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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2025.1626624</article-id>
<article-version article-version-type="Corrected Version of Record" vocab="NISO-RP-8-2008"/>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Review</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Crop management to enhance plant resilience to abiotic stress using nanotechnology: towards more efficient and sustainable agriculture</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Al-Dossary</surname><given-names>Othman</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/3219317/overview"/>
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<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Alnaddaf</surname><given-names>Lina M.</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/3063463/overview"/>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Al-Khayri</surname><given-names>Jameel M.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
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<aff id="aff1"><label>1</label><institution>Department of Agricultural Biotechnology, College of Agriculture and Food Sciences, King Faisal University</institution>, <city>Al-Ahsa</city>,&#xa0;<country country="sa">Saudi Arabia</country></aff>
<aff id="aff2"><label>2</label><institution>Department of Field Crops, College of Agriculture, Homs University</institution>, <city>Homs</city>,&#xa0;<country country="check-value">Syria</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: Othman Al-Dossary, <email xlink:href="mailto:othmand@kfu.edu.sa">othmand@kfu.edu.sa</email>; Lina M. Alnaddaf, <email xlink:href="mailto:hatem005@gmail.com">hatem005@gmail.com</email>; Jameel M. Al-Khayri, <email xlink:href="mailto:jkhayri@kfu.edu.sa">jkhayri@kfu.edu.sa</email></corresp>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2025-10-14">
<day>14</day>
<month>10</month>
<year>2025</year>
</pub-date>
<pub-date publication-format="electronic" date-type="corrected" iso-8601-date="2025-12-04">
<day>04</day>
<month>12</month>
<year>2025</year></pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1626624</elocation-id>
<history>
<date date-type="received">
<day>11</day>
<month>05</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>09</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Al-Dossary, Alnaddaf and Al-Khayri.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Al-Dossary, Alnaddaf and Al-Khayri</copyright-holder>
<license>
<ali:license_ref start_date="2025-10-14">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p>
</license>
</permissions>
<abstract>
<p>Nanotechnology has become a transformative tool in modern agriculture, playing a pivotal role in enhancing crop resilience to abiotic stresses, including drought, salinity, and extreme temperatures. As global population growth and environmental challenges place increasing pressure on agricultural systems, nanotechnology plays a crucial role in enhancing crop yields and ensuring long-term sustainability. Nanotechnology, through advanced applications, optimizes nutrient delivery, strengthens plant defense mechanisms, and enables precise monitoring of environmental conditions. These innovations enhance soil quality, regulate physiological responses in plants, and mitigate the adverse effects of environmental stressors, thereby promoting sustainable farming practices and improving food production efficiency. Nanoparticles (NPs), synthesized through green methods using plant or microbial extracts, have shown promise in enhancing stress tolerance by facilitating uptake, translocation, and intracellular movement within plants. Major factors influencing NPs efficacy include size, concentration, composition, and duration of exposure. Biosensors and nanobiosensors provide prognostic tools for real-time detection and management of plant stress. Despite their potential benefits, the use of nanotechnology in agriculture raises concerns regarding environmental and health impacts. The accumulation of NPs in soil and aquatic ecosystems may affect microbial diversity, disrupt soil enzymatic activity, and alter plant&#x2013;microbe interactions, posing risks to non-target organisms and overall ecosystem health. Moreover, variability in plant responses to NPs complicates the development of standardized application protocols. Therefore, its successful adoption relies on sustained interdisciplinary research, ethical oversight, and the development of sound policy frameworks. This integrated approach is crucial for developing resilient, efficient, and sustainable agricultural systems that can meet future challenges.</p>
</abstract>
<kwd-group>
<kwd>crop management</kwd>
<kwd>signaling pathways</kwd>
<kwd>nanotechnology</kwd>
<kwd>abiotic stress</kwd>
<kwd>sustainable agriculture</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declare financial support was received for the research, and/or publication of this article. This work was supported by the Deanship of Scientific Research, Vice Presidency for Graduate Studies and Scientific Research, King Faisal University, Saudi Arabia [Grant No. KFU253401].</funding-statement>
</funding-group>
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<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Plant Abiotic Stress</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Global food security faces unprecedented challenges, as the world population is projected to reach 10 billion by 2050, with nearly two billion people currently suffering from nutrient deficiencies and approximately eight hundred million experiencing chronic hunger. Nanotechnology emerges as a revolutionary approach to address these critical challenges, offering innovative solutions that could transform agricultural practices and enhance food production efficiency at the molecular level (<xref ref-type="bibr" rid="B15">Atanda et&#xa0;al., 2025</xref>).</p>
<p>Abiotic stresses caused by non-biological environmental factors, including salinity, drought, extreme temperatures, and heavy metal contamination, significantly impact crop yield and global nutritional security, resulting in 20&#x2013;50% annual global crop yield losses (<xref ref-type="bibr" rid="B20">Cao et&#xa0;al., 2025</xref>). Traditional agricultural practices have proven inadequate in effectively mitigating these mounting challenges, necessitating the exploration of cutting-edge technological interventions (<xref ref-type="bibr" rid="B163">Tortella et&#xa0;al., 2023</xref>).</p>
<p>The integration of nanotechnology into agriculture represents a paradigm shift, leveraging the unique physicochemical properties of nanomaterials to regulate plant physiological processes, the development of nanofertilizers, nanopesticides, and nanosensors that can optimize nutrient delivery, strengthen plant defense mechanisms, and enable real-time environmental monitoring for sustainable farming practices (<xref ref-type="bibr" rid="B13">Alnaddaf et&#xa0;al., 2025</xref>).</p>
<p>Nanoparticles can be strategically introduced into plants through diverse application methods, including seed coating, root uptake, and foliar spraying (<xref ref-type="bibr" rid="B134">Semida et&#xa0;al., 2021</xref>). Once internalized, these particles translocate through the vascular system and intercellular spaces to various plant organs, with their uptake and distribution patterns varying based on nanoparticle type and plant species (<xref ref-type="bibr" rid="B187">Zaman et&#xa0;al., 2025</xref>).</p>
<p>This targeted delivery system presents a fundamental advantage over conventional agricultural inputs, enabling controlled nutrient release with minimal quantities while maximizing crop yield and reducing environmental impact (<xref ref-type="bibr" rid="B43">Gupta et&#xa0;al., 2023</xref>).</p>
<p>Under stress conditions, particularly drought, nanoparticles play crucial roles in maintaining nutritional balance by promoting the absorption, transport, and delivery of essential nutrients such as manganese (Mn), nitrogen (N), zinc (Zn), and potassium (K) within plant tissues (<xref ref-type="bibr" rid="B4">Ahmad et&#xa0;al., 2025</xref>).</p>
<p>The transformative potential of nanotechnology extends beyond mere nutrient delivery. Simultaneously, minimizing stress-induced damage by improving soil quality and enhancing stress resistance mechanisms (<xref ref-type="bibr" rid="B144">Shoukat et&#xa0;al., 2025</xref>).</p>
<sec id="s1_1">
<label>1.1</label>
<title>Mechanisms of abiotic stress response in plants</title>
<p>Plants perceive abiotic stress via specific sensors located at the cell wall, plasma membrane, cytoplasm, mitochondria, chloroplasts, and other organelles. This stress perception leads to signal transduction pathways involving secondary messengers such as calcium ionsA (Ca<sup>+2</sup>), reactive oxygen species (ROS), and protein kinases, which amplify the stress signal throughout the plant. In addition, stress conditions alter the balance of endogenous plant hormones (<xref ref-type="bibr" rid="B3">Abdelkader et&#xa0;al., 2023</xref>). Abscisic acid (ABA) is especially essential for responses to drought and salinity, often mediating stomatal closure to prevent water loss (<xref ref-type="bibr" rid="B33">Francis et&#xa0;al., 2024</xref>). Other hormones, such as jasmonic acid and salicylic acid, also play distinct and sometimes combinatorial roles in stress adaptation. Moreover, a network of transcription factors (TFs), including NF-Y, WOX, WRKY, bZIP, and NAC, regulates stress-responsive genes, enabling rapid and targeted genomic adaptation. MicroRNAs (miRNAs) and non-coding RNAs help fine-tune gene expression in stressful environments (<xref ref-type="bibr" rid="B58">Ju&#xe1;rez-Maldonado, 2023</xref>). Epigenetic changes, including DNA methylation and histone modification, modulate genes that enable stress tolerance. In addition, abiotic stress causes an accumulation of ROS, which can damage plant cells. Plants deploy enzymatic (e.g., superoxide dismutase, catalase, ascorbate peroxidase) and non-enzymatic (e.g., ascorbate, glutathione) antioxidants to neutralize ROS and restore cellular equilibrium (<xref ref-type="bibr" rid="B57">J&#xf3;sko et&#xa0;al., 2021</xref>). As a result of being exposed to various stress factors, plants resort to a set of metabolic and physiological adjustments. These adjustments include modifying nutrient uptake and distribution to maintain a balance of carbon, nitrogen, and phosphorus. Although this balance occurred under prolonged stress, it affects growth and yield. Other adaptive strategies include accumulation of osmolytes (such as proline and sugars), modification of cell membranes, and protein stabilization processes (<xref ref-type="bibr" rid="B62">Khan et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s1_2">
<label>1.2</label>
<title>Plant dynamics of abiotic stress</title>
<p>Plants can temporarily repress growth to conserve energy or reallocate resources. These changes may be reversible when stress is alleviated, or they may lead to permanent alterations in development if the stress is prolonged. Plants often face multiple stresses simultaneously, and the interaction between stress pathways can produce unique physiological responses that differ from single-stress reactions (<xref ref-type="bibr" rid="B115">Raliya et&#xa0;al., 2015</xref>). Crosstalk between hormonal pathways is central to coordinating these complex responses. Under stress, plants frequently prioritize survival mechanisms over growth, which can result in diminished yield but increased resilience. Plants&#x2019; ability to sense, transduce, and respond to abiotic stress is vital for survival and agricultural productivity, especially in the context of climate change and global food security. Understanding and enhancing these mechanisms remain a key focus of plant science and crop improvement research (<xref ref-type="bibr" rid="B161">Thiruvengadam et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s1_3">
<label>1.3</label>
<title>The relationship between nanotechnology and plant abiotic stresses</title>
<p>Nanotechnology and plant stress are closely connected through the development and application of nanoscale materials and sensors that help detect, manage, and enhance plant tolerance to abiotic stress via two main points:</p>
<p>Detection and Monitoring: Nano-enabled biosensors and nanobiosensors allow the rapid, sensitive, and real-time detection of plant stress signals, including disease biomarkers, toxins, heavy metals, or stress-related signaling molecules, well before visible symptoms appear. These sensors use advanced nanomaterials (like carbon nanotubes, metal nanoparticles) to greatly improve the precision and speed of stress detection, thus enabling early intervention and smarter crop management (<xref ref-type="bibr" rid="B39">Goyal and Mehrotra, 2025</xref>).</p>
<p>Stress Mitigation and Tolerance: Nanomaterials, including green-synthesized nanoparticles of metals such as silver, copper, and zinc, can be delivered to plants to help mitigate effects. Nanoparticles can scavenge harmful reactive oxygen species (ROS) generated during stress, act as stress signaling inducers (boosting a plant&#x2019;s defenses), and deliver nutrients or protective compounds more efficiently than conventional forms. This can increase tolerance to environmental stresses and reduce crop yield losses (<xref ref-type="bibr" rid="B187">Zaman et&#xa0;al., 2025</xref>).</p>
<p>By integrating nanotechnology into agriculture, it is possible to both monitor plant health dynamically and intervene more effectively to raise plant resilience against stress factors. Nanotechnology thus acts as both a diagnostic tool and a direct aid in plant stress management, supporting sustainable and productive agriculture. However, issues with nanomaterial stability, cost, and potential environmental impact remain areas of continued research and debate (<xref ref-type="bibr" rid="B156">Suresh Kumar et&#xa0;al., 2025</xref>).</p>
<p>Nanotechnology has a close and growing relationship with abiotic stress management in plants, offering novel tools to detect, mitigate, and enhance tolerance to some non-biological stresses such as drought, salinity, extreme temperature, and heavy metal toxicity (<xref ref-type="bibr" rid="B187">Zaman et&#xa0;al., 2025</xref>).</p>
<p>This review comprehensively examines the role of nanotechnology in enhancing plant tolerance to abiotic stresses, addresses key research gaps in the field, and evaluates the broader implications of this emerging technology for achieving more efficient, sustainable, and resilient agricultural systems in the face of global food security challenges.</p>
</sec>
</sec>
<sec id="s2">
<label>2</label>
<title>Nanotechnology fundamentals</title>
<sec id="s2_1">
<label>2.1</label>
<title>Types of nanoparticles</title>
<p>Nanoparticles (NPs) are recognized for improving nutrient availability and enhancing plant resistance to pathogens and environmental stresses. Various classes of nanomaterials show promise in agricultural applications, including inorganic materials (e.g., silica, copper, iron, zinc, and selenium), organic materials (such as biopolymers, lipids, chitosan, peptides, and proteins), and hybrid materials. Laboratory-synthesized nanoagrochemicals, including nanofertilizers and nanopesticides, have demonstrated a performance increase of approximately 20&#x2013;30% compared to traditional products (<xref ref-type="bibr" rid="B165">Verma et&#xa0;al., 2024</xref>).</p>
