Interactions between selenium and sulphur nutrition in Arabidopsis thaliana

Selenium (Se) is an essential plant micronutrient, but is toxic at high tissue concentrations. It is chemically similar to sulphur (S), an essential plant macronutrient. The interactions between Se and S nutrition were investigated in the model plant Arabidopsis thaliana (L.) Heynh. Arabidopsis plants were grown on agar containing a complete mineral complement and various concentrations of selenate and sulphate. The Se/S concentration ratio in the shoot ([Se](shoot)/[S](shoot)) showed a complex dependence on the ratio of selenate to sulphate concentration in the agar ([Se](agar)/[S](agar)). Increasing [S](agar) increased shoot fresh weight (FW) and [S](shoot), but decreased [Se](shoot). Increasing [Se](agar) increased both [Se](shoot) and [S](shoot), but reduced shoot FW. The reduction in shoot FW in the presence of Se was linearly related to the shoot Se/S concentration ratio. These data suggest (i) that Se and S enter Arabidopsis through multiple transport pathways with contrasting sulphate/selenate selectivities, whose activities vary between plants of contrasting nutritional status, (ii) that rhizosphere sulphate inhibits selenate uptake, (iii) that rhizosphere selenate promotes sulphate uptake, possibly by preventing the reduction in the abundance and/or activity of sulphate transporters by sulphate and/or its metabolites, and (iv) that Se toxicity occurs because Se and S compete for a biochemical process, such as assimilation into amino acids of essential proteins.     

Early effects of phenolic compounds, extracted from two forest litters, on ammonium uptake and assimilation in Pinus laricio and Pinus pinaster seedlings

The early effects of low molecular weight phenolic compounds, released by Pinus laricio and Pinus pinaster litter, on ammonium uptake and its assimilation in two Pinus species were studied. In Pinus laricio seedlings, the exposure to phenols extracted from Pinus laricio litter increased not only the ammonium uptake but also the activity of the main enzymes involved in its assimilation, whereas the phenols extracted from Pinus pinaster litter had a negative effect on these metabolic processes. In Pinus pinaster seedlings, the exposure to both phenols decreased the ammonium uptake and the activity of the main enzymes involved in its assimilation. Histological analysis carried out in Pinus laricio roots showed that phenols extracted from Pinus laricio litter induced the greatest growth of cortex, element through which occurs the ions uptake in plants, whereas phenol extracted from Pinus pinaster litter inhibited cortex development. On the other hand, in Pinus pinaster seedlings the observation showed that both phenols inhibited cortex growth indicating a strict correlation between cortex development and ammonium uptake and its assimilation.     

Differential effects of AM fungal isolates on Medicago truncatula growth and metal uptake in a multimetallic (Cd, Zn, Pb) contaminated agricultural soil

Toxic metal accumulation in soils of agricultural interest is a serious problem needing more attention, and investigations on soil–plant metal transfer must be pursued to better understand the processes involved in metal uptake. Arbuscular mycorrhizal (AM) fungi are known to influence metal transfer in plants by increasing plant biomass and reducing metal toxicity to plants even if diverging results were reported. The effects of five AM fungi isolated from metal contaminated or non-contaminated soils on metal (Cd, Zn) uptake by plant and transfer to leachates was assessed with Medicago truncatula grown in a multimetallic contaminated agricultural soil. Fungi isolated from metal-contaminated soils were more effective to reduce shoot Cd concentration. Metal uptake capacity differed between AM fungi and depended on the origin of the isolate. Not only fungal tolerance and ability to reduce metal concentrations in plant but also interactions with rhizobacteria affected heavy metal transfer and plant growth. Indeed, thanks to association with nodulating rhizobacteria, one Glomus intraradices inoculum increased particularly plant biomass which allowed exporting twofold more Cd and Zn in shoots as compared to non-mycorrhizal treatment. Cd concentrations in leachates were variable among fungal treatments, but can be significantly influenced by AM inoculation. The differential strategies of AM fungal colonisation in metal stress conditions are also discussed.     

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.     

