Lineage-specific regions in Pseudomonas syringae pv tomato DC3000

SUMMARY Comparative analyses of the chromosome of Pseudomonas syringae pv tomato DC3000 with the finished, complete genomes of Pseudomonas aeruginosa PAO1, an animal pathogen, and the non-pathogenic soil inhabitant Pseudomonas putida KT2440 revealed a high degree of sequence conservation in genes involved in 'housekeeping functions'. However, divergence is present among these three fluorescent pseudomonads, yielding 'suites' of species-specific genes that may provide the genetic basis for adaptation to an ecological niche and lifestyle. For DC3000, 1053 genes located on the chromosome were specific to DC3000 and not present in PAO1 or KT2440. The majority of these DC3000-specific genes either lack a known function or are mobile genetic elements. However, these genes do share features among themselves such as association with regions of atypical trinucleotides, unusual G+C content and localization within large tracts of DC3000-specific sequence, suggestive of lateral gene transfer events. Indeed, a comparison of syntenic blocks among these three complete Pseudomonas genomes revealed that a substantial portion (533) of the DC3000-specific chromosomal genes (1053) were located in lineage-specific regions (defined as being larger than 2 kb and enriched in mobile genetic elements and/or genes specific to DC3000 in this three-way comparison). A large proportion of mobile genetic elements (199 of 318 genes; 63%), which are highly enriched in DC3000, were present within such regions. Similarly, most of the genes encoding type III secretion system virulence effectors were located in lineage-specific regions. Consistent with the plasticity of the DC3000 genome, a putative chromosomal inversion mediated by identical copies of ISPsy6 involving 2838 kb (44%) of the DC3000 genome was detected. These data suggest that a substantial portion of the differentiation of DC3000, a plant pathogen, from an animal pathogen and a soil inhabitant has involved transfer of a large number of novel genes coupled with amplification of mobile genetic elements.