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Insertions and Deletions Transversions
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Ethyl methane sulfonate (CH3SO3CH2CH3) and ethyl ethane sulfonate (CH3CH2SO3CH2CH3) are agents that cause the removal of purine rings from DNA. The multistep process begins with the ethylation of a purine ring and ends with the hydrolysis of the glycosidic (purine-deoxyribose) bond, causing the loss of the base. The molecules of the acridine dyes, such as pro avin and acridine orange ( g. 12.24), are at. Presumably, they initiate mutation by inserting into the DNA double helix, causing the helix to buckle in the region of insertion, possibly leading to base additions and deletions during DNA replication. Crick and Brenner used acridineinduced mutations to demonstrate both that the genetic
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Figure 12.23 Four possible outcomes after treatment of DNA with an alkylating agent, which removes the
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purine adenine in this example. The bases shown in red are the four bases that DNA polymerase may insert opposite the gap. After another round of DNA replication, the gap remains to generate further mutations. The inserted base forms a base pair (blue), which can be a restoration or a transition or transversion mutation.
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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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Mutation
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code was read from a xed point and that it was triplet (chapter 11).
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Misalignment Mutagenesis
Additions and deletions in DNA can also come about by misalignment of a template strand and the newly formed (progeny) strand in a region containing a repeated se-
quence. For example, in gure 12.25 we expect the progeny strand to contain six adjacent adenines because the template strand contains six adjacent thymines. Misalignment of the progeny strand results in seven consecutive adenines: six thymines replicated, plus one already replicated but misaligned. Misalignment of the template strand results in ve consecutive adenines because one thymine is not available in the template. Regions with long runs of a particular base may be very mutation prone. They may explain the hot spots observed by Benzer (see g. 12.8) and others.
Intergenic Suppression
When a critical mutation occurs in a codon, several routes can still lead to survival of the individual; simple reversion and intragenic suppression are two that we have already considered. A third route is through intergenic suppression restoration of the function of a mutated gene by changes in a different gene, called a suppressor gene. Suppressor genes are usually transfer RNA genes. When mutated, intergenic suppressors change the way in which a codon is read. Suppressor genes can restore proper reading to nonsense, missense, and frameshift mutations. Nonsense mutations convert a codon that originally speci ed an amino acid into one of the three nonsense
Figure 12.24 Structure of two acridine dyes: pro avin and
acridine orange.
Figure 12.25 Misalignment of a template or progeny strand during DNA synthesis. If
the progeny strand is misaligned after DNA replication has begun, the resulting progeny strand will have an additional base. If the template strand is misaligned during DNA replication, the resulting progeny strand will have a deleted base. These changes will show up after another round of DNA replication. (From J.W. Drake, B.W. Glickman, and L.S.
Ripley, Updating the Theory of Mutation, American Scientist, 71:621 630, 1983. Reprinted with permission of American Scientist, magazine of Sigma Xi, The Scienti c Research Society.)
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
codons. Missense mutations change a codon so that it speci es a different amino acid. Frameshift mutations, by additions or deletions of nucleotides, cause an alteration in the reading frame of codons. A frameshift mutation, caused by the insertion of a single base, can be suppressed by a transfer RNA that has an added base in its anticodon ( g. 12.26a). It reads four bases as a codon and thus restores the original reading frame. The transfer RNA produced by a nonsense suppressor gene reads the nonsense codon as if it were a codon for an amino acid; an amino acid is placed into
the protein, and reading of the messenger RNA continues. At least three suppressors of the mutant amber codon (UAG) are known in E. coli. One suppressor puts tyrosine, one puts glutamine, and one puts serine into the protein chain at the point of an amber codon. Normally, tyrosine transfer RNA has the anticodon 3 -AUG5 . The suppressor transfer RNA that reads amber as a tyrosine codon has the anticodon 3 -AUC-5 , which is complementary to amber. Hence, a mutated tyrosine transfer RNA reads amber as a tyrosine codon (fig. 12.26b). If the amber nonsense codon is no longer read as a stop signal, then won t all the genes terminating in the amber codon continue to be translated beyond their ends, causing the cell to die In the tyrosine case, two genes for tyrosine transfer RNA were found; one contributes the major fraction of the transfer RNAs, and the other, the minor fraction. It is the minor-fraction gene that mutates to act as the suppressor. Thus, most messenger RNAs are translated normally, and most amber mutations result in premature termination, although a suf cient number are translated (suppressed) to ensure the viability of the mutant cell. In general, intergenic suppressor mutants would be eliminated quickly in nature because they are inefficient the cells are not healthy. In the laboratory, we can provide special conditions that allow them to be grown and studied.
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