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Table 14.1 Elements in Phage
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Gene Products cI cII cIII cro N rex int Q FtsH pR pL pRE pRM pR pI tR1 tL1 nutR nutL
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Infection
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Repressor protein whose function favors lysogeny Enhances transcription at the pI and pRE promoters Inhibits the FtsH protease Antirepressor protein that favors lytic cycle Antiterminator acting at nutR and nutL Protects bacterium from infection by T4 rII mutants Integrase for prophage integration Antiterminator of late operon Bacterial protease that degrades cII protein Right operon Left operon Establishment of repression at repressor region Maintenance of repression at repressor region Late operon int gene Terminates after cro gene Terminates after N gene In cro gene In N gene
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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
Fourteen
Gene Expression: Control in Prokaryotes and Phages
Rarely, mutagenic chemicals or radiation cause expression of lytic cycle
Figure 14.26 Summary of regulation of phage life cycles. (1) In the initial infection, transcription begins in cro and N but terminates shortly thereafter at left and right terminators. (2) The product of the N gene allows transcription through the initial terminators; in essence, all genes can now be transcribed. (3) Lysogeny will occur if the cI protein gains access to the right and left operators; the lytic cycle will prevail if the cro-gene product gains access to those two operators.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
14. Gene Expression: Control in Prokaryotes and Phages
The McGraw Hill Companies, 2001
Transposable Genetic Elements
TRANSPOSABLE GENETIC ELEMENTS
Up until this point, we have thought of the genome in fairly conservative terms. If we map a gene today, we expect to see it in the same place tomorrow. However, our discovery of mobile genetic elements has modi ed that view to some extent. We now know that some segments of the genome can move readily from one place to another. Their moving can have an effect on the phenotype of the organism, primarily at the transcriptional level. We thus begin our discussion of mobile genetic elements here, and we conclude it in chapter 16, because mobile elements also affect the phenotypes of eukaryotes.
IS Elements
Transposable genetic elements, transposons, or even jumping genes, are regions of the genome that can move from one place to another. In some cases, transposition is conservative: the transposons move without copying themselves. They are liberated from the donor site by doublestrand breaks in the DNA. In other cases, transposition is replicative: a copy of the transposon is inserted while the original stays in place.This mechanism involves only singlestrand breaks of the DNA at the donor site. Barbara McClintock rst discovered transposable elements in corn in the 1940s (see chapter 16); they were discovered in prokaryotes in 1967, where they rst showed up as polar mutants in the galactose operon of E. coli. No
genes of the operon were expressed past the point of the polar mutation.This effect was explained by assuming that the transposon brought with it a transcription stop signal. The presence of an inserted piece of DNA in these polar mutants was veri ed by heteroduplex analysis ( g. 14.27). The rst transposable elements discovered in bacteria were called insertion sequences or IS elements. It turns out that these are the simplest transposons. The IS elements consist of a central region of about 700 to 1,500 base pairs surrounded by an inverted repeat of about 10 to 30 base pairs, the numbers depending on the speci c IS element. Presumably, the inverted repeats signal the transposing enzyme that it is at the ends of the IS element. The central region of the IS element contains a gene or genes for the transposing event (usually genes for transposase and resolvase enzymes); the relatively small IS elements carry no bacterial genes ( g. 14.28). The target site that the transposable element moves to is not a speci c sequence, as with the att site of . It becomes a direct repeat anking the IS element only after insertion, giving rise to a model of insertion ( g. 14.29). The target site is cut in a staggered fashion, leaving singlestranded ends. The IS element is then inserted between the single-stranded ends. Repair processes convert the two single-stranded tails to double-stranded segments and, hence, to direct anking repeats. When DNA is sequenced, the pattern of a direct anking repeat surrounding an inverted repeat, with a segment in the middle, signals the existence of a transposable element. Currently, we know of more than fteen families, including a total of over ve hundred known members, of IS elements.
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