Representative IMEs
IMEs 1, 2 are defined as mobile genetic elements that encode their own excision and integration regardless of their mechanism and/or specificity of integration, and that do not encode the T4SS machinery and the coupling protein but are able to hijack or subvert the mating apparatus of related or unrelated conjugative elements (such as an IncA or IncC plasmid, or an ICE), regardless of mechanism.
IMEs are autonomous in integration but nonautonomous in conjugation, and can be divided into two major subcategories: relaxosome-encoding and relaxosome-missing.
Relaxosome-encoding IMEs have attL, int, rlx/mob, nic/oriT, and attR; the relaxase targets an oriT site and the resulting relaxosome is recruited to the T4SS together with a coupling protein encoded by a co-resident conjugative element.
Relaxosome-missing IMEs have attL, int, nic/oriT, and attR, but contain no conjugal transfer genes; a co-resident conjugative element recognizes the nic/oriT site of the IME, sufficiently making its intercellular transfer and subsequent transposition.
Table 1|List of representative relaxosome-encoding IMEs
Category | Sub-category | Designation | Accession number | Synonym | Resistance genes | DR | FASTA sequence | Gene list | GBK-annotation | SQN-annotation | Manual annotation | Gene organization |
Relaxosome-encoding IME |
|
Tn4451 |
U15027 |
|
blaCARB-2, tetA(G), and floR |
GA |
|
|
|
|
|
|
Relaxosome-encoding IME |
|
Tn4555 |
U75371 |
|
sul2 |
|
|
|
|
|
|
|
Relaxosome-encoding IME |
|
Tn6523 |
AF261825 |
SGI1 |
None |
|
|
|
|
|
|
|
Relaxosome-encoding IME |
|
Tn6591 |
AE014073 |
GIsul2 |
cfxA |
|
|
|
|
|
|
|
Relaxosome-encoding IME |
|
Tn6966 |
CP064830 |
|
tetA(B), sul1, qacED1, aadA16, dfrA27, arr-3, catB3, blaOXA-1, aacA4cr, floR, sul2, aph(4)-Ia, aacC4, aphA1, aadA2, dfrA12, and dfrA12 |
|
|
|
|
|
|
|
Table 2|List of representative relaxosome-missing IMEs
Category | Sub-category | Designation | Accession number | Synonym | Resistance genes | DR | FASTA sequence | Gene list | GBK-annotation | SQN-annotation | Manual annotation | Gene organization |
Relaxosome-missing IME |
|
Tn6483 |
KY883660 |
|
qnrVC1, arr2, dfrA22e, qacED1, and sul1 |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6487 |
MF344569 |
|
None |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6562 |
CP027478 |
|
cop |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6565 |
LT969519 |
|
None |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6568 |
KU130294 |
|
None |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6573 |
CP043398 |
|
mcr-11.1 |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6588 |
CP042860 |
|
blaNDM-1, blaOXA-1, bleMBL, aacA4-CR, catB3, lnuF, sul1, aphA6, mph(E), sul2, aph(4)-Ia, aadA2, floR, dfrA12, aphA1, aacC4, qacED1, and arr3 |
TTAAT |
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6589 |
CP042857 |
|
aacC2, aadA2, ampR, blaNDM-1, bleMBL, catB3, cfr, floR, qacED1, qnrA1, rmtB, sul1, tetR(G), and tetA(G) |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6590 |
CP042861 |
|
acr3, arsC, arsH, strA, strB, and sul2 |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6722 |
CP016990 |
|
blaTEM-1, blaCTX-M-3, floR, arr3, and aacA4 |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6735 |
CP014062 |
|
ampC |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6737 |
CP054855 |
|
blaVEB-1, strA, strB, floR, dfrA14b, aadB, cmlA1, qacH, and qnrVC4 |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6779 |
CP054844 |
|
None |
None |
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6836 |
CP013067 |
|
None |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6843 |
AP019193 |
|
cop |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6844b |
CP054855 |
|
tetA(E) |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6852 |
HG530068 |
|
chrA |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6855 |
MK347425 |
|
mexCD-oprJ |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6872 |
LR134189 |
|
None |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6875 |
CP031508 |
|
None |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6877 |
CP021775 |
|
blaGES-1, and blaOXA-2 |
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6872 |
LR134189 |
|
|
|
|
|
|
|
|
|
Relaxosome-missing IME |
|
Tn6875 |
CP031508 |
|
|
|
|
|
|
|
|
|
References
1. Guedon G, Libante V, Coluzzi C et al. The obscure world of integrative and mobilizable elements, highly widespread elements that pirate bacterial conjugative systems. Genes (Basel) 2017; 8.
2. Bellanger X, Payot S, Leblond-Bourget N et al. Conjugative and mobilizable genomic islands in bacteria: evolution and diversity. FEMS Microbiol Rev 2014; 38: 720-60