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              Translation of RNA to Protein                                                                45

































                                                                FIGURE 10 Model of the E. coli ribosome, based on low-
                                                                resolution electron microscopy. The diagram shows the relative
                                                                orientation of the large and small subunits and other functional
              FIGURE 9 Electron micrograph of a thin section through two
                                                                components involved in polypeptide chain synthesis. H, head of
              neighboring epithelial cells showing endoplasmic reticulum (large
                                                                the small subunit; CP, central protuberance and L7/L12 stalk of
              arrows) to which are attached numerous polysomes. Groups of  the large subunit; EF-Tu, elongation factor required for binding
              free ribosomes (small arrows) occur in the cytoplasm; pm, plasma  aminoacyl-tRNA to the ribosomal A site; EF-G, elongation factor
              membrane. Bar represents 1000 nm. [Micrograph by I. D. J.  required for translocation of the peptidyl-tRNA from the A to the P
              Burdett, National Institute for Medical Research, London, U.K.]  site. The broken line indicates the boundary between the transla-
                                                                tional and exit domains which are involved in peptide bond forma-
                                                                tion and extrusion of the nascent polypeptide chain, respectively.
                                                                [From Cox, R. A., and Arnstein, H. R. V. (1997). In “Encyclope-
                The higher resolution studies of subunits have also re-  dia of Molecular Biology and Molecular Medicine” (R. A. Meyers,
              vealed the mode of action of several antibiotics such as  ed.), Volume 6, pp. 108–125. VCH Publishers, New York. With
              paromomycin, streptomycin, and spectinomycin, which  permission.]
              modify the decoding function of the small subunit, and
              chloramphenicol, puromycin, and vernamycin, which af-
              fect peptide bond formation.                      A. mRNA Stability
                                                                Provided all other components of the translational systems
                                                                are present in optimum amounts, control of translation
              V. TRANSLATIONAL CONTROL                          may be achieved through the availability of the relevant
                OF GENE EXPRESSION                              messenger RNAs at the site of protein synthesis. The
                                                                steady-state level of mRNA is determined by the rate of
              Cells needs to synthesize specific proteins in the required  its synthesis and degradation. Control of transcription is
              amounts at particular times and to deliver them to the cor-  of major importance for synthesis in both prokaryotes and
              rect locations. These processes depend on a great many  eukaryotes. The stability of mRNA depends on its pri-
              interactions and numerous mechanisms exist for the trans-  mary and secondary structure as well as on the presence
              lational control of gene expression. Examples of the ways  of factors such as stabilizing proteins and nucleases.
              used by cells to control the translation of mRNA are pre-  The secondary structure of mRNA is determined by its
              sented in the sections that follow. This material is intended  nucleotide sequence. For any mRNA a number of coding
              to be illustrative rather than comprehensive, and it is to be  sequences are possible because of the degeneracy of the
              expected that novel control mechanisms will continue to  genetic code. Thus, degeneracy allows for particular fea-
              be discovered.                                    tures of secondary structure (often termed cis factors) to
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