<p>Metal and metal oxide nanoparticles, such as silver and zinc nanoparticles, are extensively studied for their effects on secondary metabolism and plant growth. Specifically, zinc oxide nanoparticles (ZnO NPs) have been shown to enhance biomass accumulation and photosynthesis in Arabidopsis (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>) (<xref ref-type="bibr" rid="B69">Lala, 2021</xref>). These nanoparticles promote growth and strengthen plant resilience against abiotic stresses such as drought, cadmium exposure, and salinity. Zinc plays a critical role in the function and stability of various enzymes, supporting the development of healthy crops (<xref ref-type="bibr" rid="B165">Verma et&#xa0;al., 2024</xref>). Studies have demonstrated that ZnO NPs significantly improve rice germination rates under salt stress and increase drought tolerance in Kotschy&#x2019;s dragon head through positive effects on physiological and biochemical traits (<xref ref-type="bibr" rid="B144">Shoukat et&#xa0;al., 2025</xref>; <xref ref-type="bibr" rid="B141">Shelar et&#xa0;al., 2024</xref>). Furthermore, foliar application of ZnO NPs has resulted in higher chickpea yields than traditional bulk ZnSO<sub>4</sub> applications, illustrating their capacity to boost plant growth, biomass, and zinc accumulation in grains (<xref ref-type="bibr" rid="B18">Burman et&#xa0;al., 2013</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Mechanisms of plant resistance to abiotic stresses.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Abiotic stress</th>
<th valign="middle" colspan="3" align="center">Activate plant signaling cascades</th>
<th valign="middle" align="left">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="2" align="left">Drought</td>
<td valign="top" align="left">Enhance stress hormone accumulation example abscisic acid (ABA)</td>
<td valign="top" align="left">Stress response regulation</td>
<td valign="top" align="left">Improve water use efficiency</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B47">Heikal et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">affects calcium signaling</td>
<td valign="top" align="left">Alter Ca<sup>+ 2</sup> influx or efflux and activating stress response genes</td>
<td valign="top" align="left">Enhance drought tolerance</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B102">Nasrallah et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="middle" rowspan="3" align="left">Salinity</td>
<td valign="top" align="left">Increase antioxidant enzyme expressions</td>
<td valign="top" align="left">Reduce ROS levels and oxidative damage</td>
<td valign="top" align="left">Enhance SOD and CAT activities</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B28">El-Saadony et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Influence the salt overly sensitive (SOS) pathway</td>
<td valign="top" align="left">Stabilize SOS gene expression</td>
<td valign="top" align="left">Up regulating SOS1 expression</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B79">Mahmoud et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">activate the mitogen-activated protein kinase (MAPK) signaling pathway</td>
<td valign="top" align="left">Enhance stress-responsive gene transcription</td>
<td valign="top" align="left">Enhance oxidative stress resistance</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B55">Joshi and Joshi, 2024</xref>)</td>
</tr>
<tr>
<td valign="middle" rowspan="2" align="left">Temperature</td>
<td valign="top" align="left">Stimulate osmolyte biosynthesis</td>
<td valign="top" align="left">Aid osmotic adjustment and cellular structure protection</td>
<td valign="top" align="left">Increase protein accumulation</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B9">Al-Khayri et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Activate heat shock factors signaling</td>
<td valign="top" align="left">Increase HSPs accumulation</td>
<td valign="top" align="left">Increase cellular protection against heat damage</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B179">Yang et&#xa0;al., 2020</xref>)</td>
</tr>
<tr>
<td valign="middle" align="left">Heavy metal</td>
<td valign="top" align="left">Modulate the Halliwell&#x2013;Asada pathway for detoxifying H<sub>2</sub>O<sub>2</sub></td>
<td valign="top" align="left">Up regulate pathway enzymes</td>
<td valign="top" align="left">Reduce H<sub>2</sub>O<sub>2</sub> levels</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B114">Rajput et&#xa0;al., 2023</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Magnesium oxide nanoparticles (MgO NPs) also contribute to enhanced growth and physiological characteristics in tobacco plants. These nanoparticles increase chlorophyll content, enzyme activity, and magnesium uptake without causing phytotoxic effects (<xref ref-type="bibr" rid="B19">Cai et&#xa0;al., 2020</xref>). Additional research has shown that MgO NPs influence growth, chlorophyll content, and gene/miRNA expression in ornamental pineapple. While they enhance plant development at certain concentrations, higher doses may inhibit these benefits (<xref ref-type="bibr" rid="B105">Owusu Adjei et&#xa0;al., 2021</xref>).</p>
<p>Polymeric nanoparticles represent another critical category and include cellulose and chitosan nanoparticles. These materials facilitate the delivery of nutrients and protective agents into plant tissues, thereby improving growth and stress resistance. Notably, chitosan nanoparticles have been found to stimulate plant defense mechanisms through complex interactions with physiological pathways (<xref ref-type="bibr" rid="B143">Shinde et&#xa0;al., 2024</xref>).</p>
<p>Protein and lipid-based nanoparticles also hold multiple important applications in plant systems. These biocompatible carriers enhance the stability and effectiveness of bioactive molecules during their transport to target sites within the plant. Their application can modulate secondary metabolism and promote the biosynthesis of valuable secondary metabolites (<xref ref-type="bibr" rid="B165">Verma et&#xa0;al., 2024</xref>).</p>
<p>As elicitors, nanoparticles have gained considerable attention for their ability to induce the production of secondary metabolites under stress conditions, thereby enhancing plant resistance and metabolic activity. Additionally, they promote the generation of reactive oxygen species (ROS) and trigger the activation of secondary metabolic pathways. This function is particularly important for maximizing the production of commercially valuable bioactive compounds across various industries (<xref ref-type="bibr" rid="B69">Lala, 2021</xref>).</p>
<p>Therefore, nanoparticles are increasingly being designed to regulate plant immunity, particularly by interacting with plant-specific pattern recognition receptors (PRRs) and their associated signaling pathways. However, the rigid structure of plant cell walls poses unique challenges to immune modulation via nanoparticles. To overcome this, nanoparticles must be able to penetrate or traverse the cell wall to reach intracellular targets or interact with receptors on the plasma membrane. The formulation or surface modification of nanoparticles is thus critical for enhancing their delivery and uptake by plant cells (<xref ref-type="bibr" rid="B112">Pradeep et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Biogenic synthesis</title>
<p>Biogenic synthesis creates nanoparticles (NPs) sustainably, using plants or microbes instead of harsh chemicals. Plant extracts are a common source. Examples include neem (<italic>Azadirachta indica</italic>) and moringa (<italic>Moringa oleifera</italic>) (<xref ref-type="bibr" rid="B12">Alnaddaf et&#xa0;al., 2021</xref>). These extracts contain natural compounds, such as polyphenols and flavonoids, which act as reducing agents. They convert metal salts into NPs. These compounds also act as capping agents, stabilizing the NPs. This method is simple, cost-effective, works at room temperature, and avoids toxic by-products (<xref ref-type="bibr" rid="B95">Murali et&#xa0;al., 2021</xref>). Different plants yield NPs of different sizes and shapes. Microbes also synthesize NPs. Bacteria like <italic>Bacillus subtilis</italic> and <italic>Pseudomonas fluorescens</italic> are used. Synthesis can occur inside or outside the bacterial cell. Enzymes or metabolites reduce metal ions. Bacterial synthesis offers good size control and is scalable and eco-friendly (<xref ref-type="bibr" rid="B93">Murali et&#xa0;al., 2023</xref>). Fungi and yeast can be used too (<xref ref-type="bibr" rid="B10">Alloosh et&#xa0;al., 2021</xref>). Green NPs possess a natural biomolecule coating derived from the plant or microbe. This coating often enhances NP properties, such as biocompatibility or antimicrobial effects. Green NPs are generally safer and show higher efficacy in agriculture than chemically synthesized ones. These methods are crucial, sustainable alternatives (<xref ref-type="bibr" rid="B158">Tasnim et&#xa0;al., 2024</xref>).</p>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>Nanoparticle-plant interactions</title>
<sec id="s3_1">
<label>3.1</label>
<title>Mechanisms of nanoparticle internalization and physiological effects in plants</title>
<sec id="s3_1_1">
<label>3.1.1</label>
<title>Pathways and transport dynamics of nanoparticles in plants</title>
<p>Understanding the intricate processes by which nanoparticles (NPs) enter, move within, and exert their effects on plant systems at both cellular and subcellular levels is paramount for their effective, safe, and sustainable application in agriculture (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>) (<xref ref-type="bibr" rid="B13">Alnaddaf et&#xa0;al., 2025</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Mechanisms and signaling pathways for nanoparticle application in plants (constructed by L. M. Alnaddaf).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-16-1626624-g001.tif">
<alt-text content-type="machine-generated">Flowchart depicting mechanisms affecting processes, divided into six categories: Interaction Mechanisms, Cellular Uptake Mechanisms, Transport Pathways, Signaling Pathways, Physiological Effects, and Environmental Factors. Each category lists specific elements such as surface binding, endocytosis, xylem transport, hormone pathways, photosynthesis, and various environmental factors. Each category is color-coded and includes related subpoints.</alt-text>
</graphic></fig>
<p>The primary pathway for nanoparticle entry into plants is through the root system. Nanoparticles can traverse the epidermal cell wall of roots, typically through pre-existing small pores (ranging from 3 to 5 nm). In instances where nanoparticles are larger than these natural pores, they may induce the formation of new entry points, facilitating their absorption (<xref ref-type="bibr" rid="B151">Singh et&#xa0;al., 2024</xref>).</p>
<p>Nanoparticles can also enter plants through their leaves, primarily via stomata (microscopic pores on the leaf surface) or through microscopic cracks and imperfections in the leaf cuticle (the waxy protective layer). Various assisted delivery methods, beyond natural entry points, can enhance nanoparticle uptake. These include seed priming (where seeds are treated with nanoparticles before planting), hydroponic systems (where nanoparticles are introduced directly into the nutrient solution), and direct injection methods (<xref ref-type="bibr" rid="B26">Djanaguiraman et&#xa0;al., 2025</xref>).</p>
<p>Once inside the plant, nanoparticles can move through the apoplastic pathway. This involves movement through the non-living components, specifically the cell walls and intercellular spaces. This is generally a passive process driven by transpiration pull (<xref ref-type="bibr" rid="B67">Kumar and Biswas, 2023</xref>).</p>
<p>Additionally, nanoparticles can move via the symplastic pathway. This involves movement through the living components, specifically the cytoplasm of adjacent cells connected by plasmodesmata (cytoplasmic bridges). This movement often requires the assistance of membrane carrier proteins (<xref ref-type="bibr" rid="B26">Djanaguiraman et&#xa0;al., 2024</xref>).</p>
<p>After entering the root or leaf, nanoparticles are efficiently transported throughout the plant via its vascular system, comprising the xylem and phloem. The xylem primarily facilitates the upward transport of nanoparticles from the roots to the aerial parts alongside water and nutrients. The phloem, responsible for sugar transport, can also redistribute nanoparticles throughout the plant, including remobilization back to the roots. The efficiency and pattern of nanoparticle uptake and translocation are significantly influenced by several factors. These include the nanoparticle&#x2019;s size, shape, surface charge, and chemical composition. Additionally, plant species-specific characteristics and prevailing environmental conditions play crucial roles in determining their movement within the plant (<xref ref-type="bibr" rid="B13">Alnaddaf et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s3_1_2">
<label>3.1.2</label>
<title>Cellular interactions and physiological impacts of nanoparticles</title>
<p>Nanoparticles initially interact with plant cell walls and plasma membranes, which serve as the primary physical barriers. High concentrations of certain nanoparticles can compromise membrane integrity, potentially leading to the leakage of cellular contents and disruption of normal cellular functions (<xref ref-type="bibr" rid="B35">Gao et al., 2023</xref>). Following entry and translocation, nanoparticles can accumulate within various cellular and subcellular organelles. Common sites of accumulation include chloroplasts (affecting photosynthesis), mitochondria (impacting respiration), and vacuoles (involved in detoxification and storage) (<xref ref-type="bibr" rid="B151">Sonkar et&#xa0;al., 2023</xref>).</p>
<p>The presence of nanoparticles can induce a wide array of physiological and biochemical changes in plants. These include alterations in photosynthetic rates, nutrient uptake efficiency, enzyme activity, and overall gene expression patterns (<xref ref-type="bibr" rid="B13">Alnaddaf et&#xa0;al., 2025</xref>).</p>
<p>While some nanoparticles mitigate oxidative stress by enhancing antioxidant defense systems, others, particularly at elevated concentrations, can paradoxically induce oxidative stress by promoting the generation of reactive oxygen species (ROS). Nanoparticles have been observed to influence the synthesis, transport, and signaling pathways of key plant hormones, such as auxins, gibberellins, cytokinins, and abscisic acid. This modulation can significantly impact plant growth, development, and adaptive stress responses (<xref ref-type="bibr" rid="B109">P&#xe1;ramo et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B133">Selvakesavan et&#xa0;al., 2023</xref>).</p>
<p>Emerging research suggests that nanoparticles may interact with the plant&#x2019;s genetic material, potentially leading to changes in gene expression or, in some cases, DNA damage. Further research is needed to fully understand these complex genetic interactions and their long-term implications (<xref ref-type="bibr" rid="B11">Alnaddaf et&#xa0;al., 2023</xref>).</p>
<p>Crucially, the effects of nanoparticles are highly dose-dependent. Low and optimized concentrations often confer beneficial effects, such as enhanced growth, improved nutrient utilization, and increased stress tolerance. Conversely, higher concentrations can lead to phytotoxicity, growth inhibition, and other adverse impacts on plant health (<xref ref-type="bibr" rid="B94">Murali et&#xa0;al., 2022</xref>).</p>
