Selenium uptake in Zea mays supplied with selenate or selenite under hydroponic conditions

Background and aims

Selenium is an essential micro-nutrient for animals, humans and microorganisms; it mainly enters food chains through plants. This study proposes to explore effect of inorganic Se forms on its uptake and accumulation in Zea mays.

Methods

Zea mays was grown in a controlled-atmosphere chamber for 2 weeks in a hydroponic solution of low-concentration selenium (10 μg/L (i.e.0.12 μM) or 50 μg/L (i.e. 0.63 μM) of Se). For each concentration, four treatments were defined: control (without selenium), selenite alone, selenate alone and selenite and selenate mixed.

Results

At low concentrations, selenium did not affect the biomass production of Zea mays. However, for both concentrations, Se accumulation following a selenite-only treatment was always higher than with selenate-only. Moreover, in the selenate-only treatment, Se mainly accumulated in shoots whereas in the selenite-only treatment, Se was stocked more in the roots. Interactions between selenate and selenite were observed only at the higher concentration (0.63 μM of selenium in the nutrient solution).

Conclusions

Se form and concentration in the nutrient solution strongly influenced the absorption, allocation and metabolism of Se in Zea mays. Selenate seems to inhibit selenite absorption by the roots.     

Nitrogen-Use Efficiency in Relation to Different Forms and Application Rates of Se in Lettuce Plants

The main objective of this work was to determine whether nitrogen-use efficiency (NUE) was affected by the application of different forms and dosages of selenium (Se), to ascertain the influence of this trace element in a biofortification program in lettuce plants. The parameters analyzed were biomass, NO₃ ⁻ concentration, and total reduced N as well as those defining NUE in plants: total nitrogen accumulation (TNA), nitrogen efficient ratio (NER), nitrogen-utilization efficiency (NUtE), and nitrogen-uptake efficiency (NUpE). According to our results, application of Se affected NUE in lettuce plants. With the application of selenite as well as selenate NO₃ ⁻ uptake was reduced, thus diminishing the NUpE and the foliar concentration of this anion. In addition, selenate application at a rate of 20 μM and selenite at 5 μM induced N utilization, reflected by an increase in NER and NUtE; this result coincides with augmented biomass production. Notably, our results indicate that when Se is applied at high rates, selenite is far more phytotoxic, this being associated with a higher reduction of NUE in these plants.     

Evaluation of the Potential of Peas (Pisum sativum L.) to Be Used in Selenium Biofortification Programs Under Mediterranean Conditions

Selenium (Se), which has antioxidant, anticancer, and antiviral properties, is an essential micronutrient for humans and animals. This micronutrient is found in high quantity in legumes. Peas have an ever-increasing importance in Spain, and to increase their nutritional value, two foliar Se fertilizers: sodium selenate and sodium selenite, at five different rates: 0, 10, 20, 40, 80 g?ha^?1, were studied during the 2010/2011 crop season on semiarid Mediterranean conditions. Sodium selenate was much more effectively taken up by plants compared to sodium selenite. There was a strong linear relationship between the total Se content and Se rate in both sodium selenate and selenite. For each gram of Se fertilization as either sodium selenate or sodium selenite, the increase of total Se concentration in the grain was 148 and 19 μg Se?kg^?1 dry weight, respectively. Ingestion of 100 g of peas previously fertilized with 10 g of sodium selenate per hectare would result in an intake of 179 μg of Se. This is almost 90 % of the daily recommended dose needed to reduce the chance of some cancers and about 179 % of the minimum concentration required to prevent Se deficiency diseases in animals. The pea has shown to have a strong ability to uptake and accumulate Se under Mediterranean conditions; therefore, this would make it a very strong candidate for inclusion in biofortification programs aiming to increase Se in the food chain.     

