Translate this page into:
Ethylene-Mediated Regulation of Wheat (Triticum Aestivum) Physiology in Association with Trichoderma Spp. Under Lead-Induced Heavy Metal Stress
* Corresponding author: Dr. Wahid Mohammad Ansari, Department of Botany, Zakir Hussain Delhi College, Jawaharlal Nehru Marg, SKD Basti, Press Enclave, Ajmeri Gate, New Delhi, India. mwahidansari@zh.du.ac.in
-
Received: ,
Accepted: ,
How to cite this article: Singh L, Yadav P, Muley D, Ansari WM. Ethylene-Mediated Regulation of Wheat (Triticum Aestivum) Physiology in Association with Trichoderma Spp. Under Lead-Induced Heavy Metal Stress. J Qassim Univ Sci. doi: 10.25259/JQUS_10_2026
Abstract
Heavy metal accumulation, particularly lead, poses a significant threat to agricultural productivity and food safety, necessitating effective amelioration strategies. The non-degradable nature of lead ensures its persistence in soil and water, leading to long-term environmental and health risks through bioaccumulation in the food chain. Triticum aestivum, a vital global crop, is highly susceptible to lead stress. Lead toxicity severely disrupts plant physiological and biochemical processes, including reduced photosynthetic rates, altered stomatal conductance, decreased nutrient uptake, and impaired seed germination, ultimately leading to significant yield losses. This stress often induces oxidative damage through reactive oxygen species overproduction. Phytohormones like ethylene are crucial regulators of plant responses to abiotic stress, influencing photosynthetic efficiency and antioxidant systems. Ethylene production can be enhanced under metal stress, and its signaling pathways are implicated in plant adaptation and tolerance to toxic metals. Simultaneously, beneficial fungi such as Trichoderma spp. are known for their plant growth-promoting abilities and stress mitigation. Trichoderma can modulate plant hormone pathways, including ethylene, and some species produce ACC (1-Aminocyclopropane-1-carboxylic acid) deaminase, which reduces ethylene levels, thereby enhancing plant tolerance and root development. This research investigates the synergistic interactions of exogenously applied ethylene and Trichoderma spp. on wheat’s physiological and biochemical responses under lead stress. We hypothesize that while lead stress will severely disrupt photosynthetic proficiency and increase oxidative damage, both ethylene and Trichoderma will individually enhance stress resistance. Their co-application is predicted to synergistically boost photosynthesis, enhance antioxidant defense mechanisms, and decrease lead accumulation, offering an eco-friendly approach to alleviate heavy metal stress in wheat.
Keywords
Antioxidant defense
Ethylene
Lead toxicity
Trichoderma
Triticum aestivum
INTRODUCTION
Heavy metal toxicity and especially lead is a major issue threatening the world’s agriculture and food security through compromising the crops growth and production.[1] Lead poisoning, which is a widespread environmental problem, has a negative impact on plant metabolism due to the disturbed redox balance, photosynthesis, and cell division, which in turn reduces the biomass accumulation and growth of such crops like wheat (Triticum aestivum).[2,3] Particularly, high levels of lead may disrupt the major metabolic activities, such as electron movement, disrupt cell organelle health, increase membrane fragility, and deter photosynthetic performance, which directly affect grain quality in cereal.[4] The build-up of lead in plant tissues also leads to the decreased germination percentage and the lack of the physiological characteristics necessary to develop a plant.[2] The high rate of lead pollution, which often originates in human-made processes (mining and industrial releases), is causing lead to be deposited in the agricultural soil and irrigation water, thus causing a serious threat to the sustainability of agricultural operations.[5] Therefore, the absorption and retention of these poisonous metals by plants, especially staple foods like wheat, have dire health effects on humans because they find their way into the food chain.[6] In the case of about 36 per cent of the world population, wheat is a basic food product and therefore the contamination of it with lead becomes a central issue of societal health.[2] High levels of lead interfere with protein synthesis and enzyme activity causing inhibited protein biosynthesis and impaired cellular activity in wheat.[7] Besides, long-term intake of lead-contaminated wheat can lead to the development of lead in the human body, which increases the threat of serious health consequences including neuro-toxicity, hypertension, and renal damage in the most susceptible people.[8] This widespread environmental issue has led to the need to come up with strong mechanisms of curbing lead toxicity in wheat and improving its tolerance to heavy metal stress bearing in mind that the plant is very susceptible to high concentration of heavy metals.
