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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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Eukaryotic DNA Transcription
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Finally, transcription in the archaea, although under much simpler control than in the eukaryotes, resembles transcription in eukaryotes rather than prokaryotes. The study of the details of the transcription process its initiation, control, and termination is one of the most active and exciting areas in modern genetics.
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that had not had any of the major posttranscriptional modi cations. In essence, they were premessenger RNAs.
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Introns
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Eukaryotes have segments of DNA within genes that are transcribed into RNA but never translated into protein sequences. These intervening sequences, or introns, are removed from the RNA in the nucleus before its transport into the cytoplasm ( g. 10.28). P. Sharp and his colleagues at MIT and R. Roberts, T. Broker, L. Chow, and their colleagues at the Cold Spring Harbor Laboratory rst discovered introns in 1977. (Sharp and Roberts were awarded 1993 Nobel prizes for their work.) An example of a gene with introns appears in gure 10.29. The segments of the gene between introns, which are transcribed and translated and hence exported to the cytoplasm and expressed are termed exons. The results of intron removal are clear when a messenger RNA with its introns removed is hybridized with the original gene ( g. 10.30). The DNA forms double-stranded structures with the exons in RNA. The introns in DNA have nothing to pair with in the RNA, so they form single-stranded loops. Introns also occur in eukaryotic transfer RNA and ribosomal RNA genes. For introns to be removed, the ends of the exons must be brought together and connected in a process called splicing. At least two types of splicing occur, although they are related: self-splicing and protein-mediated splicing.
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Eukaryotic transcription results in a primary transcript. In contrast to most prokaryotic transcripts that contain information from several genes, virtually all transcripts from higher eukaryotes contain the information from just one gene. (Transcripts from several genes are found in some lower eukaryotes, such as nematode worms.) Three major changes occur in primary transcripts of RNA polymerase II before transport into the cytoplasm: modi cations to the 5 and 3 ends and removal of intervening sequences. We refer to these changes as posttranscriptional modi cations. At the 5 end of polymerase II transcripts, 7-methyl guanosine is added in the wrong direction, 5 5 ( g. 10.27). This cap allows the ribosome to recognize the beginning of a messenger RNA. At the other end, the 3 end of polymerase II transcripts, a sequence of twenty to two hundred adenine-containing nucleotides, known as a poly-A tail, is added by the enzyme poly-A polymerase. Polyadenylation takes place after the 3 end of the transcript is removed by a nuclease that cuts about twenty nucleotides downstream from the signal 5 -AAUAAA-3 . The tail adds stability to the molecule and aids in its transportation from the nucleus. When messenger RNAs were rst studied in eukaryotes, the messenger RNAs in the nucleus were found to be much larger than those in the cytoplasm and were called heterogeneous nuclear mRNAs, or hnRNAs. It now turns out that these were primary transcripts, RNAs
Self-Splicing
In 1982,Thomas Cech and his colleagues, building on the work of others, including Sidney Altman, who showed that RNA can have catalytic properties, discovered selfsplicing by RNA. (Cech and Altman were awarded 1989
Cap CH3 N+ mRNA
HN C H2N N
N Base OH O O 5 5
Base OH O
Base OH O 3
HO P P P P P
A cap of 7-methyl guanosine is added in the wrong direction (5 5 ), to the 5 end of eukaryotic mRNAs. In some cases, the 2 -OH groups on the second or second and third riboses (red) are methylated.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
10. Gene Expression: Transcription
The McGraw Hill Companies, 2001
Ten
Gene Expression: Transcription
Transcription Intervening sequence I hnRNA Gene segment A Intervening sequence II Gene segment C
Gene segment B
Modification
Richard J. Roberts (1943 ).
(Courtesy of Richard J. Roberts.)
Philip A. Sharp (1944 ).
(Courtesy of Dr. Philip A. Sharp.)
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