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Conjugation Can Be Used to Map Bacterial Genes by Mixing

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Conjugation can be used to map bacterial genes by mixing Hfr and F- cells of different genotypes and interrupting conjugation at regular intervals. The amount of time required for individual genes to be transferred from the Hfr to the F- cells indicates the relative positions of the genes on the bacterial chromosome.
-Interrupted conjugation was used to map three genes in E. coli. The donor genes first appeared in the recipient cells at the following times: gal, 10 minutes; his, 8 minutes; pro, 15 minutes. Which gene is in the middle?

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