Overview of plant stress responses
Heavy metal stress of plants, such as leads triggers complex defense mechanisms aimed at counteracting the toxicity of heavy metals; these pathways include physiological and biochemical changes.[9] These adaptations include antioxidant defense system activation, gene expression changes on stress-related proteins, and phytohormone signaling pathway modulation to ensure cell integrity and functionality.[5,10] As an example, wheat plants have evolved systems to prevent the internalization of heavy metals or to become resistant to heavy metals by increasing the activity of antioxidant enzymes, ion homeostasis control system, and the synthesis of stress proteins.[7] Also, the synthesis of secondary metabolites, such as phytochelatins and metallothioneins, used to chelate and trap lead ions, may be altered by plants and, therefore, lead to lower availability and toxicity of these lead ions in plant tissues.[11] These advanced defense systems, including vacuolar compartmentalization and ionomic redistribution via specialized transporters, slow down the translocation of lead to grains, and although spared by these processes, the primary goal of the process is cell protection and cell retention in vegetative tissues.[8] This multifaceted success of protective mechanisms ensures that the plants survive and strive to reduce the effect of lead on essential metabolic processes; still, lead can have a very strong effect on growth and productivity [Figure 1].[12] On top of that, high levels of lead in plant tissues can have adverse effects on plant growth, development, and yield of different crop species by changing the metabolic processes, enzyme activities, and stability of biological membranes.[13] These harmful effects spread to the fundamental physiological processes of photosynthesis and respiration and result in impaired energy production and the overall vitality of plants.[5] Moreover, although various studies managed to explain the different expression of any gene when stressed by heavy metal in wheat, the central key genes, proteomic, metabolomic, and ionomic changes, and their interactions are not properly understood.[7]
Role of phytohormones in stress adaptation
Phytohormones comprise key plant growth and development regulators, which enable plants to acquire their optimal physiological condition regarding their genetic potential.[14] Phytohormone signaling pathways regulation to stimulate and amplify adaptive responses is extremely vital in the context of heavy metal stress.[15] This is represented by molecular, biochemical, and physiological effects: increases the activity of metal transporters, increases the activity of non-enzymatic antioxidants, and the production of phytohormones to enhance the antioxidative defense system.[16] Among these hormones, ethylene has a central role in regulating the reaction of plants toward abiotic stresses such as heavy metals via complex cross-signaling with other hormonal processes, finally coordinating the formation of strong defense responses.[17]
Ethylene signaling pathways in plants
The ethylene signaling pathways in plants are one of the central regulatory axes of plants in the context of plant resilience to various abiotic stress induced by environmental factors, such as heavy metal toxicity. The ethylene gaseous phytohormone controls physiological and developmental processes, including root growth, stem elongation, and stomata opening. The downstream effects of ethylene may determine metabolic adaptations and antioxidant responses in relation to reactive oxygen species (ROS) signaling.[18] It has been experimentally proven that ethylene could alleviate oxidative stress by stabilizing the formation of H2O2 (Hydrogen peroxide) in plants treated with heavy metals, increasing the cellular protection system.[10] Also, exposure to metals like lead triggers the production of enzymes producing ethylene and activates genes such as ACS and EIN2, leading to stress tolerance[Figure 2].[17] The interaction between ethylene and the ROS is crucial, and the ROS produced during stress can control hormonal processes, and ethylene aids the synthesis of protective enzymes that conserve proteins and lipids.[17] Ethylene in turn has been cited as a major contributor to the overall network of plant stress responses, and in effect, it interacts with other phytohormones to ensure adaptative measures to heavy metal put up and shut are finely adjusted.[14] Its ability to regulate the antioxidant systems and cause adventitious root development highlights its adaptability to environmental stresses in plants.[18,19] Mechanisms through which ethylene coordinates these responses are worthy of thorough explanation, especially on the part of wheat, which is exposed to lead stress, and their interactions with other phytohormones, and subsequent genetic regulation [Figure 3].
