List of the NPs applications in plant abiotic stress.
Abstract
The agricultural sector faces unprecedented challenges to ensure food security as the global population soars and climate change intensifies. Abiotic stresses are well-known for diminishing agricultural output and constraining crop yield generation worldwide. While conventional methods for managing crop stress fall short of meeting global demands, the integration of nanotechnology in agriculture offers a sustainable approach, providing a cornerstone for resilient and resource-efficient crop production in the face of evolving environmental challenges. Through targeted delivery systems and tailored formulations, nanoparticles exhibit the potential to enhance plant physiological processes, nutrient uptake efficiency, and stress tolerance mechanisms. This chapter describes the potential role of nanoparticles in abiotic stress management and activation of plant defence-related genes, improving the yield and quality of crops by combating nutrient deficiency and inducing stress tolerance. Moreover, it also discusses the potent molecular mechanisms upon application of nanoparticles for inducing tolerance to various abiotic stresses. However, while nanoparticle-based approaches hold great promise, their implementation also raises concerns regarding environmental impact, toxicity, regulatory frameworks, and socioeconomic implications.
Keywords
- abiotic stresses
- nanoparticles
- drought
- salinity
- heavy metal
- molecular pathway
- sustainable agriculture
- crop plant
- food security
1. Introduction
Over the last few decades, crop production has decreased even as food demands have risen due to an increasing human population [1]. According to the reports, 2 billion people have micronutrient deficiencies, 800 million people experience chronic hunger, and 653 million people will still be undernourished in 2030 [2]. In 2011, the Food and Agriculture organization estimated a loss of 1.3 billion tons of food per year around the globe. As a result, food security will remain a major concern as the world’s population is predicted to reach 10 billion in 2050 [3]. Climate change is considered a major limiting factor, having a detrimental effect on plant growth, development, crop yield, and productivity. Abiotic stresses such as drought, salinity, soil erosion, flooding, and nutrient depletion are the byproducts of climate change and are one of the key factors for stunted plant growth, resulting in reduced yield and quality of crop plants [4]. The development of stress-tolerant plants could be a promising and practical solution to this challenge. However, conventional breeding methods have not yielded the desired results in improving crop plant tolerance to stress through inter-generic or inter-specific hybridization [5]. Thus, novel and promising agricultural strategies capable of reforming modern agriculture are required to increase food productivity while limiting pesticide and fertilizer use to reduce abiotic stress. Nanotechnology is currently an emerging and rapidly developing field of study that has gained tremendous attention recently due to its broad-spectrum applications in various sectors, particularly in plant biotechnology and agriculture. Nanotechnology, in particular, can help provide effective solutions to agricultural problems and achieve a sustainable and secure future for agriculture [6]. Nanotechnology uses nanoparticles as a tool to mitigate abiotic stress factors and to improve crop yield and food quality.
Nanoparticles (NPs) are small materials ranging in size from 1 to 100 nm. In contrast to their larger-sized counterparts, NPs have distinct and diverse physicochemical properties including high adsorption efficacy, large surface area-to-volume ratio, and effective linking and work efficiency. These qualities make them a suitable candidate for disease management strategies as bactericides/fungicides/pesticides to boost plant health. Moreover, NPs can also help to alleviate nutrient deficiency symptoms and supplement essential elements when used as macro and micro-nano fertilizers in plants [7]. Titanium dioxide (TiO2), iron oxide (Fe3O4), zinc oxide (ZnO), silicon oxide (SiO2), copper (Cu-NPs), and selenium (Se-NPs) have recently caught significant attention due to their non-threatening use in agriculture [8, 9, 10]. Different biological, physical, and chemical techniques can be used for NPs synthesis. The impact of NPs on plants varies and primarily depends upon the size, origin, time of application, and concentration [11]. Recent studies reported that the application of NPs resulted in improved plant tolerance against abiotic stresses by increasing the antioxidant activities, decreasing hydrogen peroxide (H2O2) and malondialdehyde (MDA) accumulation, and maintaining the efficiency of the photosynthetic apparatus. Moreover, they also have the ability to reach the photo system-II (PS-II) reaction center and increase the transmission of electrons and light absorption by penetrating chloroplast, resulting in improved photosynthetic efficiency and plant growth [12, 13].
Despite their extensive and commercial use in different products, concerns about NP’s toxicological and environmental impacts are circulating in the research community [14]. It has been reported that the excessive use of NPs can result in physiological abnormalities and oxidative stress in plants, leading to reduced antioxidant activities and gas exchange characteristics. Moreover, disruption of cell division, mitotic index, and root growth has also been reported [15, 16]. Most of the published literature emphasizes the positive effects of NPs on growth and crop productivity. However, they lack an in-depth understanding of NP-plant interaction at the biochemical, physiological, and molecular levels. Therefore, this chapter will focus on a detailed understanding of interaction of NPs with plants at the molecular level and help to fill in research gaps that will enable potential future use of NPs. This comprehensive review will be a valuable source of information to conduct more studies and develop new eco-friendly NPs of different metals to counter the devastating impact of abiotic stresses on plants.
