Do Antimonite and Silicon Share the Same Root Uptake Pathway by Lsi1 in Sorghum bicolor L. Moench?

A study was conducted to further develop our understanding of antimony (Sb) uptake in plants. Unlike other metal(loid)s, such as silicon (Si), the mechanisms of Sb uptake are not well understood. However, SbIII is thought to enter the cell via aquaglyceroporins. We investigated if the channel protein Lsi1, which aids in Si uptake, also plays a role in Sb uptake. Seedlings of WT sorghum, with normal silicon accumulation, and its mutant (sblsi1), with low silicon accumulation, were grown in Hoagland solution for 22 days in the growth chamber under controlled conditions. Control, Sb (10 mg Sb L-1), Si (1mM) and Sb + Si (10 mg Sb L-1 + 1 mM Si) were the treatments. After 22 days, root and shoot biomass, the concentration of elements in root and shoot tissues, lipid peroxidation and ascorbate levels, and relative expression of Lsi1 were determined. When mutant plants were exposed to Sb, they showed almost no toxicity symptoms compared to WT plants, indicating that Sb was not toxic to mutant plants. On the other hand, WT plants had decreased root and shoot biomass, increased MDA content and increased Sb uptake compared to mutant plants. In the presence of Sb, we also found that SbLsi1 was downregulated in the roots of WT plants. The results of this experiment support the role of Lsi1 in Sb uptake in sorghum plants.

Physiological and Biochemical Responses of Commercial Strawberry Cultivars under Optimal and Drought Stress Conditions.

Improving the extent of adaptation and the choice of the most tolerant cultivar is the first step to mitigating the adverse effects of limited water, especially in susceptible plants such as strawberries. To address this issue, two commercial strawberry cultivars (Camarosa and Gaviota) were compared when irrigated to match 100, 75, 50, and 25% field capacity (FC) to simulate the control, slight, moderate, and severe drought stress conditions, respectively. Drought stress induced the reduction of total chlorophyll, carotenoid, relative water content, and phenolic content significantly, whereas the activity of antioxidant enzymes, electrolyte leakage, osmolyte accumulation, and oxidative markers upsurged progressively in drought severity-dependent behavior. Gaviota produced more proline, hydrogen peroxide as a marker of membrane lipid peroxidation and disposed of by higher electrolyte leakage, significantly. On the other hand, Camarosa having higher soluble carbohydrates as well as enzymatic and non-enzymatic antioxidants could be considered a drought-tolerant cultivar. Genotypic variation between these cultivars could be used in breeding projects to promote drought-tolerant strawberries in the future.

Multiple effects of silicon on alleviation of arsenic and cadmium toxicity in hyperaccumulator Isatis cappadocica Desv.

Arsenic (As) and cadmium (Cd) belong to the group of major pollutants extremely toxic to plants. Metal hyperaccumulating plants play an important role in phytoextraction of heavy metals. Silicon (Si) plays an important role in the amelioration of heavy metal stress through physio-biochemical mechanisms, which remain poorly understood in hyperaccumulators. The main purpose of this study was to determine the impact of Si on growth and performance of As hyperaccumulator Isatis cappadocica Desv., exposed to As and Cd. Results showed that Si (especially at 1 mM level) alleviated the harmful impact of As/Cd and significantly increased the root and shoot biomass, root and shoot length and chlorophyll contents of I. cappadocica by enhancing the plant defense mechanisms. Between the two investigated harmful elements, As was accumulated in plant parts significantly more than Cd, however with considerably lower toxic growth effects. The As/Cd concentration, bioaccumulation and translocation factor and total As content both in roots and shoots of Si-supplied plant were significantly reduced as a protective mechanism, especially in Cd exposed plant. In comparison with single As/Cd treatment, Si supply reduced H2O2 content, increased total soluble protein content and enhanced glutathione S-transferase activity in shoots. The results of this study clearly showed that Si minimized As/Cd uptake and root to shoot translocation, and therefore Si cannot enhance the phytoextraction potential of this plant species. Additionally, Si-improved growth and reduced oxidative damages caused by excess of As and Cd suggested that the similar mechanisms of metal(loid) alleviation are adopted in hyperaccumulators as well as non-hyperaccumulating plants.