</sec>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Determinants of nanoparticle efficacy and various effects on plants</title>
<p>The impact of nanoparticles on plants is a complex phenomenon, highly contingent upon a multitude of factors related to the nanoparticles themselves, the specific plant species, and the prevailing environmental conditions. A thorough understanding of these influencing factors is paramount for optimizing the beneficial applications of nanotechnology in agriculture while simultaneously mitigating potential risks (<xref ref-type="table" rid="T2"><bold>Tables&#xa0;2</bold></xref>, <xref ref-type="table" rid="T3"><bold>3</bold></xref>).</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Potential effects of NPs on plant growth, physiology, nutrition and stress mitigation.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Nanoparticle type</th>
<th valign="middle" align="center">Potential effects</th>
<th valign="middle" align="left">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="top" colspan="3" align="center">Enhanced growth &amp; nutrient uptake</th>
</tr>
<tr>
<td valign="top" align="left">ZnO and Fe<sub>2</sub>O<sub>3</sub></td>
<td valign="top" align="left">Zinc oxide and iron oxide (Fe<sub>2</sub>O<sub>3</sub>) nanoparticles enhance seed germination rates by promoting enzymatic activity and boosting energy production within seeds.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B42">Guo et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">stimulate root elongation and lateral root formation<break/>increasing the availability of essential micronutrients</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B98">Nair and Chung, 2017</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">Enhance zinc ion availability, essential for enzyme function and protein synthesis, improving plant nutrition and growth, especially under nutrient-limited conditions.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B118">Rashid et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Graphene Oxide</td>
<td valign="top" align="left">Act as carriers for nutrients like magnesium, further boosting photosynthetic activity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B129">Santana et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Carbon-based nanomaterials CNTs</td>
<td valign="top" align="left">Form nanochannels in root cell membranes, facilitating water/nutrient transport and supporting robust root/shoot development in seedlings.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B56">Joshi et&#xa0;al., 2018</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Cu and Zn NPs</td>
<td valign="top" align="left">Boost enzymatic activities related to carbohydrate metabolism, ensuring sufficient energy for growth.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B80">Malik et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Fe<sub>2</sub>O<sub>3</sub> NPs</td>
<td valign="top" align="left">Improve root/shoot biomass (e.g., in wheat, rice) by providing a readily absorbable iron source, facilitating efficient iron uptake and utilization, and addressing iron deficiency.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B51">Hussain et&#xa0;al., 2019</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">TiO<sub>2</sub> NPs</td>
<td valign="top" align="left">Enhance seed germination by improving water uptake and accelerating the breakdown of stored carbohydrates. This results in faster, uniform seedling emergence.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B136">Shah et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">Improve zinc (Zn) ion availability, which is essential for enzyme function and protein synthesis</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B54">Jan et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left">Promote shoot growth by enhancing cell division and elongation in the apical meristem</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B171">Wang et&#xa0;al., 2019</xref>)</td>
</tr>
<tr>
<th valign="top" colspan="3" align="center">Enhanced photosynthesis</th>
</tr>
<tr>
<td valign="top" align="left">TiO<sub>2</sub> NPs</td>
<td valign="top" align="left">Increase light absorption and chlorophyll content, leading to higher photosynthetic rates. They interact with chloroplasts to improve the efficiency of light-dependent reactions and carbon fixation.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B25">Dias et&#xa0;al., 2018</xref>)</td>
</tr>
<tr>
<th valign="top" colspan="3" align="center">Stress mitigation</th>
</tr>
<tr>
<th valign="top" colspan="3" align="center">Reducing Oxidative Stress (ROS Scavenging)</th>
</tr>
<tr>
<td valign="top" align="left">Carbon-based Nanomaterials (CNTs)</td>
<td valign="top" align="left">Reduce oxidative stress by scavenging ROS, stabilizing metabolic processes under stress.<break/>stabilizing metabolic processes under challenging environmental conditions</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B110">Patel et&#xa0;al., 2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO and TiO<sub>2</sub></td>
<td valign="top" align="left">Reduce oxidative damage by scavenging ROS, preventing cellular degradation during water scarcity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B63">Khan et&#xa0;al., 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Au and Ag</td>
<td valign="top" align="left">Reduce heat-induced ROS accumulation, shielding plants from oxidative damage.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B176">Wu et&#xa0;al., 2017</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO and cerium<break/>oxide (CeO<sub>2</sub>)</td>
<td valign="top" align="left">Reduce ROS accumulation caused by heavy metal toxicity, minimizing oxidative damage</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B177">Xia et&#xa0;al., 2008</xref>)</td>
</tr>
<tr>
<th valign="top" colspan="3" align="center">Salinity stress</th>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left">Mitigate ion toxicity by reducing sodium uptake while increasing potassium assimilation, preserving ionic homeostasis</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B60">Khan et&#xa0;al., 2020</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Silicon (Si) NPs</td>
<td valign="top" align="left">Strengthen cell walls and improve membrane stability, mitigating salt-induced dehydration</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B174">Wei et&#xa0;al., 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">carbon-based nanomaterials</td>
<td valign="top" align="left">Promote osmolyte (e.g., proline) production, maintaining cellular turgor and enzymatic activity, enhancing resilience under salinity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B125">Safikhan et&#xa0;al., 2018</xref>)</td>
</tr>
<tr>
<th valign="top" colspan="3" align="center">Heavy metal stress</th>
</tr>
<tr>
<td valign="top" align="left">Fe<sub>2</sub>O<sub>3</sub> NPs</td>
<td valign="top" align="left">Exhibit high affinity for heavy metal ions, immobilizing them in soil and preventing plant uptake</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B149">Soliemanzadeh and Fekri, 2021</xref>)</td>
</tr>
<tr>
<th valign="top" colspan="3" align="center">Heat stress</th>
</tr>
<tr>
<td valign="top" align="left">Si NPs</td>
<td valign="top" align="left">Stimulate heat shock protein (HSP) production, protecting cellular proteins and membranes from denaturation</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B45">Haq et&#xa0;al., 2019</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
<sec id="s3_2_1">
<label>3.2.1</label>
<title>Nanoparticle properties: uptake and bioactivity</title>
<p>The concentration at which nanoparticles are applied is a critical determinant of their effects. Low, optimized concentrations often elicit beneficial responses, such as enhanced plant growth, improved nutrient uptake, and increased tolerance to various stresses. Conversely, higher concentrations can lead to phytotoxicity, induce oxidative stress, and impair essential physiological processes (<xref ref-type="bibr" rid="B145">Siddiqi et&#xa0;al., 2021</xref>).</p>
<p>Moreover, the physical size of nanoparticles profoundly influences their uptake, subsequent translocation within the plant, and their interactions with plant cells. Generally, smaller nanoparticles (e.g., those in the 3-5 nm range) are more readily absorbed by roots and exhibit greater mobility throughout the plant vascular system (<xref ref-type="bibr" rid="B61">Khan et&#xa0;al., 2022</xref>). The size exclusion limits of plant tissues, such as the pores in the cuticle and stomata, play a crucial role in regulating nanoparticle entry. Furthermore, the morphology or shape of nanoparticles can also significantly affect their cellular uptake and interactions with plant cells. Different shapes may influence their biological activity and potential toxicity (<xref ref-type="bibr" rid="B164">Tripathi et&#xa0;al., 2021</xref>).</p>
<p>The surface charge of nanoparticles is a key factor influencing their initial interaction with plant surfaces (e.g., the root epidermis) and their subsequent movement within plant tissues. Surface modifications, which can alter the charge, are often employed to enhance uptake efficiency or achieve targeted delivery (<xref ref-type="bibr" rid="B99">Nair et&#xa0;al., 2020</xref>).</p>
<p>The intrinsic chemical composition of nanoparticles dictates their inherent properties and how they interact with biological systems. Nanoparticles composed of different materials (e.g., silver, gold, zinc oxide, and titanium dioxide) can exert distinct effects on plant physiology and biochemistry (<xref ref-type="bibr" rid="B113">Rajput et&#xa0;al., 2021</xref>).</p>
<p>In addition, the presence of a coating or specific surface modifications on nanoparticles can significantly alter their stability, dispersibility in various media, uptake efficiency, and overall biological activity. These modifications are strategically used to enhance beneficial effects or reduce potential toxicity (<xref ref-type="bibr" rid="B13">Alnaddaf et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s3_2_2">
<label>3.2.2</label>
<title>Exposure parameters: duration, application method, and environmental context</title>
<p>The duration of plant exposure to nanoparticles directly influences the extent of nanoparticle uptake, their accumulation within plant tissues, and the resulting physiological responses. Prolonged exposure may lead to different outcomes compared to acute, short-term exposure (<xref ref-type="bibr" rid="B138">Sharma et&#xa0;al., 2020</xref>). The application method (e.g., foliar spray, direct soil application, seed treatment, or integration into hydroponic systems) dictates the primary entry pathways and subsequent distribution patterns within the plant (<xref ref-type="bibr" rid="B5">Ahmad et&#xa0;al., 2023</xref>). External environmental conditions, including soil pH, ambient temperature, light intensity, and the presence of other ions or compounds in the soil or growth medium, can significantly influence nanoparticle stability, bioavailability to plants, and interactive effects with plant systems (<xref ref-type="bibr" rid="B182">Yasmeen, 2023</xref>).</p>
</sec>
<sec id="s3_2_3">
<label>3.2.3</label>
<title>Plant species-specific and developmental stage-dependent responses</title>
<p>Different plant species exhibit varying degrees of sensitivity and diverse physiological responses to nanoparticles. These variations are due to differences in inherent physiological characteristics, root architecture, leaf morphology, and unique genetic makeup (<xref ref-type="bibr" rid="B190">Zhang et&#xa0;al., 2022</xref>). Plant root exudates (organic compounds released by roots into the rhizosphere) can interact with nanoparticles in the soil. This interaction can affect nanoparticle aggregation, dissolution, and ultimately, their uptake (<xref ref-type="bibr" rid="B23">Chen et&#xa0;al., 2021</xref>). The specific developmental stage of the plant at the time of nanoparticle exposure influences its susceptibility to nanoparticle effects and its capacity to respond adaptively (<xref ref-type="bibr" rid="B194">Zhou et&#xa0;al., 2023</xref>).</p>
<p>Understanding the interplay of these complex factors is essential for the rational design and development of effective and safe nano-agricultural products, as well as for accurately predicting their environmental fate and overall impact on agricultural ecosystems.</p>
</sec>
</sec>
</sec>
<sec id="s4">
<label>4</label>
<title>Mechanistic roles of NPs in abiotic stress mitigation</title>
<p>Nanoparticles play pivotal roles in alleviating plant stress through multiple interconnected mechanisms that enhance the plant&#x2019;s intrinsic defense systems and optimize key physiological processes, thereby sustaining plant health and productivity under challenging environmental conditions (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>, <xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>) (<xref ref-type="bibr" rid="B187">Zaman et&#xa0;al., 2025</xref>). Specifically, we examine how nano-formulated nutrients, in conjunction with phytohormones, influence plant growth and stress resilience. This influence occurs through the promotion of antioxidant enzyme synthesis. These nano-enabled strategies activate critical defense mechanisms, including reactive oxygen species (ROS) scavenging systems, thereby improving plant tolerance to adverse environmental conditions.</p>
<p>Our analysis demonstrates how nano-mineral nutrient management simultaneously modulates key antioxidant enzymes (superoxide dismutase, catalase, and peroxidase) and regulates phytohormonal activity (auxins, cytokinins, and gibberellins). This dual modulation is necessary for three key processes: reducing ionic toxicity, strengthening oxidative stress responses, and maintaining cellular water balance in stressed plants (<xref ref-type="bibr" rid="B85">Meng et&#xa0;al., 2025</xref>).</p>
<sec id="s4_1">
<label>4.1</label>
<title>Drought stress mitigation and associated mechanisms</title>
<p>Drought is one of the most critical abiotic challenges facing global agriculture, resulting from insufficient irrigation and reduces rainfall that leads to prolonged dry periods, severely limiting crop growth. The complex nature of drought stress makes it difficult to monitor and manage effectively through conventional approaches (<xref ref-type="bibr" rid="B111">P&#xe9;rez-Labrada et&#xa0;al., 2020</xref>).</p>
<sec id="s4_1_1">
<label>4.1.1</label>
<title>Mechanistic responses to drought</title>
<p>Recent studies demonstrate that cerium oxide nanoparticles enhance drought tolerance in sorghum by regulating abscisic acid (ABA)-related genes and promoting the activity of antioxidant enzymes (<xref ref-type="bibr" rid="B28">El-Saadony et&#xa0;al., 2022</xref>). ABA plays a vital role during drought stress by promoting stomatal closure to minimize water loss and interacting with JA/SA signaling pathways. The P5CS gene enhances proline biosynthesis for osmotic adjustment under drought, while AREB/ABF transcription factors activate ABA-responsive genes during water stress (<xref ref-type="bibr" rid="B185">Yoshida et&#xa0;al., 2015</xref>). Downregulation of TAS14 and ZFHD genes increases drought tolerance by regulating ABA biosynthesis and osmotic pressure (<xref ref-type="bibr" rid="B111">P&#xe9;rez-Labrada et&#xa0;al., 2020</xref>).</p>
