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Evaluation of nano-silicon efficiency on compatible solutes and nutrient status of Damask rose affected by in vitro simulated drought stress

Abstract

Background

Drought stress is a critical environmental factor that disturbs plant performance. However, some non-essential elements such as silicon can improve water deficit tolerance by modulating photosynthesis assimilates and compatible solutes production. Therefore, the present work was conducted to modulate polyethylene glycol (PEG)-induced water deficiency under in vitro culture in Damask rose genotypes (Maragheh and Kashan) by nano-silicon (SiO2-NPs) treatment. A completely randomized factorial experiment was used as three concentrations of SiO2-NPs (0, 50, and 100 mg L−1) and five concentrations of PEG (0, 25, 50, 75, and 100 g L−1). Then, the comparative effects of water deficiency on vegetative traits, metabolites, and nutrients were studied.

Results

The drought promoted a significant decrease in chlorophyll, fresh/dry weight, biomass, and an increase in electrolyte leakage. The amount of micro- and macronutrients were affected by drought stress and decreased in both genotypes. In contrast, the activity of polyphenol oxidase (PPO) and total phenolic compounds (TPC) along with biochemical traits was increased. Treatment with SiO2-NPs improved the leaf area index (LAI), chlorophyll, and biomass under severe water deficiency. The concentration of compatible solutes such as carbohydrates, total flavonoid content (TFC), TPC, anthocyanin, and antioxidative capacity enhanced by the application of SiO2-NPs by about twofolded. As well as an increase in PEG concentration, the absorption of nutritional elements such as P, K, Mn, Fe, Zn, and Cu was decreased. However, SiO2-NPs application especially at 100 mg L−1 increased the amount of nutrient absorption.

Conclusions

In general, the drought tolerance in Damask rose was associated mainly with its suitable manipulation of antioxidant production and orderly enhancement of nutrient adsorption, so that the effect of SiO2-NPs in improving the qualitative and quantitative characteristics of ʻKashanʼ was more than that of ʻMaraghehʼ. These results briefly highlight that the SiO2-NPs may provide greater tolerance to drought stress in Damask rose.

Graphical Abstract

Introduction

Rosa damascena Mill. is a perennial shrub related to the Rosaceae family used in the perfume, cosmetics, and food industries [1]. The main biologically active molecules isolated from various organs of R. damascena include flavonoids, glycosides (kaempferol, cyanidin 3,5, d-glycosides, and quercetin), gallic acid, terpenes, and anthocyanins. Damask rose leaves are sources of vitamins C, A, B, and K, pectin, tannins, and carotenoids [2]. R. damascena flowers have analgesic, anti-inflammatory, antibacterial, anti-depressant, and anti-viral effects, as well as a diuretic, and are used in traditional medicine as sedatives [2].

Damask rose essential oil has large amounts of alcoholic monoterpenes such as geraniol, citronellol, and phenylethanol, which are the main factors in evaluating the quality and also the odor of rose essential oils [3] which is different also between thorny genotypes than the others without thorn [4]. Increasing the demand for R. damascena essential oil requires extensive propagation of screened genotypes. This may be achieved by in vitro culture. The traditional methods of propagating Damask rose are cuttings and grafting. Some limitations of these traditional methods include seasonal dependence, high reproductive costs, laborious work, and time consumption [5].

In vitro culture techniques are useful for assessing resistance or adaptation to various stressors as extreme factors can be easily controlled [6]. The simplicity of such manipulations makes it possible to study large populations of plants and stress conditions on a small scale and in the least time. Simulation of water deficiency under in vitro culture by some materials such as PEG during the regeneration process is a good method to evaluate the effect of water deficit on morpho-physiological features [7]. PEG is an osmotic laxative that reduces the ambient water potential and has been used to mimic water deficiency without any toxicity in plants [8].

Agricultural practices, genetic factors, and environmental conditions affect crop quality [9]. Damask rose is mostly planted in several altitude regions of Iran, in which Isfahan, Fars, and Kerman have the highest area under cultivation and flower production [10].

Approximately 33% of the land area is exposed to water deficit, which poses a serious threat to plant growth and performance [11]. Water deficit stress causes different effects on the morpho-physiological responses of plants, such as changes in plant water use efficiency, lower growth rate, reduced stem length, leaf area, nutritional imbalance, and photosynthetic reactions [12]. Against oxidative stress due to water stress, antioxidative compounds including phenolics, flavonoids, and anthocyanins have a major effect in reducing the negative features of water deficiency [13]. Phenolic substances act as reactive oxygen species scavengers which accumulated in plants exposed to extreme environmental factors [14]. Dissolved sugars in plants accumulate in response to water stress [15] which can be due to the decomposition of starch [16]. The enhancement in flavonoids and phenols content under severe drought stress is because of the accumulation of soluble sugars in plants, which is attributed to the reduced transport of carbohydrates under water deficiency [17].

It is believed that silicon (Si) is not an essential element for plant growth and development, but several studies demonstrated the benefits of Si application in reducing the harmful effect of abiotic and biotic stress in plants [18]. Previous research emphasized the positive effect of Si in increased tolerance of plants to abiotic stress, especially salt and water stresses [19]. Si may cause osmotic regulation and reduce oxidative injury in plants under stress [20]. Numerous mechanisms modulated in plant growth parameters improvement by Si under drought stress including activation of photosynthetic enzymes [21], activation of antioxidative enzymes, improvement of hydraulic conductivity [22], nutrient uptake [23], root growth and water use efficiency [24], and accumulation of organic osmolytes [25]. Previous studies have shown the positive effect of silicon on plants under hydroponic condition, as Si in nutrient solution cause tolerance against dehydration of explants by modulating in hydraulic relations [26].

Nanoparticles, due to their special characteristics consisting of size, surface charge, shape, and potential interaction with plants, reduce the effect of water stress [27]. With increasing interest in silicon nanoparticles, it has been found that SiO2-NPs penetrate the roots through symplastic and apoplastic pathways [28]. Hamayun et al. [29] stated that the application of Si ameliorated the adverse effects of drought stress induced by PEG on plant growth attributes, such as stem length, root weight, and chlorophyll content.

To our knowledge, there is no research on the effect of nano-silicon (SiO2-NPs) application in Damask rose in drought stress under in vitro conditions. Therefore, the goal of the present work was to study whether the application of SiO2-NPs can reduce drought stress under in vitro culture caused by PEG, thus improving the growth of R. damascena.

Materials and methods

Collect plant material and set the experiment

One-year branches of two Damask rose genotypes of Maragheh and Kashan were collected from Maragheh (37.3892 N, 46.2534 E) and Kashan (33.9850 N, 51.4100°E) according to our previous studies [30]. For surface disinfection, the explants (cuttings) were first placed under running water for half an hour and then for 20 min in water and dishwashing liquid on a shaker. They were rinsed three times with ddH2O and then immersed in 5% NaOCl for 15 min under a flow-box and after three rinses were immersed in 70% ethanol for 180 s. Finally, after washing with distilled water, the explants were ready for culture. The one node shoot explants were recut into 1.5–2 cm length, and they were placed on the MS medium culture. The MS free hormone media [31] were considered as the establishment initial which solidified with 7.5 g of agar. The pH value was adjusted to 5.7 before autoclaving at 121 °C, 150 kPa for 20 min. Five shoot nodes were cultured, in 150 ml culture vessels containing 30 ml of establishment medium as one replication, then kept at 25 ± 2 °C and 16 h-photoperiod (light intensity, 8.85 W/m2) and 60–70% RH. Following explants were screened every day for fungal or bacterial contamination then any contaminated vessels were removed from the experiment collection. Shoots were repeatedly sub-cultured three times at a constant 3 week subculture interval. For the proliferation, the basic medium was the same as the MS medium establishment [31], except that instead of iron stock, 130 mg of the iron sequester and 332 mg of calcium chloride, as well as 0.36 mg L−1 BA and 0.03 mg L−1 IBA were added to the medium. After sterilization and transfer to the flow-box, 25 ml of medium was added to each culture vessel to be used for subculture. Finally, five explants were placed in 150 ml culture vessels containing the proliferation medium as one replication and keep in the growth chamber to be propagated, and for the next stages, small shoots propagated from these explants have been used. This project was arranged as a factorial experiment based on a completely randomized design with four replications.