Silicon alleviates antimony phytotoxicity in giant reed (Arundo donax L.).

MAIN CONCLUSION: Silicon enhances photosynthetic efficiency, biomass, and lignification of root structures possibly limiting antimony translocation and mitigating phytotoxicity in giant reed plants. Antimony (Sb) is a non-essential metalloid causing toxic effects in plants. Silicon has been reported to impart tolerance against biotic and abiotic stress in plants. Fast-growing plant, giant reed (Arundo donax L.) is a promising energy crop, can be a suitable plant for phytoremediation. However, information regarding the tolerance capacity with respect to Sb toxicity and potential of Si to mitigate the Sb phytotoxicity in giant reed are very scarce. Rhizomes of giant reed were grown for ten weeks in hydroponics exposed to Sb, Si, and their combination wherein treatment without Sb/Si served as control. Effect of these treatments on rate of net photosynthesis and photosynthetic pigments, phytoextraction ability of Sb, Si and Sb uptake, plant biomass, and lignification and suberization of roots along with localization of Sb and Si were analysed. We found that Si considerably improved the growth and biomass of giant reed under Sb toxicity. Antimony reduced the photosynthesis and decreased the content of photosynthetic pigments, which was completely alleviated by Si. Silicon amendment to Sb treated plants enhanced root lignification. Silicon enhanced lignification of root structures probably restricted the Sb translocation. However, co-localization of Sb with Si has not been observed neither at the shoot nor at the root levels. Similarly, Sb was also not detected in leaf phytoliths. These findings suggest that Si treatment promotes overall plant growth by improving photosynthetic parameters and decreasing Sb translocation from root to shoot in giant reed by improving root lignification.

Effect of silicon on root growth, ionomics and antioxidant performance of maize roots exposed to As toxicity.

Nowadays, one of the biggest challenges of plant physiology is to find out the ways how to mitigate negative impacts of abiotic stress on plants. It is the pollution of groundwater or soil by various metals and metalloids that significantly affects the quality of life. Both arsenic (As) and silicon (Si) are metalloids - while the first one is toxic in general, the latter one is considered as beneficial for plants suffering from various kinds of stresses. The aim of our work was to elucidate the growth and development of young maize (Zea mays L.) plants exposed to both of these metalloids simultaneously. Experiments were focused on the comparison of root growth and biomass allocation, changes in uptake of macro- and micronutrients, visualisation of free radicals along with monitoring of the dynamics of main antioxidant enzymes activity in roots. The results showed that increasing concentration of As (75 and 150 µM As) severely inhibited root length and the amount of biomass, and addition of Si (2.5 mM) to the medium containing As did not have a significant effect on root growth. Similarly, the application of Si did not influence the uptake of macro- and microelements into the roots (mainly Ca, P, K, Mo, Cu, Zn and Ni) which was mostly decreased due to As. On the other hand, Si significantly decreased the presence of both superoxide and hydrogen peroxide in roots that suffered from As toxicity. Although the overall growth of maize plants was not improved by Si amendment, we assume that Si might affect the functionality of key antioxidant enzymes in time, and in this way at least partially help to overcome negative effects of As on maize roots.

Mitigation of climate change and environmental hazards in plants: Potential role of the beneficial metalloid silicon.

In the last decades, the concentration of atmospheric CO2 and the average temperature have been increasing, and this trend is expected to become more severe in the near future. Additionally, environmental stresses including drought, salinity, UV-radiation, heavy metals, and toxic elements exposure represent a threat for ecosystems and agriculture. Climate and environmental changes negatively affect plant growth, biomass and yield production, and also enhance plant susceptibility to pests and diseases. Silicon (Si), as a beneficial element for plants, is involved in plant tolerance and/or resistance to various abiotic and biotic stresses. The beneficial role of Si has been shown in various plant species and its accumulation relies on the root's uptake capacity. However, Si uptake in plants depends on many biogeochemical factors that may be substantially altered in the future, affecting its functional role in plant protection. At present, it is not clear whether Si accumulation in plants will be positively or negatively affected by changing climate and environmental conditions. In this review, we focused on Si interaction with the most important factors of global change and environmental hazards in plants, discussing the potential role of its application as an alleviation strategy for climate and environmental hazards based on current knowledge.

