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An overview of fluoride stress: physiological and biochemical responses of plants.
Abstract
Original full-text study online at
https://www.tandfonline.com/doi/full/10.1080/15592324.2026.2706350#abstract
Fluoride, a hazardous air and soil contaminant, poses significant risks to agricultural productivity and plant health. Its accumulation in soil and subsequent uptake by plant roots and mesophyll cells interfere with critical morphological, physiological, and biochemical processes essential for growth and development. Fluoride toxicity affects germination, respiration, photosynthesis, mineral nutrition, enzyme activity, and reproduction, ultimately reducing crop yield. It disrupts cell signaling pathways, impairs antioxidant enzyme function (e.g., superoxide dismutase), and interacts antagonistically with calcium, a key element in fertilization and pollen tube growth. The resulting toxicity manifests as chlorosis, necrosis, growth inhibition, leaf and fruit abscission, and reduced seed production. This review provides a comprehensive evaluation of fluoride-induced stress in plants, highlighting recent advances in understanding its mechanisms and potential mitigation strategies. By synthesizing current findings, this study offers insights into developing fluoride-resistant crops and improving management practices to safeguard agricultural productivity in contaminated regions.
Graphical abstract
Keywords: Fluoride toxicity; plant responses; agricultural productivity; morpho-physiological effects; biochemical alterations
Introduction
Plants face many abiotic stresses such as salinity, alkalinity,Citation 1 fluoride toxicity,Citation2 micronutrient deficiency,Citation3 heavy metal stress,Citation 4 drought stress,Citation 5 and extreme temperatures. These environmental factors can significantly impact plant growth, development, and overall productivity.Citation 6 As a result, plants have evolved various adaptive mechanisms to cope with these stressors, including osmotic regulation, ion homeostasis, antioxidant defense systems, and metabolic adjustments.Citation 7 Additionally, advances in biotechnology have led to the development of genetically modified plants with enhanced tolerance to specific abiotic stresses, offering promising solutions for sustainable agriculture in challenging environments.Citation 8 , Citation 9 Fluoride toxicity in plants manifests as reduced growth and development, chlorosis, necrosis, leaf, flower, and fruit abscission, as well as decreased seed production.Citation 10 While significant research has focused on abiotic stresses such as salinity, drought, and heavy metal toxicity due to their widespread impact on crop productivity, fluoride toxicity remains relatively underexplored. This disparity can be attributed to the localized nature of fluoride contamination, which is primarily associated with industrial emissions, phosphate fertilizers, and groundwater pollution in specific regions. Additionally, the physiological and biochemical effects of fluoride toxicity are often subtle and gradual, making it less immediately apparent compared to more visibly damaging stresses like drought-induced wilting or salt-induced ion toxicity. Furthermore, fluoride does not have an essential physiological role in plants, unlike elements such as sodium or chloride, which complicates its study in plant stress biology.Citation 11 As a result, research on fluoride stress has lagged behind, despite its potential to disrupt key metabolic processes, limit crop yields, and pose food security risks in contaminated areas.Citation 12 Addressing this gap is crucial for developing mitigation strategies and ensuring sustainable agricultural practices in fluoride-affected regions.
This review study carefully assesses the literature regarding the morphological, physiological, and biochemical impacts of fluoride toxicity on crop plants in order to better understand and minimize the detrimental effects of fluoride contamination on agricultural ecosystems.Citation 13
Even while its concentration is thought to be relatively modest in relation to other elements, it is yet substantial enough to add to the crust’s overall makeup. It is predicted that fluoride concentrations in the mantle, which sits beneath the Earth’s crust, are 200 times higher than those in the crust.Citation 14 , Citation 15
There are several minerals that contain fluoride, but the most prevalent source is fluorite, or calcium fluoride. Cryolite, fluorspar, and apatite are some other minerals that include fluoride.Citation 16 Weathering processes naturally release fluoride into the environment from geological formations rich in fluoride-containing minerals. Fluoride gases, including silicon tetrafluoride (SiF4) and hydrogen fluoride (HF), are released into the atmosphere during volcanic eruptions. Downwind of volcanic vents (), these gases may contribute to the deposition of fluoride in soil and water bodies.Citation 17
As fluoride-rich minerals in rocks and soil weather, fluoride ions are released into surface and groundwater systems. The rate and amount of fluoride emission are influenced by the solubility of fluoride-containing minerals, which varies with geological circumstances.Citation 18 Surface water bodies, including lakes, rivers, and streams, can have differing fluoride concentrations based on the geology of their catchment areas. Fluoride leaching from nearby rocks and soil, as well as changes in aquifer recharge rates and flow patterns, all have an impact on groundwater fluoride levelsCitation 19 ().
