Nazim 2024

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Journal of Crop Health

https://doi.org/10.1007/s10343-024-01005-8

RESEARCH

Exogenous Potassium Silicate Improves Drought Tolerance in Cotton


Genotypes by Modulating Growth, Gas Exchange and Antioxidant
Metabolism
Muhammad Nazim1,2,3,4 · Xiangyi Li1,2,3,4 · Akash Tariq1,2,3,4,5,6 · Khurram Shahzad7 · Murad Muhammad1,2,4 ·
Zhang Dawei8 · Fanjiang Zeng1,2,3,4

Received: 11 April 2024 / Accepted: 10 June 2024


© Der/die Autor(en), exklusiv lizenziert an Springer-Verlag GmbH Deutschland, ein Teil von Springer Nature 2024

Abstract
Cotton is a highly sensitive crop to drought stress. Consequently, it is crucial to devise strategies that optimize crop
production in conditions of limited water availability. While potassium silicate has demonstrated effectiveness in mitigating
drought stress in various crops, its specific impact on different cotton cultivars under drought conditions remains not fully
clarified. This research aimed to assess the efficacy of six potassium silicate levels (0, 100, 200, 300, 400 and 500 mg L–1)
on four cotton genotypes (Zong main-113, Xin Nong-525, Xin lu Zhong-55, and Xin lu Zhong-66) under two field
capacity levels (80% and 50% FC) in a sand culture. Foliar applied potassium silicate significantly improved photosynthetic
efficiency, shoot biomass, root biomass, and leaf area under water stress (50% FC). The most substantial reduction in H2O2,
malondialdehyde levels, and electrolyte leakage was recorded with potassium silicate applied at a rate of 400 mg L–1. This
concentration effectively mitigated reactive oxygen species accumulation, safeguarding plants against oxidative damage at
50% FC. Furthermore, potassium silicate contributed to maintaining water status, resulting in increased leaf water content
and elevated water-soluble proteins in cotton plants. The order of drought resistance (50% FC) with the application of
potassium silicate at 400 mg L–1 was Zong Mian-113, Xin Nong-525, Xin lu Zhong-55, and Xin lu Zhong-66. The findings
could help in selection of drought resistance cultivars of cotton in water limited conditions.

Keywords Antioxidant activities · Biochemical performance · Gossypium hirsutum · K2SiO3 · Photosynthetic


pigments · Sandy soil · Water Deficit Stress

Introduction regions worldwide (El Sabagh et al. 2020; Nazim et al.


2021). Approximately 64% of the world’s soil is situated
Drought stands as a significant ecological factor that in deserts or regions with water scarcity, with 57% of
severely impacts food quality and global agricultural crop the potentially arable land designated for dry-land crops
yield productivity, predominantly in arid and semi-arid (Chen et al. 2022). Water, which constitutes 80–95% of

 Xiangyi Li
[email protected]

1 4
Xinjiang Key Laboratory of Desert Plant Roots Ecology and University of Chinese Academy of Sciences, 100049 Beijing,
Vegetation Restoration, Xinjiang Institute of Ecology and China
Geography, Chinese Academy of Sciences, 830011 Urumqi, 5
CSIC, Global Ecology Unit, CREAF-CSIC-UAB,
China
08193 Bellaterra, Barcelona, Catalonia, Spain
2
State Key Laboratory of Desert and Oasis Ecology, 6
CREAF, 08193 Cerdanyola Del Vall’es, Catalonia, Spain
Key Laboratory of Ecological Safety and Sustainable
7
Development in Arid Lands, Xinjiang Institute of Department of Soil Science, Faculty of Agriculture, Lasbela
Ecology and Geography, Chinese Academy of Sciences, University of Agriculture, Water and Marine Sciences,
830011 Urumqi, China 90150 Uthal, Balochistan, Pakistan
3 8
Cele National Station of Observation and Research for Economic Crop Research Institute, Xinjiang Academy of
Desert-Grassland Ecosystems, 848300 Cele, China Agricultural Sciences, 830091 Urumqi, China

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M. Nazim et al.

crop biomass, is integral to numerous physiological pro- enhances plant physiology by improving seed germina-
cesses crucial for plant survival (Farouk and Omar 2020; tion, maintaining cell membranes, regulating water status,
Muhammad et al. 2023). Drought profoundly affects vari- promoting osmolytes accumulation, regulating stomata, in-
ous aspects of plant life, impacting morpho-physiological, creasing water-use efficiency, boosting photosynthesis, and
physio-biochemical, and metabolic functions at all stages enhancing antioxidant defenses (Wang et al. 2021).
of growth and development (Abeed et al. 2021; Zafar et al. Considering its significant nature of this concept offers
2023), Processes like photosynthetic pigments production, numerous potential benefits to plant functions and has been
uptake of water and essential nutrients, osmotic balance, massive interest in evaluating different sources of silicon
as well as the regulation of reactive oxygen species (ROS) (Si) for the improvement of abiotic stress tolerance in
are significantly impaired under drought condition (Wahab various agricultural crops (Guntzer et al. 2012; Ahmed
et al. 2022). Plants combat drought by growing extensive et al. 2023). Consequently, foliar applied potassium sil-
roots for water uptake, reducing leaf area to minimize icate improved the tolerances of cotton to salinity stress
water loss, accumulating protective solutes, and enhancing (Ashraf et al. 2022), rapeseed (Brassica napus L.) culti-
antioxidant defenses (Seleiman et al. 2021). Cotton stands vars to drought (Shirani Rad et al. 2022), maize crop (Zea
as most important and top economic fiber crop, is derived mays L.) (Gomaa et al. 2021), (Karvar et al. 2023), Banana
from Gossypium spp., cultivated extensively worldwide, (Musa acuminata L.) (Aziz et al. 2023), Faba bean (Vi-
particularly in China, the United States (USA), India, and cia faba L.) (Fouda et al. 2022), Ajwain (Trachyspermum
Pakistan (Jabran et al. 2019). Despite its comparatively ammi L.) (Sarhan and Shehata 2023), Sorghum (Sorghum
much tolerance to abiotic stresses compared to others crop, bicolor L.) (Ahmed et al. 2011), Wheat (Triticum aes-
extreme environmental factors like droughts, salinity, and tivum L.) (Khalid et al. 2022), and other crop are reported.
temperature significantly impact both yield and fiber qual- The studies showed the imperative use of potassium silicate
ity (Esmaeili et al. 2021; Zafar et al. 2022). In China, in plant abiotic stress and their management. Nevertheless,
the largest cotton producer globally, there has been drastic conducting further research at the morpho-physiological
reduction of 70% above the last four decades (Wang and and biochemical levels would be helpful to illuminate the
Memon 2020). Similarly, worldwide cotton productivity underlying mechanisms behind potassium silicate medi-
yield losses due to drought stress range between 50 and ated drought stress adaptation in cotton crop. The existing
73% (Khan et al. 2018). Addressing these challenges is literature on the effects of potassium silicate on cotton
crucial not only to improve cotton growth and yield pro- cultivars under drought stress is limited. Therefore, this
ductivity in light of recent severe drought condition but also study hypothesized that potassium silicate could mitigate
to sustain a steady supply of raw-materials for the garment the adverse impacts of water scarcity by promoting veg-
(Clothing) textile (cotton ginning) industries (Wang et al. etative growth and inducing physiological changes. The
2016; Nadeem et al. 2023). main objective of this research was to determine the order
To alleviate drought induced adversative effects on cot- of drought resistance among four commonly grown cotton
ton, several bio-technological, genetically modified and cultivars with different potassium silicate applications.
breeding approaches, are being employed (Sultana et al.
2023; Batool et al. 2023). However, several farmers in
some countries are unable to reached these technologies Materials and Methods
due to their excessive cost, prolonged investigation dura-
tion, and rarely ethical hurdles (Nawaz et al. 2023). Under Experiment Site
these prevailing situations, the exogenous application of
nutrients such as potassium silicate may undoubtedly work The pot experiment was conducted at the Cele National Re-
as an alternate, very simple, less effort time consuming, search Observatory for Desert Ecosystem in Xinjiang under
and technique that can enhanced plant resilience to against controlled greenhouse conditions. The greenhouse temper-
adverse effect of droughts stress in plant (Nazim et al. ature was 32 ± 2 °C during the day and 23 ± 2 °C at night,
2024). Potassium silicate acts as an essential factor for with a relative humidity of around 45–50%, and a photope-
stimulating various biological processes, which ultimately riod of 14 h. The soil used was collected from the experi-
increases the plant’s growth potential and existence, par- mental field of the Cele National Research Observatory for
ticularly under stressed environments (Souri et al. 2021). Desert Ecosystem in Xinjiang. It was air-dried and then
It has Synergetic role in plant functions as a co-factor of sieved with a 2 mm sieve. The soil was identified as aeolian
above 300 plus enzymes, the regulating of gene expres- loamy sand (70.13% Sand, 0.96% Silt, 28.91% Clay) with
sion (RGE) and protein synthesis, phosphate and glucose organic carbon content of 2.99 g kg–1, total nitrogen content
metabolism, and maintaining ribosome structural integrity of 0.23 g kg–1, total phosphorus content of 0.60 g kg–1, total
(Cakmak et al. 2023). In drought stress, potassium silicate

