The heavy metal paradox in arbuscular mycorrhizas: from mechanisms to biotechnological applications

Arbuscular mycorrhizal symbioses that involve most plants and Glomeromycota fungi are integral and functional parts of plant roots. In these associations, the fungi not only colonize the root cortex but also maintain an extensive network of hyphae that extend out of the root into the surrounding environment. These external hyphae contribute to plant uptake of low mobility nutrients, such as P, Zn, and Cu. Besides improving plant mineral nutrition, arbuscular mycorrhizal fungi (AMF) can alleviate heavy metal (HM) toxicity to their host plants. HMs, such as Cu, Zn, Fe, and Mn, play essential roles in many biological processes but are toxic when present in excess. This makes their transport and homeostatic control of particular importance to all living organisms. AMF play an important role in modulating plant HM acquisition in a wide range of soil metal concentrations and have been considered to be a key element in the improvement of micronutrient concentrations in crops and in the phytoremediation of polluted soils. In the present review, we provide an overview of the contribution of AMF to plant HM acquisition and performance under deficient and toxic HM conditions, and summarize current knowledge of metal homeostasis mechanisms in arbuscular mycorrhizas.