Fluoride ions are discharged into surface and groundwater systems as a result of weathering fluoride-rich materials found in rocks and soil. The solubility of fluoride-containing minerals, which varies with geological conditions, affects the pace and amount of fluoride emission.Citation 19 , Citation 20 Based on the geology of their catchment areas, surface water bodies such as lakes, rivers, and streams can have varying quantities of fluoride. Groundwater fluoride levels are affected by aquifer recharge rates and flow patterns, fluoride leaching from surrounding rocks and soil, and other factors.Citation 21
Fluoride levels in the environment are raised by human activities such as the burning of fossil fuels, the usage of fluoride-containing goods (such as insecticides and fluoridated water), the smelting of aluminum, and the manufacturing of phosphate fertilizer.Citation 16
Figure 1. The dispersion and presence of fluoride in water sources, soil, and plant life.
Overview of fluoride stress on plants
Elevated fluoride ion levels in the surrounding environment can cause fluoride stress in plants, which can result in a variety of physiological, biochemical, and morphological alterations that might hinder plant growth and development.Citation 2 In order to mitigate the adverse consequences of fluoride stress on agricultural output and ecosystem health, it is imperative to comprehend the impacts of this stress on plants.
Fluoride (F) concentrations that are too high in soil or solution culture can negatively impact a number of plant growth and development processes, such as germination, root and shoot growth, chlorophyll degradation, and grain yield loss.Citation 22 , Citation 23 Symptoms like chlorosis, leaf tip burn, and necrosis are how these impacts show up. Elevated levels of fluoride within plant cells cause disturbances to physiological cycles, impede the movement of nutrients, and reduce water consumption, which in turn affects the establishment, growth rate, and photosynthetic activities of seedlings.Citation 24 Additionally, higher F levels lead to more reactive oxygen species (ROS) being produced, which damages membranes. Although a great deal of study has been done on how salinity (NaCl) affects plant germination and emergence, nothing is known about how F-contaminated seeds affect plant development at this point.
A small number of plant species, to varying degrees, have been the subject of studies examining the effects of F on germination, including rice, gram seed, tomato, wheat, maize, soybeans, sorghum, and beans. The entire range of fluoride’s (F) impact on different elements of plant physiology and metabolism has not been fully explored, but it is absorbed by plant roots and transported inside the plant. Although it is well known that F contamination greatly reduces plant production, its effects on the morphological traits, mineral content, and metabolic functions of plants have not been thoroughly studied. It has long been known that fluoride (F) is a metabolic inhibitor, interfering with a number of metabolic processes essential to the growth and development of plants.Citation 25 In conclusion, although it is widely known that F pollution has a negative impact on plant growth and yield, there is still much to learn and understand about the specific interactions and processes that occur at the morphological, mineral, and metabolic levels of plants. Fluoride (F) has long been recognized to be a metabolic inhibitor, interfering with several metabolic pathways necessary for plant growth and development.
Fluoride uptake and storage in plants
Plants absorb and accumulate fluoride (F) through a slow process that ultimately results in harmful reactions in the plants (). Plants are primarily exposed to F through two primary pathways: first, F deposited on leaves enters through stomata, and second, F diffuses passively into the roots through the soil and water.Citation 26
Table 1. Characteristics of plants and their tendency to accumulate fluoride ions (F–).
