Exposure of Salvia sclarea plants to excess Zn for 8 days resulted in increased Ca, Fe, Mn, and Zn concentrations, but decreased Mg, in the aboveground tissues. The significant increase in the aboveground tissues of Mn, which is vital in the oxygen-evolving complex (OEC) of photosystem II (PSII), contributed to the higher efficiency of the OEC, and together with the increased Fe, which has a fundamental role as a component of the enzymes involved in the electron transport process, resulted in an increased electron transport rate (ETR). The decreased Mg content in the aboveground tissues contributed to decreased chlorophyll content that reduced excess absorption of sunlight and operated to improve PSII photochemistry (ΦPSII), decreasing excess energy at PSII and lowering the degree of photoinhibition, as judged from the increased maximum efficiency of PSII photochemistry (Fv/Fm). The molecular mechanism by which Zn-treated leaves displayed an improved PSII photochemistry was the increased fraction of open PSII reaction centers (qp) and, mainly, the increased efficiency of the reaction centers (Fv'/Fm') that enhanced ETR. Elemental bioimaging of Zn and Ca by laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS) revealed their co-localization in the mid-leaf veins. The high Zn concentration was located in the mid-leaf-vein area, while mesophyll cells accumulated small amounts of Zn, thus resembling a spatiotemporal heterogenous response and suggesting an adaptive strategy. These findings contribute to our understanding of how exposure to excess Zn triggered a hormetic response of PSII photochemistry. Exposure of aromatic and medicinal plants to excess Zn in hydroponics can be regarded as an economical approach to ameliorate the deficiency of Fe and Zn, which are essential micronutrients for human health.
Keywords: chlorophyll; chlorophyll fluorescence imaging; hormesis; leaf heterogeneity; non-photochemical quenching (NPQ); nutrient imbalances; phenolics; photoprotection; reactive oxygen species (ROS); stress effects.