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A inserted after third A of normal (GCA repeat)
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* C A G C A G C A A G C A G C A
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Frameshift Deletion and insertion combined (return to CAG repeat) * * C G C A G C A A G C A G C A G
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Frameshift Figure 11.28
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Frameshift mutations in a gene result from the addition or deletion of one or several nucleotides (any number other than a multiple of three) in the DNA. The messenger RNA shown here normally has a CAG repeat. A single-base deletion shifts the three-base reading frame to a series of AGC repeats. A later insertion restores the reading frame. Asterisks (*) indicate points of deletion or insertion.
Once geneticists had gured out that the genetic code is in nonoverlapping triplets, they turned their attention to the sixty-four codons.They wondered which amino acid, for example, does ACU specify The work was done in two stages. In the rst stage, M. W. Nirenberg, S. Ochoa, and their colleagues made long arti cial messenger RNAs and determined which amino acids these messenger RNAs incorporated into protein. In the second stage, speci c triplet RNA sequences were synthesized. The researchers then determined the amino acid-transfer RNA complex that was bound by each sequence.
C A G C A G C A G C A G C A G C A G Original mRNA (CAG repeat) * * * C A G G C A G G C A G G C A G C A G
mRNA
Three inserted Gs restore CAG repeat in mRNA
mRNA
Frameshift Figure 11.29
Restoration
The coding frame of CAG repeats is rst shifted and then restored by three additions (insertions). Asterisks (*) indicate insertions.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
11. Gene Expression: Translation
The McGraw Hill Companies, 2001
Eleven
Gene Expression: Translation
They then made synthetic messenger RNAs by using mixtures of the various diphosphate nucleotides in known proportions. Table 11.3 gives an example. From their experiments, it was possible to determine the bases used in many of the code words, but not their speci c order. For example, cysteine, leucine, and valine are all coded by two Us and a G, but the experiment could not sort out the order of these bases (5 -UUG-3 , 5 -UGU-3 , or 5 -GUU-3 ) for any one of them. Determining the order required an extra step in sophistication that is, being able to synthesize known trinucleotides.
Severo Ochoa (1905 1993).
(Courtesy of Dr. Severo Ochoa.)
Marshall W. Nirenberg (1934 ). (Courtesy of Dr.
Marshall W. Nirenberg.)
Synthetic Codons
Once trinucleotides of known composition could be manufactured, Nirenberg and P. Leder in 1964 developed a binding assay. They found that isolated E. coli ribosomes, in the presence of high-molarity magnesium chloNonoverlapping, no punctuation codon C A G C A G C A G C A G 1 2 3 4
Synthetic Messenger RNAs
The ability to synthesize long-chain messenger RNAs resulted from the 1955 discovery of M. Grunberg-Manago and Ochoa of the enzyme polynucleotide phosphorylase, which joins diphosphate nucleotides into longchain, single-stranded polynucleotides. Unlike a polymerase, polynucleotide phosphorylase does not need a primer on which to act.This enzyme is found in all bacteria. (Its main function in the cell is probably the reverse of its use here. It most likely serves as an exonuclease, degrading messenger RNA.) In 1961, Nirenberg and J. H. Matthei added arti cially formed RNA polynucleotides of known composition to an E. coli ribosomal system and looked for the incorporation of amino acids into proteins. The system just described is called a cell-free system, a mixture primarily of the cytoplasmic components of cells, such as E. coli, but missing nucleic acids and membrane components.These systems are relatively easy to create by disrupting and then fractionating whole cells. The systems hold the advantage of containing virtually all the components needed for protein synthesis except the messenger RNAs.Their disadvantages are that they are relatively short-lived (several hours) and are relatively inef cient in translation.However,an added bene t to the E.coli cell-free system is that it will translate, albeit inef ciently, RNAs that normally are not translated in vivo because they lack translation initiation signals.This feature allowed these scientists to use arti cial messenger RNAs that contained no ShineDalgarno sequence for ribosomal binding. Nirenberg and Matthei found that when the enzyme polynucleotide phosphorylase in the E. coli cell-free system made uridine diphosphates into a poly-U messenger RNA, phenylalanine residues were incorporated into a polypeptide. Thus, the rst code word established was UUU for phenylalanine. Nirenberg and Ochoa and their associates continued the work.They found that AAA was the code word for lysine,CCC was the code word for proline, and GGG was the code word for glycine.
Overlapping, one C A G C A G C A G C A G base no punctuation codon 1 codon 2 codon 3 codon 4 codon 5 codon 6
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