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Table 12.2 Complementation Matrix of X-Linked Drosophila Eye-Color Mutants
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Note: Plus sign indicates that female offspring are wild-type; minus sign indicates that they are mutant.
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III. Molecular Genetics
12. DNA: Its Mutation, Repair, and Recombination
The McGraw Hill Companies, 2001
Mutation
sites within a gene hampered the analysis of mutational sites within a gene by means of recombination. If two mutant genes are functional alleles (involving different sites on the same gene), a distinct probability exists that we will get both mutant sites (and both wild-type sites) on the same chromosome by recombination (see g. 12.4); but, in view of the very short distances within a gene, this probability is very low. Although it certainly seemed desirable to map sites within the gene, the problem of nding an organism that would allow ne-structure analysis remained until Benzer decided to use phage T4. r II Screening Techniques. Benzer used the T4 bacteriophage because of the growth potential of phages, in which a generation takes about an hour and the increase in numbers per generation is about a hundredfold. Actually, any prokaryote or virus should suf ce, but Benzer made use of other unique screening properties of the phage that made it possible to recognize one particular mutant in about a billion phages. Benzer used rII mutants of T4.These mutants produce large, smooth-edged plaques on E. coli, whereas the wild-type produces smaller plaques whose edges are not as smooth (see g. 7.7). The screening system that Benzer employed made use of the fact that rII mutants do not grow on E. coli strain K12, whereas the wild-type can.The normal host strain, E. coli B, allows growth of both the wild-type and rII mutants. Thus, various mutants can be crossed by mixed infection of E. coli B cells, and Benzer could screen for wildtype recombinants by plating the resultant progeny phages on E. coli K12 ( g. 12.6), on which only a wildtype recombinant produces a plaque. It is possible to detect about one recombinant in a billion phages, all in an afternoon s work. This ability to detect recombinants occurring at such a low level of frequency allowed Benzer to see recombinational events occurring very close together on the DNA, events that would normally occur at a frequency too low to detect in fruit ies or corn. Benzer sought to map the number of sites subject to recombination and mutation within the rII region of T4. He began by isolating independently derived rII mutants and crossing them among themselves. The rst thing he found was that the rII region was composed of two cistrons; almost all of the mutations belonged to one of two complementation groups. The A-cistron mutations would not complement each other but would complement the mutations of the B cistron. The exceptions were mutations that seemed to belong to both cistrons. These mutations were soon found to be deletions in which part of each cistron was missing (table 12.3). Deletion Mapping. As the number of independently isolated mutations of the A and B cistrons increased, it became obvious that to make every possible pairwise cross would entail millions of crosses. To overcome this prob-
Figure 12.6 Using E. coli K12 and B strains to screen for
recombination at the rII locus of phage T4. Two rII mutants are crossed by infecting the same B-strain bacteria with both phages. The offspring are plated on a lawn of K12 bacteria in which only wild-type phages can grow. The technique thus selects only wild-type recombinants.
lem, Benzer isolated mutants that had partial or complete deletions of each cistron. Deletion mutations were easy to discover because they acted like structural alleles to alleles that were not themselves structurally allelic. In other words, if mutations a, b, and c are functional but not
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