Previous research has shown that soil-derived F uptake is not as substantial as that from atmospheric air. After absorption, F enters the shoots through apoplastic and symplastic routes via xylem tissues. Furthermore, some of F is also transferred across biological membranes by hydrogen fluoride (HF), a tiny, neutral molecule that diffuses across cell membranes about seven times more quickly than fluoride ions (F). But our understanding of the exact mechanisms underlying F uptake into plant cells is still lacking.Citation 30
Depending on how much F moves from the soil to the roots and subsequently into the shoots, different plant sections accumulate varied amounts of this element. F accumulation is often highest in the roots and lowest in the leaves, seeds, and shoots. The distribution of F within the plant is reflected in this differential accumulation pattern, which also implies that different plant tissues are susceptible to F toxicity to differing degrees.Citation 31
Objectives of the review article
The review article’s goals regarding fluoride toxicity in plants include a thorough analysis of all the different facets associated with fluoride pollution and effects on plant health. First and foremost, the review seeks to offer a comprehensive grasp of the sources of fluoride pollution in the environment, with an emphasis on the effects on plant. It then aims to explore the morphological, physiological, and biochemical responses of plants to fluoride stress in order to clarify the fundamental processes of fluoride toxicity. In addition, the review intends to appraise the significance of fluoride stress on agricultural output as well as the effects it has on numerous stages of plant growth and development, from germination to reproductive stages. It also seeks to investigate how environmental stressors like salinity and heavy metal pollution interact with fluoride stress and affect plant physiology and ecosystem dynamics as a whole.
Effect of fluoride stress on growth parameters
Plants and their numerous sections are affected by high internal fluoride (F) concentrations in almost all physiological and metabolic processes. In particular, plants are greatly influenced by high F levels during crucial growth stages like seed germination and early seedling growth.Citation 22 Plants that are exposed to high levels of F have shortened root-shoot length, decreased vigor, and hindered seed germination responses. Additionally, unbalanced nutrient intake brought on by F toxicity may result in fewer seeds being produced.Citation 32 Furthermore, excessive F concentrations inhibit biomass accumulation, which is essential for the healthful development of plants. Essentially, the disturbance of these growth characteristics indicates the severe adverse impacts of F toxicity on critical developmental processes, which in turn jeopardize the general growth and yield of plants.Citation 33
Fluoride stress reduces water intake and prevents the embryo’s cell division and expansion, which both have an impact on plant germination. As a result, the metabolic activity required for germination decreases. When plants are treated with fluoride, similar results have been shown in radish, wheat, mung beans, and Bengal gram.Citation 34 By suppressing the activity of the phytase enzyme, fluoride can also prevent the dephosphorylation of phytin molecules, which is necessary for germination. Furthermore, deficits in essential nutrients caused by fluoride can further hinder plants’ ability to respond to development. Fluoride also reduces the production of DNA, RNA, and proteins, which stops cell division and elongation and eventually stunts growth. Because of their enormous surface area and stomata, leaves and green stems are more vulnerable to fluoride deposition, which exacerbates the effects of fluoride stress on plant growth.Citation 35
Effect of fluoride stress on photosynthesis
Fluoride (F) adversely affects photosynthesis by disrupting multiple structural and biochemical components of the photosynthetic apparatus (). Chloroplasts serve as the primary sites of fluoride accumulation, making them particularly susceptible to fluoride-induced damage.Citation 24 Fluoride exposure reduces the concentrations of essential photosynthetic pigments, including chlorophyll a, chlorophyll b, carotenoids, and anthocyanins, thereby limiting light-harvesting efficiency and overall photosynthetic performance.Citation 2 Additionally, fluoride decreases the availability of Fe2+, an essential cofactor for chlorophyll biosynthesis, while enhancing chlorophyll degradation through increased chlorophyllase activity, ultimately accelerating pigment loss.Citation 28 Beyond pigment degradation, fluoride interferes with the photochemical phase of photosynthesis by inhibiting the Hill reaction and reducing electron transport, particularly at photosystem II (PSII). Replacement of Cl– ions by fluoride within the oxygen-evolving complex impairs water photolysis, leading to reduced electron flow and enhanced generation of reactive oxygen species (ROS). These alterations are accompanied by chloroplast disorganization, granal distortion, reduced stomatal conductance, and restricted CO2 assimilation, collectively diminishing photosynthetic efficiency.Citation 36 Furthermore, fluoride suppresses the activity of key carbon assimilation enzymes, including sucrose synthase and ribulose-1,5-bisphosphate carboxylase/oxygenase (RUBISCO), thereby limiting carbon fixation and carbohydrate production.Citation 37
Table 2. Effects of fluoride stress on 20 different plants and the observed toxicity effects.
