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Chapter 41 : Conjugative Transposons

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Abstract:

Among medically important gram-positive bacteria, antibiotic resistance is often determined by genes present in a class of transposable elements that Don Clewell named conjugative transposons. The discovery that conjugative transposons do not cause such duplications indicated that their mechanism of transposition was completely different from that of the other types of transposable elements. Conjugative transposons were first isolated from streptococcal strains associated with human disease. The best studied include Tn, which was found in a strain of ; Tn, which came from ; and Tn and the element originally called omega and now renamed Tn, which were both isolated from strains. In this review, the author uses the Tn designations. Although an understanding of the mechanism involved must await further experimentation, the apparent abilities of conjugative transposons to avoid restriction are an obvious advantage for broadening their host ranges. All three laboratories working on the sequences at the ends of conjugative transposons have reported that there are sometimes five and sometimes six Ts at the right end of Tn. To understand the extent of the analogy between insertion and excision of conjugative transposons and lambdoid prophages, further careful quantitation and biochemical work on the former are needed. The chapter also presents evidence supporting the hypothesis that effective mating-pair formation triggers excision of conjugative transposons in gram-positive bacteria.

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41

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Genetic Recombination
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Chromosomal DNA
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Figures

Image of Figure 1
Figure 1

Map of Tn. Locations of the genes and are shown, and their transcription directions are indicated by arrows beneath the map. There are 173 bases between the stop codon of and the end of the transposon. The region required for conjugative transfer ( ) is shaded. The question mark represents a region whose function is unknown because no mutants with insertions into it were isolated. Restriction enzyme sites are indicated above the line: S, Hc, II; Hp, II. There are no recognition sites for HI, II, IRIIIIIIIIIor I( ) or for I or II ( ).

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
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Image of Figure 2
Figure 2

Model for excision of Tn. Thick lines represent Tnand thin lines represent DNA adjacent to the transposon. Coupling sequences are indicated by the hypothetical nucleotide pairs X-Y and Q-R. A staggered cleavage of the phosphodiester backbone on the 3′ side of the coupling sequence on both strands (first line) generates molecules with 3′ single-stranded ends (second line). The target and transposon sequences are joined by their 3′ single-stranded regions to generate an excisant molecule and the transposon circle. Because there is no apparent requirement for homology between the two coupling sequences, both the excisant and the transposon circle contain heteroduplexes consisting of the base pairs originally present in the coupling sequences. Semiconservative replication resolves the heteroduplex in the excisant and generates a pair of molecules (excisant pairs), of which one has the left coupling sequence at the site of excision and the other has the right coupling sequence. (Reproduced from reference 12 with permission.)

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
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Image of Figure 3
Figure 3

Excision products from pJRS1006. The transposon sequence is delineated by dotted lines above the bases; coupling regions, which are in boldface type, are isolated by spaces except in the target. * (in the target), site of insertion of Tn:, bases internal to the transposon;, bases internal to target DNA. Underlined bases in class II excisants are derived from the coupling sequence that was brought in with the transposon. L and R, left and right ends of Tnrespectively. Sequences of the transposon circle joints, target, and class I excisants after replication are from Caparon and Scott ( ). Sequences of the other excisants are predicted.

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
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Image of Figure 4
Figure 4

Model for insertion of Tn. Symbols are the same as in Fig. 2 . (A and B) Complementary bases of the target. Staggered cleavage at the 3′ ends of the new target and the heteroduplex-containing joint of the transposon circle (A) generates molecules with 3′ single-stranded ends (B). The transposon circle and target are covalently joined by their single-stranded ends to create a new insertion with a heter-oduplex at each end (C). Replication resolves the heterodu-plexes and generates a pair of molecules in which one member is flanked by the target sequence at one end and a coupling sequence from the previous insertion at its other end. The second member of the pair would be flanked by the target sequence on the side opposite its location in the first member and would have at its other end the second of the two coupling sequences of the first insertion (D). The two forms would segregate at the first replication event. (Reproduced from reference 12 with permission.)

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
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Image of Figure 5
Figure 5

Insertion of shuttle transposon. Allelic replacement of 6 by (kanamycin resistance gene [Km]). The dark line represents chromosomal DNA of JRS4. The gray circle represents the covalently closed circular form of Tnwhich is introduced into JRS4 by mating with a strain and contains the sandwich cloned at the BsfXI site. The sandwich contains the gene (represented by a white box) cloned between the region 5′ of 6 and the region 3′ of 6. The homologous regions 5′ and 3′ of 6 on the transposon and chromosome of the recipient strain pair with each other, and recombination leads to substitution of the kanamycin resistance gene for 6 and loss of the rest of the transposon, including the gene.

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
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Image of Figure 6
Figure 6

Vectors for construction of chimeric shuttle transposons. The plasmid pVIT121 is a derivative of a pUC plasmid that replicates only in . It contains a DNA fragment that was originally located immediately to the left ( Fig. 1 ) of the XI site of Tn (labeled “L” in this figure) and a fragment originally located immediately to the right (“R”) of this site. The gene (for kanamycin resistance) is inserted between the L and R fragments. Digestion of pVIT121 with HI followed by ligation removed and generated pVIT130, which contains a HIsite that can be used for insertion of the DNA fragments to be studied. The pVIT vectors can be used to generate a shuttle transposon following the introduction of linearized DNA into a gram-positive host that contains a copy of Tnin its chromosome (pVIT121 linearized by digestion with I is shown at the bottom of the figure). Homologous recombination between the L and R segments of both the pVIT vector and the transposon resident in the chromosome result in integration of the cloned DNA into Tn. The shuttle transposon can then be introduced into the desired host by conjugation. Restriction enzyme sites: B, HIBx, XI; P, I. Ap is the vector gene encoding ampicillin resistance.

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
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References

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Tables

Generic image for table
Table 1

Frequency of Tc transconjugants when CKS102 ( ) is used as donor in plate matings

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41
Generic image for table
Table 2

Insertion targets and transposon circle joint

Citation: Scott J. 1993. Conjugative Transposons, p 597-614. In Sonenshein A, Hoch J, Losick R (ed), and Other Gram-Positive Bacteria. ASM Press, Washington, DC. doi: 10.1128/9781555818388.ch41

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