2. Effects of abiotic stress on crop plant
Climate change has led to an increase in abiotic stress conditions, which are gradually affecting crop production in various respects. Different physiological and biochemical changes occur in plants during stress, resulting in disruptions in plant growth, development, and productivity. Nanobiotechnology could be the most efficient adaptation strategy to cope with climate and abiotic stress factors, thereby achieving sustainable production [17, 18]. These stressors encompass heat, drought, high soil salinity, and heavy metal (HM) toxicity—cadmium (Cd), lead (Pb), arsenic (As), nickel (Ni), chromium (Cr), cobalt (Co), and zinc (Zn) [17]. Drought and heat stress are chief contributors to reduced crop yield around the globe. Prolonged drought and heat increase reactive oxygen species (ROS) production, affecting oxidative processes and causing reductions in stomatal opening, leaf size, water potential, and root growth, resulting in a general decrease in crop yield (Figure 1) [18, 19].
Salinity and heavy metal stress are also considered among the environmental factors that limit crop yield in many countries. Soil salinity causes low osmotic potential around the seeds, preventing water uptake and thus resulting in inhibition of seed germination. Oxidative stress in legumes induced by sodium chloride (NaCl) decreases seed nutrient quality, lowers nodulation, and inhibits plant growth. Ascorbate and carotenoids are important non-enzymatic salinity defense mechanisms, whereas proline (Pro) is a known osmoregulatory stress-related compound [20, 21].
Various factors contribute to heavy metal accumulation in agricultural soils, such as chemical fertilizers, atmospheric deposition, sewage sludge, and rapid industrial growth [22]. Heavy metals significantly impede plant growth by disrupting essential functions like food intake, antioxidant enzymes, and photosynthetic machinery. The increased generation of reactive oxygen species (ROS) during heavy metal stress worsens the decline in agricultural productivity [20, 21]. The annual economic cost of heavy metal pollution exceeds $10 billion, primarily attributed to anthropogenic and industrial waste deposition. Mitigating heavy metal-induced stress requires a comprehensive approach, including transgenic crop engineering, marker-assisted breeding, and traditional breeding. Sustainable agricultural practices are vital for enhancing plant resilience in the face of heavy metal contamination [23].
Efforts to mitigate the impact of abiotic stressors on crop productivity require integrated strategies that address drought, salinity, cold, heavy metal contamination, and other environmental challenges. Sustainable agricultural practices, advanced breeding techniques, and a focus on resilience are essential components of a comprehensive approach to ensure food security in the face of a changing climate [24, 25].
3. Mechanisms of nanoparticle-plant interaction in crop plant
Plants and nanoparticles interact in a complicated and multidimensional way that is controlled by molecular processes, physiological reactions, barrier crossing, uptake, and defense [26]. Therefore, comprehending these dynamics is crucial in assessing the possible advantages and drawbacks of employing nanoparticles in agricultural and environmental settings [27]. In addition to that, optimizing the characteristics of nanoparticles, reducing their negative impacts, and maximizing their potential for environmental remediation and sustainable agriculture techniques are also essential [28]. At specified concentrations, distinct NPs have been shown to boost the growth and development of plants through standard mechanisms [29]. A detailed mechanism of action for nanoparticles within plants is given below (Figure 2).
3.1 Uptake mechanisms
Adsorbed nanoparticles (NPs) from the soil can adhere to the surface of the roots. Subsequently, they can enter the root system by direct absorption, root junctions, or wounds. Additionally, these particles may enter plants through their above-ground sections, primarily through cuticles (a non-polar channel) and stomata or trichomes (a polar pathway) [30, 31]. Artificial exposure might happen when they are sprayed on or directly injected into the leaf [31].
3.2 Absorption and intracellular movement
The first line of defense against NPs is the cell wall of plants, wherein the cellulose permits the penetration of tiny NPs (5–20 nm) while obstructing the bigger ones [32]. After navigating the cell, there are two ways for them to navigate through tissues: the apoplast and the symplast. Apoplastic transport occurs outside the plasma membrane through the extracellular spaces, cell walls of adjacent cells, and xylem vessels while symplastic transport uses plasmodesmata and sieve plates to move water and substances between the cytoplasm of adjacent cells [33, 34]. The apoplastic pathway is essential for radial movement within plant tissues and helps nanomaterials reach the root central cylinder and the vascular tissues. When reached inside the central cylinder, nanoparticles make their way to aerial parts of the plant by using a transpiration stream [35]. However, because Caspary strips function as a resistive barrier, nanoparticles can form aggregates and concentrate in the endodermis. The nanoparticles must then enter the symplast for effective translocation to shoots [36].
