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8.1 The Genetic Code   271


                       the process known as translation. It takes place on molecular workbenches called   Figure 8.1  Gene expression: The
                       ribosomes, which are composed of proteins and ribosomal RNAs (rRNAs). Trans­  flow of genetic information from
                       lation depends on the dictionary known as the genetic code, which defines each   DNA via RNA to protein. In
                       amino acid in terms of specific sequences of three nucleotides. Translation also re­  transcription, the enzyme RNA
                       quires transfer RNAs (tRNAs), small RNA adapter molecules that place specific   polymerase copies DNA to produce an
                                                                                                RNA transcript. In translation, the
                       amino acids at the correct position in a growing polypeptide chain.      cellular machinery uses instructions in
                          The Central Dogma does not explain the behavior of all genes. As Crick   mRNA to synthesize a polypeptide,
                       himself realized, many genes are transcribed into RNAs that are never translated   following the rules of the genetic code.
                       into proteins. You will see in this chapter that many nontranslated RNAs are          DNA
                       critical to various steps of gene expression. The genes encoding rRNAs and
                       tRNAs belong to this group.
                          Four general themes emerge from our discussion of gene expression. First,             Transcription
                       the pairing of complementary bases is key to the transfer of information from
                       DNA to RNA, and from RNA to protein. Second, the polarities (directionality)
                       of DNA, RNA, and polypeptides help guide the mechanisms of gene expres­  RNA transcript: serves directly as mRNA
                       sion. Third, like DNA replication and recombination, gene expression requires   in prokaryotes; processed to become
                       an  input of energy and the participation of specific proteins, RNAs, and mac­  mRNA in eukaryotes
                       romolecular assemblies, such as ribosomes. Finally, mutations that change ge­            Translation
                       netic   information or obstruct the flow of its expression can have dramatic
                       effects on phenotype.


                                                                                                          Polypeptide






                        8.1   The Genetic Code                             more letters exist than the two symbols (dot or dash), groups
                                                                           of one, two, three, or four dots or dashes in various combi­
                                                                           nations represent individual letters. For example, the sym­
                                                                           bol for C is dash dot dash dot (– · – ·), the symbol for O is
                        learning objectives
                                                                           dash dash dash (– – –), D is dash dot dot (– · ·), and E is a
                        1.  Explain the reasoning that established a sequence of   single dot (·). Because anywhere from one to four symbols
                            three nucleotides (a triplet codon) as the basic unit of   specify each letter, the Morse code requires a symbol for
                            the code relating DNA to protein.              pause (in practice, a short interval of time) to signify where
                        2.  Summarize the evidence showing that the sequence of   one letter ends and the next begins.
                            nucleotides in a gene is colinear with the sequence of
                            amino acids in a protein.
                        3.  Define reading frame and discuss its significance to the   Triplet Codons of Nucleotides Represent
                            genetic code.                                  Individual Amino Acids
                        4.  Describe experiments that determined which codons   The language of nucleic acids is written in four nucleo­
                            are associated with each amino acid and which are    tides—A, G, C, and T in the DNA dialect; A, G, C, and U
                            stop codons.                                   in the RNA dialect—while the language of proteins is writ­
                        5.  Explain how mutations were used to verify the genetic   ten in amino acids. The first hurdle to be overcome in deci­
                            code.                                          phering how sequences of nucleotides can determine the
                        6.  Discuss evidence that the genetic code is almost   order of amino acids in a polypeptide is to determine how
                            universal, and cite some exceptions.           many amino acid “letters” exist. 
                                                                               Over lunch one day at a local pub, Watson and Crick
                                                                           produced the now accepted list of the 20 common amino
                       A code is a system of symbols that equates information in   acids that are encoded directly by DNA. They created the
                       one language with information in another. A useful anal­  list by analyzing the known amino acid sequences of a
                       ogy for the genetic code is the Morse code, which uses dots   variety of naturally occurring polypeptides. Amino acids
                       and dashes to transmit messages over radio or telegraph   present in only a small number of proteins or in only cer­
                       wires. Various groupings of the dot­dash symbols represent   tain tissues or organisms did not qualify as standard
                       the 26 letters of the English alphabet. And because many   building blocks; Crick and Watson correctly assumed
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