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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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10. Gene Expression: Transcription
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The McGraw Hill Companies, 2001
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Gene Expression: Transcription
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GA UCU 3 -poly-A tail
Single-stranded DNA Exon I UCUU Intron 1 5 end DNA-RNA hybrid Intron 3 Exon II + GA Intron + G G
Figure 10.31 Self-splicing of a ribosomal RNA precursor in Tetrahymena. An external GTP is required. Two bond transfers produce a shortened RNA and a free intron.
(b) Figure 10.30
Intron 2
Protein-Mediated Splicing (the Spliceosome)
Eukaryotic nuclear messenger RNAs also have their introns removed by way of a lariat structure, just as in type II introns, but with the help of RNA-protein particles. Figure 10.36 shows consensus sequences in nuclear messenger RNA for the majority of introns. At the left (5 ) side of the intron, the GU sequence is invariant, as is the AG at the right (3 ) side. The right-most A of the UACUAAC sequence is the branch point of the lariat and is also invariant. (In DNA nucleotides, UACUAAC is TACTAAC; therefore, that region is sometimes referred to as the TACTAAC box.) Unlike the mitochondrial group II introns, however, nuclear messenger RNAs have their introns removed with the help of a protein-RNA complex called a spliceosome, named by J. Abelson and E. Brody. The splicing apparatus in eukaryotic messenger RNAs consists of several components called small nuclear ribonucleoproteins (discovered and named by J. Steitz and colleagues), abbreviated as snRNPs and pronounced snurps. Five of these particles take part in splicing, each composed of one or more proteins and a small RNA molecule; they are designated U1, U2, U4, U5, and U6. The RNA molecules range in size from 100 to 215 bases. The snRNPs and their associated proteins are located in twenty to forty small regions in the nucleus called speckles because of their appearance in the uorescent microscope.
The mRNA of adenovirus hybridized with its DNA. Three introns are visible as single-stranded DNA loops. They form single-stranded loops because they have nothing in the RNA molecule to hybridize with. Also visible is the poly-A tail of the mRNA. (a) Electron micrograph, (b) explanatory diagram. ([a] Courtesy of Louise T. Chow and Thomas Broker.)
Thomas Cech (1947 ).
(Courtesy of Dr. Thomas Cech. Photo by Ken Abbott.)
Sidney Altman (1939 ).
(Courtesy of Dr. Sidney Altman. Photo: Michael Marsland, Yale University Of ce of Public Affairs.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
10. Gene Expression: Transcription
The McGraw Hill Companies, 2001
Eukaryotic DNA Transcription
The RNAs of these particles have been sequenced, and sequencing shows they have regions of complementarity to either sites in the exons, sites in the introns, or sites in the other snRNP RNAs (table 10.5). These sequences, together with the experimental techniques of photocrosslinking and the creation of selective mutations (using techniques of site-directed mutagenesis described in chapter 13) have given us insight into the splicing mechanism. Photocrosslinking tells us which components are in contact. Mutations change pairings of components and may disrupt the structure. The change can be rescued (the pairing restored) by making a second change in the complementary RNA. When this happens successfully, the
Joan A. Steitz (1941 ).
(Courtesy of Dr. Joan A. Steitz.)
Ribozyme
GGGAGG5
ggcccucuOH
ggcccucua5
+GOH
3 GOH g g c c c u c u OH GGGAGG5 Ga5
The intron removed from the ribosomal RNA of Tetrahymena can catalyze the removal of the 3 end of an RNA, diagrammed here as ve AMP residues (a5) from the sequence 5 -GGCCCUCUA5-3 . The intron is called the Tetrahymena ribozyme. Any sequence can be removed from an RNA as long as there is a sequence complementary to the GGGAGG-5 of the ribozyme to bring the RNA into position. In (a), the reaction needs an external guanine-containing nucleotide (GOH); substrate nucleotides are in lowercase letters. This transesteri cation requires no external energy. In (b), the secondary structure of the ribozyme is shown. GOH is the site of cleavage, and the position of the G-binding site is shown. Further structure must develop to bring the G-site to the substrate. Wavy lines represent additional structure not shown. (Reprinted with permission from Ann Marie Pyle, et al., RNA
substrate binding site in the catalytic core of the Tetrahymena ribozyme, Nature, Volume 358, 1992. Copyright 1992 Macmillan Magazines, Ltd.)
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