Transposon Tn21, Flagship of the Floating Genome

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Microbiology and Molecular Biology Reviews, September 1999, p. 507-522, Vol. 63, No. 3
1092-2172/99/$04.00+0
Copyright © 1999, American Society for Microbiology. All rights reserved.

Transposon Tn21, Flagship of the Floating Genome

Cynthia A. Liebert,¹ Ruth M. Hall,² and Anne O. Summers¹^,^*

Department of Microbiology, The University of Georgia, Athens, Georgia 30602-2605,¹ and CSIRO Molecular Science, North Ryde, New South Wales 1690, Australia²

The transposon Tn21 and a group of closely related transposons (the Tn21 family) are involved in the global disseminationof antibiotic resistance determinants in gram-negative facultativebacteria. The molecular basis for their involvement is carriageby the Tn21 family of a mobile DNA element (the integron) encodinga site-specific system for the acquisition of multiple antibioticresistance genes. The paradigm example, Tn21, also carries genesfor its own transposition and a mercury resistance (mer) operon.We have compiled the entire 19,671-bp sequence of Tn21 and assessedthe possible origins and functions of the genes it contains. Ourassessment adds molecular detail to previous models of the evolutionof Tn21 and is consistent with the insertion of the integron In2into an ancestral Tn501-like mer transposon. Codon usage analysisindicates distinct host origins for the ancestral mer operon,the integron, and the gene cassette and two insertion sequenceswhich lie within the integron. The sole gene of unknown functionin the integron, orf5, resembles a puromycin-modifying enzymefrom an antibiotic producing bacterium. A possible seventh genein the mer operon (merE), perhaps with a role in Hg(II) transport,lies in the junction between the integron and the mer operon.Analysis of the region interrupted by insertion of the integronsuggests that the putative transposition regulator, tnpM, is theC-terminal vestige of a tyrosine kinase sensor present in theancestral mer transposon. The extensive dissemination of the Tn21family may have resulted from the fortuitous association of agenetic element for accumulating multiple antibiotic resistances(the integron) with one conferring resistance to a toxic metalat a time when clinical, agricultural, and industrial practiceswere rapidly increasing the exposure to both types of selectiveagents. The compendium offered here will provide a reference pointfor ongoing observations of related elements in multiply resistantstrains emergingworldwide.

^* Corresponding author. Mailing address: Department of Microbiology, The University of Georgia, Athens, GA 30602-2605. Phone:(706) 542-2669. Fax: (706) 542-6140. E-mail: Summers{at}ARCHES.UGA.EDU.

Microbiology and Molecular Biology Reviews, September 1999, p. 507-522, Vol. 63, No. 3
1092-2172/99/$04.00+0
Copyright © 1999, American Society for Microbiology. All rights reserved.

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