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(b) Figure 14.25
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The repressor. (a) The repressor is a dimer with each subunit having helical amino- and carboxyl-terminal ends. The helical structure of each amino-terminal end binds in the major DNA groove. (b) Diagram of the interaction of amino acid residues 1 69 (blue) with oR1 (red) in the closely related phage 434. Dashed lines are hydrogen bonds. Numbers 1 to 6 and 4 and 5 are phosphate numbers. Amino acids are designated by the single-letter code ( g. 11.1). Small red circles are water molecules. (From D. W. Rodgers and S. C.
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Harrison, The complex between phage 434 repressor DNA-binding domain and operator site OR3: Structural differences between consensus and nonconsensus half-sites, Structure, 1:227 40, Dec. 15, 1993. Current Biology Ltd.)
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The right operator on the phage chromosome overlaps the pRM and pR promoters. There are three repressor recognition sites within the operator: oR1, oR2, and oR3. Preferential binding by the Cro repressor to oR3 and the CI repressor to oR1 determines whether transcription occurs to the left or the right.
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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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Lytic and Lysogenic Cycles in Phage
ends. The carboxyl- and amino-terminal ends are separated by a relatively open region, susceptible to protease attack. The alpha-helical regions of the amino-terminal ends interdigitate into the major groove of the DNA to locate the speci c sequences making up the left and right operator sequences. As described earlier for the lac operator, oR1, oR2, and oR3 each have twofold symmetry. The binding of the repressor in oR1 enhances the binding of another molecule of repressor into oR2. Together, they enhance pRM transcription, presumably through contact with RNA polymerase. The repressors also block pR transcription (see g. 14.24).
Lysogenic Versus Lytic Response
We have described the mechanism by which establishes lysogeny. How then does turn toward the lytic cycle Here, control is exerted by the cro-gene product, another repressor molecule that works at the left and right operators in a manner antagonistic to the way the CI repressor works. In other words, using the right operator as an example, cro-gene product binds preferentially to the leftmost of the three sites within oR and represses cI but enhances the transcription of cro (see g. 14.24). The cro-gene product can direct the cell toward a lytic response if it occupies the oR and oL sites before the repressor, or if the repressor is removed. From the point of view of phage , when would be a good time for the CI repressor to be removed Thinking in evolutionary terms, we would expect that a prophage might be at an advantage if it left a host s chromosome and began the lytic cycle when it sensed damage to the host. In fact, one of the best ways to induce a prophage to enter the lytic cycle is to direct ultraviolet (UV) light at the host bacterium. (Actually, this was how lysogeny was discovered, by French geneticist Andr Lwoff.) UV light causes damage to DNA and induces several repair systems. One, called SOS repair (see chapter 12), makes use of the protein product of the recA gene. Among the activities of this enzyme is to cleave the repressor in the susceptible region between domains. The cleaved repressor falls free of the DNA, making the operator sites available for the cro-gene product. The lytic cycle then follows. Initially, however, when the phage rst infects an E. coli cell, the decision for lytic versus lysogenic growth is probably determined by the cII-gene product. This protein, as we mentioned, is susceptible to a bacterial protease, which, in turn, is an indicator of cell growth. When E. coli growth is limited, its proteases tend to be limited, a circumstance that would favor lysogeny for the phage. It is the cII protein that, when active, favors lysogeny and when inactive favors the lytic cycle. Thus, under active bacterial growth, the cII protein is
more readily destroyed, it thus fails to enhance cI transcription, and lysis follows. When bacteria are not growing actively, the cII protein is not readily destroyed, it enhances cI transcription, and lysogeny results. Thus, under initial infection, the choice between lysogeny or the lytic cycle depends primarily on the cII protein, which gauges the health and activity of the host. After lysogeny is established, it can be reversed by processes that inactivate the cI protein, indicating genetic damage to the bacterium (the SOS response) or an abundance of other hosts in the environment (zygotic induction, see chapter 7). In zygotic induction, the lytic cycle is induced during conjugation, presumably when an Hfr cell sends a copy of the prophage into an F cell. At that point, without repressor present, the prophage can reassess whether to continue lysogeny or enter the lytic cycle. Not all the details regarding the CI-Cro competition are known, but an understanding of the relationship of lytic and lysogenic life cycles and the nature of DNA-protein recognition has emerged ( g. 14.26 and table 14.1).
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