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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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14. Gene Expression: Control in Prokaryotes and Phages
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The McGraw Hill Companies, 2001
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Transposable Genetic Elements
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Phenotypic and Genotypic Effects of Transposition
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Transposition can have several effects on the phenotype and genotype of an organism. If transposition takes place into a gene or its promoter, it can disrupt the expression of that gene. Depending on the orientation of a transposon, it can prevent the expression of genes. A transposon can also cause deletions and inversions. Direct repeats on a chromosome can come about, for example, by the sequential transposition of the same IS or transposon, in the same orientation. Pairing followed by recombination results in a deletion of the section between the repeats ( g. 14.34). In the case of inverted repeats, pairing followed by recombination results in an inversion of the section between the repeats.
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A well-known case of transposon orientation controlling a phenotype in bacteria occurs in Salmonella typhimurium. The agella of this bacterium occur in two types. Any particular bacterium has either type 1 or type 2 agella (called phase 1 or phase 2 agella). The difference is in the agellin protein the agella are composed of. Phase 1 agella are determined by the H1 gene and phase 2 agella are determined by the H2 gene. The change from one phase to another occurs at a rate of about 10 4 per cell division. After extensive genetic analysis, the following scheme was suggested and later veri ed using recombinant DNA techniques. The H1 and H2 genes are at separate locations on the bacterial chromosome ( g. 14.35). H2 is part of an operon that also contains the rH1 gene, the repressor of H1. The promoter of this operon lies within a transposon upstream of the operon. When the promoter is in the proper orientation, the H2 operon is expressed, resulting in phase 2 agella. The rH1 gene product represses the H1 gene ( g. 14.35a). If the inverted repeat ends of the transposon undergo recombination, the transposon is inverted (see g. 14.34), moving the promoter into an incorrect orientation for the transcription of the H2 operon. No H1 repressor is made, so the H1 gene is expressed ( g. 14.35b). As N. Kleckner has summarized, transposons can have marked effects on the phenotype by their actions in transposition and by the fact that they may carry genes valuable to the cell. However, they can also exist without any noticeable consequences. This fact has led some evolutionary geneticists to suggest that transposons are an evolutionary accident that, once created, are selfmaintaining. Since they may exist without a noticeable bene t to the host s phenotype, transposons have been
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Pairing and recombination in DNA repeats. (a) Direct repeats can result in deletion (in the form of a circle) due to a single crossover. (b) Inverted repeats can result in an inversion of the region between the repeats due to a crossover.
Arrangement of agellin genes on the Salmonella chromosome. The promoter (p) is within a transposon. In one orientation (a), the H2 operon is transcribed, which results in H2 agellin and rH1 protein, the repressor of the H1 gene. In the second orientation (b), the H2 operon is not transcribed, resulting in uninhibited transcription of the H1 gene.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
14. Gene Expression: Control in Prokaryotes and Phages
The McGraw Hill Companies, 2001
Fourteen
Gene Expression: Control in Prokaryotes and Phages
Nancy Kleckner (1947 ).
(Courtesy of Nancy Kleckner. Photo by Stu Rosner.)
proteins is the direct result of the gene product of the htpR gene, which codes for 32. The normal sigma factor is 70, the product of the rpoD gene; the heat shock genes have promoters recognized by 32 rather than 70. Heat causes the htpR gene to become active, as well as stabilizing 32. From DNA sequence data, the difference in promoters between normal genes and heat shock genes seems to lie in the 10 consensus sequence (Pribnow box). In normal genes, it is TATAAT; in heat shock protein genes, it is CCCCATXT, in which X is any base.
referred to as sel sh DNA. In recent theoretical and experimental studies, however, some scientists have suggested that transposons improve the evolutionary tness of the bacteria that have them (see chapter 20).
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