Effect of silicon on the young maize plants exposed to nickel stress.

Nickel (Ni) is involved in several physiological processes in plants but its excess in environment has many phytotoxic effects. Silicon (Si), an element required for optimal plant performance, has been shown to have beneficial effects for plants coping with various types of stresses. Here we studied the alleviative potential of Si (2.5 mM) added to hydroponically grown maize (Zea mays L.) plants under Ni (100 µM) stress. Ni decreased most of the growth parameters, total chlorophyll (Chl) and leaf relative water content (RWC), and catalase (CAT; EC 1.11.1.6) activity, while leaf water loss (LWL), contents of proline (Pro), hydrogen peroxide (H2O2) and ascorbate (AsA), membrane lipid peroxidation and activities of peroxidase (POX; EC 1.11.1.7) and superoxide dismutase (SOD; EC 1.15.1.1) were increased. Supplementation of Si to Ni-treated plants enhanced the leaf area, Chl content, RWC, CAT and POX (only in younger leaf) activities and decreased LWL, the contents of Pro (in younger leaf), H2O2 (roots) and AsA, lipid peroxidation and POX and SOD activities. We may conclude that Si mitigated the Ni-induced stress in maize by amelioration of the leaf water deficient status (Pro, RWC, LWL), enhancing membrane stability (MDA) and influencing enzymatic (SOD, POX, CAT) and non-enzymatic (Pro, AsA) defence systems. The increased Chl content and leaf area improve overall plant performance.

Comparison of the single and combined effects of arsenic and antimony on growth and physiology of giant reed (Arundo donax L.).

Fast-growing plant, giant reed (Arundo donax L.) has been gaining a lot of popularity in the phytoremediation of metal-polluted soils. However, information regarding the physiological background of tolerance and accumulation capacity of A. donax with respect to antimony (Sb), arsenic (As), and their co-contamination are very limited. Rooted stem cuttings were grown for 5 months in hydroponics exposed to Sb (10 mg L-1), As (10 mg L-1), and their combined toxicity (Sb 5 mg L-1 + As 5 mg L-1) wherein treatment without As/Sb served as control. Effect of these treatments on key photosynthetic parameters (rate of net photosynthesis, effective quantum yield of photosystem II, chlorophyll fluorescence, and photosynthetic pigments), phytoextraction ability of metalloids, nutrient uptake, root growth, and lignification were analyzed. Arsenic-containing treatments severely affected root morphology of A. donax compared to Sb/control and plants exposed to As showed intensive lignification already in young apical part of the root in the present study. Shoot concentration was found to be 11.35±0.75 Sb mg kg-1 and 8.97±0.52 As mg kg-1 compared to root concentration of 1028.3±19.1 Sb mg kg-1 and 705.3±69.9 As mg kg-1 in the treatments of Sb and As. Even though Sb and As were translocated to the shoots in relatively small amount, both metalloids significantly decreased the shoot and root growth of A. donax and negatively affected the photosynthetic parameters. Moreover, co-contamination of Sb and As proved to be severely toxic to growth and physiology of A. donax even though the magnitudes of the metalloids used were lower than those of Sb/As alone treatments. In conclusion, Sb and As caused a marked reduction in growth and physiological characteristics of A. donax, opposing its use in phytoremediation of highly contaminated soils. Tolerance capacity of plants to simultaneous presence of As and Sb in the environment is crucial for the successful implementation of phytoremediation since the co-contamination by As and Sb might reduce the efficiency of phytoremediation when using this fast-growing and high biomass-producing plant species.

Divergence of reactions to arsenic (As) toxicity in tobacco (Nicotiana benthamiana) plants: A lesson from peroxidase involvement.