<p>Silver nanoparticles (AgNPs) alleviate drought stress in plant species such as lentils by maintaining water balance and enhancing growth parameters through improved osmotic adjustment and enhanced antioxidant capacity (<xref ref-type="bibr" rid="B6">Ahmad et&#xa0;al., 2024</xref>). Iron nanoparticles (FeNPs), particularly Fe<sub>2</sub>O<sub>3</sub>, play vital roles in several plant metabolic processes, including photosynthesis, respiration, DNA synthesis, and pigment production. Fe<sub>2</sub>O<sub>3</sub> nanoparticles can mitigate oxidative stress in drought-exposed plants by reducing reactive oxygen species (ROS) levels and enhancing chlorophyll content, thereby improving photosynthetic efficiency under stress conditions (<xref ref-type="bibr" rid="B137">Shang et&#xa0;al., 2019</xref>).</p>
</sec>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Salinity stress mitigation and ionic homeostasis</title>
<p>Salinity affects approximately 20% of the world&#x2019;s arable land, arising from high sodium chloride concentrations in coastal and arid regions. This condition particularly threatens glycophytes, which are highly sensitive to salt stress, limiting both food production and crop quality (<xref ref-type="bibr" rid="B20">Cao et&#xa0;al., 2025</xref>).</p>
<sec id="s4_2_1">
<label>4.2.1</label>
<title>Mechanistic responses to salinity</title>
<p>Iron oxide nanoparticles enhance plant growth in saline environments by helping maintain ionic balance through activation of salt overly sensitive (SOS) kinases and calcium signaling pathways, which regulate gene networks for stress mitigation and ion homeostasis (<xref ref-type="bibr" rid="B84">Meel and Saharan, 2024</xref>). These pathways involve mitogen-activated protein kinases (MAPKs) that initiate defense responses and facilitate cellular repair processes.</p>
<p>Zerovalent nano-iron (nZVI) particles may release hydroxyl radicals (OH&#x2022;) capable of degrading pectins. Zinc oxide nanoparticles mitigate the excessive salinity effect on basil plants by regulating proline accumulation and enhancing antioxidant activity (<xref ref-type="bibr" rid="B20">Cao et&#xa0;al., 2025</xref>). Silver nanoparticles (AgNPs) alleviate salt stress by restoring ionic balance and improving nutrient availability, with studies showing nearly doubled growth parameters in quinoa Q6 line exposed to salinity stress (<xref ref-type="bibr" rid="B43">Gupta et&#xa0;al., 2023</xref>).</p>
</sec>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Temperature stress and physiological protection</title>
<p>Temperature represents another major risk factor affecting crop yields, with high temperatures negatively affecting respiration, transpiration, and photosynthesis. Temperature stress leads to reduced photosynthesis, shortened growth periods, and ultimately lower crop yields. Challenges intensify under climate change conditions (<xref ref-type="bibr" rid="B147">Sidhu et&#xa0;al., 2024</xref>).</p>
<sec id="s4_3_1">
<label>4.3.1</label>
<title>Mechanistic responses to temperature stress</title>
<p>Nanoparticles provide physical protection by enhancing the structural integrity of plant cell walls, making plants more resilient to thermal and environmental stresses. They also modulate levels of key phytohormones such as ethylene (ET), which regulates genes associated with cell wall reinforcement and defense activation under temperature stress conditions (<xref ref-type="bibr" rid="B147">Sidhu et&#xa0;al., 2024</xref>).</p>
</sec>
</sec>
<sec id="s4_4">
<label>4.4</label>
<title>Heavy metal stress and detoxification mechanisms</title>
<p>Heavy metals, soil erosion, floods, and nutrient deficiencies considerably affect crop yield and quality. Silicon nanoparticles (SiNPs) reduce cadmium (Cd) stress by increasing biomass and lowering oxidative stress through enhanced antioxidant enzyme activities, including superoxide dismutase (SOD), glutathione reductase (GR), catalase (CAT), and phenylalanine ammonia-lyase (PAL) (<xref ref-type="bibr" rid="B7">Ahmed et&#xa0;al., 2023</xref>).</p>
<sec id="s4_4_1">
<label>4.4.1</label>
<title>Mechanistic responses to heavy metal toxicity</title>
<p>SiNPs significantly improve both yield and quality of rice grains under lead (Pb) and Cd stress by inhibiting metal uptake and facilitating sequestration into less harmful cellular compartments (<xref ref-type="bibr" rid="B64">Khanna et&#xa0;al., 2021</xref>). Iron nanoparticles (FeNPs) stimulate growth in stressed plants by promoting chlorophyll content and photosynthesis while reducing the toxicity and metal bioavailability in contaminated soils (<xref ref-type="bibr" rid="B96">Naidu et&#xa0;al., 2023</xref>).</p>
</sec>
</sec>
<sec id="s4_5">
<label>4.5</label>
<title>Molecular signaling pathways and stress response integration</title>
<sec id="s4_5_1">
<label>4.5.1</label>
<title>ROS regulation and antioxidant defense</title>
<p>NPs can induce controlled oxidative stress by increasing ROS levels, which may initially disrupt redox homeostasis (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). However, they simultaneously enhance antioxidant defense systems by stimulating both enzymatic antioxidants (SOD, CAT, APX, GR) and non-enzymatic antioxidants (ascorbic acid, glutathione, phenols, flavonoids). Some nanoparticles, like C<sub>60</sub> fullerenes, directly scavenge excess ROS, reducing oxidative damage during abiotic stress (<xref ref-type="bibr" rid="B108">Pandey et&#xa0;al., 2018</xref>).</p>
</sec>
<sec id="s4_5_2">
<label>4.5.2</label>
<title>Hormonal regulation and gene expression</title>
<p>NPs modulate levels of key phytohormones, including ABA, jasmonic acid (JA), and salicylic acid (SA), which are central to abiotic stress signaling. JA regulates genes for defense compounds and secondary metabolite production, while SA coordinates systemic acquired resistance responses (<xref ref-type="bibr" rid="B6">Ahmad et&#xa0;al., 2024</xref>). Bionanoparticles can deliver nucleic acids into plant cells, leading to upregulation or downregulation of specific genes related to stress adaptation through interactions with cellular components that affect gene accessibility (<xref ref-type="bibr" rid="B16">Bahwirth et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s4_5_3">
<label>4.5.3</label>
<title>Secondary metabolite enhancement</title>
<p>NPs act as elicitors, triggering ROS production that activates secondary metabolic pathways, enhancing synthesis of bioactive compounds, such as flavonoids, phenolics, and terpenoids. These compounds serve dual roles in plant defense and human health benefits. Exposure to copper oxide nanoparticles (CuO) increased polyphenol content and antioxidant activity in Withania somnifera (<xref ref-type="bibr" rid="B120">Raza et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s4_5_4">
<label>4.5.4</label>
<title>Nutrient optimization and physical protection</title>
<p>As nanofertilizers, NPs enhance nutrient uptake and utilization efficiency, ensuring adequate nutrition during stress conditions while supporting overall metabolism and growth. They also aid in osmotic adjustment by promoting osmotic accumulation, helping plants maintain turgor and essential physiological functions under water-deficient conditions (<xref ref-type="bibr" rid="B126">Saleh et&#xa0;al., 2021</xref>).</p>
<p>The multifunctional nature of nanoparticles, from molecular signaling to whole-plant physiological responses, underscores their significant potential in enhancing plant resilience to abiotic stresses. Their ability to simultaneously address multiple stress factors while optimizing plant defense mechanisms positions nanotechnology as a promising tool for advancing sustainable and stress-resilient agriculture under changing climate conditions (<xref ref-type="bibr" rid="B13">Alnaddaf et&#xa0;al., 2025</xref>).</p>
</sec>
</sec>
</sec>
<sec id="s5">
<label>5</label>
<title>Agricultural applications</title>
<sec id="s5_1">
<label>5.1</label>
<title>The role of nanotechnology in promoting crop resilience against abiotic stress</title>
<p>Nanoparticles utilize their diverse and versatile properties to enhance plant tolerance to abiotic stresses, affecting all parts of the plant through multiple complementary mechanisms (<xref ref-type="table" rid="T2"><bold>Tables&#xa0;2</bold></xref>, <xref ref-type="table" rid="T3"><bold>3</bold></xref>).</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Effect of nanoparticles on different plants.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="center">Nanoparticle type</th>
<th valign="top" align="center">Concentration</th>
<th valign="top" align="center">Mode of application</th>
<th valign="top" align="center">Plant</th>
<th valign="top" align="center">Remarks</th>
<th valign="top" align="center">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">nano-Si-CDs</td>
<td valign="top" align="left">10 mg/L</td>
<td valign="top" align="left">foliar</td>
<td valign="top" align="center">Maize</td>
<td valign="top" align="left">Significantly increased fresh weight (327.1% in roots, 247.2% in shoots) and dry weight (212.0% in roots, 118.5% in shoots)</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B181">Yao et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">carbon dots (CDs)</td>
<td valign="top" align="left">10, 20, 30, and 40 mg/L</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Lettuce</td>
<td valign="top" align="left">Enhanced photosynthetic activity by improving key mechanisms, including chloroplast activity, chlorophyll content, Rubisco enzyme activity, and photosystem II (PSII) performance.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B49">Hu et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Si NPs</td>
<td valign="top" align="center">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Radish</td>
<td valign="top" align="left">Boosted fresh weight (36.0%), chlorophyll content (14.2%), and carotenoid levels (18.7%); increased reducing sugars (23.7%), total sugars (24.8%), and proteins (232.7%), enriching nutritional profile</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B178">Xu et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left">10 or 20 mg/L</td>
<td valign="top" align="left">soaking</td>
<td valign="top" align="center">Rice</td>
<td valign="top" align="left">Enhanced germination, seedling growth, water uptake, and upregulated aquaporin genes (PIP1;1, PIP2;1); elevated amylase, dehydrogenase, and catalase activity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B78">Mahakham et&#xa0;al., 2017</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">2&#x2013;9 g/L</td>
<td valign="top" align="left">foliar</td>
<td valign="top" align="center">Saffron</td>
<td valign="top" align="left">Increased flower yield, chlorophyll content, water retention, protein levels, and antioxidant enzyme activity (POX, CAT)</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B124">Rostami et&#xa0;al., 2019</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">TiO<sub>2</sub> NPs</td>
<td valign="top" align="center">&#x2013;</td>
<td valign="top" align="left">foliar</td>
<td valign="top" align="center">Vetiveria zizanioides</td>
<td valign="top" align="left">Enhanced biomass, essential oil/khusimol production, chlorophyll, PSII efficiency, nitrate reductase, and carbonic anhydrase activity</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B135">Shabbir et&#xa0;al., 2019</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CeO<sub>2</sub>:SA NPs</td>
<td valign="top" align="center">&#x2013;</td>
<td valign="top" align="left">foliar</td>
<td valign="top" align="center"><italic>Portulaca oleracea</italic> L.</td>
<td valign="top" align="left">Reduced salt stress impact on purslane through improved physiological functions and growth-related characteristics.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B46">Hassanpouraghdam et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Fe NPs 54nm</td>
<td valign="top" align="left">500mg/kg</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Arabidopsisthalian</td>
<td valign="top" align="left">Increased plant biomass, carbohydrates, and phosphorus via enhanced photosynthesis (increased stomatal opening) and improved P availability (reduced rhizosphere pH).</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B184">Yoon et&#xa0;al., 2019</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Fe NPs</td>
<td valign="top" align="left">20 mg/L</td>
<td valign="top" align="left">soaking for three days</td>
<td valign="top" align="center">Rice</td>
<td valign="top" align="left">Improved seedling growth was achieved by increasing tissue water content, boosting the activity of hydrolytic and antioxidant enzymes, strengthening cell membrane integrity and viability, and elevating chlorophyll and iron levels.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B41">Guha et&#xa0;al., 2018</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">10 mg/L</td>
<td valign="top" align="left">foliar</td>
<td valign="top" align="center">Mung bean</td>
<td valign="top" align="left">Promoted overall plant growth (resulting in longer stems and larger root volume), improved nutrient uptake (specifically phosphorus accumulation by stimulating phosphatase and phytase activity), increased photosynthetic capacity (via higher chlorophyll and protein levels), and fostered beneficial rhizosphere microbial populations.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B116">Raliya et&#xa0;al., 2016</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Fe NPs</td>
<td valign="top" align="left">25 mg/kg</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Wheat</td>
<td valign="top" align="left">Enhanced growth more substantially than in normal soil, increasing root/shoot/grain dry weights and outperforming FeSO<sub>4</sub>/Fe-EDTA</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B196">Zia-ur-Rehman et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Fe-0 NPs (35&#x2013;45 nm)</td>
<td valign="top" align="left">8&#x2013;14 m&#xb2;/g</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Sunflower</td>
<td valign="top" align="left">Alleviated stress via soil Cr immobilization &amp; upregulated antioxidant enzymes (SOD, POD, CAT, APX)</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B88">Mohammadi et&#xa0;al., 2020</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">25 ppm</td>
<td valign="top" align="left">seed priming</td>
<td valign="top" align="center">Rice</td>
<td valign="top" align="left">Reduced oxidative damage, enhanced antioxidant activity &amp; proline</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B82">Mazhar et&#xa0;al., 2022a</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">SeNPs</td>
<td valign="top" align="left">75 ppm</td>
<td valign="top" align="left">seed priming</td>
<td valign="top" align="center">Tomato</td>
<td valign="top" align="left">Lowered oxidative stress, boosted antioxidant defense &amp; ascorbate-glutathione cycle, increased bioactive compounds</td>
<td valign="top" align="left"><xref ref-type="bibr" rid="B52">Ishtiaq et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CaONPs</td>
<td valign="top" align="left">75 ppm</td>
<td valign="top" align="left">seed priming</td>
<td valign="top" align="center">canola</td>
