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ATP contains 3 phosphate groups. Adenylyl cyclase removes two of these phosphate groups. The remaining phosphate group is linked into the sugar to make cAMP. Cyclic AMP is made of a ribose sugar with oxygens at both carbons 2 and 3 (the carbons at the bottom of the pentagon). The oxygen bound to carbon 3 is also bound to the phosphorus. Similarly, the oxygen bound at carbon 5 was already bound to the phosphorus. This forms a ring where the phosphorus is linked with an oxygen to both carbons 3 and 5.
When ATP levels decrease due to depletion of glucose, some remaining ATP is converted to cAMP by adenylyl cyclase. Thus, increased cAMP levels signal glucose depletion.
Diagram of the lac operon with and without cAMP. A) In the absence of cAMP, CAP does not bind the promoter. RNA polymerase does bind to the promoter and transcription occurs at a low rate. In the presence of cAMP, CAP binds the promoter and increases RNA polymerase activity. This is shown with a circle labeled cAMP + CAP bound to the promoter. RNA polymerase is also bound to the promoter and a thick arrow indicates faster transcription. B) cAMO-CAP complex stimulates RNA polymerase activity and increases RNA synthesis. However, even in the presence of cAMP-CAP complex, RNA synthesis is blocked when repressor is bound ot he operator. This is shows as the cAMP + CAP circle as well as the RNA polymerase bound to the promoter. The repressor is bound to the operator and this blocks RNA polymerase from moving forward.
(a) In the presence of cAMP, CAP binds to the promoters of operons, like the lac operon, that encode genes for enzymes for the use of alternate substrates. (b) For the lac operon to be expressed, there must be activation by cAMP-CAP as well as removal of the lac repressor from the operator.
Conditions Affecting Transcription of the lac Operon
Glucose CAP binds Lactose Repressor binds Transcription
+ + No
+ + Some
+ + No
+ + Yes
  • What affects the binding of the trp operon repressor to the operator?
  • How and when is the behavior of the lac repressor protein altered?
  • In addition to being repressible, how else is the lac operon regulated?

Global responses of prokaryotes

In prokaryotes, there are also several higher levels of gene regulation that have the ability to control the transcription of many related operons simultaneously in response to an environmental signal. A group of operons all controlled simultaneously is called a regulon .

Alarmones

When sensing impending stress, prokaryotes alter the expression of a wide variety of operon s to respond in coordination. They do this through the production of alarmones , which are small intracellular nucleotide derivatives. Alarmones change which genes are expressed and stimulate the expression of specific stress-response genes. The use of alarmones to alter gene expression in response to stress appears to be important in pathogenic bacteria. On encountering host defense mechanisms and other harsh conditions during infection, many operons encoding virulence genes are upregulated in response to alarmone signaling. Knowledge of these responses is key to being able to fully understand the infection process of many pathogens and to the development of therapies to counter this process.

Alternate σ factors

Since the σ subunit of bacterial RNA polymerase confers specificity as to which promoters should be transcribed, altering the σ factor used is another way for bacteria to quickly and globally change what regulons are transcribed at a given time. The σ factor recognizes sequences within a bacterial promoter , so different σ factors will each recognize slightly different promoter sequences. In this way, when the cell senses specific environmental conditions, it may respond by changing which σ factor it expresses, degrading the old one and producing a new one to transcribe the operons encoding genes whose products will be useful under the new environmental condition. For example, in sporulating bacteria of the genera Bacillus and Clostridium (which include many pathogens), a group of σ factors controls the expression of the many genes needed for sporulation in response to sporulation-stimulating signals.

  • What is the name given to a collection of operons that can be regulated as a group?
  • What type of stimulus would trigger the transcription of a different σ factor?

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Source:  OpenStax, Microbiology. OpenStax CNX. Nov 01, 2016 Download for free at http://cnx.org/content/col12087/1.4
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