Using plasmodesmata and retina structures, symplastic transport is the transfer of chemicals and water between the cytoplasm of adjacent cells [37]. Since the transfer of nanomaterials to the symplast allows for their efficient translocation to the shoots, this mechanism is thought to be more significant for the transportation of nanoparticles. The symplastic transit of nanoparticles across the membrane and their accumulation influences the charge inside the cell, which modifies the electrochemical potential of the membrane. These changes can have an impact on the movement of other materials over the membrane, so to sustain turgor pressure, water and nutrient intake, and plant development, the membrane’s electrochemical potential must stay in equilibrium [38].
In the case of foliar applications, nanomaterials must cross the barrier the cuticle presents, following the lipophilic or the hydrophilic pathway [39]. Numerous other pathways also allow nanoparticles to penetrate the intracellular area [40]. They could get through by attaching to transport proteins, ion channels, aquaporins, endocytosis, or by starting the creation of new pores, some of which might be bigger than the ones that already exist [41]. The processes of nanoparticle adsorption, translocation, and accumulation are heavily influenced not only by the type of plant but also by the physical and chemical properties of the nanoparticles themselves [42].
The way NPs move inside the plant is a key indicator of their accumulation and storage in different parts of the plant. For example, if an NP took the apoplastic pathway and used the xylem for translocation, then it would most probably move from roots to shoots and aerial parts of plants but would not move downwards toward roots. Thus, application methods for these types of NPs should be carried out at the root level, ensuring a good distribution in the plant. On the other hand, if an NP translocates via phloem and takes the symplastic pathway, then it will accumulate in plant organs acting as sinks, such as fruits and grains. Thus, appropriate measures should be taken while applying NPs to plants to avoid further human or animal ingestion of nanomaterials. Moreover, translocation is not essentially limited to a specific cell type, and lateral movement of nanomaterials between xylem and phloem is also possible [43].
The crystallinity of the nanoparticles utilized is one of the most critical elements influencing accumulation [42], the hydrodynamic dimension of the NPs, the nature of the surrounding soil, such as whether it is “organic” or inorganic, and the influence of zeta potentials [44]. The plant’s large concentration of nanoparticles results in a high amount of phytotoxicity. The phytotoxic impact of NPs is determined by their size, shape, exposure period, and concentration [45].
3.3 Nanoparticle-plant interaction
Plants’ regular cellular activities can be disrupted by NPs, resulting in oxidative stress. This disruption is frequently caused by NPs interfering with the electron transport chain in mitochondria and chloroplasts, where these particles cause plants to produce ROS, resulting in an oxidative burst [46]. This increased ROS level can induce cellular damage such as lipid peroxidation, DNA damage, and protein alterations [47]. But despite its destructive nature, it is essential for various biological activities like stress tolerance. However, fortunately, plants are gifted with antioxidant defense systems to counterbalance the negative effects of high ROS levels. Enzymatic (e.g., superoxide dismutase, catalase, and guaiacol peroxidase) and non-enzymatic (e.g., ascorbate, glutathione, and carotenoids) molecules are produced in response to scavenge excess ROS [48]. NP-plant interactions can disturb hormonal homeostasis, influencing plant metabolism and stress responses. In response to NPs, many hormonal pathways may be activated or downregulated, regulating plant growth, development, and stress tolerance [49]. A study conducted on
Nanoparticles cannot work successfully unless they interfere with the molecular processes and gene expression of plants. The diverse gene expressions of micro-RNA were studied in nanoparticle-treated cells. Nanoparticles have an effect on miR-398 and miR-408, which are responsible for seed germination, root length, seedling development, and free radical scavengers [51]. The random drop in miR-164 expression, which is involved in auxin hormone signaling, is, nevertheless, linked to the nanoparticle-mediated rise in roots [52].
4. Synthesis of nanoparticles and their role in alleviating abiotic stress
As their name implies, nanoparticles have distinct physicochemical features and range in size from 1 to 100 nm. Usually, both top-down and bottom-up synthesis methods are used to synthesize nanoparticles [53]. Under the top-down method, bigger molecules break down to generate nanoparticles. Examples of top-down methods include electro-explosion, mechanical milling, chemical etching, laser ablation, and sputtering. Metal-oxide nanoparticles are mostly synthesized by [54]. In contrast, simpler materials mix and react to generate nanoparticles during bottom-up synthesis. Examples of bottom-up techniques include pyrolysis, condensation, the sol-gel process, and spinning. The bottom-up approach includes the green production of nanoparticles, typically from biological material [55]. Besides, NPs can also be synthesized by using the Green synthesis approach which involves reacting processed extracts of plants, fungi, and microorganisms with metal salt solutions, followed by reduction and capping to create metal nanoparticles coated with target biomolecules. Green-synthesized nanoparticles help improve results and facilitate efficient transportation to the target spot due to their small size [56].
The success of nanoparticle production depends on the following characteristics:
Characterizations of morphology by microscopy (scanning and transmission electron microscopy).
Characterization of structures by Zeta-size analyzers, Raman, XRD, and XPS.
UV-visible and photoluminescence optical characterizations [57].
