Effect of nanoparticles on plants under salinity stress.
Abstract
Plants are under the threat of climatic changes and there is a reduction in productivity and deterioration in quality. The application of nanoparticles is one of the recent approaches to improve plant yield and quality traits. A number of nanoparticles, such as zinc nanoparticles (ZnO NPs), iron nanoparticles (Fe2O3 NPs), silicon nanoparticles (SiO2 NPs), cerium nanoparticles (CeO2 NPs), silver nanoparticles (Ag NPs), titanium dioxide nanoparticles (TiO2 NPs), and carbon nanoparticles (C NPs), have been reported in different plant species to play a role to improve the plant physiology and metabolic pathways under environmental stresses. Crop plants readily absorb the nanoparticles through the cellular machinery of different tissues and organs to take part in metabolic and growth processes. Nanoparticles promote the activity of a range of antioxidant enzymes, including catalase (CAT), peroxidase (POD), and superoxide dismutase (SOD), in plant species, which in turn improve the growth and development under stressful conditions. The present review focuses on the mode of action and signaling of nanoparticles to the plant systems and their positive impact on growth, development, and ROS scavenging potential. The appropriate elucidation on mechanisms of nanoparticles in plants leads to better growth and yields under stress conditions, which will ultimately lead to increased agricultural production.
Keywords
- agriculture
- climatic challenges
- crop development
- food security
- nanotechnology
1. Introduction
Population of the globe is rising and is predicted to reach almost 9.6 billion by the year 2050. Sustained growth of 70–100% in global agricultural and food production is essential to feed the growing population [1]. The area under cultivation may shrink over time due to the increasing nonagricultural uses of the land and urbanization, making it difficult to increase agricultural production [2]. Plants are constantly exposed to environmental changes during their life cycle. Deteriorating soil health conditions inevitably have a detrimental impact on plant development and productivity [3]. Billions of dollars worth of crops are being destroyed each year due to abiotic stresses, such as salinity and drought. Prior to the advent of efficient selection techniques, conventional breeding was used to maintain agricultural productivity, but its effectiveness was constrained by the diversity of stress tolerance traits [2]. Determining innovative solutions to reduce abiotic stress challenges and maintain food security is therefore urgently required under these deteriorating environmental conditions [1] as most promising approach currently available is nanotechnology.
To improve abiotic stress tolerance in agricultural biosystems, Eric Drexler initially coined the term “Nanotechnology” [4]. It deals with the study of nanostructures that possess diverse physicochemical properties and biochemical activities that are dependent on their surface-to-volume ratio [5]. Different physical, chemical, and biological processes can be used to manufacture nanoparticles (NPs), and they can interact with plants in a variety of ways [6]. Crop plants readily absorb NPs, which can enter the cells and play crucial roles in metabolic and growth processes [7]. There is a surge in the use of nanobiotechnology tools in agricultural production that has the potential to boost plant metabolism since NPs promote plant growth, development, and yield to withstand environmental stresses [8]. Additionally, it has been observed that NPs promote the activity of a range of antioxidant enzymes, including catalase (CAT), peroxidase (POD), and superoxide dismutase (SOD) [9]. Extensive research revealed that NPs are crucial for plants dealing with abiotic stress conditions [10].
Nanobiotechnology will improve plant functions that will help them cope with environmental challenges [11]. Therefore, this technology is strongly encouraged due to the rising global food demand as well as the potential for positive effects on the economy and ecology [12]. Despite its extraordinary potential in the enhancement of agricultural productivity and improvement in abiotic stress tolerance, the wider application of nanotechnology at the field level is limited in agriculture. In this review article, updates on the positive effects of nanotechnology for the improvement of abiotic stresses in crops have been discussed in detail.
2. Role of nanoparticles in salinity stress
Salinity has a negative impact on crops in various physiological and biochemical processes that decrease crop production drastically [13]. Water scarcity in the soil causes low osmotic potential and ionic toxicity of Cl− and Na+ in plant cells [14]. It is also seen that salt stress results in decreased concentration of photosynthetic pigments, reduced stomatal flow, lack of efficiency of photosystem II, and increased production of ROS.