Preparation of medium containing PEG and SiO 2 -NPs treatment

PEG treatments were used to stimulate water deficiency. Therefore, PEG-6000 was purchased from Merc (Germany) and used at five levels of drought stress (0, 25, 50, 75, and 100 g L−1 with an osmotic pressure of 0, − 0.2, − 0.5, and − 0.9 MPa, respectively) as the first factor. The SiO2-NPs (size < 50 nm) were bought from NANOSANY Corporation (Mashhad, Iran), prepared at three levels (0, 50, and 100 mg L−1) and added to the MS medium in which the two genotypes (Maragheh and Kashan) were cultured as the second factor. After preparing the propagation medium, as mentioned above, the shoot explants were placed in jars containing the culture medium with the mentioned treatments under a flow-box. Therefore, five shoot explants were placed in each culture vessel and kept in the growth chamber and after about 14 days were collected to the traits assessment.

Morpho-physiological traits

After the harvest of samples, morphological traits including shoot height, number of leaves, fresh (FW) and dry weight (DW) of leaves, and biomass percentage were measured using Eq. 1. The leaf area index (LAI) was determined using Image J software:

$${\text{Biomass }}\left( \% \right) \, = \, \left[ {\left( {\text{Dry weight plant}} \right)/\left( {\text{Fresh weight plant}} \right)} \right] \, \times { 1}00$$
(1)

Determination of leaf chlorophyll

The amount of leaf chlorophyll is expressed as chlorophyll index (SPAD readings) using a portable chlorophyll meter (Instruments SPAD-502, Osaka, Japan). For this purpose, three points of the fully expanded leaves were read in each replication and the average was calculated.

Electrolyte leakage (EL)

To study the percentage of the EL of shoots, the youngest fully developed leaves were separated from each shoot and placed in a glass tube containing 20 ml of ddH2O and placed on a shaker for 24 h at room temperature, then centrifuged at 1000 rpm for 24 h. After 24 h, the electrical conductivity (EC) of each sample was measured using an EC meter (Jenway model, UK) and read as EC1. To measure the total leakage of electrolytes (EC2), the leaf samples were placed in an autoclave for 20 min at 120 °C, and after cooling the samples in the ambient temperature, their EC read as EC2. Then, the percentage of EL was evaluated according to the following formula and was recorded as a percentage [32]:

$${\text{EL }}\left( \% \right) \, = \, ({\text{EC}} {1}/{\text{EC2}}) \, \times { 1}00$$

Biochemical traits

Measurement of total soluble carbohydrates (TSC)

For the analysis of TSC, a mixture of 5 ml of 80% ethanol (v/v) and 0.1 g of leaf sample was prepared and centrifuged at 15,000 rpm for 15 min. Then, 3 ml of 0.2% anthrone reagent (0.5 g of anthrone in 250 ml of 72% sulfuric acid) was added to 100 ml of the ethanolic extract and then incubated in boiling water (95 °C) for 10 min [33] After cooling on ice bath, the sample absorbance was read spectrophotometrically (model: AA-6300, Shimadzu, Japan) at 620 nm. The TSC content was expressed in mg g−1 FW [34].

Total flavonoid content (TFC) measurement

For the estimation of TFC, the (Dewanto et al. [35] method was used. The extract (20 μl) was added to 75 μl of NaNO2 solution (5%) and vortexed for 6 min, after adding 0.15 ml of AlCl3 (10%). After 5 min, 0.5 ml of 1 M NaOH solution was added to the previous solution. The final volume reached 2.5 ml and was thoroughly mixed and the absorbance of the mixture was determined using a spectrophotometer at 510 nm. The TFC of R. damascena extracts was expressed as mg catechin per g of dry weight (mg CE g−1 DW).

Anthocyanin assay

To measurement of anthocyanin, 1 ml of the leaf extracts were mixed with 5 ml of 95% ethanol, 1.0 N HCl (85:15, v: v) for 4 h at 4 °C in a dark room [36]. The absorbance was read at 530 nm. The following formula was used to evaluate the anthocyanin content:

$${\text{Anthocyanin content}} = \left( {{\text{absorbance at 53}}0{\text{ nm }} \times {\text{ volume of extraction solution }} \times { 1}00} \right)/\left( {{\text{volume of sample }}\left( {{\text{mL}}} \right) \, \times { 98}.{2}} \right)$$

Total antioxidant activity (TAA)

TAA of R. damascena shoots was calculated by the DPPH method [37]. The extract was obtained from potassium phosphate buffer (100 mM) and added to 1 ml of 50 μM DPPH solution in methanol. The mixture was incubated in a dark room for 20 min. The reduction of DPPH absorbance at 515 nm was recorded using a spectrophotometer. DPPH radical inhibition activity or TAA was evaluated using the following equation:

$${\text{TAA}}\% \, = \, \left[ {\left( {{\text{absorbance control}} - {\text{absorbance sample}}} \right)/\left( {\text{absorbance control}} \right)} \right] \times {1}00.$$

Total phenol content (TPC) measurement

TPC was calculated by the method of Singleton et al. [38]. The mixture was 200 μl of leaf extracts (1% HCl in methanol), 800 μl of methanol, 500 μl of Folin–Ciocalteu reagent (1: 1 with water), and 2500 μl of sodium carbonate solution (20%). After the vortex of the admixture, the tubes were incubated in the darkness at room temperature for 40 min. The absorbance at 725 nm was read using a spectrophotometer. TPC was expressed as mg gallic acid per g DW (mg GAE g−1 DW).

Polyphenol oxidase (PPO) enzyme

The activity of PPO was assayed with 4-methyl catechol as a substrate by the method of Zauberman et al. [39]. The fresh leaf samples (0.5 g) were extracted with 10 ml of 0.1 M sodium phosphate buffer (pH = 6.8) and 0.2 g of polyvinylpyrrolidone (PVP). After centrifugation at 19000 rpm for 20 min, the supernatant was collected as crude enzyme extract. The activity of PPO was measured using 1 ml of 0.1 M sodium phosphate buffer (pH 6.8), 0.5 ml of 100 mM l, 4-methyl catechol, and 0.5 ml of the extract. The increase in absorbance at 410 nm was automatically recorded using a spectrophotometer. The activity of PPO was expressed as µM ml−1 min−1 mg−1 protein or unit mg−1 protein.

Measurement of shoot mineral elements

To measure nutrients, shoots were placed in an oven at 65 °C for 24 h. After the shoots dried, the samples were powdered with an electric mill. After preparing the ash from the plant samples at 550 °C, the extract was extracted using 10 ml of 2 N hydrochloric acid and distilled water to a volume of 50 ml [40]. The content of P with molybdate vanadate reagent using a spectrophotometer at 450 nm, K content (with chlorosis solution 0.87 g L−1) using the Flame photometer (model: PFP7, UK) and some micronutrients (Fe, Mn, Zn, and Cu) were read by dry digestion and combined with hydrochloric acid using the atomic absorption spectrometer.

