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.     

正交试验分析不同作物的产量、含硒量与土壤酸碱度、硒酸盐、亚硒酸盐含量的相关性

采用正交试验研究了碎米荠、韭菜、大豆、马铃薯的产量、含硒量与土壤酸碱度、硒酸盐、亚硒酸盐含量的关系。结果表明:影响作物含硒量最大的因素是作物品种,不同作物间差异极显著。在土壤中施用硒酸钠和亚硒酸钠均能够提高作物含硒量,用量均以1.0 mg/kg为宜;硒酸钠会使作物产量降低,用量过高使作物硒吸收总量下降;土壤pH值增加有利于植物对硒的吸收,但综合考虑作物产量、含硒量和硒摄入总量,土壤适合的pH值应在6.7～7.9之间。     

Effects of selenite and selenate on the antioxidant systems in Senecio scandens L

Abstract

Selenate and selenite are the most prevalent bioavailable selenium (Se) forms and most easily taken up by plants. Some studies indicate that they are differently absorbed and accumulated in plants and that selenium is toxic if accumulated at high concentrations. Toxicity is due to substitution of sulphur by selenium in cysteine and methionine aminoacids with alteration of the tertiary structure and catalytic activity of proteins and with inhibition of enzymes involved in chlorophyll biosynthesis. Moreover, the interaction between Se and thiol groups induces loss of efficiency of plant defence systems and increases the reactive oxygen species (ROS) production thus enhancing the oxidative stress. To further elucidate the role of Se in higher plants, in this study the antioxidative response to the phytotoxicity of selenite and selenate in Senecio scandens L. was evaluated. The data indicate that while selenite induces oxidative stress enhancing ROS production, lipid peroxidation and the oxidised forms of ascorbate and glutathione, selenate does not significantly affect the analysed pathways. This article outlines that the synergistic action of different antioxidant components is necessary to overcome the phytotoxicity of selenium in Senecio.     

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.     

Accumulation of an organic anticancer selenium compound in a transgenic Solanaceous species shows wider applicability of the selenocysteine methyltransferase transgene from selenium hyperaccumulators

Tolerance to high selenium (Se) soils in Se-hyperaccumulating plant species is correlated with the ability to biosynthesise methylselenocysteine (MeSeCys), due to the activity of selenocysteine methyltransferase (SMT). In mammals, inclusion of MeSeCys in the diet reduces the incidence of certain cancers, so increasing the range of crop plants that can produce this compound is an attractive biotechnology target. However, in the non-Se accumulator Arabidopsis, overexpression of SMT does not result in biosynthesis of MeSeCys from selenate because the rate at which selenate is reduced to selenite by ATP sulfurylase (ATPS) is low. This limitation is less problematic in other species of the Brassicaceae that can produce MeSeCys naturally. We investigated the potential for biosynthesis of MeSeCys in other plant families using Nicotiana tabacum L., a member of the Solanaceae. When plants were watered with 200 μM selenate, overexpression of a SMT transgene caused a 2- to 4-fold increase in Se accumulation (resulting in increased numbers of leaf lesions and areas of necrosis), production of MeSeCys (up to 20% of total Se) and generation of volatile dimethyl diselenide derived directly from MeSeCys. Despite the greatly increased accumulation of total Se, this did not result in increased Se toxicity effects on growth. Overexpression of ATPS did not increase Se accumulation from selenate. Accordingly, lines overexpressing both ATPS and SMT did not show a further increase in total Se accumulation or in leaf toxicity symptoms relative to overexpression of SMT alone, but directed a greater proportion of Se into MeSeCys. This work demonstrates that the production of the cancer-preventing compound MeSeCys in plants outside the Brassicaceae is possible. We conclude that while the SMT gene from Se hyperaccumulators can probably be utilised universally to increase the metabolism of Se into MeSeCys, the effects of enhancing ATPS activity will vary depending on the species involved.     