2.1 Zinc nanoparticles (ZnO NPs)
Salt stress causes chlorophyll concentration leads to membrane disintegration and the rate of photosynthesis is significantly decreased. It also causes injury in thylakoid and grana that results in limited starch content [15]. Lupine (
2.2 Iron nanoparticles (Fe2O3 NPs)
Iron oxide nanoparticles are a rich source of Fe for plants. When peppermint was exposed to Fe2O3 NPs, proline content, and lipid peroxidation were decreased significantly in saline soil. Antioxidant enzyme activities (guaiacol peroxidase, CAT, and SOD) declined in plants. They also increased the potassium, zinc, calcium, iron, leaf dry and fresh weight, and phosphorus [23]. Grape softwood showed a prominent increase in protein content and reduced production of hydrogen peroxide, proline, and antioxidant enzyme activities when treated with potassium silicate and Fe2O3 NPs [24]. Under salt stress, the application of Fe2O3 NPs on
2.3 Silicon nanoparticles (SiO2 NPs)
SiO2 NPs are used to help plants by forming a layer in cell walls and maintaining yield. In squash and tomato plants, the antioxidant system is enhanced and seed germination increases due to SiO2 NPs under salt stress [26]. In Basil plants, silica nanoparticles have shown promising results related to morphological and physiological traits under salt stress [27]. SiO2 NPs increased the seedling growth of lentils and seed germination and improved the defense mechanism of plants in saline conditions [28]. They help plants to cope up with salt stress by increasing the fresh weight in maize [29]. Under salt stress, the application of SiO2 NPs on soybean decreased toxic ROS production and Na + level in leaves [30]. Wheat cultivars treated with SiO2 NPs improved biological antioxidant levels and seedling growth under salt stress [31]. Application of SiO2 NPs on the strawberry plant in saline conditions increased the photosynthetic pigment and maintained the carotenoid and chlorophyll content, decreasing the effect on epicuticular wax [32].
2.4 Cerium nanoparticles (CeO2 NPs)
They can be used as fertilizer to stimulate the growth of roots, enhance the antioxidant enzyme activities, and to prevent membrane leakage and peroxidation [33]. Moreover, CeO2 NPs help to preserve cell wall and chloroplast structure [34]. Activation of CeO2 NPs as antioxidants depends upon the pH of surroundings, sub-cellular localization, surface charge, concentration, and particle size. CeO2 NPs increased the growth in
2.5 Silver nanoparticles Ag NPs
Silver nanoparticles enhanced the sodium, potassium, and chloride to regulate the osmolality level in treated plants under salt stress conditions. The stability of Ag NPs can be easily controlled in aquatic environments as compared to soil conditions [38]. Priming of seeds was carried out with Ag NPs to enhance the seed germination in wheat and the development of tomato plants [39]. The combined effect of Ag NPs with NaCl reduced the thiobarbituric acid reactive substances, electrolyte leakage, and hydrogen peroxide to control the oxidative damage in plants that is linked with the overproduction of ROS [40].
2.6 Titanium dioxide nanoparticles (TiO2 NPs)
Titanium is a transition element and the 9th most abundant element that contributes 0.33% of the earth’s outer layer [43]. It improves photosynthesis and chlorophyll pigments in plants by altering antioxidant enzyme activities [44]. Titanium has an integral role in plants’ tolerance under stressful conditions [45].