Statistical analysis

This study was conducted as a factorial experiment based on a completely randomized design with four replications and five explants in each vessel culture. Data were statistically analyzed by MSTAT-C ver. 2.1 software and the means were compared using the LSD test and at 5% probability.

Results

Effect of SiO 2 -NPs and PEG-induced water deficiency on morpho-physiological traits of Damask rose

According to Table 1, increasing in PEG level reduced explant shoot height in both genotypes by 12% and 35.9% in ʻMaraghehʼ and ʻKashanʼ, respectively. However, ʻMaraghehʼ had the highest shoot height in controls treated with 100 mg L−1 SiO2-NPs. Treatment with SiO2-NPs under drought stress to some extent prevented height reduction and its effect was greater in ʻMaraghehʼ. ʻKashanʼ had the lowest height at 75 and 100 g L−1 PEG without SiO2-NPs treatment. The most leaf area index (LAI) was related to the ʻMaraghehʼ at severe drought stress and 100 mg L−1 SiO2-NPs, while ʻKashanʼ had the least LAI at severe drought stress without SiO2-NPs treatment (Table 1). Both genotypes showed a decreasing trend in LAI as well as increasing concentration. However, the LAI of the genotypes under water deficit and well-watered conditions significantly increased with the supplementation of SiO2-NPs (Table 1). In addition, treatment with SiO2-NPs increased LAI in ʻKashanʼ more than in ʻMaraghehʼ. Water deficiency caused to a reduction in shoot FW and DW by 42.5% and 66% in ʻMaraghehʼ and 52.4% and 63.7% in ʻKashanʼ, respectively. SiO2-NPs improved shoot FW and DW of Damask explants under drought stress. Different concentrations of PEG on Damask rose biomass had a decreasing trend by 40.7% and 23.7% in ʻMaraghehʼ and ʻKashanʼ, respectively. Treatment with SiO2-NPs improved biomass percentage up to 16% and 14% in Maragheh and Kashan genotypes, respectively. Both genotypes had the highest percentage of biomass in control plants treated with 100 mg L−1 SiO2-NPs, while the lowest was related to ʻMaraghehʼ under severe drought stress (Table 1).

Table 1 Effect of PEG and SiO2-NPs on Height, Number of leaves and Leaf area index (LAI), Shoot FW and DW and Biomass of Maragheh and Kashan genotypes

EL percentage of both genotypes of Damask rose were reduced under drought stress up to 81% and 62%, respectively, in ʻMaraghehʼ and ʻKashanʼ (Fig. 1). However, treatment with SiO2-NPs reduced the content of EL by 200% and 160% in ʻMaraghehʼ and ʻKashanʼ, respectively. ʻMaraghehʼ had the lowest EL at the control and was treated with 100 mg L−1 SiO2-NPs.

Fig. 1
figure 1

Effect of SiO2-NPs application under drought stress induced by PEG on leaf electrolyte leakage (EL) of two Damask genotypes. Different letters indicate significant differences according to the LSD test at P < 0.05

Chlorophyll content showed a decreasing trend in both Damask rose genotypes as well as an increase in PEG concentration up to 30% and 41%, respectively, in ʻMaraghehʼ and ʻKashanʼ, although the total level of chlorophyll in ʻMaraghehʼ was higher than ʻKashanʼ. Treatment of plants under water deficiency with 100 mg L−1 SiO2-NPs caused to an increase in the level of leaf chlorophyll up to 17% and 30% in ʻMaraghehʼ and ʻKashanʼ, respectively (Fig. 2).

Fig. 2
figure 2

Interaction between drought stress × two Damask genotypes on leaf chlorophyll index a and SiO2-NPs × two Damask genotypes on chlorophyll index b. Different letters indicate significant differences according to the LSD test at P < 0.05

Effect of SiO 2 -NPs and PEG-induced water deficiency on biochemical traits of Damask rose

According to Table 2, Damask rose TSC was increased during different levels of PEG levels by 4.9- and 1.7-fold in ʻMaraghehʼ and ʻKashanʼ, respectively. ʻMaraghehʼ under severe water deficiency and treated with 100 mg L −1 SiO2-NPs had the most TSC, while the control plants in ʻKashanʼ had the least amount of TSC. Similar results were observed in TFC and anthocyanin which the highest being observed in Maragheh genotype treated with 100 mg L−1 SiO2-NPs under severe water deficiency, while the control explants in both genotypes had the least amount of TFC and anthocyanin. Both TFC and anthocyanin had an increasing trend with an increase in drought stress as TFC content in control explants of ʻMaraghehʼ and ʻKashanʼ increased from 1.1 (mg CE g−1 FW) to 6.9 (mg CE g−1 FW) and 5.8, respectively. However, treatment with SiO2-NPs caused more increase in stress and also control Damask explants in the same situations. SiO2-NPs application from 0 to 100 m L−1 caused more increase, more than two folded in ʻMaraghehʼ which was a little bit more than ʻKashanʼ. A similar increase was evaluated in anthocyanin content, with more than twofolded increases in both genotypes as well as increases in drought stress and SiO2-NPs level, so that by 15% and 24% enhancement in anthocyanin content were observed in ʻMaraghehʼ and ʻKashanʼ, respectively as illustrated in Fig. 3.

Table 2 Effect of PEG and SiO2-NPs on total carbohydrate content (TSC), Total Flavonoid content (TFC), Anthocyanin and TAA of Maragheh and Kashan genotypes
Fig. 3
figure 3

Effect of SiO2-NPs and PEG-induced water deficiency on morpho-physiological charactristics in leaves of Maragheh and Kashan genotypes in vitro culture conditions

In addition, our results showed an increase in TAA of the explants extracts of both Damask rose genotypes with enhancement in PEG concentration by twofolded. Under severe drought stress (100 g L−1 PEG), treatment of Damask explants with 100 mg L−1 SiO2-NPs increased TAA by 19% and 28% in ʻMaraghehʼ and ʻKashanʼ, respectively (Table 2).

TPC has heightened along with enhancement in the level of PEG in both Damask roses approximately by twofolded. Application of SiO2-NPs under severe water deficiency caused to increase in TPC in ʻKashanʼ (38%) more than in ʻMaraghehʼ (54%). However, the highest TPC was observed in ʻMaraghehʼ treated with 100 mg L−1 SiO2-NPs under severe drought stress (Fig. 4). The results of PPO activity assay also showed the same results as in TPC observations, the activity of PPO increased by fourfolded in both Damask roses, so that the ʻMaraghehʼ had the highest PPO enzyme activity. In addition, the application of SiO2-NPs led to an increase in enzyme activity, as well (Fig. 5).

