) and
gs (μmol H2O m−2 s−1) has been recorded for control condi-
tion while it was reduced for drought plants. The plants ended
up maximum A (μmol CO2 m−2 s−1), E (mmol H2O m−2 s−1)
and gs (μmol H2O m−2 s−1) at tillering relative to anthesis
stage (Fig. 2). The higher A (μmol CO2 m−2 s−1) and gs
(μmol H2O m−2 s−1) have been noted where Si was ap-
plied in foliar way while E (mmol H2O m−2 s−1) was re-
corded minimum in foliar application relative to Si
fertigation and followed by seed priming methods respec-
tively. Plants grow deprived of silicon application display
highest E (4.65 mmol H 2 O m − 2 s −1 ). The lower A
(8.58 μmol CO 2 m − 2 s − 1 ) and g s (2.69 μmol H 2 O
m−2 s−1) contents were observed without Si apply. An
insignificant result was noted for cultivars against gaseous
exchange parameters (Fig. 2).
3.3 Biochemical Parameters
The acquaintance of plants to drought stress suggested in
much higher TFA (mg g−1 FW), TSS (mg g−1 FW), LP
(m mol proline g−1 FW). High TFA was observed on the
plants than normal water conditions while the maximum un-
der regular water supply, TSP was observed while under
drought, it was minimal (mg g−1 FW), (mg g−1 FW), TSS
(mg g−1 FW), LP (m mol proline g−1 FW) at the anthesis of
rising tillers while the average TSP demonstrated at the
tillering growth stage relative to the anthesis (mg g−1 FW)
stage (Fig. 3). The use of Si as foliar recorded highest TFA
(mg g−1 FW) contents, TSS (mg g−1 FW), LP (m mol pro-
line g−1 FW) and TSP (mg g−1 FW) whereas lowest TFA
(mg g−1 FW), TSS (mg g−1 FW), LP (m mol proline g−1
FW) and TSP (mg g −1 FW) was observed without Si
application.
3.4 Yield and Yield Components
The maximum tillers count, fertile tillers, spike length (cm)
and spike/plant rather than drought. The Chakwal-50 ge-
notype recorded maximum number of tillers than Sehar-06
while Sehar-06 recorded high fertile tillers (counts), spike
length and spikelet’s spike−1 than Chakwal-50 (Fig. 4).
The silicon application methods were also significant for
this character (Table 3). The Si application with irrigation
at tillering stage recorded highest tillers production, fertile
tillers, spike length (cm) and spikelet’s spike−1 relative
control (Fig. 4). The crop was exposed to drought stress
recorded minimum 100-grain weight, BY, GY and HI rel-
ative to control (Fig. 4). The use of Si with irrigation water
recorded maximum HGW, BY and GY relative to foliar
use at anthesis stage recorded highest HI. The minimum
HGW, BY, GY and HI has been observed where applied in
plants with no Si (Fig. 4).
3.5 Nutrient Analysis
Under stressed conditions Ca (mg g-1 DW) had a lower
concentration in leaf than control treatment. The genotypes
of the wheat (Chakwal-50 and Sehar-06) were substantially
 Silicon

Fig. 1 Effect of silicon on photosynthetic pigments of wheat genotypes at two different growth stages under normal and water deficit conditions at P ≤
0.05 (The values are means of three replicates + standard error (SE)

different in the leaf Ca2+ concentration (P = .05) but signif- The varying methods of Si reported expressively for Ca
icantly higher in the wheat genotype Chakwal-50 relative to and Fe content of the leaf. The silicon application at anthesis
leaf Ca 2+ concentration (0.10 mg g −1 DW) (Fig. 5 and stage recorded highest Ca contents in leaf statistically alike
Table 4). with plants Si application with irrigation at tillering stage
Silicon

Fig. 2 Effect of silicon on gas exchange attributes of wheat genotypes at two different growth stages under normal and water deficit conditions at P ≤
0.05 (The values are means of three replicates + standard error (SE)

while highest Fe concentration was recorded in control statis-
tically equivalent to plants sprayed foliarly with Si on anthe-
sis. The lowest leaf Ca concentration had been noted in control
plants (Fig. 5). The maximum leaf accumulation of Ca
(mg g−1 DW) was reported when foliar Si was applied under
control treatment at anthesis level, at the tillering point, closely
followed by Si fertigation, while minimum leaf Ca has been
observed when Si was applied in raised beds to plants under
drought condition. However, maximum Fe concentration was
observed, the Si treatment applied where water drought
condition, and statistically similar where no Si was applied.
However, Si fertilization minimum Fe accumulation results in
leaves (0.05 mg g−1 DW) were recorded under drought con-
ditions (Fig. 5).
4 Discussions
Biochemical attributes build up in the plant under various
stresses and participate in the osmotic adjustment [2]. In this