4.1 Commonly used nanoparticles for abiotic stress
Several metals including silver (Ag), copper (Cu), gold (Au), iron (Fe), titanium (Ti), and zinc (Zn) and their oxides have been used for the green synthesis of NPs using plants and their extracts, microorganisms and membranes, and DNA of viruses including diatoms.
Recently, it has been reported that the cerium oxide (CeO) NPs application maintained the quantum yield of photosystem (PS) II and CO2 assimilation through ROS scavenging, particularly hydrogen peroxide, induced by abiotic stress. The application of titania (TiO2) NPs improved the activity of catalase (CAT), glutathione peroxidase (GPOX), and superoxide dismutase (SOD) and reduced oxidative stress in Duckweed (
Nanoparticles | Dosage (mg/L) | Duration (days) | Plant species | Abiotic stress | Application | References |
---|---|---|---|---|---|---|
Selenium dioxide | 20–40 mg | 30 | Salinity | Enhanced the plant growth and yield | [65] | |
Calcium oxide | 25 mg | 30 | Heavy metal | Increased plant growth, photosynthesis efficiency, and antioxidant enzymes | [66] | |
Silicon | 2–50 mg | 45 | Salinity | Improved oxidative stress tolerance | [67] | |
Gold | 10 mg | 07 | Salinity | Improved the plant defense systems | [68] | |
Cerium oxide | 50–100 mg | 21 | Salinity | Improved the crop yield by modulating the plant’s physiological and biochemical mechanisms | [69] | |
Cerium oxide | 50–100 mg | 30 | Salinity | Improved the plant growth by maintaining cytosolic K+/Na + ratio | [70] | |
Zinc oxide | 0.5–1 mg | 1–7 | Salinity | Enhanced plant germination and salinity tolerance by improving the activities of antioxidant enzymes | [71] | |
Iron oxide | 10 mg | 07 | Cadmium and drought stress | Increased the tissue dry weight and reduced the Cd accumulation | [63] | |
Iron oxide | 10 mg | 05 | Salinity and heavy metal | Facilitates photosynthetic pigments and restricts cadmium uptake | [72] | |
Silver | 50–75 mg | 10 | Salinity | Improved plant growth and germination | [73] | |
Iron | 5–20 mg | 15 | Heavy metal and drought stress | Improved photosynthesis and alleviated oxidative stress | [12] | |
Zinc oxide | 0.5–1 g | 07 | Heavy metal and drought stress | Increased plant growth and reduction in chromium bioavailability | [74] | |
Selenium | 100 mg | 03 | Drought | Enhanced plant growth and development | [75] | |
Titanium oxide | 2–5 ppm | 01 | Salinity | Promote plant growth and ameliorate salinity stress | [76] | |
Copper | 50, 100, 200 mg | 30 | Heavy metal | Increased biomass, antioxidant enzyme contents, and photosynthesis efficiency | [77] | |
Silica | 2–50 mg | 21 | Drought and salinity | Improved the growth and productivity of cucumber plants by balancing nutrient uptake | [78] | |
Titanium oxide | 1.5–10 ppm | 07 | Salinity | Improved growth and enhanced tolerance against salinity | [79] | |
Cerium oxide | 50–100 mg | 14 | Salinity | Enhanced plant growth by regulating photosynthesis and water use efficiency | [80] | |
Chitosan | 100 mg | 01 | Salinity | Mitigates the deleterious effects of salinity | [81] | |
Titanium oxide | 2–5 ppm | 03 | Cold | Increased plant growth and antioxidant activity | [82] | |
Titanium oxide | 50–100 mg | 30 | Heat | Enhanced plant growth and photosynthesis efficiency | [83] |
4.2 Delivery methods of nanoparticles to crop plants
The delivery of nanoparticles to crop plants can be achieved through various methods to enhance plant growth, increase nutrient uptake, and mitigate environmental stresses. Different application methods are described in detail below (Figure 3).
4.2.1 Seed coating
Nanoparticles are applied as coatings on the surface of seeds. Seeds are immersed or coated with a solution containing nanoparticles before planting. This method ensures direct contact between nanoparticles and emerging roots, promoting early interaction with the plant [84].
4.2.2 Soil drenching
Nanoparticles are applied as a solution directly to the soil around the plant root zone. Nanoparticle solutions are poured or sprayed onto the soil surface, allowing the roots to absorb the particles. Effective for delivering nanoparticles to the root system, promoting nutrient absorption and root development [85].
4.2.3 Foliar spray
Nanoparticles are applied as a spray directly to the leaves and stems of plants. In this method, a nanoparticle solution is sprayed onto the plant’s vegetative parts. Plants absorb foliar NPs usually through stomata, ion channels, protein carriers, cracks or water pores, endocytosis, stigma, wound and trichomes. This approach allows rapid uptake, controlled release of NPs at targeted locations and improved absorption and assimilation of foliar fertilizer [86].
