Rhizobia adopt many different lifestyles including survival in soil, growth in the rhizosphere, attachment to root hairs and infection and growth within legume roots, both in infection threads and in nodules where they fix nitrogen. They are actively involved in extracellular signalling to their host legumes to initiate infection and nodule morphogenesis. Rhizobia also use quorum-sensing gene regulation via N-acyl-homoserine lactone signals and this can enhance their interaction with legumes as well as their survival under stress and their ability to induce conjugation of plasmids and symbiotic islands, thereby spreading their symbiotic capacity. They produce several surface polysaccharides that are critical for attachment and biofilm formation; some of these polysaccharides are specific for their growth on root hairs and can considerably enhance their ability to infect their host legumes. Different rhizobia use several different types of protein secretion mechanisms (Types I, III, IV, V and VI), and many of the secreted proteins play an important role in their interaction with plants. This review summarizes many of the aspects of the extracellular biology of rhizobia, in particular in relation to their symbiotic interaction with legumes.
ForewordThis article will focus mostly on the aspects of extracellular biology associated with Rhizobium-legume interactions, ranging from initial attraction and attachment through to root colonization and infection. It will start with a summary overview of the essential events leading to nodule formation on legumes by almost all rhizobia. These events include exchange of signals between the two prospective symbiotic partners and the ability of the bacteria to initiate developmental changes in legumes leading to the formation of infected nitrogen-fixing nodules. The review will then focus on how the extracellular biology of rhizobia modifies and enhances these different stages of interactions with legumes, particularly those that are infected via root-hair infections. To optimize legume infection and form effective nitrogenfixing nodules, rhizobia export different surface polysaccharides and proteins, deliver proteins into plant cells and use quorum-sensing regulation of gene expression to coordinate their behaviour in ways that enhance and spread their symbiotic capacity. Rhizobia are also highly attuned to recognize and utilize various plant-made components, some of which act as signals and others as nutrients during different stages of the interaction.