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Many  Helicobacter pylori  ( Hp ) strains carry cryptic plasmids of different size and gene content, the function of which is not well understood. A subgroup of these plasmids (e.g. pHel4, pHel12), contain a mobilisation region, but no cognate type IV secretion system (T4SS) for conjugative transfer. Instead, certain  H. pylori  strains (e.g. strain P12 carrying plasmid pHel12) can harbour up to four T4SSs in their genome ( cag -T4SS,  comB ,  tfs3 ,  tfs4 ). Here, we show that such indigenous plasmids can be efficiently transferred between  H. pylori  strains, even in the presence of extracellular DNaseI eliminating natural transformation. Knockout of a plasmid-encoded  mobA  relaxase gene significantly reduced plasmid DNA transfer in the presence of DNaseI, suggesting a DNA conjugation or mobilisation process. To identify the T4SS involved in this conjugative DNA transfer, each individual T4SS was consecutively deleted from the bacterial chromosome. Using a marker-free counterselectable gene deletion procedure ( rpsL  counterselection method), a P12 mutant strain was finally obtained with no single T4SS (P12ΔT4SS). Mating experiments using these mutants identified the  comB  T4SS in the recipient strain as the major mediator of plasmid DNA transfer between  H. pylori  strains, both in a DNaseI-sensitive (natural transformation) as well as a DNaseI-resistant manner (conjugative transfer). However, transfer of a pHel12:: cat  plasmid from a P12ΔT4SS donor strain into a P12ΔT4SS recipient strain provided evidence for the existence of a third, T4SS-independent mechanism of DNA transfer. This novel type of plasmid DNA transfer, designated as  a lternate  D NaseI- R esistant (ADR) mechanism, is observed at a rather low frequency under  in vitro  conditions. Taken together, our study describes for the first time the existence of three distinct pathways of plasmid DNA transfer between  H. pylori  underscoring the importance of horizontal gene transfer for this species.

Multiple Pathways of Plasmid DNA Transfer in Helicobacter pylori