Nirenberg and Leder developed a technique for using ribosomebound tRNAs to provide additional information about the genetic code

tage of this system was that it could be used with very short synthetic mRNA molecules that could be synthesized with a known sequence. Nirenberg and Leder synthesized over 50 short mRNAs with known codons and added them individually to a mixture of ribosomes and tRNAs. They then isolated the ribosome-bound tRNAs and determined which amino acids were present on the bound tRNAs. For example, synthetic RNA with the codon GUU retained a tRNA to which valine was attached, whereas RNAs with the codons UGU and UUG did not. Using this method, Nirenberg and his colleagues were able to determine the amino acids encoded by more than 50 codons.

A third method provided additional information about the genetic code. Gobind Khorana and his colleagues used chemical techniques to synthesize RNA molecules that contained known repeating sequences. They hypothesized that an mRNA that contained, for instance, alternating uracil and guanine nucleotides (UGUG UGUG) would be read during translation as two alternating codons, UGU GUG UGU GUG, producing a protein composed of two alternating amino acids. When Khorana and his colleagues placed this synthetic mRNA in a cellfree protein-synthesizing system, it produced a protein made of alternating cysteine and valine residues. This technique could not determine which of the two codons (UGU or GUG) specified cysteine, but, combined with other methods, it made a crucial contribution to cracking the genetic code. The genetic code was fully understood by 1968 (< Figure 15.12). In the next section, we will examine some of the features of the code, which is so important to modern biology that Francis Crick has compared its place to that of the periodic table of the elements in chemistry.

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Nirenberg

A brief biography of Marshall

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