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Yanofsky demonstrated colinearity of the gene and protein. He had the advantage of working with a gene whose protein product was known.
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tural allelism and examine the mapping of mutant sites within a gene 317 324
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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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Figure 12.43 Recombination and repair can cause gene
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conversion. During recombination, heteroduplex DNA is formed, containing mismatched base pairs. Without methylation cues, repair enzymes convert the mismatch to a complementary base pair, in a random fashion that is, an AC base pair can be converted to either an AT (a allele) or a GC (a allele) base pair. Two of the four possible repair choices create 3:1 ratios of alleles rather than the expected 2:2 ratios in the offspring. The 3:1 ratio represents gene conversion.
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STUDY OBJECTIVE 4: To study mutagenesis
325 338
After a mutation, the normal phenotype, or an approximation of it, can be restored either by back mutation or suppression. Intragenic suppression occurs when a second mutation within the same gene causes a return of normal or nearly normal function. Intergenic suppression occurs when a second mutation happens, usually in a transfer RNA gene, that counteracts the original mutation. Nonsense, missense, and frameshift mutations can all be suppressed. Spontaneous mutation probably occurs primarily because of tautomerization of the bases of DNA. If a base is in the rare form during DNA replication, it can form unusual or mutant base pairings. We describe the mechanisms of action of the most common mutagens. STUDY OBJECTIVE 5: To investigate the processes of DNA repair and recombination 339 352
Heteroduplex
DNA repair processes can be divided into four categories: damage reversal, excision repair, double-strand break repair, and postreplicative repair. Photoreactivation is an example of damage reversal.Thymine dimers are undimerized by a photolyase enzyme in the presence of light energy. Excision repair removes a damaged section of a DNA strand. Repair enzymes ll in the gap. Excision repair can be divided into three types. In base excision repair, bases are removed by environmental causes or by glycosylases that sense damaged bases. AP endonuclease and an exonuclease, phosphodiesterase, or lyase then removes the base-free nucleotide. Some enzymes use base ipping to gain access to nucleotides in the double helix. In nucleotide excision repair, enzymes in the Uvr system remove a patch containing the lesion, usually a thymine dimer. In mismatch repair, enzymes of the Mut system use methylation cues to remove a progeny patch containing the mismatch. Double-strand break repair relies on one of two mechanisms. In nonhomologous end joining, the cell simply brings the broken ends back together. In homologydirected recombination, the cell repairs the broken ends using a recombinational mechanism. Postreplicative repair lls in gaps left by DNA polymerase III. Some polymerase enzymes can use lesions as templates. Otherwise, the RecA protein is central to the process. A single strand from the undamaged duplex is
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
used to ll the gap in the damaged duplex. Single-stranded DNA induces the SOS response, which temporarily eliminates LexA-mediated repression. Recombination in eukaryotes begins with a doublestranded break in one double helix, followed by invasion of one of the ends into the other double helix. Repair, ligation, and branch migration follow. The crossover points, called
Holliday junctions, need to be resolved, resulting in patches and splices. In E. coli, the RecBCD protein invades linear DNA, creating tails for invasion of the circular bacterial chromosome. Recombination results in heteroduplex DNA, which, if repaired, can lead to gene conversion. Thus, a battery of enzymes within the cell can modify DNA. These enzymes serve in DNA replication, repair, and recombination.
S O L V E D
PROBLEM 1: An investigator isolates two recessive wing
P R O B L E M S
Answer: Both processes are similar in that they entail removal of an incorrect base in a DNA double helix by an excision process followed by a repair process. The processes differ in the event that triggers them. Mismatch repair is triggered by a base pair that does not occupy the correct space in the double helix that is, by a non-Watson and Crick pairing (not AT or GC). AP repair is triggered by enzymes that recognize a missing base.
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