Fig. 3 Effect of silicon on biochemical characters of wheat genotypes at two different growth stages under normal and water deficit conditions at P ≤
0.05 (The values are means of three replicates + standard error (SE)
analysis, conditions of water stress significantly enhanced
some of the biochemical characteristics (leaf complete free
amino acid, proline and soluble sugars) excluding soluble
protein in both genotypes of wheat at both growth stages.
The decline in soluble proteins under stress from drought
may be linked with the minimum protein production [38]
and increased protein breakdown and is necessary for os-
moregulation in higher quantities. At anthesis the highest
rate was observed than the tillering stage of growth under
stress with Si added foliarly except for total proteins. In
Silicon
addition, Chakwal-50 plants reported maximum biochemi-
cal characteristics than Sehar-06. Differences between ge-
notypes may be attributed to their genetic composition for
the accumulation of biochemical parameters [39].
Due to its high-water solubility, amino acid plays a signif-
icant role in defending cells against denaturation under both
limiting water and saline conditions [40]. Proline serves as
nitrogen source (N) for the plants during adaptive condition
and protects the cellular constituents against dehydration inju-
ry [41]. It also has an essential function in cytoplasmic
Silicon
Fig. 4 Effect of silicon on yield and yield components of wheat genotypes at two different growth stages under normal and water deficit conditions at
P ≤ 0.05 (The values are means of three replicates +standard error (SE)
tolerance of many plants under water limiting condition [42].
Plant tissues that represent the concentration of sugar play a
leading part in the transmission of sugar by triggering genes
responsible for translocation under drought [43]. The reduc-
tion in proteins (soluble) is well known under condition. The
decrease in soluble protein content, as o reported by [44], was
higher at anthesis rather than at tillering stage.
Nonetheless, Si might help for osmotic modification while
it requires time to be part of the metabolic pathway [22]. In
addition, Si physiologically facilitates assimilation of ammo-
nium and confines the production of soluble N complexes,
including i.e. amides and amino acids, important for hyphae
propagation [45]. A substantial increase in amino acid content
due to Si application is accountable for modification under
conditions of drought stress in sorghum roots [46]. In the
current study, this increase in leaf proline concentration under
drought stress went on to rise through the Si usage that may be
because of the high-water retention in plants. Application of Si
showed a higher degree of polysaccharides in the rice plant
cell wall of the leaves involved in cell water interactions; those
include crop mechanical properties and water permeability
[28].
 Silicon

Fig. 4 (continued)

Higher proline content due to Si usage in sunflower and root sugar contents and quality under osmosis condition [27].
potato plants under water stress conditions, respectively. The Similarly recorded an increase in the sugar contents in sor-
similar findings were consistent stated that Si use improved the ghum. In various crops, the Si mediated improvement in total
Silicon