4.2.4 Hydroponic systems
Nanoparticles are added to the nutrient solution in hydroponic systems. Nanoparticles are dissolved in the nutrient solution, and plants absorb them through their roots in a soilless growing medium, which provides precise control over nutrient delivery and uptake [87].
4.2.5 Root injection
Nanoparticles are injected directly into the plant root system. Nanoparticle solutions are injected into the soil around the root zone or directly into the roots, which offers a targeted and efficient delivery method [88].
4.2.6 Biological vector-mediated delivery
Nanoparticles are delivered into plants using biological vectors, such as bacteria or fungi. Engineered microorganisms carrying nanoparticles are introduced to the plant, facilitating uptake, which utilizes natural biological processes for nanoparticle delivery [89].
4.2.7 Nanoparticle-embedded fertilizers
Nanoparticles are incorporated into conventional fertilizers. Nanoparticles are blended with fertilizers, allowing for simultaneous delivery of nutrients and nanoparticles during standard fertilization practices, which integrates nanoparticle application with routine agricultural practices [90].
NPs were taken up by roots and/or leaves, and they moved through two different channels: apoplastic and symplastic [91]. After passing through the first barrier, the cell walls, NPs were distributed between the plasma membrane and the walls of the cells; osmotic pressure and capillary forces may have an impact on the NPs’ subsequent migration [92]. In addition to ion channels and transport or carrier proteins like aquaporin, NPs can also enter cells by cell membrane barrier breaking or endocytosis [93]. The precise location and function of NPs in plant organelles are not exploited by any mechanistic method [94]. The translocation and uptake kinetics of NPs should be investigated further, as well as the biochemical, molecular, and physiological patterns of NPs uptake and transport kinetics, as these are crucial to comprehending the accumulation behavior of NPs in plants.
5. Molecular responses of plants to nanoparticles under abiotic stress
The complex and diverse array of molecular responses triggered in plants under abiotic stresses leads to the activation of stress phytohormones, accumulation of reactive oxygen species (ROS), kinase cascades, and stimulation of ion channels, contributing to electro-physiology and signaling [95, 96, 97]. Subsequently, a sophisticated species and stress-specific regulation of gene networks occur, targeting defense and repair mechanisms against stress-induced injuries [98]. Stress-induced proteins play roles in ROS neutralization, mitogen-activated protein (MAP) kinase, and salt overly sensitive (SOS) kinase signaling cascades [96, 99]. Recent studies reveal functional outcomes extending to ion and water transport, uptake, and transcription regulation for secondary response elements (Figure 4) [100, 101].
Nanoparticles (NPs) have proven to be useful in triggering these molecular responses when they are applied to plants facing abiotic stress conditions. Noteworthy NPs include aluminum oxide (Al2O3) NPs, silicon-NPs, zinc oxide (ZnO) NPs, multi-walled carbon nanotubes, and titanium dioxide (TiO2) NPs. Research by Zhao et al. underscores the potential of nanomaterials like silver and copper oxide, inducing stress responses and defense mechanisms and enhancing crop stress resilience [102]. Proteomic and transcriptomic approaches have helped to investigate the molecular effects of NPs on various plant species, with observed morphological and physiological effects influenced by NP dosage, type, size, and shape.
5.1 NPs mitigate salinity stress in plants
Nanoparticles (NPs) play a crucial role in mitigating salinity stress in plants by influencing cellular mechanisms and gene expression [103]. Studies on micro-RNA expression in NP-treated cells revealed the impact of NPs on miR398 and miR408, crucial for regulating seed germination, root and seedling growth, antioxidants, and free radical scavengers [104, 105]. Foliar application of Zn-NPs on rapeseed plants under salinity stress altered gene expression, reducing some genes (e.g.,
Carbon-based NPs, specifically multi-walled carbon nanotubes [107], affected genes involved in antioxidant and salt sensitivity systems in rapeseed plants under salinity stress [108]. According to Zhao et al., under salinity stress, multi-walled carbon nanotubes can change gene expression, affecting the antioxidant and salt overly sensitive 1 (SOS1) system in rapeseed plants [109]. Cerium oxide NPs decreased ROS levels and increased calcium content in plants, influencing ROS and Ca2+-mediated signaling genes [110]. Silicon-NPs demonstrated positive effects on
