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5. Gene Expression

How Is Gene Expression Increased or Decreased in Response to Environmental Change?

In prokaryotes, regulatory proteins are often controlled by nutrient availability. This allows organisms such as bacteria to rapidly adjust their transcription patterns in response to environmental conditions. In addition, regulatory sites on prokaryotic DNA are typically located close to transcription promoter sites — and this plays an important part in gene expression.

A three-part schematic shows how a repressor protein can inhibit transcription by preventing RNA polymerase from binding DNA. Part 1 shows the layout of a linear region of DNA. The operator is represented by colored shading on the DNA molecule and spans three nucleotides. The site of transcription is shaded a different color, and an arrow points from left to right above the shading to show the direction transcription proceeds. Part 2 shows the positions of an inactive repressor protein and RNA polymerase relative to a DNA molecule when transcription is occurring. Part 3 shows the positions of an active repressor protein and RNA polymerase in relation to a DNA molecule when transcription is repressed.
Transcription repression near the promoter region.

Molecules can interfere with RNA polymerase binding. An inactive repressor protein (blue) can become activated by another molecule (red circle). This active repressor can bind to a region near the promoter called an operator (yellow) and thus interfere with RNA polymerase binding to the promoter, effectively preventing transcription.

For an example of how this works, imagine a bacterium with a surplus of amino acids that signal the turning “on” of some genes and the turning “off” of others. In this particular example, cells might want to turn “on” genes for proteins that metabolize amino acids and turn “off” genes for proteins that synthesize amino acids. Some of these amino acids would bind to positive regulatory proteins called activators. Activator proteins bind to regulatory sites on DNA nearby to promoter regions that act as on/off switches. This binding facilitates RNA polymerase activity and transcription of nearby genes. At the same time, however, other amino acids would bind to negative regulatory proteins called repressors, which in turn bind to regulatory sites in the DNA that effectively block RNA polymerase binding (Figure 3).

The control of gene expression in eukaryotes is more complex than that in prokaryotes. In general, a greater number of regulatory proteins are involved, and regulatory binding sites may be located quite far from transcription promoter sites. Also, eukaryotic gene expression is usually regulated by a combination of several regulatory proteins acting together, which allows for greater flexibility in the control of gene expression.

A schematic shows three transcriptional regulator proteins on a DNA molecule. The DNA molecule is folded in on itself to form loops and each regulator protein is bound to the apex of a DNA loop and interacting with a single mediator protein bound to RNA polymerase. RNA polymerase is in turn bound to a region of DNA between the promoter sequence and the site of transcription.
 The complexity of multiple regulators

Transcriptional regulators can each have a different role. Combinations of one, two, or three regulators (blue, green, and yellow shapes) can affect transcription in different ways by differentially affecting a mediator complex (orange), which is also composed of proteins. The effect is that the same gene can be transcribed in multiple ways, depending on the combination, presence, or absence of various transcriptional regulator proteins.

As previously mentioned, enhancer sequences are DNA sequences that are bound by an activator protein, and they can be located thousands of base pairs away from a promoter, either upstream or downstream from a gene. Activator protein binding is thought to cause DNA to loop out, bringing the activator protein into physical proximity with RNA polymerase and the other proteins in the complex that promote the initiation of transcription (Figure 4).

Different cell types express characteristic sets of transcriptional regulators. In fact, as multicellular organisms develop, different sets of cells within these organisms turn specific combinations of regulators on and off. Such developmental patterns are responsible for the variety of cell types present in the mature organism.

Transcriptional regulators can determine cell types

The wide variety of cell types in a single organism can depend on different transcription factor activity in each cell type. Different transcription factors can turn on at different times during successive generations of cells. As cells mature and go through different stages (arrows), transcription factors (colored balls) can act on gene expression and change the cell in different ways. This change affects the next generation of cells derived from that cell. In subsequent generations, it is the combination of different transcription factors that can ultimately determine cell type.

Conclusion

To live, cells must be able to respond to changes in their environment. Regulation of the two main steps of protein production — transcription and translation — is critical to this adaptability. Cells can control which genes get transcribed and which transcripts get translated; further, they can biochemically process transcripts and proteins in order to affect their activity. Regulation of transcription and translation occurs in both prokaryotes and eukaryotes, but it is far more complex in eukaryotes.

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