FZL, an FZO-like protein in plants, is a determinant of thylakoid and chloroplast morphology

FZO is a dynamin-related membrane-remodeling protein that mediates fusion between mitochondrial outer membranes in animals and fungi. We identified a single FZO-like protein in Arabidopsis, FZL, a new plant-specific member of the dynamin superfamily. FZL is targeted to chloroplasts and associated with thylakoid and envelope membranes as punctate structures. fzl knockout mutants have abnormalities in chloroplast and thylakoid morphology, including disorganized grana stacks and alterations in the relative proportions of grana and stroma thylakoids. Overexpression of FZL-GFP also conferred defects in thylakoid organization. Mutation of a conserved residue in the predicted FZL GTPase domain abolished both the punctate localization pattern and ability of FZL-GFP to complement the fzl mutant phenotype. FZL defines a new protein class within the dynamin superfamily of membrane-remodeling GTPases that regulates organization of the thylakoid network in plants. Notably, FZL levels do not affect mitochondrial morphology or ultrastructure, suggesting that mitochondrial morphology in plants is regulated by an FZO-independent mechanism.