Trichoderma spp. as plant growth promoters and stress alleviators
The Trichoderma spp. are used as multifactorial stress relievers and plant growth promoters. Such fungi are biocontrol agents and symbionts to plants, which increase growth and tolerance to abiotic and biotic stressors, including heavy metal toxicity, through regulation of phytohormone production, enhancing nutrient uptake, and stimulating systemic responses to defense, and so alleviate the negative effects of heavy metal toxicity, such as lead.[20] As it is, in Arabidopsis thaliana subjected to cadmium, exogenous treatment of ACC (1-Aminocyclopropane-1-carboxylic acid), a precursor of ethylene generated by Trichoderma spp., raises ethylene concentration, boosts root growth, and superoxide dismutase activity.[17] Moreover, the ACC deaminase is an enzyme that is produced by some Trichoderma isolates, and it breaks down ethylene, leading to a decrease in ethylene levels to cause growth and stress resistance.[4] An example of this enzymatic process, which is represented by T. asperellum MAP1, turns ACC into a-ketobutyrate and ammonia to avoid the production of ethylene that otherwise would hamper the development of plants in times of stress.[20] The strains improve the growth of plants and metal tolerance, especially against lead toxicity, by reducing the buildup of ethylene.[21] Regulatory properties of Trichoderma strains as ACC deaminase-producing favor the use of such organisms as bio-inoculants to promote sustainable agriculture.[20,21] Further, Trichoderma has the ability to secrete ethylene independently, and to modulate host ethylene biosynthesis signatures, stress responses, and development itself.[22] This dual modulation makes Trichoderma an advanced controller of the plant ethylene signaling system, allowing it not only to prevent disastrous ethylene bursts but also to maintain the basal ethylene concentration, which is essential in terms of adapting to the stress of lead.[23] In addition to the modulation of ethylene, Trichoderma spp. also produce auxins (indole-3-acetic acid) that subsequently promote root growth in stressful heavy metal environments through their interaction with ethylene pathways.[24] On the same note, endophytic fungi, including certain species of Trichoderma, have been found to reduce multi-metal toxicity in crops, such as soybean, through gibberellin release, facilitating growth and making plants more tolerant.[25] Therefore, a combination of Trichoderma-ethylene regulation and the other phytohormone production pathway types leads to a whole enhancement of plant vitality and resistance to heavy metal toxicity.
Mechanisms of lead stress in wheat
The toxic effects of lead in wheat are complex and mainly centered on causing disruptions to the integrity of cells and the metabolic pathways that are important for growth and development. More precisely, it interferes with photosynthesis by causing damage to chloroplasts, inhibiting enzyme activity, and disrupting electron transport, thus reducing chlorophyll content and the overall efficiency of photosynthesis.[26] Additionally, lead toxicity triggers oxidative stress through the enhanced production of ROS, which results in lipid peroxidation, alterations in proteins, and DNA lesions.[14] This ROS accumulation leads to loss of cellular membrane integrity, damage to organelles, and eventually increases electrolyte leakage and cell death, which highly affects plant viability and yield.[4,2] Furthermore, lead toxicity disturbs nutrient uptake and transportation due to root architecture damage and disturbance in the functioning of nutrient transporters, which results in the deficiency of essential minerals required for wheat growth.[27] The occurrence of lead in soil may reduce microbial activities and root respiration, which further impairs nutrient acquisition and utilization by plants.[4] Furthermore, the inhibition of roots’ elongation and rooting architecture under lead stress might be mediated through changing the signaling mechanism of auxin and cytokinin, and further results in complications in nutrient acquisition.