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60     Chapter 3    Extensions to Mendel’s Laws


              Figure 3.15  Dominant alleles of two genes needed for purple color in sweet peas. (a) White and purple sweet pea flowers. (b) The
              9:7 ratio of purple to white F 2  plants indicates that at least one dominant allele of each gene is necessary for the development of purple color.
              © William Allen/National Geographic Creative
              (a)  Lathyrus odoratus (sweet peas)      (b)  A dihybrid cross showing reciprocal recessive epistasis



                                                       P             AA bb             aa BB

                                                      Gametes         A b               a B





                                                      F 1  (all identical)  Aa Bb      Aa Bb


                                                      F 2                       A B  A b  a B  a b

                                                                            A B  AA BB AA Bb Aa BB Aa Bb  9  A– B– (purple)

                                                                            A b  AA Bb  AA bb AaBb    Aa bb  (3) A– bb
                                                                                                      7  (3) aa B–   (white)
                                                                            a B Aa BB Aa Bb aa BB aa Bb  (1) aa bb

                                                                            a b Aa Bb Aa bb aa Bb aa bb


                                                                   Figure 3.16  A biochemical explanation for reciprocal
                genotype hh for the second gene, do not make substance H at   recessive epistasis in the generation of sweet pea color.
              all. Thus, even if these people make an enzyme that would   Enzymes specified by the dominant alleles of two genes are both
              add A or B to this polysaccharide base, they have nothing to   necessary to produce pigment. The recessive alleles of both genes
              add it onto; as a result, Bombay-phenotype individuals  appear   specify no enzymes. In aa homozygotes, no intermediate precursor
              to be type O. For this reason, homozygosity for the recessive   2 is generated, so even if enzyme B is available, it cannot produce
                                                                   purple pigment.
              h allele of the H-substance gene masks the effects of the ABO   AA, Aa            BB, Bb
              gene, making the hh genotype epistatic to any combination of
               A
                  B
              I , I , and i alleles (except for ii.).                        Enzyme A          Enzyme B
                                       B
                                                A
                                                      B
                  A person who carries I , I , or both I  and I  but is also   Colorless  Colorless     Purple
                                     A
                                                                                                        pigment
                                                                                      precursor 2
                                                                    precursor 1
              an hh homozygote for the H-substance gene may appear to
                                                            A
              be type O, but he or she will be able to pass along an I  or                        bb
               B
              I  allele in sperm or egg. An offspring receiving, let’s say,   AA, Aa
              an I  allele for the ABO gene and a recessive h allele for the                  No enzyme B
                 A
              H-substance gene from the father plus an  i allele and a   Colorless  Enzyme A  Colorless  No purple
              dominant H allele from the mother would have blood type   precursor 1   precursor 2        pigment
                          A
              A (genotype I i Hh), even though neither of the parents is
              phenotype A or AB (Fig. 3.14b).                                  aa
              White sweet pea flowers  In the first decade of the twenti-  No enzyme A          BB, Bb
              eth century, William Bateson conducted a cross between                     No
              two lines of pure-breeding white-flowered sweet peas    Colorless        colorless  Enzyme B  No purple
              (Fig. 3.15a). Quite unexpectedly, all of the F 1  progeny were   precursor 1  precursor 2  pigment
              purple (Fig. 3.15b). Self-pollination of these novel hybrids
              produced a ratio of 9 purple: 7 white in the F 2  generation.    aa                 bb
              The explanation? Two genes work in tandem to produce
              purple sweet pea flowers, and a dominant allele of each      No enzyme A        No enzyme B
              gene must be present to produce that color.           Colorless            No              No purple
                  A simple biochemical hypothesis for these results  is   precursor 1  colorless         pigment
              shown in Fig. 3.16. Because it takes two enzymes catalyzing             precursor 2
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