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Effect of Selenium, Copper and Manganese Nanocomposites in Polysaccharide Matrices on the Content of Photosynthetic Pigments in Potato Leaf Tissues
1 Laboratory of Plant-Microbe Interactions, Siberian Institute of Plant Physiology and Biochemistry, Siberian Branch of the Russian Academy of Sciences, Irkutsk, Russia
2 Department of Forest Genetics and Forest Tree Breeding, Georg-August University of Göttingen, Büsgenweg 2, Göttingen, Germany
3 Center for Integrated Breeding Research (CiBreed), Georg-August University of Göttingen, Albrecht-Thaer-Weg 3, Göttingen, Germany
4 Laboratory of Population Genetics, N.I. Vavilov Institute of General Genetics, Russian Academy of Sciences, Gubkin Str. 3, Moscow, Russia
5 Genome Research and Education Center, Laboratory of Forest Genomics, Department of Genomics and Bioinformatics, Institute of Fundamental Biology and Biotechnology, Siberian Federal University, Krasnoyarsk, Russia
6 Scientific and Methodological Center, G.F. Morozov Voronezh State University of Forestry and Technologies, Timiryazeva Str. 8, Voronezh, Russia
* Corresponding Author: Konstantin V. Krutovsky. Email:
(This article belongs to the Special Issue: Application of Nanomaterials in Plants)
Phyton-International Journal of Experimental Botany 2026, 95(5), 4 https://doi.org/10.32604/phyton.2026.078352
Received 29 December 2025; Accepted 07 April 2026; Issue published 27 May 2026
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The article presents the study of the effect of nanocomposites (NCs) based on selenium (Se), copper (Cu), and manganese (Mn) nanoparticles (NPs) embedded in a matrix of natural polysaccharides—arabinogalactan (AG), carrageenan (CAR), and starch (ST)—on the content of chlorophylls (Chls) and carotenoids in potato tissues in vitro. Potatoes were grown for 28 days on Murashige-Skoog (MS) medium with the addition of a NC, then pigments were isolated from leaf tissues, and their content was determined spectrophotometrically. Both a stimulating effect and an inhibitory effect of different NCs on the pigment content were found. Se and Cu NCs in the AG matrices (Se/AG and Cu/AG NCs) increased the Chl content both in the leaf tissues of healthy potato plants and plants infected with the phytopathogenic bacterium Clavibacter sepedonicus (Cms). Mn NCs increased the content of carotenoids and Chls in potatoes infected with Cms. A decrease in the pigment content was detected when growing potatoes on a medium with the addition of Cu NC based on a ST matrix (Cu/ST NC), which is probably due to the high content of NPs in this NC (20%). It was concluded that NCs based on an AG matrix are capable of exerting a stimulating effect on the content of potato pigments, which can be used to increase the yield of this crop.Keywords
In recent years, global climate change and the rapid advancement of agricultural technology have fueled growing interest in new materials and technologies for agricultural production. Crop yield losses can reach 40%, and the long-term use of pesticides does not fully address the problem, causing pollution, reduced soil fertility, the emergence of resistant pathogens, reduced biodiversity, and harm to humans and animals [1]. Consequently, there is an urgent need for innovative and efficient agricultural technologies to address global food production and security challenges. Nanomaterials (NM), which are currently being actively studied, serve as such advanced agents [2,3,4]. They possess solubility, high reactivity, electrical and magnetic properties, and high biological activity. Artificial NMs find application in various technical fields [5,6], medicine, pharmacology [7,8], food industry [9], and cosmetics [10]. In agriculture, the use of such substances is promising in various aspects of veterinary medicine [11], as feed additives for livestock [12], and in the field of agrochemistry as new highly effective pesticides [13,14,15], plant health agents against phytopathogens [16], and agents for increasing plant resistance to stress [17].
It is known that, when nanoparticles (NPs) are applied to plants, they may affect, in addition to the antioxidant system, photosynthetic processes, including the content of photosynthetic pigments [18,19,20]. The composition of photosynthetic pigments in the leaves of higher plants varies widely depending on the species, life form, ontogenetic stages, and environmental conditions of vegetation [21]. Chlorophyll (Chl) and carotenoids are the main photosynthetic pigments that contribute to the efficient absorption of light by plants [22]. The chlorophyll (Chl) family plays a pivotal role in light harvesting by absorbing light across distinct regions of the spectrum, thereby enabling photosynthetic organisms to adapt to diverse environmental conditions under both long-term acclimation and short-term light fluctuations. Carotenoids perform multiple essential functions in photosynthesis, contributing to light absorption and serving as key antioxidants that mitigate photodamage and photoinhibition [23].
In our previous studies, we investigated nanocomposites (NCs), defined as metal or non-metal NPs embedded within natural polymer matrices [24]. To evaluate their potential for enhancing plant health, we examined the biological activity of NCs based on selenium (Se) [25,26], manganese (Mn) [27], and copper (Cu) [28] NPs incorporated via chemical synthesis into natural polymer matrices—carrageenan (CAR), arabinogalactan (AG), and starch (ST). It was demonstrated that Se-based NCs [25] and Cu-based NCs [28] at a concentration of 0.000625%, as well as Mn-based NCs [27] at 0.00625%, significantly reduced the viability of the phytopathogenic bacteria Clavibacter sepedonicus (Cms), the causative agent of potato ring rot, and Pectobacterium carotovorum, which infects a broad spectrum of crops ranging from vegetables to woody plants [29,30].
Furthermore, at antimicrobial concentrations, these NCs stimulated the growth and development of potato plants in vitro, both healthy and Cms-infected [26,27,28]. The effects of NC treatment were evaluated using in vitro biometric parameters and biochemical indicators of the antioxidant defense system, including reactive oxygen species (ROS) levels, antioxidant enzyme activities, and lipid peroxidation intensity [26,27,28].
In addition to these studies, including assessments of NC safety for plant systems, it is essential to investigate their influence on photosynthetic pigment content. Accordingly, the present study examines the effects of NCs and their precursor compounds on the levels of photosynthetic pigments (Chl a, Chl b, and carotenoids) in potato leaf tissues.
2.1 Synthesis of Nanocomposites (NCs)
For NC synthesis, arabinogalactan (AG) was isolated from the polysaccharide fraction of Siberian larch (Larix sibirica Ledeb.) (OOO Wood Chemistry, Irkutsk, Russia). The obtained AG was subsequently purified to remove residual impurities and flavonoids by chromatography on a polyamide column. The NCs were synthesized at room temperature according to the methods described below.
AG (1.0 g) and sodium bis(2-phenylethyl) diselenophosphinate (0.136 g) were dissolved in 50 mL distilled water under magnetic stirring. The solution was incubated at 35–40°C for 3 h, followed by the addition of 30% H2O2. The reaction mixture was precipitated into a fourfold excess of acetone (or ethanol) to isolate the Se(0)-containing NC. The precipitate was washed with the same solvent, filtered, and air-dried. The yield was 97% (calculated relative to Se in the precursor), with a final Se content of 3.4%.
The resulting NC was obtained as a water-soluble orange-red powder. X-ray diffraction (D8 ADVANCE, Bruker Corporation, Billerica, Massachusetts, USA) indicated an X-ray amorphous structure. The formation of amorphous Se(0) NPs was confirmed by a characteristic absorption band at ~310 nm and the absence of crystalline reflections in the diffraction pattern. Transmission electron microscopy (TEM) revealed predominantly spherical NPs with sizes ranging from 20 to 65 nm (average ~25 nm), uniformly distributed within the AG matrix [26].
