Selenium enhances ROS scavenging systems and sugar metabolism increasing growth of sugarcane plants.

Selenium (Se) beneficial effect on plants is related to an increase in nitrogen (N) assimilation and its role as an abiotic stress mitigator by reactive oxygen species (ROS) scavenging enhanced by antioxidant metabolism. This study aimed to evaluate sugarcane (Saccharum spp.) growth, photosynthetic and antioxidant responses, and sugar accumulation in response to Se supply. The experimental design was a factorial scheme 2 × 4: two sugarcane varieties (RB96 6928 and RB86 7515) and four Se application rates (0; 5; 10 and 20 µmol L-1) applied as sodium selenate in the nutrient solution. Leaf Se concentration increased under Se application in both varieties. The enzymes SOD (EC 1.15.1.1) and APX (EC 1.11.1.11) showed increase activities under Se application on variety RB96 6928. Nitrate reductase activity increased in both varieties resulting in the conversion of nitrate into higher total amino acids concentration indicating an enhanced N assimilation. This led to an increased concentration of chlorophylls and carotenoids, increased CO2 assimilation rate, stomatal conductance, and internal CO2 concentration. Selenium provided higher starch accumulation and sugar profiles in leaves boosting plant growth. This study shows valuable information regarding the role of Se on growth, photosynthetic process, and sugar accumulation in sugarcane leaves, which could be used for further field experiments. The application rate of 10 µmol Se L-1 was the most adequate for both varieties studied considering the sugar concentration and plant growth.

Selenium increases photosynthetic capacity, daidzein biosynthesis, nodulation and yield of peanuts plants (Arachis hypogaea L.).

This study aimed to investigate the roles of selenium (Se) application on the profile of photosynthetic pigments, oxidant metabolism, flavonoids biosynthesis, nodulation, and its relation to agronomic traits of peanut plants. Two independent experiments were carried out: one conducted in soil and the other in a nutrient solution. When the plants reached the V2 growth stage, five Se doses (0, 7.5, 15, 30, and 45 µg kg-1) and four Se concentrations (0, 5, 10, and 15 µmol L-1) were supplied as sodium selenate. The concentration of photosynthetic pigments, activity of antioxidant enzymes and the concentration of total sugars in peanut leaves increased in response to Se fertilization. In addition, Se improves nitrogen assimilation efficiency by increasing nitrate reductase activity which results in a higher concentration of ureides, amino acids and proteins. Se increases the synthesis of daidzein and genistein in the root, resulting in a greater number of nodules and concentration and transport of ureides to the leaves. Se-treated plants showed greater growth, biomass accumulation in shoots and roots, yield and Se concentration in leaves and grains. Our results contribute to food security and also to increase knowledge about the effects of Se on physiology, biochemistry and biological nitrogen fixation in legume plants.

Roles of selenium in mineral plant nutrition: ROS scavenging responses against abiotic stresses.

Agronomic biofortification of crops with selenium (Se) is an important strategy to minimize hidden hunger and increase nutrient intake in poor populations. Selenium is an element that has several physiological and biochemical characteristics, such as the mitigation of different types of abiotic stress. Selenoproteins act as powerful antioxidants in plant metabolism through the glutathione peroxidase (GSH) pathway, and provide an increased activity for enzymatic (SOD, CAT, and APX) and non-enzymatic (ascorbic acid, flavonoids, and tocopherols) compounds that act in reactive oxygen species (ROS) scavenging system and cell detoxification. Selenium helps to inhibit the damage caused by climate changes such as drought, salinity, heavy metals, and extreme temperature. Also, Se regulates antenna complex of photosynthesis, protecting chlorophylls by raising photosynthetic pigments. However, Se concentrations in soils vary widely in the earth's crust. Soil Se availability regulates the uptake, transport, accumulation, and speciation in plants. Foliar Se application at the concentration 50 g ha-1 applied as sodium selenate increases the antioxidant, photosynthetic metabolism, and yield of several crops. Foliar Se application is a strategy to minimize soil adsorption and root accumulation. However, the limit between the beneficial and toxic effects of Se requires research to establish an optimal dose for each plant species under different edaphoclimatic conditions. In this review, we present the compilation of several studies on agronomic biofortification of plants with Se to ensure food production and food security to mitigate hidden hunger and improve the health of the population.