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Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

potassium content of 23.11 g kg–1, electrical conductivity of envelope and subsequently put in oven drying at temper-
0.43 dS m–1 and pH of 8.2. ature of 72 °C until a consistent weight was attained. The
weights of the dried samples were then recorded using an
Cotton Cultivars and Potassium Silicate electric balance. To assessment of leaf area (cm2) and leaf
succulence were estimated using 3rd leaf from the bottom
The seeds of four cultivars (Zong Mian-113, Xin Nong-525, of each plant was selected, following the formulas devel-
Xin lu Zhong-55, and Xin lu Zhong-66) were collected from oped by Carleton and Foote (1965). Leaf electrolyte leakage
Economic Crop Research Institute, Xinjiang Academy of (LEL) and RLWCs were measured following the methods
Agricultural Sciences, Urumqi, China. Vigorous, pests and described by Rahman et al. (2019) and (Das et al. 2022),
disease-free seeds were sterilized for 20 min with 30% (w/v) respectively.
hydrogen peroxide (H2O2), and then repeatedly washed and A portable infrared gas analyzer apparatus (LI-6400XT,
soaked abundantly with distilled water. After that, wipe the LI-COR Inc., Lincoln, NE, USA) was used in completed
water off the cotton seeds surface with a filter and set the sunlight between 11:00 A.M. and 1:00 P.M. to measure the
seeds aside. The potassium silicate (K2SiO3) 1.57:1 was photosynthetic (Pn), transpiration rate (E), leaf temperature
purchased from Shanghai YuanYe Biotechnology company (LT), and stomatal conductance (gs). Leaf extracts were got
the China. Potassium silicate was applied in concentrations by using 80% acetone to determine the levels of carotenoids
of 0, 100, 200, 300, 400, and 500 mg L–1. Two levels of (Cars.) and Chlorophylls (Chl-a, Chl-b, and total-Chls) by
field capacity (80 ± 5% and 50 ± 5%) were maintained at using the procedures outlined of (Lichtenthaler and Well-
the four-leaf stage. burn 1983).
The procedures defined by Yu et al. (2003) and Kim et al.
Seedling Establishment (2020) were used to quantify the quantities of malondialde-
hyde (MDA) and hydrogen peroxide (H2O2) in cotton leaf
The pots, each with a diameter of 15 cm and a depth of tissues.
20 cm, were filled with sandy loam soil (aeolian). The ex- Cotton fresh leaves were used in the preparation of en-
periment followed a completely randomized design (CRD) zyme extracts. The activities of antioxidant enzymes includ-
with factorial and three replications per experimental unit. ing guaiacol peroxidase (GPOD), glutathione S-transferase
Five seeds of each cultivar were sown, and after germi- (GST), catalase (CAT), and ascorbate peroxidase (APX)
nation, two seedlings were maintained in each pot. Field were then determined in accordance with the procedures
capacity levels of 80 ± 5 and 50 ± 5% were established at presented by Rahman et al. (2019).
the onset of the four-leaf stage of cotton. Foliar application Bates et al. (1973) described, Acid-ninhydrin method is
of potassium silicate was carried out 30 days after sowing a widely used method for proline content, a little renewed to
the seeds, and following the onset of drought stress. Regular determine the proline content level. First, take (0.625 g) of
weighing of pots around 1:00 pm enabled the measurement ninhydrin was collective with (15 mL) of glacial acetic acid
of water consumption through evapotranspiration from each and (10 mL) of (6 M) Orthophosphoric acid to make acid-
pot (Allen et al. 1998). Subsequently, pots were watered to ninhydrin. However, proceeds a fresh cotton leaf weigh-
predetermined weights. The plants were harvested 46 days ing about (0.1 g) were normalized in (2.5 mL) of an aque-
after germination. ous solution of (3%) sulfosalicylic acid (extraction buffer),
and then centrifuged at (12,000 rpm) for 10 min. The reac-
Data Collection tion was then stopped by placing the glass tube contain-
ing (0.5 mL) of the supernatant, (0.5 mL) of acid-ninhydrin,
Following the imposition of drought stress, gas exchange and (0.5 mL) of glacial acetic in the freezer for a further
characteristics were measured in 46-day-old plants. Analy- hour. After that, (1.5 mL) of toluene was supplementary
sis of a comprehensive set morpho-physio-biochemical pa- added to the reactant solution, which was then vortexed
rameters were conducted on the 3rd leaves (from the bot- for 10 s and left to stand at room temperature for 10 min.
tom) freshly harvested from the 46-day-old plants. The concentration of proline was estimated using a stan-
The response of different variables including growth at- dard curve, as described by (Bates et al. 1973). The water
tributes such as shoot and root length, shoot and root fresh soluble protein (WSP) content of the cotton leaf samples
weight, and shoot and root dry weight were measured at was measured using the Bradford-protein-test, as defined
maturity and selected healthy plant from each pot were by (Bradford 1976). The total free amino acid content was
harvested and shoot and root lengths were measured with calculated following the methodology described by Lee and
measuring scale. Then shoot and root fresh weights were Takahashi (1966).
taken using an electric balance and data were recorded. The
plants sample were wisely arranged by placing them in an

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M. Nazim et al.

Statistical Analysis length 38.57%, 70.58%, 94.11%, 188.01 and 129.15% as


compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas-
The collected data underwent analysis using R software sium silicates, respectively. However, in variety Xin Nong-
(Version 4.3.1) through the application of a linear mixed 525, under drought stress, potassium silicate application at
model. The cotton cultivars, potassium silicates levels and the rate of 400 mg L–1 increased the root length 48.18%,
drought levels were considered as fixed effect while repli- 96.37%, 128.51%, 257.07 and 176.22% as compared to
cation as random effect. Prior to statistical analysis, data 0, 100, 200, 300, 400 and 500 mg L–1 potassium silicates, re-
normality was verified. Mean comparisons were compared spectively (Table 1). The application of potassium silicates
conducted using least-square means, and Tukey’s multiple led to an increase in shoot length, peaking at a 400 mg L–1
comparison procedure using the “emmeans” package at p < application rate. Beyond this concentration, however, shoot
0.05. All plots were generated using the ggplot package length exhibited a declining trend. In variety, Zong main-
in R. 113, under drought stress, potassium silicate application
at the rate of 400 mg L–1 increased the shoot length by
17.38%, 21.12%, 31.77%, 57.08 and 21.82% as compared
Results to 0, 100, 200, 300, 400 and 500 mg L–1 potassium sil-
icates, respectively. However, in variety Xin Nong-525,
Root and Shoot Length under drought stress, potassium silicate application at the
rate of 400 mg L–1 increased the shoot length by 19.78%,
The normal irrigation showed higher root and shoot length 24.25%, 36.12%, 65.01 and 24.85% as compared to 0, 100,
in all varieties and potassium silicate levels. However, 200, 300, 400 and 500 mg L–1 potassium silicates, respec-
drought stress had an adverse effect on root and shoot tively (Table 1). At the higher concentration 500 mg L–1
length across all varieties shown in Table 1. Nevertheless, of potassium silicate was observed did not display such
when potassium silicates were applied, there was a notice- an obvious promotion in root and shoot length parameters
able improvement in root and shoot length for all varieties in cotton plant. At the higher concentration 500 mg L–1
under drought-stress conditions. The application of potas- of potassium silicate, phytotoxic effects were observed in
sium silicates led to an increase in root length, peaking at the form of premature yellowed and burnt leaves in cotton
a 400 mg L–1 application rate. Beyond this concentration, plant (Fig. 1).
however, root length exhibited a declining trend. In variety,
Zong main-113, under drought stress, potassium silicate
application at the rate of 400 mg L–1 increased the root

