Rice is a major dietary source of the toxic metal, cadmium (Cd). Previous studies reported that the rice transporter, OsNRAMP1, (Natural resistance‐associated macrophage protein 1) could transport iron (Fe), Cd and arsenic (As) in heterologous yeast assays. However, the in planta function of OsNRAMP1 remains unknown. Here, we showed that OsNRAMP1 was able to transport Cd and manganese (Mn) when expressed in yeast, but did not transport Fe or As. OsNRAMP1 was mainly expressed in roots and leaves and encoded a plasma membrane‐localized protein. OsNRAMP1 expression was induced by Cd treatment and Fe deficiency. Immunostaining showed that OsNRAMP1 was localized in all root cells, except the central vasculature, and in leaf mesophyll cells. The knockout of OsNRAMP1 resulted in significant decreases in root uptake of Cd and Mn and their accumulation in rice shoots and grains, and increased sensitivity to Mn deficiency. The knockout of OsNRAMP1 had smaller effects on Cd and Mn uptake than knockout of OsNRAMP5, while knockout of both genes resulted in large decreases in the uptake of the two metals. Taken together, OsNRAMP1 contributes significantly to the uptake of Mn and Cd in rice, and the functions of OsNRAMP1 and OsNRAMP5 are similar but not redundant.

Rice is a major dietary source of the toxic metal cadmium (Cd). Reducing Cd accumulation in rice grain is important for food safety. We generated transgenic rice in two cultivars overexpressing OsNRAMP5, encoding a major influx transporter for manganese (Mn) and Cd, under the control of the OsActin1 or maize Ubiquitin promoter, and tested the effect on Cd uptake and translocation. Overexpression of OsNRAMP5 increased Cd and Mn uptake into the roots, but decreased Cd accumulation in the shoots markedly, and had a relatively small effect on Mn accumulation in the shoots. Overexpressed OsNRAMP5 protein was localized to the plasma membrane of all cell types in the root tips and lateral root primordia without polarity. Synchrotron X-ray fluorescence mapping showed that the overexpression lines accumulated more Cd in the root tips and lateral root primordia compared with wild type. When grown in three Cd-contaminated paddy soils, OsNRAMP5 overexpression decreased grain Cd concentration by 49–94% compared with wild type. Overexpression of OsNRAMP5 decreased Cd translocation from roots to shoots by disrupting radial transport of Cd into the stele for xylem loading, demonstrating the effect of transporter localization and polarity on ion homeostasis.

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Iron (Fe) is essential for rice growth and humans consuming as their staple food but is often deficient because of insoluble Fe(III) in soil for rice growth and limited assimilation for human bodies, while cadmium (Cd) is non-essential and toxic for rice growth and humans if accumulating at high levels. Over-accumulated Cd can cause damage to human bodies. Selecting and breeding Fe-rich but Cd-free rice cultivars are ambitious, challenging and meaningful tasks for researchers. Although evidences show that the mechanisms of Fe/Cd uptake and accumulation in rice are common to some extent as a result of similar entry routes within rice, an increasing number of researchers have discovered distinct mechanisms between Fe/Cd uptake and accumulation in rice. This comprehensive review systematically elaborates and compares cellular mechanisms of Fe/Cd uptake and accumulation in rice, respectively. Mechanisms for maintaining Fe homeostasis and Cd detoxicification are also elucidated. Then, effects of different fertilizer management on Fe/Cd accumulation in rice are discussed. Finally, this review enumerates various approaches for reducing grain Cd accumulation and enhancing Fe content in rice. In summary, understanding of discrepant cellular mechanisms of Fe/Cd accumulation in rice provides guidance for cultivating Fe-fortified rice and has paved the way to develop rice that are tolerant to Cd stress, aiming at breeding Fe-rich but Cd-free rice.

Lead (Pb) is one of the most toxic metals affecting human health globally. Food is an important source of chronic Pb exposure in humans. How Pb is taken up by rice, a staple food for over half of the global population, remains unknown. In the present study, we investigated the role of OsNRAMP5, a member of the NRAMP (Natural Resistance-Associated Macrophage Protein) transporter family, in Pb uptake by rice roots. Heterologous expression of OsNRAMP5 in yeast increased Pb uptake and sensitivity toward Pb. Knockout of OsNRAMP5 in rice by CRISPR/Cas9 gene editing resulted in significant decreases in root uptake of Pb and accumulation in rice shoots. The maximum influx velocity (V max ) for Pb uptake of the knockout mutants was 70% lower than that of wildtype plants. When grown in Pb-contaminated paddy soil, OsNRAMP5 knockout mutants accumulated approximately 50 and 70% lower Pb concentrations in the grain and straw, respectively, than the wild type. OsNRAMP5 expression in rice roots was not affected by Pb exposure. These results indicate that OsNRAMP5 is a major transporter for Pb uptake in rice, in addition to its role in the uptake of manganese and cadmium. This study provides a mechanistic understanding of Pb uptake in rice plants and a potential strategy to limit Pb accumulation in rice grains.