2.1.2 Manganese-Containing NCs
Mn(OH)2/AG NC was synthesized by dissolving AG (2 g; Mw 18 kDa) in 5 mL distilled water, followed by the addition of MnSO4·5H2O (0.4 g) dissolved in 3 mL water. Subsequently, NH4OH (0.1 mL) and hydrazine (0.2 mL) were added under continuous magnetic stirring. The reaction mixture was stirred for 3 h at room temperature, then precipitated with excess ethanol. The precipitate was washed with ethanol and dried to yield 1.68 g of Mn(OH)2/AG NC.
The NPs are presumed to form as hydrated manganese hydroxide species (Mn(OH)2·nH2O), stabilized within the AG matrix via interactions with oxygen-containing functional groups, primarily hydroxyl moieties. TEM revealed electron-dense, predominantly spherical nanoparticles with diameters of 3–6 nm. The nanoparticles exhibited minimal aggregation and were uniformly dispersed throughout the polymer matrix, maintaining spatial separation within the bulk material. The resulting NCs were found to be paramagnetic particles [27].
To synthesize Mn(OH)2/CAR NC, kappa-carrageenan (3 g, 1100 kDa) were stirred in H2O (150 mL) and heated at 50°C until the mixture become homogeneous. MnSO4·5H2O (0.69 g) in H2O (3 mL) and NH4OH (0.3 mL) were then added. After 24 h, the reaction product was precipitated in alcohol. Thorough washing of the precipitate with alcohol yielded 2.4 g of Mn(OH)2/CAR NC. The resulting NC was a fine brown or light-brown powder, the color intensity of which depended on the mass content of Mn in the sample. The average Mn mass in Mn(OH)2/AG and Mn(OH)2/CAR NCs was 5.2 and 20.3 wt%, respectively [27].
To synthesize Cu2O/AG NC, an aqueous solution of CuCl2·2H2O (0.09 g in 2 mL H2O) was added dropwise to a solution of AG (1 g) in 6 mL distilled water under vigorous stirring. The mixture was heated to 320 K (47°C) and maintained for 30 min. Subsequently, 5 mL of an aqueous solution containing NaBH4 (0.08 g) and NaOH (0.003 g) was added as a reducing system. The reaction was continued for 3 h under vigorous stirring. The mixture was filtered, and the product was isolated from the filtrate by double reprecipitation with ethanol to remove low-molecular-weight impurities, followed by vacuum drying. The Cu content, determined by energy-dispersive X-ray (EDX) microanalysis, was 7.5% [28].
To synthesize Cu2O/ST NC, CuCl2·2H2O (0.8 g in 8 mL H2O) was added to a solution of ST (potato starch, grade Extra, 3 g) in 25 mL water under vigorous stirring. The reaction mixture was maintained at 50°C for 30 min, followed by the addition of 15 mL of an aqueous NaBH4 (0.7 g) and NaOH (0.03 g) solution. After stirring for 3 h, the mixture was filtered. The product was purified by double reprecipitation from ethanol and dried under vacuum. The copper content determined by EDX analysis was 20.3%.
Both NCs were obtained as polymer-stabilized Cu2O NPs dispersed within the respective polysaccharide matrices. The NC solutions, in which the Cu content in the final concentration was 0.000625%, were used for the experiments [28].
Potato plants of Lukyanovsky variety were used for the in vitro experiment. This variety was provided by the A.G. Lorkh Federal Potato Research Centre (Kraskovo Village, Lyubertsy City, Moscow Region, Russia). It is a mid-early table variety resistant to the golden potato cyst nematode and moderately resistant to late blight. This variety was selected because it is considered susceptible to Cms [31], so all physiological reactions are more pronounced in this variety. Micropropagation of test-tube plants was accomplished by cuttings. Control plants were grown on standard Murashige-Skoog (MS) medium. NCs were added as components of the MS medium microsalts, where MnSO4·4H2O was replaced by Mn(OH)2/AG NC, and CuSO4·4H2O was replaced by Cu2O/AG NC. Se NC was added to the MS medium. All NCs were added to the MS medium at a final NP concentration of 0.000625%. The experimental variants are presented in Table 1.
Table 1: Experiments with potato plants in vitro using different types of NPs and NC matrix with and without Cms infection.
| NP | NC Matrix | Without Cms Infection | With Cms Infection |
|---|---|---|---|
| – | – | Control (5) | Cms (5) |
| – | – | SeO2 (3) | – |
| – | – | AG (3) | – |
| Se | AG | Se/AG NC (5) | Se/AG NC + Cms (5) |
| Mn(OH)2 | AG | Mn/AG NC (5) | Mn/AG NC + Cms (5) |
| CAR | Mn/CAR NC (5) | Mn/CAR NC + Cms (5) | |
| Cu2O | AG | Cu/AG NC (5) | Cu/AG NC + Cms (5) |
| ST | Cu/ST NC (5) | Cu/ST NC + Cms (5) | |
| Mn(OH)2 + Cu2O | AG | Mn/AG NC + Cu/AG NC (3) | – |
Two hours after adding the NC, 1 mL of the Cms suspension (titer = 1 × 109 CFU/mL) was added to the potato plant fragments using a sterile syringe, avoiding contact of the bacteria with the plants. The plants were grown under controlled conditions at 26°C and 5–6 Klux illumination for 28 days. Pigment content was then determined.
2.3 Determination of Photosynthetic Pigments
Pigments were extracted with 80% acetone and quantified using a Specord S 100 spectrophotometer (Analytik Jena, Jena, Germany). Chl and carotenoid contents were calculated using the Vernon [32] and Wettstein [33] formulas per unit fresh weight of leaf tissue. A sample of wet leaf tissue was homogenized in 10 mL of a mixture of petroleum ether and acetone, followed by pigment extraction with undiluted acetone. After combining the extracts, the mixture was brought to 25 mL with acetone, and the optical density of the resulting solution was measured at 644, 662 nm, and 440 nm (to determine Chl a, Chl b, and carotenoids, respectively).
Microsoft Excel 2007 and SigmaPlot 12.5 programs were used for analysis and statistical processing of the results. The samples were tested for normality using the Shapiro-Wilk test [34]. The differences in pigment content between potato leaf tissue samples isolated from control plants in vitro, plants subjected to NCs and infected plants were estimated using the analysis of variance (ANOVA).
3.1 Effect of Nanocomposites (NCs) on Photosynthetic Pigment Content in Potato Leaf Tissue
Examples of potato plants infested with Cms and treated with Se/AG NC are shown in Fig. 1. Leaves with pronounced lesions were observed in the Cms-infested plants. In the Se/AG NC treatment alone, the plants had a rich green color. In the combined Se/AG NC + Cms treatment, the color of the plants was less bright green than under the Se/AG NC treatment alone.
Figure 1: Examples of potato plants infested with Cms and treated with Se/AG NC.
It was found that a single treatment of healthy potato plants or those infected with the phytopathogenic Cms with Se oxide at a final Se concentration of 0.00625% resulted in a deviation of the Chl a/b ratio from approximately 1:3 in the control to 1:1.7 and 1:2, respectively, while the total Chl content increased in both cases (Table 2).
This effect is likely due to the development of compensatory metabolic processes under the influence of excessive Se in the plant growth medium, leading to an increase in total Chl content. The increase in the proportion of Chl b in the total Chl (a + b) is presumably due to the oxidation of Chl a caused by ROS generated by plant cells in response to the introduction of Se+4 ions. The introduction of Se/AG NC with a final Se concentration in the nutrient medium of 0.000625% was characterized mainly by maintaining the Chl a/b ratio at a level of 3:1, including in the infected plants. At the same time, the total Chl content in the leaves of infected plants was significantly reduced (by 3–5 times) compared to healthy plants, even after treatment with the NCs. In general, it should be noted that the treatment of plants with Se(0) NPs along is characterized by the absence of any negative impact on the Chl content and the ratio of its types in plant leaves, and the Chl levels were maintained at the level of control intact plants, which again confirms the safety of these NCs for potato plants.