Table 4 Mean squares from analysis of variance (ANOVA) of the data concentration [19]. Si application under drought stress was
for effect of time and method of silicon application on calcium (mg g−1
found to be more successful than well-watered situation, which
DW) and iron (mg g−1 DW) contents of two wheat genotypes grown
under normal (100% FC) and water stress (60% FC) conditions suggests that Si is a vital element for proper growth and devel-
opment under conditions of water shortage. Supplemental Si ‘s
SOV DF Ca (mg g−1 DW) Fe (mg g−1 DW) application enhanced in water relationships, gas exchange and
FC 1 3076.55*** 0.00008ns
antioxidant activity show that a single trait cannot render a
Si 5 31.48*** 0.00703***
plant immune to water, but rather complex combinations of
different traits make a plant capable of surviving in drought
G 1 1.16ns 0.00139**
conditions [47]. Si soil mediated application is more advanta-
FC ×Si 5 40.32*** 0.00474***
geous and profoundly influencing physiological processes and
FC ×G 1 138.92*** 0.00019ns
enhancing the growth of wheat under saline condition [19].
Si × G 5 3.50ns 0.00033ns
The use of Si in well-watering conditions can enhance dry
FC × Si × G 5 5.76ns 0.00013ns
weight in wheat, and it can also boost crop growth in drought
Error 46 3.26 0.00014
conditions by maintaining high leaf areas to ensure high as-
**, *** = Significant at 0.01, 0.001 probability levels, respectively, similation efficiency, cuticle thickness is significant in reduc-
NS Non-significant at 5% probability level, Ca Calcium, Fe Iron, ing water loss through evapotranspiration [1]. Drought condi-
Si Silicon, G Genotypes, FC Field capacity levels tions significantly enhanced the growth and yield characteris-
tics especially in the world’s dry or arid regions [48]. The
current study also highlighted that drought stress reduced
soluble proteins and antioxidant enzymes activity under harsh growth and yield attributes in both wheat genotypes at either
environmental condition [32]. The use of Si under water short- tillering and/or anthesis stage (plant height, dry weight
age condition, enhanced protein contents in wheat relative to plant−1, total number of tillers, fertile tillers, spike length,
no Si use under drought [22]. Different proteins in the form of grain spike-1, spikelet spike−1, grain yield, biological yield
enzymes are gathered in cells of different plant species as a and HI). Many scientists published similar findings in aquat-
result to Si accumulation. Among others, photosynthesis, re- ically stressed soybean and wheat plants. The yield loss under
dox homeostasis, pathogen response and other protein devel- drought may have resulted from inhibiting enough transloca-
opment are mostly regulated [42]. The Si application increased tion of grain assimilates, plants fought for water contents in
the yield of wheat grain by stimulating physical characteristics, the root zone [33]. Furthermore, declining water and gas-
managed photosynthetic activity and enhanced growth by im- exchange activity (as recorded in current study) could have
proving the stability of the cell membranes and cell Si been attributable to reduced yields, since reductions in these

Fig. 5 Effect of silicon on calcium (Ca) and iron (Fe) contents of wheat genotypes at two different growth stages under normal and water deficit
conditions at P ≤ 0.05 (The values are means of three replicates + standard error (SE)

