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Arsenic poisoning affects millions of people worldwide. Human arsenic intake from rice consumption can be substantial because rice is particularly efficient in assimilating arsenic from paddy soils, although the mechanism has not been elucidated. Here we report that two different types of transporters mediate transport of arsenite, the predominant form of arsenic in paddy soil, from the external medium to the xylem. Transporters belonging to the NIP subfamily of aquaporins in rice are permeable to arsenite but not to arsenate. Mutation in OsNIP2;1 (Lsi1, a silicon influx transporter) significantly decreases arsenite uptake. Furthermore, in the rice mutants defective in the silicon efflux transporter Lsi2, arsenite transport to the xylem and accumulation in shoots and grain decreased greatly. Mutation in Lsi2 had a much greater impact on arsenic accumulation in shoots and grain in field-grown rice than Lsi1. Arsenite transport in rice roots therefore shares the same highly efficient pathway as silicon, which explains why rice is efficient in arsenic accumulation. Our results provide insight into the uptake mechanism of arsenite in rice and strategies for reducing arsenic accumulation in grain for enhanced food safety.efflux ͉ influx ͉ arsenic contamination ͉ silicon ͉ aquaporin A rsenic (As) is a human carcinogen, and there may be no threshold below which it does not cause cancer (1). More than 40 million people worldwide are at risk from drinking As-contaminated groundwater (2), and chronic inorganic As poisoning has reached a massive scale in Bangladesh and West Bengal, India (3). In these countries, As-contaminated groundwater is also widely used for irrigating crops during dry season rice production, adding Ͼ1,000 metric tons of As to soil per year in Bangladesh alone and resulting in As accumulation in soils and elevated As uptake by crops (4-6). Elevated As accumulation in rice has the potential to become a new disaster for the population in Southeast Asia (7). As concentrations in rice grain are often high enough to cause concern even in uncontaminated soils containing background levels of As, because paddy rice appears to be particularly efficient in As assimilation compared with other cereal crops (8). Worldwide market surveys show that rice grain contains considerably higher levels of inorganic As than other foods (9, 10). Human intake of As from consumption of rice can be substantial, especially for people who consume a lot of rice (11). It is therefore crucial that the mechanism of As accumulation in rice is understood to counteract this widespread contamination of the food chain.Plants take up arsenate, the predominant form of As in aerobic soils, through phosphate transporters (12, 13). However, in paddy soils, which are flooded during much of the rice growing season, arsenite becomes the predominant chemical species of As (14). It has been shown that arsenite is taken up via aquaglyceroporins in microbes (13,(15)(16)(17). Evidence from physiological studies suggests that arsenite may also be transp...

Contents Summary 777 Introduction 777 Mechanisms of arsenic uptake and efflux 778 Rhizosphere interactions 782 Arsenic metabolism in planta 783 Long‐distance translocation of arsenic 786 Arsenic hyperaccumulation 788 Conclusions 788 Acknowledgements 789 References 789 Summary Arsenic (As) is an element that is nonessential for and toxic to plants. Arsenic contamination in the environment occurs in many regions, and, depending on environmental factors, its accumulation in food crops may pose a health risk to humans. Recent progress in understanding the mechanisms of As uptake and metabolism in plants is reviewed here. Arsenate is taken up by phosphate transporters. A number of the aquaporin nodulin26‐like intrinsic proteins (NIPs) are able to transport arsenite, the predominant form of As in reducing environments. In rice (Oryza sativa), arsenite uptake shares the highly efficient silicon (Si) pathway of entry to root cells and efflux towards the xylem. In root cells arsenate is rapidly reduced to arsenite, which is effluxed to the external medium, complexed by thiol peptides or translocated to shoots. One type of arsenate reductase has been identified, but its in planta functions remain to be investigated. Some fern species in the Pteridaceae family are able to hyperaccumulate As in above‐ground tissues. Hyperaccumulation appears to involve enhanced arsenate uptake, decreased arsenite‐thiol complexation and arsenite efflux to the external medium, greatly enhanced xylem translocation of arsenite, and vacuolar sequestration of arsenite in fronds. Current knowledge gaps and future research directions are also identified.

Arsenic (As) is an environmental and food chain contaminant. Excessive accumulation of As, particularly inorganic arsenic (As(i)), in rice (Oryza sativa) poses a potential health risk to populations with high rice consumption. Rice is efficient at As accumulation owing to flooded paddy cultivation that leads to arsenite mobilization, and the inadvertent yet efficient uptake of arsenite through the silicon transport pathway. Iron, phosphorus, sulfur, and silicon interact strongly with As during its route from soil to plants. Plants take up arsenate through the phosphate transporters, and arsenite and undissociated methylated As species through the nodulin 26-like intrinsic (NIP) aquaporin channels. Arsenate is readily reduced to arsenite in planta, which is detoxified by complexation with thiol-rich peptides such as phytochelatins and/or vacuolar sequestration. A range of mitigation methods, from agronomic measures and plant breeding to genetic modification, may be employed to reduce As uptake by food crops.

