Understanding boron toxicity in aquatic plants (Salvinia natans and Lemna minor) in the presence and absence of EDTA.

Even though boron is a widely used element in various industries and a contributor to water pollution worldwide, few studies have examined the toxicity of boron in aquatic plants. EDTA is used to maintain aquatic plants cultures, however it is possible to modify the toxicity of metals. The objective of this study is to assess the toxicity of boron in aquatic plants and explore the impact of EDTA presence on the resulting toxic responses. Floating watermoss Salvinia natans and duckweed Lemna minor were exposed to concentrations ranging from 5 to 100 mg/L for 7 days and 1 to 60 mg/L for 3 days, respectively. Growth and photosynthetic activity parameters were investigated in the presence and absence of EDTA. Growth inhibitions in both aquatic plants were observed in a concentration-dependent manner, irrespective of the presence or absence of EDTA. For instance, based on the specific growth rate (leaves coverage), EC10 values for S. natans were calculated as 12.7 (9.9-15.3) mg/L and 8.0 (5.8-10.3) mg/L with and without EDTA, respectively. In the case of L. minor, EC10 values were calculated as 1.3 (0.8-1.89) mg/L and 2.0 (0.4-4.3) mg/L with EDTA without EDTA, respectively. Significant effects were also observed on the photosynthetic capacity, however there was no change in the increase of boron concentration. Generally, negligible effects of EDTA to the toxicity of boron were observed in the present study. By comparing toxicity results based on the presence and absence of EDTA, which is an essential element in the test medium, the results of this study are expected to be utilized for the ecological risk assessment of boron in aquatic ecosystems.

Boron-mediated amelioration of copper toxicity in Citrus sinensis seedlings involved reduced concentrations of copper in leaves and roots and their cell walls rather than increased copper fractions in their cell walls.

Little information is available on how boron (B) supplementation affects plant cell wall (CW) remodeling under copper (Cu) excess. 'Xuegan' (Citrus sinensis) seedlings were submitted to 0.5 or 350 µM Cu × 2.5 or 25 µM B for 24 weeks. Thereafter, we determined the concentrations of CW materials (CWMs) and CW components (CWCs), the degree of pectin methylation (DPM), and the pectin methylesterase (PME) activities and PME gene expression levels in leaves and roots, as well as the Cu concentrations in leaves and roots and their CWMs (CWCs). Additionally, we analyzed the Fourier transform infrared (FTIR) and X-ray diffraction (XRD) spectra of leaf and root CWMs. Our findings suggested that adding B reduced the impairment of Cu excess to CWs by reducing the Cu concentrations in leaves and roots and their CWMs and maintaining the stability of CWs, thereby improving leaf and root growth. Cu excess increased the Cu fractions in leaf and root pectin by decreasing DPM due to increased PME activities, thereby contributing to citrus Cu tolerance. FTIR and XRD indicated that the functional groups of the CW pectin, hemicellulose, cellulose, and lignin could bind and immobilize Cu, thereby reducing Cu cytotoxicity in leaves and roots.

Boron deficiency decreased the root activity of Ga-exposed rice seedlings by reducing iron accumulation and increasing Ga in iron plaque.

Gallium (Ga) is an emerging chemical pollutant chiefly associated with high-tech industries. Boron (B) alleviates the negative effects of toxic elements on plant growth. Thereby, the effects of B fertilization on Ga toxicity in rice seedlings was studied to clarify the role of iron plaque in the distribution of Ga, Fe, and B in Ga-treated rice seedlings in the presence or absence of B. Gallium exposure significantly reduced the biomass of rice seedlings. Boron deficiency induced a significant change in the distribution of B in Ga-treated rice seedlings compared with "Ga+B" treatments. Accumulation of Ga in roots, dithionite-citrate-bicarbonate (DCB) extracts, and shoots showed a dose-dependent manner from both +B and -B rice seedlings. Boron nutrition levels affect the distribution of Fe in roots, DCB extracts, and shoots, in which DCB-extractable Fe was significantly decreased from "Ga-B" treatments compared with "Ga+B" treatments. Root activity was significantly decreased in both Ga-exposed rice seedlings; however, B-deficient seedlings showed a severe reduction than +B rice seedlings. These results reveal that Fe plaque might be a temporary sink for B accumulation when plants are grown with proper B, wherein the re-utilization of DCB-extractable B stored in Fe plaque is mandatory for plant growth under B deficiency. Correlation analysis revealed that B deficiency decreased the root activity of Ga-exposed rice seedlings by reducing DCB-extractable Fe and increasing DCB-extractable Ga in Fe plaque. This study enhances our understanding of how B nutritional levels affect Ga toxicity in rice plants.