Application of sodium selenate to cowpea (Vigna unguiculata L.) increases shoot and grain Se partitioning with strong genotypic interactions.

BACKGORUND: Cowpea is a crop widely used in developing countries due its rusticity. Besides its rich genotypic variability, most breeding programs do not explore its potential to improve elements uptake. Selenium (Se) is a scarce element in most soils, resulting in its deficiency being common in human diets. This study aimed to evaluate the interaction between biofortification with Se and genotypic variation in cowpea, on the concentrations of Se in roots, leaves + stem and grains. METHODS: Twenty-nine cowpea genotypes were grown in a greenhouse in the absence (control) and presence of Se (12.5 µg Se kg-1 soil) as sodium selenate, in fully randomized scheme. The plants were cultivated until grains harvest. The following variables were determined: roots dry weight (g), leaves + stems dry weight (g), grains dry weight (g), Se concentration (mg kg-1) in roots, leaves + stems and grains, and Se partitioning to shoots and grains. RESULTS: Selenium application increased the Se concentration in roots, leaves + stems and grains in all genotypes. At least twofold variation in grain Se concentration was observed among genotypes. Selenium application did not impair biomass accumulation, including grain dry weight. Genotype "BRS Guariba" had the largest Se concentration in grains and leaves + stems. Genotype MNC04-795 F-158 had the largest partitioning of Se to shoots and grain, due to elevated dry weights of leaves + stems and grain, and high Se concentrations in these tissues. CONCLUSION: This information might be valuable in future breeding programs to select for genotypes with better abilities to accumulate Se in grain to reduce widespread human Se undernutrition.

Selenium biofortification enhances ROS scavenge system increasing yield of coffee plants.

There are conclusive evidences of selenium (Se) deficiency in Brazilian soils and foods. Brazil is the largest producer and consumer of coffee worldwide, which favors agronomic biofortification of its coffee. This study aimed to evaluate effects of foliar application of three formulations and six rates of Se on antioxidant metabolism, agronomic biofortification and yield of coffee beans. Seven Se concentrations (0, 10, 20, 40, 80, 100 and 160 mg L-1) were applied from three formulations of Se (sodium selenate, nano-Se 1500, and nano-Se 5000). Selenium application up to 40 mg L-1 increased the concentration of photosynthetic pigments such as chlorophylls, pheophytins and carotenoids in coffee leaves. Foliar application of Se ranging from 20 to 80 mg L-1 decreased lipid peroxidation and concentration of hydrogen peroxide, but increased superoxide dismutase, ascorbate peroxidase, catalase and glutathione reductase activities in coffee leaves. These results indicated that foliar Se application stimulates antioxidative metabolism to mitigate reactive oxygen species. Foliar application of 20 mg Se L-1 of sodium selenate increased coffee yield by 38%, and 160 mg Se L-1 of nano-Se 5000 increased dramatically coffee yield by 42%. Selenium concentration in grains ranged from 0.116 to 4.47 mg kg-1 (sodium selenate), 4.84 mg kg-1 (nano-Se 1500) and 5.82 mg kg-1 (nano-Se 5000). The results suggest the beneficial effect of Se on the increment of photosynthetic pigments, antioxidative metabolism, increased coffee yield and nutritional quality of grains. The recommended foliar Se application in this study can mitigate abiotic stressors such as high temperatures resulting in higher yield of coffee plants.

Assessment of selenium spatial distribution using µ-XFR in cowpea (Vigna unguiculata (L.) Walp.) plants: Integration of physiological and biochemical responses.

Low concentrations of selenium (Se) are beneficial for plant growth. Foliar Se application at high concentrations is toxic to plants due to the formation of reactive oxygen species (ROS). This study characterized Se toxicity symptoms using X-ray fluorescence (XRF) technique in response to foliar Se application in cowpea plants. Five Se concentrations (0, 10, 25, 50, 100 e 150 g ha-1) were sprayed on leaves as sodium selenate. The visual symptoms of Se toxicity in cowpea leaves were separated into two stages: I) necrotic points with an irregular distribution and internerval chlorosis at the leaf limb border (50-100 g ha-1); II) total chlorosis with the formation of dark brown necrotic lesions (150 g ha-1). Foliar Se application at 50 g ha-1 increased photosynthetic pigments and yield. Ultrastructural analyses showed that Se foliar application above 50 g ha-1 disarranged the upper epidermis of cowpea leaves. Furthermore, Se application above 100 g ha-1 significantly increased the hydrogen peroxide concentration and lipid peroxidation inducing necrotic leaf lesions. Mapping of the elements in leaves using the XRF revealed high Se intensity, specifically in leaf necrotic lesions accompanied by calcium (Ca) as a possible attenuating mechanism of plant stress. The distribution of Se intensities in the seeds was homogeneous, without specific accumulation sites. Phosphorus (P) and sulfur (S) were found primarily located in the embryonic region. Understanding the factors involved in Se accumulation and its interaction with Ca support new preventive measurement technologies to prevent Se toxicity in plants.

