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               36                                                                              Translation of RNA to Protein





























                                                                 FIGURE  6  Schematic  illustration  of  base  pairing  between  a
                                                                 codon and its anticodon. The diagram shows the interaction at the
                                                                 P site of the ribosome (see Fig. 9) between the initiation codon
                                                                 AUG of mRNA and the anticodon CAU of fMet-tRNA Met  and the in-
                                                                                                     f
                                                                 teraction at the A site between the codon GUA of mRNA and the
                                                                 anticodon UAC of Val-tRNA Val . The polarity of an RNA species


                                                                 runs from the 5 end to the 3 end. The fragment of tRNA is rep-
                                                                 resentative of the general structure (see Fig. 5b) placed in the
                                                                 appropriate orientation; the numbers refer to the nucleotide po-

                                                                 sitions measured from the 5 end. The interaction between the
                                                                 codon and the anticodon is antiparallel, and the three base pairs
                                                                 have a bihelical conformation.
                                                                 of the codon form conventional base pairs with the third
                                                                 and middle bases of the anticodon, respectively, but the
                                                                 third base of the codon pairs with the first base of the anti-
                                                                 codon by a less stringent interaction (e.g., base-pairing
                                                                 of G with U as well as with C), giving rise to degen-
                                                                 eracy. This so-called “wobble” considerably reduces the
                                                                 number of tRNA species required to decode the 61 sense
                                                                 codons. Thus, the protein synthesis system in the cytosol
                                                                 of eukaryotes contains only a few more than 40 different
                                                                 tRNAs, and in mitochondria 22 to 24 tRNA species are
                                                                 sufficient.
               FIGURE  5  Structure  of  phenylalanyl-transfer  RNA.  (a)  Sec-  TheattachmentofaminoacidstotRNAinvolvesthefor-


               ondary cloverleaf structure. The 5 and 3 ends of the molecule  mation of an ester bond between the alpha-carboxyl group
               are marked; the continuous line represents the sugar phosphate  of the amino acid and the 3 -hydroxyl group of the termi-

               backbone. The short lines denote base residues and the dots de-
                                                                 nal adenosine of tRNA. It requires specific enzymes, the
               note base-pairing through standard hydrogen bonds (see Fig. 1).
                                                                 aminoacyl-tRNA synthetases. There are 20 different syn-
               The D loop contains dihydrouracil residues; the T C loop contains
                                                                 thetases, each specific for one of the 20 amino acids, and
               thymine and pseudouridine. (b) Tertiary structure. The abbrevia-
               tions for unusual bases are defined in (a). The sugar phosphate  each enzyme recognizes something unique in the structure
               backbone is represented by double parallel lines. Standard base  of its cognate tRNA. The structural determinants which
               pairs  are  represented  by  short  double  lines,  and  nonstandard  ensure accuracy of this charging reaction vary for different
               base pairs by single lines. The anticodon sequence is stippled,
               and the acceptor end is shaded. [From Arnstein, H. R. V., and Cox,  tRNAs. The anticodon may play a part but sometimes even
               R. A. (1992). “Protein Biosynthesis,” Oxford University Press, Ox-  a single base elsewhere is sufficient to determine the speci-
               ford. With permission.]                           ficity of the tRNA-synthetase interaction. The accuracy of
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