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166    Chapter 5    Linkage, Recombination, and the Mapping of Genes on Chromosomes


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              Figure 5.29  Mitotic crossing-over. (a) In a y sn  / y  sn Drosophila female, a mitotic crossover between the centromere and sn can produce
              two daughter cells, one homozygous for y and the other homozygous for sn, that can develop into adjacent aberrant patches (twin spots). This
              outcome depends on a particular distribution of chromatids at anaphase (top). If the chromatids are arranged in the equally likely opposite
              orientation, only phenotypically normal cells will result (bottom). (b) Crossovers between sn and y can generate single yellow patches. However, a
              single mitotic crossover in these females cannot produce a single singed spot if the sn gene is closer to the centromere than the y gene.
                  Transient pairing
                  during mitosis            Mitotic metaphase                        Daughter cells

              (a)  Crossing-over between sn and the centromere

                                                                                sn +  y  Yellow    sn +  y
                                        sn +  y            sn +  y
                                                                                                               Twin
                                        sn   y +           sn   y +                                            spot
                      sn +  y                                                   sn   y Singed      sn  y +
                                                                                      +
                                                or

                      sn   y +                                                  sn +  y  Wild type   sn  y +
                                        sn +  y            sn   y +
                                                                                                               Normal
                                        sn   y +           sn +  y                                             tissue
                                                                                sn   y +  Wild type   sn +  y


              (b)  Crossing-over between sn and y

                                                                                sn +  y  Yellow    sn  y       Yellow
                                        sn +  y            sn   y                                              spot

                                        sn +  y +          sn   y +                                            Normal
                      sn +  y                                                     +   +                 +
                                                                                sn   y Wild type   sn  y       tissue
                                                or

                      sn   y +                                                  sn +  y  Wild type   sn  y +
                                        sn +  y            sn   y +
                                                                                                               Normal
                                        sn +  y +          sn   y                                              tissue
                                                                                      +
                                                                                sn +  y Wild type   sn  y


              Figure 5.30  Mitotic recombination during the growth of   happened during a cell division early in the growth of the
              diploid yeast colonies can create sectors. Arrows point to large,   colony, giving the resulting daughter cells a long time to
              red ade2 / ade2 sectors formed from ADE2 / ade2 heterozygotes.  proliferate. If a red sector is small, the recombination hap-
                Image courtesy of B.A. Montelone, Ph.D. and T.R. Manney, Ph.D  pened later.

                                                                   Mitotic Recombination Has Significant
                                                                   Consequences

                                                                   Problem 51 at the end of this chapter illustrates how ge-
                                                                   neticists use mitotic recombination to obtain information
                                                                   about the locations of genes relative to each other and to the
                                                                   centromere. Mitotic crossing-over has also been of great
                                                                   value in the study of development because it can generate
                                                                   animals in which different cells have different genotypes
                                                                   (see Problem 52 and also Chapter 19). Finally, as the
                                                                     Genetics and Society Box  Mitotic Recombination and
                                                                     Cancer  Formation  explains, mitotic recombination can
                                                                   have major repercussions for human health.
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