Mechanisms of selenium hyperaccumulation in plants: A survey of molecular,biochemical and ecological cues

Background

Selenium (Se) is a micronutrient required for many life forms, but toxic at higher concentration. Plants do not have a Se requirement, but can benefit from Se via enhanced antioxidant activity. Some plant species can accumulate Se to concentrations above 0.1% of dry weight and seem to possess mechanisms that distinguish Se from its analog sulfur (S). Research on these so-called Se hyperaccumulators aims to identify key genes for this remarkable trait and to understand ecological implications.

Nicotianamine synthase: Gene isolation,gene transfer and application for the manipulation of plant iron assimilation

Basic cellular processes such as electron transport in photosynthesis and respiration require the precise control of iron homeostasis. To mobilise iron, plants have evolved at least two different strategies. The non-proteinogenic amino acid nicotianamine is an essential component of both pathways.We briefly review the characterisation of the nicotianamine synthase as a member of a novel class of enzymes, the cloning of the corresponding gene coding sequences of barley, Arabidopsis and tomato as well as the molecular basis of the chloronerva mutant exhibiting severe defects in the regulation of iron metabolism.Further, we report on current experiments aiming to the application of various NAS-genes to manipulate iron assimilation in model and crop plants using transgenic sense and antisense approaches.     

The accumulation and assimilation of dimethylselenide by four plant species

Plants of Agrostis tenuis Sibth., Hordeum vulgare L., Lycopersicon esculentum Mill. and Raphanus sativus L. were grown hydroponically in sealed systems and fumigated with 8 g m^-3 [⁷⁵Se]-dimethylselenide. The accumulation of ⁷⁵Se was measured and the shoot tissues were extracted to examine the products of the ⁷⁵Se assimilation. Characteristic differences were observed between species in the accumulation of ⁷⁵Se and the transport from shoots to roots. High-voltage electrophoresis and chromatography of extracts made with 80% aqueous ethanol revealed the presence of inorganic selenite as an assimilation product as well as the selenium analogues of glutathione and methionine. Extensive incorporation of ⁷⁵Se into protein-bound selenomethionine was observed in all plant species.Abbreviation DMSe dimethylselenide     

Beryllium in the environment: Whether fatal for plant growth?

Beryllium could be a threatening heavy metal pollutant in the agro-ecosystem that may severely affect the performance of crops. Beryllium is used in various industries to make nuclear weapons and reactors, aircraft and space vehicle structures, instruments, and X-ray machines, and its entry into the environment is alarming for the productivity and sustainability of the ecosystem. In this review, we present a contemporary synthesis of the existing data regarding the toxic effects of beryllium on toxicity on biochemical and physiological processes in plants. Moreover, uptake, translocation, and assimilation of beryllium and its interaction with some essential mineral elements are also discussed. Although limited data are available regarding biochemical responses of plants to beryllium toxicity, we tried to clarify some basic physiological and biochemical steps that can be hampered by beryllium in plants. We linked our hypothetical concepts with previous evidence and provide a comprehensive summary of all possible remediation strategies that can be used for plants. Overall, we hope this review will be beneficial due to its practical implications and research directions.     

Effects of simulated herbivory on photosynthesis and N resorption efficiency in <Emphasis Type="Italic">Quercus</Emphasis><Emphasis Type="Italic">pyrenaica</Emphasis> Willd. saplings

We examined the effects of simulated folivory by caterpillars on photosynthetic parameters and nitrogen (N) resorption efficiency in Quercus pyrenaica saplings. We analyzed the differences between intact leaves in control plants, punched leaves in damaged plants, and intact leaves in damaged plants. We then established two levels of simulated folivory: low (≈13% of the leaf area of one main branch removed per plant) and high (≈26% of the leaf area of one main branch removed per plant) treatments. No differences were found in net assimilation rate and conductance between either leaf type or treatment during the most favourable period for photosynthesis. However, the N content was lower in punched than in intact leaves, and as a result PNUE was higher in damaged leaves from treated trees. In leaf-litter samples, N mass was significantly higher in punched than in intact leaves in treated plants, and LMA was significantly higher in damaged than in intact leaves of both the treated and control plants. Consequently, N resorption efficiency was around 15% lower in damaged leaves as compared with intact leaves from treated and control plants. Mechanical injury to leaves not only triggered no compensatory photosynthetic response to compensate a lower carbon uptake due to leaf area loss, but also affected the resorption process that characterizes leaf senescence.     