Genome-wide transcriptome profiling of mulberry (Morus alba) response to boron deficiency and toxicity reveal candidate genes associated with boron tolerance in leaves.

Mulberry (Morus alba) is an essential plant with countless economic benefits; however, its growth and metabolic processes are hampered by boron (B) stresses. Very little research has been performed to elucidate boron tolerance and detoxification mechanisms in this species. The M. alba cultivar, Yu-711, was exposed to five different concentrations of boric acid (H3BO3), including deficient (T1; 0 mM) moderate B deficiency (T2; 0.02 mM), sufficient (CK; 0.1 mM) and toxic (T3 and T4; 0.5 and 1 mM) levels for 18 days of growth in pots experiment. Transcriptome analysis of B deficiency and toxicity treatments was performed on mulberry leaves. The transcriptome data reveal that a total of 6114 genes were differentially expressed (DEGs), of which 3830 were up-regulated and 2284 were down-regulated. A comparative analysis between treatment groups CK-vs-T1 (deficiency) and CK-vs-T4 (toxicity) indicates that 590 and 1383 genes were down-regulated in both deficiency and B toxicity, respectively. The results show that 206 genes were differentially expressed in all treatments. B deficiency and toxicity significantly altered the expression of the key aquaporins (PIP2-1, PIP2-7, PIP2-4 and NIP3-1) and high-affinity boron transporter genes (BOR1 and BOR7). In addition, boron stress also altered the expression of antioxidants and photosynthesis-related genes. B stresses were found to alter several transcription factors including ERF1B, which is associated with the regulation of boron uptake and the synthesis and signaling of phytohormones. Unravelling the mechanisms of B tolerance and detoxification is important and would give us further insight into how B stresses affect mulberry plants.

Identification and function characterization of BnaBOR4 genes reveal their potential for Brassica napus cultivation under high boron stress.

Boron (B) is essential for plant growth, but toxic in excess. In several countries, soil toxic B levels are always a severe agricultural problem in arid and semi-arid regions. Phytoremediation of excess B containing soil is still in its infancy, while high B tolerant plants with elevated protein abundance of B efflux transporter were successfully established or explored. Brassica napus (B. napus) is one of the most important oil crops. However, B efflux transporters underlying excess B tolerance in B. napus remain unknown. Here, we reported that in Brassicaceae species, B. napus had four homologous genes of Arabidopsis AtBOR4 , which were renamed BnaBOR4.1, BnaBOR4.2, BnaBOR4.3 and BnaBOR4.4. BnaBOR4.1, BnaBOR4.2 and BnaBOR4.3 showed constitutive expression and BnaBOR4.4 appeared to be a pseudogene. BnaBOR4.2 and BnaBOR4.3 were expressed in inner cell layers and BnaBOR4.1 in the outer cell layer in root tip, and all were expressed in vascular tissue in the mature zone. B efflux activity assays in yeast demonstrated that BnaBOR4.1, BnaBOR4.2 and AtBOR4 but not BnaBOR4.3 had comparable levels of B transport activity. Structure-functional analysis between BnaBOR4.3 and BnaBOR4.2 demonstrated that amino acid residue substitution at position 297 (Ala vs Pro) and 427 (Met vs Leu) is critical for the B transport activity. Mutant BnaBOR4.3M427L partially restored the B efflux activity, and both mutants BnaBOR4.3A297P and BnaBOR4.3A297P&M427L fully restored B efflux activity, indicating that the Pro297 residue is critical for their function. Further validation of BnaBOR4 was accomplished by growing transgenic Arabidopsis plants under high B conditions. Taken together, our study identified two functional B efflux genes BnaBOR4.1 and BnaBOR4.2 in B. napus, and a key amino acid residue proline 297 associated with B efflux activity. This study highlights the potential of BanBOR4 genes for B. napus cultivation under high B stress.

Physiological and metabolic changes in response to Boron levels are mediated by ethylene affecting tomato fruit yield.