Abiotic stress | Nanoparticles | Plant species | Genes | Gene regulation and plant response | References |
---|---|---|---|---|---|
Drought | Fe, Cu, Co, and ZnO | Increased expression, regulates ABA biosynthesis and signaling pathways involving ABA hormones | [116] | ||
Fe, Cu, Co, and ZnO | improve expression in leaves, Fe-NP exhibits improved expression in both roots and leaves, ↑ drought tolerance by upregulating drought-related genes and, subsequently, ↓ ROS levels, regulating osmolytes, and increasing flavonoid biosynthesis. | [116] | |||
Fe, Cu, Co, and ZnO | Increased expression in roots with all NPs, regulate gene expression by activating transcription factors like DREB2 or ERD1. | [116] | |||
Fe, Cu, Co, and ZnO | ↑ Expression in roots with Fe- or Co-NP, part of the ABA-dependent pathway, high expression in roots results in drought tolerance. | [116] | |||
Fe, Cu, Co, and ZnO | ↑ Expression in both root and leaves, functions in the ABA-independent pathway, ↑ Expression in the cascade of genes acting directly in response to abiotic stress. | [116] | |||
Fe, Cu, Co, and ZnO | Increased expression in roots with Fe, Cu, and ZnO-NPs, regulate ABA-independent pathways & improve drought tolerance. | [116, 117] | |||
ZnO | Upregulation of CAT1 in leaves, ↑ CAT activity, ↑ stress tolerance. | [118] | |||
ZnO | ↑ Expression of proline biosynthesis in leaves, improves drought stress. | [118] | |||
ZnO | ↑ Expression in leaves encodes LEA D-11 DHN (dehydrin), part of the ABA-dependent drought tolerance pathway. | [118, 119] | |||
Drought | Silica | ↑ Expression in leaves resulting in ↑ expression of plasma membrane-localized transporters → ↑ movement of substances like abscisic acid → increased tolerance. | [119] | ||
Silica | ↑Regulation in leaves plays a role in mitigating the effects of drought tolerance. | [119] | |||
ZnO | ↑ Expression of antioxidant genes→ inhibit H2O2, resulting in improved drought tolerance. | [117] | |||
Chitosan-GSNO | ↑ Expression in drought stress, a multi-member family of cysteine-rich proteins having strong antimicrobial activity and play a key role in protection against drought. | [120] | |||
Chitosan-GSNO | Increased expression, early response gene of drought stress | [120] | |||
Salinity | Fe2O3 | Over expression of PAL and TAT genes, ↑production of RA that showed a positive correlation with MDA and scavenging of DPPH radicals. | [121] | ||
Chitosan | High expression, improve the plant growth and development under salinity by increasing the alkaloid contents in the leaf and roots. | [122] | |||
BioSeNPs | Over expression of BnPIP1-1 and BnPIP2-1, ↑ROS scavenging attributes (enzymatic and non-enzymatic antioxidants) and enhanced water potential. | [123] | |||
PEC | Upregulation of DEGs, 46% decreased ROS accumulation in roots, improved plant germination and morphological traits | [110] | |||
Fe3O4 and ZnO | ↑expression of APX and SOD genes. ↓ Regulation of the GR gene which activated the antioxidants against salinity stress. | [124] | |||
Se | Overexpression of PAL and RA genes, increased activity of SOD, CAT, and POD. | [125] | |||
Salinity | PNC | ↑Expression of nia 1, increase the production of nitric oxide up to 34%, ion of nia1 gene. | [126] | ||
ZnO | Downregulation of MYC, MPK4, and SKRD2 genes. 44% upregulation of ARP mRNA expression, which resulted in improved plant physiology and morphology. | [106] |
5.2 NPs alleviate the impact of drought stress in plants
Sessile organisms such as plants are constantly exposed to a range of abiotic elements such as drought, resulting in a substantial decrease in yield. Drought stress impairs the photosynthesis, nutrient uptake, osmotic, and antioxidant activities of plants. Photorespiration can lead to overproduction of reactive oxygen species (ROS) in drought-stressed plants, resulting in denaturation of proteins, DNA damage, and lipid peroxidation, which hinders cell growth and elongation, resulting in poor plant growth and productivity [127].
Recent studies have reported the role of metal-based and carbon-based NPs in mitigating drought stress by inducing tolerance. Metal-based NPs, such as ZnO, TiO2, Fe, and Cu-NPs, and carbon-based NMs such as graphene, fullerene, fullerol, and carbon NTs, have been extensively used to control drought stress by increasing water and nutrient uptake through stress tolerance and upregulation of genes involved in cell growth [128, 129].