Litter N retention over winter for a low and a high phenolic species in the alpine tundra

Mycorrhizal fungus colonization of roots may modify plant metal acquisition and tolerance. In the present study, the contribution of the extraradical mycelium of an arbuscular mycorrhizal (AM) fungus, Glomus mosseae (BEG 107), to the uptake of metal cations (Cu, Zn, Cd and Ni) by cucumber (Cucumis sativus) plants was determined. The influence of the amount of P supplied to the hyphae on the acquisition and partitioning of metal cations in the mycorrhizal plants was also investigated. Pots with three compartments were used to separate root and root-free hyphal growing zones. The shoot concentration of Cd and Ni was decreased in mycorrhizal plants compared to non-mycorrhizal plants. In contrast, shoot Zn and Cu concentrations were increased in mycorrhizal plants. High P supply to hyphae resulted in decreased root Cu concentrations and shoot Cd and Ni concentrations in mycorrhizal plants. These results confirm that some elements required for plant growth (P, Zn, Cu) are taken up by mycorrhizal hyphae and are then transported to the plants. Conversely, Cd and Ni were transported in much smaller amounts by hyphae to the plant, so that arbuscular mycorrhizal fungus colonization could partly protect plants from toxic effects of these elements. Selective uptake and transport of plant essential elements over non-essential elements by AM hyphae, increased growth of mycorrhizal plants, and metal accumulation in the root may all contribute to the successful growth of mycorrhizal plants on metal-rich substrates. These effects are stimulated when hyphae can access sufficient P in soil.     

Long-term effects of SO2 on plants,SO2 metabolism and regulation of intracellular pH

The impact of SO₂ on the ionic balance of plants and its implications for intracellular pH regulation was studied to find explanations for long-term effects of SO₂. When sulphur, taken up as SO₂ by the shoots of plants, is not assimilated in organic compounds, but stored as sulphate, an equivalent amount of H⁺ is produced. These H⁺ ions are not buffered chemically, but removed by metabolic processes.On the basis of knowledge on metabolic buffering mechanisms a conceptual model is proposed for the removal of shoot-generated H⁺ by (i) OH^- ions, produced in the leaves when sulphate and nitrate are assimilated in organic compounds and/or by (ii) OH^- ions produced by decarboxylation of organic anions (a biochemical pH stat mechanism). The form in which nitrogen is supplied largely determines the potential of the plant to neutralize H⁺ in the leaves during SO₂ uptake by the proposed mechanisms.In field experiments with N₂ fixing Vicia faba L. crops, the increase of sulphate in the shoots of SO₂-exposed plants was equivalent in charge to the decrease of organic anion content, calculated as the difference between inorganic cation content (C) and inorganic anion content (A), indicating that H⁺ ions produced in the leaves following SO₂ uptake were partly removed by OH^- from sulphate reduction and partly by decarboxylation of organic anions.The appearance of chronic SO₂ injury (leaf damage) in the field experiment at the end of the growing period is discussed in relation to the impact of SO₂ on the processes involved in regulation of intracellular pH. It is proposed that the metabolic buffering capacity of leaf cells is related to the rates of sulphate and nitrate reduction and the import rate of organic anions, rather than to the organic anion content in the vacuoles of the leaf cells.     

