A Novel Linear Plasmid Mediates Flagellar Variation in Salmonella Typhi

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Abstract
Unlike the majority of Salmonella enterica serovars, Salmonella Typhi (S. Typhi), the etiological agent of human typhoid, is monophasic. S. Typhi normally harbours only the phase 1 flagellin gene (fliC), which encodes the H:d antigen. However, some S. Typhi strains found in Indonesia express an additional flagellin antigen termed H:z66. Molecular analysis of H:z66+ S. Typhi revealed that the H:z66 flagellin structural gene (fljBz66) is encoded on a linear plasmid that we have named pBSSB1. The DNA sequence of pBSSB1 was determined to be just over 27 kbp, and was predicted to encode 33 coding sequences. To our knowledge, pBSSB1 is the first non-bacteriophage–related linear plasmid to be described in the Enterobacteriaceae. Flagella are whip-like structures found on the surface of bacterial cells that mediate swimming. Flagella contain a protein called flagellin, which is recognised as a danger signal by the immune system. Salmonella Typhi, the bacteria that causes typhoid fever, normally have flagella called H:d, but some strains only from Indonesia express distinct flagella, called H:z66. In this study we have located and sequenced the genes responsible for expressing these alternative flagella. Remarkably, these genes are located on a linear plasmid, an extra-chromosomal element that we have named pBSSB1. The significance of this finding is that linear plasmids are relatively common in bacterial species such as Streptomyces and Borrelia. However, such a linear element has never previously been described in enteric bacteria such as Escherichia coli and Salmonella. The identification of this novel linear plasmid in genetically tractable bacteria will facilitate future studies on the biology of linear plasmids and the pathogenicity of both flagella and Salmonella Typhi.