Fig. 4
figure 4

Interaction between drought stress × two Damask genotypes on leaf total phenolic content a and SiO2-NPs × two Damask genotypes on total phenolic content b. Different letters indicate significant differences according to the LSD test at P <0.05

Fig. 5
figure 5

Interaction between drought stress × SiO2-NPs on leaf PPO activity a, drought stress × two Damask genotypes on PPO activity b, and SiO2-NPs × two Damask genotypes on PPO activity c. Different letters indicate significant differences according to the LSD test at P < 0.05

Effect of SiO 2 -NPs and PEG-induced water deficiency on nutrient minerals of Damask rose

According to the results of Table 3, the effect of PEG-induced water deficiency and SiO2-NPs on nutritional elements of ʻMaraghehʼ and ʻKashanʼ genotypes showed a significant difference (P ≤ 0.01). Water deficiency caused a decrease in K and P content of Damask leaf by 56% and 52% in ʻMaraghehʼ and 47% and 52% in Kashan, respectively. The highest leaf P and K contents were obtained in ʻMaraghehʼ explants treated with 100 mg L−1 SiO2-NPs and without water deficit. In contrast, ʻKashanʼ showed the least content of P and K in leaves at severe water deficiency and without SiO2-NPs treatment. However, the explants supplemented with SiO2-NPs improved the K and P absorption up to 44% and 30% in ʻMaraghehʼ and 50% and 43% in ʻKashanʼ.

Table 3 Effect of PEG and SiO2-NPs on K and P of Maragheh and Kashan genotypes

The effect of PEG-induced drought stress and SiO2-NPs application had a significant difference in Zn, Mn, and Cu concentrations in Damask explants (Table 4) as along with increased PEG concentration, the amount of these elements was decreased in comparison with un-stress explants. Use of SiO2-NPs caused an improvement in microelements such as Zn (75% vs 44%) and Mn (79% vs 45%) absorption in ʻKashanʼ more than ʻMaraghehʼ, although the effect of SiO2-NPs on Cu (60% vs 56%) absorption was not so impressive. The Fe content showed a declining trend along with rising in the level of PEG up to 49% in ʻMaraghehʼ and 51% in ʻKashanʼ, although the initial content of Fe was more in ʻMaraghehʼ compared to ʻKashanʼ. Treatment with SiO2-NPs enhanced the Fe absorption up to 38% and 58% in ʻMaraghehʼ and ʻKashanʼ, respectively (Fig. 6).

Table 4 Effect of PEG and SiO2-NPs on nutritional elements of Maragheh and Kashan genotypes
Fig. 6
figure 6

Interaction between drought stress × two Damask genotypes on leaf Fe a and SiO2-NPs × two Damask genotypes on Fe b. Different letters indicate significant differences according to the LSD test at P < 0.05

Multivariate analysis of Damask Rose under normal and PEG treatments supplemented with SiO 2 -NPs

Pearson’s correlations of some growth parameters, biochemical traits, and nutrient content are presented in Fig. 7. The findings indicated that PPO, TFD, anthocyanin, carbohydrate, TPC, and TAA positively correlated with each other, and also a significant positive was observed among height, LAI, DW, FW, SPAD, Zn, MN, Fe, Cu, K, and P content. On the other hand, EL negatively correlated with height, LAI, DW, FW, SPAD, Zn, MN, Fe, Cu, K, P content, number of leaves, and biomass. Finally, a positive correlation was observed between the number of leaves and biomass (Fig. 7a).

Fig. 7
figure 7

Heat maps of Pearson correlation heat map a and loading biplot of the evaluated traits b of the growth parameters, biochemical and nutrient changes in Damask rose under PEG-induced osmotic stress treated with SiO2-NPs. Heat maps representing electrolyte leakage (EL), polyphenol oxidase (PPO), total flavonoids content (TFD), Anthocaynin, total soluble carbohydrate, total phenolics content (TPC), height, leaf area index (LAI), leaf dry weight (DW), leaf fresh weight (FW), SPAD, number of leaves, biomass and some nutrient content such as Zn, Mn, Fe, Cu, K, P

The principal component analysis (PCA) clarified that two PCA were contributing 86.1% of the total variation. The first PCA was the most effective by a variance of 56. 8% and the second PCA elucidated 27.3% of the total variance. Moreover, the biplot of traits indicated which traits were classified into three groups. The first group included PPO, TFD, anthocyanin, carbohydrates, TPC, TAA, and EL; the second group contained height, LAI, DW, FW, SPAD, Zn, Mn, Fe, Cu, K, and P content; and finally, the third group included a number of leaves and biomass (Fig. 7b).

Discussion

Drought stress, in addition to a reduction in plant growth and reproductivity, causes a change in some metabolic pathways which caused plant tolerance to stress. Drought tolerance depends on the reactions to continue primary metabolic processes and improved plant tolerance. Damask rose is also similar to other plants that reacted against to drought stress and coped to stress via changes in physiological characteristics and the content of absorption of elements. Our findings showed that along with increases in drought stress levels, the Damask rose biomass decreased as well as a reduction in LAI and plant length. These results were in line with Hessini et al. [41], who demonstrated decrease in FW and DW by approximately 29% under moderate (50% FC) water stress and 48% and 33% under severe (25% FC) water stress, respectively, relative to the well-watered R. damascena. The mechanism for Si-induced biomass increase could be attributed to the involvement of Si in cell wall deposition and nutrient absorption [42] as Hattori et al. [43] demonstrated that silicon-induced acceleration of biomass production in sorghum only when the plants were subjected to water deficit conditions. In fact, drought stress causes a decrease in cell enlargement and swelling, and as a result, growth decreases as demonstrated by reduced fresh/dry weight (Table 1). On the other hand, along with an increase in the severity of water deficiency, as leaf photosynthesis decreases, the carbohydrate needs for osmotic regulation in plants increase, and then, root growth is inevitably prevented [44].

The application of nano-silicon prevented EL (%) increase, as illustrated in Fig. 1. It is suggested that silicon is identified as an immobile element inside the plant, and as soon as it accumulates inside the cell, changed to a polymerized gel which is no longer usable for the plant. Therefore, it caused the cells to be strong and stable, thus reducing the amount of leakage of electrolytes in the plant [45]. It is demonstrated that Si is a mineral nutrient that caused to improve the water use efficiency of plants [46]. In addition, the positive effect of SiO2-NPs on leaf chlorophyll content seems to be related to its role as a cofactor in pigments biosynthesis [47] and inhibition of the activity of chlorophyllase which becomes more active under extreme conditions [48]. Furthermore, silicon accumulates in the width of the leaf and increases the strength of the leaves which caused to increase in the concentration of chlorophyll in the leaf area. Silicon has also the potential to increase the photosynthesis rate by stabilizing the chloroplast structure [45] and increasing the efficiency of photosystem II [49]. A reduction in the amount of chlorophyll with the increment along with an increase in drought severity is also reported by Hsu and Kao [50] which was in agreement with our findings in both genotypes. On the other hand, silicon caused to firmer xylem cell walls [51] which are responsible for water transportation into the plant [52]. Besides, the accumulation of silicon in the leaf, forms a layer double of silicon [53] and reduces transpiration. Recent reports showed the positive effects of bulk silica [54, 55] and nanoparticles of silicon [30] in photosynthesis parameters improvement of some plants under water deficiency.