Boron (B) is an essential nutrient for the plant, and its stress (both deficiency and toxicity) are major problems that affect crop production. Ethylene metabolism (both signaling and production) is important to plants' differently responding to nutrient availability. To better understand the connections between B and ethylene, here we investigate the function of ethylene in the responses of tomato (Solanum lycopersicum) plants to B stress (deficiency, 0 µM and toxicity, 640 µM), using ethylene related mutants, namely nonripening (nor), ripening-inhibitor (rin), never ripe (Nr), and epinastic (Epi). Our results show that B stress does not necessarily inhibit plant growth, but both B stress and ethylene signaling severely affected physiological parameters, such as photosynthesis, stomatal conductance, and chlorophyll a fluorescence. Under B toxicity, visible symptoms of toxicity appeared in the roots and margins of the older leaves through necrosis, caused by the accumulation of B which stimulated ethylene biosynthesis in the shoots. Both nor and rin (ethylene signaling) mutants presented similar responses, being these genotypes more sensitive and displaying several morphophysiological alterations, including fruit productivity reductions, in response to the B toxicity conditions. Therefore, our results suggest that physiological and metabolic changes in response to B fluctuations are likely mediated by ethylene signaling.

The effects of 100 nm pure boron nanoparticles on the primary antioxidant system of Nile tilapia (Oreochromisniloticus).

In this study, the effects of 100 nm boron nanoparticles (B-NPs) on the primary antioxidant status of Nile tilapia were researched via analysis of enzyme activities and related gene expressions. This is a new study which focuses on the relationship between B-NPs and oxidative stress that contribute to the literature in terms of its scope. Fish (n = 15) for each group were exposed to three different concentrations as 5, 25 (n2) and 125 (n3) mg/L during 96 h to see the response of the primary antioxidant system. According to the results, SOD expressions differed in all treatment groups compared to the control group (P < 0.05). CAT expressions were different in 5 and 125 mg/L groups compared to control and 25 mg/L groups (P < 0.05). GPX expressions were only different in 125 mg/L group (P < 0.05). The changes in enzyme activities of SOD and CAT were significantly different in 25 mg/L groups. GPX enzyme activities were not significant (P > 0.05). TBARS concentrations in 25 mg/L group were significantly different from those in the control and 125 mg/L groups (P < 0.05).

Role of ABA in the adaptive response of Arabidopsis plants to long-term boron toxicity treatment.

Boron (B) toxicity causes impairments in several plant metabolic and physiological processes. Under conditions of excessive B availability, this micronutrient is passively transported through the transpiration stream and accumulates in leaves, causing the development of necrotic regions in leaf tips. Some plants have developed adaptive mechanisms to minimize the toxic effects of excessive B accumulation in their tissues. Thus, for instance, in Arabidopsis it has been described an ABA-dependent decrease in the transpiration rate that would restrict B accumulation in aerial plant tissues in response to short-term B toxicity, this effect being mediated by AtNCED3 (which encodes a key enzyme for ABA biosynthesis). The present work aimed to study the possible involvement of ABA in the adjustment of plant water balance and B homeostasis during the adaptive response of Arabidopsis to prolonged B toxicity. For this purpose, Arabidopsis wild-type and the ABA-deficient nced3-2 mutant plants were subjected to B toxicity for 7 days. We show that ABA-dependent stomatal closure is determinant for the adjustment of plant water relations under conditions of prolonged B toxicity. Results suggest that, in addition to the AtNCED3 gene, the AtNCED5 gene could also be involved in this ABA-dependent stomatal closure. Finally, our results also indicate the possible role of endogenous root ABA content in the mechanism of active efflux of B via BOR4 (efflux-type B transporter) from the root to the external environment under excess B conditions.

Regulation on copper-tolerance in Citrus sinensis seedlings by boron addition: Insights from root exudates, related metabolism, and gene expression.