Although the inhibitory effects of fluoride on photosynthesis have been consistently documented, the severity of damage varies considerably among plant species and depends on fluoride concentration, exposure duration, and nutrient status. Such variability suggests that differences in antioxidant capacity, chloroplast stability, and nutrient homeostasis may contribute to species-specific fluoride tolerance. However, the molecular mechanisms governing these adaptive responses remain insufficiently understood and warrant further investigation. Representative studies describing the photosynthetic responses of different plant species to fluoride stress are summarized in .
Effect of fluoride toxicity on respiration rate
Cellular respiration is a fundamental metabolic process that provides ATP required for plant growth, nutrient uptake, maintenance, and adaptation to environmental stresses. Fluoride toxicity disrupts respiratory metabolism by altering the activity of key enzymes involved in energy production, thereby reducing metabolic efficiency and impairing normal physiological functions. However, the extent of respiratory inhibition varies considerably depending on plant species, tissue type, developmental stage, duration of exposure, and the amount of fluoride accumulated within plant tissues.Citation 53 One of the primary effects of fluoride is the inhibition of several respiratory enzymes, including ascorbate oxidase, polyphenol oxidase, succinate dehydrogenase, malate dehydrogenase, and peroxidase, resulting in reduced respiratory activity and ATP production.Citation 11 Because glycolytic enzymes differ in their sensitivity to fluoride, plants may exhibit shifts in carbon metabolism, with increased reliance on the pentose phosphate pathway (PPP) during prolonged fluoride exposure.Citation 54
Evidence also suggests that fluoride-induced metabolic adjustments activate alternative respiratory pathways associated with oxidative stress tolerance. Increased activities of glucose-6-phosphate dehydrogenase, cytochrome oxidase, catalase, and peroxidase have been reported in fluoride-stressed tissues, indicating that enhanced PPP activity may help generate reducing equivalents (NADPH) required for antioxidant defense and cellular repair.Citation 26 These metabolic responses suggest that plants attempt to compensate for impaired respiration by reallocating carbon flux toward protective pathways.
Although respiratory inhibition is a common consequence of fluoride toxicity, the underlying regulatory mechanisms remain poorly understood. Future studies integrating physiological, biochemical, and molecular approaches are required to clarify how fluoride reprograms respiratory metabolism and contributes to stress adaptation in different plant species.
Fluoride stress on ATPase enzymes
ATPases are membrane-associated enzymes that play indispensable roles in cellular energy metabolism by maintaining ion gradients, membrane potential, intracellular pH, and ATP homeostasis. Consequently, impairment of ATPase activity under fluoride stress has profound implications for plant growth, nutrient transport, and stress adaptation. Fluoride has been shown to inhibit ATP synthase and plasma membrane H+-ATPases, thereby disrupting ATP generation and reducing the energy available for active transport and other ATP-dependent physiological processes.Citation 55 Furthermore, fluoride-induced damage to plasma and tonoplast membranes compromises membrane integrity, exacerbating ion imbalance and reducing transport efficiency.
Experimental evidence indicates that fluoride suppresses plasma membrane ATPase activity in species such as Beta vulgaris, suggesting that ATPases are important cellular targets of fluoride toxicity. The inhibitory effect appears to be influenced by Mg2+ availability because fluoride readily forms Mg–F complexes that interact with ATPase catalytic sites, thereby restricting ATP hydrolysis and enzyme activity. This interaction highlights the close relationship between fluoride toxicity, magnesium homeostasis, and membrane-associated energy metabolism.Citation 56 Similar reductions in plasma membrane ATPase activity have also been reported following prolonged fluoride exposure, supporting the hypothesis that membrane ATPases are highly susceptible to fluoride-induced oxidative and structural damage.Citation 57 , Citation 58
Although and metabolic dysfunction, the precise molecular mechanisms regulating ATPase sensitivity under fluoride stress remain largely unresolved. Current evidence suggests that ATPase inhibition is not solely a consequence of direct fluoride binding but may also result from oxidative membrane damage, altered ion homeostasis, and disruption of cellular signaling networks.Citation 59 , Citation 60 Future studies integrating structural biology, membrane physiology, and molecular genetics are therefore needed to clarify the regulation of ATPases under fluoride stress and to identify potential targets for improving fluoride tolerance in crop plants. Recent reviews likewise emphasize that molecular mechanisms governing fluoride uptake, transport, and cellular responses in plants remain insufficiently understood, underscoring the need for further mechanistic investigations.