2.7 Carbon nanoparticles (C NPs)
Carbon nanoparticles have distinctive properties, including morphologically small surface area and better chemical reactivity [47]. They help in increasing seed germination and crop production under salt stress [48]. Application of C NPs on
Type of NPs | Plant species | Physiological responses of plants under salt stress | References |
---|---|---|---|
Zinc NPs (ZNO NPs) | Enhanced photosynthetic pigments, antioxidant responses, and growth. | [16] | |
Increased antioxidant enzymes and improved salinity tolerance. | [17] | ||
Increased the activities of CAT and SOD and photosynthetic pigments and decreased the accumulation of soluble sugar and proline content. | [18] | ||
Ionic regulation, osmolyte synthesis, and upregulated oxidative defense system. | [19] | ||
Enhanced content of carotenoids and chlorophyll a & b. | [20] | ||
Decreased accumulation of total soluble sugars and proline contents, osmoprotectants. | [22] | ||
Iron NPs (Fe2O3 NPs) | Decreasing lipid peroxidation and increase in leaf dry and fresh weight, potassium, calcium, iron, phosphorus, and zinc contents. | [23] | |
Grape softwood | Increased protein content and reduced production of hydrogen peroxide, proline, and antioxidant enzymes. | [24] | |
Increased antioxidant activities. | [25] | ||
Improved osmolyte synthesis, antioxidant activities, and improved Na+/ K+ ratio. | [9] | ||
Silicon NPs (SiO2 NPs) | Tomato and squash | Increased ROS and seed germination. Maintained vitamin C concentration and chlorophyll content. | [26] |
Increasing proline accumulation, antioxidant responses, leaf dry and fresh weight, and chlorophyll content. | [27] | ||
Boosting defense mechanism by ROS. | [29] | ||
Decreased ROS, lipid peroxidation, and Na+, increased antioxidant activities and K+ content. | [30] | ||
Improved chlorophyll content and seed germination. | [31] | ||
Maintained carotenoid and chlorophyll content, and decreased the effect on epicuticular wax. | [32] | ||
Chitosan | Increasing chlorophyll content, growth, photosystem II, and enhanced nitric oxide bioactivity. | [53] | |
Cerium NPs (CeO2 NPs) | Rubisco carboxylase activity stimulates growth and increased photosynthesis. | [33] | |
Increased enzymatic and nonenzymatic defense system. | [35] | ||
Efficient chloroplast and increase in biomass. | [36] | ||
Silver NPs (Ag NPs) | Increased germination with seed priming. | [39] | |
Enhanced dry and fresh biomass of plants. Increased the activity of CAT and decreased the activity of POD, and increased the accumulation of soluble sugar and proline content. | [41] | ||
Improved proline and RWC, decreased oxidative damage by increasing the antioxidant enzyme activities. | [42] | ||
Titanium dioxide NPs (TiO2NPs) | Increased seedling growth, dry and fresh weight, root and shoot length. Increased antioxidant activity. | [46] | |
Carbon NPs (C NPs) | Reestablishing ion homeostasis and redox balance. | [49] | |
Increased enzymatic and nonenzymatic defense system, increased carotenoid, and chlorophyll content. | [50] | ||
Increased proline content in roots and leaves, photosystem II activity, soluble sugar in leaves, and membrane integrity was maintained. | [51] | ||
Enhanced phosphorus and potassium contents in root and phosphorus content in the shoot. Increased activity of antioxidant enzymes. Improved chlorophyll content, free ascorbic acid, amino acid, and soluble sugars. | [52] |
3. Role of NPS in drought tolerance
Water crisis is one of the several issues that have been afflicted by climate change and global warming. Water is crucial for plant vitality as it has a role in the transportation of nutrients. Therefore, water deficit results in drought stress, which harshly affects the survival of plants and reduces agricultural production [54]. Therefore, a key solution in relation to sustainable agriculture is the identification of resistant crop varieties or improving drought tolerance in plants. Crop management and coping with various environmental challenges are possible by the new features of nanotechnology. The negative impacts of a restricted water supply on agriculture have been attempted to be reduced utilizing nano-materials. Farmers may be able to identify the effects of stress on plants at an early stage by using nano-sensors in global positioning systems that produce satellite photographs of fields [55]. Crop production in drought-prone locations may rise if soils have been given better water-retention capabilities [56]. Using nanoparticle-based plant modifications, conventional technologies may be used to improve crop plants by increasing the capacity of food crops to retain water, and nanoparticles improve the effectiveness of water consumption in the plants [57].
