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8.2 Transcription: From DNA to RNA   287


                       (a)  A collagen gene: structure and expression      Figure 8.14  Structure and expression of a typical
                                                               Poly-A      eukaryotic gene. (a) Schematic view of landmarks in a collagen
                                            ATG            Stop signal
                                                                           gene and its products. Exons are shown in red, introns in green, and
                                                                           nontranscribed parts of the gene in blue. Mature mRNAs are
                       DNA        Exon 1  Intron 1  Exon 2  Intron 2  Exon 3  processed from the primary transcript; introns are spliced out, a 5′
                             Promoter       Transcription                  methyl-G cap is added, and a poly-A tail is added to the 3′ end. The
                                            AUG            Stop  AAUAAA    5′ untranslated region (5′ UTR) lies between the 5′ end and the start
                       Primary  5'  Exon 1  Intron 1  Exon 2  Intron 2  Exon 3  3'  codon (AUG), and the 3′ untranslated region (3′ UTR) lies between
                       transcript                                          the stop codon and the poly-A tail at the 3′ end of the mRNA
                                                          AAUAAA           (orange bars). (b) The same gene at the nucleotide level. Colors are
                                             AUG      Stop                 the same as in part (a), except that the mature mRNA is shown in
                       Mature      5' meG  Exon 1  Exon 2  Exon 3  AAAAAAAA 3'  purple for emphasis. The AAUAAA poly-A addition signal in the
                       mRNA         Cap                       Poly-A tail
                                                                           mRNA is underlined. Introns can occur anywhere in the transcribed
                                                                           part of a gene, including within a codon or either of the UTRs.
                                      5' UTR              3' UTR
                                              Met......
                       Protein              N          C






































                       place in concert (Fig. 8.16). The spliceosome consists of   It might seem strange that eukaryotic genes incorpo­
                       four subunits known as small nuclear ribonucleoproteins,   rate DNA sequences that are spliced out of the mRNA
                       or snRNPs (pronounced “snurps”). Each snRNP contains   before translation and thus do not encode amino acids.
                       one or two small nuclear RNAs (snRNAs) 100–300 nucleo­  No one knows exactly why introns exist. One hypothesis
                       tides long, associated with proteins in a discrete particle.   proposes that introns make it possible to assemble genes
                       Certain snRNAs can base pair with the splice donor and   from various exon building blocks that encode modules
                       splice   acceptor sequences in the primary transcript, so   of protein function. This type of assembly would allow
                       these   snRNAs are particularly important in bringing   the shuffling of exons to make new genes, a process that
                         together the two exons that flank an intron.      appears to have played a key role in the evolution of com­
                          Given the complexities of spliceosome structure, it is   plex organisms. The exon­as­module proposal is attrac­
                       remarkable that a few primary transcripts can splice them­  tive  because  it  is  easy  to  understand  the  selective
                       selves without the aid of a spliceosome or any additional   advantage of the potential for exon shuffling. Neverthe­
                       factor. These rare primary transcripts function as ribozymes:   less, it remains a hypothesis without proof; introns may
                       RNA molecules that can act as enzymes and catalyze a spe­  have  become  established  through  means  that  scientists
                       cific biochemical reaction.                         have yet to imagine.
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