Nano priming effects on seed vigour and disease resistance in cereal crops

Introduction

Researches have shown that biochemical, physiological, and morphological processes are involved in plant systems (Kottegoda et al., 2017) [22] . When a boom in agricultural output is achieved, nanotechnology has boosted a number of plant life nodes, including seed germination, plant fresh and dry weight gain, reduction of pest and insect incidence, and activation of antioxidant systems (Haghighi et al., 2012; Siddiqui and Al-Whaibi, 2014; Wang et al., 2015, 2016a, b; Shalaby et al., 2016) [12,21] . Nanotechnology has the potential to enhance green farming methods and boost food security by allowing for easier penetration into seeds. Nanotechnology has several benefits for plants, including increased reactivity, precise targeting, and the ability to encapsulate nutrients and plant protection products for long-term release and resistance (Ocsoy et al., 2013; Anusuya and Sathiyabama, 2015; Corsi et al., 2018) [27] . Eco-friendly use of nanotechnology in agriculture could contribute to sustainable development. To assure safe usage of nanotechnology, confocal microscopy and biochemical studies of plant-nanomaterial interactions are conducted in vivo using quantum dots (Wu et al., 2017) [10] .

Seed Priming and its Effects in Agriculture

Seed is the main input used in agricultural production, hence, ensuring its quality is crucial to increasing crop productivity. According To plant involves a complex collection of responses and physiology that determine agricultural field stand. The ancient Greek civilization used seed priming (SP), a physiological and biochemical seed invigoration technique (Evenari, 1980) [11] , to quickly transition to full germination. This process involves starch hydrolysis, osmotic adjustments, and enzymatic activation mediated by imbibition (Araújo et al., 2016; Hussain et al., 2015, 2016) [5, 14] , which activates and strengthens the plant's cellular defense system against various stresses (Jisha et al., 2013) [16] . The most important requirement for sufficient crop growth that guarantees the effective use of input resources that define the final yield potential and the fundamentals of a sustainable agricultural economy is quick germination and a similar crop stand in the field (Bose et al., 2018) [6] . According to Bose et al. (2018) [6] , Singhal et al., and Kumar et al., SP is a well-known, inexpensive, readily adaptable, and highly effective method for improving germination and crop vigor that ensures better yield of significant agricultural products under both normal and stressful conditions. Partially hydrating and dehydrating seeds without allowing radicle initiation is known as SP, a pre-sowing technique. By soaking in water or other priming agents under carefully regulated conditions, this advancement of seeds (hydration) starts the premetabolic activity (Bose et al., 2018) [6] . The length of the soaking period and the reaction's termination have a significant impact on this treatment. For SP treatment, which establishes the seed's storability and metabolic activities, this treatment adhered to a set duration for soaking and constantly quick-drying to their initial moisture level. [6] . Due to the hydration and rehydration process, the nanopriming phase II takes longer than the typical seed germination process. There are two hydration processes in primed seed and only one in control. While seedling growth is inhibited during the drying process, metabolic activity which is essential for metabolite synthesis, antioxidant activity, solubilization of stored material, priming and stress memory, and repair mechanisms remains active. It improves climate acclimation, stress tolerance, and quick and consistent seedling establishment. Priming agents are the substances that are most frequently employed in priming treatments. They might include water, inorganic chemical salts, hormones, sugars, vital nutrients, plant organic sources, water or nutrients, biofertilizer, and other helpful microorganisms (Jisha et al., 2013) [16] . SP has been classified on the basis of priming agents likehydropriming (priming with water), osmopriming (priming with osmotic salts like poly-ethylene glycol (PEG), KNO3), matrix priming (priming with solid and semisolid matrices), nutripriming (priming with plant nutrients like micronutrients and macronutrients), biopriming (priming with biological compounds), hormopriming (priming with plant growth regulators (PGRs)), and nanopriming (priming with carbon or metal-based nanomaterials). Primed seeds exhibit increased crop establishment rates, enhanced seedling vigor, and greater rates of seedling emergence (Singhal and Bose, 2020). Additionally, SP helped developing seedlings overcome the detrimental effects of environmental challenges and increase their stress tolerance (Bose et al., 2018) [6] . Additionally, because SP reduces the usage of chemicals on agricultural land, it addresses a variety of agricultural pollution issues, including heavy metal toxicity. These days, using NPs for SP has proven beneficial in the aforementioned areas (Zuverza-Mena et al., 2017) [41] . By promoting cellular metabolic activities, water absorption, cell molecule and osmolyte repair, and other processes, priming seeds improves their physiological and growth performance as well as their potential for production (Canfield et al., 2010) [7] .

