codon; at the other extreme, leucine, serine, and arginine are each specified by six different codons. The different codons for a given amino acid are said to be synonymous. The code itself is termed degenerate, meaning that more than one codon can specify the same amino acid.

Synthesis of all polypeptide chains in prokaryotic and eu-karyotic cells begins with the amino acid methionine. In most mRNAs, the start (initiator) codon specifying this amino-terminal methionine is AUG. In a few bacterial mRNAs, GUG is used as the initiator codon, and CUG occasionally is used as an initiator codon for methionine in eukaryotes. The three codons UAA, UGA, and UAG do not specify amino acids but constitute stop (termination) codons that mark the carboxyl terminus of polypeptide chains in almost all cells. The sequence of codons that runs from a specific start codon to a stop codon is called a reading frame. This precise linear array of ribonucleotides in groups of three in mRNA specifies the precise linear sequence of amino acids in a polypeptide chain and also signals where synthesis of the chain starts and stops.

Because the genetic code is a comma-less, non-overlapping triplet code, a particular mRNA theoretically could be translated in three different reading frames. Indeed some mRNAs have been shown to contain overlapping information that can be translated in different reading frames, yielding different polypeptides (Figure 4-20). The vast majority of mRNAs, however, can be read in only one frame because stop codons encountered in the other two possible reading frames terminate translation before a functional protein is produced. Another unusual coding arrangement occurs because of frame-

Frame 1

Frame 1

Frame 2

5' —g|cuu| |guu| |uac| |gaa| |uua|— —I Leu H Val H Tyr H Glu H Leu I-

▲ FIGURE 4-20 Example of how the genetic code—a non-overlapping, comma-less triplet code—can be read in different frames. If translation of the mRNA sequence shown begins at two different upstream start sites (not shown), then two overlapping reading frames are possible. In this example, the codons are shifted one base to the right in the lower frame. As a result, the same nucleotide sequence specifies different amino acids during translation. Although they are rare, many instances of such overlaps have been discovered in viral and cellular genes of prokaryotes and eukaryotes. It is theoretically possible for the mRNA to have a third reading frame.

shifting. In this case the protein-synthesizing machinery may read four nucleotides as one amino acid and then continue reading triplets, or it may back up one base and read all succeeding triplets in the new frame until termination of the chain occurs. These frameshifts are not common events, but a few dozen such instances are known.

The meaning of each codon is the same in most known organisms—a strong argument that life on earth evolved only once. However, the genetic code has been found to differ for a few codons in many mitochondria, in ciliated protozoans, and in Acetabularia, a single-celled plant. As shown in Table 4-2, most of these changes involve reading of normal stop codons as amino acids, not an exchange of one amino acid for another. These exceptions to the general code probably were later evolutionary developments; that is, at no single time was the code immutably fixed, although massive changes were not tolerated once a general code began to function early in evolution.

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