Fig. 1 Effect of foliar-applied Potassium Silicate on vegetative performance of four cotton genotypes under drought stress

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Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

Shoot and Root Dry Weight as compared to 0, 100, 200, 300, 400 and 500 mg L–1
potassium silicates, respectively (Table 1). However, in
The normal irrigation showed higher shoot dry weight variety Xin Nong-525, under drought stress, potassium
(SDW) and root dry weight (RDW) in all varietal compres- silicate application at the rate of 400 mg L–1 increased the
sion and potassium silicate levels. However, drought stress ILA 16.67%, 29.17%, 41.68%, 50.01% and 33.32% as
had an adverse effect on SDW across all varieties. Never- compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas-
theless, when potassium silicates were applied, there was sium silicates, respectively (Table 1). The normal irrigation
a noticeable improvement in SDW and RDW for all vari- showed increased Carotenoids (Car) in all varietal compres-
eties under drought-stress conditions displayed in Table 1. sion and potassium silicate levels. However, drought stress
The application of potassium silicates led to an increase in had an adverse effect on Car across all varieties exhibited
SDW, peaking at a 400 mg L–1 application rate. Beyond this in Fig. 2b. Nevertheless, when potassium silicates were
concentration, however, SDW exhibited a declining trend. applied, there was a noticeable improvement in Car for all
In variety, Zong main-113, under drought stress, potassium varieties under drought-stress conditions. The application
silicate application at the rate of 400 mg L–1 increased the of potassium silicates led to an increase in Car, peaking
SDW by 14.16%, 71.67%, 78.75%, 99.61% and 84.07% as at a 400 mg L–1 application rate. Beyond this concentra-
compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas- tion, however, Car exhibited a declining trend. In variety,
sium silicates, respectively (Table 1). However, in variety Zong main-113, under drought stress, potassium silicate
Xin Nong-525, under drought stress, potassium silicate application at the rate of 400 mg L–1 increased the Car
application at the rate of 400 mg L–1 increased the SDW 33.31%, 66.67%, 116.41%, 201.07%, and 151% as com-
by 34.76%, 176.08%, 193.47%, 245.65% and 206.52% as pared to 0, 100, 200, 300, 400 and 500 mg L–1 potassium
compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas- silicates, respectively (Fig. 2b). However, in variety Xin
sium silicates, respectively (Table 1). The application of Nong-525, under drought stress, potassium silicate appli-
potassium silicates led to an increase in SDW, peaking at cation at the rate of 400 mg L–1 increased the Car 19.76%,
a 400 mg L–1 application rate. Beyond this concentration, 33.32%, 119.81%, 218.84% and 159.76% as compared to
however, RDW exhibited a declining trend. In variety, 0, 100, 200, 300, 400 and 500 mg L–1 potassium silicates,
Zong main-113, under drought stress, potassium silicate respectively (Fig. 2b).
application at the rate of 400 mg L–1 increased the RDW
by 31.43%, 114.27%, 142.85%, 177.14% and 134.27% as Leaf Temperature and Electrolyte Leakage Levels
compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas-
sium silicates, respectively (Table 1). However, in variety The normal irrigation showed decreased Leaf Temperature
Xin Nong-525, under drought stress, potassium silicate (LT) and Electrolyte leakage levels (EL) in all varietal com-
application at the rate of 400 mg L–1 increased the RDW by pression and potassium silicate levels. However, drought
35.48%, 129.03%, 161.29%, 198.08 and 151.61% as com- stress had an adverse effect on LT and EL across all vari-
pared to 0, 100, 200, 300, 400 and 500 mg L–1 potassium eties. Nevertheless, when potassium silicates were applied,
silicates, respectively (Table 1). Whereas the concentration there was a noticeable improvement in LT and EL for all va-
500 mg L–1 of potassium silicate did not show increasing rieties under drought-stress conditions. The application of
these parameters in cotton plant. potassium silicates led to a decrease in LT and EL, peaking
at a 400 mg L–1 application rate. Beyond this concentration,
Individual Leaf Area and Carotenoids Contents however, LT exhibited an increasing trend. In variety, Zong
main-113, under drought stress, potassium silicate appli-
The normal irrigation showed increased Individual Leaf cation at the rate of 400 mg L–1 decreased the LT 2.64%,
Area (ILA) in all varietal compression and potassium sil- 8.34%, 14.71%, 34.48 and 11.43% as compared to 0, 100,
icate levels indicated in Table 1. However, drought stress 200, 300, 400 and 500 mg L–1 potassium silicates, respec-
had an adverse effect on ILA across all varieties. Never- tively (Fig. 2a). However, in variety Xin Nong-525, un-
theless, when potassium silicates were applied, there was der drought stress, potassium silicate application at the rate
a noticeable improvement in ILA for all varieties under of 400 mg L–1 decreased the LT 2.38%, 7.51%, 13.17%,
drought-stress conditions showed in Table 1. The appli- 30.29%, and 10.26% as compared to 0, 100, 200, 300, 400
cation of potassium silicates led to an increase in ILA, and 500 mg L–1 potassium silicates, respectively (Fig. 2a).
peaking at a 400 mg L–1 application rate. Beyond this con- The application of potassium silicates led to a decrease in
centration, however, ILA exhibited a declining trend. In EL, peaking at a 400 mg L–1 application rate. Beyond this
variety, Zong main-113, under drought stress, potassium concentration, however, EL exhibited an increasing trend. In
silicate application at the rate of 400 mg L–1 increased the variety, Zong main-113, under drought stress, potassium sil-
ILA by 12.13%, 24.14%, 34.48%, 41.38% and 27.29% icate application at the rate of 400 mg L–1 decreased the EL

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Table 1 The effect of foliar-applied Potassium Silicate on various growth parameters parameters of four cotton genotypes under normal (80% FC) and drought stress (50% FC).
Potassium Root length (cm) Shoot length (cm) Shoot dry weight (g) Root dry weight (g) Individual Leaf Area (cm2)
silicates Control Drought stress Control Drought stress Control Drought Control Drought stress Control Drought
(mg L–1) stress stress