The expression of the
5.3 NPs mitigate extreme temperature effects in plants
The tolerance of plants to heat and cold stress involves the regulation of numerous genes, transcription factors, and proteins. The expression levels of these components, whether upregulated or downregulated, directly impact the plant’s ability to survive under challenging conditions. For instance, in rice, the foliar application of ZnO-NPs induced the expression of genes related to the antioxidative system and chilling-response transcription factors in chilling-treated seedlings [134]. In soybeans, ZnO-NPs led to the upregulation of various genes, including
Stress | NPs | Plant | Genes | Major regulated genes | References |
---|---|---|---|---|---|
Cold Stress | TiO2 | Upregulation of LRubisco; SRubisco; chlorophyll a/b-binding protein; PEPC. | [82] | ||
Cold Stress | TiO2 | Upregulation of genes, Differential expression of genes related to cellular defense, cell signaling transcriptional regulation, and chromatin modification. | [141] | ||
Heat Stress | CeO2 | Upregulation of Heat Shock Proteins (HSP70) and excessive production of hydrogen peroxide H2O2. | [142] | ||
Heat Stress | MWCNTs | Upregulation of HSP90 | [143] | ||
Flooding | AL2O3 | Differential expression of proteins related to protein synthesis/degradation; glycolysis; lipid metabolism. Upregulation of Downregulation of | [144] | ||
Flooding | Ag | Differential expression of root proteins for cell signaling and cell metabolism. | [145] | ||
Flooding | Ag | Upregulation of proteins related to protein metabolism; cell division/organization; amino acid metabolism; | [146] | ||
Cd Stress | Si | Downregulation of Upregulation of | [147] | ||
Fluoride stress | Ag | Downregulation of NADPH oxidase and | [148] |
5.4 NP intervention in alleviating the adverse effects of heavy metals in plants
Cong et al. outlined the impact of silicon nanoparticles (Si-NPs) in diminishing the absorption and toxicity of cadmium (Cd) in rice [149]. Si-NPs inhibit genes linked to the transportation and uptake of Cd from the root to the shoot, specifically targeting low-affinity cation transporter (
5.5 Flood stress mitigated by NP in crop plants
Flooding induces severe oxygen depletion in plant roots, leading to significant physiological changes [153]. In a study using
6. Nanotechnology for sustainable agriculture in crop plants
The integration of nanotechnology in agriculture offers a sustainable approach, providing a cornerstone for resilient and resource-efficient crop production in the face of evolving environmental challenges. Nano-enabled technologies, ranging from nanofertilizers to genetic engineering, present a paradigm shift in promoting plant growth and resilience [154]. The superior attributes of nanoparticles (NPs), including a high surface-area-to-volume ratio and efficient nutrient supply, distinguish them from conventional fertilizers [155]. Studies demonstrate the potential of NPs, such as iron oxide (Fe3O4) NPs and nano-silicon (Si), in enhancing crop growth under diverse stress conditions, encompassing heavy metal-contaminated soil and drought stress [156, 157]. Notably, green-synthesized silver nanoparticles (AgNPs) enhance antimicrobial activity in
Moreover, examples such as cerium oxide (CeO) NPs preserving photosystem yield [58] and titanium dioxide (TiO2) NPs boosting antioxidant enzyme activity underscore the diverse avenues through which NPs positively influence plant health [160, 161]. Leveraging the stress ameliorative potential of NPs, particularly through green synthesis methods, emerges as a promising tool to counteract negative effects induced by abiotic stresses in various crops [162].
6.1 Applications of nanotechnology for crop protection
In the context of plant abiotic stress management, nanotechnology can play a role in enhancing plant tolerance to environmental stressors. Here are some applications of this emerging technology that have been studied for their potential to mitigate abiotic stress in plants:
6.1.1 Nanofertilizers
Nanofertilizers are nutrients encapsulated or coated within nanomaterial to enable controlled release, and its subsequent slow diffusion into the soil. These fertilizers help to restore degraded soils, manage soil health, improve seed germination, enhance plant growth, and increase crop yield like nitrogen nanoparticles designed to improve the efficiency of nitrogen delivery to plants. Phosphorus nanoparticles enhance the availability and uptake of phosphorus by plants. Potassium nanoparticles improve the delivery of potassium to plants for better growth and stress resistance [155, 163].
6.1.2 Nanomediators
They act as efficient carriers for targeted delivery of genetic material, regulating gene expression to improve stress response in plants. These nanoparticles can also aid in nutrient management by facilitating the controlled release and mitigation of persistent organic pollutants, Pesticides, and HM. Facilitate the targeted delivery of genetic material for gene therapy [164].
6.1.3 Nanopesticides
Nanopesticides revolutionize agricultural practices by offering targeted and controlled release of pesticides. This reduces environmental impact by minimizing pesticide usage while effectively managing both target and non-target plant pests. Additionally, nanopesticides contribute to plant disease management, enhancing crop health and ultimately improving overall yields. As a result, these innovative formulations showcase potential advancements in sustainable and efficient pest control strategies for modern agriculture [165, 166].
6.1.4 Nanosensors
Nanosensors, incorporating nanoparticles, play a pivotal role in detecting biological molecules and assessing environmental conditions. In agriculture, they help monitor heavy metal levels in soil, ensuring optimal plant growth and preventing contamination. Nanosensors also enhance the sensitivity and selectivity of gas sensors, allowing for precise detection of gases related to environmental health. Their versatility extends to identifying pollutants and contaminants in the environment, offering a promising avenue for real-time, high-precision environmental monitoring and management [167].
6.1.5 Nanomaterials
Nanomaterials such as carbon nanotubes (CNTs), graphene nanoparticles, and metal nanoparticles are employed to enhance plant resilience to various environmental stresses. In the context of salinity stress, these nanomaterials have shown promise in increasing plant tolerance by regulating ion balance and minimizing salt-induced damage. Additionally, they play a crucial role in reducing the bioaccumulation of heavy metals in plants and mitigating environmental contamination [168, 169].