3.1 Silicon nanoparticles (SNPs)
Only a few studies have documented the biological activity of silica, an element that makes up a major portion of the Earth’s crust and is found as silicon [58]. The tolerance of Hawthorn (
3.2 Zinc nanoparticles (ZnO NPs)
Application of Zn plays a role in increasing the radical growth and seed viability along with the establishment of germinated seeds, especially in Zinc deficient areas. Soybean seeds subjected to water stress showed that nano-zinc oxide has improved seed germination [62]. A composite of ZnO, B2O3, and CuO NPs reduced the effects of drought stress on soybean [63] and increased grain production and shoot growth by 36 and 33%, respectively, and improved nitrate and phosphorus uptake. By boosting the activity of the antioxidant enzymes SOD and CAT in wheat, ZnO NPs improved drought resistance. Zn and Cu NPs also increased the antioxidative enzyme activity, decreased lipid peroxidation, stabilized the photosynthetic pigments with increased RWC, and enhanced drought tolerance in wheat [64]. CuO and ZnO NPs modified the root morphology of plants colonized by
3.3 Iron nanoparticles (Fe2O3 NPs)
Iron is a key micronutrient that is essential for plant growth and development and its reduction causes chlorosis in plants [67]. Thus, iron absorption in plants experiencing drought stress may play a crucial role in their ability to withstand drought [68]. Fe2O3 NPs applied topically to
3.4 Titanium nanoparticles (titanium dioxide TiO2 & anatase titanium dioxide AnTiO2)
Wheat seed gluten and starch contents have responded favorably for using TiO2 NPs foliar applications and improved plant height, seed and ear numbers, ear weight, gluten and starch content, yield, biomass, and harvest index under drought stress [73]. Enhanced photosynthesis and increased the maize plant’s capacity to absorb light under water deficit [74]. Applying nanoparticles and Gibberellic acid to basil plants effectively enhanced the rate of photosynthesis, enhancing their tolerance to drought stress [75]. Various doses and sizes (10–25 nm) of AnTiO2NP on the flax plant under water-scarce conditions responded favorably for growth, development, hydrogen peroxide (H2O2), malondialdehyde content, seed oil production, protein content, and photosynthetic pigments [76]. The effects of TiO2 NP on onion seedlings have also been documented as they boosted SOD activity. TiO2 NP concentrations of 40 and 50 mg/ml, lowered the CAT and POD activities and secretion of amylase was reduced. However, seed germination and seedling growth were reduced at higher concentration of TiO2 NPs, whereas the effect was enhanced at lower concentration [77].
3.5 Silver nanoparticles (Ag NPs)
Developing drought-tolerant cultivars and reducing drought stress is essential for maintaining food security [78]. Only a small amount of historical literature focused on the interaction between drought and Ag NPs. Diminishing the detrimental impacts of drought stress on
3.6 Cerium oxide nanoparticles (CeO2 NPs)
Cerium oxide NPs could help
Type of NPs | Plant species | Physiological responses of plants under drought stress | References |
---|---|---|---|
Silicon NPs (SiO2NPs) | Positive effects on photosynthesis, malondialdehyde, RWC, membrane electrolyte leakage, chlorophyll, carotenoid, carbohydrate, and proline content. | [59] | |
Reduced shoot-to-root ratio, improved root growth and the maintenance of photosynthetic rate, augmentation of water uptake efficiency, increase in leaf area index and leaf weight. | [40] | ||
Improved shoot growth, increased the leaf chlorophyll content, maintained leaf water potential in stressed plants, reduced membrane lipid peroxidation. | [61] | ||
Zinc NPs (ZnO NPs) | Increased germination, radical growth and seed viability, shoot growth and grain yield, and uptake of N and P. | [62, 63] | |
Increased antioxidant enzymes, reduced lipid peroxidation, stabilized the photosynthetic pigments, proliferation of elongated root hairs, and increased water stress tolerant gene expression. | [64] | ||
Increased RWC and MSI, improved stem and leaf anatomy, photosynthetic efficiency, growth, and yield. | [66] | ||