氮磷肥对茶树锌硒等中微量元素吸收与分配的影响

锌（Zn）和硒（Se）及其他中微量元素（铝Al,钙Ca,铁Fe,铜Cu,锰Mn）是茶叶品质的重要指标,但茶树吸收Zn、Se能力及氮（N）磷（P）肥影响中微量元素吸收与分配的过程尚不清楚。以红壤丘陵区福鼎大白茶树为研究对象,开展Zn+Se、Zn+Se+N、Zn+Se+P、Zn+Se+N+P和对照共5种处理3次重复随机化区组试验,处理第3年春季分茶叶、成熟叶、吸收根、运输根和储藏根采集植物样品,测定其元素含量。结果表明,茶树地上和地下器官Zn和Se及其他中微量元素对N、P、Zn、Se添加的响应具分异性。与对照相比,茶树地上和地下器官Zn和Se含量均显著增加,与Zn+Se相比,施N和/或P肥仅显著提高茶叶和成熟叶Se含量（P<0.05）;与对照相比,施肥处理均显著提高吸收根和运输根Al、Fe含量以及储藏根Cu含量;运输根Mn含量表现为Zn+Se+N、Zn+Se+P、Zn+Se+N+P显著高于对照,储藏根Mn含量为Zn+Se+N+P显著高于其他处理;茶树各器官Ca含量对施肥处理无显著响应。此外,茶叶和成熟叶的Zn含量与吸收根显著正相关,而Se含量则与储藏根显著正相关。茶树具有吸收和积累Zn和Se的能力,而施N、P肥有助于提高茶叶Se含量,研究结果为红壤丘陵区培育高品质锌硒茶及营建生态高值茶园提供了依据。     

Subcellular distribution of selenium during uptake and its influence on mitochondrial oxidations in germinatingVigna radiata L

The metabolic significance of Se in plants is not well documented, though the presence of many selenoenzymes in bacteria and the essentiality of Se in higher animals is established. Since germination is an active process in plant growth and metabolism, the effect of Se was investigated in germinatingVigna radiata L, a nonaccumulating Sedeficient legume. Growth and protein were enhanced in seedlings supplemented with selenium (Se) as sodium selenite in the medium up to 1 μg/mL. The pattern of uptake of⁷⁵Se in the differentiating tissues and the subcellular distribution were investigated. The percentage of incorporation of⁷⁵Se was greater in the mitochondria at the lowest level (0.5 μg/mL) of Se supplementation compared to higher levels of Se exposure. Proteins precipitated from the postmitochondrial supernatant fractions, when separated by means of polyacrylamide gel electrophoresis (PAGE), indicated a major selenoprotein in the seedlings germinated at 2.0 μg/mL Se. In seedlings grown with supplemented Se, enhanced respiratory control ratio and succinate dehydrogenase activity were observed in the mitochondria of tissues, indicative of a role for Se in mitochondrial membrane functions.     

Selenium uptake by plants as a function of soil type,organic matter content and pH

In pots containing sandy soils at two levels (pH 5 and 7) to which 0.5 mg Se L^-1 soil had been added, an increase in the proportion of clay soil or peat soil led to a decrease in the uptake of Se by spring wheat grain (Triticum aestivum L., var. Drabant) and winter rape plants (Brassica napus L., var. Emil). The effect was most pronounced for the smallest additions of clay and peat soils. Differences in Se uptake between the two pH levels were greatest in treatments where the additions of clay and peat soils were small. At the high pH, an increase in clay content from 7% to 39% resulted in a decrease in Se uptake of 79% for wheat and 70% for rape. At the low pH, the uptake decreased by 72% and 77%, respectively. At the higher pH, an increase in the content of organic matter from 1.4% to 39% resulted in decreases in Se uptake of 88% for wheat grain and 69% for rape. At the low pH, Se uptake decreased by 63% and 48%, respectively. Adding peat soil to clay soil had little effect on Se uptake. Among the limed, unmixed clay, sand and peat soils to which Se had not been added, uptake was highest from the sandy soil, i.e. 8.3 ng Se/g wheat grain and 42 ng Se/g rape. The lowest uptake rates were obtained in the clay soil, i.e. 3.0 ng Se/g for wheat grain and 9.0 ng Se/g for rape.     