植物硒素营养及其机理研究进展

土壤生境中Se的丰缺对植物的Se素营养有重要的影响。本文阐述了植物对Se的吸收,积累和运移过程,概述了Se对植物的毒性,对植物产量和品质影响及其机理研究的进展,在此基础上,提出以研究Se对土壤酶活性影响来解释Se对植物的必要性的新途径。     

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.     

Interactions of selenium hyperaccumulators and nonaccumulators during cocultivation on seleniferous or nonseleniferous soil--the importance of having good neighbors

? This study investigated how selenium (Se) affects relationships between Se hyperaccumulator and nonaccumulator species, particularly how plants influence their neighbors' Se accumulation and growth. ? Hyperaccumulators Astragalus bisulcatus and Stanleya pinnata and nonaccumulators Astragalus?drummondii and Stanleya?elata were cocultivated on seleniferous or nonseleniferous soil, or on gravel supplied with different selenate concentrations. The plants were analyzed for growth, Se accumulation and Se speciation. Also, root exudates were analyzed for Se concentration. ? The hyperaccumulators showed 2.5-fold better growth on seleniferous than on nonseleniferous soil, and up to fourfold better growth with increasing Se supply; the nonaccumulators showed the opposite results. Both hyperaccumulators and nonaccumulators could affect growth (up to threefold) and Se accumulation (up to sixfold) of neighboring plants. Nonaccumulators S.?elata and A.?drummondii accumulated predominantly (88-95%) organic C-Se-C; the remainder was selenate. S.?elata accumulated relatively more C-Se-C and less selenate when growing adjacent to S.?pinnata. Both hyperaccumulators released selenocompounds from their roots. A.?bisulcatus exudate contained predominantly C-Se-C compounds; no speciation data could be obtained for S.?pinnata. ? Thus, plants can affect Se accumulation in neighbors, and soil Se affects competition and facilitation between plants. This helps to explain why hyperaccumulators are found predominantly on seleniferous soils.     

X-ray absorption spectroscopy study shows that the rapid selenium volatilizer, pickleweed (Salicornia bigelovii Torr.), reduces selenate to organic forms without the aid of microbes

In many plant species, selenium (Se) volatilization is limited by the reduction of selenate and its chemical conversion to organic Se compounds, a process that may be facilitated by rhizosphere microbes. This study was conducted to determine if pickleweed (Salicornia bigelovii Torr.), which is characterized by having high rates of Se volatilization from selenate, is able to reduce selenate into organic forms of Se axenically, or whether it requires the presence of microbes. X-ray absorption spectroscopy analysis showed that shoots and roots of pickleweed plants supplied with 50 microM selenate accumulated Se predominantly in organic Se forms (about 65-75% of the total accumulated Se), regardless of whether the plants were grown axenically or in the presence of microbes. The results suggest that, unlike other species for which selenate reduction appears to be rate limiting. e.g. Indian mustard (Brassica juncea L.) and broccoli (Brassica oleracea L.), pickleweed is unusual in that it has an enhanced capacity to reduce selenate to organic forms that is independent of the presence of rhizosphere microbes.     

Plant availability of soil selenate additions and selenium distribution within wheat and ryegrass

Selenate fertilization is an effective way to secure selenium (Se) nutrition in Se-poor areas but the cycling of the added selenate in the soil-plant system requires further clarification. We examined the Se uptake efficiency of wheat and ryegrass and Se distribution within these plants in two pot experiments. The behaviour of added selenate in a sand soil under wheat was monitored by sequential extractions during a ten-week growing period. In addition, the relationship between Se uptake of ryegrass and the salt extractable and ligand exchangeable Se in a sand and silty clay soil were studied. The added selenate remained mainly salt soluble in the soil throughout the monitoring. Se uptake by wheat comprised 12% of the soluble Se pool in soil and extended over the whole period of growth. In wheat, over 50% of Se accumulated in grains. The Se uptake of ryegrass comprised, on average, 40% of the soil salt soluble Se. In ryegrass, over 80% of the Se accumulated in roots. The distribution pattern of Se in plants can clearly have a major influence on both the Se cycle in soil and the nutritional efficiency of Se fertilization. The simple salt extraction showed fertilization-induced changes in the soluble soil Se pool, whereas the ligand exchangeable Se fraction reflected the difference in the nonlabile Se status between the two soils.     