Total carbohydrates are also a group of compatible osmolytes that accumulate under water-deficient conditions and act as an osmoprotectant. An increase in carbohydrates due to drought stress is related to osmotic regulation and turgor maintenance which is caused to stabilizing membranes and proteins [56]. In the present work, with the increase in water deficiency, the content of carbohydrates increased from 4.12 and 4.8 mg g−1FW in control explants to 20.5 and 8.4 mg g−1 FW under 100 g L−1 PEG, respectively, in ʻMaraghehʼ and ʻKashanʼ. In general, the increase of carbohydrates during stress may be caused by the breakdown of polysaccharides, such as starch, biosynthesis of sugars via non-photosynthetic pathways, the failure to convert these compounds into other products, the reduction of transfer from leaves to other organs, or the cessation of growth, which also increases carbohydrates content [57] which is the passive reaction to plant growth prevention under drought stress [58]. In the present study, total polyphenol content, total flavonoid content, total antioxidant capacity (DPPH), and anthocyanin were significantly increased with the increase in the PEG concentration. extreme conditions, such as drought stress, caused to increase in the anthocyanin pigments in leaves that have an antioxidative role in the protection of the photosynthetic system against light oxidation [59]. Results showed a synergistic effect between drought stress and SiO2-NPs, as they strengthen each other’s effect and caused increased levels of biochemical traits under severe water deficiency and 100 mg L−1 SiO2-NPs. As a matter of fact, Verma and Dubey [60] reported that silicon has a significant effect on the metabolism of soluble sugars and the partitioning of photosynthetic substances in growing plants increases it. Therefore, it assumed that silicon keeps the carbohydrate reserve of plants under stress, for metabolic processes and maintenance of basic metabolism. On the other hand, the research on PEG-induced water deficiency in wheat showed that the reason for the increase in the levels of phenolic compounds is the increase in the activity of the biosynthetic enzymes of phenols [61]. Phenolics accumulation during stress conditions can act as a sign/alarm and start a cascade of other reactions that ultimately lead to an increase in stress tolerance [62]. An increased DPPH scavenging activity during water deficit conditions was also observed in Salvia [63] and Fraxinus [64]. Phenolic substances are derivatives of the phenylpropanoid pathway which are a part of the non-enzymatic antioxidative defense. It seems that silicon may directly or indirectly induce the genes of the biosynthesis pathway of these compounds and thereby increase the plant's resistance to drought stress [65].

The nutrient element content of Damask rose explants was remarkably influenced by water deficiency. Totally, the concentration of measured elements including K, P, Cu, Zn, Fe, and Mn sharply reduced with the increment of water deficiency. Brown et al. [66] also showed the same results relying on the reduced levels of absorbed Fe under water deficit. Pei et al. [54] reported that Si decreased Mg, K, and Ca contents in wheat under water deficiency. The same results rely on the reduction in Zn, Mn, and Cu obtained by Sarker and Oba [67] and Gunes et al. [68]. It is believed that, besides the harmful effect of water deficiency on plant growth and productivity, problems with nutrient minerals can occur as a secondary effect, because it is dependent on the moisture in the soil to move through the soil matrix and be taken up by plants [69]. Under drought stress, roots are not able to absorb most of the nutrients from the rhizosphere because of the lack of root activity and also, slow ion diffusion and water movement rates [70] Furthermore, the mineralization process depends on microorganisms and enzyme activity, which may be influenced by drought. Therefore, water deficit causes low nutrient availability in the soil and lower nutrient transport in plants [71]. It seems that the severe decrease in root length under water deficiency is the most important reason for reducing the absorption of potassium (K) in the soil by the plant. Increasing the amount of K is an important indicator in tolerance to drought stress and it seems that treatment with SiO2-NPs had the potential to increase the plant K in ʻMaraghehʼ more than ʻKashanʼ (44% vs 30%), as shown in Table 3. Besides, Mahouachi [72] found reduced amounts of K in bananas under water deficiency. Similar findings were obtained by Restrepo-Diaz et al. [73] in the leaves of olive plants under water stress, regardless of nutritional status. By reducing the amount of soil water, the mobility of K is reduced, and consequently, the availability of K by plants root is also reduced [71]. It has been reported that the reason for leaf K reduction under water deficiency may be due to the movement of this element from the leaves to the roots, because K acts as an osmotic protector. Potassium has less solubility in arid conditions and as a result is less absorbed by the plant [74]. Phosphate (P) moves through diffusion in the soil so that under water deficiency, the radii of water-filled pores decrease, tortuosity increases, and P mobility decreases [75]. Water deficit causes a reduction in P absorption and transport in plants. A decrease in available P forms and an increase in occluded P in the soil caused to reduce in P uptake and consequently induces lower foliar P content [76, 77]. Sardans and Peñuelas [77] demonstrated that a 22% reduction in soil moisture produced a 40% decrease in the accumulated aboveground P content in plants, primarily because there was a little increase in aerial biomass. Jin et al. [78] showed that water deficit caused to decrease in plant growth and development, and total P uptake. Despite 21 mg of absorbable P per kilogram of soil, which is more than plants need, immobilization of phosphorus in high acidity and its stabilization, especially in water deficiency conditions, is the major reason for the reduction of P accumulation in the leaf tissue [79].

Silicon increases root endodermal silicification and improve the water balance in cells [80]. Under stress conditions, the application of silicon offers beneficial effects for plant growth and development, promoting enhanced water uptake and alleviating oxidative damage [81]. It seems that the effect of SiO2-NPs in microelements absorption especially in the Kashan genotype which has moderate tolerance to drought stress [30], is more drastic than that in ʻMaraghehʼ. Several destinations are estimated for silicon once it has entered the plant symplast; In the roots, it is mostly found in endo- and exo-dermal tissues where it could be integrated into the cell wall by cross-linking with other wall components, such as hemicelluloses, pectins, and phenolic compounds [82]. In the shoots, high concentrations of silicic acid led to its autopolymerisation into silica [83]. Deposited silica can be found in the form of phytoliths which occur in a multitude of shoot tissues [84]. Alternatively, silica accumulates in or beneath the cuticle layer of the cell wall in epidermal cell layers and tissues that surround the vasculature [85]. In general, SiO2-NPs efficiency seems to be more pronounced in ʻKashanʼ which is considered as a moderate tolerance to water deficiency [30]. In both genotypes exposed to drought stress, nano-silicon application increased chlorophyll, indicating the synthesis of new pigments, and maintenance of chlorophyll previously existing. Although, the exact mechanisms are still under debate but it seems that silicon can relieve drought by lowering root hydraulic conductance and reduction of water loss through transpiration, activation of antioxidant capacity and enhancement of minerals. However, the positive effects of silicon treatment in the plants are obvious not only under stress conditions but also have confirmed in stress-free conditions.

Conclusion

In general, the results demonstrated that silicon nanoparticles play a key role in maintaining critical physiological and biochemical functions as well as minerals adsorption in Damask rose under water deficiency. Silicon treatment under water deficiency increased carbohydrates content compared to the lack of silicon application. It is concluded that the effect of SiO2-NPs in Kashan was more drastic in increasing the antioxidant capacity and total phenolic compounds. In addition, the effect of SiO2-NPs application in Kashan genotype was more efficient in microelements adsorption compared to Maragheh. In general, having more biomass and leaf area under drought probably makes ʻMaraghehʼ more tolerant than ʻKashanʼ but the effect of SiO2-NPs application in sensitive genotypes is more pronounced. Finally, further research should be conducted to understand the possible synergistic effect of SiO2-NPs on Damask rose various physiological events such as biostimulation elucidating the up-regulation of gene expression, crosstalk amongst phytohormones that could be altered and enzyme activity.

Availability of data and materials

Correspondence and requests for materials should be addressed to H.S.H.

References

  1. Naquvi KJ, Ansari SH, Ali M, Najmi AK. Volatile oil composition of Rosa damascena Mill. (Rosaceae). New Delhi: AkiNik Publications; 2014.

    Google Scholar 

  2. Baydar NG, Baydar H. Phenolic compounds, antiradical activity and antioxidant capacity of oil-bearing rose (Rosa damascena Mill.) extracts. Ind Crops Prod. 2013;41:375–80.

    Article  CAS  Google Scholar 

  3. Batooli H, Safaei-Ghomi J. Comparison of essential oil composition of flowers of three Rosa damascena Mill. genotypes from Kashan. J Med Plants. 2012;11:157–66.

