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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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the i gene; that is, it is formed from four identical copies of the repressor protein. Since each operator site has twofold symmetry, two repressor monomer proteins bind to each operator site.The monomer is shaped so that it ts into the major groove of the DNA to locate the exact base sequence of the operator; it then binds at that point through electrostatic forces. A tetramer can bind to two of the operator sites at the same time, presumably o1 and o3 or o1 and o2. In the process, the DNA is formed into a loop ( g. 14.5).
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Induction of the Lac Operon
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Under conditions of repression, before the operon can be turned on to produce lactose-utilizing enzymes, the repressor will have to be removed from the operator.The repressor is an allosteric protein; when it binds with one particular molecule, it changes the shape of the protein, which changes its ability to react with a second particular molecule. Here the rst molecule is the inducer allolactose and the second molecule is the operator DNA. When allolactose is bound to the repressor, it causes the repressor to change shape and lose its af nity for operator sequences ( g. 14.5). With allolactose bound to the repressor, the ability of the repressor to bind to the operator is greatly reduced, by a factor of 103. Since no covalent bonds are involved, the repressor simply dissociates from the operator. After the repressor releases from the operator, RNA polymerase can now begin transcription. The three lac operon genes are then transcribed and subsequently translated into their respective proteins. This system of control is very ef cient. The presence of the lactose molecule permits transcription of the genes of the lac operon, which act to break down the lactose. After all the lactose is metabolized, the repressor returns to its original shape and can again bind to the operator. The system is turned off. Using very elegant genetic analysis, details of this system were worked out by Fran ois Jacob and Jacques Monod, who subsequently won 1965 Nobel prizes for their efforts.
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(b) Figure 14.5
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Because the lac operator DNA sequences are palindromes, each half can bind one repressor subunit. (a) The tetrameric repressor binds to o1 and o3, causing the DNA in between to form a loop. Each of the subunits is shown in a different color. The round portion of the subunit in touch with the DNA is the N-terminal end of the repressor subunit; the C-terminal ends form tails that bind the subunits together. Also indicated are the CAP site and the 10 and 35 sequences. (b) When each of the subunits binds an allolactose molecule (black circles), the shape of the middle portion of the subunit changes, causing the subunit to fall free of the operators.
Lac Operon Mutants
Merozygote Formation
Discovery and veri cation of the lac operon system came about through the use of mutants and partial diploids of the lac operon well before DNA sequencing techniques had been developed. The structural (enzymespecifying) genes of the lac operon, z, y, and a, all have known mutant forms in which the particular enzyme does not perform its function. These mutant forms are designated z , y , and a . The alleles for normal forms of the enzymes are z , y , and a . Partial diploids in E. coli can be created through sexduction (chapter 7) because some strains of E. coli have
Fran ois Jacob (1920 ).
(Courtesy of Dr. Fran ois Jacob.)
Jacques Monod (1910 1976).
(Archives Photographiques, Mus e Pasteur.)
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 lac operon incorporated into an F factor. Since F strains can pass the F particle into F strains, lac operon diploids (also called merozygotes, or partial diploids) can be formed. By careful manipulation, various combinations of mutations can be looked at in the diploid state.
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