Microbial-enhanced Selenium and Iron Biofortification of Wheat (Triticum aestivum L.) - Applications in Phytoremediation and Biofortification

Selenium (Se) is an essential trace element for humans and other mammals. Most dietary Se is derived from crops. To develop a Se biofortification strategy for wheat, the effect of selenate fertilization and bacterial inoculation on Se uptake and plant growth was investigated. YAM2, a bacterium with 99% similarity to Bacillus pichinotyi, showed many plant growth promoting characteristics. Inoculation with YAM2 enhanced wheat growth, both in the presence and absence of selenate: YAM2-inoculated plants showed significantly higher dry weight, shoot length and spike length compared to un-inoculated plants. Selenate also stimulated wheat growth; Un-inoculated Se-treated plants showed a significantly higher dry weight and shoot length compared to control plants without Se. Bacterial inoculation significantly enhanced Se concentration in wheat kernels (167%) and stems (252%), as well as iron (Fe) levels in kernels (70%) and stems (147%), compared to un-inoculated plants. Inoculated Se-treated plants showed a significant increase in acid phosphatase activity, which may have contributed to the enhanced growth. In conclusion; Inoculation with Bacillus sp. YAM2 is a promising Se biofortification strategy for wheat and potentially other crops.     

Application of X-ray absorption near edge spectroscopy to the study of the effect of sulphur on selenium uptake and assimilation in wheat seedlings

Selenium (Se) is an essential trace element for humans and animals. A hydroponic experiment was performed to study the effects of sulphur (S) on Se uptake, translocation, and assimilation in wheat (Triticum aestivum L.) seedlings. Sulphur starvation had a positive effect on selenate uptake and the form of Se supplied greatly influenced Se speciation in plants. Compared with the control plants, Se uptake by the S-starved plants was enhanced by 4.81-fold in the selenate treatment, and selenate was readily transported from roots to shoots. By contrast, S starvation had no significant effect on selenite uptake, and selenite taken up by roots was rapidly converted to organic forms and tended to accumulate in roots. X-ray absorption near edge spectroscopy (XANES) analysis showed that organic forms of selenium, including selenocystine, Se-methyl-selenocysteine (MeSeCys), and selenomethionine-Se-oxide, were dominant in the plants exposed to selenite and accounted for approximately 90 % of the total Se. Whereas selenate remained as the dominant species in the roots and shoots exposed to selenate, with little selenate converted to selenite and MeSeCys. Besides, sulphur starvation increased the proportion of inorganic Se species in the selenate-supplied plants, but had no significant effects on Se speciation in plants exposed to selenite. The present study provides important knowledge to understand the associated mechanism of Se uptake and metabolism in plants.     

Microbial transformations of selenite by methane-oxidizing bacteria

Methane-oxidizing bacteria are well known for their role in the global methane cycle and their potential for microbial transformation of wide range of hydrocarbon and chlorinated hydrocarbon pollution. Recently, it has also emerged that methane-oxidizing bacteria interact with inorganic pollutants in the environment. Here, we report what we believe to be the first study of the interaction of pure strains of methane-oxidizing bacteria with selenite. Results indicate that the commonly used laboratory model strains of methane-oxidizing bacteria, Methylococcus capsulatus (Bath) and Methylosinus trichosporium OB3b, are both able to reduce the toxic selenite (SeO₃ ^2?) but not selenate (SeO₄ ^2?) to red spherical nanoparticulate elemental selenium (Se⁰), which was characterized via energy-dispersive X-ray spectroscopy (EDXS), X-ray absorption near-edge structure (XANES) and extended X-ray absorption fine structure (EXAFS). The cultures also produced volatile selenium-containing species, which suggests that both strains may have an additional activity that can transform either Se⁰ or selenite into volatile methylated forms of selenium. Transmission electron microscopy (TEM) measurements and experiments with the cell fractions cytoplasm, cell wall and cell membrane show that the nanoparticles are formed mainly on the cell wall. Collectively, these results are promising for the use of methane-oxidizing bacteria for bioremediation or suggest possible uses in the production of selenium nanoparticles for biotechnology.