Coffea arabica seedlings genotypes are tolerant to high induced selenium stress: Evidence from physiological plant responses and antioxidative performance.

Selenium (Se) is considered a beneficial element to higher plants based on its regulation of antioxidative system under abiotic or biotic stresses. However, the limit of beneficial and toxic physiological effects of Se is very narrow. In the present study, the antioxidant performance, nutritional composition, long-distance transport of Se, photosynthetic pigments, and growth of Coffea arabica genotypes in response to Se concentration in solution were evaluated. Five Coffea arabica genotypes (Obatã, IPR99, IAC125, IPR100 and Catucaí) were used, which were grown in the absence and presence of Se (0 and 1.0 mmol L-1) in nutrient solution. The application of 1 mmol L-1 Se promoted root browning in all genotypes. There were no visual symptoms of leaf toxicity, but there was a reduction in the concentration of phosphorus and sulfur in the shoots of plants exposed to high Se concentration. Except for genotype Obatã, the coffee seedlings presented strategies for regulating Se uptake by reducing long-distance transport of Se from roots to shoots. The concentrations of total chlorophyll, total pheophytin, and carotenoids were negatively affected in genotypes Obatã, IPR99, and IAC125 upon exposure to Se at 1 mmol L-1. H2O2 production was reduced in genotypes IPR99, IPR100, and IAC125 upon exposure to Se, resulting in lower activity of superoxide dismutase (SOD), and catalase (CAT). These results suggest that antioxidant metabolism was effective in regulating oxidative stress in plants treated with Se. The increase in sucrose, and decrease in SOD, CAT and ascorbate peroxidase (APX) activities, as well as Se compartmentalization in the roots, were the main biochemical and physiological modulatory effects of coffee seedlings under stress conditions due to excess of Se.

Selenium toxicity stress-induced phenotypical, biochemical and physiological responses in rice plants: Characterization of symptoms and plant metabolic adjustment.

Selenium (Se) at low concentration is considered benefit element to plants. The range between optimal and toxic concentration of Se is narrow and varies among plant species. This study aimed to evaluate the phenotypic, physiological and biochemical responses of four rice genotypes (BRS Esmeralda, BRSMG Relâmpago, BRS Bonança and Bico Ganga) grown hydroponically treated with sodium selenate (1.5 mM L-1). Selenium treated plants showed a dramatically decrease of soluble proteins, chlorophylls, and carotenoids concentration, resulting in the visual symptoms of toxicity characterized as leaf chlorosis and necrosis. Selenium toxicity caused a decrease on shoot and root dry weight of rice plants. Excess Se increased the oxidative stress monitored by the levels of hydrogen peroxide and lipid peroxidation. The enzymatic antioxidant system (catalase, superoxide dismutase, and ascorbate peroxidase) increased in response to Se supply. Interestingly, primary metabolism compounds such as sucrose, total sugars, nitrate, ammonia and amino acids increased in Se-treated plants. The increase in these metabolites may indicate a defense mechanism for the osmotic readjustment of rice plants to mitigate the toxicity caused by Se. However, these metabolites were not effective to minimize the damages on phenotypic traits such as leaf chlorosis and reduced shoot and root dry weight in response to excess Se. Increased sugars profile combined with antioxidant enzymes activities can be an effective biomarkers to indicate stress induced by Se in rice plants. This study shows the physiological attributes that must be taken into account for success in the sustainable cultivation of rice in environments containing excess Se.

Selenate and selenite affect photosynthetic pigments and ROS scavenging through distinct mechanisms in cowpea (Vigna unguiculata (L.) walp) plants.