    Google Scholar 

  4. Seyed Hajizadeh H, Ebadi B, Morshedloo MR, Abdi GA. Morphological and phytochemical diversity among some Iranian Rosa damascena Mill. landraces. Tehran: Islamic Azad University; 2021.

    Google Scholar 

  5. Gorji-Chakespari A, Nikbakht AM, Sefidkon F, Ghasemi-Varnamkhasti M, Valero EL. Classification of essential oil composition in Rosa damascena Mill. genotypes using an electronic nose. J Appl Res Med Aromat Plants. 2017;4:27–34.

    Google Scholar 

  6. Errabii T, Gandonou CB, Essalmani H, Abrini J, Idaomar M, Skali-Senhaji N. Growth, proline and ion accumulation in sugarcane callus cultures under drought-induced osmotic stress and its subsequent relief. African J Biotechnol. 2006;5:16.

    Google Scholar 

  7. Sakthivelu G, Devi MKA, Giridhar P, Rajasekaran T, Ravishankar GA, Nedev T, et al. Drought-induced alterations in growth, osmotic potential and in vitro regeneration of soybean cultivars. Gen Appl Plant Physiol. 2008;34:103–12.

    CAS  Google Scholar 

  8. Matheka JM, Magiri E, Rasha AO, Machuka J. In vitro selection and characterization of drought tolerant somaclones of tropical maize (Zea mays L.). Biotechnology. 2008;7:641–650.

    Article  CAS  Google Scholar 

  9. Rebey IB, Jabri-Karoui I, Hamrouni-Sellami I, Bourgou S, Limam F, Marzouk B. Effect of drought on the biochemical composition and antioxidant activities of cumin (Cuminum cyminum L.) seeds. Ind Crops Prod. 2012;36:238–45.

    Article  Google Scholar 

  10. Ahmadi Y, Khosh-Khui M, Salehi H, Eshghi S, Kamgar Haghighi AA, Karami A. Effect of Salinity Stress on Growth and Biochemical Characteristics of Three Population of Damask Rose of Iran. Iran J Hortic Sci Technol. 2019;20:87–100.

    Google Scholar 

  11. Wang Y, Zhao W, Zhang Q, Yao Y. Characteristics of drought vulnerability for maize in the eastern part of Northwest China. Sci Rep. 2019;9:1–9.

    Google Scholar 

  12. Xu H, Biswas DK, Li W-D, Chen S-B, Zhang L, Jiang G-M, et al. Photosynthesis and yield responses of ozone-polluted winter wheat to drought. Photosynthetica. 2007;45:582–8.

    Article  CAS  Google Scholar 

  13. Hichem H, Mounir D, et al. Differential responses of two maize (Zea mays L.) varieties to salt stress: changes on polyphenols composition of foliage and oxidative damages. Ind Crops Prod. 2009;30:144–51.

    Article  CAS  Google Scholar 

  14. D’Souza MR, Devaraj VR. Biochemical responses of Hyacinth bean (Lablab purpureus) to salinity stress. Acta Physiol Plant. 2010;32:341–53.

    Article  Google Scholar 

  15. Zhang J, Dell B, Conocono E, Waters I, Setter T, Appels R. Water deficits in wheat: fructan exohydrolase (1-FEH) mRNA expression and relationship to soluble carbohydrate concentrations in two varieties. New Phytol. 2009;181:843–50.

    Article  CAS  PubMed  Google Scholar 

  16. Fischer C, Höll W. Food reserves of scots pine (Pinus sylvestris L.). Trees. 1991;5:187–95.

    Article  Google Scholar 

  17. Ibrahim MH, Jaafar HZE. Photosynthetic capacity, photochemical efficiency and chlorophyll content of three varieties of labisia pumila benth. exposed to open field and greenhouse growing conditions. Acta Physiol Plant. 2011;33:2179–85.

    Article  CAS  Google Scholar 

  18. Etesami H, Jeong BR. Silicon (Si): review and future prospects on the action mechanisms in alleviating biotic and abiotic stresses in plants. Ecotoxicol Environ Saf. 2018;147:881–96.

    Article  CAS  PubMed  Google Scholar 

  19. Bauer P, Elbaum R, Weiss IM. Calcium and silicon mineralization in land plants: transport, structure and function. Plant Sci. 2011;180:746–56.

    Article  CAS  PubMed  Google Scholar 

  20. Hajizadeh HS, Asadi M, Zahedi SM, Hamzehpour N, Rasouli F, Helvaci M, et al. Silicon dioxide-nanoparticle nutrition mitigates salinity in gerbera by modulating ion accumulation and antioxidants. Folia Hortic. 2021;33:91–105.

    Article  Google Scholar 

  21. Yin L, Wang S, Liu P, Wang W, Cao D, Deng X, et al. Silicon-mediated changes in polyamine and 1-aminocyclopropane-1-carboxylic acid are involved in silicon-induced drought resistance in Sorghum bicolor L. Plant Physiol Biochem. 2014;80:268–77.

    Article  CAS  PubMed  Google Scholar 

  22. Shen X, Zhou Y, Duan L, Li Z, Eneji AE, Li J. Silicon effects on photosynthesis and antioxidant parameters of soybean seedlings under drought and ultraviolet-B radiation. J Plant Physiol. 2010;167:1248–52.

    Article  CAS  PubMed  Google Scholar 

  23. Chen W, Yao X, Cai K, Chen J. Silicon alleviates drought stress of rice plants by improving plant water status, photosynthesis and mineral nutrient absorption. Biol Trace Elem Res. 2011;142:67–76.

    Article  CAS  PubMed  Google Scholar 

  24. Lux A, Luxová M, Hattori T, Inanaga S, Sugimoto Y. Silicification in sorghum (Sorghum bicolor) cultivars with different drought tolerance. Physiol Plant. 2002;115:87–92.

    Article  CAS  PubMed  Google Scholar 

  25. Rizwan M, Ali S, Ibrahim M, Farid M, Adrees M, Bharwana SA, et al. Mechanisms of silicon-mediated alleviation of drought and salt stress in plants: a review. Environ Sci Pollut Res. 2015;22:15416–31.

    Article  CAS  Google Scholar 

  26. Gao X, Zou C, Wang L, Zhang F. Silicon decreases transpiration rate and conductance from stomata of maize plants. J Plant Nutr. 2006;29:1637–47.

    Article  CAS  Google Scholar 

  27. Tarafdar JC, Xiong Y, Wang W-N, Quinl D, Biswas P. Standardization of size, shape and concentration of nanoparticle for plant application. Appl Biol Res. 2012;14:138–44.

    Google Scholar 

  28. Shi Y, Zhang Y, Yao H, Wu J, Sun H, Gong H. Silicon improves seed germination and alleviates oxidative stress of bud seedlings in tomato under water deficit stress. Plant Physiol Biochem. 2014;78:27–36.

    Article  CAS  PubMed  Google Scholar 

  29. Hamayun M, Sohn E-Y, Khan SA, Shinwari ZK, Khan AL, Lee I-J, et al. Silicon alleviates the adverse effects of salinity and drought stress on growth and endogenous plant growth hormones of soybean (Glycine max L.). Pak J Bot. 2010;42:1713–22.

    CAS  Google Scholar 

  30. Hajizadeh HS, Azizi S, Rasouli F, Okatan V. Modulation of physiological and biochemical traits of two genotypes of Rosa damascena Mill. by SiO2-NPs under In vitro drought stress. BMC Plant Biol. 2022;22:1–16.