Boron (B) can alleviate Citrus copper (Cu)-toxicity. However, the underlying mechanism by which B mitigates Cu-toxicity is unclear. 'Xuegan' (Citrus sinensis) seedlings were exposed to 0.5 (control) or 350 (Cu-toxicity) µM Cu and 2.5 or 25 µM B for 24 weeks. Thereafter, we investigated the secretion of low molecular weight compounds [LMWCs; citrate, malate, total soluble sugars (TSS), total phenolics (TP), and total free amino acids (TFAA)] by excised roots and their concentrations in roots and leaves, as well as related enzyme gene expression and activities in roots and leaves. Cu-stress stimulated root release of malate and TFAA, which might contribute to citrus Cu-tolerance. However, B-mediated-mitigation of Cu-stress could not be explained in this way, since B addition failed to further stimulate malate and TFAA secretion. Indeed, B addition decreased Cu-stimulated-secretion of malate. Further analysis suggested that Cu-induced-exudation of malate and TFAA was not regulated by their levels in roots. By contrast, B addition increased malate, citrate, and TFAA concentrations in Cu-toxic roots. Cu-toxicity increased TP concentration in 25 µM B-treated leaves, but not in 2.5 µM B-treated leaves. Our findings suggested that the internal detoxification of Cu by LMWCs played a role in B-mediated-alleviation of Cu-toxicity.

A critical review of plant adaptation to environmental boron stress: Uptake, utilization, and interplay with other abiotic and biotic factors.

Boron (B) is an indispensable mineral nutrient for plants and is primarily taken up by roots mainly in the form of boric acid (H3BO3). Recently, research shows that B has a significant impact on plant growth and productivity due to its narrow range between deficiency and toxicity. Fertilization and other procedures to address B stress (deficiency and toxicity) in soils are generally expensive and time-consuming. Over the past 20 years, substantial studies have been conducted to investigate the mechanisms underlying B acquisition and the molecular regulation of B stress in plants. In this review, we discuss the effects of B stress on plant growth, physiology, and biochemistry, and finding on enhancing plant tolerance from the perspective of plant B uptake, transport, and utilization. We also refer to recent results demonstrating the interactions among B and other biological and abiotic factors, including nitrogen, phosphorus, aluminum, and microorganisms. Finally, emerging trends in this field are discussed.

Boron contamination and its risk management in terrestrial and aquatic environmental settings.

Boron (B) is released to terrestrial and aquatic environments through both natural and anthropogenic sources. This review describes the current knowledge on B contamination in soil and aquatic environments in relation to its geogenic and anthropogenic sources, biogeochemistry, environmental and human health impacts, remediation approaches, and regulatory practices. The common naturally occurring sources of B include borosilicate minerals, volcanic eruptions, geothermal and groundwater streams, and marine water. Boron is extensively used to manufacture fiberglass, thermal-resistant borosilicate glass and porcelain, cleaning detergents, vitreous enamels, weedicides, fertilizers, and B-based steel for nuclear shields. Anthropogenic sources of B released into the environment include wastewater for irrigation, B fertilizer application, and waste from mining and processing industries. Boron is an essential element for plant nutrition and is taken up mainly as boric acid molecules. Although B deficiency in agricultural soils has been observed, B toxicity can inhibit plant growth in soils under arid and semiarid regions. High B intake by humans can be detrimental to the stomach, liver, kidneys and brain, and eventually results in death. Amelioration of soils and water sources enriched with B can be achieved by immobilization, leaching, adsorption, phytoremediation, reverse osmosis, and nanofiltration. The development of cost-effective technologies for B removal from B-rich irrigation water including electrodialysis and electrocoagulation techniques is likely to help control the predominant anthropogenic input of B to the soil. Future research initiatives for the sustainable remediation of B contamination using advanced technologies in soil and water environments are also recommended.

What can we learn from epidemiological studies on chronic boron exposure?

Boric acid and sodium borates are classified as toxic to reproduction and development under category 1B, with the hazard statement of H360FD (May damage fertility. May damage the unborn child) in the EU-CLP regulation. This classification triggered environmental and occupational epidemiology studies on boron in China, Türkiye, and Argentina. The highest mean total daily boron exposure (DBE) levels in mining areas or processing plants in China and Türkiye were 41.2 and 47.17 mg/day, respectively. These DBE levels are higher than the proposed reference doses (RfDs) for the developmental (9.6 mg B/day) and reproductive effects (20.3 mg B/day) of boron in females and males. Despite these high exposure levels in occupational settings, boron-mediated reproductive and developmental effects have not been reported. Blood (or serum) boron concentrations were also reported in these studies. The highest mean blood-boron concentration reported for the boron-exposed male workers in Türkiye is 570.6 ng B/g (ppb). This blood-boron concentration is still much lower than the blood boron concentrations corresponding to the no observed adverse effect level (NOAEL) for both developmental and reproductive effects in rats. Given the blood boron concentrations reported in current epidemiological studies, the lack of reproductive and developmental effects in humans is not surprising. Recent epidemiological studies have proven that it will not be possible to reach the critical blood boron concentrations in humans for the reproductive and developmental effects, especially in daily life.

