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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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12. DNA: Its Mutation, Repair, and Recombination
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The McGraw Hill Companies, 2001
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Recombination
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5 3 3 5 (a) Endonuclease
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Figure 12.38 The double-strand break model of genetic
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(b) 5 3 3 3 Exonuclease
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RecA-mediated invasion
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recombination. Two homologous duplexes (a: red, blue) of the four present in a meiotic tetrad are shown. An endonuclease creates a double-stranded break in one of the duplexes (b). A 5 3 exonuclease then digests away from the break in both directions, creating 3 tails (c). RecA-mediated invasion occurs in the second duplex (d), followed by repair synthesis to close all gaps (e). Branch migration then takes place (f ). Each of the Holliday junctions is then resolved independently, either by nicks in the two outer strands (o) or the two inner strands ( i). Therefore, four resolution structures are possible (g j). In patches, the ends of each duplex are the same as the original, indicating that there may not be recombination for loci anking the point of crossover. In splices, the ends of each duplex have recombined, indicating that anking loci may have crossed over. (Reprinted from Cell, Vol. 87, Frank Stahl, Meiotic Recombination in
Yeast: Coronation of the Double-Stranded-Break Repair Model, pp. 965 968, Copyright 1996, with permission from Elsevier Science.)
(d) Repair synthesis
(e) o
Branch migration
i (f)
Resolution
oo, oo Patch (g)
ii, ii Patch (h)
( g. 12.39). As the junction points move, they create heteroduplex DNA, places where the two strands of each double helix come from different original helices. These stretches have the potential to produce mismatches where the two chromatids differed originally. To resolve the cross-linked duplexes, a second cut at each junction is required. Each of the two crossover points is a Holliday junction. If we open these junctions, we can see that each can be resolved in two different ways. (RuvC endonuclease, the protein product of the ruvC gene, resolves the Holliday junctions in E. coli; see g. 12.39. RuvC cuts the Holliday junction at the consensus sequence 5 [A or T]TT[G or C]-3 . The cut is on the 3 side of the two thymines.) Since there are two Holliday junctions per crossover, there are four potential combinations, as shown in gure 12.38g j. Some of these combinations produce patches, where no recombination takes place among loci to the sides of the hybrid piece. Other combinations produce splices, where reciprocal recombination of loci takes place at the ends. The Holliday junctions can be seen in the electron microscope ( g. 12.40). Note that homology-directed recombination to repair double-strand breaks is similar to the process shown in gure 12.38.
(i) or
o oo, ii Splice ii, oo Splice
Bacterial Recombination
In bacterial recombination, a linear molecule recombines with a circular molecule (see g. 7.15). Usually, invading DNA originates in the linear molecule. The RecBCD protein, whose subunits are the products of the recB, recC, and recD loci, initiates the rst steps in forming an invading linear DNA molecule. RecBCD is a helicase, an exonuclease, and an endonuclease.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
12. DNA: Its Mutation, Repair, and Recombination
The McGraw Hill Companies, 2001
Twelve
DNA: Its Mutation, Repair, and Recombination
Rotate to open
RuvB
RuvB
RuvC
Open arms into cross (d) Branch migration
Heteroduplex (e)
Structure with RuvA and RuvB
Figure 12.39 Branch migration at a Holliday junction. (a) Two double helices (red, blue) are connected by a
crossover. (b) The structure opens when one of the double helices rotates. (c) We further clarify the structure by separating the arms into an open cross, showing the direction in which the arms move during branch migration (arrows: the equivalent of pulling out on the left and right arms, drawing in the top and bottom arms). (d) A more realistic drawing with the RuvA and RuvB proteins, indicating one of the RuvA tetramers behind the center of the cross. A second tetramer (not shown) is located above the cross center, forming a RuvA sandwich of the cross center. The RuvB hexamers are shown on either side of the cross. (e) RuvC can resolve the cross to form either a splice or a patch, depending on which cut is made. (Reprinted with permission from Nature, Vol. 374, C. Parsons, et al.,
Structure of a Multisubunit Complex that Promotes DNA Branch Migration. Copyright 1995 Macmillan Magazines Limited.)
The RecBCD protein enters a DNA double helix from one end and travels along it in an ATP-dependent process. As it travels along the DNA, it acts as a 3 5 exonuclease, degrading one strand of the linear double helix ( g. 12.41). This process continues until RecBCD comes to a chi site, the sequence 5 -GCTGGTGG-3 ,
which appears about a thousand times on the E. coli chromosome. RecBCD s recognition of that sequence attenuates its 3 5 exonuclease activity and enhances its 5 3 exonuclease activity, begun after an endonucleolytic cleavage. From that point on, RecBCD creates a 3 overhang or tail. That tail is coated by RecA and then
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