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Phage (see g. 7.21) exhibits a complex system of controls of both early and late operons, as well as controls for the decision of lytic infection versus lysogenic integration. The genes are grouped into four operons: left, right, late, and repressor ( g. 14.18). The left and right operons contain the genes for DNA replication and recombination and phage integration. The late operon contains the genes that determine phage head and tail proteins and lysis of the host cell. The sequence of events following phage infection is relatively well known. The map of ( g. 14.18) is a circle, but the chromosome has two linear stages in its life cycle ( g. 14.19). It is packed within the phage head in one linear form, and it integrates into the host chromosome to form a prophage in another linear form ( g. 14.19). Those two linear forms do not have the same ends ( gs. 14.18 and 14.19b). The mature DNA, which is packed within the phage heads before lysis of the cells, is anked by cos sites (chapter 13). It results from a break in the circular map between the A and R loci. The prophage is integrated at the att site, and the circular map is thus broken there at integration. The homologous integration sites on both and the E. coli chromosome consist of a 15 bp core sequence (called O in both), anked by different sequences on both sides in both the bacterium and the phage ( g. 14.20). In the phage, the region is referred to as POP , where P and P (P for phage) are two different regions anking the O core on the phage DNA. In the bacterium, the region is called BOB , where B and B (B for bacterium) are two different regions anking the O core on the E. coli chromosome. Integration, which is a part of the lysogenic life cycle, requires the product of the int gene, a protein known as integrase, and is referred to as site-speci c recombination. Later excision of the prophage, during induction, when the phage leaves the host chromosome to enter the lytic cycle, requires both the integrase and the protein product of the neighboring xis gene, excisionase. After infection of the E. coli cell by a phage, the phage DNA circularizes, using the complementarity of the cos sites. Transcription begins, and within a very short time the phage is guided toward either entering the lytic cycle and producing virus progeny or entering
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LY T I C A N D LY S O G E N I C CYCLES IN PHAGE
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When a bacteriophage infects a cell, it must express its genes in an orderly fashion; some gene products are needed early in infection, and other products are not needed until late in infection. Early genes usually control phage DNA replication; late genes usually determine phage coat proteins and the lysis of the bacterial cell. A phage is most ef cient if it expresses the early genes rst and the late genes last in the infection process. Also, temperate phages have the option of entering into lysogeny with the cell; here, too, control processes determine which path is taken. One generalization that holds true for most phages is that their genes are clustered into early and late operons, with separate transcriptional control mechanisms for each. Phage is perhaps the best-studied bacteriophage. It has a chromosome of about 48,500 base pairs. Since it is a temperate phage, it can exist either vegetatively or as a prophage, integrated into the host chromosome. This phage warrants our attention because of the interesting and complex way that its life-cycle choice is determined. It is a model system of operon controls. The complexity results from having two con icting life-cycle choices. Brie y, the expression of one of the two life-cycle alternatives, lysogenic or lytic cycles, depends on whether two repressors, CI and Cro, have access to op-
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