Table 2: Content (mean ± S.D.) of Chl a and b and their ratio in the leaves of healthy and Cms-infected potato plants on the third day of incubation under the influence of Se/AG NC (6.4% Se) and its precursors SeO2 and AG, and in the control.
| Potato Plants | Chl | Control | SeO2 | AG | Se/AG NC |
|---|---|---|---|---|---|
| Healthy | Chl a, mg/g | 1.23 ± 0.09 | 0.72 ± 0.02* | 1.63 ± 0.03* | 0.95 ± 0.10 |
| Chl b, mg/g | 0.70 ± 0.05 | 0.26 ± 0.03* | 0.49 ± 0.04 | 0.29 ± 0.12 | |
| Chl (a + b), mg/g | 1.94 ± 0.02 | 0.97 ± 0.02* | 2.12 ± 0.03* | 1.2 ± 0.08 | |
| Chl a/Chl b | 1.72 | 2.80 | 3.32 | 3.4 | |
| Infected with Cms | Chl a, mg/g | 0.35 ± 0.06 | 0.26 ± 0.03* | 0.30 ± 0.12 | 0.33 ± 0.08 |
| Chl b, mg/g | 0.16 ± 0.03 | 0.09 ± 0.01* | 0.10 ± 0.03* | 0.12 ± 0.01* | |
| Chl (a + b), mg/g | 0.51 ± 0.01 | 0.35 ± 0.04 | 0.40 ± 0.09 | 0.45 ± 0.03 | |
| Chl a/Chl b | 2.24 | 2.88 | 2.9 | 2.90 |
Examples of potato plants infested with Cms and treated with Cu/AG and Cu/ST NCs are shown in Fig. 2. Treatment with Cu/AG NC resulted in an increase in the length and number of leaves on uninfected plants, as previously published [28]. The leaves were a deep green color. In the combined Cu/AG NC + Cms treatment, plants with partially dried leaves were observed, and some plants were light green. When plants were treated with Cu/ST NC, the leaves curled and turned yellow, and some leaves fell off in both uninfected and Cms-infected potatoes.
Figure 2: Examples of potato plants infested with Cms and treated with Cu/AG and Cu/ST NCs.
The results of in vitro potato incubation on nutrient media containing Cu NC are presented in Table 3.
Table 3: Effect of copper-containing NC on pigment concentration (mean ± S.D.) in potato leaf tissue in vitro.
| Experiment | Chl a, mg/g | Chl b, mg/g | Chl a + Chl b, mg/g | Carotenoids, mg/g |
|---|---|---|---|---|
| Control | 0.80 ± 0.30 | 0.55 ± 0.08 | 1.35 ± 0.33 | 1.30 ± 0.24 |
| Cms | 0.53 ± 0.10 | 0.17 ± 0.06* | 0.70 ± 0.16 | 1.22 ± 0.17 |
| Cu/AG NC | 1.00 ± 0.09 | 0.44 ± 0.01 | 1.44 ± 0.19 | 1.31 ± 0.03 |
| Cu/AG NC + Cms | 0.52 ± 0.03 | 0.37 ± 0.03 | 0.88 ± 0.06 | 1.41 ± 0.06 |
| Cu/ST NC | 0.24 ± 0.01** | 0.37 ± 0.02 | 0.61 ± 0.03** | 0.28 ± 0.01** |
| Cu/ST NC + Cms | 0.19 ± 0.03** | 0.26 ± 0.01 | 0.45 ± 0.01** | 0.20 ± 0.01** |
Infection significantly reduced Chl b content. Cu/AG NC had no effect on potato pigment content. Cu/ST NC significantly reduced Chl a and carotenoids in both infected and uninfected plants. These data indicate that Cu/ST NC has a negative effect on the amount of photosynthetic pigments in potatoes. Therefore, Cu/AG NC was used in further experiments to study the combined effect of NCs on pigment content.
3.1.3 Manganese-Containing NCs
Examples of potato plants infested with Cms and treated with Mn/AG and Mn/CAR NCs are shown in Fig. 3. As in the previous series of experiments, potato plants infected with Cms had a light green color. Mn-containing NCs had no visual negative impact on uninfected plants, and even when potatoes were infected with Cms, they maintained the bright green color of the aboveground parts.
Figure 3: Examples of potato plants infested with Cms and treated with Mn/AG and Mn/CAR NCs.
The effect of Mn NCs in vitro on the pigment content in leaf tissues of both healthy potato plants and those infected with Cms is presented in Table 4. Infection reduced the content of all potato pigments. Mn/AG NC slightly reduced Chl b content in healthy plants. However, upon infection, this NC increased Chl a and carotenoids content. Mn/CAR NC reduced Chl b content. Therefore, Mn/AG NC was used in further experiments to study the combined effect of NCs on pigment content, as it increased pigment content under biotic stress.
Table 4: Effect of manganese-containing NC on pigment concentration (mean ± S.D.) in potato leaf tissue in vitro.
| Experiment | Chl a, mg/g | Chl b, mg/g | Chl a + Chl b, mg/g | Carotenoids, mg/g |
|---|---|---|---|---|
| Control | 1.28 ± 0.02 | 0.95 ± 0.02 | 2.23 ± 0.04 | 1.79 ± 0.05 |
| Cms | 0.40 ± 0.02** | 0.29 ± 0.03** | 0.69 ± 0.05** | 1.04 ± 0.01** |
| Mn/AG NC | 0.94 ± 0.12 | 0.78 ± 0.04* | 1.72 ± 0.15* | 1.71 ± 0.10 |
| Mn/AG NC + Cms | 1.69 ± 0.01** | 0.87 ± 0.01 | 2.56 ± 0.03** | 2.00 ± 0.01** |
| Mn/CAR NC | 0.96 ± 0.20 | 0.62 ± 0.10* | 1.58 ± 0.30 | 1.58 ± 0.11 |
| Mn/CAR NC + Cms | 1.54 ± 0.18 | 0.80 ± 0.06 | 2.35 ± 0.24 | 1.97 ± 0.11 |
Considering that Mn, like Cu, is an essential element for the successful functioning of the plant photosynthetic apparatus, it was interesting to test the combined effect of Mn- and Cu-containing NCs on the concentration of photosynthetic pigments in potato leaf tissue in vitro. Cu/AG NC was chosen as the Cu-containing NC in the combined treatment, because it was found that it did not have a negative effect on pigment content, unlike Cu/ST NC (Table 3). The obtained results are presented in Fig. 4. The results showed that Mn-containing NCs increased the carotenoids content in potato tissue. This effect was negated by combined treatment of plants by Mn-containing NCs together with Cu-containing NCs.
Figure 4: Effect of copper- and manganese-containing NC on the pigment content (mean ± S.D.) in potato leaf tissues in vitro; *p ≤ 0.01 compared to control.
Plants and photosynthetic bacteria convert solar radiation into chemical energy through photosynthesis, thereby sustaining primary productivity and supporting nearly all life on Earth. Photosynthetic pigments, principally Chls and carotenoids, are integral components of the photosynthetic apparatus, capturing light energy to drive electron transport reactions [35]. Chl functions as the primary light-harvesting pigment, whereas carotenoids contribute both to light capture and to photoprotection by dissipating excess energy. Together, these pigments perform complementary roles that maintain photosynthetic efficiency and protect plant cells from oxidative damage [36].
