Combined Impact of Excess Zinc and Cadmium on Elemental Uptake, Leaf Anatomy and Pigments, Antioxidant Capacity, and Function of Photosynthetic Apparatus in Clary Sage (<i>Salvia sclarea</i> L.)

Clary sage (<i>Salvia sclarea</i> L.) is a medicinal plant that has the potential to be used for phytoextraction of zinc (Zn) and cadmium (Cd) from contaminated soils by accumulating these metals in its tissues. Additionally, it has been found to be more tolerant to excess Zn than to Cd stress alone; however, the interactive effects of the combined treatment with Zn and Cd on this medicinal herb, and the protective strategies of Zn to alleviate Cd toxicity have not yet been established in detail. In this study, clary sage plants grown hydroponically were simultaneously exposed to Zn (900 µM) and Cd (100 μM) for 8 days to obtain more detailed information about the plant responses and the role of excess Zn in mitigating Cd toxicity symptoms. The leaf anatomy, photosynthetic pigments, total phenolic and anthocyanin contents, antioxidant capacity (by DPPH and FRAP analyses), and the uptake and distribution of essential elements were investigated. The results showed that co-exposure to Zn and Cd leads to an increased leaf content of Fe and Mg compared to the control, and to increased leaf Ca, Mn, and Cu contents compared to plants treated with Cd only. This is most likely involved in the defense mechanisms of excess Zn against Cd toxicity to protect the chlorophyll content and the functions of both photosystems and the oxygen-evolving complex. The data also revealed that the leaves of clary sage plants subjected to the combined treatment have an increased antioxidant capacity attributed to the higher content of polyphenolic compounds. Furthermore, light microscopy indicated more alterations in the leaf morphology after Cd-only treatment than after the combined treatment. The present study shows that excess Zn could mitigate Cd toxicity in clary sage plants.