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In the trp operon, an attenuator region lies between the operator and the rst structural gene ( g. 14.13). The messenger RNA transcribed from the attenuator region, termed the leader transcript, has been sequenced, re-
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vealing two surprising and interesting facts. First, four subregions of the messenger RNA have base sequences that are complementary to each other so that three different stem-loop structures can form in the messenger RNA ( g. 14.14). Depending on circumstances, regions 1 2 and 3 4 can form two stem-loop structures, or region 2 3 can form a single stem-loop. When one stemloop structure is formed, the others are preempted. As
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Attenuator region of the trp operon, which contains the leader peptide gene (red). This region is transcribed into the leader transcript.
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Figure 14.14 Nucleotide sequence of part of the leader transcript of the trp attenuator region (bases 50 to 140). Stem-loops 1 2 and 3 4, or stem-loop 2 3, can form because of complementarity of the nucleotides. All possible base pairings are shown in the middle of the gure.
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(From D. L. Oxender, et al., Attenuation in the Escherichia coli tryptophan operon: Role of RNA secondary structure involving the tryptophan codon region, Proceedings of the National Academy of Sciences, 76:5524 28, 1979. Reprinted by permission.)
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III. Molecular Genetics
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14. Gene Expression: Control in Prokaryotes and Phages
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Fourteen
Gene Expression: Control in Prokaryotes and Phages
we will see, the particular combination of stem-loop structures determines whether transcription continues.
Leader Peptide Gene
The second fact obtained by sequencing the leader transcript is that there is a small gene coding information for a peptide from bases 27 to 68 ( g. 14.15). The gene for this peptide is referred to as the leader peptide gene. It codes for fourteen amino acids, including two adjacent tryptophans. These adjacent tryptophan codons are critically important in attenuator regulation. The proposed mechanism for this regulation follows.
Excess Tryptophan
Assuming that the operator site is available to RNA polymerase, transcription of the attenuator region will begin. As soon as the 5 end of the messenger RNA for the leader peptide gene has been transcribed, a ribosome attaches and begins translating this messenger RNA. Depending on the levels of amino acids in the cell,
three different outcomes can take place. If the concentration of tryptophan in the cell is such that abundant tryptophanyl-tRNAs exist, translation proceeds down the leader peptide gene. The moving ribosome overlaps regions 1 and 2 of the transcript and allows stem-loop 3 4 to form, as shown in the con guration at the far left of gure 14.16. This stem-loop structure, referred to as the terminator, or attenuator, stem, causes transcription to be terminated. Note that stem-loop 3 4, the terminator stem, followed by a series of uracil-containing bases, is a rho-independent transcription terminator (see chapter 10). Hence, when existing quantities of tryptophan, in the form of tryptophanyl-tRNA, are adequate for translation of the leader peptide gene, transcription is terminated.
Tryptophan Starvation
If the quantity of tryptophanyl-tRNA is lowered, the ribosome must wait at the rst tryptophan codon until it acquires a Trp-tRNATrp. This is shown in the con guration in the middle part of gure 14.16. The stalled ribosome will
Base sequence of the trp leader transcript and the amino acids these nucleotides code. Note the presence of adjacent tryptophan codons. (From D. L. Oxender, et al., Attenuation in the Escherichia coli tryptophan operon: Role of RNA secondary structure
involving the tryptophan codon region, Proceedings of the National Academy of Sciences, 76:5524 28, 1979. Reprinted by permission.)
Figure 14.16 Model for attenuation in the E. coli trp operon. The circle represents the ribosome attempting to translate the leader transcript of gure 14.14. Under conditions of excess tryptophan, the 3 4 stem-loop forms (the terminator stem), terminating transcription. Under conditions of tryptophan starvation, the ribosome is stalled, and stem-loop 2 3 forms, allowing continued transcription. Under general starvation, there is no translation, resulting in the formation of stem-loops 1 2 and 3 4, which again results in the termination of transcription. (From D. L. Oxender, et al., Attenuation in the Escherichia coli tryptophan operon: Role of RNA secondary
structure involving the tryptophan codon region, Proceedings of the National Academy of Sciences, 76:5524 28, 1979. Reprinted by permission.)
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