Nanopriming and Nanomaterials Used in Seed Priming

Numerous innovative applications in biotechnology and agriculture made possible by nanotechnology improve soil health, lower production inputs, and encourage sustainability in the agricultural system. Additionally, it is assisting other agricultural-related businesses such food processing and packaging, the creation of a biosensor for pathogen detection, the manufacturing of nanofoods, and nanofertilizers (Dehkourdi and Mosavi, 2013). Leaching, hydrolysis, degradation, and decomposition processes prevent plants from consuming nutrients in bulk, so it's critical to minimize these losses by utilizing this cuttingedge technology. NPs have the ability to increase plant metabolism because of their unique characteristics, which include increased surface area, high reactivity, tenable pore size, and particle structure. According to Wang et al. (2016a, b) [37,38] , nanoparticles are particulate nanomaterials with a nanoscale size of at least 1 to 100 nm. The two forms of NP that have been used as SP agents are metal-based (Zuverza-Mena et al., 2017) [41] and carbon-based (such as carbon nanotubes) (Mahakham et al., 2017) [24] . In significant crops like wheat and maize, carbon nanotube priming can boost germination, growth, water/nutrient uptake, and yield (Tiwari et al., 2014; Joshi et al., 2018) [35] . Compared to other materials, nanoparticles like zinc oxide nanoparticles (nZnO), iron nanoparticles (nFe), copper nanoparticles (nCu), silicon nanoparticles (nSi), and magnesium nanoparticles (nMg) can be quickly absorbed, transported, and accumulated in plant parts. These nanoparticles control the physiological and biochemical processes of plants, resulting in improved plant growth and development (Rastogi et al., 2017) [30] . However, the NP's effectiveness varies from plant to plant depending on its particle size, concentration, chemical structure, and surface coating (Rastogi et al., 2017) [30] . NPs are used as SP agents because they are affordable, nontoxic, biocompatible, and environmentally benign. Crop germination, seedling establishment, shoot/root elongation, fresh and dry weight, growth, chlorophyll content, photosynthetic activity, water uptake, biomass production, flowering, and antioxidant qualities can all be affected by various nanopriming agents, including silver nanopriming (AgNP) and nanotitanium dioxide (TiO2) (Dehkourdi and Mosavi, 2013). The feasibility of using nanoprimed seeds with micronutrient nanoparticles for sustainability has recently been investigated by scientists. Increased root characteristics like root length and root volume (Adhikari et al., 2013) [1] , increased drought tolerance, increased micronutrient retention in the leaves (Anita and Rao, 2014) [4] , increased antioxidant activity followed by increased jasmonate, decreased time to flowering and increased seed weight. According to Mahakham et al. (2017) [24] , aged rice seeds primed with AgNPs greatly enhanced germination performance, seedling vigor, and α-amylase activity, which guarantees more soluble sugar to support the growth of newly emerged seedlings. By interacting with the sulfhydryl and COOH-groups of proteins, metal nanoparticles reduce the efficiency of enzymes (Shaw et al., 2014) [31] . In many crops, nanopriming also increases stress tolerance. Nano-ZnO priming with the concentration 60 mg L21 improved salinity tolerance by reducing the malondialdehyde (MDA) and Na1 concentrations in Lupinus termis but reduce sodium concentration and