Chl is the central pigment of the photosynthetic machinery, responsible for photon absorption and energy transfer to the reaction centers of photosystems [37]. Upon light absorption, Chl molecules undergo electronic excitation and transfer energy to the photosystems, thereby initiating photochemical reactions. In higher plants, Chl metabolism is tightly regulated through coordinated enzymatic steps and transcriptional control. Biosynthesis begins with glutamate, which is converted to 5-aminolevulinic acid and subsequently to protochlorophyllide (Pchlide) via a multistep pathway involving enzymes such as glutamyl-tRNA reductase and magnesium chelatase (subunit H) [35,38]. Precise regulation of Chl and its intermediates is critical, as Chl metabolism influences not only photosystem assembly but also processes including programmed cell death, the “stay-green” phenotype, and chloroplast–nucleus signaling [39]. Excessive accumulation of Pchlide, a potent photosensitizer, can promote reactive oxygen species (ROS) generation, leading to growth inhibition or accelerated cell death [40]. Chl degradation is induced during stress and senescence, and substantial progress has been made in elucidating its regulatory mechanisms [41,42]. Structurally, Chl a contains a methyl group that contributes to its photochemical properties and renders it more susceptible to photodegradation relative to other pigments, while Chl b broadens the absorption spectrum and enhances light-harvesting efficiency, thereby supporting biomass accumulation [43].
Carotenoids are isoprenoid pigments synthesized de novo in all photosynthetic organisms. In plants, they confer yellow, orange, and red coloration to leaves, flowers, fruits, and certain storage organs [44]. Functionally, carotenoids participate in photosynthesis, photoprotection, pigmentation, phytohormone biosynthesis, and intracellular signaling [45]. Synthesized in plastids, they are integral to light-harvesting complexes and play a critical role in protecting the photosynthetic apparatus from oxidative stress. In addition, carotenoids serve as precursors of apocarotenoids, including vitamin A derivatives and the phytohormones abscisic acid and strigolactones, which regulate plant development and stress responses [46,47].
Carotenoid biosynthesis originates from geranylgeranyl diphosphate and proceeds through a series of enzymatically regulated reactions leading to the formation of major carotenoids such as lutein, zeaxanthin, and β-carotene. Key enzymes include phytoene synthase, phytoene desaturase, and lycopene β-cyclase. This pathway is tightly controlled at the transcriptional level and responds to environmental and light signals, with red and blue wavelengths particularly effective in promoting carotenoid accumulation [48].
Carotenoids are catabolized by carotenoid cleavage dioxygenases, generating apocarotenoids such as abscisic acid and strigolactones [49]. These metabolites function as signaling molecules that regulate plant growth, development, and adaptive stress responses. The dynamic balance between carotenoid biosynthesis and degradation ensures adequate light harvesting and photoprotection while preventing excessive accumulation that could disrupt cellular homeostasis.
Beyond their role in pigmentation, carotenoids are essential for protecting the photosynthetic apparatus from photooxidative damage. They associate with Chls in photosystem I (PSI) and photosystem II (PSII) complexes and are embedded within the thylakoid membrane lipid bilayer, where they suppress uncontrolled energy transfer to molecular oxygen and limit ROS formation. Moreover, carotenoids function as accessory pigments by absorbing wavelengths outside the principal absorption range of Chls and transferring this energy to the reaction centers, thereby enhancing overall photosynthetic performance [36].
Chl a, the main photosynthetic pigment in plants, contains a methyl (-CH3) group, whose key function is photon binding, and is more sensitive to photodegradation than other pigments [50]. Chl b plays an important role in improving light absorption efficiency and, thus, increasing energy and biomass production in plants [51]. In our experiments, Se NCs and Cu NCs increased Chl content. A similar effect was previously described in a study of strawberries (Fragaria ananassa) under greenhouse experimental conditions, where it was shown that the application of nanoselenium (at concentrations of 25, 50, 75, and 100 mg L−1) and nanocopper (at concentrations of 50 and 100 mg L−1) increased the Chl content and its fluorescence intensity [52]. Moreover, the increase in Chl a content under the treatments was more pronounced than that of Chl b. In addition, Se nanofertilizer increased Chl levels more than Cu NP-based nanofertilizer. The highest Chl values were recorded after the application of 100 mg L−1 Se nanofertilizer. CuO nanofertilizer had an obvious effect on the minimum and maximum fluorescence parameters (F0 and Fm), with significantly higher values (513 and 1713) compared to the control and Se nanofertilizer [52].
Spraying the leaves of Moldavian dragonhead (Dracocephalum moldavica L.) with CuO NPs resulted in an increase in the photosynthetic pigments Chl a and Chl b by 77% and 123%, respectively [20]. When growing rice (Oryza sativa L.) under hydroponics, it was shown that the content of photosynthetic pigments initially increased and then decreased with increasing CuO NP concentrations (0, 10, 20, and 50 mg L−1) [53]. Foliar treatment with 5 mg L−1 bimetallic NPs consisting of the micronutrients Cu and Fe increased the energy and efficiency of photosynthetic activity in rice O. sativa plants [54]. CuO and Fe2O3 NPs stimulated the growth of pomegranate Punica granatum L. variety Hegazy and provided resistance to the root-knot nematode Meloidogyne javanica [55]. An increase in the level of photosynthetic pigments, both chlorophylls and carotenoids, was observed. The authors believe that the effect of NPs partly depends on the increase in the production of photosynthetic pigments [55]. The Chl a content increased in sunflower seedlings after seed soaking in a Cu NP solution compared to the control, while the Chl b content decreased by 3–7% [56]. Bimetallic Cu-Se NPs synthesized using Aspergillus niger increased the resistance of eggplant Solanum melongen to the bacterial pathogen Ralstonia solanacearum and positively affected photosynthetic pigments, increasing the levels of chlorophyll and carotenoids [57]. In an experiment on wheat (Triticum aestivum L.) under greenhouse conditions, it was shown that photosynthetic pigments and gas exchange parameters were significantly increased when treated with Se NPs obtained by green synthesis, compared to the control [58].
Exogenous application of nanoselenium with glutathione (SeG) significantly enhanced the growth of cucumber Cucumis sativus plants and increased the content of photosynthetic pigments [59]. Se NPs (10 mg L−1), chitosan (CS) (0.1%), and CS-Se NPs (at two concentrations of 5 and 10 mg L−1) were shown to increase the content of photosynthetic pigments in leaf tissues of Moldovan lemon balm (Dracocephalum moldavica L.) [60]. Spraying the leaves of red bean (Phaseolus vulgaris L.) plants with Se NPs coated with gum arabic resulted in a significant increase in the content of photosynthetic pigments and the area of photosynthetic plant organs (cm2 plant−1) [61]. Under field conditions, spraying sweet wormwood (Artemisia annua L.) leaves with Se NPs resulted in a significant increase in photosynthetic pigments [62]. In common bean (Phaseolus vulgaris), CuO NPs were shown to improve plant vegetative growth and, in particular, increase the content of photosynthetic pigments: Chl a (2.96 mg/g), Chl b (1.93 mg/g), and total carotenoids (1.16 mg/g) [63]. Spraying tomatoes (Solánum lycopérsicum) with Se NPs promoted the accumulation of photosynthetic pigments [64]. Spraying lemon verbena (Aloysia citrodora) with Se NPs increased the content of photosynthetic pigments [65]. Se has been shown to have a beneficial effect on increasing photosynthetic pigments in Cofféa arábica leaf tissue [66]. In sesame (Sesamum indicum L.), Se NPs have been shown to increase Chl a and Chl b content [67]. Eco-friendly Se NPs synthesized via Sternbergia candida reduced salt stress in pepper (Capsicum annuum L.) plants by increasing the content of chlorophyll and carotenoids [68]. A similar stimulating effect of Se NPs on the amount of photosynthetic pigments was shown in wheat (T. aestivum) under salt stress in a field experiment [69]. Se NPs increased the content of photosynthetic pigments in the medicinal plant Echium italicum L. [70]. Spraying Se NPs on the leaves of broccoli (Brassica oleracea var. italic) mitigated the effects of drought by improving pigment stability and increasing carotenoid content [71]. In our experiments, Mn-containing NPs increased carotenoid content. In field experiments, the addition of CuO NPs to the soil significantly affected photosynthetic pigments (Chl a, Chl b, and carotenoids) in soybean (Glycine max L.). Carotenoid content was 2.5 times higher with nCuO-S treatment [43].
