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IS 26 has been shown to form cointegrates both by a copy-in mechanism involving one insertion sequence (IS) and a target and by a targeted conservative mechanism involving two ISs. IS 26 is the flagship of a group of 65 bacterial ISs in the recently redefined IS 6 /IS 26 family. Here, whether other family members can also use two mechanisms was examined using members of the IS 257 /IS 431 and IS 1216 isoform groups, which are associated with antibiotic resistance genes in staphylococci and enterococci, respectively. Transposases Tnp257 and Tnp1216 have 39% and 47% amino acid identities, respectively, with Tnp26 and are 62% identical to one another. Using a novel transposition assay, pUC-based plasmids carrying these ISs integrated into the chromosome of a temperature-sensitive polA  Escherichia coli strain grown at the restrictive temperature. In the cointegrates, the plasmid carrying IS 257 was flanked by various 8-bp target site duplications, consistent with random target selection. However, in a mating-out assay, only the targeted conservative reaction was detectable at a low frequency in a recA -negative E. coli strain, indicating that IS 257 is at least 100-fold less active than IS 26 For IS 1216 , in mating-out assays, both copy-in and targeted conservative cointegrate formation were detectable at frequencies similar to those observed for IS 26 Duplication of various 8-bp target sites was detected for the copy-in route. For both IS 257 and IS 1216 , when both of the plasmids carried an IS, the targeted conservative route occurred at a significantly higher frequency than the copy-in route, and only cointegrates formed by the conservative route were detected. IMPORTANCE IS 26 differs from other studied ISs in the reactions that it can undertake. The differences make IS 26 uniquely suited to its key role in the recruitment and spread of antibiotic resistance genes in Gram-negative bacteria. However, whether other ISs in the IS 6 /IS 26 family can perform the same reactions is not known. IS 257 /IS 431 and IS 1216 isoforms found associated with antibiotic resistance genes in the Gram-positive bacteria staphylococci, enterococci, streptococci, and clostridia are related to IS 26 However, the way that they move had not been investigated, limiting interpretation of their role in resistance gene dissemination and in the formation of cointegrates and complex resistance regions in staphylococci and enterococci. Here, they are shown to share the broad catalytic capabilities of IS 26 , demonstrating that it is likely that all members of the redefined IS 6 /IS 26 family of bacterial ISs likewise are able to use both the copy-in and conservative routes.

IS 26 Family Members IS 257 and IS 1216 Also Form Cointegrates by Copy-In and Targeted Conservative Routes