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               48                                                                              Translation of RNA to Protein




















































                      FIGURE 12  Translational control of bacteriophage synthesis. (a) Arrangements of MS2 and f2 bacteriophage cistrons.
                      (b) Arrangement of Q β  bacteriophage cistrons. (c) Replicative intermediate synthesizing new plus strands on the
                      complementary minus strand copy of the original bacteriophage RNA. The ribosome binding sites (RBS) are indicated
                      by the numbered arrows. The major RBS (1) binds ribosomes efficiently but can be blocked by ribosomal protein S1.
                      The secondary RBS (2) becomes available only after translation of at least part of the coat protein cistron by ribosomes.
                      RBS 3 is masked by the secondary structure of native bacteriophage RNA but is accessible in nascent RNA (c) or
                      in vitro when the secondary structure is destroyed. Noncoding regions are shown in black upstream from the RBS
                      sites and at the 3-end. [From Arnstein, H. R. V., and Cox, R. A. (1992). “Protein Synthesis,” Oxford University Press,
                      Oxford. With permission.]
               RNA-dependent DNA polymerase. Other example of the  ences are due to the differential and independent initiation
               operation of such a frameshift include the synthesis of  of translation at each cistron as a result of differences in
               the reverse transcriptase enzymes of several retrotrans-  the secondary structure of the mRNA initiation sites. Fur-
               posons, such as the yeast Ty1. The mechanism of chang-  thermore, in vivo there is a delay in the synthesis of coat
               ing the reading frame involves “slippery” sequences and  protein and this temporal control involves translational
               a complex folding of the mRNA into a structure termed a  repression of the cistron by ribosomal protein S1. In ad-
               pseudoknot.                                       dition, S1 functions as one of the subunits of the f2 RNA
                 In E. coli, translational control of the three cistrons of  replicase; therefore, association of the newly synthesized
               Q β ,f2,andrelatedbacteriophageRNAs(Fig.12)accounts  translation product of the f2 replicase cistron with S1 will
               for the synthesis of coat protein:replicase:A protein in  favor its dissociation from the phage RNA, thus allowing
               the approximate ratio of 20:5:1. These quantitative differ-  translation of the coat protein cistron to start.
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