Role of reactive oxygen species (ROS) in fluoride stress
Reactive oxygen species (ROS) play a central role in fluoride-induced phytotoxicity by acting as both signaling molecules and mediators of oxidative damage. Under normal physiological conditions, ROS are continuously generated at low levels in chloroplasts, mitochondria, and peroxisomes during aerobic metabolism and are efficiently scavenged by enzymatic and non-enzymatic antioxidant systems.Citation 11 Fluoride-induced inhibition of photosynthetic carbon fixation reduces the availability of oxidized NADP+, promoting electron leakage from photosystem I to molecular oxygen through the Mehler reaction and generating superoxide radicals (O2•–). In addition, peroxisomes contribute substantially to ROS production through xanthine oxidase-mediated oxidation of xanthine and hypoxanthine, as well as membrane-associated electron transport chains.Citation 35 , Citation 61 The overproduction of ROS subsequently triggers lipid peroxidation, protein oxidation, nucleic acid damage, membrane destabilization, and enzyme inactivation, ultimately impairing cellular metabolism and plant growth. Although ROS accumulation is generally regarded as a hallmark of fluoride toxicity, emerging evidence suggests that ROS also function as important signaling molecules regulating stress perception, antioxidant defense, and metabolic adaptation. The balance between ROS production and scavenging therefore determines whether ROS act as beneficial signaling intermediates or become cytotoxic.Citation 62 The major ROS generated under fluoride stress, their cellular sources, biological effects, and corresponding antioxidant defense mechanisms are summarized in .
Table 3. Major reactive oxygen species generated under fluoride stress, their cellular sources, biological effects, and plant defense responses.
Impact of fluoride toxicity on the anti-oxidative enzyme system
The excessive accumulation of reactive oxygen species (ROS) under fluoride stress activates a complex antioxidant defense system that protects plant cells against oxidative damage (). Because the mechanisms of ROS generation have already been discussed (Section 2.6), this section focuses on the antioxidant responses that maintain cellular redox homeostasis under fluoride exposure. The effectiveness of this defense system largely determines the degree of fluoride tolerance exhibited by different plant species. Among the enzymatic antioxidants, superoxide dismutase (SOD), catalase (CAT), peroxidase (POX), ascorbate peroxidase (APX), and glutathione reductase (GR) constitute the primary detoxification machinery against fluoride-induced oxidative stress.Citation 66 , Citation 67 SOD catalyzes the conversion of superoxide radicals (O2•–) into hydrogen peroxide (H2O2), which is subsequently detoxified by CAT and APX to prevent excessive H2O2 accumulation.Citation 68 , Citation 69 Meanwhile, POX contributes to H2O2 scavenging while participating in lignification and cell wall strengthening, whereas GR maintains intracellular glutathione in its reduced form, thereby sustaining the ascorbate–glutathione cycle and overall cellular redox balance.Citation 70 , Citation 71 The activities of antioxidant enzymes under fluoride stress, however, are highly species-dependent and are influenced by fluoride concentration, exposure duration, tissue type, and nutrient availability.Citation 72 Moderate fluoride exposure frequently induces antioxidant enzymes as an adaptive response, whereas prolonged or excessive fluoride accumulation may overwhelm the antioxidant system, resulting in enzyme inhibition and enhanced oxidative damage.Citation 73 These contrasting responses indicate that antioxidant activation represents a dynamic defense mechanism rather than a universal indicator of fluoride tolerance.Citation 73 Although numerous studies have reported alterations in antioxidant enzyme activities under fluoride stress, relatively little is known about the molecular regulation of these enzymes or their interaction with signaling pathways governing stress adaptation. Future investigations integrating transcriptomics, proteomics, metabolomics, and functional genomics are therefore needed to clarify the regulatory networks controlling antioxidant responses and to identify biomarkers associated with fluoride tolerance.