<td valign="top" align="left">Improved germination (30%), seedling fresh weight (34%), leaf number (16%), chlorophyll (28.9%), pod/seed production (73%), 100-seed weight (35.13%), yield (35.18%) via improved antioxidants &amp; reduced stress markers.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B83">Mazhar et&#xa0;al., 2022b</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CuNPs</td>
<td valign="top" align="left">20mg/L</td>
<td valign="top" align="left">Foliar</td>
<td valign="top" align="center">Trigonella foenum-graecum L.</td>
<td valign="top" align="left">Improved growth/biomass, promoted pigments, osmolytes, anthocyanin, shikimic acid, phenols, upregulated antioxidant enzymes</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B31">Fouda et&#xa0;al., 2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">20, 40, and 60 mg/L for 12 h</td>
<td valign="top" align="left">soaking</td>
<td valign="top" align="center">(Lupinustermis) seeds</td>
<td valign="top" align="left">Mitigated negative impacts of 150 mM NaCl stress on seedlings.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B48">Hossain et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Fe<sub>2</sub>O<sub>3</sub> NPs (40 nm)</td>
<td valign="top" align="left">5, 10, 20, 30, and 40 &#xb5;M</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">lemon balm (Melissa officinalis</td>
<td valign="top" align="left">Increased essential oil, restored chlorophyll, decreased proline, MDA, H<sub>2</sub>O<sub>2</sub></td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B89">Mohasseli et&#xa0;al., 2020</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">&#x3b3;-Fe<sub>2</sub>O<sub>3</sub> NP</td>
<td valign="top" align="left">0.5, 0.8, 1, or 2 mg/mL</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Brassica napus</td>
<td valign="top" align="left">Increased chlorophyll, growth, reduced H<sub>2</sub>O<sub>2</sub></td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B106">Palmqvist et&#xa0;al., 2017</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">nanochitosan</td>
<td valign="top" align="left">(0.05% and 0.1%)</td>
<td valign="top" align="left">seed priming</td>
<td valign="top" align="center">broad bean seeds</td>
<td valign="top" align="left">Inhibited germination/seedling growth due to cytotoxicity; NPs penetrated seed coat, accumulated in embryos, disrupting cell division/development.</td>
<td valign="top" align="left"><xref ref-type="bibr" rid="B2">Abdel-Aziz, 2019</xref></td>
</tr>
<tr>
<td valign="top" align="left">nano-biochar</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">various crops</td>
<td valign="top" align="left">Decreased cumulative runoff (10.88&#x2013;31.75%) and sediment; increased rainfall interception (18.08&#x2013;31.14%) on slopes</td>
<td valign="top" align="left"><xref ref-type="bibr" rid="B22">Chen et&#xa0;al., 2020</xref></td>
</tr>
<tr>
<td valign="top" align="left">Schiff base -Nanoclay Polymer Composites (NCPCs)</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center"/>
<td valign="top" align="left">Effectively inhibited nitrification (30&#x2013;87%); slowed nitrification &amp; reduced nitrate leaching</td>
<td valign="top" align="left"><xref ref-type="bibr" rid="B131">Saurabh et&#xa0;al., 2019</xref></td>
</tr>
<tr>
<td valign="top" align="left">slow-release nanoclay polymer composites made up of acrylic acid (AA) and acrylamide (Am)</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Rice, wheat</td>
<td valign="top" align="left">Improved N-use efficiency: Reduced N<sub>2</sub>O emissions (rice: 16.1%; wheat: 12.4%) vs. urea</td>
<td valign="top" align="left"><xref ref-type="bibr" rid="B66">Kirti et&#xa0;al., 2021</xref></td>
</tr>
<tr>
<td valign="top" align="left">macro-nutrient (K<sub>2</sub>SO<sub>4</sub>) and micro-nutrient (ZnO and SiO<sub>2</sub>)</td>
<td valign="top" align="left">salt concentrations (6 and 10 dS/L<sup>)</sup></td>
<td valign="top" align="left">hydroponic experiment</td>
<td valign="top" align="center"><italic>Medicago sativa</italic> L.</td>
<td valign="top" align="left">Improved plant height, shoot dry weight, flowers, tillers, root length, root fresh/dry weight (nano-K<sub>2</sub>SO<sub>4</sub>); enhanced proline/SOD under salinity (nano-SiO<sub>2</sub>)</td>
<td valign="top" align="left"><xref ref-type="bibr" rid="B29">El-Shal et&#xa0;al., 2022</xref></td>
</tr>
<tr>
<td valign="top" align="left">AgNPs</td>
<td valign="top" align="left">20-30 ppm</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Soybean</td>
<td valign="top" align="left">Enhanced <italic>Bradyrhizobium japonicum</italic> symbiotic efficiency (25-30%), improving nodulation &amp; N-fixation.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B146">Siddiqui et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">AgNP-treated Mesorhizobium ciceri inoculants</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Chickpea</td>
<td valign="top" align="left">Reduced <italic>Fusarium oxysporum</italic> infections (40%) while boosting growth.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B1">Abd-Alla et&#xa0;al., 2020</xref>).</td>
</tr>
<tr>
<td valign="top" align="left">Fe<sub>2</sub>O<sub>3</sub> NPs</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Wheat</td>
<td valign="top" align="left">Synergy with <italic>Bacillus megaterium</italic>: 35% higher grain Fe, 28% yield increase vs. traditional fertilization.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B193">Zhao et&#xa0;al., 2021</xref>).</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs combined with zinc-mobilizing Pseudomonas brassicacearum (20-30 nm)</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">cultivation in zinc-deficient paddies</td>
<td valign="top" align="center">Rice</td>
<td valign="top" align="left">Increased Zn uptake efficiency (45-50%), addressing malnutrition</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B188">Zhang et&#xa0;al., 2022</xref>).</td>
</tr>
<tr>
<td valign="top" align="left">graphene oxide (GO) sheets functionalized with plant growth-promoting Bacillus amyloliquefaciens</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">hydroponic</td>
<td valign="top" align="center">Lactuca sativa</td>
<td valign="top" align="left">Enhanced nutrient uptake efficiency (35-40%), reduced leaf nitrate accumulation</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B162">Torres et&#xa0;al., 2022</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">SiO<sub>2</sub> NP-coated Azospirillum brasilense inoculants</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">In drought-prone</td>
<td valign="top" align="center">Sorghum</td>
<td valign="top" align="left">Maintained 75% colonization efficiency at 40% field capacity (vs. 30% uncoated).</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B104">Nguyen et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">silica-encapsulated Halomonas venusta biofertilizers</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">Saline coastal areas</td>
<td valign="top" align="center">Rice</td>
<td valign="top" align="left">0% higher survival &amp; maintained nitrogenase activity at 8 dS/m salinity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B169">Wang L. et&#xa0;al., 2023</xref>).</td>
</tr>
<tr>
<td valign="top" align="left">chitosan nano-formulations containing Trichoderma harzianum</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Grapevine</td>
<td valign="top" align="left">Provided season-long root pathogen protection &amp; enhanced nutrient uptake</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B81">Mart&#xed;nez et&#xa0;al., 2023</xref>).</td>
</tr>
<tr>
<td valign="top" align="left">Fe2O3 NPs</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Pumpkin</td>
<td valign="top" align="left">45% accumulated in roots, &lt;1% detected in leaves after root uptake &amp; translocation.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B195">Zhu et&#xa0;al., 2008</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">fullerene (C70)</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">foliar</td>
<td valign="top" align="center"/>
<td valign="top" align="left">Transported root&#x2192;shoot and leaf&#x2192;root (via phloem)</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B74">Lin and Xing, 2008</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CuO NPs</td>
<td valign="top" align="left">100 mg/L</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Maize</td>
<td valign="top" align="left">Found in epidermis cell walls, intercellular spaces, cortical cell cytoplasm/nuclei; suggesting apoplastic transport.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B173">Wang X. et&#xa0;al., 2023</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">a high<break/>concentration</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Tomato</td>
<td valign="top" align="left">Induced tomato oxidative stress, reducing growth/biomass.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B71">Li et&#xa0;al., 2016</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center"/>
<td valign="top" align="left">Penetrated cell walls, damaging morphology</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B128">Sangour et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">kiwi</td>
<td valign="top" align="left">Caused pollen mortality via membrane damage</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B152">Speranza et&#xa0;al., 2013</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left">higher concentrations</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center"><italic>A. thaliana</italic></td>
<td valign="top" align="left">Reduced shoot/root length by destroying root apical meristem cells</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B170">Wang et&#xa0;al., 2013</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Ag NPs</td>
<td valign="top" align="left"/>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Mung beans</td>
<td valign="top" align="left">Generated ROS, causing lipid peroxidation &amp; cellular damage.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B97">Nair and Chung, 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">TiO<sub>2</sub> NPs</td>
<td valign="top" align="left">500 mg/L to<break/>4000 mg/L</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center"><italic>Brassica napus L</italic></td>
<td valign="top" align="left">Improved morphology/physiology (root length, height, biomass, gas exchange, chlorophyll, NRase) up to 4000 mg/L foliar, but also caused pollen mortality.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B73">Li et&#xa0;al., 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ionic cerium, bulk CeO<sub>2</sub>, and CeO<sub>2</sub> NPs</td>
<td valign="top" align="left">10 mg/L</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Radish</td>
<td valign="top" align="left">The nanoparticles exhibited a range of effects, including negative, positive, and instances where no significant impact was observed.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B189">Zhang W. et&#xa0;al., 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CeO<sub>2</sub> NPs (lab-synthesized 7 nm and 25 nm and<break/>commercial CeO<sub>2</sub> NPs)</td>
<td valign="top" align="left">&#x2003;-</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Latuca species</td>
<td valign="top" align="left">All three tested CeO<sub>2</sub> nanoparticle types demonstrated toxicity towards the three <italic>Lactuca</italic> species studied. Furthermore, the different CeO<sub>2</sub> nanoparticles exhibited varying levels of toxicity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B191">Zhang P. et&#xa0;al., 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CeO<sub>2</sub> NPs (~250 nm in size) and bulk CeO<sub>2</sub> particles (~2000 nm in size)</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center"><italic>Brassica napus</italic> L.</td>
<td valign="top" align="left">Application of bulk CeO<sub>2</sub> at concentrations of 10 mg/L and 100 mg/L increased plant biomass by 28% and 35%, respectively. In contrast, CeO<sub>2</sub> nanoparticles applied at these same concentrations did not produce this enhancing effect.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B76">Ma et&#xa0;al., 2015</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">CeO<sub>2</sub> NPs</td>
<td valign="top" align="left">&#x2013;</td>
<td valign="top" align="left">fertilizer</td>
<td valign="top" align="center">Rice</td>
<td valign="top" align="left">CeO<sub>2</sub> nanoparticles altered the root&#x2019;s antioxidant defense system. Specifically, at a low concentration (62.5 mg/L), they inhibited root H<sub>2</sub>O<sub>2</sub> production by 75%. Conversely, at higher concentrations (125 mg/L and 500 mg/L), they significantly elevated root H<sub>2</sub>O<sub>2</sub> levels. This increase led to lipid peroxidation and electrolyte leakage.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B122">Rico et&#xa0;al., 2013</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">50 and 100 ppm</td>
<td valign="top" align="left">Foliar</td>
<td valign="top" align="center">eggplant</td>
<td valign="top" align="left">improve physiological traits, including membrane stability index (MSI) and relative water content (RWC), enhance photosynthesis, and strengthen the anatomical structure of stems and leaves. Critically, they also increase fruit yield by 12.2% and 22.6% (at 50 ppm and 100 ppm, respectively) compared to untreated plants.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B134">Semida et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">50 or 100</td>
<td valign="top" align="left">60% ETc irrigation</td>
<td valign="top" align="center">eggplant</td>
<td valign="top" align="left">Treated plants achieve 50.8&#x2013;66.1% higher WP than fully irrigated plants without nanoparticle treatment.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B134">Semida et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">FeNPs</td>
<td valign="top" align="left">100 mg/L</td>
<td valign="top" align="left">Foliar</td>
<td valign="top" align="center">Tomato</td>
<td valign="top" align="left">Application significantly enhances growth, increasing shoot length (42%), root length (66%), fruit weight (24%), fruit number (66%), leaf number (173%), and branch number (45%). FeNPs also boost pigment content (carotenoids and lycopene) and reduce cadmium accumulation in plant tissues</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B6">Ahmad et&#xa0;al., 2024</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
<sec id="s5_1_1">
<label>5.1.1</label>
<title>Enhanced nutrient delivery and smart release systems</title>
<p>Nanotechnology enables the development of NP-fortified fertilizers that deliver nutrients in a targeted manner, minimizing nutrient runoff and reducing groundwater contamination risks (<xref ref-type="bibr" rid="B6">Ahmad et&#xa0;al., 2024</xref>). Integration with smart fertilizers allows programmed nutrient delivery based on specific environmental conditions, such as soil moisture or pH fluctuations, supplying crops with necessary nutrients at optimal times (<xref ref-type="bibr" rid="B112">Pradeep et&#xa0;al., 2024</xref>).</p>
<p>Nanofertilizers coat or encapsulate nutrient substances, allowing controlled release into the soil while helping recharge depleted soils and maintain soil health (<xref ref-type="bibr" rid="B127">Salem et&#xa0;al., 2021</xref>). Nitrogen, potassium, and phosphorus nanoparticles improve macronutrient utilization and availability, resulting in better plant growth and increased stress tolerance (<xref ref-type="bibr" rid="B64">Khanna et&#xa0;al., 2021</xref>).</p>