Selenium accumulation in lettuce germplasm

Selenium (Se) is an essential micronutrient for animals and humans. Increasing Se content in food crops offers an effective approach to reduce the widespread selenium deficiency problem in many parts of the world. In this study, we evaluated 30 diverse accessions of lettuce (Lactuca sativa L.) for their capacity to accumulate Se and their responses to different forms of Se in terms of plant growth, nutritional characteristics, and gene expression. Lettuce accessions responded differently to selenate and selenite treatment, and selenate is superior to selenite in inducing total Se accumulation. At least over twofold change in total Se levels between cultivars with high and low Se content was found. Synergistic relationship between Se and sulfur accumulation was observed in nearly all accessions at the selenate dosage applied. The change in shoot biomass varied between lettuce accessions and the forms of Se used. The growth-stimulated effect by selenate and the growth-inhibited effect by selenite were found to be correlated with the alteration of antioxidant enzyme activities. The different ability of lettuce accessions to accumulate Se following selenate treatment appeared to be associated with an altered expression of genes involved in Se/S uptake and assimilation. Our results provide important information for the effects of different forms of Se on plant growth and metabolism. They will also be of help in selecting and developing better cultivars for Se biofortification in lettuce.     

Regulation of sulphur assimilation in lettuce plants in the presence of selenium

Selenium (Se) is considered an essential trace element for animals because of its nutritional and clinical value, including its special relevance in cancer prevention, and thus Se is at present used in biofortification programmes. However, possible effects of Se application on S metabolism and plant growth are still not clear. Thus, we analysed the effect that Se application in two different forms (selenate versus selenite) exerts on the S metabolism in lettuce plants grown for 66 days. Our results indicate that the application of selenite as opposed to selenate does not affect the foliar concentration of S. With respect to different enzymes in charge of sulphate (SO₄²⁻) assimilation, the ATP-sulphurylase activity varies only with the application of different rates of selenate, while the activity of O-acetylserine(thiol)lyase (OAS-TL) and serine-acetyltransferase (SAT) increase in activity mainly when selenite is applied. Finally, the concentration of cysteine (Cys) and total thiols (SH-total), fundamentally in the selenate treatments, increased with shoot biomass. In conclusion, this study confirms that the form and application rate of Se affects S assimilation, selenate being the more suitable form to improve effectiveness of the biofortification programme with this trace element.     

Tracking Se Assimilation and Speciation through the Rice Plant – Nutrient Competition,Toxicity and Distribution

Up to 1 billion people are affected by low intakes of the essential nutrient selenium (Se) due to low concentrations in crops. Biofortification of this micronutrient in plants is an attractive way of increasing dietary Se levels. We investigated a promising method of Se biofortification of rice seedlings, as rice is the primary staple for 3 billion people, but naturally contains low Se concentrations. We studied hydroponic Se uptake for 0–2500 ppb Se, potential phyto-toxicological effects of Se and the speciation of Se along the shoots and roots as a function of added Se species, concentrations and other nutrients supplied. We found that rice germinating directly in a Se environment increased plant-Se by factor 2–16, but that nutrient supplementation is required to prevent phyto-toxicity. XANES data showed that selenite uptake mainly resulted in the accumulation of organic Se in roots, but that selenate uptake resulted in accumulation of selenate in the higher part of the shoot, which is an essential requirement for Se to be transported to the grain. The amount of organic Se in the plant was positively correlated with applied Se concentration. Our results indicate that biofortification of seedlings with selenate is a successful method to increase Se levels in rice.     

Toxicity of selenium to Lemna minor in relation to sulfate concentration

The aquatic plant Lemna minor L. was treated with sodium selenite or sodium selenate to test the toxicity of these salts in relation to high or low levels of sulfate in the culture medium. Several morphophysiological aspects, such as multiplication rate (MR), ratio of the number of fronds to number of colonies (Nfr/Ncol), frond size, cell ultrastructure, pigment content and guaiacol peroxidase (EC 1.11.1.7) activity were evaluated. Their variations might be an indirect means of evaluating the degree of susceptibility or tolerance of this plant to selenium (Se). Sodium selenite or sodium selenate treatments at concentrations ranging from 1 to 256 μ M generally decreased the investigated parameters. Moreover, the sulfate concentration influenced the toxicity of both Se salts. In general, with treatments in a medium containing a high sulfate (HS) content, sodium selenite appeared more toxic than sodium selenate, whereas in a low sulfate (LS) medium, sodium selenate seemed more toxic. MR was significantly increased at 1–4 μ M selenite and LS or 8 μ M selenate and HS levels, suggesting that Se may be an essential nutrient for this plant.     

