General Principles of Gene Regulation

One of the major themes of molecular genetics is the central dogma, which stated that genetic information flows from DNA to RNA to proteins (see Figure 10.17a) and provided a molecular basis for the connection between genotype and phenotype. Although the central dogma brought coherence to early research in molecular genetics, it failed to address a critical issue: How is the flow of information along the molecular pathway regulated?

Consider E. coli, a bacterium that resides in your large intestine. Your eating habits completely determine the nutrients available to this bacteria: it can't seek out nourishment when nutrients are scarce; nor can it move away when confronted with unpleasant changes. E. coli makes up for its inability to alter the external environment by being internally flexible. For example, if glucose is present, E. coli uses it to generate ATP; if there's no glucose, it utilizes lactose, arabinonse, maltose, xylose, or any of a number of other sugars. When amino acids are available, E. coli uses them to synthesize proteins; if a particular amino acid is absent, E. coli produces the enzymes needed to synthesize that amino acid. Thus, E. coli responds to environmental changes by rapidly altering its biochemistry. This biochemical flexibility, however, has a high price. Producing all the enzymes necessary for every environmental condition would be energetically expensive. So how does E. coli maintain biochemical flexibility while optimizing energy efficiency?

The answer is through gene regulation. Bacteria carry the genetic information for many proteins, but only a subset of this genetic information is expressed at any time. When the environment changes, new genes are expressed, and proteins appropriate for the new environment are synthesized. For example, if a carbon source appears in the environment, genes encoding enzymes that take up and metabolize this carbon source are quickly transcribed and translated. When this carbon source disappears, the genes that encode them are shut off. This type of response, the synthesis of an enzyme stimulated by a specific substrate, is called induction.

Multicellular eukaryotic organisms face a different dilemma. Individual cells in a multicellular organism are specialized for particular tasks. The proteins produced by a nerve cell, for example, are quite different from those produced by a white blood cell. The problem that a eukaryotic cell faces is how to specialize. Although they are quite different in shape and function, a nerve cell and a blood cell still carry the same genetic instructions.

A multicellular organism's challenge is to bring about the specialization of cells that have a common set of genetic instructions. This challenge is met through gene regulation: all of an organism's cells carry the same genetic information, but only a subset of genes are expressed in each cell type. Genes needed for other cell types are not expressed. Gene regulation is therefore the key to both unicellular flexibility and multicellular specialization, and it is critical to the success of all living organisms.

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