Potential application of melatonin in reducing boron toxicity in rice seedlings through improved growth, cell wall composition, proline, and defense mechanisms.

Melatonin (MT) has been demonstrated to provide defense against both biotic and abiotic stressors. Boron toxicity (BT) can significantly limit the growth and production of plants. However, few studies have been conducted on whether MT is effective in attenuating B toxicity in different plants. In order to evaluate the efficacy of exogenous MT treatment in reducing the negative impact of BT on rice seedlings, this study examined the influence of MT on growth, antioxidant capacity, cell wall composition, and proline metabolism in rice seedlings under hydroponics. Four treatments were established: MT (50 µM), MT + BT (50 µM MT + 800 µM B), BT (800 µM), and CK (control) in a completely randomized design. The results indicate that BT had a significant detrimental effect on the shoot length, root length, and root and shoot fresh weights of rice seedlings by 11.96%, 27.77%, 25.69%, and 18.67%, respectively as compared to the control treatment. However, exogenous MT application increased these parameters and reduced B accumulation in aboveground parts (14.05%) of the plant. Exogenous MT also increased the endogenous melatonin content and antioxidant enzyme activities (64.45%, 71.61%, 237.64%, and 55.42% increase in superoxide dismutase, ascorbate peroxidase, and peroxidase activities, respectively), while decreasing reactive oxygen species levels and oxidized forms of glutathione and ascorbic acid. Additionally, MT enhanced the biosynthesis of proline by decreasing proline dehydrogenase (ProDH) and increasing the GSH (glutathione) and ASA (ascorbic acid) contents. Exogenous MT also increased cell wall components that can increase B adsorption to the cell wall. Overall, these findings suggest that MT application can be a potential solution for strengthening the stress tolerance of rice seedlings, particularly under conditions of B toxicity. In regions where soil contains high levels of boron, the use of MT could enhance rice crop yields and quality.

Combined effects of microplastics and excess boron on Microcystis aeruginosa.

Both microplastics (MPs) and excess boron (B) have adverse effects on microalgae. However, the combined toxic effects of MPs and excess B on microalgae have not been studied. The aim of this study was to investigate the combined effects of excess B and three types of surface-modified microplastics, including plain polystyrene (PS-Plain), amino-modified polystyrene (PS-NH2), and carboxyl-modified polystyrene (PS-COOH), on chlorophyll a content, oxidative damage, photosynthetic activity and microcystin (MC) production in Microcystis aeruginosa. The results showed that PS-NH2 inhibited the growth of M. aeruginosa, with the maximum inhibition rate of 18.84 %, while PS-COOH and PS-Plain showed stimulatory effects with the maximum inhibition rates of -2.56 % and - 8.03 % respectively. PS-NH2 aggravated the inhibition effects of B, while PS-COOH and PS-Plain alleviated the inhibition effects. Furthermore, the combined exposure of PS-NH2 and excess B had a much greater effect on oxidative damage, cell structure, and production of MCs in algal cells than the combined effects of PS-COOH and PS-Plain. The charges on microplastics affected both B adsorption on microplastics and the aggregation of microplastics with algal cells, indicating that the charge on microplastics is a dominant factor influencing the combined effects of microplastics and excess B on microalgae. Our findings can provide direct evidence for the combined effects of microplastics and B on freshwater algae and improve the understanding of the potential risks of microplastics in aquatic ecosystems.

Boron bio-mining by high boron-tolerant native microalgae in Turkey and boron toxicity in the aquatic environment.

