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The EF-Ts/EF-Tu cycle. EF-Ts and EF-Tu are required for a transfer RNA to attach to the A site of the ribosome. At top center, we have EF-Tu attached to a GDP. The GDP is then displaced by EF-Ts, which in turn is displaced by GTP. A transfer RNA attaches and is brought to the ribosome. If the codon-anticodon t is correct, the transfer RNA attaches at the A site with the help of the hydrolysis of GTP to GDP Pi, allowing the EF-Tu to release. The EF-Tu is now back where we started. Since EF-Tu has a strong af nity for GDP, the role of EF-Ts is to displace the GDP, and later to be replaced by GTP.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
11. Gene Expression: Translation
The McGraw Hill Companies, 2001
Eleven
Gene Expression: Translation
BOX 11.2
ntibiotics, substances living organisms produce that are toxic to other living organisms, are of interest to us for two reasons: They have been extremely important in ghting the diseases that strike human beings and farm animals, and many are useful tools for analyzing protein synthesis. Some antibiotics impede the process of protein synthesis in a variety of ways, often poisoning bacteria selectively; the effectiveness of antibiotics normally derives from the metabolic differences between prokaryotes and eukaryotes. For example, an antibiotic that blocks a 70S bacterial ribosome without affecting an 80S human ribosome could be an excellent antibiotic. About 160 antibiotics are known.
Biomedical Applications
Antibiotics
PUROMYCIN Puromycin resembles the 3 end of an aminoacyl-tRNA ( g. 1). It is bound to the A site of the bacterial ribosome, where peptidyl transferase creates a bond from the nascent peptide attached to the transfer RNA in the P site to puromycin. Elongation can then no longer occur.The peptide chain is released prematurely, and protein synthesis at the ribosome terminates. Experiments with puromycin helped demonstrate the existence of the A and P sites of the ribosome. It was found that puromycin could not bind to the ribosome if translocation factor EF-G were absent. With EF-G, translocation took place, and puromycin could then bind to the ribosome. Puromycin s ability to bind only after translocation indicates that a second site on the ribosome becomes available after translocation. STREPTOMYCIN, TETRACYCLINE, AND CHLORAMPHENICOL Streptomycin, which binds to one of the proteins (protein S12) of the 30S subunit of the prokaryotic ribosome, inhibits initiation of protein synthesis. Streptomycin also causes misread-
ing of codons if chain initiation has already begun, presumably by altering the conformation of the ribosome so that transfer RNAs are less rmly bound to it. Bacterial mutants that are streptomycin resistant, as well as mutants that are streptomycin dependent (they cannot survive without the antibiotic), occur. Both types of mutants have altered 30S subunits, speci cally changed in protein S12. Tetracycline blocks protein synthesis by preventing an aminoacyltRNA from binding to the A site on the ribosome. Chloramphenicol blocks protein synthesis by binding to the 50S subunit of the prokaryotic ribosome, where it blocks the peptidyl transfer reaction. Chloramphenicol does not affect the eukaryotic ribosome. However, chloramphenicol, as well as several other antibiotics, is used cautiously because the mitochondrial ribosomes within eukaryotic cells are very similar to prokaryotic ribosomes. Some of the antibiotics that affect prokaryotic ribosomes thus also affect mitochondria.As was mentioned, the similarity between bacteria and mitochondria implies that mitochondria have a prokaryotic origin. (Similarities between cyanobacteria and chloroplasts also support the idea that chloroplasts have a prokaryotic origin.) THE TROUBLE WITH ANTIBIOTICS Over the years, antibiotics have virtually eliminated certain diseases from the industrialized world. They have also made modern surgery possible by preventing most serious infections
that tend to follow operations.Antibiotics have been so successful that, in the 1980s, many pharmaceutical companies drastically cut back the development of new antibiotics. However, a disaster was in the making as we overprescribed antibiotics to people and farm animals: bacteria are not prepared to take this onslaught without ghting back. Mutation takes place all the time at a low but dependable rate.Thus, resistant bacteria are constantly arising from sensitive strains. We can select for penicillin- and streptomycinresistant strains of bacteria in the laboratory by allowing the antibiotic to act as a selective agent, removing all but the resistant individuals. The same sort of arti cial selection that we can apply in the lab applies every time a person or animal takes an antibiotic. We may be at a point now where the ability of bacteria to develop resistance, and to pass that resistance to other strains, has put us on the verge of disaster. The process of evolution works amazingly fast in bacteria because of their ubiquity, large population sizes, and ability to transfer genetic material between individuals. We may shortly nd ourselves as we were before World War II, when simple infections in hospitals were often lethal. Right now, only one antibiotic can keep the common and potentially deadly infectious bacterium Staphylococcus under control: vancomycin. Several types of disease-causing bacteria have already evolved a tolerance to vancomycin. The answer to this potentially disastrous problem is to develop new antibiotics and reduce the irresponsible use of antibiotics in people and animals. Hopefully, the warning bell has sounded.At least a dozen new antibiotics that show promise are in the early stages of development by pharmaceutical companies.
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