Nano-Seed Priming under Heavy Metal and Abiotic Stress: A Critical Assessment of Efficacy, Toxicity Thresholds, and Application Safety

Introduction
Abiotic stresses such as drought, salinity, extreme temperatures, and heavy metal pollution are among the most serious factors limiting global crop production and threatening food security[1], [2], [3]. Although biotic stresses also contribute to yield losses, a large proportion of plant damage is indirectly linked to abiotic stress conditions that weaken plant defense systems and increase susceptibility to pathogens[4], [5]. Heavy metal contamination, largely driven by industrial and anthropogenic activities, causes significant economic losses worldwide and poses long-term risks to agricultural sustainability[6], [7]. These challenges call for further development of climate-smart and sustainable agricultural practices to improve cropping system resilience as well as increase productivity stability across variable climatic conditions [8], [9]. Thus, nanotechnology to be more specific nano-enabled seed priming (NESP) has emerged as a promising pre-sowing approach for enhancing germination and early stage seedling adaptability to stress (individually or in combination)[1], [10], [11]. The application of nanoparticles offers a small size and high surface area that enable these materials to penetrate and interact efficiently with seed tissues, thereby affecting early physiological processes [12]. Pre-Nanoparticle-based seed priming has previously been reported to increase water uptake, induce antioxidant defense systems, activate starch-degrading enzymes, and stimulate metabolic activation during germination, improving seedling quality during abiotic stress conditions [1], [13]. These effects may also extend into a cushion of immediate nutrient supply, which helps support initial development within a harsh environment. Nano-priming effects on plants, however, depend strictly on the type, concentration, size, surface charge, and solubility of nanoparticles, controlling their penetration through the seed coat and their localization in seed tissues[14], [15]. There is increasing literature reporting beneficial effects of nano-priming;however, a few major knowledge gaps remain. Lack of standardized dosage ranges among different types of nanoparticles, and cellular and molecular interactions of nanoparticles with plant tissue are not well-understood, especially after their release in soil environments [16], [17]. Moreover, nanoparticles may pose environmental risks, including  the fact that nanoparticles can lead to environmental hazards such as mean residence times in soils, bio-accumulation along the food chain, and disruption of microbial communities[1], [18],[19]. Nanoparticles can also cause oxidative stress, genotoxic effects, inhibition of enzymes, and inhibition of photosynthetic processes at high doses, making it important to define safe dose limits[1], [20]. In other words, information regarding the transport pathways and environmental fate of nanoparticles, as well as their interactions with soil microorganisms, needs to be understood before their use in agriculture is recommended at large scale [1], [16]. Addressing these issues will allow nano-enabled seed priming to be developed as a precision-based and environmentally responsible tool for improving crop establishment and stress tolerance under heavy metal and abiotic stress conditions[1].
Types of Nanoparticles Used in Seed Priming and Their Effects under Stress
Different types of nanoparticles have been tested for seed priming to improve germination and early seedling growth under heavy metal and abiotic stress. Although many studies report positive effects, the results are not always the same because plant responses depend on nanoparticle type, dose, and plant species[1], [21]:

1. Iron-Based Nanoparticles
   Iron-based nanoparticles are widely used in seed priming because iron plays a key role in plant metabolism [1]. At low to moderate concentrations, these nanoparticles improve photosynthesis, increase the activity of antioxidant enzymes, and reduce oxidative damage caused by stress. As a result, seedlings generally show better tolerance to drought, salinity, and heavy metal stress. In contrast, when iron nanoparticles are applied at higher concentrations, they may disrupt the internal redox balance of plant cells, leading to reduced growth. Therefore, the positive effects of iron nanoparticles in seed priming depend strongly on using appropriate concentrations (Figure 1) [6], [22], [23].
