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Tamarin: Principles of Genetics, Seventh Edition
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II. Mendelism and the Chromosomal Theory
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7. Linkage and Mapping in Prokaryotes and Bacterial Viruses
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The McGraw Hill Companies, 2001
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Linkage and Mapping in Prokaryotes and Bacterial Viruses
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No growth
met + bio + thr + leu +
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Figure 7.12 Lederberg and Tatum s cross showing that E. coli undergoes genetic
recombination.
F Factor
In bacteria, conjugation is a one-way transfer, with one strain acting as donor and the other as recipient. Sometimes donor cells, if stored for a long time, lose the ability to be donors, but they can regain the ability if they are mated with other donor strains. This discovery led to the hypothesis that a fertility factor, F, made any strain that carried it a male (donor) strain, termed F .The strain that did not have the F factor, referred to as a female or F strain, served as a recipient for genetic material during conjugation. Research supports this hypothesis. The F factor is a plasmid, a term originally coined by Lederberg to refer to independent, self-replicating genetic particles. Plasmids are usually circles of double-stranded DNA. (Plasmids are at the heart of recombinant DNA technology, which is discussed in detail in chapter 13.) They are auxiliary circles of DNA that many bacteria carry. They are usually much smaller than the bacterial chromosome. Researchers found that the transfer of the F factor occurred far more frequently than the transfer of other genes from the donor. That is, during conjugation, about one recombinant occurred in 107 cells, whereas transfer of the F factor occurred at a rate of about one conversion of F to F in every ve conjugations. An E. coli strain was then discovered that transferred its genetic material at a rate about one thousand times that of the normal F strain. This strain was called Hfr, for high frequency of recombination. Several other phenomena occurred with this high rate of transfer. First, the ability to transfer the F factor itself dropped to almost zero in this strain. Second,
Strain A
Strain B
Filter Figure 7.13 The U-tube experiment. Alternating suction and
pressure force liquid and macromolecules back and forth across the lter.
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
not all loci were transferred at the same rate. Some loci were transferred much more frequently than others. Escherichia coli cells are normally coated with hairlike pili ( mbriae). F and Hfr cells have one to three additional pili (singular: pilus) called F-pili, or sex pili. During conjugation, these sex pili form a connecting bridge between the F (or Hfr) and F cells ( g. 7.14). Once a connection is made, the sex pilus then contracts to bring the two cells into contact. DNA transfer takes place through a nick in either the plasmid (in F cells) or the bacterial chromosome (in Hfr cells). A single strand of the DNA double-stranded donor DNA then passes from the F or Hfr cell to the F cell across the cell membranes. DNA replication in both the donor and recipient cells reestablishes double-stranded DNA in both. The F factor itself has the genes for sex-pilus formation and DNA transfer to a conjugating F cell. At least twenty-two genes are involved in the transfer process, including genes for the pilus protein, nicking the DNA, and regulation of the process. In the transfer process of conjugation, the donor cell does not lose its F factor or its chromosome because only
a single strand of the DNA double helix is transferred; the remaining single strand is quickly replicated. (The process of DNA replication is described in chapter 9.) For a short while, the F cell that has conjugated with an Hfr cell has two copies of whatever chromosomal loci were transferred: one copy of its own and one transferred in. With these two copies, the cell is a partial diploid, or a merozygote. The new foreign DNA (exogenote) can be incorporated into the host chromosome (endogenote) by an even number of breakages and reunions between the two, just as in transformation. The unincorporated linear DNA is soon degraded by enzymes. The conjugation process is diagrammed in gure 7.15.
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