Summary• Selenite can be a dominant form of selenium (Se) in aerobic soils; however, unlike selenate, the mechanism of selenite uptake by plants remains unclear.• Uptake, translocation and Se speciation in wheat (Triticum aestivum) supplied with selenate or selenite, or both, were investigated in hydroponic experiments. The kinetics of selenite influx was determined in short-term (30 min) experiments. Selenium speciation in the water-extractable fraction of roots and shoots was determined by HPLC-ICPMS.• Plants absorbed similar amounts of Se within 1 d when supplied with selenite or selenate. Selenate and selenite uptake were enhanced in sulphur-starved and phosphorus-starved plants, respectively. Phosphate markedly increased K m of the selenite influx. Selenate and selenite uptake were both metabolically dependent. Selenite was rapidly converted to organic forms in roots, with limited translocation to shoots. Selenomethionine, selenomethionine Se-oxide, Se-methyl-selenocysteine and several other unidentified Se species were detected in the root extracts and xylem sap from selenite-treated plants. Selenate was highly mobile in xylem transport, but little was assimilated to organic forms in 1 d. The presence of selenite decreased selenate uptake and xylem transport.• Selenite uptake is an active process likely mediated, at least partly, by phosphate transporters. Selenite and selenate differ greatly in the ease of assimilation and xylem transport.

Abstract. The cellular compartmentation of elements was analysed in the Zn hyperaccumulator Arabidopsis halleri (L.) O'Kane & Al-Shehbaz (=Cardaminopsis halleri) using energy-dispersive X-ray microanalysis of frozen-hydrated tissues. Quantitative data were obtained using oxygen as an internal standard in the analyses of vacuoles, whereas a peak/background ratio method was used for quanti®cation of elements in pollen and dehydrated trichomes. Arabidopsis halleri was found to hyperaccumulate not only Zn but also Cd in the shoot biomass. While large concentrations of Zn and Cd were found in the leaves and roots,¯owers contained very little. In roots grown hydroponically, Zn and Cd accumulated in the cell wall of the rhizodermis (root epidermis), mainly due to precipitation of Zn/Cd phosphates. In leaves, the trichomes had by far the largest concentrations of Zn and Cd. Inside the trichomes there was a striking sub-cellular compartmentation, with almost all the Zn and Cd being accumulated in a narrow ring in the trichome base. This distribution pattern was very dierent from that for Ca and P. The epidermal cells other than trichomes were very small and contained lower concentrations of Zn and Cd than mesophyll cells. In particular, the concentrations of Cd and Zn in the mesophyll cells increased markedly in response to increasing Zn and Cd concentrations in the nutrient solution. This indicates that the mesophyll cells in the leaves of A. halleri are the major storage site for Zn and Cd, and play an important role in their hyperaccumulation.

Arsenic (As) exposure from consumption of rice can be substantial, particularly for the population on a subsistence rice diet in South Asia. Paddy rice has a much enhanced As accumulation compared with other cereal crops, and practical measures are urgently needed to decrease As transfer from soil to grain. We investigated the dynamics of As speciation in the soil solution under both flooded and aerobic conditions and compared As accumulation in rice shoot and grain in a greenhouse experiment. Flooding of soil led to a rapid mobilization of As, mainly as arsenite, in the soil solution. Arsenic concentrations in the soil solution were 7-16 and 4-13 times higher under the flooded than under the aerobic conditions in the control without As addition and in the +As treatments (10 mg As kg -1 as arsenite or arsenate), respectively. Arsenate was the main As species in the aerobic soil. Arsenic accumulation in rice shoots and grain was markedly increased under flooded conditions; grain As concentrations were 10-15-fold higher in flooded than in aerobically grown rice. With increasing total As concentrations in grain, the proportion of inorganic As decreased, while that of dimethylarsinic acid (DMA) increased. The concentration of inorganic As was 2.6-2.9 fold higher in the grain from the flooded treatment than in that from the aerobic treatment. The results demonstrate that a greatly increased bioavailability of As under the flooded conditions is the main reason for an enhanced As accumulation by flooded rice, and growing rice aerobically can dramatically decrease the As transfer from soil to grain.

China faces great challenges in protecting its soil from contamination caused by rapid industrialization and urbanization over the last three decades. Recent nationwide surveys show that 16% of the soil samples, 19% for the agricultural soils, are contaminated based on China’s soil environmental quality limits, mainly with heavy metals and metalloids. Comparisons with other regions of the world show that the current status of soil contamination, based on the total contaminant concentrations, is not worse in China. However, the concentrations of some heavy metals in Chinese soils appear to be increasing at much greater rates. Exceedance of the contaminant limits in food crops is widespread in some areas, especially southern China, due to elevated inputs of contaminants, acidic nature of the soil and crop species or cultivars prone to heavy metal accumulation. Minimizing the transfer of contaminants from soil to the food chain is a top priority. A number of options are proposed, including identification of the sources of contaminants to agricultural systems, minimization of contaminant inputs, reduction of heavy metal phytoavailability in soil with liming or other immobilizing materials, selection and breeding of low accumulating crop cultivars, adoption of appropriate water and fertilizer management, bioremediation, and change of land use to grow nonfood crops. Implementation of these strategies requires not only technological advances, but also social-economic evaluation and effective enforcement of environmental protection law.