</sec>
<sec id="s5_1_2">
<label>5.1.2</label>
<title>Rhizosphere interactions and soil enhancement</title>
<p>NPs interact multifacetedly with the rhizosphere, contacting microbes, minerals, and organic matter that indirectly affect plant roots. Their biological inertness allows persistence in soil for extended periods, causing significant changes in microflora populations, soil fertility, and plant physiology (<xref ref-type="bibr" rid="B84">Meel and Saharan, 2024</xref>).</p>
<p>When incorporated into soil, NPs enhance aeration and water-holding capacity, facilitating better gas exchange and water absorption. They stimulate soil enzyme activity, sustaining soil health and boosting organic processes essential for nutrient cycling (<xref ref-type="bibr" rid="B28">El-Saadony et&#xa0;al., 2022</xref>). NPs also promote plant growth-promoting microorganisms (PGPM) such as Rhizobium and Bradyrhizobium, which form symbiotic associations that fix nitrogen and mobilize nutrients (<xref ref-type="table" rid="T4"><bold>Table&#xa0;4</bold></xref>).</p>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>The role of plant growth-promoting microorganisms in drought tolerance.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Plant growth-promoting rhizobacteria (PGPR)</th>
<th valign="top" colspan="2" align="left">Mechanisms of rhizobacteria-induced drought tolerance in plants</th>
<th valign="top" align="left">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" rowspan="3" align="left"><italic>Lactobacillus paracasei</italic>, <italic>Rhodococcus jostii</italic> RHA1, <italic>Microbacterium</italic> sp. 3J1, <italic>Arthrobacter siccitolerans</italic> 4J27, <italic>Rhodococcus</italic> sp. 4J2A2, <italic>Rhodococcus opacus</italic> PD630, and <italic>Pseudomonas putida KT2440</italic></td>
<td valign="top" align="left">Overproduce varieties stable proteins and osmolytes</td>
<td valign="top" align="left">Balance the available water or reduce water loss</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B183">Yerbury et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B107">Palud et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B70">LeBlanc et&#xa0;al., 2008</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Increase gene expression that codes for enzymes involved in DNA synthesis</td>
<td valign="top" align="left">Reduce plant damage</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B36">Garc&#xed;a-Fontana et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B101">Narv&#xe1;ez-Reinaldo et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B14">Alvarez et&#xa0;al., 2004</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Activate drought-sensitive genes</td>
<td valign="top" align="left">During drought period, plants decrease evaporation and enhance water absorption via reducing leaves size and extending roots further into the soil.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B172">Wang et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B166">V&#xed;lchez et&#xa0;al., 2016</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s5_1_3">
<label>5.1.3</label>
<title>Advanced agricultural applications</title>
<p>Nanopesticides: Developed to improve pest control efficiency by delivering pesticides precisely where and when needed, minimizing environmental impact. When combined with digital farming technologies, they enable precise application methods that reduce wastage and enhance pest management (<xref ref-type="bibr" rid="B15">Atanda et&#xa0;al., 2025</xref>).</p>
<p>Nanosensors: Vital tools engineered to detect biological molecules and heavy metal concentrations, helping maintain optimal growth conditions and prevent contamination. They increase sensitivity and selectivity in gas detection, enabling real-time monitoring of environmental health indicators (<xref ref-type="bibr" rid="B143">Shinde et&#xa0;al., 2024</xref>).</p>
<p>Nanomediators: Advanced carriers that enable precise delivery of genetic material, allowing plants to regulate gene expression and respond effectively to stresses. They facilitate controlled nutrient release and reduce residual organic pollutants (<xref ref-type="bibr" rid="B114">Rajput et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B86">Mgadi et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s5_1_4">
<label>5.1.4</label>
<title>Seed priming and stress memory enhancement</title>
<p>Recent studies explore NPs&#x2019; potential to improve stress memory in seeds through brief exposure to stress stimuli, triggering faster induction of stress-related signaling pathways upon subsequent exposures. This results in enhanced germination and growth under stressful conditions (<xref ref-type="bibr" rid="B62">Khan et&#xa0;al., 2023</xref>).</p>
<p>Metal nanoparticles (Ag, Cu, Ti, Au, Zn, Fe) and their oxides, synthesized via eco-friendly methods, significantly increase germination rates and promote plant growth (<xref ref-type="bibr" rid="B123">Riseh and Vazvani, 2024</xref>). For example, nano-primed stevia seedlings exhibited 106% increased germination percentage and 128.12% increased germination speed, with seedling dry weights increasing by 283%, 168.9%, and 220% for roots, shoots, and total biomass, respectively (<xref ref-type="bibr" rid="B27">Eevera et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B82">Mazhar et&#xa0;al., 2022a</xref>).</p>
<p>Multi-walled carbon nanotubes (MWCNTs) at 100 &#x3bc;g/mL enhanced germination and seedling growth in barley, soybean, and maize, while silicon nanoparticles (SiNPs) at 20 ppm improved growth and photosynthetic efficiency in common beans (<xref ref-type="bibr" rid="B96">Naidu et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s5_1_5">
<label>5.1.5</label>
<title>Sustainable agriculture and environmental benefits</title>
<p>Nanotechnology reduces adverse effects of conventional farming by improving pesticide and fertilizer efficiency, minimizing chemical applications, and consequently decreasing risks of soil erosion, nutrient loss, and pesticide pollution. This promotes biodiversity conservation and enhances agricultural ecosystem sustainability (<xref ref-type="bibr" rid="B103">Nawaz et&#xa0;al., 2023</xref>).</p>
<p>The technology offers complementary solutions to conventional breeding limitations by supporting genetic improvement through targeted modifications and nanomaterials that promote plant resistance, leading to crops capable of withstanding climate change and abiotic stresses (<xref ref-type="bibr" rid="B163">Tortella et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s5_1_6">
<label>5.1.6</label>
<title>Long-term sustainability considerations</title>
<p>The accumulation of NPs in soil ecosystems requires careful assessment of long-term sustainability. While improving plant resilience, they influence microbial diversity, soil enzymatic activities, and soil&#x2013;plant interactions (<xref ref-type="bibr" rid="B77">MacDonald and Mohan, 2025</xref>). Extensive long-term field studies are essential to evaluate NP behavior under different soil types and climatic conditions, ensuring nanotechnology benefits are realized without compromising ecosystem health (<xref ref-type="bibr" rid="B155">Sundararajan et&#xa0;al., 2023</xref>).</p>
</sec>
</sec>
<sec id="s5_2">
<label>5.2</label>
<title>Traditional vs. nanotechnology approaches in agriculture: a comprehensive comparison</title>
<p>The global agricultural system faces unprecedented challenges, including climate change, soil degradation, water scarcity, and the need to feed a growing population while minimizing environmental impact (<xref ref-type="bibr" rid="B30">FAO (Food and Agriculture Organization), 2021</xref>). This demands innovative solutions that can address multiple constraints simultaneously. While traditional agricultural methods have proven reliable over decades, their limitations in terms of efficiency and environmental impact are increasingly apparent. Simultaneously, nanotechnology emerges as a promising alternative, offering precision and enhanced efficacy in crop stress management (<xref ref-type="bibr" rid="B100">Nandeha et&#xa0;al., 2025</xref>).</p>
<p>Traditional methods, including the use of conventional fertilizers, pesticides, and selective breeding, have supported agricultural productivity for decades. These methods demonstrate consistent performance across diverse environmental conditions and crop systems, with urea fertilizer application increasing global cereal production by an estimated 48% since 1961 (<xref ref-type="bibr" rid="B153">Stewart et&#xa0;al., 2004</xref>).</p>
<p>Furthermore, traditional pesticides and fertilizers operate within well-established regulatory frameworks that encompass comprehensive safety databases spanning decades of use. This regulatory maturity provides farmers and consumers with confidence in application and safety profiles.</p>
<p>Traditional approaches suffer from significant efficiency limitations. Research consistently demonstrates low nutrient use efficiency in conventional systems, with nitrogen use efficiency (NUE) in cereal crops typically ranging from 30 to 50% addition to significant losses through volatilization, leaching, and denitrification (<xref ref-type="bibr" rid="B119">Raun and Johnson, 1999</xref>). Similarly, traditional approaches have shown limited improvements in water use efficiency, achieving only (10-25)% improvements under stress conditions.</p>
<p>In contrast, Nanotechnology addresses many limitations of traditional methods through precision in nutrient delivery. Nanoparticles offer size-dependent cellular uptake mechanisms, with studies demonstrating that nanoparticles between 10 and 40 nm in diameter exhibit optimal cellular penetration in plant tissues (<xref ref-type="bibr" rid="B132">Schwab et&#xa0;al., 2016</xref>). This precision is exemplified by zinc oxide nanoparticles (ZnO-NPs) at 25 nm, which showed 85% higher zinc uptake efficiency in tomato plants compared to conventional zinc sulfate, with targeted accumulation in chloroplasts where zinc is most needed for photosynthetic enzymes (<xref ref-type="bibr" rid="B115">Raliya et&#xa0;al., 2015</xref>).</p>
<p>Unlike traditional single-purpose applications, single nanoparticle formulations can address multiple stress factors simultaneously through engineered multi-functionality (<xref ref-type="bibr" rid="B87">Mittal et&#xa0;al., 2020</xref>). Cerium oxide nanoparticles (CeO<sub>2</sub>-NPs) demonstrate this dual functionality as both antioxidants and photosynthetic enhancers. In drought-stressed soybean plants, CeO<sub>2</sub>-NPs (3nm) increased antioxidant enzyme activity by 45% while maintaining photosynthetic efficiency at 80% of well-watered controls (<xref ref-type="bibr" rid="B176">Wu et&#xa0;al., 2017</xref>).</p>
<p>Nanotechnology enables a significant reduction in chemical inputs while maintaining or improving efficacy (<xref ref-type="bibr" rid="B32">Fraceto et&#xa0;al., 2016</xref>). Nano-encapsulated herbicides require 60-80% lower active ingredient concentrations compared to conventional formulations while achieving equivalent weed control, reducing environmental contamination risk (<xref ref-type="bibr" rid="B68">Kumar et&#xa0;al., 2019</xref>). Additionally, the integration of nanosensors enables continuous monitoring and responsive treatment systems (<xref ref-type="bibr" rid="B37">Giraldo et&#xa0;al., 2019</xref>), with carbon nanotube-based sensors embedded in plant leaves detecting nitric oxide levels in real-time and triggering automated delivery of protective nanoparticles when stress thresholds are exceeded (<xref ref-type="bibr" rid="B173">Wang X. et&#xa0;al., 2023</xref>).</p>
<p>Direct comparisons reveal substantial advantages for nanotechnology approaches. Studies comparing nano-fertilizers to conventional alternatives showed average nutrient use efficiency improvements of (35-65) % across major nutrients (<xref ref-type="bibr" rid="B75">Liu and Lal, 2015</xref>). Water Use Efficiency improvement is even more dramatic, with nanoparticle applications achieving (25-60) % improvement compared to the traditional method. For instance, silicon nanoparticles (Si-NPs) applied to wheat under drought stress improved grain yield by 45% compared to 18% improvement with conventional silicon fertilizer (<xref ref-type="bibr" rid="B34">Frazier et&#xa0;al., 2014</xref>).</p>
<p>In addition, nanotechnology is demonstrating superior performance in enhancing plant stress tolerance. Iron oxide nanoparticles (Fe<sub>2</sub>O<sub>3</sub>NPs) enhanced salt tolerance in barley, maintaining 75% of normal yield under less than 150 mM NaCl stress compared to 45% with conventional treatments (<xref ref-type="bibr" rid="B92">Mukhtiar et&#xa0;al., 2024</xref>). However, these benefits come with complexity, as the same treatments can have variable effects across different plant systems.</p>
<p>High production costs are a significant barrier to the adoption of nanotechnology. Nano-fertilizer production costs range from 2,000 to 8,000 $ per ton compared to 400 to 800 $ for conventional fertilizers (<xref ref-type="bibr" rid="B154">Subramanian et&#xa0;al., 2015</xref>). This economic gap is more pronounced for specialized applications, with silver nanoparticles for antimicrobial applications costing approximately $50-200 per gram, making large-scale agricultural applications economically prohibitive (<xref ref-type="bibr" rid="B139">Sharma et&#xa0;al., 2018</xref>).</p>
<p>The global nano-fertilizer market is projected to reach $4.8 billion by 2025, growing at a 12.3% CAGR, driven by precision agriculture adoption (<xref ref-type="bibr" rid="B40">Grand View Research, 2020</xref>). This growth trajectory suggests increasing commercial viability as production scales increase and costs decrease.</p>
<p>Rather than viewing nanotechnology as a complete replacement for traditional methods, an integrated approach offers optimal benefits. This strategy involves using nanoparticles for high-value crops or specific stress conditions where cost-benefit ratios are favorable, combining nano-enhanced precision delivery with conventional breeding and management practices, and implementing phase-wise adoption with continuous monitoring and adaptive management.</p>
</sec>
<sec id="s5_3">
<label>5.3</label>
<title>Nanotechnology-enhanced sensing and modeling for precision plant stress management</title>
<p>Several non-AI predictive technologies are crucial for detecting and managing plant abiotic stress. Biosensors and nanobiosensors are primary examples. Nanomaterials possess unique and tunable properties, such as optical characteristics, electrical conductivity, and shock resistance. These properties enable the creation of flexible and highly sensitive detection mechanisms capable of measuring a broad spectrum of physiological and environmental parameters relevant to plant stress (<xref ref-type="bibr" rid="B91">Mukherjee et&#xa0;al., 2024</xref>). These sophisticated analytical systems integrate a biological sensing element with a transducer, enabling the detection of specific biomarkers that indicate stress in plants. Nanotechnology significantly enhances the effectiveness of these biosensors by boosting their sensitivity and specificity, thereby facilitating the early and precise identification of both abiotic and biotic stresses (<xref ref-type="bibr" rid="B117">Rana et&#xa0;al., 2024</xref>).</p>
