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17 FROM GENE TO PROTEIN Genes specify proteins via transcription and translation The products of Gene Expression: A Developing Story Basic Principles of Transcription and Translation The genetic code The genetic code must have evolved very early in the history of life The products of Gene Expression: It was hypothesized earlier that “one gene–one enzyme”. As researchers learned more about proteins, they slightly modified this hypothesis: • While most enzymes are proteins, many proteins are not enzymes (e.g. keratin, insulin, ect…). Proteins that are not enzymes are still, nevertheless, gene products. Scientists began to think in terms of one gene–one protein. • Also, many proteins are comprised of two or more polypeptide chains, each chain specified by its own gene (e.g. globulin chains of hemoglobin). • As a result of this new information, the previous hypothesis has been restated as one gene-one polypeptide. Basic Principles of Transcription and Translation • Genes provide the instructions for making specific proteins. • The bridge between DNA and protein synthesis is RNA. • RNA is chemically similar to DNA, except that it contains ribose as its sugar and substitutes the nitrogenous base uracil for thymine. – An RNA molecules almost always consists of a single strand. • In DNA or RNA, the four nucleotide monomers act like the letters of the alphabet to communicate information. • The specific sequence of hundreds or thousands of nucleotides in each gene carries the information for the primary structure of a protein, the linear order of the 20 possible amino acids. • To get from DNA, written in one chemical language, to protein, written in another, requires two major stages, transcription and translation. • Transcription is the synthesis of RNA under the direction of DNA. • During transcription, a DNA strand provides a template for the synthesis of a complementary RNA strand. • This process is used to synthesize any type of RNA from a DNA template. • Transcription of a gene produces a messenger RNA (mRNA) molecule. • Translation is the actual synthesis of polypeptide under the direction of mRNA. • During translation, the cell must translate the base sequence of an mRNA molecule into the amino acid sequence of a polypeptide. • Translation occurs at ribosomes. • The basic mechanics of transcription and translation are similar in eukaryotes and prokaryotes. • Because bacteria lack nuclei, transcription and translation are coupled. • Ribosomes attach to the leading end of a mRNA molecule while transcription is still in progress. Fig. 17.3a • In a eukaryotic cell, almost all transcription occurs in the nucleus and translation occurs mainly at ribosomes in the cytoplasm. • In addition, before the primary transcript (pre-mRNA) can leave the nucleus, it is modified in various ways during RNA processing before the finished mRNA is exported to the cytoplasm. Fig. 17.3b • To summarize, genes program protein synthesis via genetic messenger RNA. • The molecular chain of command in a cell is : DNA RNA •Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings protein. The Genetic Code Codons: Triplets of Bases • If the genetic code consisted of a single nucleotide or even pairs of nucleotides per amino acid, there would not be enough combinations (4 and 16 respectively) to code for all 20 amino acids. • Triplets of nucleotide bases are the smallest units of uniform length that can code for all the amino acids. • In the triplet code, three consecutive bases specify an amino acid, creating 43 (64) possible code words. • The genetic instructions for a polypeptide chain are written in DNA as a series of three-nucleotide words. • During transcription, one DNA strand, the template strand, provides a template for ordering the sequence of nucleotides in an RNA transcript. • The complementary RNA molecule is synthesized according to base-pairing rules, except that uracil is the complementary base to adenine. • During translation, blocks of three nucleotides, codons, are decoded into a sequence of amino acids. Fig. 17.4 • During translation, the codons are read in the 5’ 3’ direction along the mRNA. • Each codon specifies which one of the 20 amino acids will be incorporated at the corresponding position along a polypeptide. • Because codons are base triplets, the number of nucleotides making up a genetic message must be three times the number of amino acids making up the protein product. • It would take at least 300 nucleotides to code for a polypeptide that is 100 amino acids long. Cracking the Genetic Code • The first codon was deciphered in 1961 by Marshall Nirenberg of the National Institutes of Health. • Marshall Nirenberg determined the first match, that UUU coded for the amino acid phenylalanine. • He created an artificial mRNA molecule entirely of uracil and added it to a test tube mixture of amino acids, ribosomes, and other components for protein synthesis. • This “poly(U)” translated into a polypeptide