Effect of byproduct,nitrogen fertilizer,and zeolite on phosphate rock dissolution and extractable phosphorus in acid soil

The relative plant availability of selenate versus selenite depends on the concentrations of competing ions, specifically sulfate and phosphate, respectively. In solution culture, the concentration of phosphate is typically 100- to 1000-fold greater than in soil solution, an artifact that could lead to underestimation of the phytoavailability of selenite. A nutrient solution study was conducted to compare the availability of selenite and selenate to perennial ryegrass (Lolium perenne L. cv. Evening Shade) and strawberry clover (Trifolium fragiferrum L. cv. O'Conner) at basal concentrations of SO₄ (0.5 mM) and PO₄ (2 μM) similar to those found in soil solution. Concentrations up to 5 mM SO₄ and 200 μM PO₄ allowed quantitative comparison of the efficacy of the competing ions. In both species, selenite was more phytotoxic than selenate, especially for shoot growth. Selenate was less toxic, and tended to preferentially inhibit root growth. Translocation percentages were much higher with selenate (≥84%) than with selenite (≤47%). A 10-fold increase in sulfate decreased uptake from selenate by >90% in both species. In ryegrass, 10-fold increases in phosphate caused 30% to 50% decreases in Se accumulation from selenite, but in clover such decreases only occurred in the roots. Sulfate-selenate antagonisms were thus stronger than phosphate-selenite antagonisms. Nonetheless, conventional nutrient solutions with millimolar phosphate will significantly underestimate Se availability from selenite, and moderate levels of sulfate salinity can inhibit selenate uptake sufficiently to reverse the apparent relative availability of the two forms of Se. This revised version was published online in June 2006 with corrections to the Cover Date.     

Selenium Species and Their Distribution in Freshwater Fish from Argentina

The distribution and speciation of selenium (Se) in freshwater fish (muscle and liver tissue) from lakes in Argentina was investigated. Three introduced species, brown trout (Salmo trutta), rainbow trout (Oncorhynchus mykiss) and brook trout (Salvelinus fontinalis), and one native species, creole perch (Percichthys trucha), were investigated. Values for total selenium in muscle ranged from 0.66 to 1.61 μg/g, while in the liver, concentrations were much higher, from 4.46 to 73.71 μg/g on a dry matter basis. Separation of soluble Se species (SeCys₂, selenomethionine (SeMet), SeMeSeCys, selenite and selenate) was achieved by ion exchange chromatography and detection was performed by inductively coupled plasma–mass spectrometry. The results showed that in fish muscle, from 47 to 55 % of selenium was soluble and the only Se species identified was SeMet, which represented around 80 % of soluble Se, while in the liver, the amount of soluble Se ranged from 61 to 76 % and the percentage of species identified (SeMet and SeCys₂) was much lower and ranged from 8 to 17 % of soluble Se.     

Proteomic analysis of selenium embryotoxicity in cultured postimplantation rat embryos