In our experiments, a decrease in pigments was also recorded under the influence of NPs. This effect was demonstrated by Cu/ST NC, as well as by combined Cu- and Mn-containing NCs. The high NP content of 20% in the NC apparently had a negative impact on plant cells, causing oxidative stress. Under the combined effect of the two NCs, Chl b was more sensitive than Chl a and carotenoids. Effect of studied NCs on potato photosynthetic pigments in vitro is briefly summarized in Table 5.
Table 5: Summary of the effect of NC on potato photosynthetic pigments in vitro.
| Effect | Se NC | Cu NC | Mn NC | Cu/AG NC + Mn NC |
|---|---|---|---|---|
| Pigment content | Se/AG NC did not affect chlorophyll content in both healthy and infected plants. |
| Mn/AG NC causes an increase in the amount of chlorophylls and carotenoids during infection. | Decreased Chl b content |
In the study presented here, we did not separately examine the possible effect of free Se, Cu, and Mn ions, which could hypothetically be formed during treatment with Se-, Cu-, and Mn-containing NCs, since the study presented here was preceded by a series of studies devoted to the biological effect of these metal-containing NCs (MCNCs) on potatoes in vitro [26,27,28]. We have previously published the results of several studies of the effect of these studied MCNCs on the biometric and biochemical characteristics of potato plants in vitro [26,27,28]. In particular, at the initial stage of the study, we compared the effect of Se NC with the effect of Se in ionic form. For example, as one of the controls for the study of the effect of Se NCs on the biometric and biochemical characteristics of potato plants in vitro, the effect of SeO2 along was compared with the effect of Se NC [72]. It was found that SeO2 had a negative effect on plant viability compared to the Se NC treatment. In another study, we also investigated the effect of sodium bis(2-phenylethyl) diselenophosphinate (BIS), a precursor of Se NC, on potato viability in vitro [26]. It was also found that when Se-containing NC and BIS were used at equal concentrations for plant treatment, BIS suppressed plant growth and development, while Se NC had a positive effect on all studied parameters.
Furthermore, in one of our earlier studies [73], we studied the content of free elemental Se from Se NC in potato tissues in vitro using energy dispersive X-ray microanalysis (EDXMA). It was found that the Se content in potato leaf tissues was 0.01–0.03% of air-dry weight, indicating insignificant Se penetration into potato tissues from Se NC and its insignificant accumulation in the plant body after Se NC treatment [73].
Similar studies of the effect of free Cu and Mn ions on potatoes in vitro were also carried out in early studies of Cu and Mn NCs and showed a positive and stronger effect of these metals in the composition of NCs, and not in the form of free ions [27,28]. Therefore, we have every reason to believe that it is the studied NCs that: (1) reduce oxidative stress in plant tissues, which was reflected in a decrease in the amount of reactive oxygen species, modulation of the activity of antioxidant enzymes (catalase, peroxidase), a decrease in lipid peroxidation products—diene conjugates and malondialdehyde [26,27,28,72]; (2) activate proteins involved in pathogenesis, which is also confirmed by new data on an increase in the level of PR gene expression in potato tissues in vitro when they are treated with Se-, Cu-, and Mn-containing NCs, obtained by us recently and not yet published; (3) have an antibacterial effect on phytopathogenic bacteria, which we have previously shown in [74]; (4) have an effect on the hormonal status of plants (we are currently conducting research in this direction).
The results presented in this article demonstrate the absence of a negative effect of Se, Cu, and Mn NCs based on AG matrix on the content of photosynthetic pigments in potato leaf tissues in vitro. Se/AG and Cu/AG NCs increased the Chl content in both healthy plants and those infected with the pathogen Cms. Mn/AG NC increased the carotenoid content. Cu/ST NC decreased the amount of pigments in healthy plants and those infected with Cms. The combined effect of Mn/AG and Cu/ST NCs led to a decrease in Chl b. Thus, it can be concluded that NCs based on AG do not have a negative effect on the pigment system of plants, which is in agreement with our other studies, they can be considered as safe agents for plant enhancement and protection from phytopathogenic bacteria.
Acknowledgement:
Funding Statement: This study was supported by the federal program № 0277-2021-0004 (121031300011-7) of the Ministry of Science and Higher Education of the Russian Federation with the basic project “Study of molecular mechanisms of physiological processes and allelopathy in plant-microbe relationships”.
Author Contributions: Alla I. Perfileva: conceptualization, investigation, project administration, visualization, writing—original draft, writing—review & editing, funding acquisition. Konstantin V. Krutovsky: investigation, supervision, visualization, writing—review & editing. All authors reviewed and approved the final version of the manuscript.
Availability of Data and Materials: The data that support the findings of this study are available from the first author (A.I.P.) upon reasonable request.
Ethics Approval: Not applicable.
Conflicts of Interest: The authors declare no conflicts of interest.
