Mechanisms underlying mercury detoxification in soil-plant systems after selenium application: a review.

Feasible countermeasures to mitigate mercury (Hg) accumulation and its deleterious effects on crops are urgently needed worldwide. Selenium (Se) fertilizer application is a cost-effective strategy to reduce Hg concentrations, promote agro-environmental sustainability and food safety, and decrease the public health risk posed by Hg-contaminated soils and its accumulation in food crops. This holistic review focuses on the processes and detoxification mechanisms of Hg in whole soil-plant systems after Se application. The reduction of Hg bioavailability in soil, the formation of inert HgSe or/and HgSe-containing proteinaceous complexes in the rhizosphere and/or roots, and the reduction of plant root uptake and translocation of Hg in plant after Se application are systemically discussed. In addition, the positive responses in plant physiological and biochemical processes to Se application under Hg stress are presented to show the possible mechanisms for protecting the plant. However, application of high levels Se showed synergistic toxic effect with Hg and inhibited plant growth. The effectiveness of Se application methods, rates, and species on Hg detoxification is compared. This review provides a good approach for plant production in Hg-contaminated areas to meet food security demands and reduce the public health risk.

Detoxification difference of cadmium between the application of selenate and selenite in native cadmium-contaminated soil.

Cadmium (Cd) has strong mobility and could cause toxicity to plants, and selenium (Se) can effectively detoxify Cd stress. However, differences in the detoxification effects of different species and dosages of exogenous Se on Cd and its mechanism are still unclear. In this study, a pot experiment was conducted to determine the effects of different rates of selenite and selenate application on radish growth, the uptake and translocation of Cd, and the fractions of Cd transformation in native Cd-contaminated soil. Results indicated that the decrease in radish biomass in selenate treatment was significantly greater than that in selenite treatment at a high Se application rate (2.5 mg·kg-1) (p < 0.05). In contrast to selenite treatments, the application of selenate significantly increased the translocation of Cd from radish roots to shoots (p < 0.05). Cadmium concentration and its bioaccumulation factor in radish decreased gradually with increasing selenite application rates, while these values decreased at low Se rate (1 mg·kg-1) and increased at high Se rate for selenate treatment. Different Se application rates resulted in Cd fractions distributions to change in soil. Therefore, the application of selenite treatment had a greater detoxification effect on Cd in soil than that in selenate treatment, and the double toxic effect was observed between Se and Cd in high selenate treatment (2.5 mg·kg-1). Combined with human health risk asseeement, the application of 2.5 mg·kg-1 selenite could be a good approach for detoxification in native Cd-contaminated soil used in this study.

Influence of processing methods and exogenous selenium species on the content and in vitro bioaccessibility of selenium in Pleurotus eryngii.

Understanding the effects of processing on the Se content and bioaccessibility in food is critical in guiding the development of Se-enriched products. In this study, Se-enriched Pleurotus eryngii was obtained by applying different Se supplements to the substrate. Selenium content and its bioaccessibility among raw and processed fruit bodies were compared. The application of exogenous Se had no significant effect on the yield of P. eryngii, while amendment Se yeast could slightly promote the growth of P. eryngii. The enrichment ability of P. eryngii among different Se supplements declined in the order of Na2SeO3 > Se yeast > Na2SeO4. However, the processing treatments resulted in 6.6%-45.9% Se loss. The Se bioaccessibility of P. eryngii was 78.4%-89.7%. Frying treatment reduced Se bioaccessibility in samples, whereas boiling treatment enhanced it. Therefore, Se yeast and boiling treatment are recommended as the ideal Se supplement and processing method for Se-enriched P. eryngii.

Insights into uptake, accumulation, and subcellular distribution of selenium among eight wheat (Triticum aestivum L.) cultivars supplied with selenite and selenate.

Selenium (Se)-enriched wheat can be improved by altering Se sources and selecting wheat cultivars. Such improvement can affect subcellular distribution and speciation of Se in wheat. Thus, a pot experiment was conducted to investigate Se uptake and distribution when Se was applied as selenite or selenate at low and high rates (1 and 10 mg kg-1, respectively). Moreover, Se's impact on the grain and biomass yield of eight wheat cultivars was also investigated. The subcellular distribution and speciation of Se were also explored to elucidate Se metabolism and micro-distribution pattern in wheat. Results showed that biomass and grain yield were decreased with the application of both selenite and selenate in almost all the cultivars, regardless of the Se rate. Application high Se rate resulted in a significant (p < 0.05) decrease in grain yield and biomass compared with low rate of Se. Compared with the low rate of selenite application, the grain and the biomass yield of ZM-9023 significantly (p < 0.05) increased by about 15% for low rate of selenate application. In addition, both selenite and selenate treatment increased the uptake of Se in each part of wheat, compared with the control. Selenium was mostly accumulated in the grain and root of wheat under selenite treatment, while more Se accumulation was found in leaves and straw for selenate application. Further investigation on the subcellular distribution of Se showed that the proportion of Se in soluble fraction was significantly (p < 0.05) higher in wheat leaves than that in organelle fraction and cell walls (46%-66%). Meanwhile, Se6+ was the main species found in soluble fraction, whereas SeMet and MeSeCys were the species predominantly stored in organelle fraction. In conclusion, wheat cultivar ZM-9023 is the most Se-rich potential cultivar, and the isolation of Se in the soluble fraction plays an important role in Se tolerance and accumulation.