Leptin and metabolic control of reproduction

Leptin treatment prevents the effects of fasting on reproductive processes in a variety of species. The mechanisms that underlie these effects have not been elucidated. Progress in this area of research might be facilitated by viewing reproductive processes in relation to mechanisms that maintain fuel homeostasis. Reproduction, food intake, and fuel partitioning can be viewed as homeostatic responses controlled by a sensory system that monitors metabolic signals. These signals are generated by changes in intracellular metabolic fuel availability and oxidation rather than by changes in the amount of body fat or by changes in any aspect of body composition. Leptin might be viewed as either a mediator or as a modulator of the intracellular metabolic signal. Consistent with its purported action as a mediator of the metabolic signal, leptin synthesis and secretion are influenced acutely by changes in metabolic fuel availability, and these changes might lead to changes in reproductive function. The effects of leptin treatment on reproduction are blocked by treatments that inhibit intracellular fuel oxidation. Metabolic signals that inhibit reproduction in leptin-treated animals might act via neural pathways that are independent of leptin's action. Alternatively, both leptin and metabolic inhibitors might interact at the level of intracellular fuel oxidation. In keeping with the possibility that leptin modulates the metabolic signal, leptin treatment increases fuel availability, uptake, and oxidation in particular tissues. Leptin might affect reproduction indirectly by altering fuel oxidation or other peripheral processes such as gastric emptying. Reproductive processes are among the most energetically expensive in the female repertoire. Because leptin increases energy expenditure while simultaneously inhibiting energy intake, it may have limited use as a long-term treatment for infertility.     

Shading and the capture of localized soil nutrients: nutrient contents,carbohydrates, and root uptake kinetics of a perennial tussock grass

Summary The ability to exploit spatial and temporal heterogeneity in soil resources can be one factor important to the competitive balance of plants. Competition above-ground may limit selective plant responses to below-ground heterogeneity, since mechanisms such as root proliferation and alterations in uptake kinetics are energy-dependent processes. We studied the effect of shading on the ability of the perennial tussock grassAgropyron desertorum to take up nutrients from enriched soil microsites in two consecutive growing seasons. Roots of unshaded plants selectively increased phosphate uptake capacity in enriched soil microsites (mean increases of up to 73%), but shading eliminated this response. There were no changes in ammonium uptake capacity for roots in control and enriched patches for either shaded or unshaded plants. The 9-day shade treatments significantly reduced total nonstructural carbohydrate (TNC) concentrations for roots in 1990, but had no apparent effect on root carbohydrates in 1991 despite dramatic reductions in shoot TNC and fructan concentrations. Enrichment of the soil patches resulted in significantly greater phosphate concentrations in roots of both shaded and unshaded plants, with less dramatic differences for nitrogen and no changes in potassium concentrations. In many respects the shaded plants did surprisingly well, at least in terms of apparent nutrient acquisition. The effects of aboveground competition on nutrient demand, energy requirements, and belowground processes are discussed for plants exploiting soil resource heterogeneity.     

Cadmium uptake and bioaccumulation in selected cultivars of durum wheat and flax as affected by soil type

Accumulation of cadmium (Cd) in crop plants is of great concern due to the potential for food chain contamination through the soil-root interface. Although Cd uptake varies considerably with plant species, the processes which determine the accumulation of Cd in plant tissues are affected by soil factors. The influence of soil type on Cd uptake by durum wheat (Triticum turgidum var. durum L.) and flax (Linum usitatissimum L.) was studied in a pot experiment under environmentally controlled growth chamber conditions. Four cultivars/lines of durum wheat (Kyle, Sceptre, DT 627, and DT 637) and three cultivars/lines of flax (Flanders, AC Emerson, and YSED 2) were grown in two Saskatchewan soils: an Orthic Gray Luvisol (low background Cd concentration; total/ABDTPA extractable Cd: 0.12/0.03 mg kg^-1, respectively) and a Dark Brown Chernozem (relatively high background Cd concentration; total/ABDTPA Cd: 0.34/0.17 mg kg^-1 respectively). Plant roots, stems, newly developed heads, and grain/seeds were analyzed for Cd concentration at three stages of plant growth: two and seven weeks after germination, and at plant maturity. The results showed that Cd bioaccumulation and distribution within the plants were strongly affected by both soil type and plant cultivar/line. The Cd concentration in roots leaves and stems varied at different stages of plant growth. However, all cultivars of both plant species grown in the Chernozemic soil accumulated more Cd in grain/seeds than plants grown in the Orthic Gray Luvisol soil. The different Cd accumulation pattern also corresponded to the levels of ABDTPA extractable and metal-organic complex bound soil Cd found in both soils. Large differences were found in grain Cd among the durum wheat cultivars grown in the same soil type, suggesting the importance of rhizosphere processes in Cd bioaccumulation and/or Cd transport processes within the plant. Distribution of Cd in parts of mature plants showed that durum grain contained up to 21 and 36% of the total amount of Cd taken up by the plants for the Orthic Gray Luvisol and Chernozemic soils, respectively. These results indicate the importance of studying Cd speciation, bioaccumulation and cycling in the environment for the management of agricultural soils and crops.     