Figure 2. Fluoride’s adverse effects on plants: toxicity and ramifications.
Impact of fluoride toxicity on anabolic processes
Anabolic metabolism encompasses the biosynthesis of carbohydrates, proteins, nucleic acids, lipids, and other cellular constituents that are essential for plant growth and development. Fluoride toxicity disrupts these biosynthetic pathways primarily by reducing ATP production, impairing carbon assimilation, disturbing nutrient homeostasis, and inducing oxidative stress, thereby limiting the availability of energy and metabolic precursors required for cellular biosynthesis.Citation 74 The suppression of anabolic metabolism under fluoride stress extends beyond photosynthesis. Reduced carbohydrate production limits substrate availability for the synthesis of structural polysaccharides, while inhibition of protein synthesis restricts the formation of metabolic enzymes and structural proteins required for growth. Fluoride-induced deficiencies of essential nutrients such as Fe, Mg, Ca, and P further impair chlorophyll biosynthesis, amino acid metabolism, nucleic acid synthesis, and membrane formation, thereby reducing overall metabolic activity.Citation 75 In addition, oxidative damage to cellular macromolecules diverts metabolic resources from biosynthetic processes toward cellular repair and antioxidant defense, further suppressing plant growth and biomass accumulation.
Although photosynthesis provides the primary carbon source for anabolic metabolism, increasing evidence suggests that fluoride-induced inhibition of biosynthetic pathways cannot be explained solely by reduced photosynthetic efficiency. Instead, fluoride exerts coordinated effects on cellular energy metabolism, nutrient utilization, enzyme activities, and redox homeostasis, collectively restricting plant anabolic capacity. Nevertheless, the molecular regulation of anabolic pathways under fluoride stress remains poorly understood, and future studies employing transcriptomic, proteomic, and metabolomic approaches are needed to identify the key regulatory networks involved in metabolic adaptation and fluoride tolerance.Citation 76 Although fluoride-induced suppression of anabolic metabolism has been consistently reported, the relative contribution of impaired energy metabolism, nutrient imbalance, and oxidative stress to the inhibition of individual biosynthetic pathways remains unclear. Future studies integrating physiological, transcriptomic, proteomic, and metabolomic approaches are therefore required to elucidate the regulatory mechanisms underlying fluoride tolerance.
Impact of fluoride toxicity on the catabolic processes
Fluoride toxicity disrupts catabolic metabolism by altering energy-generating pathways that are essential for maintaining plant growth, nutrient acquisition, and cellular homeostasis. Cellular respiration, a central catabolic process responsible for ATP production, is particularly sensitive to fluoride stress, although its response depends on fluoride concentration, duration of exposure, plant species, and developmental stage. Mild fluoride exposure may transiently stimulate oxygen consumption, whereas prolonged or excessive exposure generally suppresses respiratory activity and reduces metabolic efficiency.Citation 53 Fluoride-induced alterations in respiration are associated with changes in the activity of several key metabolic enzymes. Inhibition of enzymes involved in glycolysis and the tricarboxylic acid (TCA) cycle restricts ATP generation, leading to impaired energy availability for biosynthetic and transport processes. Furthermore, respiratory responses vary among tissues due to differences in fluoride accumulation and metabolic activity, explaining the inconsistent findings reported across plant species.Citation 77 Under fluoride stress, plants may partially compensate for reduced respiratory efficiency by redirecting carbon metabolism toward the pentose phosphate pathway (PPP). Increased activities of enzymes such as glucose-6-phosphate dehydrogenase, cytochrome oxidase, catalase, and peroxidase have been reported in fluoride-exposed tissues, indicating activation of alternative metabolic pathways that generate reducing power (NADPH) required for antioxidant defense and cellular repair. However, the molecular mechanisms regulating this metabolic reprogramming remain poorly understood and require further investigation.Citation 78
