Tripartite Symbiosis Between Legumes, Arbuscular Mycorrhizal Fungi and Nitrogen Fixing Rhizobia: Interactions and Regulation.

Legume plants can interact with nitrogen-fixing rhizobia bacteria and arbuscular mycorrhizal fungi (AMF) simultaneously, forming a tripartite symbiotic association. Co-inoculation studies performed on a variety of legumes have shown that rhizobia and AMF influence each other when they co-occur in tripartite association and affect host plant nutrition and performance. Although single plant-microbe interactions have been extensively studied, our understanding in the field of tripartite interactions is insufficient and current knowledge cannot predict the symbiotic outcome, which appears to depend on many parameters. In this review we examine the current state of research on the legume-rhizobium-AMF tripartite symbiosis. We investigate the dynamic interaction between the two microsymbionts and the effect of one microbe on the other, both at the physiological and the molecular levels, and the result of dual inoculation on host plant growth, fitness and response to stresses. Rhizobia and AMF interact both extraradically and intraradically, effects on microbe and host plant gene expression levels are observed, AMF positively regulates nodulation, while rhizobia can affect AMF root colonisation either positively or negatively. Factors observed to regulate the establishment and function of the tripartite symbiosis, such as the rhizobia-AMF combination, host plant identity and environmental conditions are discussed.