Extraordinarily high leaf selenium to sulfur ratios define 'Se-accumulator' plants

BACKGROUND AND AIMS: Selenium (Se) and sulfur (S) exhibit similar chemical properties. In flowering plants (angiosperms) selenate and sulfate are acquired and assimilated by common transport and metabolic pathways. It is hypothesized that most angiosperm species show little or no discrimination in the accumulation of Se and S in leaves when their roots are supplied a mixture of selenate and sulfate, but some, termed Se-accumulator plants, selectively accumulate Se in preference to S under these conditions. METHODS: This paper surveys Se and S accumulation in leaves of 39 angiosperm species, chosen to represent the range of plant Se accumulation phenotypes, grown hydroponically under identical conditions. RESULTS: The data show that, when supplied a mixture of selenate and sulfate: (1) plant species differ in both their leaf Se ([Se](leaf)) and leaf S ([S](leaf)) concentrations; (2) most angiosperms show little discrimination for the accumulation of Se and S in their leaves and, in non-accumulator plants, [Se](leaf) and [S](leaf) are highly correlated; (3) [Se](leaf) in Se-accumulator plants is significantly greater than in other angiosperms, but [S](leaf), although high, is within the range expected for angiosperms in general; and (4) the Se/S quotient in leaves of Se-accumulator plants is significantly higher than in leaves of other angiosperms. CONCLUSION: The traits of extraordinarily high [Se](leaf) and leaf Se/S quotients define the distinct elemental composition of Se-accumulator plants.     

The effect of take-all disease on gas-exchange rates and biomass in two winter wheat lines with different drought response

There have been no studies of the effect of take-all on leaf gas-exchange rates, despite the fact that take-all severely restricts plant water and nutrient uptake, which results in significant biomass and grain yield reduction. Here we describe the effect of inoculation with Gaeumannomyces graminis (Sacc.) var. tritici (Ggt) on carbon assimilation rate (A) and biomass production of wheat plants grown under two water regimes. We show that the impact of Ggt inoculation on plant growth and leaf A may be through reduced photosynthetic capacity of the leaves and not water stress per se. The nature of this reduced photosynthetic capacity remains uncertain but may involve nutrient deficiency and different enzymes produced by the fungus. In each of the 3 years the experiment was conducted, Ggt significantly reduced A, i.e. at anthesis by 18% in 2000, 15% in 2001, and 12% in 2002. In agreement with other field studies, Ggt reduced tiller number and production of all plant components, mostly root dry mass and grain mass per plant. Highly significant negative correlations were found between disease rating and A in all years, showing that at disease ratings equal or higher than 3 (on a scale from 1 to 4) A could practically be zero. While A decreased, intercellular CO₂ concentration increased or did not change, and stomatal conductance was relatively high. In addition, A was more reduced under high than under low soil moisture content. These results support the idea that water stress per se did not contribute to the observed reduction of A. The mechanism of photosynthetic capacity reduction due to the Ggt root-rotting fungus is of interest as it may lead to the molecular mechanisms of plant resistance and ultimately to the development of take-all resistant plants.