References
1. Zhang H , Zheng T , Wang Y , Li T , Chi Q . Multifaceted impacts of nanoparticles on plant nutrient absorption and soil microbial communities. Front Plant Sci. 2024; 15: 1497006. doi:10.3389/fpls.2024.1497006. [Google Scholar] [CrossRef]
2. Kumari A , Gupta AK , Sharma S , Jadon VS , Sharma V , Chun SC , et al. Nanoparticles as a tool for alleviating plant stress: mechanisms, implications, and challenges. Plants. 2024; 13( 11): 1528. doi:10.3390/plants13111528. [Google Scholar] [CrossRef]
3. de Paiva Pinheiro SK , da Silva Pontes M , Miguel TBAR , Grillo R , de Souza Filho AG , de Castro Miguel E . Nanoparticles and plants: a focus on analytical characterization techniques. Plant Sci. 2024; 348: 112225. doi:10.1016/j.plantsci.2024.112225. [Google Scholar] [CrossRef]
4. Acharya C , Mishra S , Chaurasia SK , Pandey BK , Dhar R , Pandey JK . Synthesis of metallic nanoparticles using biometabolites: mechanisms and applications. Biometals. 2025; 38( 1): 21– 54. doi:10.1007/s10534-024-00642-w. [Google Scholar] [CrossRef]
5. Hughes KJ , Ganesan M , Tenchov R , Iyer KA , Ralhan K , Diaz LL , et al. Nanoscience in action: unveiling emerging trends in materials and applications. ACS Omega. 2025; 10( 8): 7530– 48. doi:10.1021/acsomega.4c10929. [Google Scholar] [CrossRef]
6. Sohail A . Progress in nanomaterials: synthesis, characterization, and applications. Next Nanotechnol. 2025; 8: 100263. doi:10.1016/j.nxnano.2025.100263. [Google Scholar] [CrossRef]
7. Ma X , Tian Y , Yang R , Wang H , Allahou LW , Chang J , et al. Nanotechnology in healthcare, and its safety and environmental risks. J Nanobiotechnol. 2024; 22( 1): 715. doi:10.1186/s12951-024-02901-x. [Google Scholar] [CrossRef]
8. Soni AK , Jha RK . Nanotechnology’s applications and potential in various fields. Cureus. 2024; 16( 4): e59234. doi:10.7759/cureus.59234. [Google Scholar] [CrossRef]
9. Zhou XQ , Hayat Z , Zhang DD , Li MY , Hu S , Wu Q , et al. Zinc oxide nanoparticles: synthesis, characterization, modification, and applications in food and agriculture. Processes. 2023; 11( 4): 1193. doi:10.3390/pr11041193. [Google Scholar] [CrossRef]
10. Gupta V , Mohapatra S , Mishra H , Farooq U , Kumar K , Ansari MJ , et al. Nanotechnology in cosmetics and cosmeceuticals—a review of latest advancements. Gels. 2022; 8( 3): 173. doi:10.3390/gels8030173. [Google Scholar] [CrossRef]
11. Kazemi M . Revolutionizing veterinary medicine: the role of nanoparticles in advancing animal health, nutrition and disease management. Vet Med Sci. 2025; 11( 5): e70528. doi:10.1002/vms3.70528. [Google Scholar] [CrossRef]
12. Kholif AE , Anele A , Chahine M , Anele UY . Applications of nanotechnology in ruminant animal production: advances, challenges, and future prospects. Nanomaterials. 2025; 15( 23): 1773. doi:10.3390/nano15231773. [Google Scholar] [CrossRef]
13. Jain S , Mathur J . Nanoformulations for plant growth promotion. In: Radhakrishnan EK , Aswani R , Visakh PM , editors. Nanoformulations for agricultural applications. Beverly, MA, USA: Scrivener Publishing; 2025. p. 119– 52. [Google Scholar]
14. Bouhadi M , Javed Q , Jakubus M , Elkouali M , Fougrach H , Ansar A , et al. Nanoparticles for sustainable agriculture: assessment of benefits and risks. Agronomy. 2025; 15( 5): 1131. doi:10.3390/agronomy15051131. [Google Scholar] [CrossRef]
15. Qin Q , Wang Q , Adeel M , Li Z , Liu X , Zhang T , et al. Multidimensional regulation mechanism and research progress of nanofertilizers on vegetable quality. J Agric Food Chem. 2026; 74( 1): 127– 44. doi:10.1021/acs.jafc.5c11405. [Google Scholar] [CrossRef]
16. Wassie KB , Yemata G . Metal oxide nanoparticles as promising agents for triggering defense mechanisms in plants against bacterial diseases. Discov Nano. 2025; 20( 1): 127. doi:10.1186/s11671-025-04288-6. [Google Scholar] [CrossRef]
17. Cao Y , Turk K , Bibi N , Ghafoor A , Ahmed N , Azmat M , et al. Nanoparticles as catalysts of agricultural revolution: enhancing crop tolerance to abiotic stress: a review. Front Plant Sci. 2025; 15: 1510482. doi:10.3389/fpls.2024.1510482. [Google Scholar] [CrossRef]
18. Haisel D , Cyrusová T , Vaněk T , Podlipná R . The effect of nanoparticles on the photosynthetic pigments in cadmium-zinc interactions. Environ Sci Pollut Res Int. 2019; 26( 4): 4147– 51. doi:10.1007/s11356-018-04060-7. [Google Scholar] [CrossRef]
19. Hashemi S , Asrar Z , Pourseyedi S , Nadernejad N . Investigation of ZnO nanoparticles on proline, anthocyanin contents and photosynthetic pigments and lipid peroxidation in the soybean. IET Nanobiotechnol. 2019; 13( 1): 66– 70. doi:10.1049/iet-nbt.2018.5212. [Google Scholar] [CrossRef]
20. Nekoukhou M , Fallah S , Pokhrel LR , Abbasi-Surki A , Rostamnejadi A . Foliar enrichment of copper oxide nanoparticles promotes biomass, photosynthetic pigments, and commercially valuable secondary metabolites and essential oils in dragonhead (Dracocephalum moldavica L.) under semi-arid conditions. Sci Total Environ. 2023; 863: 160920. doi:10.1016/j.scitotenv.2022.160920. [Google Scholar] [CrossRef]
21. Fernández-Marín B , Gulías J , Figueroa CM , Iñiguez C , Clemente-Moreno MJ , Nunes-Nesi A , et al. How do vascular plants perform photosynthesis in extreme environments? An integrative ecophysiological and biochemical story. Plant J. 2020; 101( 4): 979– 1000. doi:10.1111/tpj.14694. [Google Scholar] [CrossRef]
22. Lokstein H , Renger G , Götze JP . Photosynthetic light-harvesting (antenna) complexes—structures and functions. Molecules. 2021; 26( 11): 3378. doi:10.3390/molecules26113378. [Google Scholar] [CrossRef]
23. Simkin AJ , Kapoor L , Doss CGP , Hofmann TA , Lawson T , Ramamoorthy S . The role of photosynthesis related pigments in light harvesting, photoprotection and enhancement of photosynthetic yield in planta. Photosynth Res. 2022; 152( 1): 23– 42. doi:10.1007/s11120-021-00892-6. [Google Scholar] [CrossRef]
24. Trofimov BA , Sukhov BG , Aleksandrova GP , Medvedeva SA , Grishchenko LA , Mal’kina AG , et al. Arabinogalactan-based nanocomposites with magnetic, optical, catalytic and biologically active properties. Dokl Akad Nauk. 2003; 393( 5): 634– 5. [Google Scholar]
25. Perfileva AI , Nozhkina OA , Graskova IA , Sidorov AV , Lesnichaya MV , Aleksandrova GP , et al. Synthesis of selenium and silver nanobiocomposites and their influence on phytopathogenic bacterium Clavibacter michiganensis subsp. sepedonicus. Russ Chem Bull. 2018; 67( 1): 157– 63. doi:10.1007/s11172-018-2052-4. [Google Scholar] [CrossRef]
26. Perfileva AI , Nozhkina OA , Ganenko TV , Graskova IA , Sukhov BG , Artem’ev AV , et al. Selenium nanocomposites in natural matrices as potato recovery agent. Int J Mol Sci. 2021; 22( 9): 4576. doi:10.3390/ijms22094576. [Google Scholar] [CrossRef]
