The conjugative, chromosomally integrating element R391 is the archetype of the IncJ class of mobile genetic elements. Originally found in a South African Providencia rettgeri strain, R391 carries antibiotic and mercury resistance traits, as well as genes involved in mutagenic DNA repair. While initially described as a plasmid, R391 has subsequently been shown to be integrated into the bacterial chromosome, employing a phage-like integration mechanism closely related to that of the SXT element from Vibrio cholerae O139. Analysis of the complete 89-kb nucleotide sequence of R391 has revealed a mosaic structure consisting of elements originating in bacteriophages and plasmids and of transposable elements. A total of 96 open reading frames were identified; of these, 30 could not be assigned a function. Sequence similarity suggests a relationship of large sections of R391 to sequences from Salmonella, in particular those corresponding to the putative conjugative transfer proteins, which are related to the IncHI1 plasmid R27. A composite transposon carrying the kanamycin resistance gene and a novel insertion element were identified. Challenging the previous assumption that IncJ elements are plasmids, no plasmid replicon was identified on R391, suggesting that they cannot replicate autonomously.Horizontal gene transfer, the intraspecies and interspecies exchange of genetic information, plays an important role in the evolution of bacteria (10,24,26,39,65). Three major mechanisms, transformation, transduction, and conjugation (9), provide bacterial populations with access to a "horizontal gene pool," enabling them to rapidly respond to environmental challenges (16, 34).The most important contributor to horizontal gene transfer is the heterogeneous group of mobile genetic elements that includes plasmids, insertion (IS) elements, transposons, integrons, phages, and genomic islands (10, 17). They are "selfish" elements that promote their own maintenance and distribution and, in addition, can function as vectors for accessory DNA elements. These accessory elements commonly consist of genes that are nonessential for survival but confer a phenotype, which is advantageous under particular environmental conditions. Prominent examples of such traits are antibiotic and heavy metal resistances, degradation of xenobiotic compounds, symbiosis and virulence determinants, resistance to radiation, and increased mutation frequency (15).A group of related conjugative DNA elements has been identified as carriers of antibiotic resistance genes in members of the ␥-Proteobacteria from dispersed global locations. R391, the archetype of this group, was originally detected in a South African Providencia rettgeri isolate (8); for many years, it was believed to be a plasmid and was assigned to a new incompatibility group, IncJ. Subsequently, other IncJ elements, conferring the same phenotype as that conferred by R391 (19, 20) (32), and Shewanella putrefaciens (pMERPH; mercury resistance) from the United Kingdom (48). Previous studies have exa...

Although widely studied in gram-positive Streptococci and in the gram-negative Bacteroides, there is a scarcity of information on the occurrence and nature of conjugative transposon-like elements in the well-studied Enterobacteriaceae. In fact, some of the major reviews on conjugative transposons prior to 1996 failed to mention their occurrence in this group. Recently, their presence has been reported in Salmonella, Vibrio and Proteus species, and in some cases such as the SXT element in Vibrio and the IncJ group element CTnR391, there has been some molecular characterization. The elements thus far examined appear to be larger than the common gram-positive conjugative transposons and to be mosaic in structure, with genes derived from several sources. Recent evidence suggests that in the Enterobacteriaceae the elements may be related to enteric pathogenicity islands. The evolution, distribution and role of these elements in the Enterobacteriaceae is discussed.