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Only species belonging to the Fabid clade, limited to four classes and ten families of Angiosperms, are able to form nitrogen-fixing root nodule symbioses (RNS) with soil bacteria. This concerns plants of the legume family ( Fabaceae ) and  Parasponia (Cannabaceae)  associated with the Gram-negative proteobacteria collectively called rhizobia and actinorhizal plants associated with the Gram-positive actinomycetes of the genus  Frankia . Calcium and calmodulin-dependent protein kinase ( CCaMK ) is a key component of the common signaling pathway leading to both rhizobial and arbuscular mycorrhizal symbioses (AM) and plays a central role in cross-signaling between root nodule organogenesis and infection processes. Here, we show that  CCaMK  is also needed for successful actinorhiza formation and interaction with AM fungi in the actinorhizal tree  Casuarina glauca  and is also able to restore both nodulation and AM symbioses in a  Medicago truncatula ccamk  mutant. Besides, we expressed auto-active  CgCCaMK  lacking the auto-inhibitory/CaM domain in two actinorhizal species:  C. glauca (Casuarinaceae),  which develops an intracellular infection pathway, and  Discaria trinervis  ( Rhamnaceae ) which is characterized by an ancestral intercellular infection mechanism. In both species, we found induction of nodulation independent of  Frankia  similar to response to the activation of  CCaMK  in the rhizobia-legume symbiosis and conclude that the regulation of actinorhiza organogenesis is conserved regardless of the infection mode. It has been suggested that rhizobial and actinorhizal symbioses originated from a common ancestor with several independent evolutionary origins. Our findings are consistent with the recruitment of a similar genetic pathway governing rhizobial and  Frankia  nodule organogenesis.

The Independent Acquisition of Plant Root Nitrogen-Fixing Symbiosis in Fabids Recruited the Same Genetic Pathway for Nodule Organogenesis