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Normally, a gene used to transfect mice is incorporated randomly in the mouse genome. However, in about one in one thousand experiments, the gene replaces the normal gene by a process similar to meiotic recombination (homologous recombination; see chapter 12). With this
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process in mind, geneticists have been able to select for homologous recombination; by transfecting with defective genes, they have created mice without working copies of a particular gene.The mice produced are called knockout mice, and they give geneticists the opportunity to study the phenotype of an animal that lacks a particular gene. The geneticist rst creates a vector with the modi ed gene in question. In addition, anking regions to that gene are added so that homologous recombination can occur. Finally, two antibiotic genes are introduced so that selection for successful transfection takes place. Within the anking regions, the gene for neomycin resistance (neor ) is inserted; its product inactivates the antibiotic neomycin ( g. 13. 22). Outside of the anking regions,
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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13. Genomics, Biotechnology, and Recombinant DNA
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Thirteen
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Genomics, Biotechnology, and Recombinant DNA
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the gene for thymidine kinase (tk) is inserted. This gene phosphorylates the drug gancyclovir; the phosphorylated gancyclovir is a nucleotide analogue that is incorporated during DNA synthesis, killing the cell. Thus, the combination of the tk gene and gancyclovir is lethal; without the tk gene, gancyclovir is harmless. If cells are exposed to both drugs, neomycin and gancyclovir, normal cells will be killed by neomycin, cells with the tk gene will be killed by gancyclovir, and only the cells with the neor gene but lacking the tk gene will survive. These alternative outcomes allow us to select the cells in which homologous recombination took place ( g. 13.22). Cells that did not incorporate the vector will die from the effects of neomycin (they are neomycin sensitive). Cells that randomly took up the vector DNA by nonhomologous recombination will contain the neor and tk genes and will be killed by gancyclovir. However, cells that underwent homologous recombination will contain the neor gene but lack the tk gene; these cells will therefore survive in the presence of both antibiotics ( g. 13.22). Geneticists can isolate embryonic stem cells from mice, cells that can produce any mouse tissue. The cells are transfected and then grown in tissue culture in the
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presence of neomycin and gancyclovir, and only cells that undergo homologous recombination will survive. These cells are then injected into early-stage mouse embryos to become part of the developing mouse.The mice that develop will be chimeric; some will have incorporated the transfected cells into the germ line and become heterozygotes for the disabled gene. Finally, when mice like this are mated, one fourth of their offspring will be homozygous for the disabled gene. Thus, knockout mice, have been created through this ingenious technique. Knockout mice are especially useful for studying development and immunology. For example, if the gene for the Mullerian-inhibiting substance is knocked out, males are infertile because they develop female reproductive organs. This experiment led to insight into the genetic path for sex determination (see chapter 5). Hundreds of knockout experiments are published each year.
Reporter Systems
We conclude this section by discussing two reporter systems, systems used to indicate that a transfection experiment was successful. Plants can be transfected with
... ...
(a) Homologous recombination
neo r
... ...
Host chromosome (homologous region)
(b) Nonhomologous recombination
neo r
Vector
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X* neo r tk
... ...
Host chromosome (nonhomologous region)
Figure 13.22 Creating a knockout mouse. A vector is created that has a disabled (nonfunctional) form
of the gene in question (X*; red). Next to the gene in question is the neomycin resistance gene (neor; green); both genes are surrounded by regions (blue) that ank the normal gene on its chromosome. Finally, outside the anking regions in the vector is the gene for thymidine kinase (tk; yellow). In homologous recombination (a), involving crossovers in the homologous anking regions, the disabled gene and the neomycin resistance gene replace the normal gene on the cell s chromosome. In nonhomologous recombination (b), almost the entire vector is incorporated into the host chromosome, including the thymidine kinase gene. Techniques then allow for the selective growth of cells with the rare homologous recombination event. One fourth of the offspring of heterozygous chimeric mice will be knockouts for the gene in question.
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