Variety Zong main-113


0 26.84 ± 1.37 b 20.74 ± 2.03 a 40.47 ± 1.95 b 36.07 ± 1.21 a 1.63 ± 0.07 c 1.13 ± 0.08 a 0.55 ± 0.51 a–c 0.35 ± 0.31 a 34 ± 1.73 c 29 ± 1.24 a
100 34.16 ± 2.13 d 28.06 ± 2.32 c 46.74 ± 2.05 f 42.34 ± 2.23 c 1.79 ± 0.03 d 1.29 ± 0.04 b 0.66 ± 0.61 a–c 0.46 ± 0.34 ab 38 ± 2.01 f 33 ± 1.69 b
200 41.48 ± 2.73 g 35.38 ± 3.11 e 48.09 ± 2.65 h 43.69 ± 2.37 d 2.44 ± 0.05 i 1.94 ± 0.06 e 0.95 ± 0.92 a–c 0.75 ± 0.68 a–c 41 ± 1.18 h 36 ± 1.97 d
300 46.36 ± 1.24 h 40.26 ± 2.81 f 51.11 ± 2.19 i 47.53 ± 2.31 e 2.52 ± 0.02 j 2.02 ± 0.04 f 1.05 ± 0.81 a–c 0.85 ± 0.62 a–c 44 ± 1.87 i 39 ± 1.65 e
400 65.88 ± 3.03 l 59.78 ± 2.43 k 61.06 ± 2.41 j 56.66 ± 1.67 g 2.76 ± 0.06 k 2.26 ± 0.05 g 1.17 ± 11 bc 0.97 ± 0.78 a–c 46 ± 2.07 j 41 ± 1.36 g
500 53.68 ± 2.53 j 47.58 ± 1.93 i 48.34 ± 2.17 l 43.94 ± 2.29 k 2.58 ± 0.04 l 2.08 ± 0.07 h 1.02 ± 0.94 c 0.82 ± 0.71 a–c 42 ± 1.67 k 37 ± 1.43 h
Variety Xin Nong-525
0 21.29 ± 1.12 b 15.19 ± 2.61 a 36.07 ± 2.14 b 31.67 ± 2.22 a 0.96 ± 0.04 c 0.46 ± 0.02 a 0.51 ± 0.44 a–c 0.31 ± 0.21 a 29 ± 1.56 c 24 ± 1.48 a
100 28.61 ± 3.03 d 22.51 ± 2.21 c 42.34 ± 2.47 f 37.94 ± 2.65 c 1.12 ± 0.06 d 0.62 ± 0.03 b 0.62 ± 0.37 a–c 0.42 ± 0.38 ab 33 ± 2.04 f 28 ± 1.77 b
200 35.93 ± 2.31 g 29.83 ± 2.67 e 43.69 ± 2.16 h 39.29 ± 1.97 d 1.77 ± 0.07 i 1.27 ± 0.06 e 0.91 ± 0.84 a–c 0.71 ± 0.61 a–c 36 ± 1.59 h 31 ± 1.88 d
300 40.81 ± 2.63 h 34.71 ± 1.83 f 47.51 ± 1.41 i 43.11 ± 208 e 1.85 ± 0.06 j 1.35 ± 0.04 f 1.01 ± 0.87 a–c 0.81 ± 0.54 a–c 39 ± 1.65 i 34 ± 1.27 e
400 60.33 ± 2.73 l 54.23 ± 1.97 k 56.66 ± 1.87 j 52.26 ± 1.93 g 2.09 ± 0.05 k 1.59 ± 0.03 g 1.13 ± 1.01 bc 0.93 ± 0.74 a–c 41 ± 1.77 j 36 ± 1.85 g
500 48.13 ± 1.77 j 42.03 ± 2.09 i 43.94 ± 2.28 l 39.54 ± 2.31 k 1.91 ± 0.04 l 1.41 ± 0.09 h 0.98 ± 0.84 c 0.78 ± 0.52 a–c 37 ± 1.19 k 32 ± 1.78 h
Variety Xin lu Zhong-55
0 23.54 ± 1.54 b 17.44 ± 1.45 a 38.37 ± 2.51 b 33.97 ± 2.38 a 1.23 ± 0.06 c 0.73 ± 0.03 a 0.54 ± 0.51 a–c 0.34 ± 0.33 a 31 ± 1.76 c 26 ± 1.28 a
100 30.86 ± 2.86 d 24.76 ± 2.11 c 44.64 ± 2.17 f 40.24 ± 2.08 c 1.39 ± 0.07 d 0.89 ± 0.04 b 0.66 ± 0.56 a–c 0.46 ± 0.35 ab 35 ± 1.53 f 30 ± 1.67 b
200 38.18 ± 3.03 g 32.08 ± 2.89 e 45.99 ± 2.24 h 41.59 ± 2.39 d 2.04 ± 0.08 i 1.54 ± 0.06 e 0.94 ± 0.74 a–c 0.74 ± 0.66 a–c 38 ± 1.25 h 33 ± 1.76 d
300 43.06 ± 2.43 h 36.96 ± 2.34 f 49.81 ± 2.36 i 45.41 ± 2.28 e 2.12 ± 0.05 j 1.62 ± 0.07 f 1.05 ± 0.91 a–c 0.85 ± 0.62 a–c 41 ± 1.76 i 36 ± 1.84 e
400 62.58 ± 1.83 l 56.48 ± 2.31 k 58.96 ± 1.68 j 54.56 ± 2.14 g 2.36 ± 0.06 k 1.86 ± 0.05 g 1.17 ± 1.01 bc 0.97 ± 0.72 a–c 43 ± 1.16 j 38 ± 1.69 g
500 50.38 ± 2.05 j 44.28 ± 2.62 i 46.24 ± 2.16 l 41.84 ± 1.85 k 2.18 ± 0.08 l 1.68 ± 0.09 h 1.01 ± 0.97 c 0.81 ± 0.67 a–c 39 ± 1.45 k 34 ± 1.43 h
Variety Xin lu Zhong-66
0 22.14 ± 2.76 b 16.04 ± 2.87 a 37.27 ± 2.23 b 32.87 ± 2.41 a 1.08 ± 0.07 c 0.58 ± 0.03 a 0.52 ± 0.36 a–c 0.32 ± 0.11 a 30 ± 176 c 25 ± 1.89 a
100 29.46 ± 1.86 d 23.36 ± 2.53 c 43.54 ± 2.53 f 39.14 ± 1.87 c 1.24 ± 0.04 d 0.74 ± 0.05 b 0.63 ± 0.53 a–c 0.43 ± 0.32 ab 34 ± 1.29 f 29 ± 1.76 b
200 36.78 ± 1.78 g 30.68 ± 2.78 e 44.89 ± 2.29 h 40.49 ± 2.49 d 1.89 ± 0.06 i 1.39 ± 0.04 e 0.92 ± 0.76 a–c 0.72 ± 0.64 a–c 37 ± 2.07 h 32 ± 1.44 d
300 41.66 ± 2.56 h 35.56 ± 2.87 f 48.73 ± 1.86 i 44.34 ± 2.37 e 1.97 ± 0.07 j 1.47 ± 0.05 f 1.02 ± 0.92 a–c 0.82 ± 0.57 a–c 40 ± 2.09 i 35 ± 1.68 e

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400 61.18 ± 1.74 l 55.08 ± 2.16 k 57.86 ± 2.19 j 53.46 ± 2.24 g 2.21 ± 0.06 k 1.71 ± 0.03 g 1.14 ± 0.87 bc 0.94 ± 0.59 a–c 42 ± 1.94 j 37 ± 1.83 g
500 48.98 ± 2.19 j 42.88 ± 1.38 i 45.14 ± 2.27 l 40.74 ± 2.35 k 2.03 ± 0.05 l 1.53 ± 0.07 h 0.99 ± 0.98 c 0.79 ± 0.53 a–c 38 ± 1.68 k 33 ± 1.54 h
Data represent the means ± Standard deviation (n = 3). Means within Treatments (Irrigation regimes, Potassium silicate application, and genotypes) having different letters are significantly different
at P ≤ 0.05
M. Nazim et al.
Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

300, 400 and 500 mg L–1 potassium silicates, respectively


(Fig. 3c).

