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III. Molecular Genetics
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11. Gene Expression: Translation
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Folded polypeptide (a) (c)
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Figure 11.27 The change in structure of GroEL with GroES attached explains how the chaperonin can unfold a partially folded polypeptide to allow it to refold in a different way. (a) A space- lling model of GroEL is shown without (left) and with (right) GroES. GroEL s rings are blue and magenta, and GroES is green. (b) The same structures are seen in a cutaway view. (c) This diagram shows how the attachment of GroES causes the top part of the top ring of GroEL to pull apart an improperly folded polypeptide. ([a & b]: Reprinted from Bernd Bakau
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and Arthur L. Horwich, The Hsp70 and Hsp60 Chaperone Machines in Cell,
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vol. 92, 351 366. Copyright 1988, with permission from Elsevier Science.)
THE GENETIC CODE
Researchers in the mid-1950s assumed that the genetic code consisted of simple sequences of nucleotides specifying particular amino acids.They sought answers to questions such as: Is the code overlapping Are there nucleotides between code words (punctuation) How many letters make up a code word (codon) Logic, along with genetic experiments, supplied some of the answers, but only with the rapidly improving techniques of biochemistry did they eventually decode the genetic language.
code would allow for 4 4 4 64 codons, which are more than enough to specify twenty amino acids.
Evidence for the Triplet Nature of the Code
The experimental manipulation of mutant genes, primarily by Francis Crick and his colleagues, reinforced the triplet code concept. In these experiments, a chemical mutagen, the acridine dye pro avin, was used to cause inactivation of the rapid lysis (rIIB) gene of the bacteriophage T4. Pro avin inactivates the gene by either adding or deleting a nucleotide from the DNA (see chapter 12). The rII gene controls the plaque morphology of this bacteriophage growing on E. coli cells. Rapid-lysis mutants produce large plaques; the wild-type form of the gene, rII , results in normal plaque morphology. Figure 11.28 shows the consequences of adding or deleting a nucleotide. From the point of addition or deletion onward, a frameshift causes codons to be read in different groups of three. If a deletion is combined with an addition to produce a double-mutant gene, the frameshift occurs only in the region between the two mu-
Triplet Nature of the Code
Several lines of evidence seemed to indicate that the nature of the code was triplet (three bases in messenger RNA specifying one amino acid).If codons contained only one base, they would only be able to specify four amino acids since there are only four different bases in DNA (or messenger RNA).A couplet code would have 4 4 16 two-base words, or codons, which is still not enough to specify uniquely twenty different amino acids. A triplet
Tamarin: Principles of Genetics, Seventh Edition
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The Genetic Code
tants. If this region is small enough or does not contain coding for vital amino acids, the function of the gene may be restored. Two deletions or two insertions combined will not restore the reading frame. However, Crick and his colleagues found that the combination of three additions or three deletions did restore gene function.This nding led to the conclusion that the genetic code was triplet,beNormal (CAG repeat) C A G C A G C A G C A G C A G First A deleted (AGC repeat) * C G C A G C A G C A G C A G C mRNA
cause a triplet code would be put back into the reading frame by three additions or three deletions ( g. 11.29).
Overlap and Punctuation in the Code
Questions still remained: was the code overlapping Did it have punctuation Several logical arguments favored a nopunctuation, nonoverlapping model ( g. 11.30). An overlapping code would be subject to two restrictions. First, a change in one base (a mutation) could affect more than one codon and thus affect more than one amino acid. But studies of amino acid sequences almost always showed that only one amino acid was changed, which argued against codon overlap. Second, certain restrictions affected which amino acids occurred next to each other in proteins. For example, the amino acid UUU coded could never be adjacent to the amino acid coded by AAA because one or both (depending on the number of bases overlapped) of the overlap codons UUA and UAA would always insert other amino acids between them. Overlap, then, seemed to be ruled out since every amino acid appears next to every other amino acid in one protein or another. Punctuation between codons was also tentatively ruled out. The messenger RNA in the tobacco necrosis satellite virus has just about enough codons to specify its coat protein with no room left for a punctuating base or bases between each codon.
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