[28] Lead, when accumulated above permissible levels, can also reduce cation exchange in roots, thus restricting the uptake of vital elements such as K+, Ca 2+, Mg 2+, Fe 2+, and Zn 2+.[2] This impairment of mineral uptake consequently compromises the overall nutritional status and metabolic efficiency of the wheat plant, ultimately impacting crop yield and quality.[5] The detrimental effects of lead are further compounded by its ability to decrease the soil’s osmotic potential, leading to reduced water absorption and inducing osmotic stress, which subsequently triggers morphological changes in plants.[29] Such morphological alterations often include stunted growth, reduced biomass, chlorosis, and necrosis, collectively contributing to significant yield losses in lead-contaminated agricultural systems.[4,11,30,31] Furthermore, lead’s high affinity for protein N- and S-ligands damages the photosynthetic system, reducing both photosynthetic and respiratory rates by diminishing mitochondrial cristae density and subsequently lowering oxidative phosphorylation potential.[11] These physiological disturbances collectively contribute to a significant decline in overall plant productivity and biomass accumulation under lead stress conditions.[11,26] The impact of lead extends to disrupting enzyme activities essential for metabolic processes, thereby compromising cellular metabolism and overall plant functionality.[30] The toxicity of lead significantly impairs chlorophyll synthesis, leading to compromised photosynthesis, reduced respiration, and the inhibition of ATP synthase, ultimately initiating oxidative stress that can result in plant mortality.[6] These pervasive disruptions underscore the urgent need for effective remediation strategies to alleviate lead toxicity in agricultural systems and ensure sustainable wheat production.[32,11,33,26]
Antioxidant defense and reactive oxygen species management
Lead is the main cause of heavy-metal stress that significantly increases reactive oxygen species (ROS) production in plants causing oxidative stress and dysfunction of cellular processes.[34] Such an oxidative burst leads to the mobilization of a strong antioxidant defense system, enzymatic and non-enzymatic, to neutralize such harmful substances in order to maintain homeostasis in the cell.[4] Enzymatic antioxidants (superoxide dismutase, catalase, and diverse peroxidases) play a central role in the scavenging of ROS, whereas the non-enzymatic antioxidant (ascorbate, glutathione, and tocopherols) are supplementary agents against oxidative stress.[11,26] A fine balance between the antioxidant defense and ROS creation is something that plant life cannot be without during lead toxicity circumstances because any disproportion can trigger serious physiological injuries and limited growth.[35,26] It has been reported that lead toxicity induces the changes in the activity of many antioxidant enzymes and non-enzymatic substances and reduces the ability of the plant to eliminate the ROS.[26] To provide an example, with an increase in lead levels, the activity of antioxidant enzymes significantly decreases, and at the same time, the levels of malondialdehyde and ROS increase, which is an indicator of increased oxidative stress and lipid peroxidation.[36] On the contrary, minimal amounts of heavy metals, such as lead, may sometimes induce metabolic processes and stimulate growth-related enzymes.[4] However, when ROS is not controlled, it may eventually result in oxidative stress, which inevitably causes cellular damage and programmed cell death.[9] Plants have developed more advanced means to overcome the adverse consequences of lead-induced free radicals and to avoid their intracellular retention through the means of activating antioxidant defense.[37] This defense system is important since stress observed leads to expression and activity of antioxidant enzymes in wheat, hence the plant is better prepared to withstand the resultant oxidative stress.[1] The process of dismutation superoxide into molecular oxygen and hydrogen peroxide with superoxide radicals is catalyzed by superoxide dismutase, which is deemed essential to prevent the accumulation of highly reactive superoxide species.[11] This cytotoxic agent is then neutralized by catalase and peroxidases into water and oxygen and prevents additional damage of the cellular components.[11] There is an essential role played by the intricate enzymatic detoxification cascade, as well as non-enzymatic antioxidants, in redox-homeostasis maintenance of cells under heavy-metal stress.[38,39]
