The Plasmid Mobilome of the Model Plant-Symbiont Sinorhizobium meliloti: Coming up with New Questions and Answers

Abstract

Rhizobia are Gram-negative Alpha- andBetaproteobacteria living in the underground that have theability to associate with legumes for the establishment ofnitrogen-fixing symbioses.Sinorhizobium melilotiinparticular—the symbiont ofMedicago,Melilotus, andTrigonellaspp.—has for the past decades served as a model organism forinvestigating, at the molecular level, the biology, biochemistry,and genetics of a free-living and symbiotic soil bacterium ofagricultural relevance. To date, the genomes of seven differentS. melilotistrains have been fully sequenced and annotated,and several other draft genomic sequences are also available(http://www.ncbi.nlm.nih.gov/genome/genomes/1004).The vast amount of plasmid DNA thatS. melilotifrequently bears(up to 45% of its total genome), the conjugative ability of some ofthose plasmids, and the extent of the plasmid diversity hasprovided researchers with an extraordinary system to investigatefunctional and structural plasmid molecular biology within theevolutionary context surrounding a plant-associated modelbacterium. Current evidence indicates that the plasmidmobilome inS. melilotiis composed of replicons varying greatlyin size and having diverse conjugative systems and propertiesalong with different evolutionary stabilities and biological roles.While plasmids carrying symbiotic functions (pSyms) are knownto have high structural stability (approaching that ofchromosomes), the remaining plasmid mobilome (referred to asthe non-pSym,functionally cryptic,oraccessorycompartment)has been shown to possess remarkable diversity and to be highlyactive in conjugation. In light of the modern genomic andcurrent biochemical data on the plasmids ofS. meliloti,the current article revises their main structural components,their transfer and regulatory mechanisms, and their potentialas vehicles in shaping the evolution of the rhizobial genome.