<p>Biosensors and nanobiosensors play a crucial role in the timely identification of stress factors. Their early detection capabilities are due to proactive interventions, which are critical in mitigating potential crop losses before they become severe. The incorporation of nanomaterials in biosensor development allows for a significant increase in sensitivity and overall efficacy (<xref ref-type="bibr" rid="B157">Tang et&#xa0;al., 2024</xref>). This enables the detection of even trace amounts of stress-related biomarkers, providing a more accurate and nuanced understanding of plant stress (<xref ref-type="bibr" rid="B140">Sheikh et&#xa0;al., 2024</xref>). Nanobiosensors are versatile and capable of detecting a wide array of biological derivatives. This includes the identification of bacteria, viruses, and various chemical toxins that can impact agricultural produce, thereby contributing to food safety and quality control. These tools are often integrated with continuous monitoring platforms, allowing for real-time data collection on plant physiological responses to environmental changes (<xref ref-type="bibr" rid="B21">Chaturvedi et&#xa0;al., 2025</xref>).</p>
<p>Some models used to study the behavior of NPs and their effects on plants foe example 1- Computational Modeling: Simulating NP transport in plant tissues (e.g., using finite element analysis). 2- Physiologically Based Pharmacokinetic (PBPK) Models: Predicting NP fate within plants based on plant physiology (<xref ref-type="bibr" rid="B50">Huang et&#xa0;al., 2024</xref>). 3- Quantitative Structure-Activity Relationships (QSAR): Relating NP properties (size, charge, coating) to biological effects (uptake, toxicity, and efficacy). 4- Kinetic Modelling: Predicting NP dissolution rates and ion release over time in the rhizosphere. 5- Stochastic Models: Assessing variability in NP behavior and plant responses under field conditions (<xref ref-type="bibr" rid="B186">Yu et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B160">Thiruvengadam et&#xa0;al., 2025</xref>; <xref ref-type="bibr" rid="B130">Sarabandi et&#xa0;al., 2025</xref>). These technologies empower farmers and researchers to implement targeted management strategies, advancing toward a more responsive and efficient agricultural system without relying on complex AI models for prediction. Recent studies indicate that the use of single-walled carbon nanotube (SWNT) nano-sensors for real-time detection of SA and H<sub>2</sub>O<sub>2</sub>, elucidating stress-specific biochemical waveforms, and enabling early interventions to promote stress tolerance. Other nano-sensors can measure nutrient levels, pathogen attacks, or pollutant stress, further supporting crop resilience under abiotic (drought, salinity, heat) stress conditions. While promising, challenges remain regarding nanosensor stability, long-term effects on plants, potential environmental impacts, and cost-effectiveness for large-scale field use.</p>
</sec>
</sec>
<sec id="s6">
<label>6</label>
<title>The challenges of integrating nanotechnology in agriculture</title>
<sec id="s6_1">
<label>6.1</label>
<title>Environmental concerns and long-term safety assessment of nanoparticles in agriculture</title>
<p>The rapid advancement of agricultural nanotechnology has generated considerable enthusiasm for its potential to address global food security challenges while improving environmental sustainability (<xref ref-type="bibr" rid="B159">Thakur and Yadav, 2025</xref>). However, the unique physicochemical properties that make nanoparticles effective in agricultural applications also raise concerns about their environmental fate, ecological impacts, and long-term safety implications. The limited understanding of nanoparticle behavior in complex environmental systems necessitates a precautionary approach that balances innovation with environmental protection (<xref ref-type="bibr" rid="B44">Gupta, 2021</xref>).</p>
<p>Current knowledge gaps regarding the long-term environmental consequences of nanoparticle applications in agriculture highlight the urgent need for comprehensive risk assessment frameworks, enhanced monitoring systems, and adaptive regulatory approaches. This assessment examines the current state of knowledge regarding environmental risks while identifying critical research priorities and policy needs for sustainable implementation of agricultural nanotechnology (<xref ref-type="table" rid="T5"><bold>Table&#xa0;5</bold></xref>) (<xref ref-type="bibr" rid="B159">Thakur and Yadav, 2025</xref>).</p>
<table-wrap id="T5" position="float">
<label>Table&#xa0;5</label>
<caption>
<p>Nanotoxicity in plants: mechanisms and its effects.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Nanoparticle type</th>
<th valign="top" align="left">Toxicity mechanisms</th>
<th valign="top" align="left">Physiological/morphological effects</th>
<th valign="top" align="left">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">ZnO NPs</td>
<td valign="top" align="left">ROS overproduction, disruption of antioxidant systems, DNA damage; accumulation via apoplastic/symplastic paths</td>
<td valign="top" align="left">Inhibited germination, stunted root/shoot, chlorosis, lipid peroxidation, genotoxicity</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B59">Kang et&#xa0;al., 2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">TiO<sub>2</sub> and other metallic NPs</td>
<td valign="top" align="left">Light-induced ROS, chloroplast dysfunction, membrane damage</td>
<td valign="top" align="left">Reduced photosynthesis, cellular structural damage</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B192">Zhang et&#xa0;al., 2025</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Silicon NPs</td>
<td valign="top" align="left">Block uptake of trace metals; modulation of antioxidant defenses and gene expression</td>
<td valign="top" align="left">Enhanced stress tolerance, reduced metal toxicity</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B90">Mukarram et&#xa0;al., 2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Nanoplastics</td>
<td valign="top" align="left">Foliar penetration, disruption of phyllosphere microbes; elevated oxidative enzymes, altered metabolism</td>
<td valign="top" align="left">Inhibited growth, altered gene expression, microbial community shifts</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B142">Shi et&#xa0;al., 2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Combined NPs and Heavy Metals</td>
<td valign="top" align="left">Synergistic or antagonistic interactions affecting ROS, ion homeostasis</td>
<td valign="top" align="left">Reduced heavy metal uptake, improved enzyme activity, biomass recovery</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B150">Soni et&#xa0;al., 2024</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">Carbon-based Quantum Effects</td>
<td valign="top" align="left">Membrane interaction, DNA methylation changes, ROS induction</td>
<td valign="top" align="left">Stomatal closure, germination delay, transcriptomic/proteomic changes</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B38">Gowtham et&#xa0;al., 2024</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s6_2">
<label>6.2</label>
<title>Soil ecotoxicity and ecosystem impact assessment</title>
<sec id="s6_2_1">
<label>6.2.1</label>
<title>Microbial community disruption and functional alterations</title>
<p>Soil microbial communities represent the foundation of terrestrial ecosystem functioning, mediating critical processes, including nutrient cycling, organic matter decomposition, and plant-microbe interactions. Nanoparticles can significantly alter soil microbial diversity and functionality through direct toxic effects, modification of soil chemistry, and interference with microbial communication systems (<xref ref-type="bibr" rid="B67">Kumar et&#xa0;al., 2025</xref>).</p>
<p>Silver nanoparticles (AgNPs) demonstrate broad-spectrum antimicrobial activity that, while beneficial for pathogen control, can indiscriminately affect beneficial soil microorganisms, including nitrogen-fixing bacteria such as Rhizobium species, mycorrhizal fungi (Glomus spp.), and decomposer organisms essential for nutrient mineralization (<xref ref-type="bibr" rid="B148">Sillen et&#xa0;al., 2015</xref>). Laboratory studies indicate that AgNPs at concentrations of 1-10 mg/kg soil reduce bacterial diversity by 15-40% within 30 days of application, with effects persisting for several months after initial exposure (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
</sec>
<sec id="s6_2_2">
<label>6.2.2</label>
<title>Enzymatic activity suppression</title>
<p>Key soil enzymes, including dehydrogenase (an indicator of overall microbial activity), phosphatase (phosphorus mineralization), and &#x3b2;-glucosidase (carbon cycling), show reduced activity ranging from 20-60% inhibition following exposure to metal oxide nanoparticles at environmentally relevant concentrations. Copper oxide nanoparticles (CuO NPs) demonstrate particular concern for nitrogen cycling, inhibiting nitrification processes by 25-50% and affecting nitrogen availability for plant uptake (<xref ref-type="bibr" rid="B17">Bouhadi et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s6_2_3">
<label>6.2.3</label>
<title>Mycorrhizal association disruption</title>
<p>Titanium dioxide nanoparticles (TiO<sub>2</sub> NPs) interfere with arbuscular mycorrhizal fungi (AMF) colonization, reducing plant nutrient uptake efficiency by 10-30% and compromising the symbiotic relationships essential for sustainable agriculture. These effects are particularly concerning given the critical role of mycorrhizal associations in phosphorus acquisition and plant stress tolerance (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
</sec>
<sec id="s6_2_4">
<label>6.2.4</label>
<title>Soil chemical properties and geochemical alterations</title>
<p>Nanoparticles can fundamentally alter soil chemistry through various mechanisms, including ion release, surface reactions, and modification of soil colloid properties. Zinc oxide nanoparticles (ZnO NPs) dissolve preferentially in acidic soils, releasing Zn<sup>2+</sup> ions that increase soil pH and affect the bioavailability of other nutrients, including phosphorus, iron, and manganese (<xref ref-type="bibr" rid="B72">Li et&#xa0;al., 2025</xref>).</p>
<p>Iron-based nanoparticles can reduce soil pH through oxidation reactions and electron transfer processes, potentially mobilizing heavy metals and altering phosphorus availability through changes in adsorption-desorption equilibria. These geochemical alterations may have long-lasting effects on soil fertility and plant nutrient availability, requiring careful monitoring and management (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
</sec>
<sec id="s6_2_5">
<label>6.2.5</label>
<title>Organic matter interactions</title>
<p>Nanoparticles demonstrate strong binding affinity for soil organic matter (SOM) through electrostatic interactions, hydrophobic associations, and coordination bonding. This binding can either stabilize organic carbon by protecting it from microbial decomposition (positive effect) or interfere with natural decomposition processes by blocking enzyme access or altering microbial community structure (negative effect), with significant long-term implications for soil carbon sequestration and fertility maintenance (<xref ref-type="bibr" rid="B65">Khort et&#xa0;al., 2021</xref>).</p>
</sec>
</sec>
<sec id="s6_3">
<label>6.3</label>
<title>Environmental persistence and bioaccumulation assessment</title>
<sec id="s6_3_1">
<label>6.3.1</label>
<title>Nanoparticle fate and transformation in soil systems</title>
<p>Understanding the environmental fate of nanoparticles requires a comprehensive evaluation of their persistence, transformation, and mobility in soil systems under varying environmental conditions (<xref ref-type="bibr" rid="B175">Wei et&#xa0;al., 2024</xref>). Metal nanoparticles, particularly silver and copper-based formulations, demonstrate high persistence in soil with half-lives ranging from 6 months to several years depending on soil pH, organic matter content, moisture levels, and microbial activity (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
<p>Metal oxide nanoparticles, including TiO<sub>2</sub> and ZnO, demonstrate extreme environmental persistence with minimal degradation over 2-5 year monitoring periods, raising concerns about long-term accumulation and potential ecological effects. Carbon-based nanoparticles, including fullerenes and carbon nanotubes, show variable persistence ranging from months to years, with degradation rates influenced by soil organic content, microbial diversity, and environmental conditions (<xref ref-type="bibr" rid="B187">Zaman et&#xa0;al., 2025</xref>).</p>
<p>Nanoparticles undergo various transformation processes in soil environments that affect their bioavailability, toxicity, and environmental fate. Dissolution processes result in the release of ionic forms, particularly relevant for Ag, Cu, and Zn nanoparticles, which may exhibit different toxicological profiles compared to their nano-form counterparts (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
<p>Aggregation and agglomeration processes lead to the formation of larger particle clusters that alter surface area-to-volume ratios, reducing bioavailability while potentially affecting soil physical properties. Surface coating with natural organic matter modifies nanoparticle surface properties, potentially reducing toxicity while affecting transport and bioavailability (<xref ref-type="bibr" rid="B65">Khort et&#xa0;al., 2021</xref>).</p>
</sec>
<sec id="s6_3_2">
<label>6.3.2</label>
<title>Bioaccumulation potential and trophic transfer</title>
<p>Plant uptake and translocation studies demonstrate variable patterns across plant species and nanoparticle types, with implications for food chain contamination and human exposure. Root accumulation typically represents the highest concentration point, with 50-90% of absorbed nanoparticles retained in root tissues rather than translocated to aerial plant parts (<xref ref-type="bibr" rid="B53">Isibor et&#xa0;al., 2024</xref>).</p>
<p>Shoot translocation remains limited for most nanoparticle types, with only 5-30% of absorbed nanoparticles detected in above-ground tissues, though this varies significantly with nanoparticle size, surface coating, and plant species. Grain and fruit accumulation generally shows low but detectable levels (0.1-5% of absorbed nanoparticles) in edible tissues, raising questions about long-term dietary exposure implications (<xref ref-type="bibr" rid="B8">Alizadeh et&#xa0;al., 2025</xref>).</p>
<p>Soil invertebrates, including earthworms (<italic>Eisenia fetida</italic>) and springtails (<italic>Folsomia candida</italic>), demonstrate significant nanoparticle accumulation with bioconcentration factors ranging from 1.2 to 3.5, indicating potential for trophic transfer to higher levels. Limited studies on secondary consumers suggest potential transfer to birds and small mammals, though transformation and reduced bioavailability may limit bioaccumulation compared to persistent organic pollutants (<xref ref-type="bibr" rid="B168">Waalewijn-Kool et&#xa0;al., 2014</xref>).</p>
</sec>
</sec>
<sec id="s6_4">
<label>6.4</label>
<title>Phytotoxicity assessment at environmentally relevant concentrations</title>
<sec id="s6_4_1">
<label>6.4.1</label>
<title>Field application and physiological disruption</title>