Boron (B) is one of the most important trace elements. Turkey has the largest B reserves in the world with 3 billion tons of B. Therefore, the toxicity of B is more important in Turkey. In this study, effective B removal was provided by different types of microalgae and then B recovery in culture media by bio-mining was detected. The water of Lake Mogan in Gölbasi/Ankara/Turkey was used as the microalgae culture medium as a cost-reducing factor. The effects of light, temperature, NaNO3 and K2HPO4 stresses on B removal were determined. The highest B removal was 17.19% at 25 µmol/m2s light intensity and 25 °C for Phormidium animale in Lake Mogan culture medium. Boron removal of Scenedesmus sp. was 36.42% at 25 µmol/m2s light intensity, at 25 °C, at 1.5 g/L NaNO3 and 40 mg/L K2HPO4 concentrations in BG11 medium on the 15th day. The chl (a + b) concentration of Scenedesmus sp. was 1.63 µg/mL at 75 µmol/m2s light intensity and the chl (a) concentration of P. animale was 2.01 µg/mL at 25 µmol/m2s light intensity. Considering all parameters, Scenedesmus sp. and P. animale are recommended as effective biomaterials for the B removal process.

Physiological and iTRAQ-based quantitative proteomics analyses reveal the similarities and differences in stress responses between short-term boron deficiency and toxicity in wheat roots.

BACKGROUND: Boron (B) is a trace element that is essential for normal wheat development, such as root growth. In wheat, roots are important organs that absorb nutrients and water. However, at present, there is insufficient research on the molecular mechanism underlying how short-term B stress affects wheat root growth. METHODS AND RESULTS: Here, the optimal concentration of B for wheat root growth was determined, and the proteomic profiles of roots under short-term B deficiency and toxicity were analyzed and compared by the isobaric tag for relative and absolute quantitation (iTRAQ) technique. A total of 270 differentially abundant proteins (DAPs) that accumulated in response to B deficiency and 263 DAPs that accumulated in response to B toxicity were identified. Global expression analysis revealed that ethylene, auxin, abscisic acid (ABA), and Ca2+ signals were involved in the responses to these two stresses. Under B deficiency, DAPs related to auxin synthesis or signaling and DAPs involved in calcium signaling increased in abundance. In striking contrast, auxin and calcium signals were repressed under B toxicity. Twenty-one DAPs were detected under both conditions, including RAN1 that played a core role in the auxin and calcium signals. Overexpression of RAN1 was shown to confer plant resistance to B toxicity by activating auxin response genes, including TIR and those identified by iTRAQ in this research. Moreover, growth of the primary roots of tir mutant was significantly inhibited under B toxicity. CONCLUSION: Taken together, these results indicate that some connections were present between RAN1 and the auxin signaling pathway under B toxicity. Therefore, this research provides data for improving the understanding of the molecular mechanism underlying the response to B stress.

Interaction between Boron and Other Elements in Plants.

Boron (B) is an essential mineral nutrient for growth of plants, and B deficiency is now a worldwide problem that limits production of B deficiency-sensitive crops, such as rape and cotton. Agronomic practice has told that balanced B and other mineral nutrient fertilizer applications is helpful to promote crop yield. In recent years, much research has reported that applying B can also reduce the accumulation of toxic elements such as cadmium and aluminum in plants and alleviate their toxicity symptoms. Therefore, the relation between B and other elements has become an interesting issue for plant nutritionists. Here we summarize the research progress of the interaction between B and macronutrients such as nitrogen, phosphorus, calcium, potassium, magnesium, and sulfur, essential micronutrients such as iron, manganese, zinc, copper, and molybdenum, and beneficial elements such as sodium, selenium, and silicon. Moreover, the interaction between B and toxic elements such as cadmium and aluminum, which pose a serious threat to agriculture, is also discussed in this paper. Finally, the possible physiological mechanisms of the interaction between B and other elements in plants is reviewed. We propose that the cell wall is an important intermediary between interaction of B and other elements, and competitive inhibition of elements and related signal transduction pathways also play a role. Currently, research on the physiological role of B in plants mainly focuses on its involvement in the structure and function of cell walls, and our understanding of the details for interactions between B and other elements also tend to relate to the cell wall. However, we know little about the metabolic process of B inside cells, including its interactions with other elements. More research is needed to address the aforementioned research questions in future.

Phytohormones in plant responses to boron deficiency and toxicity.

