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Not only are the three lac genes (z, y, a) induced together, but they are adjacent to one another in the E. coli chromosome; they are, in fact, transcribed on a single, polycistronic messenger RNA ( g. 14.2). Induction involves the protein product of another gene, called the regulator gene, or i gene (lacI). Although the regulator gene is located adjacent to the three other lac genes, it is a totally independent transcriptional entity. The regulator speci es a protein, called a repressor, that interferes with the transcription of the genes involved in lactose metabolism.
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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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Lac Operon (Inducible System)
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Figure 14.1 The enzyme -galactosidase hydrolytically cleaves lactose into glucose and galactose (a). The enzyme can also convert lactose to allolactose (b).
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The lac operon is transcribed as a multigenic (polycistronic) mRNA. The z, y, and a indicate the lacZ, lacY, and lacA loci. The mRNA transcript is then translated as individual proteins. The lac operon regulator gene is denoted as i; the o stands for operator and the p for promoter. Both the operon and the regulator gene have their own promoters. (Source: Data from R. C. Dickson, et al., Genetic regulation: The lac
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control region, Science, 187:27 35, January 10, 1975.)
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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 Operator
For the repressor protein to exert its in uence over transcription, there must be a control element (receptor site) located near the beginning of the -galactosidase (lacZ) gene.This control element is a region referred to as the operator, or operator site ( g. 14.2).The operator site is a sequence of DNA that the product of the regulator gene, the repressor, recognizes. When the repressor is bound to the operator, it either interferes with RNA polymerase binding or prevents the RNA polymerase from achieving the open complex (see chapter 10). In either case, transcription of the operon is prevented ( g. 14.3). The repressor is released when it combines with an inducer, a derivative of lactose called allolactose (see g. 14.1).
Note that the promoter not only is recognized by RNA polymerase but also has other controlling elements in the immediate vicinity of the initiation site of transcription. We can now de ne an operon as a sequence of adjacent genes all under the transcriptional control of the same promoter and operator. The nucleotide sequence of the lac operator region is shown in gure 14.4.The operator in gure 14.3 is referred to as the primary operator, o1, centered at 11. Two other operator sequences have been found. One, o2, is centered at 412.The third overlaps the C-terminal end of the i gene, is centered at 82, and is referred to as o3. The structure of the repressor and its interaction with the operator sites was worked out recently with X-ray crystallography. The functional repressor is a homotetramer of the protein product of
The repressor. By binding to the operator, the repressor either prevents RNA polymerase from binding to the promoter and transcribing the lac operon as shown, or prevents the polymerase from achieving the open con guration. In either case, transcription of the lac operon is prevented. When the repressor is not present, transcription takes place. The functional repressor is a tetramer.
i E. coli chromosome z
p o3
CAP site
Cap site 35 sequence 10 sequence
ShineDalgarno sequence
o1 Repressor binding
DNA sequence
5 5 80 70 60 50 40 30 20 10 +1 +10 +20 +30
The lac operon promoter and operator regions. The CAP site is described later. The base sequence corresponds to the diagram above it. The terminal amino acids of the i gene are shown, as well as the initial amino acids of the lacZ gene. In addition, we picture the Shine-Dalgarno sequence of the DNA, the repressor-binding region (centered at around 10 of the gene), the 10 and 35 sequences of the promoter, and primary (o1) and secondary (o3) operator sites (see text). (Data from R. C. Dickson, et al., Genetic
regulation: The lac control region, Science, 187:27 35, January 10, 1975.)
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