<p>Field-scale concentrations of nanoparticles often differ significantly from laboratory test conditions, requiring careful evaluation of effects at environmentally relevant exposure levels. Even at low concentrations (0.1-10 mg/L in soil solution), nanoparticles can induce oxidative stress through increased reactive oxygen species (ROS) production, leading to cellular damage and reduced plant performance (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
<p>Membrane integrity disruption affects nutrient and water uptake efficiency, with consequences for plant growth and stress tolerance under field conditions. Photosynthetic efficiency reductions of 5-25% have been documented at nanoparticle concentrations of 1-50 mg/kg soil, with implications for carbon fixation and overall plant productivity.</p>
<p>Root development inhibition, including reduced elongation and branching, affects nutrient acquisition capacity and may compromise plant establishment and growth under field conditions (<xref ref-type="table" rid="T5"><bold>Table&#xa0;5</bold></xref>) (<xref ref-type="bibr" rid="B26">Djanaguiraman et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s6_4_2">
<label>6.4.2</label>
<title>Species-specific sensitivity and agricultural implications</title>
<p>Crop sensitivity to nanoparticles varies significantly among species and cultivars, requiring species-specific risk assessment approaches. Leguminous crops, including soybean and pea, demonstrate high sensitivity due to their dependence on rhizobial nitrogen-fixing symbionts that may be disrupted by nanoparticle exposure.</p>
<p>Leafy vegetables, including lettuce and spinach, show rapid nanoparticle uptake through extensive root systems, potentially leading to higher accumulation levels in edible tissues. Cereal crops, including wheat and rice, demonstrate moderate sensitivity with significant species-specific variations in uptake, translocation, and toxic response thresholds (<xref ref-type="bibr" rid="B121">Raza et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s6_4_3">
<label>6.4.3</label>
<title>Field-scale reality versus laboratory assessments</title>
<p>Laboratory studies often employ nanoparticle concentrations 10-1000 times higher than realistic field applications, potentially overestimating toxicity risks. However, localized concentration hotspots near application sites can reach levels of concern, with soil surface accumulation showing 2-10 times higher concentrations in the top 5 cm compared to deeper soil layers.</p>
<p>Rhizosphere concentrations may be 1.5-5 times higher than bulk soil levels due to root exudate interactions and localized accumulation processes. Seasonal variations in nanoparticle concentrations result from leaching, plant uptake, microbial degradation, and reapplication cycles, requiring long-term monitoring to understand exposure patterns (<xref ref-type="bibr" rid="B180">Yang et&#xa0;al., 2025</xref>).</p>
</sec>
</sec>
</sec>
<sec id="s7">
<label>7</label>
<title>Regulation and sustainability</title>
<sec id="s7_1">
<label>7.1</label>
<title>Regulatory frameworks and policy gap analysis</title>
<sec id="s7_1_1">
<label>7.1.1</label>
<title>Current regulatory inadequacies</title>
<p>Existing pesticide and fertilizer regulations prove inadequate for nanomaterials due to their focus on chemical composition rather than size-dependent properties that determine nanoparticle behavior and effects. Traditional regulations fail to account for unique nano-scale properties, including high surface area-to-volume ratios, quantum effects, and enhanced reactivity that distinguish nanoparticles from their bulk counterparts (<xref ref-type="bibr" rid="B17">Bouhadi et&#xa0;al., 2025</xref>).</p>
<p>Standard ecotoxicity testing protocols may not capture nano-specific effects such as particle-cell interactions, intracellular uptake mechanisms, and subcellular localization that influence toxicity profiles. Current regulatory approval processes rely on short-term studies spanning weeks to months rather than the long-term assessments (years to decades) necessary to evaluate environmental persistence and chronic exposure effects (<xref ref-type="bibr" rid="B180">Yang et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s7_1_2">
<label>7.1.2</label>
<title>International regulatory inconsistencies</title>
<p>The European Union has implemented the most comprehensive regulatory approach through the REACH (Registration, Evaluation, Authorization, and Restriction of Chemicals) regulation, which requires the submission of nano-specific data for nanomaterials. However, implementation remains inconsistent across member states, and enforcement mechanisms require strengthening (<xref ref-type="bibr" rid="B43">Gupta et&#xa0;al., 2023</xref>).</p>
<p>The United States Environmental Protection Agency (EPA) continues developing nano-specific guidance documents, but implementation remains inconsistent across different agricultural applications and regulatory pathways. Developing countries face significant challenges in building regulatory capacity for nanotechnology oversight, which can create international trade implications, including the lack of harmonized standards creating trade barriers, regulatory shopping where manufacturers seek the least restrictive jurisdictions, and insufficient coordination on global risk assessment approaches (<xref ref-type="bibr" rid="B180">Yang et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s7_1_3">
<label>7.1.3</label>
<title>Critical regulatory development needs</title>
<p>Priority regulatory developments include nano-specific risk assessment protocols that account for size, surface area, reactivity, and environmental fate characteristics unique to nanomaterials. Standardized characterization requirements should mandate comprehensive physical-chemical characterization, including particle size distribution, surface area, surface charge, and coating composition for all nano-enabled agricultural products (<xref ref-type="bibr" rid="B87">Mittal et&#xa0;al., 2020</xref>).</p>
<p>Long-term environmental monitoring requirements should establish post-market surveillance systems for tracking environmental concentrations, ecosystem effects, and bioaccumulation patterns over extended periods. Lifecycle assessment integration should consider environmental impacts across production, use, and disposal phases of nanomaterial lifecycles (<xref ref-type="bibr" rid="B180">Yang et&#xa0;al., 2025</xref>).</p>
</sec>
</sec>
<sec id="s7_2">
<label>7.2</label>
<title>Risk mitigation strategies and sustainable implementation</title>
<sec id="s7_2_1">
<label>7.2.1</label>
<title>Design-based risk reduction approaches</title>
<p>Biodegradable nanoparticle development focuses on creating environmentally degradable formulations that maintain agricultural efficacy while reducing persistence concerns. Targeted delivery systems enable precision application strategies that minimize environmental exposure while maximizing agricultural benefits.</p>
<p>Surface modification techniques can reduce toxicity while maintaining efficacy through coatings that control release rates, reduce bioavailability, or enhance biodegradation. Temporal application strategies involve timing applications to minimize ecological impact during sensitive periods such as pollinator activity, spawning seasons, or critical growth phases (<xref ref-type="bibr" rid="B156">Suresh Kumar et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s7_2_2">
<label>7.2.2</label>
<title>Environmental monitoring and adaptive management</title>
<p>Environmental monitoring frameworks should include baseline assessment protocols for characterizing pre-application environmental conditions, real-time monitoring systems for continuous assessment of soil and water quality, and biological indicator programs using sensitive species as early warning systems.</p>
<p>Adaptive management protocols should establish clear criteria for modifying application practices based on monitoring results, stakeholder engagement processes for incorporating community concerns and scientific findings, and regular review cycles for updating risk assessments based on new scientific evidence (<xref ref-type="bibr" rid="B85">Meng et&#xa0;al., 2025</xref>).</p>
</sec>
</sec>
<sec id="s7_3">
<label>7.3</label>
<title>Recommendations for responsible development</title>
<sec id="s7_3_1">
<label>7.3.1</label>
<title>Integrated risk-benefit framework</title>
<p>The implementation of the precautionary principle necessitates adopting conservative measures until comprehensive safety data is available. Concurrently, stakeholder engagement ensures the involvement of farmers, environmental groups, consumers, and regulatory agencies in the decision-making process. This can be achieved by transparent communication, which involves openly sharing research findings, risk assessments, and acknowledgments of uncertainty with all parties involved (<xref ref-type="bibr" rid="B187">Zaman et&#xa0;al., 2025</xref>).</p>
</sec>
<sec id="s7_3_2">
<label>7.3.2</label>
<title>Research priority identification</title>
<p>Critical research needs include long-term field studies spanning multiple years and diverse environmental conditions, mechanistic toxicity studies elucidating cellular and molecular mechanisms of nanoparticle effects, and ecosystem-level studies examining community and population-level responses to chronic exposure (<xref ref-type="bibr" rid="B85">Meng et&#xa0;al., 2025</xref>).</p>
<p>Standardized testing protocol development should establish harmonized methods for nanoparticle characterization, environmental fate assessment, and ecological risk evaluation. International cooperation initiatives should promote shared research programs, harmonized regulatory standards, and technology transfer to developing countries (<xref ref-type="bibr" rid="B24">Das and Ingti, 2025</xref>).</p>
</sec>
</sec>
<sec id="s7_4">
<label>7.4</label>
<title>Long-term environmental sustainability assessment</title>
<sec id="s7_4_1">
<label>7.4.1</label>
<title>Ecosystem-level impact evaluation</title>
<p>Chronic exposure to nanoparticles could induce changes at the ecosystem level. These changes may include a reduction in biodiversity due to the preferential toxicity of nanoparticles to sensitive species, shifts in functional groups that affect decomposer and nutrient cycling communities, and the development of resistance in exposed organisms. Consequently, these alterations have the potential to disrupt essential ecosystem services, such as pollination, biological pest control, and nutrient cycling, which are critical for maintaining sustainable agricultural systems (<xref ref-type="bibr" rid="B167">Vita et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s7_4_2">
<label>7.4.2</label>
<title>Cumulative and interactive effects assessment</title>
<p>Nanoparticles interact synergistically with other environmental stressors, including climate change, where changes in temperature and moisture affect nanoparticle behavior and toxicity profiles. Chemical pollution interactions may result in additive or synergistic effects when nanoparticles combine with pesticides, heavy metals, or other contaminants.</p>
<p>Habitat fragmentation reduces ecosystem resilience to nanoparticle stress, while agricultural intensification creates multiple stressor scenarios where nanoparticles interact with conventional agricultural inputs (<xref ref-type="bibr" rid="B85">Meng et&#xa0;al., 2025</xref>).</p>
</sec>
</sec>
</sec>
<sec id="s8" sec-type="conclusions">
<label>8</label>
<title>Conclusion and future perspectives</title>
<p>Nanotechnology presents a promising tool for advancing sustainable agriculture. It enhances plant tolerance to abiotic stresses, such as salinity and drought, improves nutrient uptake efficiency, and bolsters natural defense mechanisms. Furthermore, nanobiosensors enable the early detection of environmental stressors, while green synthesis methods offer an environmentally friendly production pathway for nanoparticles. However, a thorough understanding of nanoparticle interactions with plant systems and the environment, particularly those involving complex biological processes, remains essential.</p>
<p>A comparison with traditional approaches reveals significant advantages for nanotechnology in terms of efficiency, precision, and a reduced environmental footprint. Nevertheless, major challenges such as high production costs, safety concerns, and regulatory gaps must be addressed for its successful implementation. Evidence suggests that relying on any single approach is insufficient to meet modern agricultural demands; instead, the future lies in the intelligent integration of nanotechnology with traditional practices to develop productive and sustainable farming systems.</p>
<p>Realizing this potential while minimizing risks requires a responsible, research-driven strategy. This must include broad stakeholder engagement, adaptive management practices, and robust regulatory frameworks to ensure safe and equitable adoption. Environmental concerns and long-term safety issues demand urgent scientific attention and proactive policy development. Implementing comprehensive risk-assessment frameworks, establishing robust environmental monitoring programs, and fostering international cooperation will be critical to ensuring that nanotechnology benefits agriculture without compromising human or ecosystem health.</p>
<p>Looking ahead, research priorities should focus on developing sustainable synthesis methods, designing smart delivery systems, and elucidating the molecular mechanisms of nanoparticle-plant interactions. Integrating nanotechnology with fields such as genomics, proteomics, and metabolomics will provide a holistic understanding of plant responses to stress, which is essential for developing climate-resilient crops. Through this interdisciplinary, precautionary, and integrated approach, nanotechnology can significantly boost crop yields, even on marginal lands, thereby contributing to global food security while protecting environmental health.</p>
</sec>
</body>
<back>
<sec id="s9" sec-type="author-contributions">
<title>Author contributions</title>
<p>OA-D: Funding acquisition, Writing &#x2013; review &amp; editing. LA: Writing &#x2013; review &amp; editing. JA-K: Writing &#x2013; review &amp; editing.</p></sec>
<ack>
<title>Acknowledgments</title>
<p>The authors extend their appreciation for the support of the Deanship of Scientific Research, Vice Presidency for Graduate Studies and Scientific Research, King Faisal University, Saudi Arabia [Grant No. KFU253401].</p>
</ack>
<sec id="s11" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p></sec>
<sec id="s12" sec-type="correction-statement">
<title>Correction note</title>
<p>A correction has been made to this article. Details can be found at: <ext-link xlink:href="https://doi.org/10.3389/fpls.2025.1728599" ext-link-type="uri">10.3389/fpls.2025.1728599</ext-link>.</p></sec>
<sec id="s13" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p></sec>
<sec id="s14" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p></sec>
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<fn id="n1" fn-type="custom" custom-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/755940">Rajesh Kumar Singh</ext-link>, Guangxi Academy of Agricultural Science, China</p></fn>
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<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/399986">Murali M</ext-link>, University of Mysore, India</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1567146">Muhammad Ather Nadeem</ext-link>, University of Sargodha, Pakistan</p></fn>
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