7. Challenges and risks associated with nanobiotechnology
Although nanobiotechnology harbors immense potential for future agricultural technology revolutions, particularly considering climate change and growing populations, it also presents numerous challenges and associated risks. This requires more study and research on nanomaterials and their properties, as they behave very differently than in bulk form. Some materials are suspected of being toxic at the nanoscale due to significant, yet unknown, hazardous properties related to their unique physiochemical features. This can put manufacturers, formulators, handlers, applicators, and consumers at greater risk. Thus, nanoparticles can cause heterogeneous effects as they can leave toxic effects on non-target organisms upon contact, i.e., nanoparticles can come into direct contact with humans and cause toxic effects on them. The hazardous effects of nanoparticles on non-target organisms include dermal absorption of nanoparticles and translocation to go deep into the lungs and brain through inhalation and crossing the brain barriers, respectively.
Nanoparticles can enter various human body parts and cause harm by disrupting cellular pathways, enzymatic actions, and organ functions. Nanomaterial disposal could result in the formation of a new class of non-biodegradable pollutants in the environment [170, 171]. The potential risks associated with nanomaterials lead to the challenges of basic and applied nanobiotechnology in different fields including crop production. The major possible challenges are the following [170, 171]:
High-quality mass production of nano-based products at an economical cost.
The development of a personalized nanomaterial production system to fulfill local needs.
Environmental and human health safety and protection during the use and disposal of nanomaterials.
The challenge is to overcome the gap between basic and applied nano-based research.
The production cost, severe risks, and practical knowledge gaps are also considered as chief concerns/challenges in nanobiotechnology applications.
Hence, a collaborative effort, based on comprehensive scientific research and technological advancements, between regulatory bodies and workforce research should be made to cater to the above-mentioned challenges and associated risks. This would help to provide the necessary information to devise appropriate guidelines for comprehensive risk management and applications of nanobiotechnology for sustainable crop production.
8. Conclusion and future prospects
The impact of climatic variations in recent years has contributed significantly to reduce crop productivity and gave birth to various environmental stress factors. Abiotic stresses are the key player responsible for reduced crop yield, quality and stunted plant growth around the globe. Efforts to cater to challenges associated with abiotic stress led to the development of nanobiotechnology, which encompasses small-size nanoparticles that aid in enhancing the farming sector and managing global food demand. Several NPs, such as TiO2, SiO2, and AgNPs, can reduce the negative effects of abiotic stress by activating the genetic makeup of plant defense mechanisms via the induction of ROS production and phytotoxicity. Their small size allows them to penetrate easily into plant tissues, after which they positively influence plant morphological, physiological, and biochemical processes, promote plant development, and improve crop productivity in plants under various abiotic stresses. Furthermore, NPs have a large surface area that improves the absorption and delivery of various targeted nutrients. Despite the positive effects of nanoparticles, there are some risks and challenges associated with them that need to be addressed. These risks will aid in exploring their dimensions and help to improve crop productivity by understanding the underlying interaction pathways at the molecular level.
It is important to conduct future research exploring different uses of nanoparticles to enhance crop plant productivity. Nanoenzymes including Mn3O4-NPs gain much attention due to their potential role in mitigating stress conditions in plants. Mechanisms responsible for nanopriming-induced seed germination, breaking seed dormancy, and their interactions with seeds need to be investigated. A better understanding of plant–NP interaction will enable researchers to design tailor-made nanomaterials targeting agricultural challenges. Moreover, targeted delivery approaches using nanomaterials will prove to be another emerging way to enhance plant productivity. Recently, a study conducted with this approach used nanomaterials to convert chloroplasts into a “chloroplast factory,” resulting in improved plant photosynthesis under low light conditions [172]. In addition to that, the use of nanomaterial CRISPR-Cas genome editing in cargo delivery will also improve the efficiency of genetic engineering to enhance plant stress tolerance [173]. However, the management of biosafety hazards associated with the use of nanoparticles demands the development of strict policies and procedures so that we can utilize and benefit from their potential role in enhancing crop tolerance to abiotic stresses.
Authors contribution
All authors have contributed equally to writing and drafting this article. Hafiza Fizzah Zulfiqar also contributed to designing the images in this article.
Abbreviations
heavy metal | |
cadmium | |
lead | |
arsenic | |
nickel | |
chromium | |
cobalt | |
zinc | |
reactive oxygen species | |
nanoparticles | |
copper oxide | |
deoxyribose nucleic acid | |
ribose nucleic acid | |
ultraviolet | |
photosystem | |
titania | |
catalase | |
glutathione peroxidase | |
superoxide dismutase | |
mitogen-activated protein | |
salt overly sensitive | |
zinc oxide | |
mitogen-activated protein kinase | |
myelocytomatosis oncogene | |
auxin-repressed protein | |
ethylene responsive element binding protein | |
heat shock transcription factor | |
low-affinity cation transporter | |
natural resistance-associated macrophage protein 5 | |
nanoscale zero-valent iron | |
random amplified polymorphic DNA | |
cerium oxide |
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