Secondary metabolite involvement in lead detoxification
Secondary metabolites have been found largely useful in alleviating lead toxicity due to their strong non-enzymatic antioxidant capability to scavenge reactive oxygen species and alleviate oxidative damage.[40] According to recent transcriptomic studies, secondary metabolism has a dual mechanism as it functions simultaneously as a scavenging system and as a signaling pathway which activates the production of phytochelatins and Srich thiolate-rich peptide, which regulate metal homeostasis.[41,30] Moreover, the presence of a symbiotic relationship with the mycorrhizal fungi improves soluble phenolics concentration in roots that combines with glutathione to create an effective physiological defense mechanism against lead toxicity.[4] This microbial-dependent regulation additionally introduces control over the biosynthesis of the endogenous phytohormones. Beneficial symbionts modify the synthesis of growth regulators such as indole-3-acetic acid and compounds linked to stress, such as salicylic acid, to maintain normal growth in the presence of lead.[42] Microbial associations also exhibit the ability to receive particular metabolites that are secreted by fungal symbionts in the rhizosphere, enabling the uptake of these microbially released phytohormones to activate calcium-dependent protein kinases and trigger downstream stress-responsive genes, osmotins, and heat shock proteins.[42] Although these metabolic pathways are promising to understand, further research is necessary in this area to have a better idea about the phenomenon that takes place in the symbiotic relationships and their impact on the tolerance of lead by the plant.[11] Further studies should hence be aimed at explaining the molecular playing field of plant-microbe interactions to streamline phytoremediation of lead-contaminated soils.[11,10] Certain categories of these compounds, such as flavonoids, alkaloids, and phenols, have been found to regulate plant development and pigment formation, and at the same time carry out antioxidant activity to suppress certain enzyme activities during stress periods.[40]
Crosstalk between hormonal and microbial pathways
The complexity of the interaction between plant hormonal signal transduction and a positive reciprocal relationship with specific beneficial microbes, specifically with Trichoderma species, provides a paradigm shift in understanding improved plant tolerance to heavy-metal stress. Modulating ethylene by Trichoderma in association with other phytohormones can coordinate the plant responses to result in toxicity, in which growth and defense systems are up-regulated.[32] This includes Trichoderma-mediated changes in hormonal balance, including changes in levels of auxin and cytokinin, which in turn act in concert with ethylene to adjust root development and stress-adaptation. The highly intricate interaction of Trichoderma and plant hormones also goes beyond affecting the water balance and mineral nutrient uptake, which is grossly affected by lead toxicity.[26] This complex of interactions between hormones and microbes allows for a more active and integrated physiological reaction, which allows plants to be more tolerant and, possibly, to correct the lead-polluted conditions [Table 1]. These synergistic interactions, the potential to exploit Trichoderma plant-symbioses as a sustainable biotechnological intervention to reduce the adverse effects of heavy-metal stress on crop productivity, are highlighted.[4,11] Understanding these complicated relationships would be crucial in developing sustainable agricultural measures that would maximize crop production and guarantee food security in areas with heavy contamination of heavy metals.