Selenium (Se) is a beneficial element to higher plants. Application of Se at low concentrations enhances the antioxidant metabolism reducing the reactive oxygen species (ROS) generated by plant membrane cells. This study aimed to evaluate how the application of Se in the forms sodium selenate and sodium selenite regulates ROS scavenging in field-grown cowpea plants. Seven Se application rates (0; 2.5; 5; 10; 20; 40 and 60 g ha-1) of each of the two Se forms were applied to plants via the soil. Photosynthetic pigments concentration, gas exchange parameters, lipid peroxidation by malondialdehyde (MDA) concentration, hydrogen peroxide concentration, activity of catalase (CAT, EC:1.11.1.6), glutathione reductase (GR, EC:1.6.4.2), ascorbate peroxidase (APX, EC:1.11.1.11) and Se concentration in leaves and grains were evaluated. In general, Se application led to a decrease in chlorophyll a concentration whilst leading to an increase in chlorophyll b, indicating conservation of total chlorophyll concentration. Application of 2.5 g ha-1 of Se as selenate provided a notable increase in total chlorophyll and total carotenoids compared to the other application rates. Selenate and selenite application decreased lipid peroxidation. However, each Se source acted in a different pathway to combat ROS. While selenate showed more potential to increase activity of APX and GR, selenite showed a higher potential to increase CAT activity. The negative correlation between CAT and GR is indicative that both pathways might be activated under distinct circumstances. The more prominent activity of CAT under high rates of selenite resulted in a negative correlation of this enzyme with chlorophyll a and carotenoids. Both selenate and selenite application increased sucrose and total sugars concentration in leaves of cowpea plants. Overall, these results indicate that application of Se in cowpea under field conditions stimulates distinct pathways to scavenge ROS. This could prove beneficial to mitigate oxidative stress during plant development.

Selenium toxicity in upland field-grown rice: Seed physiology responses and nutrient distribution using the µ-XRF technique.

Selenium (Se) is an essential element for human and animal, although considered beneficial to higher plants. Selenium application at high concentration to plants can cause toxicity decreasing the physiological quality of seeds. This study aimed to characterize the Se toxicity on upland rice yield, seed physiology and the localization of Se in seeds using X-ray fluorescence microanalysis (µ-XRF). In the flowering stage, foliar application of Se (0, 250, 500, 1000, 1500, 2000 g ha-1) as sodium selenate was performed. A decrease in rice yield and an increase in seed Se concentrations were observed from 250 g Se ha-1. The storage proteins in the seeds showed different responses with Se application (decrease in albumin, increase in prolamin and glutelin). There was a reduction in the concentrations of total sugars and sucrose with the application of 250 and 500 g Se ha-1. The highest intensities Kα counts of Se were detected mainly in the endosperm and aleurone/pericarp. µ-XRF revealed the spatial distribution of sulfur, calcium, and potassium in the seed embryos. The seed germination decreased, and the electrical conductivity increased in response to high Se application rates showing clearly an abrupt decrease of physiological quality of rice seeds. This study provides information for a better understanding of the effects of Se toxicity on rice, revealing that in addition to the negative effects on yield, there are changes in the physiological and biochemical quality of seeds.

Selenium protects rice plants from water deficit stress.

Selenium (Se) is essential to humans and animals due to its antioxidant properties. Although it is not considered an essential nutrient for higher plants. Many studies show that Se in low concentrations (up to 0.5 mg kg-1) provides beneficial effects to non-hyperaccumulating plants by participating in antioxidant defense systems and enhancing tolerance to abiotic stress. Therefore, this study aimed to evaluate the effects of Se application rates on rice plants under different soil water conditions. The experiment was conducted on an Oxisol using four Se rates (0, 0.5, 1.0 and 2.0 mg kg-1) and two soil water conditions (irrigated and water deficit). Selenium application via soil up to 0.5 mg kg-1 increased the plant height, chlorophyll index, sulfur and copper accumulation in shoots, carbon dioxide assimilation, superoxide dismutase (EC 1.15.1.1) activity and decreased the hydrogen peroxide concentration in rice leaves. The accumulation of Se in shoot biomass and Se concentration in seeds increased linearly with the applied rates. Water deficit strongly decreased the plant growth and yield. However, rice plants treated with Se showed higher net photosynthesis, water use efficiency and antioxidant system. This study provides useful information about the roles of Se in protecting rice plants from water deficit stress.