To evaluate the multiplicity of reactions to toxic metalloid arsenic (As) with specific emphasis on the role of plant peroxidases, a model plant Nicotiana benthamiana was cultivated in in vitro conditions at various doses of As (applied as As5+ up to 80 µM). After 28-day cultivation, several physiological characteristics such as plant growth, photosynthetic pigment concentration, As concentration, peroxidase (POX) expression levels, and POX activity were evaluated. A newly sequenced gene for POX has been identified, that belongs to the Class III plant extracellular peroxidases, and its relationship to the genus Solanum as the most relative species has been confirmed. In the control and selected As treatments (20As, 50As, and 80As), newly identified POX expression and POX activity were continuously detected during the whole cultivation period. The plant reactions to As stress were distinguished into three groups: low As, moderate As, and high As. A tight relationship was found between the photosynthetic pigments and POX expression. Accumulation of As in roots and shoots showed correlations with POX activities. The results showed that the diversity of reactions depends on As dose and time exposure and indicate an interface of peroxidase functional role with other physiological processes in plants suffering from As toxicity.

Aluminum toxicity and aluminum stress-induced physiological tolerance responses in higher plants.

Aluminum (Al) precipitates in acidic soils having a pH < 5.5, in the form of conjugated organic and inorganic ions. Al-containing minerals solubilized in the soil solution cause several negative impacts in plants when taken up along with other nutrients. Moreover, a micromolar concentration of Al present in the soil is enough to induce several irreversible toxicity symptoms such as the rapid and transient over-generation of reactive oxygen species (ROS) such as superoxide anion (O2•-), hydrogen peroxide (H2O2), and hydroxyl radical (•OH), resulting in oxidative bursts. In addition, significant reductions in water and nutrient uptake occur which imposes severe stress in the plants. However, some plants have developed Al-tolerance by stimulating the secretion of organic acids like citrate, malate, and oxalate, from plant roots. Genes responsible for encoding such organic acids, play a critical role in Al tolerance. Several transporters involved in Al resistance mechanisms are members of the Aluminum-activated Malate Transporter (ALMT), Multidrug and Toxic compound Extrusion (MATE), ATP-Binding Cassette (ABC), Natural resistance-associated macrophage protein (Nramp), and aquaporin gene families. Therefore, in the present review, the discussion of the global extension and probable cause of Al in the environment and mechanisms of Al toxicity in plants are followed by detailed emphasis on tolerance mechanisms. We have also identified and categorized the important transporters that secrete organic acids and outlined their role in Al stress tolerance mechanisms in crop plants. The information provided here will be helpful for efficient exploration of the available knowledge to develop Al tolerant crop varieties.

Multiple effects of silicon on alleviation of nickel toxicity in young maize roots.

Although beneficial metalloid silicon (Si) has been shown to alleviate the toxicity of various heavy metals, there is a lack of knowledge about the role of Si in possible alleviation of phytotoxicity caused by excess of essential nickel (Ni). In the present study we investigated the growth and biomass production, reactive oxygen species (ROS) formation and activities of selected antioxidants, as well as combined effect of Ni and Si on the integrity of cell membranes and electrolyte leakage in young maize roots treated for 24, 48 and 72 h with excess of Ni and/or Si. By histochemical methods we also visualized Ni distribution in root tissues and compared the uptake of Ni and Si with the development of root apoplasmic barriers. Ni enhanced the root lignification and suberization and shifted the development of apoplasmic barriers towards the root tip. Similarly, localization of Ni correlated with lignin and suberin deposition in root endodermis, further supporting the barrier role of this tissue in Ni uptake. Si reversed the negative impact of Ni on root anatomy. Additionally, improved cell membrane integrity, and enhanced ascorbate-based antioxidant system might be the mechanisms how Si partially mitigates the deleterious effects of Ni excess in maize plants.

Silicon crosstalk with reactive oxygen species, phytohormones and other signaling molecules.

