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C A U C A U C A U C A U mRNA His (a) His His His
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C A U C A U C A U C A U C mRNA Ile (b) Figure 11.8 Ile Ile Ile
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Translation Initiation
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The subunits of the ribosome (30S and 50S) usually dissociate from each other when not involved in translation. To begin translation, an initiation complex forms, consisting of the following components in prokaryotes: the 30S subunit of the ribosome, a messenger RNA, the Met charged N-formyl methionine tRNA (fMET-tRNAf ), and
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(a) In the normal reading of the messenger RNA, these codons are read as repeats of CAU, coding for histidine. (b) A shift in the reading frame of the messenger RNA causes the codons to be read as AUC repeats coding for isoleucine.
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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11. Gene Expression: Translation
The McGraw Hill Companies, 2001
Information Transfer
3 A C C A C C G A U G U C C C G U C G G A C A C X G7 G G G U A G A A U
5 G G C U A C 3
A G C
Met tRNA f
A C G C U A A G A U A D G C U C C GO G C G A G G C A D D A C C C A A A U C C G G C C U A C G G C U G A C T C C U U G7 A C G A A G C C C A A U
5 C G C G G G G U G A G A G C C U C G G U C G G G CO U A C A D G G C C U G
Anticodon
D T G7 GO CO X Figure 11.10
Odd bases = Dihydrouridine = Pseudouridine = Ribothymidine = 7-Methylguanylic acid = 2 -O -methylguanylic acid = 2 -O -methylcytidylic acid = 3-(3-Amino-3-carboxypropyl) uridine
(b) tRNA m
Anticodon
The two tRNAs for methionine in E. coli. (a) The initiator tRNA. (b) The interior tRNA.
three initiation factors (IF1, IF2, IF3). Initiation factors (as well as elongation and termination factors) are proteins loosely associated with the ribosome.They were discovered when ribosomes were isolated and then washed, losing the ability to perform protein synthesis. The components that form the initiation complex interact in a series of steps. It is known that IF3 binds to the 30S ribosomal subunit, allowing the 30S subunit to bind to messenger RNA ( g. 11.11, step 1). Meanwhile, a complex forms with IF2, the charged N-formyl methionine Met tRNA (fMET-tRNAf ) and GTP (guanosine triphosphate; g. 11.11, step 2). It is IF2 that brings the initiator transfer RNA to the ribosome. IF2 binds only to the charged initiator transfer RNA, and, without IF2, the initiator transfer RNA cannot bind to the ribosome. The nal step in
initiation-complex formation is bringing together the rst two components ( g. 11.11, step 3). The hydrolysis of GTP to GDP Pi (inorganic phosphate, PO4 3 see g. 9.8) produces conformational changes; these changes allow the initiation complex to join the 50S ribosomal subunit to form the complete ribosome and then allow the initiation factors and GDP to be released. Frequently, the hydrolysis of a nucleoside triphosphate (e.g., ATP, GTP) in a cell occurs to release the energy in the phosphate bonds for use in a metabolic process. However, in the process of translation, the hydrolysis apparently changes the shape of the GTP so that it and the initiation factors can be released from the ribosome after the 70S particle has been formed.Thus, hydrolysis of GTP in translation is for conformational
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
11. Gene Expression: Translation
The McGraw Hill Companies, 2001
Eleven
Gene Expression: Translation
Step 1
30S Ribosome subunit mRNA
fMet Step 2 IF2 + GTP +
fMet-tRNA Met f
Step 3
Combine end products of step 1 and step 2
50S GDP+ Pi 50S + Initiation complex GTP
IF2 IF3 Complete 70S ribosome
The prokaryotic 70S ribosome forms in a three-step process. In the rst step, the 30S ribosomal subunit and the mRNA combine. In the second step, the initiator tRNA combines with IF2. In the nal step, the components from steps 1 and 2 combine to form the initiation complex, followed by the formation of the 70S ribosome.
change rather than covalent bond formation. IF1 helps the other two initiation factors bind to the 30S ribosomal subunit or stabilizes the 30S initiation complex. The process in eukaryotes is generally similar, but more complex.The eukaryotic initiation factor abbreviations are preceded by an e to denote that they are eukaryotic (eIF1, eIF2, etc.). At least eleven initiation factors are involved, including a specific cap-binding protein, eIF4E.
The ribosome apparently recognizes the prokaryotic messenger RNA through complementarity of a region at the 3 end of the 16S ribosomal RNA and a region slightly upstream from the initiation sequence (AUG) on the messenger RNA.This idea, the Shine-Dalgarno hypothesis ( g. 11.12), is named after the people who rst suggested it. The sequence (AGGAGGU) of complementarity between the messenger RNA and the 16S ribosomal RNA is referred to as the Shine-Dalgarno sequence. Although
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