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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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Mutation
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Figure 12.2 The complementation test de nes allelism. Are two mutations (a1, a2) allelic if they affect the
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same trait To nd out, mutant homozygotes are crossed to form a heterozygote. (a) If the mutations are allelic, then both copies of the gene in the heterozygote are mutant, resulting in the mutant phenotype. (b) If the mutations are nonallelic, then there is a wild-type allele of each gene present in the heterozygote, resulting in the wild-type phenotype. (The two loci need not be on the same chromosome.)
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structural change (involving the same base pairs) and are thus structural alleles. If a small amount of recombination occurs that generates wild-type offspring, then the two alleles are not mutations at the same point ( g. 12.4). Alleles that were functional but not structural were rst termed pseudoalleles because it was believed that loci were
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Figure 12.3 A heterozygote of two recessive mutations can
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have either the trans or cis arrangement. In the trans position, functional alleles produce a mutant phenotype. (Red marks represent mutant lesions.) In the cis position, functional alleles produce a wild-type phenotype. The cis-trans position effect thus reveals functional alleles.
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made up of subloci. Fine-structure analysis led to the understanding that a locus is a length of genetic material divisible by recombination rather than a bead on a string. Eye-color mutants of Drosophila melanogaster can be studied by complementational analysis. The white-eye locus has a series of alleles producing varying shades of red.This locus is sex linked, at about map position 3.0 on the X chromosome. (Several other eye-color loci on the X chromosome are not relevant to this cross e.g., prune and ruby.) If an apricot-eyed female is mated with a white-eyed male, the female offspring are all heterozygous and have mutant light-colored eyes ( g. 12.5). Thus, apricot and white are functional alleles: they do not complement (table 12.2). To determine whether apricot and white are structural alleles, light-eyed females are crossed with white-eyed males, and the offspring are observed for the presence of wild-type or light-eyed males.Though their rate of appearance is less than 0.001%, this is signi cantly above the background mutation rate. The conclusion is that apricot and white are functional, but not structural, alleles.
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
Functional but not structural alleles
Functional and structural alleles
A locus
Heterozygous F1 in trans position Recombination Recombination
Double mutant Gametes Wildtype +
Single mutant
Single mutant
Figure 12.4 Functional alleles may or may not be structurally
allelic. (Red marks represent mutant sites.) Functional alleles that are not also structural alleles can recombine between the mutant sites, resulting in occasional wild-type (and double mutant) offspring. Structural alleles (which are also always functional alleles) are defective at the same base pairs and cannot form either wild-type or double mutant offspring by recombination.
Fine-Structure Mapping
After Beadle and Tatum established in 1941 that a gene controls the production of an enzyme that then controls a step in a biochemical pathway, Benzer used analytical techniques to dissect the ne structure of the gene. Finestructure mapping means examining the size and number of sites within a gene that are capable of mutation and recombination. In the late 1950s, when biochemical techniques were not yet available for DNA sequencing, Benzer used classical recombinational and mutational techniques with bacterial viruses to provide reasonable estimates on the details of ne structure and to give insight into the nature of the gene. He coined the terms muton for the smallest mutable site and recon for the smallest unit of recombination. It is now known that both muton and recon are a single base pair. Before Benzer s work, genes were thought of as beads on a string. The very low rate of recombination between
Figure 12.5 Crosses demonstrating that apricot and white
eyes are functional, but not structural, alleles in Drosophila. Light-eyed females are heterozygous for both alleles. When testcrossed, they produce occasional offspring that are wildtype (X allele) or light-eyed (Xw allele). This indicates a crossover between the two mutant sites (white and apricot) in the heterozygous females, producing, reciprocally, an allele with both mutational sites and the wild-type.
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