     

Uptake of Selenium and its Antioxidant Activity in Ryegrass When Applied as Selenate and Selenite Forms

Selenium (Se) is an essential micronutrient for animal and human nutrition, but whether it is essential to plants remains controversial. However, there are increasing experimental evidences that indicate a protective role of Se against the oxidative stress in higher plants through Se-dependent glutathione peroxidase (GSH-Px) activity. The effects of the Se chemical forms, selenite and selenate, the rate of their application on shoot Se concentration and their influence on the antioxidative system of ryegrass (Lolium perenne cv. Aries), through the measurement of GSH-Px activity and lipid peroxidation, were evaluated in an Andisol of Southern Chile. Moreover, a soil–plant relationship for Se was determined and a simple method to extract available Se from acid soils is proposed. In a 55-day experiment ryegrass seeds were sown in pots and soil was treated with sodium selenite or sodium selenate (0–10 mg Se kg⁻¹). The results showed that the Se concentration in shoots increased with the application of both selenite and selenate. However, the highest shoot Se concentrations were obtained in selenate-treated plants. For both sources of Se, there was a significant positive correlation between the shoot Se concentration and the GSH-Px activity; and the Se-dependence of this enzymatic activity was related especially with the chemical form of applied Se rather than the Se concentration in plant tissues. Furthermore, the lipid peroxidation, as measured by Thiobarbituric Acid Reactive Substances (TBARS), decreased at low levels of shoot Se concentration, reaching the lowest level at approximately 20 mg Se kg⁻¹ in plants and then increased steadily above this level. In addition, the acid extraction method used to evaluate available Se in soil showed a positive good correlation between soil Se and shoot Se concentrations irrespective of chemical form of Se applied.     

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.     

Influence of arbuscular mycorrhizal fungi on growth and selenium uptake by garlic plants

The aim of this study is to investigate the effects of arbuscular mycorrhizal fungi (AMF) on garlic plants growth and the uptake of selenium (Se). Garlic plants were grown in the pots inoculated with Glomus fasciculatum and G. mosseae and maintained in a greenhouse. Three weeks after planting, the pots had received different concentrations of Se (5, 10, 15, 20, 25 mg kg^?1 of soil) in the form of selenium dioxide (SeO₂) at 3 weeks intervals up to 12 weeks. For physiological and biochemical analysis, the samples were randomly collected from five plants of each experiment. Maximum AM infection, spore population and plant biomass were observed in the roots of mycorrhizal-mediated plants without Se, and they were gradually declined in both mycorrhizal and non-mycorrhizal (NM) plants with increasing concentrations of Se. Among the two Glomus species tested, G. fasciculatum-mediated plants showed higher AM infection, spore population and plant biomass than G. mosseae. No differences were observed for the uptake of Se in mycorrhizal plants and NM plants. However, NM plants uptake more Se than mycorrhizal plants. Higher contents of total chlorophyll and sugars were observed in plants inoculated with G. fasciculatum without Se and they were decreased in the presence of Se. In contrast, increased amount of glutathione peroxidase was observed at increasing concentrations of Se up to 20 mg kg^?1. High-performance liquid chromatography data revealed that SeO₂ converted to organic form of Se as γ-glutamyl-Se-methylselenocysteine. These results are basis for further investigations on the role of AMF on plant growth and uptake of Se in crop plants.     

