Microstructural and physiological responses to cadmium stress under different nitrogen levels in Populus cathayana females and males

Although increasing attention has been paid to the relationships between heavy metal and nitrogen (N) availability, the mechanism underlying adaptation to cadmium (Cd) stress in dioecious plants has been largely overlooked. This study examined Cd accumulation, translocation and allocation among tissues and cellular compartments in Populus cathayana Rehder females and males. Both leaf Cd accumulation and root-to-shoot Cd translocation were significantly greater in females than in males under a normal N supply, but they were reduced in females and enhanced in males under N deficiency. The genes related to Cd uptake and translocation, HMA2, YSL2 and ZIP2, were strongly induced by Cd stress in female roots and in males under a normal N supply. Cadmium largely accumulated in the leaf blades of females and in the leaf veins of males under a normal N supply, while the contrary was true under N deficiency. Furthermore, Cd was mainly distributed in the leaf epidermis and spongy tissues of males, and in the leaf palisade tissues of females. Nitrogen deficiency increased Cd allocation to the spongy tissues of female leaves and to the palisade tissues of males. In roots, Cd was preferentially distributed to the epidermis and cortices in both sexes, and also to the vascular tissues of females under a normal N supply but not under N deficiency. These results suggested that males possess better Cd tolerance compared with females, even under N deficiency, which is associated with their reduced root-to-shoot Cd translocation, specific Cd distribution in organic and/or cellular compartments, and enhanced antioxidation and ion homeostasis. Our study also provides new insights into engineering woody plants for phytoremediation.