Boron (B) is an essential element for plant growth. Many agricultural soils around the globe have either insufficient or excessive amounts of available B, with major implications for crop production. Understanding major limitations imposed by B nutritional disorders may allow breeding crops for improved B use efficiency as well as make them more resilient to excessive B, thus reducing yield penalties. It has become apparent that B-related physiological disorders are mediated in large part by their impact on plant hormone production and signaling. The aim of this review is to summarize current knowledge of the roles of hormones in plant responses to B and their impact on plant growth and development. The most significant effect of B deficiency is the inhibition of root elongation. Boron deficiency promotes the redistribution of auxin in the root elongation zone. Together with cytokinin signals and ethylene, this redistribution and modulation of auxin content triggers inhibition of the root cell elongation. Under B deficiency, root development is also regulated by brassinosteroids and jasmonic acid. Excess B can induce the production of reactive oxygen species (ROS). Abscisic acid and salicylic acid are both produced in response to B toxicity, and both can induce the antioxidant defense system to detoxify ROS. Another adaptation to B toxicity involves changes in the expression levels and activity of aquaporins in roots, thus reducing the uptake of water and delivery of B into the transpiration stream. In addition, abscisic acid mediates stomatal closure to further limit transpiration and the consequent accumulation of B in leaves.

Mechanistic insight on boron-mediated toxicity in plant vis-a-vis its mitigation strategies: a review.

Boron (B) is an essential micronutrient, crucial for the growth and development of crop plants. However, the essential to a toxic range of B in the plant is exceptionally narrow, and symptoms develop with a slight change in its concentration in soil. The morphological and anatomical response, such as leaf chlorosis, stunted growth, and impairment in the xylem and phloem development occurs under B-toxicity. The transport of B in the plant occurs via transpiration stream with the involvement of B-channels and transporter in the roots. The higher accumulation of B in source and sink tissue tends to have lower photosynthetic, chlorophyll content, infertility, failure of pollen tube formation and germination, impairment of cell wall formation, and disruption of membrane systems. Excess B in the plant hinders the uptake of other micronutrients, hormone transport, and metabolite partitioning. B-mediated reactive oxygen species production leads to the synthesis of antioxidant enzymes which help to scavenge these molecules and prevent the plant from further oxidative damage. This review highlights morpho-anatomical, physiological, biochemical, and molecular responses of the plant under B toxicity and thereby might help the researchers to understand the related mechanism and design strategies to develop B tolerant cultivars.

The physio-biochemical and molecular responses and mechanism of B uptake under its toxic condition have been illustrated. The spatial distribution of boron under its toxic condition and its accumulation in the plant might be regulated with sugar alcohols (polyols). This review throws light on the elevated level of B in the soil-plant system and provides management strategies for alleviating B toxicity in the plant.

High boron stress leads to sugar beet (Beta vulgaris L.) toxicity by disrupting photosystem â ¡.

This sugar beet acts as a soil remediator in areas where there are high levels of boron (B) in the soil, since it has a high requirement of boron (B) for growth, and has strong resistance to high B levels. Although B toxicity in different plants has been widely researched, little is known about the response of photosystem II (PSII) activity in sugar beet leaves to B toxicity at present. To clarify the growth and photosynthetic physiological response of sugar beet to B toxicity, the effects of different concentrations of H3BO3 (0.05, 1.5, 2.5,3.5 mM) on the growth, photosynthetic characteristics and antioxidant defense system of sugar beet seedlings were investigated by hydroponic experiments. In the present study, high B stress inhibited the growth of sugar beet and significantly decreased the biomass of the plants. There was a remarkable increase in the accumulation of B in the shoots, which affected photosynthesis and decreased the photosynthetic pigments. As B toxicity increased, leaf PSII activities and maximum photochemical efficiency of PSII (Fv/Fm) showed a tendency to decrease; at the same time, the photosynthetic performance index based on absorbed light energy (PIABS) decreased as well. Meanwhile, the energy allocation parameters of the PSII reaction center were changed, the light energy utilization capacity and the energy used for electron transfer were reduced and the thermal dissipation was increased at the same time. Furthermore, B toxicity decreased catalase (CAT) activity, increased peroxidase (POD) and superoxide dismutase (SOD) activities, and increased malondialdehyde (MDA) accumulation. According to the results obtained in this study, high B concentrations reduced the rate of photosynthesis and fluorescence, thus weakened antioxidant defense systems, and therefore inhibited the growth of sugar beet plants. Thus, in high B areas, sugar beet possesses excellent tolerance to high B levels and has a high B translocation capacity, so it can be used as a phytoremediation tool. This study provides a basis for the feasibility of sugar beet resistant to high B environments.