Assessing the potential availability of selenium in the soil-plant system with manure application using diffusive gradients in thin-films technique (DGT) and DOM-Se fractions extracted by selective extractions.

Knowledge of the Se fractionation and the role of dissolved organic matter (DOM) in soil is the key to understanding Se mobility and its bioavailability in the soil-plant system. In this study, single extractions using phosphate-buffer (PBS), sequential extraction procedures (SEP), and diffusive gradients in thin-films (DGT) were used to measure Se bioavailability in soil supplemented with selenite and organic amendment (cow and chicken manures). Selenium fraction was isolated into DOM-Se fractions, such as hydrophilic acid-bound Se (HY-Se), fulvic acid-bound Se (FA-Se), humic acid-bound Se (HA-Se), and hydrophobic organic neutral-bound Se (HON-Se), by a rapid batch technique using XAD-8 resin (AMBERLITE XAD™, USA). Simultaneous application of either cow or chicken manure with selenite could result in the decrease of Se availability in the soil. Isolating Se available fraction into DOM-Se fractions showed that low-molecular-weight DOM-Se as an available fraction and even HY-Se as a less available fraction (OM-Se) were likely the major sources for Brassica juncea (L.) Czern. et Coss uptake in soil. Moreover, knowledge of the DOM-Se composition, especially the low-molecular-weight DOM-Se fractions, is important for assessing the bioavailability of Se in soil, the results of which are more accurate than the chemical extraction method. The high value of Pearson correlation coefficients between CDGT-Se and Se concentrations in shoots, tubers and roots of Brassica juncea (L.) Czern. et Coss in cow and chicken manures treatment were 0.95 and 0.99, 0.96 and 0,96, and 0.89 and 0.97 (p < 0,05), respectively, indicating that DGT-Se can reflect the Se uptake ability by plants and can be used to predict the bioavailability of Se when manure and selenite are simultaneously applied.

DOM derivations determine the distribution and bioavailability of DOM-Se in selenate applied soil and mechanisms.

Straw amendment and plant root exudates modify the quality and quantities of soil dissolved organic matter (DOM) and then manipulate the fractions of soil selenium (Se) and its bioavailability. Two typical soils with distinct pH were selected to investigate the effect of different contributors on DOM-Se in soil. The mechanisms relying on the variation in DOM characteristics (quality, quantity and composition) were explored by UV-Vis, ATR-FTIR and 3D-EEM. Straw amendment significantly (p < 0.05) suppressed the selenate bioavailability. The reduction in wheat Se content was greater in krasnozems than in Lou soil, as more HA fraction appeared in krasnozems. The root exudates of wheat mainly elevated the low molecular hydrophilic compounds (Hy) in soil, which contributed to the SOL-Hy-Se fractions and thus grain Se in soils (p < 0.01). However, straw amendment promoted DOM transforming from small molecules (Hy and FA) to aromatic large molecules (HA), when accompanied with the reduction and retention of Se associated with these molecules. As a result, selenium bioavailability and toxicity reduced with DOM amendment and DOM-Se transformation.

Understanding boosting selenium accumulation in Wheat (Triticum aestivum L.) following foliar selenium application at different stages, forms, and doses.

There are a lack of systematic studies comparing the effects of foliar-applied selenium (Se) with different Se sources at different growth stages in wheat. Herein, we biofortified wheat via the foliar application of selenite and selenate at different rates and different stages under field conditions. Results showed that foliar-applied selenate and selenite had no significant effect either on wheat biomass or grain yield (p < 0.05). Selenium distribution in different parts of wheat plant ranked decrease as leaf > root > grain > glume > stem with selenite treatment, and it appeared in the decline order as leaf > grain > glume > stem > root with selenate treatment. These results suggested that biofortification with selenate caused, relatively to selenite, a higher accumulation of Se in grains. Foliar application of Se of either selenate or selenite at pre-filling stage was superior in improving the Se concentration of wheat grains than application at pre-flowering stage. Meanwhile, organic Se comprised about 72-93% of total Se in wheat grains, which was reduced by 5.8% at high Se rate (100 g ha-1), irrespective of the forms of Se or stages applied. The organic Se proportion in wheat grains was 9% higher with the selenate treatment than with the selenite treatment. Selenomethionine (SeMet) was the main organic species (67-86%) in wheat grains, followed by selenocysteine (SeCys2). In summary, our results indicate that Se biofortification of wheat is most effective with 20 g ha-1 selenate foliar-applied at pre-filling stage.

