THE ARABINOSE OPERON: GENES INVOLVED IN ARA OPERON EXPRESSION
The ara operon contains three structural genes: araB, araA, and araD. Because the cell would prefer to utilize glucose instead of arabinose as an energy source, the ara operon contains a CAP-binding site like the one discussed earlier. The operon also contains an inducerbinding site (araI) and an operator-binding site (araO). The organization of the ara operon is shown in figure 16.18. Transcription of the ara operon is controlled by a regulatory protein that is encoded by the araC gene, which is not part of the ara operon.
THE ARABINOSE OPERON: THE DUAL ACTION OF THE ARAC PROTEIN
The AraC protein can bind to the araI site and also araO under specific conditions. When arabinose is present in the medium, it is bound by the AraC protein. The AraC–arabinose complex binds to the araI site. When the CAP–cAMP complex is also bound to the promoter region, transcription of the ara operon is induced (fig. 16.19 right). Because binding of the AraC protein to the inducer site activates transcription of the ara operon, the AraC protein functions as a positive regulator.
When arabinose is not present, the cell has no need to synthesize enzymes involved in catabolizing this sugar. The AraC protein, in the absence of arabinose, still binds to the araI sequences. Thus, binding of the inducer (arabinose) does not affect the ability of the AraC protein to bond araI. However, in the absence of the bound inducer, the AraC protein can also physically interact with other AraC proteins and assist in their binding to araO. Thus, in the absence of bound arabinose, the AraC proteins at araI and araO interact with each other to bend the DNA into a tight conformational loop, which suppresses transcription of the ara operon (fig. 16.19 left).
Although the lac operon contains one protein that is a positive regulator and a second protein that is a negative regulator, the ara operon contains a single protein that exhibits both positive and negative control. In the presence of arabinose, the AraC protein binds the sugar and the AraC–arabinose complex binds only the inducer sequence to activate transcription. The AraC–arabinose complex cannot bind the operator sequence. In the absence of arabinose, the AraC protein binds to both the inducer and operator sequences, and the interaction between the proteins in these two regions forms a loop that negatively regulates ara operon transcription.