BACKGROUND: Developmental toxicity of selenium (Se) is a nutritional, environmental and medicinal concern. Here, we investigated Se embryotoxicity by proteomic analysis of cultured rat embryos. METHODS: Rat embryos at day 9.5 or 10.5 of gestation were cultured for 48 or 24 h, respectively, in the presence of sodium selenate (100 or 150 µM) or sodium selenite (20 or 30 µM). Proteins from the embryo proper and yolk sac membrane were analyzed by two‐dimensional electrophoresis for quantitative changes from those in control embryos. Proteins with quantitative changes were identified by mass spectrometric analysis. RESULTS: Growth inhibition and morphological abnormalities of cultured embryos were observed in all the Se treatment groups. By the analysis of the embryo proper, actin‐binding proteins were identified as proteins with quantitative changes by selenate: increased phosphorylated‐cofilin 1, increased phosphorylated‐destrin, decreased drebrin E, and decreased myosin light polypeptide 3. Many proteins showed similar changes between selenate and selenite, including increased ATP‐synthase, decreased acidic ribosomal phosphoprotein P0, and decreased pyrroline‐5‐carboxylate reductase‐like. In the yolk sac membrane, antioxidant proteins were identified for protein spots with quantitative changes by selenite: increased peroxiredoxin 1 and increased glutathione S‐transferase. CONCLUSION: The identified proteins with quantitative changes by selenate or selenite were considered to be candidate proteins involved in Se embryotoxicity: the actin‐binding proteins for selenate embryotoxicity, proteins with the similar changes for the common Se embryotoxicity and antioxidant proteins for modification of Se embryotoxicity by redox‐related treatments. These proteins may also be used as biomarkers in developmental toxicity studies. Birth Defects Res (Part B), 2008. © 2008 Wiley‐Liss, Inc.     

Selenium in cereals: improving the efficiency of agronomic biofortification in the UK

Wheat, despite its relatively low selenium (Se) concentration in the UK, is still an important dietary Se source and its biofortification by use of Se fertiliser may be an efficient means to increase the relatively low Se status of the population. We need to know more about the fate of Se applied to the soil and how to ensure the efficiency of Se application, and the three studies reported in this issue of Plant and Soil are timely and informative. Selenium in soil, both globally and locally, is notoriously variable; however, the soils in these studies yielded wheat grain Se concentrations in the narrow range of 16–44 ng/g. The low plant Se levels reported here are not surprising, given that selenite is the dominant Se form in these soils. A regression equation (which used total and extractable Se and extractable S as variables) explained a high proportion of the variance in grain Se concentration. Sulphur application (a common practice on UK wheat growing soils) had variable effects on grain Se concentration, depending on soil S status, pH and possibly other factors. A fertiliser methodology study investigated ways to optimise Se application for the purpose of biofortification. It was calculated that an application of a modest 10 g Se/ha as selenate would increase the grain Se concentration of UK wheat from around 30 ng/g to 300 ng/g. The national Se fertiliser program in Finland shows that this increase would have a large effect on population Se status. However, Se recovery in grain at this application rate is only 14%, and it can be argued that large-scale agronomic biofortification of cereals with Se would be somewhat wasteful of a relatively scarce trace element. Selenium’s effects and interactions in soil, plants, animals and humans are complex and often surprising and will keep researchers busy well into the future.     

Evaluation of Lactobacilli from Human Sources for Uptake and Accumulation of Selenium

The trace mineral selenium (Se) is currently in demand for health supplements for human and animal nutrition. In the present study, 25 isolates of Lactobacillus species of human origin from Indian population were screened for their ability to uptake and accumulate Se in a more bioavailable form. Total Se accumulated by cells was measured by inductively coupled plasma emission spectrometry (ICPES) after wet digestion of Se-enriched cultures. Ten out of 25 isolates grew luxuriantly, as red pigmented colonies, on medium amended with different concentrations of sodium selenite. All the strains when grown in a culture medium supplemented with 10 μg/mL sodium selenite (corresponding to 4.5 μg/mL Se) resulted in Se accumulation in the range of 129.5 to 820.0 μg/g of dry weight (d.w.) as measured using the ICPES method. Elemental Se produced due to reduction of sodium selenite by bacteria was seen as irregular globules under the scanning electron microscope (SEM). Out of all the tested cultures, Lactobacillus reuteri NCDC77 was found to have the greatest ability to uptake Se (28.8 % of inorganic Se in medium) and total Se accumulated was up to 820 μg/g d.w., significantly higher than that of control (742.5 μg/g d.w.). The findings of present study indicate that lactobacilli from human sources have the ability for uptake and accumulation of Se, and the prolific strain has the potential to be explored as an alternative source of organic dietary Se.     

