Exploring the importance of sulfate transporters and ATP sulphurylases for selenium hyperaccumulation\u2014a comparison of Stanleya pinnata and Brassica juncea (Brassicaceae)

Selenium (Se) hyperaccumulation, the capacity of some species to concentrate Se to levels upwards of 0.1% of dry weight, is an intriguing phenomenon that is only partially understood. Questions that remain to be answered are: do hyperaccumulators have one or more Se-specific transporters? How are these regulated by Se and sulfur (S)? In this study, hyperaccumulator Stanleya pinnata was compared with related non-hyperaccumulator Brassica juncea with respect to S-dependent selenate uptake and translocation, as well as for the expression levels of three sulfate/selenate transporters (Sultr) and three ATP sulphurylases (APS). Selenium accumulation went down ~10-fold with increasing sulfate supply in B. juncea, while S. pinnata only had a 2\u20133-fold difference in Se uptake between the highest (5 mM) and lowest sulfate (0 mM) treatments. The Se/S ratio was generally higher in the hyperaccumulator than the non-hyperaccumulator, and while tissue Se/S ratio in B. juncea largely reflected the ratio in the growth medium, S. pinnata enriched itself up to 5-fold with Se relative to S. The transcript levels of Sultr1;2 and 2;1 and APS1, 2, and 4 were generally much higher in S. pinnata than B. juncea, and the species showed differential transcript responses to S and Se supply. These results indicate that S. pinnata has at least one transporter with significant selenate specificity over sulfate. Also, the hyperaccumulator has elevated expression levels of several sulfate/selenate transporters and APS enzymes, which likely contribute to the Se hyperaccumulation and hypertolerance phenotype

Effects of selenium biofortification on crop nutritional quality

Selenium (Se) at very low doses has crucial functions in humans and animals. Since plants represent the main dietary source of this element, Se-containing crops may be used as a means to deliver Se to consumers (biofortification). Several strategies have been exploited to increase plant Se content. Selenium assimilation in plants affects both sulphur (S) and nitrogen (N) metabolic pathways, which is why recent research has also focused on the effect of Se fertilization on the production of S- and N- secondary metabolites with putative health benefits. In this review we discuss the function of Se in plant and human nutrition and the progress in the genetic engineering of Se metabolism to increase the levels and bioavailability of this element in food crops. Particular attention is paid to Se biofortification and the synthesis of compounds with beneficial effects on health

Selenium Biofortification in Radish Enhances Nutritional Quality via Accumulation of Methyl-Selenocysteine and Promotion of Transcripts and Metabolites Related to Glucosinolates, Phenolics, and Amino Acids

Two selenium (Se) fertilization methods were tested for their effects on levels of anticarcinogenic selenocompounds in radish (Raphanus sativus), as well as other nutraceuticals. First, radish was grown on soil and foliar selenate applied 7d before harvest at 0, 5, 10 and 20 mg Se per plant. Selenium levels were up to 1,200 mg Se/kg DW in leaves and 120 mg Se/kg DW in roots. The thiols cysteine and glutathione were present at 2-3 fold higher levels in roots of Se treated plants, and total glucosinolate levels were 35% higher, due to increases in glucoraphanin. The only seleno-aminoacid detected in Se treated plants was Se-methyl-SeCys (100 mg/kg FW in leaves, 33 mg/kg FW in roots). The levels of phenolic aminoacids increased with selenate treatment, as did root total nitrogen and protein content, while the level of several polyphenols decreased. Second, radish was grown in hydroponics and supplied with 0, 5, 10, 20, or 40 \uf06dM selenate for one week. Selenate treatment led to a 20-30% increase in biomass. Selenium concentration was 242 mg Se/kg DW in leaves and 85 mg Se/kg DW in roots. Cysteine levels decreased with Se in leaves but increased in roots; glutatione levels decreased in both. Total glucosinolate levels in leaves decreased with Se treatment due to repression of genes involved in glucosinolates metabolism. Se-methyl-SeCys concentration ranged from 7-15 mg/kg FW. Aminoacid concentration increased with Se treatment in leaves but decreased in roots. Roots of Se treated plants contained elevated transcript levels of sulfate transporters (Sultr) and ATP sulfurylase, a key enzyme of S/Se assimilation. No effects on polyphenols were observed. In conclusion, Se biofortification of radish roots may be achieved via foliar spray or hydroponic supply. One to ten radishes could fulfill the daily human requirement (70 \uf06dg) after a single foliar spray of 5 mg selenate per plant or one week of 5-10 \uf06dM selenate supply in hydroponics. The radishes metabolized selenate to the anticarcinogenic compound Se-methyl-selenocysteine. Selenate treatment enhanced levels of other nutraceuticals in radish roots, including glucoraphanin. Therefore, Se biofortification can produce plants with superior health benefit

Selenium Hyperaccumulators Facilitate Selenium-Tolerant Neighbors via Phytoenrichment and Reduced Herbivory

