This study examines the genetic contexts and evolutionary steps responsible for the formation of the widely spread transposon Tn carrying and which confers resistance to beta-lactam and aminoglycoside antibiotics in Gram-negative bacteria. The and genes were found in several transposons. They were first observed within an IS bounded 3.7 kb transposon (Tn on several plasmids located within a 4.7 kb module. Truncated and expanded variations of Tn were found across other plasmids, as well as in other Gram-negative bacteria (including ). Moreover, and were in much larger resistance-heavy transposons including the ISAba1-bounded 24.6 kb (here called Tn), found in an chromosome. A novel ISKpn12-bounded transposon was also observed to contain and which was found interrupting Tn along with an IS pseudo-compound transposon to form a 24.9 kb resistance island in an plasmid. Multiple mobile genetic elements are involved in the formation of transposon structures that circulate and . Among these, IS and ISAba1 appear to have played a major role in the formation and spread of these elements in the species.
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