    Article  Google Scholar 

  31. Murashige T, Skoog F. A revised medium for rapid growth and bio assays with tobacco tissue cultures. Physiol Plant. 1962;15:473–97.

    Article  CAS  Google Scholar 

  32. Sekozawa Y, Sugaya S, Gemma H, Iwahori S. Cold tolerance in’Kousui’Japanese pear and possibility for avoiding frost injury by treatment with n-propyl dihydrojasmonate. HortScience. 2003;38:288–92.

    Article  CAS  Google Scholar 

  33. Irigoyen JJ, Einerich DW, Sánchez-Diaz M. Water stress induced changes in concentrations of proline and total soluble sugars in nodulated alfalfa (Medicago sativd) plants. Physiol Plant. 1992;84:55–60.

    Article  CAS  Google Scholar 

  34. Dubois M, Gilles KA, Hamilton JK, Rebers PA, t, Smith F. Colorimetric method for determination of sugars and related substances. Anal Chem. 1956;28:350–6.

    Article  CAS  Google Scholar 

  35. Dewanto V, Wu X, Adom KK, Liu RH. Thermal processing enhances the nutritional value of tomatoes by increasing total antioxidant activity. J Agric Food Chem. 2002;50:3010–4.

    Article  CAS  PubMed  Google Scholar 

  36. Wagner GJ. Content and vacuole/extravacuole distribution of neutral sugars, free amino acids, and anthocyanin in protoplasts. Plant Physiol. 1979;64:88–93.

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  37. Abe N, Murata T, Hirota A. Novel DPPH radical scavengers, bisorbicillinol and demethyltrichodimerol, from a fungus. Biosci Biotechnol Biochem. 1998;62:661–6.

    Article  CAS  PubMed  Google Scholar 

  38. Singleton VL, Orthofer R, Lamuela-Raventós R. Analysis of total phenols and other oxidation substrates and antioxidants by means of folin-ciocalteu reagent. Methods Enzymol. 1999;299:152–178. https://doi.org/10.1016/S0076-6879(99)99017-1

    Article  CAS  Google Scholar 

  39. Zauberman G, Ronen R, Akerman M, Weksler A, Rot I, Fuchs Y. Post-harvest retention of the red colour of litchi fruit pericarp. Sci Hortic. 1991;47:89–97.

    Article  CAS  Google Scholar 

  40. Wallinga I, Van Vark W, Houba VJG, der Lee JJ. Soil and plant analysis, series of syllabi part 7, plant analysis procedure. Netherlands: Wageningen Agric Univ Wageningen; 1989.

    Google Scholar 

  41. Hessini K, Wasli H, Al-Yasi HM, Ali EF, Issa AA, Hassan FAS, et al. Graded moisture deficit effect on secondary metabolites, antioxidant, and inhibitory enzyme activities in leaf extracts of Rosa damascena Mill. var. trigentipetala. Horticulturae. 2022;8:177.

    Article  Google Scholar 

  42. Zarafshar M, Akbarinia M, Askari H, Hosseini SM, Rahaie M, Struve D. Toxicity assessment of SiO2 nanoparticles to pear seedlings. Int J Nanosci Nanotechnol. 2015;11:13–22.

    Google Scholar 

  43. Hattori T, Inanaga S, Araki H, An P, Morita S, Luxová M, et al. Application of silicon enhanced drought tolerance in Sorghum bicolor. Physiol Plant. 2005;123:459–66.

    Article  CAS  Google Scholar 

  44. Lu Z, Neumann PM. Water-stressed maize, barley and rice seedlings show species diversity in mechanisms of leaf growth inhibition. J Exp Bot. 1998;49:1945–52.

    Article  CAS  Google Scholar 

  45. Liang Y, Sun W, Zhu Y-G, Christie P. Mechanisms of silicon-mediated alleviation of abiotic stresses in higher plants: a review. Environ Pollut. 2007;147:422–8.

    Article  CAS  PubMed  Google Scholar 

  46. Sattar A, Cheema MA, Sher A, Ijaz M, Wasaya A, Yasir TA, et al. Foliar applied silicon improves water relations, stay green and enzymatic antioxidants activity in late sown wheat. SILICON. 2020;12:223–30.

    Article  CAS  Google Scholar 

  47. Misra A, Srivastava AK, Srivastava NK, Khan A. Zn-acquisition and its role in growth, photosynthesis, photosynthetic pigments, and biochemical changes in essential monoterpene oil (s) of Pelargonium graveolens. Photosynthetica. 2005;43:153–5.

    Article  CAS  Google Scholar 

  48. Feng J, Shi Q, Wang X, Wei M, Yang F, Xu H. Silicon supplementation ameliorated the inhibition of photosynthesis and nitrate metabolism by cadmium (Cd) toxicity in Cucumis sativus L. Sci Hortic. 2010;123:521–30.

    Article  CAS  Google Scholar 

  49. Al-aghabary K, Zhu Z, Shi Q. Influence of silicon supply on chlorophyll content, chlorophyll fluorescence, and antioxidative enzyme activities in tomato plants under salt stress. J Plant Nutr. 2005;27:2101–15.

    Article  Google Scholar 

  50. Hsu S-Y, Kao CH. Differential effect of sorbitol and polyethylene glycol on antioxidant enzymes in rice leaves. Plant Growth Regul. 2003;39:83–90.

    Article  CAS  Google Scholar 

  51. Ma JF, Mitani N, Nagao S, Konishi S, Tamai K, Iwashita T, et al. Characterization of the silicon uptake system and molecular mapping of the silicon transporter gene in rice. Plant Physiol. 2004;136:3284–9.

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  52. McElrone AJ, Pockman WT, Martinez-Vilalta J, Jackson RB. Variation in xylem structure and function in stems and roots of trees to 20 m depth. New Phytol. 2004;163:507–17.

    Article  PubMed  Google Scholar 

  53. da Silva Lobato AK, Guedes EMS, Marques DJ, de Oliveira Neto CF. Silicon: a benefic element to improve tolerance in plants exposed to water deficiency. In: Akinci S, editor. Responses org to water stress. London: InTech; 2013.

    Google Scholar 

  54. Pei ZF, Ming DF, Liu D, Wan GL, Geng XX, Gong HJ, et al. Silicon improves the tolerance to water-deficit stress induced by polyethylene glycol in wheat (Triticum aestivum L.) seedlings. J Plant Growth Regul. 2010;29:106–15.

    Article  CAS  Google Scholar 

  55. Zhang C, Moutinho-Pereira JM, Correia C, Coutinho J, Gonçalves A, Guedes A, et al. Foliar application of Sili-K®increases chestnut (Castanea spp.) growth and photosynthesis, simultaneously increasing susceptibility to water deficit. Plant Soil. 2013;365:211–25.

    Article  CAS  Google Scholar 

  56. Bohnert HJ, Nelson DE, Jensen RG. Adaptations to environmental stresses. Plant Cell. 1995;7:1099.

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  57. Ehdaie B, Alloush GA, Madore MA, Waines JG. Genotypic variation for stem reserves and mobilization in wheat: II. postanthesis changes in internode water-soluble carbohydrates. Crop Sci. 2006;46:2093–103.

    Article  CAS  Google Scholar 

  58. Esmaili S, Tavallali V, Amiri B. Nano-silicon complexes enhance growth, yield, water relations and mineral composition in tanacetum parthenium under water deficit stress. SILICON. 2021;13:2493–508.