27. Khutsishvili SS , Perfileva AI , Nozhkina OA , Ganenko TV , Krutovsky KV . Novel nanobiocomposites based on natural polysaccharides as universal trophic low-dose micronutrients. Int J Mol Sci. 2021; 22( 21): 12006. doi:10.3390/ijms222112006. [Google Scholar] [CrossRef]
28. Khutsishvili SS , Perfileva AI , Kon’kova TV , Lobanova NA , Sadykov EK , Sukhov BG . Copper-containing bionanocomposites based on natural raw arabinogalactan as effective vegetation stimulators and agents against phytopathogens. Polymers. 2024; 16( 5): 716. doi:10.3390/polym16050716. [Google Scholar] [CrossRef]
29. Perfileva AI , Kharasova AR , Nozhkina OA , Sidorov AV , Graskova IA , Krutovsky KV . Effect of nanopriming with selenium nanocomposites on potato productivity in a field experiment, soybean germination and viability of Pectobacterium carotovorum. Horticulturae. 2023; 9( 4): 458. doi:10.3390/horticulturae9040458. [Google Scholar] [CrossRef]
30. Perfileva AI , Krutovsky KV . Effect of manganese- and selenium-containing nanocomposites on soybean resistance to Pectobacterium carotovorum and microbial landscape of soybean seedlings. Plant Growth Regul. 2024; 104( 1): 389– 402. doi:10.1007/s10725-024-01175-5. [Google Scholar] [CrossRef]
31. Romanenko AS , Riffel AA , Graskova IA , Rachenko MA . The role of extracellular pH-homeostasis in potato resistance to ring rot pathogen. J Phytopathol. 1999; 147( 11–12): 679– 86. doi:10.1046/j.1439-0434.1999.00450.x. [Google Scholar] [CrossRef]
32. Vernon LP . Spectrophotometric determination of chlorophylls and pheophytins in plant extracts. Anal Chem. 1960; 32( 9): 1144– 50. doi:10.1021/ac60165a029. [Google Scholar] [CrossRef]
33. von Wettstein D . Chlorophyll-letale und der submikroskopische formwechsel der plastiden. Exp Cell Res. 1957; 12( 3): 427– 506. doi:10.1016/0014-4827(57)90165-9. [Google Scholar] [CrossRef]
34. Shapiro SS , Wilk MB . An analysis of variance test for normality (complete samples). Biometrika. 1965; 52( 3–4): 591– 611. doi:10.1093/biomet/52.3-4.591. [Google Scholar] [CrossRef]
35. Li Y , Cao T , Guo Y , Grimm B , Li X , Duanmu D , et al. Regulatory and retrograde signaling networks in the chlorophyll biosynthetic pathway. J Integr Plant Biol. 2025; 67( 4): 887– 911. doi:10.1111/jipb.13837. [Google Scholar] [CrossRef]
36. Nguyen TNP , Sung J . Light spectral-ranged specific metabolisms of plant pigments. Metabolites. 2024; 15( 1): 1. doi:10.3390/metabo15010001. [Google Scholar] [CrossRef]
37. Li X , Zhang W , Niu D , Liu X . Effects of abiotic stress on chlorophyll metabolism. Plant Sci. 2024; 342: 112030. doi:10.1016/j.plantsci.2024.112030. [Google Scholar] [CrossRef]
38. Tripathy BC , Pattanayak GK . Chlorophyll biosynthesis in higher plants. In: Eaton-Rye JJ , Tripathy BC , Sharkey TD , editors. Photosynthesis: plastid biology, energy conversion and carbon assimilation. Dordrecht, The Netherlands: Springer; 2012. p. 63– 94. [Google Scholar]
39. Tanaka A , Tanaka R . Chlorophyll metabolism. Curr Opin Plant Biol. 2006; 9( 3): 248– 55. doi:10.1016/j.pbi.2006.03.011. [Google Scholar] [CrossRef]
40. Reinbothe C , El Bakkouri M , Buhr F , Muraki N , Nomata J , Kurisu G , et al. Chlorophyll biosynthesis: spotlight on protochlorophyllide reduction. Trends Plant Sci. 2010; 15( 11): 614– 24. doi:10.1016/j.tplants.2010.07.002. [Google Scholar] [CrossRef]
41. Zhu X , Chen J , Qiu K , Kuai B . Phytohormone and light regulation of chlorophyll degradation. Front Plant Sci. 2017; 8: 1911. doi:10.3389/fpls.2017.01911. [Google Scholar] [CrossRef]
42. Tanaka A , Ito H . Chlorophyll degradation and its physiological function. Plant Cell Physiol. 2025; 66( 2): 139– 52. doi:10.1093/pcp/pcae093. [Google Scholar] [CrossRef]
43. Yusefi-Tanha E , Fallah S , Pokhrel LR , Rostamnejadi A . Role of particle size-dependent copper bioaccumulation-mediated oxidative stress on Glycine max (L.) yield parameters with soil-applied copper oxide nanoparticles. Environ Sci Pollut Res Int. 2024; 31( 20): 28905– 21. doi:10.1007/s11356-024-33070-x. [Google Scholar] [CrossRef]
44. Quian-Ulloa R , Stange C . Carotenoid biosynthesis and plastid development in plants: the role of light. Int J Mol Sci. 2021; 22( 3): 1184. doi:10.3390/ijms22031184. [Google Scholar] [CrossRef]
45. Sun T , Rao S , Zhou X , Li L . Plant carotenoids: recent advances and future perspectives. Mol Hortic. 2022; 2( 1): 3. doi:10.1186/s43897-022-00023-2. [Google Scholar] [CrossRef]
46. Stra A , Almarwaey LO , Alagoz Y , Moreno JC , Al-Babili S . Carotenoid metabolism: new insights and synthetic approaches. Front Plant Sci. 2023; 13: 1072061. doi:10.3389/fpls.2022.1072061. [Google Scholar] [CrossRef]
47. Demmig-Adams B , Hodges AK , Polutchko SK , Adams WW III . Zeaxanthin and other carotenoids: roles in abiotic stress defense with implications for biotic defense. Plants. 2025; 14( 17): 2703. doi:10.3390/plants14172703. [Google Scholar] [CrossRef]
48. Stanley L , Yuan YW . Transcriptional regulation of carotenoid biosynthesis in plants: so many regulators, so little consensus. Front Plant Sci. 2019; 10: 1017. doi:10.3389/fpls.2019.01017. [Google Scholar] [CrossRef]
49. Nisar N , Li L , Lu S , Khin NC , Pogson BJ . Carotenoid metabolism in plants. Mol Plant. 2015; 8( 1): 68– 82. doi:10.1016/j.molp.2014.12.007. [Google Scholar] [CrossRef]
50. Rico CM , Peralta-Videa JR , Gardea-Torresdey JL . Chemistry, biochemistry of nanoparticles, and their role in antioxidant defense system in plants. In: Nanotechnology and plant sciences. Cham, Switzerland: Springer International Publishing; 2015. p. 1– 17. [Google Scholar]
51. Xiao Y , Ma J , Xian J , Peijnenburg WJGM , Du Y , Tian D , et al. Copper accumulation and physiological markers of soybean (Glycine max) grown in agricultural soil amended with copper nanoparticles. Ecotoxicol Environ Saf. 2022; 229: 113088. doi:10.1016/j.ecoenv.2021.113088. [Google Scholar] [CrossRef]
52. El-Bialy SM , El-Mahrouk ME , Elesawy T , Omara AE , Elbehiry F , El-Ramady H , et al. Biological nanofertilizers to enhance growth potential of strawberry seedlings by boosting photosynthetic pigments, plant enzymatic antioxidants, and nutritional status. Plants. 2023; 12( 2): 302. doi:10.3390/plants12020302. [Google Scholar] [CrossRef]
53. Wang SQ , Sun YB , Huang QQ , Xu YM , Dong RY , Meng QY . Effects of nano-copper oxide on physiobiochemical properties of Brassica chinensis L. and its heavy metal accumulation under cadmium stress. Huan Jing Ke Xue Huanjing Kexue. 2023; 44( 9): 5204– 13. [Google Scholar]