Exogenous applications of silicon (Si) can initiate cellular defence pathways to enhance plant resistance to abiotic and biotic stresses. Plant Si accumulation is regulated by several transporters of silicic acid (e.g. Lsi1, Lsi2, and Lsi6), but the precise mechanisms involved in overall Si transport and its beneficial effects remains unclear. In stressed plants, the accumulation of Si leads to a defence mechanism involving the formation of amorphous or hydrated silicic acid caused by their polymerization and interaction with other organic substances. Silicon also regulates plant ionic homeostasis, which involves the nutrient acquisition, availability, and replenishment in the soil through biogeochemical cycles. Furthermore, Si is implicated in modulating ethylene-dependent and jasmonate pathways, as well as other phytohormones, particularly under stress conditions. Crosstalk between Si and phytohormones could lead to improvements in Si-mediated crop growth, especially when plants are exposed to stress. The integration of Si with reactive oxygen species (ROS) metabolism appears to be a part of the signaling cascade that regulates plant phytohormone homeostasis, as well as morphological, biochemical, and molecular responses. This review aims to provide an update on Si interplays with ROS, phytohormones, and other signaling molecules that regulate plant development under stress conditions.

Structural modifications of plant organs and tissues by metals and metalloids in the environment: A review.

At the dawn of the industrial revolution, the exorbitant use of heavy metals and toxic elements by mankind unfurls a powerful and complex web of hazard all around the world that significantly contributed to unprecedented trends in environmental degradation. Plants as sessile organisms, that cannot escape from the stress directly, have adapted to this environment via concurrent configurations of several traits. Among them the anatomy has been identified as much more advanced field of research that brought the explosion of interest among the expertise and its prodigious importance in stress physiology is unavoidable. In conjunction with various other disciplines, like physiology, biochemistry, genomics and metabolomics, the plant anatomy provides a large data sets that are paving the way towards a comprehensive and holistic understanding of plant growth, development, defense and productivity under heavy metal and toxic element stress. Present paper advances our recent knowledge about structural alterations of plant tissues induced by metals and metalloids, like antimony (Sb), arsenic (As), aluminium (Al), copper (Cu), cadmium (Cd), chromium (Cr), lead (Pb), manganese (Mn), mercury (Hg), nickel (Ni) and zinc (Zn) and points on essential role of plant anatomy and its understanding for plant growth and development in changing environment. Understanding of anatomical adaptations of various plant organs and tissues to heavy metals and metalloids could greatly contribute to integral and modern approach for investigation of plants in changing environmental conditions. These findings are necessary for understanding of the whole spectra of physiological and biochemical reactions in plants and to maintain the crop productivity worldwide. Moreover, our holistic perception regarding the processes underlying the plant responses to metal(loids) at anatomical level are needed for improving crop management and breeding techniques.

Aluminum toxicity in plants and its possible mitigation in acid soils by biochar: A review.

Toxicity of aluminum (Al) is a serious problem for agricultural plants, especially due to excessive soil acidification caused by continuous intensive agriculture and modified environmental conditions related with global climate change. Decreased root elongation and shoot growth, reduced biomass production, nutrient imbalance and altered physiological and metabolic processes are responsible for lower yield and crop quality and therefore, decreased variability and productivity of the land. Recently, biochar is gaining popularity for ameliorating metal toxicity in soils. However, there is a lack of comprehensive information regarding the effects of biochar and its functioning. Multiple mechanisms are involved in ameliorating Al toxicity in which inherent properties of biochar influencing Al adsorption, absorption, complexation, cation exchange and electrostatic interaction are considered to play major roles. Modification of biochar to enhance these mechanisms might hold the key for long term solution. Present review indicates gaps for further research. Long term field studies are needed to understand the effects of biochar on Al toxicity.

Alleviation mechanisms of metal(loid) stress in plants by silicon: a review.

Silicon (Si), although not considered as an essential element for plants in general, can ameliorate the phytotoxicity induced by excess metal(loid)s whether non-essential (e.g. Cd, Pb, Cr, Al, As, and Sb) or essential (e.g. Cu, Ni, and Zn). The Si-enhanced resistance allowing plants to cope with this type of abiotic stress has been developed at multiple levels in plants. Restriction of root uptake and immobilization of metal(loid)s in the rhizosphere by Si is probably one of the first defence mechanism. Further, retention of elements in the root apoplasm might enhance the resistance and vigour of plants. At the cellular level, the formation of insoluble complexes between Si and metal(loid)s and their storage within cell walls help plants to decrease available element concentration and restrict symplasmic uptake. Moreover, Si influences the oxidative status of plants by modifying the activity of various antioxidants, improves membrane stability, and acts on gene expression, although its exact role in these processes is still not well understood. This review focuses on all currently known plant-based mechanisms related to Si supply and involved in amelioration of stress caused by excess metal(loid)s.

