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280    Chapter 8    Gene Expression: The Flow of Information from DNA to RNA to Protein


              The Genetic Code Is Almost,                          broad outlines of information flow between gene and
              but Not Quite, Universal                             protein, these results did not explain exactly how the cel-
                                                                   lular machinery accomplishes gene expression. This
              We now know that virtually all cells alive today use the   question is our focus as we present in the next sections
              same basic genetic code. One early indication of this uni-  the details of transcription and translation.
              formity was that a translational system derived from one
              organism could use the mRNA from another organism to
              convert genetic information to the encoded protein. Rabbit   essential concepts
              hemoglobin mRNA, for example, when injected into frog
              eggs or added to cell-free extracts from wheat germ, di-  •  The nearly universal genetic code consists of 64 codons,
              rects the synthesis of rabbit hemoglobin proteins. More   each one composed of three nucleotides. Sixty-one
                                                                       codons specify amino acids, while three—UAA, UAG,
              recently, comparisons of DNA and protein sequences have   UGA—are stop codons. The code is degenerate in that
              revealed a perfect correspondence according to the genetic   more than one codon can specify one amino acid.
              code between codons and amino acids in almost all organ-  •  The codon AUG specifies methionine; it also serves
              isms examined.                                           as the initiation codon establishing the reading frame
                                                                       that groups nucleotides into successive, nonoverlapping
                                                                       codon triplets.
              Conservation of the genetic code
              The universality of the code is an  indication that it   •  Missense mutations change a codon so that it specifies
                                                                       a different amino acid; frameshift mutations alter the
              evolved very early in the history of life. Once it emerged,   reading frame for all codons following the mutation; and
              the code remained constant over billions of years, in part   nonsense mutations change a codon for an amino acid
              because evolving organisms would have little tolerance   into a stop codon.
              for change. A single change in the genetic code could
              disrupt the production of hundreds or thousands of pro-
              teins in a cell—from the DNA polymerase essential for
              replication to the RNA polymerase required for gene ex-
              pression to the tubulin proteins that compose the mitotic
              spindle—and such a change would therefore be lethal.  8.2   Transcription:
                                                                   From DNA to RNA

              Exceptional genetic codes
              Researchers were thus quite amazed to observe a few ex-  learning objectives
              ceptions to the universality of the code. In some species of
              the single-celled eukaryotic protozoans known as ciliates,   1.  Describe the three stages of transcription: initiation,
              the codons UAA and UAG, which are nonsense codons in      elongation, and termination.
              most organisms, specify the amino acid glutamine; in other   2.  Compare transcription initiation in prokaryotes and
              ciliates, UGA, the third stop codon in most organisms,    eukaryotes.
              specifies cysteine. Ciliates use the remaining nonsense   3.  List three ways by which eukaryotes process mRNA
                codons as stop codons.                                  after transcription.
                  Other systematic changes in the genetic code exist in
              mitochondria,  the  semiautonomous,  self-reproducing
              organelles within eukaryotic cells that are the sites of   Transcription is the process by which the polymerization of
              ATP formation. Each mitochondrion has its own chro-  ribonucleotides, guided by complementary base pairing,
              mosomes and  its own  apparatus  for gene expression   produces an RNA transcript of a gene. The template for the
              (which we describe in detail in Chapter 15). In the mito-  RNA transcript is one strand of that portion of the DNA
              chondria of yeast, for example, CUA specifies threonine   double helix that constitutes the gene.
              instead of leucine. Yet another exception to the code is
              seen in certain prokaryotes that sometimes use the tri-
              plet UAG to specify insertion of the rare amino acid   RNA Polymerase Synthesizes a Single-
              pyrrolysine (see Fig. 7.28c and also Problem 57 at the   Stranded RNA Copy of a Gene
              end of this chapter.)
                  The experimental evidence presented so far helped   Figure 8.10 depicts the basic components of transcription
              define a nearly universal genetic code. But although   and illustrates key events in the process as it occurs in the
              cracking the code made it possible to understand the   bacterium  E. coli. This figure divides transcription into
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