Effects of selenium combined with zinc amendment on zinc fractions and bioavailability in calcareous soil.

Selenium (Se) and zinc (Zn) are two important trace elements for human being and animals. The interaction between Se and Zn on the bioavailability of Zn in soil is still unclear. Therefore, pot experiments exposed to different dosages of zinc sulfate (ZnSO4) (0, 20, and 50 mg/kg soil) and sodium selenite (Na2SeO3) (0, 0.5, 1.0, and 2.5 mg/kg soil) were conducted to investigate the effects of selenite application on Zn bioavailability in calcareous soil and its related mechanisms. The total Zn content of different tissues (roots and shoots) of pak choi (Brassica chinensis L.) and the changes in Zn fraction distribution in soil before planting and after harvest were determined, and the mobility factor (MF) and distribution index (DI) of Zn in soils were calculated. In addition, the Pearson correlation and path analysis were conducted to clarify the relationships between Zn fractions in soil and the Zn uptake of pak choi. Results showed that Se amendment elevated soil Zn bioavailability at appropriate levels of Se and Zn. When 1.0 and 2.5 mg/kg of Se and 20 mg/kg of Zn were applied in soil, the proportion of exchangeable Zn (Ex-Zn) and Zn weakly bound to organic matter (Wbo-Zn) to the total content of Zn was significantly increased by 28.14%-82.52% compared with that of the corresponding single Zn treatment. Therefore, the Zn concentration in the shoots of pak choi was significantly increased by 27.2%-31.1%. High Zn (50 mg/kg) and Se co-amended treatments showed no significantly beneficial effect on the bioavailability of Zn. In addition, the potential available Zn content in soil (weakly bound to organic matter and carbonate bound Zn) and MF and DI values were all positively correlated with the Zn concentrations in pak choi, indicating that these indexes can be used to predict the bioavailability of Zn in soil. This study can provide a good reference for Se and Zn biofortification of plants in calcareous soil.

Assessment of speciation and in vitro bioaccessibility of selenium in Se-enriched Pleurotus ostreatus and potential health risks.

Due to the two-dimensional effect of selenium (Se) to health, which form of Se is most effective for increasing the bioaccessible Se content in P. ostreatus and whether these products have potential health risks are worth considering. Three Se supplements were applied at different application rates into substrates for cultivating P. ostreatus. The total content and speciation of Se in P. ostreatus fruit bodies were analyzed, and the bioaccessibility of Se was determined via an in vitro physiologically based extraction test (PBET). Results showed that P. ostreatus had the highest utilization efficiency with selenite, followed by Se yeast and selenate. Organic Se (46%-90%) was the major Se speciation in P. ostreatus regardless applied Se species. Although the Se bioaccessibility of the gastrointestinal digestion of P. ostreatus was high (70%-92%), the estimated daily intake and target hazard quotient values are all within the safe ranges. Se-enriched P. ostreatus can be safely used as a dietary source of Se for increasing Se intake.

Assessing the uptake of selenium from naturally enriched soils by maize (Zea mays L.) using diffusive gradients in thin-films technique (DGT) and traditional extractions.

A generally accepted method to predict selenium (Se) bioavailability of long-term contaminated soils has not yet been established, even if risk assessments in selenosis areas are crucial. In this study, a set of methods were tested to assess the bioavailability of Se to field maize. Fifty maize (Zea mays L.) samples and corresponding soils were collected from a selenosis area (Ziyang, China). The diffusive gradients in thin-films (DGT) technique and the traditional chemical extraction methods, including seven single-step extraction procedures and a five-step sequential extraction were used to predict the bioaccumulation of Se in plant. The result verified the presence of 50% of total Se in the form of residual Se fraction, followed by organic-bound and Fe-Mn oxide-bound Se fractions in soil. In addition, Se6+, Se4+, and Se2- were all detected in the solution extracted by H2O, KCl, phosphate-buffered solution (PBS), NaHCO3, ethylenediaminetetraacetic acid-2Na (EDTA-2Na) and ammonium bicarbonate-diethylenetriaminepentaacetic acid (AB-DTPA), but Se6+ was not extracted by NaOH. The Se extracted by single-step extraction methods was weakly correlated with the Se uptake by plants with relatively high Se concentration (>3 mg·kg-1). The abilities of the tested methods to predict Se bioavailability in naturally Se-enriched soils declined in the following order: DGT > soil solution > PBS > KCl > H2O > NaHCO3 > EDTA > DTPA > NaOH. The ratio of CDGT to soil solution Se (Csoln) totaled 0.13, indicating an extremely low Se supply from the soil solid phase to the soil solution. Se measured by DGT was mainly derived from the soluble and exchangeable Se fractions that can accurately reflect the plant-absorbed Se pool. Therefore, the DGT technique is highly applicable in the simultaneous prediction of Se bioavailability in naturally Se-enriched soils.

