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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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Gene Expression: Transcription
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Promoter DNA
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Transcription
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UA A Trailer
Transcribed piece of prokaryotic RNA and its DNA template region. Note the promoter and terminator regions on the DNA and the leader and trailer regions on the RNA. The initiation (AUG) and nonsense (UAA) codons for protein synthesis are shown. These signals are read at the ribosome at the time of translation.
RIBOSOMES AND RIBOSOMAL RNA
Ribosomes are organelles in the cell, composed of proteins and RNA (ribosomal RNA, or rRNA), where protein synthesis occurs. In a rapidly growing E. coli cell, ribosomes can make up as much as 25% of the mass of the cell. Ribosomes, as well as other small particles and molecules, are measured in units that describe their rate of sedimentation during density-gradient centrifugation in sucrose. This technique gives information on size and shape (due to the speed of sedimentation) while simultaneously isolating the molecules. Isolation by centrifugation in sucrose is a relatively gentle isolation technique; the molecules still retain their biological properties and can be used for further experimentation. In the 1920s, physical chemist T. Svedberg developed ultracentrifugation, giving his name to the unit of sedimentation: the Svedberg unit, S. In sucrose density-gradient centrifugation, the gradient is formed by layering on decreasingly concentrated sucrose solutions. In a related technique, cesium chloride density-gradient centrifugation, mentioned in chapter 9, the gradient develops during centrifugation. The sucrose centrifugation is stopped after a xed time, whereas in the cesium chloride technique, the system spins until it reaches equilibrium. The sucrose method tends to be more rapid. Samples can be isolated from a sucrose gradient by punching a hole in the bottom of the tube and collecting the drops in sequentially numbered containers. The rst (lowest-numbered) containers will contain the heaviest molecules (with the highest S values). Ribosomes in all organisms are made of two subunits of unequal size.The sedimentation value is 50S (Svedberg units) for the large one in E. coli and 30S for the smaller one. Together they sediment at about 70S. Eukaryotic ribosomes vary from 55S to 66S in animals and 70S to 80S in fungi and higher plants. Most of our discussion will be con ned to the well-studied ribosomes of E. coli.
Each ribosomal subunit comprises one or two pieces of ribosomal RNA and a xed number of proteins. The 30S subunit of E. coli has twenty-one proteins and a 16S molecule of ribosomal RNA, and the 50S subunit has thirty-four proteins and two pieces of ribosomal RNA one 23S and one 5S section ( g. 10.14). Advances in understanding ribosomal structure have come about after protein chemists isolated and puri ed all the proteins of the ribosome. This allowed researchers to experiment on the proper sequence needed to assemble the subunits and also allowed them to develop immunological techniques to show the positions of many proteins in the completed ribosomal subunits. In E. coli, all three ribosomal RNA segments are transcribed as a single long piece of RNA that is then cleaved and modi ed to form the nal three pieces of RNA (16S, 23S, and 5S). The region of DNA that contains the three ribosomal RNA molecules also contains genes for four transfer RNAs ( g. 10.15). There appear to be about ve to ten copies of this region in each chromosome of E. coli. The occurrence of the three ribosomal RNA segments on the same piece of RNA ensures a nal ratio of 1:1:1, the ratio needed for ribosomal construction.
TRANSFER RNA
During protein synthesis (see g. 10.3), a messenger RNA, carrying the information transcribed from the gene (DNA), is bound to the ribosome. Amino acids are brought to the ribosome attached to transfer RNAs. The code is read in sequences of three nucleotides, called codons. The nucleotides of the codon on messenger RNA are complementary to and pair with a sequence of three bases the anticodon on a transfer RNA. Each different transfer RNA carries a speci c amino acid.Thus, the transfer RNA recognizes the speci city of the genetic code ( g. 10.16).
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