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In addition to the F factor found frequently in bacteria, a variety of other plasmids occur, including the R and Col plasmids. The R plasmids carry genes for resistance to various antibiotics, and the Col plasmids have genes that are responsible for producing proteins called colicins, which are toxic to strains of E. coli ( g. 17.17). Plasmids containing genes for Col-like toxins speci c for other bacterial species are also known. Col and R plasmids can exist in two states. In one state, the plasmid has
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Prokaryotic Plasmids
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In chapters 7 and 13, we discussed the role of plasmids in the study of prokaryotic genetics and in recombinant DNA work. They are mentioned again here because they represent extrachromosomal genetic systems, primarily
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Electron micrograph of replication of Col E1 circular plasmid. The arrows mark the branch points of the theta structure. Magni cation 90,000 . (Source: J. I.
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Tomizawa, Y. Sakakibara, and T. Kakefuda, Replication of Colicin E1 plasmid DNA in cell extracts: Origin and direction of replication, Proceedings of The National Academy of Sciences, USA, 71 [1974]:2260 64.)
a sequence of genes called the transfer operon (tra), which makes the plasmids similar to F factors in that they can transfer their genes from one bacterium to the next. In the other state, the plasmids lack this operon and cannot transfer their loci to another cell. Thus, Col and R plasmids are actually made of two parts: the loci for antibiotic resistance or colicin production and the part responsible for infectious transfer. In R plasmids, the infectious transfer part is called the resistance transfer factor (RTF). The occurrence of resistance plasmids was observed in Japan in the late 1950s, when it was discovered that bacteria were simultaneously acquiring resistance to several antibacterial agents. When cultures of Shigella, a dysentery-causing bacterium, were exposed to streptomycin, sulfonamide, chloramphenicol, or tetracycline, the bacteria exhibited resistance not only to the one particular agent they were exposed to but to one or more of the others as well. The plasmid responsible for this multiple resistance was named R222. The Col plasmids contain loci that produce proteins that are toxic, for various reasons, to strains of bacteria not carrying the plasmids. Colicins attack sensitive bacterial cells at bacterial surface receptors. They have been classi ed into twenty or more categories according to the types of receptors they attack. Some colicins may enter the cell directly, but others do not. For example, colicin K appears to kill sensitive cells by inhibiting DNA, RNA, and protein synthesis, although not directly entering the cell. Colicin E3, however, acts as an intracellular ribonuclease that cleaves off about fty nucleotides from the 3 end of the 16S ribosomal RNA within the ribosome. The cleavage inactivates the sensitive cell s ribosomes and is, of course, lethal.
Since many R plasmids, Col plasmids, and F factors, as well as host chromosomes, have insertion sequences (chapter 14), a good deal of exchange occurs among the plasmids, and many are able to integrate into the host chromosome. Although their mobility makes it easier to map and study plasmids, it also poses a human health problem. Resistance to various antibacterial agents is easily transferred among enterobacteria worldwide.This can even occur outside of host organisms (people) where pollution or sewage is found. In addition, resistance found in relatively harmless enterobacteria, such as E. coli, can easily pass to more pathogenic bacteria, such as Shigella and Salmonella. Since we are selecting for resistance every time we use antibacterial drugs, we should not use these drugs indiscriminately. For some time, health workers have been concerned about excessive medical use of antibacterial drugs as well as about the large quantities of antibiotics used in animal feed.
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