Plasmid Rms149, the archetype of Pseudomonas plasmid incompatibility group IncP-6, was identified in Pseudomonas aeruginosa as an agent conferring resistance to streptomycin, sulfanilamide, gentamicin, and carbenicillin in 1975. It has been classed as a broad-host-range plasmid due to its ability to replicate in both Escherichia coli (where it is designated IncG) and Pseudomonas species, although both species are ␥-proteobacteria. To provide reference information on this Inc group, we have determined the complete sequence of Rms149 and found that, although the genome comprises 57,121 bp, it is essentially a small mobilizable plasmid carrying multiple mobile elements, which make up 79% (>45 kb) of its genome. A replicon has been identified which encodes a single polypeptide with moderate identity to other replication proteins. The region encoding this protein can replicate in Pseudomonas putida and E. coli. This sequence is directly downstream of a putative partitioning region highly similar to that of pRA2. A functional IncQ-type mobilization region is also present. Thus, the backbone appears to be a novel combination of modules already identified in other plasmid systems. Analysis of the segments that fall outside this core of stable inheritance and transfer functions show that this plasmid has been subject to multiple insertion events and that the plasmid appears to carry a considerable load of DNA that no longer should be phenotypically advantageous. The plasmid therefore functions not just as a vehicle for spread of selective traits but also as a store for DNA that is not currently under selection.The key features of a plasmid are the ability to replicate autonomously and to be maintained in a cell lineage without a high rate of segregational loss. An optional but frequently encountered property is the ability to transfer or to be mobilized from one bacterium to another. From these properties it follows that genes that become associated with a plasmid may spread rapidly from one genetic background to another. Some plasmids appear to consist of nothing more than functions that confer the above core abilities, and these have been termed cryptic plasmids.To succeed as such a selfish element, a plasmid must be stable and confer minimal burden on its host, or at least overcome competitive losses by horizontal transfer (42). Conversely, a plasmid carrying genes that promote the growth of its host, relative to competitor bacteria, will benefit by increased propagation. However, since maintenance of a plasmid is generally found to place a metabolic or phenotypic load on the cell, universally useful genes will be selectively reacquired by the chromosome over evolutionary time. Therefore, successful plasmids typically carry payload genes, favorable in some environments but not others (12), or genes with a transient plasmid association that are spreading through a microbial community. In passing through different strains or species, plasmids are exposed to different genetic contexts and provide the opportunity for comb...