Interactions between fluoride toxicity and mineral nutrition
The interaction between fluoride toxicity and mineral nutrition is complex and bidirectional, as fluoride not only disrupts the uptake and utilization of essential mineral nutrients but is itself influenced by nutrient availability. Among the essential elements, calcium (Ca), magnesium (Mg), iron (Fe), zinc (Zn), manganese (Mn), phosphorus (P), and nitrogen (N) play important roles in determining the severity of fluoride toxicity through their involvement in ion transport, enzyme activation, and metabolic regulation.Citation 79 The interaction between fluoride toxicity and mineral nutrition is complex and bidirectional, as fluoride not only disrupts the uptake and utilization of essential mineral nutrients but is itself influenced by nutrient availability. Among the essential elements, calcium (Ca), magnesium (Mg), iron (Fe), zinc (Zn), manganese (Mn), phosphorus (P), and nitrogen (N) play important roles in determining the severity of fluoride toxicity through their involvement in ion transport, enzyme activation, and metabolic regulation.Citation 11 Adequate calcium nutrition stabilizes plasma membranes, preserves membrane permeability, and reduces fluoride-induced cellular damage, whereas calcium-deficient plants exhibit greater susceptibility to fluoride toxicity due to enhanced fluoride uptake and impaired membrane integrity.Citation 80–82 Furthermore, calcium deficiency may disrupt pollen germination, pollen tube elongation, and reproductive development, thereby exacerbating fluoride-induced yield losses.
Magnesium also plays an important role because it serves as a cofactor for numerous enzymes involved in photosynthesis and energy metabolism. Fluoride can interfere with magnesium-dependent enzymatic reactions through the formation of Mg–F complexes, thereby inhibiting ATP-dependent metabolic processes and reducing enzyme activity.Citation 53 Although several studies have suggested that fluoride primarily affects the physiological activity of magnesium rather than its tissue concentration, disruption of magnesium function may contribute to impaired photosynthesis, respiration, and overall metabolic performance.Citation 83 Fluoride toxicity also alters the uptake and distribution of micronutrients such as iron (Fe), manganese (Mn), and zinc (Zn). Deficiencies of these nutrients frequently accompany fluoride exposure, resulting in chlorosis, reduced chlorophyll synthesis, impaired enzyme activity, and diminished photosynthetic efficiency.Citation 84 In addition, interactions with macronutrients, including nitrogen (N), phosphorus (P), and potassium (K), indirectly influence fluoride absorption and translocation within plants. Moderate supplies of nitrogen, phosphorus, and calcium may enhance fluoride uptake because of increased root activity, whereas nutrient deficiencies or excessive calcium can reduce fluoride accumulation by restricting its mobility or promoting precipitation within the rhizosphere.Citation 85 Collectively, these findings demonstrate that fluoride toxicity cannot be considered independently of plant mineral nutrition. Rather than acting through a single mechanism, fluoride disrupts nutrient homeostasis by modifying ion transport, membrane function, enzyme activities, and metabolic processes, ultimately reducing plant growth and productivity.Citation 86 However, the interactions between fluoride and mineral nutrients remain highly species-specific and are influenced by soil properties, nutrient availability, and environmental conditions. Future studies integrating ionomics, molecular transport biology, and nutrient signaling are therefore required to elucidate the regulatory mechanisms governing fluoride–mineral interactions and to develop nutrient-based strategies for improving fluoride tolerance in crop plants. summarizes the interactions between fluoride toxicity and essential mineral nutrients, highlighting their collective effects on nutrient homeostasis, enzyme activity, photosynthesis, and plant growth.
Figure 3. Highlighting the interaction of essential minerals and fluoride.