Selenium Improves Physiological Parameters and Alleviates Oxidative Stress in Shoots of Lead-Exposed <Emphasis Type="Italic">Vicia faba</Emphasis> L. <Emphasis Type="Italic">minor</Emphasis> Plants Grown Under Phosphorus-Deficient Conditions

The goal of the study was to investigate the effects of exogenous selenium (Se) on the tolerance of faba bean plants to lead (Pb) stress under P-deficient conditions. The bean plants were grown for 2 weeks on Hoagland solution supplied with Pb (0, 50 μM) and Se (0, 1.5, or 6 μM), separately or simultaneously. It was shown that Pb did not affect shoot growth but caused major damage in the leaves, which was accompanied by Pb accumulation in these tissues. The exposure of the shoots to Pb led to significant changes in the biochemical parameters: the MDA content, glutathione peroxidase (GSH-Px), guaiacol peroxidase (GPOX), and catalase (CAT) activity increased. Furthermore, Pb intensified O ₂ ^?? and H₂O₂ production. Both the Se concentrations used increased the chlorophyll b, chlorophyll a+b, and carotenoid content in the faba bean plants. Selenite also generally enhanced CAT, GPOX, and GSH-Px activities and the T-SH level. Our results imply that the degree of disturbances caused by Pb could be partially ameliorated by Se supplementation. Selenite at a lower dose alleviated Pb toxicity by decreased H₂O₂ and O ₂ ^?? production and decreased the GSH-Px, GPOX, and CAT activities. The beneficial effect of the higher selenite concentration could be related to reduction of lipid peroxidation in the shoots of the Pb-treated plants. However, the effect of Se on the Pb-stressed plants greatly depended on the selenite dose in the nutrient solution.     

Responses of an American eel brain endothelial-like cell line to selenium deprivation and to selenite,selenate, and selenomethionine additions in different exposure media

The effect of selenium deprivation and addition on the American eel brain endothelial cell line (eelB) was studied in three exposure media: complete growth medium (L15/FBS), serum-free medium (L15), and minimal medium (L15/ex). L15/ex contains only galactose and pyruvate and allowed the deprivation of selenium on cells to be studied. In L15/ex, without any obvious source of selenium, eelB cells survived for at least 7 d, formed capillary-like structures (CLS) on Matrigel, and migrated to heal wounds. Three selenium compounds were added to cultures: selenite, selenate, and selenomethionine (SeMet). Adding selenite or selenate to eelB cell cultures for 24 h caused dose-dependent declines in cell viability, regardless of the exposure media. Although varying with exposure media and viability end point, selenite was approximately 70-fold more cytotoxic than selenate. By contrast, 24 h exposures to either dl- or l-SeMet in the three media caused little or no cytotoxicity. However for 7 d exposures in L15/ex, dl- and l-SeMet were very cytotoxic, even at the lowest tested concentration of 31 μM. By contrast in L15 and L15/FBS, cytotoxicity was only observed with 500 and 1000 μM l-SeMet. In L15/FBS, eelB continued to migrate and form CLS in the presence of SeMet but at 500 μM, cell migration appeared stimulated. As judged from a colony-forming assay over 14 d in L15/FBS, 500 and 1000 μM dl- and l-SeMet inhibited cell proliferation. Overall, the responses of eel cells to selenium depended on the selenium form, concentration, and exposure media, with responses to SeMet being most dependent on exposure media.     

