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The cI gene, with the aid of the cII-gene product, is transcribed from a promoter known as pRE, the RE standing for repression establishment ( g. 14.23). Once cI is transcribed, it is translated into a protein called the repressor, which interacts at the left and right operators, oL and oR, of the left and right operons. When these operators are bound by cI protein, transcription of the left and right operons (and therefore also the late operon) ceases. There are several rami cations of the repression. First, lysogeny can be initiated because the int gene has been transcribed at the early stage of infection. Second, since CII and CIII are no longer being synthesized, cI transcription from the pRE promoter stops. However, cI can still be transcribed because there is a second promoter, pRM (RM stands for repression maintenance), that allows low levels of transcription of the cI gene.
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The cIII-protein product inhibits a host cell protease, called FtsH, that would break down the cII-gene product. The cII-gene product binds at two promoters, enhancing their availability to RNA polymerase, just as the CAP-cAMP product enhances the transcription of the lac operon. The cII protein binds at the promoters for cI transcription and for int transcription ( g. 14.22). At this point, the phage can still choose between either the lytic or the lysogenic cycles. Integrase (the product of the int gene)
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cIII protein breaks down FtsH protease int pI
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Figure 14.22 The cII-gene product of phage binds to the cI promoter ( pRE) and the int promoter ( pI), enhancing transcription of those genes. The cIII protein breaks down the FtsH protease that would normally break down the cII protein.
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Early regulation region of phage . Two promoters, pRE and pRM, transcribe the cI and rex genes. The left operator overlaps the left promoter, and the right operator overlaps both the right promoter and the maintenanceof-repression promoter.
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
The cI gene can further control its own concentration in the cell. When the right and left operators were sequenced, each was discovered to have three sites of repressor recognition ( g. 14.24). On the right operator, for example, the right-most site (oR1) was found to be most ef cient at binding repressor. When repressor was bound at this site, the right operon was repressed, and transcription of cI was enhanced (again, in a way similar to enhancement of transcription by the binding of CAPcAMP at the CAP site in the lac operon). Excess repressor, when present, however, was also bound by the other two sites within oR. The foregoing process results in the repression of the cI gene itself. Hence, maintenance levels of CI can be kept within very narrow limits. A third rami cation of repression is the prevention of superinfection. That is, bacteria lysogenic for phage are protected from further infection by other phages because repressor is already present in the cell. Thus, the excess of repressor controls new invading phages. (We say that bacterial cells lysogenic for phage are immune from infection by additional phage.) These bacteria are also protected from infection by T4 phage with rII mutants. The rex-gene product, the product of the other gene in the repressor operon, controls this protection. The promoters for maintenance and establishment of repression differ markedly in their control of repressor gene expression. When pRE is active, a very high level of repressor is present, whereas pRM produces only a low level of repressor. The level of repressor is due to the length of the leader RNA transcribed on the 5 side of the cI gene. The pRE promoter transcribes a very long leader RNA and is very ef cient at translation of the cI region. In contrast, the pRM promoter begins transcription at the initiation codon of the protein.This leaderless messenger RNA is translated very inef ciently into CI. The repressor is a dimer of two identical subunits ( g. 14.25). Each subunit is composed of two domains, or
Amino acids COOH COOH 132 236 (COOH end)
93 131
1 92
(NH2 end)
DNA (a)
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