    Article  CAS  Google Scholar 

  59. He F, Mu L, Yan G-L, Liang N-N, Pan Q-H, Wang J, et al. Biosynthesis of anthocyanins and their regulation in colored grapes. Molecules. 2010;15:9057–91.

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  60. Verma S, Dubey RS. Effect of cadmium on soluble sugars and enzymes of their metabolism in rice. Biol Plant. 2001;44:117–23.

    Article  CAS  Google Scholar 

  61. Tian X, Lei Y. Nitric oxide treatment alleviates drought stress in wheat seedlings. Biol Plant. 2006;50:775–8.

    Article  CAS  Google Scholar 

  62. Andersen OM, Markham KR. Flavonoids: chemistry, biochemistry and applications. Boca Raton: CRC Press; 2005.

    Book  Google Scholar 

  63. Bettaieb I, Hamrouni-Sellami I, Bourgou S, Limam F, Marzouk B. Drought effects on polyphenol composition and antioxidant activities in aerial parts of Salvia officinalis L. Acta Physiol Plant. 2011;33:1103–11.

    Article  CAS  Google Scholar 

  64. Štajner D, Orlovic S, Popovic BM, Kebert M, Galic Z. Screening of drought oxidative stress tolerance in serbian melliferous plant species. African J Biotechnol. 2011;10:1609–14.

    Google Scholar 

  65. Dragišić Maksimović J, Bogdanović J, Maksimović V, Nikolic M. Silicon modulates the metabolism and utilization of phenolic compounds in cucumber (Cucumis sativus L.) grown at excess manganese. J Plant Nutr Soil Sci. 2007;170:739–44.

    Article  Google Scholar 

  66. Brown CE, Pezeshki SR, DeLaune RD. The effects of salinity and soil drying on nutrient uptake and growth of Spartina alterniflora in a simulated tidal system. Environ Exp Bot. 2006;58:140–8.

    Article  CAS  Google Scholar 

  67. Sarker U, Oba S. Drought stress enhances nutritional and bioactive compounds, phenolic acids and antioxidant capacity of amaranthus leafy vegetable. BMC Plant Biol. 2018;18:1–15.

    Article  Google Scholar 

  68. Gunes A, Pilbeam DJ, Inal A, Coban S. Influence of silicon on sunflower cultivars under drought stress, I: growth, antioxidant mechanisms, and lipid peroxidation. Commun Soil Sci Plant Anal. 2008;39:1885–903.

    Article  CAS  Google Scholar 

  69. Taiz L, Zeiger E. Fisiologia vegetal. Castelló: Universitat Jaume I; 2006.

    Google Scholar 

  70. Dubey RS, Pessarakli M. Physiological mechanisms of nitrogen absorption and assimilation in plants under stressful conditions. In: Pessarakli M, editor. Handbook of plant and crop physiology. Boca Raton: CRC Press; 2001.

    Google Scholar 

  71. Hu Z-Y, Zhu Y-G, Li M, Zhang L-G, Cao Z-H, Smith FA. Sulfur (S)-induced enhancement of iron plaque formation in the rhizosphere reduces arsenic accumulation in rice (Oryza sativa L.) seedlings. Environ Pollut. 2007;147:387–93.

    Article  CAS  PubMed  Google Scholar 

  72. Mahouachi J. Growth and mineral nutrient content of developing fruit on banana plants (Musa acuminata AAA, ‘grand nain’) subjected to water stress and recovery. J Hortic Sci Biotechnol. 2007;2007(82):839–44.

    Article  Google Scholar 

  73. Restrepo-Diaz H, Benlloch M, Navarro C, Fernández-Escobar R. Potassium fertilization of rainfed olive orchards. Sci Hortic. 2008;116:399–403.

    Article  CAS  Google Scholar 

  74. Osuagwu GGE, Edeoga HO, Osuagwu AN. The influence of water stress (drought) on the mineral and vitamin potential of the leaves of Ocimum gratissimum L. Recent Res Sci Technol. 2010;2:27–33.

    CAS  Google Scholar 

  75. Faye I, Diouf O, Guisse A, Sene M, Diallo N. Characterizing root responses to low phosphorus in pearl millet Pennisetum glaucum (L.) R. Br. Agron J. 2006;98:1187–94.

    Article  CAS  Google Scholar 

  76. Marschner H. Mineral nutrition of higher plants academic press san diego. 2nd ed. CA: Acad Press San Diego; 1995.

    Google Scholar 

  77. Sardans J, Peñuelas J. Increasing drought decreases phosphorus availability in an evergreen mediterranean forest. Plant Soil. 2004;267:367–77.

    Article  CAS  Google Scholar 

  78. Jin J, Wang G, Liu X, Pan X, Herbert SJ, Tang C. Interaction between phosphorus nutrition and drought on grain yield, and assimilation of phosphorus and nitrogen in two soybean cultivars differing in protein concentration in grains. J Plant Nutr. 2006;29:1433–49.

    Article  CAS  Google Scholar 

  79. Devau N, Le Cadre E, Hinsinger P, Jaillard B, Gérard F. Soil pH controls the environmental availability of phosphorus: experimental and mechanistic modelling approaches. Appl Geochem. 2009;24:2163–74.

    Article  CAS  Google Scholar 

  80. Farooq M, Wahid A, Kobayashi N, Fujita D, Basra SMA. Plant drought stress: effects, mechanisms and management. Sustain Agric. 2009. https://doi.org/10.1007/978-90-481-2666-8_12.

    Article  Google Scholar 

  81. Pulz AL, Crusciol CAC, Lemos LB, Soratto RP. Silicate and limestone effects on potato nutrition, yield and quality under drought stress. Rev Bras Ciência Do Solo. 2008;32:1651–9.

    Article  CAS  Google Scholar 

  82. Fleck AT, Schulze S, Hinrichs M, Specht A, Waßmann F, Schreiber L, et al. Silicon promotes exodermal casparian band formation in Si-accumulating and Si-excluding species by forming phenol complexes. PLoS ONE. 2015. https://doi.org/10.1371/journal.pone.0138555.

    Article  PubMed  PubMed Central  Google Scholar 

  83. Yoshida S, Ohnishi Y, Kitagishi K. Histochemistry of silicon in rice plant: III. the presence of cuticle-silica double layer in the epidermal tissue. Soil Sci Plant Nutr. 1962;8:1–5.

    Google Scholar 

  84. Shakoor SA, Bhat MA, Mir SH. Phytoliths in plants: a review. J Bot Sci. 2014;3:10–24.

    Google Scholar 

  85. Peleg Z, Saranga Y, Fahima T, Aharoni A, Elbaum R. Genetic control over silica deposition in wheat awns. Physiol Plant. 2010;140:10–20.

    Article  CAS  PubMed  Google Scholar 

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Acknowledgements

The present study was carried out by the use of facilities and materials at the University of Maragheh and the paper is published as the second part of a research project No. 1024 supported by the University of Maragheh, research affairs office.

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The current research has received no funding from agencies in the public, commercial, or not-for-profit sectors.

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Formal analysis, experiment design, investigation, methodology conceptualization, validation and resources, HSH, SA, FR, review and editing, visualization, writing–original draft preparation, data curation, HSH, FR and OK. All authors read and approved the final manuscript.

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Correspondence to Hanifeh Seyed Hajizadeh.

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Seyed Hajizadeh, H., Azizi, S., Rasouli, F. et al. Evaluation of nano-silicon efficiency on compatible solutes and nutrient status of Damask rose affected by in vitro simulated drought stress. Chem. Biol. Technol. Agric. 10, 22 (2023). https://doi.org/10.1186/s40538-023-00397-5

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