54. Dey S , Nath S , Alam Ansari T , Biswas A , Barman F , Mukherjee S , et al. Application of green synthesized bimetallic nZVI-Cu nanoparticle as a sustainable alternative to chemical fertilizers to enhance growth and photosynthetic efficiency of rice seedlings. Plant Physiol Biochem. 2023; 201: 107837. doi:10.1016/j.plaphy.2023.107837. [Google Scholar] [CrossRef]
55. Abdel-Wahab DA , El-Zawahry AMI , Hamada AM , Abdel-Salam MM , Samy AM . Nanoparticles boost pomegranate growth and defense, suppressing root-knot nematodes. Front Plant Sci. 2025; 16: 1560126. doi:10.3389/fpls.2025.1560126. [Google Scholar] [CrossRef]
56. Kachel-Górecka M , Sokal K , Stryjecka M . Effect of nanometals and pulsed electric field (PEF) on the germination capacity of seeds and antioxidative properties of seedlings of sunflower. Plants. 2025; 14( 16): 2512. doi:10.3390/plants14162512. [Google Scholar] [CrossRef]
57. Attia MS , Hasanin MS , Hashem AH , Abdel-Kader SA , Aloufi AS , Baazeem A , et al. Aspergillus niger mediated biosynthesis of bimetallic copper-selenium nanoparticles and their roles in enhancing resistance to Ralstonia solanacearum. Biotechnol J. 2025; 20( 9): e70123. doi:10.1002/biot.70123. [Google Scholar] [CrossRef]
58. El-Saadony MT , Saad AM , Najjar AA , Alzahrani SO , Alkhatib FM , Shafi ME , et al. The use of biological selenium nanoparticles to suppress Triticum aestivum L. crown and root rot diseases induced by Fusarium species and improve yield under drought and heat stress. Saudi J Biol Sci. 2021; 28( 8): 4461– 71. doi:10.1016/j.sjbs.2021.04.043. [Google Scholar] [CrossRef]
59. Jia Y , Chen L , Kang L , Fu X , Zheng S , Wu Y , et al. Nano-selenium and glutathione enhance cucumber resistance to Botrytis cinerea by promoting jasmonic acid-mediated cucurbitacin biosynthesis. ACS Nano. 2024; 18( 31): 20576– 90. doi:10.1021/acsnano.4c05827. [Google Scholar] [CrossRef]
60. Azimi F , Oraei M , Gohari G , Panahirad S , Farmarzi A . Chitosan-selenium nanoparticles (Cs-Se NPs) modulate the photosynthesis parameters, antioxidant enzymes activities and essential oils in Dracocephalum moldavica L. under cadmium toxicity stress. Plant Physiol Biochem. 2021; 167: 257– 68. doi:10.1016/j.plaphy.2021.08.013. [Google Scholar] [CrossRef]
61. Abouelhamd N , El Lateef Gharib FA , Amin AA , Ahmed EZ . Impact of foliar spray with Se, nano-Se and sodium sulfate on growth, yield and metabolic activities of red kidney bean. Sci Rep. 2023; 13( 1): 17102. doi:10.1038/s41598-023-43677-8. [Google Scholar] [CrossRef]
62. Logvinenko L , Golubkina N , Fedotova I , Bogachuk M , Fedotov M , Kataev V , et al. Effect of foliar sodium selenate and nano selenium supply on biochemical characteristics, essential oil accumulation and mineral composition of Artemisia annua L. Molecules. 2022; 27( 23): 8246. doi:10.3390/molecules27238246. [Google Scholar] [CrossRef]
63. El-Sayed ER , Mohamed SS , Mousa SA , El-Seoud MAA , Elmehlawy AA , Abdou DAM . Bifunctional role of some biogenic nanoparticles in controlling wilt disease and promoting growth of common bean. AMB Express. 2023; 13( 1): 41. doi:10.1186/s13568-023-01546-7. [Google Scholar] [CrossRef]
64. Liu Y , Liu R , Cheng L , Yu S , Nie Y , Zhang H , et al. Improvement by application of three nanomaterials on flavor quality and physiological and antioxidant properties of tomato and their comparison. Plant Physiol Biochem. 2023; 201: 107834. doi:10.1016/j.plaphy.2023.107834. [Google Scholar] [CrossRef]
65. Ghanbari F , Bag-Nazari M , Azizi A . Exogenous application of selenium and nano-selenium alleviates salt stress and improves secondary metabolites in lemon verbena under salinity stress. Sci Rep. 2023; 13( 1): 5352. doi:10.1038/s41598-023-32436-4. [Google Scholar] [CrossRef]
66. de Brito Mateus MP , Tavanti RFR , Tavanti TR , Santos EF , Jalal A , Dos Reis AR . Selenium biofortification enhances ROS scavenge system increasing yield of coffee plants. Ecotoxicol Environ Saf. 2021; 209: 111772. doi:10.1016/j.ecoenv.2020.111772. [Google Scholar] [CrossRef]
67. Ahmad I , Chen C , Younas Z , Yousaf T , Mashwani ZUR . Seed and foliar application of nano-selenium improves sesame triacylglycerols and oil yield via photosynthetic pigment and enzymatic and chemical antioxidant enhancement revealed by spectrophotometric, UHPLC-analysis and chemometric modeling. Front Plant Sci. 2024; 15: 1431877. doi:10.3389/fpls.2024.1431877. [Google Scholar] [CrossRef]
68. Kanber S , Yildiztekin M , Firat Baran M . Eco-friendly synthesis of selenium nanoparticles via Sternbergia candida: enhancing antioxidant defense and mitigating salt stress in pepper (Capsicum annuum L.) plants. ChemistryOpen. 2025; 14( 6): e202400341. doi:10.1002/open.202400341. [Google Scholar] [CrossRef]
69. Islam SS , Haque MM , Sohag MA , Haque SN , Hoque MR , Chandra Sarker B , et al. Exogenous application of selenium nanoparticles boosted the drought tolerance in wheat by increasing morpho-physiochemical and yield attributes. Scientifica. 2025; 2025( 1): 7789912. doi:10.1155/sci5/7789912. [Google Scholar] [CrossRef]
70. Gheisary B , Fattahi M . Selenium and zinc oxide nanoparticles stimulate product quality, phenolic content, antioxidant activity, and shikonin production in Italian bugloss (Echium italicum L) plantlets under in vitro conditions. BMC Plant Biol. 2025; 25( 1): 1465. doi:10.1186/s12870-025-07461-2. [Google Scholar] [CrossRef]
71. Vicas SI , Rosan CA , Padilla-Contreras D , Cavalu SD , Zsiros R , Borza IM , et al. Foliar application of nanoselenium enhances drought tolerance in Brassica oleracea var. italica through antioxidant reinforcement and pigment stabilization. Life. 2026; 16( 1): 70. doi:10.3390/life16010070. [Google Scholar] [CrossRef]
72. Papkina AV , Perfileva AI , Zhivet’yev MA , Borovskii GB , Graskova IA , Klimenkov IV , et al. Complex effects of selenium-arabinogalactan nanocomposite on both phytopathogen Clavibacter michiganensis subsp. sepedonicus and potato plants. Nanotechnol Russia. 2015; 10( 5–6): 484– 91. doi:10.1134/s1995078015030131. [Google Scholar] [CrossRef]
73. Nozhkina OA , Perfileva AI , Graskova IA , Dyakova AV , Nurminsky VN , Klimenkov IV , et al. The biological activity of a selenium nanocomposite encapsulated in carrageenan macromolecules with respect to ring rot pathogenesis of potato plants. Nanotechnol Russia. 2019; 14( 5–6): 255– 62. doi:10.1134/s1995078019030091. [Google Scholar] [CrossRef]
74. Perfileva AI , Zakharova OV , Graskova IA , Krutovsky KV . Effect of selenium, copper and manganese nanocomposites in arabinogalactan matrix on potato colonization by phytopathogens Clavibacter sepedonicus and Pectobacterium carotovorum. Plants. 2024; 13( 24): 3496. doi:10.3390/plants13243496. [Google Scholar] [CrossRef]
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