A new insight on structural and some functional aspects of peri-endodermal thickenings, a specific layer in Noccaea caerulescens roots.

BACKGROUND AND AIMS: Cell walls of the peri-endodermis, a layer adjacent to the endodermis in alpine pennycress (Noccaea caerulescens) roots, form C-shaped peri-endodermal thickenings (PETs). Despite its specific position close to the endodermis, the assumed similarity of PETs to phi thickenings in many other species, and the fact that N. caerulescens is a well-studied heavy-metal-hyperaccumulating plant, the PET as a root trait is still not understood. METHODS: Here, we characterized PET cell walls by histochemical techniques, Raman spectroscopy, immunolabelling and electron microscopy. Moreover, a role of PETs in solute transport was tested and compared with Arabidopsis thaliana plants, which do not form PETs in roots. KEY RESULTS: Cell walls with PETs have a structured relief mainly composed of cellulose and lignin. Suberin, typical of endodermal cells, is missing but pectins are present on the inner surface of the PET. Penetrating dyes are not able to cross PETs either by the apoplasmic or the symplasmic pathway, and a significantly higher content of metals is found in root tissues outside of PETs than in innermost tissues. CONCLUSIONS: Based on their development and chemical composition, PETs are different from the endodermis and closely resemble phi thickenings. Contrarily, the different structure and dye impermeability of PETs, not known in the case of phi thickenings, point to an additional barrier function which makes the peri-endodermis with PETs a unique and rare layer.

Silicification of Root Tissues.

Silicon (Si) is not considered an essential element, however, its tissue concentration can exceed that of many essential elements in several evolutionary distant plant species. Roots take up Si using Si transporters and then translocate it to aboveground organs. In some plant species, root tissues are also places where a high accumulation of Si can be found. Three basic modes of Si deposition in roots have been identified so far: (1) impregnation of endodermal cell walls (e.g., in cereals, such as Triticum (wheat)); (2) formation of Si-aggregates associated with endodermal cell walls (in the Andropogoneae family, which includes Sorghum and Saccharum (sugarcane)); (3) formation of Si aggregates in "stegmata" cells, which form a sheath around sclerenchyma fibers e.g., in some palm species (Phoenix (date palm)). In addition to these three major and most studied modes of Si deposition in roots, there are also less-known locations, such as deposits in xylem cells and intercellular deposits. In our research, the ontogenesis of individual root cells that accumulate Si is discussed. The documented and expected roles of Si deposition in the root is outlined mostly as a reaction of plants to abiotic and biotic stresses.

Silicon Uptake and Localisation in Date Palm (Phoenix dactylifera) - A Unique Association With Sclerenchyma.

Date palm (Phoenix dactylifera) can accumulate as much as 1% silicon (Si), but not much is known about the mechanisms inherent to this process. Here, we investigated in detail the uptake, accumulation and distribution of Si in date palms, and the phylogeny of Si transporter genes in plants. We characterized the PdNIP2 transporter following heterologous expression in Xenopus oocytes and used qPCR to determine the relative expression of Si transporter genes. Silicon accumulation and distribution was investigated by light microscopy, scanning electron microscopy coupled with X-ray microanalysis and Raman microspectroscopy. We proved that PdNIP2-1 codes for a functional Si-permeable protein and demonstrated that PdNIP2 transporter genes were constitutively expressed in date palm. Silicon aggregates/phytoliths were found in specific stegmata cells present in roots, stems and leaves and their surfaces were composed of pure silica. Stegmata were organized on the outer surface of the sclerenchyma bundles or associated with the sclerenchyma of the vascular bundles. Phylogenetic analysis clustered NIP2 transporters of the Arecaceae in a sister position to those of the Poaceae. It is suggested, that Si uptake in date palm is mediated by a constitutively expressed Si influx transporter and accumulated as Si aggregates in stegmata cells abundant in the outer surface of the sclerenchyma bundles (fibers).