Iron NPs (Fe2O3 NPs) | Improved crop yield at the flowering stage and enhanced oil percentage. | [69] | |
Enhanced plasma membrane proton pump ATPase activity, plant biomass, chlorophyll content, and CO2 assimilation. | [71, 72] | ||
Titanium NPs (Titanium dioxide TiO2 | Enhancement in seed gluten and starch contents, plant height, seed number, and biomass. | [73] | |
Improved sunlight absorbance, synthesis of photosynthetic pigments, and photosynthesis. | [74] | ||
Increased plant drought resistance and improve photosynthetic mechanism. | [75] | ||
Increased chlorophyll and carotenoids, enhancing flax development and yield, declining malondialdehyde and H2O2 content. | [76] | ||
Increased SOD activity, seed germination, and seedling growth. | [77] | ||
Silver NPs (Ag NPs) | Increased germination rate, root length, root fresh, and dry weight. | [68] | |
Cerium Oxide NPs (CeO2 NPs) | Enhanced catalytic scavenging of ROS, reduced O2− (41%), H2O2 (36%), increased yield, pollen germination, and CO2 absorption. | [80] |
4. Nanoparticles and other environmental stresses
Titanium oxide nanoparticles play a substantial role in the mitigation of light stress in crops as they catalyze the oxidation-reduction reaction, which then forms superoxide anion radicals and hydroxides. Oxidative stress is induced by ultraviolet (UV) light and has a negative impression on the growth of the plant. UV-B produces H2O2 and superoxide radicals and enhanced the leakage of electrolytes and lipid peroxidation, which leads to reduce the photosynthesis rate and normal leaf structure is also deteriorated [81]. In wheat plants, Silicon NPs increase antioxidant activities for the regulation of oxidative stress after UV-B exposure [82]. Herbicides are used to control weeds in agroecosystems. A methyl viologen herbicide, Paraquat is used extensively to control weeds in rice. Multiwall carbon nanotubes can modulate the toxicity of Paraquat [50], which promotes lateral root growth and photosynthesis in
5. Effect of nanoparticles on antioxidant and molecular mechanism of plants
Nanoparticles have an impact on plants’ antioxidant system at the molecular level as they increase the capability of plants to tolerate oxidative stress. When
The molecular mechanism of plants can be studied by using the model plant species.
Type of nanoparticles | Plant species | Impact on antioxidant system and ROS | References |
---|---|---|---|
Zinc NPs (ZnO NPs) | Decreased malondialdehyde content and increased SOD, CAT, POD, and APX activities under salinity stress. | [91] | |
Silver NPs (Ag NPs) | Significant increase in proline content, antioxidant enzyme activities, flavonoid contents, and phenolics. | [42] | |
Titanium NPs (Titanium dioxide TiO2 | Enhance antioxidant stress tolerance. | [92] | |
Enhance antioxidant enzyme activities contribute to reduction in hydrogen peroxide and malondialdehyde content under salinity. | [45] | ||
Silicon NPs (SiO2NPs) | Reduce the detrimental impact of Pb under lead stress by altering vitamin C, antioxidant enzyme activation, and flavonoids and increase plant capabilities to endure abiotic stresses. | [93] | |
Cerium oxide NPs (CeO2 NPs) | Scavenge ROS in isolated chloroplasts protect plant photosynthesis from detrimental effects Of ROS accumulation during abiotic stresses. | [87] |
6. Mechanism of nanoparticle (NPs) absorption in plants
Absorption and translocation of NPs in plants are one of the latest disciplines of study. The most commonly used NPs to enhance abiotic stress tolerance in plants are metal-based (MB) and carbon-based (CB). Among MB, the most widely studied nanomaterials are metal and metal oxides, such as copper, silver, titanium, iron, and zinc; while the most explored CB nanomaterials are carbon nanotubes (CNTs), fullerene (C70), and fullerol (C60(OH)20) [94]. The impact of NPs on plants is determined by a number of variables, including availability, uptake, translocation, and accumulation. The plant cell wall restricts the entry of foreign elements; therefore, effective techniques are needed to introduce advantageous NPs and make them available to plants.