| Plant species | Stress/treatment | Major findings | Reference |
|---|---|---|---|
| Triticum aestivum L. | Lead stress | Lead exposure significantly increased ROS generation, lipid peroxidation, and reduced growth and chlorophyll content. | [1] |
| Triticum aestivum L. | Lead toxicity | Lead caused inhibition of photosynthetic activity and disturbance in mineral nutrient uptake. | [4] |
| Triticum durum Desf. | Lead contamination | Lead accumulation reduced biomass production and altered metal partitioning between roots and shoots. | [13] |
| Triticum aestivum L. | Lead + polyamine treatment | Polyamine supplementation enhanced antioxidant enzyme activity and mitigated oxidative stress. | [12] |
| Arabidopsis thaliana (L.) Heynh. | Ethylene signaling under metal stress | Ethylene signaling regulates stress-responsive gene expression and growth–defense balance. | [18] |
| Triticum aestivum L. | Lead-induced oxidative stress | Wheat tolerance correlated with increased antioxidant enzyme activity and membrane stability. | [5] |
| Triticum aestivum L. | Lead uptake and translocation. | Lead accumulation disrupted nutrient homeostasis and reduced photosynthetic efficiency. | [26] |
| Triticum aestivum L. | Lead + endophytic fungi | Endophytic inoculation reduced Lead translocation and enhanced antioxidative defense. | [2] |
| Triticum aestivum L. | Temporal Lead exposure | Growth-stage–dependent Pb accumulation patterns identified in wheat | [8] |
| Oryza sativa L. | Lead stress | Lead exposure induced oxidative damage and altered expression of metal transporter genes | [43] |
| Zea mays L. | Lead toxicity | Lead stress suppressed photosystem II efficiency and reduced nutrient assimilation. | [35] |
| Brassica juncea (L.) Czern. | Lead accumulation | High Lead accumulation associated with enhanced phytochelatin synthesis | [39] |
| Glycine max (L.) Merr. | Lead stress | Lead altered nitrogen metabolism and reduced nodulation efficiency | [36] |
| Solanum lycopersicum L. | Lead exposure | Lead reduced fruit yield and disrupted antioxidant homeostasis | [38] |
| Oryza sativa L. | Lead + microbial inoculation | Beneficial microbes reduced Lead uptake and improved antioxidant capacity | [21] |
ROS: Reactive oxygen species
Current challenges and research gaps
No empirical studies so far have been conducted that investigated the combination of exogenous ethylene and Trichoderma species as applied to lead tolerance in wheat. The indirect evidence suggests that there is a likely synergistic effect, as the two agents induce overlapping stress-response transduction. Despite the excesses that have been made in understanding lead toxicity, the roles of ethylene signaling and microbial interventions remain not fully established, hence restricting the practical application of the processes in wheat production. Further studies will be needed to shed more light on the exact molecular mechanisms through which Trichoderma spp. Regulate the ethylene signaling to promote greater lead tolerance. Specifically, the transcription of the genes of detoxification and antioxidant protection needs to be examined in more detail. Besides, additional demarcation of the transcriptional activities organized by the Trichoderma-ethylene interaction, which determines the heavy-metal sequestration and efflux, will also be necessary to design biofortification approaches. There should be optimization of application modes and inoculum concentrations of Trichoderma to maximize the utility of the organism at different environmental factors, type of soil, and degree of contamination. Along with this, it is necessary to conduct functional genomics and proteomics research to be able to discover novel biomarkers of lead tolerance and to have means of genetic engineering to be able to select more robust wheat cultures.
Future applications and prospects
The comprehensive understanding of molecular mechanisms will allow developing novel biotechnological programmes, such as the genetic engineering of wheat to increase lead removal ability, improve stress tolerance, and, consequently, sustain food production and food security. Using newer genome editing technologies like CRISPR/Cas9, targeting induced local lesions in genomes (TILLING), and genome-wide associated studies (GWAS) will allow further subdivision of the complex molecular processes responsible of plant to metal interactions and help to develop plants more resistant to metals. Field trials and integrative multi-omics models that involve measurements on any type of biomass, evaluation of pollutant deposition, and physiological control should be examined via rigorous field testing, and through the combination of these outcomes, the interactions between growth, contamination, and resilience in contaminated crops would be fully comprehended. Such a comprehensive measure will produce operational data to use in rolling out sustainable remediation measures and strengthening the plants in metal-stressed environments.
CONCLUSION
This study reveals that the ethylene signaling and formation of Trichoderma colonization are potential paradigms to alleviate lead-induced stress, which highlights the possibilities of biotechnology advancements in the field of sustainable agriculture. The proposed framework is not just a response to the urgent task of the heavy-metal presence but also is in line with the larger environmental sustainability and food-security goals because it enhances plant resistance and utilizes Phyto transformation as a response method. All these objectives will entail the development of multidisciplinary solutions that will integrate biological, biotechnological, and physicochemical approaches to alleviate heavy-metal contamination and protect environmental health.
Acknowledgement
P. Yadav acknowledges the receipt of Junior Research Fellowship and Senior Research Fellowship from the University Grants Commission (UGC), Govt. of India. We sincerely apologies to our contemporaries whose work could not be discussed in this article due to space restrictions.
