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62     Chapter 3    Extensions to Mendel’s Laws


              Figure 3.18  Dominant epistasis may also result in a 13:3 phenotypic ratio. (a) In the F 2  generation resulting from a dihybrid
              cross between white leghorn and white wyandotte chickens, the ratio of white birds to birds with color is 13:3. This ratio emerges because
              at least one copy of A and the absence of B is needed to produce color. (b) Enzyme A, encoded by allele A, is needed to synthesize
              pigment. Allele a encodes no enzyme. Pigment deposition in the feathers depends on protein b encoded by allele b, the normal (wild-type)
              allele of a second gene. The mutant dominant allele B, however, encodes an abnormal version of the protein that prevents pigment
              deposition, even when the normal protein b is present.
              (a)  B is epistatic to A                             (b)  Biochemical explanation for dominant epistasis in the
                               White        White                     generation of chicken feather color
                               leghorn      wyandotte                                        BB, Bb
                 P             AA BB          aa bb                         AA, Aa
                                                                                            Protein B
                                                                   Colorless Enzyme A                No pigment
                 Gametes        A B           a b                  precursor        Pigment          deposited

                                                                              aa
                 F  (all identical)  Aa Bb   Aa Bb                                           BB, Bb
                  1
                                                                          No enzyme A
                                                                                             Protein B
                                                                   Colorless         No              No pigment
                 F 2                                               precursor         pigment         deposited
                                           A B   A b  a B   a b
                                                                              aa
                                      A B AA BB AA Bb Aa BB Aa Bb
                      (9) A– B–                                                                bb
              13                  (white)                                 No enzyme A
                      (3) aa B–
                                      A b AA Bb AA bb  Aa Bb Aa bb                           Protein b
                      (1)  aa bb                                   Colorless         No              No pigment
               3      A– bb (colored)                              precursor         pigment         deposited
                                      a B Aa BB Aa Bb aa BB aa Bb
                                                                                               bb
                                                                            AA, Aa
                                      a b Aa Bb Aa bb aa Bb aa bb
                                                                                            Protein b
                                                                   Colorless Enzyme A                 Pigment
                                                                   precursor        Pigment           deposited





                  The squash genes A and B have not been identified at   pathway known to underly the 13:3 ratio for chicken
              the molecular level, and the biochemical pathway in which   feather color is shown in Fig. 3.18b.
              they interact is unknown. However, based on knowledge of
              similar phenomena in other plants, a likely biochemical   Important points regarding epistasis
              pathway underlying the 12:3:1 phenotypic ratio is shown in
              Fig. 3.17b.                                          Several important points emerge  from the examples of
                                                                     recessive and dominant epistasis we have discussed:
              Chicken  feather  color  A variant ratio indicating domi-  ∙  Epistasis is an interaction between alleles of different
              nant epistasis is seen in the feather color of certain chick-  genes, not between alleles of the same gene.
              ens (Fig. 3.18a). White leghorns have a doubly dominant   ∙  In dihybrid crosses, the F 2  phenotypic ratios result-
              AA BB genotype for feather color; white wyandottes are   ing from epistasis depend on the functions of the
              homozygous recessive for both genes (aa bb). A cross be-  specific alleles and the particular biochemical path-
              tween these two pure-breeding white strains produces an   ways in which the genes participate.
              all-white dihybrid (Aa Bb) F 1  generation, but birds with
              color in their feathers appear in the F 2 , and the ratio of   In the Labrador retriever and sweet pea examples of
              white to colorful is 13:3 (Fig. 3.18a). We can explain this   recessive epistasis, the completely dominant alleles of both
              ratio by assuming a kind of dominant epistasis in which B   genes specify normally functional protein, while the reces-
              is epistatic to A; the A allele produces color only in the   sive alleles are either nonfunctional or specify weakly
              absence of B; and the a, B, and b alleles produce no color.   functional protein. Nevertheless, the phenotypic ratios
              The interaction is characterized by a 13:3 ratio because the   among the F 2  of a dihybrid cross differ in the Labradors
              9 A– B–, 3 aa B–, and 1 aa bb genotypic classes combine   and peas because the underlying biochemical pathways
              to produce only one phenotype: white. The biochemical   are not identical. Likewise, the two dominant epistasis
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