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Table 14.3 Relationship Between N-Terminal
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Amino Acid and Half-Life of E. coli -Galactosidase Proteins with Modi ed N-Terminal Amino Acids
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N-Terminal Amino Acid Met, Ser, Ala, Thr, Val, Gly Ile, Glu Tyr, Gln Pro Phe, Leu, Asp, Lys Arg
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Source: Data from Bachmair, et al., Science, 234:179 86, 1986.
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Protein Degradation
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A nal control affecting the amount of gene product in a cell is control of the rate at which proteins degrade. The normal life spans of proteins vary greatly. For example, some proteins last longer than a cell cycle, whereas others may be broken down in minutes. Several models have been suggested for control of protein degradation, including the N-end rule and the PEST hypothesis. According to the N-end rule, the amino acid at the amino-, or N-terminal, end of a protein is a signal to proteases that control the average length of life of a protein.
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Half-Life 20 hours 30 minutes 10 minutes 7 minutes 3 minutes 2 minutes
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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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The McGraw Hill Companies, 2001
Fourteen
Gene Expression: Control in Prokaryotes and Phages
Aspartate transcarbamylase. Left: the enzyme bound with cytidine triphosphate (CTP). Right: the enzyme without CTP. Notice the difference in shape with and without CTP. With CTP, the enzyme literally closes up. (Courtesy of Evan R. Kantrowitz.)
S U M M A R Y
STUDY OBJECTIVE 1: To study the way in which in-
ducible and repressible operons work 406 414
Most bacterial genes are organized into operons, which can either be repressed or induced. Transcription begins in inducible operons, such as lac, when the metabolite that the operon enzymes act upon appears in the environment. The metabolite (or a derivative), the inducer, combines with the repressor (the product of the independent regulator gene) and renders the repressor nonfunctional. In the absence of the inducer, the repressor binds to the operator, a segment between the promoter and the rst gene of the operon. When in place, the repressor blocks transcription. After combining with the inducer, the repressor diffuses from the operator, and transcription proceeds. All operons responsible for the breakdown of sugars in E. coli are inducible. In the presence of glucose, other inducible sugar operons (such as the arabinose and galactose operons) are repressed, even if their sugars appear in the environment. This process is called catabolite repression. Cyclic AMP and a catabolite activator protein (CAP) enhance the
transcription of the nonglucose sugar operons. Glucose lowers the level of cyclic AMP in the cell and thus prevents the enhancement of transcription of these other operons. Repressible operons, such as the trp operon in E. coli, have the same basic traits as an inducible operon polycistronic transcription controlled by an operator site between the promoter and the rst structural gene. However, the repressor protein, controlled by an independent regulator gene, is functional in blocking transcription only after it has combined with the corepressor. This corepressor is the end product of the operon s pathway or some form of the end product (tryptophan in the trp operon). STUDY OBJECTIVE 2: To examine attenuator control in
bacteria 415 418
Amino acid-synthesizing operons often have an attenuator region. The ability of a ribosome to translate a leader peptide gene determines the secondary structure of the messenger RNA transcript. If the ribosome can translate the leader peptide gene, there must be adequate quantities of
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
14. Gene Expression: Control in Prokaryotes and Phages
The McGraw Hill Companies, 2001
Solved Problems
the amino acid present, and a terminator stem and loop form in the messenger RNA, causing termination of transcription. In the trp operon of bacilli, a protein that binds tryptophans serves the same purpose. STUDY OBJECTIVE 3: To analyze the control of the life cy-
complex. Their presence can cause inversions or deletions. The agellar phase in Salmonella is controlled by the orientation of a transposon. STUDY OBJECTIVE 5: To look at other transcriptional and
cle of phage
418 424
posttranscriptional mechanisms of control of gene expression in bacteria and phages 430 434
Af nity of early and late operons in phages for different sigma factors are another form of transcriptional control, as seen in phage T4 transcription and the transcription of heat shock proteins. Translational control can be exercised through a gene s position in an operon (genes at the beginning are transcribed most frequently), through redundancy of the genetic code, or through a stringent response that shuts down most transcription during starvation. Posttranslational control is primarily regulated by feedback inhibition.The N-terminal amino acid or particular regions within the proteins program the rate of protein degradation.
Control of gene expression in phage is complex. The decision for lytic versus lysogenic response is determined by competition between two repressors, CI and Cro. STUDY OBJECTIVE 4: To determine the way in which transposable genetic elements transpose and control gene expression in bacteria 425 430 Transposons are mobile genetic elements; copies of them can be inserted at other places in the genome. Their ends are inverted repeats. Upon insertion, they are anked by short, direct repeats. They can be simple (IS elements) or
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