Author’s contribution
MWA and PY: Manuscript - design; LS: Literature review, draft; MWA, DM and PY: Finalize the manuscript.
Ethical approval
Institutional Review Board approval is not required.
Declaration of patient consent
Patient consent is not required as no patients are involved in the study.
Financial support and sponsorship
Nil.
Conflicts of interest
There are no conflicts of interest.
Use of artificial intelligence (AI)-assisted technology for manuscript preparation
The authors confirm that there was no use of artificial intelligence (AI)-assisted technology for assisting in the writing or editing of the manuscript, and no images were manipulated using AI.
References
- Lead-induced oxidative stress and role of antioxidant defense in wheat (Triticum aestivum l.) Physiol Mol Biol Plants. 2020;26:793-802.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Bio-stimulating effect of endophytic Aspergillus flavus AUMC 16068 and its respective ex-polysaccharides in lead stress tolerance of triticum aestivum plant. Sci Rep. 2024;14:11952.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Impact of gibberellic acid GA3, quantum dot biochar, and rhizosphere bacteria on fenugreek plant growth and stress responses under lead stress. Sci Rep. 2024;14:29612.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Lead toxicity in cereals: Mechanistic insight into toxicity, mode of action, and management. Front Plant Sci. 2021;11:587785.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Adaptive mechanisms of wheat cultivars to lead toxicity through enhanced oxidative defense, ionomic redistribution, and anatomical modifications. J Plant Physiol. 2024;303:154370.
- [CrossRef] [PubMed] [Google Scholar]
- Enhancing wheat productivity and reducing lead uptake through biochar, bentonite, and rock phosphate integration. Sustainability. 2025;17:3491.
- [CrossRef] [Google Scholar]
- Multi-omics uncover the mechanism of wheat under heavy metal stress. Int J Mol Sci. 2022;23:15968.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Temporal dynamics of lead uptake in wheat: Identifying key growth stages for mitigation. J Agriculture Food Res. 2025;24:102430.
- [CrossRef] [Google Scholar]
- Comprehensive review of multiomics applications and remediation of plant heavy metal toxicity. Stress Biol. 2025;5:58.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Decoding oxidative stress regulation in food crops exposed to heavy metals: Interdisciplinary strategies for sustainable mitigation. Discov Sustain. 2025;6
- [CrossRef] [Google Scholar]
- Lead toxicity in plants: Mechanistic insights into toxicity, physiological responses of plants and mitigation strategies. Plant Signal Behav. 2024;19:2365576.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Physiological and biochemical responses of wheat (Triticum aestivum L.) plants to polyamines under lead stress. Innovare J Agric Sci. 2021;1:1-10.
- [CrossRef] [Google Scholar]
- Lead contamination affects the primary productivity traits, biosynthesis of macromolecules and distribution of metal in durum wheat (Triticum durum L.) Saudi J Biol Sci. 2021;28:4946-53.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Heavy metals and ethylene: Shaping plant responses through signaling. Planta. 2025;262:9.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Impact of heavy metal pollution in the environment on the metabolic profile of medicinal plants and their therapeutic potential. Plants (Basel). 2024;13:913.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Physiological and biochemical responses to heavy metals stress in plants. Int J Second Metabolite. 2024;11:169-90.
- [Google Scholar]
- Mechanisms of selected plant hormones under heavy metal stress. Pol J Environ Stud. 2021;30:497-50.
- [Google Scholar]
- To fight or to grow: The balancing role of ethylene in plant abiotic stress responses. Plants (Basel). 2021;11:33.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Uncovering the mechanisms of salicylic acid-mediated abiotic stress tolerance in horticultural crops. Front Plant Sci. 2023;14:1226041.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Molecular mechanisms of the 1-aminocyclopropane-1-carboxylic acid (ACC) deaminase producing trichoderma asperellum MAP1 in enhancing wheat tolerance to waterlogging stress. Front Plant Sci. 2021;11:614971.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Bacterial ACC deaminase: Insights into enzymology, biochemistry, genetics, and potential role in amelioration of environmental stress in crop plants. Front Microbiol. 2023;14:1132770.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Trichoderma: A game changer in the modern era of plant disease management. In: Challenges in plant disease detection and recent advancements Challenges in plant disease detection and recent advancements. IntechOpen; 2024.
