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Table 7.3 Genotypes of Hfr and F Cells Used
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in an Interrupted Mating Experiment
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*Limiting percentage for az, tonAr, lac, and galB loci.
Figure 7.16 Frequency of Hfr genetic characters among
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recombinants after interrupted mating. As time proceeds, new alleles appear and then increase in frequency. Interruption of the mating limits the frequency of successful passage.
(From F. Jacobs and E. L. Wollman, Sexuality and the Genetics of Bacteria, Academic Press, 1961.)
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
7. Linkage and Mapping in Prokaryotes and Bacterial Viruses
The McGraw Hill Companies, 2001
Sexual Processes in Bacteria and Bacteriophages
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age does not increase with additional time. The limiting percentage is lower for loci that enter later, a fact explained by the assumption that even without the food blender, mating is usually interrupted before completion by normal agitation alone.
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Mapping and Conjugation
Jacob and Wollman, working with several different Hfr strains, collected data that indicated that the bacterial chromosome was circular. The strains were of independent origin, and the results were quite striking (table 7.4). If we ponder this table for a short while, one fact becomes obvious: The relative order of the loci is always the same. What differs is the point of origin and the direction of the transfer. Jacob and Wollman proposed that normally the F factor is an independent circular DNA entity in the F cell, and that during conjugation only the F factor is passed to the F cell. Since it is a small fragment of DNA, it can be passed entirely in a high proportion of conjugations before the cells separate. Every once in a while, however, the F factor becomes integrated into the chromosome of the host, which then becomes an Hfr cell. The point of integration can be different in different strains. However, once the F factor is integrated, it determines the initiation point of transfer for the E. coli chromosome, as well as the direction of transfer. The F factor is the last part of the E. coli chromosome to be passed from the Hfr cell. This explains why an Hfr, in contrast to an F , rarely passes the F factor itself. In the original work of Lederberg and Tatum, the one recombinant in 107 cells most likely came from a conjugation between an F cell and an Hfr that had formed spontaneously from an F cell. Integration of the F factor is diagrammed in gure 7.18.The F factor can also reverse this process and loop out of the E. coli chromosome. (Sometimes the F factor loops out incorrectly, as in gure 7.19, forming an F [F-prime] factor. The passage of this F factor to an F cell is called F-duction or sexduction. Not really useful in mapping, the process has proved exceptionally useful in studies of gene expression because of the formation of stable merozygotes, which we will examine in chapter 14.) We could now diagram the E. coli chromosome and show the map location of all known loci. The map units would be in minutes, obtained by interrupted mating. However, at this point, the map would not be complete. Interrupted mating is most accurate in giving the relative position of loci that are not very close to each other. With this method alone, a great deal of ambiguity would arise as to the speci c order of very close genes on the chromosome. The remaining sexual process in bacteria, transduction, provides the details that interrupted mating or transformation don t explain.
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Figure 7.17 Conjugation in E. coli. Hfr chromosome is blue;
F chromosome is red; and new DNA replication is black. As time proceeds, alleles from the Hfr enter the F cell in an orderly, sequential fashion. After the cells separate, two crossovers can bring Hfr alleles into the F chromosome. The F factor (orange) is the last part of the Hfr chromosome to enter the F cell.
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