Plants require sunlight, water, CO 2 , and essential nutrients to drive photosynthesis and fulfill their life cycle. The photosynthetic apparatus resides in chloroplasts and fundamentally relies on transition metals as catalysts and cofactors. Accordingly, chloroplasts are particularly rich in iron (Fe), manganese (Mn), and copper (Cu). Owing to their redox properties, those metals need to be carefully balanced within the cell. However, the regulation of transition metal homeostasis in chloroplasts is poorly understood. With the availability of the arabidopsis genome information and membrane protein databases, a wider catalogue for searching chloroplast metal transporters has considerably advanced the study of transition metal regulation. This review provides an updated overview of the chloroplast transition metal requirements and the transporters involved for efficient photosynthesis in higher plants.
Transition Metal Transporters Are Central to Chloroplast Metal Regulation and PhotosynthesisChloroplasts harbor three types of membranes: a double (inner and outer) envelope and a thylakoid membrane. The envelope and thylakoid are separated by the stroma compartment. The membranes, and the transporters embedded therein, are selective barriers enabling exchange of ions and metabolites between the cytosol, the stroma, and the thylakoid lumen compartment. Chloroplast functioning and efficient photosynthesis for plant growth rely on the metal cofactormediated electron transport chain, in particular the transition metals Fe, Mn, and Cu [1]. However, as ions, their chemical properties can lead to generation of undesired reactive oxygen species (ROS) [2], which, together with their different binding affinities to proteins [3], greatly challenge the use of metals in oxygenic photosynthesis. To avoid harmful generation of ROS, plants chelate metal ions by organic ligands, such as nicotianamine, phenolic compounds, or organic acids (e.g., citrate, malate, ascorbate) [4]. Thus, the transport, homeostasis, and regulation of the individual transition metals are fundamental to optimizing chloroplast functioning.