Chlorophyll a, b and Total Chlorophyll

The normal irrigation showed increased Chlorophyll con-


tents (Chl) in all varietal compression and potassium sil-
icate levels. However, drought stress had an adverse ef-
fect on Chl across all varieties presented in (Fig. 4a–c).
Nevertheless, when potassium silicates were applied, there
was a noticeable improvement in Chl for all varieties un-
der drought-stress conditions. The application of potassium
silicates led to an increase in Chl, peaking at a 400 mg L–1
application rate. Beyond this concentration, however, Chl
exhibited a declining trend. In variety, Zong main-113, un-
der drought stress, potassium silicate application at the rate
of 400 mg L–1 increased the Chl a 7.44%, 32.91%, 59.26%,
114.23%, and 76.06% as compared to 0, 100, 200, 300, 400
and 500 mg L–1 potassium silicates, respectively (Fig. 4a).
However, in variety Xin Nong-525, under drought stress,
potassium silicate application at the rate of 400 mg L–1
increased the Chl a 11.12%, 49.54%, 88.89%, 172.17%,
and 110.61% as compared to 0, 100, 200, 300, 400 and
500 mg L–1 potassium silicates, respectively (Fig. 4a). In va-
riety, Zong main-113, under drought stress, potassium sili-
cate application at the rate of 400 mg L–1 increased the Chl b
33.31%, 66.82%, 116.63%, 114.68%, and 98.57% as com-
pared to 0, 100, 200, 300, 400 and 500 mg L–1 potassium
silicates, respectively (Fig. 4b). However, in variety Xin
Nong-525, under drought stress, potassium silicate applica-
tion at the rate of 400 mg L–1 increased the Chl b 57.14%,
114.29%, 198.69%, 242.86% and 171.43% as compared to
0, 100, 200, 300, 400 and 500 mg L–1 potassium silicates,
respectively (Fig. 4b). In variety, Zong main-113, under
drought stress, potassium silicate application at the rate
of 400 mg L–1 increased the Total Chls 39.67%, 79.52%,
146.03%, 194.21%, and 117.34% as compared to 0, 100,
200, 300, 400 and 500 mg L–1 potassium silicates, respec-
tively (Fig. 4c). However, in variety Xin Nong-525, un-
der drought stress, potassium silicate application at the rate
of 400 mg L–1 increased the Total Chls 35.75%, 62.31%,
89.51%, 128.41%, and 109.16% as compared to 0, 100,
200, 300, 400 and 500 mg L–1 potassium silicates, respec-
Fig. 2 a–c Impact of foliar-applied Potassium Silicate on Leaf Temp. (a), tively (Fig. 4c).
Carotenoids (b) and Proline (c) of four cotton genotypes under normal
(CK) and drought stress (DS)
Hydrogen Peroxide and Malondialdehyde

23.53%, 49.63%, 90.91%, 198.66%, and 74.83% as com- The normal irrigation showed decreased hydrogen peroxide
pared to 0, 100, 200, 300, 400 and 500 mg L–1 potassium sil- (H2O2) and malondialdehyde (MDA) in all varietal com-
icates, respectively (Fig. 3c). However, in variety Xin Nong- pression and potassium silicate levels (Fig. 3a, b). However,
525, under drought stress, potassium silicate application at drought stress had an adverse effect on H2O2 and MDA
the rate of 400 mg L–1 decreased the EL 12.53%, 41.18%, across all varieties. Nevertheless, when potassium silicates
71.43%, 139.86, and 59.87% as compared to 0, 100, 200, were applied, there was a noticeable improvement in H2O2

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M. Nazim et al.

tion at the rate of 400 mg L–1 decreased the H2O2 10.06%,


7.47%, 12.93%, 21.94%, and 9.64% as compared to 0, 100,
200, 300, 400 and 500 mg L–1, respectively (Fig. 3a). How-
ever, in variety Xin Nong-525, under drought stress, potas-
sium silicate application at the rate of 400 mg L–1 decreased
the H2O2 1.97%, 5.84%, 9.96%, 16.57% and 7.46% as com-
pared to 0, 100, 200, 300, 400 and 500 mg L–1 potassium
silicates, respectively (Fig. 3a). In variety, Zong main-113,
under drought stress, potassium silicate application at the
rate of 400 mg L–1 decreased the MDA 25.53%, 51.29%,
118.24%, 178.91%, and 155.19% as compared to 0, 100,
200, 300, 400 and 500 mg L–1 potassium silicates, respec-
tively (Fig. 3b). However, in variety Xin Nong-525, under
drought stress, potassium silicate application at the rate of
400 mg L–1 decreased the H2O2 14.28%, 41.17%, 59.51%,
132.17% and 96.13% as compared to 0, 100, 200, 300, 400
and 500 mg L–1 potassium silicates, respectively (Fig. 3b).

Proline and Free Amino Acid

The normal irrigation showed increased proline and free


amino acid (FAA) in all varietal compression and potas-
sium silicate levels. However, drought stress had an adverse
effect on proline and FAA across all varieties. Nevertheless,
when potassium silicates were applied, there was a notice-
able improvement in proline and FAA for all varieties un-
der drought-stress conditions showed in Fig. 2c and 5a.
The application of potassium silicates led to an increase in
proline and FAA, peaking at a 400 mg L–1 application rate.
Beyond this concentration, however, proline and FAA ex-
hibited a declining trend. In variety, Zong main-113, under
drought stress, potassium silicate application at the rate of
400 mg L–1 increased the proline 35.43%, 58.29%, 82.36%,
105.38%, and 23.53% as compared to 0, 100, 200, 300, 400
and 500 mg L–1 potassium silicates, respectively (Fig. 2c).
However, in variety Xin Nong-525, under drought stress,
potassium silicate application at the rate of 400 mg L–1 in-
creased the proline 59.29%, 97.86%, 138.57%, 178.33%,
and 38.43% as compared to 0, 100, 200, 300, 400 and
500 mg L–1 potassium silicates, respectively (Fig. 2c). In
variety, Zong main-113, under drought stress, potassium
silicate application at the rate of 400 mg L–1 increased the
FAA 11.07%, 14.05%, 19.18%, 24.73%, and 7.38% as com-
Fig. 3 a–c Impact of foliar-applied Potassium Silicate on H2O2 (a), pared to 0, 100, 200, 300, 400 and 500 mg L–1 potassium sil-
Malondialdehyde (b) and Electrolyte leakage levels (c) of four cotton icates, respectively (Fig. 5a). However, in variety Xin Nong-
genotypes under normal (CK) and drought stress (DS) 525, under drought stress, potassium silicate application at
the rate of 400 mg L–1 decreased the FAA 8.91%, 12.79%,
and MDA for all varieties under drought-stress conditions 19.38%, 26.74%, and 1.94% as compared to 0, 100, 200,
showed in Fig. 3a, b. The application of potassium silicates 300, 400 and 500 mg L–1 potassium silicates, respectively
led to a decrease in H2O2 and MDA, peaking at a 400 mg L–1 (Fig. 5a).
application rate. Beyond this concentration, however, H2O2
and MDA exhibited a decreasing trend. In variety, Zong
main-113, under drought stress, potassium silicate applica-

K
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Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

Fig. 4 a–c Impact of foliar-applied Potassium Silicate on chlorophyll-


a (a), chlorophyll b (b) and Total chlorophyll content (c) of four cotton Fig. 5 a–c Impact of foliar-applied Potassium Silicate on FAA (a),
genotypes under normal (CK) and drought stress (DS) GPOD (b) and RLWC (c) of four cotton genotypes under normal (CK)
and drought stress (DS)

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M. Nazim et al.

