title: Identification and structure of the Rhizobium galegae common nodulation genes: evidence for horizontal gene transfer.
abstract: Rhizobia are soil bacteria able to fix atmospheric nitrogen in symbiosis with leguminous plants. In response to a signal cascade coded by genes of both symbiotic partners, a specific plant organ, the nodule, is formed. Rhizobial nodulation (nod) genes trigger nodule formation through the synthesis of Nod factors, a family of chitolipooligosaccharides that are specifically recognized by the host plant at the first stages of the nodulation process. Here, we present the organization and sequence of the common nod genes from Rhizobium galegae, a symbiotic member of the RHIZOBIACEAE: This species has an intriguing phylogenetic position, being symbiotic among pathogenic agrobacteria, which induce tumors instead of nodules in plant shoots or roots. This apparent incongruence raises special interest in the origin of the symbiotic apparatus of R. galegae. Our analysis of DNA sequence data indicated that the organization of the common nod gene region of R. galegae was similar to that of Sinorhizobium meliloti and Rhizobium leguminosarum, with nodIJ downstream of nodABC and the regulatory nodD gene closely linked to the common nod operon. Moreover, phylogenetic analyses of the nod gene sequences showed a close relationship especially between the common nodA sequences of R. galegae, S. meliloti, and R. leguminosarum biovars viciae and trifolii. This relationship in structure and sequence contrasts with the phylogeny based on 16S rRNA, which groups R. galegae close to agrobacteria and separate from most other rhizobia. The topology of the nodA tree was similar to that of the corresponding host plant tree. Taken together, these observations indicate that lateral nod gene transfer occurred from fast-growing rhizobia toward agrobacteria, after which the symbiotic apparatus evolved under host plant constraint.
authors.affiliation: Department of Applied Chemistry and Microbiology, Institute of Biotechnology, University of Helsinki, Biocenter 1, FIN-00014 Helsinki, Finland. leena.suominen@helsinki.fi
authors.forename: L
authors.initials: L
authors.lastname: Suominen
authors.forename: C
authors.initials: C
authors.lastname: Roos
authors.forename: G
authors.initials: G
authors.lastname: Lortet
authors.forename: L
authors.initials: L
authors.lastname: Paulin
authors.forename: K
authors.initials: K
authors.lastname: Lindström
mesh_terms: D001426:Bacterial Proteins / Q000235:genetics*; D004269:DNA, Bacterial / Q000737:chemistry / Q000235:genetics; D019143:Evolution, Molecular; D007887:Fabaceae / Q000382:microbiology; D023061:Gene Order; D022761:Gene Transfer, Horizontal; D008969:Molecular Sequence Data; D009876:Operon / Q000235:genetics; D010802:Phylogeny; D010946:Plants, Medicinal; D012336:RNA, Ribosomal, 16S / Q000235:genetics; D015183:Restriction Mapping; D012231:Rhizobium / Q000235:genetics*; D016961:Rhizobium leguminosarum / Q000235:genetics; D017422:Sequence Analysis, DNA; D016962:Sinorhizobium meliloti / Q000235:genetics; D013045:Species Specificity; D013559:Symbiosis
chemical_list: D001426:Bacterial Proteins; D004269:DNA, Bacterial; D012336:RNA, Ribosomal, 16S