SummaryBackgroundSoil surrounding selenium (Se) hyperaccumulator plants was shown earlier to be enriched in Se, impairing the growth of Se-sensitive plant species. Because Se levels in neighbors of hyperaccumulators were higher and Se has been shown to protect plants from herbivory, we investigate here the potential facilitating effect of Se hyperaccumulators on Se-tolerant neighboring species in the field.ResultsWe measured growth and herbivory of Artemisia ludoviciana and Symphyotrichum ericoides as a function of their Se concentration and proximity to hyperaccumulators Astragalus bisulcatus and Stanleya pinnata. When growing next to hyperaccumulators, A. ludoviciana and S. ericoides contained 10- to 20-fold higher Se levels (800–2,000 mg kg−1 DW) than when growing next to nonaccumulators. The roots of both species were predominantly (70%–90%) directed toward hyperaccumulator neighbors, not toward other neighbors. Moreover, neighbors of hyperaccumulators were 2-fold bigger, showed 2-fold less herbivory damage, and harbored 3- to 4-fold fewer arthropods. When used in laboratory choice and nonchoice grasshopper herbivory experiments, Se-rich neighbors of hyperaccumulators experienced less herbivory and caused higher grasshopper Se accumulation (10-fold) and mortality (4-fold).ConclusionsEnhanced soil Se levels around hyperaccumulators can facilitate growth of Se-tolerant plant species through reduced herbivory and enhanced growth. This study is the first to show facilitation via enrichment with a nonessential element. It is interesting that Se enrichment of hyperaccumulator neighbors may affect competition in two ways, by reducing growth of Se-sensitive neighbors while facilitating Se-tolerant neighbors. Via these competitive and facilitating effects, Se hyperaccumulators may affect plant community composition and, consequently, higher trophic levels

Selenium biofortification differentially affects sulfur metabolism and accumulation of phytochemicals in two rocket species (Eruca sativa mill. and diplotaxis tenuifolia) grown in hydroponics

Biofortification can be exploited to enrich plants in selenium (Se), an essential micronutrient for humans. Selenium as selenate was supplied to two rocket species, Eruca sativa Mill. (salad rocket) and Diplotaxis tenuifolia (wild rocket), at 0-40 \u3bcM in hydroponics and its effects on the content and profile of sulphur (S)-compounds and other phytochemicals was evaluated. D. tenuifolia accumulated more total Se and selenocysteine than E. sativa, concentrating up to ~300 mg Se kg 121 dry weight from 10-40 \u3bcM Se. To ensure a safe and adequate Se intake, 30 and 4 g fresh leaf material from E. sativa grown with 5 and 10-20 \u3bcM Se, respectively or 4 g from D. tenuifolia supplied with 5 \u3bcM Se was estimated to be optimal for consumption. Selenium supplementation at or above 10 \u3bcM differentially affected S metabolism in the two species in terms of the transcription of genes involved in S assimilation and S-compound accumulation. Also, amino acid content decreased with Se inE. sativa but increased in D. tenuifolia and the amount of phenolics was more reduced in D. tenuifolia. In conclusion, selenate application in hydroponics allowed Se enrichment of rocket. Furthermore, Se at low concentration (5 \u3bcM) did not significantly affect accumulation of phytochemicals and plant defence S-metabolites

Influence of sulfate supply on selenium uptake dynamics and expression of sulfate/selenate transporters in selenium hyperaccumulator and nonhyperaccumulator Brassicaceae

Summary Stanleya pinnata not only hyperaccumulates selenium (Se) to 0.5% of its dry weight, but also exhibits higher tissue Se‐to‐sulfur (S) ratios than other species and its surroundings. To investigate the mechanisms underlying this Se enrichment, we compared S. pinnata with the nonhyperaccumulators S. elata and Brassica juncea for selenate uptake in long‐ (9 d) and short‐term (1 h) assays, using different concentrations of selenate and competitor sulfate. Different sulfate pre‐treatments (0, 0.5, 5 mM, 3 d) were also tested for effects on selenate uptake and sulfate transporters' expression. Relative to nonhyperaccumulators, S. pinnata showed higher rates of root and shoot Se accumulation and less competitive inhibition by sulfate or by high‐S pretreatment. The selenate uptake rate for S. pinnata (1 h) was three‐ to four‐fold higher than for nonhyperaccumulators, and not significantly affected by 100‐fold excess sulfate, which reduced selenate uptake by 100% in S. elata and 40% in B. juncea. Real‐time reverse transcription PCR indicated constitutive upregulation in S. pinnata of sulfate transporters SULTR1;2 (root influx) and SULTR2;1 (translocation), but reduced SULTR1;1 expression (root influx). In S. pinnata, selenate uptake and translocation rates are constitutively elevated and relatively sulfate‐independent. Underlying mechanisms likely include overexpression of SULTR1;2 and SULTR2;1, which may additionally have evolved enhanced specificity for selenate over sulfate