physiological activities, crop yields particularly in wheat are
severely limited [35]. In response to drought, an important
difference was suggested between drought tolerant
(Chakwal-50) and responsive (Sehar-06) genotypes, which
is consistent with old findings [49]. The Sehar-06 (drought
sensitive) genotype gave good results relative to Chakwal-50
which in water deficit conditions was found to be more recep-
tive to Si application. The conservation of photosynthetic pig-
ments and increased osmolytes deposition and production of
antioxidant enzymes and Chakwal-50 they played a crucial
role in the increase of grain yield. The exogenous silicon
application enhanced yield traits of many crops [36]. The
findings of the current study supported the findings that
under natural and drought conditions, exogenous Si appli-
cation substantially enhanced wheat growth and yield
characteristics. Application of Si with irrigation has been
better output than other approaches, however. Si foliar
spray has been identified as the second-best approach to
augment the growth and development of both genotypes
in wheat. If application enlarged, the yield of wheat grain
by to impel physical characteristics and maintaining pho-
tosynthesis cycle and increased growth by boosting cell
membrane stability and Si concentration within cells
[13]. Si application has been found to be more effective
when compared with drought stress to towel watered con-
dition which indicates that Si is a required element and
occurrence of its appropriate quantity in growth medium
plant survival guarantees under drought conditions.
Supplemental Si′s substantial enhances in water relation-
ships, gas exchange and antioxidant activity show that a
single characteristic cannot make a plant water-resistant,
but rather complex combinations of different characteristics
allow a plant to survive in drought conditions [1]. Si in
well-watering conditions might increase wheat dry matter,
and it can also boost crop growth in drought conditions by
holding high leaves area to ensure higher assimilation effi-
ciency, cuticle thickness is helpful in reducing water loss
through transpiration [47].
5 Conclusion
Current controlled research study concluded that the exoge-
nous application of Si at the anthesis stage helpful in wheat
growth and altered its physiological process by improving the
yield under water limited conditions. Si application in wheat
through foliar at anthesis stage and fertigtaion at tillering stage
were performed better in enhancing the growth, physiological
process and yield under drought stress condition. So the ap-
plication of Si is good practice in wheat by improving its
tolerance against drought stress with better growth and yield
response.
Silicon
Author Contributions MAD and MYA plan the research and MAD con-
duct research under supervision of MYA; ZA and ANS write the intro-
duction part; NAM and WNJ write the manuscript; AM and MN help to
statically analysis and graphical representation; FN read the manuscript as
proofreading; MA and MYA help in English editing and final formatting
according to journal style.
Compliance with Ethical Standards
Conflict of Interest The authors declare that they have no conflict of
interest.
References
1. Ahmad Z, Waraich EA, Barutçular C, et al (2020) Enhancing
drought tolerance in wheat through improving morpho- physiolog-
ical and antioxidants activities of plants by the supplementation of
foliar silicon. https://doi.org/10.32604/phyton.2020.09143
2. Waraich EA, Rashid F, Ahmad Z, Ahmad R, Ahmad M (2020)
Foliar applied potassium stimulate drought tolerance in canola un-
der water deficit conditions. J Plant Nutr 0:1–12. https://doi.org/10.
1080/01904167.2020.1758132, 43
3. Ahmad Z, Anjum S, Iqbal MA, Saeed-ur-Rehman H (2018) Foliar
applied potassium enhances fibre quality, water foliar applied po-
tassium enhances fibre quality, water relations and yield of cotton. J
Agric Res 56:17–25
4. Gao L, Caldwell CD, Jiang Y (2018) Photosynthesis and growth of
camelina and canola in response to water deficit and applied nitro-
gen. Crop Sci 58:393–401. https://doi.org/10.2135/cropsci2017.07.
0406
5. Ahmad Waraich E, Ahmed Z, Ahmad Z et al (2020) Alterations in
growth and yield of Camelina induced by different planting densi-
ties under water deficit stress. Phyton (B Aires) 89:587–597. https://
doi.org/10.32604/phyton.2020.08734
6. Seyahjani EA, Yarnia M, Farahvash F et al (2020) Influence of
rhizobium, pseudomonas and mycorrhiza on some physiological
traits of red beans (Phaseolus vulgaris l.) under different irrigation
conditions. Legum Res. https://doi.org/10.18805/LR-454
7. Mohtashami R, Movahhedi Dehnavi M, Balouchi H, Faraji H
(2020) Improving yield, oil content and water productivity of dry-
land canola by supplementary irrigation and selenium spraying.