increased leaf water potential in wheat (Wang et al., 2016a, b) [37,38] . According to Ali et al. (2019) [2] , these nanoparticles can also reduce the buildup of other heavy metals in crops like rice. According to Mahakham et al. (2017) [24] , nano priming promoted the induction of aquaporin genes and reactive oxygen species (ROS) generation in germinated seeds, both of which are crucial for improving seed germination. In spinach seeds, iron pyrite (FeS2/Py) priming enhanced the quantity of leaves, leaf size, chlorophyll, and mineral content (Ca, Mn, and Zn) (Srivastava et al., 2014). Additionally, it increased grain output in rice, bio mass production in fenugreek and alfalfa, and the seed yield of oilseed crops like sesame and mustard. Additionally, defensive responses connected to jasmonates can be induced by nanoiron oxide. By increasing the activities of sodium dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX), nano-ZnO priming affected an efficient ROS scavenging system that maintained cellular homeostasis and increased the activity of phosphoglucomutase and cytoplasmic invertase enzymes involved in sucrose biosynthesis under salinity. A novel approach to iron biofortification in wheat was suggested by Nano SP with iron oxide (25-600 ppm), which increases iron absorption, deposition, and translocation in plants as well as iron buildup in grain (Sundaria et al., 2019) [32] . According to Munir et al. (2018) [25] , ZnO nanopriming also raises Zn concentration in wheat. According to Rizwan et al. (2019) [14] , iron oxide and zinc oxide both enhance plant development and lessen oxidative damage during cadmium stress. Nanotechnology's shining future in precision agriculture has the potential to usher in a second sustainable green revolution. [16] . This involves increasing amylase activity to produce more soluble sugars, as documented in rice (Sharma et al., 2021), canola (Khan et al., 2024) [20] , and chickpea seeds (Das and Dutta, 2022) [9] . Nanopriming regulates the balance of abscisic acid (ABA) and gibberellin (GA) by increasing the production of GA and cytokinin-related genes while decreasing ABA levels (Abdel-Aziz and Rizwan, 2019) [14] . A study found that priming rice seeds with selenium (Se) and salicylic acid (SA) at low temperatures increased GA3amylase activity, content, and expression of OsGA3ox1 and OsGA20ox1. There is a decrease in both ABA content and expression of OsNCED1. Induction enhances GA-mediated α-amylase by raising GA3 levels while lowering ABA expression (Nie et al., 2022) [26] . The current work did not explore whether nanopriming uses the same GA signaling mechanism. Nanoparticles need to get through the seed coat, which is made up of physicochemically stable sclerenchyma. According to Schwab et al., this structure works as a barrier to protect seeds from external environmental threats. Zinc oxide nanoparticles (ZnO NPs) and titanium dioxide nanoparticles (nano-TiO 2) have been proven in experiments to penetrate seed coats and induce cell differentiation by triggering enzymes that break down seed dormancy. Seed priming increases hydroxyl radical (•OH) production, weakening seed cell walls and improving germination rates (Mahakham et al., 2017) [24] . Nanoparticles deliver critical nutrients to plant cells, including zinc, iron, potassium, and calcium. These nutrients boost plant chlorophyll content, speed up photosynthesis, and promote leaf growth, leading to higher yields. Figure 4. illustrates the intrinsic mechanism of nanopriming on plant cells.