Leaf Relative Water Contents and Photosynthetic compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas-
Rate sium silicates, respectively (Fig. 6b). In variety, Zong main-
113, under drought stress, potassium silicate application at
Normal irrigation showed increased leaf relative water con- the rate of 400 mg L–1 increased the Tr 27.06%, 35.59%,
tents (LRWC) and photosynthetic rate (Pn) in all vari- 78.68%, 121.35%, and 48.74% as compared to 0, 100,
etal compression and potassium silicate levels. However, 200, 300, 400 and 500 mg L–1 potassium silicates, respec-
drought stress had an adverse effect on LRWC and Pn tively (Fig. 6c). However, in variety Xin Nong-525, un-
across all varieties. Nevertheless, when potassium silicates der drought stress, potassium silicate application at the rate
were applied, there was a noticeable improvement in LRWC of 400 mg L–1 increased the Tr 42.74%, 66.93%, 159.16%,
and Pn for all varieties under drought-stress conditions. 256.43%, and 48.92% as compared to 0, 100, 200, 300, 400
The application of potassium silicates led to an increase and 500 mg L–1 potassium silicates, respectively (Fig. 6c).
in LRWC and Pn, peaking at a 400 mg L–1 application rate.
Beyond this concentration, however, LRWC and Pn exhib- Catalase and Glutathione S-Transferase
ited a declining trend. In variety, Zong main-113, under
drought stress, potassium silicate application at the rate of The normal irrigation showed increased catalase (CAT),
400 mg L–1 increased the LRWC 4.88%, 19.53%, 29.27%, glutathione S-transferase (GST) in all varietal compression
43.91/%, and 34.39% as compared to 0, 100, 200, 300, 400 and potassium silicate levels. However, drought stress had
and 500 mg L–1 potassium silicates, respectively (Fig. 5c). an adverse effect on CAT and GST across all varieties. Nev-
However, in variety Xin Nong-525, under drought stress, ertheless, when potassium silicates were applied, there was
potassium silicate application at the rate of 400 mg L–1 in- a noticeable improvement in CAT and GST for all varieties
creased the LRWC 11.46%, 22.86%, 34.29%, 48.57%, and under drought-stress conditions presented in Fig. 7a, b.
37.14% as compared to 0, 100, 200, 300, 400 and 500 The application of potassium silicates led to an increase in
potassium silicates, respectively (Fig. 5c). In variety, Zong CAT and GST, peaking at a 400 mg L–1 application rate.
main-113, under drought stress, potassium silicate appli- Beyond this concentration, however, CAT and GST exhib-
cation at the rate of 400 mg L–1 increased the Pn 21.57%, ited a declining trend. In variety, Zong main-113, under
79.18%, 92.39%, 148.58%, and 30.58% as compared to drought stress, potassium silicate application at the rate
0, 100, 200, 300, 400 and 500 mg L–1 potassium silicates, re- of 400 mg L–1 increased the CAT 1.49%, 3.76%, 6.09%,
spectively (Fig. 6a). However, in variety Xin Nong-525, un- 10.61%, and 1.97% as compared to 0, 100, 200, 300, 400
der drought stress, potassium silicate application at the rate and 500 mg L–1 potassium silicates, respectively (Fig. 7a).
of 400 mg L–1 increased the Pn 30.49%, 111.93%, 118.79%, However, in variety Xin Nong-525, under drought stress,
210.04%, and 43.15% as compared to 0, 100, 200, 300, 400 potassium silicate application at the rate of 400 mg L–1
and 500 mg L–1 potassium silicates, respectively (Fig. 6a). increased the CAT 4.91%, 8.38%, 22.65%, 40.39%, and
2.71% as compared to 0, 100, 200, 300, 400 and 500 mg L–1
Stomatal Conductance and Transpiration Rate potassium silicates, respectively (Fig. 7a). In variety, Zong
main-113, under drought stress, potassium silicate appli-
The normal irrigation showed increased stomatal conduc- cation at the rate of 400 mg L–1 increased the GST 9.46%,
tance (gs) and transpiration rate (Tr) in all varietal compres- 18.83%, 31.81%, 35.41%, and 12.89% as compared to
sion and potassium silicate levels. However, drought stress 0, 100, 200, 300, 400 and 500 mg L–1 potassium silicates,
had an adverse effect on gs and Tr across all varieties. Nev- respectively (Fig. 7b). However, in variety Xin Nong-525,
ertheless, when potassium silicates were applied, there was under drought stress, potassium silicate application at the
a noticeable improvement in gs and Tr for all varieties under rate of 400 mg L–1 increased the GST 9.65%, 15.66%,
drought-stress conditions shown in Fig. 6b, c. The applica- 29.18%, 32.89%, and 3.82% as compared to 0, 100, 200,
tion of potassium silicates led to an increase in gs and Tr, 300, 400 and 500 mg L–1 potassium silicates, respectively
peaking at a 400 mg L–1 application rate. Beyond this con- (Fig. 7b).
centration, however, gs and Tr exhibited a declining trend.
In variety, Zong main-113, under drought stress, potassium Guaiacol Peroxidase and Ascorbate Peroxidase
silicate application at the rate of 400 mg L–1 increased the
gs 51.67%, 91.48%, 143.09%, 167.54%, and 119.17% as The normal irrigation showed increased Guaiacol perox-
compared to 0, 100, 200, 300, 400 and 500 mg L–1 potas- idase (GPOD), ascorbate peroxidase (APX) in all vari-
sium silicates, respectively (Fig. 6b). However, in vari- etal compression and potassium silicate levels. However,
ety Xin Nong-525, under drought stress, potassium sili- drought stress had an adverse effect on GPOD and APX
cate application at the rate of 400 mg L–1 increased the across all varieties. Nevertheless, when potassium silicates
gs 82.78%, 157.52%, 249.19%, 331.51%, and 241.67% as were applied, there was a noticeable improvement in GPOD

K
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Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

Fig. 6 a–c Impact of foliar-applied Potassium Silicate on Pn (a), gs (b) Fig. 7 a–c Impact of foliar-applied Potassium Silicate on CAT (a),
and Tr (c) of four cotton genotypes under normal (CK) and drought GST (b) and APX (c) of four cotton genotypes under normal (CK)
stress (DS) and drought stress (DS)

and APX for all varieties under drought-stress conditions. 25.68%, and 8.33% as compared to 0, 100, 200, 300, 400
The application of potassium silicates led to an increase in and 500 mg L–1 potassium silicates, respectively (Fig. 5b).
GPOD and APX, peaking at a 400 mg L–1 application rate. However, in variety Xin Nong-525, under drought stress,
Beyond this concentration, however, GPOD and APX ex- potassium silicate application at the rate of 400 mg L–1 in-
hibited a declining trend. In variety, Zong main-113, under creased the GPOD 5.58%, 12.27%, 22.13%, 28.66%, and
drought stress, potassium silicate application at the rate of 9.12% as compared to 0, 100, 200, 300, 400 and 500 mg L–1
400 mg L–1 increased the (GPOD) 5.13%, 11.37%, 20.24%, potassium silicates, respectively (Fig. 5b). In variety, Zong

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M. Nazim et al.

Fig. 8 Principal component analysis (PCA) assimilating with foliar application of K2SiO3, four cotton genotypes and irrigation level based on
morpho-physiological and agronomic characteristics

main-113, under drought stress, potassium silicate applica- 300, 400 and 500 mg L–1 potassium silicates, respectively
tion at the rate of 400 mg L–1 increased the APX 5.95%, (Fig. 7c).
27.45%, 34.57%, 44.05%, and 14.28% as compared to
0, 100, 200, 300, 400 and 500 mg L–1 potassium silicates, PCA, Correlation and Heatmap
respectively (Fig. 7c). However, in variety Xin Nong-525,
under drought stress, potassium silicate application at the To examine the differences and correlations between several
rate of 400 mg L–1 increased the APX 5.41%, 27.03%, morpho-physiological and biochemical characteristics of
39.18%, 49.32%, and 9.46% as compared to 0, 100, 200, cotton under foliar-applied potassium silicate and drought
stress, a principal component analysis (PCA) was carried

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Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

