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               34                                                                              Translation of RNA to Protein


                                                                 B.  Prokaryotic mRNA
                                                                 In organisms that do not have a nucleus (prokaryotes), pre-
                                                                 mRNA usually undergoes little or no modification so that
                                                                 pre-mRNAandmRNAareverysimilarifnotidentical.Be-
                                                                 cause pre-mRNA is colinear with DNA, DNA and proteins
                                                                 are usually colinear in these organisms. Gene expression
                                                                 in prokaryotes usually involves the co-transcription of sev-
                                                                 eral adjacent genes, and translation of mRNA sequences

                                                                 into polypeptides may begin at the 5 end of mRNA while

                                                                 transcription is still in progress at the 3 end.

                                                                 C. Eukaryotic mRNA
                                                                 In cells with a nucleus (eukaryotes) the genetic informa-
                                                                 tion is stored mainly in the nucleus and to a minor degree
               FIGURE 2  Structure of the peptide bond. (a) Two L-amino acids  in some organelles (mitochondria and chloroplasts). The
               with different side chains, R 1  and R 2 ; (b) A dipeptide formed from  description that follows pertains only to nuclear genes.
               the two amino acids shown in (a). [From Cox, R. A., and Arnstein,  Eukaryotic genes are more complicated than prokaryotic
               H. R. V. (1995). In “Encyclopedia of Molecular Biology and Mole-
               cular Medicine” (R. A. Meyers, ed.), Volume 6, pp. 108–125. VCH  genes because the coding region is often discontinuous:
               Publishers, New York. With permission.]           the coding sequences or exons are interrupted by inter-
                                                                 vening sequences (introns). Thus, genes and proteins are
                                                                 usually not colinear in eukaryotes. In the nucleus, a com-
               involves base-pairing between three bases (designated a  plicated set of splicing reactions removes all the introns
               codon) in the mRNA and the three-base anticodon of a  from pre-mRNA and fuses the exons into a continuous
               transfer RNA (tRNA). In a separate reaction, each tRNA  coding sequence. Other processing steps involve adding a
               is first linked to a particular amino acid and thus the pairing  “cap” to the 5 end of the mRNA and a poly(A) “tail” to

               of mRNA with tRNA determines the sequence of amino  the 3 end. After completion of these nuclear maturation

               acids in the resulting protein.                   steps the mRNA is transported to the cytoplasm where it is





















                      FIGURE 3  Structure of typical mRNAs. (a) Prokaryotic mRNA. At the beginning of the transcript there are two ad-

                      ditional phosphate residues linked by pyrophosphate bonds to the 5 phosphate group of the terminal nucleotide

                      (see Fig. 1). The 5 untranslated sequence often contains a ribosome binding site (the Shine–Dalgarno sequence)
                      which increases the efficiency of translation. AUG and UAA are representative initiation and termination codons,
                      respectively. Cistrons (gene sequences) are separated by short (typically 12 to 24 nucleotides) noncoding inter-

                      cistronic sequences. (b) Eukaryotic mRNA. Typically, the processed transcripts are monocistronic. The 5 end is
                      usually modified by the addition of 7-MeG, known as a cap structure, which is linked by two pyrophosphate groups


                      to the terminal nucleotide of mRNA. The coding sequence is located between 5 and 3 untranslated sequences.

                      In the majority of cases the 3 end is modified by the addition of 25 to 250 adenylate residues, termed the poly(A)
                      tail. [From Arnstein, H. R. V., and Cox, R. A. (1992). “Protein Biosynthesis,” Oxford University Press, Oxford. With
                      permission.]
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