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To further characterize the  fosB -carrying plasmids of 19 vancomycin-resistant enterococci, the complete sequences of the  fosB - and  vanA -containing plasmids of  Enterococcus faecium  (pEMA120) and  E. avium  (pEA19081) were obtained by single-molecule, real-time sequencing. We found that these two plasmids are essentially identical (99.99% nucleotide sequence identity), which proved the possibility of interspecies transmission. Comparative analysis of the plasmids revealed that the backbone of pEMA120 is 99% similar to a conjugative  fosB -negative  E. faecium  plasmid, pZB18. There is a  traE  disrupted in the transfer region of pEMA120, in comparison to pZB18 with an intact  traE . The difference of their transfer frequencies between pEMA120 and pZB18 suggests this interruption of  traE  might affect conjugative transfer. Two copies of the  fosB  gene linked to a  tnpA  gene, forming an IS L3 -like transposon, were found at separate locations within pEMA120, which had not been reported previously. These two  fosB -carrying transposons were confirmed to form circular intermediates by inverse PCR. The hybridization of plasmid DNA digested by  Bsa I, having restriction site within the  fosB  sequence, demonstrated that the presence of multiple copies of  fosB  per plasmid is common. The total copy number of the  fosB  gene as revealed by qRT-PCR did not correlate with fosfomycin MICs or growth rates at sub-MICs of fosfomycin in different transconjugants. From susceptibility tests, the  fosB  gene, regardless of the copy number, conferred high fosfomycin MICs that ranged from 16384 to 65536 μg/ml. This first complete nucleotide sequence of a plasmid carrying two copies of  fosB  in VRE suggests that the  fosB  gene can transfer to multiple loci of plasmids by the IS L3  family transposase TnpA, possibly in the form of circular intermediates, leading to the dissemination of high fosfomycin resistance in VRE.

frontiers in microbiology

Identification of Novel Conjugative Plasmids with Multiple Copies of fosB that Confer High-Level Fosfomycin Resistance to Vancomycin-Resistant Enterococci