Agric Water Manag 232:106046. https://doi.org/10.1016/j.agwat.
2020.106046
8. Ahmad Z, Anjum S, Waraich EA, Ayub MA, Ahmad T, Tariq
RMS, Ahmad R, Iqbal MA (2018) Growth, physiology, and bio-
chemical activities of plant responses with foliar potassium appli-
cation under drought stress–a review. J Plant Nutr 41:1734–1743
9. Mahmood A, Kanwal H, Kausar A et al (2019) Seed priming with
zinc modulate growth, pigments and yield of chickpea (Cicer
arietinum l.) under water deficit conditions. Appl Ecol Environ
Res. https://doi.org/10.15666/aeer/1701_147160
10. Shahsavari N (2019) Effects of zeolite and zinc on quality of canola
(Brassica napus L.) under late season drought stress. Commun Soil
Sci Plant Anal. https://doi.org/10.1080/00103624.2019.1604729
11. Sharma M, Gupta SK, Majumder B, Maurya VK, Deeba F, Alam
A, Pandey V (2017) Salicylic acid mediated growth, physiological
and proteomic responses in two wheat varieties under drought
stress. J Proteome 163:28–51. https://doi.org/10.1016/j.jprot.2017.
05.011
12. El Sabagh A, Hossain A, Barutçular C et al (2019) Effects of
drought stress on the quality of major oilseed crops: implications
and possible mitigation strategies - a review. Appl Ecol Environ
Res 17:4019–4043. https://doi.org/10.15666/aeer/1702_40194043
Silicon
13. Zhang W, Yu X, Li M, Lang D, Zhang X, Xie Z (2018) Silicon
promotes growth and root yield of Glycyrrhiza uralensis under salt
and drought stresses through enhancing osmotic adjustment and
regulating antioxidant metabolism. Crop Prot 107:1–11. https://
doi.org/10.1016/j.cropro.2018.01.005
14. Sallam A, Alqudah AM, Dawood MFA, Baenziger PS, Börner A
(2019) Drought stress tolerance in wheat and barley: advances in
physiology, breeding and genetics research. Int J Mol Sci 20
15. Guo R, Shi L, Jiao Y, Li MX, Zhong XL, Gu FX, Liu Q, Xia X, Li
HR (2018) Metabolic responses to drought stress in the tissues of
drought-tolerant and drought-sensitive wheat genotype seedlings.
AoB Plants 10. https://doi.org/10.1093/aobpla/ply016
16. Barnawal D, Bharti N, Pandey SS, Pandey A, Chanotiya CS, Kalra A
(2017) Plant growth-promoting rhizobacteria enhance wheat salt and
drought stress tolerance by altering endogenous phytohormone levels
and TaCTR1/TaDREB2 expression. Physiol Plant 161:502–514.
https://doi.org/10.1111/ppl.12614
17. Ma J, Li R, Wang H, Li D, Wang X, Zhang Y, Zhen W, Duan H,
Yan G, Li Y (2017) Transcriptomics analyses reveal wheat re-
sponses to drought stress during reproductive stages under field
conditions. Front Plant Sci 8. https://doi.org/10.3389/fpls.2017.
00592
18. Fani E, Hassibi P, Meskarbashee M et al (2019) Effect of drought
stress and silica spraying on some physiological and functional
traits of canola cultivars. Bulg J Agric Sci 25:62–66
19. Souri Z, Khanna K, Karimi N, Ahmad P (2020) Silicon and plants:
current knowledge and future prospects. J Plant Growth Regul.
https://doi.org/10.1007/s00344-020-10172-7
20. Cao B-L, Wang L, Gao S et al (2017) Silicon-mediated changes in
radial hydraulic conductivity and cell wall stability are involved in
silicon-induced drought resistance in tomato. Protoplasma 254:
2295–2304. https://doi.org/10.1007/s00709-017-1115-y
21. Chen D, Wang S, Yin L, Deng X (2018) How does silicon mediate
plant water uptake and loss under water deficiency? Front Plant Sci
9. https://doi.org/10.3389/fpls.2018.00281
22. Laane HM (2017) The effects of the application of foliar sprays
with stabilized silicic acid: an overview of the results from 2003-
2014. Silicon. 9:803–807. https://doi.org/10.1007/s12633-016-
9466-0
23. Laane HM (2018) The effects of foliar sprays with different silicon
compounds. Plants 7(2):45
24. Agostinho FB, Tubana BS, Martins MS, Datnoff LE (2017) Effect
of different silicon sources on yield and silicon uptake of rice grown
under varying phosphorus rates. Plants. 6. https://doi.org/10.3390/
plants6030035
25. Artyszak A (2018) Effect of silicon fertilization on crop yield quan-
tity and quality—A literature review in Europe. Plants 7(3):54
26. Schaller J, Turner BL, Weissflog A, Pino D, Bielnicka AW,
Engelbrecht BMJ (2018) Silicon in tropical forests: large variation
across soils and leaves suggests ecological significance.
Biogeochemistry. 140:161–174. https://doi.org/10.1007/s10533-
018-0483-5
27. Gou T, Yang L, Hu W, Chen X, Zhu Y, Guo J, Gong H (2020)
Silicon improves the growth of cucumber under excess nitrate stress
by enhancing nitrogen assimilation and chlorophyll synthesis. Plant
Physiol Biochem 152:53–61. https://doi.org/10.1016/j.plaphy.
2020.04.031
28. Gao M, Zhou J, Liu H, Zhang W, Hu Y, Liang J, Zhou J (2018)
Foliar spraying with silicon and selenium reduces cadmium uptake
and mitigates cadmium toxicity in rice. Sci Total Environ 631-632:
1100–1108. https://doi.org/10.1016/j.scitotenv.2018.03.047
