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3 5 (c) Figure 12.40 A Holliday intermediate structure, equivalent 3 5 3 5 (d) 5
3 RecA-mediated invasion
to the structure seen in gure 12.39c. Each arm is about 1 micron long. (H. Potter and D. Dressler. DNA recombination: In
vivo and in vitro studies, Cold Spring Harbor Symposium on Quantitative Biology, Volume XLIII, 1979, pp. 969 85.)
3 Bacterial 5 chromosome Branch migration
3 5
5 3 5 Repair
invades the circular bacterial chromosome to initiate a crossover event ( g. 12.41). After this pairing, the unpaired segments of the double helix of the bacteria and the exogenote are both degraded. Finally, DNA ligase seals the circular double helix. The resulting hybrid DNA will then be open to mismatch repair that can restore either original base pairs or base pairs from the invading DNA (see below).
5 3 (e)
3 5 5 3 (f) Degradation of linear DNA 5 3 (g) Figure 12.41 RecBCD enters a linear DNA double helix (red) 3 5 3 5
Hybrid DNA
The result of bacterial recombination or meiotic recombination with branch migration is a length of hybrid DNA. This hybrid DNA, also called heterozygous DNA or heteroduplex DNA, has one of two fates, if we assume a difference in base sequences in the two strands. Either the heteroduplex can separate unchanged at the next cell division, or the cell s mismatch repair system can repair it ( g. 12.42). Without appropriate methylation cues, the mismatch repair system can convert the CA base pair to either a CG or a TA base pair. If TA were the original bacterial base pair, conversion to CG would be a successful recombination, whereas return of the CA to TA would be restoration rather than recombination. Recombination in yeast, or any other eukaryote, generates two heteroduplexes. The repair process can cause gene conversion ( g. 12.43), the alteration of progeny ratios indicating that one allele was converted to another, a phenomenon seen in up to 10% of yeast asci. The mismatched AC will be changed to an AT or a GC base pair; the mismatched TG base pair will be changed to TA or CG. The result of the repair, as shown at the bottom of gure 12.43, can be gene conversion in which an ex-
at one end and travels along it, digesting the 3 strand. When the protein encounters a chi site (green), it cuts the other strand and begins acting as a 3 5 exonuclease, creating a 3 overhang (b, c). The 3 overhang can then invade a double helix mediated by RecA. Repair and degradation of the linear DNA results in hybrid DNA in the bacterial chromosome, which can be xed by the mismatch repair system.
pected ratio of 2:2 (a a a a ) is converted to a 3:1 ratio (a a a a ) or a 1:3 ratio (a a a a ). If the heteroduplexes are not repaired, then a single cell generates both kinds of offspring after one round of DNA replication. Thus, the colony from the cell will be half wild-type (a ) and half mutant (a ). We see this phenomenon only in an Ascomycete fungus, such as yeast, in which all the products of a single meiosis remain together.
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
Figure 12.42 Fate of a heteroduplex DNA. Recombination results in heteroduplex DNA with mismatched bases. Replication without repair produces two different daughter molecules. Repair converts the mismatched base pair to one or the other normal base pair.
S U M M A R Y
STUDY OBJECTIVE 1: To look at the nature of mutation in
prokaryotes
316 317
In 1943, Luria and Delbr ck demonstrated that bacterial changes are true mutations similar to mutations in higher organisms. They showed that a high variability occurs in the number of mutants in small cultures as compared with the number of mutants in repeated subsamples of a large culture. Mutations thus occur spontaneously; they are caused by mutagens, which include chemicals and radiation. This chapter is concerned primarily with point mutations rather than changes in whole chromosomes or chromosomal parts. STUDY OBJECTIVE 2: To analyze functional and struc-
Allelism is de ned by the cis-trans complementation test. Complementation implies independent loci, or nonallelic genes. Lack of complementation implies allelism. Functional alleles that differ from each other at the same nucleotides are also called structural alleles. Benzer used T4 phages to do ne-structure studies using complementation testing and deletion mapping. STUDY OBJECTIVE 3: To verify the colinearity of gene
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