Trace metals at the frontline of pathogen defence responses in non-hyperaccumulating plants

This review evaluates basic and recent findings on the role of trace metals in immunity against pathogen attack in plants that are not metal hyperaccumulators. Metal hyperaccumulation is an exclusive evolutionary trait contributing to efficient plant defence against biotic stress. The defence can be based on direct metal toxicity or the joint effects of accumulated metal and organic compounds, the latter being based on integrated signalling networks. While the role of metals in biotic stress defence of hyperaccumulators has been intensively studied, their role in the pathogen immunity of non-accumulator plants is far less understood. New findings show that in metal non-hyperaccumulating plants, localized hot spots of zinc, manganese, and iron increase plant immunity, while manipulation of nutrient availability may be used for priming against subsequent pathogen attack. Recent findings on the role of metals in plant-pathogen interactions are discussed considering the narrow line between deficiency and toxicity, host-pathogen nutrient competition and synergistic effects of simultaneous metal and biotic stress. We discuss the suitability of the direct-defence and joint-effects hypotheses in non-hyperaccumulating plants, and the involvement of metals as active centres of immunity-related enzymes. We also consider future challenges in revealing the mechanisms underlying metal-mediated plant immunity.