Differential Effects of Senescence on the Phloem Exports of Cadmium and Zinc from Leaves to Grains in Rice during Grain Filling

Abstract

In rice, non-essential toxic cadmium (Cd) and the essential nutrient zinc (Zn) share similar transport pathways, which makes it challenging to differentially regulate the allocation of these elements to the grain. The phloem is the main pathway for the loading of these elements into rice grains. It has long been accepted that tissue senescence makes the nutrients (e.g., Zn) stored in leaves available for further phloem export toward the grain. Whether senescence could drive the phloem export of Cd remains unclear. To this end, the stable isotopes 111Cd and 67Zn were used to trace the phloem export and the subsequent allocation of Cd and Zn from the flag leaves, where senescence was accelerated by spraying abscisic acid. Furthermore, changes upon senescence in the distribution of these elements among the leaf subcellular fractions and in the expression of key transporter genes were investigated. Abscisic acid-induced senescence enhanced the phloem export of Zn but had no impact on that of Cd, which was explained by the significant release of Zn from the chloroplast and cytosol fractions (concentrations decreased by ~50%) but a strong allocation of Cd to the cell wall fraction (concentration increased by ~90%) during senescence. Nevertheless, neither Zn nor Cd concentrations in the grain were affected, since senescence strengthened the sequestration of phloem-exported Zn in the uppermost node, but did not impact that of phloem-exported Cd. This study suggests that the agronomic strategies affecting tissue senescence could be utilized to differentially regulate Cd and Zn allocation in rice during grain filling.