containing a single amino acid, phenyalanine, in a long chain. • Other more elaborate techniques were required to decode mixed triplets such a AUA and CGA. • By the mid-1960s the entire code was determined. • 61 of 64 triplets code for amino acids. • The codon AUG not only codes for the amino acid methionine but also indicates the start of translation. • Three codons do not indicate amino acids but signal the termination of translation. Fig. 17.5 • The genetic code is redundant but not ambiguous. • There are typically several different codons that would indicate a specific amino acid. • However, any one codon indicates only one amino acid. • [If you have a specific codon, you can be sure of the corresponding amino acid, but if you know only the amino acid, there may be several possible codons.] • Both GAA and GAG specify glutamate, but no other amino acid. • Codons synonymous for the same amino acid often differ only in the third codon position. • In summary, genetic information is encoded as a sequence of nonoverlapping base triplets, or codons, each of which is translated into a specific amino acid during protein synthesis. • To extract the message from the genetic code requires specifying the correct starting point. • This establishes the reading frame and subsequent codons are read in groups of three nucleotides. • For example, the sequence of amino acids - Trp - Phe - Gly - Arg - Phe - can be assembled in the correct order only if the mRNA codons UGGUUUGGCCGUUUU are read in the correct sequence and groups. • The cell reads the message in the correct frame as a series of non-overlapping three-letter words: UGG UUU - GGC - CGU - UUU. CHAPTER 17 FROM GENE TO PROTEIN Section B: The Synthesis and Processing of RNA 1. Transcription is the DNA-directed synthesis of RNA: a closer look 2. Eukaryotic cells modify RNA after transcription Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 17.2 Transcription is the DNA-directed synthesis of RNA: a closer look • Messenger RNA is transcribed from the template • • strand of a gene. RNA polymerase separates the DNA strands at the appropriate point and links the RNA nucleotides as they base-pair along the DNA template. Like DNA polymerases, RNA polymerases can add nucleotides only to the 3’ end of the growing polymer. • Genes are read 3’ 5’, creating a 5’ 3’ RNA molecule. • Specific sequences of nucleotides along the DNA mark where gene transcription begins and ends: • RNA polymerase attaches and initiates transcription at the promotor. • The terminator signals the end of transcription. • The promotor, terminator and the nucleotides in between are called a transcription unit. • Bacteria have a single type of RNA polymerase that synthesizes all RNA molecules. • In contrast, eukaryotes have three RNA polymerases (I, II, and III) in their nuclei. • RNA polymerase II is used for mRNA synthesis. • Transcription can be separated into three stages: initiation, elongation, and termination. Fig. 17.6a A. RNA Polymerase Binding and Initiation of Transcription • The presence of a promotor sequence determines which strand of the DNA helix is the template. • Within the promotor is the starting point for the transcription of a gene. • The promotor also includes a binding site for RNA polymerase several dozen nucleotides upstream of the start point. • In prokaryotes, RNA polymerase can recognize and bind directly to the promotor region. • In eukaryotes, proteins called transcription factors recognize the promotor region, especially a TATA box, and bind to the promotor. • After they have bound to the promotor, RNA polymerase binds to transcription factors to create a transcription initiation complex. • RNA polymerase then starts transcription. Fig. 17.7 B. Elongation of the RNA Strand • As RNA polymerase moves along the DNA, it untwists the double helix, 10 to 20 bases at time. • The enzyme adds nucleotides to the 3’ end of the growing strand. • Behind the point of RNA synthesis, the double helix re-forms and the RNA molecule peels away. Fig. 17.6b • A single gene can be transcribed simultaneously by several RNA polymerases II, following each other, at a time. • A growing strand of mRNA trails off from each polymerase. • The length of each new strand reflects how far along the template the enzyme has traveled from the start point. • The congregation of many polymerase molecules simultaneously transcribing a single gene increases the amount of mRNA transcribed from it. • This helps the cell make the encoded protein in large amounts. C. Termination of Transcription • Transcription proceeds until after the RNA polymerase transcribes a terminator sequence in the DNA. • In prokaryotes, RNA polymerase stops transcription right at the end of the terminator. • Both the RNA and DNA is then released. • In eukaryotes, at a point 10-35 nucleotides downstream from the terminator sequence (AAUAAA signal), the pre-mRNA is released from the. • At a point about 10 to 35 nucleotides past this sequence, the pre-mRNA is cut from the enzyme.