Effects of different water management on absorption and accumulation of selenium in rice

Rice is the staple food for more than half of the world's population, but selenium (Se) is low in many rice growing countries. Water management model affects rice soil pH and Eh, and then affects the bioavailability of Se in soil. A pot experiment was conducted to investigate the effects of water management on soil Se species, dynamics and selenium uptake by rice plants. Sodium selenite was added to the soil so that the soil selenium content reached 0.5 mg kg^?1 to study the effects of 3 different water management modes on soil selenium uptake by rice plants. These three modes are flood irrigation (F), aerobic irrigation (A) and alternate flood and aerobic irrigation (AFA). The results showed that flooded irrigation treatment increased the soil soluble selenium concentration, and the selenium in soil solution mainly existed in the form of selenite and selenomethionine selenium oxide. The content of selenium in grain was 2.44 and 1.84 times that of flooded irrigation treatment under A and AFA respectively. The content of selenium in straw was 1.32 and 1.58 times that of flooded treatment under A and AFA respectively. After rice grain enzyme hydrolysis, HPLC-ICP-MS analysis showed that Selenomethionine was the main selenium speciation in rice grains. This study showed that aerobic flooded treatment is one of the most effective ways to increase selenium content in rice field.     

Uptake and transport of selenite and selenate by soybean seedlings of two genotypes

In an attempt to address the role of biological behavior on Se uptake by soybean crop and the genotype effects, experiments with time and concentration sequences of Se uptake by seedlings in Hoagland solution are conducted using selenite and selenate respectively. Two soybean cultivars Tong-ai 405 (TA) and Qidong Green-skin (QG) are used as different genotypes. In presence of selenite, Se uptake by both roots and shoots exhibited a linear increase with the growing time at 5 M and with the solution Se concentrations. However, in presence of selenate, the linear response to growing time is only valid before 24 h of growing. While root Se uptake is much slower under selenate than under selenite in the time sequence experiment, shoot Se levels are similar between the two different Se form treatments. Nevertheless, in the experiment of concentration sequence, either root Se or shoot Se responses linearly to solution Se concentration regardless of the Se forms supplied. A big discrepancy of root Se level with a similarity of shoot Se between the two cultivars is observed in the concentration sequence experiment. This supports a much faster passive uptake of selenite but more or less an active uptake of selenate by soybean seedlings. Comparatively, cultivars TA have a consistently higher Se concentration than QG both in roots and shoots under selenate, while no difference of concentration ratio of shoot to root is recognized between them. The higher Se level in seed grains, therefore, may be accounted for not by Se transport form root to shoot but by greater ability of Se uptake and retention under selenate by the former cultivar. Therefore, not only forms of Se supply but also genotype difference affects the Se bioavailability by different soybean cultivars. This should be taken into account for screening the high Se-efficiency plants or cultivars to improve the Se supply of the food chain.     

Overexpression of ATP Sulfurylase in Indian Mustard Leads to Increased Selenate Uptake, Reduction, and Tolerance

In earlier studies, the assimilation of selenate by plants appeared to be limited by its reduction, a step that is thought to be mediated by ATP sulfurylase. Here, the Arabidopsis APS1 gene, encoding a plastidic ATP sulfurylase, was constitutively overexpressed in Indian mustard (Brassica juncea). Compared with that in untransformed plants, the ATP sulfurylase activity was 2- to 2.5-fold higher in shoots and roots of transgenic seedlings, and 1.5- to 2-fold higher in shoots but not roots of selenate-supplied mature ATP-sulfurylase-overexpressing (APS) plants. The APS plants showed increased selenate reduction: x-ray absorption spectroscopy showed that root and shoot tissues of mature APS plants contained mostly organic Se (possibly selenomethionine), whereas wild-type plants accumulated selenate. The APS plants were not able to reduce selenate when shoots were removed immediately before selenate was supplied. In addition, Se accumulation in APS plants was 2- to 3-fold higher in shoots and 1.5-fold higher in roots compared with wild-type plants, and Se tolerance was higher in both seedlings and mature APS plants. These studies show that ATP sulfurylase not only mediates selenate reduction in plants, but is also rate limiting for selenate uptake and assimilation.