Uptake kinetics and interaction of selenium species in tomato (Solanum lycopersicum L.) seedlings.

Selenite and selenate are two main selenium (Se) forms absorbed by plants. The comparative effects of selenite and/or selenate on Se uptake and translocation in plants in spite of their coexistence in the environment are still unclear. Therefore, tomato (Solanum lycopersicum L.) seedlings were grown in a hydroponic solution with exogenous selenite, selenate, or selenite and selenate mixed, and Se concentrations in shoots, roots, and xylem sap were measured after harvest. Results showed that selenite (> 0.1 mg Se L-1) could cause phytotoxicity more easily than selenate (> 1 mg Se L-1) under hydroponic conditions. And the absorbability rate of tomato to selenate was higher than that to selenite when Se application level was 0.0175-0.2998 mg L-1, while the opposite result was observed in other Se concentrations. More Se accumulated in roots and Se(VI) in the xylem sap decreased when both Se forms supplied. This study demonstrated that the difference between selenite and selenate on Se uptake and translocation in tomatoes depended on exogenous Se concentration. And selenite could inhibit the absorption and translocation of selenate when supplied with both Se forms.

Effect of selenium-enriched organic material amendment on selenium fraction transformation and bioavailability in soil.

To exploit the plant byproducts from selenium (Se) biofortification and reduce environmental risk of inorganic Se fertilizer, pot experiment was conducted in this study. The effects of Se-enriched wheat (Triticum aestivum L.) straw (WS + Se) and pak choi (Brassica chinensis L.) (P + Se) amendment on organo-selenium speciation transformation in soil and its bioavailability was evaluated by pak choi uptake. The Se contents of the cultivated pak choi in treatments amended with the same amount of Se-enriched wheat straw and pak choi were 1.7 and 9.7 times in the shoots and 2.3 and 6.3 times in the roots compared with control treatment. Soil respiration rate was significantly increased after all organic material amendment in soil (p < 0.05), which accelerated the mineralization of organic materials and thus resulted in soluble Se (SOL-Se), exchangeable Se (EX-Se), and fulvic acid-bound Se (FA-Se) fraction increasing by 25.2-29.2%, 9-13.8%, and 4.92-8.28%, respectively. In addition, both Pearson correlation and cluster analysis showed that EX-Se and FA-Se were better indicators for soil Se availability in organic material amendment soils. The Marquardt-Levenberg Model well described the dynamic kinetics of FA-Se content after Se-enriched organic material amendment in soil mainly because of the mineralization of organic carbon and organo-selenium. The utilization of Se in P + Se treatment was significantly higher than those in WS + Se treatment because of the different mineralization rates and the amount of FA-Se in soil. Se-enriched organic materials amendment can not only increase the availability of selenium in soil but also avoid the waste of valuable Se source.

Selenium distribution in the Chinese environment and its relationship with human health: A review.

This paper reviewed the Se in the environment (including total Se in soil, water, plants, and food), the daily Se intake and Se content in human hair were also examined to elucidate Se distribution in the environment and its effects on human health in China. Approximately 51% of China is Se deficiency in soil, compared with 72% in the survey conducted in 1989. Low Se concentrations in soil, water, plants, human diet and thus human hair were found in most areas of China. The only significant difference was observed between Se-rich and Se-excessive areas for Se contents in water, staple cereal, vegetables, fruits, and animal-based food, no remarkable contrast was found among other areas (p>0.05). This study also demonstrated that 39-61% of Chinese residents have lower daily Se intakes according to WHO/FAO recommended value (26-34µg/day). Further studies should focus on thoroughly understanding the concentration, speciation, and distribution of Se in the environment and food chain to successfully utilize Se resources, remediate Se deficiency, and assess the Se states and eco-effects on human health.

Role of organic acids on the bioavailability of selenium in soil: A review.

Organic Acids (OAs) are important components in the rhizosphere soil and influence Se bioavailability in soil. OAs have a bidirectional contrasting effect on Se bioavailability. Understanding the interaction of OAs with Se is essential to assessing Se bioavailability in soil and clarifying the role of OAs in controlling the behavior and fate of Se in soil. This review examines the mechanisms for the (im)mobilization of Se by OAs and discusses the practical implications of these mechanisms in relation to sequestration and bioavailability of Se in soil.