Fig. 9 Corr. plot study of Cot-


ton plants subjected to drought
stress and foliar applied K2SiO3
treatments parameters, with an
emphasis on the Morpho-phys-
iology and biochemistry, com-
ponents of cotton. A stronger
relation is indicated by the
darker colors (red and blue,
signifying positive and negative
correlations, respectively)

out. Treatments were scattered among various biplots, and (400 mg L–1) was applied. This improvement was observed
vectors were used to depict the parameters represented in in all (growth and physio-biochemical) parameters. Interest-
Fig. 8. The 87.4% variability of the total variation from the ingly, the Zong main-113 variety performed better in both
22 parameters was covered by the first two PCs (PC1 75.1% controlled and stressful environments compared to control,
and PC2 12.3%). The first PC was related to all the ana- when treated without foliar spray under drought stress con-
lyzed parameters except, H2O2, LT, MDA and EL% which ditions, the Xin Nong-525 variety exhibited a negative re-
were attributed to PC2. All the foliar-applied treatments of lationship with all metrics (Fig. 10).
potassium silicate of cotton under drought stress showed
variability amongst the treatments. In a drought with nega-
tive eigenvalues, the amount of H2O2, CAT, LT and MDA Discussion
were increased (Fig. 8). Similarly, Potassium silicate with
drought had a synergistic impact that decreased H2O2, CAT, Cotton is considered as moderate resistant crop to water
LT and MDA (Fig. 8). These parameters GPOD, FFA, pro- shortage owing to its deep root systems. The current study
line, and LRWC were significantly and positively related to revealed the mechanism by which potassium silicate con-
each other in the PCA graphs, also strengthen these assump- fronted the negative effects of drought in cotton during the
tions. According to PCA graphs showed in Fig. 8, param- five-leaf (vegetative) stage. Foliar application of potassium
eters Chl, Tr, ILA, and gs also had significant and positive silicate at the critical five-leaf vegetative stage of cotton un-
relations with one another. These relations are linked to cot- der drought stress showed its potential in enhancing drought
ton plants’ defensive mechanisms and the impacts that arise tolerance. To understand the mechanism of potassium sili-
when they are under stress. All growth, physiological, and cate-mediated crop tolerance, we examined several morpho-
biochemical attributes exhibited a positive linear correlation physio-biochemical traits that are vital for cotton survival in
according to Pearson correlation, with the except of a neg- water-limited conditions. A significant reduction in cotton
ative correlation with H2O2, LT, MDA and EL% showed in plant growth and biomass, as well as the withering, color
(Fig. 9). The results of the heatmap matrix demonstrated of yellowing, and dehydrating of leaves, indicated drought
that, in comparison to the control, the Zong main-113 va- induced damage in cotton plants (Fig. 1). Our findings sug-
riety performed better under normal conditions when KSi gest that potassium silicate plays a vital role in counteract-

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M. Nazim et al.

ing protective barrier in leaf apoplasts (Arif et al. 2021;


Nazim et al. 2024). However, the results showed that the
application of potassium silicate (400 mg L–1) positively
increased growth and biomass attributes showed (Fig. 6).
Cotton growth and physio-biochemical alterations are en-
hanced by foliar-applied Potassium silicate (0–400 mg L–1)
shown in (Table 1). Younis et al. (2020) reported that by us-
ing K2SiO3, wheat seedlings under water stress showed im-
proved leaf thickness, dry mass, RWC, and LA. Similarly,
Silicon increased the amount of chlorophyll, RWC, and wa-
ter usage efficiency in Maize Seedlings under drought stress
(Sharf-Eldin et al. 2023). Previous studies have revealed
that drought stress negatively impacts molecular, biochem-
ical, and physiological processes, leading to impaired plant
development and reduced crop yield (Qamar et al. 2020;
Nazim et al. 2021). When plants are stressed by drought,
their ability to produce dry matter is limited due to con-
straints on photosynthesis and stomatal closure (Khatun
et al. 2021).
Since, chlorophyll contents have a direct impact on the
plant growth and their development (Zafar et al. 2023). The
application of Potassium silicate (400 mg L–1) significantly
improved the Chl. a, Chl. b, Carotenoids contents, and gas
Fig. 10 showed a heatmap matrix of many factors in four genotypes exchange parameters in our study, both under water stress
of cotton treated with foliar application of K2SiO3 under normal and condition (Fig. 2a). Severe drought stress causes to stom-
drought stress atal closure, reduced transpiration, and decreased photosyn-
thesis, which collectively result in lower plant growth and
ing these adverse effects, promoting healthier growth even reduced cell water potential (Boora et al. 2023). Applied
under water stress conditions. Foliar applied Potassium sil- Potassium silicate also improved the leaf chlorophyll con-
icate enhanced the plant growth and biomass production, tent, nutrient intake, increased photosynthetic rate, and thin
decreased indications of yellowing symptoms and wither- down cell membranes in a variety of plants (Teixeira et al.
ing of the leaves and helped cotton plants to overcome the 2020). Researchers have used silicon to increase photosyn-
adverse effects of drought in cotton plants. The significant thesis in recent trials of drought stress and have shown
increase in root biomass was in Potassium silicate-treated a comparable increased (Adrees et al. 2022; Raza et al.
plants indicates that Potassium silicate facilitated greater 2023). Applied potassium silicate is likely to lessen the ef-
water and nutrient absorption. Thereby instigating to im- fects of drought stress on photosynthetic pigments by natu-
proved growth characteristics and increased photosynthetic rally producing more cytokinin, which restores chlorophyll
activity under drought conditions, prominence the poten- synthesis (Mohammadi et al. 2024), and enhances chloro-
tial of potassium silicate in promoting drought resilience in plast ultrastructure associated with plant growth improve-
cotton (Hafez et al. 2021; Nazim et al. 2023). ment (Wahab et al. 2022). In addition, foliar spray of K2SiO3
The notable increases in root and shoot biomass, result- (0.5%) increased the nutrients in plants that promote photo-
ing from foliar-applied potassium silicate, enhanced cotton synthesis and yield of Sugarcane under the drought condi-
resistance to drought stress. According to Ibrahim et al. tions as reported by (Anitha et al. 2023). These findings are
(2020) foliar sprays of K2SiO3 (2 mM) comparatively less consistent with our study, which also revealed the positive
damage caused by water stress and increased shoot (SDW) of K2SiO3 on stress tolerance and plant productivity. It is ev-
and root dry weight (RDW), relative water content (RWC), ident that drought caused reduced Gaseous exchange such
leaf area index (LAI), were reduced by deficit irrigation. as Pn, Tr, gs, and RWC of leaves in cotton plants (Fig. 5c).
Ayub et al. (2018) also reported that exogenously applied Meshram et al. (2022) revealed that significant relationship
K2SiO3 (4 mmol) significantly improved plant growth and between transpiration rate, total chlorophyll concentration,
biomass production in both shoots and roots under both and photosynthesis, which is essential for cotton growth
stress and non-stress conditions. Furthermore, the applied under drought stress. Our study supports these findings,
of KSi (0.1–0.6 mM) increased plant height because of its emphasizing the role of physiological parameters in im-
protective properties against stress factors, by strengthen- proving cotton’s drought tolerance. The potential of potas-

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Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

Fig. 11 Generalized process


of Potassium Silicate induced
drought tolerance in cotton

sium silicate helping in plants to retain more water during In our experiment, proline levels 82.36% significantly in-
drought mmight be the cause of these improved gas ex- creased under water stress by foliar applied potassium sili-
change parameters, water use efficiency and productivity cate (400 mg L–1). However, silicon (Si) application did not
inconformity to results reported in maize cropby (Gomaa influence proline content in either genotype under drought
et al. 2021). Consistent with our findings, (Gong et al. 2005; stress. Similar to our findings, proline concentration has
Sah et al. 2022) suggested that the increased relative water been observed to increase in cotton leaves under copper
content (RWC) of leaves is attributed to the formation of stress (Ali et al. 2023). Verma et al. (2021) increased leaf
a dual-layered silica-cuticle structure on the leaf epidermal area and increased photosynthetic leaf gas exchange ap-
tissue. The initial symptom of drought stress on the plant pear to be directly correlated. Foliar applied Potassium sil-
is the reduction in water content that subsequently caused icate greatly increased the activity of CAT, GST, and APX
growth and physiological disorder (Nazim et al. 2024). (Fig. 7) whereas substances such as H2O2 (a), malondi-
Photosynthetic pigments are essential components of en- aldehyde (b) and electrolyte leakage levels (c) significantly
ergy metabolism in plants, playing a vital role in their adap- decreased (Fig. 3). Moreover, silicon was applied to cot-
tation and survival under adverse environmental conditions, ton plants to maintain and repair their cell membranes,
such as drought. These adaptations have been observed by which resulted in a notable reduction in reactive oxygen
Shiade et al. (2023) and are linked to the large-scale ac- species (Nazim et al. 2024). In a similar vein, numerous
cumulation of free amino acids (Fig. 5a). Proline typically studies have shown that silicon increased the antioxidant
accumulates in the cytosol and aids in cytoplasmic osmotic activity of various crops (Alharbi et al. 2022). Synthesis
adjustment under drought conditions (Batool et al. 2023), and accumulation of various organic osmolytes proline, to-

