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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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Gene Expression: Control in Prokaryotes and Phages
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enes are transcribed into RNA, which, for the most part, is then translated into protein. Control mechanisms are exercised along the way. Without some control of gene expression, an Escherichia coli cell, for example, would produce all its proteins in large quantities all the time, and all the cells in a eukaryotic organism would be identical. Although most control mechanisms are negative (preventing something from happening), controls can also be positive (causing some action to occur or enhancing some action). This chapter is devoted to analyzing control processes in prokaryotes and phages; in chapter 16, we examine control processes in eukaryotes. In the process leading from a sequence of nucleotides in DNA to a protein, control is exerted in many places. In general, control of gene expression can take place at the levels of transcription, translation, or protein functioning. The most ef cient place to control gene expression is at the level of transcription. One of the best-understood mechanisms exerts control of transcription, regulating the production of messenger RNA according to need. E. coli messenger RNAs are short-lived in vivo: They degrade enzymatically within about two minutes. A complete turnover (degradation and resynthesis) in the cell s messenger RNA occurs rapidly and continually, and this rapid turnover is a prerequisite for transcriptional control, a central feature of the regulation of prokaryotic gene expression.
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ronment or if it is overproducing tryptophan, the cell stops the manufacture of tryptophan until a need arises again. A repressible system is a system of enzymes whose presence is repressed, stopping the production of the end product when it is no longer needed. Repressible systems are repressed by an excess of the end product of their synthetic (anabolic) pathway. The best-studied inducible system is the lac operon in E. coli. Since the term operon refers to the control mechanism, we will defer a de nition until we describe the mechanism.
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LAC OPERON (INDUCIBLE SYSTEM)
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Lactose Metabolism
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Lactose (milk sugar a disaccharide) is a -galactoside that E. coli can use for energy and as a carbon source after it is broken down into glucose and galactose. The enzyme that performs the breakdown is -galactosidase ( g. 14.1). (The enzyme can additionally convert lactose to allolactose, which, as we will see, is also important.) There are very few molecules of -galactosidase in a wild-type E. coli cell grown in the absence of lactose. Within minutes after adding lactose to the medium, however, this enzyme appears in quantity within the bacterial cell. When the synthesis of -galactosidase (encoded by the lacZ, or z gene) is induced, the production of two additional enzymes is also induced: -galactoside permease (encoded by the lacY, or y gene) and -galactoside acetyltransferase (encoded by the lacA, or a gene). The permease is involved in transporting lactose into the cell. The transferase is believed to protect the cell from the buildup of toxic products created by -galactosidase acting on other galactosides. By acetylating galactosides other than lactose, the transferase prevents -galactosidase from cleaving them.
THE OPERON MODEL
Not all of the proteins prokaryotes can produce are needed in all circumstances in the same quantities. For example, some metabolites, such as sugars, which the cell breaks down for energy and as a carbon source, may not always be present in the cell s environment. If a given metabolite is not present, enzymes for its breakdown are not useful, and synthesizing these enzymes is wasteful. If the cell produces enzymes for the degradation of a particular carbon source only when this carbon source is present in the environment, the enzyme system is known as an inducible system. Inducible enzymes are synthesized when the environment includes a substrate for those enzymes. The enzymes will then catabolize (break down) the substrate. On the other hand, the enzymes in many synthetic pathways are in low concentration or absent when an adequate quantity of the end product of the pathway is already available to the cell. That is, if the cell encounters an abundance of the amino acid tryptophan in the envi-
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