2. Zinc Oxide Nanoparticles
   Zincoxide nanoparticles (ZnO NPs) are commonly used because they can both reduce stress effects and supply plants with an important micronutrient. When applied at low concentrations, ZnO nanoparticles enhance antioxidant defenses, support osmotic balance (Figure 2), and improve the movement of nutrients within seedlings. These effects help promote seed germination and early growth under salinity and drought stress conditions [15], [21] , [22], [24]. However, ZnO nanoparticles also have a very narrow therapeutic index between the dosage which is beneficial and toxic [25]. High sulfate levels often cause oxidative stress, damage to the membrane, and the blocking of germination [26]. This clear dose-dependent response makes ZnO NPs efficient but also dangerous, especially in the absence of standardized priming protocols [25].
3. Titanium Dioxide Nanoparticles
   Titanium dioxide nanoparticles (TiO₂ NPs) are well known for their photocatalytic properties. When used at appropriate levels, they can improve photosynthetic efficiency and enhance energy use during the early stages of plant development (Figure 3). Research has shown that TiO₂ NPs can enhance seed germination by enhancing uptake of water and promoting the metabolism of carbohydrates [27], [28], [29]. However, at improper concentrations, TiO₂nanoparticles are also capable of causing adverse effects, such as genetic instability and metabolic disorder [30], [31]. As a result, their application in seed priming also demands strict regulation of dosage and exposure time to prevent undesirable physiological responses [26], [28].
   .
4. Silicon-Based Nanoparticles
   Silicon-The reliability in seed priming under heavy metal and abiotic stress is highest with silicon (SiO₂ NPs, silicon nanocarriers) based nanoparticles (Figure 4). Such nanoparticles improve seed hydration, increase the integrity of the cell wall, and induce antioxidant defence system [1], [32], [33]. Notably, silicon nanoparticles have been constantly found to lower the absorption as well as usage of hazardous heavy metals like cadmium and lead [34], [35].
5. Carbon-Based Nanoparticles
   Carbon-based nanomaterials such as carbon nanotubes and carbon nanodots affected seed germination primarily by increasing water absorption and oxygen permeability through seed coats (Figure 1). Which can enhance germination rate and early seedling growth of seedlings under stress conditions. On the other hand, these carbon nanomaterials also provoke concerns about their toxicity, persistence, and long-term environmental fate [36], [37]. Excessive levels or unsuitable particle size may result in cytotoxicity and reduce root growth [38]. Thus, their application in seed priming is still encouraging but should be judiciously assessed [13].
6. Biopolymer Nanoparticles (Chitosan)
   These include MWCNTs and carbon particles (Figure5). It has been reported that they can inhibit carbohydrate decomposition reactions, improve oxygen diffusion through membranes, and improve plants’ water uptake [39], [40]. While they have been shown to significantly increase germination percentages and promote the growth of plants even at high concentrations, excessive amounts can be harmful at an early developmental stage of plants [41], [42].
   Comparative Efficacy and Toxicity Assessment of Nanoparticles
   Though numerous types of nanoparticles are able to enhance seed germination and seedling development under stress, the defensible utility of such results will ultimately rest on the effectiveness-phytotoxicity balance [15]. Nanoparticle classes clearly differ with regard to safety margins and uniformity of beneficial effects, as evidenced by a comparative analysis of published studies. Among the different types of nanoparticles, silicon-based nanoparticles present the best ratio of effectiveness(Table 2): safety,regularly enhancing germination, stress tolerance, and the accumulation of heavy metals across several crop species [22], [43], [44]. Iron nanoparticles also appear as promising, but necessitate a tighter control of the dose to prevent a redox imbalance. In contrast, zinc oxide, silver, and titanium dioxide nanoparticles are believed to have lower safety windows with only small changes in concentration inducing a shift in plant response toward stimulation or toxicity [22], [30], [45], [46]. While carbon-based nanomaterials may provide rapid physiological benefits within the timeframe of germination, their potential for persistence in the environment and long-term ecological impact is far from resolved. In fact, biopolymer nanoparticles, especially chitosan, are a middle ground because they are biodegradable and can induce stress in plants. But have the proper dosage if they just work. In contrast, this comparison emphasizes that nano-seed priming should not be regarded as a homogeneous technology since the performances are dependent on the kind of nanoparticle used and/or application conditions [47], [48], [49].Consequently, nanoparticle choice and anticipated agronomic setting should both be material-specific performance and toxicity-inspired thresholds and limitations [13], [38]. These examples of a comparative framework enable the engineering of seed priming methods to maximize benefits to a target species while minimizing undesirable environmental and physiological side effects [13], [15], [50].
