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3.2 Extensions to Mendel for Two-Gene Inheritance   63


                       Figure 3.19  Redundant genes result in a 15:1 phenotypic ratio. (a) Normal maize leaves (AA BB) and a leaf lacking both
                       dominant alleles A and B (aa bb). (b) In maize, either dominant allele A or B is sufficient for normal leaf development. Only the absence
                       of both dominant alleles (aa bb) results in malformed, thin leaves. The result is a 15:1 ratio in the dihybrid cross F 2 .
                       a: © Dr. Michael J. Scanlon, Cornell University
                              (a)  Normal maize leaves and mutants lacking  (b)  A and B are redundant for maize leaf development
                                    two redundant gene functions
                                                                       P             AA bb         aa BB



                                                                       Gametes        A b           a B


                                                                         (all identical)
                                                                       F 1           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–
                                                                            (3)  A– bb (normal)
                                                                    15
                                                                            (3)  aa B–      A b AA Bb AA bb  Aa Bb Aa bb
                                                                     1      aa bb (skinny)
                                                                                            a B Aa BB Aa Bb aa BB aa Bb

                                                                                            a b Aa Bb Aa bb aa Bb aa bb
                                     AA BB              aa bb



                         examples (squash and chicken colors) yield different   malformed only in the absence of both A and B (aa bb),
                       F 2   phenotypic ratios because of differences in the biochem-  the F 2  phenotypic ratio signifying redundant gene action
                       ical pathways involved.                             is 15:1 (Fig. 3.19b). 
                                                                               The proteins (A and B) encoded by the dominant al-
                        ∙  Recessive epistasis usually indicates that the domi-
                          nant alleles of the two genes function in the same   leles act in parallel, redundant pathways that recruit precur-
                                                                           sor cells to become part of the leaf (Fig. 3.20). That is, if
                          pathway to achieve a common outcome. In the      either pathway functions, the leaves will develop their nor-
                          Labrador retriever example, B and E both function    mal broad shape.
                          to generate black hairs.
                        ∙  Dominant epistasis usually indicates that the domi-  Often, as in this case, redundant genes specify nearly
                          nant alleles of the two genes have antagonistic func-  identical proteins that perform the same function. Why
                                                                           does the organism have two genes that do the same thing?
                          tions. Both in the cases of squash and chicken color,   One answer is that redundant genes often arise by chance
                          the dominant allele of gene B prevents deposition of a   evolutionary processes that duplicate genes, as will be
                          pigment whose synthesis depends on the dominant     explained in Chapter 10.
                            allele of gene A.

                                                                           Summary: A Variety of Different
                       Redundancy: One or More Genes in a                  Biochemical Pathways Can Produce
                       Pathway Are Superfluous                             Any Given Altered Mendelian Ratio
                       In maize, two genes, A and B, control leaf development.   So far we have seen that when two independently assorting
                       Normal broad leaves develop as long as the plant has ei-  genes interact to determine a trait, the 9:3:3:1 ratio of the
                       ther a dominant A allele or a dominant B allele (A− B−,    four Mendelian genotypic classes in the F 2  generation can
                       A− bb, or aa B–). However, the leaves of plants that have   produce a variety of phenotypic ratios, depending on the
                       neither dominant allele (aa bb) are skinny because they   nature of the gene interactions. The result may be four, three,
                       contain too few cells (Fig. 3.19a). Given that leaves are   or two phenotypes, composed of different combinations of
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