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


              Figure 5.3  Designations of parental and recombinant relate to past history. Figure 5.2 has been redrawn here as Cross
              Series A for easier comparison with Cross Series B, in which the dihybrid F 1  females received different allelic combinations of the white and
              yellow genes. Note that the parental and recombinant classes in the two cross series are the opposite of each other. The percentages of
              recombinant and parental types are nonetheless similar in both experiments, showing that the frequency of recombination is independent of
              the arrangement of alleles.
                                  Cross Series A                                      Cross Series B
                  P                                                  P
                            w    y +          w +  y                            w +  y +          w   y


                            w    y +                                            w +  y +
                  F 1                                                F 1
                            w    y +          w    y +                          w +  y +          w +  y +

                            w +  y                                              w   y
                  F males                                            F males
                   2
                                                                      2
                   w    y +    w +  y      w   y        w +  y +      w +  y +     w  y        w +  y      w   y +

                  Parental    Parental   Recombinant  Recombinant     Parental    Parental  Recombinant  Recombinant
                         ~ 99%                     ~ 1%                      ~ 99%                    ~ 1%


              find out what kinds and ratios of gametes these F 1  females   illustration of the linkage concept. The two genes are so
              produce, we need to look at the telltale F 2  males.  tightly coupled that the parental combinations of alleles—
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                  This time, as Cross Series B in Fig. 5.3 shows, w  y / Y   w y  and w  y (in Cross Series A of Fig. 5.3) or w  y  and
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              and w y  / Y are the recombinants that account for little   w y (in Cross Series B)—are reshuffled to form recombi-
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              more than 1% of the total, while w y / Y and w  y  / Y are   nants in only 1 out of every 100 gametes formed. In other
              the parental combinations, which again add up to almost   words, the two parental allele combinations of these tightly
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              99%. You can see that no preferred association of w  and y   linked genes are inherited together 99 times out of 100.
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              or of y  and w exists in this cross. Instead, a comparison of
              the two experiments with these particular X chromosome   Gene-pair-specific variation in the
              genes demonstrates that the observed frequencies of the   degree of linkage
              various types of progeny depend on how the arrangement
              of alleles in the F 1  females originated. We have redrawn   Linkage is not always this tight. In Drosophila, a mutation for
              Fig. 5.2 as Cross Series A in Fig. 5.3 so that you can make   miniature wings (m) is also found on the X chromosome. A
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              this comparison more directly. Note that in both experi-  cross of red-eyed females with normal wings (w  m  / w  m )
              ments, it is the parental classes—the combinations origi-  and white-eyed males with miniature wings (w m / Y) yields
              nally present in the P generation—that show up most   an F 1  generation containing all red-eyed, normal-winged flies.
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              frequently in the F 2  generation. The reshuffled recombi-  The genotype of the dihybrid F 1  females is w  m  / w m. Of the
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              nant classes occur less often. It is important to appreciate   F 2  males, 67.2% are parental types (w  m  and w m), while the
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              that the designation of parental and recombinant gametes   remaining 32.8% are recombinants (w m  and w  m).
              or progeny of a doubly heterozygous F 1  female is opera-  This preponderance of parental combinations among
              tional, that is, determined by the particular set of alleles she   the F 2  genotypes reveals that the two genes are linked: The
              receives from each of her parents.                   parental combinations of alleles travel together more often
                  When genes assort independently, the numbers of pa-  than not. But compared to the 99% linkage between the w
              rental  and  recombinant  F 2   progeny  are  equal  because  a   and y genes for eye color and body color, the linkage of w
              doubly heterozygous F 1  individual produces an equal num-  to m is not that tight. The parental combinations for color
              ber of all four types of gametes. By comparison, two genes   and wing size are reshuffled in roughly 33 (instead of 1)
              are considered linked when the number of F 2  progeny with   out of every 100 gametes.
              parental genotypes exceeds the number of F 2  progeny with
              recombinant genotypes. Instead of assorting independently,   Linkage of autosomal traits
              the genes behave as if they are connected to each other   Linked autosomal genes are not inherited according to the
              much of the time. The genes for eye and body color that   9:3:3:1 Mendelian ratio expected for two independently
              reside on the X chromosome in Drosophila are an extreme     assorting, noninteracting genes, each with one completely
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