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Heteroduplex analysis revealing a transposon. (a) Two plasmids were hybridized, one with and one without a transposon. (b) The transposon is seen as a single-stranded loop (red); it has nothing to pair with in the heteroduplex. ([a] Courtesy of
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Tamarin: Principles of Genetics, Seventh Edition
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14. Gene Expression: Control in Prokaryotes and Phages
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An IS element (IS5) inserted into a target site in a bacterial chromosome creates a direct repeat on either side of the IS element. The explanation is shown in gure 14.29. An inverted repeat (red) is seen as the same sequence read inwards from outside on the upper strand (left) and the lower strand (right). (Reprinted with
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permission from Nature, Vol. 297, M. Kroger and G. Hoborn, Structural Analysis of Insertion Sequence IS5. Copyright 1982 Macmillan Magazines Limited.)
Insertion of an IS element (IS5 of g. 14.28) results in a direct anking repeat surrounding the transposon in the host chromosome. This occurs because the insertion takes place at a point in which a staggered cut is made in the host DNA, leaving complementary regions on either side of the transposon. Repair replication results in two copies of the anking sequence.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
14. Gene Expression: Control in Prokaryotes and Phages
The McGraw Hill Companies, 2001
Transposable Genetic Elements
Composite Transposons
After the discovery of IS elements, a more complex type of transposable element, a composite transposon, was discovered. A composite transposon consists of a central region surrounded by two IS elements.The central region usually contains bacterial genes, frequently antibiotic resistance loci. For example the composite transposon Tn10 contains the genes for transposase and resolvase, as well as the bacterial gene for -lactamase, which confers resistance to ampicillin ( g. 14.30). Arrangements of composite transposons can vary quite a bit. The IS elements at the two ends can be identical or different; they can be in the same or different orientations; they can be similar to known IS elements or different from any freely existing IS elements. In the latter case, they are called ISlike elements. Two IS elements can transpose virtually any region between them. In fact, composite transposons most likely came into being when two IS elements became located near each other. We can see this very clearly in a simple experiment. In gure 14.31, there is a small plasmid constructed with transposon Tn10 in it. The re-
verse transposon, consisting of the two IS elements and the plasmid genes, or the normal transposon, could each transpose.
Mechanism of Transposition
Transposition can come about by several mechanisms; however, it does not use the normal recombination machinery of the cell (see chapter 12). One model, by J. Shapiro, explains the fact that many transposons in the
J. A. Shapiro (1943 ).
(Courtesy of Dr. J. A. Shapiro.)
Tn10
A composite transposon consists of a central region anked by two IS elements. Transposon Tn10 contains the transposase and resolvase enzyme genes as well as the bacterial gene -lactamase, which protects the cell from the antibiotic ampicillin.
Tn10
Other genes Figure 14.31
Two IS elements in a plasmid can transpose either of the two regions between them. In the case shown, either the Tn10 transposon or the reverse transposon ( other genes ) is transposed.
Transposition or IS Tn10 Normal transposition IS IS
New composite transposon Other genes IS
Reverse transposition
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
14. Gene Expression: Control in Prokaryotes and Phages
The McGraw Hill Companies, 2001
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Gene Expression: Control in Prokaryotes and Phages
process of transposition go through a cointegrate state ( g. 14.32), in which there is a fusion of two elements. During the process of transposition (in this case from one plasmid to another), an intermediate cointegrate stage is formed, made up of both plasmids and two copies of the transposon. Then, through a process called resolution, the cointegrate is reduced back to the two original plasmids, each now containing a copy of the transposon. A diagram of Shapiro s mechanism appears in gure 14.33. At rst, staggered cuts are made in the donor and recipient DNA molecules ( g. 14.33a and b). Then nonhomologous ends are joined so that only one strand of the transposon connects them ( g. 14.33c). This process is presumably controlled by the transposon-coded transposase enzyme. Repair-DNA replication now takes place to ll in the single-stranded segments.The result is a cointegrate of the two plasmids with two copies of the transposon. The last step is a recombination event at a homologous site within the two transposons. This is catalyzed by a resolvase enzyme, which resolves the cointegrate into the original two plasmids, each with a copy of the transposon ( g. 14.33e).
Figure 14.33 The Shapiro mechanism of transposition. Staggered cuts are made at the site of transposon insertion and at either side of the transposon itself (a and b). Nonhomologous single strands join, resulting in two singlestranded copies of the transposon in the cointegrate (c). Repair replication of these single strands produces two copies of the transposon (d). A crossover at the transposon resolves the cointegrate into two plasmids, each with a copy of the transposon (e). Figure 14.32
Transposition frequently goes through an intermediate cointegrate stage. In this case, the transposon is copied from one plasmid to another, with an intermediate stage consisting of a single large plasmid.
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