B. Nanopriming promotes the expression of genes involved in seed imbibition, plant defense, and antioxidation

Nanopriming does not alter the plant's phenology or genomics. Johnson and Puthur (2021) [17] found that stresstolerant memory is sustained by transcription, translation, and epigenetic mechanisms. This molecular change allows plants to respond more quickly and efficiently to future stressors. Plants produce reactive oxygen species (ROS) that signal other messengers like ethylene (ET), salicylic acid (SA), nitric oxide (NO), jasmonic acid (JA), and oleoresinol stanols (BRs). These messengers can regulate secondary metabolism (Chung et al., 2018) [8] . Signaling proteins and transcription factors have a crucial role in maintaining "stress memory". Priming-induced genes can be regulated at various levels, including RNA-induced transcription silencing, RNA-dependent DNA methylation (RdDM), changes in nucleosome or histone variant positioning, DNA methylation, and histone modifications. Yan et al. ( 2022) [42] discovered that priming rice with nanosilver (Ag NPs) raised reactive oxygen species (ROS) levels and activated hormone signaling pathways. This priming also increased the expression of APETALA2 (AP2) domain-containing protein, dof zinc finger domaincontaining protein, and WRKY. Transcription factors increase protection by regulating the genes of defenserelated proteins such as HSP, GST, FBP, and LEA. They also control glutathione and cytokinin-O-glucosyltransferase (CGTs) to enhance seed germination. During the suckering stage of seed germination, the seed absorbs significant amounts of water, which can activate aquaporin genes (Salam et al., 2022a). El-Badri et al. ( 2022) [10] found that selenium nanoparticles (Se NPs) can boost the expression of aquaporin genes (BnPIP1-1 and BnPIP2-1), leading to increased water absorption. The researchers examined the impact of zinc oxide nanoparticles (ZnO NPs) on two rapeseed cultivars. El-Badri et al. (2021) [43] discovered that ZnO NPs increased seed germination by upregulating BnCAM and affecting the seed imbibition gene BnPER. Nanopriming induces moderate ROS generation, which improves plant resilience to abiotic stress. This promotes the synthesis of secondary metabolites. Nile et al. suggest that serial analysis of gene expression (SAGE) can help understand the impact of nanoparticles on plants. Ulhassan et al. (2023) [36] discovered that silicon nanoparticles (Si NPs) can increase chlorophyll synthesis genes (BnCAO, BnCHLG, BnPOR), inhibit the expression of senescence associated genes (BnSAG12), and increase the expression of defense genes (BnPAL, BnCAD, BnPPO, BnMT-1), reducing chlorophyll degradation and leaf senescence under chromium stress. The literature reveals that nano-priming approaches have favourable effects by regulating ROS and enhancing the enzymatic antioxidant system. Non-enzymatic antioxidants work alongside enzymatic antioxidants to regulate ROS levels. This could explain why their levels spiked when antioxidant defenses were compromised after a hazardous event. The molecular mechanisms of enzymatic and nonenzymatic antioxidant systems regulating ROS levels in response to NPs remain unclear. Figure 5 illustrates how nano-priming modulates antioxidant defenses and regulates ROS levels. Seed priming with nanoparticles has been shown to improve plant defense against fungal, bacterial, and nematode pathogens. For example, in cumin, priming with chitosan and silver nanoparticles enhanced antioxidant enzyme activities (SOD, CAT, peroxidase, and polyphenol oxidase), thereby reducing Fusarium wilt. Similarly, selenium nanoparticle-primed tomato seeds exhibited strong resistance to late blight through increased accumulation of lignin, callose, and hydrogen peroxide, along with elevated levels of defense-related enzymes such as SOD, LOX, β-1,3-glucanase, and PAL. Pathogen infection in tomato plants by organisms like Xanthomonas campestris, Pseudomonas syringae, Alternaria solani, and Fusarium oxysporum negatively affected plant growth and biochemical parameters, including pigments and enzyme activities. However, seed priming with titanium dioxide nanoparticles improved growth, restored biochemical attributes, and reduced disease severity. Likewise, silicon dioxide nanoparticles were effective under greenhouse conditions in reducing diseases caused by pathogens such as Pectobacterium, Meloidogyne, and Rhizoctonia, while enhancing plant growth, photosynthesis, and defense enzyme activities, ultimately lowering disease incidence and nematode proliferation. Field studies further demonstrated that advanced nanoparticle formulations, such as triazole-conjugated silver nanoparticles, reduced the incidence of bakanae disease in rice. However, the comparative effectiveness of seed priming versus foliar application of nanomaterials remains inconsistent across studies.

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