Getting to the Route: The Evolution of Nitrogen-Fixing Nodules.

Root nodule symbiosis allows for plant acquisition of reactive nitrogen through fixation of atmospheric molecular dinitrogen by nitrogen-fixing bacteria. Nodulation is a complex trait, with diverse modes of bacterial infection and nodule morphologies across species, reflecting evolutionary adaptation. Understanding ancient forms of this trait may carry advantages for its current utilization, since basal states likely reflect the least complexity. In this review we focus on the evolution of nodule development, particularly on events that have led to increased complexity of this symbiosis in later adaptations. We hypothesize that the ancestral form of nodulation comprises of an evolutionary coupling of nutrient-dependent lateral root development with apoplastic intercellular bacterial growth, alongside the acquisition or evolution of an ancestral chitinaceous signaling molecule by the microbial symbiont. Uncovering the evolutionary adaptations underpinning the extant diversity of this trait allows for a better understanding of the simplest ancestral state.