Phylogeny and Expression of NADPH Oxidases during Symbiotic Nodule Formation

The mutualistic interaction between gram-negative soil bacteria and the roots of legumes leads to the establishment of nodules, where atmospheric nitrogen is fixed. Nodulation is a multistep process with numerous essential players. Among these are reactive oxygen species (ROS), which are mainly generated by Nicotinamide Adenine Dinucleotide Phosphate (NADPH) oxidases. In plants, these enzymes are known as respiratory burst oxidase homologs (RBOHs). In legumes, these proteins are encoded by a multigene family with members that are differentially expressed in various tissues and organs at distinct developmental stages. RBOHs have critical roles at several stages of nodulation: in the early signaling pathway triggered by nodulation factors in the root hairs, during both the progression of infection threads and nodule ontogeny, and in nitrogen fixation and senescence. Data from the literature along with the analysis conducted here imply that legumes use different RBOHs for different stages of nodulation; these RBOHs belong to the same phylogenetic subgroup, even though they are not strictly orthologous. Accordingly, the regulation of activity of a given RBOH during the nodulation process probably varies among legumes.