Impact of fluoride toxicity on fruit yield
Fluoride’s interference with calcium, a crucial element for fertilization, can result in various harmful effects on fruit yield and quality. Plants deficient in calcium are especially vulnerable to fluoride damage, which often manifests as leaf injury.Citation 87 This interference can also disrupt pollen germination and growth, ultimately impacting the fertilization process. Certain fruits, such as peaches, exhibit high sensitivity to fluoride exposure. Fluoride-induced conditions like “black tip” or “suture red spot” can develop in affected fruits, characterized by premature ripening, reddening of the suture area externally and internally, and splitting of the flesh along the suture line.Citation 88
Prolonged exposure to heightened levels of fluoride can lead to a significant increase in the occurrence of abnormal fruit in peaches and deformation in strawberries. Moreover, fluoride has been observed to disrupt fruiting in tomatoes and beans by interfering with fertilization and seed development processes. The stage of flowering is notably sensitive to fluoride toxicity, with the carpels, responsible for seed production, being more vulnerable compared to the pollen-producing anthers.Citation 89 Exposure to fluoride during flowering alters the surface of the stigma, crucial for pollen reception, thereby affecting the growth of pollen tubes and subsequent fertilization events. The accumulation of fluoride on both the surface and within the stigma and style disrupts the normal calcium gradient, essential for guiding pollen tube growth and facilitating fertilization. This interference ultimately hampers the successful development of seeds and subsequent fruit formation.Citation 90 The detrimental effects of fluoride on fertilization, seed formation, and fruit yield underscore the intricate interplay between fluoride exposure and vital reproductive processes in plants. These findings highlight the importance of understanding and mitigating fluoride-induced disruptions to ensure optimal plant reproductive health and yield.Citation 90
Potential mitigation strategies and research gaps
Addressing fluoride toxicity in crops requires a multifaceted approach, integrating soil amendments, breeding for tolerance, and biotechnological interventions.Citation 91 Liming and phosphate fertilizers can reduce fluoride bioavailability by forming stable complexes, while organic amendments like biochar enhance soil buffering capacity.Citation 92 Additionally, silicon and calcium supplementation have shown promise in mitigating fluoride uptake by competing with fluoride ions at absorption sites.Citation 93 From a genetic perspective, identifying and engineering fluoride-tolerant crop varieties through molecular breeding or gene-editing technologies could offer long-term solutions.Citation 94 However, discrepancies in fluoride toxicity thresholds across species, variations in soil interactions, and the limited understanding of plant fluoride detoxification pathways highlight significant research gaps. Future studies should focus on elucidating the molecular mechanisms of fluoride tolerance and developing sustainable management strategies for affected agricultural regions.
Conclusion
Fluoride toxicity poses a significant global challenge, impacting both plant and animal health. While extensive research has focused on its effects on animals and human health, the implications for crop plants require greater attention. Understanding the physiological, biochemical, and molecular mechanisms underlying fluoride tolerance is crucial for developing effective mitigation strategies. The adverse effects of fluoride contamination in air and soil on crop growth and yield have serious implications for global agriculture.
To address this challenge, future research should focus on identifying key molecular pathways involved in fluoride detoxification and tolerance, leveraging biotechnological advancements such as CRISPR-based gene editing and transgenic approaches to develop fluoride-resistant crop varieties. Additionally, the application of beneficial microbes, biochar, and soil amendments like calcium and silicon could offer practical strategies for minimizing fluoride uptake in plants. Integrating these approaches will contribute to the development of sustainable agricultural practices in fluoride-affected regions, ensuring long-term food security and environmental stability. Recent advances in transcriptomics, proteomics, metabolomics, and functional genomics have opened new opportunities to unravel the molecular mechanisms underlying fluoride uptake, transport, detoxification, and stress adaptation. Future research integrating these multi-omics approaches with CRISPR/Cas-mediated genome editing and molecular breeding will facilitate the identification of key genes, transporters, signaling pathways, and regulatory networks associated with fluoride tolerance, thereby accelerating the development of fluoride-resilient crop cultivars.
Ethics approval
The ethical statement is not applicable in this study as this is a review paper, and we are using secondary published information.
Acknowledgments
This work was supported by the Deanship of Scientific Research, Vice Presidency for Graduate Studies and Scientific Research, King Faisal University, Saudi Arabia (KFU263861).
Disclosure statement
No potential conflict of interest was reported by the author(s).
Data availability statement
Data included in study/supplementary material/referenced in study.
Additional information
Funding
References
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