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Constitutive mutants are mutants in which the three lac operon genes are transcribed at all times that is, they are not turned off even in the absence of lactose. Inspection of gure 14.3 shows that constitutive production of the enzymes can come about in several ways. A defective repressor, produced by a mutant regulator gene, will not turn the system off, nor will a mutant op-
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erator that will no longer bind the normal repressor. The regulator constitutive mutants are designated i ; the operator constitutive mutants are designated oc. Both types of mutants produce the same phenotype: constitutive expression of the three lac operon genes. When a new mutant is isolated, it is possible to determine whether it is caused by a regulator or operator mutation. For example, we can determine the exact location of a mutation on the bacterial chromosome by standard mapping techniques (see chapter 7) or, more recently, by DNA sequencing (see chapter 13). Alternatively, the Jacob and Monod model predicts different modes of action for the two types of mutations. In merozygotes, a constitutive operator mutation affects only the operon it is physically a part of. Operator muta-
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(a) A lac operon in E. coli with a mutation of the regulator gene (i ). Transcription and translation of this gene yield a defective repressor; the cell thus has constitutive production of the lac operon. In (b), the wild-type regulator gene is introduced in an F factor; there is both a bacterial chromosome and an F factor, each containing a regulator gene. (The F operon carries a mutant z allele, allowing us to keep track of the transcriptional control of the chromosomal operon only.) In this case, the phenotype is now normal (inducible) because enough repressor is produced by the F allele (i ), by transcription and translation, to bind to both operators. RNA polymerase is shown as solid spheres on the DNA; the wild-type repressor is shown as a green square; the mutant repressor, which cannot bind to the operator, is shown as a red diamond.
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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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Lac Operon (Inducible System)
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tions are therefore called cis-dominant. However, a constitutive i-gene mutation, since it works through an altered protein, is recessive to a wild-type regulator gene in the same cell, regardless of which operon (chromosomal or F factor) the mutation is on. Constitutive regulator mutations are, therefore, trans-acting. (If two mutations are on the same piece of DNA, they are in the cis con guration. If they are on different pieces of DNA, they are in the trans con guration.) Trans-acting mutations usually work through a protein product that diffuses through the cytoplasm. Cis-acting mutants are changes in recognition sequences on the DNA. In gure 14.6a, the bacterium has a regulator constitutive mutation (i ); the cell has constitutive production
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of the operon. If the wild-type regulator is introduced in an F plasmid ( g. 14.6b), the normal (inducible) phenotype is restored because the F i allele is dominant to the chromosomal mutation the i regulates both the chromosomal and F operons. Hence, both operons are inducible. We don t need to be concerned about the other components of the F plasmid because it carries a z allele; only the activity of the chromosomal operon will be observed. In gure 14.7a, however, the chromosomal operon carries an operator constitutive mutation; the cell also has constitutive production of the operon. When a wild-type operator is introduced into the cell in an F plasmid ( g. 14.7b), the cell still has the constitutive phenotype because the operator allele on the F
Figure 14.7 (a) A lac operon in E. coli with a mutation of the operator (oc). The cell has a constitutive phenotype; the operator cannot bind the wild-type repressor protein, and thus transcription is continuous, even in the absence of lactose. The phenotype is unchanged even when a wild-type operator is introduced into the cell in an F factor (b); there is both a bacterial chromosome and an F factor, each containing an operator. (The F operon carries mutant regulator and z alleles, allowing us to keep track of the transcriptional control of the chromosomal operon only.) The F operator does not change the phenotype of the cell because the wild-type operator exerts no control over the chromosomal operator, which exerts a cis-dominant effect; another operator on another operon has no effect. RNA polymerase is shown as solid spheres on the DNA; the wild-type repressor is shown as a green square; the mutant repressor, which cannot bind to the operator, is shown as a red diamond.
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