Dissecting the promotional effect of zinc on cadmium translocation from roots to shoots in rice

It is often expected that Zn decreases Cd accumulation in plants due to competition for the same transporters. Here, we found that increasing Zn supply markedly increased the root-to-shoot translocation of Cd in rice. RNA sequencing showed that high Zn up-regulated expression of genes involved in glutathione biosynthesis and metabolism and the Zn/Cd transporter gene OsHMA2, but down-regulated expression of genes related to Zn uptake. Knockout of the iron or Zn transporter genes OsIRT1, OsIRT2, or OsZIP9 did not affect the Zn promotional effect on Cd translocation. Knockout of the manganese/Cd transporter gene OsNRAMP5 greatly reduced Cd uptake but did not affect the Zn promotional effect. Variation in the tonoplast transporter gene OsHMA3 affected Cd translocation but did not change the Zn promotional effect. Knockout of the Zn/Cd transporter gene OsHMA2 not only decreased Cd and Zn translocation, but also abolished the Zn promotional effect. Increased expression of OsHMA2 under high Zn conditions supports the hypothesis that this transporter participates in the promotional effect of Zn on Cd translocation. The results also show that OsIRT1, OsIRT2, and OsZIP9 made only small contributions to Cd uptake under low Zn conditions but not under high Zn conditions, whereas the dominant role of OsNRAMP5 in Cd uptake diminished under low Zn conditions.