29. Khan W-D, Aziz T, Iqbal M, Ramzani PMA, Rasool K, Rahman
SU, Anwar MN (2017) WITHDRAWN: soil and foliar silicon ap-
plication enhances maize grain yield, its proximate composition,
and antioxidant mechanisms under saline field conditions. Plant
Physiol Biochem. https://doi.org/10.1016/j.plaphy.2017.08.023
30. Deus ACF, de Mello PR, de Cássia Félix Alvarez R et al (2020)
Role of silicon and salicylic acid in the mitigation of nitrogen defi-
ciency stress in Rice plants. Silicon. 12:997–1005. https://doi.org/
10.1007/s12633-019-00195-5
31. Ali S, Rizwan M, Hussain A et al (2019) Silicon nanoparticles
enhanced the growth and reduced the cadmium accumulation in
grains of wheat (Triticum aestivum L.). Plant Physiol Biochem.
https://doi.org/10.1016/j.plaphy.2019.04.041
32. Merwad ARMA, Desoky ESM, Rady MM (2018) Response of
water deficit-stressed Vigna unguiculata performances to silicon,
proline or methionine foliar application. Sci Hortic (Amsterdam)
228:132–144. https://doi.org/10.1016/j.scienta.2017.10.008
33. Ahmad Z, Waraich EA, Ahmad R, Shahbaz M (2017) Modulation
in water relations, chlorophyll contents and antioxidants activity of
maize by foliar phosphorus application under drought stress. Pak J
Bot 49:11–19
34. Waraich EA, Ahmad Z, Ahmad R, Saifullah, Ashraf MY (2015)
Foliar applied phosphorous enhanced growth, chlorophyll contents,
gas exchange attributes and PUE in wheat (Triticum aestivum L.). J
Plant Nutr 38:1929–1943. https://doi.org/10.1080/01904167.2015.
1043377
35. Waraich EA, Ahmed Z, Ahmad Z, et al (2020) Alterations in
growth and yield of Camelina induced by different planting densi-
ties under water De fi cit stress. 1–11. https://doi.org/10.32604/
phyton.2020.08734
36. Nawaz F, Ahmad R, Ashraf MY, Waraich EA, Khan SZ (2015)
Effect of selenium foliar spray on physiological and biochemical
processes and chemical constituents of wheat under drought stress.
Ecotoxicol Environ Saf 113:191–200. https://doi.org/10.1016/j.
ecoenv.2014.12.003
37. Nawaz F, Ashraf MY, Ahmad R, Waraich EA, Shabbir RN,
Bukhari MA (2015) Supplemental selenium improves wheat grain
yield and quality through alterations in biochemical processes un-
der normal and water deficit conditions. Food Chem 175:350–357.
https://doi.org/10.1016/j.foodchem.2014.11.147
38. Coskun D, Britto DT, Huynh WQ, Kronzucker HJ (2016) The role
of silicon in higher plants under salinity and drought stress. Front
Plant Sci 7. https://doi.org/10.3389/fpls.2016.01072
39. Akinwale RO, Awosanmi FE, Ogunniyi OO, Fadoju AO (2017)
Determinants of drought tolerance at seedling stage in early and
extra-early maize hybrids. Maydica 12;62(1):9
40. Ahmad R, Waraich EA, Nawaz F, Ashraf MY, Khalid M (2016)
Selenium (se) improves drought tolerance in crop plants - a myth or
fact? J Sci Food Agric 96:372–380
41. Laxa M, Liebthal M, Telman W, Chibani K, Dietz KJ (2019) The
role of the plant antioxidant system in drought tolerance.
Antioxidants 8(4):94
42. Hemmati M, Delkhosh B, Rad AHS, Mohammadi GN (2019)
Effect of the application of foliar selenium on canola cultivars as
influenced by different irrigation regimes. Tarim Bilim Derg.
https://doi.org/10.15832/ankutbd.424899
43. Choudhury FK, Rivero RM, Blumwald E, Mittler R (2017)
Reactive oxygen species, abiotic stress and stress combination.
Plant J 90:856–867. https://doi.org/10.1111/tpj.13299
44. Ahmad M, El-Saeid MH, Akram MA et al (2016) Silicon fertiliza-
tion – a tool to boost up drought tolerance in wheat (Triticum
aestivum L.) crop for better yield. J Plant Nutr 39:1283–1291.
https://doi.org/10.1080/01904167.2015.1105262
45. Kim YH, Khan AL, Waqas M, Lee IJ (2017) Silicon regulates
antioxidant activities of crop plants under abiotic-induced oxidative
stress: a review. Front Plant Sci 8:1–7. https://doi.org/10.3389/fpls.
2017.00510
46. Asgharipour MR, Mosapour H (2016) A foliar application silicon
enchances drought tolerance in fennel. J Anim Plant Sci 26:1056–
1062
 Silicon

47. Ahmad Z, Waraich EA, Barutçular C et al (2020) Enhancing drought 49. Schillinger WF (2019) Camelina: long-term cropping systems re-
tolerance in wheat through improving morphophysiological and an- search in a dry Mediterranean climate. F Crop Res 235:87–94.
tioxidants activities of plants by the supplementation of foliar silicon. https://doi.org/10.1016/j.fcr.2019.02.023
Phyton (B Aires). https://doi.org/10.32604/phyton.2020.09143
48. Waraich EA, Ahmad R, Ahmad R et al (2020) Comparative study Publisher’s Note Springer Nature remains neutral with regard to jurisdic-
of growth, physiology and yield attributes of camelina (Camelina tional claims in published maps and institutional affiliations.
sativa L.) and canola (Brassica napus L.) under different irrigation
regimes. Pak J Bot 52:1537–1544. https://doi.org/10.30848/
PJB2020-5(2)