   A General Approach
   This review is an outcome of the scrutinized investigation of all available studies on the application of nanoparticles as a seed priming material under the impact of heavy metal and abiotic stress [51], [52]. Available studies were compared based on nanoparticle type, dosage, and seed treatment approach. We restricted analyses to studies that measured physiological responses to stress-wave responses and that reported changes in uptake. [1], [32], [38](Table 3).
   Entry of Nanoparticles into Seeds and Their Interaction with Plant Cells
   The success of nano-enabled seed priming as a paradigm goes beyond growth stimulation, but also entails the capability of nanoparticles to cross the outer seed layers, move inside the seeds, and interact with plant tissues.Studies show that nanoparticles are able to pass through natural micropores in the seed coat or enter cells via active transport mechanisms, thereby reaching the embryo. After being taken into the seed, nanoparticles could then affect the development of the embryo or the subsequent growth of the seedling [1], [51].
   Once in plant cells, nanoparticles can interfere with essential metabolic processes through the following means:
   ● Controlling levels of reactive oxygen species (ROS) with regulatory potential [1].
   ● Activating or inhibiting specific enzymes [1].
   ● Improving germination and growth-related pathways [1].
   ● Modifying hormone-related stress response pathways [1].
   The size, chemical composition, and surface characteristics of nanoparticles, and the duration of their retention on the seed surface, govern the transport of nanoparticles inside the seed and thus their behavior during seed priming. There havebeen some advances in this area, but the mechanisms underlying seed priming with nanoparticles remain elusive. This will provide a better insight into these processes that’ll help in the development of effective and safe priming strategies [13].
   Characterization of Nanoparticles
   A fundamental step in knowing how nanoparticles behave in the short and long run during seed priming and planting is the characterization of these nanoparticles due to their physicochemical properties. Other extensively studied features are particle size and shape, zeta potential, solubility, ion release rate, and crystal structure[53]. A range of approaches is implemented to examine these characteristics, including Transmission Electron Microscopy (TEM), Scanning Electron Microscopy (SEM) [54], [55].Fitted with Energy-Dispersive X-ray Spectroscopy (SEM-EDS) and spectroscopic techniques. Additionally, the elemental composition of a plant can be measured using methods such as X-ray Fluorescence (XRF), which can also track the movement of nanoparticles through various plant tissues. Meaningful Knowledge: The Aim to Bridge Between Nanoparticle Properties and Biological Effects.These Tools Establish Links Between NP Properties and Effects [17]
   Physiological and Biochemical Indicators Used in Reviewed Studies
   In the reviewed studies, different physiological and biochemical indicators were employed to assess the impacts of nano-seed priming [38], [56]. Physiological parameters consisted of germination percent, root and shoot length, seedling dry weight, seedling vigour index, photosynthetic efficiency, and stomatal behavior [57], [58]. Biochemical responses were primarily determined as the activity of antioxidant enzymes (catalase, peroxidase, and superoxide dismutase), and malondialdehyde (MDA) levels as an indicator of oxidation damage [59], [60]. Glucosemia resistance, in addition to changes in absolute sugar and protein content, changes in the expression of genes associated with stress responses were also reported in many studies. [61].
   Role of Carbon-Based Nanomaterials
   Carbon-based nanomaterials, especially carbon nanotubes, have gained the attention of researchers, and their impact on seed physiology is very prominent; numerous studies have been reported on the topic [37], [62]. As mentioned [8], these materials have reported to decrease hydrogen peroxide accumulation, serve as membrane protectant and promote water uptake during germination. Thus, it was reported that carbon nanomaterials could cause root injury by damaging root cell membranes at very high concentrations under extreme conditions [1], [63]. In summary, the application of carbon-based nanomaterials has great potential for enhancing early growth stress tolerance, while the negative effects of the application should be avoided by careful dose control [37], [64].
