Type 4 plant metallothioneins - players in zinc biofortification?

Food security is defined as uninterrupted access to food that meets people's dietary needs. One essential trace element of a complete diet is zinc, which is vital for various processes, including growth, development, and the immune response. The estimated global prevalence of zinc deficiency is around 30%. Meat and meat products provide an abundant and also bioavailable source of zinc. However, in developing countries, access to meat is restricted, and in developed countries, meat consumption has declined for ethical and environmental reasons. The potential for zinc deficiency arises from (i) low concentrations of this element in plant-based diets, (ii) poor zinc absorption from plant-based food in the human intestine, and (iii) the risk of uptake of toxic metals together with essential ones. This review summarises the current knowledge concerning type 4 metallothioneins, which represent promising targets for zinc biofortification. We describe their place in the zinc route from soil to seed, their expression patterns, their role in plants, and their three-dimensional protein structure and how this affects their selectivity towards zinc. This review aims to provide a comprehensive theoretical basis for the potential use of type 4 plant metallothioneins to create zinc-biofortified crops.