- [Google Scholar]
- Assessment of Tunisian trichoderma isolates on wheat seed germination, seedling growth and fusarium seedling blight suppression. Microorganisms. 2023;11:1512.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Harnessing Trichoderma spp. for sustainable plant disease management: Mechanisms, metabolites and application strategies—A review. J Appl Pharm Sci. 2024;34:304-17.
- [Google Scholar]
- Metagenomics for mitigation of heavy metal toxicity in plants. In: Current omics advancement in plant abiotic stress biology Current omics advancement in plant abiotic stress biology. Elsevier; 2024. p. :383-96.
- [Google Scholar]
- Pb uptake, accumulation, and translocation in plants: Plant physiological, biochemical, and molecular response: A review. Heliyon. 2024;10:e27724.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Phytostimulants in sustainable agriculture. Front Sustain Food Syst. 2022;6
- [CrossRef] [Google Scholar]
- Deciphering lead tolerance mechanisms in a population of the plant species Biscutella auriculata l. from a mining area: Accumulation strategies and antioxidant defenses. Chemosphere. 2020;261:127721.
- [CrossRef] [PubMed] [Google Scholar]
- Global lead contamination in soils, sediments, and aqueous environments: Exposure, toxicity, and remediation. J Trace Elements. 2025;14:100259.
- [CrossRef] [Google Scholar]
- Impacts of metals and metalloids on plant physiology: A review. Middle East J Agric Res. 2022;11:838-931.
- [Google Scholar]
- Attenuation of lead toxicity by promotion of tolerance mechanism in wheat roots by lipoic acid. Cereal Res Commun. 2018;46:424-35.
- [CrossRef] [Google Scholar]
- Plant growth-promoting bacteria enhance photosynthesis, nodulation and root system architecture in lentil under lead toxicity. J Cent Eur Agric. 2022;23:582-91.
- [Google Scholar]
- Biochemical responses in Salvia limbata CA Mey. towards mercury and lead induced in vitro oxidative stress. Sci Rep. 2025;15:32119.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Signaling and crosstalk of rhizobacterial and plant hormones that mediate abiotic stress tolerance in plants. Front Microbiol. 2023;14:1171104.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Deciphering lead tolerance mechanisms in maize: A thorough analysis. Int J Biol Sci. 2025;7:38-42.
- [CrossRef] [Google Scholar]
- Effect of industrial wastewater on wheat germination, growth, yield, nutrients and bioaccumulation of lead. Sci Rep. 2020;10:11361.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Silicon and iron nanoparticles protect rice against lead (Pb) stress by improving oxidative tolerance and minimizing Pb uptake. Sci Rep. 2024;14:5986.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Effect of heavy metal stress on phenolic compounds accumulation in winter wheat plants. Molecules. 2022;28:241.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Heavy metal induced stress on wheat: Phytotoxicity and microbiological management. RSC Adv. 2020;10:38379-403.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Lead uptake, toxicity and mitigation strategies in plants. Plant Arch. 2021;21:712-21.
- [CrossRef] [Google Scholar]
- Lead induces oxidative stress in Pisum sativum plants and changes the levels of phytohormones with antioxidant role. Plant Physiol Biochem. 2019;137:121-9.
- [CrossRef] [PubMed] [Google Scholar]
- Phosphate solubilizing aspergillus Niger PH1 ameliorates growth and alleviates lead stress in maize through improved photosynthetic and antioxidant response. BMC Plant Biol. 2024;24:642.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]
- Melatonin alleviates lead-induced stress in rice through physiological regulation and molecular defense mechanisms. Sci Rep. 2025;15:34788.
- [CrossRef] [PubMed] [PubMed Central] [Google Scholar]