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M. Nazim et al.

tal free amino acids, (Salajegheh et al. 2020), and total plied under water stress conditions significantly increased
soluble sugars are widely recognized to have an integral total free amino acids and soluble sugars, enhancing os-
role in plant adaptation to drought, mainly by maintaining motic adjustment and root water uptake in both genotypes.
turgor pressure (Fatemi et al. 2021). Thus, potassium sili- This application of Sodium Silicate (2 mM) mediated im-
cate treatment increases SOD, glutathione reductase (Shi- provement aligns with similar findings in peanut genotypes
rani Rad et al. 2022), and catalase activities (Oraee et al. under water stress (Patel et al. 2021).
2024), slightly balancing the negative effects of drought on This study is inventive in regulating Cotton drought toler-
plants. Cotton growth and development are impeded dur- ance in response to foliar applied Potassium silicate (KSi).
ing droughts because of fluctuations in the nutrition bal- Elevated concentrations of MDA and proline in cotton
ance (Ahmad et al. 2019). Similarly, a direct comparison seedling under water stress conditions serve as biochemi-
with water stress plants revealed a strong correlation be- cal attributes for drought stress sensitivity. The increased
tween proline content and the activities of SOD and glu- anti-oxidative enzymes activities, such as CAT, APX, and
tathione reductase following foliar applied potassium sil- SOD, appear to be effective mechanisms for maintaining
icate (400 mg L–1). This suggests that potassium silicate photosynthetic performance in stress-tolerance. Thus, the
may aid in mitigating water loss by significantly enhancing increased expression of physio-biochemical attributes is an
proline accumulation (Fig. 2c). Nonetheless, higher pho- essential strategy to improve cotton drought tolerance in
tosynthetic efficiency might reverse cellular metabolism to response to water deficits, ultimately with the foliar appli-
more growth-promoting attributes and lessen the accumu- cation of potassium silicate in the hyper-arid Taklamakan
lation of free amino acids (Shafiq et al. 2019). According Desert, (Fig. 11) depicts a generalized process of potassium
to Kim et al. (2017) studies suggest that early exposure to silicate-induced drought tolerance in cotton.
drought or water deficit creates a ‘stress imprint’ in plants,
which enhances their ability to scavenge reactive oxygen
species (ROS) and minimize oxidative damage when sil- Conclusion
icon is applied by Khan et al. (2023). Alternatively, it’s
possible that even in drought conditions, Potassium sili- The study concluded that among the different application
cate-supplemented plants utilized more free amino acids to rates of potassium silicate (0, 100, 200, 300, 400, and
boost their biomass (Table 1), which prevented the accumu- 500 mg L–1), 400 mg L–1 was the most effective in miti-
lation of free amino acids in the plants. Plants store many gating drought stress at 50% field capacity across all cot-
low-molecular-weight compounds when there is a shortage ton cultivars tested (Zhongmian-113, Xin Nong-525, Xinlu
of water, one of which is to keep the osmotic stability. Zhong-55, and Xinlu Zhong-66). In plots treated with potas-
Furthermore, Yang et al. (2021) reported that high ROS sium silicate, a lower quantity of antioxidants indicated re-
formation during drought stress results in oxidative dam- duced drought stress compared to untreated plots. The or-
age to plant DNA, lipids, and proteins, which inhibits plant der of drought tolerance at the 400 mg L–1 application rate
growth and development. Similarly, other researchers found was Zhongmian-113, Xin Nong-525, Xinlu Zhong-55, and
that the biomass, grain, and yield of wheat are reduced by Xinlu Zhong-66. Thus, in water-limited areas, potassium
drought stress (Ahmad et al. 2018; Ullah et al. 2024). These silicate application can be an effective strategy for achiev-
results are aligning with recent research specifying Silicon ing higher cotton productivity with the appropriate selection
enhanced antioxidant defense mechanisms in peanuts geno- of cotton cultivars. These findings provide a foundation for
types subjected to drought conditions (Patel et al. 2021). future long-term research on potassium silicate and its ef-
Our results demonstrated that under both irrigation regimes, fects on a broader range of cotton cultivars.
the application of Potassium silicate (400 mg L–1) enhanced Supplementary Information The online version of this article (https://
accumulation could help plants maintain optimal leaf tur- doi.org/10.1007/s10343-024-01005-8) contains supplementary mate-
gor pressure and stomatal conductance for CO2 absorption rial, which is available to authorized users.
and play a crucial role in adaptation and survival of plants
Acknowledgements This research was supported by the Xinjiang Key
in undesired environmental condition. Nevertheless, potas- Research and Development Programme (Project No. 2022B02040-2)
sium silicate reduced the damaging effects of drought on the and a scholarship from the University of Chinese Academy of Sciences
growth and its constituent parts also regulated Gaseous ex- (UCAS). We extend our heartfelt gratitude for their invaluable support
change characteristics against the harmful effects of singlet and commitment to academic advancement.
oxygen (1O2), thereby safeguarding the chloroplast from Author Contribution MN: Conceptualization, Data curation, Formal
photooxidative damage (Karvar et al. 2023), because it al- analysis, Investigation, Methodology, Software, Validation, Visu-
tered transpiration, increased the rate of photosynthesis, and alization, writing—original draft, Writing—review & editing. XL:
increased osmolytes and antioxidants of plants (Ibrahim Conceptualization, Funding acquisition, Resources, Supervision, Val-
idation, Visualization, Writing—review & editing. AT: Visualization,
et al. 2020). Our results showed that Potassium Silicate ap-

K
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
Exogenous Potassium Silicate Improves Drought Tolerance in Cotton Genotypes by Modulating Growth, Gas Exchange and...

Writing—review & editing. KS: Formal analysis, Software, Writ- tential through foliar applied silicon. Silicon 14:11243–11252.
ing—review & editing. MM: Formal analysis, Writing—original draft. https://doi.org/10.1007/s12633-022-01849-7
ZD: Methodology, Resources. FZ: Supervision, Writing—review & Ayub Q, Khan M, Khan A et al (2018) Effect of gibberellic acid
editing. and potassium silicate on physiological growth of Okra (Abel-
moschus esculentus L.) under salinity stress. Pure Appl Biol
7:8–19. https://doi.org/10.19045/bspab.2018.70002
Aziz HA, Sharaf M, Omar M et al (2023) Improvement of selected
Declarations morphological, physiological, and biochemical parameters of
banana (Musa acuminata L.) using potassium silicate under
Conflict of interest M. Nazim, X. Li, A. Tariq, K. Shahzad, M. Muham- drought. Int J Exp Bot 92:1019–1036. https://doi.org/10.32604/
mad, Z. Dawei and F. Zeng declare that they have no competing phyton.2023.026769
interests. Bates LS, Waldren RP, Teare ID (1973) Rapid determination of free
Ethical standards This article does not contain any studies with human proline for water-stress studies. Plant Soil 39:205–207. https://
or animal subjects. Ethics approval and consent to participate: Not doi.org/10.1007/BF00018060
applicable. Consent for publication: Not applicable. Batool M, El-Badri AM, Hassan MU et al (2023) Drought stress in
brassica napus: effects, tolerance mechanisms, and management
strategies. J Plant Growth Regul 42:21–45. https://doi.org/10.
1007/S00344-021-10542-9
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