Production and detoxification of H2O2 in lettuce plants exposed to selenium

Selenium is considered an essential element for animals. Despite that it has not been demonstrated to be essential for higher plants, it has been attributed with a protective role against reactive oxygen species in plants subjected to stress. In this study, lettuce plants ( Lactuca sativa cv. Philipus) received different application rates (5, 10, 20, 40, 60, 80 and 120 μM) of selenite or selenate, with the aim of testing the effect of Se on the production and detoxification of H₂O₂ in non-stressed plants. The results indicate that the form selenate is less toxic than selenite; that is, the plants tolerated and responded positively to this element, and even increasing in growth up to a rate of 40 μM for the form selenate. On the contrary, the application of selenite triggered a higher foliar concentration of H₂O₂ and a higher induction of lipid peroxidation [malondialdehyde content and lipoxygenase activity] in comparison to that observed after the selenate application. Also, the plants treated with selenate induced higher increases in enzymes that detoxify H₂O₂, especially ascorbate peroxidase and glutathione (GSH) peroxidase, as well as an increase in the foliar concentration of antioxidant compounds such as ascorbate and GSH. These data indicate that an application of selenate at low rates can be used to prevent the induction in plants of the antioxidant system, thereby improving stress resistance.     

Accumulation of red elemental selenium nanoparticles and their biological effects in Nicotinia tabacum

The uptake, accumulation and biological effects of red nano-sized elemental selenium (nanoSe) in comparison to selenate were investigated in plant system at the first time. The data clearly indicated that red nanoSe was taken up by tobacco callus cultures and rooted tobacco plantlets. The roots of regenerated plantlets accumulated selenium in very high concentrations, 2,947 ± 99 mg/kg DW, from the medium containing 530 μM nanoSe. The biological effects of nanoSe were different from the selenate ion in plant tissue culture. NanoSe (265–530 μM concentration range) stimulated the organogenesis and the growth of root system significantly (~40 %) while selenate did not show these effects at any concentration moreover inhibited both callus growth and root regeneration totally in 265–530 μM concentrations.     

Soil factors affecting selenium concentration in wheat grain and the fate and speciation of Se fertilisers applied to soil

UK crops have a low selenium (Se) status, therefore Se fertilisation of wheat (Triticum aestivum L.) at 10 field sites was investigated and the effect on the content and speciation of Se in soils determined. Soil characterisation was carried out at each field site to determine the soil factors that may influence wheat grain Se concentrations in unfertilised plots. Soil samples were taken after harvest from each treatment to determine the fate and speciation of selenate fertiliser applied to soil. Wheat grain Se concentrations could be predicted from soil Se concentration and soil extractable sulphur (S) using the following regression model: Grain Se?=?a?+?b(total soil Se)?+?c(extractable soil Se) - d(extractable soil S), with 86 % of the variance being accounted for, suggesting that these properties control Se concentrations in grain from unfertilised plots. Extractable soil Se concentrations were low (2.4 – 12.4 µg kg^?1) and predominantly consisted of selenite (up to 70 % of extractable Se) and soluble organic forms, whereas selenate was below the detection limit. Little of the added Se, in either liquid or granular form was left in the soil after crop harvest. Se fertilisation up to 20 g ha^?1 did not lead to a significant Se accumulation in the soil, suggesting losses of Se unutilised by the crop.     

Impact of sulphur fertilisation on crop response to selenium fertilisation

UK wheat (Triticum aestivum L.) has a low selenium (Se) concentration and agronomic biofortification with Se is a proposed solution. A possible limitation is that UK wheat is routinely fertilised with sulphur (S), which may affect uptake of Se by the crop. The response of wheat to Se and S fertilisation and residual effects of Se were determined in field trials over 2 consecutive years. Selenium fertilisation at 20 g ha^?1 as sodium selenate increased grain Se by four to seven fold, up to 374 µg Se kg^?1. Sulphur fertilisation produced contrasting effects in 2 years; in year 1 when the crop was not deficient in S, grain Se concentration was significantly enhanced by S, whereas in year 2 when crop yield responded significantly to S fertilisation, grain Se concentration was decreased significantly in the S-fertilised plots. An incubation experiment showed that addition of sulphate enhanced the recovery of selenate added to soils, probably through a suppression of selenate transformation to other unavailable forms in soils. Our results demonstrate complex interactions between S and Se involving both soil and plant physiological processes; S can enhance Se availability in soil but inhibit selenate uptake by plants. Furthermore, no residual effect of Se fertiliser applied in year 1 was found on the following crop.