Different factors like size, chemical content, and plant species affect the entry of NPs, which is further influenced by their stability, transport and absorption, toxicity, and accumulation [95]. Particle size, surface charge, and the hydrophobicity of the plant surface all play important roles in their absorption [96]. Additionally, the absorption rate and translocation in plants are directly correlated to the structure of the nanomaterial utilized [97]. All of these elements highlight the requirement for developing and enhancing laboratory techniques to comprehend the NPs physicochemical qualities [98] as they undergo biotransformation in the soil, which has a direct impact on their toxicity and bioavailability. Foliar spraying or incubating isolated cells, roots, pollen, seeds, and protoplast with NPs, direct injection, irrigation of plants with NPs, delivery by biolistic, and hydroponic treatment have all been employed in previous research to make NPs available to the plant cells [99]. Bioaccumulation defines the uptake of NPs by plant roots and travels through apoplastic and symplastic routes to the cortex and pericycle [100].
Nanoparticles entered through the stomata, cuticle, stigma, trichomes, wounds, and lenticels and move through the phloem. They reach the xylem and phloem through the root tip meristem, where the Casparian strips continue to the shoot but have not fully developed. Endocytosis allows NPs to enter cells even when the cell wall, cell membrane, and Casparian strips block their uptake and transport. Additionally, transporters like aquaporins and carrier proteins facilitate their easier entry into cells [101]. The capacity of roots to absorb nutrients can change if NPs accumulate on the surface of the roots. Parenchymatic intercellular gaps in seeds enable NPs to be directly absorbed before being diffused into the cotyledon. Stomata allow for the internalization of NPs larger than 10 nm, which are then delivered to the plant’s vascular system
7. Mechanism of translocation and accumulation of NPs in plants
Mechanism of NPs translocation in different plant cells and organelles has been clarified [96]. Plant cell wall serves as a barrier that manages NPs uptake and establishes solubilization needed to enable their translocation. NPs with a size range of 40–50 nm can easily pass through the cell wall [104]. Composition of NPs affects their mobility through cell membrane or cell wall and also encourages their adsorption to radical exudates. Positively charged NPs have better adherence to cell walls. Their coating and morphology have a big impact on how they behave inside plants and the rhizosphere [105]. After penetration through cells, they go through the shoots [101] and roots are transmitted to various aerial tissues and the seeds [6]. Gold NPs were only collected in the shoots of
Negatively charged gold nanoparticles are easily translocated from plant roots to shoots. The most stable are SiO2 and TiO2, as they remain present in plant tissues after their uptake. When
Different processes have been identified by the translocation of CeO2 and ZnO NPs into
Type of nanoparticles | Plant species | Conc (mg/l *mg/kg) | Accumulation (mg/kg) | References | |
---|---|---|---|---|---|
Roots | Shoots | ||||
Copper NPs | 1000 | 1544.1 | 17.27 | [109] | |
250 | 3773 | — | [110] | ||
125 | — | 18.46 | [111] | ||
1500 | 190.4 | — | [112] | ||
20 | 5.82 | 19.06 | [113] | ||
100 | 800 | — | [114] | ||
Silver NPs | 4000 | 2102 | 11.35 | [108] | |
1000 | 20 | 5 | [115] | ||
250 | — | 50 | [116] | ||
Zinc NPs | 1000 | — | 250 | [117] | |
100 | 10 | 30 | [107] | ||
Titanium NPs | 1000 | — | 250 | [117] | |
Magnesium NPs | 1000 | 103 | 131 | [118] |
8. Conclusion and prospective
Various abiotic factors are damaging the plants in terms of growth and development leading to a reduction in yield and deteriorating the quality. Applications of NPs have been proven very protective as they stimulate germination, growth, and improve yield. They increase tolerance in plants as they enhance the uptake of water and nutrients. They are capable to metabolize starch reserves in plant cells. They stimulate the process of photosynthesis and alter levels of phytochromes and modulate oxidative stress.
Although various studies suggest the beneficial roles of NPs in plants but the molecular basis of the actual mechanism is still unknown. Further elucidation on this mechanism may generate the smart NPs for the production of crops sustainable to the environment. In addition, their interaction with signaling molecules is also required to be explored. The economic stability of the use of NPs in agriculture is important as silver and gold nanoparticles costs very expensive. Understanding their mode of action, toxicity limits, signaling, and translocation; hence cheaper NPs may be used as an alternative.
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