   Molecular and Hormonal Regulation
   Several studies have shown that seed priming with nanoparticles can influence molecular and hormonal pathways involved in germination and early seedling growth[21], [52][1].These effects include activation of genes related to nutrient uptake, stimulation of antioxidant defense systems, and regulation of key plant hormones such as gibberellins and abscisic acid [1], [65]. When nanoparticles are able to pass through the seed coat and reach the embryo, they often accelerate the shift from dormancy to active growth. This process is commonly associated with improved photosynthetic performance and increased biomass production under stress conditions [1], [51].
   Methodological Summary
   These methodologies indicate that nano-seed priming generally improved germination, early growth, enzymatic activity, and stress tolerance[1], [13]. Nevertheless, such effects are highly particle-type-, particle concentration- and surface property-dependent [66]. To make sure that the outcomes are reproducible and environmentally safe, experimental design and detailed nanoparticle characterization arerequired [67]. More importantly, long-term studies are necessary to clarify the environmental destinies of the nanoparticles and to utilize them sustainably in agriculture [68], [69].
   Synthesis of Evidence
   Our review of the findings of all the studies in this domain shows that nano-seed priming can have a positive effect on the seed germination and early seedling growth under heavy metal and abiotic stress, but this response is not uniform for all the conditions tested. This strategy relies on the nature of the nanoparticle, the applied dose, and the type of plants[21], [52], [70]. This shows that nano-seed priming is not anyone’s magic bullet, as its effects only occur within a defined degree of existence[38]. One of the most clear patterns seen across many studies is a dose-response. At lower concentrations, nanoparticles frequently increase physiological and biochemical factors (e.g., antioxidant activity, water uptake, and stress signalling) during the germination stage and increase seedling vigor. On the other hand, at high concentrations, they often induce adverse effects, such as oxidative stress, membrane damage, and inhibited growth[71], [72], [73]. While this pattern receives widespread reportage, there is scant delineation of a safe dose threshold in most studies, which renders them less than practically useful[74]. Nanoparticle types also differ from each other. Silicon-based and biopolymer nanoparticles usually have a more stable and safer response, whereas the safety range for metal and metal-oxide nanoparticles is narrow, so that a minor dose adjustment can turn the plant response from beneficial to detrimental[17], [72], [75]. While carbon-based nanomaterials can promote early physiological reactions, we still do not know if they persist in plants or ecosystems. Another important limitation of authors in the available literature is the lack of standardization for the various existing definitions[17], [32], [76]. Comparison of results among different studies is not feasible due to differences in nanoparticle preparation and characterization techniques, as well as in seed treatment methods and parameters measured[1], [62]. Moreover, many findings havebeen based on short-range laboratory experiments and may not reflect field conditions or longer-term environmental interactions. In conclusion, our evidence suggests that nano-seed priming is a precise technique rather than a general agronomic practice[38], [77]. Before large-scale employment is recommended that the safe dose ranges must be defined, and greatly standardized experimental protocols applied, as well as the plant response and environmental impacts must be better understood.[1], [17], [38]
   Conclusion
   In spite of this, nano-enabled seed priming to ameliorate seed germination and early seedling establishment of heavy metal as well as other abiotic stress-affected seeds holds great promise, but is still subject to major challenges because of their highly context-dependent beneficiary exposure systems [38], [51], [78]. The current review synthesis revealed that the positive effects of nano-priming are primarily under the control of three main factors: nanoparticle type, dose, and plant genotype [52], [78].Among all the materials characterized, silicon-based NPs have reliably provided the most reliable balance between relieving and inducing stress by enhancing antioxidant defenses and suppressing metal translocation [22], [44].For comparison notes, nanostructures based on metals and metal oxides, such as ZnO, Ag, and TiO₂, have a narrow therapeutic index: low concentration gradients can shift the biological response from stimulation to toxicity [20], [22][22].Conclusions: Nano-seed priming is not a magic bullet, agronomically based solution, but a precision-based solution. This requires conducting the relevant due diligence on dosage refinement, the application of standardized characterization protocols, and long-term environmental risk assessment prior to widespread agronomic release [1], [79].

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